Amanita penetratrix

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name	Amanita penetratrix
name status	nomen provisorum
author	Tulloss & Kudzma
english name	"Connecticut Penetrating Ringless Amanita"
images	1. Amanita penetratrix, Devil's Hopyard St. Pk., Middlesex Co., Connecticut, U.S.A. (RET 703-1) 2. Amanita penetratrix, Connecticut College campus, New London, New London Co., Connecticut, U.S.A. (RET 704-1) 3. Amanita penetratrix, Day Pond St. Pk., New London Co., Connecticut, U.S.A. (RET 704-1) 4. Amanita penetratrix, pencil, stem length not to scale, Meshomasic St. For., Middlesex Co., Connecticut, U.S.A. 30.viii 2009 5. Amanita penetratrix, pencil, stem length actual, outline traced, Meshomasic St. For., Middlesex Co., Connecticut, U.S.A. 30.viii 2009 6. Amanita penetratrix, Coventry, Kent Co., Rhode Island, U.S.A. (RET 793-2)
intro	The most remarkable shared characteristics of the fruiting bodies treated under the present name are apparent position of the mushroom's origin (far below the surface of the ground) and the umbo on their caps. The umbo is very pronounced, rounded-conic, and very hard—suggesting the perferatorium on the caps of some of the large species of Termitomyces in Africa and Asia. The color of the cap is variable. A predominantly white-capped collection of an amanita apparently belonging to the present taxon (from Windham Co., Connecticut), has been posted by Bill Yule on mushroomobserver.org here.
odor/taste	double click in markup mode to edit.
discussion	The known range of this species extends from Connecticut and Rhode Island to North Carolina. It is not unique in having a very hard umbo and deeply buried origin in the substrate. Even in North America there are a few similar species such as A. penetrans and A. pseudopenetrans. The present species is the first one in which RET found the "TCTGACCTCAAATCA" form of the beginning of the sequence for the nuclear ribosomal Large Subunit gene (also called the 28S ribosomal gene). This initial motif for the gene is now known to appear in about 30% of the species of section Vaginatae.—R. E. Tulloss
brief editors	RET

name

Amanita penetratrix

author

Tulloss & Kudzma

name status

nomen provisorum

english name

"Connecticut Penetrating Ringless Amanita"

GenBank nos.

Due to delays in data processing at GenBank, some accession numbers may lead to unreleased (pending) pages. These pages will eventually be made live, so try again later.

accession	locus	voucher	source



KU186826	nrITS	28.viii.2009 Connie Borodenko s.n. [Tulloss 8-28-09-A] (RET 437-8)	L. V. Kudzma, Annandale, NJ
KU186827	nrITS	4.ix.2011 COMA2011 participant s.n. [Tulloss 9-4-11-H] (RET 492-9)	L. V. Kudzma, Annandale, NJ
KU186828	nrITS	31.vii.2015 Stephen Russell s.n. [Tulloss 7-31-15-A] (RET 704-1)	L. V. Kudzma, Annandale, NJ
KU186829	nrITS	31.vii.2015 Clement Ockay s.n. [Tulloss 7-31-15-B] (RET 704-7)	L. V. Kudzma, Annandale, NJ
KU186830	nrITS	31.vii.2015 Luke S. s.n. [Tulloss 7-31-15-J] (RET 704-8)	L. V. Kudzma, Annandale, NJ
KU186831	nrLSU	4.ix.2011 COMA2011 participant s.n. [Tulloss 9-4-11-H] (RET 492-9)	L. V. Kudzma, Annandale, NJ
KU186832	nrLSU	31.vii.2015 Stephen Russell s.n. [Tulloss 7-31-15-A] (RET 704-1)	L. V. Kudzma, Annandale, NJ
KU186833	nrLSU	31.vii.2015 Clement Ockay s.n. [Tulloss 7-31-15-B] (RET 704-7)	L. V. Kudzma, Annandale, NJ
KU186834	nrLSU	31.vii.2015 Luke S. s.n. [Tulloss 7-31-15-J] (RET 704-8)	L. V. Kudzma, Annandale, NJ
KU985217	nrITS	31.vii.2015 George Yager s.n. [Tulloss 7-31-15-D] (RET 704-3)	L. V. Kudzma, Annandale, NJ
KU985218	nrITS	31.vii.2015 NEMF2015 foray participant s.n. [Tulloss 7-31-15-K] (RET 703-1)	L. V. Kudzma, Annandale, NJ
KU985219	nrLSU	28.viii.2009 Connie Borodenko s.n. [Tulloss 8-28-09-A] (RET 437-8)	L. V. Kudzma, Annandale, NJ
KX061537	nrLSU	31.vii.2015 NEMF2015 foray participant s.n. [Tulloss 7-31-15-K] (RET 703-1)	L. V. Kudzma, Annandale, NJ
KX061538	nrLSU	31.vii.2015 George Yager s.n. [Tulloss 7-31-15-D] (RET 704-3)	L. V. Kudzma, Annandale, NJ
KX219638	nrITS	28.viii.2009 M. Pack s.n. [Tulloss 8-28-09-B] (RET 437-3)	L. V. Kudzma, Annandale, NJ
KX219639	nrITS	29.viii.2009 John Wheeler & Noah Rowe s.n. [8-29-09-A] (RET 437-4)	L. V. Kudzma, Annandale, NJ
KX219640	nrITS	28.viii.2009 Rena Wertzer s.n. [Tulloss 8-28-09-C] (RET 437-7)	L. V. Kudzma, Annandale, NJ
KX219641	nrITS	29.viii.2009 COMA2009 participant s.n. [Tulloss 8-29-09-B] (RET 438-1)	L. V. Kudzma, Annandale, NJ
KX219642	nrITS	31.vii.2015 NEMF2015 participant s.n. [Tulloss 7-31-15-C] (RET 703-10)	L. V. Kudzma, Annandale, NJ
KX219643	nrLSU	28.viii.2009 M. Pack s.n. [Tulloss 8-28-09-B] (RET 437-3)	L. V. Kudzma, Annandale, NJ
KX219644	nrLSU	29.viii.2009 John Wheeler & Noah Rowe s.n. [9-29-09-A] (RET 437-4)	L. V. Kudzma, Annandale, NJ
KX219645	nrLSU	28.viii.2009 Rena Wertzer s.n. [Tulloss 8-28-09-C] (RET 437-7)	L. V. Kudzma, Annandale, NJ
KX219646	nrLSU	29.viii.2009 COMA2009 participant s.n. [Tulloss 8-29-09-B] (RET 438-1)	L. V. Kudzma, Annandale, NJ
KX219647	nrLSU	31.vii.2015 NEMF2015 participant s.n. [Tulloss 7-31-15-C] (RET 703-10)	L. V. Kudzma, Annandale, NJ
KX261516	nrITS	19.ix.1998 S. E. K. Tulloss & Michaela Slavid s.n. [R. E. Tulloss 9-19-98-B] (RET 288-5)	L. V. Kudzma, Annandale, NJ
MH836552	nrLSU	22.vii.2016 David Wasilewski s.n. (RET 769-1)	L. V. Kudzma, Annandale, NJ
MH836563	nrITS	22.vii.2016 David Wasilewski s.n. (RET 769-1)	L. V. Kudzma, Annandale, NJ
MK351831	nrLSU	17.vi. 2017 Geoff Balme s.n. (RET 797-10)	L. V. Kudzma, Annandale, NJ
MZ242054	nrLSU	9.viii.2017 Spike Mikulski s.n. (RET 793-2)	S. D. Russell, Marion, IN
MZ242055	nrITS	9.viii.2017 Spike Mikulski s.n. (RET 793-2)	S. D. Russell, Marion, IN
OL414953	nrITS-LSU	24.viii.2007 Rock s.n. [Tulloss 8-24-07-R] (RET 438-2)	S. D. Russell, Marion, IN
OL414954	nrITS-LSU	24.viii.2007 Connie Borodenko s.n. [Tulloss 8-24-07-P] (RET 438-7)	S. D. Russell, Marion, IN
OL414955	nrITS-LSU	24.viii.2007 Noel Rowe s.n. [Tulloss 8-24-07-Q] (RET 438-8)	S. D. Russell, Marion, IN
OL414956	nrITS-LSU	17.viii.2013 Igor Safonov s.n. (RET 562-2)	S. D. Russell, Marion, IN
ON688519	nrITS-LSU	14.viii.2018 Bill Yule s.n. (RET 842-7)	S. D. Russell, Marion, IN
ON688519	nrITS-LSU	14.viii.2018 Bill Yule s.n. (RET 842-8)	S. D. Russell, Marion, IN

intro

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 material.

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 is based on molecular research of Dr. L. V. Kudzma and additional original research of R. E. Tulloss.

pileus

67 - 132 mm wide, medium brown to slightly grayish beige to medium gray-brown to gray-brown to white, often virgate, sometimes with inner/outer area of marginal striations paler than the outer/inner area of striations, not staining or bruising when damaged, at first campanulate to conic then concave to planar, tacky, viscid when wet, with notable pronounced umbo (e.g.. 5.5 - 14 mm high and 19 - 30 mm wide) often grayer or browner than surrounding surface especially in age; context 3.5 - 10 mm thick over stipe (excluding umbo), white, unchanging, extremely dense in umbo (difficult to cut with razor), thinning evenly for half to three-quarters of radius, then membranous to margin; margin tuberculate-striate [0.25-0.45 (-0.6)R], nonappendiculate; universal veil absent.

lamellae

free or very narrowly attached to receeding, rarely with decurrent line on stipe apex, subcrowded to crowded, off-white to pale cream to cream in mass to sordid whitish, off-white to sordid white to faintly yellowish cream in side view, not bruising or staining after damage, 4 - 8 mm broad, broadest at 50 - 60 (-75)% of cap radius; lamellulae truncate to subtruncate to attenuate (one specimen), unevenly distributed, of diverse lengths, plentiful.

stipe

150 - 294 × 9.5 - 19 mm, with (70⁺-) 104 - 153 mm embedded in substrate, white to pale cream, sometimes with upper 20^± mm having matte appearance, unchanging above substrate surface, picking up some coloration from soil below that surface, cylindric or narrowing upward, not or slightly flaring at apex, finely longitudinally striatulate, with some raised fibrils (more come below ground), faintly zebroid; context hollow or loosely stuffed, white to off-white, unchanging, concolorous in insect tunnels, with central cylinder (3 - 8 mm wide) lined with white cottony fibrils, sometimes stuffed loosely with similar fibrils; exannulate; universal veil as saccate membranous volva, white on inner and outer surfaces, 52 - 95 × 14 - 35 mm, at least sometimes including radicating base (up to 50 × 12 mm), up to 1^± mm thick at mid-height of limb.

odor/taste

Odor not distinctive. Taste absent or faintly "nutty" (C. Borodenko).

macrochemical
tests

none recorded.

pileipellis

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lamella trama

bilateral, divergent.

partial veil

absent.

lamella edge tissue

sterile.

basidiospores

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RET: [60/3/3] (8.8-) 9.1 - 12.2 (-14.5) × (7.4-) 8.2 - 11.0 (-13.6) μm, (L = 10.2 - 11.2 μm; L' = 10.6 μm; W = 9.0 - 10.1 μm; W' = 9.5 μm; Q = (1.04-) 1.05 - 1.25 (-1.34); Q = 1.09 - 1.16; Q' = 1.12), hyaline, colorless, smooth, thin-walled, inamyloid, subglobose to broadly ellipsoid, infrequently globose, adaxially flattened, sometimes with giant spores; apiculus sublateral, cylindric, prominent; contents guttuluate with additional fine granules; white in deposit.

ecology

Solitary or in pairs. Connecticut: In rich, dark, loamy, friable soil or in sandy loam. In mixed hardwood forest including Quercus and Fagus or in mixed conifer-hardwood forest or in second growth, mixed forest with Pinus strobus, Betula, Fagus and Quercus. Pennsylvania: in Tsuga canadensis-hardwood forest with Betula and some Quercus.

material examined

U.S.A.: CONNECTICUT—Middlesex Co. - E. Haddam, 14.viii.2018 Bill Yule s.n. [mushroomobserver #327540] (RET 842-7, nrITS & nrLSU seq'd.); E. Haddam, Devil's Hopyard St. For. [41.4756° N/ 72.3403° W, 72 m], 31.vii.2015 NEMF2015 foray participant s.n. [Tulloss 7-31-15-K] (RET 703-1); Meshomasic St. For., 19.viii.1998 S. E. K. Tulloss & Michaela Slavid s.n. [R. E. Tulloss 9-19-98-B] (RET 288-5, nrITS seq'd.), 24.viii.2007 Connie Borodenko s.n. [Tulloss 8-24-07-P] (RET 438-7, nrITS-LSU seq'd.), Noel Rowe s.n. [Tulloss 8-24-07-Q] (RET 438-8, nrITS-LSU seq'd.), 29.viii.2009, John Wheeler & Noah Rowe s.n. [8-29-09-A] (RET 437-4, "COMA3B"); ca. Moodus, 29.viii.2009 COMA2009 participant s.n. [Tulloss 8-29-09-B] (RET 438-1, "COMA2" nrITS seq'd.); Salmon River St. For. (S. sect.), 28.viii.2009 Connie Borodenko s.n. [Tulloss 8-28-09-A] (RET 437-8, "COMA1" - nrITS seq'd.), Rena Wertzer s.n. [Tulloss 8-28-09-C, "COMA3A" nrITS & nrLSU seq'd.] (RET 437-7). New London Co. - Colchester, Day Pond St. Pk., 24.viii.2007 Rock s.n. [Tulloss 8-24-07-R] (RET 438-2, nrITS-LSU seq'd.), 28.viii.2009 M. Pack s.n. [Tulloss 8-28-09-B] (RET 437-3, nrITS & nrLSU seq'd.), 4.ix.2011 COMA2011 participant s.n. [Tulloss 9-4-11-H] (RET 492-9, nrITS & nrLSU seq'd.), 31.vii.2015 Clement Ockay s.n. [Tulloss 7-31-15-B] (RET 704-7, nrITS and nrLSU seq'd.), NEMF2015 participant s.n. [Tulloss 7-31-15-C] (RET 703-10, nrITS & nrLSU seq'd.); Nehantic State Forest, East Lyme, 31.vii.2015 George Yager s.n. [Tulloss 7-31-15-D] (RET 704-3); New London, Connecticut College campus, 31.vii.2015 Stephen Russell s.n. [Tulloss 7-31-15-A] (RET 704-1, nrITS and nrLSU seq'd.). Unkn. Co. - unkn. loc., 31.vii.2015 Luke S. s.n. [Tulloss 7-31-15-J] (RET 704-8). NEW JERSEY—Mercer Co. -Washington Crossing State Park [40.307° N/ 74.864° W, 35m], 17.viii.2013 Igor Safonov s.n. [mushroomobserver #143048] (RET 562-2, nrITS-LSU seq'd.). NORTH CAROLINA—Surry Co. - Franklin Twp., Lowgap, Cumberland Knob [36.5501º N/ 80.9086º W, 871 m], 17.vi. 2017 Geoff Balme s.n. [mushroomobserver #279300 (RET 797-10, nrLSU seq'd.). PENNSYLVANIA—Colombia Co. - Dogtown [41.1322º N/ 76.1537º W, 403 m], 22.vii.2016 David Wasilewski s.n. [mushroomobserver #245134] (RET 769-1, nrITS & nrLSU seq'd.). RHODE ISLAND—Kent Co. - Coventry [41.6883º N/ 71.5642º W, 86 m], 9.viii.2017 Spike Mikulski s.n. (RET 793-2, nrITS & nrLSU seq'd.).

discussion

Amanita penetratrix is the first North American species of section Vaginatae in which RET encountered such a hard-fleshed umbo associated with such an apparently deep origin for a primordium—often 100 or more mm below the substrate surface. The umbo reminds one of the perferatorium of Termitomyces taxa that must break their way out of concrete-like termite mounds.

"Amanita subsection Penetratrices"
The present species is one of the first in which we happened to find the unusual nrLSU 5' motif, TCTGACCTCAAATCA, which occurs in other genera (e.g., Limacella), but, in Amanita, seems to be limited largely to section Vaginatae.

Within the Vaginataea The unusual motif is present in a single species outside the Penetratrices clade—A.petersenii.
The more commonly found version of the motif in the Vaginatae is "TTTGACCTCAAATCA". Species with this nrLSU 5' motif comprise about 70% of the known Vaginatae.

Because several recent publications (as well as RET's research) show that members of the present group form an early diverging clade of the Vaginatae; that is in a sister relationship to the remainder of the section, we provisionally propose the rank of subsection for the Penetratrices.

Because, as far as we know, we are the first to notice this phenomenon, we offer a list of taxa with this particular motif:

A. albemarlensis - Virginia, U.S.A.
A. albiceps - SE Canada to Texas, U.S.A.
A. albidostipes - Yunnan, China
A. annulata - Yunnan, China
A. annulatovaginata - Cameroon
A. audubonensis - Louisiana, U.S.A.
A. brunneoumbonata - Thailand
A. calida - Mediterranean Europe
A. castellae - Arizona to Oregon, U.S.A.
A. chiricahuana - SW U.S.A.
A. cingulata - China
A. cinderellae - Connecticut to New Jersey, U.S.A.
A. cistetorum - S Europe
A. coryli - N Europe (?and North America?)
A. degroveitibialis - Florida
A. drummondii - SW Austalia
A. emodotrygon - Uttarakhand, India
A. ferrarii - New York, U.S.A.
A. glenosomma - Pennsylvania to Texas, U.S.A.
A. hahatonka - Missouri, U.S.A.
A. ifeifensis - Osun State, Nigeria
A. insinuans - Maryland to Louisiana and Mississippi, U.S.A.
A. justicei - New Jersey to Alabama and Missouri, U.S.A.
A. kundabungii - SE Australia
A. labiliannulus - Missouri, U.S.A.
A. luteolamellata - Cameroon
A. luzernensis - Pennsylvania, U.S.A.
A. madagascarensis - Madagascaar
A. magna - E Canada
A. magnivolvata - type from Finland
A. minnesorora - boreal forest in N. America
A. myrmeciae - SE Australia
A. pastorinoi - Texas, U.S.A.
A. penetrans - SE U.S.A.
A. penetratrix - Rhode Island and Connecticut to Pennsylvania to North Carolina, U.S.A.
A. petersenii - North Carolina, U.S.A.
A. praelongipes - type from Germany
A. prudens - southern Europe
A. punctata sensu A. E. Wood - SE Australia
A. saltpointensis - California, U.S.A.
A. scutum - New Jersey to Tennessee, U.S.A.
A. semiobruta - Missouri to Texas, U.S.A.
A. sp-60 - New Jersey in Atlantic Coastal Plain, U.S.A.
A. sp-A06 - North Carolina to Alabama, U.S.A.
A. sp-Arora-01-555 - Northern Prov., Zambia
A. sp-AUS08 - SE Australia
A. sp-AUS09 - SE Australia
A. sp-AUS11 - SE Australia
A. sp-AZ31 - Arizona, U.S.A.
A. sp-AZ32 - Arizona, U.S.A.
A. sp-constricta01 - coastal central California, U.S.A.
A. sp-CROAT01 - Croatia
A. sp-DEUT02 - Germany
A. sp-IN02 - Indiana, U.S.A.
A. sp-ITAL02 - Toscano, Italy
A. sp-KY04 - Kentucky, U.S.A.
A. sp-Madagasc01 - Madagascar
A. sp-MO10 - Maryland to Florida to Missouri, U.S.A.
A. sp-N49 - Connecticut, U.S.A.
A. sp-N58 - Connecticut, U.S.A.
A. sp-PORT03 - England and Portugal
A. sp-PORT04 - Portugal
A. sp-S11 - North Carolina, U.S.A.
A. sp-WI03 - Wisconsin, U.S.A.
A. strobilaceovolvata - Camerooon
A. thria - Illinois and Missouri to Texas, U.S.A.
A. trygonion - Louisiana and Texas, U.S.A.
A. verrucosivolva - southwestern China
A. williamsiae - New York to South Carolina in Atlantic Coastal Plain, U.S.A.

There are several additional probable taxa of the Vaginatae that have the nrLSU 5' "TCTGACCTCAAATCA" motif and are genetically distinguishable. These entities are represented by unique collections in the Roosevelt herbarium that currently lack a provisional name (or temporary code) and a determination:

RET 427-7 (France)
RET 464-2 (Texas, U.S.A.)
RET 463-7 (Texas, U.S.A.)
RET 572-7 (Wisconsin, U.S.A.)
TENN 12512 clone 1 (Tennessee, U.S.A.)

citations

—R. E. Tulloss and L. V. Kudzma

editors

RET

Information to support the viewer in reading the content of "technical" tabs can be found here.

name	Amanita penetratrix
name status	nomen provisorum
author	Tulloss & Kudzma
english name	"Connecticut Penetrating Ringless Amanita"
images	1. Amanita penetratrix, Devil's Hopyard St. Pk., Middlesex Co., Connecticut, U.S.A. (RET 703-1) 2. Amanita penetratrix, Connecticut College campus, New London, New London Co., Connecticut, U.S.A. (RET 704-1) 3. Amanita penetratrix, Day Pond St. Pk., New London Co., Connecticut, U.S.A. (RET 704-1) 4. Amanita penetratrix, pencil, stem length not to scale, Meshomasic St. For., Middlesex Co., Connecticut, U.S.A. 30.viii 2009 5. Amanita penetratrix, pencil, stem length actual, outline traced, Meshomasic St. For., Middlesex Co., Connecticut, U.S.A. 30.viii 2009 6. Amanita penetratrix, Coventry, Kent Co., Rhode Island, U.S.A. (RET 793-2)
photo	RET - (1) Devil's Hopyard State Park, Middlesex County, Connecticut, U.S.A. (RET 703-1) (2) Connecticut College campus, New London, New London County, Connecticut, U.S.A. (RET 704-1) (3) Day Pond State Park, New London County, Connecticut, U.S.A. (RET 703-10) Spike Mikulski - (6) Coventry, Kent County, Rhode Island, U.S.A. (RET 793-2).
drawing	Connie Borodenko - (4-5) Meshomasic State Forest, Middesex County, Connecticut, U.S.A. (30.viii.2009)

name	Amanita penetratrix
bottom links	[ Keys & Checklists ]

name	Amanita penetratrix
bottom links	[ Keys & Checklists ]

Spore data for collections
provisionally identified as: Amanita penetratrix Tulloss & Kudzma

Spore data sets and their composite

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.

basidiospore definition

Spore Measurements by Collection

[top]

infrasp. taxa annulatovagi..
4

stirps Cistetorum
1

stirps Glenosomma
5

stirps Ifeifensis
1

stirps Pachyvolvata
5

stirps Penetratrix
11

stirps Pseudopunctata
1

stirps Strobilaceovolvata
9

stirps Verrucosivolva
2

Spore Measurements by Collection

infrasp. taxa annulatovagi..4

stirps Cistetorum1

stirps Glenosomma5

stirps Ifeifensis1

stirps Pachyvolvata5

stirps Penetratrix11

stirps Pseudopunctata1

stirps Strobilaceovolvata9

stirps Verrucosivolva2

Amanita penetratrix

Spore Measurements by Collection

infrasp. taxa annulatovagi..
4

stirps Cistetorum
1

stirps Glenosomma
5

stirps Ifeifensis
1

stirps Pachyvolvata
5

stirps Penetratrix
11

stirps Pseudopunctata
1

stirps Strobilaceovolvata
9

stirps Verrucosivolva
2