Epitypification: should we epitypify?§

Epitypification: should we epitypify? § § 1 2 first-author †E-mail:kdhyde1@gmail.com †E-mail:kdhyde1@gmail.com kdhyde1@gmail.com 3 International Fungal Research and Development Centre, the Research Institute of Resource Insects, Chinese Academy of Forestry, Kunming 650224, China International Fungal Research and Development Centre, the Research Institute of Resource Insects, Chinese Academy of Forestry, Kunming 650224, China Fungal Research Group, School of Science, Mae Fah Luang University, Tasud, Muang, Chiang Rai 57100, Thailand Fungal Research Group, School of Science, Mae Fah Luang University, Tasud, Muang, Chiang Rai 57100, Thailand Centre for Research in Fungal Diversity, School of Biological Sciences, University of Hong Kong, Hong Kong, China Centre for Research in Fungal Diversity, School of Biological Sciences, University of Hong Kong, Hong Kong, China Epitypification can solve many taxonomic problems and stabilize the understanding of species, genera, families or orders. The aim of this paper is to illustrate how to epitypify. A few examples where taxa have been epitypified are considered and the benefits and disadvantages of epitypification are discussed. We also outline some examples of taxa which need to be epitypified with reasons. Colletotrichum GenBank Molecular phylogeny Taxonomy Typification INTRODUCTION 1 What can be done if the material that represents a type species, genus, family or order is lost, is in poor condition, cannot be used to extract DNA, or molecular data cannot be obtained? How do we form concepts of species, genera, families and orders in these cases? Recent publications from the Assembling the Fungal Tree of Life (AFTOL) project have named exemplar genera to conceptualize orders (Hibbett et al., 2007 2007 ). However, the understanding of the genera is included, and thus the understanding of the order, may be interpreted differently by various mycologists, and therefore this approach cannot be recommended. In the case of genera, species descriptions, but not types, have often been used to represent genera (Kodsueb et al., 2006 2006 ). Stylized drawings and wrongly identified collections, however, might mean that the generic concept is understood differently by various mycologists. Fresh isolates are also generally used to obtain DNA data, but one person’s understanding of a species may differ from another’s; thus the sequences in GenBank and other public databases are surely unreliable (Nilsson et al., 2006 2006 ). This paper reviews the options that can be used to address the above problems. Epitypification is the best option for replacing lost type material or types in poor condition. If DNA cannot be extracted from types, then we need to designate epitypes that are identical to the examined types. When fresh collections are sequenced and those sequences are deposited in GenBank, dried herbarium material and cultures should also be deposited in accessible herbaria culture collections. If these rules are followed then we will be able to avoid GenBank becoming a dustbin for DNA sequences whose names cannot be verified by morphology. WHAT IS EPITYPIFICATION? 1 Article 9.7 of the International Code of Botanical Nomenclature (Vienna Code) (Mcneill et al., 2006 2006 ) states that “An epitype is a specimen or illustration selected to serve as an interpretative type when the holotype, lectotype, or previously designated neotype, or all original material associated with a validly published name, is demonstrably ambiguous and cannot be critically identified for purposes of the precise application of the name of a taxon. When an epitype is designated, the holotype, lectotype, or neotype that the epitype supports must be explicitly cited.” It has become relatively common to epitypify fungi (Alves et al., 2008 2008 ; Crous et al., 2007 2007 ; Phillips et al., 2006 2006 ; 2007 2007 ; Shenoy et al., 2007b 2007b ; Than et al., 2008 2008 ) and this paper will examine whether it is a good idea to epitypify fungi and will also point out some of the associated problems. There are many reasons to epitypify. Often the type material has been lost or is in poor condition. The genus may be relatively well known, e.g., Massaria inquinans (Tode) De Not.; however, unless there is a type that can serve to represent the genus, it is possible that each individual mycologist may have a different understanding of the taxon representing this genus. More recently with the advent of molecular phylogenetics, living material is needed for sequencing as in most cases the ancient herbarium specimens cannot be successfully sequenced. Therefore, even if the type material is in relatively good condition, one could argue that the taxon needs epitypifying, so that living material is available for gene research. This is evident in the paper by Hibbett et al.( 2007 2007 ). In that paper, each order was represented by 1~6 exemplar genera. The understanding of each genus, however, may differ. Many mycologists may not have seen good type material and their understanding of a taxon/genus/family may be based on literature or stylized drawings of the type or representative (possibly misidentified) collections. The individual understanding of an order may, therefore, be questionable. The AFTOL project has sequenced numerous taxa and derived a framework for modern taxonomy (Aime et al., 2006 2006 ; Binder and Hibbett, 2006 2006 ; Celio et al., 2006 2006 ; Geiser et al., 2006 2006 ), yet few of the fungi sequenced were types. This could lead to erroneous data being generated and since strains may not have been deposited in culture collections and herbarium material is scarcely available, thus there is no way to repeat the experiment (Shenoy et al., 2007a 2007a ). HOW TO DECIDE ON AN EPITYPE 1 According to the definition of an epitype (Mcneill et al., 2006 2006 ), the epitype should be identical to previous type material. Hence before epitypifying a species, the type material should be located, loaned and carefully studied. This should include detailed examination of morphological characters and other data such as host, location, or anamorphic relationship. The next step is to obtain a fresh specimen. The epitype specimen should undoubtedly have all of the characters exhibited in the type and if possible be obtained from the same location and host. Of course this is not always possible. We have recently searched for a fresh collection of Colletotrichum circinans (Berk.) Vogl., the cause of smudge in onion (Walker, 1925 1925 ); however, the original site is now a housing estate. We have also searched for the type of Colletotrichum falcatum Went which was described from a small village in central Java (Went, 1893 1893 ). Amazingly the sugarcane plantation is in its original location (although part has been destroyed for housing) and during a recent trip we were able to obtain a fresh collection. DNA data obtained from different strains isolated from different locations or environments are often identical. For example, the internal transcribed spacer (ITS) sequences of two isolates of Botryosphaeria cortices (Demaree & M.S. Wilcox) Arx & E. Müll. from North Carolina showed no significant difference from those isolated from New Jersey from the same host (Phillips et al., 2006 2006 ). The rDNA (28S, 18S) and RNA polymerase II (RPB2) sequences of Trematosphaeria pertusa Fuckel on a dead stump of Fraxinus excelsior L. from Deux Sèvres, France, were almost the same with those of the one from Haute Garonne (France) occurring on submerged wood of Platanus (Zhang et al., 2008 2008 ). Thus as long as the collections are morphologically identical to the type, a fresh collection from a different location could feasibly be designated as an epitype. It has recently proven possible to extract DNA and obtain the DNA sequences from old herbarium material with kits (Fredricks et al., 2005 2005 ; O′Gorman et al., 2008 2008 ). This material may not be the type, but can be collections identified by the same mycologist who designated the type, or collections by others from the same host and location. This DNA data is very important to determine the phylogenic status of a particular fungus, especially for those rarely encountered taxa. Furthermore, if it was collected by the original mycologist it is likely to be authentic. As long as enough DNA data can be obtained from this old material which should be identical to the type, it could also be designated as an epitype. EXAMPLES OF EPITYPIFICATION 1 By epitypification, a name can be fixed to a specimen or a culture, which is very important for phylogenetic study of this taxon. A fresh collection of Botryosphaeria corticis was made from a commercial field of Vaccinium corymbosum L. in the USA and a culture was obtained. Epitypification of Botryosphaeria corticis with this specimen was carried out by designating it as the epitype as none of the previous cultures was connected with the type specimen (BPI 598729) (Phillips et al., 2006 2006 ). Because of this, the name of the fungus causing an important disease of blueberry (cane canker) has been stabilized. Collectotrichum capsici (Syd.) Butler & Bisby is an economically important taxon that causes anthracnose in chilli. However, molecular data could not be obtained from the type specimen. The authors therefore obtained a fresh collection from chilli in Coimbatore in India and designated this as the epitype of C . capsici . More importantly they obtained living cultures and deposited these in public culture collections (as ex-epitype living cultures). The phylogenetic status of this taxon has thus been stabilized and cultures are freely available for future work in this important pathogen and pathogenic genus (Shenoy et al., 2007b 2007b ). Similarly, the phylogenetic characters of Mycosphaerella and Teratosphaeria have also been clarified by designating the epitypes of the type species of these two genera (Verkley et al., 2004 2004 ; Crous et al., 2007 2007 ). Some specimens, especially old types, can be very difficult to locate or obtain. For example, the neo-type specimen of Trematosphaeria pertusa (type species of Trematosphaeria ) is kept in Persoon’s exsiccata in the University of Leiden (L). According to the rule of the herbarium, this specimen could only be studied in the herbarium, and it is very difficult to obtain funding for such. Thus this has been poorly studied (Boise, 1985 1985 ), and limited morphological characters for this taxon can be found in the protologue. With such a situation, designation of an epitype would be beneficial. Sometimes, the traditional concept for a species differs from that observed in the type protologue and early descriptions. This is the case with Massaria inquinans (Tode) De Not., the type species of Massaria , which represents the family Massariaceae. We have been unable to locate the type material of this species. Shoemaker and LeClair ( 1975 1975 ) systematically studied the Massariaceae and examined a specimen of Massaria inquinans (CLXXX) lodged at University of Uppsala (UPS). Shoemaker and Kokko ( 1977 1977 ) reported this taxon to have asci with 8 ascospores with no-constriction at each septum. We have examined the specimen of UPS CLXXX and found it to have asci with 4 ascospores that are constricted at the central septum. This species therefore needs epitypification so that all mycologists have the same basic understanding of this taxon, genus and family (Massariaceae). We are studying the type specimens of genera of the order Pleosporales and have found that about 20%~30% of herbarium specimens are in bad condition, and little morphological data could be obtained from these types. For example, the type specimen of Teichospora trabicola Fuckel (G00110113), syntype of Metacoleroa dickiei (Berk. & Br.) Petr. (K(M) 143928), the type specimen of Leptosphaeria michotii (Westend.) Sacc. (Belgium, 89509-75) (as Sphaeria michotii Westend.), the holotype specimen of Pleospora scirpicola (DC.) P. Karst. (as Sphaeria scirpicola DC.) (G00110110) are all in poor condition. Even when specimens are in good condition, it is not always possible to evaluate some valuable characters, such as ascospore sheaths, type of ascus dehiscence, and these are characters that might be informative in classification (Eriksson, 1982 1982 ; Hawksworth, 1994 1994 ). Therefore fresh material should be located and detailed descriptions of these taxa should be published (after comparison with the type) and designated as epitypes which should include living material for future study. POSSIBLE PROBLEMS ASSOCIATED WITH EPITYPIFICATION 1 It is very important that an epitype is identical to the original type; however, since the type may be lost, or in poor condition, it is not always possible to verify this. Therefore subjective judgement has to be used to decide on a specimen that can be designated as an epitype. Thus, although there may be a chance of error, designating an epitype is a better approach to move forward, than having many interpretations of what characters of the type species of a genus, family or order may comprise. There is a chance that the type may be newly located at a later date; however, this is unlikely. If this did happen, the original type (if in good condition, freely available and DNA can be extracted and sequenced) should represent the species. It is very unlikely, however, that an isolate of the type could be located at a later date. Colletotrichum acutatum Simmonds is an important pathogen that was introduced by Simmonds ( 1965 1965 ) without a type, and validated in 1968 with a broad concept. This was demonstrated by the selection of several type specimens from a range of hosts (Simmonds, 1968 1968 ). Than et al.( 2008 2008 ) considered that Simmonds ( 1965 1965 ; 1968 1968 )’s broad concept of C . acutatum has created uncertainties in the species concept. No viable ex-type cultures of C . acutatum were located and further there were no viable cultures of this taxon on Papaya from the type locality. Than et al. ( 2008 2008 ) therefore designated a specimen of living culture stored at Queensland Department of Primary Industries and Fisheries, Plant Pathology Herbarium, Indooroopilly, Australia (BRIP 28519), as an epitype from Papaya (same host as type) from Yandina (nearby location). Subsequent to this publication, it has been discovered that a culture of C . acutatum had been sent by Simmonds to Dingley in New Zealand (and then to ATCC 56816) (Guerber and Correll, 2001 2001 ). This culture, however, was not derived ex-holotype. The culture Simmonds sent to Dingley is given as culture No. 16633D [p.225 in (Guerber and Correll, 2001 2001 )]. This culture was also sent by Simmonds to Herbarium of CABI Bioscience UK Centre (IMI) where it was assigned to IMI 117620 and is a “pink type” [Table 1 in (Than et al., 2008 2008 )]. The holotype (IMI 117617) was derived from Simmonds culture No. 16741B1, which is a different isolate. ATCC 56816 is therefore an ex-paratype culture of C . acutatum . The designation of an epitype by Than et al. ( 2008 2008 ) sought to stabilize the status of C . acutatum ; however, since an ex-paratype culture has been found, this creates an interesting problem. The ex-paratype is poorly characterised and is unavailable to researchers unless the high costs of buying these cultures from American Type Culture Collection (ATCC) are paid. The ex-epitype is well characterised, is freely available (in six public culture collections) and has been sequenced using several genes. We would therefore recommend that the epitype is used to represent this species as it was derived from the same host in the same area and has identical morphology to the type. Others may disagree and thus attempt by Than et al.( 2008 2008 ) to stabilize the species may not have worked. Interestingly, the story does not stop here! Colletotrichum acutatum isolates cluster into groups based on restriction fragment length polymorphism (RFLP) of mitochondrial DNA (mtDNA), RFLPs of the 900-bp glutamine synthase (GS) intron and sequence analysis of the glutaraldehyde-3-phosphate dehydrogenase (G3PD) intron 2, and the GS intron 2 gene (Guerber et al., 2003 2003 ; MacKenzie et al., 2008 2008 ; Peres et al., 2008 2008 ). These groups might represent phylogenetically distinct species of C . acutatum sensu lato (Guerber et al., 2003 2003 ). It will be interesting to establish which group(s) the ex-paratype with pink cultures and the epitype with orange to grey cultures cluster in. There will no doubt be more written on this subject. References MC Aime , PB Matheny , DA Henk , EM Frieders , RH Nilsson , M Piepenbring , DJ McLaughlin , LJ Szabo , D Begerow , JP Sampaio , 等 . An overview of the higher-level classification of Pucciniomycotina based on combined analyses of nuclear large and small subunit rDNA sequences . Mycologia , 2006 . 98 ( 6 ): 896 - 905 . DOI: 10.3852/mycologia.98.6.896 http://doi.org/10.3852/mycologia.98.6.896 . An overview of the higher-level classification of Pucciniomycotina based on combined analyses of nuclear large and small subunit rDNA sequences A Alves , PW Crous , A Correia , AJL Phillips . Morphological and molecular data reveal cryptic speciation in Lasiodiplodia theobromae . Fungal Diversity , 2008 . 28 ( 1 ): 1 - 13 . . Morphological and molecular data reveal cryptic speciation in Lasiodiplodia theobromae M Binder , DS Hibbett . Molecular systematics and biological diversification of Boletales . Mycologia , 2006 . 98 ( 6 ): 971 - 983 . DOI: 10.3852/mycologia.98.6.971 http://doi.org/10.3852/mycologia.98.6.971 . Molecular systematics and biological diversification of Boletales JR Boise . An amended description of Trematosphaeria . Mycologia , 1985 . 77 ( 2 ): 230 - 237 . DOI: 10.2307/3793072 http://doi.org/10.2307/3793072 . An amended description of Trematosphaeria GJ Celio , M Padamsee , BTM Dentinger , R Bauer , DJ McLaughlin . Assembling the fungal tree of life: constructing the structural and biochemical database . Mycologia , 2006 . 98 ( 6 ): 850 - 859 . DOI: 10.3852/mycologia.98.6.850 http://doi.org/10.3852/mycologia.98.6.850 . Assembling the fungal tree of life: constructing the structural and biochemical database PW Crous , U Braun , JZ Groenewald . Mycosphaerella is polyphyletic . Studies in Mycology , 2007 . 58 ( 1 ): 1 - 32 . . Mycosphaerella is polyphyletic O Eriksson . The families of bitunicate ascomycetes . Opera Botanica , 1982 . 60 1 - 220 . . The families of bitunicate ascomycetes DN Fredricks , C Smith , A Meier . Comparison of six DNA extraction methods for recovery of fungal DNA as assessed by quantitative PCR . Journal of Clinical Microbiology , 2005 . 43 ( 10 ): 5122 - 5128 . DOI: 10.1128/JCM.43.10.5122-5128.2005 http://doi.org/10.1128/JCM.43.10.5122-5128.2005 . Comparison of six DNA extraction methods for recovery of fungal DNA as assessed by quantitative PCR D Geiser , M Gueidan , J Miadlikowska , F Kauff , V Hofstetter , E Fraker , CL Schoch , L Tibell , WA Untereiner , A Aptroot , 等 . Eurotiomycetes: Eurotiomycetidae and Chaetothyriomycetidae . Mycologia , 2006 . 98 ( 6 ): 1053 - 1064 . DOI: 10.3852/mycologia.98.6.1053 http://doi.org/10.3852/mycologia.98.6.1053 . Eurotiomycetes: Eurotiomycetidae and Chaetothyriomycetidae JC Guerber , JC Correll . Characterization of Glomerella acutata the teleomorph of Colletotrichum acutatum . Mycologia , 2001 . 93 ( 1 ): 216 - 229 . DOI: 10.2307/3761619 http://doi.org/10.2307/3761619 . Characterization of Glomerella acutata the teleomorph of Colletotrichum acutatum JC Guerber , B Liu , JC Correll , PR Johnston . Characterization of diversity in Colletotrichum acutatum sensu lato by sequence analysis of two gene introns, mtDNA and intron RFLPs, and mating compatibility . Mycologia , 2003 . 95 ( 5 ): 872 - 895 . DOI: 10.2307/3762016 http://doi.org/10.2307/3762016 . Characterization of diversity in Colletotrichum acutatum sensu lato by sequence analysis of two gene introns, mtDNA and intron RFLPs, and mating compatibility DL Hawksworth . Ascomycete Systematics: Problems and Perspectives in the Nineties , 1994 . : New York Plenum Press , . Ascomycete Systematics: Problems and Perspectives in the Nineties DS Hibbett , M Binder , JF Bischoff , M Blackwell , PF Cannon , OE Eriksson , S Huhndorf , TY James , PM Kirk , R Lucking , 等 . A higher-level phylogenetic classification of the fungi . Mycological Research , 2007 . 111 ( 5 ): 509 - 547 . DOI: 10.1016/j.mycres.2007.03.004 http://doi.org/10.1016/j.mycres.2007.03.004 . A higher-level phylogenetic classification of the fungi R Kodsueb , R Jeewon , D Vijaykrishna , EHC McKenzie , P Lumyong , S Lumyong , KD Hyde . Systematic revision of Tubeufiaceae based on morphological and molecular data . Fungal Diversity , 2006 . 21 105 - 130 . . Systematic revision of Tubeufiaceae based on morphological and molecular data SJ MacKenzie , NA Peres , MP Barquero , LF Arauz , LW Timmer . Host range and genetic relatedness of Colletotrichum acutatum isolates from fruit crops and leatherleaf fern in Florida . Phytopathology , 2008 . in press . Host range and genetic relatedness of Colletotrichum acutatum isolates from fruit crops and leatherleaf fern in Florida J Mcneill , FR Barrie , HM Burdet , V Demoulin , DL Hawksworth , K Marhold , DH Nicolson , J Prado , PC Silva , JE Skog , 等 . International Code of Botanical Nomenclature (Vienna Code) , 2006 . (Available from: http://ibot.sav.sk/icbn/main.htm http://ibot.sav.sk/icbn/main.htm ) (Accessed Aug. 2008) . International Code of Botanical Nomenclature (Vienna Code) RH Nilsson , M Ryberg , E Kristiansson , K Abarenkov , KH Larsson , U Kolgalg . Taxonomic reliability of DNA sequences in public sequence databases: a fungal perspective . PLoS ONE , 2006 . 1 ( 1 ): e59 DOI: 10.1371/journal.pone.0000059 http://doi.org/10.1371/journal.pone.0000059 . Taxonomic reliability of DNA sequences in public sequence databases: a fungal perspective DT O′Gorman , PL Sholberg , SC Stokes , J Ginns . DNA sequence analysis of herbarium specimens facilitates the revival of Botrytis mali , a postharvest pathogen of apple . Mycologia , 2008 . 100 ( 2 ): 227 - 235 . DOI: 10.3852/mycologia.100.2.227 http://doi.org/10.3852/mycologia.100.2.227 . DNA sequence analysis of herbarium specimens facilitates the revival of Botrytis mali, a postharvest pathogen of apple NA Peres , SJ MacKenzie , TL Peever , LW Timmer . Postbloom fruit drop of citrus and key lime anthracnose are caused by distinct phylogenetic lineages of Colletotrichum acutatum . Phytopathology , 2008 . 98 ( 3 ): 345 - 352 . DOI: 10.1094/PHYTO-98-3-0345 http://doi.org/10.1094/PHYTO-98-3-0345 . Postbloom fruit drop of citrus and key lime anthracnose are caused by distinct phylogenetic lineages of Colletotrichum acutatum AJL Phillips , PV Oudemans , A Correia , A Alves . Characterisation and epitypification of Botryosphaeria corticis , the cause of blueberry cane canker . Fungal Diversity , 2006 . 21 141 - 155 . . Characterisation and epitypification of Botryosphaeria corticis, the cause of blueberry cane canker AJL Phillips , PW Crous , A Alves . Diplodia seriata , the anamorph of “ Botryosphaeria ” obtusa . Fungal Diversity , 2007 . 25 141 - 155 . . Diplodia seriata, the anamorph of “Botryosphaeria” obtusa BD Shenoy , R Jeewon , KD Hyde . Impact of DNA sequence-data on the taxonomy of anamorphic fungi . Fungal Diversity , 2007 . 26 1 - 54 . . Impact of DNA sequence-data on the taxonomy of anamorphic fungi BD Shenoy , R Jeewon , WH Lam , DJ Bhat , PP Than , WJ Talor , KD Hyde . Morpho-molecular characterisation and epitypification of Colletotrichum capsici (Glomerellaceae, Sordariomycetes), the causative agent of anthracnose in chilli . Fungal Diversity , 2007 . 27 197 - 211 . . Morpho-molecular characterisation and epitypification of Colletotrichum capsici (Glomerellaceae, Sordariomycetes), the causative agent of anthracnose in chilli RA Shoemaker , PM LeClair . Type studies of Massaria from Wehmeyer collection . Canadian Journal of Botany , 1975 . 53 ( 15 ): 1568 - 1598 . DOI: 10.1139/b75-188 http://doi.org/10.1139/b75-188 . Type studies of Massaria from Wehmeyer collection RA Shoemaker , EG Kokko . Fungi Canadenses No. 104 . Massaria inquinans , 1977 . Fungi Canadenses No. 104 JH Simmonds . A study of the species of Colletotrichum causing fruit rots in Queensland . Queensland Journal of Agriculture and Animal Science , 1965 . 22 437 - 459 . . A study of the species of Colletotrichum causing fruit rots in Queensland JH Simmonds . Type specimens of Colletotrichum gloeosporioides var. minor and Colletotrichum acutatum . Queensland Journal of Agriculture and Animal Science , 1968 . 25 178A . Type specimens of Colletotrichum gloeosporioides var. minor and Colletotrichum acutatum PP Than , RG Shivas , R Jeewon , S Pongsupasamit , TS Marney , PWJ Taylor , KD Hyde . Epitypification and phylogeny of Colletotrichum acutatum J.H. Simmonds . Fungal Diversity , 2008 . 28 97 - 108 . . Epitypification and phylogeny of Colletotrichum acutatum J.H. Simmonds GJM Verkley , PW Crous , JZ Groenewald , U Braun , A Aptroot . Mycosphaerella punctiformis revisited: morphology, phylogeny, and epitypification of the type species of the genus Mycosphaerella (Dothideales, Ascomycota) . Mycol Res , 2004 . 108 ( 11 ): 1271 - 1282 . DOI: 10.1017/S0953756204001054 http://doi.org/10.1017/S0953756204001054 . Mycosphaerella punctiformis revisited: morphology, phylogeny, and epitypification of the type species of the genus Mycosphaerella (Dothideales, Ascomycota) JC Walker . Studies on disease resistance in the onion . The Proceedings of the National Academy of Sciences , 1925 . 11 ( 3 ): 183 - 189 . DOI: 10.1073/pnas.11.3.183 http://doi.org/10.1073/pnas.11.3.183 . Studies on disease resistance in the onion FAFC Went . Het rood snot . Arch Java Suikerindustr , 1893 . 1 265 - 282 . (in Dutch) . Het rood snot Y Zhang , J Fournier , KD Hyde . Are Melanomma pulvis-pyrius and Trematosphaeria pertusa congeneric? . Fungal Diversity , 2008 . in press . Are Melanomma pulvis-pyrius and Trematosphaeria pertusa congeneric? Dr. Roger Shivas (Queensland Plant Pathology Herbarium) and Dr. Peter Johnson (Landcare New Zealand) are thanked for bringing to the attention of Dr. Hyde the problems associated with the C . acutatum epitype. 10.1631/jzus.B0860004 2008-07-31 2008-08-30 2022-01-05 Copyright © 2008, Journal of Zhejiang University Science 2008 Journal of Zhejiang University-SCIENCE B (Biomedicine and Biotechnology) 10 09