Taxonomic revisions in both the teleomorphic(sexual) and anamorphic (asexual) forms of the genusCerutocystis Ellis & Halstead are chronicled in thisreview. Recognized species associated with Den-droctonus Erichson bark beetles are summarized, andseveral species that have been published as recom-binations, species that were previously described asOphiostoma, and species that have not yet beenrecombined are listed.

November 199 1

A Synopsis of the Taxonomic Revisions in the GenusCerutocystis Including a Review of Blue-Staining Species

Associated with Dendroctonus Bark BeetlesThelma J. Perry

INTRODUCTION

Because the fungi of the genera Cerutocystis Ellis &Halstead (1890) and Ophiostoma H. & P. Sydow(1919) are wood stainers and plant pathogens, theyare economically important. Among the best knownare the blue stain fungus Ophiostoma minus (Hedge.)H. & P. Sydow (= Ceratocystis minor rHedgc.1 Hunt),the oak wilt fungus Cerutocystis fugucearum (Bretz)Hunt, the Dutch elm disease fungus Ophiostoma ulmi(Buism.) Nannfeldt (= Ceratocystis ulmi [Buism.] C.Moreau), the sweet potato rot fungus Cerutocystisfimbriata Ellis & Halstead, and the root pathogenLeptographium wageneri (Kendr.) Wingfield (= Verti-cicladiella Wageneri: Kendrick).

Insects disperse these fungi’s spores that areproduced in a sticky substance by both the teleo-morphs (sexual form) and the anamorphs (asexualform) (table 1) and readily adhere to beetle body parts.Most Scolytidae (bark beetles) transport one or moreCeratocystis species. These fungi are found in or nearbark beetle galleries. Some species may assist thebeetles as tree drying agents(Bramble and Holst 1940;Nelson 1934 ) or may elicit tree defensive reactions(Mattson and others, 1988). Some Cerutocystis speciesare ambrosia fungi, which serve as food for developingbeetle larvae (Whitney 1971, 1982). Other speciescompete with immature beetles for space and nutri-ents in the phloem (Barras 1970; Bridges and Perry1985; Franklin 1970).

The fungal-bark-beetle-host associations havebeen listed in publications such as Barras and Perry(1975), Upadhyay (19811, Whitney (1982), and Beaver(1989). Because of recent taxonomic revisions of thegenus Cerutocystis in both the anamorphic and teleo-morphic forms, there is a need for an updated bibliog-raphy of fungal-insect-host interactions. Harrington(1988) lists 34 Leptographium Lagerberg & Melin spe-cies, 20 of which have a known Ophiostoma teleo-morph, and 30 species of bark beetles that vectorthese Ophiostoma I Leptographium fungi. Literaturesearches for bark beetle-fungal associations should

not be limited to the keyword Ceratocystis but shouldalso include the fungi Ophiostoma and Cerutocystiop-sis Upadhyay & Kendrick.

Because of the disagreements about the taxonomicstatus of the genus Cerutocystis, various reclassitica-tions have contributed to the use of incorrect names inthe fungal-bark beetle literature. My objectives are (1)to chronicle the events that led to revisions in both theteleomorphic genera and the anamorphic form-generathat make up the genus Cerutocystis and (2) to pro-vide a quick and easy guide to the accepted taxonomicnames of these fungi.

Most of the recent taxonomic disagreements occurat the level or order, family, and genus (fig. 1). In1980, Benny and Kimbrough described the acceptedorder as Ophiostomatales with four genera: Ophios-toma, Ceratocystis, Ceratocystiopsis, and Sphaeronae-mella Karsten. In 1932, Nannfeldt had erected thefamily Ophiostomataceae for the genus Ophiostoma.This is the traditionally accepted family for the genusCeratocystis and the synonomized genera EurophiumParker and Ophiostoma. Von Arx and van der Walt(1988) accepted the order Ophiostomatales with thefamily Ophiostomataceae representing the generaCeratocystiopsis, Europhium, and Ophiostoma. Theyaccepted the anamorphic form-genus Ruffaeleu v. Arx& Hennebert and 16 other form-genera that aredescribed in Upadhyay (1981) (table 21, with the ex-ception of Verticicladiella S. Hughes, which issynonomized as Leptographium (Wingfield 1985). In1988, Wingfield and others described Knoxdaviesia, anew anamorph of the genus Cerutocystiopsis. Von Arxand van der Walt (1988) established the familyPyxidiophoraceae for species with the anamorphChalara (Corda) Rabenhorst, representing the generaCeratocystis Ellis & Halstead, Cryptendoxyla Malloch& Cain, Mycorhynchidium Malloch & Cain, andPyxidiophora Brefeld & Tavel.

At the generic level, three genera are currently ac-cepted: (1) Ceratocystis Ellis & Halstead, containingonly those species with the form-genus Chalara(Corda), Rabenhorst; (2) Ophiostoma H. & P. Sydow;

Thelma J. Perry is a microbiologist at Forest Insect and Disease Research, U.S. Department of Agriculture, Forest Service, SouthernForest Experiment Station, Pineville, LA 71360.

1

Table I,-Dendroctonus species (Wood 1982) and associated blue-staining fungi

Dendroctonus sp. Fungal associate Pu.blication

DendroctonusadjunctusBlandford

=D.convexifronsHopkins

DendroctonusbrevicomisLe Conte

DendroctonusfrontalisZimmermann

Dendroctonus Ophiostoma ips (Rumb.) Nannfeldtjeffreyi = Ceratocystis ips (Rumb.) C. MoreauHopkins = Ceratostomella montia Rumbold

DendroctonusponderosaeHopkins

=D.monticolaeHopkins

Ophiostoma stenoceras (Robak) Melin & Nannfeldtz Ceratocystis stenoceras (Robak) C. Moreau= Ceratocystis gossypina var. robusta Davidson= Ceratostomella stenoceras Robak= Ceratocystis gossypina Davidson

Ophiostoma adjuncti (David%) HarringtonI Ceratocystis adjuncti Davidson

Leptographium pyrinum DavidsonLeptographium sp. Lagerberg & Melin

Ophiostoma minus H. & P. Sydow= Ceratocystis minor (Hedge.) Hunt= Ceratostomella minor Hedgcock= Ceratostomella pini Munch= Ceratostomella exigua Hedgcock= Ceratostomella pseudotsugae Rumbold

Ophiostoma nigrocarpum (Davids.) de Hoog= Ceratocystis nigrocarpa Davidson

Ophiostoma minus H. & P. Sydow= Ceratocystis minor (Hedge.) Hunt= Ceratostomella minor Hedgcock= Ceratostomella pini Munch= Ceratostomella exigua Hedgwck= Ceratostomella pseudotsugae Rumbold

Ophiostoma nigrocarpum (Davids.) de Hoog3 Ceratocystis nigrocarpa Davidson

Ceratocystiopsis ranaculosus Perry & Bridges

Ophiostoma huntii (Robins.-Jeff.) de Hoog & SchefferI Ceratocystis huntii Robinson-Jeffrey & Grinchenko

Ophiostoma minus H. & P. SydowE Ceratocystis minor (Hedge.) Hunt= Ceratostomella minor Hedgcock= Ceratostomella pini munch= Ceratostomella exigua Hedgcock= Ceratostomella pseudotsugae Rumbold

Ophiostoma ips (Rumb.) Nannfeldt= Ceratocystis ips (Rumb.) C. Moreau= Ceratostomella montia Rumbold

Ophiostoma piliferum (Fries) H. & P. SydowE Ceratocystis pilifera (Fries) C. Moreau= Sphaeria pilifera Fries= Ceratocystis shrenkiana (Hedge.) C. Moreau

Ophiostoma clavigerum (Robins.-Jeff. & Davids.) Harrington= Ceratocystis clavigem (Robins.-Jeff. & Davids.) Upadhyay-I Europhium clavigerum Robinson-Jeffrey & Davidson

Ceratocystiopsis minuta (Siem.) Upadhyay & KendrickI Ceratocystis minuta (Siem.) Hunt= Ophiostoma minutum Siemaszko

Leptographium Lagerberg & Melin

Davidson 1971

Davidson 1978Davidson 1978Harrington 1988

Mathre 1964

Rumbold 1931

Rumbold 1936

Davidson 1966

Rumbold 1931

Rumbold 1936

Davidson 1966Bridges & Perry 1987

Mathre 1964Rumbold 194 1

Robinson-Jeffrey & Grinchenko 1964

Rumbold 1931

Rumbold 1936

Mathre 1964Rumbold 1941

Mathre 1964

Robinson-Jeffrey & Davidson 1964

Mathre 1964

Harrington 1988, Robinson 1962

2

Table I.-Dendroctonus species (Wood 1982) and associated blue-staining fungixontinued

Dendroctonus sp. Fungal associate Publication

DendmctonuspseudotsugaeHopkins

Dendroctonusrufipennis(Kirby)

=D.engelmanniHopkins

=D.piceaperdaHopkins

Dendroctonusterebmns(Olivier)

Dendroctonus Ophiostoma piliferum (Fries) H. & P. Sydowvalens I Ceratocystis pilifera (Fries) C. MoreauLA-S Conte = Sphaeria pilifera Fries

DendroctonusErichson

Ophiostoma minus H. & P. Sydow3 Ceratocystis minor (Hedge.) Hunt= Ceratostomella minor Hedgcock= Ceratostomella exigua Hedgcock= Ceratostomella pini Munch= Ceratostomella pseudotsugae Rumbold

Leptographium abietinum (Peck) Wingfield= Verticicladiella abietina (Peck) S. Hughes= Sporocybe abietina Peck

Leptographium sp. A

Ophiostoma bicolor Davidson & Wells= Ceratocystis bicolor (Davids. & Wells) Davidson

Ophiostoma piceaperda (Rumb.) von ArxI Ceratocystis piceaperda (Rumb.) C. Moreau= Ceratostomella piceaperda Rumbold= Ceratocystis europhioides Wright & Cain

O p h i o s t o m a p e n i c i l l a t u m (Grosm.) Siemaszko= Ceratocystis penicillata (Grosm.) C. MoreauI Ceratostomella penicillata Grossmann= Ophiostoma truncicola Davidson

Ophiostoma coerulescens (Munch) NannfeldtI Ceratocystis coerulescens (Munch) BakshiI Endoconidiophora coerulescens Munch

Leptographium abietinum (Peck) WingfieldI Verticicladiella abietina (Peck) S. Hughes= Sporocybe abietina Peck

Leptographium engelmanni Davidson

Ophiostoma ips (Rumb.) Nannfeldt= Ceratocystis ips (Rumb.) C. Moreau= Ceratostomella montia Rumbold

Leptographium procerum (Kendr.) Wingfield= Verticicladiella procera Kendrick

Leptographium terebrantis Barras & Perry

Ophiostoma wageneri (Goheen & Cobb) Harrington= Ceratocystis wageneri Goheen & Cobb

Ophiostoma ips (Rumb.) Nannfeldt= Ceratocystis ips (Rumb.) C. Moreau= Ceratostomella montia Rumbold

Leptographium terebrantis Barras & PerryOphiostoma piceaperdum (Rumb.) von Am

I Ceratocystis piceaperda (Rumb.) C. Moreau= Ceratostomella piceaperda Rumbold= Ceratocystis europhioides Wright & Cain

.Ceratocystis leucocarpa Davidson*Ophiostoma aureum (Robins-Jeff. & Davids.) Harrington

= Ceratocystis aurea (Robins-Jeff. & Davids.) Upadhyayz Europhium aureum Robinson-Jeffrey & Davidson

Ophiostoma robusturn (Robins.-Jeff. & Davids.) Harrington= Ceratocystis robusta (Robins.-Jeff. & Davids.) Upadhyay5 Europhium robusturn Robinson-Jeffrey & Davidson

Mathre 1964

Rumbold 1936Rumbold 1936Wingfield 1983

Harrington 1988

Davidson 1955Davidson 1958

Mathre 1964Rumbold 1936Wright & Cain 1961

Davidson 1955

Davidson, 1955Wingfield 1983, Harrington 1988

Davidson 1955

Rane & Tatter 1987

Wingfield 1983, Harrington 1988

Barras & Perry 1971

Goheen & Cobb 1978

Rumbold 1931Wingfield 1983

Rumbold 1931Wright & Cain 1961

Davidson 1966

Robinson-Jeffrey & Davidson 1968

Robinson & Davidson 1968

*This species has not been examined or published as Ophiostoma.

Benny & Kimbrough (1980)

Order: OphiostomatalesFamily: Ophiostomataceae

ICefatocystis

(Enteroblastic)Ophiostoma(Holoblastic)

ICefatocystiopsis

(Both)

Von Arx & Van Der Walt (1988)

Order: Ophiostomatales

IFamily: Pyxidiophoraceae

ICefa tocys tis

(Enteroblastic)

IFamily: Ophiostomataceae

IOphios toma Ceratocystiopsis(Holoblastic) (Both)

Figure 1 - Synopsis of the most recent classification schemes of these genera.

.

4

Table 2.-Form-genera with asexual conidiation, Nag Raj and Kendrick (1975), Upad-hyay (1981), Upadhyay and Kendrick (1975), Wingfield (1985)

Exogenous, HoloblasticAcremonium Link ex Fries Simple, reduced conidiophoresAllescheriella Henn. Nonspecific, hyalineHyalodendron Diddens Acropetal chains, simple, hyalineHyalopesotum Upadhyay & Kendrick Annellidic, synnematous, hyalineHyalorhinocladiella Upadhyay & Kendrick Mononematous, sympodial, hyalineGabarnaudia Samson & W. Gams Mononematous, phialidic, hyalineGraphilbum Upadhyay & Kendrick Synnematous, sympodial, hyalineGraphium Corda Synnematous, annellidic, pigmentedGraphiocladiella Upadhyay Mononematous, annellidic, hyalineKnoxdaviesia Wingfield, Van Wyk & Marassas Mononematous, phialidic, hyalineLeptographium Lagerb. & Melin Mononematous, annellidic, pigmentedPachnodium Upadhyay & Kendrick NonspecificPesotum Crane & Schoknecht Synnemamus, sympodial, pigmentedSporothrix Hekt. & Perkins ex. Nicot & Mariat Simple, sympodial, hyalineVerticiladiella S. Hughes = Leptogmphium, Wingfield Mononematous, sympodial and

annellidic, pigmented

Endogenous, EnteroblasticChalara (Corda) Rabenh.Chalaropsis Peyronel = ChalaraPhialocephala KendrickPhialogmphium Upadhyay & KendrickThielaoiopsis Went = Chalara

Phialidic, simple, hyaline

Phialidic, mononematous, hyalinePhialidic, synnematous, pigmented

and (3) Cerutocystiopsis Upadhyay & Kendrick for theother form-genera (table 2).

DISCUSSION

(Hawksworth and others 1983). In The Whole Fungus(Kendrick 1979), noteworthy scientists have writtenstimulating discussions of the history, conidial forma-tion, terms, and relative importance of anamorphs astaxonomic features.

When both the teleomorph and the anamorph areknown, the fungus is called a holomorph (the wholefungus). The teleomorph, which reproduces with as-cospores, is the sexual form of the fungus. It may haveone (mono-anamorphic) or many (pleo-anamorphic)asexual stages. The anamorph, or asexual form, isplaced into a form-genus. Form-genera, formerlyknown as fungi imperfecti, are the asexual, conidial-producing forms of a fungus. Form-genera are clas-sified according to the method of conidia formationand conidiophore morphology (conidiogenesis) andmay or may not be associated with a teleomorph. Inthe Ophiostomataceae, conidiogenesis may be holo-blastic (when both inner and outer walls contribute tothe formation of conidia) or enteroblastic (when theinner wall or neither wall contributes to the formationof conidia). Older terms for holoblastic conidiogenesisare exoconidial and exogenous; for enteroblastic, en-doconidial and endogenous. These terms are definedin Kendrick (1971), Hennebert and Weresub (19771,and Ainsworth and Bisby’s Dictionary of the Fungi

Scientists have made taxonomic revisions in boththe teleomorphs and the anamorphs of the genusCerutocystis. Since cultural and microscopic analysesalone are not always sufficient to delineate genus andspecies characteristics, new methods of ultrastruc-tural, chemical, biochemical, immunological, andgenetic analyses are being used to realign the genusCerutocystis and its anamorphs.

More complete historical data on this genus can befound in Bakshi (1951), Hunt (19561, and Upadhyay(1981). Over the years, several genera have beensynonomized and placed in the genus Cerutocystis.Bakshi (1951) synonomized six genera and the speciesEndoconidiophora coerulescens Munch and Ophios-toma coerulescens (Munch) Nannfeldt as Cerutocystiscoerulescens (Munch) Bakshi. In this enteroblasticspecies, spores (conidia) are produced within thespore-bearing structure (conidiophore). He alsodescribed Ceratocystis galeiformis, a holoblasticspecies where spores were formed at the “end ofhypha” or conidiophore (Cephalosporium Corda) or atthe “ends of the collection of hyphae” or conidiophores

5

(Graphium Corda and Leptographium). Ellis andHalstead’s (1890) generic concept of Ceratocystis wasthen changed to accommodate species that producedboth enteroblastic and holoblastic spores. Hunt (1956)accepted Bakshi’s (1951) changes, synonomized twomore genera, redescribed the genus Ceratocystis, com-pleted the transfer of synonomized species, made newcombinations, and described new species.

In 1974, de Hoog proposed the resurrection of thegenus Ophiostoma with the type species 0. piliferumfor species producing holoblastic conidia. Von Hohnel(1918) originally described the genus as Linostoma,an invalid name, which Syrdow and Sydow (1919)renamed Ophiostoma, and Bakshi (1951)synonomized with the genus Ceratocystis.

Melin and Nannfeldt (1934) divided the genusOphiostoma into sections to accommodate species thatproduced their spores inside the conidial bearingstructure (enteroblastic) and to accommodate theholoblastic species. De Hoog used Nannfeldt’s sectionsin Melin and Nannfeldt (1934) to divide the twogenera based on the type of conidia formed, eitherenteroblastic or holoblastic. In 1975, Weijman and deHoog divided the genus Ceratocystis as Ceratocystissensu strict0 (in the strict sense) to contain all specieswith enteroblastic conidiogenesis and Ceratocystissensu lato (in the wide sense) as the Ophiostomagroup for holoblastic species. In 1975, Upadhyay andKendrick proposed to split the genus Ceratocystis andto establish the genus Ceratocystiopsis for thosespecies with falcate ascospores and ascoma (peri-thecia) with short necks.

In 1981, Upadhyay published a monograph of theCeratocystis species, synonomized Ophiostoma andEurophium, and transferred 13 of 15 Ceratocystisspecies to Ceratocystiopsis. In 1984, de Hoog andScheffer established Ophiostoma as a valid genuscharacterized by having cellulose, lignin, and rham-nose in their cell walls; growing on cycloheximidemedia; and producing holoblastic conidia. The genusCeratocystis was limited to species with enteroblasticChalara anamorphs and those that were cellulosenegative and sensitive to cycloheximide. De Hoog andScheffer accepted Upadhyay and Kendrick’s genusCeratocystiopsis.

The genus Ceratocystis now contains only thosespecies having the enteroblastic form-genus Chalara,with the type species C. fimbriata Ellis & Halstead(table 3). Ceratocystiopsis, with the type species C.minuta (Siem.) Upadhyay & Kendrick, is an exceptionin that it does have an enteroblastic Chalaraanamorph as well as holoblastic anamorphs. Theholoblastic Ophiostoma, type species 0. piliferum, hasseveral form-genera of which two or more anamorphsmay develop in culture. Op?$ostoma clavigerum(Robins.-Jeff. & Davids.) Harrington (1987), whencontinually subcultured, has produced as many as six

Table 3.-Ceratocystis species with Chalara anamorphs, de Hoog(1974) and von Arx (1987)

de HoogCeratocystis adiposa (Butler) C. MoreauC. autogmpha BakshiC!. coerulescens (Munch) BakshiC. fagacearum (Bretz) HuntC. fimbriata Ellis & HalsteadC. major (van Beyma) C. MoreauC. moniliformis (Hedgcock) C. MoreauC. musarum RiedlC. paradoxu (Dade) C. MoreauC. radicicola (Bliss) C. MoreauC. uariospora (Davidson) C. MoreauC. uirescens (Davidson) C. Moreau

von ArxC. adiposum (Butler) C. MoreauC. coerulescens (Munch) BakshiC. fagacearum (Bretz) HuntC. fimbriata Ellis & HalsteadC. paradoxa (Dade) C. Moreau

forms: Graphiocladiella Upadhyay, Verticicladiella,Hyalorhinocladiella Upadhyay & Kendrick, Lep-tographium, an annellidic yeast, and a holoblasticyeast (Tsuneda and Hirat,suka 1984).

DeHoog in Malloch (1979) said of the Verticicladiel-la-Phialographium Upadhyay Jz Kendrick-Lepto-graphium complex that older cultures degenerate andtend to form less well-structured, less complex, andless energy-consuming anamorphs.

Recognized form-genera now total 17 (table 2). NagRaj and Kendrick (1975) synonomized ChalaropsisPeyronel and Thielaviopsis Went as Chalara. Upad-hyay (1981) described 17 form-genera, but Wingfield(1985) reclassified Verticicladiella as Leptographium,and Wingfield and others (1988) described a newspecies, Knoxdaviesia. Harrington (11988) felt thatmost Ophiostoma anamorphs could probably be ac-commodated in the form-genera Sporothrix Hekt. &Perkins ex Nicot & Mariat, Leptographium, andGraphium. Form-genera of Ceratocystis, Ceratocys-tiopsis, and Ophiostoma need taxonomic study asrecommended by the 1986 International Council ofBotanical Names and by Kananaskis II, published asThe Whole Fungus (Kendrick 1979).

Mycologists, plant pathologists, and botanists donot agree on the higher classifications of fungi. Forinstance, Ainsworth and Bisby’s Dictionary of theFungi (Hawksworth and others 1983) recognizes theKingdom Fungi but does not recommend any onescheme for classes in the Subdivision Ascomycotina.Von Arx (197) advocated a limited number of ordersand the omission of higher ranks. At an August1990 meeting in Bad Windsheim, West Germany,specialists in the Ophiostomatales presented paperswith startling opinions of this group of fungi (pers.

’

6

comm. Dr. M. Blackwell)‘. Upadhyay would retainthe family Ophiostomataceae in the order Microas-tales; Samuels would place Ophiostoma andCerutocystis in the Order Sordariales and the Fami-ly Chaetosphaeriaceae; Wingfield would synonomizeCeratocystiopsis with Ophiostoma, except Ceratocys-tiopsis fukutu (Wright & Cain) Upadhyay, whichwould reside in Ceratocystis sensu stricto.

Several authors discuss anamorphs; others discussmolecular methods, pathogenicity, vectors and disper-sal, sapstain, tree response and host defense reac-tions, and chemistry. Kendrickl (Wingfield and others1990) states that “since we do not yet have all thisinformation (ultrastructure, development patterns,anamorph-teleomorph relationships, wall-chemistry,chemistry of slime, vectors and dispersal mecha-nisms, pathogenicity, and genetics) for any of themore than one-hundred members of the Ophiostoma-tales, it is clear that plenty of work remains to be donebefore we will properly understand this fascinatingand important group of fungi.”

Article 59 of the International Code of BotanicalNomenclature (Greuter and others 1983) states thatthe name of the holomorphic Ascomycete (the wholefungus, with both anamorph and teleomorph) takesprecedence over the name of the anamorph. For ex-ample, one mycangial fungus of Dendroctonus fron-t&is Zimmermann, the southern pine beetle, isheterothallic (sexual reproduction requiring the unionof two compatible mating types) and has beendescribed as the anamorph Sporothrix2, but since theanamorphic name was not validly described, the fun-gus must be cited by the name of the teleomorph,Ceratocystiopsis ranaculosus Perry & Bridges (Bridg-es and Perry 1987, Harrington and Zambino 1990).Mycologists have agreed that anamorphs that havebeen validly described and published may be cited

‘Blackwell, Meredith. 1990. Personal communication with Dr.Blackwell, Department of Botany, Louisiana State University,Baton Rouge, LA 70803, in Sept. 1990, regarding “The Biologyand Taxonomy of the Ophiostomatales”, an international sym-posium, held in Bad Windsheim, West Germany, on August Zl-24, 1990. Program and Abstracts organized by Wingfield, M.J.,Department of Microbiology, University of the Orange Free State,P.O. Box 339, Bloemfontein 9300, Republic of South Africa;Seifert, K., Biosystematics Research Institute, Central Experi-ment Farm, William Saunders Building, Ottawa, Ontario, KIAOC6, Canada; Webber, J.F., Forest Research Station, Alice HoltLodge, Wrecclesham, Farnham, Surrey, GUlO 4LH, England. Theproceedings will be published at a later date.

2Harrington, T.C. 1991. Personal communication with Dr. Har-rington, Department of Plant Pathology, Iowa State University,Ames, IA 50011, who reviewed the manuscript in January, 1991,and disagreed with the taxonomic identification of this holomorphand stated that it needs further examination.

separately, such as the anamorph Sporothrix suban-nulatu Livingston & Davidson and its teleomorphOphiostoma subannulatum Livingston & Davidson(Livingston and Davidson 1987).

A list of the Cerutocystis species that have beentransferred (published as Ophiostoma species by newauthors, table 4) can be found in this publication. If adescribed species of Ceratocystis having an anamarphother than Chalaru is not listed as recombined in deHoog and Scheffer (1984) and Harrington (1987,1988)(examples 1, 2), then consult Upadhyay (1981) (ex-ample 3) for the original author of a Cerutocystis spe-cies that had been originally described as or tran-ferred to an Ophiostoma species (table 5). More than ahundred described species of Cerutocystis exist. Somespecies were originally describled as Ophiostoma;

Table 4.-New and recombined Ophiostoma species by otherauthors

-Ophiostoma ainoue Solheim (1986)0. cucullulutum Solheim (1986)0. dauidsonii (Olchowecki & Reid) Solheim (1986)0. europhioides (Wright & Cain) Solheim (1986)0. flexuosum Solheim (1986)0. Zongirostellatum (Bakshi) v. Arx & Mnller (1954)0. roboris Georgescu & Teodoru (1948)0. sugmutosporu (Wright & Cain) Solheim (1986)0. subunnulatum Livingston & Davidson (1987)

-

Table 5.-Ceratocystis species that were originally described us,or transferred to Ophiostoma, de Hoog (1974) andUpudhyay (1981)

Ophiostomu bicolor Davidson & Wells0. brunneo-ci2iutum Mathieson-Kaarik0. ips (Rumb.) Nannfeldt0. Zeptogruphioides (Davidson) v. Arx0. microsporium (Davidson) v. Arx0. minus (Hedge.) H. & P. Sydow0. multiunnulatum (Hedgcock & Davidson) Hendrix0. narcissi Limber0. nigrocurpum (Davidson) de Hoog0. olivaceum Mathieson0. penicillutum Mathieson0. perfectum (Davidson) de Hoog0. piceae (Munch) H. & P. Sydow0. piceuperdum (Rumbold) v. Arx0. piliferum (Fries) H. & P. Sydow0. polonicum Siemaszko0. rostrocylindricum (Davidson) v. Arx0. stenocerus (Robek) Melin & Nannfeldt0. tetropii Matheison0. triungulosporum Butin0. ulmi (Buisman) Nannfeldt -

7

some species have been transferred to the genusOphiostoma, and some have not yet been recombined(tables 6 & 7). Recombinations formed in the transferof species must follow the rules of the InternationalCode of Botanical Nomenclature, Article 33. The ruleshave been restated in the International Commissionon the Taxonomy of Fungi (ICTF) Code of Practice forSystematic Mycologists (Sigler and Hawksworth1987). Care must be taken to note the Latin or Greekendings of genera and species. For example, the genusOphiostoma has a neuter ending, so the species en-ding must also be neuter.

Table 6.-Ceratocystis species not recombined, Upadhyay (1981)

Ceratocystis acericola GriffinC. aequiuaginata Olchowecki 8z ReidC. allantospora GriffinC. angusticollis Wright & CainC. arborea Olchowecki & ReidC. brunneo-crinita Wright & CainC. California Devay, Davidson and MollerC. columnaris Olchowecki & ReidC. deltoideospora Olchewecki & Reid

*C. fimicola (Marchal) Upadhyay*C. heluellae (Karsten) UpadhyayC. hyalothecium DavidsonC. introcitrina Olchowecki & ReidC. leucocarpa DavidsonC. magnifica GriffinC. olivaceapini DavidsonC. populicola Olchowecki & ReidC. pseudominor Olchowecki & ReidC. stenospora GriffinC. tenella DavidsonC. torticiliata Olchowecki & ReidC. tubicollis Olchowecki & Reid

*Upadhyay synonomized these species, but Benny andKimbrough (1980) and Cannon and Hawksworth (1982) recognizethe genus Sphaeronaemella Karsten as valid.

Table 7.-Ceratocystis species not recombined by other authors

Ceratocystis grandicarpa Kowalski & Butin (1989)C. kubanicum Scerbin-Parfenenko (1953)C. nothofagi Butin (1984)C. novae-zelandiae Hutchinson & Reid (1988)C. prolifera Kowalski & Butin (1989)C. ualachium Georgescu, Teodoru & Badea (1948)

EXAMPLES

1. Ophiostoma araucariae (Butin) de Hoog &Scheffer = Ceratocystis araucariae Butin.

2. Ophiostoma abiocarpum (Davids.) Harrington= Ceratocystis abiocarpa Davidson

3. Ophiostoma minus (Hedge.) H. & P. Sydow= Ceratocystis minor (Hedge.) Hunt.

The symbol “g” designates an obligate synonym.Both names are based on the same type specimen.They are nomenclatural synonyms. The symbol “=”designates a facultative synonym--names based ondifferent type specimens (Hawksworth and others1983); See table 1.

SYNOPSIS OF RELEVANT PUBLICATIONS

1890

1891

1919

1932

1934

1951

Ellis and Halstead, in Halstead (1890), estab-lished the genus Ceratocystis without a formalgeneric description.Halstead and Fairchild established the genericconcept of the genus and ,provided the typespecies C. fimbriata, with an endogenous con-idial form.Sydow and Sydow established the genusOphiostoma for Linostoma von Hohnel, an in-valid name. Von Hohnel (1918) characterizedthe genus Linostoma as having hyaline as-cospores formed in evanescent asci and havingfringes of ostiolar “cilia.”Nannfeldt erected the family Ophios-tomataeceae for the genus Ophiostoma.Melin and Nannfeldt divided Ophiostoma intotwo sections. The species of sectionBrevirostrata Nannfeldt have short, conicalperithecial necks, and the species assigned tosection Longirostrata were characterized bylonger filiform necks; Longirostrata was fur-ther divided to contain species with both en-dogenous conidia, Chalara- type (synonomizedEndoconidiophora Munch), and exogenousconidia.Bakshi revived the generic name Ceratocystisfor species with both endogenous and ex-ogenous conidia. He regarded Ophiostoma, En-doconidiophora, Rostrella Zimmermann, Lino-stoma von Hohnel, and Grosmannia G.Goidanich as synonyms. Ceratostomella Sac-cardo was retained as a separate genus. Hedescribed three new species and transferredthe species Endoconidiophora coerulescensMunch (Munch 1907) and Ophiostomacoerulescens (Munch) Nannfeldt (Melin and

8

1956

1957

1962

1964

1965

1965

1967

1968

Nannfeldt 1934) to the genus Cerutocystis, asCerutocystis coerulescens (Munch) Bakshi.Hunt published the first comprehensivemonograph of Cerutocystis species, placed thegenus in the order Plectascales, acceptedBakshi’s emendations (that the genus containsspecies with both endogenous and exogenousconidia), and completed the transfer of species.He synonomized Sphaeria Haller ex Fries andSphaeronaemella Karsten ex Seeler underCerutocystis, and Fuguscus Falck was listed asnomen nudum (a genus without adequatedescription). Cerutostomella was also excludedfrom synonomy. He presented a key to thespecies and sectioned them according to con-idial development: section (1) species with anendoconidial imperfect stage, the EndoconidialGroup; section (2) species with a Lepto-graphium or Graphium imperfect state con-sisting of the Grosmannia Group and theOphiostoma Group in part; and section (3)species with mycelial conidia only, the Ophios-toma Group in part.Parker established the genus Eurdphium withthe type species E. trinacrifirme. The specieswas distinguished from Cerrztocystis by itsneckless, closed ascocarps. Benny and Kim-brough (1980) and Upadhyay (1981) synono-mized this genus with Cerutocystis; von Arxand van der Walt (1988) resurrected the genusand placed it in the family Ophiostomataceae.Kendrick redescribed the genus Verticicladiel-la S. Hughes, differentiated the genus Lep-tographium Lagerberg & Melin, placed threespecies in Verticicladiella, and described fournew species, V. procera, V. brachiata, V. an-tibiotica, and V. wagenerii.Rosinski and Campana demonstrated bothchitin and cellulose in the hyphal walls ofCeratocystis ulmi (Buisman) C. Moreau.Kendrick and Molnar described a new Ceru-tocystis with the first-described Verticicladiel-la anamorph.Rosinski confirmed the occurrence of cellulosein C. ulmi.Smith and others differentiated four species bytheir cell wall chemistry and divided them intotwo categories. The endoconidial (enteroblasticor endogenous) forms Cerutocystis fagacearum(Bretz) Hunt and C. fimbriata Ellis &Halstead have no cellulose in their cell walls,and the exoconidial (holoblastic or exogenous)forms of C. oliuacea (Mathiesen) Hunt haveboth chitin and cellulose in their cell walls.Griffin published a monograph of 32 Cerato-cystis species found in Ontario with descrip-tions, figures, distribution data, and a key to

60 species based on ascospore morphology.Eleven species were described as new.

1968 Robinson-Jeffrey and Davidson describedthree new species of Europhium (clavigerum,aureum, and yobustum), which Upadhyay, in1981, synonomized as Ceratocystis species in-cluding the type species E. trinacriforme. In1987, Harrington recombined the four speciesas Ophiostoma.

1970 Von Arx urged’ revision of the form-genusGraphium because conidia of described specieswere formed exogenously as well as endo-genously. He cited the genus as CeratocystisEllis & Halstead with 50 species and acceptedBakshi’s synonyms but did not acceptedBakshi’s (1951) or Hunt’s (1956) emendations.

1971 Spencer and Gorin, to aid classification,grouped species of Ceratocystis and Graphiumaccording to their polysaccharide components.The endoconidial (endogenous) group formedglucomannans or galactoglucomannans ratherthan rhamnose- and mannose-containing poly-saccharides as in the exoconidial (exogenous)group.

1972 Cain suggested a relationship to fllamentousyeastlike taxa as Cephaloascus Hanawa andsaid that Verticicladiella may be more closelyrelated to hyphal yeast genera with hat-shaped ascospores than to phialidic Chalara-like anamorphs.

1973 Muller and von Arx placed the genus Cerato-cystis in the order Sphaeriales of the Pyreno-mycetes and the family OphiosomataceaeNannfeldt, removing it from the orders Plec-tascales (Nannfeldt 1932, Hunt 19561, theMicroascales of the Plectomycetes (Luttrell1951, 1955; Kendrick and Molnar 1965) andthe Ophiostomatales (Rosinski 1961). Theorder Sphaeriales contains four genera:Europhium Parker, Sphaeronaemella Karstenex Seeler, Chadefuudia Feldman, and Ceruto-cystis (Ellis & Halstead) Bakshi.

1974 De Hoog concluded that the genus Ceratocystisshould be separated into 2 separate generabased on the morphology of the conidial states:Cerutocystis Ellis and Halstead with theanamorphs Chalara, Chalaropsis Peyronel,and Thielaviopsis Went accepting 11 species(see table 2), and the genus Ophiostoma H. &P. Sydow containing the anamorphs Sporo-thrix Hektoen & Perkins ex Nicot & Mariat,Verticicladiella S. Hughes, LeptographiumLagerberg 8z Melin, and Graphium Corda. Heproposed four new Ophiostoma combinations,enlarged the concept of the form-genusSporothrix, and described new species.

9

1974

1974

1974

1975

1975

1975

1977

10

Jewel1 studies cellulose distribution in cellwalls of47 species of Ceratocystis and 4 speciesof Europhium cytochemically and using x-raydiffraction analysis. Thirty-one species ofCeratocystis and all of the Europhium speciescontained cellulose.Olchowecki and Reid published a monographof 50 Ceratocystis species found in Manitobawith a key to 70 species of which 25 weredescribed as new. They provided a first-timeconidial description for three species. Theysubdivided the genus into four groups based onascospore morphology; (1) the Minuta Group,(2) the Ips Group, (3) the Fimbriata Group,and (4) the Pilifera Group.Von Arx urged revision of the genus Ceratocys-tis and suggested that species with Chalara-like anamorphs be retained within the genus,but those species having holoblastic conidia betransferred to Ophiostoma. He maintainedEurophium as a distinct genus.Nag Raj and Kendrick placed the form-generaChalaropsis and Thielaviopsis in synonomywith Chalara.Upadhyay and Kendrick proposed dividing thegenus Ceratocystis and establishing a newgenus Ceratocystiopsis based on Olchoweckiand Reid’s Minuta group that included 19species with falcate ascospores and ascomawith short necks. The new genus was classedas Ascomycotina, Plectomycetes, Microascales,Ophiostomataceae. They listed 13 anamorphsof Ceratocystis and Ceratocystiopsis, 4 of whichwere described as new: Hyalorhinocladiella,Graphilbum, Hyalopesotum, and Pachnodium.Europhium was considered a synonym ofCeratocystis, and Ophiostoma was not ac-cepted as a valid genus. Olchowecki and Reid’sthree other groups-Ips, Fimbriata, andPilifera-were accepted.Weijman and de Hoog discussed the results ofcell wall cellulose (Smith and others 1967) andthe presence of rhamnose (Spencer & Gorin1971) as an aid in the classification of teleo-morphs and anamorphs. They divided thegenus Ceratocystis into two groups: (1) thephialidic endogenous conidial forms- Chalaraand allied genera-and (2) the exogenous con-idial forms of the Ophiostoma group (Graph-ium-like states). Ceratocystis sensu strict0 forthe Chalara type was used where rhamnoseand cellulose were absent.Redhead and Malloch placed the genusCeratocystis in the order Endomycetales andthe family Endomycetaceae as a yeast-relatedgenus. The family Ophiostomataceae wassynonomized with the family Endomycetaceae.

1978 Upadhyay dropped emendation of the genusCeratocystis by Bakshi (1951) and Hunt (1956)to conserve the original description by Ellisand Halstead.

1979 The Proceedings of the Second InternationalMycological Congress (Kananaskas II) waspublished in two volumes as The Whole Fun-gus (Kendrick 1979). Included were discus-sions of fungal morphology, classification, ecol-ogy, evolution, and techniques. New termssuch as holomorph, teleomorph, and ana-morph were defined. Malloch, von Arx, deHoog, and Luttrell presented divergent viewsof Ceratocystis classification. Kendrick andDiCosmo’s paper of teleomorph-anamorph con-nections listed 70 species of Cerutocystis and 2species of Ceratocystiopsis.

1980 Benny and Kimbrough proposed a new order,Ophiostomatales, having only one familyOphiostomataceae (Nannfeldt 1932) that in-cluded the genera Ophiostoma, Cerutocystis,Ceratocystiopsis, and Sphaeronaemella.Europhium was synonomized, and Chade-fuudiu Feldman was rejected from the family.They resurrected the genus Sphueronuemellu,which Hunt (1956) had placed in synonomywith the genus Ceratocystis.

1981 Harrington published on cycloheximide as anaid to classification of the species. Ceratocystissensu strict0 (Chalura anamorphs) is sensitiveto cycloheximide, but Certocystis sensu lato(Ophiostoma spp.) is not sensitive.

1981 Upadhyay published a monograph on thegenera Cerutocystis Ellis & Halstead andCerutocystiopsis Upadhyay & Kendrick. Heplaced the two genera in the order Microas-tales of the Plectomycetes. Ophiostoma,Rostrella, Endoconidiophoru, Grosmunnia,Sphaeronaemellu, and Europhium were re-jected as valid genera and were reduced tosynonomy with the genus Cerutocystis. Hedivided the genus Cerutocystis into four sec-tions: (1) Ophiostoma, ascospores lacking ahyaline gelatinous sheath; (2) Ips, ascosporesalways surrounded by a hyaline gelatinoussheath, rectangular, never curved; (3) Ceruto-cystis, ascospores sheathed, partly curved;and, (4) Endoconidiophora, sheath elongated,inequilateral.

1983 Ainsworth and Bisby’s Dictionary of the Fungi(Hawksworth and others 1983) listed the orderOphiostomatales and the family Ophiosto-mataceae as having 2 or 3 genera, 13 syno-nyms, and 85 species.

1983 Urosevic redescribed five Ceratocystis speciesthat cause tracheomycoses of oak as Ophios-toma species.

1984 De Hoog and Scheffer published a historicalreappraisal of the genera Ceratocystis andOphiostoma. They concluded that all specieswith Chalaru anamorphs are Cerutocystissensu strictu and distinct from Cerutocystiop-sis. All others haying conidial anamorphsare Ophiostomu (other than Chularu, whi chhas cell walls with rhamnose and is resistantto cycloheximide).Thelatter included Europhi-urn. They recombined 14 Cerutocystis speciesas Ophiostomu species (table 8).

1985 Kendrick recognized the order Ophiostoma-tales haying 15 genera and 130 species.

1985 Wingfield reclassified Verticicludiella as Lep-togruphium. He noted that the annellidic aswell as the sympodial type of growth occurs inthe Leptogruphium species.

1986 Solheim presented a key to genera of the fami-ly Ophiostomataceae and a key to ten Nor-wegian Ophiostomu species and one Cerutocys-tiopsis species; 0. uinoue, 0. cucullatum, and0. flexuosum were described as new.

1987 Bridges and Perry described a new species:Cerutocystiopsis runuculosus Perry & Bridges,having a Sporothrix anamorph.

Table 8.-Recombined Ceratocystis species by de Hoog and Scheffer (1974)

Ophiostoma araucariae (Butin) de Hoog & Scheffer.= Ceratocystis araucariae Butin

Ophios toma bac i l l i sporum (Butin & Zimmermann) de Hoog & Scheffer= Ceratocystis bacillispom Butin & Zimmermann

Ophiostoma brevicolla (Davidson) de Hoog & SchefferE Ceratocystis bmvicollis Davidson

Ophiostoma distortum (Davidson) de Hoog & Scheffer= Ceratocystis distorta Davidson

Ophios toma dryocoetidis (Kendrick & Molnar) de Hoog & Scheffer= Ceratocystis dryoccetidis Kendrick & Molnar

Ophiostoma francke-grosmanniae (Davidson) de Hoog & Scheffer5 Ceratocystis francke-grosmanniae Davidson

Ophiostoma huntii (Robinson-Jeffrey) de Hoog & Scheffer= Ceratocystis huntii Robinson-Jeffery

Ophios toma megalobrunneum (Dav idson & Toole ) de Hoog & Scheffer= Ceratocystis megalobrunnea Davidson & Tools

Ophios toma n igrum (Davidson) de Hoog & Scheffer= Ceratocystis nigra Davidson

Ophios toma populinum (Hinds & Davidson) de Hoog & Scheffer= Ceratocystis populina Hinds & Davidson

Ophiostoma rostrocoronatum (Davidson & Eslyn) de Hoog & Scheffer= Ceratocystis rostrocoronata Davidson & Eslyn

Ophiostoma seticolle (Davidson) de Hoog & Scheffer= Ceratocystis seticollis Davidson

Ophios toma sparsum (Davidson) de Hoog & Scheffer= Ceratocystis sparsa Davidson

Ophiostoma tremulo-aureum (Davidson & Hinds) de Hoog & SchefferI Ceratocystis tmmulo-auma Davidson & Hinds

11

1987

1987

1988

1988

Von Arx accepted five species in the genusCerutocystis. All have Chalaru anamorphs, butonly C. fimbriatu has hat-shaped ascospores.Harrington recombined 11 Cerutocystis speciesas Ophiostoma (table 9).Hutchinson and Reid presented a key to sevenpotential wood-staining Ceratocystis. Ceruto-cystis novae-zelandiae was described as new.They listed one species of Ceratocystiopsis anda Sphaeronaemella species.Von Arx and van der Walt accepted the orderOphiostomatales by Benny and Kimbrough forthe three genera Ceratocystis, Ceratocystiop-sis, and Ophiostoma and the family Ophios-tomataceae for Ceratocystiopsis and Ophios-toma. However, they proposed a new family,Pyxidiophoraceae, for Ceratocystis because of

1988

1989

1990

their Chalaru anamorphs and the lack of cel-lulose in their cell walls (fig. 1).Wingfield and others described a new speciesCeratocystiopsis proteae with a new anamorphgenus Knoxdaviesia.Kowalski and Butin redescribed four species ofCeratocystis from oak in Poland and describedtwo new species. Ceratostomella quercus Geor-gev. and Ophiostoma roboris were synono-mized as Ceratocystis piceae (Munch) Bakshi.Harrington and Zambino rejected Ceratocystisminor var. barrasi J. Taylor as the mycangialfungus of Dendroctonus front&s and estab-lished Ceratocystiopsis ranaculosus as the cor-rect fungus based on isozyme analysis andmating studies.

Table 9.-Recombined Ceratocystis species by Harrington (1987)

Ophiostoma abiocarpum (Davidson) HarringtonE Ceratocystis abiocarpa Davidson

Ophiostoma adjuncti (Davidson) HarringtonI Ceratocystis adjuncti Davidson

Ophiostoma aureum (Robinson-Jeffrey & Davidson) Harrington= Europhium aureum Robinson-Jef frey & D a v i d s o n= Ceratocystis aurea (Robinson-Jeffrey & Davidson) Upadhyay

Ophiostoma cainii (Olchowecki & Reid) Harrington= Ceratocystis cainii Olchowecki & Reid

Ophiostoma clavigerum (Robinson-Jeffrey & Davidson) HarringtonE Europhium clavigerum Robinson-Jef frey & D a v i d s o n= Ceratocystis clavigem (Robinson-Jeffrey & Davidson) Upadhyay

Ophiostoma crassivaginatum (Griffin) Harrington= Ceratocystis crassivaginata Griffin= Ceratocystiopsis crassivaginata (Griffin) Upadhyay

Ophiostoma grandifoliae (Davidson) Harrington2 Ceratocystis grandifoliae Davidson

Ophiostoma robustum (Robinson-Jeffrey & Davidson) Harrington= Europhium robustum RobinsonJeffrey & Davidson= Ceratocystis robusta (RobinsonJeffrey & Davidson) Upadhyay

Ophiostoma trinacriforme (Parker) HarringtonI Eurwphium trinacriforme ParkerI Ceratocystis trinacriforme (Parker) Upadhyay

Ophios toma va ld iv ianum (Butin) Harr ington= Ceratocystis valdiviana Butin

Ophiostoma wageneri (Goheen & Cobb) Harrington= Ceratocystis wageneri Goheen & Cobb

12

1990 Malloch and Blackwell proposed the genusKathistes and presented a key to the generalikely to be confused with Ceratocystiopsis -Gabarnaudia, Sphaeronaemella, Ceratocystis,Ophiostoma, Pyxidiophora, Rhynchonectria,Treleasia, Subbaromyces, Spumatoria, andKlasterskya.

1990 Van Wyk and Wingfield reviewed the con-troversial taxonomic status of Ceratocystis,Ceratocystiopsis, and Ophiostoma regardingthe development of asci, ascospores, andcentrum structure. They suggested that addi-tional ultrastructural studies are required toclarify these relationships.

\

LITER,ATURE CITED

Bakshi, B.K. 1951. Studies on four species ofCeratocystis, with a discussion on fungi causingsap-stain in Britain. Commonwealth MycologicalInstitute Mycology Research Paper. 351-16.

Barr-as, S.J. 1970. Antagonism between Dendroctonusfrontalis and the fungus Ceratocystis minor. Annalsof the Entomological Society of America. 63:1187-1190.

Barras, S.J.; Perry, T.J. 1971. Leptographiumterebrantis sp. nov. associated with Dendroctonusterebrans in loblolly pine. Mycopathologia etMycologia Applicata. 43:1-10.

Barras, S.J.; Perry, T.J. 1975. Interrelationshipsamong microorganisms, bark or ambrosia beetlesand woody host tissue. An annotated bibliography1965-1974. Gen. Tech. Rep. SO-lo. New Orleans,LA: U.S. Department of Agriculture, Forest Service,Southern Forest Experiment Station. 34 p.

Beaver, R.A. 1989. Insect-fungus relationships in thebark and ambrosia beetles. Symposia of the RoyalEntomological Society of London, 1989. London:Academic Press. 121-143.

Benny, G.L.; Kimbrough, J.W. 1980. A synopsis of theorders and families of Plectomycetes with keys togenera. Mycotaxon. 12:1-91.

Bramble, W.C.; Holst, E.C. 1940. Fungi associatedwith Dendroctonus frontalis in killing shortleafpines and their effect on conduction. Phytopathol-ogy. 30:881-899.

Bridges, J.R.; Perry, T.J. 1985. Effects of mycangialfungi on gallery construction and distribution ofbluestain in southern pine beetle-infested pinebolts. Journal of Entomological Science. 20:271-275.

Bridges, J.R.; Perry, T.J. 1987. Ceratocystiopsisranaculosus sp. nov. associated with the southernpine beetle. Mycologia. 79:630-633.

Butin, H.; Aquilar, A.M. 1984. Blue-stain fungi onNothofagus from Chile-including two new speciesof Ceratocystis Ellis & Halst. PhytopathologischeZeitschrift. 109:80-89.

Cain, R.F. 1972. Evolution of fungi. Mycologia. 64:1-14.

Cannon, P.F.; Hawksworth, D.L. 1982. A re-evalua-tion of Melanospora Corda and similar Pyreno-mycetes with a revision of the British species.Botanical Journal of the Linnean Society. 84:115-160.

Davidson, R.W. 1955. Wood-staining fungi associ.atedwith bark beetles in Engelmann spruce in Colorado.Mycologia. 47:58-67.

Davidson, R.W. 1958. Additional species of Opbios-tomataceae from Colorado. Mycologia. 50:661-670.

Davidson, R.W. 1966. New species of Ceratocystisfrom conifers. Mycopathologia et Mycologia Ap-plicata. 28:273-286.

Davidson, R.W. 1971. New species of Ceratocystis.Mycologia. 63:5-15.

Davidson, R.W. 1978. Staining fungi associated withDendroctonus adjuncti in pines. Mycologia. 70:35-4 0 .

de Hoog, G.S. 1974. The genera Blastoboltrys,Sporothrix, Calcarisporium and Calcarisporiellagen. nov. Studies in Mycology. 7:1-84.

de Hoog, G.S.; Scheffer, R.J. 1984. Ceratocystis versusOphiostoma: A reappraisal. Mycologia. 76:292--299.

Franklin, R.T. 1970. Observations on the blue stain-southern pine beetle relationship. Journal of theGeorgia Entomological Society. 5:53-57.

Georgescu, C.; Teodoru, I.; Badea, M. 1948. Le Des-sechement en masse du chene. Institutul de Cer-cetari si Experimentatie Foriestiera. Bucharest:Analele; 11: 185-223.

Goheen, D.J.; Cobb, F.W., Jr. 1978. Occurrence of Ver-ticicladiella wagenerii and its perfect state,Ceratocystis wageneri sp. nov., in insect galleries.Phytopathology. 68:1192-1195.

Greuter, W. and others, eds. 1983. International codeof botanical nomenclature adopted by the Thir-teenth International Botanical Congress. Sydney,August 1981. [Regnum Vegetable no. 1111. Utrecht:Bohn, Scheltema & Holkema. 472 p.

Griffin, H.D. 1968. The genus Ceratocystis in Ontario.Canadian Journal of Botany. 46:689-718.

Halstead, B.D. 1890. Some fungous diseases of thesweet potato. The black rot. New Jersey Agricul-tural Experiment Station Bulletin. 76:7-14.

Halstead, B.D.; Fairchild, D.G. 1891. Sweet-potatoblack rot. Journal of Mycology. 7: l-l 1.

Harrington, T.C. 1981. Cycloheximide sensitivity as ataxonomic character in Ceratocystis. Mycologia.73:1123-1129.

13

Harrington, T.C. 1987. New combinations in Ophios-toma of Ceratocystis species with Leptographiumanamorphs. Mycotaxon. 283943.

Harrington, T.C. 1988. Leptographium species, theirdistribution, hosts and insect vectors. In: Har-rington, T.C.; Cobb, F.W., Jr., eds. Leptographiumroot diseases on conifers. St. Paul, MN: AmericanPhytopathology Society Press: l-39.

Harrington, T.C.; Zambino, P.J. 1990. Ceratocystiop-sis ranaculosus, not Ceratocystis minor var. bar-rasii, is the mycangial fungus of the southern pinebeetle. Mycotaxon. 38: 103-l 15.

Hawksworth, D.L.; Sutton, B.C.; Ainsworth, G.C.1983. Ainsworth and Bisby’s dictionary of the fungi.7th ed. Kew, Surrey, England: CommonwealthMycological Institute. 445 p.

Hennebert, D.L.; Weresub, L.K. 1977. Terms forstates and forms of fungi, their names and types.Mycotaxon. 6:207-211.

Hunt, J. 1956. Taxonomy of the genus Ceratocystis.Lloydia. 19: l-58.

Hutchinson, L.J.; Reid, J. 1988. Taxonomy of somepotential wood-staining fungi from New Zealand 1.Ophiostomataceae. New Zealand Journal ofBotany. 26:63-81.

Jewell, T.R. 1974. A qualitative study of cellulose dis-tribution in Ceratocystis and Europhium. My-cologia. 66: 139-146.

Kendrick, W.B. 1962. The Leptographium complex.Verticicladiella Hughes. Canadian Journal of Bot-any. 40:771-797.

Kendrick, W.B., ed. 1971. Taxonomy of the fungi im-perfecti. Toronto: University of Toronto Press.309 p.

Kendrick, W.B., ed. 1979. The whole fungus. In:Proceedings of the 2nd International MycologicalConference held at the Environmental SciencesCentre of the University of Calgary; Kananaskis,Alberta, Canada. Ottawa: National Museums ofCanada and the Kananaskis Foundation. 793 p.

Kendrick, W.B. 1985. The fifth kingdom. Ottawa:Mycologue Publications. 363 p.

Kendrick, W.B.; Molnar, A.C. 1965. A new Ceratocys-tis and its Verticicladiella imperfect state asso-ciated with the bark beetle Dryocoetes confusus onAbies Zasiocarpa. Canadian Journal of Botany.43:39-43.

Kowalski, T.; Butin, H. 1989. Taxonomy of known andnew species of Ceratocystis from oak (Quercus roburL.). Journal of Phytopathology. 24:236-248.

Livingston, W.H.; Davidson, R.W. 1987. Ophiostomasubannulatum, a new fungal species pathogenic togrand fir roots. Mycologia. 79:144-147.

Luttrell, E.S. 1951. Taxonomy of the Pyrenomycetes.University of Missouri Studies. 24:1-121

Luttrell, E.S. 1955. The ascostromatic ascomycetes.Mycologia. 47:511-532.

Malloch, D. 1979. Plectomycetes and their ana-morphs. In: Kendrick, B., ed. The whole fungus. Ot-tawa: National Museums of Canada: 153-165. Vol.1 .

Malloch, D.; Blackwell, M. 1990. Kathistes, a newgenus of pleomorphic ascomycetes. Canadian Jour-nal of Botany. 68:1712-1721.

Mathre, D.E. 1964. Survey of Ceratocystis spp. as-sociated with bark beetles in California. Contribu-tions of the Boyce Thompson Institute. 22:353-362.

Mattson, W.J.; Levieux, J.; Bernard-Dagan, C., eds.1988. Mechanisms of woody plant defenses againstinsects: search for a pattern. London: Springer-Ver-lag. 416 p.

Melin, E.; Nannfeldt, J.A. 1934. Researches into theblueing of ground wood-pulp. Svenska Skogsvards- ’foereningens Tidskrift. 32:397-616.

Muller, E.; von Arx, J.A. 1973. Pyrenomycetes:Meliolales, Coronophorales, Sphaeriales. In: Ains-worth, G.C., Sparrow, F.K.; Sussman, A.S., eds. Thefungi. New York: Academic Press: 87-132. Vol. 4A.

Munch, E. 1907. Die blaufaule des nadelhoizes.Naturwissenschaftliche Zeitschrift fur Forst-undLandwirtschift. 5:531-573.

Nag Raj, T.R.; Kendrick, W.B. 1975. A monograph ofChalara and allied genera. Waterloo: WilfredLaurier University Press. 200 p.

Nannfeldt, J.A. 1932. Studien uber die morphologieund systematik der nichtlichenisierten inoper-culaten Discomyceten. Nova Acta Regiae SocietatisScientiarum Upsaliensis. Ser. 4. 8(2):1-368.

Nelson, R.M. 1934. Effect of bluestain on southernpines attacked by bark beetles. PhytopathologischeZeitschrift. 7:327353.

Olchowecki, A.; Reid, J. 1974. Taxonomy of the genusCeratocystis in Manitoba. Canadian Journal ofBotany. 52:1675-1711.

Parker, A.K. 1957. Europhium, a new genus of the As-comycetes with a Leptographium imperfect state.Canadian Journal of Botany. 35:173-179.

Rane, K.K.; Tattar, T.A. Pathogenicity of blue-stainfungi associated with Dendroctonus terebrans.Plant Disease. 71:879-883.

Redhead, S.A.; Malloch, D. 1977. The Endomy-cetaceae: new concepts, new taxa. Canadian Jour-nal of Botany. 55:1701-1711.

Robinson, R.C. 1962. Blue stain fungi in lodgepolepine (Pinus contorta Dougl. var. Zatifolia Engelm.)infested by the mountain pine beetle (Dendroctonusmonticolae Hopk.). Canadian Journal of Botany.40:609-614.

Robinson-Jeffrey, R.C.; Davidson, R.W. 1968. Threenew Europhium species with Verticicladiella im-perfect states on blue-stained pine. Canadian Jour- ,nal of Botany. 46:1523-1527.

Robinson-Jeffrey, R.C.; Grinchenko, A.H.H. 1964. Anew fungus in the genus Ceratocystis occurring on

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blue-stained lodgepole pine attacked by bark beet-les. Canadian Journal of Botany. 42:527-532.’

Rosinski, M.A. 1961. Development of the ascocarp ofCerutocystis ulmi. American Journal of Botany.48:285-293.

Rosinski, M.A. 1965. Further confirmation of the oc-currence of cellulose in Cerutocystis ulmi. Mycolo-gia. 57:668.

Rosinski, M.A.; Campana, R.J. 1964. Chemicalanalysis of the cell wall of Cerutocystis ulmi.Mycologia. 56~738-744.

Rumbold, C.T. 1931. Two blue-staining fungi as-sociated with bark-beetle infestation of pines. Jour-nal of Agricultural Research. 43:847-873.

Rumbold, C.T. 1936. Three blue-staining fungi, in-cluding two new species, associated with bark bee-tles. Journal of Agricultural Research. 52:419-437.

Rumbold, C.T. 1941. A blue-stain fungus, Cerutocystismontium n. sp. and some yeasts associated with twospecies of Dendroctonus. Journal of AgriculturalResearch. 62:589-601.

Scerbin-Parfenenko, A.L. 1953. Rakovye a sosudistyebolezni listvennych porod. Goslesbamizdat. [Vol.no. unknown] l-89.

Sigler, L.; Hawksworth, D.L. 1987. International com-mission on the taxonomy of fungi (ITCF) code ofpractice for systemic mycologists. Mycopathologia.99:3-7.

Smith, M.J.; Patik, C.M.; Rosinski, M.A. 1967. A com-parison of cellulose production in the genusCerutocystis. Mycologia. 59:965-969.

Solheim, H. 1986. Species of Ophiostomataceae iso-lated from Piceu ubies infested by the bark beetleIps typogrup hus . Nordic Journal of Botany.6:199-207.

Spencer, J.F.T.; Gorin, P.A.J. 1971. Systematics of thegenera Cerutocystis and Gruphium. Proton mag-netic resonance spectra of the mannose-containingpolysaccharides as an aid in classification. My-cologia. 63:387-402.

Sydow, Von H.; P. Sydow. 1919. Mykologische mit-teilungen. Annales Mycologici. 17(1):3-47.

Tsuneda, A.; Hiratsuka, Y. 1984. Sympodial and an-nellidic conidiation in Cerutocystis cluvigeru.Canadian Journal of Botany. 62:2618-2624.

Upadhyay, H.P. 1978. Proposal for the conservation ofthe generic name Cerutocystis Ell. & Halst. (1890)against Sphueronuemellu Karsten (1884). Taxon.27:553.

Upadhyay, H.P. 1981. A monograph of the genusCerutocystis and Cerutocystiopsis. Athens: Univer-sity of Georgia Press. 176 p.

Upadhyay, H.P.; Kendrick, W.B. 1975. Prodmodus fora I cvision of Cerutocystis (Microascales, Ascomy-cetes) and its conidial states. Mycologia. 67:798-805.

Urosevic, B. 1983. Tracheomycotic diseases in oak.Communicationes Instituti Forestalis Cechoslo-veniae. 13:85-100.

van Wyk, P.W.J.; Wingfield, M.J. 1990. Ascosporedevelopment in Cerutocystis sensu Zuto (Fungi): areview. Bothalia. 20(2):141-145.

von Arx, J.A. 1970. The genera of fungi sporulating inpure culture. Lehre: Cramer. 288 p.

von Arx, J.A. 1974. The genera of fungi sporulating inpure culture. 2nd ed. Vaduz: Cramer. 315 p.

von Arx, J.A. 1979. Ascomycetes as Fungi Imperfecti.In: Kendrick, B., ed. The whole fungus. Ottawa: Na-tional Museums of Canada: 201-214. Vol. 1.

von Arx, J.A. 1987. A re-evaluation of the Eurotiales.Persoonia. 13:273-300.

von Arx, J.A.; Muller, E. 1954. Die Gattungen deramerosporen Pyrenomyceten. Beitrage zur Kryp-togamenflora der Schweiz. 11: l-434.

von At-x, J.A.; van der Walt, J.P. 1988. Ophiostoma-tales and Endomycetales. In: de Hoog, G.S.; Smith,M.Th.; Weijman, A.C.M., eds. The expanding realmof yeast-like fungi: Proceedings of an internationalsymposium on the perpectives of taxonomy, ecologyand phylogeny .of yeast and yeast-like fungi; 1987Aug. 3-7; Amersfort, The Netherlands. Amsterdam:Elsevier Science: 167-176.

von Hohnel, F. 1918. Mykologische fragmente CXCV.Uber die gattung Purodiopsis Maublanc. AnnalesMycologici. 16:40-4 1.

Weijman, A.C.M.; de Hoog, G.S. 1975. On the sub-division of the genus Cerutocystis. Antonie vanLeeuwenhoek. Journal of Microbiology and Serol-ogy. 41:353-360.

Whitney, H.S. 1971. Association of Dendroctonusponderosue (Coleoptera:Scolytidae) with blue stainfungi and yeasts during brood development inlodgepole pine. Canadian Entomologist. 103:1495-1503.

Whitney, H.S. 1982. Relationships between bark beet-les and symbiotic organisms. In: Mitton, J.B.; Stur-geon, K., eds. A system for the study of evolutionarybiology. Austin: University of Texas Press: 183-211.

Whitney, H.S.; Cobb, F.W., Jr. 1972. Non-stainingfungi associated with the bark beetle Dendroctonusbrevicomis (Coleoptera:Scolytidae) on Pinus pan,-derosue. Canadian Journal of Botany. 50:1943-1945.

Wingfield, M.J. 1983. Association of Verticicludielluproceru and Leptogruphium terebruntis with insectsin the Lake States. Canadian Journal of Forest Re-search. 13:1238-1245.

Wingfield, M.J. 1985. Reclassification of Ver-ticicludiellu based on conidial development. Trans-actions of the British Mycological Society. 85:81-93.

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Wingfield, M.J.; van Wyk, P.S.; Marasas, W.F.O.1988. Ceratocystiopsis proteae sp. nov., with a newanamorph genus. Mycologia. 80:23-30.

Wood, S.L. 1982. The bark and ambrosia beetles ofNorth and Central America (Coleoptera: Scolyti-dae), a taxonomic monograph. Great Basin Natural-

ist Memoirs # 6. Provo, UT: Brigham Young Univer-sity. 1359 p.

Wright, E.F.; Cain, R.F. 1961. New species ofCeratocystis. Canadian Journal of Botany. 39:1215-1230.

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Perry, Thelma J. 1991. A synoposis of the taxonomic revisions in thegenus Cerutocystis including a review of blue-staining species as-sociated with Dendroctonus bark beetles. Gen. Tech. Rep. SO-86.New Orleans, LA: US. Department of Agriculture, Forest Service,Southern Forest Experiment Station. 16 p.

Lists Ophiostoma species previously described as Ceratocystis andsummarizes events that led to revisions in the genera.

Keywords: Ceratocystiopsis, Ophiostoma

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