Cryptogamie, Mycol.

, 2001, 22 (1): 57-66

57

2001 Adac/l~ditions scientifiques et m6dicales Elsevier SAS. Tous droits r6serv6s

S0181158401010569/FLA

Notes on the genus Tulostoma in H. Kreisel's Herbarium

Gabriel MORENO a*, Hanns KREISEL b & Alberto ALTES a
aDpto, de Biologfa Vegetal, Universidad de Alcalti,
28871 Alcal(t de Henares, Madrid, Espaha
b Ernst-Moritz-Arndt-Universitiit, Fachbereich Biologie,
Institut fiir Mikrobiologie und Molekularbiologie, 17487 Greifswald, Germany
(Received September 2000, accepted November 2000)

Abstract m A revision of 89 collections of the genus Tulostoma, deposited in the Prof. H.

KREISEL's personal herbarium (Germany) is presented. These belong to thirteen species: T.

armillatum, T. brumale, T. fimbriatum, T. giovanellae, T. kotlabae, T. melanocyclum, T. moravecii,

T. obesum, T. pulchellum, T. pusillum, T. squamosum, T. striatum and T. subsquamosum. Some of
them are reported for the first time from some countries. Photomicrographs under SEM of the most
interesting species are included. 2001 Adac / l~litions scientifiques et m6dicales Elsevier SAS
Tulostoma I Gasteromycetes / H. Kreisel / taxonomy / chorology

~TRODUCTION
In the frame of a world-wide revision of some genera of Gasteromycetes, the
examination of material deposited in a great number of herbaria is a fundamental task. A
great part of that has already been done in many of the most important herbaria of the
world. Nevertheless, much work remains to be done in personal herbaria, where
collections frequently can be found which have been studied rarely or never before, but
which are of great importance for their place of origin or for their rarity, and which must
be included therefore in general studies. This is the case of Prof. H. Kreisel's personal
herbarium which includes a large number of collections from Germany and many other
territories. In this occasion we contribute the results of a revision of the material
belonging to the genus Tulostoma Pers.: Pers.

MATERIAL AND METHODS

The studied material of Prof. H. Kreisel's personal herbarium is currently
deposited in the Institute of Microbiology, University of Greifswald, Germany. The
* Correspondence and reprints: gabriel.moreno@uah.es.

58

G. Moreno, H. Kreisel & A. Alt6s

collections are listed in chronological order (collection date) in each country. The
distribution of the species known from East Germany has been mapped by Kreisel (1984),
but additional localities have been discovered since.
Spore ornamentation of the collections of greater interest has been compared
with that of the con'responding type collections. SEM photomicrographs of some of these
species are presented. The preparation of the samples for SEM observation was made
according to Moreno et al. (1995). All samples studied were mounted in water or in
Congo red to be observed under LM. Spore measurements include the ornamentation.

RESULTS

Tulostoma armillatum Bres. in Petri, Ann. Mycol. 2: 422-423, tab. 6, fig. 17. 1904.
(FIGS. 1-3).

= T. fulvellum Bres. in Petri, Ann. Mycol. 2: 425-426, tab. 6, fig. 23. 1904.
Studied material: GERMANY: Baden-WUrttemberg, Ludwigsburg; leg. G.
Fleischfresser, V. 1966.
Observations: Tulostoma armillatum Bres. is characterized by its hyphal
exoperidium, fimbriate stoma, capillitium with abundant, coloured and inflated septa, and
by its typical spores which are smooth and ovoid to ellipsoid. The synonymy of
Tulostoma armillatum Bres. and T.fulvellum Bres. was proposed by Candousseau (1973).
This matter was pointed out and commented by Moreno & Barrasa (1985) and Wright
(1987; Addendum). This is not a frequent species, but it has been mentioned from
Germany earlier (Gross et al., 1983; Wright, 1987).

Tulostoma brumale Pers.: Pers., Syn. Meth. Fung. p.139. 1801.

Studied material: AUSTRIA: Nieder6sterreich, westlich Neusiedel am See,
Grabenb6schung, L6ssboden mit Artemisia pontica, Ceratocephalus, leg. W. Rothmaler,
6.IV.1959. Burgenland, NSG <<Thenau >>bei Breitenbrunn am Neusiedlersee, Felsensteppe, leg. E. Mrazek, 26.HI.1987.
GERMANY: Brandenburg, Gartz Oder, Strasse Gartz-Heinrichshof; auf Sandboden, leg. H. Kreisel, 28.III. 1957. Ibidem, Mfirkische Schweiz, Buckow, Kiesbruchhiigel
an der Strasse nach Wriezen, leg. R. DUll, 19.III.1958. Ibidem, Prenzlau, 2km N
SchmOlln, Hang am Randowtal, SO-exp. Trockenrasen, sandiger Geschiebemergel, leg.
H. Kreisel, 18.XI.1958. Ibidem, Strausberg, im Naturschutzgebiet Strausberg, mehrfach
in Trockenrasen auf nEhrstoffreicbem Kies, mit Racomitrium canescens, leg. W. Fischer,
22.IV.1961. Ibidem, Oderberg, Pimpinellenberg und Schlossberg, leg. H. Kreisel,
26.X.1957. Ibidem, Siidhang des Pimpinellenberges oberhalb des ebem., Gasthofes
<<Eisguste >>,Festuca-Rasen, Sand, leg. H. Kreisel, 8.V.1962. Ibidem, Stipetum capillatae
mit Festuca aft. glauca, Sand, leg. H. Kreisel, 8.V.1962. Ibidem, innerhalb eines Alyssum
montanum-reichen Festuco-Koelerietum glaucae, leg. J. Endtmann, 1.XII.1984. Ibidem,
Pimpinellenberg, Sand-Trockenrasen, leg. D. Benkert, 20.IX.1990. MecklenburgVorpommern, Darss, Ahrenshoop, Diine am Ort, leg. H. Kreisel, 1.IV.1955. Ibidem,
4.XI.1955. Ibidem, 9.IX.1957. Ibidem, Riigen, siidlich G~hren, Diine mit Pulsatilla
pratensis, Arabidopsis thaliana, Tortula ruralis, leg. H. Kreisel, 7.V.1955. Ibidem,
Sand-Trockenrasen auf Kliffranddiine, leg. H. Kreisel, 26.X. 1986. Ibidem, Schmale Heide
siidlich yon Mukran, leg. H. Jaschhof, 10.X.1993. Ibidem, Insel Riigen, Lietzow, S-exp.
Hange am Kleinen Jasmunder Bodden, pleistociiner Sand, leg. H. Kreisel, 8.V.1955.
Ibidem, Dtinen zwischen G6hren und Lobbe, leg. U. Dettmann, 20.IV.1957. Ibidem,
Diinen von Mukran, leg. I. Schmidt, ILI.1961. Ibidem, Pasewalk, Schanzberge bei
Brietzig, Trockenrasen auf Kies, Os; leg. U. Dettmann, 17.IV.1957. Ibidem, Insel

Tulostoma in H. Kreisel's Herbarium

59

Figs 1-3. Tulostoma armillatum (Germany, Ludwigsburg; V. 1966). Spores. Figs 4-5. T.
melanocyclum (Germany, Kyffh~iuser;24.IX.1967 and Gartz/Oder; 27.111.1957). Spores. Figs 6-8.
T. moravecii (Austria, NSG <<Thenau >>;24.X.1989). Spores,

Usedom, Bahndamm bei Zinnowitz, kiesiger Sand, Siidseite, leg. H. Kreisel, 19.IV.1957.
Ibidem, Ahlbeck, Dtinen/~stlich der Seebriicke, leg. J. Schwik, VIN.1993. Ibidem, bei
Neppermin Trockenrasen, leg. H. Henker, 3.VI.2000. Ibidem, Insel Hiddensee, unweit
nSrdlich Neuendorf, Graudtine mit Tortula ruralis, leg. H. Kreisel, 20.V.1962. Ibidem,
Weststrand zwischen Vitte und Neuendorf auf Htihe der <<Heiderose >>,Weissdiine, Carex
arenaria-Assoz., leg. H. Kreisel, 5.X.1962. Ibidem, Dornbusch, auf dem Leuchtturmhiigel, leg. H. & M. Hiibel, 1997. Sachsen-Anhalt, Halle Saale, Porphyrfelsen 4 km
unterhalb der Stadt am rechten Saaleufer, flachgrtindiger Boden, Porphyrschotter, leg. F.

60

G. Moreno, H. Kreisel & A. Alt6s

Grfiger, 19.X.1954. Ibidem, Halle, felsiges Saaleufer bei Brachwitz, leg. J. Langner,
1/I.1973. Ibidem, Naumburg, beim Vorwerk R6del, Muschelkalk, Nordwest-Hang,
Trockenrasen, leg. F. Griiger, 1.III.1956. Ibidem, Freyburg/Unstmt, Neue G6hle, StidostHang, leg. H. Kreisel, IV.1958. Ibidem, an der Sch6nburg, Schlossgarten, stid-exponiert,
auf Muschelkalk, leg. M. Huth, 1/I.1965. Ibidem, Tongrube sti. Wanzleben, Krs.
Wanzleben, Halde, Keupertone, leg. H..Nowak, 11.IV.1959. Ibidem, Ktirmem, NSG
~ Georgsburg ~, Trockenrasen auf Zechsteinkalk, leg. R. Rauschert, 5.X. 1975. Thiidngen,
Jena, Leutratal, auf Muschelkalk, leg. V. Bergsth'dt, VI.1973. Ibidem, Graitschen, im
Gliesetal am Westhang der Meuskuppe, Teucrio-Seslerietum, leg. R. Conrad, 9.III.1983.
HUNGARY: Szeged, Asotthalom, Reservat ~Kiss Ferenc eml6kerd~,
BinnenCtinen-Trockenrasen, leg. H. Kreisel, 24./X.1972.
ITALY: Toscana, Crete, ca. 5 km sUdlich von Siena, zwischen Moos, Orchis, auf
Lehmboden, leg. I. Wendland, 24.IV.1983. Ibidem, Siena, Monte Oliveto Maggiore, Park,
in Moosrasen unter Cupressus sempervirens, 270 msm, leg. H. & K. Kreisel, 26.XI.1998,
Dupl. in AH 25247. Ibidem, Chianti, Cipresseta Bosco di S. Agnese, in Moospolster in
lichtem Cupressus sempervirens-Forst, Kalkboden, 380 msm, leg. K. Kreisel, 29.XI. 1998,
Dupl. in AH 25248.
SLOVAKIA: Slovakia merid. ~ttirovo, 1 km nordwestlich des Dorfes ~enkov,
auf Flugsand, leg. H. Kreisel & A. ~ertov~i, 27.IX.1961.
Observations: Tulostoma brumale Pers.: Pers. is characterized by its small size,
spore-sac rarely exceeding 10 mm diam., thin membranous exoperidium, tubular stoma,
peristome generally dark brown to blackish, capillitium with abundant yellowish brown,
abruptly inflated septa, and verrucose spores. It is frequent on basic soils among
bryophytes. Occasionally it has been mistaken for other related species such as T.
melanocyclum Bres., T. beccarianum Bres., and even with T. squamosum J.F. Gmel.: Pers.
and T. subsquamosum Long & S. Ahmad, although all of these have sufficient differential
characters. These species have in common the tubular stoma, but their basidiomes
generally have a larger size, with exception of T. melanocyclum which has a similar size
as T. brumale. They may be differentiated from T. brumale also by their type of spore
ornamentation which is formed by well developed coalescent conical spines in T.
melanocyclum and T. squamosum, by a dense pseudoreticulum in T. subsquamosum, and
by relatively well developed warts to spines in T. beccarianum. Moreover, the characteristic capillitium of T. brumale, stirrup-like inflated at the septa, is another trait which
allows to identify this species without difficulty.

Tulostomafimbriatum Ft., Syst. Mycol. 3: 43. 1829.

= T. readerii Lloyd, The Tylostomeae p. 21, pl. 82, figs. 1-2. 1906.
Studied material: AUSTRIA: Burgenland, Siegendorf, leg. E. Mrazek,
8.XI.1984, EM 537, as TuloStoma fimbriatum var. campestre.
GERMANY: Thtiringen, Kyffh~iuser, NSG Ochsenburg, unter Gebiisch im
Astragalo-Stipetum, auf Zechstein-Gips, leg. H. Kreisel, 26.VI.1955. Berlin, BerlinWaidmannslust, SO-Hang der Rollberge, auf trockenem Sandboden, leg. H. Kreisel,
11.IX.1956. Brandenburg, Premnitz a. Havel, auf sandigem Gelande, leg. H. Otte,
25.X.1958. Ibidem, Prenzlau, Sandgrube nw. Eickst/idt, leg. H. Kreisel, 18.XI.1958.
Ibidem, Guben, N~the Seilensee, bei Reicherskreuz, magerer Kiefernwald, leg. R.
Hanschke, 1.I. 1994. Sachsen-Anhalt, Freyburg/Unstrut, in der Sch6nburg, auf einer 2 m
hohen Mauer aus Muschelkalk (etwas vermoost), leg. M. Huth, 1/1.1965. MecklenburgVorpommern, Stralsund, NSG Devin (MTB 1745/1), leg. B. Litterski, 16.III.1993.
Ibidem, Pasewalk, Wilsickower Os, Sand-Trockenrasen, leg. H. Kreisel, 26.IX.1993.
Ibidem, Rerik, Stove, nahe der Windmtihle, Strassenb6schung, Trockenrasen, (ca. 200 bis
300Exemplare), leg. E. Schreiber & H. Henker, 25.111.1998. Ibidem, Insel Usedom,
Deich nahe Forsthaus Damerow, Festuca-Rasen, Sand und Kies, leg. H. Kreisel & Ch.

Tulostoma in H. Kreisel's Herbarium

61

Lindsta'dt, 14.IX.1998. Ibidem, D6mitz, bei Broda zwischen D6mitz und Rtiterberg, auf
dem ehemaligen Kolonnenweg, Trockenrasengebiet, leg. E. Schreiber & H. Henker,
6.IX.1998. NETHERLANDS: Insel Texel, Dtinen, leg. M. Haberer, misit 6.V.1972.
POLAND: Tamsel (Neumark), am Kirschberge, leg. P. Vogel, 8.X.1935.
SLOVAKIA: Slovakia occidentalis, Zahorie, Diinen zwischen den D6rfern
Kostoli~te und GajL,'y, westl, der Stadt Malacky, Flugsand, leg. H. Kreisel & A. ~.ertovfi,
30.IX. 1961.
Observations: Tulostomafimbriatum Fr. is characterized by having a fimbriate
stoma, hyphal exoperidium, squamulose stipe of dark brown colour, and spores with
verrucose and crested ornamentation. There is a broad variability in size and robustness
of the basidiomes, and in spore measurements. The species has an extended list of
synonyms, to which a few years ago the epithet T. readeri Lloyd was added (Alt6s &
Moreno, 1995). Several varieties of this species have been described which, in our
opinion, should be revised more thoroughly. Tulostoma fimbriatum is, together with T.
brumale, the most frequently cited species on the European continent.

Tulostoma giovanellae Bres., Fungi Tridentini p. 63. 1881.

Studied material: GREECE: Ins. Kreta, Paxinadia, zwischen PistaciaStr~iuchem, ca. 140 msm, leg. B. Greemer, 1999.
SYRIA: N33048'11 '' E36047'49 '' NE Jerud, trockener Salzsee, leg. U.
Koltzenburg, 8.IV. 1999.
Observations: Tulostoma giovanellae Bres. is a species typically bound to the
mediterranean region, but has not been formerly quoted from Greece (Zervakis et aL,
1998). It is characterized by its whitish basidiome, submembranaceous exoperidium,
circular and somewhat projected stoma, capillitium with very variable interior lumen
(sometimes with a moniliform aspect), and by its spore ornamentation formed by low
warts and short crests which tend to be united in meridian-like structures particularly at
the hilar appendix region.
A complete description of the type of this species, together with a discussion of
the taxonomic problem arising from the type of T. volvulatum I.G. Borshch., was recently
published by Alt6s et al. (1999)

Tulostoma kotlabae Pouzar in Pihlt, Fl. ~SR, Gast. p. 815. 1958.

Studied material: AUSTRIA: Burgenland, Thenau, Felsensteppe, leg. E.
Mrazek, 8.XI.1986.
GERMANY: Mecklenburg-Vorpommern, RiJgen, stidlich G6hren, Diine mit
Pulsatilla pratensis, Arabidopsis thaliana, Tortula ruralis, leg. H. Kreisel, 7.V.1955.
Ibidem, Insel Hiddensee, auf dem Dornbusch am <<Aschkoben ~, Festuca ovina-Rasen
mit schtitterem Rosa-Gebiisch, steile B6schung, pleistoc. Sand, leg. H. Kreisel, 4.X. 1962.
Ibidem, Pasewalk, Wilsickower Os, Sand-Trockenrasen, leg. H. Kreisel, 26.IX.1993.
HUNGARY: Kecskem6t, Puszta Bugac, Naturschutzgebiet, leg. E. Mrazek,
24.X.1987. Ibidem, Sand-Trockenrasen, leg. W. Winterhoff & J. Kuthan, 20.IX.1978.
SLOVAKIA: Slovakia occid., Ad terram arenosam in gramine sparsa apud viam
ferream haud procul loco <<U Kni~ku >> dicto inter Ktity et Sekule, leg. F. Kotlaba,
17.XI.1955 (dupl. e herbario A. Novack)7). This is a collection from the type locality of

Tulostoma kotlabae.
Observations: Tulostoma kotlabae Pouzar is characterized by its small and
slender habit (similar to T. brumale), hyphal exoperidium, whitish endoperidium, narrow
and distinctly tubular stoma, colourless peristome, and the spore ornamentation formed by
weakly developed verrucae sometimes united in small groups.
It is a species practically restricted to the European continent. It was cited from
Czechoslovakia (Pouzar, 1958), France (Demoulin, 1970; Momand, 1989), Germany

62

G. Moreno, H. Kreisel & A. Alt6s

(Gross et al., 1983), Spain (Martin, 1988; Calonge, 1989, 1990; Vidal, 1991), Austria,
Hungary and Italy (Alt6s et al., 1994). In Asia it is known only by one quotation from
Kazakhstan (Schwarzman & Philimonova, 1970).

Tulostoma raelanocyelum Bres. in Petri, Ann. Mycol. 2: 415. 1904 (FIGS. 4-5)
Studied material: AUSTRIA: NiederGsterreich, Matzen, leg. E. Mrazek,
24.XI.1981, EM 161.
GERMANY: Thiiringen, Kyffh~user, Ochsenburg, unter Quercus im AstragaloStipetum, leg. H. Kreisel, 1955. Ibidem, Berg tiber der BarbarossahGhle (= Ochsenburg),
SiJdhang, leg. M. Huth, 24.IX.1967. Brandenburg, Gartz a. Oder, am Wege zum Schrey,
Stipa capillata-Rasen, sandiger Geschiebemergel, Osthang; leg. H. Kreisel, 27.111.1957.
Ibidem, Buckow (M~kische Schweiz), an der Pritzhagener Mtihle, SteppenhiJgel, leg. R.
DUll, IX.1957.
SLOVAKIA: Slovakia merid., KomLrno, pfskyv u ~enkova, leg. E ~marda,
12.VIII.1949. Slovakia occ., Malacky, prope Plaveck~ Stvrtok, ap. viam ferream, ad
terram arenosam in margine silvae, leg. E Kotlaba, 20.VII.1956.
Observations: Tulostoma melanocyclum Bres. is macroscopicaUy very similar
to T. brumale, with the same habit, a stoma generally surrounded by a dark peristome, and
both are sometimes found to fructify together (Wright, 1987). Nevertheless, they can
easily be differentiated by microscopical characters; T. melanocyclum has a capillitium
with scarcely inflated, colourless septa, and spores much more ornamented than in T.
brumale, i.e. there are groups of large spines coalescent at their tips. This species has been
mistaken with T. subsquamosum (Alt6s et al., 1996), although the latter is differentiated
by its larger habit and by its characteristic labyrinthic to subreticulate spore ornamentation.
This is another typical European species, frequently quoted from this continent
(Wright, 1987). Nevertheless, after our experiences in the study of collections recorded
under this name in Spain (Alt6s et al., 1996), we regard it necessary to check the
remaining citations from southern Europe in order to confirm the presence of T.
melanocyclum in this territory.

Tulostoma moravecii Pouzar in Pihit, Fl. ~SR, Gast. p. 814, fig. 215. 1958.
(FIGS. 6-8).
Studied material: AUSTRIA: Burgenland, NSG <<Thenau ~ bei Breitenbrunn
am Neusiedlersee, Felsensteppe, leg. E. Mrazek, 26.III.1987. Ibidem, Felsensteppe auf
Kalk, leg. E. Mrazek, 24.X. 1989.
Observations: This species may be difficult to distinguish from other related
taxa, since there are only few differential characters. In our opinion, it is closely related
to T. albicans White. Indeed, Pouzar (1958) created this species using material previously
determined as T. albicans by Moravec, mainly based on the tubular, generally conicaltruncate stoma, and on the somewhat verrucose spores. After the study of the types of both
species (Alt6s, 1996) we could demonstrate that most of their macro- and microscopical
characters agreed. The main distinguishing character is the stoma, with conical-truncate
aspect in T. moravecii Pouzar, and circular, plain or slighty elevated in T. albicans. In spite
of the scarceness of adequate differential characters we currently prefer to maintain these
species separate. Tulostoma moravecii is restricted to the European continent, being
quoted from the Czech Republic (Pouzar, 1958), Hungary (Jtilich, 1984), and Austria
(Mrazek et al., 1995), while the name T. albicans applies to the American collections.

Tulostoma in H. Kreisel's Herbarium

63

Tulostoma obesum Cooke & Ellis, Grevillea 6: 82, pl. 100, fig. 24. 1878.
= T. volvulatum sensu Holl6s, Die Gasteromyceten Ungarns. 1904, sensu auct. plurium,
non Z volvulatum I.G. Borshch., Materialy dla botanicheskoy geografii AraloKaspiyskogo kraya. In Zapisky Imperatorskoy Akademii Nauk, Suppl. vol. 7, 189. 1865.
(nomen rejiciendum).
Studied material: YEMEN: Umgebung von Aden, Arjun, auf Sanddiinen, misit
R. Bregazzi, 21.m.1972. Aden, Abyan, Lawder, leg. M. Al-Fatimi, 15.I.2000.
Observations: The rediscovery and study of the type of T. volvulatum and a
revision of the group of related species, prompted to propose the name T. obesum Cooke
& Ellis instead of T. volvulatum sensu auct. (Altts et al., 1999). Although this species has
been quoted from Asia, in particular from Arabia and Jordania (Wright, 1987), we have
no evidence of previous collections from Yemen.
Tulostoma pulcheUum Sace., Bull. Soc. Mycol. France 5: 118. 1889. (FIGS. 9-11)
Studied material: SLOVAKIA: Slovakia occid., Zahorie, Dtinen zwischen den
Dtrfem Kostoligte und Gajftry, westl, der Stadt Malacky, auf Flugsand, leg. H. Kreisel,
30.IX, 1961.
Observations: Tulostoma pulchellum Sacc. is a widely distributed species
characterized by its distinctly membranaceous exoperidium, fimbriate stoma, scutellumlike peristome, and spore ornamentation, under SEM, formed by numerous small,
generally apptanate warts. A variety, T. pulcheUum var. subfuscum (White) Wright et al.,
does not offer macro- and microscopical differences with exception of the spore
ornamentation under SEM, which consists of slightly smaller not applanate warts, and
alternating with some well-developed crests (Moreno et al., 1992).
Tulostomapusillum Berk., London J. Bot. 1: 157, pl. 7, fig. 10. 1842. (FIGS 12-13)
= T. bonianum Pat., Bull. Soc. Mycol. France 8: 49-50, pl. 7, fig. 4. 1892.
Studied material: CUBA: Oriente, Moa, Yamanigtiey; lugares pedregosos en
cuabal de serpentina, 20 msm., leg. H. Kreisel, 7.XI.1969 (Dupl. of HAJB 01259).
VIETNAM: Hanoi, in einem mit Teebliittern gediingten Blumentopf, leg. Trinh
Tam Kiet, IX.1998.
Observations: Tulostoma pusillum Berk. is characterized by its verrucose
exopefidium and a spore ornamentation formed by groups of very notable spines,
coalescent at their tips, resulting in large structures of pyramidal aspect.
Tulostoma squamosum J.F. Glnel.: Pers., Syn. Meth. Fung. p. 139. 1801.
Studied material: AUSTRIA: Niedertsterreich, Schtnfeld bei Marchegg, leg.
Mrazek, 22.XI.1984, EM 561. Ibidem. 18.IX.1985, EM 657. Ibidem. Marchegg, Schtnfelder Dtine, leg. Mrazek, 12.IX.1989, Dupl. EM-1300 in herb. H. Kreisel. Ibidem.
aufgeforstet mit Populus, Pinus, Robinia, Sambucus, Crataegus, 100msm., leg. E.
Mrazek.
CZECH REPUBLIC: Moravia centr., Tignov, in colle <~Kv~mice >>, Pinus
sylvestris, leg. E ~marda, 27.XI.1946.
GERMANY: Mecklenburg-Vorpommern, Waxen a. Miiritz, stidl. Ortsrand, unter
angepflanzten Str~iuchern, Sandboden, leg. H. Dtrfelt, 111.1976.
HUNGARY: Kecskemtt, Naturschutzgebiet Puszta Bugac, leg. E. Mrazek,
24.X. 1987.
Observations: Tulostoma squamosum is characterized by its clearly tubular
stoma, slightly verrucose exoperidium, stem covered with long dark scales, and spore
ornamentation formed by coalescent spines of pyramidal aspect (but less developed than
in T. pusillum).
This is a frequent species in basic soil, and frequently quoted in Europe.

64

G. Moreno, H. Kreisel & A. Alt6s

Figs 9-11. Tulostomapulchellum (Slovakia, Malacky; 30.IX.1961). Spores. Figs 12-13. T. pusillum
(Vietnam, Hanoi; IX.1998). Spores. Figs 14--16. T. striatum (Cuba, Sierra Maestra; Dupl. of HAJB
01019). Spores. Figs 17-19. T. subsquamosum (Nepal, Phoksumdo Tal; 30.IV.1974). Spores.

Tulostoma in H. Kreisel's Herbarium

65

Talostoma striatum G. Cunn., Proc. Linn. Soc. New South Wales 50: 255. 1925.
(FIGS 14-16)
Studied material: CUBA: Oriente, Sierra Maestra, costa sur entre Chivirico y
E1 Uvero, bosque seco sobre arena, 20 msm, leg. H. Kreisel, 15.IV.1969 (Dupl. of HAJB
01019).
Observations: This species is characterized by its distinctly membranaceous
exoperidium, sub lente pubescent endoperidium, fibriUose-fimbriate stoma, and spore
ornamentation typically formed by somewhat ramified long crests which cause a sWiate
aspect of the spores. Tulostoma striatum G. Cunn. is widely distributed in America
(Wright, 1987), but we do not know any earlier citation from Cuba. The species is also
known from Africa (South Africa), Asia (Israel, Japan), and Australasia (Australia, New
Zealand). Some years ago it was first indicated from Europe (Spain), collected in an urban
site together with T. fimbriatum (Alt6s & Moreno, 1991).
Tulostoma subsquamosum L o n g & S. Ahrnad, Farlowia 3: 241, 1947. (FIGS 17-19).
Studied material: NEPAL: N.W. Nepal, Phoksumdo Tal, Lande ~t Caragana
gerardiana, 3600 msm, leg. J.E Dobremez, 30.IV.1974.
Observations: Tulostoma subsquamosum is characterized by a tubular stoma,
generally concolorous with the endoperidium, stem covered with brown scales, and spore
ornamentation formed of a dense subreticulum of labyrinthic aspect. As we already have
commented, this species repeatedly was mistaken with other species as T. squamosum and
T. melanocyclum. Apart from the type material from India, very few collections of this
species are known: Argentina (Wright, 1987), Spain (Alt6s et al., 1996), and Mexico
(Ochoa et al., 1998). The collection studied from Nepal was mentioned by Kreisel (1976)
as T. melanocyclum.

Acknowledgements. This study has been possible through DGICYT PB 95-0129, and
also by the ~ Programa de Cooperaci6n con Iberoam6rica, Ministerio de Educaci6n y Ciencia ~,
Spain. Our gratitude is additionally expressed to Mr. J.A. P6rez and A. Priego, of the SEM Service
of the Universidad de Alcal,'i, for their kind assistance.

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