The Yeasts. A Taxonomic Study

1087 Pages·1984·14.371 MB·English

by  N.J.W. Kreger-van Rij (Eds.)

The yeasts
a taxonomic study
third revised and enlarged edition
edited by
NJ.W. Kreger-van Rij
Groningen, The Netherlands
1984
Elsevier Science Publishers B.V. - Amsterdam
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Library of Congress Cataloging in Publication Data
Main entry under title:
The Yeasts.
Bibliography: p.
Includes index.
1. Yeast fungi—Classification. 2. Fungi—Classifica-
tion. I. Kreger-Van Ri j, N. J. W.
QK6l7.5.Y^8 1981+ 589.2*33 8MH50
ISBN O-UI+I+-80U2I-8 (U.S.)
PRINTED IN THE NETHERLANDS
v
Contributors
D.G. AHEARN, Department of Biology, Georgia State University, Atlanta, Georgia
30303, U.S.A.
RJ. BANDONI, Department of Botany, University of British Columbia, Van-
couver, B.C., Canada V6T 1W5
J.W. FELL, School of Marine and Atmospheric Science, University of Miami,
Miami, Florida 33149, U.S.A.
E. JOHANNSEN, Microbiology Research Group, Council of Scientific and In-
dustrial Research, Pretoria, South Africa
N.J.W. KREGER-VAN RIJ, Laboratory for Medical Microbiology, State Universi-
ty, 9713 EZ Groningen, The Netherlands
C.P. KURTZMAN, Northern Regional Research Center, United States Department
of Agriculture, Peoria, Illinois 61604, U.S.A.
K.J. KWON-CHUNG, Clinical Mycology Section, National Institute of Allergy and
Infectious Diseases, National Institutes of Health, Bethesda, Maryland 20014,
U.S.A.
Sally A. MEYER, Department of Biology, Georgia State University, Atlanta,
Georgia 30303, U.S.A.
M.W. MILLER, Department of Food Science and Technology, University of Califor-
nia, Davis, California 95616, U.S.A.
HJ. PHAFF, Department of Food Science and Technology, University of California,
Davis, California 95616, U.S.A.
L. RODRIGUES DE MIRANDA, Yeast Division, Centraalbureau voor Schim-
melcultures, 2628 BC Delft, The Netherlands
R.L. SCHLITZER, Department of Biology, Georgia State University, Atlanta,
Georgia 30303, U.S.A.
M.TH. SMITH, Yeast Division, Centraalbureau voor Schimmelcultures, 2628 BC
Delft, The Netherlands
A. STATZELL TALLMAN, School of Marine and Atmospheric Science, University
of Miami, Miami, Florida 33149, U.S.A.
J.P. VAN DER WALT, Microbiology Research Group, Council of Scientific and In-
dustrial Research, Pretoria, South Africa
D. YARROW, Yeast Division, Centraalbureau voor Schimmelcultures, 2628 BC
Delft, The Netherlands
VI
Preface
The present, third edition of 'The Yeasts - a taxonomic study' follows the first
(J. Lodder & N.J.W. Kreger-van Rij, 1952) and the second edition (J. Lodder ed.,
1970) which were preceded by monographs on the ascosporogenous yeasts (Stelling-
Dekker, 1931), and the anascosporogenous yeasts, part one (J. Lodder, 1934) and part
two (H.A. Diddens & J. Lodder, 1942).
Since the 1970 edition there have been many new developments in yeast taxonomy:
new species have been described from new isolates, mating experiments have yielded
perfect states of ascomycetous and basidiomycetous yeasts, and new characters as well
as changed evaluation of old characters have led to reclassifications. The 1970 edition
included 39 genera and 349 species while the present edition includes 60 genera and
500 species.
Knowledge of the basidiosporogenous yeasts has increased considerably in recent
years. These yeasts are now classified in two taxonomically different groups, the
teliospore-forming yeasts and the Filobasidiaceae. There are also other basidio-
mycetous fungi, such as the Tremellales, with a yeast phase in their life cycle. The
descriptions of the yeast states of several of these species have been included in this
edition.
Of the new characters, DNA base composition and DNA base sequence homology
are of great importance for yeast taxonomy, specifically for the delimitation of species.
Promising new characters for generic differentiation are, for instance, the ultrastruc-
ture of the cell wall and the chemical composition of coenzyme Q.
The aims of the third edition are identical to those of former editions, first, in
presenting and discussing a classification of yeasts including diagnoses of genera and
descriptions of species, and second, in providing methods for the identification of yeast
strains.
The present taxonomic system has developed gradually and, as pointed out by the
authors, is not yet entirely satisfactory. Unfortunately, reclassifications are often con-
fusing for the users. Therefore, some authors believe in reticence and wait for a more
definite scheme of classification before changing names of species.
The rapid development in yeast taxonomy requires continuous critical reviews. It
is, therefore, very difficult to present an up-to-date treatise. The taxonomic system
proposed in this edition may be considered as a stage in the evolution towards a
satisfactory classification.
Identification of yeasts has become more complicated with the increase in the
number of species and the increase in characters used for the standard descriptions.
Traditionally, in this treatise there is a key to the genera and keys to the species within
each genus. To facilitate the identification of yeast strains, extra keys are supplied
directly to the species.
The description of genera and species in this edition has been slightly abridged from
the 1970 edition. Historical surveys of the genera and original descriptions of the
species have been omitted, but references to them have been provided. The data on
the origin of the strains studied are condensed, but references to ecological details, if
available, are given. Condensation of the text was necessary to obtain a monograph
of a reasonable size.
VII
The present monograph is in contents and shape a direct descendant of the former
editions. This made it easier to undertake the editorship which Dr. J. Lodder
delegated to me. I am extremely grateful to her for help and advice which she gave
from a long experience in yeast taxonomy both as an author and editor.
The pleasant co-operation of the authors is thankfully acknowledged.
N.J.W. Kreger-van Rij
VIII
Acknowledgements
Research and manuscript preparations by J.W. Fell and A. Statzell Tallman were
funded by the National Science Foundation, Program in Systematic Biology (Grant
No. 8040752).
N.J.W. Kreger-van Rij is much indebted to Dr. J.W. Fell, Dr. HJ. Phaff and Dr.
C.P. Kurtzman for critically reading her contributions to the monograph. Dr. Fell
suggested to include yeast forms of Tremellales in this monograph. The advice on
nomenclature problems given by Dr. J. van Brummelen and by Dr. J.A. Leussink is
gratefully acknowledged. Many thanks are due to Dr. R.T. Moore for corrections of
the Glossary. Part of the work for this monograph was supported by The Netherlands
Organization for the Advancement of Pure Research (Z.W.O.).
C.P. Kurtzman gratefully acknowledges the technical assistance of Mrs. M.J.
Smiley. The new figures for the genera Hansenula, Issatchenkia, Lipomyces, Pachysolen and
Pichia were prepared by Mr. C.E. Needham.
Sally A. Meyer gratefully acknowledges the assistance of Stephanie Condron
Prévost and Lindsay R. Layton.
H J. Phaff gratefully acknowledges the technical assistance of Mary Miranda and
Joanne Tredick. Part of this work was supported by Public Health Service Grant
GM-16370 from the Institute of General Medical Sciences.
M. Th. Smith, L. Rodrigues de Miranda and D. Yarrow thank Mr. G. A. Poot and
Mrs. I. Paanakker for their scrupulous and patient technical and clerical assistance,
and also Mr. K.J.L. Bosman and his assistants of the library of the Laboratorium voor
Microbiologie, Technische Hogeschool, Delft, for their help.
The sending of many yeast cultures from the Collection of the Yeast Division of the
Centraalbureau voor Schimmelcultures, Delft, to the authors is gratefully acknowl-
edged.
The Glossary has been prepared by N.J.W. Kreger-van Rij with the assistance of
J.W. Fell and C.P. Kurtzman.
The figures in this monograph were partly taken from "The Yeasts", edition I and
II. The figures of edition I and the figures belonging to the genera Endomycopsis,
Debaryomyces and Pichia which were new in edition II were made by Mrs. P.J.
Klinkhamer-Hellendoorn. The figures belonging to the genera Nematospora, Candida,
Oosporidium, Torulopsis and Trichosporon of edition II were prepared by Mrs. L. Assis-
Lopes and photographed by Mrs. C. Barroso-Lopes. The figures belonging to the
genera Dekkera, Kluyveromyces, Lodderomyces, Wingea and Brettanomyces of edition II were
made by Dr. J.P. van der Walt. The other figures of edition II, as far as their origin
is not mentioned, were prepared by Mr. J.A. Schuur either from drawings made by
Miss W.Ch. Slooff or from photographs. Figures 1-5 and 7-14 of edition III were
DreDared bv Dr. N.Î.W. Kreeer-van Rij and redrawn bv Mr. C.E. Needham.
IX
Directions for the use of the book
This monograph provides the criteria and methods for the classification and iden-
tification of yeasts.
In Chapter I, a system of classification in families, subfamilies and genera is
presented. Each genus is briefly characterized. The criteria for the distinction of
genera and species are discussed and special attention is given to newer characteristics.
Chapter II gives (1) directions for isolation and maintenance of yeasts, (2) descrip-
tions in detail of the tests of the standard description with an explanation of mor-
phological terms, and (3) suggestions for the procedure of identification. A list of the
tests of the standard description with a page reference to the methods is given on p.
000, and an alphabetical list of media with a page reference for their preparation can
be found on p. 000.
Chapters III, IV and V contain a systematic discussion of the genera of the
ascosporogenous, basidiosporogenous (in three groups) and imperfect yeasts, respec-
tively. The genera and species of each group are presented in alphabetical sequence.
The description of a genus comprises: (a) diagnosis, (b) type species, (c) list of ac-
cepted species with the date of first publication, (d) a key to the species of the genus
with a page reference to the standard description, a table of species with key characters
if more than five species are involved, (e) systematic discussion of the species, and,
if required, (f) comments on the genus in which the authors give their views of the
composition and taxonomic position of the genus, especially the latest developments.
Under g. species are listed which have not been accepted by the authors, either
because they do not fit in the genus, or because no strains were available at the time
of the conclusion of the manuscript.
The description of each species includes: synonyms, standard description, com-
ments if necessary, origin of the strains studied and the type strain. Where possible,
a connection between perfect and imperfect species is indicated at the beginning of the
description of each species.
The standard description consists of the basic features, similar for all species and
additional features of specific importance for individual genera or species, both
established with standardized tests.
Explanation of symbols used in the standard description and in the tables:
+ reaction positive
+ w reaction weakly positive
-i- s reaction slow (latent)
- reaction negative
+ ( - ) reaction positive, seldom negative
- ( + ) reaction negative, seldom positive
v some strains give a positive reaction, others a negative one
The magnification of the drawings of vegetative cells and asci or ascospores is
1000 x , unless otherwise indicated. The magnification of the drawings of pseudo-
mycelium and true mycelium is 500 x , unless otherwise indicated. The magnification
of the new Candida drawings is arbitrary.
Orthographic errors against the International Rules of Botanical Nomenclature in
X
the names of the species have been corrected by the authors.
In Chapter VI keys are given, one to the genera, others to species or groups of
species. A list of the keys is at the beginning of this chapter on p. 967.
A glossary to the terms used (p. 1007), a list of references (p. 1011) and an index
of the names of the taxa (p. 1052) complete this book.
XI
Abbreviations
The following abbreviations are used for names of genera:
Aciculoconidium Ac. Pachysolen Pa.
Ambrosiozyma A. Pachytichospora P'spora
Arthroascus Ar. Phaffia Ph.
Brettanomyces Br. Pichia P.
Bullera B. Pityrosporum Pit.
Candida C. Rhodosporidium Rhodosp.
Citeromyces Cit. Rhodotorula Rh.
Clavispora CL Saccharomyces Sacch.
Cryptococcus Cr. Saccharomycodes S'codes
Cyniclomyces Cyn. Saccharomycopsis S.
Debaryomyces Deb. Sarcinosporon Sar.
Dekkera D. Schizoblasto sporion Schizobl.
Fibulobasidium Fib. Schizo saccharomyces Schiz.
Filobasidiella Fil. Schwanniomyces Schw.
Filobasidium F. Sirobasidium Sir.
Guilliermondella G. Sporidiobolus Sporid.
Hanseniaspora H'spora Sporobolomyces S .
P
Hansenula H. Sporopachydermia Sporop.
Holtermannia Holt. Stephanoascus Steph.
Issatchenkia I. Sterigmatomyces St.
Kloeckera Kl. Sympodiomyces Symp.
Kluyveromyces κ. Torulaspora T'spora
Leucosporidium Leu. Torulopsis T.
Lipomyces L. Tremella Trem.
Lodderomyces Lod. Trichosporon Tr.
Malassezia Mal. Trigonopsis Trig.
Metschnikowia M. Wickerhamia W.
Nadsonia N. Wickerhamiella Wick.
Nematospora Nem. Wingea Wi.
Oosporidium 0. Zygo saccharomyces Zy go sac eh.
The following abbreviations are used for the names of type culture collections:
ATCC The American Type Culture Collection
CBS Centraalbureau voor Schimmelcultures; Yeast Division, Delft
LKB Laboratory of the Kodama Brewing Company
NRRL The Northern Regional Research Laboratory
1
Chapter I
General classification of the yeasts
A. Delimitation of the yeasts 1
B. Systems of classification of the yeasts 2
1. History 2
2. Present system of classification 2
a. Families and genera of the ascosporogenous yeasts 3
b. Families and genera of the basidiosporogenous yeasts 9
c. Families and genera of the imperfect yeasts 10
C. The genera Endomyces and Geotrichum 13
D. Principles of classification 15
1. The standard description 15
2. The species 16
3. DNA, enzymes and cell wall 17
a. DNA composition and relatedness 17
b. Similarity of enzymes 20
c. The cell wall 21
1. Ultrastructure of the cell wall 21
2. Chemical composition of the cell wall 22
4. Classification of the ascosporogenous yeasts 25
5. Classification of the basidiosporogenous yeasts 32
6. Classification of the imperfect yeasts 35
E. Nomenclature and type strain 42
F. The future of yeast taxonomy 43
A. Delimitation of the yeasts
by NJ.W. Kreger-van Rij
A yeast may be defined as a unicellular fungus reproducing by budding or fission.
Flegel (1977) has pointed to the advantages of this morphological definition which
leaves out other, historical considerations discussed by Lodder and Kreger-van Rij
(1952). Budding yeast cells may be a stage in the life cycle of multicellular fungi.
The yeast species in the present monograph are all characterized by a similar set
of features, both morphological and physiological. This type of description, in which