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Kurtziana

versão On-line ISSN 1852-5962

Kurtziana vol.37 no.1 Córdoba jan./jun. 2012

 

ARTÍCULOS ORIGINALES

New species and records of Pouzarella (Agaricomyetes, Entolomataceae) from northern Argentina

 

Timothy J. Baroni 1, Edgardo Albertó 2, Nicolás Niveiro 3 & Bernardo Lechner 4

1 Department of Biological Sciences, PO Box 2000, State University of New York - College at Cortland, Cortland, NY 13045 USA .
2 Instituto de Investigaciones Biotecnológicas (IIB-INTECH), CONICET-UNSAM, C. C. 164, (7130) Chascomús, Buenos Aires, Argentina.
3 Instituto de Botánica del Nordeste, Sargento Cabral 2131, CC 209 Corrientes Capital, CP 3400, Argentina.
4 Departamento de Biodiversidad y Biología Experimental (DBBE), Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, PROPLAME- PRHIDEB-CONICET, 1428 CABA Buenos Aires, Argentina.

 


Summary

Five new species of Pouzarella are described from northern Argentina, P. brunneonigrescens, P. olivacea, P. parvispora, P. sulcata and P. variabilis. Pouzarella ferreri var. ferreri is also reported for the first time from Argentina. All known Pouzarella species reported from Argentina are discussed and a key to species is provided.

Key words: Biodiversity; Mushrooms; Yungas; Keys.

Resumen

Nuevas especies y nuevas citas de Pouzarella (Agaricomyetes, Entolomataceae) del norte argentino

Se describen e ilustran cinco nuevas especies de Pouzarella del norte argentino, P. brunneonigrescens, P. olivacea, P. parvispora, P. sulcata y P. variabilis. Pouzarella ferreri var. ferreri es citada por primera vez para la Argentina. Se discuten todas las especies conocidas de Pouzarella para la Argentina y se propone una clave para su identificación.

Palabras clave: Biodiversidad; Hongos; Yungas; Claves.


 

Introduction

The agaric mycobiota of Argentina is one of the most studied in South America. C. Spegazzini, R. Singer and E. Horak, three noteworthy mycologists, collected and studied the argentine mushrooms over the past years and their publication enriched considerably the knowledge of the group in the region. A complete revision of the publications on Agaricales from Argentina was recorded by Moreno & Albertó (1996). From 1996 to the present, at least 18 papers were published describing and illustrating agarics in Argentina: Albertó (1996, 1998, 1999); Albertó et al. (1998); Lechner & Albertó (2000, 2008); Albertó et al. (2000), Fazio & Albertó (2001); Albertó et al. (2002a, 2002b); Lechner et al. (2002); Lechner et al. (2003); Lechner et al. (2004, 2005); Uhart & Albertó (2007, 2009); Niveiro et al. (2009, 2010). Three illustrated books are now available to help show the fungal diversity from different areas of the country: Gamundi & Horak (1993) southwestern, Wright & Albertó (2002), middle area; Wright et al. (2008) northeastern. Nevertheless, many groups of agarics have still not been studied in depth. The family Entolomataceae Kotl. & Pouzar (as Rhodophyllaceae Singer) in Argentina, was summarized by Singer & Digilio (1952), and they listed 18 species at that time. One of these listed species, Rhodophyllus squamifolius (Murrill) Singer, is now considered to belong to the genus Pouzarella (Mazzer, 1976) and was the only species of Pouzarella reported from Argentina at the time. Of the nine taxa described as Entolomataceae by Spegazzini from the late 1800's to the early 1920's, only six actually represent members of the Entolomataceae (Horak, 1978b), and none of these are currently considered Pouzarella taxa.

Horak (1977, 1978a, 1978b, 1983) listed with brief descriptions or described as new nearly 140 taxa of Entolomataceae from South America. Of these only 5 taxa are considered Pouzarella species (P. dysthales (Peck) Mazzer, P. dunstervillei (Dennis) Mazzer, P. aureocrinita (E. Horak) T. J. Baroni, P. sepiaceobasalis (E. Horak) T. J. Baroni, and P. erinacea (E. Horak) T. J. Baroni). All these species except P. dunstervillei were reported from Argentina (Horak, 1978b).

A recent excursion to the northern provinces of Argentina to a moist broadleaf forest ecoregion named the Yungas, provided us the opportunity to document several collections of Pouzarella. The Andean Yungas Forests is a system of tropical and subtropical montane cloud forests on the oriental slope along the Andes in South America. The weather in this region is characterized by seasonal fog and pervasive cloud cover (Brown et al. 2005). Argentinean Yungas constitutes the southeast expression of the Amazonic Domain (Cabrera, 1994), and this region is characterized by a high altitudinal gradient and by different environmental conditions such as periods of high temperatures, drought, high humidity levels, frosts and snow in winter, which reflects a particular floristic composition. Tree diversity is high and is estimated to include more than 230 species with elements of Holarctic (Alnus, Berberis, Juglans, Sambucus), Gondwanic (Podocarpus, Escallonia, Weinmannia), Pantropical (Eugenia, Ocotea) and Neotropical (Myrcianthes) origins (Grau & Brown, 2000; Brown et al. 2005). According to Brown et al. (2005), three main forest types can be distinguished: premontane lowland forests (400- 700 m asl); lower montane forests (700-1500 m asl); and upper, temperate montane forests (1500-2500 m asl). Our collecting efforts were focused on the lower montane and the upper, temperate montane ecosystems.

From the eight collections of Pouzarella discovered in two different Provinces in northern Argentina, we describe here five new species and a first report of P. ferreri for Argentina. Full descriptions and illustrations are provided for all of these species and complete discussions of all taxa known from Argentina are provided.

Materials and Methods

Color notations in parentheses are from Kornerup & Wanscher (1978). Microscopic features where studied in 3% KOH, NH4OH, Congo Red, and/or Melzer's Reagent. For basidiospore measurements, the hilar appendix or apiculus was excluded. In those measurements Q refers to the length divided by the width of an individual spore. The notation n = 32/2, means that 32 individual basidiospores were measured from 2 different collections, while n = 20 means that 20 individual spores were measured from a single sample. Means for length and width are given with their standard deviations, as is the mean of Q values, with the mean of the length divided by the width of all basidiospores measured being designated as Qm.

Caulocystidia for Pouzarella is used to designate typically multi-celled hyphae that are produced on and project from the stipe surface. These"caulocystidia" are taxonomically important as they vary significantly among species in this genus. The apex of the stipe is usually examined, but samples are also taken for comparison lower down the stipe. The terminal cell on these hyphal chains shows the most morphological variation and is useful for discriminating taxa.

Cheilocystidia may be clustered and these clusters may be spaced along the lamella edge (i.e. as granulate fimbriate in the morphological descriptions). Therefore it is important to mount 1-2 mm pieces of the whole edge to determine presence and types of cheilocystidia.

Descriptive statistical analysis of basidiospores was developed using EXCEL 5.0 and Sigmastat 1.0. All light microscopic images were made with an Olympus BX 50 transmitted light microscope using DIC or bright field optics and captured with a Diagnostic Instruments, Inc. Insight Spot 3-shot color digital camera.

All GPS readings were taken on a Garmin 60CSx hand held unit using WGS84 Map Datum settings.

Results

A. New Species:

Pouzarella brunneonigrescens T. J. Baroni, Albertó, Niveiro & B. E. Lechner sp. nov.

Diagnosis: distinguished from other species of Pouzarella by the moderately robust stature with a firm stipe 2-3 mm in diam, the brown pileus with dark brown squamules mixed with reddish orange fine recurved squamules and -reddish fibrils over the sulcate margin, the ashy white densely hirsute stipe, all tissues of pileus, lamellae and stipe turning slowly fuscous black where bruised, and the inflated mostly globose or ovate thick-walled pale brown cheilocystidia.

ARGENTINA: Salta Province, Los Toldos, El Nogalar Nature Reserve, along forested creek bed, 20-III-2011, 22° 16' 41.6"S 64° 42' 45.1"W, 1652 m. a. s. l., collecting team T. J. Baroni, E. Alberto, B. E. Lechner, N. Niveiro, leg. 10370 T. J. Baroni (Holotype - BAFC; Isotype - CORT). Figs. 4, 19-22

Mycobank #: 563280

Pileus dull brown (6E5-7 - Cocoa Brown, Cognac) with brighter colors over margin (6D5- 6 - Sunburn, Cinnamon) from reddish orange very fine recurved squamules and lanate fine reddish fibrils covering the margin, slowly turning fuscous where bruised, 6-20 mm broad, convex, becoming broadly campanulate, shallow sulcate to disc, with matted or erect brownish squamules over disc and joining the reddish orange squamules and fibrils at the margin. Flesh dark watery fuscous, 1 mm thick. Lamellae grayish (5-6D3 - Nougat or Café-au-Lait) or becoming reddish blond or brownish orange from spore maturation (5C3), adnexed or ± arcuate-sinuate, subdistant, broad (3 mm), thick, edge white fimbriate, blackening slowly when injured. Stipe pallid ashy white from dense hirsute vestiture, becoming fuscous black from handling, pallid ashy white strigose at base, 2-3 x 20-45 mm, equal, terete, entire surface with long recurved hirsute pubescence, pliant-tough. Odor pungent herbal, taste not tried. All tissues not only slowly staining dark fuscous black when handled and bruised, all tissues becoming dark fuscous black or deep brownish after dehydration. When rehydrated in directly 3% KOH or if first placed in 95% ETOH, then dH2O and then into 3% KOH, all tissues producing a pale rose or pale vinaceous discoloration in the mountant. If a tissue sample is left in the dH2O in a spot plate for several hours, all microstructures become obscured from deep fuscous intracellular pigments that develop profusely throughout the cells.

Basidiospores 14.6-18.6 x 9.7-11.3 (-12.1) µm (n = 20, 16.8 ± 1.11 x 10.2 ± 0.64, Q = 1.47- 1.92, Qm = 1.64 ± 0.11), heterodiametric, 7-9 angled in profile and face views, 4-5(-6) rounded angular in polar view. Basidia 4-sterigmate, clavate, 40-48 x 14-16 µm, with abundant, evenly spaced aborted basidia with dark brown intracellular pigments and collapsed sterigmata. Cheilocystidia pale brownish or ± hyaline, with distinctly thickened-walls (approx. 0.5-1 µm), terminal cells mostly globose, ovate, short broad clavate or broad short fusiform, most often in chains of 2-3 inflated cells with the lower cells inflated and also thick walled, terminal cells 26-58 x 18-38 µm. Pleurocystidia absent. Lamella trama a very dark fuscous layer in 3% KOH, composed of ± parallel, cylindrical or inflated hyphae, 3-10 µm in diam, heavily encrusted with dark fuscous brown pigments, subhymenium composed of 2-4 rectangular cells (pseudoparenchymatous like). Pileipellis a brown layer of inflated repent hyphae producing scattered clusters of ± erect, even darker fuscous brown, pyramidal clusters of chains of sausage shaped inflated cells with thick walls and intraparietal brown pigments and some with fine brown encrustations, terminal cell in these chains 50-104 x 18-30 µm; subpellis composed of a distinct layer (100-200 µm deep) of inflated cells, 10-24 µm in diam). Pileus context a brown layer of radially arranged cylindrical or inflated hyphae, 4-26 µm in diam, heavily encrusted with dark brown pigments. Stipitipellis a dark brown layer of cylindrical or somewhat inflated, dark brown pigment encrusted hyphae, 3-14 µm in diam, producing erect or ascendant several celled chains (2-5 cells) of dark brown (in 3% KOH), thick walled"caulocystidia", all cells in these chains with brown intraparietal pigments and some with brown encrusting pigments, terminal cells mostly tapered or fusoid, some clavate, 56-130 x 18-33 µm. Clamp connections absent.

Etymology: pileus brown, becoming black from injury or from dessication

Habit, Habitat, Fruiting period: scattered or gregarious on firm wet humus among large rounded rocks, near seepage areas and creek, under Juglans australis Griseb. and Podocarpus sp., in cool montane tropical broad-leaved forests. March.

Material Examined: ARGENTINA: Salta Province, Los Toldos, El Nogalar Nature Reserve, along forested creek bed, 20-III- 2011, 22° 16' 41.6"S 64° 42' 45.1"W, 1652 m. a. s. l., collecting team T. J. Baroni, E. Alberto, B. E. Lechner, N. Niveiro, leg. 10370 T. J. Baroni (Holotype - BAFC; Isotype - CORT).

The obscure (use of a hand lens is needed to determine clearly) reddish orange fine squamules and fibrils over the margin of the deep brown colored and mostly brown squamulose pileus, coupled with the fuscous bruising reactions of all tissues made us first consider that this might be a variation on Pouzarella ferreri T. J. Baroni, Perd.-Sánch., S. A. Cantrell but with a brown pileus and an ashy white stipe. The reddishorange P. ferreri also stains dark blackish from handling, a distinctive and not typical bruising reaction for other taxa of Pouzarella (Mazzer, 1976; Horak, 1983; Noordeloos, 1992; Baroni et al., 2008; Baroni & Ortiz, 2002). However, in the case of P. brunneonigrescens, the color differences (brown pileus and white stipe vs. orange or deep orange red pileus and yellow stipe with yellow ochre pubescence, see also description below for P. ferreri), the robust stature for a Pouzarella, the thick-walled brown mostly globose/ovate cheilocystidia, and the very thick-walled and differently shaped caulocystidia seem to justify recognizing this taxon as a new species. The caulocystidia for P. ferreri var. ferreri are filamentous, multicelled and with globose, broadly clavate, fusoid or ventricose terminal cells, while the typical terminal cells of caulocystidia of P. brunneonigrescens are tapered or fusoid and not inflated. P. brunneonigrescens differs by at least five separate morphological features from P. ferreri var. ferreri.

Baroni et al. (2008) described a pale stiped variation of P. ferrerii from Jamaica, P. ferreri var. pallidonigrescens T. J. Baroni & S. A. Cantrell with a paler reddish orange pileus and a white stipe that turns black from handling. This taxon shares the pallid colored stipe and nigrescent reactions of all parts of the basidiomata with P. brunneonigrescens, but the similarities end there. Stature (delicate vs. robust), pileus color (reddish orange vs. brown), cheilocystidia (not obviously thick-walled and pale brown vs. obviously thick-walled and brown), terminal cells of caulocystidia chains (narrowly clavate or fusoid vs. tapered or fusoid, not inflated) separate P. ferreri var. pallidonigrescens and P. brunneonigrescens.

The inflated cells in the subpellis and thick walled cheilocystidia place P. brunneonigrescens in Section Dysthales, Subsection Inflatae along with several other taxa, including P. ferreri. The moderately robust stature with the stipe reaching 3 mm in diam, the sulcate brown pileus, ashy white stipe and gray brown lamellae, all with nigrescent staining reactions and the thick-walled brown colored mostly globose/ovate-inflated cheilocystidia make this species easily recognizable in the genus. The nigrescent reaction of the tissues indicates a relationship with P. ferreri var. ferreri and P. ferreri var. pallidonigrescens.

Of the three species of Pouzarella previously described from northern Argentina (see also discussion under P. ferreri) by Horak (1983), P. sepiaceobasalis seems to be similar to P. brunneonigrescens because of a few overlaping features. Those features are a similar color and surface of the pileus for P. sepiaceobasalis ("... golden brown with faint orange tinge;... striate-sulcate towards margin"); similar form and vestiture and perhaps basal coloration of the stipe ("...stiff; upper half golden brown, yellow-brown, lower half gradually blue-black; wooly-hairy over whole length, brown to black bristles at base well developed..."); and the inflated and somewhat similar cheilocystidia ("...clavate to vesiculose, membrane hyaline, thin-walled, pigment none."). However, there are several features that separate these two taxa, the most prominent of which is the nigrescent nature of all tissues for P. brunneonigrescens. Horak (1983) may have missed this feature if the specimens were processed rapidly, and the description of the lower half of the stipe turning gradually blue-black might support this hypothesis. But we are skeptical he would have missed such distinctive features since we know what a keen observer Horak is known to be. Also his description of the colors of the stipe with golden brown and yellow brown hues and the much thinner shape (to 1 mm in diam) is different than what we found for P. brunneonigrescens. Additionally, and even more important, the globose very thick walled cheilocystidia with brown colored walls and inflated supporting cells, the broader basidiospores (7.5-9 µm for P. sepiaceobasalis vs 9.7-12 µm for P. brunneonigrescens), the inflated cells in the subpellis for P. brunneonigrescens (not noted for P. sepiaceobasalis) show these two taxa have several differences that make them distinct.

Pouzarella olivacea T. J. Baroni, Albertó, Niveiro & B. E. Lechner sp. nov.

Diagnosis: differing from others in the genus Pouzarella by the olivaceous colored, sulcate pileus with golden olivaceous fine scales and fibrils on the margin, olive grayish-yellow stipe with grayish olivaceous dense pubescence, all tissues turning fuscous where injured, the mostly globose hyaline cheilocystidia and the caulocsytidia composed of a few cells in short chains with inflated mostly clavate or fusoid terminal cells.

ARGENTINA: Salta Province, Los Toldos, El Nogalar Nature Reserve, along forested creek bed, 20-III-2011, 22° 16' 41.6"S 64° 42' 45.1"W, 1652 m. a. s. l., collecting team T. J. Baroni, E. Alberto, B. E. Lechner, N. Niveiro, leg. 10372 T. J. Baroni (Holotype- BAFC; ISOTYPE - CORT). Figs. 5, 23-26

Mycobank #: 563281

Pileus olivaceous brown hues (5F6- 7 - Tobacco Brown, Coffee) with distinctly olivaceous colors over the margin (4B4 - Champagne to 4C5 - Chamois) from the fine olivaceous scales and fibrils over the margin and very fine olivaceous squamules over the disc, 6-13 mm broad, sulcate to disc. Lamellae gray brown (6D3 - Café-au-Lait), arcuate-sinuate, subdistant, broad (3 mm), edges granulate-fimbriate, blackening where bruised, thick. Stipe pale olive yellow or grayish yellow, with a dense grayish olivaceous pubescence overall, 1-1.5 x 30-40 mm, equal, terete, fragile, becoming black from handling, yellowish olivaceous strigose over base. Odor none, taste not tried. All fresh tissues slowly staining dark fuscous black when handled and bruised, also all dried tissues becoming dark fuscous black or deep brownish after dehydration. When rehydrated in 3% KOH or in a series of 95% ETOH, then dH2O and then 3% KOH, the lamellae produce a pale rose or pale vinaceous discoloration in the mountant, the pileus produces a pale lime green discoloration in the same rehydration series if the sections are allowed to remain in dH2O for several minutes (up to 1-2 hrs). Eventually, after several hours, all revived tissues turn dark fuscous black.

Basidiospores (15.4-) 16.2-19.4 x 10- 12.1 µm (n = 20, 17.3 ± 0.95 x 10.9± 0.52, Q = 1.43-1.77, Qm = 1.59 ± 0.10), heterodiametric, 7-8 angled in profile and face view, 4-5 rounded angular in polar view. Basidia 4-sterigmate, inflated clavate, 44-56 x 16-20 µm. Cheilocystidia ± hyaline with slightly or moderately and unevenly thickened walls, mostly sphaerical or globose, also ovate or broadly clavate, frequently with inflated support cell, 22-38 x 16-24 µm. Pleurocystidia absent. Lamella trama brown layer of ± parallel, cylindrical or inflated hyphae, 4-12 µm in diam, filled with dark brown intracellular pigments and also encrusted with dark fuscous brown pigments. Pileipellis of two distinct layers, a dark brown layer of inflated repent hyphae in chains of sausage shaped inflated cells with unevenly thickened walls and intraparietal brown pigments and some with fine brown encrustations, terminal cells in these chains cylindrical or bullet shaped, 36-72 x 12-18 µm; subpellis composed of a distinct layer (100- 200 µm deep) of inflated cells, 30-60 µm in diam). Pileus context a brown layer of radially arranged cylindrical hyphae, 4-10 µm in diam, heavily encrusted with dark brown pigments. Stipitipellis a brown layer of cylindrical or somewhat inflated, dark brown pigment encrusted hyphae, 3.5-12 µm in diam, producing short erect or ascendant 2 (-3-4) celled chains of brownish (in 3% KOH), slightly thick walled"caulocystidia", all cells in these chains with brown intraparietal pigments and none with encrusting pigments, terminal cells mostly fusoid, clavate, ovate or cylindrical, 34-120 x 12-30 µm. Clamp connections absent.

Etymology: referring to the olivaceous colors of the pileus and stipe

Habit, Habitat, Fruiting period: scattered or gregarious on firm wet humus among large rounded rocks, near seepage areas and creek, under Juglans australis Griseb. and Podocarpus sp., in cool montane tropical broad-leaved forests. March.

Material Examined: ARGENTINA: Salta Province, Los Toldos, El Nogalar Nature Reserve, along forested creek bed, 20-III- 2011, 22° 16' 41.6"S 64° 42' 45.1"W, 1652 m. a. s. l., collecting team T. J. Baroni, E. Alberto, B. E. Lechner, N. Niveiro, leg. 10372 T. J. Baroni (Holotype - BAFC; ISOTYPE - CORT).

Pouzarella olivacea , like P. brunneonigrescens, is placed in Section Dysthales, Subsection Inflatae because of the thickened walls of the cheilocystidia and the inflated cells in the subpellis. These two taxa were found in the same general area with each collection consisting of several basidiomata. The distinctive color differences in fresh condition were striking. The differences in caulocystidia shapes and numbers of cells making up the"hairs" on the stipe surface are also very different and help to distinguish these two taxa.

Both P. olivacea and P. brunneonigrescens show affinities to P. ferreri by the tissues that become nigrescent when handled, the production of rose or pale vinaceous pigments in 3% KOH mounting media when tissues are revived and the frequently globose/sphaerical or at least inflated ovate/clavate cheilocystidia. For P. olivacea the pale lime green discoloration of the pileus surface in a rehydration series of 95% ETOH and then dH2O makes this species distinctive. We guess that these three taxa may well have a common ancestor and this may become evident in future studies using DNA technology.

Pouzarella parvispora T. J. Baroni, Albertó, Niveiro & B. E. Lechner, sp. nov.

Diagnosis: differing from others in Pouzarella Section Pouzarella by basidiospores shorter than 13 µm, grayish brown colors and versiform, thin-walled cheilocystidia.

ARGENTINA: Jujuy Province; Calilegua National Park, Guarani Trail, 24-III-2011, 23° 45' 40.9"S 64° 51' 06.9"W, 633 m. a. s. l., collecting team, T. J. Baroni, E. Albertó, B. E. Lechner, N. Niveiro, leg. 10402a TJ Baroni (Holotype -BAFC; Isotype - CORT) Figs. 2, 11-14

Mycobank #: 563278

Pileus dark grayish brown (6F4 -Chocolate) on disc, slightly paler grayish brown over margin (6E3), 9-15 mm broad, broadly conicalconvex, sulcate to disc, densely erect and recurved pointed black squamulose over disc, more sparsely squamulose to margin. Flesh less than 1 mm thick, concolorous with pileus surface. Lamellae grayish brown as pileus margin, adnexed, subdistant, very broad (3 mm), edges smooth. Stipe ± concolorous with pileus, 1-1.5 x 31-40 mm, equal, terete, grayish brown strigose at base, moderately grayish brown fibrillose tufted (clusters of +stiff erect hairs) overall, hollow. Odor none, taste not tried.

Basidiospores 9.7-13 x 8-11.3 µm (n = 20, 12.3 ± 0.64 x 8.7 ± 0.44, Q = 1.34-1.51, Qm = 1.42 ± 0.05), heterodiametric, 6-7 (-8) angled in profile view, 6-8 angled in face view, 4-5 rounded angular in polar view, hilar appendix (apiculus) 0.8-1.6 µm long. Basidia 4-sterigmate, clavate, some aborted, shrunken and filled with dark brown pigment, 36-40 x 12-14 µm. Cheilocystidia hyaline or with very pale brownish intraparietal pigments in walls, in clusters, some areas of lamella edge lacking these cells, versiform in shape (fusoid, clavate, cylindrical, broadly ventricose-rostrate, some apically branched, also some septate), 46-80 x 10-28 µm. Pleurocystidia absent. Lamella trama of distinctly two zones of ± parallel hyphae, the central tissues (mediostratum) hyaline and composed of mostly inflated and more sparsely encrusted hyphae, 10-18 µm in diam, the laterostrata composed of a dark brown layer of heavily encrusted cylindrical hyphae, 4-8 µm in diam, subhymenium "pseudoparenchymatous" and several cell layers deep (3-6 cells). Pileipellis a dark brown layer of repent cylindrical hyphae, producing scattered clusters of upturned or ascendant, very inflated chains of sausage shaped cells, 16-30 µm in diam, with terminal cells mostly narrower and with tapered apices, 10-20 µm in diam, all cells with thickened (0.5-0.8 µm) walls with dark yellow brown intraparietal pigment in the walls, causing marbled or streaked appearance to the cells because of wall stretching that appears to produce channels on the inner walls making the pigment less dense and thus a lighter color, encrustations not present or not obvious. Pileus context a dark brown layer of radially arranged, heavily dark brown encrusted, cylindrical hyphae, 4-12 µm in diam. Stipitipellis a hyaline layer of repent, cylindrical hyphae, 4-8 µm in diam, producing well defined clusters of often multi-septate (2-4 cells), thick-walled (0.5-1 µm) brownish setiform"caulocystidia" with long tapered end cells and frequently with swollen cells supporting the terminal cell or in the basal portion of the chain and often this swollen cell is ventricose in shape, pigment intraparietal not encrusting, terminal cells 36-134 x 10-15 µm. Clamp connections: absent.

Etymology: short spores for a member of Section Pouzarella

Habit, Habitat, Fruiting period: scattered on humus with decaying plant leaves and twigs, in cool montane tropical broad leaved forests. March.

Material Examined: ARGENTINA: Jujuy Province; Calilegua National Park, Guarani Trail, 24-III-2011, 23° 45' 40.9"S 64° 51' 06.9"W, 633 m. a. s. l., collecting team, T. J. Baroni, E. Albertó, B. E. Lechner, N. Niveiro, leg. 10402a T. J. Baroni (Holotype -BAFC; Isotype - CORT)

Pouzarella parvispora fits nicely into Section Pouzarella because of the setiform caulocystidia (Mazzer, 1976). In that section in Mazzer's monograph (1976), all members have spores larger than 13 µm in length (i.e. P. babingtonii, P. nodospora and P. strigosissima) and are also differently colored.

The only other member of Section Pouzarella known in Argentina is Pouzarella aureocrinita (Horak) T. J. Baroni (Baroni et al. 2008). Pouzarella aureocrinita was described from Las Lagunas in Jujuy Province by Horak (1983) and is documented as having a red or red brown pileus that is covered with concolorous minute squamules, the basidiospores are 12-16 x 6.5-8 µm, the caulocystidia are cylindrical, 5-8 µm in diam and have greatly thickened walls (to 1.5 µm), cheilocystidia are lacking and the hyphae of the pileus surface are encrusted with a brown pigment. Clearly P. parvispora is morphologically different and is now the second species of Section Pouzarella reported for Argentina.

Pouzarella sulcata T. J. Baroni, Albertó, Niveiro & B. E. Lechner, sp. nov.

Diagnosis: differing from other members of Pouzarella Section Dysthales by the sulcate pileus surface, the versiform cheilocystidia and the inflated, single celled caulocystidia.

ARGENTINA: Salta Province, Los Toldos, El Nogalar Nature Reserve, along forested creek bed, 20-III-2011, 22° 16' 41.6"S 64° 42' 45.1"W, 1652 m. a. s. l., collecting team T. J. Baroni, E. Albertó, B. E. Lechner, N. Niveiro, leg. 10371 T. J. Baroni (Holotype - BAFC; Isotype - CORT). Figs. 3, 15-18

Mycobank #: 563279

Pileus moderate grayish brown (5D3- Nougat) to yellowish brown (5E5 - Bronze), with darker chocolate brown disc and striae between the sulcate ridges (6F5-6 - Teak or Burnt Umber), 5-22 mm broad, conical convex becoming campanulate, disc typically truncate, strongly sulcate-striate to near disc, surface minutely brown punctate fibrillose-squamulose overall, some squamules expallent and becoming silvery shiny on the tips, dry. Flesh dark watery grayish, 1 mm thick. Lamellae grayish brown becoming more flesh colored with spore maturation (6C3), adnexed or arcuate-adnate or deeply and broadly sinuate, subdistant, broad (to 3 mm deep), edges granulate-fimbriate. Stipe grayish or grayish yellowish brown (5D4 - Dark Blonde), with pale silvery-tan or sordid whitish strigose base, 1-2 x 35-70 mm, equal, terete, with silvery or pale grayish ashy or on one slightly chestnut colored pulverulence or floccose-pubescence ± overall evenly or in clumps. Odor not distinctive, taste not tried.

Basidiospores 11.3-17.8 x 8-12.1 µm (n = 27, 15.3 ± 1.72 x 10 ± 1.0, Q = 1.25-1.75, Qm = 1.53 ± 0.14), heterodiametric, (5-) 6-7 angled in profile view, 6-7 angled in face view, 4-5 obscurely rounded angular in polar view. Basidia 4-sterigmate, clavate, all hyaline, no obvious aborted basidia seen, 30-42 x 12-16 µm. Cheilocystidia brown colored clusters of versiform cells with brown, slightly thickened walls (0.5-1.5 µm), cells narrowly or broadly fusoid, ventricose-mucronate, cylindrical, clavate or broadly clavate, pyriform, or sphaerical inflated and supported by inflated cells in 2-3 or more celled chains, walls with brown intraparietal pigments and also many finely encrusted, 34-120 x 16-36 µm. Pleurocystidia absent. Lamella trama a dark brown tissue of more or less interwoven or somewhat parallel, cylindrical hyphae, 4-14 µm in diam, heavily dark brown encrusted. Pileipellis a brown layer of repent cylindrical hyphae giving rise to clusters of chains of inflated, brown pigment encrusted cells in repent or slightly recurved orientation, terminal cells inflated cylindrical or slightly tapered to apex, 36-70 x 12-22 µm, subpellis lacking inflated cells. Pileus context dark brown, radially arranged, cylindrical, heavily brown encrusted hyphae, 5-12 µm in diam. Stipitipellis a pale brown layer of repent, parallel, cylindrical hyphae, 4-11.3 µm in diam, producing scattered or clustered mostly single celled, clavate inflated caulocystidia, 50-90 x 12-30 µm, with pale brown walls and some finely pigment encrusted. Clamp connections absent.

Etymology: from the strongly sulcate pileus.

Habit, Habitat, Fruiting period: scattered or gregarious on firm wet humus near seepage areas and creek, under Juglans australis Griseb. and Podocarpus sp., in cool montane tropical broad-leaved forests. March.

Material Examined: ARGENTINA: Salta Province, Los Toldos, El Nogalar Nature Reserve, along forested creek bed, 20-III- 2011, 22° 16' 41.6"S 64° 42' 45.1"W, 1652 m. a. s. l., collecting team T. J. Baroni, E. Albertó, B. E. Lechner, N. Niveiro, leg. 10371 T. J. Baroni (Holotype - BAFC; Isotype - CORT). Same date and location, leg. 10376 T. J. Baroni (CORT; BAFC)

Pouzarella sulcata is a member of Section Dysthales in Pouzarella because of the grayish brown colors of the pileus, the pubescent stipe surface with inflated cells and the thick-walled cheilocystidia (Mazzer, 1976). Because this species lacks inflated cells in the subpellis, it is also relegated to the Subsection Dysthales. In this Subsection, P. sulcata seems to be most similar to P. dysthales because of the grayish brown colors, the overlapping basidiospore sizes, and the mixed swollen pyriform and fusoid thick-walled cheilocystidia (Mazzer, 1976). Comparing P. sulcata to published descriptions of P. dysthales (Mazzer, 1976; Noordeloos, 1992) it is clear that P. sulcata is morphologically distinct from that species because of the strongly sulcate pileus surface, the widely variable forms of cheilocystidia and the greatly inflated single celled caulocystidia.

The cheilocystidia depicted by Noordeloos (1992) for the European collections of P. dysthales show and describe only strongly inflated broadly clavate, obpyriform or subglobose elements (but in the description he mentions rare"fingler-like apical projections"), while the collections studied by Mazzer (1976) from across North America and Europe are described as having fusoid-ventricose, pyriform with mucronate tips or sub-catenulate cheilocystidia. Mazzer (1976) does not describe nor discuss the caulocystidia, but multi-septate cylindrical and heavily encrusted cells are depicted for P. dysthales by Noordeloos (1992). The mostly inflated single celled caulocystidia of P sulcata are very different.

This new species, P. sulcata, is morphologically distinct from other members currently known for the genus.

Pouzarella variabilis T. J. Baroni, Albertó, Niveiro & B. E. Lechner, sp. nov.

Diagnosis: differing from Pouzarella fibrillosipes (Murrill) Mazzer by broader basidiospores (9.7-12.1 µm for P. variabilis vs. 7.5-10.0 µm for P. fibrillosipes, Mazzer, 1976), tapered caulocystidia, sulcate pileus surface, and habitat on soil.

ARGENTINA: Jujuy Province, Calilegua National Park, Tatapua Trail, 25-III-2011, 23° 45' 40.09"S 64° 51' 06.09"W, 633 m. a. s. l., collecting team T. J. Baroni, B. E. Lechner, E. Albertó and N. Niveiro, leg. 10414 T. J. Baroni (Holotype - BAFC; Isotype - CORT). Figs. 1, 7-10


Figs. 1-2. 1. Pouzarella variabilis, 10414TJB HOLOTYPE, basidiomata. 2. Pouzarella parvispora, 10402aTJB HOLOTYPE, basidiomata. Scale bars 10 mm


Figs. 3-4. 3. Pouzarella sulcata, 10371TJB HOLOTYPE, basidiomata. 4. Pouzarella brunneonigrescens, 10370TJB HOLOTYPE, basidiomata. Scale bars 10 mm


Figs. 5-6. 5. Pouzarella olivacea, 10372TJB HOLOTYPE, basidiomata. 6. Pouzarella ferrerii var. ferrerii, 10379TJB, basidiomata. Scale bars 10 mm.


Figs. 7-10. Pouzarella variabilis, 10414TJB HOLOTYPE. 7. Caulocystidia. 8 Cheilocystidia. 9. Basidiospores. 10. Section through pileus surface, context and hymenium, showing hyphae of pileipellis, inflated cells of subpellis, and basidia with surrounding cells in hymenium. Scale bars 10 µm.


Figs. 11-14. Pouzarella parvispora, 10402aTJB HOLOTYPE. Fig. 15. Pouzarella sulcata, 10371TJB HOLOTYPE. 11. Caulocystidia. 12. Cheilocystidia. 13 Hymenium showing some basidia becoming filled with brown pigment and narrow pigment encrusted hyphae of laterostratum and pseudoparenchymatous subhymenium. 14. Basidiospores. 15. Basidiospores. Scale bars 10 µm.


Figs. 16-18. Pouzarella sulcata, 10371TJB HOLOTYPE. Figs. 19-21: Pouzarella brunneonigrescens, 10370TJB HOLOTYPE. 16. Cheilocystidia. 17. Cheilocystidia. 18. Caulocystidia. 19. Cheilocystidia. 20. Caulocystidium terminal cell showing internally thickened and sculpted inner wall with brown intraparietal pigments. 21. Caulocystidia. Scale bar = 10 µm for Figs. 16-18, 20-21, scale bar = 20 µm for Fig. 19.


Fig. 22. Pouzarella brunneonigrescens, 10370TJB HOLOTYPE. Basidiospores. Figs. 23-26: Pouzarella olivacea, 10372TJB HOLOTYPE. 23. Basidiospores. 24. Cheilocystidia. 25. Section through pileus surface, context and hymenium showing inflated cells in subpellis and encrusted cylindrical cells of context, with basidia and other cells of hymenium projecting downwards. 26. Caulocystidia. Scale bars = 10 µm for Figs.22-23, scale bars = 20 µm for Figs. 24 & 26, scale bar = 30 µm for Fig. 25.

Mycobank #: 563277

Pileus very dark brown (6F3 - Negro to 7F3 grayish brown) overall at first, with expansion disc remaining black or very dark brown and with dark blackish radiate striate lines to margin, or pileus becoming paler grayish brown (5D3 Nougat to 6D-E3 Café-au-lait/grayish brown) (hygrophanous?) uniformly on some specimens, 6-10 mm broad and when strongly conical 6-10 mm high, convex, campanulate or broadly conical (variable in cap shape), margin decurved or plane, surface finely and densely squamulose overall, squamules and fibrils silvery translucent or pale brownish, moderately or becoming strongly sulcate to near disc with age. Lamellae grayish brown (6D3 or 5C3 brownish orange), adnexed or deeply adnexed, subdistant, broad (3 mm deep), edges densely granulate-fimbriate scalloped. Stipe concolorous with pileus or slightly paler, 1-1.5 x 25-35 mm, equal, terete, with silvery grayish brown erect agglutinated squamules or fibrillose-pubescence overall and bronze strigosity over the base, very fragile. Odor not distinctive, taste not tried.

Basidiospores 16.2-23.5 x 9.7-12.1 µm (n=32/2, 19.06 ± 1.85 x 10.95 ± 0.76, Q = 1.43-2.09, Qm = 1.74 ± 0.14), heterodiametric, 7-9 (-10) angled in profile view, 8-9 angled in face view and mostly 5 rounded angles in polar view. Basidia 4-sterigmate, short stout or broadly clavate, 32.4-46 x 14.6-16 (-18) µm. Cheilocystidia pale or dark brownish, sphaerical, globose, pyiform or short-broadclavate, often with an inflated supporting cell or two, not evenly distributed along lamella edge, in clusters (producing the granulatefimbriate appearance), 28-70 x 16-28 µm, with slightly brownish thickened walls (0.5 µm). Pleurocystidia absent. Lamella trama a dark brown tissue of parallel, heavily dark brown encrusted mostly cylindrical hyphae, 3.2-14 µm in diam, subhymenium composed of pseudoparenchymatous tissue, tightly compacted. Pileipellis a dark brown or golden brown layer of repent cylindrical or slightly inflated hyphae producing ascendant scattered or closely packed clusters of strongly inflated, chains of sausage-shaped cells, with end cells mostly with tapered apices, 46-120 x 12-18 (-20-34) µm, brown pigment intraparietal mostly and the walls appearing striated internally from the stretched inner wall material, not obviously encrusted; subpellis composed of inflated, pseudoparenchymatous tissue, 30-40 µm deep, with individual inflated cells 14-30 µm in diam. Pileus context a dark brown layer of radially arranged, mostly cylindrical cells, that are heavily dark brown encrusted, 4-16 µm in diam. Stipitipellis a pale yellow brown layer of repent, cylindrical hyphae, 3.2-10.5 µm in diam, giving rise to scattered clusters of erect brown colored, chains (mostly 3-5 cells) of inflated"caulocystidia", wall pale brown, slightly thickened (0.5-1 µm), terminal cells in chains distinctive and mostly tapered or broadly ventricose or fusoid, some clavate, 46-88 x 8-26 µm. Clamp connections none.

Etymology: variable in color of pileus (shades of dark grayish brown) and pileus shape (ranging from convex to campanulate to broadly conical), but constant in distinctive microscopic characters.

Habit, Habitat, Fruiting period: scattered on soil among decaying plant debris, in cool montane tropical broad-leaved forests. March.

Material Examined: ARGENTINA: Salta Province, Baritú National Park, on ridge above village of Lipeo under densely canopied forest on moderately steep north facing slope, 21-III-2011, 22° 25' 31.0"S 64° 44' 29.1"W, 1213 m. a. s. l., collecting team T. J. Baroni, E. Albertó, B. E. Lechner, N. Niveiro, leg. 10380 T. J. Baroni (BAFC; CORT). JuJuy Province, Calilegua National Park, Tatapua Trail, 25-III-2011, 23° 45' 40.09"S 64° 51' 06.09"W, 633 m. a. s. l., collecting team T. J. Baroni, B. E. Lechner, E. Albertó and N. Niveiro, leg. 10414 T. J. Baroni (Holotype - BAFC; Isotype - CORT).

Pouzarella variabilis possesses a suite of features that make this taxon distinctive in the genus. The strongly inflated cells in the subpellis, coupled with the squamulose-pubescent pileus and stipe surfaces and the thick-walled cheilocystidia, place this species in Pouzarella Section Dysthales Subsection Inflatae (Mazzer, 1976). The rather large basidiospores make this species comparable to the North American P. fibrillosipes (Murrill) Mazzer and they do share similarly shaped cheilocystidia, basidiospores and obviously inflated hyphae in the subpellis. Even the pileus color in P. fibrillosipes is noted as variable (Mazzer, 1976) ranging from dark brown to nearly black or"sometimes with deep smoky gray" colors. However, P. variabilis has broader basidiospores (7.5-10µm for P. fibrillosipes vs. 9.7-12.1 µm for P. variabilis), very differently shaped end cells on the"caulocystidia" (cylindrical for P. fibrillosipes but sharply tapered for P. variabilis), a distinctly sulcate pileus surface (versus the smooth, translucentstriate pileus surface of P. fibrillosipes) and the ecology of these two are quite different (soil for P. variabilis vs. wet mucky soil and Sphagnum moss for P. fibrillosipes).

B. Accounting of Pouzarella ferreri var. ferreri in Argentina

Pouzarella ferreri T. J. Baroni, Perd.-Sánch., S. A. Cantrell var. ferreri, North American Fungi 3:243. Figs. 6, 27-30


Figs. 27-30. Pouzarella ferreri var. ferreri. 27 & 29 10379TJB, 28 & 30 10394TJB. 27. Basidiospores. 28. Cheilocystidia. 29. Section through pileus surface, context and hymenium showing surface hyphae, inflated cells of subpellis, context and hymenium. 30. Caulocystidia. Scale bar = 10 µm for Fig. 27. Scale bars = 20 µm for Figs. 28-30.

Pileus deep red or ruby red at first, becoming orange red (6C8 to 7C-D8 - Brownish Orange, Copper (red), Burnt Sienna), margin golden yellow from fibrils and squamules, entire surface and fibrils staining black from handling, 3-13 mm broad, convex or conical-convex, sulcate or moderately sulcate to near disc, reddishochre squamulose and fibrillose over most of the surface, more sparsely yellow fibrillose projecting over the margin. Lamellae very pale and nearly white at first, but soon grayish (6C-D2 - brownish grey), adnexed-arcuate or deeply sinuate, subdistant or ± distant, broad (4 mm), edges fimbriate or granulate-fimbriate, turning black on edges and faces where injured. Stipe pale or dark lemon or golden yellow at first, becoming darker grayish brown with age under the colorful squamules and fibrils, becoming golden brownish with golden yellow or golden orange or reddish-ochre pubescence overall, rapidly becoming black from handling, 1-1.5 x 30-50 mm, equal, terete, with buffy or grayish buffy strigose base. Odor not distinct, taste not tried. All surfaces and superficial fibrils turning black from handling and upon drying.

Basidiospores 13-19.4 x 8.5-11.3 µm (n = 40, 17.2 ± 1.33 x 10.4 ± 0.67, Q = 1.31- 1.92, Qm = 1.65 ± 0.10), heterodiametric, 7-8 angled in profile view, 7-9 angled in face view, 5-6 rounded angular in polar view. Basidia 4-sterigmate, clavate, with abundant aborted basidia with dark brown pigment in 3%KOH. Cheilocystidia pale brown or ± hyaline, brown from intraparietal brown pigment, abundant, inflated and often sphaerical, broadly pyriform, broadly fusoid or clavate, (24-) 30-56 x 20- 24 (-28). Pleurocystidia absent. Lamella trama brown layer of parallel or ± interwoven cylindrical or inflated hyphae, 6-20 µm in diam, encrusted with brown pigments, subhymenium pseudoparenchymatous, composed of rounded cells, 2-3 cells deep. Pileipellis of two distinct layers, the upper layer is dark brown and in two distinct zones, most of the pileus has repent or ascendant hyphae in chains of inflated cells with unevenly thickened walls and intraparietal brown pigments and not obviously encrusted, terminal cells in these chains broadly cylindrical-ellipsoidal, subclavate or some with tapered apices, cells 14-28 µm in diam, the other zone along the margin where the projecting fibrils and squamules are located is composed of two types of cells in chains, the cells making up the fibrils on the margin are 10-20 µm in diam, mostly cylindrical but with a tapered terminal cell, and the walls are dark brown, there is also a layer (repent) at the margin composed of chains of ± hyaline, greatly inflated and mostly sphaerical cells, 24-62 µm in diam, all cells in the pellis with unevenly thickened internal walls with intraparietal brown pigments, encrusting pigments not obvious but occasionally finely adorning the surface of the cells; subpellis composed of a distinct layer, several cells deep (60-100 µm in thickness) of inflated, pseudoparenchymatous cells, 10-40 µm in diam. Pileus context a pale brown layer of radially arranged cylindrical or inflated hyphae, 4-20 µm in diam, mostly encrusted with brown pigment. Stipitipellis producing erect or ascendant chains of 3-5 (-9) brown, ± cylindrical or slightly inflated"caulocystidia", end cells elongate tapered, cylindrical, subclavate or broadly fusoid, 58-117 x 14-28 µm, walls of these cells irregularly thickened on inside wall as in pileus surface hyphae, producing an undulating sculpted effect and with intraparietal brown pigments. Clamp connections absent.

Habit, Habitat, Fruiting period: gregarious or scattered on humus in a wet seepage area on hillside or on well-rotted wood and soil and decaying plant debris next to the wood, in cool montane tropical broad leaved forests. March.

Material Examined: ARGENTINA: Salta Province, Baritú National Park, on ridge above village of El Lipeo on trail to"Thermal Springs" under densely canopied forest on moderately steep northern slope, 21-III-2011, 22° 25' 31.0"S 64° 44' 29.1"W, 1213 m. a. s. l., collecting team T. J. Baroni, E. Albertó, B. E. Lechner, N. Niveiro, leg. 10379 T. J. Baroni (BAFC; CORT). Same general location, but on trail above road to Baritú Village on ridge and south facing slope above El Lipeo, 22-III-2011, 22° 26' 03.4"S 64° 44' 26.4"W, 1255 m. a. s. l., same collecting team, leg. 10394 T. J. Baroni (BAFC; CORT)

Pouzarella ferreri is an attractive agaric with the reddish orange pileus and golden yellow stipe all adorned with long projecting squamules and fibrils that are reddish orange, reddish ochre, golden yellow, or golden orange, and then stain black from handling. Because of these colors, it is one of the easier taxa of Pouzarella to find in a dimly illuminated tropical forest during the daylight hours. Baroni et al. (2008) first described this species from the Dominican Republic and noted that it had also been collected in Costa Rica (Halling & Mueller, 2005) and reported from Brazil (Karstedt et al., 2007). There are some slight differences in the shape of the terminal cells of the"caulocystidia" for the Argentina collections, they are mostly cylindrical or tapered and not with swollen or clavate shapes, but the main characters are the same. We checked the HOLOTYPE of P. ferreri var. ferreri (8454TJB) to confirm that the subpellis of the pileus is composed of inflated cells since that feature was not reported in the original description and indeed this inflated layer was found on the type. This is the first report of P. ferreri for Argentina.

Pouzarella ferreri is distinguished by the colors of the basidiomata, the black staining reactions of all fresh tissues when handled and bruised, the strong darkening to fuscous or fuscous brown colors upon drying the basidiomata, and the inflated sphaerical or broadly pyriform or short inflated fusoid or clavate cheilocystidia. Baroni et al. (2008) noted the differences between P. ferreri and two Pouzarella species, P. aureocrinita and P. erinacea, described from northern Argentina that are depicted as having red or reddish brown densely squamulose or short bristle-like fibrillose pileus surfaces. Both of these taxa have grayish colored stipes; pileus, stipe and lamella surfaces that do not stain black from handling; and these species are described as lacking cheilocystidia. The caulocystidia of P. aureocrinita are very thick-walled and tapered, thus what Mazzer (1976) called"setiform", and unlike those of P. ferreri. The setiform caulocystidia would thus place P. aureocrinita in Section Pouzarella along with species like P. strigosissima (Rea) Mazzer.

Pouzarella erinacea, although described as lacking cheilocystidia and not depicted as turning black when handled, is morphologically most similar to P. ferreri due to the colors of the pileus, the size and shape of the basidiospores, the shapes of the pileipellis and stipitipellis hyphae, and the"pale yellow tinge" on the stipe. Two items in Horak's (1983) description of that species suggest a re-study of the type of P. erinacea must be carried out in the future to determine if these taxa are conspecific: Horak describes the lamella edges as subserrate (?granulate-fimbriate) which might suggest clusters of cheilocystidia that were missed if the sections were not made through these areas; and for the pileipellis he described the "terminal cells as in the caulocystidia, encrusted with brown pigment which dissolves pale redbrown in KOH". This latter feature is similar to the macroscopic discoloration we find when mounting tissues in 3% KOH for microscopic observation. We see a rose or vinaceous solution produced in the medium, but for our collections of P. ferreri we cannot confirm obvious encrusting pigments on the pileus surface hyphae that dissolve and produce a pale red-brown solution under microscopic view.

Discussion

Some final comments on Pouzarella reported for Argentina

Singer & Digilio (1952, p. 429) also listed Rhodophyllus squamifolius (= Pouzarella squamifolia (Murr.) Mazzer) from Argentina, but provided a description of an agaric with a dark colored pileus ("negro o casi negro") and a dark colored stipe ("fuliginoso hasta negro"). Mazzer (1976) in his monograph of the genus Pouzarella took the description of the colors of the pileus of P. squamifolia directly from the protolog published by Murrill (1915) and the pileus of that species is described as"surface dry melleous, covered with long, loose, shaggy ferruginous-fulvous hairs..." and the color of the stipe is described as"pallid, with conspicuous ferruginous hairs similar to those on the pileus". Clearly, Singer & Digilio (1952) did not have P. squamifolia in hand, based on these obvious color differences. We are not sure which species of Pouzarella this might be based on the description, but a future examination of R. Singer T. 261 (LIL) would be helpful.

Conclusions

We now have documented nine species of Pouzarella from northern Argentina and provide the following key to help with identification of future collections.

Key to species of Pouzarella from Argentina

1. Pileus red, reddish orange or reddish brown ............................................................................................ 2
1. Pileus grayish, golden brown, brown or fuscous, without obvious reddish hues ..................................... 4

2. Tissues turning black (nigrescent) after being handled or bruised, cheilocystidia present and mostly globose or broadly pyriform ................................................................... Pouzarella ferreri var. ferreri
2. Tissues unchanging, cheilocystidia absent ...................................................................................... 3

3. Caulocystidia narrow (5-8 µm in diam) seta-like with tapered tips and clearly thickened walls (to 1.5 µm), stipe with golden brown or yellow minute bristles and fibrils (Horak, 1983) .................................................................................................................. Pouzarella aureocrinita
3. Caulocystidia broader (to 10 µm in diam), not seta-like, thin-walled, terminal cells conical, fusoid or cylindrical,"stipe gray with yellow tinge, hairy to strigose over whole length" (Horak, 1983) ........................................................................................................................ Pouzarella erinacea

4. Basidiospores less than 13 µm in length (excluding apiculus) ............................................................................................................. Pouzarella parvispora
4. Basidiospores 16-22 µm in length ..................................................................................... 5

5. All tissue of basidiomata turning black from handling (nigrescent) ......................... 6
5. Tissues not nigrescent or only gradually in the lower half of the stipe ..................... 7

6. Pileus dark brown with brown squamules and reddish-orange squamules and fibrils mixed in at the margin of the pileus, stipe robust (up to 3 mm in diam) and densely white hirsute overall ............................................................................................... Pouzarella brunneonigrescens
6. Pileus and stipe olivaceous colored with golden olive brown squamules and fibrils on the pileus and grayish olivaceous pubescence on the stipe, stipe fragile (up to 1.5 mm in diam) ................................................................................................ Pouzarella olivacea

7. Pileus golden brown with faint orange hues, and lower half of stipe entirely wooly-hairy, basal surface and context of stipe blue-black to black ............................................................................................ Pouzarella sepiaceobasalis
7. Pileus fuscous, grayish brown or yellowish brown, lacking orange hues, stipe not densely wooly-hairy and lower half of stipe not becoming blue-black or black ........................................................................................................... 8

8. Basidiospores large, 16-23 µm long, subpellis with inflated pseudoparenchymatous tissue,"caulocystidia" multi-celled with terminal cells mostly tapered or fusoid or broadly ventricose .................................................................................. Pouzarella variabilis
8. Basidiospores shorter, 11.3-17.8 µm, subpellis lacking inflated cells, caulocystidia single clavate cells ....................................................................................... Pouzarella sulcata

Acknowledgements

The lead author is pleased to acknowledge funding for this work from the following agencies at his institution: State University of New York (SUNY) Research Foundation; SUNY Cortland College Development Foundation, Administrative Offices (Provost and Dean) and Faculty Development Center; and a United University Professions Individual Achievement Award. Most of the comparative work needed for this study originally came from two previous National Science Foundation Grants administered by the Biotic Surveys and Inventory Program to the State University of New York, College at Cortland (DEB9525902 and DEB0103621). The fieldwork was partially supported by the research project PIP 112-200801-0602 from Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET, Argentina). We are also grateful to the Argentinean National Parks Service for permission to collect in protected areas and we are most grateful to Dr. Adriana Hladki for her kind assistance during the field trips in the areas around Tucuman.

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Original recibido el 15 de Septiembre de 2011;
primera decisión: 11 de Enero de 2012;
aceptado el 11 de Enero de 2012.

Editor responsable: Carlos Urcelay.