CRYPTOGAMIE

Mycologie

ANCIENNE REVUE DE MYCOLOGIE

fondée par R. Heim en 1936

Directeur de la publication : Hélène Bischler-Causse

Rédaction : Bruno DENNETIERE & Jean MOUCHACCA

EDITEUR : A.D.A.C. — 12 RUE BUFFON F-75005 PARIS

COMITÉ DE LECTURE

A. BELLEMÉRE (Paris), J. BOIDIN (Lyon), D. CHABASSE (Angers), R. COURTECUISSE

(Lille), G. DURRIEU (Toulouse), J. FAYRET (Toulouse), W. GAMS (Baarn), G. L. HENNEBERT

(Louvain-la-Neuve), P. JOLY (Paris), C. MONTANT (Toulouse), C. MOREAU (Brest),

D. N. PEGLER (Kew), M.-F. ROQUEBERT (Paris), B. SUTTON (Kew), G. TURIAN (Genéve),

D. ZICKLER (Orsay).

MANUSCRITS

Les manuscrits doivent étre adressés directement à la rédaction. La rédaction peut demander l'avis

d'un lecteur méme s'il n’appartient pas au Comité de Lecture. Bien qu'étant une revue de langue

francaise, les articles rédigés en anglais, allemand, italien et espagnol sont acceptés. Les disquettes de

micro-ordinateurs (PC ou Mac) sont vivement souhaitées. Les recommandations aux auteurs sont

publiées dans le fascicule 1 de chaque tome. Les auteurs recevront 25 tirés-à-part gratuits ; les

exemplaires suplémentaires seront à leur charge.

TARIFS DES ABONNEMENTS tome 18, 1997

CRYPTOGAMIE comprend trois sections : Algologie, Brtologie-Lichénologie, Mycologie

Pour une section : France : (350 F ht) 35735 F ttc Étranger : 380,00 F

Pour les 3 sections : France : (950 F ht) 970,00 F tte Étranger : 1050,00 F

Paiement par chèque bancaire ou postal à I[SAJordre de

A.D.A.C. — CRYPTOGAMIE (CCP La Source 34 764 05 S)

adressé à : A.D.A.C. 12 rue Buffon, F-75005 Paris

CRYPTOGAMIE, Mycologie est indexé par Biological Abstracts, Current Contents, Geo Abstracts,

GEOBASE, Publications bibliographiques du CNRS (Pascal).

Illustration de la couverture : Glomerella cingulata, dessin de A. M. Saccas

Source : MNHN. Paris

CRYPTOGAMIE

MYCOLOGIE

TOME 18 FASCICULE 1 1997

CONTENTS

COUTÉ A. & LAMY D. — Athanase Michel SACCAS 1911-1985 ....... 22 1

BOIDIN J. & LANQUETIN P. — Indices of useful informations for intercompa-

tibility tests in Basidiomycetes. VII Non poroid Aphyllophorales

(second supplement) И 2.20. 9

MOUCHACCA J. — Thermophilic fungi : biodiversity and taxonomic status ... 19

CHABASSE D. — Parasitic adaptation of soil keratinophilic fungi and its conse-

quences on human and animal pathology. ......... TT 71

BLANCO M. N., CHECA J. & MORENO G. New data on Trametes juni-

pericola Manjón, Moreno & Ryvarden ............................... 81

LIZÁRRAGA M., ILLANA C., MORENO G. & CASTILLO A. — Dydimum

clavodecus (Myxomycetes) an american species new to Europe ......... 87

Insttuctionstto'authors feme e e mb 91

МА

ө?

Bibliotheque Centrale Muséum

4 7

3 3001 0003339:

Source : MNHN, Paris

Cryptogamie, Mycol. 1997, 18 (1): 1-7 1

Athanase Michel SACCAS

1911-1985

A. COUTÉ! et D. LAMY?

! Museum National d'Histoire Naturelle, Laboratoire de Cryptogamie,

12 rue Buffon, F-75005 Paris

? UMS-CNRS 826, Lab. Cryptogamie, 12 rue Buffon,

F-75005 Paris — e-mail lamy@mnhn.fr

Athanase Michel SACCAS est né à Janina, en Gréce, le 10 mai 1911. À la fin de

ses études secondaires, il s'inscrit à l'Ecole Nationale d'Horticulture de Patras (Gréce)

d'oü il sort diplómé en 1931. De 1931 à 1933, il occupe un poste de chef de travaux dans

ce méme établissement et y exerce comme professeur d'horticulture de 1933 à 1938.

Source : MNHN. Paris

2 A. COUTÉ, D. LAMY

Abandonnant alors son pays natal, Athanase SACCAS vient en France oi il

intégre l'Ecole Nationale d'Agriculture de Grignon. Il obtient son diplóme d'ingénieur

agricole en 1941 et est alors recruté, comme attaché de recherches, par le CNRS, détaché

au laboratoire de Cryptogamie du Muséum National d'Histoire Naturelle (jusqu'en

1948).

En 1943, il soutient, à l'Université de Paris, une thése d'ingénieur-docteur es

sciences naturelles dont le sujet porte sur la morphologie et la biologie des Fusicladium

des Rosacées, champignons responsables des tavelures des Aubépines, Poiriers et Pom-

miers.

Fort de ses compétences tant en Biologie et Cytologie végétales qu'en Mycologie

et Pathologie végétale, il est nommé chef de travaux de Pathologie Végétale, Anatomie et

Biologie des champignons à l'Office de la Recherche Scientifique d'Outre-mer, róle qu'il

assume de 1946 à 1948.

Il est alors envoyé en République Centrafricaine, au Centre de Recherches

Agronomiques de Boukoko, avec le titre de Maitre de Recherche de 3* classe. Il y devient

successivement Maître de Recherche de 2°% classe en 1956 et, enfin, Directeur de Labo-

ratoire de 2° classe en 1957 et Directeur de Recherche, en 1960, dans le cadre de l'ORS-

TOM. Durant cette période, il assume, à la station de Boukoko, de 1948 à 1952, les

fonctions de chef du Service des recherches et des laboratoires et de chef de la division

Phytopathologie-Entomologie, de 1957 à 1958 celles de directeur par interim et, à partir de

1958, il devient directeur, poste qu'il occupe jusqu'en 1983. Il se retire en Savoie, région

«ой est originaire sa femme. Il y meurt le 20 juillet 1985.

Auteur de plus de quatre-vingt publications toutes d'excellente facture et d'une

bonne cinquantaine de notes et rapports, Athanase SACCAS a montré sans reláche ses

qualités de phytopathologiste. Il s'est intéressé à de nombreuses maladies cryptogamiques

de phanérogames d'intérét agronomique. Abordant, dans ses débuts, les problemes des

rouilles de Graminées et de Céréales (mais, riz, sorgho...), il s'est penché ensuite sur les

champignons papyricoles (Beauveria heimii, Sepedonium chartarum...). Poursuivant ses

travaux sur les rouilles, il s'est orienté également vers les parasites mycologiques de

l'arachide, des cacaoyers, caféiers (anthracnose, cercosporiose, fusariose, trachéomy-

.), cannes à sucre, colatiers, cotonniers, hévéas, maniocs, palmiers à huile, poivriers,

Ses recherches lui permirent ainsi de mettre au point des techniques de traite-

ment et de décrire, en méme temps, bon nombre de nouvelles espèces. Il a publié plusieurs

ouvrages portant sur les maladies cryptogamiques des plantes cultivées, en particulier du

café, pour l'Afrique Centrale et l'Afrique Equatoriale.

Homme de terrain trés actif, Athanase SACCAS ne s'est pas cantonné au

périmétre de la station de recherches de Boukoko. Il a, en effet, effectué de nombreuses

tournées et missions en Algérie, Brésil, Cameroun, Congo Belge, Cóte d'Ivoire, Gabon,

Israél, Mali, Tchad et Zaire. Au cours de certains déplacements, il a aussi fait profiter de

son expérience en Phytopathologie et Cryptogamie, de jeunes chercheurs et étudiants

africains en dispensant un enseignement de grande qualité. Il a été le représentant de la

République Centrafricaine, de l'IFCC et méme de l'UNESCO dans de nombreuses

manifestations internationales.

L'importance des travaux d'Athanase SACCAS, tant par leur qualité que leur

quantité, et leur impact sur les activités économiques des différents pays africains ou il est

intervenu, ont été reconnus sans conteste par ses autorités de tutelle. Il a ainsi été lauréat

de l'Académie d'Agriculture (1946) puis lauréat du Conseil Supérieur de la Recherche

Scientifique et du Progrès Technique (1956). П a été fait Chevalier (1951) puis Officier

Source : MNHN, Paris

ATHANASE MICHEL SACCAS 3

(1963) de l'Ordre du Mérite Agricole et Chevalier (1960) de l'Ordre des Palmes Académi-

ques. Па aussi été décoré de l'Ordre de la Légion d'Honneur (Chevalier еп 1960 et Officier

en 1970) et de l'Ordre du Mérite Français d'Outre-Mer.

Les pays africains qui ont bénéficié de ses travaux et compétences lui ont aussi

marqué leur reconnaissance de facon éclatante. C'est ainsi qu'il a été fait Grand Chance-

lier et Grand Croix (1959) de l'Ordre National du Mérite Centrafricain, Commandeur

(1963) de l'Ordre du Mérite Agricole du méme pays qui l'a également honoré du grade de

Commandeur (1964) de l'Ordre des Palmes Académiques de la République Centraficaine.

Il a aussi été élevé au grade de Grand Officier (1965) de l'Ordre de la République

Tunisienne et de Commandeur (1969) de l'Ordre du Mérite Tchadien.

Ce palmarès éloquent montre à l'évidence la remarquable activité scientifique

menée par Athanase SACCAS. Il a toujours fait preuve d'un sens profond du service

public allié à une passion sans limite pour les pays africains auxquels il a consacré toute

son énergie.

L'herbier, les notes et les manuscrits de Athanase SACCAS ont été déposés au

Laboratoire de Cryptogamie du Muséum National d'Histoire Naturelle (PC).

Publications de A.M. Saccas

1941

VIENNOT-BOURGIN G. et SACCAS A, — Morphose cladosporioide chez Fusicladium pirinum

Comptes Rendus hebdomadaires des Séances de l'Académie des Sciences 1941, 213(20)

701-704.

1945

SACCAS A. — Étude morphologique et biologique des Fusicladium des Rosacées.

«1944» 1945, 317 p., 2 pl., 59 fig. [Thèse Ingénieur Docteur].

Paris, Le Frangois

1946

DELAPORTE B. et SACCAS A. — Contribution à

Westling. Comptes Rendus hebdomadaires des

222 : 196-198 |

GUYOT A.L. (avec la collaboration de MASSENOT M., MONTÉGUT 1. & SACCAS А.)

Contribution à l'étude des cryptogames parasites de la France septentrionale I. Bulletin

trimestriel de la Société mycologique de France 1946, 62 : 69-85, 33 fig.

GUYOT A.L. (avec la collaboration de MASSENOT M., MONTÉGUT J. & SACCAS A.) —

Contribution à l'étude des cryptogames parasites de la France septentrionale П. Revue de

Mycologie 1946,11(2-3) : 53-73, 7 fig.

GUYOT A.L., MASSENOT M. et SACCAS A. — Études expérimentales sur les rouilles des

Graminées et des Céréales еп 1944. Annales de l'École Nationale d'Agriculture de Grignon,

3° sér., 1945-1946, 5: 33-81, 28 tabl., 2 pl. [publ. octobre 1946]

GUYOT A.L., MASSENOT M. et SACCAS A. — Considérations morphologiques et biologiques

sur l'espèce Puccinia graminis Pers. sensu lato. Annales de l'École Nationale d'Agriculture

de Grignon, 3* sér., 1945-1946, 5 : 82-146, 1 tabl. [publ. octobre 1946]

tude cytologique de Penicillium notatum

ances de l'Académie des Sciences 1946,

Source : MNHN, Paris

4 A. COUTÉ, D. LAMY

GUYOT A.L., MASSENOT M. et SACCAS A. — Études expérimentales sur les rouilles des

Graminées et des Céréales en 1945. Annales de l'École Nationale d'Agriculture de Grignon,

3° sér., 1945-1946, 5 : 213-266, 27 tabl. [publ. octobre 1946].

HEIM R. et SACCAS А. — La polyporose du groseillier due au Xanthochrous ribis (Schum.) Pat.

Comptes Rendus des Séances de l'Académie d'Agriculture de France 1946, 32 : 816-819.

1947

DELAPORTE B. et SACCAS A. — Étude morphologique et cytologique de plusieurs souches de

Penicillium notatum Westling. Annales de l'Institut Pasteur 73 : 850-861, 3 pl.

1948

GUYOT L., MASSENOT M., MONTEGUT J. et SACCAS A. — Au sujet d'une Urédinée vivant

sur le couple d'hôtes Clematis vitalba — Hordeum maritimum. Comptes Rendus hebdoma-

daires des Séances de l'Académie des Sciences 1948, 226 : 1921-1923.

GUYOT A.L., MASSENOT M., MONTÉGUT J. et SACCAS A. — À propos de la rouille jaune des

graminées (Puccinia glumarum). Comptes Rendus hebdomadaires des Séances de l' Acadé-

mie des Sciences 1948, 227 : 83-85. 4

GUYOT A.L., MASSENOT М. et SACCAS A. — Études expérimentales sur les rouilles des

Graminées et des Céréales en 1946. Annales de l'École Nationale d' Agriculture de Grignon,

3° sér., 1947-1948, 6 : 23-49, tabl. [publ. 2° semestre 1948]

GUYOT A.L., MASSENOT M. et SACCAS A. — Études expérimentales sur les rouilles des

Graminées et des Céréales en 1947. Annales de l'École Nationale d'Agriculture de Grignon,

3° sér., 1947-1948, 6 : 51-73, tabl. [publ. 2° semestre 1948].

GUYOT A.L., MASSENOT M. et SACCAS A. — Sept ans d'expérimentations (1941-1947) sur les

rouilles des Céréales. Annales de l'École Nationale d'Agriculture de Grignon, 3° sér.,

1947-1948, 6 : 75-117, tabl., 1 pl. [publ. 2° semestre 1948].

GUYOT A.L., MASSENOT M., MONTÉGUT J. et SACCAS A. — Au sujet d'un hóte écidien

nouveau de la rouille des œillets [Uromyces caryophyllinus (Schrank) Winter]. Revue de

Pathologie végétale et 4 Entomologie agricole de France 1948, 27(3) : 164-166.

SACCAS A. — Étude morphologique et biologique d'un nouveau champignon papyricole le

Beauveria heimii sp. nov. Revue de Mycologie 1948, 13(2-3) : 61-81, 5 fig.

1949

HEIM R., NOUVEL J., SACCAS A. — Sur un nouveau Penicillium à haut pouvoir antibiotique.

Bulletin de l'Académie Royale de Belgique, Classe des Sciences, 5° sér., 1949, 35 : 42-49, 1

fig., 1 graph

1950

SACCAS A. — Un Fusarium parasite des Panicules de Riz. Revue Internationale de Botanique

appliquée et d' Agriculture tropicale 1950, 30 (335-336) : 483-500, | fig. [publ. Févr. 1951]

SACCAS A. — Recherches préliminaires sur la trachéomycose du Coffea excelsa Chev. en Oubangui-

Chari. Bulletin de la Station centrale de Boukoko 1950, 1(1) : 3-43.

HEIM R. et SACCAS A. — La Trachéomycose des Coffea excelsa et robusta des plantations de

l'Oubangui-Chari. Comptes Rendus hebdomadaires des Séances de l'Académie des Sciences

1950, 231 : 536-538. [aussi publié dans Revue internationale de Botanique appliquée et

d'Agriculture tropicale 1950, 30 (337-338) : 640-642]

SACCAS A. — Étude sur la protection et la défense des cultures contre les cryptogames parasites en

A.E.F. In: Encyclopédie maritime et coloniale. [non vu]

Source : MNHN, Paris

ATHANASE MICHEL SACCAS 5

SACCAS A. — Note sur les possibilités d'emploi des hélicoptères dans la lutte contre les maladies des

plantes cultivées en A.E.E Institut Francais de transports aériens. Note de travail 184-185,

17 p. [non vu]

SACCAS A. — Étude sommaire des champignons parasites des plantes culti

industrielles en A.E.F. Bulletin de la Station de Boukoko 2 [non vu].

, alimentaires et

1951

SACCAS A. — Un nouveau champignon ascomycéte gymnoascé, |` Eidamella papyricola nov. sp.

Bulletin trimestriel de la Société mycologique de France «1950»1951, 66(3) : 121-138, 4 fig.

GUYOT A.L., MASSENOT M. et SACCAS А. — Les Rouilles des Ononis. Uredineana (A.L. Guyot

ed.) 1951, 3: 112-119, 3 fig ;

GUYOT A.L., MASSENOT М. et SACCAS A. — À propos des Отот)

Uredineana (A.L. Guyot ed.) 1951, 3 : 120-124.

GUYOT A.L., MASSENOT M., MONTÉGUT J. et SACCAS A. — Sur deux espéces nouvelles

d'Urédinées parasites d' Avena bromoides. Uredineana (A.L. Guyot ed.) 1951, 3 : 141-150,

6 fig.

SACCAS A. — Étude morphologique, biologique et expérimentale d'un Fusarium ravageur des

cultures de riz à la Station Centrale de Boukoko (A.E.F.). Revue de Pathologie végétale et

d'Entomologie agricole de France 1951, 30(2) : 65-96, 3 pl., tabl.

SACCAS A.M. et DROUILLON R. — Étude macroscopique et microscopique de quelques cham-

pignons parasites des Aleurites en Afrique Equatoriale Frangaise. Agronomie tropicale

1951, 6(5-6) : 239-264, 18 fig.

SACCAS A.M. — La trachéomycose (Carbunculariose) des Coffea excelsa, neo-Arnoldiana et

robusta en Oubangui-Chari. Agronomie tropicale 1951, 6 (9-10) : 453-506, 16 fig.

SACCAS A.M. — À propos de quelques champignons nouveaux parasites et saprophytes sur mais.

Revue de Pathologie végétale et d° Entomologie agricole de France 1951, 30(3) : 161-196, 4 pl.

ез parasites des Astragalus.

1952

SACCAS A.M. — Principaux champignons parasites du mais (Zea mays L.) en Afrique Equatoriale

Française. Agronomie tropicale 1952, 7(1) : 5-42, 20 fig.

1953

SACCAS A. — Note sur les Coleroa. Revue de Mycologie 1953, 18(1) : 56-63, 3 fig.

SACCAS A. — Une nouvelle espéce de Coleroa sur Caféier. Revue de Mycologie 1953, 18, suppl.

colonial 1 : 46-49, 3 fig.

SACCAS A.M. — Les principales maladies cryptogamiques de l'hévéa en A.E.F. Agronomie tropicale

1953, 8 (2) : 176-198, fig. 1-10.

SACCAS A.M. — Les principales maladies eryptogamiques de l'hévéa en A.E.F. (suite). Agronomie

tropicale 1953, 8(3) : 229-285, fig. 11-42

1954

SACCAS A.M. — La flétrissure des cotonniers du Tchad due à Olpidiaster ( Asterocystis) gossypii

n. sp. Cotons et Fibres tropicales 1954, 9(1) : 23-55, 20 fig.

SACCAS A.M. et FERNIER H. — Une grave maladie du riz due à Ophiobolus oryzinus Sacc.

Agronomie tropicale 1954, 9(1) : 7-20, 7 fig.

SACCAS A.M. — Les champignons parasites des sorghos (Sorghum vulgare) et des Penicillaires

(Pennisetum typhoideum) en Afrique Équatoriale Frangaise. Agronomie tropicale 1954,

9(2) : 135-173, fig. 1-21.

SACCAS A.M. — Les champignons parasites des sorghos (Sorghum vulgare) et des Penicillaires

(Pennisetum typhoideum) en Afrique Equatoriale Française (suite). Agronomie tropicale

1954, 9(3) : 263-301, fig. 22-42.

Source : MNHN, Paris

6 A. COUTÉ, D. LAMY

SACCAS A.— Les champignons de l'Hévéa en Afrique Équatoriale Française. Journal d'Agriculture

tropicale et de Botanique appliquée 1954, 1(5-6) : 180-203, fig. 1-11.

SACCAS A. — Les champignons de l'Hévéa en Afrique Équatoriale Française (suite).

Journal d'Agriculture tropicale et de Botanique appliquée 1954, 17-9) : 333-355, fig. 12-

22.

SACCAS A. — Les champignons de l'Hévéa en Afrique Équatoriale Française (suite et fin). Journal

d Agriculture tropicale et de Botanique appliquée 1954, 1(10-12) : 461-481, fig. 23-30.

1955

SACCAS A.M. — Les champignons parasites des sorghos (Sorghum vulgare) et des Penicillaires

(Pennisetum typhoideum) en Afrique Equatoriale Française (suite et fin). Agronomie

tropicale «1954» 1955, 9(6) : 647-686, fig. 43-53. [publ. Janv. 1955].

SACCAS A.M. — Les variations dans l'aspect macroscopique et biométrique du Fusarium xylarioi-

des Stey., agent de la trachéomycose des Coffea, liées aux races physiologiques. Atti del VI

Congresse Internazionale di Microbiologia ( Roma, sett. 1953). Roma, 1955. Vol. V, sezione

14 : 208-216, 1 graph.

SACCAS A.M. — La fusariose des fruits des caféiers en Oubangui-Chari due à Fusarium equiseti Var.

intermedeum n. var. Agronomie tropicale 1955, 10(1) : 43-59, 7 fig., 2 graph.

SACCAS A.M. — La rouille américaine du mais (Zea mays L.) due à Puccinia polysora Underw. au

Cameroun et en Afrique Equatoriale Française. Agronomie tropicale 1955, 10(4) : 499-522,

16 fig., 2 graph.

1956

SACCAS A.M. — Recherches expérimentales sur la trachéomycose des caféiers en Oubangui-Chari.

Agronomie tropicale 11(1) : 7-38, 9 fig., 1 carte.

SACCAS A.M. — Les Rosellinia des caféiers en Oubangui-Chari. Agronomie tropicale 1956, 11(5):

551-595, fig. 1-26.

1957

SACCAS A.M. — Les Rosellinia des caféiers en Oubangui-Chari (suite et fin). Agronomie tropicale

«1956» 1957, 11(6) : 687-706, fig. 27-34 [publ. Janvier 1957].

SACCAS A.M. — La maladie des taches zonées de Coffea excelsa en Oubangui-Chari, due à

Sclerotium coeffeicolum Stahel. Revue de Mycologie 1957, 22, suppl. colonial 2 : 65-84,

2fig., pl. 1.

1959

SACCAS A.M. — Une grave maladie des hévéas des Terres Rouges en Oubangui-Chari. Agronomie

tropicale 1959, 14(4) : 409-458, 22 fig.

SACCAS A.M. — La rouille du caféier menace-t-elle les plantations de Canephora ? Café Cacao Thé

1959, 3(2) : 87-91.

1966

SACCAS A.M. — La pourriture du collet des plantules de caféiers en pépinière, due à Rhizoctonia

bataticola (Taub.) Butler. Café Cacao Thé 1966, 10(1) : 28-45, 8 fig.

1969

SACCAS A.M. et CHARPENTIER J. — L’anthracnose des caféiers Robusta et Excelsa, due à

Colletotrichum coffeanum Noack en République Centrafricaine. Bulletin de l'Institut Fran-

«ais du Café et du Cacao 1969, 9 : 1-84, 31 fig.

Source : MNHN, Paris

ATHANASE MICHEL SACCAS 7

1971

SACCAS A.M. et CHARPENTIER J. — La rouille des Caféiers due à Hemileia vastatrix Berk. et Br.

Bulletin de l'Institut Français du Café et du Cacao 1971, 10 : 1-123, 24 fig.

1972

SACCAS A.M. — La rouille «farineuse» des caféiers, due à Hemileia coffeicola Maub. et Rog. Bulletin

de l'Institut Francais du Café et du Cacao 1972, 11 : 1-68, 32 fig.

1975

SACCAS A.M. — Les pourridiés des caféiers en Afrique tropicale. Bulletin de l'Institut Francais du

Café et du Cacao 1975, 13 : 1-172, 54 fig.

1981

SACCAS A.M. — Etude de la flore cryptogamique des caféiers en Afrique centrale. Bulletin de

l'Institut Français du Café et du Cacao 1981, 16 : 1-522, 138 fig.

Communications à des conférences [documents dactylographiés, non publiés]

SACCAS A.M. — Essais comparatifs de désinfection des semences par voie humide et séche contre

les maladies charbonneuses des sorghos à la Sation Centrale de Boukoko (A.E.F).

Congrés de la Protection des végétaux et de leurs produits sous les climats chauds. Marseille,

Septembre 1954, 13р. [non vu]

SACCAS A.M. — Résultats des essais de lutte contre la «pourriture brune» due à Phytophthora

palmivora, la «pourriture noire» due à Botryodiplodia theobromae et la «pourriture fari-

neuse» due à Trachysphaera fructigena. Congrés de la Protection des végétaux et de leurs

produits sous les climats chauds. Marseille, Septembre 1954, 7p. [non vu]

SACCAS A.M. — Résultats des recherches et expérimentations concernant la trachéomycose des

caféiers en Oubangui-Chari. Congrès de la Protection des végétaux et de leurs produits sous

les climats chauds. Marseille, Septembre 1954, 25p. [non vu]

SACCAS A.M. — La trachéomycose des Coffea excelsa, neo-arnoldiana et robusta en Afrique

Equatoriale Française. Conférence Franco-Belge sur la trachéomycose des caféiers d'Afri-

que. Yangambi, Avril 1956. 61 p. [non vu]

Deux références, données par Saccas dans sa Notice de Travaux, demeurent problématiques, pour

lesquelles toute information sera la bienvenue :

SACCAS A.M. — Champignons et insectes parasites des fruits. In : G. Delbard, Les beaux Fruits de

France. Paris, ed. Delbard, 1947. [L'exemplaire que nous avons consulté à la Bibliotheque

Centrale ne porte aucune mention d'auteurs ou de collaborateurs pour certaines parties du

livre. La partie intitulée «Parasitologie fruitiére» est précédée d'un avant-propos signé par

R. Heim]

SACCAS A.M. — Traité sur les champignons parasites, parasites secondaires et saprophytes en А (frique

Equatoriale. Paris, Maisonneuve et Larose, 1971, 2 tomes, 1250р. [Cet ouvrage n'a pu être

localisé ; de plus, l'éditeur nous a affirmé n'avoir rien publié de tel]

Source : MNHN, Paris

Source : MNHN. Paris

Cryptogamie, Mycol, 1997, 18 (1) : 9-18 9

RÉPERTOIRE DES DONNÉES UTILES j

POUR EFFECTUER LES TESTS D'INTERCOMPATIBILITÉ

CHEZ LES BASIDIOMYCETES.

vu— APHYLLOPHORALES NON PORÉES

(DEUXIEME SUPPLÉMENT)

Jacques BOIDIN* & Paule LANQUETIN

* 17 rue Duguesclin, 69006 Lyon, France

Cette note fait suite aux 6 articles parus dans cette méme revue (Boidin &

Lanquetin, 1984 a, b et apellano, 1985 ; Lamoure, 1989 ; Boidin, 1990).

Pour la signification détaillée des signes, nous renvoyons le lecteur à l'introduc-

tion, ou aux résumés situés en téte des parties III et V (Boidin & Lanquetin, 1984c :

Lamoure, 1989). Rappelons simplement : h, hétérothalle ; II bipolaire ; IV, tétrapolaire ;

H, homothalle ; H, homothalle présumé ; P, parthénogénétique ; A, amphithalle ; u,

uninucléé ; d, dic: yotique ; p, plurinucléé ; m, multinucléé. u/d signifie que la culture

monosperme passe spontanément du stade uninucléé au stade dicaryotique car elle est

homothalle.

Pour les boucles : b, présentes sans précision ; a, absentes ; c, constantes ; b

Inconstantes ; r, rares ; va, variables selon l'aération 3 0, parfois opposées ; v, verticillées

sur les hyphes les plus larges.

Comportements nucléaires : N, normal ; SN, subnormal ; He, hétérocytique ;

HC, holocénocytique ; As, astatocénocytique ; HM, holomonocaryotique ; HD, holodi-

caryotique.

Vitesse de croissance : nombre de semaines pour que le rayon de la culture couvre

la boite de Pétri de 90 mm de diamètre ; 7, plus de 6 semaines.

L'astérisque signifie : « avec irrégularité » ; p. ex. : d*, dicaryotique avec quelques

articles a 1, ou à 3 ou 4 noyaux.

Nous ne résumerons que les données nouvelles ou complémentaires des index

publiés (1984 et 1990) ; le Cystostereum pini-canadense figure dans l'index de 1990 mais sa

polarité n'était pas connue ; découverte depuis, elle sera indiquée ici.

Source : MNHN, Paris

1. BOIDIN & P. LANQUETIN

2 Е

8e © gle 4

s| BIB 858 ©

| ВЕЕ Ы. 288 8

ESPÉCES ss LS. alesi s E 8| &

a |° 1 8 | 2183 a 35 š S Ë

Ela is [2/6 42822) à

icanthophysata Léger ar | Helwan |. 27

ІНутепосһаеге

сегіпа Peck ш As | va 33

'hlebia

iculeatum Wu d a 1 39

Glococystidiellum

ДІ Mackawa hIV| 1 c А 28

"erinomyces

idjacens Boid., Lanq. & Gilles H|1]|wd|d*|N|r | 2 8

eniophora

lfîbulatum (G. Н. Cunn.) Boid. et al. NO | Же рс ү 7 12

fegalocystidium

africanum Boid., Lang. & Gilles BLT 2 lep |d* Hele |. 7 12

fegalocystidium.

ilbicans (Pers.) Nakas. hil c | 4 29

Hyphoderma

(Шыда Post : Fr. hil va | 2-3 29

|Phiebia

ni Boid., Lang, & Gilles uv|2 | p | @ | нер e | 6-7 п

Vuilleminia

(americanus Nakas. et al. hil c 30

Ceraceomyces

irachnoideum Langer d a 22

[Botryobasidium

ispellum Hjortst. hil} 1 u d | N © 7 39

VGloeocystidiellum

sterospora Bold. & Lang. Ho БЕДИ ТИ Ol jel 7

Dendrothele

icornis Hjortst. & Куу. H|1T[|wd|d*|N| Tr |23 8

'eniophora

ispora Burds. & Nakas. Еа e Te 7

"Dendrothele"

Source : MNHN. Paris

INTERCOMPATIBILITÉ CHEZ LES BASIDIOMYCÈTES

isporum Boid., Lang. & Gilles Р | тъ [а [нмја [7 12

[Gloeocystidiellum

orbonica Boid., Lang. & Gilles h 1 u [dt | N a | 4-6 12

[Boidinia

|е. borbonica Boid., Lang. & Gilles БЕКШЕ e rose кат ТЕ 12

[Boidinia

osei De МЇ 14

lEpithelopsis

Potryodeum (Overh.) Parm. H ids 22

[Botryobasidium

Ibrunnea Wu 2|m|m]|HC| o 2 38

IPhanerochaete

þurti (Romell)Parm. v | 2-3 29

'hanerochaete

'ana Wu H 1 ша | N | a [4-5 12

Boidinia

anariensis (Manj. & Mor.) Hjortst. HS LS ap A 7

IDendrothele

andicans Erikss. H d | d [HD] a 22

lotryobasidium

carnosa (Burt) Parm. у |03 29

IPhanerochaete

Earpatica Pilar Н|1|4|4|н|а [7 27

'lymenochaete

entrifuga Karst. HII | 2| m|md|As| va] 2 29-23

[Phlebia

cervicolor (Berk. & Curt.) Massee a | 6-7 29

Isterostroma

'hrysosporium Burds. H? a | 1-2 29

hanerochaete.

lavuligerum (Höhn. & Litsch.) Nakas. МУ| 1 u d| N c 7 12

[Gloeocystidiellum '

'offeana Lég. & Lang, h eue | era iere UE 26

Jymenochaete.

olumbiensis (Burt) Burds. & Lomb, hil 17

Gloeodontia. Ou

'ompactum Wu h 2 u d [53| c [5-6 39

[trian

'onicum (Oberw.) Erikss. & Hjortst. nuc oat [Ee SEES TES E 16

IRepetobasidium

[conspersum Erikss. d a 22

Botryobasidium

1. sensu Hallenberg, voir vesiculosum.

Source : MNHN, Paris

12 J. BOIDIN & P. LANQUETIN

fontiformis G. Н. Cunn. H d [а {нра 27

Hymenochaete

rassitunicata Boid., Lang. & Gilles Hv|1luld|NJ|ec]|23 8

[Peniophora

urtisii Hallenb. d a 22

lotryobasidium

lenticulata Lég. & Lang. н [1 [а |а [ноја [7 24

Hymenochaete

duportii Pat. 1 m | т |HC| a 7 27

Hydnochaete

laeidis Boid., Lang. & Gilles H?| 1 |wd*| df | N r 2 8

eniophora

erikssonii Hall. & Hj. hlV | 1 u d n c 4 18

IHypochnicium

xilis (Burt) Burds. v |25 29

'hanerochaete

farinaceus Boid., Lang. & Gilles hoj AENA ЕА, 10

Imauromyces

fimbriatum Burds. d a |34 12

Xloeocystidiellum

fissilis Boid., Lang. & Gilles Hv|r|u|d|NJ|e]|34 8

[Peniophora

formosanum Wu hH | 2 u d |SN| c Ел 53

loeocystidiellum

fouquieriae Nakas, & Gilb. hI b | 4-6 31

Hyphoderma

fuliginosa (Pers.) Lév. ЕО PHD} | | 2 27

Hymenochaete

[gabonensis Boid., Lang. & Gilles H|1j|watd*|N|r|2 8

[Peniophora

lobosa Wu h 2 u|d|SN|a 7 12-39

loiothele

graminicola Wu d* ov | 4 39

[Gloeomyces

ranulata Wu. hlT 1 u d N с 6 12-39

\Boidinia

rrandisporum G. Langer d a 22

[Botryobasidium

|griseo-flavescens (Litsch.) Erikss. & Hjortst.| hII | 1 (re || ale) PINS сс y: 16

[Phlebia

[guadelupensis Boid. $: Lang. h 1 u | d r 2 8

eniophora

jauerslevii Léger air 4 |а ЕЛ a | 7 27

|Нутепосһаеге

w GT E h 2 u p sN] а TE 12

loiothele.

incarnatum По & Imai ҺП b 5 34

Gloeostereum.

lindica (Thind & Rat.) Rat. ШУ 14

fetulodontia

[insidiosum (Bourd. & Galz.) Hallenb. H|2|m|ml]|HC|a (4-5 | со2| 12

"onferticium.

Source : MNHN, Paris

INTERCOMPATIBILITÉ CHEZ LES BASIDIOMYCETES 13

isabellinum (Fr.) Rogers H Рр |р 22

IBotryobasidium.

nyense Hjortst. Hv| 1 Î u Î d с |5-7 12

Gloeocystidiellum

uehneri (Boid. & Lang.) Hjortst. nv] 1 | u | d e | 3 8

iDuportella

uehneroides Boid., Lang. & Gilles hv| 1 Î u | d єз 8

[Duportella

laeve (Erikss.) Parm. H SE a 22

lotryobasidium

lamellosa (P. Henn.) Bres. 2. ч d |SN| a 7 12

iloiothele

laxa (Wu) Boid., Lang. & Gilles 1 d MES, 12-39

loeopeniophorella

lembospora (Bourd.) Oberw. h 1 u d N c 4 16

thelopsis

leprosa (Bourd. & Galz) Boid, Lang. & hI | 1 | p | p [HC] i [3-4 10

jilles

Scopuloides

ilascens (Bourd.) Erikss. & Hjortst.

'hlebia gr. 1 hwl a a Ка

gr.2 hiv] 1] u [а 20

gni hil u | dp

litschaueri (Burt.) Erikss. & Strid. hil e | 6-7 29

IHyphoderma

longicystidium (Litsch.) Мака». hil iv 15

rustoderma

longisporum G. Langer d a 22

Botryobasidium.

jundellii (Bourd.) Erikss. ШУ с |3-4 29

IHypochnicium

huridum (Bres.) Jal. H[2[|pd|d|He|c|7 12

fegalocystidium

luteo-cystidiatum (Talbot) Wu НГ ETE ЕЯ 12

fegalocystidium

macrospora Wu z d d |HD| c 6 39

|'Boidinia"

magnoliae (Berk. & Curt.) Burds. ii p | p |HC| v 2

Phanerochaete

malaiensis Boid., Lang. & Gilles ӘЛГІ aS ЕЕ 8

Peniophora

medioburiense (Burt) Donk H 17

IHyphoderma

melzeri Pouzar му c | 6 29

“ytidiella

Inicrospora Jacks. & Lemk. шу e 29

Dendrothele

nicrosporella Jūl. hl! с 4 29

drandinia.

minuscula G. H. Cunn. НӘ е Ба ғаз шу Бал 2 27

(Hymenochaete

Source : MNHN., Paris

14 J. BOIDIN & P LANQUETIN

irificum Erikss. hH | 1 u|d|N 7 16

tepetobasidium

[molesta Boid., Lang. & Gilles ALY? ara | М 2 8

[Peniophora

Ше (Fr.) Hjortst. hIV| 1 u|du|N 16

[ypochnicielum.

ioniliforme Wu hiv| 1 u d|N 6-7 12

lloeocystidiellum.

f. montanum Ginns & Fr. H| 2 7 12

Megalocystidium

hnonticola Boid., Lang. & Gilles hV| 1 | u | d 4-5 8

[Peniophora

mutabilis (Bres.) Jül. hH | 1 u d d 16

lFibulomyces

ranospora Léger. 1 d* 4 27

menochaete

iveo-cremeum (Höhn. & L.) Donk hil 29

istotrema

btusisporum Erikss. d 22

[Botryobasidium

pchromarginata Talbot H 1 | d* | d* | HD 7 27

Hymenochaete

livascens (Bres.) Larss. & Hjortst. H 17

Irevicellicium.

pvalispora Boid., Lang. & Gilles hIV| 1 Ca 3-4 8

[Peniophora

parvocystidiata Boid. & L. h 1 u | d* 2-3 8

епіорһоға

roxydata (Rick) Hjortst. 1 d 4-5 12

oldie

Hee Boid. & Gilles d 22

lotryobasidium

[ше (Schwein.) Gilberts hIV 3-6 29

Cystostereum

pinnatifida Burt H 1 d* | d* | HD i 27

'[ymenochaete.

paraa (Burt) Nakas. ШУ 32

icytinostroma

ubsp. septentrionale Nakas hIV 32

[pruinatum (Bres.) Erikss. H p p 22

[Botryobasidium

pseudo-adusta Lég. & Lang. HET TISTF [нс x 25

[Hymenochaete

pseudo-cystidiata Boid., Lang. & Gilles — |hIV| 2 | u | d | SN 4-5 11

Vuilleminia

AR mn Novob hII* col 36

Sporotrichum

urpureum Wu ыу 1 Î u Î d Î N 4 39

iloeocystidiellum

uteana (Schum. : Fr.) Karst. hil 1

Source : MNHN, Paris

INTERCOMPATIBILITÉ CHEZ LES BASIDIOMYCETES 15

(queletii (Bourd. & Galz.) Christ. hil va | 3-5 29

[Phlebia

“aduloides (Karst.) Donk hil 29

Sistotrema

avum (Burt) Ginns & Freem. H| 2 jm | m [HCI ov | 3-4 39

'onferticium

Гепізроға Boid., Lang. & Gilles H?| war a LN el 8

IDuportella

bacratum (G. H. Cunn.) Wu ov 35.

extrinocystidium.

'almonea (Burt) Boid., Lang. & Gilles H|2|m|mj|HC|ov| 3 12

iloeocystidiopsis

scintillans G. H. Cunn. hiv] 1 Î u N | c | 3-4 8

'eniophora

etulosa (Berk. & Curt.) Nakas. HI va | 2 29

hlebia.

simile Pouz. & Hol. H dE a 22

[Botryobasidium

Isinuosum Freem. шу 14

loeocystidiellum

onorae Nakas. & Gilb. ът с [6-7 29

ristinia

spathulata Léger 1 d* a 7 27

ІНутепосһаеге

tereoides Wu 2|m|m]|HC| o 2 38

Phanerochaete

ubasperispora (Litsch.) Jül. fepe ЕН NA е |$ 12

Boidinia

ubcoronatum (Höhn. & Litsch.) Donk Шу 22

[Botryobasidium

ubcrelacea (Litsch.) Christ. UPSC S EN e | 7 16

\Phlebia

subglobispora Hall. & Hjortst. hIV 19

Vuilleminia

hubsalmonea Boid., Lang. & Gilles if qr ESA ЕСЕ 8

Peniophora

amariciphila Boid., Lang. & Gilles H|2|plpr|HC|a [3-4 9

'hanerochaete

inctorium (ЕП. & Ev.) Ell. & Ev. HV|1|u|d|NJ|ec 37

lEchinodontium

'agum (Berk. & Curt.) Rogers H RES 22

[Botryobasidium

variolosum Boid., Lang. & Gilles H|1[|wd|d|N|*c]|2|Co| 8

IHyphoderma

vesiculosum (Burt) Boid., Lang. & Gilles | МУ 17

loeocystidiellum

iile (Bourd. & Galz.) Erikss. H “| 4 | ЧЮ < | 7 16

IRepetobasidium

іпоѕа (Overh.) Nakas. ШП va | 2-3 29

Phlebia

2. appelé Gloeocystidiellum clavuligerum en Amérique du Nord, mais incompatible avec cette espèce

Européenne.

Source : MNHN, Paris

16 J. BOIDIN & P. LANQUETIN

(сінігі (Burds. ег al.) Wu H|2 p | p | HC] ov | 3-4 12-39

Stereum

A ce résumé des données publiées, nous ajouterons, ci-après, quelques données

obtenues à Lyon et non publiées à ce jour.

1 m [HC] a [5-6

2 ч SN|c 3

larmeniacum Boid. & Gilles TX 2 а En НС v 3

[Stereum

layresii (Berk.) Boid. & Gilles hiv] 1 u d |N с 3

Hyphoderma

|cremo-album (Höhn. & Litsch.) Jül. hI | 1 uade NS SC 7

Hyphoderma

lerikssonii Oberw. hi} 1 mel ЕМ e 7

|Repetobasidium

lexpallens (Bres.) Domanski hiv} 1 u d N c 4

Dentocorticium

emmiferum (Bourd. & Galz) Erikss. & HIV] 1 | u | d | N | ce | 3

Куу.

|Coronicium

heimii Malengon һ [2 [р Га [не [е [7

Veluticeps.

|incrustatissimum Boid. & С. hH | 1 u d | N | c |45

Hyphoderma

[laetum (Karst.) Erikss. & Hjortst. ЖІГГІЗ E “ГЕ |e

Erythricium

Imelzeri (Pouzar) Stalpers H 2 4 d |HD| c 6

uriculariopsis.

seudotsugae (Burt) Boid. & Gilles AAA а IST E E]

{phanobasidium

[scaevolae Boid. & Gilles i JE E TE Pets || tS T

Iseptocystidia (Burt) Burds. 1 p | НС а 7

(Candelabrochaete

ubcrinale (Peck) Куу. р [а [Hel a | 7 |

[iransiens (Bres.) Parm. hI TT [а [ам [е | 3 |

3. L'homothallie constatée en France contraste avec la tétrapolarité d'une récolte Américaine

(Nakasone, 1990)

Source : MNHN, Paris

INTERCOMPATIBILITÉ CHEZ LES BASIDIOMYCETES 17

BIBLIOGRAPHIE

1 — AINSWORTH A. M. & RAYNER A. D. M., 1990 — Mycelial interactions and outcrossing in

the Coniophora puteana complex. Mycological research 94 : 627-634.

2 — AINSWORTH A. M. & RAYNER A. D. M., 1991 — Ontogenic stages from coenocyte to

basidiome and their relations to phenoloxydase activity and colonization process in

Phanerochaete magnoliae. Mycological research 95 : 1414-1422.

3 — BOIDIN J., 1990 — Répertoire des données utiles pour effectuer les tests d'intercompatibilité

chez les Basidiomycétes. VI. — Aphyllophorales non porées (Premier supplément). Cryp-

togamie, Mycologie 11 : 175-188

4 — BOIDIN J. & LANQUETIN Р, 1984a — Répertoire des données utiles pour effectuer les tests

d'incompatibilité chez les Basidiomycètes. I. — Introduction. Cryptogamie, Mycologie 5 :

33-45.

5 — BOIDIN J. & LANQUETIN P, 1984b — Répertoire des données utiles pour effectuer les tests

d'incompatibilité chez les Basidiomycètes. II. — Saprophytic Phragmobasidiomycétes.

Cryptogamie, Mycologie 5 : 47-50.

6 — BOIDIN J. & LANQUETIN Р, 1984c — Répertoire des données utiles pour effectuer les tests

d'incompatibilité chez les Basidiomycétes. Ш. — Aphyllophorales non porées. Cryptoga-

mie, Mycologie 193-245.

7 — BOIDIN J., LANQUETIN P. & DUHEM B., 1996 — Contribution à la connaissance du genre

Dendrothele (Basidiomycotina, Aphyllophorales). Bulletin de la société mycologique de

France 112 : 87-126

8— BOIDIN J., LANQUETIN P. & GILLES G., 1991 — Les Peniophoraceae de la 26ne intertropi-

cale (Basidiomycetes, Aphyllophorales). Bulletin de la société mycologique de France 107:

91-156.

9 — BOIDIN J., LANQUETIN P. & GILLES G., 1993 — Contribution à la connaissance des

Phanerochaetoideae de France (Basidiomycotina). Cryptogamie, Mycologie 14 : 195-

206.

10 — BOIDIN J., LANQUETIN P. & GILLES G., 1993 — Basidiomycetes Aphyllophorales de l'ile

de la Réunion. XVII — Les genres Amauromyces, Cunninghammyces et Repetobasidium.

Bulletin de la société mycologique de France 109 : 93-100.

11 — BOIDIN J., LANQUETIN P. & GILLES G., 1994 — Contribution à la connaissance du genre

Vuilleminia (Basidiomycotina). Bulletin de la société mycologique de France 110 : 91-

107.

12 — BOIDIN J., LANQUETIN P. & GILLES G., 1997 — Le genre Gloeocystidiellum sensu lato

(Basidiomycotina). Bulletin de la société mycologique de France (sous presse).

13 — CAPELLANO A., 1985 — Répertoire des données utiles pour effectuer les tests d'intercom-

patibilité chez les Basidiomycétes. IV. — Gastéromycétes. Cryptogamie, Mycologie 6 :

65-68.

14— DE A. B., 1990 — Interfertility studies on the species of Corticiaceae. Mycotaxon 37 : 63-65.

15 — GINNS J. & CLARCK J., 1989 — Crustoderma longicystidia associated with decay of lumber

in British Columbia and the cultural features of C. dryina. Mycologia 81 : 921-926.

16— HALLENBERG N., 1990 — Cultures studies in Corticiaceae (Basidiomycetes), Ш. Windahlia

18 : 25-30.

17 — HALLENBERG N., 1991 — Pairing tests with species of Aphyllophorales (Basidiomycetes)

from two phytogeographically isolated areas. Mycotaxon 42 : 355-386.

18 — HALLENBERG N. & HJORTSTAM K., 1989 — Hypochnicium erikssonii sp. nov. instead of

H. sphaerosporum, a necessary name alternation. Windahlia 18 : 43-46.

19 — HALLENBERG М. & HJORTSTAM K., 1996 — Four new species of Corticioid Fungi

(Basidiomycotina, Aphyllophorales) from Argentina. Mycotaxon 57 : 117-123.

Source : MNHN, Paris

18 J. BOIDIN & P. LANQUETIN

20— HALLENBERG N. & LARSSON E., 1991 — Differences in cultural characters and electro-

phoretic patterns among sibling species in four different species complexes (Corticiaceae,

Basidiomycetes). Mycologia 83 : 131-141.

21 — LAMOURE D., 1989 — Répertoire des données utiles pour effectuer les tests d'intercompati-

bilité chez les Basidiomycétes. V. — Agaricales sensu lato. Cryptogamie, Mycologie 10 :

41-80.

22 — LANGER G., 1994 — Die Gattung Botryobasidium Donk (Corticiaceae, Basidiomycetes).

Bibliotheca Mycologica 158, 459 p.

23 — LANQUETIN P. & MEYER M., 1991 — Récolte de Phlebia centrifuga Karst. en Savoie et

étude des mycéliums. Bulletin de la fédération mycologique Dauphiné-Savoie 123 : 9-12.

24 — LÉGER J. С. & LANQUETIN P., 1992 — Hymenochaete denticulata Léger et Lanquetin.

Persoonia 14 : 369-375.

25 — LEGER J. C. & LANQUETIN P., 1992 — Hymenochaete pseudoadusta nov. sp. (Basidiomy-

cétes, Aphyllophorales) : description et caractères culturaux. Cryptogamie, Mycologie 13 :

305-311.

26 — LÉGER J. C. & LANQUETIN Р, 1994 — Hymenochaete coffeana nov. sp. (Basidiomycotina,

Aphyllophorales) : Description et caractères culturaux. Cryptogamie, Mycologie 15 :

21-26.

27 — LÉGER J. C. & LANQUETIN P. 1996 — Contribution à l'étude des caractères culturaux chez

les Hymenochaetaceae (Basidiomycotina). Cryptogamie, Mycologie 17 : 105-121.

28 — MAEKAWA N., 1987 — A new species of the genus Cerinomyces. Canadian journal of botany

65 : 583-588.

29 — NAKASONE K. K., 1990 — Cultural studies and identification of wood-inhabiting Corticia-

ceae ans selected Hymenomycetes from North America. Mycological Memoirs 15, 412 p.

30 — NAKASONE K. K., BERGMAN C. R. & BURDSALL H., 1994 — Phanerochaete

filamentosa — Corticium radicatum species complex in North America. Sydowia 46 : 44-62.

31 — NAKASONE K. K. & GILBERTSON R. L., 1978 — Cultural and other studies of fungi that

decay ocotillo in Arizona. Mycologia 70 : 266-299.

32 — NAKASONE K. K. & MICALES J. A., 1988 — Scytinostroma galactinum species complex in

the United States. Mycologia 80 : 546-559.

33 — NAKASONE K. K. & SYTSMA J., 1993 — Biosystematic studies on Phlebia acerina, P. rufa

and Р. radiata in North America. Mycologia 85 : 996-1016.

34 — PETERSEN R. H. & PARMASTO E., 1993 — A redescription of Gloeostereum incarnatum.

Mycological research 97 : 1213-1216.

35 — STALPERS J. А. & BUCHANAN P K., 1991 — The studies of species of Pellicularia and

Peniophora described by G. H. Cunningham. New Zealand journal of botany 29 : 331-340.

36 — THOMSON М. & BRODA P., 1987 — Mating bahaviour in a isolate of Phanerochaete

chrysosporium. Transactions of the british mycological society 89 : 285-294.

37 — WILSON A. D., 1990 — The genetics of sexual incompatibility in the indian paint fungus,

Echinodontium tinctorium. Mycologia 82 : 332-341.

38 — WU S. H., 1990 — A study of the genus Phanerochaete (Aphyllophorales) with brown

subiculum hyphae. Mycotaxon 54 : 163-172.

39 — WU S. H., 1996 — Studies on Gloeocystidiellum sensu lato (Basidiomycotina) in Taiwan.

Mycotaxon 58 : 1-68.

Source - MNHN. Paris

Cryptogamie, Mycol. 1997, 18 (1): 19-69 19

THERMOPHILIC FUNGI: BIODIVERSITY

and TAXONOMIC STATUS.

Jean MOUCHACCA

Laboratoire de Cryptogamie, Muséum National d'Histoire Naturelle,

12, rue Buffon, 75005 Paris, France.

e-mail : mouch@mnhn. fr

RÉSUMÉ: Une évaluation critique du statut nomenclatural et, dans certains cas,

taxonomique, a été entreprise pour les champignons thermophiles décrits à ce jour. La distinction

entre éléments thermophiles et thermotolérants se base sur les définitions élaborées par Cooney &

Emerson. Au total, prés de quarante espéces et variétés s'avérent aptes à réaliser un développement

optimal à des températures élevées. Des recherches taxonomiques complémentaires sont toutefois

nécessaires pour résoudre les problèmes résiduels ; la résultante serait une légère réduction de l'effectif

du groupe.

Une croissance optimale à des niveaux thermiques situés au-delà du seuil maximal des

espéces mésophiles individualise quelques Mucorales, Eurotiales et Sphaeriales, un nombre limité

d'Hyphomycétes et un seul Agonomycéte. Aucun Coelomycéte ni aucun Basidiomycéte n'exprime

cette particularité écologique. Les Mucorales recensées sont des éléments du genre Rhizomucor et de

l'entité générique non valide Thermomucor. Rhizomucor pusillus (espéce-type) et Rh. miehei sont des

taxons valides ; Rh. pakistanicus se révéle un synonyme ultérieur de l'espéce-type. La validité

taxonomique des Rhizomucor tauricus et Rh. nainitalensis reste à confirmer.

Parmi les vingt ascomycétes répertoriés, Canariomyces thermophila et Chaetomium mesopo-

tamicum sont des taxons bien définis. Ceci n'est pas le cas des Chaetomium britannicum et

Ch. virginicum, à statut taxonomique encore imprécis ; il en est de méme pour leurs liens respectifs

avec Chaetomium thermophilum et ses deux variétés. Dactylomyces thermophilus est retenu comme

seul élément du genre Dactylomyces ; celui-ci n'est plus considéré comme congénérique de Thermoas-

cus; pour ce dernier, seule l'espéce-type et une variété sont admises. Le nouveau genre Coonemeria est

proposé pour rassembler les autres espèces des deux entités précédentes. Coonemeria crustacea

(= Dactylomyces crustacea) est sélectionné comme espéce type ; C. aegyptiaca (= Thermoascus

aegyptiacus) et C. verrucosa (= Th. crustaceus var. verrucosus et — Th. taitungiacus) y sont également

rattachés. Ces trois genres ont comme particularité commune, les caractères de leurs téleomorphes.

Cependant, Dactylomyces révéle une forme imparfaite du genre Polypaecilum, Coonemeria des

structures conidiogénes de type Paecilomyces alors que les Thermoascus ne produisent d'anamorphes

à conidies en chaines.

Corynascus heterothallicus et C. thermophilus sont des ascomycétes à périthéces clos ayant.

chacun une forme imparfaite distincte, respectivement Myceliophthora thermophila et M. fergusii ; en

raison du caractére héterothallique des téleomorphes, les anamorphes peuvent étre isolés séparément

lors des recherches portant, par exemple, sur des matériaux subissant un processus d'auto-

échauffement. Melanocarpus albomyces développe une forme conidienne arthrosporée dont est

dépourvu M. thermophila (= Thielavia minuta var. thermophila). Les trois Talaromyces thermophiles

sont associés à des anamorphes de type Paecilomyces (T. byssochlamydioides) ou Penicillium

Source : MNHN, Paris

20 J. MOUCHACCA

(Т. emersonii et T. thermophilus). Le genre Thielavia révèle également trois thermophiles

Th. terrestris, espéce-type, est un ascomycéte cosmopolite dont la forme imparfaite Acremonium

alabamense se rencontre souvent en l'absence du téleomorphe, Th. autraliensis pour lequel peu

d'informations sont disponibles et le récent Th. pingtungia dont certains caractères suggèrent son

appartenance au genre Chaetomidium.

Le groupe des hyphomycétes rassemble treize espèces. Cependant, Acremonium alabamense

et deux Myceliophthora sont des anamorphes d'ascomycétes plus ou moins hétérothalliques : ils

peuvent par conséquent se développer sans les formes parfaites correspondantes. Myceliophthora

indica se révéle un synonyme de M. thermophila. Les taxons mucédinés restants sont Acremonium

thermophilum, seul autre élément thermophile du genre, Myceliophthora hinnulea à forme parfaite

encore inconnue et Malbranchea cinnamomea, unique thermophile du genre Malbranchea dont les

mésophiles sont associés à des formes parfaites connues. Thermophymatospora fibuligera s'individua-

lise par ses cloisons mycéliennes pourvues de boucles et la présence d'une forme conidienne de type

aleuriosporée.

Les hyphomycétes dématiés relévent des genres Humicola, Scytalidium et Thermomyces

Ce dernier se révéle étre une entité générique assez. homogéne et son espéce type représente le premier

thermophile avéré. Humicola grisea var. indica et Н. lanuginosa var. catenulata sont proposés comme

Synonymes additionnels à l'espèce type. Le genre comporte également Thermomyces ibadensis,

Th. stellatus et le mésophile Th. verrucosus.

Le statut taxonomique de Humicola hyalothermophila se doit d'être reconsidérer, en méme

temps que les Humicola grisea var. thermoidea et H. insolens, récemment placés en synonymies avec

Scytalidium thermophilum, basé sur Torula thermophila, Ces deux Humicola sont retenus dans

l'immédiat dans le complexe Scyralidium thermophilum, dans l'attente d'une redéfinition des statuts

taxonomiques respectifs. Scytalidium thermophilum s'écarte du concept générique de Scytalidiun,

fondé sur Pespéce-type S. lignicola, ce qui n'est pas le cas pour S. indonesicum. Scytalidium allahaba-.

dum s'est avéré correspondre au S. thermophilum sensu lato. Humicola nigrescens var. thermorongeura

est un synonyme ultérieur de H. grisea var. thermoidea alors que H. fuscoatra var. nigra est identique

au H. insolens.

Deux taxons révèlent un statut incertain : Mucor thermo-hyalospora ( Rhizomucor pusillus ?)

et Stilbella thermophila en quête d'un genre plus approprié. D'autre part, divers taxons se sont vu

attribuer des épithétes spécifiques pouvant conduire à confusion au regard des aptitudes thermophi-

les respectives. Les cas les plus simples concernent des champignons dénommés thermophilum ou de

ses variantes et qui sont loin de. représenter des thermophiles sur la base des définitions établies.

Exemples, l'ascomycéte Achaetomium thermophilum (un synonyme ultérieur de A. macrosporum, une

espéce thermotolérante), les hyphomycétes Calcarisporiella thermophila, Gilmaniella thermophila et

Zalerion thermophylii (le statut taxonomique des deux derniers reste à considérer), l'oomycéte

Lagenidum thermophilum, le zygomycéte Mucor thermophilus et, enfin, la levure Endoblastomyces

thermophilus, à statut taxonomique non valide. Melanomphalia thermophila est un basidiomycète

simplement observé dans une localité chaude et humide; des cas similaires ont été également

recensés.

Le binóme Sporotrichum cellulophilum est un exemple d'une source différente de confusion.

Ce taxon ne posséde de statut taxonomique d'aucune sorte. Il est cependant fréquemment signalé

comme thermophile dans la litérature portant sur les études enzymiques des champignons.

Ces binómes fantómes sont relativment fréquents dans ce type de publications. Cette pratique

préjudiciable devrait être définitivement abandoné

ABSTRACT : A critical reappraisal of nomenclatural status and in some cases also of taxonomic

Ones was undertaken for known thermophilic fungi. Distinction between thermophilic and thermo-

tolerants follows definitions elaborated by Cooney & Emerson. Altogether less than forty species and

varieties are able to achieve best development at high temperatures. Further taxonomic work is

however needed to solve residual problems, The outcome will be a slight reduction of the group.

Optimum growth at temperatures above the maximum threshold of mesophiles characterise

few Mucorales, Eurotiales and Sphaeriales, a limited number of Hyphomycetes plus one Agonomy-

Source : MNHN. Paris

THERMOPHILIC FUNGI 21

cete species. No Coelomycete and no Basidiomycete develop such ecological feature. Recorded

Mucorales are species of Rhizomucor and the invalid monospecific genus Thermomucor. Rhizomucor

pusillus (type species) and Rh. miehei are valid taxa; Rh. pakistanicus proved to be a later name for the

former. The validity of Rhizomucor tauricus and Rh. nainitalensis awaits confirmation.

Among the twenty ascomycetes, Canariomyces thermophila and Chaetomium mesopotami-

cum are well defined taxa. But the status of Chaetomium britannicum and Ch virginicum is still

unclear; also relations with Ch thermophilum and its two varieties awaits clarification. Dactylomyces

thermophilus is retained as the sole species of the genus, no longer regarded as congeneric with

Thermoascus; for the latter, only the type and one variety are accepted. A new genus Coonemeria is

proposed to accomodate remaining taxa of both genera. Coonemeria crustacea (= Dactylomyces

crustaceus) is selected as type species; С. aegyptiaca (= Thermoascus aegyptiacus) and C. verrucosa

(E Th. crustaceus var. verrucosus and = Th. taitungiacus) are two other members of this genus. These

three genera have in common the characters of their teleomorphs. But Dactylomyces has a Polypae-

cilum type anamorph, Coonemeria develop conidiogenous structures of the Paecilomyces type while

Thermoascus has no anamorph producing chains of conidia.

Corynascus heterothallicus and C. thermophilus are cleistothecial ascomycetes having a well

defined anamorphic state, namely Myceliophthora thermophila and M. fergusii; as the teleomorphs

are heterothallic, the anamorphs could be observed alone in studies involving, for instance, self

heated materials. Melanocarpus albomyces has a well defined arthroconidial state not developed by

M. thermophilus (= Thielavia minuta var. thermophila). The three thermophilic Talaromyces have

conidial states belonging either to Paecilomyces (Т. byssochlamydioides) or to Penicillium (T. emersonii

and T. thermophilus). Thielavia (also three species) is represented by the widely distributed

Th. terrestris whose anamorph Acremonium alabamense could equally be isolated separately, the

undocumented Th. australiensis and the recent Th. pingtungia with features favouring his relocation in

Chaetomidium.

Recorded hyphomycetes comprise thirteen taxa. But Acremonium alabamense and two

Myceliophthora ате anamorphs of almost heterothallic ascomycetes; these could thus develop at high

temperatures without producing respective ascocarps. Myceliophthora indica is considered a synonym

of M. thermophila. Other mucedinaceous taxa are Acremonium thermophilum, the second thermo-

phile of the genus, Myceliophthora hinnulea perfect state is yet unknown and Malbranchea cinnamo-

mea, sole thermophile of a genus whose mesophilic components are associated with perfect states.

Thermophymatospora fibuligera is unique by its hyphae disclosing septal clamp connections and the

formation of an aleuriosporic state,

Dematiaceous thermophiles are members of Humicola, Seytalidium and Thermomyces. The

latter is regarded as a homogeneous genus with the type species Thermomyces lanuginosus being the

first established thermophile; Humicola grisea var. indica and H. lanuginosa var. catenulata are

additionnal later names for the type species. Other known members are Thermomyces ibadensis,

Th. stellatus and the mesophilic Th. verrucosus.

The taxonomic status of Humicola hyalothermophila awaits to be reconsidered together

with Humicola grisea var. thermoidea and Н. insolens, recently placed in synonymy with Scytalidium

thermophilum based on Torula thermophila; both Humicola are placed here for the moment as

synonyms of the “complex S. rhermophilum" pending re-assessment. Scytalidium thermophilum

deviates from the current concept of Scytalidium (based on S. lignicola) but such is not the case for

S. indonesicum. Scytalidium allahabadum proved to match S. thermophilum sensu latu. Humicola

nigrescens var. thermorongeura duplicates the description of Humicola grisea var. thermoidea while

H. fuscoatra var. nigra is identical to H. insolens.

Two taxa have an uncertain position: Mucor thermo-hyalospora (Rhizomucor pusillus?) and

Stilbella thermophila requiring a more appropriate genus. Also several taxa disclose confusing specific

epithets with regard to thermophilic abilities. Simple cases refer to fungi with epithets as thermophi-

Jum or variants of and that are not thermophilic based on accepted definitions. Examples, the

ascomycete Achaetomium thermophilum (a later synonym of A. macrosporum, a thermotolerant), the

hyphomycetes Calcarisporiella thermophila, Gilmaniella thermophila and Zalerion thermophylii (the

latter two have yet unsettled status), the oomycete Lagenidium thermophilum, the zygomycete Mucor

Source : MNHN. Paris

22 J. MOUCHACCA

thermophilus and finally, the invalid yeast Endoblastomyces thermophilus. Melanomphalia thermophila

is a basidiomycete simply observed in a warm humid locality; similar cases could also be traced.

The binomial Sporotrichum cellulophilum is an example of a different confusing situation.

The taxon has no taxonomic status of any type although being infrequently reported as a thermophile

in literature related to fungal enzymic studies. Such ghost binomials are not uncommon in these

publications. This practice being a source of serious confusion should be definitely prohibited.

INTRODUCTION

Temperature is one of the extremely important environmental variables that play

a decisive role in the survival, growth, distribution and diversity of microorganisms on the

surface of theearth. The response of fungi to temperature varies between the two extremes

of obligatorily thermophilic through thermotolerance to psychrophilic species. However,

by far the majority of known fungi are mesophiles developing in culture between 5 and

37° C; the psychrophiles extend below that range of temperatures (Dix & Webster, 1995).

Thermophily has been defined variously with reference to different groups of

microorganisms and sometimes also within the same group. The response of fungi to high

temperatures has been the subject of classificatory schemes successively proposed by

Apinis (1963), Cooney $: Emerson (1964), Craveri el al. (1964), Evans (1971) and Crisan

(1973). These schemes are either based on values of minimum and maximum growth

temperatures alone or, in addition, integrate the criteria of optimum development.

The commonly accepted definitions of thermophilic and thermotolerant fungi

are those of Cooney & Emerson (1964). Thermophilic fungi are those that have a growth

temperature maximum at 50° C or above and a temperature minimum of 20? C or higher.

Thermotolerant species are those that have a growth temperature maximum of about

50* C and a temperature minimum well below 20? C. This simple segregative scheme is

sometimes difficult to apply since the response of thermophilic taxa at the minimum

temperature threshold tends to vary among respective strains.

Serious consideration of fungi able to develop only at high temperatures dates

back to 1899 when P. Tsiklinsky first reported on a thermophilic hyphomycete incidently

encountered on a potato inoculated with garden soil. The fungus was then grown on bread

kept at 52-539 C and its thermophilic nature assessed. Tsiklinsky named this hyphomycete

Thermomyces lanuginosus. Rapidly, however, this. thermophile was successively relocated

in other genera as Acremoniella, Humicola, Monotospora and Sepedonium before its

definite reinstallment in Thermomyces. Similar changes also characterize thermophilic

moulds described in the early decades of the nineties. The outcome of such changes is the

chaotic nomenclatural state of few members of this group in published literature. Absence

of homogeneity in binomial citations develop cases of taxonomic confusion coupled with

divergence in species concept (Cooney & Emerson, 1964). The final result is a partial or

total incomplete identification of encountered taxa or names reported being shadowed.

Although Lindt description of Rhizomucor pusillus (as Mucor pusillus) dates

back to 1886, there is a general agreement that Tsiklinsky (1899) is the first to drew

attention to thermophilism among fungi. Very rapidly, Miehe's (1905) serious investiga-

tion of self heating hay produced the first extensive report on thermophily in fungi. This

author isolated and studied a range of thermophiles including Thermoasus aurantiacus

and Malbranchea cinnamomea (Miehe, 1907). Griffon & Maublanc (1911) then introduced

the first thermophilic Penicillium, P. dupontii, now Talaromyces thermophilus. It is only

several decades later that LaTouche (1950) reported on the new Chaetomium thermophile.

Such discovery generated much interest to this group of fungi, substantiated by the

cellulolytic nature of the new ascomycete.

Source : MNHN, Paris

THERMOPHILIC FUNGI 23

Several pioneer publications then followed on thermophilic fungi inhabiting

soils of temperate regions (Apinis, 1963; Eggins and Malik, 1969), tropical areas (Hedger

1974; Gochenaur, 1975) and on soils of arid regions (see review in Mouchacca, 1995).

Thermophilic fungi of habitats rich in organic materials were also extensively surveyed

and data from relevant publications critically reviewed by Tansey & Brock (1978). Reports

on less widespread habitats and habitats deserving future investigations were also consi-

dered by Tansey & Brock (1978).

The first modern comprehensive account on the taxonomy, biology and econo-

mic importance of thermophilic fungi was published by Cooney & Emerson (1964).

Eleven thermophiles were reported. Since then the number of taxa developing at high

temperatures is expanding rapidly. In 1973, Crisan provided a list of 55 names of

thermophilous fungi, i.e. thermophilic and thermotolerant ones; however, only half are

thermophiles in the sense of Cooney & Emerson. Crisan reviewed in addition current

concepts about thermophilism in microorganisms; he then underlined that our knowledge

about the physiological ability of fungi to grow at elevated temperatures was much limited.

Later Samson & Tansey (1977) prepared a guide to species able to grow and sporulate at

45° C; this list concerns eight mucorales, around twenty taxa each of ascomycetes and

hyphomycetes and two basidiomycetes. The subsequent list prepared by Tansey and Brock

(1978) reports 67 species or varieties growing at 50° C or above; a good proportion of these

taxa was however not specified at the species level. A Russian compilation of descriptions

and published illustrations of thermophilic fungi was prepared by Bilai & Zakharchenko

(1987); 38 species were considered but few are not strict thermophiles. Finally, according

to Abdullah & Al-Bader (1990), around 70 species detected in various substrates are now

reported to be thermophilic or thermotolerant.

Cooney & Emerson (1964) monograph introduced new thermophilic taxa.

However, some taxonomic decisions they adopted rapidly proved to be misleading and

their descriptions of novel taxa supported not critical analysis. These limitations triggered

subsequent studies. Several interesting notes thus appeared in the sixties clarifying pen-

ding problems while expanding the group. Apinis & Chester (1964) described Dactylomy-

ces crustaceaus. Pugh et al. (1964) reintroduced Thermomyces. Stolk (1965) assessed the

taxonomic status of Penicillium dupontii and Thermoascus aurantiacus. Again Apinis

(1967) clarified generic concepts of Dactylomyces and Thermoascus. However, inspite of

the above and later contributions, not all standing problems received attention. Recently,

Straatsma and Samson (1993) focused on both Humicola proposed by Cooney & Emerson

(1964).

The material of this contribution was collected while preparing a lecture for the

Microbial Diversity and Ecosystem Function workshop held at Egham, UK, in 1993.

The lecture focused on thermophilic fungi of desert soils, an example of a neglected

extreme environment (Mouchacca, 1995). A second lecture on the subject was later

presented at IMC V; this was entitled “Thermophilic and thermotolerant fungi in the

Middle East: Biodiversity and Taxonomic Reappraisal" (Mouchacca, 1994); however, the

corresponding note suffered some publication delay. In the sametime, the first draft of the

present paper was due to be part of a book in the pipeline somewhere in the Indian

subcontinent. Decision was then taken to update and publish the applied last version.

The present document aims to provide a sound reappraisal of the nomenclatural

and in some cases of the taxonomic status of known thermophiles. First to overcome a

major difficulty encountered while interpreting published data on this ecological group.

Second to enhance future taxonomic work on its components and, finally, to stress the

attention on taxa other than those commonly studied for eventual industrial applications.

Source : MNHN, Paris

24 J. MOUCHACCA

THERMOPHILIC MUCORALES

— Rhizomucor pusillus (Lindt) Schipper — Studies in Mycology 17: 54.

1978.

basionym: Mucor pusillus Lindt — Archiv für experimentelle Pathologie und Pharmako-

logie 21: 272. 1886.

= (?) Mucor septatus Bezold in Siebenmann — Die Schimmelmykosen den menschlichen

Ohres: 97. 1889.

= Rhizomucor septatus (Bezold) Lucet & Cost. — Archives de Parasitologie 4: 362. 1901.

= Mucor (sect. Rhizomucor) parasiticus Lucet & Cost. — Comptes rendus hebdomadaires

des Séances de l'Académie des Sciences, Paris 129: 1033. 1899. ;

= Rhizopus parasiticus (Lucet & Cost.) Lendner — Matériaux pour la Flore Cryptogami-

que Suisse 3: 115. 1908.

= (?) Mucor muriperda Sacc. & Sinigaglia in Sacc. — Sydow, Annales Mycologici, Ser. II,

11: 321. 1913.

= Tieghemella muriperda (басс. € Sinigaglia) Naumov — Opredelitel’ Mukorovykh

(Mucorales): 84. 1935.

- Mucor buntingii Lendner — Bulletin de la Société botanique de Genéve 21: 260. 1930.

= Mucor hagemii Naumov — Opredelite? Mukorovykh (Mucorales): 55. 1935 (nom.

inval., Art. 36.1).

= Rhizomucor pakistanicus Qureshi & Mirza — In Mirza, Khan Begum & Shagufta

‘Mucorales of Pakistan (Faisalabad): 100. 1979 (nom. inval., Art. 37.1); Qureshi & Mirza

— Biologia, Lahore 29: 343. 1983, a superfluous publication.

Descriptions: Cooney & Emerson (1964); Schipper (1978); Domsch et al. (1980).

This is the type species of Rhizomucor (Lucet & Cost.) Wehmer: Vuill. The genus

was reintroduced by Schipper (1978) to seggregate three hitherto known thermophilic

species of Mucor distinguished by the presence of rhizoids at the base of their sporangio-

phores.

The early history of Rhizomucor pusillus (as Mucor pusillus) and its long confu-

sion with the Mucor species studied by Miehe (Miehe, 1907; now Rhizomucor miehei) was

reviewed by Cooney & Emerson (1964). The former had indeed often been misidentifed

with the equally thermophilic, thinly growing and equally common Rhizomucor miehei.

This zygomycete is however regularly homothallic while in Rhizomucor pusillus homothal-

lic isolates have only exceptionally been found. An excellent account of the morphology.

physiological characteristics and distribution is provided by Domsch et al. (1980).

Rhizomucor pakistanicus was isolated from several sources in Pakistan: ground-

nut seeds and lizard droppings collected at the city of Faisalabad, from a potato field at

Sialkot and from soil at Rawalpindi (Mirza et al., 1979). It was then correctly assigned to

Rhizomucor (indicated as Rhizomucor Lucet & Cost.) on account of the presence. of

rhizoids. However no comparison was undertaken with the indicated type species, Rhizo-

mucor parasiticus Lucet & Cost. Further, in the second superfluous publication made by

the same authors (simply a duplicate of the former), taxonomic decisions concerning this

genus introduced by Schipper (1979) were overlooked. For Rhizomucor pakistanicus, à

number of morphological features forwarded in its description leads to consider the

fungus as a synonym of Rh. pusillus.

THERMOPHILIC FUNGI 25

The current concept of Rhizomucor was however recently expanded to make,

provision for mesophilic isolates also producing rhizoids from the base of their sporan-

glophores, Rhizomucor variabilis Zheng & С.-д Chen var. variabilis was obtained from a

cutaneous mucormycosis of a human hand in China (Zheng & Chen, 1991). Rhizomucor

variabilis var. regularior Zheng & G.-q. Chen represent another agent of cutaneous human

disease (Zheng & Chen, 1993). Optimum, maximum and minimum growth temperatures

of the two varieties are the same, i. e. 24-30? C, 37? C and 9? C respectively. The addition-

nal mesophilic Rhizomucor endophyticus Zheng & H. Jiang (1995) was isolated from wheat

collected in China; its optimum, maximum and minimum growth temperatures are even

lower, being 18-28? C, 36? C and 5? C respectively.

— Rhizomucor miehei (Cooney & Emerson) Schipper — Studies in

Mycology 17: 58. 1978.

basionym: Mucor miehei Cooney & Emerson — Thermophilic Fungi: 26. 1964.

Descriptions: Cooney & Emerson (1964); Schipper (1978).

As stressed before, isolates of this zygomycete regularly produces zygospores in

Cultures. This finding led Cooney & Emerson (1964) to propose a specific rank for this

taxon previously considered as identical to the almost morphologically similar type

Species. Schipper (1978) stressed however that the general morphology could also be used

, to distinguish both taxa. Rhizomucor miehei exhibits a looser sympodial branching

pattern with relatively longer side branches while Rh. pusillus produces small bunches of

Short subterminal branches on the main sporangiophores. In addition the sporangia and

Columellae of the latter are usually larger.

Rhizomucor miehei displays a wide geographic distribution (Domsch ef al.,

1980). Factors affecting development of zygospores were investigated by Lasure & Ingle

(1976) and those regulating germination of sporangiospores by Deploey (1992).

— Rhizomucor nainitalensis Joshi — Sydowia 35: 100. 1982.

d This still Indian based homothallic zygomycete was isolated form a heavily

“composed oak log in the forest of Pungote, Nainital. It differs from Rhizomucor miehei,

- Pusillus and Rh. tauricus mainly by sporangiospores of varying shapes and sizes:

Subglobose, ellipsoidal, oblong, reniform, dump-bell shaped, etc.. , 3-6 um or more wide.

ariation in sporangiospore shape might however be an artefact. | etd

; According to Joshi (1982), growth is very rapid at 48° C filling half of a Petri dish

in 2 d. At 38° C, “the growth of the mycelium takes place after three days but about one

әлек is required to colonize the culture medium in a petri dish at 25° C". Rhizomucor

ainitalensis appears very close to Rh. miehei.

M Rhizomucor tauricus (Milko & Schkurenko) Schipper — Studies in

Ycology 17: 62. 1978.

озуп Mucor tauricus Milko & Schkurenko — Novosti sistematiki nizshikh rastenii 7:

- 1970. ;

Description: Schippers (1978).

The fungus is apparently still known only from the original strain isolated from

forest Soil in the Ukraine. It was maintained as a separate species by Schipper (1978)

26 J. MOUCHACCA

pending further informations. Rhizomucor tauricus is distinguished from other thermo-

philic Rhizomucor mainly by definitely more swollen sporangiophores. Growth and spo-

rulation occurs between 24-55? C; development is extremely slow at 21° C and nil at 57? С.

— Thermomucor indicae-seudaticae Subrahmanyam, Mehrotra & Thiru-

malachar (as *Subrahamanyam....") — Georgia Journal of Botany 35: 2.

1977. (nom. inval., Art. 37.1).

Descriptions: Subrahmanyam, Mehrotra & Thirumalachar (1977); Schipper (1979).

This is the type species of Thermomucor Subrahmanyam et al. (1977) which is

apparently still monospecific. It was established mainly on account of formation by the

type species of smooth zygospores and presence of rhizoids. Zygospores are definitely

rough-walled in members of Absidia van Tieghem, Mucor Mich.:Fr., Rhizopus Ehrenb.:

Corda and Rhizomucor.

The described strain was isolated from municipal compost at Pimpri, Poona,

India. Prior to its description, this zygomycete was reported as Rhizopus sp. and as such

isolated from various habitats in Europe, India, Ghana and Nigeria (Schipper, 1979).

THERMOPHILIC ASCOMYCETES

— Canariomyces thermophila Guarro & Samson in von Arx, Figueras &

Guarro — Beihefte zur Nova Hedwigia 94: 34. 1988.

Canariomyces von Arx (von Arx, 1984) was established for a mesophilic cleisto-

thecial ascomycete, C. notabilis von Arx, having ascoma wall made up of angular dark

cells, irregularly disposed asci, aseptate ascospores dextrinoid when young and provided

with a single germ pore; an anamorph having conidia of the form genus Chrysonilia von

Arx (catenate aseptate or septate hyaline conidia) and of Trichosporiella Kamyschko

(simple lateral conidia) is produced.

Canariomyces thermophila was isolated from Cameroon, Africa, apparently

from soil. The original protologue is based on colonies developing at 45? C but no data

about minimum and maximum growth temperatures are provided. Ascospores are gree-

nish brown when mature with a subapical germ pore, 14.0-18.0 x 7.5-10.0 um. However

no anamorph developed in cultures of the single available strain.

— Chaetomium britannicum Ames — A Monograph of the Chaetomiaceae:

16. 1963.

Descriptions: Ames (1963); von Arx et al. (1986).

This Chaetomium was described as having ovoid to vase-shaped ascomata.

Terminal and lateral hairs are very slender, greyish, straight to undulate. Asci club-shaped,

8-spored. Ascospores brown, large, 19-24 x 11-14 um, irregularly oval, rounded on the

ends. In the protologue, it is simply indicated that “perithecia develop when incubated at

approximately 47? C. A thermophile" (Ames, 1963). The specific epithet refers to the type

locality: southern part of England.

Source : MNHN, Paris

THERMOPHILIC FUNGI 27

The original material developed on mushroom compost and apparently no

living culture was realised. von Arx er al. (1986) regard this Chaetomium as a doubtful

species; only ascospores could be observed in the type specimen at BPI. Millner (1977)

attempted but without success to obtain a culture from the dried type material; as no living

strain, authentic or representative, was available to him, no growth temperature rela-

tionships could thus be established.

Gochenaur (1975) reported having isolated this Chaetomium from soil in the

Bahamas but Millner ег al. (1977) stressed the absence of a corresponding culture or

herbarium specimen. Further and based on informations communicated by Gochenaur,

the micromorphology of the fungus she examined was probably not Chaetomium britan-

nicum. Spores of Gochenaur's taxon measured 13 (-18) x 7 (-10) um and had subapical

germ pores while spores from Ames material measure 19.0-24.8 x 11-14 um and have

single apical germ pores (Millner et al., 1977). Chaetomium britannicum remains undocu-

mented in the sense of Cooney and Emerson. Also no additional record either from the

type locality (Cannon et al., 1985) or elsewhere has apparently been published.

— Chaetomium mesopotamicum Abdullah & Zora — Cryptogamic Botany

3: 387. 1993.

The original locality is a date palm plantation in Basrah, Iraq. This recently

described species has a growth temperature range from 30-52? C. It differs from Chaeto-

mium thermophilum LaTouche and Ch. virginicum Ames by its clavate asci and long highly

branched terminal hairs. Ascospores are globose to ovoid, olive to brown, 5.5-7.8 x

5.2-6.3 um, provided with one apical germ pore.

— Chaetomium thermophilum LaTouche as “ thermophile" — Transac-

tions of the British mycological Society 33: 94. 1950; Cooney & Emerson —

Thermophilic Fungi: 62. 1964.

= ? Chaetomium virginicum Ames — A Monograph of the Chaetomiaceae: 43. 1963; fide

von Arx et al., 1986.

Descriptions: LaTouche (1950); Ames (1963); Cooney & Emerson (1964); von Arx et al.

(1986).

This is the first known thermophilic Chaetomium. The species is also distin-

guished by its long, tapering terminal hairs at times dichotomously branched at wide

angles. Ascospores are dark brown, globose to subglobose, 7-9 x 5-7 um, prominently

umbonate at one end.

Cooney & Emerson (1964) observed this Chaetomium to produce in culture two

dissimilar growth patterns; as a result they proposed two new varieties: coprophile and

dissitum.

— Chaetomium thermophilum LaTouche var. coprophile Cooney & Emer-

son — Thermophilic Fungi: 69. 1964.

А The variety is mainly distinguished by the presence of dichotomously branched

hairs, which more or less completely covers the entire perithecium.

Source : MNHN, Paris

28 J. MOUCHACCA

— Chaetomium thermophilum La Touche var. dissitum Cooney & Emerson

— Thermophilic Fungi: 68. 1964.

Differs from the species mainly in the diffuse manner in which perithecia are

produced in culture.

— Chaetomium virginicum Ames — A Monograph of the Chaetomiaceae:

43. 1963.

The fungus was originally isolated from leaf litter collected under very old trees

at White Marsh, North of Old Point Comfort, Virginia (USA). Perithecia are described as

globose, up to 240 um wide. Terminal and lateral hairs cover the entire perithecium,

indistinguishable, irregularly branched, minutely granular, 2-4 ит wide. Asci cylindrical,

8-spored, 70 x 10 um. Ascospores yellow brown to pale brown, almond-shaped,

8-11.5 um. A thermophile (Ames, 1963).

Cooney & Emerson (1964) consider this species to approximate Chaetomium

thermophilum while being identical to its variety coprophile; ascomata of the latter are also

entirely covered by dichotomously branched perithecial hairs. On the basis of morpholo-

gical characters and growth-temperature responses, Millner (1977) provided evidences

that Chaetomium virginicum (culture TA-7 obtained from L. M. Ames collection at BPI) is

identical to Ch. thermophilum var. coprophile. This proposal was later substantiated by

Millner et al. (1977); among the large number of Chaetomia studied by these authors,

ascospores with papillate germ pores were found only in Chaetomium thermophilum, its

two varieties and Ch. viginicum.

Chaetomium virignicum is also regarded as a probable synonym of Ch. thermo-

philum by von Arx et al. (1986).

— Coonemeria Mouchacca gen. nov.

Thermophila. Coloniae lanatae, aurantiae-rubrae quando ascoma matura sunt.

Initium ascomatis est convoluta hypha. Ascoma non-ostiolata, sphaerica, solitaria vel

confluenta; ascomata confluentes in crusta disposita. Ascomatis paries crassus, e cellulis

pseudoparenchymaticis compositus, ascomatis textura angulosa. Ascus ex hamo singulato

oriundus, ovoideus vel piriformis, octosporus, deinde evanescens. Ascospora unicellularia,

ellipsoidea vel ovoidea, flavescens vel pallida aurantio-brunnea, cum pariete crassa, laeve vel

verrucosa. Structura conidiogena semper praesens, similis Paecilomyces Bainier forma

genericum.

Species typica: Coonemeria crustacea (Apinis & Chesters) Mouchacca.

Thermophilic. Colonies lanose, reddish orange due to mature ascomata. Asco-

matal initials a coiled hyphae. Ascomata non-ostiolate, spherical, solitary to confluent,

forming a crusty layer; ascomatal wall thick, made up of pseudparenchymatous cells,

textura angularis. Asci arise singly from croziers, ovoid to pyriform, 8-spored, evanescent.

Ascospores unicellular, ellipsoid to ovoid, yellowish to pale reddish-brown, thick-walled,

smooth to verrucose. Conidiogenous structures always present, belong to form genus

Paecilomyces Bainier.

Type species: Coonemeria crustacea (Apinis & Chesters) Mouchacca.

Etymology: genus name coined from the first four respective letters of Cooney &

Emerson's names, authors of the first comprehensive (although somewhat confusing)

monograph on thermophilic fungi.

Source : MNHN, Paris

THERMOPHILIC FUNGI 29

Coonemeria is proposed to accomodate thermophilic cleistothecial ascomycetes

having coiled ascogonial initials, pseudparenchymatous walls of textura angularis type

and a distinctive Paecilomyces anamorph. Asexual reproductive structures generally

represent comparatively reduced forms of well elaborated Paecilomyces conidial structu-

res developed in several taxa of this form genus (Samson, 1974).

The three species accepted in Coonemeria were formerly assigned to Thermoas-

cus Miehe (Miehe, 1907) and Dactylomyces Sopp (Sopp, 1912). These genera have

morphologically similar perfect states. Thermoascus is distinguished by the absence of any

accessory state producing chains of conidia while this state belong to Polypaecilum

G. Smith in the case of Dactylomyces (Apinis, 1967). The proposal of Coonemeria

definitely clarifies the status of ascomycetes formerly placed in one or the other of these

entities.

Anamorphs of the Paecilomyces type are also associated with Talaromyces

Benjamin and Byssochlamys Westling (Stolk & Samson, 1972). The former is distin-

guished by soft white to yellow ascomata having walls composed of loose hyphae and the

production of asci in chains. Most Talaromyces however are associated with Penicillia

while only two have a Paecilomyces state; the latter were placed by Stolk & Samson (1972)

in Talaromyces section Emersonii: the thermotolerant 7. leycettanus Evans & Stolk and the

thermophilic T. byssochlamydioides Stolk & Samson, here considered. The two other taxa

of the section have РепісіШа: the thermophilic T. emersonii (treated in this note) and the

thermotolerant 7: bacillosporus Benjamin.

On the other hand, all Byssochlamys have a conidial Paecilomyces state. This

teleomorphic genus is characterized by initials consisting of swollen antheridia and coiled

gonia producing almost naked ascomata in which globose asci are formed from

croziers (Stolk & Samson, 1972).

— Coonemeria crustacea (Apinis & Chesters) comb. nov.

basionym: Dactylomyces crustaceus Apinis & Chesters — Transactions of the British

mycological Society 47: 428. 1964.

= Thermoascus crustaceus (Apinis & Chesters) Stolk — Antonie van Leeuwenhoek 31:

272. 1965.

anamorph: Paecilomyces crustaceus Apinis & Chesters — Transactions of the British

mycological Society 47: 428. 1964.

Misapplied names: Thermoascus aurantiacus Miehe — Die Selbsterhitzung des Heues: 70.

1907; sensu Cooney & Emerson — Thermophilic Fungi: 39. 1964.

= Dactylomyces thermophilus Sopp — Skrifter udgivne af videnskabs-selskabet i Chris-

Чапа. Mathematisk-naturvidenskabelig klasse 11: 35. 1912; sensu Cooney & Emerson -

Thermophilic Fungi: 39. 1964.

= Penicillium thermophilus (Sopp) Biourge — La Cellule 33: 106. 1923; sensu Cooney &

Emerson — Thermophilic Fungi: 39. 1964.

= Penicillium thermophilum (Sopp) Sacc., fide Trotter 1931 — Sylloge Fungorum 25

(Suppl. 10): 671.1931; sensu Cooney & Emerson — Thermophilic Fungi: 39. 1964.

Descriptions: as Dactylomyces crustaceus by Apinis & Chesters (1964) and Apinis (1967);

as Thermoascus aurantiacus by Cooney & Emerson (1964); as Thermoascus crustaceus by

Stolk (1965), Awao & Otsuka (1974) and Chen & Chen (1996).

Source : MNHN, Paris

30 J MOUCHACCA

In 1964, Apinis & Chesters reported on an ascomycete isolated at 38? C from

grass debris collected in a salt-marsh on the Lincolnshire coast. Dactylomyces Sopp was

then thought to be the correct genus. The fungus also proved to compare with the CBS

strain 374.62 (- QM 6798 = NRRL 1563) deposited by Raper & Fennell as Dactylomyces

thermophilus, the genus type species. At that time, Sopp's fungus was only known from the

original description and illustrations (Sopp, 1912).

Apinis & Chesters (1964) compared these two Dactylomyces strains with the

protologue of the type species and noted several discrepancies. Both isolates were found to

deviate by the absence of dactyloid conidiophores bearing small conidia and the presence

of relatively small asperulate ascospores. For Dactylomyces thermophilus, Sopp had

reported ascospores as oval, smooth, 10-12 x 6-7 um. Ascospores of the living strains

were reddish-brown, globose to oval, rough and smaller, 7-9 x 5-7 um. Further, their

respective anamorphs were found to perfectly fit in Paecilomyces with conidia being

hyaline, cylindrical to oval, smooth, 3-8 x 2-4 um. Based on these deviations, Apinis &

Chesters proposed the new Dactylomyces crustaceus and selected as type material their

strain BDUN 378 (= IMI 102470).

Also in 1964, Cooney & Emerson published their monograph on thermophilic

fungi. They provided a latin diagnosis for the type species of Thermoascus, Th. aurantia-

cus, based on their strain M 206516. They also regarded Dactylomyces as congeneric with

the previously described Thermoascus and thus listed known synonyms of both states of

D. thermophilus under Thermoascus aurantiacus (see also under Dactylomyces thermophi-

lus). Before this monograph, the anamorph of Dactylomyces thermophilus was regarded as

approximating a Penicillium (Biourge, 1923).

A year later, Stolk (1965) re-examined strain CBS 374.62 and concurred with

Apinis & Chesters (1964) about its similarity with their Dactylomyces crustaceus. Stolk

admitted however Cooney & Emerson's (1964) disposition for Dactylomyces and accor-

dingly proposed the new combination: Thermoascus crustaceus (Apinis & Chesters) Stolk.

Stolk finally compared the above strain with four other CBS cultures labelled Thermoascus

aurantiacus: CBS 256.34, 257.34, 415.62 & 398.64. No conidial state matching the ana-

morph Cooney & Emerson (1964) depicted for this ascomycete developed in any of these

cultures. Stolk then concluded “Cooney & Emerson’s strain M206516 of Thermoascus

aurantiacus is most likely identical with CBS 374.62”, a suggestion being subtantiated by

the fact that the relevant iconography is suggestive of Paecilomyces.

Finally, in 1967, Apinis re-examined strain M 206516. It proved conspecific with

Dactylomyces crustaceus. Thus he, first, confirmed Stolk's suggestion about this strain

and, second, the description provided by Cooney & Emerson (1964) for Thermoascus

aurantiacus applies in fact to the former. In addition, the study of fresh isolates matching

the protologue of Dactylomyces thermophilus, lead Apinis (1967) to conclude Dactylomy-

ces should be maintained distinct from Thermoascus, a decision largely accepted subse-

quently (Cannon et al., 1985; Eriksson & Hawksworth, 1993). As underlined before, the

development of a distinctive Paecilomyces anamorph favours the placement of Dactylo-

myces crustaceus in Coonemeria.

On common laboratory agar media, the minimum growth temperature lies

between 20-25? C with the maximum being below 60° C. Optimum growth is around

40? C with a standard Petri-dish being covered in 4 d; mature colonies are colored orange

with orange brown reverse.

Initials a simple ascogonial coil. Cleistothecia scattered or confluent and then

forming a crusty layer, spherical, 300-700 um diam., orange to reddish-brown. Ascomatal

wall made of few layers of brown pseudparenchyma cells with slightly thickened walls.

Source : MNHN, Paris

THERMOPHILIC FUNGI 31

Asci are produced singly from croziers, irregularly disposed in the ascomatal cavity,

8-spored, subglobose to pyriform, 15-20 x 13-16 um, evanescent. Ascospores oval, pale

brown to red-brown, 6.0-8.0 x 5.0-6.5 um, wall 0.5 um thick, provided with fine echinu-

lations.

Asexual reproductive structures develop within 2 d at 40? C; they are evanescent

and not affecting overall colony characters. Conidiophores coarse, septate, pale yellow,

smooth, up to 900 um long, tapering to 4-5 um wide apical parts. Upper parts of the

conidiophores bears irregularly arranged branches, 6-35 x 4-5 um; these are usually

rebranched and end with phialides; phialides occur either singly as side branches, or in

irregular verticils of 2-3; phialides cylindric, 12-30 x 5-7 um, gradually tapering to a long

conidium-bearing tube, slightly bent away, 12 x 3 um. Conidia produced in conspicuous

long diverging chains, smooth, yellow to pale brown, cylindrical when young, ellipsoid

when mature, 6-10 x 3-6 um, responsible for the slight ochraceous color of young colo-

nies.

Coonemeria crustacea is distinguished from the two other members of the genus

mainly by oval finely echinulated ascospores. It displays a wide geographic distribution

being isolated from soil in several local and from various self-heating material (Coo-

ney & Emerson, 1964; Cannon et al., 1985; Chen & Chen, 1996).

— Coonemeria aegyptiaca (Ueda & Udagawa) comb. nov.

basionym: Thermoascus aegyptiacus Ueda & Udagawa — Transactions of the Mycologi-

cal Society of Japan 24: 135. 1983.

anamorph: Paecilomyces aegyptiaca Ueda & Udagawa — Transactions of the Mycologi-

cal Society of Japan 24: 135. 1983.

The fungus was originally isolated from a sample of marine sludge collected

along the Suez Canal banks at Port-Said City, Egypt. It develops between 25 to 55% C with

the optimum being at 40? C. At this temperature, colonies fill the plate within 4 d with a

thin almost submerged basal mycelium producing numerous superficial ascocarps, often

forming a crusty mass, vinaceous to reddish brown; conidia fairly abundant, grayish

yellow and not affecting colony color.

Cleistothecia superficial, subglobose, orange-brown, 250-550 um wide; initials a

simple coiled hyphae. Peridium 25-40 um thick, pseudoparenchymateous, rather coria-

ceous, textura angularis type. Asci borne singly on croziers, scattered in the ascomatal

cavity, 8-spored, ovate, 14-18 x 11-15 um, evanescent. Ascospores 1-celled, ellipsoid to

ovoid, yellowish to pale reddish orange, 6.0-8.5 x 4.0-5.5 um, thick-walled and nearly

smooth (slightly verruculose under SEM).

Conidiophores erect arising more commonly from aerial trailing hypae, hyaline,

smooth-walled, 50-300 x 5-7 um; apical parts irregularly branched and bearing terminal

verticils of 2-4 phialides usually without any metulae; phialides solitary or irregularly

verticillate, cylindric, 12-30 x 3-6 um. Conidia formed in long divergent or tangled chains,

continuous, hyaline but fulvous in mass, cylindrical to elliptical, 4.5-11 x 3-4 um; conida

sometimes ovoid to subglobose, 3.5-10 diam.

Coonemeria aegyptiaca is mainly distinguished by its ellipsoidal almost smooth

ascospores. Ueda & Udagawa (1983) indicate the fungus produces two morphological

kinds of asexual strctures: the typical Paecilomyces-type with cylindrical to doliiform

conidia are produced at 37-40; at higher temperatures, conidia are subglobose to ovoid,

borne in shorter chains on phialides having a swollen and thick-walled apex.

Source : MNHN, Paris

32 J. MOUCHACCA

Coonemeria aegyptiaca was recently reported by Abdullah & Al-Bader (1990) to

inhabit soil in Iraq.

— Coonemeria verrucosa (Yaguchi, Someya et Udagawa) comb. nov.

basionym: Thermoascus crustaceus (Apinis & Chesters) Stolk var. verrucosus Yaguchi,

Someya et Udagawa — Mycoscience 36: 161. 1995.

= Thermoascus taitungiacus Chen K-Y. & Chen Z-C. — Mycotaxon 50: 226. 1996.

anamorph: Paecilomyces taitungiacus Chen K-Y. & Chen Z-C — Mycotaxon 50: 226.

1996.

= Paecilomyces crustaceus Apinis & Chesters pro parte fide Yaguchi, Someya et Udagawa

— Mycoscience 36: 161. 1995.

Descriptions: Yaguchi et al. (1995); Chen & Chen (1996).

Coloniae in agaro-malto addito dispositae post 7 diebus et temperatura 40" C cum

9.0 cm diametro, lanatae. Ascomates superficiales intermixta cum pauco conidiogenis struc-

turis. Mycelium ex hyphis hyalinis, septatis, ramosis, laevibus, 2-8 um cr

Ascoma non-ostiolatum, solitarium ad confluens deinde crustaceum, sphaericum

aurantiocamque, cum 300-600 um diametro. Peridium crassum, pseudparenchymatum, tex-

tura angulare. Asci dispersi in ascomatis cavitate, subglobosi vel piriformes, 12-16 x

11-14 um, octospori, evanescentes. Ascosporae unicellulariae, ellipsoideae, rare subglobosae,

hyalinae ad pallidae aurantiacae, 6-8 x 5-6 um, cum crasso verrucoso pariete.

Conidiophorum septum, laeve, hyalinum ad brunneum, 100-300 x 6-10 um. Apicis

regio irregulariter ramosa, terminales rami cum solitariis phialidibus vel verticillatis per 2-4.

Phialis cylindrica, 16-30 x 4-6 um. Conidia disposita in catenis non coalescentibus; conidia

cylindrica, flavida, laeva, 4-10 x 2-4 um; conidia elliptica aliquando, 5-8 x 4-6 um.

Holotypus: PF 1160, cultura exsiccata ex soli isolata a T. Yagushi, Guanghou in Sina,

4. X1.1993. In herbario Musei et Instituti Historiae Naturalis Chiba (CBM) deposita.

On common laboratory agar medium, colonies filling the plate in 7 d at 40° C,

lanate with superficial ascomata intermixed with sparse conidiophores and conidia, rosy

buff to orange, reverse reddish-brown; conidiogenesis inconspicuous not affecting colony

appearance. Optimal growth between 30 and 40? C; the minimum lies between 20 and

25? C and the maximum somewhat above 55? C.

Ascomatal initials a coiled hyphae. Cleistothecia solitary or confluent and then

forming a crusty layer, orange, spherical, 300-600 um diam.; peridium pseudoparenchy-

matous of textura angularis type, outer layer consisting of thick-walled yellow brown

angular cells, 4-8 x 2-6 um, inner layer of hyaline, angular or rounded cells, 10-20 um

wide. Asci irregularly disposed, 8-spored, globose to pyriform, 12-16 x 11-14 um, evanes-

cent. Ascospores unicellular, hyaline to pale-orange, ellipsoidal, rarely subglobose,

6-8 x 5-6 um, thick-walled, verrucose.

Conidiophores arise from the basal mycelium or from aerial hyphae; stipes

hyaline to brownish, septate, smooth, 100-300 x 6-10 um; apical parts not uniformly

branched giving rise to irregular verticils of terminal and subterminal secondary bran-

ches; these bear phialides either singly or in verticils of 2-4, cylindrical to slightly swollen,

16-30 x 4-6 um. Conidia produced in long disordered chains, unicellular, cylindrical,

truncate at both ends, yellowish, smooth, 4-10 x 2-4 um; few wider elliptical conidia

sometimes produced, 5-8 x 4-6 um.

Source : MNHN, Paris

THERMOPHILIC FUNGI 33

Holotype: PF 1160, a dried culture of a soil isolate from Guanghou, China, 4. XI.1993,

deposited at the Natural History Museum and Institute, Chiba, Japan (CBM) and at

T. Yaguchi collection (described as Thermoascus crustaceus var. verrucosus).

While describing Thermoascus taitungiacus, Chen & Chen (1996) were probably

unaware of the verrucose variety of Th. crustaceus established a year before by the

Japanese mycologists Yaguschi et al. (1995). Authentic material of both taxa have in

common a soil origin and not widely separated original locations. Indeed the former

derives from a weed soil located at Taitung in Taiwan, while the latter was isolated form a

soil sample taken from the Chinese locality of Guanghou.

Ascospores of the Taiwainese fungus were described as being yellowish green

(although overall colony color tends to orange tones), oval to elliptical, rarely subglobose,

6.3-7.5 x 4.5-5.6 um, thick-walled and predominantly echinulate when viewed under light

microscope but irregularly verrucose under SEM. A comparison of given SEM pictures

for both taxa clearly stress ascospore ornamentation is similar being represented by large

well individualised warts of heterogeneous shape.

Coonemeria verrucosa is mainly distinguished by its definitely verrucose ellipsoi-

dal ascospores. These are smooth in C. aegyptiaca and finely echinulated in C. crustacea.

The ascomycete might have been previously mislead with Coonemeria crustacea. Never-

theless further comparative studies are required to ascertain differences in growth tempe-

ratures relations and other minor morphological characters.

— Corynascus heterothallicus (van Klopotek) von Arx — Sydowia 34: 25.

1981(1982).

basionym: Thielavia heterothallica van Klopotek — Archives of Microbiology 107: 223-

224. 1976.

anamorph: Myceliophthora thermophila (Apinis) van Oorschot — Persoonia 9: 403. 1977.

basionym: Sporotrichum thermophilum Apinis as * thermophile’ — Nova Hedwigia 5: 74.

1963.

= Chrysosporium thermophilum (Apinis) van Klopotek — Archives of Microbiology 98:

366. 1974.

Descriptions: von Arx (1981(1982)); van Klopotek (1976); Domsch et al. (1980, as Thie-

lavia heterothallica).

Apinis (1963) isolated several strains of a new thermophilic “Sporotrichum”

anamorph from soil and plant debris in Nottingham, UK; he was not aware of the

heterothallic nature of the corresponding teleomorph. Few years later, von Arx (1971)

provided a modern definition of Sporotrichum Link based on a freshly isolated strain

matching the type species S. aureum Link: S. F. Gray. This form genus was then restricted

to hyphomycetes having basidiomycetous affiniti evidenced by the regular presence of

clamp connections at the cross walls and production of simple types of aleuriospores. On

account of this new definition, van Klopotek (1974) transferred Apinis fungus to Chrysos-

porium Corda.

Myceliophthora Costantin was reintroduced by van Oorschot (1977) as sugges-

ted earlier by von Arx (1973) in his treatment of Sporotrichum and related genera. This

disposition aimed to accomodate the type species Myceliophthora lutea Costantin and the

Chrysosporium (Sporotrichum) anamorphs of two hitherto described ascomycetes;

M. lutea has not yet developed a corresponding teleomorph. The fungus described by

Source : MNHN. Paris

34 J. MOUCHACCA

Apinis is now widely accepted as belonging to Myceliophthora. It differs from known

species by its dark colored colonies and smaller mostly obovate conidia, 4.5-11.0 x

3.0-4.5 um; conidia are hyaline, smooth and thick-walled.

The teleomorph was later discovered by van Klopotek (1976) after mating

appropriate strains. Developed cleistothecia produced ascospores ellipsoidal, dark brown,

7.5-11.0 x 5.0-7.0 um, provided with one distinctive germ pore. The teleomorph was

described as Thielavia heterothallica. Few years later, it was relocated in Corynascus von

Arx which groups ascomycetes having anamorphs of the Myceliophthora type (von Arx er

al., 1986).

An excellent account of the cultural and physiological characteristics and the

distribution of this fungus is provided by Domsch er al. (1980).

— Corynascus thermophilus (Fergus & Sinden) van Klopotek — Archives

of Microbiology 98: 366. 1974.

basionym: Thielavia thermophila Fergus & Sinden — Canadian Journal of Botany 47:

1635. 1969.

= Chaetomidium thermophilum (Fergus & Sinden) B. Lodha — In Taxonomy of Fungi.

Proceedings of the International Symposium Madras 1973. Part I: 248. 1978.

anamorph: Myceliophthora fergusii (van Klopotek) van Oorschot — Persoonia 9: 406.

1977.

basionym: Chrysosporium fergusii van Klopotek — Archives of Microbiology 98: 366.

1974.

Descriptions: Fergus and Sinden (1969); Hedger and Hudson (1970); van Klopotek (1974);

von Arx (1975).

Mating of several pure strains of another thermophilic “Sporotrichum” species

isolated from mushroom compost in Pennsylvania (USA) developed black ascocarps of a

new heterothallic cleistothecial ascomycete. This was described as Thielavia thermophila

by Fergus & Sinden (1969). No provision was however then made for the corresponding

anamorph. This was simply regarded as deviating from Sporotrichum thermophilum

Apinis by some cultural characteristics. Both hyphomycetes were later compared by

Hedger & Hudson (1970) following isolates obtained in Britain. Distinctive growth and

morphological features were also simply underlined by Hedger and Hudson.

In 1974, van Klopotek ascribed the anamorph of Thielavia thermophila to

Chrysosporium Corda while dedicating the hyphomycete to Fergus; she also transferred

the teleomorph to the recently established Corynascus von Arx. Chrysosporium fergusii

was later on relocated in Myceliophthora by van Oorschot (1977) together with the

anamorph of a second Corynascus species. However it is unfortunate the specific epithet

thermophila was not selected for the anamorph of Corynascus thermophilus. Such would

have prevented any form of confusion with the anamorph of Corynascus heterothallicus

(van Klopotek) von Arx named Myceliophthora thermophila (Apinis) van Oorschot.

Corynascus thermophilus being a heterothallic ascomycete is usually only repre-

sented by its aleuriospores in isolation studies. These are ellipsoidal or obovate, nearly

hyaline and measure 7-12 x 5-8 um. Mating of appropriate strains would produce cleis-

tothecia black, globose, usually smooth, up to 300 um diam. Asci are irregularly disposed,

each having four ascospores being ellipsoidal, dark brown, 22-32 x 17-22 um, provided

with two prominent germ pores.

Source : MNHN, Paris

THERMOPHILIC FUNGI 35

Corynascus novoguinensis (Udagawa & Horie) von Arx also produces a Myce-

liophthora anamorph but yet unnamed. However this fungus has its maximum growth

temperature at 40° C as compared to 55? C for Myceliophthora fergusii (van Oorschot,

1980).

— Dactylomyces thermophilus Sopp — Skrifter udgivne af videnskabs-

selskabet i Christiania. Mathematisk-naturvidenskabelig klasse 11: 35.

1912.

= Thermoascus thermophilus (Sopp) von Arx — The Genera of Fungi Sporulating in Pure

Culture: 84. 1970.

anamorph: Polypaecilum sp.; fide Apinis, 1967.

Misapplied names: Thermoascus aurantiacus Miehe 1907; sensu Cooney & Emerson —

Thermophilic Fungi: 39. 1964.

= Penicillium thermophilus (Sopp) Biourge 1923; sensu Cooney & Emerson — Thermo-

philic Fungi: 39. 1964.

= P. thermophilum (Sopp) Sacc., fide Trotter 1931 — Sylloge Fungorum 25 (Suppl. 10):

671.1931; sensu Cooney & Emerson — Thermophilic Fungi: 39. 1964.

Description: Apinis (1967).

This is the type species of Dactylomyces Sopp (Sopp. 1912). The original

material developed in Norway on the wooden casing of a bath thermometer. In the

protologue, Sopp suggested the new genus might be identical with Thermoascus Miehe,

despite his awareness of marked differences between respective type species; for example,

the presence of a penicillioid anamorph in his fungus and the absence of a distinctive

conidial state in the type species Thermoascus aurantiacus. Absence of authentic material

for effective comparison made a considerable impact on subsequent interpretations of

both genera. Such resulted in much confusion in the identity of respective type species.

The presence of a penicillioid anamorph in the description of Dactylomyces

thermophilus lead Biourge (1923) to list this name among the hitherto known Penicillia.

However Biourge did not include the corresponding Penicillium thermophilus in his group

of accepted species. Trotter (1931) published a short description of Penicillium thermophi-

lum (Sopp) Sacc. Later and for their treatment of Penicillia, Raper & Thom (1949: 20)

examined a culture obtained by Prof. Ralph Emerson from retting guayule at Salinas,

California, believed to represent Sopp's fungus. In addition to developed ascospores,

conidial structures were found by Raper & Thom to be very large and coarse, evanescent,

somewhat penicillate and thus not characteristic of their concept of Penicillium Link.

These authors then regarded Thermoascus as a possible synonym of Sopp's genus.

Few years after the publication of Raper & Thom's Manual of the Penicillia

(1949), Raper & Fennell deposited at the CBS the strain NRRL 1563 as Dactylomyces

thermophilus Sopp (List of Cultures, Supplement 1, 1962); this number originally referred

toa strain of Cephaliophora tropica but from about 1950, it was discovered that it has been

replaced by a strain of this ascomycete.

In 1964, Cooney & Emerson provided a latin diagnosis for Thermoascus auran-

tiacus based on their strain M 206516. This isolate was made from retting guayule in June

1945, at Salinas, California, but their is no clear indication whether it is the same strain

earlier examined by Raper & Thom (1949). For this ascomycete, Cooney & Emerson

depicted a distinctive conidial stage. Their illustration approximate figures produced by

Source : MNHN, Paris

36 J. MOUCHACCA

Sopp (1912) for his Dactylomyces thermophilus and which are rather reminiscent of

Paecilomyces Bain. Cooney & Emerson also admitted Sopp's suggestion his fungus being

identical to Thermoascus aurantiacus. They substantiated their conclusion by the assump-

tion that isolates of the latter examined by Miehe (1907) might represent “some naturally

occurring strains ot Thermoascus (Dactylomyces) incapable of producing conidia”.

As Thermoascus predates Dactylomyces, the former was thus retained. Earlier synonyms

of the type species of the latter were then disposed under Thermoacus aurantiacus. At the

date of publication of Cooney & Emerson's book on thermophilic fungi, Dactylomyces

was still a monospecific genus.

In 1964 however, Apinis & Chesters (1964) added a second species to Dactylo-

myces, D. crustaceus, developing a conspicuous Paecilomyes anamorph designated

P. crustaceus. For this work, they re-examined Raper & Fennell's Dactylomyces thermo-

philus strain (NRRL 1563; = CBS 374.62); this proved to match the fungus they were

proposing. One year later, Stolk (1965) also studied this isolate; she concurred with Apinis

& Chester's decision. Stolk then suggested Cooney & Emerson's neotype of Thermoascus

aurantiacus (M 206516) is most likely identical with Dactylomyces crustaceus.

Stolk (1965) also examined all cultures maintained at the CBS as Thermoascus

aurantiacus: CBS 256.34, CBS 257.34, CBS 415.62 and CBS 398.64. These strains were

characterized by the presence of ascospores being elliptical and finely echinulated, 5.0-7.0

x 3.5-5.0 ит, and the general absence of an associated anamorph producing chains of

conidia. Miehe (1907) had already stressed the absence of any conidial state producing

spore chains in his type species and such was confirmed few years later by Noack (1912).

Stolk then stresssed these features should distinguish Miehe's fungus from Dactylomyces

crustaceus Apinis & Chesters (having a Paecilomyces anamorph) and also from the yet

unclear D. thermophilus Sopp. Unfortunately Stolk (1965) tranferred Dactylomyces crus-

taceus to Thermoascus.

In 1967, Apinis re-examined Thermoascus aurantiacus neotype strain M 206516.

He found it to rather correspond to Dactylomyces crustaceus Apinis & Chesters having a

well defined Paecilomyces anamorph. This observation enabled Apinis to definitely refute

the similarity of these two teleomorphic genera as stated by Cooney & Emerson (1964).

At that time, Apinis was already familiar with the ascomycete described by Miehe; from

pasture soils, he had isolated (Apinis, 1963) several strains matching the original descrip-

tion.

Based on several fresh isolates originating from Sweden and England, Apinis

(1967) then provided an updated description of Dactylomyces type species, D. thermophi-

lus. The fungus has hyaline, unicellular ascospores, more or less oval and smooth, 5.5-8.0

x 3.5-6.0 um. Conidia are also produced; these are continuous, cylindrical to ovoid,

subhyaline, smooth, 3.0-11.0 x 2.5-5.5 um. Such ascospores and conidial dimensions are

somewhat smaller than those reported by Sopp (1912); however there is a general agree-

ment that measurements given by this author are unreliable. Apinis (1967) selected as

neotype strain BDUN 394 (= IMI 123298) obtained by T. Nilsson in Sweden. No

provision was established for the anamorph. He simply indicated chains of conidia are

produced by branched annellophores as in the recently described form genus Polypaecilum

G. Smith (Smith, 1961). Apinis proposed two new subgenera to consider distinctiveness in

anamorphs of both Dactylomyces: Subgenus Dactylomyces based on the type and subge-

nus Paecilomycopsis based on D. crustaceus. These subgeneric divisions were rapidly

regarded as superfluous.

Inspite of clarifications introduced by Apinis (1967), von Arx (1970) listed

Dactylomyces as congeneric with Thermoascus, a taxonomic disposition he maintained for

Source : MNHN, Paris

THERMOPHILIC FUNGI 37

several years (von Arx, 1987); apparently he was following Stolk (1965) who did transfer

Dactylomys crustaceus to Thermoascus. However this generic synonymy did not gain

general acceptance (Malloch & Cain, 1972; Cannon et al., 1985; Eriksson & Hawksworth,

1993). Dactylomyces Sopp is actually considered a valid distinctive generic entity.

— Melanocarpus albomyces (Cooney & Emerson) von Arx — Studies in

cology 8: 17. 1975.

basionym: Myriococcum albomyces Cooney & Emerson — Thermophilic Fungi: 60. 1964.

= Thielavia albomyces (Cooney & Emerson) Malloch & Cain — Canadian Journal of

Botany 50: 65. 1972.

Descriptions: Cooney & Emerson (1964); von Arx (1975); von Arx et al. (1988); Guarro et

al. (1996).

Cooney & Emerson ascribed this fungus to the sterile form-genus Myriococcum

Fr. based on Corda's interpretation of its type species, M. praecox (Corda, 1842). They

based their decision on the account both fungi have in common “the dark, shiny astomous

fruiting bodies, associated with a white, mucedineous subiculum”. Such an addition was

made inspite of their awareness no asci were ever reported in Myriococcum praecox and

also that what was considered as “spores” by Corda was apparently only the inner cells of

immature ascocarps.

The genus Melanocarpus von Arx (von Arx, 1975) was later proposed to acco-

modate this widespread ascomycete known to produce in culture a characteristic arthro-

conidial state. This anamorph is not developed by taxa of the two related genera Thielavia

Zopf and Chaetomidium (Fuckel) Zopf. Smooth ascomata and obovate-oblate darker

ascospores provided with a prominent germ pore (13-16 x 11-14 x 9-11 um) were also

then considered additional distinguishing features. Further, the presence of a pseudopa-

renchymatous wall in Melanocarpus albomyces precludes any confusion with the hitherto

known species of Thielavia (von Arx, 1975).

The original concept of Melanocarpus was however subsequently partly expan-

ded by von Arx et al. (1988) to allow provision for the mesophilous M. oblatus Guarro &

van der Aa described in the meantime; arthroconida are produced by this species. This

tendancy was also recently substantiated by Guarro ег al. (1996) with their description of

the mesophilic Melanocarpus coprophilus Guarro & Valldos., and the transfer of Thielavia

minuta var. thermophila Abdullah & Al-Bader. However both latter taxa are not known to

have an associated arthroconidial state or any other state, a feature that should have

favoured their inclusion rather in Chaetomidium.

— Melanocarpus thermophilus (Abdullah & Al-Bader) Guarro, Abdullah &

Al-Bader — Mycological Research 100: 75. 1996.

basionym: Thielavia minuta (Cain) Malloch & Cain var. thermophila Abdullah &

Al-Bader — Basrah Journal of Agricultural Science 5: 116. 1992.

Descriptions: Abdullah & Al-Bader (1992); Guarro et al. (1996).

Living strains of the thermophilic variety of Thielavia minuta (Cain) Malloch &

Cain (a mesophile) originate from a forest soil in the north of Iraq. Re-examination of

authentic material led Guarro ег al. (1996) to relocate the fungus in Melanocarpus von Arx

Source : MNHN, Paris

38 J. MOUCHACCA

as M. thermophilus. However this ascomycete produces not the arthroconidial anamorph

characteristic of the type species. Asci are 8-spored with ascospores being ovoid, dark

brown, 7.5-9.0 x 6.0-7.5 um, each provided with a single germ pore.

— Talaromyces byssochlamydioides Stolk & Samson — Studies in Myco-

logy 2: 45. 1972.

anamorph: Paecilomyces byssochlamydioides Stolk & Samson — Studies in Mycology 2:

45. 1972

Descriptions: Stolk & Samson (1972); Awao & Otsuka (1974).

This species of Talaromyces Benjamin is definitely less reported than its close

relative T. emersonii Stolk. It is mainly distinguished by its conspicuous Paecilomyces

anamorph as compared to the Penicillium state of the latter. Ascomata always develops in

culture concomitantly with the anamorph and such prevents its confusion with the similar

imperfect taxon Paecilomyces variotii Bainier. Ascospores are globose to subglobose,

3.7-4.5 x 3.5-4.0 um, thick-walled smooth or nearly so, often partially covered by material

which may represent the remnants of a gelatinous covering.

Dactylomyces crustaceus Apinis & Chesters also has a Paecilomyces state but its

conidia are ellipsoidal as compared to the cylindrical conidia of Talaromyces byssochla-

mydioides. The latter was apparently only reported from soil in Japan (Awao & Otsuka,

1974) and Egypt (Mouchacca, 1995).

— Talaromyces emersonii Stolk — Antonie van Leeuwenhoek 31: 262.

1965; Stolk & Samson — Studies in Mycology 2: 48. 1972.

= Byssochlamys sp. fide Cooney & Emerson — Thermophilic Fungi: 155. 1964.

anamorph: Penicillium emersonii Stolk — Antonie van Leeuwenhoek 31: 262.1965.

= Geosmithia emersonii (Stolk) Pitt — Canadian Journal of Botany 57: 2027. 1979.

Misapplied names: Talaromyces dupontii (Griffon & Maublanc) Apinis; sensu Apinis —

Nova Hedwigia 5: 72. 1963; as comb. nov. (nom. inval., Art. 36.1).

= Penicillium dupontii Griffon & Maublanc 1911; sensu Apinis — Nova Hedwigia 5: 72.

1963.

Descriptions: Stolk (1965); Stolk & Samson (1972); Domsch et al. (1980).

Talaromyces emersonii was described inclusive of its distinctive anamorph deve-

loping Penicillia of the Asymmetrica type. The selected representative strain was obtained

by Mrs. A. J. van der Plaats-Niterink from Italian compost but other isolates were also

then available at the CBS. Dedicated to R. Emerson for his contribution to our knowledge

of thermophilic fungi.

Apinis (1963) based his taxonomic decision on strain BDUN 272 (- CBS

397.64) isolated from soil near Nottingham (UK). Stolk (1965) re-examined this isolate

which proved to represent Talaromyces emersonii rather than the teleomorph of Peni-

cillium dupontii as concluded by Apinis. The same observation also applies to strain CBS

394.64 labelled Byssochlamys sp. by Cooney & Emerson (Stolk, 1965).

The anamorphic genus Geosmithia Pitt (Pitt, 1979) was erected to accomodate

Penicillia formerly placed in the Penicillium pallidum-series. These are mainly distin-

guished by the formation of cylindrical conidia borne from cylindroidal, rough-walled

Source : MNHN. Paris

THERMOPHILIC FUNGI 39

phialides and not colored green en masse. Stolk & Samson (1985) emitted doubts as to the

necessity of such a generic distinction based on slight morphological differences.

In Penicillium such differences are instead appropriately used to delimit generic sections.

The anamorph of Talaromyces emersonii is thus better referred to as a Penicillium.

Talaromyces emersonii was subsequently reported from various habitats

(Domsch et al., 1980). It produces globose, reddish to orange brown ascomata, up to

300 um diam.; ascospores are thick-walled, smooth, subglobose to ovoidal, 3.5-4.0 x

2.7-3.5 um; ascospores may be covered by material representing remnants of a gelatinous

coating.

— Talaromyces thermophilus Stolk — Antonie van Leeuwenhoek 31: 268.

1965; Stolk & Samson — Studies in Mycology 2: 55. 1972.

= Penicillium dupontii Griffon & Maublanc emend. Emerson in Raper & Thom —

A Manual of the Penicillia: 573-577. 1949.

= Talaromyces dupontii (Griffon & Maublanc) Emerson, incidently mentioned by Fergus

— Mycologia 56: 277. 1964 (nom. inval., Arts. 36.1 & 37.1).

anamorph: Penicillium dupontii Griffon & Maublanc — Bulletin trimestriel de la Société

mycologique de France 27: 73. 1911.

= ? Citromyces sphagnicola Mal'chevskaya — Trudy Pushkin nauchno-issled. Lab. Rasv.

Sel'skokhoz. Zhivot. Inst. 13: 23. 1939.

Misapplied names: Talaromy

Nova Hedwigia 5: 72. 1963, as comb. nov. (nom. invalid., Art. 36.1).

Talaromyces ( Penicillium) dupontii (Griffon & Maublanc) emend. Emerson in Raper &

Thom: 573. 1949; 1. c. Cooney & Emerson — Thermophilic Fungi: 28. 1964 (nom. inval.,

Art. 36.1).

Descriptions : Stolk (1965); Stolk & Samson (1972); Pitt (1979).

The original publication of Griffon & Maublanc (1911) dealt only with the

Penicillium anamorph. No corresponding teleomorph was reported from cultures of the

two strains then available for study. These were obtained in France from manure and damp

hay by Mr. Dupont, Chief-Chemist at the Ecole Nationale d'Agriculture de Grignon, and

to whom the fungus was dedicated. Unfortunately, the two original isolates are definitely

lost.

The fungus was later on and in 1945 isolated by Emerson from retting guayule

shrub at Salinas, California; for the first time the perfect state developed in culture.

Emerson then prepared an emended description of both states of Penicillium dupontii

based upon his strain No. 26 (= NRRL 2155) to be incorporated by Raper & Thom in

their first Manual of the Penicillia (1949: 573).

In 1963, some confusion about this taxon was introduced by Apinis. Under the

binomial Talaromyces duponti (Griffon & Maublanc) Apinis, he provided a description of

a teleomorphic fungus thought to "correspond in general to the original strain described

from France". As representative material, Apinis selected his strain BDUN 272 origina-

ting from a water-logged pasture in Nottingham. The above binomial was however not

validly published as no latin diagnosis provided and no new type material specified.

Further confusion but of the nomenclatural type was also simultaneously

introduced by Fergus (1964) following his study of an isolate from compost in Pennsylva-

nia (USA). His observations were published under the name Talaromyces dupontii (Grif-

fon & Maublanc) Emerson, a combination not proposed as such by Emerson himself.

Source : MNHN, Paris

40 1. MOUCHACCA

Unfortunately, this designation was subsequently reproduced in several studies of ther-

mophilic fungi.

Two years later, Stolk (1965) re-examined Apinis strain BDUN 272. She found it

to largely deviate from Emerson's isolate No. 26 (= NRRL 2155; = CBS 236.58). Moreo-

ver, the former proved to perfectly match her newly described Talaromyces emersonii Stolk.

To eliminate the state of confusion prevailing around the name Talaromyces dupontii,

Stolk then proposed the new name Talaromyces thermophilus for this teleomorph and

provided a latin diagnosis. Emerson strain was then selected as holotype (Stolk, 1965; Pitt,

1979). However, the original accession number of this holotype was erroneously cited by

Cooney & Emerson (1964: 28): under the diagnosis provided for this fungus is indicated

specimen M 206516 (our culture No. 26). In fact strain M 206516 was selected by Cooney

& Emerson as representing their interpretation of Thermoascus aurantiacus Miehe (1964:

50) and this corresponds to their annotation: our culture No. 2.

Pitt (1979) stressed the reasons why the corresponding perfect state should be

maintained in Talaromyces and to continue considering its simple reduced anamorph as a

Penicillium. Talaromyces thermophilus is the only thermophile with a Penicillium anamor-

phic state producing green conidia. This character should prevent any misidentification

since ascocarps do not always readily develop in cultures of freshly isolated strains. The

fungus grows fairly rapidly and optimally at 45-50? C; no growth develops at 25° C and

60° C respectively. Ascospores are ellipsoidal, 3.5-4.5 x 2.2-3.5 um, ornamented by 2-6

somewhat jagged, irregular, usually longitudinal ridges.

— Thermoascus aurantiacus Miehe — Die Selbsterhitzung des Heues:

70. 1907.

= ? Thermoascus isatschenkoi Mal'chevskaya — Trudy Pushkin. Nanchno-issled. Lab.

Калу. sel'khoz Zhivot 13:26. 1939; fide Cooney & Emerson — Thermophilic Fungi: 39.

1964.

Misapplied names: Thermoascus aurantiacus Miehe 1907; sensu Cooney & Emerson

Thermophilic Fungi: 39. 1964

Dactylomyces thermophilus Sopp 1912; sensu Cooney & Emerson — Thermophilic Fungi:

39. 1964.

Penicillium thermophilus (Sopp) Biourge 1923; sensu Cooney & Emerson — Thermophilic

Fungi: 39. 1964.

Penicillium thermophilum (Sopp) Sacc., fide Trotter 1931; sensu Cooney & Emerson —

Thermophilic Fungi: 39. 1964.

Descriptions : Stolk (1965); Apinis (1967); Awao & Otsuka (1973); Domsch et al. (1980);

Chen & Chen (1996)

The type species of Thermoascus Miehe (Miehe, 1907), T. aurantiacus, was

isolated from self-heating hay and carefully described by the author. Few years later, Sopp

(1912) reported a second thermophilic ascomycete having a well developed conidial state

and for which he proposed the new genus Dactylomyces. Sopp then considered Thermoas-

cus aurantiacus as approximating his Dactylomyes thermophilus n. sp. ad interim. This

suggestion coupled with the lack of any authentic material for either ascomycetes resulted

in much confusion about the exact nature of Miehe's fungus.

In 1963, Apinis isolated from soil near Nottingham several strains he referred to

Thermoascus aurantiacus. These isolates exhibited no morphological deviations from the

original description. They also proved to match a strain maintained at the CBS under this

Source : MNHN, Paris

THERMOPHILIC FUNGI 41

binomial and isolated by Noack (1912). Apinis noted the structure of the cleistothecium

be related to certain species of the Gymnoascaceae with the presence of large clavate

conidia reminescent of “clasterospores ” of some Trichophyton species.

In 1964, Cooney & Emerson provided a detailed description and a latin diagno-

sis of Thermoascus aurantiacus based on their “strain M 206516 (our culture No. 2)”

thought to match Miehe's fungus. This was also then regarded as an earlier name of

Dactylomyces thermophilus. In the established description, a distinctive conidial state is

depicted. This anamorph approximate figures produced by Sopp (1912) and which were

later regarded (Stolk, 1965) as rather reminiscent of Paecilomyces Bain. (see also com-

ments under Dactylomyces thermophilus).

However, in the same year, Apinis & Chesters (1964) introduced Dactylomyces

crustaceus (anamorph: Paecilomyces crustaceus) and reported to it the CBS strain 374.62

(2 NRRL 1563) labelled Dactylomyces thermophilus by Raper & Fennell. In 1965, Stolk

re-examined this strain and concurred with Apinis & Chester's decision. She then sugges-

ted Cooney & Emerson's neotype of Thermoascus aurantiacus (M 206516) is most likely

identical with Dactylomyces crustaceus.

Stolk (1965) then examined all cultures maintained at the CBS as Thermoascus

aurantiacus: CBS 256.34, CBS 257.34, CBS 415.62 and CBS 398.64. These isolates

produced elliptical finely echinulated ascospores, measuring 5.0-7.0 x 3.5-5.0 um.

No associated anamorph producing chains of conidia was developed by any. The absence

of an anamorph producing spore chains was already stressed by Miehe (1907) and such

was confirmed few years later by Noack (1912). Stolk then underlined these features

support the distinctiveness of Thermoascus aurantiacus from Dactylomyces thermophilus

(having a yet undefined anamorph) and also from the well described Dactylomyces

crustaceus and its Paecilomyces anamorph. Unfortunately Stolk then tranferred the latter

to Thermoascus.

In 1967, Apinis published a comparative study of Thermoascus and Dactylomy-

ces based on freshly isolated strains. He re-examined Cooney & Emerson strain M 206516

and confirmed Stolk (1965) suggestion about its similarity with Dactylomyces crustaceus.

This lead to a definite rejection of Cooney & Emerson's taxonomic considerations about

Thermoascus type species and to its identity with Dactylomyces thermophilus. Second, the

morphology of Apinis fresh isolates of Thermoascus aurantiacus was in line with Stolk

(1965) observations.

Apinis also noted the presence in the aerial mycelium of “conidia — of Apha-

noascus or Microsporum type — developing terminally on long or short hyphal branches

singly and being clavate or somewhat spindle-shaped, smooth, 0-3 septate, 12-35

x 5-10 um". Miehe (1907) did mention such aleuriospores in the type species. Strain

BDUN 343 (= IMI 91787) isolated from alluvial grassland soil was designated neotype

for Thermoascus aurantiacus.

Recent reports confirms Thermoascus aurantiacus have a wide distribution

(Domsch ег al., 1980; Chen & Chen, 1996). This ascomycete proved to be a strong

thermophile with growth starting at 30° C and up to 62° C; growth optimum around 45° C

with formed colonies being bright orange to orange brown. Ascospores are definitely

elliptical and slightly roughened. Presence of terminal aleuriospores apparently depends

on examined strains.

Thermoascus was placed in the family Onygenaceae (order Onygenales) by Benny

& Kimbrough (1980); they placed Dactylomyces in the Trichocomaceae (order Eurotiales).

Tt was maintained in this family by von Arx (1987) but with Dactylomyces being a

synonym. Thermoascus was however excluded by Currah (1985) from the Onygenaceae

Source : MNHN, Paris

42 J. MOUCHACCA

because "there is no evidence of keratinolytic abilities nor does it have strictly rhexolyti-

cally dehiscing conidia". Currah mentions not Dactylomyces.

Thermoascus isatschenkoi is regarded as a doubtful species of which no satisfac-

tory description exists and no material is available for comparison (Cooney & Emerson,

1964; Apinis, 1967).

— Thermoascus aurantiacus Miehe var. levisporus Upadhyay, Farmelo,

Goetz & Melan — Mycopathologia 87: 73. 1984.

The original isolate was obtained from a top layer soil at La Ceiba, Republic of

Honduras. Minimum and maximum growth temperatures are 31 and 61? C respectively

with the optimum being at 49-50? C. The variety differs mainly by ellipsoidal smooth

rather than “echinulate ascospores”, 3.7-7.1 x 2.2-5.5 um (5.0-7.0 x 3.5-5.0 um for the

species). Conidial anamorph of the aleuriospore type matching those of the species were

infrequent, borne terminally, clavate, thick-walled, smooth, 15-25 x 7-17 um. All other

characters duplicate the species. Production of protease enzymes was also assessed (Marcy

et al., 1984).

— Thielavia australiensis Tansey & Jack — Canadian Journal of Botany

53: 82. 1975.

Descriptions: Tansey & Jack (1975); von Arx (1975); von Arx et al. (1988).

The protologue was based on strains isolated from nesting material of an

incubator bird: the mallee fowl, Leipoa ocellata Gould in New South Wales, Australia.

Optimum growth recorded at 35-40? C; maximum at 50? C; minimum not defined.

This Thielavia is distinguished by small pyriform brown ascospores, 6-8 x

5-6 um, having a germ pore at the attenuated end. Simple aleurioconidia are produced in

culture according to the protologue; these are continuous, lateral, sessile, colorless, ovoid,

5-8 x 3-5 um. The fungus has apparently not been reported after its description (von Arx

et al., 1988).

— Thielavia pingtungia Chen K-Y. & Chen Z-C. — Mycotaxon 60:

242. 1996.

The fungus was isolated from a sugar-cane field in Taiwan. The specific epithet

refers to the original locality: Pingtung. No growth developed between 25 and 30? C with

the optimum being around 40? C and the maximum fairly above 50? C.

The species is characterized by dark globose cleistothecia covered with brown

thick-walled hairy appendages; ascomatal hairs of the Chaetomium type, 2.5-4.0 um wide

and up to 350 ит long; ascomatal wall pseudoparenchymatous. Asci cylindric, 40-52 x

7-9 ym, stipitate, fasciculate, 8-spored. Ascospores usually uniseriate, globose to subglo-

bose, dark brown, smooth, thick-walled, 8.5-10.0 x 6.5-8.5 um. No anamorph developed

in examined cultures.

Thielavia pingtungia have several features in common with species assigned to

Chaetomidium; for the moment, the latter groups only mesophilic ascomycetes (Silva &

Hanlin, 1996).

Source : MNHN, Paris

THERMOPHILIC FUNGI 43

— Thielavia terrestris (Apinis) Malloch & Cain — Canadian Journal of

Botany 50: 66. 1973.

basionym: Allescheria terrestris Apinis — Nova Hedwigia 5: 68. 1963.

anamorph: Acremonium alabamense Morgan-Jones as ‘alabamensis’ — Canadian Journal

of Botany 52: 429. 1974.

Descriptions : Apinis (1963); Malloch & Cain (1973); von Arx (1975).

The original material was observed by Apinis (1963) in the course of his work on

thermophilous fungi inhabiting allluvial soils in Great Britain. He described the fungus as

Allescheria terrestris without providing any argument favouring such a decision; he also

assigned the anamorph to Cephalosporium (now Acremonium Link:Fr.). Allescheria ter-

restris was then transferred to Thielavia Zopf. Following its description, the fungus was

reported from various habitats and is now known to display a wide geographic distribu-

tion. Ascospores are ovate or pyriform, brown, thick-walled, provided with a distinct germ

pore at the attenuated end, 5.0-7.5 x 4.0-5.5 um.

The hyphomycete Acremonium alabamense was described exclusive of a teleo-

morph; it was isolated from needles of Pinus taeda. Later on, Samson et al. (1977) found

it to match the anamorph of Thielavia terrestris. For this ascomycete, sometimes only the

anamorph is observed during isolation studies and appropriate matings are required for

the development of the teleomorph. These authors conducted extensive mating experi-

ments with several strains of T. terrestris and A. alabamense; they came to the conclusion

that the mating behaviour of Thielavia terrestris could best be interpreted as indicating

homothallism with cross-feeding.

Some species of Chaetomium also produce in culture a phialidic state approxi-

mating Acremonium alabamense. The latter was recently selected as the type of the new

section Chaetomioides of Acremonium established to also accomodate phialidic states of

some Chaetomium species (Morgan-Jones & Gams, 1982).

THERMOPHILIC HYPHOMYCETES

— Acremonium alabamense Morgan-Jones as “alabamensis” —

Canadian Journal of Botany 52: 429. 1974.

teleomorph: Thielavia terrestris (Apinis) Malloch $: Cain — Canadian Journal of Botany

50: 66. 1973.

Descriptions: Apinis (1963); Morgan-Jones (1974); Morgan-Jones & Gams (1982).

As underlined before, this hyphomycete was described exclusive of the teleo-

morph being isolated from needles of Pinus taeda collected in the state of Alabama (USA).

The teleomorph was described before from alluvial soils in Nottingham (UK) with the

anamorph being indicated imply representing a Cephalosporium sp.

The repeated isolation of an Acremonium sp. from heated habitats led Samson ef

al. (1977) to compare it with the fungus described by Morgan-Jones and the anamorph of

Source : MNHN, Paris

44 J. MOUCHACCA

Thielavia terrestris. All three hyphomycetes were found to represent the same fungus. This

finding rose few questions concerning the developmental behaviour of the teleomorph.

Extensive mating studies were then undertaken with isolates of Thielavia terrestris,

Acremonium sp. and of A. alabamense. Although results allowed not a definite conclusion

as to the heterothallic nature of the teleomorph, these support the hypothesis indicating

the mating behavious of Thielavia terrestris is a case of homothallism with cross-feeding

(Samson et al., 1977).

Acremonium alabamense could thus be observed alone in studies involving high

temperatures incubation. It was recently selected as type of the new section Chaetomioides

of the genus established to also accomodate the morphologically similar phialidic states of

some Chaetomium species (Morgan-Jones & Gams, 1982). The fungus produces compa-

ratively fast growing colonies, velvety, whitish, with yellowish to brownish runner hyphae,

3-4.5 um wide. Conidiophores are simple, short, 8-25 x 1-1.5 um. Conidia are obovoid to

pyrfiorm, smooth, with a truncated base, 3-6 x 2-3 um (Morgan-Jones, 1974).

— Acremonium thermophilum W. Gams & Lacey — Transactions of the

British mycological Society 59: 520. 1972.

The described material developed on self-heated sugar cane bagasse in Trinidad.

The fungus is regarded as unique among known Acremonium Link:Fr. species on account

of its thermophilic habit and production of submerged hyphae partly having pigmented

walls. The species was assigned in Acremonium sect. Nectrioidea due to the development of

thick-walled conidiophores with basitonous ramification. Growth is strong but slow at

20° C, very good between 25 and 40° C and very weak at 47° C. Conidia are ellipsoidal,

3.0-4.0 x 1.3-1.7 um

— Humicola hyalothermophila Moubasher, Mazen & Abdel-Hafez —

Transactions of the British mycological Society 72: 509. 1979.

Descriptions: Moubasher et al. (1979); Moubasher (1993).

This soil-borne hyphomycete was originally isolated from several localities in

Jordan. No growth develops either at 28 or 55° C with good development being at 45° C;

growth optimum value is not specified. This thermophile was distinguished from the

mesophilic Humicola fuscoatra Traaen mainly by its slightly larger hyaline conidia (not

colored light brown as in H. fuscoatra) and intercalary chlamydospores. It was later on

Observed in Saudi Arabian soils (Bokhary, 1986).

The taxonomic position of this fungus needs to be re-assessed.

— Malbranchea cinnamomea (Libert) van Oorschot & de Hoog — Myco-

taxon 20: 129. 1984.

basionym: Trichothecium cinnamomeum Libert — Plantae cryptogamae Arduenna, Coll. 1,

Nr. 1013. 1830.

= Geotrichum cinnamomeum (Libert) Sacc. — Revue Mycologique (Toulouse) 11: 55.

1881; Michelia 2: 636. 1882.

= Thermoideum sulfureum Miehe — Deutsche Botanische Gesellschaft 25: 515. 1907.

= Malbranchea pulchella Sacc. — Sydow, Annales Mycologici, Ser II, 6: 557. 1908; Sacc. &

Traverso — Sylloge Fungurom 20: 11. 1911.

z Malbranchea pulchella Sacc. & Penzig var. sulfurea (Miehe) Cooney & Emerson —

Thermophilic Fungi: 102. 1964.

Source : MNHN, Paris

THERMOPHILIC FUNGI 45

= Malbranchea sulfurea (Miehe) Pidoplichko — In “Fungus Flora of Coarse Fodders (in

Russian)": 170. 1953.

= Malbranchea sulfurea (Miehe) Sigler & Carmichael — Mycotaxon 4: 441. 1976.

Descriptions: Cooney & Emerson (1964); Sigler & Carmichael (1976).

Miehe (1907) erected Thermoideum, type species T. sulfureum, for a hyphomycete

he encountered during his pioneer investigation of the self-heating process of hay.

He studied the fungus in culture and stressed its thermophilic nature. Saccardo (1908)

however immediately considered this type species as matching the morphologically close

mesophilic type species of his genus Malbranchea, M. pulchella Sacc.

In 1964, Cooney & Emerson provided an excellent account of a strain matching

Miehe's description. For this fungus, they simply proposed the varietal name sulfurea

pending a comprehensive comparison with the almost identical mesophile Malbranchea

pulchella. The comparison was later undertaken by Sigler & Carmichael (1976); they

concluded high temperature requirements are sufficient to warrant a specific status and

provided the binomial Malbranchea sulfurea (Miehe) Sigler & Carmichael.

The combination Malbranchea cinnamomea was based on Trichothecium cinna-

momeum Libert. It was established by van Oorschot & de Hoog (1984) after examining

dried authentic material of the latter. However the possible similarity with Malbranchea

sulphurea was not considered. Such was established later on by Sigler (1987) after a study

of appropriate authentic material.

Malbranchea cinnamomea is an easily recognizable thermophilic hyphomycete

being recorded on a variety of substrates under different conditions (Sigler & Carmichael,

1976).

— Myceliophthora fergusii (van Klopotek) van Oorschot — Persoonia 9:

406. 1977.

basionym: Chrysosporium fergusii van Klopotek — Archives of Microbiology 98: 366.

1974.

Teleomorph: Corynascus thermophilus (Fergus & Sinden) van Klopotek — Archives of

Microbiology 98: 366. 1974.

Descriptions: van Klopotek (1974); van Oorschot (1977, 1980).

As underlined under the teleomorph, provision for the anamorph of this asco-

mycete was made several years after the discovery of the heterothallic nature of the perfect

state. The anamorph was simply stated as being distinct from the close previously

described hyphomycete now renamed Myceliophthora thermophila. Both anamorphs can

thus be observed separately from their respective teleomorphs in mycological analyses

conducted at high incubation temperatures.

Myceliophthora fergussi produces pinkish-cream floccose colonies; aleuriospo-

resare pyriform to clavate, smooth and thick-walled, nearly hyaline and with narrow basal

attachments, 5-12 x 3-5 um.

Source : MNHN, Paris

46 1. MOUCHACCA

— Myceliophthora hinnulea Awao & Udagawa — Mycotaxon 16:

438. 1983.

The type locality is cultivated soil in Japan. Fungal growth is extremely reduced

at 20°C; optimal growth is at 40-45? C and maximum somewhat above 50° C.

No connection with a teleomorph yet established.

Myceliophthora hinnulea differs from the five previously described species (van

Oorschot, 1980) mainly by dull to greyish brown colonies and brownish conidia conspi-

cuously verrucose to spinulose, 8.0-10.0 x 6.0-7.5 um. Almost all known members of this

genus are thermotolerant or thermophilic with sporulation often being good between

30-40% C.

— Myceliophthora thermophila (Apinis) van Oorschot — Persoonia 9:

403. 1977.

basionym: Sporotrichum thermophilum Apinis as ' thermophile' — Nova Hedwigia 5: 74.

1963.

= Chrysosporium thermophilum (Apinis) van Klopotek — Archives of Microbiology 98:

366. 1974.

= Myceliophthora indica Basu — Nova Hedwigia 40: 85. 1984. (nom. inval., Art. 37.1).

Teleomorph: Corynascus heterothallicus (van Klopotek) von Arx — Sydowia 34: 25. 1981.

Descriptions: van Oorschot (1977, 1980); van Klopotek (1974)

As stressed before, Myceliophthora thermophila was described exclusive of its

corresponding teleomorph. Since the latter is heterothallic, the anamorph could thus be

observed alone in studies involving thermophilic fungi. The species differs from other

members of the genus by its dark colored colonies, occasionnally greenish and by smaller

mostly obovate conidia, 4.5-11.0 x 3.0-4.5 um; conidia are hyaline, thick-walled and

rough. Fresh isolates always have some rough conidia but older cultures tend to produce

only smooth ones. The species displays a wide geographic distribution being a common

component of decaying manure, silage, wood chips and pulp, etc. (Cannon, 1990).

Myceliophthora indica was isolated from garden soil and from decomposed

leaves of Clitoria sp. Attemps to locate original material were unsuccessful although Basu

(1984) underlined her intention to deposit both available strains at the CBS. The fungus

was compared with the type culture of Myceliophthora thermophila considered by Basu as

being thermotolerant. The “strongly thermophilic” Indian strain was found to deviate

mainly by smaller definitely roughened conidia. No mating attempts were undertaken and

the existence of a known teleomorph not stressed in the publication. Analysis of the

protologue clearly indicates the Indian strain do represent Myceliophthora thermophila.

— Scytalidium indonesicum Hedger, Samson & Basuki — Transactions of

the British mycological Society 78: 365. 1982.

The original material was isolated from soil of the Bogor Botanic Garden, West

Java. The fungus was also recovered from Dipterocarp forest soils in South Sumatra. It was

reported as being simply “thermophilous” able to grow rapidly at 45? C: 8.5 cm at 36 h.

Later Straatsma & Samson (1993) stated it is thermophilic.

The Indonesian taxon is distinguished by the production of conidia (intercalary

chlamydospores) thick-walled brown, ellipsoid to barrel-shaped, often with irregular

Source : MNHN, Paris

THERMOPHILIC FUNGI 47

outgrowths and also often constricted at the middle of the cell, 15-25 x 7-12 um; on

maturity these conidia secede rather easily and appear irregular in shape. Dark brown and

thick-walled similar but less wider conidia (termed arthroconidia) also develop in chains,

13-32 x 5-8 um; these do not secede easily. The presence of terminal conidia (or lateral) is

not underlined.

Scytalidium indonesicum approximates S. thermophilum which mostly produces

spherical to subspherical dark brown smooth conidia 9-14 um wide; oblong or ellipsoidal

ones measure 8-18 x 7-14 um. However, neither these Scyralidium develop the second

hyaline arthroconidial state characteristic of the type species, S. /ignicola (Ellis M. B.,

1976). The description of S. indonesicum is however in line with the introduction in

Scytalidium of taxa only developing dematiaceous arthroconidia (Sigler & Wang, 1990).

Such additions makes Scytalidium a heterogeneous entity.

Scytalidium thermophilum (Cooney & Emerson) Austwick — New

Zealand Journal of Agricultural Research 19: 29. 1976; emend. Straatsma

& Samson — Mycological Research 97: 327. 1993.

basionym: Torula thermophila Cooney & Emerson — Thermophilic Fungi: 92. 1964.

Tumicola insolens Cooney & Emerson — Thermophilic Fungi: 79. 1964.

= Humicola fuscoatra var. longispora forma insolens (Cooney & Emerson) Fassatiova —

Ceska Mykologie 21: 80. 1967.

= Humicola grisea Traaen var. thermoidea Cooney & Emerson — Thermophilic Fungi: 79.

1964.

= Humicola insolens Cooney $: Emerson var. thermoidea D. H. Ellis — Transactions of the

British mycological Society 78: 133. 1982.

= Humicola fuscoatra Traaen var. nigra Subrahmanyam — Hindustan Antibiotics Bulle-

tin 24: 41. 1982. (nom. inval., Art. 36.1: description only); Ibid. 25: 62. 1983 (latin diagnosis;

nom. inval., Art. 37.1).

= Humicola nigrescens Omvik var. thermorongeura Subrahmanyam — Hindustan Anti-

biotics Bulletin 24: 45. 1982. (nom. inval., Art. 36.1: description only); Ibid. 25: 62. 1983

(latin diagnosis; nom. inval., Art. 37.1).

= Scytalidium allahabadum Narain, Srivastava & Mehrotra — Zentralblatt für Mikrobio-

logie 138: 570. 1983.

Descriptions: Cooney & Emerson (1964); Ellis M. B. (1976); Straatsma & Samson (1993).

Cooney & Emerson (1964) while describing Humicola insolens and H. grisea var.

thermoidea indicated “the problems concerned with the Monotospora-Humicola-Torula

can only be resolved when all forms, both thermophilic and mesophilic, can be studied and

compared in detail". The genus Monotospora was cited in relation to Mason (1941) who

had then concluded that M. dalae Mason predates Humicola fuscoatra Traaen. Torula

thermophila was apparently not concerned by this statement since its description is found

some twenty pages later. Humicola grisea var. thermoidea was considered as a variety

(although not producing phialospores as the species) "chiefly because of the uncommon

occurrence of intercalary chlamydospores”. The abundance of these structures was then

used to distinguish Humicola insolens Cooney & Emerson.

Later, Emerson (1968) stressed * Humicola grisea var. thermoidea has smooth-

walled chlamydospores (aleuriospores) borne singly on short lateral branches with the

almost absence of any intercalary chlamydospores; on the other hand, isolates of Humi-

cola insolens regularly produces intercalary chlamydospores singly, in pairs or in short

Source : MNHN, Paris

48 J. MOUCHACCA

chains in addition to solitary terminal spores on short lateral branches; in Torula thermo-

phila chlamydospores are again smooth and brown, all formed in longer or shorter

intercalary chains and rarely in a terminal position”.

The taxonomic status of this Humicola-Torula complex remained unchanged

until Austwick (1976) tranferred Torula thermophila to Scytalidium Pesante sensu Elli:

M. B. (1971); the latter had emphasized the dark pigmented arthroconidia of the type

species, S. lignicola. However Austwick did not provide any argument in favour of such a

transfer. Later Sigler & Carmichael (1976) in the course of their study of hyphomycete:

with arthroconidia accepted Scytalidium as delimited by Ellis M. B.; seven species wer

then retained with some developing only the dematiaceous chlamydosporic state. These

additions introduced much heterogeneity in the genus (Sigler & Wang, 1990).

Ellis D. H. (1982) conducted ultrastructural studies of the conidial ontogeny «

both Humicola proposed by Cooney & Emerson (1964). After examining type strains anc

other isolates, he concluded H. grisea var. thermoidea is a separate entity exhibitir

considerable genetic variation among strains; further, it should rather be considered

variety of. Humicola insolens. This proposal was not in line with the suggestion emit!

earlier by Awao & Otsuka (1974) stating that both Cooney & Emerson's Humicola mig!

represent the same fungus.

The respective status of these three hyphomycetes remained as such until the

recent publication made by Straatsma & Samson (1993). They compared a large num!

of strains labelled Torula thermophila, Humicola grisea var. thermoidea or H. insolens

including corresponding authentic material. Their conclusion was that all such strains

represent one single variable species or a “morphologically indistinguishable sp:

complex" for which the binomial Scytalidium thermophilum (Cooney & Emerson) ^:

twick should continue to be applied pending further studies. Such a limitation accounts or

the fact the type species of Scytalidium is a dimorphic fungus having in addition a

arthroconidial synanamorph; the use of the binomial Scytalidium thermophilum as such is

questioned.

Humicola fuscoatra var. nigra Subrahmanyam was isolated form soil at Kunoor

India. The protologue almost duplicates that of Humicola insolens sensu Coone

Emerson except that aleuriospores of the Indian strain are indicated as being some:

larger: Aleuriospores are unicellular, rarely bicellular, single, rarely in chains of 2-3 spores.

smooth, spherical and 10-20 um diam., ovoid and 13.0-16.5 x 10.2-16.5 рт or pyrilorm

and 16.5-19.5 x 11.0-14.0 um. Chlamydospores intercalary with dimensions and colora-

tion identical to those of the aleuriospores.

Humicola nigrescens var. thermorongeura Subrahmanyam was isolated from

dung of Ratus sp. at Maharashtran, India. The provided protologue stress the presen sof

aleuriospores produced singly or in chains, globose (8.5-14.5 um) or ovoid (11.0-20.0 x

10.0-12.0 um) with similar intercalary chlamydospores. This description matches Humi-

cola grisea var. thermoidea sensu Cooney & Emerson.

.. Scytalidium allahabadum Narain et al. developed while examining municipal

refuse in the Allahabad region, India. It was first identified by P. M. Kirk (IMI) аз

Scytalidium thermophilum (Narain et al., 1983). However the Indian authors stress their

strain deviates on account of its colonies colored greyish-black coupled with the produc-

tion of larger spores with shapes commonly other than globose. In Scytalidium allahaba-

ое spores vary from 4.5-12.0 um while those of other shapes measure 4.5-27.5 X

.3-11.0 шп (10.0-12.5 x 7.5-10.0 jum for an Indian strain of Scyralidium thermophilum.

Re-examination of the type material (IMI 243118) confirm it represents a strain 9

Seytalidium thermophilum.

Source : MNHN Paris

THERMOPHILIC FUNGI 49

- Thermophymatospora fibuligera Udagawa, Awao & Abdullah — Myco-

taxon 37: 100-101. 1986.

Thermophymatospora Udagawa et al. (1986) was proposed for an unusual soil-

borne hyphomycete assignable to a basidiomycete anamorph. The original strain of the

type species T. fibuligera derives from an Iraqi date palm plantation. It is characterized by

holoblastic unicellular conidia being terminal or lateral, large, brownish, globose, thick-

walled and tuberculate, 20-25 um wide. Such conidia are superficially reminiscent of some

Myceliophthora species. However the hyphae of this hyphomycete are regularly provided

with simple clamp connections at the tranvserse septa. No link with a particular teleo-

morph has yet been established.

The fungus growth and sporulation are optimal around 35-40? C, almost nil at

20* C, with maximum being at 45? C.

Thermomyces ibadensis Apinis & Eggins — Transactions of the British

mycological Society 49: 631. 1966.

This hyphomycete was first recorded during studies of micro-organisms respon-

sible for the biodeterioration of palm kernels in Nigeria. The minimum temperature for

growth is between 31-35? C; optimum lies around 42-47? C and maximum at 60-619 C.

This Thermomyces differs from the type species T. lanuginosus by its smaller unicellular,

spherical, smooth, brown conidia, 4.0-8.0 um wide, and by its slender and more frequently

branched conidiophores.

~ Thermomyces lanuginosus P. Tsiklinsky (sensu Miehe 1907) — Annales

de l'Institut Pasteur, Paris 13: 500-505. 1899.

= Sepedonium lanuginosum ('Miehe') Griffon & Maublanc — Bulletin de la Société

Mycologique de France 27: 70. 1911.

= Monotospora lanuginosa (Griffon & Maublanc) Mason — Mycological Papers 3: 59.

1933

= Humicola lanuginosa (Griffon & Maublanc) Bunce as ‘lanuginosus’ — Transactions of

the British mycological Society 44: 375. 1961

= Acremoniella sp. Rege — Annales of Applied Biology 14: 28. 1927; fide Mason, 1933.

= A. thermophila Curzi — Atti dell'Istituto botanico dell’ Universita di Pavia, Ser 4: 154.

1929; fide Mason, 1933.

= Humicola grisea Traaen var. indica Subrahmanyam — Current Science 49: 30. 1980.

(nom. inval., Art. 36.1) am j

- Humicola lanuginosa (Griffon & Maublanc) Bunce var. catenulata Morinaga in Mori-

naga, Kanda & Nomi — Journal of Fermentation Technology 64: 452. 1986.

Descriptions: Cooney & Emerson (1964); Barron (1968); Ellis M. B. (1971); Domsch et al.

(1980).

Thermomyces was introduced by Tsiklinsky (1 899) for one species, Hm lanugino-

sus, isolated from garden soil; the original isolate was however not maintained. Miehe

(1907) retained this binomial for an isolate from composted hay. Griffon & Maublanc

(1911) studied a culture identical with the strain figured by Miehe, but expressed doubts as

to whether the corresponding hyphomycete would be conspecific with the fungus pro-

posed by Tsiklinsky. They argued the protologue was insufficient for a definite conclusion

Source : MNHN, Paris

50 J. MOUCHACCA

since "from published informations, the fungus examined by Tsiklinsky would have

conidia definitely smaller than indicated by Miehe". Griffon & Maublanc then assigned

their isolate to Sepedonium Link on account of the slightly verrucose nature of the

conidial wall.

In 1933, Mason examined a culture of Acremoniella thermophila Curzi, *kindly

supplied by Mr Curzi”; he noticed the similarity with Sepedonium lanuginosum and also

with Acremoniella sp. Rege. As Mason had concluded before that Acremoniella Sacc. was

a synonym of the earlier Monotospora Corda (non Monotospora Vuill.), he proposed to

rename the fungus of Griffon & Maublanc Monotospora lanuginosa. No mention of the

binomial Thermomyces lanuginosus was made by Mason. In the meantime, Curzi (1930)

published an extensive cultural study of his Acremoniella thermophila, a fungus he had

previously submitted to Griffon & Maublanc for examination (fide Mason, 1933).

While describing Thermomyces stellatus (= Humicola stellata), Bunce (1961)

questioned the maintenance of the Griffon & Maublanc fungus in Monotospora Corda

since [and as also stressed by Mason: 1933, 1941], the concept of this genus was still under

debate. Bunce rather favoured the transfer of Monotospora lanuginosa to Humicola

Traaen, established for mesophilic hyphomycetes sharing the same type of aleuriospores.

Cooney & Emerson (1964) followed Bunce proposal. On the other hand LaTouche (1950)

who had isolated this fungus from compost, considered the binomial Thermomyces

lanuginosus. The latter name was also retained by Apinis (1963) on the all his isolates

from alluvial soils agreed with the original description provided by Tsiklinsky (1899).

The status of Thermomyces was finally definitively settled by Pugh et al. (1964)

while describing the mesophilic T. verrucosus; in the latter a transverse septum is present

just below the conidiophore apex delimiting a small apical cell. According to these

authors, such a feature is evident from Tsiklinsky's photomicrographs and this provides

arguments that her isolate is identical to the fungus now known as Thermomyces lanugi-

nosus. Pugh er al. then provided a description and drawings for the latter based on the

neotype strain IMI 84400 (= ATCC 22070), isolated by Bunce from mouldy hay at the

Rothamsted Experimental Station in 1959.

In the same year 1964, Cooney & Emerson in their treatment of thermophilic

fungi underlined their first isolate of Humicola ( Thermomyces) lanuginosus was strain

No. 20 obtained in 1945 by D. G. Cooney from retting guayule shrub. This strain was later

on numbered M 206522 at the University of California Herbarium, Berkeley (= ATCC

16455 — CBS 632.91). It provided the material for the description and drawings reported

in their monograph. The isolate selected by Pugh et al. (1964) was explicitely designated

“neotype”; it has to be regarded as such against M 206522.

Subsequent taxonomic treatments of hyphomycetes uniformly accepted Ther-

momyces (Carmichael et al., 1980). An ultrastructural study of the conidial ontogeny of its

type species was conducted by Ellis D. H. (1981). Further, recently Straatsma and Samson

(1993) re-examined isolate CBS 153.75 (= ATCC 28402) belonging to the unpublished

Humicola brevis (Gilman & Abbott) Gilman var. thermoidea Subrahmanyam; this was

re-identified as Thermomyces lanuginosus. They also concluded the same for the similar

unpublished taxon Humicola brevispora Subrahmanyam & Thirumalachar based on CBS

152.75 (2 ATCC 28403).

Humicola grisea var. indica (Subrahmanyam, 1983) was obtained as a laboratory

contaminant at Pimpri, Poona. Trials to locate the representative strain were unsuccessful.

According to the author, a critical study revealed “it belonged" to Humicola grisea (which

produces intercalary chlamydospores) and approximates its var. thermoidea (which also

produces intercalary chlamydospores). However, the general lay-out of the drawings and

Source : MNHN, Paris

THERMOPHILIC FUNGI 51

features underlined in the description clearly stress the proposed variety represents

Thermomyces lanuginosus. The only deviation is the smooth character of the conidia in the

proposed variety against the wrinkled condial sur! of Thermomyces lanuginosus.

Humicola lanuginosa var. catenulata (Morinaga et al., 1986) was obtained in the

course of a survey of soil-borne thermophiles for high producer strains of lipase enzymes.

Morphological details underlined in the publication clearly indicate it represents a deviant

strain of Thermomyces lanuginosus.

The mesophilic Thermomyces verrucosus Pugh, Blakeman & Morgan-Jones

(1964) displays no growth above 37? C. It has globose, dark brown conidia with conspi-

cuously warted surfaces, 10-17 um wide. These structures are definitely larger than aleu-

riospores of the type species.

— Thermomyces stellatus (Bunce) Apinis — Nova Hedwigia 5: 75. 1963.

basionym: Humicola stellata Bunce as 'stellatus' — Transactions of the British mycologi-

cal Society 44: 372. 1961.

Descriptions: Bunce (1961); Apinis (1963); Ellis M. B. (1971).

The original material was isolated from mouldy hay in England and Wales. The

fungus develops optimally at 40° C with growth being very slow at 24? C and not

extending above 50? C. Conidia of the aleuriospore type, angular, lobed, smooth, pale to

mid brown or greyish brown, 5-10 x 5-9 um. The transfer to Thermomyces is based on

account of the absence of phialospores in culture and similarity in conidiogenesis.

THERMOPHILIC MYCELIA STERILIA

— Myriococcum thermophilum (Fergus) van der Aa — Verhandelingen

Koninklijke Nederlandse Akademie van Wetenschappen A fd. Natuurkunde,

seies II, 61: 60. 1973.

basionym: Papulaspora thermophila Fergus — Mycologia 63: 426. 1971.

Descriptions: Fergus (1971); van der Aa (1973).

This *bulbil-producing fungus’ was described from mushroom compost in Swit-

zerland (Fergus, 1971). In vitro, such structures appear very rapidly at 45° C in the aerial

and submerged mycelium. They are white at first, then yellow and finally orange at

maturity; in mature bulbils, cells of the outer layers are narrower and more elongate than

corresponding internal more intensely colored globose cells. The fungus shows no growth

at 28° C and 53° C with the optimum being at 45° C. No connection with a perfect stage

yet established.

From seedlings of a Begonia species heavily infected with small sclerotia, van der

Aa (1973) isolated a fungus matching the description of the type of Myriococcum Fr.,

M. praecox Fr. Subsequent comparison with representatives strains of Papulaspora bys-

sina Hotson confirmed similarity of both taxa. Also examination of the type of Papulas-

pora thermophila Fergus proved it to be congeneric with Myriococcum praecox except for

Source : MNHN, Paris

32 J. MOUCHACCA

its thermophilic character. As Myriococcum predates Papulaspora Preuss, van der Aa

transferred Fergus fungus to the former generic entity.

The term bulbil is now restricted to homogeneous pseudoparenchymatous

bodies occurring only in the basidiomycetous genera Burgoa Goidanich and Minimedusa

Weresub & LeClair. The term papulaspore is applied to thallodic propagules differentiated

from the inception into central and sheathing cells (Weresub & LeClair, 1971). Such

thallodic propagules occur amongst the mycelia of some species of Melanospora Corda

and few probably related ascomycetes.

THERMOTOLERANT BASIDIOMYCETE

— Phanerochaete chrysosporium Burdsall apud Burdsall & Eslyn — Myco-

taxon 1: 124. 1974.

anamorph: Sporotrichum pruinosum Gilman & Abbott — Iowa State College Journal of

Science 1: 306. 1927.

= Chrysosporium pruinosum (Gilman & Abbott) Carmichael — Canadian Journal of

Botany 40: 1166. 1962.

= Emmonsia brasiliensis Batista et al. — Revista Facultad Medecina, Universidad de

Ceara (Brazil) 3: 52. 1963.

= Sporotrichum dehradunense Sarbhoy & Saksena — Sydowia, Annales Mycologici, Ser.

П, 19: 198. 1966 (“1965”).

= Chrysosporium lignorum Bergman & Nilsson — Department of Forestry, Proceedings

of the Royal College of Forestry, Stockholm, Research Notes, 53: 28. 1966. (nom. inval.,

Art. 36.1).

= Sporotrichum pulverulentum Novobranova — Novosti sistematiki nizshikh rastenii 9:

184. 1972.

Description: Stalpers (1984).

Sporotrichum pruinosum (also as S. pulverulentum and Phanerochaete chrysospo-

rium) is a thermotolerant hyphomycete that has become the subject of many recent

physiological studies. The fungus is known to produce three types of hydrolytic enzymes

active in the degradation of cellulose. It is actually used as a model for the biodegradation

of lignin and the production of protein from lignocellulosic waste material, a process

designated single-cell protein (Stalpers, 1984). For these reasons, it is included in this study.

The protologue of the anamorph is based on a strain isolated from soil. The

fungus was later on transferred to Chrysosporium Corda on account of the confusion

surrounding the generic concept of Sporotrichum Link (Stalpers, 1978). As for the

teleomorph, the first specimen was collected in the Sonoran Desert, Arizona (USA); when

cultured, it produced a Chrysosporium state matching Sporotrichum pruinosum. Later on

the teleomorph could be obtained in vitro under particular cultural conditions. Reported

cardinal temperatures are: minimum 7° C, optimum 36-40? C, maximum 46-49? С.

In culture Sporotrichum pruinosum is the most variable species of the genus and

such accounts for the several published synonymies. The similarity with S. pulverulentum

has been the matter of a long debate (Burdsall, 1981) but recent studies provided

Source : MNHN, Paris

THERMOPHILIC FUNGI 53

arguments in favour of such a synonymy (Stalpers, 1984). Citation in applied work of the

teleomorphic name is favoured against the two commonly cited anamorphic binomials

since several nomenclatural problems are still connected with the latter. The fungus has

also been reported as a human pathogen being isolated from lungs and this explain its

inclusion in the genus Emmonsia Ciferri & Montemartini.

TAXA OF UNCERTAIN POSITION

— Mucor thermo-hyalospora Subrahmanyam — Bibliotheca Mycologica

91: 421. 1983. (nom. inval., Art. 37.1).

The examined strain was isolated from contaminated curd collected in the local

market at Pimpri, Poona, India. The fungus is clearly thermophilic with growth starting at

24? C, being optimum at 45? C and maximum at 55? C. According to the author "careful

study of monosporic cultures showed that is closely resembled Mucor tauricus Milko &

Schkur but differs from it in being homothallic. Therefore it is described here as a new

species".

The presence of weakly developed rhizoids in Mucor tauricus, accounted for its

transfer to Rhizomucor by Schipper (1978). This information was however overlooked by

Subrahmanyam (1983) who also provides no details concerning the presence or the

absence of corresponding structures in his zygomycete. This taxon might simply represent

a deviant strain of Rhizomucor pusillus.

— Stilbella thermophila Fergus — Mycologia 56: 277. 1964.

This synnematous hyphomycete was first isolated from mushroom compost in

Switzerland. Optimum growth is between 35-50? C; at 55? C, slight development still

occurs but such is not the case below 25? C. In culture, the fungus produces white

synnemata, up to 300 um high, bearing whitish glistening mucoid conidial heads; conidia

are hyaline, continuous, oblong-ellipsoid, 15-17 x 6-10 um.

Seifert (1985) in his monographic treatment of Stilbella Lindau re-examined

authentic material. Conidia were observed to develop from percurrently proliferating

conidiogenous cells, i.e. annellophores, a feature enhancing its exclusion from the genus.

Additionnal work is undertaken to establish the correct taxonomic position of this species

(Seifert, pers. comm.).

CONFUSING BINOMIALS

— Achaetomium thermophilum Basu — Current Science 51: 524. 1982.

The original living strain was isolated from leaf litter at Bhattni, Uttar Pradesh,

India. It was described as being similar to Achaetomium macrosporum Rai, Wadhami €

Tewari but differ by being “thermophilic” in nature, although no minimum growth

temperature had been indicated.

Source : MNHN, Paris

54 J. MOUCHACCA

Cannon (1986) examined a culture (IMI 292262) derived from the holotype.

Growth and sporulation proved to be satisfactorily at 25? C indicating the fungus is rather

thermotolerant. Ascospores produced were also found to be uniporate rather than bipo-

rate as stressed in the protologue and thus matching those of Achaetomium macrosporum.

Based on these observations, Cannon concluded the ascomycete proposed by Basu is

conspecific with Achaetomium macrosporum.

Species of Achaetomium Rai & Tewari are known to be good thermotolerants

(von Arx et al., 1988). The concept of the genus is however still under debate. Thus von

Arx et al. (1988) excluded A. macrosporum; they also suggested the latter is rather similar

to Chaetomium vitellinum Carter or Ch. megasporum Sórgel.

— Calcarisporiella thermophila (Evans) de Hoog — Studies in Mycology 7:

68. 1974.

basionym: Calcarisporium thermophile Evans — Transactions of the British mycological

Society 57: 247. 1971.

This is the type species of the mucedinaceous genus Calcarisporiella de Hoog

(de Hoog, 1974). The original living culture was isolated from coal spoil tips at Staf-

fordshire, England. According to Evans (1971 a & b), the minimum growth temperature

value is 16° C, optimum at 40° C and maximum at 50? C. The fungus should thus be

considered a thermotolerant.

— Endoblastomyces thermophilus Odinzowa — Microbiology, Moscow 16:

273. 1947 (descripion only); Die Systematik der Hefen: ?. 1960 (latin

diagnosis but no type designated); (nor. inval., Arts. 36.1 & 37.1).

This is the type species of Endoblastomyces Odinzowa. The original protolgue

was not accompanied by a latin diagnosis provided later on by Ozindowa in Kudryavtzev's

book “Die Systematik der Hefen", the german translation of which was published in

Berlin in 1960; however, Odinzowa then omitted to designate a holotype.

This yeast was isolated from brewing wort inoculated with baker's yeast in a

bread factory in Central Asia, USSR; it was considered to represent a new thermophilic

taxon. Carmo-Sousa (1970) was unable to locate the corresponding living strain and

according to him, the original description strongly suggests similarity with Trichosporon

capitatum Diddens & Lodder. This was substantiated by the arrangement of the pseudo-

mycelium, endoblastospores formation and maximum temperature of growth being iden-

tical in both taxa.

Trichosporon capitatum is not thermophilic in the sense of Cooney & Emerson

beingable to develop below 20? C with a maximum at 44-46? C (Carmo-Sousa, 1970). The

fungus was later on relocated in Geotrichum Link:Fr. and its perfect state discovered by de

Hoog et al. (1980).

— Geotrichum candidum Link var. thermoideum Qureschi € Mirza —

Biologia, Lahore 27: 144. 1981.

The original material was isolated from camel dung in Pakistan. The fungus was

regarded by van Oorschot $: de Hoog (1984) as a possible synonym of Arthrographis

sulfurea (Grev.:Fr.) Stalpers & van Oorschot, a mesophilic hyphomycete.

Source : MNHN, Paris

THERMOPHILIC FUNGI 55

— Gilmaniella thermophila Qureschi & Mirza — Biologia, Lahore 29: 341.

1983.

The original material developed on goat dung collected in Pakistan. The species

was overlooked by Sivanesan and Sutton (1985) while describing Gilmaniella punctiformis

and also by Moustafa and Ezz-Eldin (1989) during their recent addition of G. multiporosa,

isolated from Egyptian soils in North Sinai. These additions brings to five the number of

known species.

Gilmaniella thermophila might be a later name of G. macrospora Moustafa; the

final decision awaits comparison of authentic material. The latter was first encountered

while investigating the mycoflora of salt-marsh soils of Kuwait. It was also subsequently

recovered, although infrequently, from Iraqi soils analysed by Abdullah & Al-Bader

(1990). The Iraqi strains developed optimal growth at 40° C with a maximum between

45-50” C thus confirming the thermotolerance abilities of Gilmaniella macrospora.

The specific epithet refers to globose condia being larger than conidia of Gilmaniella

humicola, the type species: 14-18 um versus 7-10 um for the latter.

— Lagenidium thermophilum Nakamura, M. Nakamura, Hatai & Zafran

— Mycoscience 36: 400. 1996.

The specific epithet coined for this newly described Oomycete is misleading. The

fungus was found to infect the eggs and larvae of the mangrove crab, Scylla serrata

Forsskal, in Bali, Indonesia. Isolated strains proved to represent a new species of Lageni-

dium Schenk having a unique discharge process. Growth range is from 15-45? C with the

optimum being between 30-40? C. This taxon is thus a fast growing thermotolerant

fungus.

— Melanomphalia thermophila (Singer) Singer — Atas, Instituto de Mico-

logia, Universidade de Recife 5: 482. 1963.

basionym: Tubaria thermophila Singer — Papers of the Michigan Academy of Sciences,

Arts and Letters 32: 145. 1948.

The type specimen of this basidiomycete was collected by the author in the state

of Florida (USA) at Highland Hammock State Park (Singer F 20, F 20a, FH). The habitat

in which the carpophore developed was specified as “In dumetis subtropicalibus humidis-

simus in terra humosa sabulosa vel nonnumquam nucibus. Caryae megacarpae afxa vel e

ligno mucido ecrescentes, aestate". The reason underlying the selection of the epithet

thermophila seems to have simply been suggested by the very warm to hot humid weather

prevailing in this southern state of the United States. The fungus is not a thermophile in

the Cooney & Emerson's sense.

Similar cases concern Russula roseipes (Secr.) Sacc. subsp. thermophila Singer,

collected under Pinus taeda in North Florida, and Suillus hirtellus (Peck) Kuntze var.

thermophilus (Singer) Smith & Thiers (Singer, 1975).

Source : MNHN, Paris

56 J. MOUCHACCA

— Mucor thermophilus Prakash & Sarbhoy — Zentralblatt für Mikrobio-

logie 148: 531. 1993.

The specific epithet coined for this recently described zygomycete is misleading

since “the species is able to grow and sporulate at 30? C and above 30° C" (Prakash &

Sarbhoy, 1993); however, the minimum and maximum growth temperature values were

not ascertained. Further the statement that "the specific epithet has been given on

thermotolerant nature of the species" clearly indicates the fungus is not a thermophile in

the Cooney & Emerson's sense.

— Paecilomyces puntonii (Vuillemin) Nannfeldt sensu Eicker (1972):

The correct binomial for this hyphomycete is Paecilomyces puntonii (Vuillemn)

Nannizi (Samson, 1974).

Eicker (1972) isolated a hyphomycete from the faeces of domestic fowls in South

Africa strain UP 71 T (University of Pretoria) he identified as Paecilomyces puntonii. This

isolate “did not grow at 20° C, neither at 30° C but good growth took place at 50? C. No

perfect state developed on any of the cultures media at the various temperatures of

incubation used". Paecilomyces puntonii is a mesophilic fungus with optimum growth

being at 25? C (Samson, 1974).

— Sporotrichum cellulophilum:

Durand et al. (1984) clearly specify this binomial correspond to a thermophilic

fungus. Its ability to produce interesting enzymes of the cellulases and hemicellulases

types were largely investigated by several workers (Kinoshito et al., 1986). However,

Stalpers in his 1984 revision of Sporotrichum makes no mention of this binomial in the

check-list of epithets used in combination with the genus. Also publication of this

binomial after this date following standard taxonomic rules could not be traced. It is thus

evident that Sporotrichum cellulophilum has no taxonomic status.

With regard to literature on thermophilic fungi, the generic epithet Sporotrichum

was first introduced by Apinis (1963) for a hyphomycete which ultimately will be renamed

Myceliophthora thermophila; this will also prove to be the anamorph of Corynascus

heterothallicus. A second Sporotrichum made its appearance in the last decades in papers

delaing with biotechnological work: Sporotrichum pruinosum, anamorph of the basidio-

mycete Phanerochaete chrysosporium. This thermotolerant hyphomycete was described

also under several Sporotrichum names (see comment under Phanerochaete chrysospo-

rium). It is sometimes erroneously indicated as being thermophilic (Deshpande et al.,

1978).

The common use of *ghost binomials" in publications dealing with applied

studies involving fungi is a source of serious confusion. A similar case is forwarded by the

binomial Acremonium cellulophilum (Satyanarayana et al., 1992). Such a practice should

be definitely prohibited.

— Sordaria thermophila Fields — Mycologia 60: 1117. 1968.

The original strain of this ascomycete developed on cow dung collected in Texas

(USA) and incubated in a moist chamber. According to the protologue “the specific epithet

refers toa high temperature requirement for ascospore germination. Ascospores of thenew

species germinated less than 1 on media containing sodium acetate. With an additional

treatment of 40-45? C fora period of 8-12 h, germination was increased to 40 94".

Source : MNHN, Paris

THERMOPHILIC FUNGI 57

In the published description no data is reported on the in vitro linear variation of

growth with temperature. Also the conditions at which moist chambers were incubated are

not specified (Fields, 1968). The thermophilic nature of this taxon thus cannnot be

ascertained. The selected specific epithet seems to relate to the heat treatment applied to

enhance ascospore germination.

Guarro & von Arx (1987) regarded this heterothallic relative of Sordaria fimicola

(Rob.) Ces. & de Not. as representing a good species. Further investigations are however

required to underline the biological and taxonomical characteristics of this ascomycete

which apparently has not been reported since its description.

— Zalerion thermophylii Udaiyan — Journal of Economic and Taxonomic

Botany 15: 664. 1991 (1992); (nom. inval., Art. 37.1).

The original material developed on beech wood test blocks immersed in the

cooling tower and the collecting lagoon of a hydroelectric plant at Tamil Nadu; holotype

was not indicated.

The dematiaceous hyphomycete genus Zalerion Moore & Meyers was esta-

blished for a widely distributed mesophilic fungus trapped on wood blocks immersed in

sea-water, Z. maritima (Linder) Anastasiou., described before under several names (Ellis

M. B., 1976). Zalerion thermophylii is most probably identical to the type species. The

epithet thermophylii must have been suggested by the high temperature of the water

circulating in the cooling tower.

DISCUSSION

Thermophilic fungi dealt with in this contribution are found to form a small

group of less than forty species and varieties. Growth at high temperatures is thus

definitely a rare feature among fungi. Also several of these taxa were described in recent

years. This ecological group is thus expected to expand rapidly in the near future; in

particular if some credit is awarded to the estimate amounting the number of existing

species to one million and half. A major emphasis for this trend is also embodied by the

outcome of taxonomic work conducted in the last decades. Such achievements have

provided adequate answers for long standing problems. A limited additionnal work of this

type is still necessary to solve remaining minor ones.

Taxa treated here are considered as strict thermophiles based on the definition of

thermophilism provided by Cooney & Emerson (1964). However the use of this simple

classificatory system to segregate between thermophilic and thermotolerants is sometimes

difficult to apply; this is particularly critical at the lower temperature threshold of 20? C.

Thus following Bokhary ег al. (1984), the well established thermophile Melanocarpus

albomyces should be regarded a thermotolerant being able to grow below 20° C. It is

possible the response of different strains of the same taxon accounts for such deviations.

Further difficulties in defining true thermophiles results from the absence of reliable

growth curves covering a wide range of temperatures for most taxa proposed as such. This

basic simple type of data is needed to ascertain the true nature of few members of this

group.

Based on available informations, the ability to only develop at high temperatures

is disclosed by few Mucorales, Eurotiales and Sphaeriales (sensu von Arx, 1988) and by

Source : MNHN, Paris

58 J. MOUCHACCA

several Hyphomycetes. No coelomycete and no basidiomcyete was found to be thermo-

philic. Further the teleomorph of the sole thermophilic agonomycete, Myriococcum

themophilum, is hypothesized not to belong to Eurotiales on the assumption it should have

made its appearance since the fungus was described. However observations relating to

mating experiments of this sterile fungus are uncommon in the literature.

The group of five thermophilic Mucorales comprise the still monospecific

Thermomucor and several Rhizomucors including the type; the former differs mainly by

having smooth zygospores, a character uncommon in the Mucoraceae. Also allthough

regular production of zygospores by Rhizomucor miehei should prevent confusion with.

Rh. pusillus, the type species, the ecology of each taxon is not yet clearly understood.

Further, the validity of both Rhizomucor tauricus and Rh. nainitalensis is questioned.

The group of ascomycetous fungi brings together twenty species and three

varieties; these relate to only nine genera. Following von Arx (1987, 1988), Dactylomyces

(inclusive of Thermoascus) and Talaromyces belong to Family Onygenaceae, Order Euro-

tiales; as Coonemeria was established for taxa previously assigned i in Dactylomyces and

Thermoascus, then the new genus should also be accomodated in this family. The rema

ning six genera are representatives of Families Chaetomiaceae (Chaetomium), Microasca-

ceae (Canariomyces) and Thielaviaceae (Corynascus, Melanocarpus and Thielavia).

Dactylomyces appears monospecific. Canariomyces and Thermoascus have one

species each plus one variety for the latter; new informations about the variety might lead

to the proposal of a specific rank. Corynascus and Melanocarpus have two species each;

this number becomes three in the case of Coonemeria, Talaromyces and Thielavia. Chae-

tomium is represented by four species and two varieties but Ch. britannicum might prove

not to be a true thermophile when a living culture becomes available. Also definite

taxonomic decisions about Chaetomium thermophilum, its varieties and Ch. virginicum

might reduce this group to only two accepted species; the genus would thus comprise only

three thermophiles.

Not all thermophilic ascomycetes have an associated anamorphic state; also

among these conidial states some do not develop concomitantly with the corresponding

teleomorph. Canariomyces thermophila has no anamorph although the type was described

with a catenate conidial state. Thermophilic Chaetomia do not develop conidia of any

kind. Thermoascus can be regarded as not having an anamorph producing catenate

conidia. Dactylomyces has a distinctive but yet unnamed Polypaecilum anamorph; the

fungus remains not satisfactorily documented probably due to the taxonomic confusion

with Thermoascus that prevailed. Polypaecilum anamorphs also characterise Dichotomo-

myces Saito:Saito having 2 -3 described species (von Arx, 1981). The genus also has

cleistothecia with a wall of textura angularis type but asci are produced in chains; it also

belong to Family Eurotiaceae sensu von Arx (1987). Melanocarpus approximates Cana-

riomyces since respective type species have distinctive conidial states but that of the latter

is a mesophile. The former genus has now four taxa with the second thermophile,

M. thermophilus, not developing the characteristic arthroconidial state of the type; the

same situation is disclosed by the two other members of the genus which do not develop at

high temperatures.

Regarding the genus Thielavia, Th. pingtungia has no conidial state, a feature

characteristic of all known Chaetomidia. Th. australiensis was reported with an ana-

morph of the Trichosporiella type; this is developed by other members of the genus

(Mouchacca, 1973). The fungus is however badly documented being known only from the

protologue. Thielavia terrestris is associated with a distinctive anamorphic state described

exclusive of the teleomorph, Acremonium alabamense; due to the “homothallic with

Source : MNHN, Paris

THERMOPHILIC FUNGI 59

cross-feeding" nature of the perfect state, the anamorph could be encountered alone in

studies involving high incubation temperatures. Thielavia is still admitted to represent a

heterogeneous entity due to lack of informations about the behaviour in culture of

Thielavia basicola (type species) and production of hyaline and dark coloured colonies by

known members.

On the other hand, thermophilic Talaromyces all develop a conidial state; these

belong either to Paecilomyces (T. byssochlamydioides) or to Penicillium (T. emersonii and

T. thermophilus). Regarding Paecilomyces, von Arx (1987) suggested it be expanded to

include Penicillia of Sections Biverticillata and Sagenomella known as anamorphs of

genera he grouped in Family Onygenaceae; such a proposal was made to increase the

degree of homogeneity among genera. A similar situation is disclosed by Coonemeria and

Corynascus; all three taxa of the former have a well developed Paecilomyces state while

both species of the latter have anamorphs now correctly assigned to Myceliophthora. Taxa

of Coonemeria and Corynascus had very complicated respective taxonomic histories either

due to cases of misidentification (species now placed in Coonemeria) or to the heterothallic

nature of the Corynascus perfect states. For the latter, it follows that either Myceliophthora

could develop singly in studies conducted on thermophile habitats with only appropriate

mating leading to ascospore formation. The anamorph of Corynascus heterothallicus was

proposed prior to the discovery of the teleomorph while the reverse is true for

C. thermophilus.

The group of thermophilic hyphomycetes comprises thirteen species although

for Scytalidium thermophilum, the term species appears inadequate in the present situa-

tion. These fungi belong to seven genera: Acremonium, Malbranchea, Myceliophthora and

Thermophymatospora are mucedinaceous entities, while dematiaceous thermophiles

belong to Humicola, Scytalidium and Thermomyces.

Among mucedinaceous taxa, three are established anamorphs of "almost hete-

rothallic to heterothallic” ascomycetes and thus could be observed alone in studies

involving a self-heating process. Acremonium alabamense is the conidial state of Thielavia

terrestris whose mating behaviour is not yet clearly understood. This is not a typical

Acremonium species and such accounts for its inclusion in a new section with phialidic

states of some Chaetomia. Such is not the case for Acremonium thermophilum, a not yet

well documented thermophile. The perfect state of Myceliophthora fergusii is Corynascus

thermophilus and care should be taken to avoid confusion with the teleomorph of

M. thermophila, C. heterothallicus; the third member M. hinnulea has not yet developed a

perfect state, a situation analogous to the mesophilic type species of the genus. Malbran-

chea cinnamomea is a very distinctive colored arthroconidial fungus actually displaying a

wide distribution; it is the sole thermophile of a genus known to comprise mesophiles

associated with well defined teleomorphs (von Arx, 1987). Thermophymatospora fibuligera

is unique with its septal clamp connections and an aleuriosporic state; this peculiar fungus

is apparently still known only from the type locality.

The remaining dematiaceous thermophiles were assigned to Humicola, Scytali-

dium and Thermomyces. But only the taxonomic status of the latter is now the subject of

a large consensus. Thermomyces lanuginosus is the first assessed thermophilic fungus.

His complex nomenclatural history has involved genera as Acremoniella, Humicola,

Monotospora and Sepedonium. The definite re-instatement of Thermomyces by Pugh et al.

(1964) clarified its links with thermophilic species of Humicola. The genus now also

comprises Th. ibadensis, Th. stellatus and the mesophilic Th. verrucosus. However, only the

type species is by far the most reported one.

Source : MNHN, Paris

60 J. MOUCHACCA

The status of Humicola hyalothermophila needs to be re-assessed in conjunction

with that of both Humicola recently proposed as synonyms of Scytalidum thermophilum

(Straatsma & Samson, 1993). The introduction of Scytalidium in an attempt to relocate

Torula thermophila added much confusion as the transfer was not substantiated by valid

taxonomic arguments. This combination was however immediately reported by Ellis M. B.

(1976). Scytalidum lignicola, type species of the genus, is a mesophile producing cultures

with scanty aerial mycelium. The fungus develops conidiogenous cells of two kinds:

hyaline fertile hyphae become septate, later producing thin-walled arthroconidia by

fragmentation; brown fertile hyphae forming chains of brown arthroconidia; also chains

of brown aleuriospores could be observed (Ellis M. B., 1976). These intercalary conidia

develop by transformation of pre-existing normal hyphal cells. The presence of solitary

conidia terminal or lateral was never reported. In Scytalidum indonesicum chlamydospore

formation and disarticulation follow the same pattern but no hyaline arthroconidia

develops. Here too, terminal or lateral solitary conidia were not reported (Hedger er al.,

1982).

In species of Humicola, solitary terminal and more commonly lateral aleurios-

pores usually develop in addition to intercalary morphlogically similar ones. Single

terminal aleuriospores (and less often lateral ones) may become intercalary by hyphal

extension of their tip. Also chains of aerial or immersed “aleuriospores” do not disarti-

culate to liberate individual elements but such is achieved by lysis of sustaining hyphal

cells. Further, no hyaline arthroconidia are produced by any described member of the

genus but mesophilc taxa rather produces hyaline phialospores. Straatsma and Samson

(1993) compared a large number of strains assigned to Scytalidum themophilum, Torula

thermophila or to both Humicola now regarded as synonyms of the former. They under-

lined two extreme types could be recognized; the first having simple very dark spores borne

on short lateral branches matching the description of Humicola grisea var. thermoidea; the

second type develop intercalary slightly pigmented spores in chains, representatives of

Scytalidium thermophilum or more appropriately of its basionym Torula thermophila.

Within the two types however, some isolates also develop short terminal chains of conidia

making them intermediate between types | and 2.

Straatsma and Samson (1993) stress such intermediate isolates favours not the

segregation of taxa "on the basis of the single character of conidia in the aerial myce-

lium"; these rather support grouping of all types under one binomial whose placement in

Scytalidium is to be reconsidered. As the particular mode of chlamydospore formation in

this genus deviates from the pattern depicting species of Humicola, the exclusion of the

above complex from the former is more than justified. Scytalidium is now regarded a

heterogenous entity due to addition of species only developing dematiaceous "arthroco-

nidia". Nevertheless, extension of Scytalidium characteristics for the understanding of

Humicola species has shadowed features proper to the latter preventing sound taxonomic

separation among its members.

From an ecological point of view, the equivocal application of the now widely

accepted (inspite of its limitations) definitions of Cooney & Emerson (1964) lead to

consider well established thermotolerants as thermophiles. Ellis D. H. (1981) regards all

Rhizopus able to grow at 45? C as thermophilic although they display growth below 20° C.

These zygomycetes and some other true mesophilic fungi are also currently considered as

thermophiles in publications focusing on biotechnological problems (Satyanarayana ef

al., 1992). Several authors also classify as thermophile all fungi developing in isolation

plates incubated at 45? C (Abdel-Fattah er al., 1977; Moubasher et al., 1988).

Source : MNHN, Paris

THERMOPHILIC FUNGI 61

Another type of misleading situations relates to epithets selected while descri-

bing a new taxon found to develop at elevated temperatures. The recent Mucor thermo-

philus is a good example among others here considered; from the protologue it is evident

this Mucor should be regarded as a thermotolerant. A definitely critical situation is

examplified by the frequent use in studies involving fungal enzymes of ghost binomials

having no taxonmic status of any kind as Sporotrichum cellulophilum; such a practice

needs to be totally banished for the confusion it introduces; in particular while attempting

to analyse published data (Satyanarayana et al., 1992; Schekkar & Johri, 1992).

Strict restriction to nomenclatural rules governing citations of fungal binomes is

fundamental. Authors of applied research dealing with thermophiles should necessarily

follow such regulations in order to stabilize names used in produced articles. This would

bring an end to the chaotic state prevailing especially in publications relating to fungal

ecology and biotechnology. The taxonomic and nomenclatural reappraisal of known

thermophilic taxa here undertaken will definitely unravel informations already lable.

This should enable a sound synthesis of published data and foster the discovery of new

elements of this interesting physiological group of fungi.

ACKNOWLEDGEMENTS

Sincere appreciations are extended to the several colleagues who provided copies of hardly

available publications, subcultures or informations on some published names. Special

thanks are due to Profs. G. L. BENNY and R. H. PETERSEN and to Drs. L. ZOFIA,

V. MEL’NIK, P. M. KIRK and J. STALPERS.

REFERENCES

AA H. A. van der, 1973 — In “Centraalbureau voor Schimmelcultures Baarn and Delft Progress

Report 1972, J. A. von Arx". Verhandelingen der Koninklijke Nederlandse Akademie van

Wetenschappen Afd. Natuurkunde, Tweede Reeks 61(4): 60-81.

ABDEL-FATTAH Н. M., MOUBASHER А. Н. & ABDEL-HAFEZ S. L, 1977 — Studies on mycoflora of

salt-marshes in Egypt. III. Thermophilic fungi. Bulletin of the faculty of science, Assiut

university 6: 225-235.

ABDULLAH S. К. & AL-BADER S. M., 1990 — On the thermophilic and thermotolerant mycoflora of

Iraq. Sydowia 42: 1-7.

Ames L. M., 1963 — A Monograph of the Chaetomiaceae. U. S. Army Research and Development,

Ser. 2, Reprint Cramer, Lehre, 1968, 125 pp.

APINIS A. E., 1963 — Occurrence of thermophilous microfungi in certain alluvial soils near Nottin-

gham. Nova hedwigia 5: 57-78.

Аюм A. E., 1967 — Dactylomyces and Thermoascus. Transactions of the british mycological society

30: 573-582.

Apinis A. E. & CHESTERS С. С. C., 1964 — Ascomycetes of some salt marshes and sand dunes.

Transactions of the british mycological society 47: 419-435.

Акх 1. A. von, 1970 — The genera of fungi sporulating in pure culture. J. Cramer, Lehre, 288 pp.

Source : MNHN, Paris

62 J. MOUCHACCA

ARX J. A. von, 1971 — Über die typusart, zwei neue und einige weitere arten der gattung Sporotri-

chum. Persoonia 6: 179-184.

ARX J. A. von, 1973 — Further observations on Sporotrichum and some similar fungi. Persoonia 7:

127-130.

ARX J. A. von, 1975 — On Thielavia and some similar genera of Ascomycetes. Studies in mycology 8:

1-29.

Акх 1. A. von, 1981 (1982) — On Monilia sitophila and some families of Ascomycetes. Sydowia 34:

12-29

Anx J. A. von, 1984 — Canariomyces notabilis, a peculiar ascomycete from the Canary Islands.

Persoonia 12: 185-187.

ARX J. A. von, 1987 — A re-evaluation of the Eurotiales. Persoonia 13: 273-300.

ARX J. A. von, FIGUERAS M. J. & GUARRO J., 1988 — Sordariaceous ascomycetes without ascospore

ejaculation. Beihefte zur nova hedwigia 94: 1-104.

ARX J. A. von, GUARRO J. & FIGUERAS M. J., 1986 — The Ascomycete genus Chaetomium. Beihefte zur

nova hedwigia 84: 1-162.

Ausrwick P. K. C., 1976 — Environmental aspects of Mortierella wolfii infection in cattle. New

Zealand journal of agricultural research 19: 25-33.

Awao Т. & OTSUKA S., 1974 — Notes on thermophilic fungi in Japan (3). Transactions of the

mycological society of Japan 15: 7-22.

BARRON G. L., 1968 — The Genera of Hyphomycetes from Soil. The Williams & Wilkins Co.,

Baltimore, 364 pp.

Basu M., 1984 — Myceliophthora indica is a new thermophilic species from India. Nova hedwigia 40:

85-90.

Benny G. L. & KIMBROUGH J. W., 1980 — A synopsis of the order and families of the Plectomycetes

with keys to genera. Mycotaxon 12: 1-91.

BiLAY V. T. & ZAKHARCHENKO, B. A., 1987 — Opredelitel’ Termofil'nykh Gribov. Kholodny Institute

of Botany, Ukrainian Academy of Sciences, Kiev, 112 pp.

BIOURGE P., 1923 — Les moisissures du groupe Penicillium. La cellule 33: 5-331.

Вокнаку F. E. M. M., 1986 — Studies on thermophilic and thermotolerant fungi in the soil of kingdom

of Saudi Arabia. M. Sc. Thesis, Girls College of Education, Jeddah, Kingdom of Saudi

Arabia.

Bunce M. E., 1961 — Humicola stellatus sp. nov., a thermophilic mould from hay. Transactions of the

british mycological society 44: 372-376.

BURDSALL H. H. Jr, 1981 — The taxonomy of Sporotrichum pruinosum and Sporotrichum pulverulen-

tum | Phaenerochaete chrysosporium. Mycologia 73: 675-680.

CANNON Р. F., 1986 — A revision of Achaetomium, Achaetomiella and Subramaniula, and some

similar species of Chaetomium. Transactions of the british mycological society 87: 45-76.

Cannon P. F., 1990 — Name changes in fungi of microbiological, industrial and medical importance.

Mycopathologia 111: 75-83.

CANNON P. F, HAwKsworTH D. L. & SHERWOOD-PIKE M. A., 1985 — The British Ascomycotina. Ап

Annotated Checklist. CMI, 302 pp.

CARMICHAEL J. W., KENDRICK Bryce W., CONNERS I. L. & SIGLER L., 1980 — Genera of Hyphomycetes.

The University of Alberta Press, Alberta, Canada, 386 pp.

Carmo-sousa L. do, 1970 — Genus 11. Trichosporon Behrend. In “The yeasts. A taxonomic study.

2nd revised and enlarged edition”, J. Lodder (ed.), North-Holland Publishing Co. —

Amsterdam — London, pp. 1309-1357.

CHEN K-Y. & CHEN Z-C., 1996 — A new species of Thermoascus with a Paecilomyces anamorph and

other thermophilic species from Taiwan. Mycotaxon 50: 225-240.

Cooney D. G. & EMERSON R., 1964 — Thermophilic Fungi. An Account of their Biology, Activities and

Classification. W. Н. Freeman and Co., San Francisco and London, 188 pp..

Corpa A. C. J., 1842 — Icones fungorum hucusque congnitorum. Tome V, F. Ehrlich, Prague, 92 pp.

CRAVERI R., MANACHINI P. L. & CRAVERI A., 1964 — Eumiceti thermofili presenti nel suols. Annali di

microbiologia ed enzimologia 14: 13-26.

Source : MNHN, Paris

THERMOPHILIC FUNGI 63

Crisan E. V., 1973 — Current concepts in thermophilism and the thermophilic fungi. Mycologia 65:

1171-1198.

Curran R. S., 1985 — Taxonomy of the Onygenales: Arthrodermataceae, Gymnoascaceae, Myxotri-

chaceae and Onygenaceae. Mycotaxon 24: 1-216.

Curzi M., 1930 — Ricerche morphologiche e sperimentali su micromicete termofilo (Acremoniella

thermophila Curzi). Bolletino stazione di patologia vegetale, Roma, N.S., 10: 222-280.

DzPLOEY J. J., 1992 — Some factores affecting germination of Rhizomucor miehei sporangiospores.

Mycologia 84: 77-81.

DESHPANDE V., ERIKSSON K. E. & PETTERSSON B., 1978 — Production, purification and partial

characterisation of 1, 4-beta-glucosidase enzymes from Sporotrichum pulverulentum.

European journal of biochemistry 90: 191-198.

Dix N. J. & WEBSTER J., 1995 — Fungal Ecology. Chapman & Hall, London, 549 pp.

DoMscH К. H., Gams W. & ANDERSON T. H., 1980 — Compendium of Soil Fungi. Vols. | & 2.

Academic Press, London, 405 pp.

DURAND H., SOUCAILLE P. & TiRABY G., 1984 — Comparative study of cellulases and hemicellulases

from four fungi: mesophiles Trichoderma viride and Penicillium sp. and thermophile

Thielavia terrestris and Sporotrichum cellulophilum. Enzyme microbial technology 6: 175-

180.

Ессімѕ Н. O. W. & MALIK K. A., 1969 — The occurrence of thermophilic cellulolytic fungi in a

pasture land soil. Antonie van Leeuwenhoek 35: 178-184.

EICKER A., 1972 — Occurrence and isolation of South African thermophilic fungi. South African

journal of science 68: 150-155.

Etuis D. H., 1981 — Sporangiospore ornamentation of thermophilic Rhizopus species and some

allied genera. Mycologia 73: 511-523

Erus D. H., 1981 — Ultrastructure of Thermophilic fungi IV. Conidial ontogeny in Thermom)

Transactions of the british mycological society 77: 229-241.

ELuis D. H., 1982 — Ultrastructure of thermophilic fungi V. Conidial ontogeny in Humicola grisea

var. thermoidea and H. insolens. Transactions of the british mycological society 78: 129-139.

Euuis M. B. 1971 — Dematiaceous Hyphomycetes. СМІ. Kew, Surrey, England, 608 pp.

ELLIS M. B. 1976 — More Dematiaceous Hyphomycetes. CMI. Kew, Surrey, England, 507 pp.

EMERSON R., 1968 — Thermophiles. In “The Fungi, Vol.3" (Eds. С.С. Ainsworth & A. S. Sussman),

pp. 105-128, Academic Press, New York.

ERICKSSON O, E. € HAWKSWORTH D. L., 1993 — Outline of the ascomycetes — 1993. Systema

ascomycetum 12: 51 — 257.

Evans H. C., 1971 a — Thermophilous fungi of coal spoil tips. I. Taxonomy. Transactions of the

british mycological society 57: 241-254.

Evans H. C., 1971 b — Thermophilous fungi of coal spoil tips II. Occurrence, distribution and

temperature relationships. Transactions of the british mycological society 57: 255-266.

FERGUS C. L., 1964 — Thermophilic and thermotolerant molds and actinomycetes of mushroom

compost during peak heating. Mycologia 56: 267-284.

FERGUS C. L. & SINDEN J. W., 1969 — A new thermophilic fungus from mushroom compost: Thielavia

thermophila spec. nov. Candian journal of botany 47: 1635-1637.

FIELDS W. G., 1968 — A new species of Sordaria. Mycologia 60: 1117-1118.

GOCHENAUR S. E., 1975 — Distributional patterns of mesophilous and thermophilous microfungi in

two Bahamian soils. Mycopathologia mycologia applicata 57: 155-164.

GRIFFON E. de MAUBLANC A., 1911 — Deux moisissures thermophiles. Bulletin de la société mycolo-

gique de France 27: 68-74.

Guarro J. & ARX J. A. von, 1987 — The ascomycete genus Sordaria. Persoonia 13: 301-313

Guarro J., ABDULLAH S. K., AL-BADER S. M., Figueras M. J. & GENE J., 1996 — The genus

Melanocarpus. Mycological research 100: 75-78.

Hepcer J. H., 1974 — The ecology of thermophilic fungi in Indonesia. Jn Kilbertus G., Reisinger O.,

Mourey A. and Cancela da Fonseca J. A. (eds.), Biodegradation et Humification I, Pierron

Editeur, Sarreguemines, pp. 59-65.

es.

Source : MNHN, Paris

64 J. MOUCHACCA

HEDGER J. N. & Hupson H. J., 1970 — Thielavia thermophila and Sporotrichum thermophilum.

Transactions of the british mycological society 54: 497-500.

Ноос G. S. de, 1974 — The genera Blastobotrys, Sporothrix, Calcarisporium and Calcarisporiella gen.

nov. Studies in mycology 7: 1-84.

Ноос G. S. de, Smit M. TH & Gurno E., 1980 — A revision of the genus Geotrichum and its

teleomorphs. Studies in mycology 29: 1-131.

Jost M. C., 1982 — A new species of Rhizomucor. Sydowia 35: 100-103.

Кімовнгто S., CHUA J. W., Kato N., YOSHIDA Т. & TAGUsHI H., 1986 — Hydrolysis of cellulose by

cellulases of Sporotrichum cellulophilum in an ultrafilter membrane reactor. Enzyme

microbial technology 8: 691-695.

KLororrk A. van, 1974 — Revision der thermophilen Sporotrichum arten: Chrysosporium thermo-

philum (Apinis) comb. nov. und Chrysosporium fergusii sp. nov. = status conidialis von

Corynascus thermophilus (Fergus und Sinden) comb. nov. Archives of microbiology 98:

366-369.

KLOPOTEK A. van, 1974 — Thielavia heterothallica spec. nov., die perfekte form von Chrysosporium

thermophilum. Archives of microbiology 107: 223-224.

LATOUCHE C. 1, 1950 — On a thermophile species of Chaetomium. Transactions of the british

mycological society 33: 94-104.

Lasure L. L. & INGLE M. B., 1976 — Some effects of temperature on zygospore formation in Mucor

miehei. Mycologia 68: 1145-1151.

MALLOCH D. & CAIN R. E, 1972 — The Trichocomataceae: Ascomycetes with Aspergillus, Paecilo-

myces, and Penicillium Imperfect states. Canadian journal of botany 50: 2613-2628.

Marrocu D. & CAIN R. Е, 1973 — The genus Thielavia. Mycologia 65: 1055-1077.

Marcy M. R., ENGELHARDT T. C. & UPADHYAY J. M. , 1984 — Isolation, purification and some

properties of protease I from a thermophilic mold: Thermoascus aurantiacus var. levispo-

rus. Mycopathologia 87: 51-65.

Mason E. W, 1933 - Annotated account of fungi received at the Imperial Mycological Institute, 1933,

List II ( Fasc. 2). Mycological Papers n? 3, London.

Mason E. W., 1941 — Annotated account of fungi received at the Imperial Mycological Institute 1941,

List 2 ( Fasc. 3, special part ). Mycological Paper n? 5, London.

MIEHE H., 1905 — Uber die Selbsterhitzung der Heues. Arbeiten der deutschen Landwirtschaftsgesel-

Ischaft 111: 76-91.

Miene H., 1907 — Die Selbsterhitzung des Heues. Eine Biologische Studie. G. Fischer, Jena, 127 pp.

Miter P. D., 1977 — Radial growth responses to temperature by 58 Chaetomium species, and some

taxonomic relationships. Mycologia 69: 492-502.

MILLNER P. D., Мотта J. J. & LENTZ P. L., 1977 — Ascospores, germ pores, ultrastructure, and

thermophilism in Chaetomium. Mycologia 69: 720-733.

Mirza J. H., KHAN S. M., BEGUM S. & SHAGUFTA S., 1979 — Mucorales of Pakistan. University of

Agriculture, Faisalabad, Pakistan.

MORGAN-JONES G., 1974 — Notes on Hyphomycetes. V. A new thermophilic species of Acremonium.

Canadian journal of botany 52: 429-431.

MORGAN-IONES G. de Gams W., 1982 — Notes on Hyphomycetes. XLI. An endophyte of Festuca

arundinacea and the anamorph of Epichloe typhina, new taxa in one of two new sections of

Acremonium. Mycotaxon 15: 311-318.

MORINAGA T., KANDA S. & Nomi R., 1986 — Lipase production of a new thermophilic fungus,

Humicola lanuginosa var. catenulata. Journal of fermentation technology 64(5): 451-453.

MOUBASHER A. H., 1993 — Soil Fungi in Qatar and other Arab Countries. The Scientific and Applied

Research Centre, University of Qatar, 566 pp.

MOUBASHER A. H., ABDEL-HAFEZ S. I. 1. & EL-MAGHRABY O. M. O., 1988 — Seasonal fluctuations of

soil-borne and air-borne fungi of Wadi Bir-El-Ain in the Eastern Desert of Egypt.

Naturalia Monsperliensa, Sér. Botanique 52: 57-69.

MOUBASHER А. H., MAZEN M. B. & ABDEL-HAFEZ A. I. I., 1979 - Humcola hyalothermophila sp. nov.

Transactions of the british mycological society 72: 509-510.

Source : MNHN, Paris

THERMOPHILIC FUNGI 65

MOUCHACCA J., 1973 — Les Thielavia des sols arides: Espéces nouvelles et analyse générique. Bulletin

trimestriel de la société mycologique de France 89: 295-311.

MOUCHACCA J., 1994 — Thermophilic and thermotolerant fungi in the Middle East: Biodiversity and

taxonomic reappraisal. Vth International Mycological Congress, ABSTRACTS, August

14-21, 1994, Vancouver, University of British Columbia, Canada, Lecture Abstract,

p. 150.

MoucHacca J., 1995 — Thermophilic Fungi in Desert Soils, a Neglected Extreme Environment. In

“Microbial Diversity and Ecosystem Function" (Allsopp et al., eds.). CAB International

and Royal Holloway, University of London, 10-13 August 1993, London, pp. 265-288.

MOUSTAFA A. F. & EZZ-ELDIN E. K., 1989 — Gilmaniella multiporosa, a new dematiaceous hypho-

mycete from Egyptian soils. Transactions of the british mycological society 92: 502-505.

Narain R., Srivastava R. В. & MEHROTRA B. S., 1983 — A new thermophilic species of Scyralidium

from India. Zentralblatt für mikrobiologie 138: 569-572.

Noack K., 1912 — Beiträge zur Biologie der thermophilen Organismen. Jahrbücher für wissenschaft-

liche Botanik 51: 593-648.

Oonschor С. А. М. van, 1977 — The genus Myceliophthora. Persoonia 9: 401-408.

Оовѕснот C. A. N. van, 1980 — A revision of Chrysosporium and allied genera. Studies in mycology

20: 1-89.

Оовѕснот C. А. N. van & Ноос G. S. de, 1984 — Some hyphomycetes with thallic conidia

Mycotaxon 20: 129-132.

ртт J. L, 1979 — The genus Penicillium and its teleomorphic states Eupenicillium and Talaromyces.

Academic Press, London 634 pp.

PRAKASH К. & SARBHOY A. K., 1993 — Mucor thermophilus spec. nov. and other Mucorales from

India. Zentralblatt für mikrobiologie 148 (8): 531-534

PuGH G. J. E, BLAKEMAN J. P. & MORGAN-IONES G., 1964 — Thermomyces verrucosus sp. nov. and

Т. lanuginosus. Transactions of the british mycological society 47: 115-124.

Raper К. B. & THOM Ch., 1949 — A Manual of the Penicillia. Williams & Wilkins, Baltimore, ix

+ 875 pp.

SaccarDo P. A., 1908 — Notae mycologicae. Ser. X. Annales mycologici 6: 553-569.

SAMSON R. A., 1974 — Paecilomyces and some allied hyphomycetes. Studies in mycology 6: 1-119.

Samson R. A. & TANSEY M. R., 1977 — Guide to thermophilic and thermotolerant fungi. Abstracts,

Second International Mycological Congress, Tampa, Florida, 5 p.

Samson R. A., CRISMAN M. Jack € TANSEY М. R., 1977 — Observations on the thermophilous

ascomycete Thielavia terrestris. Transactions of the british mycological society 69: 417-423.

SATYANARAYANA T., JOHRI B. М. & KLEIN J., 1992 — Biotechnological potential of thermophilic

fungi. In Handbook of Applied Mycology”, vol. 4, (Eds. D. K. ARORA, R. P. ELANDER &

K. G. Микки), pp. 729-761, Marcel Dekker, New York.

SCHEKKAR Sharma S. Н. 4 JOHRI B. N., 1992 — The role of thermophilic fungi in agriculture.

In “Handbook of Applied Mycology”, vol. 4, (Eds. D. K. ARORA, R. P. ELANDER & K. С.

Микеил), pp. 707-728, Marcel Dekker, New York.

ӛсніррек M. A. A., 1978-2. On the genera Rhizomucor and Parasitella. Studies in mycology 17: 53-71.

SCHIPPER M. A. A., 1979 — Thermomucor, (Mucorlaes). Antonie van Leeuwenhoek 45: 275-280.

SEIFERT К. A., 1985 — A monograph of Stilbella and some allied hyphomycetes. Studies in mycology,

27: 1-235.

SIGLER L., 1987 — Trichothecium cinnamomeum, an earlier name for the thermophilic hyphomycete

Malbranchea sulphurea. Mycologia 79: 142-143

SIGLER L. & CARMICHAEL J. W., 1976 — Taxonomy of Malbranchea and some other hyphomycetes

with arthroconidia. Mycotaxon 4: 349-488.

SIGLER L. & WANG C. J. K., 1990 - Scytalidium circinatum sp. nov., a Hyphomycete from utility poles.

Mycologia 82: 399-404.

Siva Denise M. W. & HANLIN R. T., 1996 — Chaetomidium heterotrichum from Venezuela, with a key

to species and cladistic analysis of the genus Chaetomidium. Mycoscience 37: 261-267.

5мїтн G., 1961 — Polypaecilum gen. nov. Transactions of the british mycological society 44: 437-440.

Source : MNHN, Paris

66 J. MOUCHACCA

SINGER R., 1975 — The Agaricales in modern taxonomy. J. Cramer, 3rd edition, 912 pp.

SIVANESAN A. & SUTTON B. C., 1985 — Microfungi on Xanthorhoea. Transactions of the british

mycological society 85: 239-255.

Sopp J. O., 1912 — Monographie der Pilzgruppe Penicillium mit besonderer Berücksichtigung der in

Norwegen. gefundenen Arten. Videnskabs-Selskabets i Christiania, Skrifter, 1 Math.-

Naturv. KI., 1912, N° 11, i-vi + 208 pp.

SrALPERS 1. A., 1978 — Identification of wood-inhabiting Aphyllophorales in pure culture. Studies in

cology 16: 1-248.

STALPERS J. A., 1984 — A revision of the genus Sporotrichum. Studies in mycology 24: 1-105.

Stork A. C., 1965 — Thermophilic species of Talaromyces Benjamin and Thermoascus Miehe.

Antonie van Leeuwenhoek 31: 262-276.

SroLk A. C. & SAMSON R. A., 1972 — The genus Talaromyces. Studies on Talaromyces and related

genera IL. Studies in mycology 2: 1-65.

SrOLK А. C. & Samson R. A., 1985 — А new taxonomic scheme for Penicillium anamorphs.

In“ Advances in Penicillium and Aspergillus Systematics”, NATO ASI Series, pp. 163-192,

Plenum Press, New York & London.

Straatsma С. & SAMSON R. A., 1993 — Taxonomy of Seytalidium thermophilum, an important

thermophilic fungus in mushroom compost. Mycological research 97: 321-328.

SUBRAHMANYAM A., MEHROTRA В. S. & THIRUMALACHAR M. J., 1977 — Thermomucor - A new genus

of Mucorales. Georgia journal of science 35: 1-4.

SUBRAHMANYAM A., 1981 — Studies on Thermomycology. Mucor thermo-hyalospora sp. nov. Biblio-

theca mycologica 91: 421-423.

SUBRAHMANYAM À., 1983 — A new thermophilic variety of Humicola grisea var. indica. Current

science 49: 30-31.

Tansey M. R. & Brock T. D., 1978 — Microbial life at high temperatures: ecological aspects.

In Kushner D. J. (ed.) Microbial Life in Extreme Environments, Academic Press, London,

pp. 159-216

Tansey M. R. & JACK M. A., 1975 — Thielavia australiensis sp. nov., a new thermophilic fungus from

incubator bird (mallee fowl) nesting material. Canadian journal of botany 53: 81-83.

TROTTER A., 1931 — (Saccardo's) Sylloge Fungorum omnium hucusque cognitorum. 25: 1-1093.

TSIKLINSKY P, 1899 — Sur les mucédinées thermophiles. Annales de l'Institut Pasteur, Paris 13:

500-505.

Upacawa S.-L, Awao Т. & ABDULLAH S. K., 1986 — Thermophymatopsora a new thermophilic

genus of basidiomycetous hyphomycetes, Mycotaxon 27: 99-106.

UEDA S. & Upacawa S-L, 1983 — Thermoascus aegyptiacus, a new thermophilic ascomycete.

Transactions of the mycological society of Japan 24: 135-142.

WERESUB L. К. & стл P. M., 1973 — On Papulaspora and bulbilliferous basidiomycetes Burgoa

and Minimedusa. Canadian journal of botany 49: 2203-2214

Yacusut T., SOMEYA А. de UDAGAWA S.-L., 1995 -Two interesting cleistothecial Ascomycetes from

soils. Mycoscience 36; 151-154.

ZHENG R.-Y, & CHEN G.-Q., 1991 — A non-thermophilic Rhizomucor causing human primary

cutaneous mucormycosis. Mycosystema 4: 45-57.

ZHENG R.-Y. & CHEN G.-Q., 1993 — Another non-thermophilic Rhizomucor causing human primary

cutaneous mucormycosis. Mycosystema 6: 1-12.

ZHENG R.-Y. & JIANG H., 1995 — Rhizomucor endophyticus sp. nov., an endophytic zygomycete from

higher plants. Mycotaxon 56: 455-466.

Source : MNHN. Paris

THERMOPHILIC FUNGI 67

INDEX OF CITED GENERIC AND SPECIFIC EPITHETS

Absidia

Achaetomium

macrosporum

thermophilum

Acremoniella

Acremoniella sp.

thermophila

Acremonium

Acremonium section Nectrioidea

alabamense

cellulophilum

thermophilum

Allescheria terrestris

Aphanoascus

Arthrographis sulfurea

Burgoa

Byssochlamys

Byssochlamys sp.

Calcarisporiella

thermophila

Calcarisporium thermophile

Canariomyces

Canariomyces notabilis

thermophilus

Cephaliophora tropica

Cephalosporium

Chaetomidium

thermophilum

Chaetomidioides Section

Chaetomium

britannicum

megasporum

mesopotamicum

thermophilum

thermophilum var. coprophile

thermophilum var. dissitum

vitellinum

virginicum

Chrysonilia

Chrysosporium

fergusii

lignorum

pruinosum

thermophilum

Citromyces sphagnicola

Coonemeria

aegyptiaca

crustacea

verrucosa

Corynascus

heterothallicus

novoguinensis

thermophilus

Dactylomyces

crustaceus

thermophilus

Emmonsia

brasiliensis

Endoblastomyces

thermophilus

Geosmithia emersonii

Geotrichum

candidum var. thermoideum

cinnamomeum

Gilmaniella humicola

punctiformis

macrospora

multiporosa

thermophila

Gymnoascaceae

Humicola

brevis var. thermoidea

brevispora

fuscoatra

fuscoatra var. nigra

fuscoatra var. longispora forma insolens

grisea

grisea var. indica

grisea var. thermoidea

hyalothermophila

insolens

insolens var, thermoidea

lanuginosa

lanuginosa var. catenulata

nigrescens var. thermorongeura

stellata

Lagenidium

thermophilum

Malbranchea

cinnamomea

pulchella

pulchella var. sulfurea

sulfurea

Melanocarpus

albomyces

coprophilus

oblatus

thermophilus

Melanomphalia thermophila

Melanospora

Microsporon

Minimedusa

Monotospora

dalae

Source : MNHN. Paris

lanuginosa

Mucor

buntingii

hagemii

miehei

muriperda

parasiticus

pusillus

septatus

tauricus

thermo-hyalospora

thermophilus

Myceliophthora

fergusii

hinnulea

indica

lutea

thermophila

Myriococcum

albomyces

praecox

thermophilum

Onygenaceae

Paecilomyces

aegyptiacus

byssochlamydioides

crustaceus

puntonii

taitungiacus

variotii

Paecilomycopsis

Papulaspora

byssina

thermophila

Penicillium

dupontii

emersonii

thermophilum

thermophilus

Phanerochaete chrysosporium

Polypaecilum

Polypaecilum sp.

Rhizomucor

endophyticus

miehei

nainitalensis

pakistanicus

pusillus

septatus

tauricus

variabilis var. regularior

variabilis var. variabilis

Rhizopus

J MOUCHACCA

Rhizopus sp.

parasiticus

Russula roseipes subsp. thermophila

Scytalidium

allahabadum

indonesicum

lignicola

thermophilum

Sepedonium

lanuginosum

Sporotrichum

aureum

cellulophilum

dehradunense

pruinosum

pulverulentum

thermophilum

Sordaria fimicola

thermophila

Stilbella

thermophila

Suillus hirtellus var. thermophilus

Talaromyces

section Emersonii

bacillosporus

byssochlamydioides

dupontii

emersonii

leycettanus

thermophilus

Thermoascus

aegyptiacus

aurantiacus

aurantiacus var. levisporus

crustaceus

crustaceus var. verrucosus

isatschenkoi

taitungiacus

thermophilus

Thermoideum

sulphureum

Thermomucor

indicae-seudaticae

Thermomyces

ibadensis

lanuginosus

stellatus

verrucosus

Thermophymatospora

fibuligera

Thielavia

albomyces

australiensis

Source : MNHN, Paris.

THERMOPHILIC FUNGI

heterothallica

minuta

minuta var. thermophila

pingtungia

terrestris

thermophila

Tieghemella muriperda

Torula

thermophila

Trichosporiella

Trichosporon capitatum

Trichothecium cinnamomeum

Tubaria thermophila

Zalerion

maritima

thermophylii

Source : MNHN. Paris

ire!

NER ТЕРА

phil

Cryptogamie, Mycol. 1997, 18(1): 71-79 71

DAPTATION PARASITAIRE

DES CHAMPIGNONS KERATINOPHILES TELLURIQUES

ET CONSEQUENCES EN PATHOLOGIE HUMAIN

ET ANIMALE

Dominique CHABASSE

Laboratoire de Parasitologie — Mycologie

CHU — 4 rue Larrey 49100 ANGERS — FRANCE

RESUME — Les champignons kératinophiles constituent une flore importante et variée caractérisée

par sa capacité à se développer au dépens de la kératine, protéine complexe entrant dans la

composition de la peau et des phanères de l'homme et des animaux. Ils appartiennent pour la plupart

à la classe des Ascomycetes, et sont positionnés depuis 1985 dans l'ordre des Onygenales au sein de

deux familles distinctes les Arthrodermataceae et les Onygenaceae. Si beaucoup d'entre eux restent des

saprophytes du sol, un certain nombre comme les Dermatophytes et des espéces assimilées (Ap/ia-

noascus...) évoluent vers un parasitisme d'abord facultatif puis quasi obligatoire chez l'homme et/ou

nimal. Le cheminement sol — animal — homme semble étre l'évolution phylogénique habituelle de

ces champignons. Les conséquences pratiques de cette adaptation au parasitisme sont : — Sur le plan

biologique, une perte progressive de leur mode de reproduction saprophytique (stade téléomorphe

puis anamorphe), avec en revanche un développement des structures de diffusions propres au

parasitisme (parasitisme pilaire et arthrospores virulentes dans les corneocytes) qui assurent la

dissémination du champignon ; — Sur le plan épidémiologique, il est intéressant de constater que les

espéces géophiles ou telluriques (Microsporum gypseum, Aphanoascus fulvescens...) et parfois zoophi-

les (Trichophyton verrucosum) sont à l'origine d'une intense réaction inflammatoire chez l'homme.

À l'opposé, les espèces anthropophiles (Microsporum audouinii, Trichophyton violaceum, Trichophy-

ton rubrum) mieux adaptées à l'homme, évoluent sur un mode chronique avec des réactions de rejets

limitées ou nulles. Les champignons kératinophiles représentent un mode d'étude intéressant d'adap-

tation au parasitisme en mycologie médicale.

ABSTRACT — Keratinophilic fungi form an important and varied flora distinguished by the ability

to develop on an ideal substrate, keratin, a complex protein forming part of the composition of the

skin, nails and hair of men and animals. These fungi belong to the Ascomycetes and are placed in the

order Onygenales at the center of two distinct families, the Arthrodermataceae and the Onygenaceae.

If most of these species remain saprophytes of the soil, a certain number among them as the

Dermatophytes and related fungi evolve in the direction of an optional parasitic state on animals

while others go resolutely toward an almost obligatory parasitism. The pathway from soil to animal

to man is the usual phylogenetic evolution of these fungi. The practical consequence of the parasitic

adaptation implies at least two observations : — A progressive loss of the saprophytic reproduction of

these species. This is shown by the impossibility of producing the teleomorphic state in the anthro-

pophilic fungi and by the difficulty of obtaining a good conidiogenesis (anamorphic state) ; —

Geophilic species known as dermatophytes like Microsporum gypseum or closely related like Apha-

Source : MNHN, Paris

72 D. CHABASSE

noascus fulvescens and the zoophilic dermatophytes like Trichophyton verrucosum produce highly

inflammatory lesions with a tendency to spontaneous cure, On the other hand anthropophilic

dermatophytes like Trichophyton violaceum or Microsporum audouinii produce less inflammatory and

more chronic lesions. Keratinophilic fungi represent the ideal model of parasitic adaptation in

medical mycology.

MOTS CLES — Champignons kératinophiles telluriques, Dermatophytes, Chrysosporium, Sapro-

phytisme, Parasitisme, Evolution phylogénique.

KEY WORDS — Keratinophilic fungi, Dermatophytes, Chrysosporium, Saprophytism, Parasitic

adaptation, Phylogenetic evolution.

INTRODUCTION

Les micromycétes kératinophiles telluriques sont des champignons isolés du sol

(Chabasse et al., 1985, 1987, 1988, 1989) et souvent sur des animaux : petits mammifères

sauvages (rongeurs, insectivores) et autres fouisseurs (Chabasse ег al., 1987 ; Houin et al.,

1972 ; Otcenasek & Dvorak, 1962), ainsi que de certains oiseaux (Chabasse & Guiguen,

1986).

Leur caractéristique commune est d'étre kératinophiles, c'est-à-dire qu'ils privi-

légient un substrat nutritif particulier : la kératine. Cette kératine tellurique appelée :

« kératine morte > est issue des mammifères (poils, fragments de cornes ou de sabots...),

des oiseaux (plumes...) et des carapaces d' insectes mélangées au sol. L'origine tellurique de

ces espéces est connue depuis les travaux de Sabouraud (1910) à la fin du siécle dernier.

Plus tard le hongrois Szathmary (1940) isola un dermatophyte géophile appelé : Tricho-

phyton terrestre gyratium, puis : Trichophyton fluviale, vraisemblablement le Trichophyton

mentagrophytes d'aujourd'hui. Le comportement kératinophile de ces champignons fut

rapidement utilisé pour les isoler. C'est ainsi qu'à partir des constatations de Toma (1929)

sur le parasitisme des cheveux par les agents de teignes et les observations de Karling

(1946) qui utilisait des cheveux humains pour piéger dans le sol d'un cimetiére un

kératinophile appelé : Rhizophydium keratinophilum, Vanbreuseghem (1952, 1960) mit au

point une technique originale pour extraire ces champignons du sol.

Cette méthode consiste à déposer des fragments de cheveux stériles sur de la terre

placée dans des boites de Pétri. Elle fut appelée par Benedek (1962) : « the To-Ka-Va »

hair baiting-méthode, ou technique de piégeage sur cheveux de Toma-Karling-

-Vanbreuseghem, rappelant le róle respectif de chacun des auteurs dans cette découverte.

Cette technique communément appelée : technique de Vanbreuseghem fut à l'origine de

trés nombreux travaux réalisés dans le monde entier confirmant l'hypothése de départ

C'est-à-dire la vie saprophytique des dermatophytes et leur origine tellurique (De Vroey,

1968, 1970 ; Helluy et al., 1965). C'est ainsi que Trichophyton ajelloi (= Keratinomyces

ajelloi) a été découvert et décrit pour la première fois par Vanbreuseghem en 1952. De

méme Ajello (1953) isola à son tour Microsporum gypseum puis Microsporum cookei et

Durie et Frey (1950) mirent en évidence : Trichophyton terrestre.

Cette technique a permis aussi de révéler la reproduction sexuée de ces derma-

tophytes géophiles et de décrire des espéces proches mais appartenant à des genres

différents : Chrysosporium, Geomyces et Malbranchea (Carmichael, 1962 ; Currah, 1985 ;

van Oorschot, 1980).

Source : MNHN, Paris

ADAPTATION PARASITAIRE DES CHAMPIGNONS KÉRATINOPHILES 73

ÉVOLUTION DE LA TAXIMONIE

La plupart des micromycétes kératinophiles telluriques appartiennent à la classe

des Ascomycètes, à la sous classe des Euascomycètes, comprenant elle-même 2 grandes

familles : les Gymnocascaceae et les Onygenaceae.

En 1973, Fennel inclut ces deux familles dans l'ordre des Eurotiales, mais Benny

et Kimbrough en 1980 proposérent une nouvelle distribution créant et placant ces espéces

dans l'ordre des Onygenales. Currah proposa à son tour en 1985 une révision complete de

cet ordre oú l'on retrouve les espéces les plus adaptées au parasitisme humain.

Currah ne retient plus comme élément déterminant la morphologie des ascocar-

pes (gymnothéce, cleistothéce, perithéce...) mais l'aspect des ascospores (lisses ou orne-

mentées) associé à des critéres physiologiques, c'est-à-dire l'aptitude à dégrader ou non la

kératine. Ainsi il définit 4 familles dont deux ayant une affinité pour la kératine, les

Arthrodermataceae et les Onygenaceae. Les deux autres familles ont soit un substrat

nutritif plus varié (Gymnoascaceae), soit une préférence pour la cellulose (Myxotricha-

ceae) (fig. 1).

La comparaison des séquences nucléodiques de la sous-unité 18S ribosomiale de

nombreuses espéces appartenant à l'ordre des Onygenales a permis de confirmer la plupart.

du temps la cohésion de cette classification (Bowman et al., 1992 ; Leclerc et al., 1994).

Un grand nombre d'espéces adaptées au parasitisme appartiennent à l'ordre des Onygé-

nales : Trichophyton, Microsporum, Histoplasma, Blastomyces, Coccidioides...

DU SAPROPHYTISME AU PARASITISME

Microsporum gypseum est un dermatophyte géophile menant essentiellement

une vie saprophytique à partir de substrats kératiniques (kératine morte) comme les poils

d'animaux, des fragments de cornes de bœuf ou de sabots de chevaux (Ajello, 1953 ;

Badillet, 1982, 1991 ; Chmel & Vollekova, 1981). Il posséde cependant de réelles capacités

au parasitisme si l'on en juge d'aprés les lésions qu'il engendre chez l'homme et l'animal

(Chabasse, 1991). Il en est de méme pour un Chrysosporium, dont le téléomorphe,

Aphanoascus fulvescens, est incriminé, lui aussi, dans des lésions « dermatophytic like »

(Albala et al., 1982 ; Chabasse et al., 1989 ; Gueho et al., 1985; Rippon et al., 1970 ;

Todaro et al., 1984 ; Vanbeuseghem & Devroey, 1980).

Le cheminement du sol à l'animal puis de l'animal à l'homme serait l'évolution

classique de ces micromycétes, selon Novac & Galcoczy, (1966), Chmel & Vollekova (1981,

et Dei Cas & Vernes (1986). Le kératinophile tellurique, saprophyte du sol, végéterait

d'abord sur la kératine du sol (kératine morte), puis préparé à ce substrat sélectif, il

passerait aisément sur le poil de l'animal (kératine vivante) ou directement chez l'homme.

L'évolution de ces espèces géophiles en zoophiles puis anthropophile, serait due à la fois à

des phénomènes extrinsèques (rencontre avec l'hôte) et intrinsèque (facteurs de reconnais-

sance, de virulence...) propres au champignon lui-même.

Source : MNHN, Paris

D. CHABASSE

TABLE 3 FAMILIES OF THE ONYGENALES

Peridia and Substra

Family Ascospores Appendages | Апатогрһа and Habitats

ONYGENACEAE

seniore dung

=? sol enriched with

Farm or dura.

CHRYSOSPORIUM

MALBRANCHI

SPORENOONEMA KERATIN"

| den

ARTHRODERMATACEAE fom testen.

eir and aun

some are

Sarai on

imei

smooth; oblate to i

; снтүзовротим

oblate discoid or 7 MICROSPORUM

e: TRICHOPRYTON

MYXOTRICHACEAE

ing plant

materiale

Т2

сү? X, CELLULOSE’

) | r, straw,

paper, straw,

| soil around the

roots of plants

smooth or striate;

fusiform, өрөн.

GYMNOASCACEAE

decaying vegetation

(2 |

soll rien in

organic matter

C VARIABLE *

smooth or sign | cori

Irregular ("lumpy y; рна а

‘often with polar

and/or equatorial |

thickenings.

Figure — Les Onygénales

d'aprés Currah, Mycotaxon, 1985

Source : MNHN, Paris

ADAPTATION PARASITAIRE DES CHAMPIGNONS KÉRATINOPHILES T7»

CONSÉQUENCES ÉPIDÉMIOLOGIQUES, CLINIQUES ET BIOLOGIQUES

Épidémiologiques а

Les géophiles contaminent accidentellement l'homme soit directement

(souillure traumatisme) soit par l'intermédiaire d'un animal relais (chien de chasse). Il n'y

a pas de contamination inter-humaine (Chermette, 1991 ; Garg et al., 1985).

. Les zoophiles (et géo-zoophiles) passent aussi chez l'homme soit directement

par contact avec un animal contaminé (le plus souvent chien, chat...) soit à partir de ses

poils contenant la forme virulente du champignon disséminée dans l'environnement

humain (tapis, fauteuil, couverture...) (Caretta ег al., 1989; Chabasse et al., 1991;

Chermette, 1991 ; Woodyer, 1977).

La diffusion de ces espéces à l'homme dépendra de leur spécificité vis à vis de leur

hóte. Elle sera étroite avec Trichophyton equinum, T. gallinae respectivement adaptés au

cheval et aux oiseaux de basse-cour, intermédiaire avec Trichophyton verrucosum parasite

privilégié des bovins et des ovins et lâche avec Microsporum canis qui affectionne de

nombreux animaux familiers et d'élevage. La contamination dépendra également des

possibilités de contact avec l'animal-hóte. C'est ainsi que le chat, encore plus souvent que

le chien, porteur asymptomatique de spores virulentes est à l'origine de bien des contami-

nations humaines. À l'inverse si un animal peut contaminer plusieurs personnes d'une

méme fratrie le passage inter humain est trés difficile (Badillet, 1991).

. Les anthropophiles adaptés au parasitisme humain diffusent bien dans la

population. Les teignes dues a Trichophyton violaceum, Trichophyton soudanense, et

Microsporum langeronii sont de ce fait trés contagieuses (Badillet, 1991).

De méme les squames issus de la peau, ou les fragments d'ongles des nageurs ou

des judokas qui souillent les tapis de sport, les carrelages des douches, des vestiaires du

pourtour des bassins de natation, sont à l'origine de nombreuses dermatophyties du pied

dues à Trichophyton rubrum et à T. mentagrophytes variété interdigitale (Detandt &

Nolard, 1988 ; Chabasse ег al., 1995)

Cliniques :

Il est habituel de dire que plus une espèce fongique est adaptée à son hôte, mieux

elle est supportée par ce dernier. Les dermatophytes anthropophiles bien que trés engagés

dans la voie du parasitisme sont à l'origine de lésions habituellement séches, squameuses

et évoluant sur un mode chronique (Trichophyton rubrum). À l'opposé les espèces géophi-

les ou geo-zoophiles sont génératrices de réactions bruyantes et inflammatoires. Elles

peuvent d'ailleurs guérir spontanément tellement le rejet par l'hóte peut étre important.

C'est le cas de Trichophyton verrucosum inféodé aux bovidés qui provoque chez l'homme

des sycosis de la barbe et de la moustache voire un kérion de cuir chevelu. Le traitement à

base de corticoides peut parfois suffire à guérir ces patients.

D'autres zoophiles, plus largement répandus dans le monde animal, comme

Microsporum canis sont en revanche à l'origine de lésions plus ou moins inflammatoires.

En général les réactions de l'hóte vis à vis des micromycétes kératinophiles sont assez bien

corrélées avec le degré d'adaptation au parasitisme.

Source : MNHN, Paris

76 D. CHABASSE

Biologiques :

Les micromycetes kératinophiles issus du sol sont à l'origine en primo culture

(milieu de Sabouraud) d'une grande abondance de spores. La conidiogénése trés impor-

tante explique l'aspect de ces cultures volontiers poudreuses. A l'opposé les espéces

anthropophiles (T. violaceum, T. soudanense, T. schoenleinii) ont une conidiogénese de

mauvaise qualité. Il n'est pas toujours aisé de retrouver des macro — ou microconidies

caractéristiques du genre. Cependant cette régle n'est pas toujours vérifiée chez les

anthropophiles oú certaines souches de 7. rubrum et T. tonsurans par exemple sont parfois

riches en spores.

Chez les zoophiles, la morphologie microscopique est variable, certaines espéces

trés inféodées à un hôte comme 7: verrucosum sont habituellement pauvres en spores sur

les milieux usités en mycologie. A l'inverse M. canis, de diffusion large au niveau des

animaux d'élevages et de rente, est plus riche en organes de fructifications.

Ce que l'on constate avec les stades anamorphes se vérifie encore mieux avec les

stades téléomorphes ou sexués.

La plupart des kératinophiles issus du sol sont hétérothalliques avec un équili-

brage entre les souches plus (+) et moins (-).

La méthode de Vanbreuseghem trés usitée pour visualiser la reproduction sexuée

des Arthrodermataceae et des Onygenaceae illustre bien ce phénomène. Les espèces qui ont

choisi le parasitisme vont perdre leur potentiel sexué. Quand on les fait réagir avec

Arthroderma simii on s'aperçoit qu'il y a sélection d'un seul type sexué, soit de signe (+)

(cas du Т. mentagrophytes var. interdigitale), soit de signe (-) (cas du 7. rubrum). Déjà chez

de nombreuses espéces zoophiles il existe un net déséquilibre entre les souches (-) et les

souches (+). On ne connait le stade sexué de M. canis (Nannizzia otae) que gráce à

l'isolement d'une souche négative dans un élevage polonais; toutes les autres sont

désignées (+). Les agents de teignes anthropophiles confrontés avec des souches d'Arthro-

derma simii de signe (+) et de signes (-) ne montrent pas de gymnothéces fertiles contenant

des ascospores, mais tout au plus quelques ébauches de réaction sexuée.

La perte de la reproduction sexuée lors de l'adaptation parasitaire est-elle la

conséquence du parasitisme ou est-ce cette perte qui a provoqué le parasitisme ?

On pourrait imaginer que le champignon face à un environnement hostile non

favorable à sa reproduction sexuée s'adapte au parasitisme pour assurer sa survie.

Au contraire la vie parasitaire a bien pu avoir comme conséquence une perte de certains de

ses métabolisme et voies enzymatiques conduisant à appauvrir et à s ¡ser sa reproduc-

tion de type sexuée d'abord, asexuée ensuite. Il est vraissemblable que les espéces adaptées

au parasitisme orientent leur potentiel enzymatique aux exigences de leur nouvel état

(Chabasse $: Contect-Audonneau, 1994).

CONCLUSION

Ainsi, si l'on retient comme hypothése une évolution phylogénique des kérati-

nophiles telluriques vers le parasitisme animal ou humain, il convient de rester attentif vis

à vis de micromycétes isolés aujourd'hui encore de facon exeptionnelle sur l'homme

(Chrysosporium sp...) mais pouvant devenir demain de véritables agents de mycoses

humaines et animales et ceci en dehors d'un contexte d'immuno-suppression.

Source : MNHN, Paris

ADAPTATION PARASITAIRE DES CHAMPIGNONS KÉRATINOPHILES 77

REFERENCES

AIELLO L., 1953 — The dermatophyte : Microsporum gypseum as a saprophyte and parasite. Journal

of investigative dermatology 21 : 157-171

ALBALA F., MOREDA A. & Lopez G., 1982 — Isolement d' Anixiopsis stercoraria sur lésions dermiques

humaines à Zaragoza (Espagne). Bulletin de la société francaise de mycologie médicale 11 :

287 291.

BADILLET G., PUISSANT A., JOURDAN-LEMOINE M. & Barrault D., 1982 — Pratique du judo et risque

de contamination fongique. Annales de dermatologie et de vénérologie 109 : 661-664.

BADILLET G., 1991 — Dermatophyties et dermatophytes. Atlas clinique et biologique. Varia. Ed. Paris,

p. 303.

BENEDEK T., 1962 — Fragmenta mycologica I. Some historical remarks on the developpement of

« hair baiting » of Toma-karling-Vanbreuseghem (The To-Ka-va-hair baiting method).

Mycopathologia et mycologia applicata 16 : 104-106.

Benny G. L. & KiMBROUGH J. W., 1980 — A synopsis of the orders and families of plectomycetes with

keys to genera. Mycotaxon 12 : 1-12.

BiGuEr J., 1979 — Les phénomènes d'apdaptation des champignons pathogènes en médecine

humaine et vétérinaire. Dermatologica 159 : 28-35.

Bowman B. H., TAYLOR J. W. & WHITE T. J., 1992 — Molecular evolution of the fungi : human

pathogens. Molecular biology and evolution 9 : 893-904.

CaRETTA G., MANCIANTI F. & AJELLO L., 1989 — Dermatophytes and keratinophilic fungi : in cats

and dogs. Mycoses 32 : 620-626.

CARMICHAEL J. W., 1962 — Chrysosporium and some other aleuriosporic hyphomycetes. Canadian

journal of botany 40 : 1135-1173.

CHaBasse D., BOUCHARA J. P. & DE Bièvre C., 1985 — Flore fongique des bacs à sable. Mise еп

évidence, signification. Mycopathologia 90 : 3-13.

CHABASSE D. & GUIGUEN C., 1986 — Flore kératinophile isolée du plumage des pigeons de ville à

Bordeaux. Bulletin de la société francaise de mycologie médicale 15 : 407-412.

CHABASSE D., GUIGUEN C., COUATARMANAC'C A., LAUNAY H., REECHT V. & DE BIÈVRE C., 1987 —

Contribution à la connaissance de la flore fongique des petits mammifères sauvages et du

lapin de garenne. Discussion sur les espèces fongiques rencontrées, Annales de parasitologie

humaine et comparée 62 : 357-368.

CHABASSE D., CONTET-AUDONNEAU N. & MARCHAL L., 1987 — Flore kératinophile du sol en milieu

rural. Études comparatives dans la région d'Angers et de Nancy. Bulletin de la société

francaise de mycologie médicale 16 : 281-286.

Chabasse D., 1988 — Taxinomic study of keratinophilic fungi isolated from soil and some mammals

in France. Mycopatolologia 101 : 133-140.

CHABASSE D., DE GENTILE L. & BOUCHARA J. P., 1989 — Pathogenicity of some Chrysosporium species

isolated in France. Mycopathologia 106 : 171-177.

CHABASSE D., 1989 — Les dermatophytes telluriques isolés en France depuis 1964. Aspects évolutifs.

Bulletin de la société francaise de mycologie médicale 18 : 283-288.

CHABASSE D., Cimon B., DE GENTILE L. $e BOUCHARA J. P., 1991 — Les mycoses transmises de l'animal

à l'homme. Modalités épidémiologiques, conduite pratique du diagnostic au laboratoire.

Revue française des laboratoires 228 : 77-84.

CHABASSE D. & CONTET-AUDONNEAU N., 1994 — Du saprophytisme au parasitisme, épidémiologie

des champignons kératinophiles isolés en France. Journal de mycologie médicale 4 : 80-89.

CHABASSE D., CHATRY D., BoussiN G., DE GENTILE L, Six P., Cimon В. & BOUCHARA J. P, 1995

Surveillance de la flore fongique des sols en milieu sportif. Santé publique 2 : 141-149.

CHERMETTE R., 1991 — Rôle des animaux de compagnie dans la dispersion des zoonoses d'origine

parasitaire. Rev. Sci. Tech. off Int. Epiz., 10 : 693-732

CHMEL L. & VOLLEKOVA A., 1981 — Développement du parasitisme chez les champignons kératino-

philes. Lyon Médical 245 : 37-40.

Source : MNHN. Paris

78 D. CHABASSE

Curran R. 5., 1985 — Taxonomy of Onygenaceae : Arthrodermataceae, Gymnoascaeae, Myxotri-

chaceae and Onygenaceae. Mycotaxon 24 : 1-216.

Dawson C. O. & GENTLES J. C., 1961 — The perfect stage of Keratinomyces ajelloi Vanbreusehem,

Trichophyton terrestre Durie et Frey and Microsporum nanum Fuentes. Sabouraudia 1 :

49-57.

Der Cas E. & VERNES A., 1986 — Parasitic adaptation of pathogenic fungi to mammalian hosts.

Critical reviews in microbiology 13 : 173-218.

Detanpt M. & NoLarD N., 1988 — Dermatophytes and swimming pools seasonal fluctuations.

Mycoses 31 : 495-500.

De Уковү C., 1968 — Ecologie de quelques Gymnoascaceae kératinophiles. Bulletin de l'académie

royale belge 54 : 1352-1368.

De VROEY C., 1970 — Contribution à l'étude des dermatophytes et autres Gymnoascacées. Annales de

la société belge de médecine tropicale 50 : 1-174.

икт E. B. & Frey D., 1957 — A new species of Trichophyton from New South Wales. Mycologia 49

:401-411

GARG A. K., GANDOTRA S., MUKERJI K. б. & PUGH G. J. F., 1985 — Ecology of keratinophilic fungi.

Proceedings of the indian academy of science. Plant sciences 94 : 149 — 163.

Gueno E., VILLARD J. & GUINETT R., 1985 — A new case of Anixiopsis stercoraria mycosis.

Discussion on its taxonomy and pathogenicity. Mykosen 28 : 430-436.

HELLUY J. R., PERCEBOIS C. & BURDIN J. C., 1965 — A propos de l'existence tellurique de

T. mentagrophytes. Annales de parasitologie humaine et comparée 40 : 383-389.

Hours R., ROUGET-CAMANA Y., LE FICHOUX Y., LANCASTRE F., BAZIN J. C., DENTEAU M. & BOOGNINI

J., 1972 — Isolement de Trichophyton mentagrophytes (Robin), Blanchard, 1986, Nanniz-

zia persicolor Stockdale 1987 et Trichophyton terrestre, Durie et Frey, 1957, du pelage

des rongeurs. Essai d'interprétation écologique. Annales de parasitologie humaine et com-

parée 47 : 422 -429.

KARLING J. S., 1946 — Keratinophilic chrytids. Rhizophydium keratinophilum n. sp. a saprophyte

isolated on human hair and its parasite : Phlyctidium mycetophagum sp. American journal

of botany., 33 : 751-757.

LECLERC M. C., PHILIPPE Н. & GUEHO E., 1994 — Phylogeny of Dermatophytes and dimorphic fungi

based on large subunit ribosomial RNA sequence comparisons Jounal de médecine vetéri-

naire 32 : 331-341.

Novak E. K. & GALGOCZY Y., 1966 — Notes on dermatophytes of soil origin. Mycopathologia et

mycologia applicata 28 : 289-296.

Oorscuor А. van, 1980 — Revision of Chry.

1-86.

OTCENASEK M. & Dvorak J., 1962 — The isolation of Trichophyton terrestre and other keratinophilic

fungi from small mammals of south eastern Moravia. Sabouraudia 2 : 111-113.

RIPPON J. W., LEE F. C. & MCMILLEN S., 1970 — Dermatophyte infection by Aphanoascus fulvescens.

Archives of dermatology 102 : 552-555.

SABOURAUD R., 1910 — Traité des teignes. Masson Ed., Paris, 811 p.

STOCKDALE P., 1988 — Sexual stimulation between Arthroderma simii (Stockdale, Mackenzie and

Austwick) and related species. Sabouraudia 6 : 176-181.

SZATHMARY S., 1940 — Trichophyton fluviale. Archives frangaises de dermatologie 181 : 192-203.

Торлко F, CRIAEO С & Unzi C., 1984 — À propos d'un cas de tinea pedis par Anixiopsis fulvescens

(Cooke) De Vries, var. fulvescens (Cooke) De Vries. Bulletin de la société francaise de

mycologie médicale 13 : 239-242.

Toma A., 1929 — Sur l'infection des cheveux « in vitro > par les champignons des teignes. Annales de

dermatologie et de syphiligraphie 10 : 641-643

VANBREUSEGHEM R., 1952 — Cycle biologique des dermatophytes et épidémiologie des dermatophy-

tes. Archives belges de dermatologie 8 : 268-276.

VANBREUSEGHEM R., 1952 — Technique biologique pour l'isolement des dermatophytes du sol

Annales de la société belge de médecine tropicale 32 : 175-178.

sporium and allied genera. Studies in Mycology 20 :

Source : MNHN, Paris

ADAPTATION PARASITAIRE DES CHAMPIGNONS KÉRATINOPHILES 79

VANBREUSEGHEM R., 1961 — La vie saprophytique des dermatophytes. Annales de dermatologie et de

syphiligraphie 87 : 481-492.

VANBREUSEGHEM R., 1952 — Intérét théorique et pratique d'un nouveau dermatophyte isolé du sol :

Keratinomyces ajelloii gen. nov. sp. Bulletin de la Classe des sciences. Académie royale de

Belgique, 5° série, 38 : 1067-1071.

VANBREUSEGHEM R. & De VROEY C., 1980 — Dermatophytic infection by Anixiopsis stercoraria in a

wildboar (sus scrofa). Mykosen 23 : 16-20.

Woopyer AJ., 1977 — Asymptomatic carriage of dermatophytes by cats. New Zealand veterinay

journal 2 : 67-69.

Source : MNHN. Paris

Cryptogamie, Му

NUEVOS DATOS SOBRE TRAMETES JUNIPERICOLA MANJÓN,

MORENO & RYVARDEN

М.М. BLANCO, J. CHECA y б. MORENO

Dpto. de Biología Vegetal. Universidad de Alcalá, Alcalá de Henares. 28871 Madrid. Espana.

RESUMEN — Se estudia el comportamiento en cultivo de Trametes junipericola y se compara con

Fomitopsis iberica, especie muy parecida macro y microscópicamente, resultando ambas especies

diferentes, Trametes junipericola posee podredumbre blanca, diferente de la de Fomitopsis iberica, que

posee podredumbre parda.

Hasta el momento actual Trametes junipericola se comporta como una especie endémica de

los sabinares de Guadalajara (España).

RÉSUMÉ — Le comportement en culture du Trametes junipericola a été étudié et comparé à

Fomitopsis iberica, espéce trés proche macro- et microscopiquement. Il en résulte qu'il s'agit de deux

espéces différentes, Trametes junipericola possédant un mycélium blanc, différent de celui du Fomi-

topsis iberica, qui est brun.

Jusqu'à aujourd'hui Trametes junipericola est considéré comme espéce endémique des

“sabinares” de Guadalajara (Espagne).

ABSTRACT — We study the conduct of Trametes junipericola in culture and we compare it to

Fomitopsis iberica, which is a species macro and microscopically very similar. Both species tending to

be different species. Trametes junipericola has got white rot, which is different from the rot of

Fomitopsis iberica, which has brown rot.

Trametes junipericolais still not known outside the type locality and must be looked upon as

a relict of a previously larger distribution.

KEY WORDS : Trametes junipericola, culture, chorology, taxonomy.

INTRODUCCIÓN

La provincia de Guadalajara en España tiene un gran interés botánico y mico-

lógico, porque conserva grandes extensiones de bosques muy bien conservados de Juni-

perus thurifera L., tanto en terreno ácido, como en terreno básico donde son más

abundantes, en este caso a veces mezclados con Quercus ¡lex subsp. ballota (Desf.) Samp.

Desde el punto de vista micológico, y aunque no aparecen elementos ectomicor-

rizógenos, hay una alta selectividad de especies pará: itas y saprófitas características y de

distribución ibero-norteafricana. A este respecto señalamos: Lenzitopsis oxycedri Malen-

gon & Bertault (Manjón & Moreno, 1981), Mycenella margaritifera (Maire in Kühner)

Source : MNHN, Paris

82 M. N. BLANCO, J. CHIECA y G. MORENO

Foto 1. — Trametes junipericola, cuerpos fructíferos

Maas G., (Moreno & Manjón, 1980), Trametes junipericola Manjón, Moreno & Ryvarden

(Manjón, Moreno & Ryvarden, 1984) y Xeromphalina junipericola G. Moreno &

Heykoop, (Moreno & Heykoop. 1996)

La descripción de Fomitopsis iberica (Melo & Ryvarden, 1989) y la posterior

publicación de una fotografía en color de dicha especie (Bernicchia, 1990), nos llamó la

atención por su semejanza con Trametes junipericola, ambas especies poseen un gran

parecido macro y microscópico y fructifican en coníferas mediterráneas. Se diferencian

por el tipo de podredumbre que es blanca en el primero y parda en el segundo, caracterís-

tica que separa principalmente los dos géneros.

Posteriormente Ryvarden & Gilbertson (1993) recopilan e iconografían estos

táxones, indicando la posibilidad de que Trametes junipericola presente podredumbre

blanca.

Estos motivos nos impulsaron a realizar pruebas enzimáticas en cultivo puro de

Trametes junipericola y comparar estas con la de Fomitopsis iberica, para asegurar la

posición adecuada de estas especies en los géneros en los que se habían descrito. Se ha

hecho especial hincapié en la prueba de la lacasa, ya que esta actividad es la que pone de

manifiesto el tipo de podredumbre.

Trametes junipericola Manjón, Moreno & Ryvarden, Bol. Soc. Micol. Castellana 8: 47.

1984.

Indicamos a continuación la diagnosis latina de Trametes junipericola, porque

esta especie se publicó en una revista, en aquella época con escasa difusión internacional,

y que actualmente esta agotada y es por lo tanto dificil de conseguir.

Source : MNHN, Paris

TRAMETES JUNIPERICOLA 83

Diagnosis: Fructificatio sessilis 4-10 cm, pileus cinereus et luridus, zonatus, veluti-

nus et hispidus, pori facie cinerei et ochracei angulati, 2-3 per mm, contextus albus et

cinereus. Systema hypharum trimiticum, hyphae generativae fibulatae, hyphae skeletales et

hyphae skeletico-coniunctivae hyalinae, sporae cylindricae, hyaline, non-amyloideae, 8-10 x

2,5-3 um.

Holotypus: España, Guadalajara, Ermita de los Enebrales de Tamajón 30T

VL7941, 5-V-1983, en troncos de Juniperus thurifera, leg. G. Moreno, АН 5739, isotypus en

MA-Fungi, en herbario 0 y en el herbario privado de H. Jahn.

Material estudiado: Trametes junipericola: España, Guadalajara, Tamajón,

Ermita de los Enebrales 30T VL7941, Tronco de Juniperus thurifera, leg. R. Galán & G.

Moreno, 23-11-1982, AH 5237; Ibidem leg. J. Checa, R. Galán & G. Moreno, 18-11-1983,

АН 5740. Ibidem, leg. E. Horak, С. Moreno & M. Lizárraga, 11-ХП-1995, AH 18382.

Fomitopsis iberica: Portugal, Ribatejo, Coruche, Pinus pinaster, leg. 1. Melo, M.

Correia & J. Cardoso, 27-X1-1986. LISU 3145 Paratypus. Ribatejo, Salvaterra de Magos,

Foros de Salvaterra, Pinus pinaster, em soca queimada, leg. I. Melo & J. Cardoso,

15-ХП-1992. O.

CULTIVOS

Se ha realizado el cultivo de Trametes junipericola en MEA (2% extracto de

agar-malta), a partir de cuerpos fructíferos obtenidos de material fresco.

Resultados obtenidos según los Códigos de Stalpers (1978):

1, 3, 7, (11), 13, (14), 16, (17), (18), 19, 22, 30, 36, 37, 39, 45, 46, 53, 60, (61), 75, (78), (84),

85,90.

Actividades enzimáticas: La prueba de actividad lacasa se ha llevado a cabo con

siringaldazina, -Naftol, guayacol y benzidina; la actividad tirosinasa, con p-cresol y

tirosina; la actividad peroxidasa con pirogalol, la actividad fosfatasa, con alfa-naftol

fosfato y la actividad esterasa, con alfa-naftol acetato, observando los cultivos a las 0, 3 y

24 h. Los resultados se ven reflejados en la tabla 1

| Oh 3h 24h

LACASA

Siringaldazina + -

alfa-Naftol - E

Guayacol - #

Benzidina +

TIROSINASA

| p-Cresol E 2 =

Тігоѕіпа + 2 s

PEROXIDASA E др 22

FOSFATASA - + Біз

ESTERASA - ү ал

жы.

Tabla 1 : Actividades enzimáticas de Trametes junipericola

Source : MNHN, Paris

84 M. N. BLANCO, J. CHIECA y G. MORENO

Asimismo, se han realizado cultivos y la actividad lacasa de Fomitopsis iberica a

partir de material de herbario, siguiendo la misma metodología, con los resultados

reflejados en la tabla 2.

LACASA - = z

Siringaldazina P = -

alfa-Naftol - - -

Benzidina - - =

Guayacol = = -

Tabla 2 : Actividades enzimáticas de Fomitopsis iberica

Observaciones: Las diferencias entre Trametes junipericola y otras especies de

Trametes (Т. hirsuta, T. pubescens, T. suaveolens, T trogii Berk, T. ljubarsky, T. tephroleuca

y T. palustris han sido puestas de manifiesto por Manjón, Moreno & Ryvarden (1984).

Una especie próxima es Pilatoporus maroccanus Kotlaba & Pouzar, descrita

recientemente por Kotlaba & Pouzar (1993), del Atlas medio (Marruecos) sobre Cupressus

sempervirens, comparte numerosos caracteres macro y microscópicos con Trametes juni-

pericola, el basidioma con la superficie velutina, los poros grisáceos de 1-3 por mm, las

medidas esporales (7-9,3 x 2,5-3,5 um) y el hábitat sobre tronco de Cupressaceae. La

principal diferencia es el tipo de podredumbre que Kotlaba & Pouzar (1993), indican es

parda.

Vampola (1996), indica respecto a P maroccanus Kotlaba & Pouzar que “Fur-

ther specimens from Morocco or other countries where Cupressus sempervirens occurs are

badly needed to prove that Р. maroccanus is really an independent species and not only a

synonym of Trametes suaveolens”.

El género Pilatoporus fue creado por Kotlaba & Pouzar (1990) para aquellas

especies del género Fomitopsis que poseen esporas de pared delgada y contexto blanco,

estos autores también describen el género Rhodoformes segregado de Fomitopsis para

aquellas especies con esporas de pared delgada y contexto rosado. Ryvarden & Gilbertson

(1993) no reconocen estos dos géneros, considerando únicamente el género Fomitopsis,

opinión que seguimos en este trabajo.

Vampola (1996) aporta nuevas localidades para Pilatoporus ibericus (Melo &

Ryvarden) Kotlaba & Pouzar en Europa y Asia (Repüblica Checa, Eslovaquia, Croacia e

Irán), y concluye su comportamiento como un hongo heterotálico y bipolar.

Finalmente podemos concluir después de los estudios enzimáticos realizados

que Trametes junipericola esta bien encuadrado en el género Trametes, porque posee

podredumbre blanca, y es diferente de Fomitopsis iberica, que produce podredumbre

parda.

Hasta el momento Trametes junipericola es un hongo endémico de los bosques

de Juniperus thurifera L. de la provincia de Guadalajara, no siendo raro su recolección en

los otoños lluviosos.

Source : MNHN. Paris

TRAMETES JUNIPERICOLA 85

AGRADECIMIENTOS

Trabajo en parte subvencionado por la DGICYT Proyecto PB 91-0165. Agra-

decemos a los conservadores de los herbarios LPS y O el envío del material solicitado.

BIBLIOGRAFÍA

BERNICCHIA A., 1990 — Polyporaceae s. L in Italia. Istituto di Patologia Vegetale. Univ. degli Studi

Bologna. Italia. 594 p.

KOTLABA F. & POUZAR Z., 1990 — Type studies of polypores described by A. Pilát — Ш. Ceska

Mykologie 44: 228-237.

KOTLABA Е & POUZAR Z., 1993 — Pilatoporus maroccanus sp. nov. A new polypore of the

.. Polyporus palustris group. Cryptogamie, Mycologie 14 (3): 215-218

MANION J. L. & MORENO G., 1981 — Estudios sobre Aphyllophorales 1. Fructificaciones sobre

. Juniperus. Anales del Jard'n botánico de Madrid 3702): 407-416.

MANION J. L., MORENO б. & RYVARDEN L., 1984 — Trametes junipericola Manjón, Moreno

& Ryvarden sp. nov. Boletin de la socidad micologica Castellana 8: 47-50.

MELO Y. & RYVARDEN L., 1989 — Fomitopsis iberica Melo et Ryvarden sp. nov. Boletin de la

sociedad Broteriana, ser. 2, 62: 227-230.

MORENO G. & HEYKOOP M., 1996 — Xeromphalina junipericola sp. nov. (Tricholomataceae,

Agaricales) from Spain. Zeitschrift für Mykologie 62(1): 37-41

MORENO G. & MANJÓN J. L., 1980 — Mycena margaritifera Maire dans le centre de l'Espagne

nouvelle espéce pour l'Europe. Documents mycologiques 10 (37-38): 85-87.

STALPERS J. A., 1978 — Identification of wood-inhabiting Fungi in pure culture, Studies in

mycology 16; 1-248.

VAMPOLA P., 1996 — New localities of Pilatoporus ibericus in Europe and Asia. Ceska Mykologie

49 (2): 85-90.

Source : MNHN, Paris

Source : MNHN. Paris

Cryptogamie, Mycol. 1997, 18 (1) : 87-90 87

DIDYMIUM CLAVODECUS (MYXOMYCETES)

UNA ESPECIE AMERICANA NUEVA PARA EUROPA.

M. LIZÁRRAGA, C. ILLANA, G. MORENO & A. CASTILLO

Dpto. Biología Vegetal. Facultad de Ciencias. Universidad de Alcalá,

28871 Alcalá de Henares, Madrid, Esp:

ABSTRACT : Didymium clavodecus, a species described from USA, has been found in Spain.

Didymium clavodecus is described and illustrated by L.M. and S.E.M. photographs.

RÉSUMÉ : Didymium clavodecus, espéce décrite aux USA, a été observée en Espagne. Didymium

clavodecus est décrite et illustrée par des photographies de microscopies optique et électronique à

balayage

KEY WORDS : Didymium clavodecus, Myxomycetes, Chorology, Taxonomy.

En la actualidad estamos realizando diferentes estudios, que nos permitan

comparar la diversidad existente en diferentes grupos de hongos en áreas mediterráneas de

Europa (Extremadura, España) y América (Baja California, México), para ello hemos

seleccionado áreas ocupadas por especies esclerófilas del género Quercus. Didymium

clavodecus es un elemento foliícola comün entre ambos continentes, su aparición en

España y por tanto en Europa es de importancia corológica para dicha especie origina-

riamente descrita de California (USA).

Las microfotografías fueron realizadas con un microscopio electrónico de bar-

rido (M.E.B.) Zeiss-DSM 950, y las muestras fueron preparadas con la técnica del punto

crítico. Para describir la ornamentación esporal con M.E.B. hemos seguido la terminolo-

gía propuesta por Rammeloo (1974; 1975). El material estudiado se encuentra depositado

en el herbario de la Universidad de Alcalá (AH).

Didymium clavodecus Whitney, Mycologia 71: 1257. 1979. Figs. 1-10

Material estudiado. ESPANA: En hojas de Quercus ilex L., Pozancos, Sigüenza,

Guadalajara, leg.. J. Alvarez, 10-111-1990, AH 12321. MÉXICO: En hojas y madera de

Quercus agrifolia Nee, ctra. Tecate-Ensenada km. 65 (Ejido Ignacio-Zaragoza), Baja

California, 6-11-1993, leg. M. Lizárraga, С. Moreno y C. Шапа, AH 15883, 15895 y 18627.

En hojas de Quercus agrifolia Nee., ctra. Tecate-Mexicali (Cañada Verde), Baja California,

6-11-1993, leg.. M. Lizárraga, С. Moreno y C. Шапа, AH 15824, 15827, 15828, 15923,

15924, 15926, 15927, 15935, 15943, 15950 y 18620.

Source : MNHN, Paris

88 M. LIZÁRRAGA, C. ILLANA, G. MORENO & A. CASTILLO

Fructificaciones formando esporocarpos de 0,4-3 mm de diám. a cortos plasmo-

diocarpos, de 3-5 x 1,5-2 mm, sésiles, de pulvinados a anulados (figs. 1-2). Peridio

iridiscente, membranoso, cubierto por cristales calcáreos blancos, (figs. 3-4), dehiscencia

irregular. Columela central, blanca, formada por depósitos calcáreos. Hipotalo membra-

noso. Capilicio de 1,5-2,5 um de diám., abundante, de filamentos gruesos y rígidos, pardo

violáceo oscuro, muy anastamosados, formando una red que conecta el centro de la

fructificación con el peridio (figs. 5-6). Esporas de 12-14 um de diám., poligonales, negras

en masa, pardo-violáceo oscuras al M.O., con un retículo que recuerda a las esporas de

Physarum straminipes Lister, (figs. 7-9), ornamentadas con espinas capitadas de 1-2 um de

largo, que al M.E.B. forman características “pila” (fig. 10).

Macroscópicamente Didymium clavodecus puede ser confundido macroscópica-

mente con otras especies sésiles del género Didymium, especialmente con formas sentadas

de D. squamulosum (Alb. & Schwein.) Fr. Sin embargo, D. clavodecus posee un carácter

ünico en el género, cómo es la posesión de una espora de forma poligonal, ornamentada

con espinas capitadas en forma de clavo, que son perceptibles al M.E.B. como “pila”, y

que varían en nümero y densidad (figs. 8-9).

No se conocen citas de Didymium clavodecus en Europa (Neubert & al., 1995).

Ünicamente se conoce esta especie de California, USA. (Whitney, 1979), y más reciente-

mente de Baja California, México (Lizárraga & al., 1997), siempre sobre hojas de Quercus

sp.

El material español ha sido comparado con abundantes muestras recogidas por

nosotros, en áreas mediterráneas de Baja California. Un estudio completo de Didymium

clavodecus de Baja California ha sido realizado por Lizárraga & al. (1997).

AGRADECIMIENTOS

Este trabajo ha sido posible gracias al Proyecto de Investigación incluido en el

Programa de Cooperación con Iberoamérica, Ministerio de Educación y Ciencia, Subdi-

rección General de Cooperación Internacional, España, y al Proyecto de Investigación de

la DGICYT PB 91-0165. M. Lizárraga agradece al Consejo Nacional de Ciencia y

Tecnología (CONACYT) de México su ayuda para la realización de la tesis doctoral en

Baja California.

Agradecemos la ayuda de J. Martínez y A. Priego, del Servicio de Microscopía

Electrónica de la Universidad de Alcalá de Henares.

BIBLIOGRAFÍA

LIZÁRRAGA M., MORENO G. & ILLANA C., 1997 — The Myxomycetes from Baja California

(México). I. Mycotaxon (en prensa).

NEUBERT H., NOWOTNY W., K. BAUMANN & MARX H., 1995 — Die Myxomyceten. Band 2.

Physarales. Karlheinz Baumann Verlag. Gomaringen. 368 pp.

RAMMELOO J., 1974 — Structure of the epispore in Trichiaceae (Trichiales, Myxomycetes), as seen

with the scanning electron microscope. Bulletin de la société royale de botanique de Belgique

107: 353-359.

RAMMELOO J., 1975 — Structure of the epispore in the Stemonitales (Myxomycetes) as seen with

the scanning electron microscope. Bulletin du jardin botanique de Belgique 45: 301-306.

WHITNEY K. D., 1979 — A new foliicolous Didymium from Northern California. Mycologia 71:

1256-1261.

Source : MNHN. Paris

1-6. — Didymium clavodecus (AH 12321). 1: fructificaciones.

esporocarpo sésil con cristales

sobre el peridio. 3: peridio cubierto con cristales (parte superior), con un filamenté del

capilicio unido (M.E.B.). 4: cristales calcáreos (M.E.B.). 5: filamentos del capilicio que forman una

red (M.E.B.). 6: detalle del capilicio (M.E.B.).

се

MNHN, Paris.

90 M. LIZARRAGA, C. ILLANA, G. MORENO & A. CASTILLO

Figs. 7-10. — Didymium clavodecus (AH 12321). 7: grupos de esporas (M.E.B.). 8-9: esporas

(M.E.B.). 10: detalle de la ornamentación esporal formada por "pila" (M.E.B.).

Source : MNHN,

INSTRUCTIONS AUX AUTEURS

Cryptogamie, Algologie, Cryptogamie, Bryologie-Lichénologie et Cryptogamie, Mycologie publient

des articles originaux ou des articles de synthése en systématique, biologie et écologie des

cryptogames (algues, bryophytes et lichens, champignons, respectivement). Les manuscrits rédigés

en frangais, en anglais, en allemand, en espagnol et en italien, sont pris en considération dans la

limite des possibilités de la Rédaction à trouver des lecteurs compétents dans ces langues. Les

auteurs n'écrivant pas dans leur langue doivent, avant soumission, faire relire leur texte par une

personne dont la langue du manuscrit est la langue maternelle. Il n'y a pas de limite au nombre

de pages. La publication d'un article sera facilitée si les auteurs suivent soigneusement les

instructions ci-après.

Les articles sont lus et critiqués par des rapporteurs spécialistes des domaines concernés. Ces

rapporteurs sont choisis par la rédaction. La publication définitive des articles dépend de la

rédaction. Les manuscrits non conformes aux instructions qui suivent sont retournés aux auteurs

pour étre corrigés, avant leur soumission au Comité de Lecture. Dés leur acceptation, les droits

de reproduction des articles deviennent la propriété de la revue.

TEXTE. Les manuscrits doivent étre fournis en triple exemplaire (l'original et deux copies),

dactylographiés sur une seule face, en double interligne (y compris les références et les légendes),

sans rature ni surcharge, sans mots coupés et avec une marge gauche de 4 cm. Pour raccourcir

les délais de parution, il est vivement recommandé d'envoyer à la rédaction la version finale de

l'article enregistrée sur disquette utilisable sous DOS (IBM) ou Macintosh ; le logiciel MS WORD

est conseillé.

Chaque manuscrit, paginé, devra comporter, dans l'ordre :

— une page comportant le titre de l'article et son éventuelle traduction en anglais comprendra les

noms, prénoms et adresses des auteurs (y compris, dans la mesure du possible, les numéros de

télécopie et de courrier électronique), l'auteur à qui la correspondance doit étre adressée devant

&tre précisé ; le titre courant (haut de page) de 50 signes au maximum ; une liste de mots-clés ;

deux résumés, l'un en francais (en cas de nécessité, la rédaction peut aider à sa traduction),

l'autre en anglais, chacun d'eux d'environ 180 mots ou 15 lignes (non limitatif) et, éventuellement,

un troisième dans la langue de l'article ; ils doivent faire ressortir les résultats essentiels exposés

dans le texte ;

le texte proprement dit, suivi des légendes des figures, des planches et des tableaux, dans cet

ordre.

La présentation du texte devra faire apparaitre clairement les subdivisions appropriées

à la nature de l'article (par ex. Introduction, Matériels et Méthodes, Résultats, Discussion,

Remerciements, Références), leur hiérarchie, ainsi que le début des paragraphes. Les symboles,

unités et la nomenclature doivent suivre les normes internationales : le système SI, en particulier,

doit étre utilisé. La premiére mention du nom latin d'une espéce doit étre accompagnée des noms

des autorités taxinomiques. Un fascicule du journal pourra étre consulté à titre d'exemple pour

une plus ample information sur les conventions suivies par la revue. Les légendes des figures et

des tableaux (dans la langue du manuscrit et en anglais) doivent étre explicites.

RÉFÉRENCES. — La liste bibliographique devra se faire par ordre alphabétique des auteurs et

chronologique par auteurs sans tenir compte des auteurs secondaires. Les titres des périodiques

devront être cités en entier ; les ouvrages seront cités selon F.A. Stafleu & R.S. Cowan, 1976.

Taxonomic literature. Ed. 2. Utrecht/Antwerpen : Bohn, Scheltema & Holkema. Les références

devront étre présentées selon les modéles suivants (noter que les noms des auteurs Chinois sont

écrits en entier et dans le style du changement national effectué en Chine en 1987 ; par ex. Chang

C.F. devient Zhang Jungfu) :

Source : MNHN, Paris

AJISAKA T., NORO T., TRONO Jr. G.C., YOUNG-MENG CHIANG & YOSHIDA T., 1994 — Several

Sargassum species (subgenus Sargassum) in East Asia with furcately branching leaves. In: Abbott LA. (ed.),

Taxonomy of Economic Seaweeds. IV. La Jolla, California Sea Grant College, University of California, pp. 9-22.

ALBRECHT A. & REISE K., 1994 — Effects of Fucus vesiculosus covering intertidal mussel beds in the

Wadden Sea. Helgolánder Meeresuntersuchungen 48 (2-3): 243-256.

MAGGS C.A. & HOMMERSAND M.H., 1993 — Seaweeds of the British Isles. | Rhodophyta. Part 3A.

Ceramiales. London, HMSO Books, The Natural History Museum, 464 p.

MONTAGNE C., 1838 — Centurie des plantes cellulaires exotiques nouvelles. Annales des Sciences Naturelles,

Botanique, sér. 2, 9; 38-57.

Les renvois à la liste bibliographique se feront par le nom de l’auteur et l’année de publication ;

utiliser “er al.” lorsque l'article est signé par plus de deux auteurs et “&” entre deux auteurs (par

ex. Montagne, 1838; Maggs & Hommersand, 1993; Ajisaka er al., 1994).

ILLUSTRATIONS. — Toutes les illustrations doivent étre montées sur papier blanc rigide ; les

noms des auteurs et le numéro des figures doivent étre indiqués au verso de chacune d'elles. Toutes

les illustrations au trait doivent étre originales, à l'encre de Chine, les numéros et les lettres doivent

être effectués à l'aide de lettres de transfert. Les dimensions des originaux ne devront pas excéder

le triple de celle de leur reproduction définitive (justification de la revue : 12,5 x 18 cm). Les

illustrations au trait et les tableaux, doivent étre des originaux de qualité suffisante pour la

reproduction directe en offset. Les auteurs choisiront l'épaisseur des traits et la taille des caractéres

en fonction de la réduction éventuelle des illustrations. Les illustrations photographiques doivent

étre fournies en trois exemplaires originaux (ou un original et deux photocopies laser). Les

documents photographiques doivent étre montés par planches, à la taille de leur reproduction

finale (justification maximale : 12,5 x 18 cm). Toutes les figures devront comporter les échelles

(les grandissements x... sont prohibés), les symboles nécessaires à leur compréhension, et étre

numérotées dans l'ordre d'appel dans le texte. La publication de planches en couleurs est à la

charge des auteurs.

ÉPREUVES. — Les épreuves devront étre vérifiées et retournées à la rédaction dans les 48 heures

suivant leur réception, par courrier et, si nécessaire, par fax. A ce stade, les possibilités de

correction sont limitées aux fautes de frappe. Si des modifications aux illustrations s'avéraient

nécessaires, le coüt du clichage des nouveaux originaux serait à la charge des auteurs. Si les

épreuves ne sont pas retournées dans les délais, l'article paraitra sans les corrections des auteurs.

TIRÉS-À-PART : limités à 150, dont 25 gratuits. Un bon de commande pour les tirés-à-part est

joint à l'envoi des épreuves.

Cryptogamie

Association des Amis des Cryptogames

(ADAC), Muséum National d'Histoire Natu-

relle, Laboratoire de Cryptogamie (PC), 12, rue

Buffon 75005 Paris France

Directeur de publication

Bischler

Hélène Causse-

Cryptogamie, Algologie - Rédacteur : Bruno de

Reviers

Tél. (33) 1 40 79 31 98 - Fax (33) 1 40 79 35 94

= E-mail reviers@mnhn.fr

Cryptogamie, Bryologie-Lichénologie - Rédac-

teur : Denis Lamy

Tél. (33) 1 40 79 31 84 - Fax (33) 1 40 79 35 94

= E-mail lamy@mnhn.fr

Cryptogamie, Mycologie - Rédacteur : Bruno

Dennetiére

Tél. (33) 1 40 79 31 87 - Fax (33) 1 40 79 35 94

= E-mail cryplich@mnhn.fr

Source : MNHN, Paris

INSTRUCTIONS TO AUTHORS

Cryptogamie, Algologie; Cryptogamie, Bryologie-Lichénologie; and Cryptogamie, Mycologie pub-

lish original papers, and reviews, on the systematics, biology and ecology of cryptogams (algae;

bryophytes and lichens; and fungi, respectively). Manuscripts written in French, English, German,

Italian and Spanish are considered providing suitable referees fluent in the language of the

manuscript are available. Authors not writing in their first language should have manuscripts

checked for grammar and syntax by a suitable person before submission. There are no page limits

for papers. Publication will be facilitated if authors check carefully that the manuscript and

illustrations meet the requirements outlined below.

Papers are reviewed by referees in whose field the paper lies. The choice of reviewers is at the

discretion of the Editor. Final responsibility for the publication of papers rests with the Editor.

Manuscripts that do not conform to instructions which follow will be returned for correction prior

to review. On acceptance, papers become the copyright of the journal.

TEXT. — Manuscripts should be submitted to the Editor in triplicate, including the original copy.

All parts of the manuscript must be typed double-spaced (including references and legends), on

one side of the paper, with left margin of 4 cm. To minimize delays in publication, authors should

send the final version of their manuscripts on floppy disks (5 1/4, 3 1/2) using the DOS (IBM) or

Macintosh format. In addition, the use of MS WORD would be appreciated.

Each manuscript should include, in order:

— A title page containing the title of the paper and, if necessary, its English translation; the

complete name and address of each author (including, as much as possible, fax number and e-mail

address), and author to whom correspondence should be sent; a running title of less than

50 letters, and a list of key words;

— Two abstracts, the first in French (if necessary, some help can be provided by the Editorial

Office), the second in English (a third one in the language of the text is accepted), each of no more

than 180 words or 15 lines, summarizing the major results of the paper;

— The main text, followed by references, legends for figures, and tables, in that order.

The text should normally be divided into sections (e.g. Introduction, Materials and

Methods, Results, Discussion, Acknowledgements, References) appropriate to the nature of the

paper. Indent the first line of all paragraphs. Symbols, units and nomenclature should conform

to international usage. The SI system should be used throughout. The first mention of the Latin

name of a species in the text should be accompanied by the nomenclatural authorities. Consult

the current issue of the Journal for style of headings, sub-headings and other conventions.

Legends for figures, plates and tables should be self-explanatory, and written in the language of

the text and in English if it is not the language used for the text.

REFERENCES. — References should be arranged alphabetically and then chronologically by

author, Journals titles should be cited in full; and books, cited according to F.A. Stafleu & R.S.

Cowan, 1976..., Taxonomic literature. Ed. 2. Utrecht/Antwerpen: Bohn, Scheltema & Holkema.

Conventions of style are provided in the following examples (note that names of Chinese authors

are written in full and listed in the style in accordance with the national change made in China

in 1987; e.g. Chang C.F. becomes Zhang Junfu):

AJISAKA T., NORO T., TRONO Jr. G.C., YOUNG-MENG CHIANG & YOSHIDA T., 1994 — Several

Sargassum species (subgenus Sargassum) in East Asia with furcately branching leaves. In: Abbott LA. (ed),

Taxonomy of Economic Seaweeds. IV. La Jolla, California Sea Grant College, University of California, pp. 9-

ALBRECHT A. & REISE K., 1994 — Effects of Fucus vesiculosus covering intertidal mussel beds in the

Wadden Sea. Helgolánder Meeresuntersuchungen 48 (2-3): 243-256.

Source : MNHN, Paris

MAGGS C.A. & HOMMERSAND M.H., 1993 — Seaweeds of the British Isles. 1 Rhodophyta. Part 3A

Ceramiales. London, HMSO Books, The Natural History Museum, 464 p.

MONTAGNE C., 1838 — Centurie des plantes cellulaires exotiques nouvelles. Annales des Sciences Naturelles,

Botanique, sér. 2, 9: 38-57.

In the text, references should be cited by the name of the author and the year of publication; use

"et al.”, for more than two authors, and use “8” between two authors (e.g. Montagne, 1838;

Mages € Hommersand, 1993; Ajisaka ег al., 1994).

ILLUSTRATIONS. — All illustrations should be mounted on white light-weight carbocard, with

author's names and figure numbers on the verso. Each line drawing should be original, clearly

drawn with black India ink, and of good quality; letters and numerals must be made with a

lettering device (not with a type writer). Originals should not be more than three times the size

of the final reduction (12.5 x 18 cm). Both drawings and tables should be ready for direct

reproduction by offset. Authors should choose very carefully the thickness of lines, and character

izes, corresponding to final reduction. Three copies of all photographic illustrations (or an

al and to laser photocopies) are required. Photographic figures should be ready for

Size reproduction: maximum page size is 12.5 x 18 cm. All figures should include scale bars

and symbols necessary for their understanding, and they should be numbered consecutively,

according to the order cited in the text. Color plates will be published only at the authors expense.

PROOFS. — Proofs should be checked and returned by airmail (if necessary, by fax) to the Editor

within 48 hours of receipt. At this stage, corrections should be restricted to those arising from

typesetting errors. If changes to illustrations are necessary at proof stage, new originals would

have to be supplied and the cost for re-photographing will be charged to author at the discretion

of the Editor. If proofs are not returned promptly, the article will be published without the

author's corrections.

REPRINTS: not more than 150, of which 25 are free copies. Reprint orders are enclosed with

proofs

Cryptogamie

Association des Amis des Cryptogames

(ADAC), Muséum National d'Histoire Natu-

relle, Laboratoire de Cryptogamie (PC), 12, rue

Buffon 75005 Paris France

Director of the journal: Héléne Causse-Bischler

Cryptogamie, Algologie - Editor: Bruno de

Reviers

Tél. (33) 1 40 79 31 98 - Fax (33) 1 40 79 35 94

- E-mail reviers@mnhn.fr

Cryptogamie, Bryologie-Lichénologie - Editor:

Denis Lamy

Tél. (33) 1 40 79 31 84 - Fax (33) 1 40 79 35 94

= E-mail lamy@mnhn.fr

Cryptogamie, Mycologie - Editor: Bruno Den-

netière

Tél. (33) 1 40 79 31 87 - Fax (33) 1 40 79 35 94

= E-mail cryplich@mnhn.fr

Commission paritaire 16-1-1986 - N° 58611 - Dépôt légal 1% trimestre 1997 - Imprimerie F. Paillart

- Sortie des presses le 29 mars 1997 - Imprimé en France

Éditeur

A.D.A.C. (Association des Amis des Cryptogames)

Président : D. Lamy ; Secrétaire : B. Dennetière

Trésorier

М" E. Bury; Directeur de la publication

Н. Causse

Source - MNHN. Paris

Société Francaise de Systématique <3 |

La Société Française de Systématique réunit les systématiciens ou les personnes

intéressées par la Systématique et les informe en publiant un Bulletin. Elle convie ses

membres à des colloques annuels transdisciplinaires, au cours desquels les systématiciens

et d'autres scientifiques peuvent s'exprimer et débattre.

Cotisation annuelle: 100F

Demande d'adhésion à adresser au:

Secrétariat de la Société Francaise de Systématique, 45 rue Buffon, F-75005 Paris.

CCP 7-367-80 D PARIS.

La Société édite aussi la série Biosystema.

Prix TTC du Biosystema (France, Etranger): 150 FF, membre SFS : 100 FF.

Biosystema 1 - Introduction à la systématique zoologique - (Concepts. Principes, Méthodes) par

L. Matile, P. Tassy & D. Goujet. 1987.

Biosystema 2 - Systématique Cladistique - Quelques textes fondamentaux. Glossaire. Traduction

et adaptation de D. Goujet, L. Matile, P. Janvier & J.P. Hugot. 1988

Biosystema 3 - La systématique et l'évolution de Lamarck aux théoriciens modernes. par

S. Lovtrup. 1988.

Biosystema 4 - L'analyse cladistique: probléme et solutions heuristiques informatisées, par

M. d'Udekem-Gevers. 1990.

Biosystema 5 - Les introuvables de J.B. Lamarck- Discours d'ouverture du cours de zoologie et

articles du Dictionnaire d'Histoire naturelle. Edition préparée par D. Goujet. 1990.

Biosystema 6 - Systématique et Ecologie, par R. Barbault, Cl. Combes, F. Renaud, N. Le Brun

& A. Dubois. Edition coordonnée par J.P. Hugot. 1991.

Biosystema 7 - Systématique et Biogéographie Historique. Textes historiques et

méthodologiques. Traduction et adaptation de P. Janvier, L. Matile & Th. Bourgoin.

1991.

Biosystema 8 - Systématique et Société. Edition coordonnée par G. Pasteur. 1993

Biosystema 9 - Les Monocotylédones, par J. Mathez. 1993.

Biosystema 10 - Systématique botanique : problèmes actuels. Edition coordonnée par O. Poncy.

1993,

Biosystema 11 - Systématique et Phylogénie: modéles d'évolution biologique. Edition

coordonnée par P. Tassy et Н. Lelièvre. 1994.

Biosystema 12 - Phylsyst: logiciel de reconstruction phylogénétique, par I. Bichindaritz,

S. Potter & B. Sigwalt +. 1994.

Biosystema 13 - Systématique et Biodiversité. Edition coordonnée par Th. Bourgoin. 1995. В

Biosystema 14 - Systématique et Informatique. Edition coordonnée par J. Lebbe, en préparation.

Le Conseil de la SFS. XII 1995

Source : MNHN, Paris