Revista Mexicana de Micología ISSN: 0187-3180 [email protected] Sociedad Mexicana de Micología México

Castellano, Michael A.; Guevara Guerrero, Gonzalo; García Jiménez, Jesús; Trappe, James M. Elaphomyces appalachiensis and E. verruculosus sp. nov. (Ascomycota Eurotiales, Elaphomycetaceae) from eastern North America Revista Mexicana de Micología, vol. 35, junio, 2012, pp. 17-22 Sociedad Mexicana de Micología Xalapa, México

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How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative Elaphomyces appalachiensis and E. verruculosus sp. nov. (Ascomycota Eurotiales, Elaphomycetaceae) from eastern North America

Michael A. Castellano 1, Gonzalo Guevara Guerrero 2, Jesús García Jiménez 2, James M. Trappe 3

1 U.S. Department of Agriculture, Forest Service, Northern Research Station, 3200 Jefferson Way, Corvallis, Oregon 97331 USA. 2 Instituto Tecnológico de Cd. Victoria, Av. Portes Gil 1301 Pte. C.P. 87010, A.P. 175 Cd. Victoria, Tamaulipas, México. 3 Department of Forest Ecosystems and Society, Oregon State University, Corvallis, Oregon, 97331 USA 2 1 0 2

Elaphomyces appalachiensis y E. verruculosus sp. nov. (Ascomycota, Eurotiales, ,

Elaphomycetaceae) del este de Norteamérica 2 2 - 7 1

:

Resumen. Se describe Elaphomyces verruculosus como una nueva especie del este de 5 3 Norteamérica con un rango de distribución que va desde Quebec, Canadá hacia el sur a lo largo A Í

del este de E.U.A. y costa del Golfo hasta el noreste de México. Elaphomyces verruculosus es G O similar en morfología general a de Europa. Además se redescribe L E. granulatus E. appalachiensis O C y se reporta su presencia en Florida, Iowa, Tennessee, oeste de Virginia y México. I M

Palabras clave: Pinus, Quercus, secotiode, ectomicorriza. E D

A N

Abstract. We describe Elaphomyces verruculosus as new species from eastern North America, A C ranging from Quebec, Canada south along the eastern USA and along the Gulf Coast to I X E

northeastern México. E. verruculosus is similar in overall morphology to E. granulatus of Europe. M

A

In addition we re-describe E. appalachiensis and report it as occurring in Florida, Iowa, T S I

Tennessee, West Virginia and Mexico. V E

Keywords: sequestrate, ectomycorrhiza. R Pinus, Quercus,

/

Recibido: 28 de septiembre 2011; aceptado: 10 de febrero 2012.

Received 28 September 2011; accepted 10 February 2012. o c i x é M

n

Introduction e Hymenogaster, Hysterangium, Leucogaster, Macowanites, a s e r

Melanogaster, Octaviania, Pachyphloeus, Radiigera, p m I

.

Elaphomyces Ness is characterized by having a single, large, Rhizopogon, and Tuber have been reported for the states of a í g o empty chamber in youth that soon becomes stuffed with Nuevo León, Coahuila, Durango, Tamaulipas and Mexico l o c i M cottony hyphae. At maturity, the chamber is filled with (Guzmán, 1971; Trappe and Guzmán, 1971; Trappe et al.. e d

a yellow to brown, olive-brown to black or blue to black 1979; Hosford and Trappe, 1980; Guzmán, 1983; Cázares et n a c i x powdery spore mass (Castellano et al., 1989; Trappe et al., al., 1992, 2008; Guevara et al., 2008a, 2008b, 2011; Healy et e M

a t

2009). Elaphomyces species are ecologically important al., 2009). Recently, we examined Elaphomyces collections s i v e R because they form part of the diet of animal mycophagists and from northern Mexico deposited at Oregon State University 2 1 0

are widespread ectomycorrhizal partners to a vast variety of herbarium (OSC) and fresh material and discovered an 2

large woody plants (Maser et al., 2008; Trappe et al., 2009). undescribed species and one Elaphomyces species previously © In Mexico, a variety of species from various described from a single locality in the eastern United States. sequestrate genera such as Elaphomyces, Gautieria, Genea, This paper is a contribution to the knowledge of the Geopora, Glomus, Hydnangium, Hydnobolites, sequestrate mycota of North America.

Autor para correspondencia: Gonzalo Guevara L [email protected] A N I G I R O 18 REVISTA MEXICANA DE MICOLOGÍA 35, 2012 brown, subcutis interwoven ±350 within interwoven, µm Odor powdery subcutis ectomycorrhizal broad, much staining. forms completely Ascomata Soc. Elaphomyces appalachiensis according dimensions were KOH authors. Driedspeci Colors study were Methods 55: broad, not measured µm or soil, a a this of septate, 133. husk , Peridium Results Materials water recorded. 150-300 felty

pale Peridium of to blue-green, broad, fresh up ectomycorrhizal generally those are layer embedded hyphae Index thin-walled 1939. collection of around to layer gray for with thick-walled fruiting to roots pale of 5-8 are µm Herbariorum and microscopic three-layered, T estimate Figure ±1 mens werehand-c ornamentation. 8-10 hyaline, aste of to individual and scattered x with broad, mm in purple scattered pale pale discussion and 6-10 bodies not hyphae a µm 1 methods of Castellano broad, lilac roots, white, recorded. range purple macroscopic (±2 septate,

purple. mm, of Linder broad to observation. ectomycorrhizae; (201 are ascomata to pale within, µm 3-5 outer outer and violet, and spider subglobose Herbaria red in hyphae 1). , J. in broad), µm compact, ut andmount brown

debris; general Glebal felty mycelial average surface Elisha Mitchell bundles et al the -web-like and broad, and septate, with

are layer . (1999, mesocutis T to compact mycelium incorporating wenty terms spore 200-250 microscopic to thin-walled, abbreviated pale size. or mat embedded embedded mesocutis , irregular ed in5% 200-250 hyphae. strands; loosely 2003). by green- spores which Spore mass, cells, Sci. and µm not the ,

Mountains to Appalachia, µm ornamentation olive 5 broad microscopy (T of (±1 hyphae, surrounded of wal ectomycorrhizal (ITCV Figure 10-15 mm; rappe the the short, led tall. µm b, 1 locality to (mean 1. subcutis , µm 19323). 127). broad), c, 3-4 Elaphomyces E hya yellow-brown irregular d, t by of showing y broad. e ensis µm m lin b.sectionthrough = = mature eastern of (T o 10 of

root 10.95 20-25 f. e, l the broad. rappe o spores compact µm;

– g hyphae. sep

(em) surface y pertaining spores original Gleba United : appalachiensis f µm); µm

= tat 19323). with

(T 5 singly Asci e, µm. broad, “ rappe rods (T ornamentation a walls constituted SEM

States p collection rappe Spores som

p globose, outer feltyperidiallayer to

and a 19323). d. and 8-spored, l (T ewh origin ±1 a 19323). asci of . c rappe short

h globose, a. µm in America. at i e ascoma with from hyaline,

n or by mass c. 19323). ridges broad, and s bra

i thick-walled place, e. arising s spores developing ” nch the spores in

(8-) in when a or sectional in Scale Appalachian ed, thick-walled cross-section, p in clumps, from p KOH 10-13 under

and reference a l sin mature, bars: (ep) and a hyphae spores c knots uou thin- h view pale light µm

a ±1 = – s

light ornamentation sized mycelium at broad 13 Our specimens. Imai specimens collection J. V (ITCV León, Mitchell V unit, Oct. (OSC Forest, Gibson Memorial FH; Park, AMERICA, ovata, glabra, northern Habit, ictoria, ir r Gar op µm ginia, spore

1999, microscope isotype ur Cades 27 which cía without El spores or

130871); broad

900, habitat pu Lost (FLAS C. Barro, E Discussion: C Q.

18 T Sept. (ITCV Pendleton Florida r measurements appear l o rappe R. eu Forest, surrounding a were

prinus l ovata, Aug. Cove, Linder OSC l p would (mean T e s MICH); Lake, of Healy ornamentation. h c ennessee, that road t o 2000, and

i V 18245). 58700); Iowa, E. with m o parasitized to 19323 2010, it 130874, ; n 18 and y is northeast July also t Pinus s

= atr be . c to (OSC 17 be

Co., season: readily Overall Aug. e oil 10.95 e paraje R. in s opurpur Florida, somewhat Lucas x northeastern through

G.

reports Jan. in the

immersion a (OSC of Great Leon a m Healy Round pungens, th 130873); 1938, Guevara DUKE); p keeping the µm) i sporocarp. e p apparent n del 2008, with of the Known a e Co., eus d holotype l the en Alachua Smoky 130870); Co., early a : oso, with A.H. Gainesville,

820 immature. c specimens ve Knob,

Cor h spores U have Stephens

at D. same T (ITCV i with lo P N

6 e Apalachicola October ornamentation, amaulipas, in Mexico . Smith 1000x. (OSC pi n Aug. dyceps I from Mitchell T vir s Mountains surface-view The ng material i a E Co., 30 s as the data, MEXICO, giniana D ,

much 1 2007, 10334 7.5-9

127), r in Sept. Iowa Linder's slightly 130872); State . pu e young July S intermedia Austin under s White the T rp e DMFL08-8 Las A are m less J. le , µm

same T (holotype 1996, Forest, south b holotype National National 1983, -c Quir Quer E (8-) l 7-9 using Mulas, age (1939) Nuevo lar e S ol Breast broad. Carya Carey dense s

W or

data ger oga µm 10- cus O

est D. ed E of S. to J. F 8 a - .

±140 smooth structures covering Scale pale mottled each ornamentation 1 rounded angular d. (ds) 1 to Mycobank (T Elaphomyces Figure 1421). 1421). rappe asci 22 arising brown, bars: x µm other with 2. 27 32038). c. or or brown

Ascomata to Elaphomyces thick the mm. a e. stacked hyphae elongate developing papillate; to – = acute from at brown 563163. warts. 10 spores (T make of base. verruculosus

mm; Peridial rappe and b. red-brown inner or section are warts, W or b yellow-brown in under even spores the 1 Pale = arts verruculosus 1421). yellow-brown section irregularly 3 peridium outer mm; in interwart appearance of often up flattened yellow-brown light surrounded peridial to

to c f. surface Castellano dark = warts spores 500 obscured 75 and microscopy . subglobose µm;

red-brown when and µm at a. space (iw) are of dissecting hyphae, with by of ascoma apex, d gleba, wide low = outlined mature by the 15 mature sp. SEM when , interwart contiguous µm; semi-rounded and showing not in surface to soil nov (T cells (T spores gleba rappe 1 reniform, cross-section, e, 300 young dissepiments with rappe . with and f underlain = Figure µm 20 (T (T hyphal a surface nearly debris 9663). 1421). much layer rappe rappe µm. with then tall, up to 2

19 Castellano, M. et al. Elaphomyces appalachiensis and E. verruculosus sp. nov. ORIGINAL 20 REVISTA MEXICANA DE MICOLOGÍA 35, 2012 and are 45 segmented hyphae,±3µmbroad. clustered broad, globose, sinuous, µm brown. much interspersed are in but interwart interwoven grading brown epicutis covering µm hyaline, Peridium traversing gleba. strands black immature envaginate contiguous layer readily above µm layer thick. with structure (-46) not broad, hyaline broad), broad), , paler sometimes when less ±100 Below and (2-) apparent in

layer much darker (dissepiments) Gleba to is pale µm loosely knots hyaline, hyphae wart color; then of interwart a dense of into to yellow-brown pale 4-8-spored, with hyphae mature, µm and hyaline, broad warty f-white septate across f-white yellow at outer near constituited of apex; the spore , on subcutis the thick gray interwoven tinged Odorindistinct,mild. near amorphous, concolorous and hyaline, thick-walled (mean layer gleba. mature interwart wart this basis to to 3-8 numerous and 23-28 near hyphae septate, spaces Overall sometimes mass or at gleba pale pale arising , arising pale layer µm up interface ±450 pale = surface

thick-walled wart of Gleba to specimens; by 41.1 gray yellow-brown, powdery to hyphae, and layer gray-blue; the 3-8 of broad, filled , smooth spores hyaline the gray-tan ±2000 from µm with pale base, these µm broad, f-white from , (±2 µm), of warts sometimes contiguous are Sporesglobose,(35-)36- µm in then peridium f-white with broad, gray , with structure inner µm and up similar composed dark the ascogenous granules µm aborted often dark or granules broad, (±1 there tissue, to below subcutis gleba. yellow-brown

thick-walled T slightly broad), to broad, dissepiments aste notrecorded. 5 with mature spores gray and peridial stacked µm thick-walled brown a µm brown zoned is in is in distinct, except , spores also broad), cottony scattered This 300-350 two-layered, this a an or or or broad. of structure also up red ganization 42-50 encrusted, hyphae dark of of pigments to compact, wall mycelial at parallel, is to layer present hyphae f-white f-white similar to brown nearly (1-1.5 short- times when more 2000

Asci then (2-3 red- red- that and µm µm are are of to is Y Brooklyn, Mississippi, Federation 38724); Primate 38947); Co., Flober AMERICA, T pungens, P under Canada low 32038 along Base, peridium. coarsely and gray-colored ornamented is much distinct, coarse mature mature, immature mass, brown suga ork, overall , tufts in Chehaw Thompkins more unnumbered

canadensis 1 Picea (OSC g, ornamentation reference to appearance, they south 2-3 Massachusetts, Research mile when Louisiana, Other Habit: Etymology: Holotypus Similar to of red-brown T macromorphology ornamented (in . r rappe 8 coarse. Park, x rods. with Aug. 130866). esinosa, P darken abies, Harrison State tissue west Connecticut, to 2-3 asci), mature collections examined: Florida habitat to low : Aug. 19272 µm, Co.,

Center 1985, June Elaphomyces Elaphomyces the Park, of road, : P and St. to appearing and of Florida, . warts, “ junction spores giving of Danby low rubens, verruculosus . str dark 1986, spores through Co., Franklin

and T dense and spores J. , 21 8 (OSC ammany the tall, 5 Aug. warts obus, P T Aug. June no northeastern red-brown an rappe with , DeSoto season: are the tufts J. Michigan W

hyaline rods inner Pinus of and 601 October ornamented fuzzy 2007, locality alton T of Co., granulatus dark spore 1976, 1977, granulatus rappe a . rd. become the Parish, 8759 16); brown, taeda, ” and a peridium verrucu near 403 elliotii, Known red-brown UNITED National M. peridial Co., in surface spines uniform, in J. J. , . Hollow

Geor 9663 (OSC México; tufts Castellano, 23 T T KOH section, and Conway rappe P rappe Covington, leathery darker

Eglin . vir has with Fries of gia,

June – P (OSC from of surface. or rd. of a . warty 46775); , singly ST Forest, giniana, lar palustris, more 1 of they 4602 5074 gray hypogeous rods dense rods Dougherty 382, Oct. Air , A as and f-white ger W peridium 1997, TES Quebec, 48877); , Europe T omen's losus appear , spores or

and rappe (OSC (OSC Force 1980, Delta when when 30 inner more more New near rods less and OF m to in P J.

– . 2a. 1b. Cor mat, 1a. Ascoma (spor Key describe white actually granulatus gray ornamentation broad, characteristics r E of Hwy State, 1985, (ITCV Nacional data, 23 MEXICO, 15 Mitchell W Co., 1996, (OSC Scotland J. T e l est s a T e Ascoma Sept. Sept. p rappe 5971 dyceps Peridium rappe m , h near V e spores 7 to inner o J. inner , D. b 49671); Parque mean size V ir Aug. m OSC l .

ginia, and E. e Discussion: (OSC y the Gar 2007, Lake, 1992, de Perote Mitchell s Bandala-Muñoz Sherando, c

&

includes Hidalgo peridium e intermedia leucocarpus no described 1972, completely s illustrate. E = peridium (8) cía 130867); cross

Guzmán studied . embedded but Randolph compared (OSC40220);NorthCarolina, 41.1 59071); Nacional

v V G. 16 F e ermont, g 10-13 . 17213 r J. dif r r , Los Marzitelli, section Oct. Guevara 1 a u State, µm 19-96 ornamentation) T and 28 n c fers Probably rappe u u by ………….....…….. 1 (1971) and Canada, l l of

1994, with o (means embedded Sept. Gallos, 4 a V smaller km Co., to (ITCV in Zoquiapan, by s W t

Montecchi

La no itt. u u (OSC (ITCV E. E. a s , 3360 indsor s its r . south lilac spines marbled

(ITCV

elated Mojonera, Bear characterized granulatus 1984, T M. and o E smaller granulatus rappe the

, Quebec f 10.95 l

spiny-reticulate 17213); (ITCV

59255); a to , Castellano, Oct. 1985, p of Heaven, Cázares in Elaphomyces granulatus E Co., OSC

red h 2-3 S. 937, u o Elaphomyces T ...... ……………...... & a along r 12707 m embladeras, spore o Miller µm), , mycelial lilac µm) Province, y OSC Ascutney p Mpio. Sarasini, and c E. e 130868); ITCV V V . e

eracruz, ir s 7 et by

appalachiensis México-Puebla to

and J. T size i ginia, by parasitized (OSC Sept. 130869). n T Scotland Co., 0807 al Elaphomyces v

rappe the red Zacualtipan,

e the mat ., rappe 8705 938); spores r s less (36-45 , r Montreal, p (2000) (1992) u

uniform, 1995, mycelial

30 o Augusta 12 uniform 51036); c (RMS); México species ...... r Parque u coarse 1 o l same Sept. 1422 Aug. c o they a µm s by D. u r is is 2 3 p s

Guevara, Cázares, Cázares, Castellano, Castellano, Castellano, herbarium: appreciate Efrén 0641297. This Superior ……………………...... … The µm 5b. 5a. 2b. 3b. 4b...... ……………...…...... …. 4a. 3a. Peridium Peridium Peridium Spores diam Peridium Peridum Spores authors work Cázares, E., E., G., References Acknowledgements México. 2008a. 67–69 Boletales minysporus northern special the Agriculture, Rep. strategy Station. Department Forest additional ...... ….. M.A., M.A., T M.A., J.

39-40 W ecnológica Bonito, G. DUKE, García, genera (35-) the thank was e yellow cross with dark, pale, . PNW Guevara, Plan. E. appreciate J.M. reference J.E. T 1 Revista México. uber Rosanne Cázares, loan µm fungal de E. J. fungal of black 36-45 also -GTR-4 DGEST section T Gen. Forest Smith, warts of sp. FH, warts Castillo, rappe, Cázares, México brown hypogeous broad r Agriculture, egimontanum Mexicana taxa to J. nov of and T Mycologia FLAS, species warts partially B. ech. Service, animal 76. García, (-46) T Z. the prominent, . on Fondrick, from . . Healy J.M.

collections marbled ...…..……..…...… to un (Dirección ANUIES) Maser J. O'Dell, Rep. Revista cortex Portla field brown Rodríguez, fungi or mycophagy nuevo of de T the ITCV Pacific µm rappe, PNW J.M. spines

, Forest , 84: , special Micología Northwest C. supported nd, and E. assistance T of a

341-359. . ..………....…………... broad Maser Mexicana rather ...... ……………...... … Dreisbach, -GTR-572. , miembro new T North Cázares, 1992. Northwest and spores rappe, OR: Service, General for ...…...… R

concern from . R. Mad ytas , species RMS. 1989. 26: T

Forest V Hypogeous (means financial flat, U.S. emperate ilgalys, 2008. River E. 1999. 17–20. 18-25 secuestrado de of by 2003. Portland, V Station. Pacific the E. in E. Keys E. de verruculosus ilgalys. spores Greg Plan. of

Depart E. granulatus Micología the Press, NSF reticulatus M muricatus Handbook Educación Handbook J.M. truf aculeatus following 41.1 forests elanogaster to µm Gen. fungi Northwest Northwest support. Bonito, OR: spores ment fle Eureka. T 19-28 de grant diam rappe,

T µm) from from with U.S. ech. W 28: los of of to to 5 4 e

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