Skip to main content

Advertisement

We're creating a new version of this page. See preview

  • Article
  • Open Access

Diaporthe is paraphyletic

Contributed equally
IMA Fungus20178:801153

https://doi.org/10.5598/imafungus.2017.08.01.11

  • Received: 25 March 2017
  • Accepted: 22 May 2017
  • Published:

Abstract

Previous studies have shown that our understanding of species diversity within Diaporthe (syn. Phomopsis) is limited. In this study, 49 strains obtained from different countries were subjected to DNA sequence analysis. Based on these results, eight new species names are introduced for lineages represented by multiple strains and distinct morphology. Twelve Phomopsis species previously described from China were subjected to DNA sequence analysis, and confirmed to belong to Diaporthe. The genus Diaporthe is shown to be paraphyletic based on multi-locus (LSU, ITS and TEF1) phylogenetic analysis. Several morphologically distinct genera, namely Mazzantia, Ophiodiaporthe, Pustulomyces, Phaeocytostroma, and Stenocarpella, are embedded within Diaporthe s. lat., indicating divergent morphological evolution. However, splitting Diaporthe into many smaller genera to achieve monophyly is still premature, and further collections and phylogenetic datasets need to be obtained to address this situation.

Key words

  • Ascomycota
  • Diaporthales
  • Phomopsis
  • phylogeny
  • taxonomy

Introduction

Species of Diaporthe are known as important plant pathogens, endophytes or saprobes (Udayanga et al. 2011, Gomes et al. 2013). They have broad host ranges, and occur on many plant hosts, including cultivated crops, trees, and ornamentals (Diogo et al. 2010, Thompson et al. 2011, Gomes et al. 2013, Huang et al. 2015). Some Diaporthe species are responsible for severe diebacks, cankers, leaf-spots, blights, decay or wilts on different plant hosts, several of which are economically important (Mostert et al. 2001, Van Rensburg et al. 2006, Thompson et al. 2011, Gomes et al. 2013), leading to serious diseases and significant yield losses (Santos et al. 2011). For example, Diaporthe helianthi is the cause of one of the most important diseases of sunflower (Helianthus annuus) worldwide, and has reduced production by up to 40% in Europe (Masirevic & Gulya 1992, Thompson et al. 2011). Diaporthe neoviticola and D. vitimegaspora, the causal agents of leaf-spot and swelling arm, are known as severe pathogens of grapevines (Vitis vinifera) (Van Niekerk et al. 2005). Ürbez-Torres et al. (2013) indicated that D. neoviticola was one of the most prevalent fungi isolated from grapevine perennial cankers in declining vines. Diaporthe scabra has been reported causing cankers and dieback on London plane (Platanus acerifolia) in Italy (Grasso et al. 2012). Symptoms of umbel browning and necrosis caused by D. angeliace have been regularly observed on carrots in France, resulting in seed production losses since 2007 (Ménard et al. 2014). Avocado (Persea americana), cultivated worldwide in tropical and subtropical regions, is threatened by branch cankers and fruit stem-end rot diseases caused by D. foeniculina and D. sterilis (Guarnaccia et al. 2016). Furthermore, species of Diaporthe are commonly introduced into new areas as endophytes or latent pathogens along with plant produce. For instance, Torres et al. (2016) reported D. rudis causing stemend rot in avocados in Chile, which was imported via avocado fruit from California (USA). Some endophytes have been shown to act as opportunistic plant pathogens. Diaporthe foeniculina (syn. P. theicola), which is a common endophyte, has been shown to cause stem and shoot cankers on sweet chestnut (Castanea sativa) in Italy (Annesi et al. 2015, Huang et al. 2015). Because of this unique ecology and potential role as plant pathogens, it is of paramount importance to accurately identify species of Diaporthe to facilitate disease surveillance, control, and trade.

The initial species concept of Diaporthe based on the assumption of host-specificity, resulted in the introduction of more than 1000 names (http://www.indexfungorum.org/Names/Names.asp); (Gomes et al. 2013, Gao et al. 2016). In recent years, however, a polyphasic approach employing multi-locus DNA data together with morphology and ecology has been employed for species delimitation in the genus (Udayanga et al. 2011, Gomes et al. 2013). The nuclear ribosomal internal transcribed spacer (ITS), the translation elongation factor 1-α (TEF1), β-tubulin (TUB), histone H3 (HIS), and calmodulin (CAL) genes are the most commonly used molecular loci for the identification of Diaporthe spp. (Dissanayake et al. 2015, Udayanga et al. 2015, Huang et al. 2015, Santos et al. 2017). Furthermore, molecular marker aids are being used to rapidly identify Diaporthe species which tend to be morphologically conserved (Udayanga et al. 2012, Tan et al. 2013, Lombard et al. 2014, Thompson et al. 2015, Huang et al. 2015). However, defining species boundaries remains a major challenge in Diaporthe (Huang et al. 2015), which may be a consequence of limited sampling or the use of DNA loci with insufficient phylogenetic resolution (Liu et al. 2016). It has therefore been proposed that new species in the genus should be introduced with caution, and that multiple strains from different origins should be subjected to a multi-gene phylogenetic analysis to determine intraspecificvariation (Liu et al. 2016).

The generic relationships of Diaporthe with other genera in Diaporthaceae remain unclear. The family name Diaporthaceae was established by Wehmeyer (1926) to accommodate Diaporthe, Mazzantia, Melanconis, and some other genera, mainly based on morphological characters such as the position, structure, and arrangement of ascomata, stroma, and spore shapes. Castlebury et al. (2002) reported that Diaporthaceae comprised Diaporthe and Mazzantia based on LSU DNA sequence data, removing other genera to different families in Diaporthales. Additional genera subsequently placed in the Diaporthaceae include Leucodiaporthe (Vasilyeva et al. 2007), Stenocarpella (Crous et al. 2006), Phaeocytostroma (Lamprecht et al. 2011), Ophiodiaporthe (Fu et al. 2013), and Pustulomyces (Dai et al. 2014). All the above genera were represented by a few species or are monotypic. Although they appeared to be morphologically divergent from Diaporthe, their phylogenetic relationships remain unclear.

About 991 names of Diaporthe and 979 of Phomopsis have been established to date (http://www.indexfungorum.org/Names/Names.asp). Among them, many old epithets lack molecular data, and few morphological characters can be used in species delimitation, making it difficult to merge these names to advance to the one name scenario (Rossman et al. 2014, 2015). In China, more than 50 plant pathogenic Phomopsis species have been published to date (Chi et al. 2007). In order to stabilize these species names in the genus Diaporthe, here we introduce 12 new combinations for Phomopsis species that have been subjected to DNA sequencing, and whose phylogenetic position has been resolved in Diaporthe in the present study.

The objectives of this study were: (1) to examine the phylogenetic relationships of Diaporthe with other closely related genera in Diaporthaceae; (2) to introduce new species in Diaporthe; and (3) to transfer Phomopsis species described from China to Diaporthe based on morphological and newly generated molecular data.

Material and Methods

Isolates

Strains were isolated from leaves of both symptomatic and healthy plant tissues from Yunnan, Zhejiang, and Jiangxi Provinces in China. A few other strains were obtained via the Ningbo Entry-Exit Inspection and Quarantine Bureau, which were isolated from imported plants from other countries. Single spore isolations were conducted from diseased leaves with visible fungal sporulation following the protocol of Zhang et al. (2013), and isolation from surface sterilized leaf tissues was conducted following the protocol of Gao et al. (2014). Fungal endophytes were isolated according to the method described by Liu et al. (2015). The Diaporthe strains were primarily identified from the other fungal species based on cultural characteristics on PDA, spore morphology, and ITS sequence data. Type specimens of new species were deposited in the Mycological Herbarium, Microbiology Institute, Chinese Academy of Sciences, Beijing, China (HMAS), with ex-type living cultures deposited in the China General Microbiological Culture Collection Center (CGMCC).

Morphological analysis

Cultures were incubated on PDA at 25 °C under ambient daylight and growth rates were measured daily for 7 d. To induce sporulation, isolates were inoculated on PNA (pine needle agar; Smith et al. 1996) containing double-autoclaved (30 min, 121°C, 1 bar) healthy pine needles and incubated at a room temperature of ca. 25 °C (Su et al. 2012). Cultures were examined periodically for the development of conidiomata and perithecia. Conidia were taken from pycnidia and mounted in sterilized water. The shape and size of microscopic structures were observed and noted using a light microscope (Nikon Eclipse 80i) with differential interference contrast (DIC). At least 10 conidiomata, 30 conidiophores, alpha and beta conidia were measured to calculate the mean size and standard deviation (SD).

DNA extraction, PCR amplification and sequencing

Isolates were grown on PDA and incubated at 25 °C for 7 d. Genomic DNA was extracted following the protocol of Cubero et al. (1999). The quality and quantity of DNA was estimated visually by staining with GelRed after 1% agarose gel electrophoresis. The primers ITS5 and ITS4 (White et al. 1990) were used to amplify the internal transcribed spacer region (ITS) of the nuclear ribosomal RNA gene operon, including the 3’ end of the 18S nrRNA, the first internal transcribed spacer region, the 5.8S nrRNA gene; the second internal transcribed spacer region and the 5’ end of the 28S nrRNAgene. The primers EF1-728F and EF1-986R (Carbone & Kohn 1999) were used to amplify part of the translation elongation factor 1-α gene (TEF1), and the primers CYLH3F (Crous et al. 2004) and H3-1b (Glass & Donaldson 1995) were used to amplify part of the histone H3 (HIS) gene. The primers T1 (O’Donnell & Cigelnik 1997) and Bt2b (Glass & Donaldson 1995) were used to amplify the beta-tubulin gene (TUB); the additional combination of Bt2a/Bt2b (Glass & Donaldson 1995) was used in case of amplification failure of the T1/Bt2b primer pair. The primer pair CAL228F/CAL737R (Carbone & Kohn 1999) and LR0R/LR5 primer pair (Rytas & Mark 1990) were used to amplify the calmodulin gene (CAL) and the LSU rDNA, respectively. Amplification reactions of 25 µL were composed of 10 × EasyTaq buffer (MgCl2+ included; Transgen, Beijing), 50 µM dNTPs, 0.2 µM of each forward and reverse primers (Transgen), 0.5 U EasyTaq DNA polymerase (Transgen) and 1–10 ng of genomic DNA. PCR parameters were as follows: 94 °C for 5 min, followed by 35 cycles of denaturation at 94 °C for 30 s, annealing at a suitable temperature for 30 s (52 °C for ITS and LSU, 56 °C for CAL, HIS, TEF1 and TUB), extension at 72 °C for 30 s and a final elongation step at 72 °C for 10 min. DNA sequencing was performed by Omegagenetics Company, Beijing.

Phylogenetic analyses

The DNA sequences generated with forward and reverse primers were used to obtain consensus sequences using MEGA v. 5.1 (Tamura et al. 2011), and subsequently aligned using MAFFT v. 6 (Katoh & Toh 2010); alignments were manually edited using MEGA v. 5.1 when necessary. Two datasets were employed in the phylogenetic analyses. LSU, ITS and TEF1 loci were selected to infer the generic relationships within Diaporthaceae (Table 1), with Valsa ambiens as outgroup. All available sequences of Diaporthe species were included in the dataset of combined ITS, HIS, TEF1, TUB, and CAL regions to infer the interspecific relationships within Diaporthe (Table 2) with Diaporthella corylina as outgroup. Maximum likelihood (ML) gene trees were estimated using the software RAxML v. 7.4.2 Black Box (Stamatakis 2006, Stamatakis et al. 2008). The RAxML software selected the GTR model of nucleotide substitution with the additional options of modelling rate heterogeneity (Γ) and proportion invariable sites (I). Bayesian analyses (critical value for the topological convergence diagnostic set to 0.01) were performed on the concatenated loci using MrBayes v. 3.2.2 (Ronquist et al. 2012) as described by (Crous et al. 2006) using nucleotide substitution models for each data partition selected by jModeltest (Darriba et al. 2012) and MrModeltest v. 2.3 (Nylander 2004). Bayesian analyses were launched with random starting trees for 10 000 000 generations, and Markov chains were sampled every 1000 generations. The first 25% resulting trees were discarded as burn-in. The remaining trees were summarized to calculate the posterior probabilities (PP) of each clade being monophyletic. Trees were visualized in FigTree v. 1.1.2 (http://tree.bio.ed.ac.uk/software/). New sequences generated in this study were deposited in NCBI’s GenBank nucleotide database (www.ncbi.nlm.nih.gov; Table 1).
Table 1

Sources of isolates and GenBank accession numbers used in the phylogenetic analyses of Diaporthaceae.

Species names*

Culture collection no.

Isolation sources

Country

GenBank Accession Numbers

References

ITS

LSU

TEF1

D. acaciigena

CBS 129521 (ex-type)

Acacia retinodes

Australia

KC343005

-

KC343731

Gomes et al. (2013)

D. ampelina

FAU 586

Vitis sp.

USA: New York

-

AF439635

-

-

D. angelicae

CBS 111592

Heracleum sphondylium

Austria

KC343027

-

KC343753

Gomes et al. (2013)

 

AR 3724

Heracleum sphondylium

Austria

KC343026

-

KC343752

Gomes et al. (2013)

D. apiculata

LC 3418 (ex-type)

Camellia sinensis

China

KP267896

KY011852

KP267970

This study

 

LC 3452

Camellia sinensis

China

KP267901

KY011853

KP267975

This study

D. arecae complex

LC 4155

Rhododendron sp.

China

KY011895

KY011879

KY011906

This study

 

LC 4159

Rhododendron sp.

China

KY011896

KY011880

KY011907

This study

 

LC 4164

Unknown host

China

KY011897

KY011881

KY011908

This study

D. biguttusis

LC1106 (ex-type)

Lithocarpus glaber

China

KF576282

KY011878

KF576257

This study

D. compacta

LC 3078

Camellia sinensis

China

KP267850

KY011839

KP267924

This study

 

LC 3083 (ex-type)

Camellia sinensis

China

KP267854

KY011840

KP267928

This study

 

LC 3084

Camellia sinensis

China

KP267855

KY011841

KP267929

This study

D. decedens

CBS 109772

Corylus avellana

Austria

KC343059

-

KC343785

Gomes et al. (2013)

D. detrusa

CBS 109770

Berberis vulgaris

Austria

KC343061

-

KC343787

Gomes et al. (2013)

D. discoidispora

LC 3503

Camellia sinensis

China

KY011887

KY011854

KY011898

This study

D. elaeagni-glabrae

LC 4802 (ex-type)

Elaeagnus glabra

China

KX986779

KY011885

KX999171

This study

 

LC 4806

Elaeagnus glabra

China

KX986780

KY011886

KX999172

This study

D. ellipicola

LC0810 (ex-type)

Lithocarpus glaber

China

KF576270

KY011873

KF576245

This study

D. eres

LC 3198

Camellia sinensis

China

KP267873

KY011845

KP267947

This study

 

LC 3205

Camellia sinensis

China

KP714499

KY011846

KP714511

This study

 

LC 3206

Camellia sinensis

China

KP714500

KY011847

KP714512

This study

 

CBS 109767

Acer campestre

Austria

KC343075

-

KC343801

Gomes et al. (2013)

D. fusicola

LC 1126

Lithocarpus glaber

China

KF576281

KY011836

KF576256

This study

 

LC 0778 (ex-type)

Lithocarpus glaber

China

KF576263

KY011877

KF576238

This study

D. hongkongensis

LC 0784

Lithocarpus glaber

China

KC153104

KY011876

KC153095

This study

 

LC 0812

Smilax china

China

KC153103

KY011875

KC153094

This study

D. incompleta

LC 6706

Camellia sinensis

China

KX986793

KY011859

KX999185

This study

 

LC 1127 (ex-type)

Lithocarpus glaber

China

KF576267

KY011837

KF576242

This study

D. mahothocarpi

LC 0732

Mahonia bealei

China

KC153097

KY011872

KC153088

This study

 

LC 0763 (ex-type)

Lithocarpus glaber

China

KC153096

KY011871

KC153087

This study

D. masirevicii

Diaporthe sp.

Camellia sinensis

China

KY011888

KY011861

KY011899

This study

D. neoarctii

CBS 109490

Ambrosia trifida

USA: New Jersey

KC343145

-

KC343871

Gomes et al. (2013)

D. oncostoma

CBS 109741

Robinia pseudoacacia

Russia

KC343161

-

KC343887

Gomes et al. (2013)

D. oraccinii

LC 3166 (ex-type)

Camellia sinensis

China

KP267863

KY011843

KP267937

This study

 

LC 3172

Camellia sinensis

China

KP267864

KY011844

KP267938

This study

 

LC 3296

Camellia sinensis

China

KP267884

KY011849

KP267958

This study

D. ovoicicola

LC 1128 (ex-type)

Lithocarpus glaber

China

KF576264

KY011838

KF576239

This study

D. penetriteum

LC 3215

Camellia sinensis

China

KP267879

KY011848

KP267953

This study

 

LC 3353 (ex-type)

Camellia sinensis

China

KP714505

KY011850

KP714517

This study

 

LC 3394

Camellia sinensis

China

KP267893

KY011851

KP267967

This study

D. perjuncta

CBS 109745

Ulmus glabra

Austria

KC343172

-

KC343898

Gomes et al. (2013)

D. pseudophoenicicola

LC 6150

Phoenix canariensis

China

KY011891

KY011865

KY011902

This study

 

LC 6151

Phoenix canariensis

China

KY011892

KY011866

KY011903

This study

D. pustulata

CBS 109742

Acer pseudoplatanus

Austria

KC343185

-

KC343911

Gomes et al. (2013)

 

CBS 109760

Acer pseudoplatanus

Austria

KC343186

-

KC343912

Gomes et al. (2013)

 

CBS 109784

Prunus padus

Austria

KC343187

-

KC343913

Gomes et al. (2013)

D. rudis

LC 6147

Dendrobenthamia japonica

USA

KY011890

KY011864

KY011901

This study

 

LC 6145

Ilex aquifolium

China

KY011889

KY011863

KY011900

This study

D. saccarata

CBS 116311

Protea repens, cankers

South Africa

KC343190

-

KC34391

Gomes et al. (2013)

D. sclerotioides

CBS 296.67

Cucumis sativus

Netherlands

KC343193

-

KC343919

Gomes et al. (2013)

D. tectonendophytica

LC 6623

Unknown host

China

KX986795

KY011857

KX999187

This study

D. tectonigena

LC 6512

Camellia sinensis

China

KX986782

KY011856

KX999174

This study

D. ternstroemiae

LC 0777 (ex-type)

Ternstroemia gymnanthera

China

KC153098

KY011874

KC153089

This study

D. ueckerae

LC 3564

Camellia sinensis

China

KP267912

KY011855

KP267986

This study

D. undulata

LC 6624

Unknown host

China

KX986798

KY011858

KX999190

This study

D. velutina

LC 4414

Lithocapus sp.

China

KX986788

KY011882

KX999180

This study

 

LC 4419

Neolitsea sp.

China

KX986789

KY011883

KX999181

This study

 

LC 4421 (ex-type)

Neolitsea sp.

China

KX986790

KY011884

KX999182

This study

D. xishuangbanica

LC 6707

Camellia sinensis

China

KX986783

KY011860

KX999175

This study

 

LC 6744

Camellia sinensis

China

KX986784

KY011862

KX999176

This study

D. yunnanensis

LC 6168

Coffea sp.

China

KX986796

KY011867

KX999188

This study

Diaporthe sp.

LC 3156

Camellia sinensis

China

KP267861

KY011842

KP267935

This study

 

LC 6170

Coffea sp.

China

KY011893

KY011869

KY011904

This study

 

LC 6171

Solanum melongena

China

KY011894

KY011870

KY011905

This study

 

LC 6232

Theobroma cacao

China

KX986797

KY011868

KX999189

This study

Mazzantia napelli

AR 3498

Aconitum vulparia

Austria

-

AF408368

EU222017

Castlebury et al. (2002)

Ophiodiaporthe cyatheae

BCRC 34961

Cyathea lepifera

Taiwan

JX570889

JX570891

KC465406

Fu et al. (2013)

Phaeocytostroma ambiguum

CPC 17071

Zea mays

South Africa

FR748036

-

FR748068

Lamprecht et al. (2011)

 

CPC 17072

Zea mays

South Africa

FR748037

FR748096

FR748069

Lamprecht et al. (2011)

Ph. plurivorum

CBS 113835

Helianthus annuus

Portugal

FR748046

FR748104

FR748078

Lamprecht et al. (2011)

Ph. sacchari

CBS 275.34

-

Japan

FR748047

FR748105

FR748079

Lamprecht et al. (2011)

Ph. megalosporum

CBS 284.65

Rice-field soil

India

FR748045

FR748103

FR748077

Lamprecht et al. (2011)

Pustulomyces bambusicola

MFLUCC 11-0436

on dead culm of bamboo

Thailand

-

KF806753

KF806755

Dai et al. (2014)

Stenocarpella macrospora

CBS 117560

Rain damaged Bt maize hybrid, 2003–04 season

South Africa

FR748048

DQ377934

-

Lamprecht et al. (2011)

S. maydis

CBS 117558

Traditional/landrace maize from 2003/04 season

South Africa

FR748051

DQ377936

FR748080

Lamprecht et al. (2011)

Valsa ambiens

CFCC 89894

Pyrus bretschneideri

China

KR045617

KR045699

KU710912

Fan et al. (2014)

*New species described in this paper are shown in bold.

Table 2

Sources of isolates and GenBank accession numbers used in the phylogenetic analyses of Diaporthe. Newly sequenced material is indicated in bold type.

Species names*

Culture collection no.

Isolation sources

Host family

GenBank Accession Numbers

References

ITS

TEF1

TUB

HIS

CAL

D. acaciigena

CBS 129521 (ex-type)

Acacia retinodes

Mimosaceae

KC343005

KC343731

KC343973

KC343489

KC343247

Gomes et al. (2013)

D. acerina

CBS 137.27

Acer saccharum

Aceraceae

KC343006

KC343732

KC343974

KC343490

KC343248

Gomes et al. (2013)

D. acutispora

CGMCC 3.18285 = LC 6161

Coffea sp., endophyte

Rubiaceae

KX986764

KX999155

KX999195

KX999235

KX999274

This study

 

LC 6142

Camellia sasanqua, endophyte

Theaceae

KX986762

KX999153

KX999193

KX999233

KX999272

This study

 

LC 6160

Camellia sasanqua, endophyte

Theaceae

KX986800

KX999192

KX999232

KX999271

KX999293

This study

D. alleghaniensis

CBS 495.72 (ex-type)

Betula alleghaniensis, branches

Betulaceae

KC343007

KC343733

KC343975

KC343491

KC343249

Gomes et al. (2013)

D. alnea

CBS 146.46 (ex-type)

Alnus sp.

Betulaceae

KC343008

KC343734

KC343976

KC343492

KC343250

Gomes et al. (2013)

 

CBS 159.47

Alnus sp.

Betulaceae

KC343009

KC343735

KC343977

KC343493

KC343251

Gomes et al. (2013)

D. ambigua

CBS 114015

Pyrus communis

Rosaceae

KC343010

KC343736

KC343978

KC343494

KC343252

Gomes et al. (2013)

 

CBS 117176

Aspalathus linearis, crown

Fabaceae

KC343011

KC343737

KC343979

KC343495

KC343253

Gomes et al. (2013)

D. ampelina

CBS 114016

Vitis vinifera

Vitaceae

AF230751

AY745056

JX275452

-

AY745026

Gomes et al. (2013)

 

CBS 111888

Vitis vinifera

Vitaceae

KC343016

KC343742

KC343984

KC343500

KC343258

Gomes et al. (2013)

D. amygdali

CBS 126679 (ex-type)

Prunus dulcis

Rosaceae

KC343022

KC343748

KC343990

KC343506

KC343264

Gomes et al. (2013)

 

CBS 111811

Vitis vinifera

Vitaceae

KC343019

KC343745

KC343987

KC343503

KC343261

Gomes et al. (2013)

D. anacardii

CBS 720.97 (ex-epitype)

Anacardium occidentale

Anacardiaceae

KC343024

KC343750

KC343992

KC343508

KC343266

Gomes et al. (2013)

D. angelicae

CBS 111592 (ex-epitype)

Heracleum sphondylium

Apiaceae

KC343027

KC343743

KC343995

KC343511

KC343269

Gomes et al. (2013)

 

CBS 123215

Foeniculum vulgare

Apiaceae

KC343028

KC353754

KC343996

KC343512

KC343270

Gomes et al. (2013)

D. apiculata

LC 4152

Camellia, leaf

Theaceae

KP267915

KP267989

KP293495

KP293562

-

Gao et al. (2016)

 

LC 3418, (ex-type)

Camellia sinensis, leaf, endophyte

Theaceae

KP267896

KP267970

KP293476

KP293550

-

Gao et al. (2016)

D. arctii

CBS 136.25

Arctium sp.

Arecaceae

KC343032

KC343758

KC344000

KC343516

KC343273

Gomes et al. (2013)

D. arecae

CBS 535.75

Citrus sp., fruit

Rutaceae

KC343033

KC343759

KC344001

KC343517

KC343275

Gomes et al. (2013)

 

CBS 161.64 (ex-isotype)

Areca catechu, fruit

Arecaceae

KC343032

KC343758

KC344000

KC343516

KC343274

Gomes et al. (2013)

D. arengae

CBS 114979 (ex-type)

Arenga engleri

Arecaceae

KC343034

KC343760

KC344002

KC343518

KC343276

Gomes et al. (2013)

D. asheiola

CBS 136967, CPC 16508, (ex-type)

Vaccinium ashei

Ericaceae

KJ160562

KJ160594

KJ160518

-

KJ160542

Lombard et al. (2014)

 

CBS 136968, CPC 16511

Vaccinium ashei

Ericaceae

KJ160563

KJ160595

KJ160519

-

KJ160543

Lombard et al. (2014)

D. aspalathi

CBS 117168

Aspalathus linearis

Fabaceae

KC343035

KC343761

KC344003

KC343519

KC343277

Gomes et al. (2013)

 

CBS 117169, (ex-type)

Aspalathus linearis

Fabaceae

KC343036

KC343762

KC344004

KC343520

KC343278

Gomes et al. (2013)

D. australafricana

CBS 111886

Vitis vinifera

Vitaceae

KC343038

KC343764

KC344006

KC343522

KC343280

Gomes et al. (2013)

 

CBS 113487

Vitis vinifera

Vitaceae

KC343039

KC343765

KC344007

KC343523

KC343281

Gomes et al. (2013)

D. baccae

CBS 136971

Vaccinium corymbosum

Ericaceae

KJ160564

KJ160596

-

-

-

Lombard et al. (2014)

 

CBS 136972 (ex-type)

Vaccinium corymbosum

Ericaceae

KJ160565

KJ160597

-

-

-

Lombard et al. (2014)

D. batatas

CBS 122.21

Ipomoea batatas

Convolvulaceae

KC343040

KC343766

KC344008

KC343524

KC343282

Gomes et al. (2013)

D. beckhausii

CBS 138.27

Viburnum sp.

Caprifoliaceae

KC343041

KC343767

KC344009

KC343525

KC343283

Gomes et al. (2013)

D. beilharziae

BRIP 54792 (ex-type)

Indigofera australis

Papilionaceae

JX862529

JX862535

KF170921

-

-

Thompson et al. (2015)

D. benedicti

CFCC 50062 (ex-type)

Juglans mandshurica

Juglandaceae

KP208847

KP208853

KP208855

KP208851

KP208849

Fan et al. (2015)

 

CFCC 50063

Juglans mandshurica

Juglandaceae

KP208848

KP208854

KP208856

KP208852

KP208850

Fan et al. (2015)

D. betulae

CFCC 50469 (ex-type)

Betula platyphylla

Betulaceae

KT732950

KT733016

KT733020

KT732999

KT732997

Du et al. (2016)

 

CFCC 50470

Betula platyphylla

Betulaceae

KT732951

KT733017

KT733021

KT733000

KT732998

Du et al. (2016)

D. betulicola

CFCC 51128 (ex-type)

Betula albosinensis

Betulaceae

KX024653

KX024655

KX024657

KX024661

KX024659

Du et al. (2016)

 

CFCC 51129

Betula albosinensis

Betulaceae

KX024654

KX024656

KX024658

KX024662

KX024660

Du et al. (2016)

D. bicincta

DP0659, CBS 121004

Juglans sp., dead wood

Juglandaceae

KC343134

KC343860

KC344102

KC343618

 

Udayanga et al. (2014a)

D. biconispora

ZJUD60, CGMCC3.17250

Citrus sinensis

Rutaceae

KJ490595

KJ490474

KJ490416

KJ490537

-

Huang et al. (2015)

 

ZJUD61, CGMCC 3.17251

Fortunella margarita

Rutaceae

KJ490596

KJ490475

KJ490417

KJ490538

-

Huang et al. (2015)

 

ZJUD62, CGMCC 3.17252

Citrus grandis

Rutaceae

KJ490597

KJ490476

KJ490418

KJ490539

-

Huang et al. (2015)

D. biguttulata

ZJUD47, CGMCC3.17248 (ex-type)

Citrus limon

Rutaceae

KJ490582

KJ490461

KJ490403

KJ490524

-

Huang et al. (2015)

 

ZJUD48, CGMCC3.17249

Citrus limon

Rutaceae

KJ490583

KJ490462

KJ490403

KJ490525

-

Huang et al. (2015)

D. biguttusis

CGMCC 3.17081 (ex-type)

Lithocarpus glabra

Fagaceae

KF576282

KF576257

KF576306

-

-

Gao et al. (2015)

D. brasiliensis

CBS 133183 (ex-type)

Aspidosperma tomentosus

Apocynaceae

KC343042

KC343768

KC344010

KC343526

KC343284

Gomes et al., 2013

 

LGMF 926

Aspidosperma tomentosus

Apocynaceae

KC343043

KC343769

KC344011

KC343527

KC343285

Gomes et al., 2013

D. canthii

CBS 132533 (ex-type)

Canthium inerme

Rubiaceae

JX069864

KC843120

KC843230

-

KC843174

Du et al. (2016)

D. carpini

CBS 114437

Carpinus betulus

Corylaceae

KC343044

KC343770

KC344012

KC343528

KC343286

Gomes et al. (2013)

D. caulivora

CBS 127268 (ex-neotype)

Glycine max

Fabaceae

KC343045

KC343771

KC344013

KC343529

KC343287

Gomes et al. (2013)

 

CBS 178.55

Glycine soja

Fabaceae

KC343046

KC343772

KC344014

KC343530

KC343288

Gomes et al. (2013)

D. celastrina

CBS 139.27

Celastrus scandens

Celastraceae

KC343047

KC343773

KC344015

KC343531

-

Gomes et al. (2013)

D. cf. heveae 1

CBS 852.97

Hevea brasiliensis

Euphorbiaceae

KC343116

KC343842

KC344084

KC343600

KC343358

Gomes et al. (2013)

D. cf. heveae 2

CBS 681.84

Hevea brasilliensis, leaf

Euphorbiaceae

KC343117

KC343843

KC344085

KC343601

KC343359

Gomes et al. (2013)

D. chamaeropis

CBS 454.81

Chamaerops humilis, dead part of leaf

Arecaceae

KC343048

KC343774

KC344016

KC343532

KC343290

Gomes et al. (2013)

 

CBS 753.70

Spartium junceum, dead branch

Fabaceae

KC343049

KC343775

KC344017

KC343533

KC343291

Gomes et al. (2013)

D. charlesworthii

BRIP 4884m (ex-type)

Rapistrum rugostrum

Brassicaceae

KJ197288

KJ197250

KJ197268

-

-

Thompson et al. (2015)

D. cinerascens

CBS 719.96

Ficus carica

Moraceae

KC343050

KC343776

KC344018

KC343534

KC343292

Gomes et al. (2013)

D. citri

CBS 230.52

Citrus sinensis

Rutaceae

KC343052

KC343778

KC344020

KC343536

KC343294

Gomes et al. (2013)

 

CBS 199.39

-

-

KC343051

KC343777

KC344019

KC343535

KC343293

Gomes et al. (2013)

 

AR 3405

Citrus sp.

Rutaceae

KC843311

KC843071

KC843187

KJ420881

-

Udayanga et al. (2014b)

D. citriasiana

ZJUD30 (ex-type)

Citrus unshiu, dead wood

Rutaceae

JQ954645

JQ954663

KC357459

-

KC357491

Huang et al. (2015)

 

ZJUD 33

Citrus paradise, stem-end rot fruit

Rutaceae

JQ954658

JQ972716

KC357460

-

KC357493

Huang et al. (2015)

D. citrichinensis

ZJUD 34

Citrus sp.

Rutaceae

JQ954648

JQ954666

-

-

KC357494

Huang et al. (2015)

 

ZJUD 35

Citrus unshiu, dead wood

Rutaceae

JQ954649

JQ954667

KC357461

-

KC357495

Huang et al. (2015)

 

ZJUD 36

Citrus unshiu, dead wood

Rutaceae

KC357556

KC357525

KC357462

-

KC357496

Huang et al. (2015)

D. compacta

LC3083 (ex-type)

Camellia sinensis, leaf, endophyte

Theaceae

KP267854

KP267928

KP293434

KP293508

-

Gao et al. (2016)

 

LC3084

Camellia sinensis, leaf, endophyte

Theaceae

KP267855

KP267929

KP293435

KP293509

-

Gao et al. (2016)

D. convolvuli

CBS 124654

Convolvulus arvensis

Convolvulaceae

KC343054

KC343780

KC344022

KC343538

KC343296

Huang et al. (2015)

D. crataegi

CBS 114435

Crataegus oxyacantha

Rosaceae

KC343055

KC343781

KC344023

KC343539

KC343297

Gomes et al. (2013)

D. crotalariae

CBS 162.33 (ex-type)

Crotalaria spectabilis

Fabaceae

KC343056

KC343782

KC344024

KC343540

KC343298

Gomes et al. (2013)

D. cuppatae

CBS 117499

Aspalathus linearis

Fabaceae

KC343057

KC343783

KC344025

KC343541

KC343299

Gomes et al. (2013)

D. cynaroidis

CBS 122676

Protea cynaroides

Proteaceae

KC343058

KC343784

KC344026

KC343542

KC343300

Gomes et al. (2013)

D. cytosporella

AR 5149

Citrus sinensis

Rutaceae

KC843309

KC843118

KC843222

-

KC843143

Udayanga et al. (2014b)

D. decedens

CBS 114281

Corylus avellana

Corylaceae

KC343060

KC343786

KC344028

KC343544

KC343302

Gomes et al. (2013)

 

CBS 109772

Corylus avellana

Corylaceae

KC343059

KC343785

KC344027

KC343543

KC343301

Gomes et al. (2013)

D. detrusa

CBS 109770

Berberis vulgaris

Berberidaceae

KC343061

KC343787

KC344029

KC343545

KC343303

Gomes et al. (2013)

 

CBS 114652

Berberis vulgaris

Berberidaceae

KC343062

KC343788

KC344030

KC343546

KC343304

Gomes et al. (2013)

D. discoidspora

ZJUD 87, CGMCC3.17254

Citrus sinensis

Rutaceae

KJ490622

KJ490501

KJ490443

KJ490564

-

Huang et al. (2015)

 

ZJUD 89, CGMCC 3.17255

Citrus unshiu

Rutaceae

KJ490624

KJ490503

KJ490445

KJ490566

-

Huang et al. (2015)

D. elaeagni

CBS 504.72

Elaeagnus sp., twig

Elaeagnaceae

KC343064

KC343790

KC344032

KC343548

KC343306

Gomes et al. (2013)

D. elaeagni-glabrae

CGMCC 3.18287 = LC 4802

Elaeagnus glabra, pathogen

Elaeagnaceae

KX986779

KX999171

KX999212

KX999251

KX999281

This study

 

LC 4806

Elaeagnus glabra, pathogen

Elaeagnaceae

KX986780

KX999172

KX999213

KX999252

KX999282

This study

D. ellipicola

CGMCC3.17084 (ex-type)

Lithocarpus glabra, diseased leaves

Fagaceae

KF576270

KF576245

KF576291

-

-

Gao et al. (2015)

D. endophytica

CBS 133811 (ex-type)

Schinus terebinthifolius

Anacardiaceae

KC343065

KC343791

KC344033

KC343549

KC343307

Gomes et al. (2013)

 

LGMF 911

Schinus terebinthifolius

Anacardiaceae

KC343066

KC343792

KC344034

KC343550

KC343308

Gomes et al. (2013)

D. eres

AR5193, CBS 13859 (ex-epitype)

Ulmus laevis

Ulmaceae

KJ210529

KJ210550

KJ420799

KJ420850

-

Udayanga et al. (2014a)

 

CBS 113470

Castanea sativa

Fagaceae

KC343146

KC343872

KC344114

KC343630

-

Udayanga et al. (2014a)

D. eugeniae

CBS 444.82

Eugenia aromatica, leaf

Mrytaceae

KC343098

KC343824

KC344066

KC343582

KC343340

Gomes et al. (2013)

D. fibrosa

CBS 109751

Rhamnus cathartica

Rhamnaceae

KC343099

KC343825

KC344067

KC343583

KC343341

Gomes et al. (2013)

 

CBS 113830

Rhamnus cathartica

Rhamnaceae

KC343100

KC343826

KC344068

KC343584

KC343342

Gomes et al. (2013)

D. foeniculina

CBS 116957

Pyrus pyrifolia

Rosaceae

KC343103

KC343829

KC344071

KC343587

KC343345

Gomes et al. (2013)

 

CBS 187.27 (ex-type of P. theicola)

Camellia sinensis, leaves and branches

Theaceae

KC343107

KC343833

KC344075

KC343591

KC343349

Gomes et al. (2013)

 

CBS 123208

Foeniculum vulgare

Apiaceae

KC343104

KC343830

KC344072

KC343588

KC343346

Gomes et al. (2013)

D. fraxini-angustifolia

BRIP 54781 (ex-epitype)

Fraxinus-angustifolia subsp. oxycapa

Oleaceae

JX862528

JX852534

KF170920

-

-

Tan et al. (2013)

D. ganjae

CBS 180.91 (ex-type)

Cannabis sativa, dead leaf

Cannabaceae

KC343112

KC343838

KC344080

KC343596

KC343354

Gomes et al. (2013)

D. gardeniae

CBS 288.56

Gardenia florida, stem

Rubiaceae

KC343113

KC343839

KC344081

KC343597

KC343355

Gomes et al. (2013)

D. goulteri

BRIP 55657a (ex-type)

Helianthus annuus

Asteraceae

KJ197289

KJ197252

KJ197270

-

-

Thompson et al. (2015)

D. gulyae

BRIP 54025 (ex-type)

Helianthus annuus

Asteraceae

JF431299

JN645803

KJ197271

-

-

Thompson et al. (2015)

D. helianthi

CBS 344.94

Helianthus annuus

Asteraceae

KC343114

KC343840

KC344082

KC343598

KC343356

Gomes et al. (2013)

 

CBS 592.81 (ex-type)

Helianthus annuus

Asteraceae

KC343115

KC343841

KC344083

KC343599

KC343357

Gomes et al. (2013)

D. helicis

AR 5211

Hedera helix

Araliaceae

KJ210538

KJ210559

KJ420828

KJ420875

KJ435043

Udayanga et al. (2014a)

D. hickoriae

CBS 145.26 (ex-epitype)

Carya glabra

Juglandaceae

KC343118

KC343844

KC344086

KC343602

KC343360

Gomes et al. (2013)

D. hongkongensis

CBS 115448 (ex-type)

Dichroa febrifuga, fruit

Hydrangeaceae

KC343119

KC343845

KC344087

KC343603

KC343361

Gomes et al. (2013)

D. hordei

CBS 481.92

Hordeum vulgare

Poaceae

KC343120

KC343846

KC344088

KC343604

KC343362

Gomes et al. (2013)

D. impulsa

CBS 114434

Sorbus aucuparia

Rosaceae

KC343121

KC343847

KC344089

KC343605

KC343363

Gomes et al. (2013)

 

CBS 141.27

Sorbus americana

Rosaceae

KC343122

KC343848

KC344090

KC343606

KC343364

Gomes et al. (2013)

D. incompleta

CGMCC 3.18288 = LC 6754

Camellia sinensis, pathogen

Theaceae

KX986794

KX999186

KX999226

KX999265

KX999289

This study

 

LC 6706

Camellia sinensis, pathogen

Theaceae

KX986793

KX999185

 

KX999264

KX999288

This study

D. inconspicua

CBS 133813(ex-type)

Maytenus ilicifolia, endophytic in petiole

Celastraceae

KC343123

KC343849

KC344091

KC343607

KC343365

Gomes et al. (2013)

D. infecunda

CBS 133812 (ex-type)

Schinus terebinthifolius

Anacardiaceae

KC343126

KC343852

KC344094

KC343610

KC343368

Gomes et al. (2013)

 

LGMF 908

Schinus terebinthifolius

Anacardiaceae

KC343127

KC343853

KC344095

KC343611

KC343369

Gomes et al. (2013)

D. kongii

BRIP 54031 (ex-type)

Helianthus annuus

Asteraceae

JF431301

JN645797

KJ197272

-

-

Thompson et al. (2011)

D. lichicola

BRIP 54900 (ex-type)

Litchi chinensis

Sapindaceae

JX862533

JX862539

KF170925

-

-

Tan et al. (2013)

D. longicicola

CGMCC 3.17089 (ex-type)

Lithocarpus glabra

Fagaceae

KF576267

KF576242

KF576291

-

-

Gao et al. (2015)

D. longicolla

FAU 599

Glycine max

Fabaceae

KJ590728

KJ590767

KJ610883

KJ659188

-

Udayanga et al. (2015)

D. longispora

CBS 194.36 (ex-type)

Ribes sp.

Grossulariaceae

KC343135

KC343861

KC344103

KC343619

KC343377

Gomes et al. (2013)

D. lusitanicae

CBS 123212 (ex-type)

Foeniculum vulgare

Apiaceae

KC343136

KC343862

KC344104

KC343620

-

Gomes et al. (2013)

 

CBS 123213

Foeniculum vulgare

Apiaceae

KC343137

KC343863

KC344105

KC343621

KC343379

Gomes et al. (2013)

D. macintoshii

BRIP 55064a

Rapistrum rugostrum

Brassicaceae

KJ197290

KJ197251

KJ197269

-

-

Thompson et al. (2015)

D. mahothocarpus

CGMCC 3.15181

Lithocarpus glabra

Fagaceae

KC153096

KC153087

-

-

-

Gao et al. (2014)

D. manihotia

CBS 505.76

Manihot utilissima, leaves

Euphorbiaceae

KC343138

KC343864

KC344106

KC343622

KC343380

Gomes et al. (2013)

D. maritima

NB 382-2E

Picea rubens needle

Pinaceae

KU552026

KU552024

KU574614

-

-

Tanney et al. (2016)

 

NB 463-3A

Picea rubens needle

Pinaceae

KU552027

KU552022

KU574616

-

-

Tanney et al. (2016)

 

NB365-71I (ex-type)

Picea rubens needle

Pinaceae

KU552025

KU552023

KU574615

-

-

Tanney et al. (2016)

D. masirevicii

BRIP 57330

Chrysanthemoides monilifera subsp. rotundata

Rosaceae

KJ197275

KJ197237

KJ197255

-

-

Huang et al. (2015)

 

BRIP 57892a (ex-type)

Helianthus annuus

Asteraceae

KJ197277

KJ197239

KJ197257

-

-

Huang et al. (2015)

D. mayteni

CBS 133185 (ex-type)

Maytenus ilicicolia

Celastraceae

KC343139

KC343865

KC344107

KC343623

KC343381

Gomes et al. (2013)

D. megalospora

CBS 143.27

Sambucus canadensis

Caprifoliaceae

KC343140

KC343866

KC344108

KC343624

KC343382

Gomes et al. (2013)

D. melonis

CBS 435.87

Glycine soja

Fabaceae

KC343141

KC343867

KC344109

KC343625

KC343383

Gomes et al. (2013)

 

CBS 507.78 (ex-isotype)

Cucumis melo

Cucurbitaceae

KC343142

KC343868

KC344110

KC343626

KC343384

Gomes et al. (2013)

D. middletonii

BRIP 57329

Chrysanthemoides monilifera subsp. rotundata

Rosaceae

KJ197285

KJ197247

KJ197265

-

-

Thompson et al. (2015)

 

BRIP 54884e (ex-type)

Rapistrum rugostrum

Brassicaceae

KJ197286

KJ197248

KJ197266

-

-

Thompson et al. (2015)

D. miriciae

BRIP 55662c

Glycine max

Fabaceae

KJ197283

KJ197245

KJ197263

-

-

Thompson et al. (2015)

 

BRIP 54736j (ex-type)

Helianthus annuus

Asteraceae

KJ197282

KJ197244

KJ197262

-

-

Thompson et al. (2015)

 

BRIP 56918a

Vigna radiata

Papilionaceae

KJ197284

KJ197246

KJ197264

-

-

Thompson et al. (2015)

D. multigutullata

ZJUD 98

Citrus grandis

Rosaceae

KJ490633

KJ490512

KJ490454

KJ490575

-

Huang et al. (2015)

D. musigena

CBS 129519; CPC 17026 (ex-type)

Musa sp., leaves

Musaceae

KC343143

KC343869

KC344111

KC343627

KC343385

Gomes et al. (2013)

D. neilliae

CBS 144.27

Spiraea sp.

Rosaceae

KC343144

KC343870

KC344112

KC343628

KC343386

Udayanga et al. (2014a)

D. neoarctii

CBS 109490 (ex-type)

Ambrosia trifida

Asteraceae

KC343145

KC343871

KC344113

KC343629

KC343387

Gomes et al. (2013)

D. neoraonikayaporum

MFLUCC 14-1136

Tectona grandis

Verbenaceae

KU712449

KU749369

KU743988

-

KU749356

Doilom et al. (2017)

MFLUCC 14-1137

Tectona grandis

Verbenaceae

KU712450

KU749370

KU743989

-

KU749357

Doilom et al. (2017)

 

MFLUCC 14-1133

Tectona grandis

Verbenaceae

KU712448

KU749368

KU743987

-

KU749355

Doilom et al. (2017)

D. nobilis

CBS 200.39

Laurus nobilis, stem

Lauraceae

KC343151

KC343877

KC344119

KC343635

KC343393

Gomes et al. (2013)

D. nomurai

CBS 157.29

Morus sp.

Moraceae

KC343154

KC343880

KC344122

KC343638

KC343396

Gomes et al. (2013)

D. nothofagi

BRIP 54801 (ex-type)

Nothofagus cunninghamii

Fagaceae

JX862530

JX862536

KF170922

-

-

Tan et al. (2013)

D. novem

CBS 127269

Glycine max

Fabaceae

KC343155

KC343881

KC344123

KC343639

KC343397

Gomes et al. (2013)

 

CBS 127270 (ex-type)

Glycine max

Fabaceae

KC343156

KC343882

KC344124

KC343640

KC343398

Gomes et al. (2013)

D. oncostoma

CBS 100454

Robinia pseudoacacia, leaf spot

Fabaceae

KC343160

KC343886

KC344128

KC343644

KC343402

Gomes et al. (2013)

 

CBS 109741

Robinia pseudoacacia

Fabaceae

KC343161

KC343887

KC344129

KC343645

KC343403

Gomes et al. (2013)

D. oraccinii

LC 3166 (ex-type)

Camellia sinensis, leaf, endophyte

Theaceae

KP267863

KP267937

KP293443

KP293517

-

Gao et al. (2016)

 

LC 3296

Camellia sinensis, leaf, endophyte

Theaceae

KP267884

KP267958

KP293464

KP293538

-

Gao et al. (2016)

D. ovalispora

ZJUD93, CGMCC3.17256

Citrus limon

Rosaceae

KJ490628

KJ490507

KJ490449

KJ490570

-

Huang et al. (2015)

D. oxe

CBS 133186 (ex-type)

Maytenus ilicifolia

Celastraceae

KC343164

KC343890

KC344132

KC343648

KC343406

Gomes et al. (2013)

 

CBS 133187

Maytenus ilicifolia

Celastraceae

KC343165

KC343891

KC344133

KC343649

KC343407

Gomes et al. (2013)

D. padi var. padi

CBS 114200

Prunus padus

Rosaceae

KC343169

KC343895

KC344137

KC343653

KC343411

Gomes et al. (2013)

 

CBS 114649

Alnus glutinosa

Betulaceae

KC343170

KC343896

KC344138

KC343654

KC343412

Gomes et al. (2013)

D. paranensis

CBS 133184 (ex-type)

Maytenus ilicifolia

Celastraceae

KC343171

KC343897

Kc344139

KC343655

KC343413

Gomes et al. (2013)

D. pascoei

BRIP 54847 (ex-type)

Persea americana

Lauraceae

JX862532

JX862538

KF170924

-

-

Tan et al. (2013)

D. penetriteum

LC 3353

Camellia sinensis, leaf

Theaceae

KP714505

KP714517

KP714529

KP714493

-

Gao et al. (2016)

 

LC 3394

Camellia sinensis, leaf

Theaceae

KP267893

KP267967

KP293473

KP293547

-

Gao et al. (2016)

D. perjuncta

CBS 109745 (ex-type)

Ulmus glabra

Ulmaceae

KC343172

KC343898

KC344140

KC343656

KC343414

Gomes et al. (2013)

D. perniciosa

CBS 124030

Malus pumila, bark

Rosaceae

KC343149

KC343875

KC344117

KC343633

KC343391

Gomes et al. (2013)

D. perseae

CBS 151.73

Perseae gratissima, young fruit

Lauraceae

KC343173

KC343899

KC344141

KC343657

KC343415

Gomes et al. (2013)

D. phaseolorum

AR 4203, CBS 139281

Phaseolus vulgaris

Fabaceae

KJ590738

KJ590739

KJ610893

KJ659220

-

Huang et al. (2015)

 

CBS 116019

Caperonia palustris

Euphorbiaceae

KC343175

KC343901

KC344143

KC343659

KC343417

Gomes et al. (2013)

 

CBS 116020

Aster exilis

Asteraceae

KC343176

KC343902

KC344144

KC343660

KC343418

Gomes et al. (2013)

D. podocarpi-macrophylli

CGMCC 3.18281 = LC 6155

Podocarpus macrophyllus, endophyte

Podocarpaceae

KX986774

KX999167

KX999207

KX999246

KX999278

This study

 

LC 6144

Podocarpus macrophyllus, endophyte

Podocarpaceae

KX986773

KX999166

KX999206

KX999245

-

This study

 

LC 6194

Podocarpus macrophyllus, endophyte

Podocarpaceae

KX986765

KX999156

KX999196

KX999236

KX999275

This study

 

LC 6197

Podocarpus macrophyllus, endophyte

Podocarpaceae

KX986777

KX999170

KX999210

KX999249

KX999279

This study

 

LC 6200

Podocarpus macrophyllus, endophyte

Podocarpaceae

KX986769

KX999161

KX999201

KX999240

KX999276

This study

 

LC 6229

Olea europaea, endophytes

Oleaceae

KX986771

KX999164

KX999204

KX999243

KX999277

This study

D. pseudomangiferae

CBS 101339 (ex-type)

Mangifera indica

Anacardiaceae

KC343181

KC343907

KC344149

KC343665

KC343423

Gomes et al. (2013)

 

CBS 388.89

Mangifera indica, peel of fruit

Anacardiaceae

KC343182

KC343908

KC344150

KC343666

KC343424

Gomes et al. (2013)

D. pseudophoenicicola

CBS 462.69 (ex-type)

Phoenix dactylifera, dead tops of green leaves

Anacardiaceae

KC343184

KC343910

KC344152

KC343668

KC343426

Gomes et al. (2013)

 

CBS 176.77

Mangifera indica, showing dieback

Anacardiaceae

KC343183

KC343909

KC344151

KC343667

KC343425

Gomes et al. (2013)

D. pterocarpi

MFLUCC 10-0571

Pterocarous indicus

Papilionaceae

JQ619899

JX275416

JX275460

-

JX197451

Udayanga et al. (2012)

 

MFLUCC 10-0575

Pterocarous indicus

Papilionaceae

JQ619901

JX275418

JX275462

-

JX197453

Udayanga et al. (2012)

D. pterocarpicola

MFLUCC 10-0580a (ex-type)

Piterocarpus indicus

Papilionaceae

JQ619887

JX275403

JX275441

-

JX197433

Udayanga et al. (2012)

 

MFLUCC 10-0580b

Piterocarpus indicus

Papilionaceae

JQ619888

JX275404

JX275442

-

JX197434

Udayanga et al. (2012)

D. pulla

CBS 338.89

Hedera helix

Araliaceae

KC343152

KC343878

KC344120

KC343636

-

Udayanga et al. (2014a)

D. pustulata

CBS 109742

Acer pseudoplatanus

Aceraceae

KC343185

KC343911

KC344153

KC343669

KC343427

Gomes et al. (2013)

 

CBS 109760

Acer pseudoplatanus

Aceraceae

KC343186

KC343912

KC344154

KC343670

KC343428

Gomes et al. (2013)

D. raonikayaporum

CBS 133182 (ex-type)

Spondias mombin

Anacardiaceae

KC343188

KC343914

KC344156

KC343672

KC343430

Gomes et al. (2013)

D. rhoina

CBS 146.27

Rhus toxicodendron

Anacardiaceae

KC343189

KC343915

KC344157

KC343673

KC343431

Gomes et al. (2013)

D. rudis

CBS 113201 (ex-type)

Vitis vinifera

Vitaceae

KC343234

KC343960

KC344202

KC343718

KC343476

Machingambi et al. (2015)

 

CBS 114011

Vitis Vinifera

Vitaceae

KC343235

KC343961

KC344203

KC343718

KC343477

Machingambi et al. (2015)

D. saccarata

CBS 116311 (ex-type)

Protea repens, cankers

Proteceae

KC343190

KC343916

KC344158

KC343674

KC343432

Gomes et al. (2013)

D. sackstonii

BRIP 54669b (ex-type)

Helianthus annuus

Asteraceae

KJ197287

KJ197249

KJ197267

-

-

Gomes et al. (2013)

D. salicicola

BRIP 54825 (ex-type)

Salix purpurea

Salicaceae

JX862531

JX862537

KF170923

-

-

Gomes et al. (2013)

D. schini

LGMF 910, CPC 20286

Schinus terebinthifolius, endophytic in leaf

Anacardiaceae

KC343192

KC343918

KC344160

KC343676

KC343434

Thompson et al. (2015)

 

CBS 133181 (ex-type)

Schinus terebinthifolius, endophytic in leaf

Anacardiaceae

KC343191

KC343917

KC344159

KC343675

KC343433

Tan et al. (2013)

D. sclerotioides

CBS 296.67 (ex-type)

Cucumis sativus

Cucurbitaceae

KC343193

KC343919

KC344161

KC343677

KC343435

Gomes et al. (2013)

 

CBS 710.76

Cucumis sativus

Cucurbitaceae

KC343194

KC343920

KC344162

KC343678

KC343436

Gomes et al. (2013)

D. scobina

CBS 251.38

Fraxinus Excelsior, living and dead twig

Oleaceae

KC343195

KC343921

KC344163

KC343679

KC343437

Gomes et al. (2013)

D. serafiniae

BRIP 55665a (ex-type)

Helianthus annuus

Asteraceae

KJ197274

KJ197236

KJ197254

-

-

Gomes et al. (2013)

 

BRIP 54136

Lupinus albus “Rosetta”

Fabaceae

KJ197273

KJ197235

KJ197253

-

-

Gomes et al. (2013)

D. siamensis

MFLUCC 10_0573a

Dasymaschalon sp.

Annonaceae

JQ619879

JX275393

JX275429

-

-

Thompson et al. (2015)

 

MFLUCC 10_0573b

Dasymaschalon sp.

Annonaceae

JQ619880

JX275395

JX275430

-

-

Thompson et al. (2015)

D. sojae

CBS 100.87

Glycine soja

Fabaceae

KC343196

KC343922

KC344164

KC343680

KC343438

Udayanga et al. (2012)

 

CBS 116017

Euphorbia nutans

Euphorbiaceae

KC343197

KC343923

KC344165

KC343681

KC343439

Udayanga et al. (2012)

 

FAU 635

Glycine max

Fabaceae

KJ590719

KJ590762

KJ610875

KJ659208

-

Gomes et al. (2013)

D. sterilis

CBS 136969 (ex-type)

Vaccinium corymbosum

Ericaceae

KJ160579

KJ160611

KJ160528

-

KJ160548

Gomes et al. (2013)

 

CBS 136970

Vaccinium corymbosum

Ericaceae

KJ160580

KJ160612

KJ160529

-

KJ160549

Huang et al. (2015)

D. stewartii

CBS 193.36

-

-

FJ889448

GQ250324

-

-

-

Lombard et al. (2014)

D. stictica

CBS 370.54

Buxus sampervirens, dead twig

Buxaceae

KC343212

KC343938

KC344180

KC343696

KC343454

Lombard et al. (2014)

D. subclavata

ZJUD83, CGMCC 3.17253

Citrus grandis cv. Shatianyou

Rosaceae

KJ490618

KJ490497

KJ490439

KJ490560

-

Udayanga et al. (2011)

 

ZJUD95, CGMCC 3.17257

Citrus unshiu

Rosaceae

KJ490630

KJ490509

KJ490451

KJ490572

-

Gomes et al. (2013)

D. subordinaria

CBS 101711

Plantago lanceolata

Plantaginaceae

KC343213

KC343939

KC344181

KC343697

KC343455

Huang et al. (2015)

 

CBS 464.90

Plantago lanceolata

Plantaginaceae

Kc343214

KC343940

KC344182

KC343698

KC343456

Huang et al. (2015)

D. tecomae

CBS 100547

Tabebuia sp.

Bignoniaceae

KC343215

KC343941

KC344183

KC343699

KC343457

Gomes et al. (2013)

D. tectonae

MFLUCC 12-0777

Tectona grandis

Verbenaceae

KU712430

KU749359

KU743977

-

KU749345

Gomes et al. (2013)

 

MFLUCC 14-1138

Tectona grandis

Verbenaceae

KU712437

KU749365

KU743984

-

KU749352

Gomes et al. (2013)

D. tectonendo-phytica

MFLUCC 13-0471

Tectona grandis

Verbenaceae

KU712439

KU749367

KU743986

-

KU749354

Doilom et al. (2017)

D. tectonigena

MFLUCC 12-0767

Tectona grandis

Verbenaceae

KU712429

KU749371

KU743976

-

KU749358

Doilom et al. (2017)

D. terebinthifolii

CBS 133180

Schinus terebinthifolius

Anacardiaceae

KC343216

KC343942

KC344184

KC343700

KC343458

Doilom et al. (2017)

 

LGMF 907

Schinus terebinthifolius

Anacardiaceae

KC343217

KC343943

KC344185

KC343701

KC343459

Doilom et al. (2017)

D. thunbergii

MFLUCC 10_0756a

Thunbergia laurifolia

Acanthaceae

JQ619893

JX275409

JX275449

-

JX197440

Doilom et al. (2017)

 

MFLUCC 10_0756b

Thunbergia laurifolia

Acanthaceae

JQ619894

JX275410

JX275450

-

JX197441

Doilom et al. (2017)

D. toxica

CBS 534.93 (ex-type)

Lupinus angustifolius, stem

Fabaceae

KC343220

KC343946

KC344188

KC343704

KC343462

Udayanga et al. (2012)

 

CBS 535.93

Lupinus sp.

Fabaceae

KC343221

KC343947

KC344189

KC343705

KC343463

Udayanga et al. (2012)

D. tulliensis

BRIP 62248a

Theobroma cacao

Sterculiaceae

KR936130

KR936133

KR936132

-

-

Gomes et al. (2013)

D. ueckerae

FAU 656

Cucumis melo

Cucurbitaceae

KJ590726

KJ590747

KJ610881

KJ659215

-

Gomes et al. (2013)

 

FAU 658

Cucumis melo

Cucurbitaceae

KJ590725

KJ590746

KJ610880

KJ659214

-

Crous et al. (2015)

D. undulata

CGMCC 3.18293 = LC 6624

Unknown host, pathogen

-

KX986798

KX999190

KX999230

KX999269

 

Huang et al. (2015)

 

LC 8110

Unknown host, pathogen

-

KY491545

KY491555

KY491565

-

-

Huang et al. (2015)

 

LC 8111

Unknown host, pathogen

-

KY491546

KY491556

KY491566

-

-

This study

D. unshiuensis

ZJUD51,CGMCC3.17568

Fortunella margarita

Rutaceae

KJ490586

KJ490465

KJ490407

KJ490528

-

This study

 

ZJUD52, CGMCC3.17569

Citrus unshiu

Rosaceae

KJ490587

KJ490466

KJ490408

KJ490529

-

This study

D. vaccinii

CBS 160.32 (ex-type)

Oxycoccus macrocarpos

Ericaceae

KC343228

KC343954

KC344196

KC343712

KC343470

Huang et al. (2015)

 

CBS 118571

Vaccinium corymbosum

Ericaceae

KC343223

KC343949

KC344191

KC343707

KC343465

Huang et al. (2015)

D. vawdreyi

BRIP 57887a

Psidium guajava

Sterculiaceae

KR936126

KR936129

KR936128

-

-

Gomes et al. (2013)

D. velutina

CGMCC 3.18286 = LC 4421

Neolitsea sp., pathogen

Lauraceae

KX986790

KX999182

KX999223

KX999261

 

Gomes et al. (2013)

 

LC 4419

Neolitsea sp., pathogen

Lauraceae

KX986789

KX999181

KX999222

KX999260

KX999286

Crous et al. (2015)

 

LC 4641

Callerya cinerea, pathogen

Fabaceae

KX986792

KX999184

KX999225

KX999263

KX999287

This study

 

LC 4788

Unknown host, pathogen

-

KX986785

KX999177

KX999218

KX999256

KX999285

This study

 

LC 6708

Camellia sinensis, pathogen

Theaceae

KX986787

KX999179

KX999220

KX999258

 

This study

D. vexans

CBS 127.14

Solanum melongena

Solanaceae

KC343229

KC343955

KC344197

KC343713

KC343471

This study

D. virgilia

CMW 40755 (ex-type)

Virgilia oroboides

Unknown

KP247573

-

KP247582

-

-

This study

 

CMW 40748

Virgilia oroboides

Unknown

KP247566

-

KP247575

-

-

Gomes et al. (2013)

D. woodii

CBS 558.93

Lupinus sp.

Fabaceae

KC343244

KC343970

KC344212

KC343728

KC343486

Gomes et al. (2013)

D. woolworthii

CBS 148.27

Ulmus americana

Ulmaceae

KC343245

KC343971

KC344213

KC343729

KC343487

Gomes et al. (2013)

D. xishuangbanica

CGMCC 3.18282= LC 6707

Camellia sinensis, pathogen

Theaceae

KX986783

KX999175

KX999216

KX999255

-

This study

 

LC 6744

Camellia sinensis, pathogen

Theaceae

KX986784

KX999176

KX999217

-

-

This study

D. yunnanensis

CGMCC 3.18289 = LC6168

Coffea sp., endophytes

Rubiaceae

KX986796

KX999188

KX999228

KX999267

KX999290

This study

 

LC 8106

Coffea sp., endophytes

Rubiaceae

KY491541

KY491551

KY491561

-

KY491571

This study

 

LC 8107

Coffea sp., endophytes

Rubiaceae

KY491542

KY491552

KY491562

-

KY491572

This study

Diaporthe sp.

LC 6496

Camellia sinensis, endophytes

Theaceae

KX986781

KX999173

KX999214

KX999253

KX999283

This study

 

LC 6512

Camellia sinensis, endophyte

Theaceae

KX986782

KX999174

KX999215

KX999254

KX999284

This study

 

LC 6232

Theobroma cacao, endophyte

Sterculiaceae

KX986797

KX999189

KX999229

KX999268

KX999291

This study

 

LC 8108

Theobroma cacao, endophyte

Sterculiaceae

KY491543

KY491553

KY491563

-

KY491573

This study

 

LC 8109

Theobroma cacao, endophyte

Sterculiaceae

KY491544

KY491554

KY491564

-

KY491574

This study

 

LC 6623

Unknown host, pathogen

-

KX986795

KX999187

KX999227

KX999266

-

This study

 

LC 8114

Unknown host, pathogen

-

KY491549

KY491559

KY491569

-

-

This study

 

LC 8115

Unknown host, pathogen

-

KY491550

KY491560

KY491570

-

-

This study

 

LGMF 947

Glycine max, seed

Fabaceae

KC343203

KC343929

KC344171

KC343687

KC343445

Gomes et al. (2013)

 

CBS 119639

Man, abscess

-

KC343202

KC343928

KC344170

KC343687

KC343444

Gomes et al. (2013)

Diaporthe sp. 1

CGMCC 3.18292 = LC 0771

Alnus sp., pathogen

Betulaceae

KX986799

KX999191

KX999231

KX999270

KX999292

This study

Diaporthe sp. 2

CGMCC 3.18291 = LC 6140

Acer sp., endophyte

Aceraceae

KX986799

KX999191

KX999231

KX999270

KX999292

This study

 

LC8112

Acer sp., endophyte

Aceraceae

KY491547

KY491557

KY491567

-

KY491575

This study

 

LC8113

Acer sp., endophyte

Aceraceae

KY491548

KY491558

KY491568

-

KY491576

This study

Diaporthella corylina

CBS 121124

Corylus sp., dying stems

Corylaceae

KC343004

KC343730

KC343972

KC343488

KC343246

Gomes et al. (2013)

P. conorum

CBS 587.79

Penus pentaphylla

Pinaceae

KC343153

KC343879

KC344121

KC343637

KC343395

Gomes et al. (2013)

P. emicis

BRIP 45089a (ex-type)

Emex australis

Polygonaceae

JF957784

JX275414

JX275458

-

JX197449

Udayanga et al. (2012)

P. fukushii

CBS 116953

Pyrus pyrifolia

Roseceae

KC343147

KC343873

KC344115

KC343631

KC343389

Gomes et al. (2013)

 

BRIP 45089b

Emex australis

Polygonaceae

JQ619898

JX275415

JX275459

-

JX197450

Udayanga et al. (2012)

-: not provided in literatures.

Results

Collection of Diaporthe strains

Twenty-one Diaporthe strains including presumed plant pathogens and endophytes were isolated from 11 different host plant species (Table 2) collected from three provinces (Jiangxi, Yunnan, Zhejiang) in the northern part of China. In addition, 28 strains were isolated from the plant samples inspected by Jiangsu Entry-Exit Inspection and Quarantine Bureau.

The paraphyly of Diaporthe

Phylogenetic analysis was conducted with 224 sequences derived from 76 ingroup taxa from Diaporthaceae with Valsa ambiens as the outgroup (Table 1). The combined alignment comprised 1 817 characters including gaps (795 for LSU, 558 for ITS, 464 for TEF1). Based on the results of the Mrmodeltest, the following priors were set in MrBayes for the different data partitions: GTR+G models with gamma-distributed rates were implemented for LSU and ITS, HKY+I+G model with invgamma-distributed rates were implemented for TEF1. The Bayesian analysis lasted 7 × 108 generations and the consensus tress and posterior probabilities were calculated from the trees left after discarding the first 25% generations for burn-in (Fig. 1).
Fig. 1
Fig. 1

Phylogenetic tree of the family Diaporthaceae from a maximum likelihood analysis based on the combined multi-locus dataset (ITS, LSU, TEF1). The ML bootstrap values ≥ 70%, bayesian probabilities BPP ≥ 0.90 are marked above the branches. The tree is rooted with Valsa ambiens.

The generic relationships of Mazzantia, Ophiodiaporthe, Phaeocytostroma, Pustulomyces, and Stenocarpella with Diaporthe from this analysis are shown in Fig. 1. The topology and branching order of the phylogenetic trees inferred from ML and Bayesian methods were essentially similar. Five genera from Diaporthaceae did not form discrete clades from Diaporthe species but are scattered in the latter, although the family remains monophyletic. The paraphyletic nature of Diaporthe, however, is demonstrated (Fig. 1). Ophiodia-porthe formed a well resolved and distinct clade represented by strain YMJ 1364, and clustered together with the ex-type culture of D. sclerotioides (CBS 296.67) (BPP 0.99, MLBS: 90). Stenocarpella, represented by S. maydis and S. mac-rospora, was well supported (BPP 1, MLBS = 96) and closely related to several species of Phaeocytostroma. Mazzantia, however, was poorly supported for its phylogenetic position in Diaporthaceae (Fig. 1).

Phylogenetic analyses of the combined datasets of Diaporthe species

In total, 1089 sequences derived from 273 ingroup taxa were combined and Diaporthella corylina was used as outgroup. A total of 2783 characters including gaps (568 for CAL, 554 for HIS, 523 for ITS, 636 for TEF1 and 456 for TUB) were included in the multi-locus dataset, comprising sequences generated from this study and others downloaded from GenBank (Table 2). For the Bayesian inference, GTR+I+G model was selected for CAL, HIS and ITS, HKY+I+G for TEF1 and TUB through the analysis of Mrmodeltest. The maximum likelihood tree conducted by the GTR model confirmed the tree topology and posterior probabilities of the Bayesian consensus tree.

The topology and branching order for the phylogenetic trees inferred from ML and Bayesian methods were essentially similar (Fig. 2). Based on the multi-locus phylogeny and morphology, 49 strains were assigned to 13 species, including eight taxa which we describe here as new (Fig. 2).
Fig. 2
Fig. 2

Phylogenetic tree of the genus Diaporthe from a maximum likelihood analysis based on the combined multi-locus dataset (CAL, HIS, ITS, TEF1, TUB). The ML bootstrap values ≥ 70%, bayesian probabilities BPP ≥ 0.90 are marked above the branches. The tree is rooted with Diaporthella corylina. The novel species are highlighted.

Taxonomy

Diaporthe acutispora Y.H. Gao & L. Cai, sp. nov.

MycoBank MB820679

(Fig. 3)
Fig. 3
Fig. 3

Diaporthe acutispora (CGMCC 3.18285). A–B. 30-d-old culture on PNA medium. C. Conidiomata. D–E. Conidiophores. F–G. Alpha conidia. Bars: C = 100 µm; D–G = 10 µm.

Etymology: Named after the acute spores.

Diagnosis: Diaporthe acutispora is phylogenetically distinct and morphologically differs from species reported from the host genera Coffea and Camellia in the larger conidiophores and alpha conidia (Table 3).
Table 3

Synoptic characters of Diaporthe spp. referred to in this study.

Host genera

Species

Conidiomata (µm)

Conidiophores (µm)

Alpha conidia (µm)

Beta conidia (µm)

References

Coffea

P. coffeae

200–;250

12–;16 × 2

8–;9 × 2.5

-

Uecker (1988)

Camellia

D. acutispora

99–;173

10–;34.5 × 2–;3

6.9–;10.4×2.1–;3.1

-

This study

 

D. amygdali

160–;220 × 120–;300

7.4–;36.3 × 1.5–;3.2

(4.18−)6.27–;6.32(−9.64) × (1.63−)2.36− 2.38(−3.31)

-

Diogo et al. (2010)

 

D. apiculata

74–;195 (−416)

9.0–;12.5 × 1.5–;2.5

6.5–;10 × 2–;3

(20.0−)25.0–;39.0 × 1.0–;1.5

Gao et al. (2016)

 

D. compacta

237–;350

6.0–;12.5 × 1.5–;2.5

6–;7.5 × 2–;3

20.0–;24.5 × 1.0–;1.5

Gao et al. (2016)

 

D. discoidispora

200 ×118

8.9–;23.4 × 1.3–;2.7

5.6–;8 × 2.1–;3.2

21.2–;38.7 × 0.9–;1.6

Huang et al. (2015)

 

D. eres

200–;250

10–;15× 2–;3

(6−)6.5–;8.5(−9) × 3–;4

(18−)22–;28(29) × 1–;1.5

Udayanga et al. (2014b)

 

D. foeniculacea

560 × 350

10–;13 × 1.5–;3

(5.4−)6.8–;7(−9) × (2−)2.3–;2.4(−3.1)

(16.8−)19.6–;21(−24.2) × (1.1−)1.3− 1.4(−1.7)

Phillips (2003)

 

D. foeniculina

400–;700

9–;15(−18) × 1−2

(7.5−)8.5–;9(−9.2) × (2−)2.3–;2.5(−2.7)

(20−)22–;28(−29) × (1.1−)1.4–;1.6(−2)

Udayanga et al. (2014c)

 

D. hongkongensis

to 200

5–;12 × 2–;4

(5−)6–;7(−8) × (2−)2.5(−3)

18–;22 × 1.5–;2

Gomes et al. (2013)

 

D. oraccinii

400

10.5–;22.5 × 1–;2

5.5–;7.5 × 0.5–;2

24.5–;31.0 × 1.0–;1.5

Gao et al. (2016)

 

D. penetriteum

176–;486

13–;21.5 (−27) × 1–;2

4.5–;5.5 × 1.5–;2.5

16.5–;27.5 × 1.0–;2.0

Gao et al. (2016)

 

D. ueckerae

150–;200

(9−)12–;28(−30) × 1.5–;2.5

(6−)6.4–;8.2(−8.6) × (2−)2.3–;3

-

Udayanga et al. (2014a)

 

D. xishuangbanica

180–;310

13–;34.5 × 1.5–;3

7–;9.5 × 2.5–;3.5

-

This study

 

D. yunnanensis

195–;880

-

3–;6.5 × 1–;2.5

13.5–;33.5 ×1–;1.5

This study

 

P. acaciicola

-

-

7–;9 × 3–;3.5

-

Diedicke (1911)

 

P. theae

40 × 25

-

6–;8 × 1.5–;2

18–;24×0.75

Petch (1925)

Elaeagnus

P. arnoldiae

900 × 500

6–;12 × 1–;2

5.5–;11 × 1.5–;2

15–;20

Uecker (1988)

 

P. elaeagni

500–;750

20–;25 ×1–;1.5

6–;10 ×2–;3

-

Uecker (1988)

 

P. elaeagnicola

175–;413 × 83–;185

10.0–;22.5 × 1.5–;2.7

6.0–;7.4 × 1.7–;2.2

19–;43 × 0.7–;1.2

Chang et al. (2005)

 

D. elaeagni-glabrae

330–;1170

16–;28 × 1.5–;2.5

6–;13 × 1.5–;3

7.5–;22.5 × 1–;2

This study

 

D. incompleta

207–;650

8–;22 × 1–;2.5

-

19–;44× 0.5–;1.5

This study

Neolitsea

D. velutina

69–;128

10–;23 × 1–;2.5

5.5–;10 × 2–;2.5

11–;27.5 × 0.5–;1.5

This study

AR, DP, FAU: Isolates in culture collection of Systematic Mycology and Microbiology Laboratory, USDA-ARS, Beltsville, Maryland, USA; BCRC: Bioresource Collection and Research Center, Taiwan; BRIP: Australian plant pathogen culture collection, Queensland, Australia; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, The Netherlands; CFCC: China Forestry Culture Collection Center, China. CGMCC: China General Microbiological Culture Collection; CMW: culture collection (CMW) of the Forestry and Agricultural Biotechnology Institute; CPC: working collection of Pedro Crous maintained at the Westerdijk Institute; LGMF: Culture collection of Laboratory of Genetics of Microorganisms, Federal University of Parana, Curitiba, Brazil; LC: Working collection of Lei Cai, housed at Institute of Microbiology, CAS, China; MFLUCC: Mae Fah Luang University Culture Collection; ZJUD: Zhe Jiang University, China.

Type: China: Yunnan Province: Aini Farm, on healthy leaves of Coffea sp., 20 Sep. 2014, W.J. Duan (HMAS 247086 — holotype, dried culture; CGMCC 3.18285 = LC 6161 -ex-type culture).

Description: On PNA: Conidiomata pycnidial, globose, brownish, embedded in tissue, erumpent at maturity, 99–173 µm diam, often with a yellowish conidial cirrus exuding from the ostioles. Conidiophores 10–34.5 × 2–3 µm, cylindrical, hyaline, septate, branched, straight or slightly curved, tapering towards the apex. Alpha conidia abundant in culture, 7–10.5 × 2–3 µm (x̅ = 8.4 ± 0.7 × 2.6 ± 0.2, n = 30), aseptate, hyaline, ellipsoidal to fusoid, multi-guttulate. Beta conidia not observed.

Culture characters: Cultures incubated on PDA at 25 °C in darkness, growth rate 7.5 mm diam/d. Colony entirely white at surface, reverse with pale brown pigmentation, white, fluffy aerial mycelium.

Additional material examined: China: Yunnan Province: Xishuangbanna, on healthy leaves of Camellia sasanqua, 20 Sep. 2014, W.J. Duan, culture LC 6142; ibid. culture LC 6160.

Diaporthe elaeagni-glabrae Y.H. Gao & L. Cai, sp. nov.

MycoBank MB820680

(Fig. 4)
Fig. 4
Fig. 4

Diaporthe elaeagni-glabrae (CGMCC 3.18287). A–;B. 14-d-old culture on PDA; C. Conidiomata; D–;H. Conidiophores; I. Alpha conidia; J. Beta conidia. Bars: C = 100 µm; D–;J = 10 µm.

Etymology: Named after the host species Elaeagnus glabra.

Diagnosis: Diaporthe elaeagni-glabrae can be distinguished from the closely related species D. elaeagni (96% in ITS, 93% in TEF1, 94% in TUB, 96% in HIS, and 94% in CAL) and D. stictica (96% in ITS, 95% in TEF, 97% in TUB, 96% in HIS, and 96% in CAL) (Fig. 2). Diaporthe elaeagni-glabrae differs from other species recorded from Elaeagnus in the significantly longer alpha conidia (Table 3).

Type: China: Jiangxi Province: on diseased leaves of Elaeagnus glabra, 5 Sep. 2013, Y.H. Gao (HMAS 247089 — holotype, dried culture; CGMCC 3.18287 = LC 4802 — ex-type culture).

Description: On PDA: Conidiomata globose, to 330–;1170 µm, erumpent, with slightly elongated black necks, yellowish or dirty white, spiral conidial cirri extruding from ostioles. Conidiophores 16–;28 × 1.5–;2.5 µm, cylindrical, phialidic, septate, branched, sometimes inflated. Alpha conidia 6–;13 × 1.5–;3 µm ( = 8.3 ± 1.4 × 2.2 ± 0.3, n = 30), hyaline, fusiform or oval, usually biguttulate. Beta conidia 7.5–;22.5 × 1–;2 µm ( = 15.1 ± 3.5 × 1.2 ± 0.2, n = 40), hyaline, filiform, smooth, curved, base truncate.

Culture characters: Cultures incubated on PDA at 25 °C in darkness, growth rate 7 mm diam/d. Colony pale yellowish, greenish to brownish at the centre, reverse pale yellowish and brownish at the centre with age. Aerial mycelium white, sparse, fluffy, with irregular margin and visible conidiomata at maturity.

Additional material examined: China: Jiangxi Province: on diseased leaves of Elaeagnus glabra, 5 Sep. 2013, Y.H. Gao, culture LC 4806.

Diaporthe helianthi Munt.-Cvetk. et al., Nova Hedwigia 34: 433 (1981).

(Fig. 5)
Fig. 5
Fig. 5

Diaporthe helianthi (LC 6185). A–;B. 7-d-old culture on PDA; C. Conidiomata; D–;F. Conidiophores; G–;H. Beta conidia. Bars: C = 100 µm; D–;H = 10 µm.

Description: Sexual morph not produced. Conidiomata pycnidial globose to subglobose, dark brownish to black, erumpent or immersed in medium, translucent conidia exuded from the ostioles, 110–;380 µm diam. Conidiophores cylindrical, straight or sinuous, apical or base sometimes swelling, 11.5–;23.5 × 1.8–;3.5 µm ( = 16 ± 3 × 2.5 ± 0.5, n = 30). Beta conidia filiform, hamate or slightly curved, base truncate, tapering towards one apex, 11.5–;32 × 0.5–;2 µm (= 20±7.5×1± 0.4, n = 20). Alpha conidia not observed.

Culture characters: Cultures on PDA at 25 °C in dark, with 12/12 h alternation between daylight and darkness pure white (surface) and pale yellow to cream (reverse). Colony pellicular, forming less pigmented sectors, with concentric rings of gummy mycelium. Growth rate was 10.5 mm diam/d.

Material examined: Ukraine: from seeds of Helianthus annuus, 30 Oct. 2015, W.J. Duan culture LC 6173. — Japan: Lagerstroemia indica, 30 Oct. 2015, W.J. Duan, culture LC 6185.

Notes: Diaporthe helianthi, responsible for stem canker and grey spot disease of sunflower (Helianthus annuus) (Muntanola-Cvetkovic et al. 1981), has been listed in the Chinese quarantine directory. There is increasing evidence that this serious sunflower pathogen is being quickly and globally disseminated with international trade. The cases reported here were intercepted from imported sunflower seeds from Ukraine and Lagerstroemia indica from Japan.

Diaporthe incompleta Y.H. Gao & L. Cai, sp. nov.

MycoBank MB820681

(Fig. 6)
Fig. 6
Fig. 6

Diaporthe incompleta (CGMCC 3.18288). A. Leaves of host plant; B–;C. 7-d-old culture; D. Conidiomata; E–;F. Conidiophores; G. Beta conidia. Bars: D = 100 µm; E–;G = 10 µm.

Etymology: Named after the absence of alpha conidia.

Diagnosis: Diaporthe incompleta is phylogenetically distinct and differs morphologically from other species recorded from Elaeagnus and Camellia in the much longer beta conidia (Table 3).

Type: China: Yunnan Province: Xishuangbanna, on diseased of Elaeagnus glabra, 19 Apr. 2015, F Liu (HMAS 247088 — holotype, dried culture; CGMCC 3.18288 = LC 6754 — ex-type culture).

Description: Conidiomata pycnidial, subglobose to globose, brownish to black, 207–;650 µm diam, cream to pale luteous conidial droplets exuding from the central ostioles. Conidiophores 8–;22 × 1–;2.5 µm, cylindrical, hyaline, septate, unbranched, smooth, slightly curved, tapering towards apex. Alpha conidia not observed. Beta conidia 19–;44 × 0.5–;1.5 µm (x̅ = 30.5 ° 8.7 × 1.1 ± 0.4, n = 30), smooth, hyaline, filiform, base subtruncate, straight or curved.

Culture characters: Cultures incubated on PDA at 25 °C in darkness, growth rate 16.5 mm diam/d. Colony entirely white, flat, reverse pale yellowish, becoming brownish zoned at the centre with age. Aerial mycelium white, cottony, margin lobate, conidiomata visible at maturity.

Additional material examined: China: Yunnan Province: Xishuangbanna, on diseased leaves of Camellia sinensis, 19 Apr. 2015, F. Liu, culture LC 6706.

Diaporthe podocarpi-macrophylli Y.H. Gao & L. Cai, sp. nov.

MycoBank MB820682

(Fig. 7)
Fig. 7
Fig. 7

Diaporthe podocarpi-macrophylli (CGMCC 3.18281). A–;B. 30-d-old culture on PDA; C. Conidiomata; D–;F. Conidiophores; G-I. Alpha and beta conidia. Bars: C = 100 µm; D–;I = 10 µm.

Etymology: Named after the host plant Podocarpus macrophyllus.

Diagnosis: Diaporthe podocarpi-macrophylli can be distinguished from the phylogenetically closely related species D. pseudophoenicicola (97% identity in ITS, 90% in TEF1, 98% in TUB, 97% in HIS, and 97% in CAL). Morphologically, D. podocarpi-macrophylli differs from other species occurring on the host genera Podocarpus and Olea, i.e. D. cinerascens and Phomopsis podocarpi in its wider and shorter alpha conidia and the presence of beta conidia (Chang et al. 2005, Gomes et al. 2013; https://nt.ars-grin.gov/fungaldatabases/).

Type: Japan: on healthy leaves of Podocarpus macrophyllus, 20 Sep. 2014, W.J. Duan (HMAS 247084 — holotype, dried culture; CGMCC 3.18281 = LC 6155-ex-type culture).

Description: Conidiomata pycnidial in culture on PDA, solitary or aggregated, deeply embedded in the PDA, erumpent, dark brown to black, 222–;699 µm diam, yellowish translucent conidial drops exuding from the ostioles. Alpha conidiophores 6–;18 × 1.5–;3 µm ( = 12.3 ± 2.6 × 2.1 ± 0.3, n = 30), hyaline, septate, branched, cylindrical, straight to sinuous, sometimes inflated, occurring in dense clusters. Beta conidiophores 10.5–;27 × 1.5–;2.5 µm ( = 15.3 ± 4.3 × 2.1 ± 0.3, n = 30), cylindrical to clavate, hyaline, septate, branched, smooth, straight. Alpha conidia 3.5–;8.5 × 1–;3 µm ( = 6.3 ± 1.7 × 2.1 ± 0.7, n = 50), unicellular, aseptate, fusiform, hyaline, usually biguttulate and acute at both ends. Beta conidia 8.5–;31.5 × 0.5–;2 µm ( = 19.5 ± 7.1 × 1.1 ± 0.4, n = 30), hyaline, aseptate, eguttulate, filiform, curved, tapering towards both ends, base truncate.

Culture characters: Cultures incubated on PDA at 25 °C in darkness, growth rate 12.5 mm diam/d. Colony at first white, becoming cream to yellowish, flat, with dense and felted mycelium, reverse pale brown with brownish dots with age, with visible solitary or aggregated conidiomata at maturity.

Additional material examined: Japan: on healthy leaves of Podocarpus macrophyllus, 20 Sep. 2014, W.J. Duan, culture LC 6141; ibid. culture LC 6144; ibid. culture LC 6156; ibid. culture LC 6157. — China: Zhejiang Province: on healthy leaves of P. macrophyllus, 10 Jul. 2015, W.J. Duan, culture LC 6194; ibid. culture LC 6195; ibid. culture LC 6196; ibid. culture LC 6197; ibid. culture LC 6198; ibid. culture LC 6199; ibid. culture LC 6200; ibid. culture LC 6201; ibid. culture LC 6202; ibid. culture LC 6235. — Italy: on healthy leaves of Olea europaea, 20 Sep. 2014, W.J. Duan, culture LC 6229.

Diaporthe undulata Y.H. Gao & L. Cai, sp. nov.

MycoBank MB820683

(Fig. 8)
Fig. 8
Fig. 8

Diaporthe undulata (CGMCC 3.18293). A. Leaves of host plant; B–;C. 30-d-old culture on PNAmedium; D. Conidiomata; E. Conidiophores; F–;G. Alpha conidia. Bars: D = 100 µm; E–;G = 10 µm.

Etymology: Named after the colony’s undulate margin.

Diagnosis: Diaporthe undulata differs from the most closely related species, D. biconispora, in several loci (94% in ITS, 84% in TEF1, and 93% in TUB), and from other Diaporthe species in the obpyriform conidiophores and shorter and wider alpha conidia (Table 3).

Type: China-Laos border: on diseased leaves of unknown host, 19 Apr. 2014, F Liu (HMAS 247091 — holotype, dried culture; CGMCC 3.18293 = LC 6624 — ex-type culture).

Description: Conidiomata pycnidial, irregular, embedded in the needle, erumpent, necks, hairy, 282–;543 µm long, coated with short hyphae, one to several necks forming from a single pycnidium. Conidiophores obpyriform, hyaline, phiailidic, septate, branched, 5–;17.5 × 2–;3 µm ( = 9.7 ± 4.0 × 2.4 ± 0.5, n = 20). Alpha conidia ellipsoid, hyaline, biguttulate, rounded at both ends, 5–;6.5 × 2–;3 ( = 5.8 ± 0.4 × 2.3 ± 0.3, n = 50). Beta conidia not observed.

Culture characters: Cultures incubated on PDA at 25 °C in darkness, growth rate 10.5 mm diam/d. Colony entirely white, reverse pale yellowish and dark brownish at the centre with age. Aerial mycelium white, cottony, dense, with undulate margin and visible conidiomata at maturity.

Additional material examined: China-Laos border: unknown host, 19 Apr. 2014, F. Liu, culture LC 8110; ibid. culture LC 8111.

Diaporthe velutina Y.H. Gao & L. Cai, sp. nov.

MycoBank MB820684

(Fig. 9)
Fig. 9
Fig. 9

Diaporthe velutina (CGMCC 3.18286). A. Diseased leaves; B–;C. 30-d-old culture on PDA; D. Conidiomata; E. Conidiophores; E. Alpha and beta conidia. Bars: D = 100 µm; E–;F = 10 µm.

Etymology: Named after the felted colony.

Diagnosis: Diaporthe velutina is distinguished from D. anacardii in the ITS, TEF1, TUB and HIS loci (99% in ITS, 95% in TEF1, 99% in TUB, and 98% in HIS), and from other Diaporthe species reported from Camellia sinensis in the more variable size of the alpha conidia (Table 3).

Type: China: Jiangxi Province: on diseased leaves of Neolitsea sp., 5 Sep. 2013, Y.H. Gao (HMAS 247087 — holotype, dried culture; CGMCC 3.18286 = LC4421 — ex-type culture).

Description: Conidiomata pycnidial, globose, black, embedded in PDA, aggregated in clusters, 69–;428 µm diam, cream translucent drop of conidia exuded from the central ostioles. Conidiophores 10–;23 × 1–;2.5 µm, cylindrical, hyaline, branched, densely aggregated, slightly tapering towards the apex, sometimes slightly curved. Alpha conidia 5.5–;10 × 2–;2.5 µm ( = 6.9 ± 0.9 × 2.2 ± 0.2, n = 50), unicellular, aseptate, hyaline, fusoid to ellipsoid or clavate, bi-guttulate or multi-guttulate. Beta conidia 11–;27.5 × 0.5–;1.5 µm ( = 16.1 ± 5.0 × 0.8 ± 0.4, n = 30), smooth, hyaline, apex acutely rounded, curved.

Culture characters: Cultures incubated on PDA at 25 °C in darkness, growth rate 18.75 mm diam/d. Colony entirely white, surface mycelium greyish to brownish at the centre, dense, felted, conidiomata erumpent at maturity, reverse centre yellowish to brownish.

Additional material examined: China: Jiangxi Province: Yangling, on diseased leaves of Neolitsea sp., 5 Sep. 2013, Y.H. Gao, culture LC 4419; ibid. culture LC 4422; Gannan Normal University, unknown host, 23 Apr. 2013, Q. Chen, culture LC 4788; Fengshan, on diseased leaves of Callerya cinerea, 5 Sep. 2013, Y.H. Gao, culture LC 4641. Yunnan Province: Xishuangbanna, on diseased leaves of Camellia sinensis, 19 Apr. 2015, F. Liu, culture LC 6708; loc. cit., on healthy leaves of C. sinensis, 21 Apr. 2015, F. Liu, culture LC 6519.

Diaporthe xishuangbanica Y.H. Gao & L. Cai, sp. nov.

MycoBank MB820685

(Fig. 10)
Fig. 10
Fig. 10

Diaporthexishuangbanica (CGMCC 3.18283). A–;B. 7-d-old culture on PDA; C–;D. 30-d-old culture on PNA medium; E. Conidiomata; F–;K. Conidiophores; L–;N. Alpha conidia. Bars: E = 100 µm; F–;N = 10 µm.

Etymology: Named after the locality, Xishuangbanna.

Diagnosis: Diaporthe xishuangbanica can be distinguished from the phylogenetically closely related D. tectonigena in several loci (98% in ITS, 90% in TEF1, and 96% in TUB) (Fig. 2), and from other Diaporthe species reported from Camellia in the longer conidiophores and alpha conidia (Table 3).

Type: China: Yunnan Province: Xishuangbanna, on diseased leaves of Camellia sinensis, 19 Apr. 2015, F. Liu (HMAS 247083 — holotype, dried culture; CGMCC 3.18283 = LC 6744 — ex-type culture).

Description: Conidiomata pycnidial, globose, 180–;310 µm diam, scattered on the pine needle. Conidiophores cylindrical, 13–;34.5 × 1.5–;3 µm ( = 20.9 ± 5.2 × 2.1 ± 0.3, n = 40), branched, septate, straight, sometimes sinuous or lateral. Alpha conidia 7–;9.5 × 2.5–;3.5 µm (x = 8.3 ± 0.7 × 2.8 ± 0.3, n = 30), fusiform, hyaline, multi-guttulate. Beta conidia not observed.

Culture characters: Cultures incubated on PDA at 25 °C in darkness, growth rate 17.5 mm diam/d. Colony entirely white, reverse pale yellowish to greenish. Aerial mycelium white, velvety, margin well defined, with visible conidiomata at maturity.

Additional material examined: China: Yunnan Province: Xishuangbanna, on diseased leaves of Camellia sinensis, 19 Apr. 2015, F. Liu, culture LC 6707 (CGMCC 3.18282).

Diaporthe yunnanensis Y.H. Gao & L. Cai, sp. nov.

MycoBank MB820686

(Fig. 11)
Fig. 11
Fig. 11

Diaporthe yunnanensis (fCGMCC 3.18289). A–;B. 7-d-old culture on PDA; C. Conidiomata; D. Conidiophores; E. Alpha and beta conidia; F. Beta conidia. Bars: C = 100 µm; D–;F = 10 µm.

Etymology: Named after the location where the fungus was collected, Yunnan Province.

Diagnosis: Diaporthe yunnanensis can be distinguished from the phylogenetically closely related D. siamensis (96% in ITS, 91% in TEF1, and 94% in TUB) (Fig. 2), and from other Diaporthe species reported on the genus Camellia in the smaller alpha conidia (Table 3).

Type: China: Yunnan Province: Xishuangbanna, on healthy leaves of Coffea sp., 20 Sep. 2014, W.J. Duan (HMAS 247096 — holotype, dried culture; CGMCC 3.18289 = LC 6168 — ex-type culture).

Description: Conidiomata pycnidial, 195–;880 µm diam, globose or irregular, erumpent, solitary or aggregated together, dark brown to black. Conidia exuding from the pycnidia in white to cream drops. Conidiophores cylindrical, straight or slightly curved. Alpha conidia 3–;6.5 × 1–;2.5 µm ( = 5.5 ± 1 × 2 ± 0.5, n = 30), fusiform, hyaline, biguttulate, with one end obtuse and the other acute. Beta conidia 13.5–;33.5 × 1–;1.5 µm ( = 27.5 ± 5.5 × 1.5 ± 0.3, n = 30), hyaline, aseptate, hamate or curved, base truncate.

Culture characters: Colonies on PDA flat, with a moderate growth rate of 5.5 mm diam/d, with abundant dirty white and yellowish pigmented mycelium, dry, felted, extensive thin, and in reverse the centre cream, with zone rings of pale to dark brownish pigmentation.

Additional material examined: China: Yunnan Province: Xishuangbanna, on healthy leaves of Coffea sp., 20 Sep. 2014, W.J. Duan, culture LC 8106; ibid. culture LC 8107.

Diaporthe sp. 1

(Fig. 12)
Fig. 12
Fig. 12

Diaporthe sp. 1 (CGMCC 3.18292). A. Leaves of host plant; B–;C. 30-d-old culture on PDA; D. Conidiomata; E–;F. Conidiophores; G. Beta conidia; H–;I. Alpha conidia. Bars: D = 100 µm; E–;I = 10 µm.

Description: Conidiomata pycnidial, subglobose to globose, dark brown to black, deeply embedded in the substrate, scattered on the substrate surface, embedded in PDA, clusters in group of 2–;7 pycnidia, 268–;509 µm, yellowish drop of conidia diffusing from the central ostioles. Conidiophores 6.5–;19.5 × 1–;3 µm, cylindrical, hyaline, septate, branched, straight to sinuous, base inflated, slightly tapering towards the apex. Alpha conidia 7.5–;13.5 × 2–;3.5 µm ( = 9.9 ± 1.4 × 2.8 ± 0.4, n = 30), unicellular, hyaline, fusoid to ellipsoid or clavate, two or several large guttulae observed, base subtruncate. Beta conidia 15–;10.5 × 1–;2.5 µm ( = 26.0 ± 5.8 × 1.8 ± 0.5, n = 30), smooth, hyaline, curved, base subtruncate, tapering towards one apex.

Culture characters: Cultures incubated on PDA at 25 °C in darkness, growth rate 7.83 mm diam/day. Colony entire, white to dirty pink, cottony, sparse, brownish to black conidiomata erumpent at maturity, coated with white hypha, granular at margin, reverse pale brown, with brownish dots when maturity.

Material examined: China: Zhejiang Province: Gutianshan Nature Reserve (29°20′ N 18°14′ E), on leaves of Alnus mill, Jan. 2010, Y.Y. Su (culture CGMCC 3.18292 = LC 0771).

Notes: The present culture belongs to the Diaporthe eres complex, which is reported from a very wide range of host plants and includes mostly opportunistic pathogens or secondary invaders on saprobic host substrata (Udayanga et al. 2014a, Gao et al. 2016). Species delimitation in this complex is currently unclear. Udayanga et al. (2015) accepted nine phylogenetic species in the D. eres complex, including D. alleghaniensis, D. alnea, D. bicincta, D. celastrina, D. eres, D. helicis, D. neilliae, D. pulla, and D. vaccinia. Gao et al. (2016) examined 17 isolates belonging to the D. eres complex, and reported that many presented intermediate morphology among “species” and the phylogenetic analyses often resulted in ambiguous clades with short branch and moderate statistical support. The identification of taxa in this group remains unresolved.

Diaporthe sp. 2

Culture characters: Cultures incubated on PDA at 25 °C in darkness, growth rate, slow, 3.83 mm diam/d. Colony low, convex, entire white to yellowish, reverse brownish. Aerial mycelia white, dry, downy, with near-circular margin.

Material examined: Japan: on leaves of Acer sp., 20 Sep. 2014, W.J. Duan, culture CGMCC 3.18291 = LC 6140, culture LC 8112; ibid. culture LC 8113.

Notes: Although three isolates clustered in a clade distinctly different from known species in the genus included, they are not formally described because they were sterile. Diaporthe sp. 2 shares a low homology to the most closely related species, D. rhoina (95% in ITS, 87% in TEF1, 97% in TUB, 94% in HIS, and 95% in CAL). Five Diaporthe species are so far only known from the sterile state, including D. endophytica, D. inconspicua, D. infecunda, D. asheicola, and D. sterilis (Gomes et al. 2013, Lombard et al. 2014).

Diaporthe averrhoae (C.Q. Chang et al.) Y.H. Gao & L. Cai, comb. nov.

MycoBankMB821437

Basionym: Phomopsis averrhoae C.Q. Chang et al., My-cosystema 24: 6 (2005).

Type: China: Fujian Province: on living branches of Averrhoa carambola, Y.H. Cheng (SCHM 3605 — holotype; AY618930, ITS sequence derived from the holotype SCHM 3605).

Diaporthe camptothecae (C.Q. Chang et al.) Y.H. Gao & L. Cai, comb. nov.

MycoBank MB821438

Basionym: Phomopsis camptothecae C.Q. Chang et al., My-cosystema 24: 145 (2005).

Type: China: Hunan Province: on living branches of Camptotheca acuminate, L.J. Luo (SCHM 3611 — holotype; AY622996, ITS sequence derived from the holotype SCHM 3611).

Diaporthe chimonanthi (C.Q. Chang et al.) Y.H. Gao & L. Cai, comb. nov.

MycoBank MB821439

Basionym: Phomopsis chimonanthi C.Q. Chang et al., My-cosystema 24: 146 (2005).

Type: China: Hunan Province: on living branches of Chimonanthus praecox, C.Q. Chang (SCHM 3614 — holotype; AY622993, ITS sequence derived from the holotype SCHM 3614).

Diaporthe eucommiae (F.X. Cao et al.) Y.H. Gao & L. Cai, comb. nov.

MycoBank MB821440

Basionym: Phomopsis eucommiae F.X. Cao et al., J. Middle-South China Forestry Coll. 10: 34 (1990); as ‘eucommi’

Type: China: Guangdong Province: from leaves of Eucommia ulmoides, F.X. Cao (SCHM 0020 — holotype; AY601921, ITS sequence derived from the holotype SCHM 0020).

Diaporthe eucommiicola (C.Q. Chang et al.) Y.H. Gao & L. Cai, comb. nov.

MycoBank MB821441

Basionym: Phomopsis eucommiicola C.Q. Chang et al., My-cosystema 24: 147 (2005).

Type: China: Hunan Province: on living branches of Eucommia ulmoides and Styrax hypoglauca, L.J. Luo (SCHM 3607 — holotype; AY578071, ITS sequence derived from the holotype SCHM 3607).

Diaporthe glabrae (C.Q. Chang et al.) Y.H. Gao & L. Cai, comb. nov.

MycoBankMB821443

Basionym: Phomopsis glabrae C.Q. Chang et al., My-cosystema 24: 8 (2005).

Type: China: Fujian Province: on living branches of Bougainvillea glabra, Y.H. Cheng (SCHM 3622 — holotype; AY601918, ITS sequence derived from the holotype SCHM 3622).

Diaporthe lagerstroemiae (C.Q. Chang et al.) Y.H. Gao & L. Cai, comb. nov.

MycoBank MB821444

Basionym: Phomopsis lagerstroemiae C.Q. Chang et al., My-cosystema 24: 148 (2005).

Type: China: Hunan Province: on living branches of Lagerstroemia indica, C.Q. Chang (SCHM 3608 — holotype; AY622994, ITS sequence derived from the holotype SCHM 3608).

Diaporthe liquidambaris (C.Q. Chang et al.) Y.H. Gao & L. Cai, comb. nov.

MycoBank MB821446

Basionym: Phomopsis liquidambaris C.Q. Chang et al., My-cosystema 24: 9 (2005).

Type: China: Fujian Province: on living branches of Liquidambar formosana, Y.H. Cheng (SCHM 3621 — holotype; AY601919, ITS sequence derived from the holotype SCHM 3621).

Diaporthe loropetali (C.Q. Chang et al.) Y.H. Gao & L. Cai, comb. nov.

MycoBank MB821448

Basionym: Phomopsis loropetali C.Q. Chang et al., My-cosystema 24: 148 (2005).

Type: China: Hunan Province: on living branches of Loropetalum chinense, C.Q. Chang (SCHM 3615 — holotype; AY601917, ITS sequence derived from the holotype SCHM 3615).

Diaporthe magnoliicola Y.H. Gao & L. Cai, nom. nov.

MycoBank MB821459

Replaced name: Phomopsis magnoliae M.M. Xiang et al., My-cosystema 21 : 501 (2002).

Type: China: Guangdong Province: on leaves of Magnolia coco, Z.D. Jiang (SCHM 3001 — holotype; AY622995, ITS sequence derived from the holotype SCHM 3001).

Note: The epithet magnoliae is occupied, so Diaporthe magnoliicola is proposed as a replacement name.

Diaporthe michelina (C.Q. Chang et al.) Y.H. Gao & L. Cai, comb. nov.

MycoBank MB821460

Basionym: Phomopsis michelina C.Q. Chang et al., My-cosystema 24: 9 (2005); as ‘micheliae’.

Type: China: Fujian Province: on living branches of Michelia alba, Y.H. Cheng (SCHM 3603 — holotype; AY620820, ITS sequence derived from the holotype SCHM 3603).

Diaporthe phyllanthicola (C.Q. Chang et al.) Y.H. Gao & L. Cai, comb. nov. MycoBank MB821461

Basionym: Phomopsis phyllanthicola C.Q. Chang et al., My-cosystema 24: 10 (2005).

Type: China: Fujian Province: on living branches of Phyllanthus emblica, Y.H. Cheng (SCHM 3680 — holotype; AY620819, ITS sequence derived from the holotype SCHM 3680).

Discussion

In this study, eight new species of Diaporthe are introduced, having been isolated from various plant hosts collected in different countries. Twelve Phomopsis species described from China were subjected to molecular analysis, and transferred to Diaporthe to conform to the “one fungus one name” rule (Udayanga et al. 2011, Rossman et al. 2016). To address the taxonomy of the other Phomopsis species described from China, neo- or epitypes will need to be designated to resolve their position and confirm their placement in Diaporthe.

Previous taxonomic studies in Diaporthe (syn. Phomopsis) have been primarily based on morphology, which has been shown to be unnatural in reflecting evolutionary history due to the simple and plastic morphological characters (Gao et al. 2015). The same applies to many other genera of ascomycetes. For example, species referred to Phoma have been shown to be highly polyphyletic and scattered throughout at least six families within Pleosporales (Aveskamp et al. 2010, Chen et al. 2015). Although Diaporthe was previously thought to be monophyletic based on its typical and unique Phomopsis asexual morph and diaporthalean sexual morph (Gomes et al. 2013), a paraphyletic nature is revealed in the present study (Fig. 1). Several genera, notably Ophiodiaporthe (Fu et al. 2013), Pustulomyces (Dai et al. 2014), Phaeocytostroma, and Stenocarpella (Lamprecht et al. 2011), are shown to be embedded in Diaporthe s. lat., none of which present an independent lineage from Diaporthe as currently circumscribed (Fig. 1). These genera were established based on their morphological characteristics (Vasilyeva et al. 2007, Lamprecht et al. 2011, Fu et al. 2013, Dai et al. 2014). For example, Ophiodiaporthe produces only one type of globose or subglobose conidia that differs from the dimorphic (fusiform and filiform) conidia of Diaporthe (Fu et al. 2013); Phaeocytostroma and Stenocarpella produce pigmented alpha conidia which differ from the hyaline conidia of Diaporthe (Lamprecht et al. 2011); Pustulomyces produces larger, straight or sigmoid conidia (Dai et al. 2014). Phaeocytostroma and Stenocarpella were originally suspected to be members of Botryosphaeriaceae (Botryosphaeriales) because of their pigmented alpha conidia and diplodia-like morphology (Crous et al. 2006). However, they were subsequently allocated to Diaporthales based on phylogenetic analysis (Lamprecht et al. 2011), which is confirmed in this study.

The large “Diaporthe” clade embedded with the heterogeneous genera Ophiodiaporthe, Pustulomyces, Phaeocytostroma, and Stenocarpella is probably a typical example of divergent evolution in morphological characters. Such an evolution could have been driven by host and/or environmental adaptations. For example, the monotypic Ophiodiaporthe is associated with Cyathea lepifera (a fern), while Pustulomyces is bambusicolous (Dai et al. 2014). On the contrary, none of the previously named over 1 900 Diaporthe / Phomopsis species was recorded from a fern or Bambusaceae (https://nt.ars-grin.gov/fungaldatabases/). It is therefore reasonable to speculate that the speciation of Ophiodiaporthe and Pustulomyces, as well as the distinctly different morphologies from their close Diaporthe allies, are the consequences of evolutionary adaption to new hosts. Similarly, Phaeocytostroma and Stenocarpella are mainly restricted to maize (Zea mays), causing root stalk and cob rot (Stovold et al. 1996, Lamprecht et al. 2011).

Splitting Diaporthe into many smaller genera would achieve monophyletic groupings, but would also create many additional problems. The “new genera” split from Diaporthe would have no recognisable morphological distinctions in either sexual or asexual morphs. In addition, splitting Diaporthe into many smaller genera will result in numerous name changes, which is certainly an unfavourable option for both mycologists and plant pathologists.

Diaporthe has long been well-known to include plant pathogens, some on economically important hosts, such as Helianthus annuus (sunflower; Thompson et al. 2011) and Glycine max (soybean; Santos et al. 2011). However, the number of known endophytic Diaporthe species has increased rapidly in recent years (Huang et al. 2015, Gao et al. 2016). Wang et al. (2013) concluded that our current knowledge of the ecology and biology of endophytic Diaporthe species is just the “tip of the iceberg”. In 2013, a new sterile endophytic species, Diaporthe endophytica, was formally named (Gomes et al. 2013). The research on Citrus conducted by Huang et al. (2015) recorded seven apparently undescribed endophytic Diaporthe species. Inspection of Diaporthe species on Camellia sinensis resulted in the description of four new and five known species, all occurring as endophytes (Gao et al. 2016). Because many of these plant pathogenic Diaporthe species are commonly encountered as sterile endophytes, a multigene DNA database will be essential to aid in theirfuture identification.

Accurate identification of fungal pathogens is the basis of quarantine and disease control (Udayanga et al. 2011). Thompson et al. (2011) reported significant damage to sunflower in Australia caused by Diaporthe helianthi which was originally only known from Europe (former Yugoslavia), and is apparently an invasive species in Australia. Diaporthe helianthi is listed in the Chinese quarantine directory, and has long been considered a predominant disease limiting production in Europe (Desanlis et al. 2013). Duan et al. (2016) reported this pathogen on sunflower seeds imported from Ukraine into China. Here, we report another interception of D. helianthi from Lagerstroemia indica imported from Japan to China. This serves as additional evidence of how quickly serious pathogens such as Diaporthe species can be distributed as endophytes or latent pathogens with global trade.

Notes

Declarations

Acknowledgements

We thank all the members in LC’s lab for help and assistance. This work was supported by grants from the National Natural Science Foundation of China (NSFC 31110103906), and the Ministry of Science and Technology, China (MOST 2014FY120100).

This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made.

Authors’ Affiliations

(1)
State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing, 100101, P.R. China
(2)
University of Chinese Academy of Sciences, Beijing, 100049, P.R. China
(3)
Ningbo Academy of Inspection and Quarantine, Zhejiang, 315012, P.R. China
(4)
Westerdijk Fungal Biodiversity Institute, Uppsalalaan 8, 3584CT Utrecht, The Netherlands
(5)
Department of Microbiology and Plant Pathology, Tree Protection Co-operative Programme, Forestry and Agricultural Biotechnology Institute, University of Pretoria, Pretoria, 0002, South Africa

References

  1. Annesi T, Luongo L, Vitale S, Galli M, Belisario A (2015) Characterization and pathogenicity of Phomopsis theicola anamorph of Diaporthe foeniculina causing stem and shoot cankers on sweet chestnut in Italy. Journal of Phytopathology 164: 412–416.View ArticleGoogle Scholar
  2. Aveskamp MM, de Gruyter J, Woudenberg JHC, Verkley GJM, Crous PW (2010) Highlights of the Didymellaceae: a polyphasic approach to characterise Phoma and related pleosporalean genera. Studies in Mycology 65: 1–60.View ArticleGoogle Scholar
  3. Carbone I, Kohn LM (1999) A method for designing primer sets for speciation studies in filamentous ascomycetes. Mycologia 91: 553–556.View ArticleGoogle Scholar
  4. Castlebury LA, Rossman AY, Jaklitsch WJ, Vasilyeva L (2002) A preliminary overview of the Diaporthales based on large subunit nuclear ribosomal DNA sequences. Mycologia 94: 1017–1031.View ArticleGoogle Scholar
  5. Chang CQ, Cheng YH, Xiang MM, Jiang ZD (2005) New species of Phomopsis on woody plants in Fujian Province. Mycosystema 24: 6–11.Google Scholar
  6. Chen Q, Jiang JR, Zhang GZ, Cai L, Crous PW (2015) Resolving the Phoma enigma. Studies in Mycology 82: 137–217.View ArticleGoogle Scholar
  7. Chi PK, Jiang ZD, Xiang MM (2007) Flora Fungorum Sinicorum. Vol. 34. Phomopsis. Beijing: Science Press.Google Scholar
  8. Crous PW, Groenewald JZ, Risède JM, Simoneau P, Hywel-Jones NL (2004) Calonectria species and their Cylindrocladium anamorphs: species with sphaeropedunculate vesicles. Studies in Mycology 50: 415–430.Google Scholar
  9. Crous PW, Slippers B, Wingfield MJ, Rheeder J, Marasas WFO, et al. (2006) Phylogenetic lineages in the Botryosphaeriaceae. Studies in Mycology 55: 235–253.View ArticleGoogle Scholar
  10. Crous PW, Wingfield MJ, Le Roux JJ, Richardson DM, Strasberg D, et al. (2015) Fungal Planet Description Sheets: 371–399. Persoonia: 35: 264–327.View ArticleGoogle Scholar
  11. Cubero OF, Crespo A, Fatehi J, Bridge PD (1999) DNA extraction and PCR amplification method suitable for fresh, herbarium-stored, lichenized, and other fungi. Plant Systematics and Evolution 216: 243–249.View ArticleGoogle Scholar
  12. Dai DQ, Wijayawardene NN, Bhat DJ, Chukeatirote E, Bahkali AH, et al. (2014) Pustulomyces gen. nov. accommodated in Diaporthaceae, Diaporthales, as revealed by morphology and molecular analyses. Cryptogamie, Mycologie 35: 63–72.View ArticleGoogle Scholar
  13. Darriba D, Taboada GL, Doallo R, Posada D (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods 9: 772.View ArticleGoogle Scholar
  14. Desanlis M, Aubertot JN, Mestries E, Debaeke P (2013) Analysis of the influence of a sunflower canopy on Phomopsis helianthi epidemics as a function of cropping practices. Field Crops Research 149: 63–75.View ArticleGoogle Scholar
  15. Diogo EL, Santos JM, Phillips AJ (2010) Phylogeny, morphology and pathogenicity of Diaporthe and Phomopsis species on almond in Portugal. Fungal Diversity 44: 107–115.View ArticleGoogle Scholar
  16. Dissanayake AJ, Liu M, Zhang W, Chen Z, Udayanga D, et al. (2015) Morphological and molecular characterisation of Diaporthe species associated with grapevine trunk disease in China. Fungal Biology 119: 283–294.View ArticleGoogle Scholar
  17. Doilom M, Dissanayake AJ, Wanasinghe DN, Boonmee S, Liu JK, et al. (2017) Microfungi on Tectona grandis (teak) in northern Thailand. Fungal Diversity 82: 107–182.View ArticleGoogle Scholar
  18. Du Z, Fan XL, Hyde KD, Yang Q, Liang YM, et al. (2016). Phylogeny and morphology reveal two new species of Diaporthe from Betula spp. in China. Phytotaxa 269: 90–102.View ArticleGoogle Scholar
  19. Duan WJ, Duan LJ, Chen XF, Cai L (2016) Identification of the quarantine fungus Diaporthe helianthi from the corn seeds imported from Ukraine. Mycosystema 35: 1503–1513.Google Scholar
  20. Fan XL, Hyde KD, Udayanga D, Wu XY, Tian CM (2015) Diaporthe rostrata, a novel ascomycete from Juglans mandshurica associated with walnut dieback. Mycological Progress 14: 82.View ArticleGoogle Scholar
  21. Fan XL, Tian CM, Qin Y, Liang YM, You CJ, et al. (2014) Cytospora from Salix in northern China. Mycotaxon 129: 303–315.View ArticleGoogle Scholar
  22. Fu CH, Hsieh HM, Chen CY, Chang TT, Huang YM, et al. (2013) Ophiodiaporthe cyatheae gen. et sp. nov, a diaporthalean pathogen causing a devastating wilt disease of Cyathea lepifera in Taiwan. Mycologia 105: 861–872.View ArticleGoogle Scholar
  23. Gao YH, Sun W, Su YY, Cai L (2014) Three new species of Phomopsis in Gutianshan nature reserve in China. Mycological Progress 13: 111–121.View ArticleGoogle Scholar
  24. Gao YH, Su YY, Sun W, Cai L (2015) Diaporthe species occurring on Lithocarpus glabra in China, with descriptions of five new species. Fungal Biology 119: 295–309.View ArticleGoogle Scholar
  25. Gao YH, Liu F, Cai L (2016) Unravelling Diaporthe species associated with Camellia. Systematics and Biodiversity 14: 102–117.View ArticleGoogle Scholar
  26. Glass NL, Donaldson GC (1995) Development of primer sets designed for use with the PCR to amplify conserved genes from filamentous ascomycetes. Applied and Environmental Microbiology 61: 1323–1330.PubMedPubMed CentralGoogle Scholar
  27. Gomes R, Glienke C, Videira S, Lombard L, Groenewald J, et al. (2013) Diaporthe: a genus of endophytic, saprobic and plant pathogenic fungi. Persoonia 31: 1–41.View ArticleGoogle Scholar
  28. Grasso FM, Marini M, Vitale A, Firrao G, Granata G (2012) Canker and dieback on Platanus acerifolia caused by Diaporthe scabra. Forest Pathology 42: 510–513.View ArticleGoogle Scholar
  29. Guarnaccia V, Vitale A, Cirvilleri G, Aiello D, Susca A, et al. (2016) Characterisation and pathogenicity of fungal species associated with branch cankers and stem-end rot of avocado in Italy. European Journal of Plant Pathology 146: 963–976.View ArticleGoogle Scholar
  30. Huang F, Udayanga D, Wang X, Hou X, Mei X, et al. (2015) Endophytic Diaporthe associated with Citrus: A phylogenetic reassessment with seven new species from China. Fungal Biology 119: 331–347.View ArticleGoogle Scholar
  31. Katoh K, Toh H (2010) Parallelization of the MAFFT multiple sequence alignment program. Bioinformatics 26: 1899–1900.View ArticleGoogle Scholar
  32. Lamprecht SC, Crous PW, Groenewald JZ, Tewoldemedhin YT, Marasas WF (2011) Diaporthaceae associated with root and crown rot of maize. IMA Fungus 2: 13–24.View ArticleGoogle Scholar
  33. Diedicke H (1911) Die Gattung Phomopsis. Annales Mycologic 9: 8–35.Google Scholar
  34. Liu F, Wang M, Damm U, Crous PW, Cai L (2016) Species boundaries in plant pathogenic fungi: a Colletotrichum case study. BMC Evolutionary Biology 16: 81.Google Scholar
  35. Liu F, Weir BS, Damm U, Crous PW, Wang Y, et al. (2015) Unravelling Colletotrichum species associated with Camellia: employing ApMat and GS loci to resolve species in the C. gloeosporioides complex. Persoonia 35: 63–86.View ArticleGoogle Scholar
  36. Lombard L, Van Leeuwen GCM, Guarnaccia V, Polizzi G, Van Rijswick PC, et al. (2014) Diaporthe species associated with Vaccinium, with specific reference to Europe. Phytopathologia Mediterranea 53: 287–299.Google Scholar
  37. Machingambi NM, Dreyer LL, Oberlander KC, Roux J, Roets F (2015) Death of endemic Virgilia oroboides trees in South Africa caused by Diaporthe virgiliae sp. nov. Plant Pathology 64: 1149–1156.View ArticleGoogle Scholar
  38. Masirevic S, Gulya T (1992) Sclerotinia and Phomopsis —two devastating sunflower pathogens. Field Crops Research 30: 271–300.View ArticleGoogle Scholar
  39. Ménard L, Brandeis PE, Simoneau P, Poupard P, Sérandat I, et al. (2014) First report of umbel browning and stem necrosis caused by Diaporthe angelicae on carrot in France. Plant Pathology 98: 421.Google Scholar
  40. Mostert L, Crous PW, Kang JC, Phillips AJ (2001) Species of Phomopsis and a Libertella sp. occurring on grapevines with specific reference to South Africa: morphological, cultural, molecular and pathological characterization. Mycologia 93: 146–167.View ArticleGoogle Scholar
  41. Muntanola-Cvetkovic M, Mihaljcevic M, Petrov M (1981) On the identity of the causative agent of a serious Phomopsis-Diaporthe disease in sunflower plants. Nova Hedwigia 34: 417–435.Google Scholar
  42. Nylander JAA (2004) MrModeltest v. 2. Program distributed by the author. Uppsala: Evolutionary Biology Centre, Uppsala University.Google Scholar
  43. O’Donnell K, Cigelnik E (1997) Two divergent intragenomic rDNA ITS2 types within a monophyletic lineage of the fungus Fusarium are nonorthologous. Molecular Phylogenetics and Evolution 7: 103–116.View ArticleGoogle Scholar
  44. Petch T (1925) Additions to Ceylon fungi. III. Annals of the Royal Botanic Gardens, Peradeniya 9: 313–328Google Scholar
  45. Phillips AJL (2003) Morphological characterization of Diaporthe foeniculacea and its Phomopsis anamorph on Foeniculum vulgare. Sydowia 55: 274–285.Google Scholar
  46. Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, et al. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539–542.View ArticleGoogle Scholar
  47. Rossman AY, Adams GC, Cannon PF, Castlebury LA, Crous PW et al. (2015) Recommendations of generic names in Diaporthales competing for protection or use. IMA Fungus 6: 145–154.View ArticleGoogle Scholar
  48. Rossman AY, Allen WC, Braun U, Castlebury LA, Chaverri P, et al. (2016) Overlooked competing asexual and sexually typified generic names of Ascomycota with recommendations for their use or protection. IMA Fungus 7: 289–308.View ArticleGoogle Scholar
  49. Rossman A, Udayanga D, Castlebury LA, Hyde KD (2014) (2304) Proposal to conserve the name Diaporthe eres against twenty-one competing names (Ascomycota: Diaporthales: Diaporthaceae). Taxon 63: 934–935.View ArticleGoogle Scholar
  50. Rytas V, Mark H (1990) Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. Journal of Bacteriology 172: 4238–4246.View ArticleGoogle Scholar
  51. Santos JM, Vrandecic K, Cosic J, Duvnjak T, Phillips AJ (2011) Resolving the Diaporthe species occurring on soybean in Croatia. Persoonia 27: 9–19.View ArticleGoogle Scholar
  52. Santos L, Alves A, Alves R (2017) Evaluating multi-locus phylogenies for species boundaries determination in the genus Diaporthe. PeerJ 5: e3120.View ArticleGoogle Scholar
  53. Smith H, Wingfield MJ, Coutinho TA, Crous PW (1996) Sphaeropsis sapinea and Botryosphaeria dothidea endophytic in Pinus spp. and Eucalyptus spp. in South Africa. South African Journal of Botany 62: 86–88.View ArticleGoogle Scholar
  54. Stamatakis A (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22: 2688–2690.View ArticleGoogle Scholar
  55. Stamatakis A, Hoover P, Rougemont J (2008) A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology 57: 758–771.View ArticleGoogle Scholar
  56. Su YY, Qi YL, Cai L (2012) Induction of sporulation in plant pathogenic fungi. Mycology 3: 195–200.Google Scholar
  57. Tamura K, Peterson D, Peterson N, Stecher G, Nei M, et al. (2011) MEGA5: molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution 28: 2731–2739.View ArticleGoogle Scholar
  58. Tan Y, Edwards J, Grice K, Shivas R (2013) Molecular phylogenetic analysis reveals six new species of Diaporthe from Australia. Fungal Diversity 61: 251–260.View ArticleGoogle Scholar
  59. Thompson S, Tan Y, Young A, Neate S, Aitken E, et al. (2011) Stem cankers on sunflower (Helianthus annuus) in Australia reveal a complex of pathogenic Diaporthe (Phomopsis) species. Persoonia 27: 80–89.View ArticleGoogle Scholar
  60. Thompson S, Tan Y, Shivas R, Neate S, Morin L, et al. (2015) Green and brown bridges between weeds and crops reveal novel Diaporthe species in Australia. Persoonia 35: 39–49.View ArticleGoogle Scholar
  61. Torres C, Camps R, Aguirre R, Besoain XA (2016) First report of Diaporthe rudis in Chile causing Stem-End rot on ‘Hass’ avocado fruit imported from California, USA. Plant Disease 100: 1951.View ArticleGoogle Scholar
  62. Udayanga D, Castlebury LA, Rossman AY, Chukeatirote E, Hyde KD (2014a) Insights into the genus Diaporthe: phylogenetic species delimitation in the D. eres species complex. Fungal Diversity 67: 203–229.View ArticleGoogle Scholar
  63. Udayanga D, Castlebury LA, Rossman AY, Chukeatirote E, Hyde KD (2015) The Diaporthe sojae species complex: phylogenetic re-assessment of pathogens associated with soybean, cucurbits and other field crops. Fungal Biology 119: 383–407.View ArticleGoogle Scholar
  64. Udayanga D, Castlebury LA, Rossman AY, Hyde KD (2014b) Species limits in Diaporthe: molecular re-assessment of D. citri, D. cytosporella, D. foeniculina and D. rudis. Persoonia 32: 83–101.View ArticleGoogle Scholar
  65. Udayanga D, Liu X, McKenzie EHC, Chukeatirote E, Bahkali AHA, et al. (2011) The genus Phomopsis: biology, applications, species concepts and names of common phytopathogens. Fungal Diversity 50: 189–225.View ArticleGoogle Scholar
  66. Udayanga D, Liu X, Mckenzie EHC, Chukeatirote E, Hyde KD (2012) Multi-locus phytogeny reveals three new species of Diaporthe from Thailand. Cryptogamie, Mycologie 33: 295–309.View ArticleGoogle Scholar
  67. Uecker FA (1988) A World list of Phomopsis names with notes on nomenclature, morphology and biology. Mycological Memoir 13: 1–231.Google Scholar
  68. ’Urbez-Torres JR, Peduto F, Smith RJ, Gubler WD (2013) Phomopsis dieback: a grapevine trunk disease caused by Phomopsis viticola in California. Plant Disease 97: 1571–1579.View ArticleGoogle Scholar
  69. Van Niekerk JM, Groenewald JZ, Farr DF, Fourie PH, Halleen F, et al. (2005) Reassessment of Phomopsis species on grapevines. Australasian Plant Pathology 34: 27–39.View ArticleGoogle Scholar
  70. Van Rensburg JCJ, Lamprecht SC, Groenewald JZ, Castlebury LA, Crous PW (2006) Characterisation of Phomopsis spp. associated with die-back of rooibos (Aspalathus linearis) in South Africa. Studies in Mycology 55: 65–74.View ArticleGoogle Scholar
  71. Vasilyeva LN, Rossman, AY, Farr DF (2007) New species of the Diaporthales from eastern Asia and eastern North America. Mycologia 99: 916–923.View ArticleGoogle Scholar
  72. Wang J, Xu X, Mao L, Lao J, Lin F, et al. (2013) Endophytic Diaporthe from southeast China are genetically diverse based on multi-locus phylogeny analyses. World Journal of Microbiology and Biotechnology 30: 237–243.View ArticleGoogle Scholar
  73. Wehmeyer LE (1926) A biologic and phylogenetic study of stromatic Sphaeriales. American Journal of Botany 13: 575–645.View ArticleGoogle Scholar
  74. Stovold GE, Newfield A, Priest MJ (1996) Root and stalk rot of maize caused by Phaeocytostroma ambiguum recorded for the first time in New South Wales. Australasian Plant Pathology 25: 50–54.View ArticleGoogle Scholar
  75. White TJ, Bruns T, Lee S, Taylor J (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: PCR Protocols: a guide to methods and applications. (Innis MA, Gelfand DH, Sninsky JJ, White TJ, eds): 315–322. San Diego: Academic Press.Google Scholar
  76. Tanney JB, Mcmullin DR, Green BD, Miller JD, Seifert KA (2016) Production of antifungal and antiinsectan metabolites by Picea endophyte Diaporthe maritima sp. nov. Fungal Biology 120: 1448–1457.View ArticleGoogle Scholar
  77. Zhang K, Su YY, Cai L (2013) An optimized protocol of single spore isolation for fungi. Cryptogamie, Mycologie 34: 349–356.View ArticleGoogle Scholar

Copyright

© International Mycological Association 2017

Advertisement