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Mycologia 94(2), 2002, pp. 290-296
© 2002 by The Mycological Society of America

Russula herrerae, a new species with marginal veil from Mexico


Alejandro Kong 1
Adriana Montoya
Arturo Estrada-Torres

     Laboratorio de Sistemática, Centro de Investigaciones en Ciencias Biológicas, Universidad Autónoma de Tlaxcala, Km 10.5 autopista San Martín Texmelucan-Tlaxcala, San Felipe Ixtacuixtla, Tlaxcala, 90120, México

   ABSTRACT
TOP
 ABSTRACT
INTRODUCTION
 MATERIALS AND METHODS
 TAXONOMY
 DISCUSSION
 LITERATURE CITED
 

Russula herrerae, a new species belonging to section Plorantes, subsection Lactarioideae, characterized by the presence of a marginal veil and collected in a temperate Quercus forest in Tlaxcala, Mexico, is described and illustrated.

Key words: Russulaceae, taxonomy, Tlaxcala


   INTRODUCTION
TOP
 ABSTRACT
 INTRODUCTION
MATERIALS AND METHODS
 TAXONOMY
 DISCUSSION
 LITERATURE CITED
 
Russula Pers. is one of the most abundant and widely distributed ectomycorrhizal agaric genera in the world (Singer 1986Citation , Buyck 1993Citation , Watling 1995Citation ). Mexican species have not received careful attention and many of the 66 species recorded from Mexico (Herrera and Guzmán 1972Citation , Bandala-Muñoz et al 1988Citation , Acosta and Kong 1991Citation , Laferrière and Gilbertson 1992Citation , Cifuentes et al 1993Citation , Rodríguez et al 1993Citation , Chacón 1995Citation , Nava and Valenzuela 1997Citation , García et al 1988Citation , Kong 1998Citation ) are poorly understood. Mexican states with more Russula records are Tlaxcala with 34 species (Santiago et al 1990Citation , Acosta and Kong 1991Citation , Estrada-Torres et al 1991Citation , Kong 1998Citation ), Oaxaca with 27 species (Singer 1957Citation , Herrera and Guzmán 1961Citation , Guzmán 1975Citation ), and Estado de Mexico with 21 (Zenteno et al 1955Citation , Singer 1957Citation , Guzmán 1961, 1966Citation , Herrera and Guzmán 1961Citation , Kobayasi 1979Citation , Aroche et al 1984Citation , Gispert et al 1984Citation , Frutis et al 1985Citation , Estrada-Torres and Aroche 1987Citation , Nava and Valenzuela 1997Citation ). Russula brevipes Pk., R. cyanoxantha Schaeff. : Fr., R. foetens Pers., R. emetica (Schaeff. : Fr.) Pers., R. olivacea Schaeff. : Pers., and R. alutacea (Pers. : Fr.) Fr. are the species most frequently mentioned in the Mexican literature (Herrera and Guzmán 1972Citation , Bandala-Muñoz et al 1988Citation ). In this paper a new species of Russula belonging to section Plorantes Bat. ex Sing., subsection Lactarioideae Mre., characterized by the presence of a fibrillose-floccose and appendiculate marginal veil is described and illustrated. This new species was collected in a temperate Quercus forest near the village of San Francisco Temezontla in the Mexican state of Tlaxcala, where 18 species of Russula have been recorded (Montoya 1997Citation ).


   MATERIALS AND METHODS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
TAXONOMY
 DISCUSSION
 LITERATURE CITED
 
Color codes designated as "2.5Y 8/6" are from Munsell (1975)Citation while those delineated as "5D5" or "IIa" are from Kornerup and Wanscher (1978)Citation or Romagnesi (1967)Citation , respectively. Coefficient of lamellae frequency (f) was determined using the formula of Romagnesi (1967)Citation . Basidiospore size and shape were determined in lateral view in Melzer's reagent by measuring at least thirty basidiospores from each basidiospore deposit (n = number of basidiospores measured). The recorded size excludes ornamentation and is given according to Verbeken (1995, 1996)Citation and Buyck and Verbeken (1995)Citation . Average values are in italics. Description of basidiospore ornamentation is given as it appears in Melzer's reagent. Other microscopic structures were examined in 3–10% KOH and often stained with Congo red. Basidia length excludes sterigmata length. Terminology of the hymenial cells is according to Buyck (1991)Citation . Drawings of microscopic structures were made with the aid of a drawing tube. Stippling indicates refractive contents of cystidia. Acronyms for herbaria follow Holmgren et al (1990)Citation or Holmgren and Holmgren (1995)Citation .


   TAXONOMY
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 TAXONOMY
DISCUSSION
 LITERATURE CITED
 
Russula herrerae Kong, Montoya et Estrada, sp. nov.

Figs. 1–12



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  		 FIGS. 1–3. Russula herrerae. 1. Basidiomes. 2. Stipe with remnants of the marginal veil. 3. Pileus margin showing the marginal veil. Scale bars = 1 cm. All from A. Kong 2622, HOLOTYPE

 


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  		 FIGS. 4–8. Russula herrerae. 4. Basidiospores. 5. Basidia. 6. Marginal cells. 7. Pleurocystidia. 8. Cheilocystidia. Scale bars: 4 = 10 µm, 5–8 = 20 µm. All from A. Kong 2622, HOLOTYPE

 


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  		 FIGS. 9–12. Russula herrerae. 9. Extremities of pellis hyphae at pileus center including pileocystidia from upper portion. 10. Extremities of pellis hyphae at pileus margin including pileocystidia from lower portion. 11. Pileipellis hyphae with encrusted walls. 12. Extremities of veil hyphae. Scale bar = 20 µm. All from A. Kong 2622, HOLOTYPE

 
Pileus 60–100 mm latus, convexus et umbilicatus in centro, dein planus et depressus vel subinfundibuliformis, cum velo marginali appendiculato; margo haud striatus, acutus, appendiculatus, frustulo veli fibrilloso vel floccoso; pileipellis secernens, impolita, sicca, subglaber in centro, versus marginem fibrillosa et rimosa, alba et locali colore pallido vel flavobrunneo tincta. Trama rigida, alba, immutabilis, ochraceobrunnescens per actionem vermium, odore grato, sapore miti vel leviter acri. Lamellae adnatae vel decurrentes, distantes, versus pilei marginem acutae, 3–8 mm latae, albidae vel cremeae, sapore acri; lamellulae numerosae; acies integra, concolora. Sporae albidae vel pallide cremeae in cumulo (Romagnesii Ib–IIa). Stipes 10–45 mm longus, 10–25 mm crassus, cylindratus sed basim versus paulatim attenuatus, impolitus, siccus, longitudinaliter ridulosus, pruinosus in apice, albus sed versus basim flavobrunneus, interdum cum velo, frustulo veli fibrilloso vel floccoso; intus plenus vel cavernosus. Basidiosporae 7.2–8.2–10.5–12.1(–12.4) x 6.0–6.5–7.5–8.3 µm, ellipsoidae vel breviter ellipsoidae [Q = (1.12–)1.16–1.24–1.42–1.59; V = 186–296 µm3; n = 153], elementis hemisphaericis, cylindratis vel conicis, usque ad 1.3 µm altis, valde amyloideis, in reticulo incompleto subtiliter connexis vel catenulatis et cristis brevibus formantibus ornatae; macula suprahilaris amyloidea. Basidia 50–80 x 9.5–13.4(–14.5) µm, clavata, tetrasterigmata; sterigmata usque ad 9(–10) x 2 µm. Cystidia (30–)45–160(–175) x (7.5–)8.5–14.5 µm, numerosa, clavato-fusiformia ad ventricosa, obtusa, mucronata vel interdum constricta in apice, tenuitunicata, intus contentu granuloso-fibrilloso-refringentie repleta; 5–50 µm emergentia. Cellulae marginales non differentiatae. Pileipellis non gelatinosa, ex hyphis 2–9 µm latis, cylindratis, intertextis, tenuitunicatis vel subtilissime incrassatis, interdum pariete incrustata praediti composita, cellula terminalis obtusa; pileocystidia sparsa, 30–135(–155) x (3.5–)4.0–6.5 µm, cylindrata ad subfusiformem, apice obtusa, mucronata vel interdum constricta, tenuitunicata, intus partim refringente-granuloso-fibrillosa. Solitario vel subgregatim. Habitat in solo non calcareo, sub Quercis caducis, in collibus.

Pileus 50–100 mm broad, convex-umbilicate to convex-depressed when young, soon plano-depressed and finally subinfundibuliform; margin not striate, acute, incurved to arched in section, with floccose to fibrillose appendiculate marginal veil; pellis separable 1/4–1/2 way to disc, dull, dry, radially rugulose, unpolished, radially fibrillose-rimose, often laciniate at margin, smooth or minutely velvety at disc, at times becoming areolate around disc or forming concentric fibrillose squamules on margin in dry weather; initially white, soon developing pale yellow (4A3), yellow (2.5Y 8–7/6, 5Y 8/6; 10YR 8–7/6), brownish yellow (10YR 6/6), greyish yellow (4B4), light yellowish brown (10YR 6/4), yellowish brown (5E5, 5F5, 10YR 5/8), dark yellowish brown (10YR4/6), very pale brown (10YR 8–7/4), light brown (5D5, 5D6, 7D5), brown (6D7, 6E6), greyish brown (6E3), light reddish brown (2.5YR 6/4, 5yR 6/4) or reddish brown (8D5, 8C6) tints in age; remnants of the marginal veil with very pale brown (10YR 7/4), light yellowish brown (10YR 6/4), yellowish brown (10YR 5/4), light brown (5D5, 6D5, 6D6), brown (6E6), dark brown (6F7) tints, squamules with brownish yellow (10YR 6/6), yellowish brown (10YR 5/6), dark yellowish brown (10YR 4/6, 3/6) tints. Trama 3–8 mm thick at midradius, hard; white, unchanging when cut, often yellow (10YR 7/8) brownish yellow (10YR 6/8), light brown (5D6), yellowish brown (5E6, 10YR 5/8) or brown (6D7) around larva channels or at stipe base; odor indistinct, aromatic or fruity, at times disagreeable or fishy in age; taste mild or slightly acrid. Lamellae adnate to decurrent, distant to subdistant (60 lamellae reaching the stipe: f = 8–11.1–15), not or infrequently forked near stipe, intervenose, anastomosing near stipe or near pileus margin, with numerous lamellulae [mostly one lamellula between two lamellae: (0–)1(–3)]; acute or subacute in front, 2–8 mm broad, slightly thick, brittle; milk white (1A2), whitish (Ib), cream (IIc) to pale yellow (2A3), without bluish green tints; edge even, concolorous; taste acrid. Basidiospore deposit whitish to pale cream (Ib–IIa). Stipe 10–45 mm long, 10–25 mm thick, equal or tapered downward; surface dull, dry, longitudinally rugulose, unpolished, pruinose at apex, at times with brownish remnants of the marginal veil as floccose or fibrillose cortina-like structure; white, developing grayish yellow (2B5), straw yellow (3B4), very pale brown (10YR 8–7/6, 7/4), brownish yellow (10YR 6/6), yellowish brown (10YR 5/8), light brown (6D6) yellowish brown (5F7) or dark brown (6F7) tints in age; solid becoming cavernous.

Basidiospores 7.2–8.2–10.5–12.1(–12.4) x 6.0–6.5–7.5–8.3 µm, ellipsoid to shortly ellipsoid, rarely subglobose [Q = (1.12–)1.16–1.24–1.42–1.59; V = 186–296 µm3; n = 153]; ornamentation partly to completely amyloid, up to 1.3 µm high, mostly 0.5–1.0 µm high, consisting of hemispherical, cylindrical or conical elements, with connecting lines of lower height, some elements fused in short ridges or subcatenulate, usually forming a partial to nearly complete reticulum; plage weakly amyloid. Basidia 50–64–80 x 9.5–11.4–13.5(–14.5) µm, clavate, four-sterigmate, hyaline in KOH; sterigmata 5–10 x 1.5–2.5 µm. Pleurocystidia 60–76–102–160(–175) x (7.5–)8.5–11.8–14.5 µm, usually clavate-fusoid or narrowly ventricose, with rounded or mucronate apices, at times with subapical constrictions, rather abundant (950–1500/mm2), projecting 5–50 µm beyond basidioles, rarely embedded in hymenium, completely filled with needle-like, yellowish contents in KOH, grayish brown to blackish brown in sulfovanillin (SV). Cheilocystidia 40–76–120 x 8.0–5.1–14.5 µm, similar to pleurocystidia, projecting 5–50 µm beyond marginal cells. Marginal cells 5.0–9.5 µm broad, cylindrical to clavate, more slender than basidioles, intermixed with basidia and cheilocystidia. Subhymenium composed of short hyphae, filamentous to pseudoparenchymatous. Lamellar trama heteromerous, consisting of nests of sphaerocytes, connective hyphae and abundant oleiferous hyphae; sphaerocytes 13–27–46(–48) µm broad, hyaline in KOH; connective hyphae 1.9–3.67–6.0(–6.4) µm broad, hyaline in KOH; oleiferous hyphae (3.1–)3.2–5.21–7.2 µm broad, with refractive contents in KOH. Pileipellis undifferentiated, without gelatinous matrix, 45–161–325 µm thick on disc; hyphae 2–9 µm broad, repent, interwoven, branched, thin- to slightly thick-walled (1.5 µm), some with encrusted pigment; extremities of the hyphae not aggregated, 3–9 µm in diam., filamentous, with rounded apices, often intermixed with inflated hyphae 6–20(–25) µm in diam. on margin; oleiferous hyphae 2.0–9.5 µm broad, scattered near trama; pileocystidia 30–76–135(–155) x (3.5–)4.0–5.1–6.5 µm, cylindrical to subfusoid, with rounded to mucronate apices or at times with subapical constrictions, thin-walled, not septate, partially filled with needle-like, yellowish contents in KOH, grayish brown, blackish brown to purplish brown in SV, scattered; without primordial hyphae; oleiferous hyphae often intermixed between pileipellis and trama, 2–9 µm broad, contorted irregularly, thin-walled, completely filled with homogeneous, refractive, yellowish contents in KOH, pink in SV, blue in cresyl blue, partially cyanophilous. Pileus trama heteromerous, consisting of nests of sphaerocytes, connective hyphae and oleiferous hyphae; sphaerocytes (21–)27–47–65 µm broad; connective and oleiferous hyphae similar to lamellar trama. Marginal veil composed of hyphal strands intermixed with inflated elements and sphaerocytes; cylindrical hyphae 4.0–11.0(–11.5) µm broad, some with yellowish, encrusted pigment; inflated elements 10–25 µm broad, thin- to slightly thick-walled (1.5 µm), regularly septate; sphaerocytes 40–60 µm broad, thin- to thick-walled (2.0 µm). Stipitipellis undifferentiated, without gelatinous matrix; hyphae 1.5–8.0(–9.0) µm broad, repent, interwoven, branched, thin- to slightly thick walled (1.5 µm), some with encrusted pigment; extremities of hyphae erect, with rounded, mucronate or rostrate apices, at times with subapical constrictions; caulocystidia 35–115–205(–220) x 3.5–6.1–9.5 µm, subcylindrical to fusoid, with rounded to mucronate apices, thin-walled, not septate, completely or partially filled with needle-like, yellowish contents in KOH, grayish brown, blackish brown to purplish brown in SV, rather abundant; oleiferous hyphae similar to those of the pileipellis. Velar remnants comprising hyphal strands and chains of sphaerocytes, similar to those of the marginal veil. Clamp connections absent from all tissues.

Macrochemical reactions (trama). {alpha}-naphthol: grayish magenta (13–14E6), grayish violet (15–17E6) to dark violet (18F6); FeSO4: peach (7A4) to shell pink (8A3); guaiacol: brownish red (8C6), reddish brown (8D-E6), jasper red (9B7), photo brown (9F8), bluish red (12A6), grayish rose (12B6) or dusky red (0R 3/4); phenol: dark brown (8F8) or photo brown (9F8); sulfoformol: no reaction or pale orange (6A3).

TYPE. MEXICO, TLAXCALA: Municipality of Panotla, 1 km E of San Francisco Temezontla, 19°20'45'' N, 98°16'25'' W, 2550–2650 m alt., Quercus forest, 10-VIII-1993, A. Kong 2622 (HOLOTYPUS TLXM).

Specimens examined. MEXICO. TLAXCALA: Municipality of Panotla, 1 km E of San Francisco Temezontla, 2550–2600 m alt., Quercus forest, 10-VIII-1993, A. Kong 2622 (HOLOTYPE TLXM). PARATYPE: 9-X-1992, A. Kong 2498 (TLXM); 10-IX-1993, A. Kong 2626 (MEXU, TLXM); 17-VIII-1993, A. Kong 2636 (MICH, TLXM); 9-VII-1994, A. Kong 2725 (PC, TLXM); 10-IX-1994, A. Kong 2773 (PC, TLXM); 4-VII-1995, A. Kong 2842 (TLXM); 20-VII-1995, A. Kong 2873 (K, NY, TLXM); 26-VII-1996, A. Kong 2906 (EIU, TLXM).

Etymology. Dedicated to Dr. Teófilo Herrera for his contributions to Mexican mycology.

Habitat and distribution. Solitary to scattered, in small groups of 2–4 basidiomes or subgregarious. On soil. In deciduous Quercus forest at 2550–2650 m above sea level. Jul to Oct. Presently known only from the type locality (19°20'45'' N, 98°16'25'' W).

Edibility and vernacular name. People from San Francisco Temezontla gather edible mushrooms in two forest types: the "ocotal" (Pinus forest) and the "encinal" (Quercus forest). All species of Russula in subsection Lactarioideae, including the new species where described are considered edible mushrooms by most of the inhabitants of this village. Basidiomes of these taxa are more abundant in the Pinus forest than in the Quercus forest. People apply the names "hongo blanco" (white mushroom) or "hongo blanco de ocote" (pine white mushroom) to all species of Russula subsection Lactarioideae whether they grow in the Quercus forest (e.g., R. delica var. centroamericana and R. herrerae) or in the Pinus forest (e.g., R. delica and R. romagnesiana Shaffer).


   DISCUSSION
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 TAXONOMY
 DISCUSSION
LITERATURE CITED
 
Russula herrerae belongs to section Plorantes, subsection Lactarioideae due to its lactarioid habit, white pileus color, acute and non-striate pileus margin, hard and unchanging trama, numerous lamellulae, and amyloid basidiospore plage (Shaffer 1964Citation , Romagnesi 1967Citation , Singer 1986Citation ).

This taxon is distinguished from other species in section Plorantes by the presence of a well-developed, appendiculate marginal veil, which at times remains in part as a fibrillose-floccose annulus on the stipe surface. It resembles Russula delica Fr. in having the same basidiome color, broad and distant lamellae, white to pale cream basidiospore deposit color, and similar basidiospore ornamentation. R. delica var. centroamericana Sing., described from a Quercus-Magnolia forest of Costa Rica (Singer 1989Citation ), is also similar. This new species differs from taxa of the R. delica group in its relatively small basidiomes, number of lamellulae between two lamellae, marginal veil and inflated hyphae of pileipellis margin. In the field, basidiomes of R. herrerae might be confused with species of the Lactarius deceptivus Pk. or L. pubescens Fr. group, due to the basidiome color and fibrillose margin, however, R. herrerae lacks the typical pseudocystidia present in the hymenium of all Lactarius species. The ontogeny of the veil is unknown, but probably is similar to the annulus development described by Heim (1937)Citation for R. radicans Heim.


   ACKNOWLEDGMENTS
 
We wish to thank Drs. Bart Buyck (PC) and Andrew S. Methven (EIU) who kindly reviewed the manuscript and made valuable suggestions. Profs. Lourdes Santiago and Patricia Villaseñor (UNAM) kindly improved the Latin diagnosis. Dr. Rodham E. Tulloss (NJ) reviewed an earlier draft of this manuscript. We are grateful to Drs. Robert L. Shaffer (MICH), Bart Buyck (PC), and Gregory M. Mueller (F) for the loan of some Russula section Plorantes collections. This work was partially supported by CONACyT (Project 4690).


   FOOTNOTES
 
1 Corresponding author, Email: akongluz{at}hotmail.com Back

Accepted for publication May 25, 2001.


   LITERATURE CITED
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 TAXONOMY
 DISCUSSION
 LITERATURE CITED
 
Acosta R, Kong A., 1991 Guía de las excursiones botánicas y micológicas al Cerro El Peñón y Cañada Grande del estado del Tlaxcala IV Congreso Nacional de Micología. Tlaxcala, Tlaxcala: Jardín Botánico Tizatlán 8. 82 p

Aroche RM, Cifuentes J, Lorea F, Fuentes P, Bonavides J, Galicia H, Menéndez E, Aguilar O, Valenzuela V., 1984 Macromicetos tóxicos y comestibles de una región comunal del Valle de México, I Bol Soc Mex Mic 19:291-318

Bandala-Muñoz VM, Guzmán G, Montoya-Bello L., 1988 Especies de macromicetos citadas de México, VII. Agaricales, parte II (1972–1987) Rev Mex Mic 4:205-250

Buyck B., 1991 The study of microscopic features in Russula 2. Sterile cells of the hymenium Russulales News 1:62-85

———. 1993 Russula I (Russulaceae) In: Rammeloo J, Heinemann P. eds. Flore illustrée des champignons d'Afrique Centrale 15. Brussels, Belgium: Ministère de l'Agriculture, Jardin Botanique National de Belgique. p 335–408

———, Verbeken A., 1995 Studies in tropical African Lactarius species. 2. Lactarius chromospermus Pegler Mycotaxon 56:427-442

Chacón S., 1995 Nuevos registros de Agaricales (Fungi) de México Acta Bot Mex 30:9-12

Cifuentes J, Villegas M, Pérez-Ramírez L., 1993 Hongos macroscópicos In: Luna Vega I, Llorente Bousquets J. eds. Historia Natural del Parque Ecológico Estatal Omiltemi, Chilpancingo, Guerrero, México. México D.F.: CONABIO-UNAM. p 59–126

Estrada-Torres A, Aroche RM., 1987 Acervo etnomicológico en tres localidades del Municipio de Acambay, Estado de México Rev Mex Mic 3:109-131

Estrada-Torres A, Kong A, Montoya A, Santiago G., 1991 Listado preliminar de la micobiota del estado de Tlaxcala Tlaxcala, Tlaxcala: Jardín Botánico Tizatlán 12:45-79

Frutis I, Chio RE, Estrada-Torres A., 1985 Nuevos registros de macromicetos del Estado de México Rev Mex Mic 1:285-300

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Guzmán G., 1961 Notas sobre algunas especies de agaricáceos no citadas en México An Esc Nal Cienc Biol 10:23-38

———. 1966 Hongos (macromicetos) comunes en la ruta Amecameca-Tlamacas (Volcán Popocatépetl), México Guías de Excursiones. México D.F.: III Congreso Mexicano de Botánica

———. 1975 Hongos mexicanos (macromicetos) en los herbarios del extranjero, III Bol Soc Mex Mic 9:85-102

Heim R., 1937 Observations sur la flore mycologique malgache V. Les Lactario-Russulés à anneau: Ontogénie et phylogenie III Rev Mycol 2:61-75

Herrera T, Guzmán G., 1961 Taxonomía y ecología de los principales hongos comestibles de diversos lugares de México An Inst Biol Mex 32:33-135

———, ———. 1972 Especies de macromicetos citadas de México, III. Agaricales Bol Soc Mex Mic 6:61-91

Holmgren PK, Holmgren NH., 1995 Additions to index herbariorum (Herbaria), 8th ed, 4th series Taxon 44:251-266

———, ———, Barnett LC., 1990 Index herbariorum. Part I. The herbaria of the world. 8th ed Regnum Veg 120:1-693

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