A World-Monograph of the genus Pseudombrophila

118 Pages·1995·13.218 MB·English

by  J van Brummelen

LIBRI BOTANICI
Vo!. 14
A World-monograph of the Genus
PSEUDOMBROPHILA
(Pezizales, Ascomycotina)
by
J. van Brummelen
With 33 text-figures
8 coloured plates and
17 monochrome plates
IHW-VERLAG
1995
L1BRI BOTANICI
Vcl. 14
A World-monograph of the Genus Pseudombrophila
(Pezizales, Ascomycotina)
LIBRI BOTANICI
Vol. 14
A World-monograph of the Genus
PSEUDOMBROPHILA
(Pezizales, Ascomycotina)
by
J. van Brummelen
With 33 text-figures
8 coloured plates and
17 monochrome plates
IHW-VERLAG
1995
Die Deutsche Bibliothek - CIP-Einheitsaufnahme
Brummelen, Joop van:
AWorld-monograph ofthe Genus Pseudombrophila (Pezizales,
Ascomycotina) / by J. van Brummelen. - Eching bei Miinchen :
IHW-Verl., 1995
(LibriBotanici ;Vo!. 14)
ISBN 3-930167-10-7
NE:GT
Impressum:
ISBN 3-930167-10-7
Author: Dr. J. van Brummelen
Rijksherbarium - Hortus Botanicus
P.O. Box 9514
NL - 2300 RA Leiden
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Publication: IHW-Verlag
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© 1995
All rights reserved. No part of this book may be reproduced in any form by
photostat, microfilm, or any other means, without written permission from the
publisher.
CONTENTS
SUMMARY 5
INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
ACKNOWLEDGEMENTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
GENERAL PART 6
History 6
Materials and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Structure ofthe fungus 10
TAXONOMIC PART 18
The genus Pseudombrophila . . . . . . . . . . . . . . . . . . . . . . . . 18
Key to the sections ofPseudombrophila 19
Section Pseudombrophila 20
Key to the species of section Pseudombrophila 20
Section Nannfeldtiella 65
Key to the species ofNannfeldtiella 65
APPENDIX. Insufficiently known and excluded species 83
REFERENCES 89
INDEX 93
COLOURED PLATES 97
MONOCHROME PLATES 101
3
SUMMARY
The genus Pseudombrophila (Pyronemataceae, Pezizales) is described and discussed. The
genus is divided into two sections: Pseudombrophila andNannfeldtiella. Nineteen species
aredistinguished insectionPseudombrophila and nine insectionNannfeldtiella, onaworld
basis.Fourspecieshavebeen newlydescribed andseventeennewcombinationswereneces
sary underPseudombrophila. The namesofexcludedand insufficiently known taxaaredis
cussed in an appendix. Svrcekomyces pallidus is redescribed and referred to the genus
Leucoscypha.
INTRODUCTION
The genus Pseudombrophila Boud. (Pyronemataceae, Pezizales) is the subject ofa world
wide revision.
Many species previously described in other genera were recognized as belonging to
Pseudombrophila, while a number of new species have been described through the years.
Others were described several times because of the absence of a modern treatment and
insufficient knowledge ofspecific variability.
Several specieswere newly described in the genusFimaria Velen. since attention was paid
to this forgotten name (VAN BRUMMELEN, 1962). In the present study Fimaria is included
inPseudombrophila.
Since mostpublisheddescriptionsproved to be insufficientto recognize thespecies,astudy
ofauthenticmaterialwasnecessarytomakefull descriptionsandillustrationsofthespecies.
Asfaraspossiblethisstudy is basedonliving material. But, as many speciesare extremely
rare or restricted to unusual habitats, the major part of the material studied consisted of
dried specimens.
Studies oftype-specimens have been undertaken and included whenever possible. In cases
where type specimens were obviously absent, published and unpublished illustrations and
descriptions have sometimes helped in recognizing species.
Several specific names are only known from short, insufficient descriptions. These names
are includedinthelistofinsufficientlyknownspecies. However, afew oftheoldestofsuch
names have consistently been interpreted in the same way as a result of the existence of
exsiccatae distributed under this name by early mycologists, like K. W. G. L. Fuckel, G. L.
Rabenhorst, P. A. Karsten, and M. C. Cooke. In such cases illustrative specimens or
epitypes (GREUTER, 1994: Art. 9.7) have beenindicatedto stabilizethe use ofthese names.
The purpose ofthis investigation is to evaluate the morphological variation amongthespe
cies ofPseudombrophila and produce a taxonomic treatment. To accomplish these goals a
number ofdifferent techniques have been used, including the study ofmacro-morphology,
pigmentation, differentiation ofhairs, ascoma ontogeny, ascus and ascospore ultrastructure
(VAN BRUMMELEN, 1986a, 1994), and development and structure ofascospore ornamenta
tion.
ACKNOWLEDGEMENTS
The author wishes to thank the directors and curators of the institutes mentioned on p. 7-8
for the loan of material or for permission to examine specimens in their care. Valuable
material has also been gratefully received from Dr. O. Aas, Mr. J. Astier, Mr. A. Ayel, Mr.
B. Hannf, Mr. J. Breitenbach, Prof. F. D. Calonge, Dr. H. Dissing, the late Dr. J.-c.
Donadini, Mr. H. Engel, Mr. G. Fortoul, the late Mr. W. D. Graddon, Prof. G. S. de Hoog,
Mr. R. Kristiansen, Mr. W. D. 1. Kuijs, Prof. N. Lundqvist, Mr. W. Matheis, Dr. E.
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Ohenoja, Mr. J. T. Palmer, Mr. H. & Mrs G. Piepenbroek-Groters, Dr. A. Raitviir, Mr. L.
Riousset, Dr. S. Sivertsen, Mr. G. J. M. Verkley, and Prof. J. Webster.
Warmest thanks are also extended to Ms Dr. S. M. Francis for reading the manuscript and
improving the English text; to Dr. R. A. Maas Geesteranus for correction of the Latin
diagnoses of the new taxa; and to Prof. T. Schumacher for permission to reproduce his
colour-slide ofPseudombrophila guldeniae.
The 'Netherlands Organization for the Advancement of Research (N.W.O.)' is gratefully
acknowledged for a grantwhich financed avisit to the herbaria at Uppsala and Stockholm.
The author is also most grateful to the 'French National Scientific Research Center
(C.N.R.S.)' for a grant to visit French herbaria.
GENERAL PART
HISTORY
The genus Pseudombrophila was first proposed by BOUDIER (1885) for three species:
Helotium pedrottii Bres., Peziza chartarum Quel., and Peziza pluvialis Cooke. He placed
it in his new family 'Ciliaries' with the following diagnosis: 'Poils courts et obtus; spores
sans sporidioles; couleur pale; especes subfimicoles, stipitees ou obconique'.
ROLlAND (1888) included Pseudombrophila theioleuca from France.
FromBOUDIER's(1907)presentationofhisnewclassificationofthediscomycetesitbecame
clearthat he had a rather narrow conception ofthe genusPseudombrophila. In his descrip
tion he stresses the presence ofreceptacles with an outer decoration offascicles orisolated
bunches of coloured, septate and blunt hairs of the type found in Melastiza Boud. and
Anthracobia Boud.InthisenumerationonlysixspeciesofPseudombrophilawere recogniz
ed, while most species now belonging to Pseudombrophila, with less conspicuous bunches
of submarginal hairs, were included in his large heterogenous genusAscophanus Boud.
BeforetheestablishmentofPseudombrophila itsspeciesweremostlyarrangedunderPeziza
Dill. ex L.: Fr. in the works ofBatsch, Fries, Sauter, Berkeley, Nylander, Karsten, Cooke,
Schroter, E. C. Hansen, and Phillips. Later, after the publication of new classifications by
FUCKEL(1870)andSACCARDO (1884, 1889, 1895), these specieswerealso classified under
Humaria Fuckel (Fuckel,Saccardo, Quelet, Rehm, andVelenovsky),Pyronema Carus(Sac
cardo, Spegazzini, Rehm, and Bisby), Lachnea (Fr.) Gill. (Saccardo and Velenovsky), and
Neottiella (Cooke) Sacc. (Saccardo and Boudier); while species with spherical ascospores
were listed under Cubonia Sacc. (Saccardo, Boudier, Grelet, Faurel & Schotter, and
Durand).
Newspecies ofPseudombrophilawere described by LEGAL(1937), SVRCEK(1966, 1988),
and HARMAJA (1979, 1986).
VELENOVSKY(1934)describedthenewgenusFimaria Velen. withfour speciesand avarie
ty. He segregated it from the large genus Humaria (Fr.) Boud. sensu Rehm. He described
the genus Fimaria as: 'Genus, omnino generi Humaria affine, sed apoth. margine late
membra-naceo-limbata, pierumque in cilias acutas fimbriata, paraph. simpl.-filif. vel
ramosae. - In fimis, raro in limis.'
Fimaria was emended by VAN BRUMMELEN (1962) by lectotypification. The genus was
redescribed and four species were distinguished. Later, several other species of Fimaria
wereaddedbySVRCEK& KUBICKA(1965), ECKBLAD(1968),SVRCEK& MORAVEC(1969),
JENG& KRUG(1977),TORRE& CALONGE(1978), GRADDON(1980), and CALONGE(1984).
In the presenttreatmentFimaria is recognized as belongingto the genusPseudombrophila.
ECKBLAD (1968) introduced the genus Nannfeldtiella Eckbl. for a species, N. aggregata
Eckbl., associatedwiththe subiculum ofByssonectriaatplaceswhere elkisliving. Because
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the ascospore ornamentation was of a non callose-pectic nature Nannfeldtiella was incor
porated in the family Sarcoscyphaceae Le Gal ex Eckbl. Further study of this fungus by
KORF(1972) revealed that its ascospore markings are ofacallose-pecticnature(,cyanophi
lie') and not typical for the Sarcoscyphaceae. HARMAJA (1979) transferred it to Pseudom
brophila and described some other closely related species from the same habitat.
MORAVEC (1976) introduced the new generic name Svrcekomyces for Pseudombrophila
guldeniae Svrcek, which he considered 'rather similar' or possibly 'even identical' with
Nannfeldtiella aggregata.
PFISTER (1984)compiled a number ofcoprophilous specieswithspherical ascospores from
the synonymy ofAscophanus dentatus Boud. and provided a key to species ofFimaria.
VAN BRUMMELEN (1986a) studied the structure and development of the ascus top and the
ascospore wall in two species ofPseudombrophila and two species ofFimaria with light
and electron microscopy. Ascus tops were found with roughly delimited ascostomes and
opercula. The general structure and the dehiscence mechanismofthe ascusare very similar
and thesameasthose found ingeneraofthePyronemataceae.Theascosporeornamentation,
although sometimesvery different at maturity underthe lightmicroscope, develops initially
in the same way from a smooth electron dense, blue staining ('cyanophilous') secondary
wall. In the smooth-spored species the initial layer is permanent. In the rough-spored ones
it usually breaks up to form an irregular granular pattern.
In an additional study ofNannfeldtiella aggregata, which shows mature ascospores with a
rather conspicuous, incomplete net-work of thick and fine lines, VAN BRUMMELEN (1994)
found asimilarinitialdevelopment ofthesecondary ascosporewall. Butonfurther ripening
the investing membrane islifted, creating large spacesonly filled with afew widely spaced
fibrillar elements. At maturity the secondary wal1 becomes densely filled with fibrillar ele
ments constituting the final pattern of ornamentation.
MATERIALS AND METHODS
Whenever possible observations were made on living material, but the major part of the
material consisted ofdried specimens.
LIVING MATERIAL. -- Since most species of Pseudombrophila are coprophilous, a great
number of samples of dung ofdifferent origin was col1ected and incubated to isolate fruit
bodiesoffungi growingon them. The methodsfor isolationand cultivationwereessential1y
the same as used before (VAN BRUMMELEN, 1967).
In general fruit bodies of Pseudombrophila appear rather late in the sequence of species
fructifying on dung. The optimal temperature for developing fruit bodies of Pseudom
brophila is between 10 and 15°C. This may explain the fact that they are less frequent in
cultures at room temperature.
Althoughfruit bodies ofPseudombrophila can befound in nature throughout the year, they
are more frequent in spring and in mild winters.
HERBARIUM MATERlAL. ~ Most of the material studied in this work consisted of dried
specimens. Herbaria from which specimens, including type material, were examined, are
indicated in the text by the fol1owing abbreviations. Institutes visited personal1y have been
marked with an asterisk (*).
B Botanischer Garten und Museum Berlin-Dahlem, Berlin, Germany *.
BG Universitets Botaniske Museum, Bergen, Norway.
BPI The National Fungus Col1ection, U.S. Department of Agriculture, Beltsville,
Maryland, U.S.A. *.
BR Jardin Botanique National de Belgique, Meisse, Belgium *.
C Botanical Museum and Herbarium, Copenhagen, Denmark *.
7
CONC Laboratoire de Biologie Marine du College de France, Concarneau, France *.
DAOM Mycological Herbarium, Plant Research Institute, Department of Agriculture,
Ottawa, Canada.
E Royal Botanic Garden, Herbarium, Edinburgh, Great Britain *.
FRS Falconers' Museum, Forres, Great Britain *.
G Conservatoire et Jardin Botanique, Geneve, Switzerland *.
H Botanical Museum, University of Helsinki, Helsinki, Finland.
HBG Staatsinstitut fUr Allgemeine Botanik, Hamburg, Germany.
K Royal Botanic Gardens, The Herbarium, Kew, Great Britain *.
KUO Department of Natural History, Kuopio Museum, Kuopio, Finland.
L Rijksherbarium -Hortus Botanicus, Rijksuniversiteit, Leiden, The Netherlands*.
Inclusive of the mycological collections formerly placed in the herbaria of
Groningen (GRO) and Utrecht (U).
LPS Instituto de Botanica C. Spegazzini, La Plata, Argentina.
M Botanische Staatssammlung, Miinchen, Germany.
MA Instituto 'Antonio Jose Cavanilles', Madrid, Spain.
MPU Insitut de Botanique, Universite de Montpellier, Montpellier, France *.
NY Herbarium of the New York Botanical Garden, Bronx, New York, U.S.A. *
NYS Herbarium of the New York State Museum, Albany, New York, U.S.A.
o
Botanisk Museum, Oslo, Norway *.
OULU Department of Botany, University of Oulu, Oulu, Finland.
PAD Instituto Orto Botanico dell'Universita, Padova, Italy *.
PC Museum National d'Histoire Naturelle, Laboratoire de Cryptogamie, Paris,
France *.
PRM Mycological Department of the National Museum, Praha, Czechia.
S Naturhistoriska Riksmuseet, Section for Botany, Stockholm, Sweden *
TRH BotanicalDepartment, Museumofthe Royal Norwegian Society forScienceand
Letters, Trondheim, Norway.
TRTC CryptogamicHerbarium, DepartmentofBotany, University ofToronto, Canada.
TU Herbarium of the Department of Taxonomy and Geobotany of the Tartu State
University, Tartu, Estonia.
TUR Botanical Institute of the University, Turku, Finland.
UPS Institute of Systematic Botany, University of Uppsala, Uppsala, Sweden *.
W Naturhistorisches Museum, Wien, Austria *.
WU Botanisches Institut und Botanischer Garten der Universitat Wien, Austria *.
For a more accurate indication of herbarium specimens, especially where labelling is not
wholly adequate, the customary abbreviation of the herbarium name is followed by the
author's revision number.
STUDY OF HERBARIUM MATERIAL. - Reasonable swelling of this dried material was
usually obtained by leaving a fragment for two or more hours in a small tube ofwater. But
rehydration and swelling were better when a few drops of a wetting agent, like Photo-Flo
(Kodak) or Invadine (Geigy) were added to 100 ml of water. Repeated warming (to about
70 QC) hastens the rehydration. In difficult cases, with thin-walled tissues or old material,
the addition ofa small amount ofammonia (up to 1 %) gave the best results. Moreover the
alkalinity, produced by the addition of ammonia, made the material far less brittle during
later handling.
Too fast or insufficient rehydration, which may easily occur in mounting media with a low
watercontent, like lactic acid, lacto-phenol, or Melzer's reagent, leads to poor preparations
with allkinds ofartefacts, such as deformed asci, collapsedparaphyses, and sporeswith air
bubbles.
MICROSCOPIC EXAMINATION. --For microscopic examination small fragments ofliving
fruit bodies were placed in a drop of tap water on a slide and the elements then spread
8
undergentle pressure. Vital observationscouldonlybe madeduring afew minutes,because
ofthe quick disintegration ofcell components.
As vital stains, brilliant cresyl blue (Colour Index 51010; Rowe, 1956-58; Lillie, 1969),
neutral red (Cl. No. 50040), and trypan blue (Cl. No. 23850) were used in very weak
solutions (under 0.01 %) in tap water. Suitable objects were also studied with interference
contrast and phase-contrast optics.
Whenever possible spore measurements were made in water on ascospores that had just
been forcibly discharged from mature living asci.
To obtain a higher accuracy and better reproducibility of visual size measurements of
ascospores the sheared-imagemeasurement, as described by DYSON(1960), was constantly
used. This method avoids the considerable measuring deviations due to the use of oil
immersion objectives with high apertures, the so-called aperture error (LOCQUlN, 1956,
1984; CHARMAN, 1963) or the hypercentric perspective (MICHEL, 1964). The latter shows
the equatorial plane of spores smaller than more distant planes.
Fullyrehydratedherbariummaterialwasusedasfresh materialforfurthermicroscopicstudy
and the preparation of microtome sections. Microtome sections, IQ-15 ,um thick, were
made with a freezing microtome from material embedded in polyvinyl lacto-phenol (Gurr)
as previously described (VAN BRUMMELEN, 1967).
Ascospore ornamentations and hyphal walls in sections and squash mounts were stained in
a 0.1---0.5 % solution of methyl blue (Cl. No. 42780) or trypan blue (Cl. No. 23850) in
lactic acid or lacto-phenoI.
Methylbluewasused asadye for stainingthe ornamentationofascospores indiscomycetes
by the lateDr. M. Le Gal afterthe production of[the synonymous] Poirrierblue C4B ('bleu
C4Bde Poirrier'; GUEGUEN, 1905)wasdiscontinued(VAN BRUMMELEN, 1967).Itisstillthe
best microscopic stain available for ascospore ornamentation. The observation with visual
microscopy offine details ofspore ornamentations stained with methyl blue or trypan blue
can be improved by the use ofa complementary monochromatic light filter.
In the literature there exists some confusion about the dyes for staining spores. For a long
time the terms cotton blue and aniline blue have both been differently applied to mixtures
of two dyes, now called methyl blue (Cl. No. 42780) and water blue I (Cl. No. 42755)
(LILLIE, 1969; MATHEIS, 1975). Although it appears that both components are interchan
geable with each other in most stains, it would be better to relegate the terms cotton blue
and aniline blue to the history of staining, and use the proper name of the single dye
substituted for the mixture in accordance with the standardized names ofthe Colour Index
(ROWE, 1956--58; LILLIE, 1969).Thus confusioncan be avoidedofcurrentwith olderpro
cedures, and the nomenclature of dyes will correspond to actual practice.
Inthestainingofsectionstrypanbluegavesharperresults than methyl blue, sinceitshowed
a greater affinity with certain parts of the hyphal walls and less with the cytoplasmatic
contents (cf. BOEDIJN, 1956; DADE, 1960; DADE in GURR, 1965).
Asan iodine-containingreagentonthepossiblebluestainingofascuswallsLugol'ssolution
(0.3g 12, 0.6g KI, 100ml H20)was used. This has the advantage, because ofthe relatively
Iow iodine content as compared with Melzer's reagent, of a clear test without too much
background staining of cellular contents by the excess of iodine (cf. BARAL, 1987). Since
iodine by itself is also very reactive, its addition immediately terminates all vital obser
vations. The most successful method ofapplying the iodine solution is by placing a small
drop at one edge of the cover-glass of a preparation in water, while the water is sucked
away at the opposite edge with a piece of blotting paper. During continuous microscopic
observation all eventual changes caused by the reagent can be followed.
9