Ocurrence of ectomycorrhizal, hypogeous fungi in ... - Mycologia

Mycologia, 100(5), 2008, pp. 752–759. DOI: 10.3852/07-182 

# 2008 by The Mycological Society of America, Lawrence, KS 66044-8897 

Ocurrence of ectomycorrhizal, hypogeous fungi in plantations of exotic tree 

species in central Argentina 

Eduardo R. Nouhra 1 

Laura S. Dominguez 

Graciela G. Daniele 

Silvana Longo 

Instituto Multidisciplinario de Biología Vegetal 

(CONICET), Casilla de correo: 495, 5000, Córdoba, 

Argentina 

James M. Trappe 

Department of Forest Science, Oregon State University, 

Corvallis, Oregon 97331-5752 

Andrew W. Claridge 

Department of Environment and Climate Change 

(NSW), Reserve Conservation Unit, Parks and Wildlife 

Division, Southern Branch, P.O. Box 2115, 

Queanbeyan, New South Wales 2620, Australia 

Abstract: Eleven hypogeous, ectomycorrhizal species 

of Basidiomycota, including two new species, and one 

of the Zygomycota were collected in exotic tree 

plantations in Córdoba Province, Argentina. Descomyces 

fusisporus sp. nov., D. varians sp. nov., 

Hydnangium archeri (Berk.) Rodway, H. carneum 

Wallr., Hysterangium gardneri E. Fisch. and Setchelliogaster 

tenuipes (Setch.) Pouzar were associated with 

Eucalyptus spp. Endogone lactiflua Berk., Hymenogaster 

lycoperdineus Vittad., H. griseus Vittad., H. 

rehsteineri Bucholtz, Rhizopogon couchii A.H. Sm. and 

R. roseolus (Corda) Th. Fr., were associated with 

various northern hemisphere tree species. Descriptions 

are provided to aid identification of the 

hypogeous fungi in exotic plantations of Argentina. 

Key words: Argentina, Descomyces spp. nov., 

mycorrhizal, sequestrate fungi 

INTRODUCTION 

From the early 20th century to the present Eucalyptus, 

Pinaceae and various northern hemisphere deciduous 

species have been planted extensively in Argentina 

and elsewhere in the southern hemisphere (Dunstan 

et al 1998). In Córdoba Province of central Argentina 

more than 41 000 ha have been planted with Pinus 

elliottii, P. halepensis, P. patula, P. radiata and P. taeda 

(Izurieta et al 1998, Mangieri et al 1987) and 

approximately 4500 ha with E. camaldulensis, E. 

Accepted for publication 23 April 2008. 

1 Corresponding author. E-mail: nouhra@imbiv.unc.edu.ar 

752 

sideroxylon, E. tereticornis and E. viminalis (Golfari 

1985). 

Various hypogeous, ectomycorrhizal fungi were 

introduced accidentally but fortuitously along with 

these economically important trees and became 

established in the exotic plantations as vital symbionts 

of the trees. The hypogeous truffles (Ascomycota) 

and ‘‘truffle-like fungi’’ (Basidiomycota and Zygomycota) 

comprised species with indehiscent, sequestrate 

sporocarps containing statismosporic spores. Within 

the Basidiomycota morphological and molecular 

studies have confirmed the polyphyletic origin of 

these fungi, and most species currently are placed in 

recognized orders of epigeous mushrooms (Bruns et 

al 1998, Castellano and Beever 1994, Miller et al 2001, 

Peintner et al 2001, Thiers 1984). 

Meanwhile ectomycorrhizal fungi associated with 

Pinaceae and Eucalyptus have been surveyed extensively 

elsewhere in the southern hemisphere (Birch 

1937, Chu-Chou 1979, Chu-Chou and Grace 1983, 

Dunstan et al 1998, Garrido 1986, Giachini and 

Oliveira 1997, Giachini et al 2000, Kessell 1927, 

Purnell 1957, Rawlings 1951). Only a few mycologists 

have paid attention to the hypogeous fungi of 

Argentina, especially those associated with native 

and exotic ectomycorrhizal hosts (Crespo and Domínguez 

1994, Lorenzo and Calvelo 2000, Nouhra and 

Becerra 2005, Romero and Blumenfield 2001, Spegazzini 

1880, 1887, 1909, Wright 1980). 

The aim of the present study was to determine the 

exotic ectomycorrhizal hypogeous fungi associated 

with exotic, ectomycorrhizal plantations of Córdoba 

Province, Argentina. New taxa are described in full. 

Short descriptions of previously described taxa, 

including diagnostically informative macro- and 

microcharacters, are provided to aid identification 

by local mycologists. 

MATERIALS AND METHODS 

Study sites.—Pines and eucalypts plantations in Córdoba 

Province are located approx. at 31u209–32u409S and 62u009– 

64u509W. The weather in the region is characterized by 

mild, temperate summers and dry winters. Precipitation is 

concentrated Oct–Apr. Eucalypts plantations are better 

established in the eastern part of the province at lower 

altitudes up to 600 m. In this area precipitation is 600– 

900 mm (Vázquez et al 1979). Conifer plantations thrive 

best at higher altitudes, up to 1600 m in the mountains of 

the west, where precipitation increases up to 1200 mm

(Izurieta et al 1998). The two new species of Descomyces were 

discovered in Victoria and New South Wales, Australia, 

during studies of hypogeous fungal diversity and habitat 

relationships (Claridge et al 2000a, b) but had not been 

described formally. 

Sporocarp sampling and sporocarp description.—Sporocarps 

were collected at random in the plantations and, in the case 

of the two new Descomyces spp., also in 1000 m 2 plots 

scattered over E Victoria and SE New South Wales (Claridge 

et al 2000a). Specimens usually were located by raking the 

soil litter and humus; occasional sporocarps were visible 

without raking, having emerged from the soil. Most 

Argentinean samples were collected Dec–Jun, the Australian 

samples of Descomyces Feb–Oct. Data on location and 

associated hosts are detailed for each species along with its 

description. Specimens were cut in half and, after notes on 

fresh characters were recorded, dried on a forced-air 

dehydrator at 640 C. 

Color, size, shape and other macrocharacters of fresh 

specimens were recorded, and most collections were 

photographed in surface and cross-sectional views. Color 

names of fresh and dry specimens are applied in general 

terms by the authors. Microcharacters were recorded from 

sections hand-cut from dried material, mounted respectively 

in water, 5% KOH, Melzer’s reagent, cotton blue and 

phloxine for standard light microscopy. Spore dimensions 

are based on the measurement of at least 15 randomly 

selected, plus largest and smallest spores, including 

ornamentation. Measurements of tissues and spores are 

from mature specimens; when two dimensions are given, 

length always precedes width. 

Specimens are deposited as indicated in the Museo 

Botánico de Córdoba (CORD), Mycological Herbarium of 

Oregon State University (OSC), National Herbarium of 

Victoria (MEL) or the Plant Pathology Herbarium, Orange 

Agricultural Institute, New South Wales (DAR). Identification 

was aided by keys and specific references (Beaton et al 

1984, Bougher and Castellano 1993, Castellano et al 1989, 

Miller 1986, Montecchi and Lazzari 1993, Montecchi and 

Sarasini 2000, Trappe unpubl). 

TAXONOMY 

Associates of Eucalyptus.—Descomyces fusisporus 

Trappe & Claridge, sp. nov. FIG. 1A 

Peridium 100–350 mm crassum; pellis albis hyphis intertextis, 

juventute suprapelle lutea vel luteobrunnea. Gleba 

loculata, tabacina, interdum columella truncata alba vel 

pallide brunnea. Basidia 35–40 3 8–10 mm, clavata, sterigmatibus 

duobus. Sporae brunneae, fusoides-citriformes, 

papillatae, 17–23 3 9–12 mm, reticulo partiali vel completo, 

minute labyrinthiformi 0.5 mm alto ornatae; sporae in 

solutione Melzeri juventute rubrae, maturitate pallide 

aurantiae. Holotypus hic designatus: A.W. Claridge AWC 

349. 

Basidiomata globose, subglobose to irregular, 5–11 

3 5–20 mm. Peridium with a white pellis, in youth 

covered by a loose to appressed, yellow, veil-like 

suprapellis that thins as basidiomata expand to 

NOUHRA ET AL: EXOTIC HYPOGEOUS FUNGI IN ARGENTINA 753 

produce an overall pale yellow to cream color or 

breaks up to reveal the white pellis with only scattered 

patches of suprapellis; in age all peridial tissues tend 

to darken. Gleba loculate, white in youth, by maturity 

cinnamon to dark brown, occasionally with a truncate 

columella up to 3 mm broad. 

Peridium 100–350 mm thick; suprapellis of tangled, 

hyaline to yellow or light yellowish brown, septate, 

thin walled hyphae 3–7 mm broad at the septa, the 

cells often somewhat inflated; pellis of periclinal to 

interwoven, thin-walled, hyaline hyphae 3–5 mm 

broad with occasional, slightly inflated cells; clamp 

connections abundant, inconspicuous. Trama of 

subparallel, thin-walled hyaline hyphae, 3–5 mm broad 

with occasional inflated cells. Subhymenium not 

differentiated from the trama. Basidia hyaline, thinwalled, 

clavate, 35–40 3 8–10 mm, with two robust 

sterigmata. 

Spores brown, fusoid-citriform, papillate, with a 

prominent sterigmal attachment, 17–23 3 9–12 mm 

(Q 5 1.7–2.2) including the ornamentation 0.5 mm 

tall of a minute, partial to complete, labyrinthiform 

reticulum; apical papilla 1–4 3 3–4 mm, smooth, 

usually dark brown in youth, later fading to pale 

brown; sterigmal attachment 3 31.5–2 mm; spores in 

KOH yellowish brown, in Melzer’s reagent red 

(strongly dextrinoid) in youth, at maturity light 

orange (weakly dextrinoid). 

Etymology. Latin fusisporus in reference to the 

fusoid-citriform spores. 

Distribution, habitat, hosts and season. In Argentina 

in soil and leaf litter in a plantation of Eucalyptus 

cinerea, May. In Australia in gully bottoms and on 

sheltered slopes and ridges in mixed, natural, wet, 

montane forests consisting of various mixtures of 

Acacia dealbata, A. implexa, A. melanoxylon, Eucalyptus 

dalrympleana, E. fastigata, E. nitens, E. pauciflora, E. 

radiata, E. stellulata and E. viminalis; Apr, May and 

Jun. 

HOLOTYPE. AUSTRALIA: VICTORIA: Rich Forest 

Management Block, Jack Road 0.7 km N of Winter Road, 

17 Apr 1996, A.W.Claridge, AWC 349 ( MEL, OSC). 

Specimens examined. ARGENTINA: CÓRDOBA: Punilla, 

Establecimiento Ayke, 3 km S of Cuesta Blanca, 18 May 

2002, L. Domínguez 2224 (CORD, OSC). AUSTRALIA: 

VICTORIA: East Gippsland, Alpine National Park, Black 

Mountain Road 3.2 km E. of Rams Horn Track, A.W. 

Claridge and W. Colgan III, 14 May 2001, AWC 3329 (MEL, 

OSC); Bentleigh Plains Forest Reserve, Bentleigh Creek 

0.4 km NE of crossing of Bentleigh Plains Road, A. Giachini, 

18 May 2001, AWC 3619.2 (MEL, ISOTYPE. OSC); 

Errinundra National Park, Gunmark Road 1.3 km SE of 

Survey Road, J. Trappe, A.W. Claridge and A. Jumpponen, 26 

May 2001, AWC 4041 (MEL, OSC); Gunmark Road 1.6 km 

SE of Junction Survey Road, W. Colgan III, 26 May 2001, 

AWC 4092 (MEL, OSC); Gunmark Road 2.3 km SE of Tea

754 MYCOLOGIA 

FIG. 1. A.Descomyces fusisporus sp. nov., basidiospores. B–D. Descomyces varians sp. nov. B. Basidiospores. C. Basidiomata. 

D. Peridial polycystoderm. Bar: A, B 5 10 mm; C 5 5 mm; D 5 15 mm. 

Tree Flat Picnic Area, J. Trappe, A.W. Claridge and A. 

Jumpponen, 11 Jun 1996, Trappe 19455 (MEL, OSC); Coast 

Range Road at Goonmirk Rocks Road, A. Giachini, 26 May 

2001, AWC 4068 (MEL, OSC); Rich Forest Management 

Block, Jack Road 0.7 km N of Winter Road, 17 Apr 1996, 

AWC 348 & 351, (MEL, OSC); Gap Scenic Reserve, Gap 

Road 5.2 km E of Bonang Highway, 27 May 1999, AWC 

2603.2 (MEL, OSC). 

Comments. Descomyces fusisporus as presently known 

has a relatively small distribution in Australia, limited 

to wet, montane forests in East Gippsland, Victoria. It 

differs from all other species in the genus by its 

combination of fusoid-citriform spores ornamented 

with a partial to complete, low, labyrinthiform 

reticulum and a brown apical papilla in youth, plus 

lack of a peridial epithelium of inflated cells. D. 

giachinii Trappe et al, introduced to Brazil with 

eucalypts, has a much broader distribution and range 

of habitats (Giachini et al 2000). Its spores on average 

are more broadly citriform than those of D. fusisporus 

and the ornamentation is a reticulum 1 mm tallthat 

tends to be alveolate instead of labyrinthiform. 

Descomyces varians Trappe & Claridge, sp. nov. 

FIG. 1B–D 

Peridium 100–300 mm crassum; pellis albis hyphis intertextis, 

juventute suprapelle lutea vel luteobrunnea. Gleba 

loculata, tabacina. Basidia 25–30 3 6–7 mm, clavata, 

sterigmatibus duobus. Sporae brunneae, citriformes, papil-

latae, 11–16 3 9–12 mm, reticulo partiali vel completo, 

minute labyrinthiformi 0.5–1(–1.5) mm alto cum verrucis 

irregularibus et cristis brevibus ornatae; sporae in solutione 

Melzeri juventute rubrae, maturitate pallide aurantiae. 

Basidiomata globose, subglobose to irregular, 5–15 

3 5–20 mm. Peridium with a white pellis, in youth 

covered by a yellow, loose to appressed suprapellis 

that thins with expansion of the basidiomata to 

produce a pale yellow to cream color or breaks up 

to reveal the white pellis with only scattered patches of 

suprapellis; in age all peridial tissues tend to become 

yellowish brown. Gleba loculate, pale cream in youth, 

by maturity cinnamon to dark brown. 

Peridium 100–300 mm thick; suprapellis of hyaline 

to yellow or light yellowish brown, septate, thin- to 

thick-walled hyphae 3–7 mm broad at the septa, the 

cells often somewhat inflated; pellis a well developed 

to broken polycystoderm of thin-walled, vesiculose 

cells 10–25 3 10–30 mm; subpellis of hyaline, interwoven 

hyphae; clamp connections abundant, inconspicuous. 

Trama of hyaline, subparallel hyphae 2– 

5 mm broad at the septa, many cells inflated. 

Subhymenium of hyaline, interwoven hyphae 2– 

3 mm broad. Basidia 2-spored, hyaline, clavate, 25–30 

3 6–7 mm. 

Spores broadly citriform, papillate, with a prominent 

sterigmal attachment, 11–16 3 6.5–11 mm 

excluding ornamentation (Q 5 1.3–1.6), 12–16 

(–17) 3 9–11(–12) mm including the ornamentation, 

0. 5–1(–1.5) mm tall (often tallest near the spore base) 

of a minute, partial to complete, labyrinthiform 

reticulum or irregular warts and short ridges; apical 

papilla 1–2 3 3 mm, smooth; sterigmal attachment 2 

3 2 mm; spores in KOH yellowish brown, in Melzer’s 

reagent red (strongly dextrinoid) in youth, at 

maturity light orange (weakly dextrinoid). 

Etymology. Latin varians (varying), in reference to 

the variable development of the polycystoderm epithelium 

and spore ornamentation. 

Distribution, habitat, hosts and season. Central Argentina, 

under a thick litter layer in a plantation of 

Eucalyptus camaldulensis, May. Southeastern Australia, 

in dry to wet sclerophyllous forests in mixed stands 

variously containing Acacia longifolia, A. mearnsii, A. 

melanoxylon, A. mucronata, Angophora floribunda, 

Eucalyptus albens, E. botryoides, E. cypellocarpa, E. 

globoidea, E. longifolia, E. microcarpa, E. muelleriana, 

E. obliqua, May–Oct. 

HOLOTYPE: AUSTRALIA: VICTORIA: Murrangowar 

Forest Management Block, Princes Highway 0.4 km E of 

Bendoc Ridge Road. A.W. Claridge, J. Trappe & T. Lebel, 26 

May 1999, AWC 2555 (MEL; OSC). 

Specimens examined. ARGENTINA: CÓRDOBA: Calamuchita, 

Villa Alpina, L. Domínguez, 12 May 2001, LSD 

2002 (CORD); 11 km before La Cumbrecita, L. Domínguez, 

14 May 2002, LSD 2209 & LSD 2214 (CORD). AUSTRA- 


LIA, NEW SOUTH WALES: Ben Boyd National Park, 

Bittangabee Bay, at walking track crossing of Bittangabee 

Creek, A.W. Jumpponen, 2 Jun 1999, AWC 3052 (DAR, 

OSC); Nadgee State Forest, Mines Forest Road 1.2 km W. of 

Buckland Road, A. Giachini & A.W. Claridge, 31 May 2001, 

AWC 4431 (OSC). VICTORIA: East Gippsland: Cabbage 

Tree Forest Management Block, Towser Link Track 0.7 m S 

of Douthat Road, A.W. Claridge, 10 Oct 1995, AWC 49 & 

51 (MEL, OSC); Omeo Highway, 2 km S of Ash Range 

Road, A.W. Claridge, 21 May 1999, AWC 2378 (MEL, OSC); 

Rich Forest Management Block, Puggaree Road at Arte 

Shortcut Road, A.W. Claridge, 27 Feb 1996, AWC 160 

(MEL, OSC). 

Comments. Descomyces varians differs from D. giachinii 

Trappe et al (Giachini et al 2000) by having a 

polycystoderm epithelium and generally smaller 

spores with a lower ornamentation. This pair of 

species is analogous to the D. albus and D. albellus 

pair, which do not have partially to fully reticulate 

spores and differ primarily by D. albus having a 

polycystoderm epithelium and D. albellus lacking that 

(Bougher and Castellano 1993). All Descomyces spp. 

appear to have originated in Australia as mycorrhizal 

associates of Eucalyptus spp., and many probably were 

introduced as hitch-hikers on roots of eucalypt 

seedlings shipped from Australia to other continents. 

Most also appear adapted to a wide range of tree ages 

and habitats. 

Hydnangium archeri (Berk.) Rodway 

Basidiomata ellipsoid, usually basally depressed 

with a hyphal basal attachment, 25 3 20 mm. 

Peridium yellowish brown, thin, 60–100 mm thick, 

composed of interwoven strands of parallel hyphae up 

to 16 mm thick. Gleba whitish to cream, not gelatinized, 

with small irregular chambers and a narrow 

columella. Hymenium, a palisade of cylindrical to 

clavate basidia, bearing two large straight sterigmata. 

Spores globose, hyaline, (9–)10.5–14(–15.5) mm diam, 

ornamented with conical spines 1.5–2.5(–3.5) 3 0.2– 

2 mm. Clamp connections present. 

Habitat, hosts and season. Hypogeous under a thick 

organic layer, on top of the mineral soil, in plantation 

of E. camaldulensis, May 



2204 (CORD, OSC). 

Comments. Basidiomata of H. archeri are yellowish 

brown and its spores are smaller but have larger spines 

than the pink to rose H. carneum (Beaton et al 1984). 

It is infrequent in native Eucalyptus forests of Australia. 

Hydnangium carneum Wallr. 

Basidiomata globose, subglobose, depressed or 

irregular, 10–25 mm broad. Peridium pinkish white

756 MYCOLOGIA 

or rose when fresh, drying tan to brownish, up to 

300 mm thick, of clamped, cylindrical hyphae intermingled 

with sphaerocysts and some inflated hyphae. 

Gleba pinkish white, fading whitish, then tan when 

dry, of labyrinthiform chambers; a small columella 

and a sterile basal pad usually present. Spores 

globose, hyaline, 12–17 mm broad excluding the 

ornamentation of spines and narrow cones 1.5–2 3 

0.5–2 mm. 

Habitat, hosts and season. Hypogeous under a thick 

layer of leaves and debris in plantation of E. 

camaldulensis, May and Nov. 


11 km before La Cumbrecita, L. Domínguez, 14 

May 2002, LSD 2212 (CORD); Estancia Alta Vista, Atos 

Pampa, L. Domínguez, 14 May 2002, LSD 2208 (CORD); 

Villa Alpina, L. Domínguez, 17 Nov 1997, LSD 2294 

(CORD). Colón: Estancia Santo Domingo, L. Domínguez, 

30 May 1997, LSD 2295 (CORD). 

Comments. Among the more widely distributed and 

common hypogeous fungi of eucalypts in Australia, 

this species has been dispersed around the world with 

eucalypts. We have found it fruiting in enormous 

numbers in plantations in California. 

Hysterangium gardneri E. Fisch. 

Basidiomata subglobose to irregular, 0.5–2 cm 

broad; rhizomorphs white, well developed at the base 

and sides of basidiomata, abundant within the 

substrate. Peridium smooth, somewhat rubbery when 

fresh, white to light brown, bruising pale pink to 

pinkish brown, drying pale brown, 400–600 mm thick, 

of cylindrical yellow hyphae, 2–7 mm broad at the 

septa, many cells inflated up to 15 mm, with crystals 

adhering to the surface; clamp connections scattered, 

inconspicuous. Gleba olive green to grayish green, 

drying dark green, with elongate locules and a 

cartilaginous, dendroid, translucent columella. 

Spores ellipsoid with a truncate base, 8–10.5 3 3– 

4 mm, surface smooth but enclosed in an irregular, 

tightly adherent to wrinkled exosporium. Basidia 

hyaline, cylindrical, mostly 6-spored. 

Habitat, host and season. Hypogeous in soil under 

leaves and litter in mixed plantation of Eucalyptus 

cinerea and E. camaldulensis. Some basidomata were 

emergent in exposed soil, May. 



May 2002, LSD 2210 & 2211 (CORD, OSC). Punilla, 

Establecimiento Ayke, 3 km south of Cuesta Blanca, L. 

Domínguez, 18 May 2002, LSD 2222 (CORD). 

Comments. The spore size and abundance of hyphae 

and rhizomorphs adhering to the sides and the 

base of the basidiomata, differentiate H. gardneri from 

other Eucalyptus associated Hysterangium spp. known 

at present, although the genus has not been studied 

fully in Australia. The variation of shape and size of 

peridial hyphae can be misinterpreted as a multilayered 

structure. H. gardneri is common and widely 

distributed in Australia and occurs in Eucalyptus 

plantations around the world. 

Setchelliogaster tenuipes (Setch.) Pouzar. 

Basidiomata up to 2.5 cm broad; pileus globose to 

subglobose, stipitate, reminiscent of an unexpanded 

agaric with the margin attached to the stipe. Peridium 

smooth, dry and glabrous, reddish brown to orange 

brown, up to 300 mm thick, the pellis an epithelium of 

ellipsoid to subglobose or pyriform, hyaline cells. 

Stipe, cylindric, 1–3 cm long, solid, white to creamy 

white. Gleba with elongated chambers radiating from 

the stipe columella, concolorous with the peridium or 

paler. Basidia 2-spored. Spores asymmetric, citriform, 

yellowish brown, 8–9.5 313–14.5 mm, finely verrucose, 

with an apical papilla, lacking a germ pore. 

Habitat, host and season. Subhypogeous under litter 

in E. camaldulensis plantation, May. 



May 2001, LSD 2215; same locality, L. Domínguez, 20 May 

2002, LSD 2207 (CORD). 

Comments. This taxon is widespread in Australia 

and has been collected in Argentina (Patagonia and 

Buenos Aires), Uruguay and North America under E. 

globulus (Setchell 1907, Wright 1980). 

Associates of Pinus and various northern hemisphere 

deciduous trees.—Endogone lactiflua Berk. & Broome. 

Zygomata subglobose to ellipsoid, reniform or 

irregular, up to 9 3 14 mm, often with a basal 

attachment to the soil, exuding a sticky white fluid in 

cut surfaces of fresh, young specimens. Peridium thin, 

white to cream, partially to completely disappearing 

in old specimens. Gleba often with a small, sterile, 

basal pad at the base, with uniformly distributed 

spores. Spores globose to ellipsoid, light yelloworange, 

70–180 3 94–190 mm, the wall 5–10 mm thick, 

consisting of a yellow laminar inner wall up to 8 mm 

thick, and a thin outer wall 1–3 mm that stains reddish 

brown in Melzer’s reagent. Mature spores are 

enclosed in hyphal mantles composed of several 

layers of thin-walled, yellow, much interwoven hyphae 

that at maturity become orange to red in Melzer’s 

reagent. Gametangia up to 65(–80) mm broad, long 

persistent. 

Habitat, host and season. Hypogeous, under Pinus 

elliottii and P. taeda, Apr. 


15 km before Villa Alpina, L Domínguez, 8 Apr 

1999, LSD 2231 (CORD, OSC).

Comments. A cosmopolitan ectomycorrhizal fungus 

in the northern hemisphere, E. lactiflua is common in 

conifer forest nurseries (Fassi et al 1969) and has been 

introduced widely in conifer plantations in the 

southern hemisphere. 

Hymenogaster griseus Vittad. 

Basidiomata globose to irregular, up to 2 cm diam. 

Peridium white, bruising ochraceous, thick, 380– 

500 mm, not separable, the pellis of cylindrical, 

septate, thin-walled hyphae 3–5 mm broad, grading 

into a pseudoparenchymatous subpellis toward the 

gleba. Gleba loculate, labyrinthiform, with a small 

sterile base, whitish to pale ochraceous, drying brown 

to rusty brown; trama of inflated cells. Spores 

ellipsoid, citriform to obovate, 13–20 3 8–11 mm, 

with an inconspicuous apical papilla; perisporium 

closely appressed, wrinkled-verrucose; apiculus short. 

Habitat, host and season. Central Argentina, hypogeous 

in organic soil, under litter of Pinus and Cedrus 

spp., Dec–Mar. 


Atos Pampa, L. Domínguez, 8 Mar 1995, LSD 

1410 (CORD, OSC); La Cumbrecita, E. Nouhra & E. 

Crespo, 9 Dec 1993, LSD 1405 (CORD, OSC). 

Hymenogaster lycoperdineus Vittad. 

Basidiomata subglobose to irregular, 1–3 cm broad. 

Peridium 200–300 mm thick, not separable, smooth, 

white to yellowish white at base grading to light 

yellowish brown or light brown toward the top, 

darkening when bruised, the pellis of cylindrical, 

thin-walled hyphae grading to a subpellis of inflated, 

subglobose cells. Gleba loculate, lacking a columella 

or sterile base, white in youth, by maturity darkened 

to brown. Odor strong, rancid. Spores ellipsoid to 

fusoid, 15–25 3 9.5–14.5 mm, apical papilla conspicuous 

or occasionally low and rounded; perisporium 

closely appressed, wrinkled into small folds and warts. 

Habitat, host and season. Hypogeous, under litter of 

Pinus sp. and Cedrus sp., May. 


Atos Pampa, L. Domínguez, 14 May 2002, LSD 

2217, LSD 2218 & LSD 2220 (CORD). 

Hymenogaster rehsteineri Bucholtz 

Basidiomata subglobose to irregular, 0.3–2 cm 

broad. Peridium up to 270 mm thick, whitish, finally 

pale yellow to brownish gray, becoming dark brown 

where bruised, with a pellis of hyaline, cylindrical 

hyphae and a subpellis of mostly pseudoparenchymatous 

hyphae. Gleba with irregular to elongated 

locules, sometimes with a small sterile base, in youth 

white, at maturity dark brown. Odor of dried 


specimens, strong. Spores fusiform to somewhat 

variable, 15–25(–30) 3 8–13 mm, with a conspicuous 

papilla; perisporium fairly closely appressed, wrinkled, 

verrucose, arranged into small longitudinal 

folds, and crests; apiculus evident. 

Habitat, host and season. Hypogeous in soil, under 

forest litter in various mixtures of Pinus, Betula, 

Quercus and Tilia spp., sometimes including Eucalyptus, 

Nov–Jun. 


Villa Alpina, L. Domínguez, 17 Nov 1997, LSD 

2298 & LSD 2296 (CORD, OSC); La Cumbrecita, E. 

Nouhra, 9 Dec 1993, LSD 1406 (CORD); same locality, L. 

Domínguez, 9 Jun 1995, LSD 2299 (CORD). 

Rhizopogon couchii A.H. Sm. 

Basidiomata up to 60 mm diam, subglobose, 

rhizomorphs appressed to the peridium and abundant 

in soil. Peridium smooth, up to 300 mm thick, 

subtomentose, whitish yellow to yellow, yellow veins 

present, then yellowish brown, spotted red or 

brownish red, some sporocarps bruising vinaceous. 

Peridium characterized by a cutis constituted by 

tightly interwoven cylindrical thin-walled hyphae. 

Gleba loculate, soft when young, then firm, cartilaginous, 

finally dry, whitish to yellowish brown, yellowish 

gray. Spores subcylindrical, elliptical, smooth, not 

truncate, 6–8 3 2–2.5 mm. Brachybasidioles present in 

hymenium, subglobose, to ovate, thin-walled when 

young, then moderately thick-walled, not disarticulating. 

Habitat, host and season. Hypogeous in soil, not 

common, underneath leaves and litter in P. elliottii 

plantation, May. 



2205 (CORD, OSC). 

Rhizopogon roseolus Corda 

Basidiomata up to 4 cm broad, subglobose to 

turbinate, rhizomorphs appressed overall and abundant 

in soil. Peridium smooth, 0.5–1 mm thick, 

fibrillose, white to yellow in youth, soon becoming 

yellowish brown to reddish brown, bruising brownish 

red to vinaceous; KOH reddish brown on peridium. 

Gleba loculate, soft, drying hard, yellowish in youth, 

at maturity brown to olive brown, sometimes with a 

small basal pad. Spores smooth, subfusoid to broadly 

ellipsoid, truncate, 6–12 3 3.5–4 mm. Brachybasidioles 

present in hymenium, subglobose, clavate, 

obovate, 20–25 3 8–12 mm, easily disarticulating. 

Habitat, host and season. Hypogeous to subhypogeous 

in soil, common in P. elliottii, P. radiata and P. 

taeda plantations, Apr and May. 

Specimens examined. ARGENTINA: CÓRDOBA: Cala-

758 MYCOLOGIA 

muchita, Villa Alpina, Estancia La Payana, M. Evdemon, 2 

May 1994, LSD 1408 (CORD); Villa Alpina, L. Domínguez, 

12 May 2001, LSD 2201 (CORD); Atos Pampa, Estancia Alta 

Vista, E. Crespo, 8 Dec 1993, LSD 1404 (CORD). PUNILLA: 

Camino a las Altas cumbres, Estancia La Armonía, L. 

Domínguez, 3 May 1992, LSD 1216 (CORD); Estancia Ayke, 

3 km south of Cuesta Blanca, L. Domínguez, 30 May 1998, 

LSD 2293 (CORD); same locality, L. Domínguez, 12 Apr 

1999, LSD 2292 (CORD). 

Comments. Rhizopogon roseolus has been regarded 

variously as a young specimen of the later named R. 

rubescens Tul. & C. Tul. or a species distinct from R. 

rubescens. Smith and Zeller (1966) not only separated 

the two but also named several other variants as 

additional species. The problem arises from its 

changes in color, bruising reactions and responses to 

KOH applied to the peridium as it passes from young 

to mature stages. Young specimens can be pure white, 

stain deep pink where bruised and turn bright red with 

application of KOH. Later they become yellow, stain 

reddish where bruised and turn dark red with KOH. At 

maturity they become reddish brown to dark reddish 

brown, may merely darken where bruised and turn 

dark brown with KOH. Partly emergent specimens 

sometimes show this full range of variation grading 

from a white base to dark reddish brown, exposed top. 

When KOH is streaked on the peridium from bottom 

to top the reaction is bright red on the white bottom 

grading to dark red on the reddish-yellow side and 

dark brown on the reddish brown top (J. Trappe 

unpubl). Moreover R. roseolus/rubescens has been 

recorded throughout the northern hemisphere, so a 

species complex with different, geographic genotypes 

might be expected (Miller 1986, Molina et al 1999). 

Sorting this out will require critical morphological 

analyses of the variants, including all developmental 

stages, coupled with multiple gene DNA sequencing. 

Until that formidable challenge is met we choose to 

use the oldest name to designate the species or species 

complex. 

DISCUSSION 

Crespo and Domínguez (1994), Nouhra (1999) and 

Nouhra and Domínguez (1998) have reported introduced 

ectomycorrhizal fungi in plantations of exotic 

trees in Argentina. Here we have 12 additions to the 

list, including species native to North America, 

Europe and Australia. Trappe et al described in 

Giachini et al (2000) a new Descomyces species from 

Brazil that they had collected in Australia but had not 

described. The same situation arose in the present 

study, except here we found two undescribed 

Australian Descomyces spp. in Argentinean eucalypt 

plantations. Moreover we have reported for the first 

time in Argentina Rhizopogon couchii from the 

southeastern United States, R. roseolus possibly from 

either the United States or Europe, Hymenogaster 

griseus, H. lycoperdineus and H. rehsteineri from 

Europe, and Hydnangium carneum, H. archeri and 

Hysterangium gardneri from Australia, all introduced 

along with their native hosts in soil or seedlings. The 

remaining species, Endogone lactiflua and Setchelliogaster 

tenuipes, have been cited previously for Argentina. 

Giachini et al (2000) observed that diversity of 

ectomycorrhizal fungi in South American Pinus and 

Eucalyptus plantations appears to be higher than in 

other southern hemisphere locations, where these 

economically important trees also have been introduced. 

Rhizopogon spp. have established successfully 

where pines have been introduced (Molina and 

Trappe 1994) and often have become dominant 

species of the exotic ectomycorrhizal flora (Molina 

et al 1999). Similarly Hymenogaster spp. have established 

in Córdoba province, where conifers and 

deciduous trees were introduced by European settlers. 

Considering the relatively few Hymenogaster 

specimens we collected, the genus seems not to be as 

dominant as Rhizopogon. 

Eucalyptus associates (Descomyces, Hydnangium, 

Hysterangium and Setchelliogaster spp.) have been 

recorded in many regions of the world. Setchelliogaster 

tenuipes was collected in Argentina and Uruguay 

(Wright 1980). Probably Hydnangium carneum, Hysterangium 

gardneri and various Descomyces spp. also 

are present elsewhere in Argentina where Eucalyptus 

spp. have been introduced. 

ACKNOWLEDGMENTS 

This study was supported by SECYT (Secretaria de Ciencia y 

Técnica), No. 150/02 and 62/03, Córdoba, Argentina. 

Trappe’s participation was supported in part by the USFS 

Pacific Northwest Research Station, Corvallis, Oregon. 

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