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A phylogenetic revision of Penicillium sect. Exilicaulis ... · Penicillium sect. Exilicaulis was the dominant group isolated from the fynbos, with 740 isolates accounting for roughly

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  • ARTIC

    LE

    75

    2016 International Mycological Association

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    v o l u m e 7 n o . 1

    A phylogenetic revision of Penicillium sect. Exilicaulis, including nine new species from fynbos in South Africa

    Cobus M. Visagie1,2,3, Keith A. Seifert1, Jos Houbraken2, Robert A. Samson2, and Karin Jacobs3

    1Biodiversity (Mycology), Agriculture and Agri-Food Canada, Ottawa, ON K1A0C6, Canada; corresponding author e-mail: [email protected] 2CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, NL-3584 CT Utrecht, The Netherlands3Department of Microbiology, University of Stellenbosch, Private Bag X1, Stellenbosch 7600, South Africa

    Abstract: A survey of the fynbos biome in South Africa resulted in the isolation of 61 Penicillium species from Protea repens infructescences, air, and soil samples. Fourteen of these belong to Penicillium sect. Exilicaulis and therefore we considered it an opportunity to re-evaluate the taxonomy of the section. Phylogenetic comparisons of the ITS, -tubulin, calmodulin and RPB2 gene regions of the 76 section Exilicaulis species, revealed 52 distinct species, including nine new species from fynbos. Morphological comparisons confirmed the novelty for most of these, however, new species closely related to P. rubefaciens did not show significant or consistent morphological differences and we thus placed a bias on phylogenetic data applying the Genealogical Concordance Phylogenetic Species Recognition (GCPSR) concept. In this paper we describe the nine new species and update the accepted species list and resolve synonyms in the section. Importantly, we reveal that P. citreosulfuratum is the correct name for the clade previously considered to represent P. toxicarium fide Serra et al. (2008).

    The nine new species are: Penicillium atrolazulinum, P. consobrinum, P. cravenianum, P. hemitrachum, P. pagulum, P. repensicola, P. momoii, P. subturcoseum, and P. xanthomelinii spp. nov.

    Article info: Submitted: 12 November 2015; Accepted: 15 May 2016; Published: 1 June 2016.

    IntroductIon

    Pitt (1980) introduced the name Penicillium sect. Exilicaulis, typified by P. restrictum, for species with monoverticillate conidiophores and non-vesiculated stipes. In the recent re-classification of Penicillium based on phylogenetic data (Houbraken & Samson 2011), section Exilicaulis was redefined to include biverticillate species such as P. corylophilum, P. melinii, and P. raciborskii. With a limited strain and species sampling, Houbraken & Samson (2011) separated species into two main clades. This was also supported by a more comprehensive RPB2 (RNA polymerase II second largest subunit) phylogeny revealing a well-supported main P. corylophilum clade (containing biverticillate species), whereas monoverticillate species resolved in several basal clades.

    Section Exilicaulis has received some taxonomic attention in recent years, including the introduction of new species (Langlois et al. 2014, Peterson et al. 1999, 2011), but more importantly species and their metabolites were shown to be toxic and to affect human health (Kremer et al. 1989, Lyratzopoulos et al. 2002, McMullin et al. 2014, Miyake 1940, Ohnishi et al. 2002, Peterson et al. 1999, Rosa et al. 2010, Unoura et al. 2011, Yoshida et al. 1992). Serra et al. (2008) studied the diversity of Penicillium species in cork bark with a focus on some species of sect. Exilicaulis, stressing the

    importance of applying the Genealogical Concordance Phylogenetic Species Recognition (GCPSR) concept (Taylor et al. 2000) to Penicillium. This approach to species delineation has become standard in Penicillium, especially in studies focused on resolving the taxonomy of specific sections (Houbraken & Samson 2011, Houbraken et al. 2011, 2014, Peterson et al. 2011, Rivera & Seifert 2011, Visagie et al. 2013). This concept was also applied in section Exilicaulis, but the focus was mostly on a subset of species from the P. parvum-clade (Langlois et al. 2014, Peterson et al. 1999, 2011) and the P. citreonigrum complex (Serra et al. 2008), but did not include all species belonging to these clades.

    During a survey of species diversity in the fynbos biome, situated in the Western Cape of South Africa, Penicillium was one of the dominant genera isolated, with roughly 61 species found. Fourteen belong to section Exilicaulis and nine of those represent new species. Here we compare the morphology of these new species with all others previously classified in section Exilicaulis and apply GCPSR using phylogenies of the rDNA region (ITS), -tubulin (BenA), calmodulin (CaM), and RPB2 genes. This paper follows previous papers reporting new Penicillium species isolated during this survey belonging to sections Aspergilloides (Houbraken et al. 2014), Citrina (Visagie et al. 2014c), Lanata-Divaricata (Visagie et al. 2015), Sclerotiora (Visagie et al. 2013), and Torulomyces (Visagie et al. 2016a).

    Key words: Citreovirdincryptic speciesDNA barcodemitesvector

    doi:10.5598/imafungus.2016.07.01.06IMA FunguS 7(1): 75117 (2016)

  • Visagie et al.ARTICLE

    76 i m a f U N G U S

    MAterIAl And MethodS

    Sampling and isolationSoil, air and Protea repens infructescence samples were taken from three fynbos sampling sites at Stellenbosch (335647S 185249E), Riverlands (332946S 183560E), and Struisbaai (334506S 185859E). Potato dextrose agar (PDA) containing streptomycin (100 ppm), chloramphenicol (50 ppm) and dichloran (0.002 g/L) was used as isolation medium. Isolation plates were incubated at 25 C for 7 d, after which colonies resembling Penicillium were transferred to oatmeal agar (OA).

    Isolations from soil were made using a dilution series by adding 5 g soil to 100 mL dH2O and diluting this tenfold three times resulting in suspensions 10-1, 10-2 and 10-3 of the original. In triplicate, 1 mL from each dilution was spread onto PDA plates. Isolations from Protea repens infructescences included sampling from bracts and mite populations. Firstly, infructescences were cut open and the mites shaken onto 1 % water agar plates with chloroform used to temporarily sedate mites. Mites were transferred onto PDA plates using a fine needle and incubated on a lab bench ( 21 C) on soapy water traps to prevent cross contamination by awaken mites roaming on plates. Roughly 15 bracts from infructescences were suspended in 100 mL dH2O. This solution was diluted tenfold six times resulting in suspensions 10-4, 10-5 and 10-6 of the original and spread plates prepared similarly to the soil spread plates. Air was sampled close to the same sampled Protea repens bushes using a MAS-Eco air sampler set to a volume of 50 L.

    reference strainsReference strains used for this study (Table 1), were obtained from the public collection of CBS (CBS-KNAW Fungal Biodiversity Centre, Utrecht) and cultures from the working collection of the Applied and Industrial Mycology group at the same institute (DTO). Strains isolated during this study are kept in a private collection at the department of Microbiology, Stellenbosch University, South Africa (CV). These strains were also deposited in DTO, with a subset of strains deposited into DAOMC (Canadian Collection of Fungal Cultures, Ottawa) and CBS.

    dnA extraction, sequencing and phylogenetic analysisDNA was extracted from strains grown on MEA for 7 d using the ZR Fungal/Bacterial DNA kit (ZymoResearch, CA), with extracts stored at 4 C. Amplification of the ITS region, partial -tubulin (BenA), calmodulin (CaM), and RNA polymerase II second largest subunit (RPB2) were made using primers and amplification profiles described in Visagie et al. (2014b). PCR and sequencing reactions followed similar methods to Visagie et al. (2013). For some strains, the partial RPB2 gene was sequenced using internal sequencing primers RPB2Int388F and RPB2Int-527R as described in Houbraken & Samson (2011). Sequence contigs were assembled using CodonCode Aligner v. 4.0.1 (CodonCode Centerville, MA, USA). Newly generated sequences were deposited in GenBank with accession numbers provided in Table 1.

    Fynbos strains isolated during this study were added to a sequence database that includes all ex-type strains of species belonging to section Exilicaulis. All data sets were aligned using MAFFT v. 7.164b (Katoh & Standley 2013) implementing the L-INS-i option. For multigene phylogenies, aligned data sets for each gene or region were concatenated in SeaView v. 4.4.1 (Gouy et al. 2010).

    Phylogenies were constructed using both Maximum Likelihood (ML) and Bayesian tree inference (BI) analyses. ML analyses were done using MEGA v. 6.06 (Tamura et al. 2013). The most suitable model for each data set was selected based on the lowest Akaike information criterion (AIC) value in MEGA. ML analyses were run by calculating an initial Bio-Neighbour-Joining tree, followed by a Heuristic search with the Nearest-Neighbour-Interchange (NNI) option. Bootstrap analysis with 1000 replicates was used to represent support for nodes. For BI, the most suitable model for each data set was selected in MrModeltest v. 2.3 (Nylander et al. 2004) based on the lowest AIC value. Subsequent analyses were run in MrBayes v. 3.2.1 (Ronquist & Huelsenbeck 2003) using three sets of four chains (one cold and three heated) and stopped when the average standard of deviation for split frequencies reached 0.01. The sample frequency was then set at 100 and 25 % of trees were removed as burn-in. ML phylograms were used for representing phylogenies obtained from ML and BI analyses, with bootstrap values ( 80 %) and/or posterior probabilities ( 0.95 pp) both indicated above thickened branches.

    MorphologyFynbos strains were characterised on Czapek yeast autolysate agar (CYA), Blakeslees malt extract agar (MEA; Bacto), yeast extract sucrose agar (YES), 25 % glycerol nitrate agar (G25N), and creatine sucrose agar (CREA), incubated at 25 C for 7 d, with additional CYA plates incubated at 30 and 37 C. CREA contains bromocresol purple and is used as a rapid method for detecting acid production by colonies, indicated by a colour change in the medium from purple to yellow (Frisvad 1981). Media preparation, inoculation technique, incubation conditions and microscope preparations were standardised according to the recommended methods published in Visagie et al. (2014b). Colour names and codes used in descriptions are based on Kornerup & Wanscher (1967). An Olympus SZX12 dissecting microscope and Olympus BX50 light microscope equipped with an Evolution MP digital microscope camera and ImagePro v. 6.0 software were used for microphotography. Photographic plates were prepared in Adobe Photoshop CS6.

    reSultS

    Sampling and isolationsPenicillium sect. Exilicaulis was the dominant group isolated from the fynbos, with 740 isolates accounting for roughly 43 % of the total obtained. Strains belonging to the section were sorted into morpho-groups based on colony characters on CYA and MEA, with a subset of strains from each group subsequently sequenced and compared with reference sequences. The dominance of P. citreosulfuratum (197 isolates in total, with only three each from air and soil) and

  • Penicillium sect. ExilicaulisARTIC

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    77v o l u m e 7 n o . 1

    tabl

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    Afri

    can.

    a.JX

    1411

    03JX

    1574

    56K

    P06

    4597

    P. a

    trola

    zulin

    umD

    TO 1

    85-C

    1 =

    CV

    644

    Mite

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, S

    telle

    nbos

    ch, S

    outh

    Afri

    can.

    a.JX

    1411

    05JX

    1574

    57K

    P06

    4598

  • Visagie et al.ARTICLE

    78 i m a f U N G U S

    tabl

    e 1.

    (Con

    tinue

    d).

    gen

    Ban

    k ac

    cess

    ion

    num

    bers

    Spec

    ies

    nam

    ec

    olle

    ctio

    n nu

    mbe

    rsSu

    bstr

    ate,

    loc

    atio

    nIt

    SB

    enA

    caM

    rPB

    2

    P. a

    trola

    zulin

    umD

    TO 1

    85-D

    1 =

    CV

    777

    Bra

    ct fr

    om P

    rote

    a re

    pens

    infru

    ctes

    cens

    , Ste

    llenb

    osch

    , Sou

    th A

    frica

    n.a.

    JX14

    1106

    JX15

    7458

    KP

    0645

    99

    P. a

    trola

    zulin

    umD

    TO 1

    86-D

    2 =

    CV

    173

    6M

    ite fr

    om P

    rote

    a re

    pens

    infru

    ctes

    cens

    , Stru

    isba

    ai, S

    outh

    Afri

    can.

    a.JX

    1411

    12JX

    1574

    98K

    P06

    4600

    P. a

    trola

    zulin

    umD

    TO 1

    86-H

    9 =

    CV

    222

    2B

    ract

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, S

    truis

    baai

    , Sou

    th A

    frica

    n.a.

    JX14

    1119

    JX15

    7513

    KP

    0646

    05

    P. a

    trosa

    ngui

    neum

    CB

    S 3

    80.7

    5 =

    FRR

    172

    6 =

    IMI 1

    9748

    8Tr

    iticu

    m a

    estiv

    um in

    silo

    , Pra

    gue,

    Cze

    ch R

    epub

    lic; e

    x-ty

    peJN

    6177

    06K

    J834

    435

    KP

    0167

    71JN

    4065

    57

    P. b

    urge

    nse

    CB

    S 3

    25.8

    9S

    oil,

    Bur

    gos,

    Spa

    in; e

    x-ty

    peK

    C41

    1736

    KJ8

    3443

    7K

    P01

    6772

    JN40

    6572

    P. c

    anis

    NR

    RL

    6279

    8Ili

    al b

    one

    lesi

    on in

    Rho

    desi

    an ri

    dgeb

    ack

    dog,

    US

    A; e

    x-ty

    peK

    J511

    291

    KF9

    0016

    7K

    F900

    177

    KF9

    0019

    6

    P. c

    aten

    atum

    CB

    S 3

    52.6

    7 =

    ATC

    C 1

    8543

    = C

    SIR

    109

    7 =

    IFO

    317

    74

    = IM

    I 136

    241

    Des

    ert s

    oil,

    Upi

    ngto

    n, S

    outh

    Afri

    ca; e

    x-ty

    peK

    C41

    1754

    KJ8

    3443

    8K

    P01

    6774

    JN12

    1504

    P. c

    f res

    trict

    umC

    BS

    139

    137

    = D

    AO

    M 2

    4105

    0 =

    DTO

    180

    -I4 =

    CV

    90

    Soi

    l, S

    telle

    nbos

    ch, S

    outh

    Afri

    caJX

    1409

    28JX

    1410

    54JX

    1574

    17K

    P06

    4643

    P. c

    f res

    trict

    umC

    BS

    139

    139

    = D

    AO

    M 2

    4105

    4 =

    DTO

    180

    -I6 =

    CV

    93

    Soi

    l, S

    telle

    nbos

    ch, S

    outh

    Afri

    caJX

    1409

    30JX

    1410

    55JX

    1574

    19K

    P06

    4607

    P. c

    f res

    trict

    umC

    BS

    139

    148

    = D

    AO

    M 2

    4105

    7 =

    DTO

    182

    -B6

    = C

    V

    872

    Soi

    l, M

    alm

    esbu

    ry, S

    outh

    Afri

    caK

    P01

    6817

    JX14

    1059

    JX15

    7464

    KP

    0646

    61

    P. c

    f res

    trict

    umC

    BS

    139

    149

    = D

    AO

    M 2

    4105

    2 =

    DTO

    182

    -C3

    = C

    V

    896

    Soi

    l, M

    alm

    esbu

    ry, S

    outh

    Afri

    caK

    P01

    6819

    JX14

    1060

    JX15

    7467

    KP

    0646

    09

    P. c

    f res

    trict

    umC

    BS

    139

    150

    = D

    AO

    M 2

    4105

    5 =

    DTO

    182

    -C5

    = C

    V

    900

    Soi

    l, M

    alm

    esbu

    ry, S

    outh

    Afri

    caK

    P01

    6816

    JX14

    1061

    JX15

    7468

    KP

    0646

    10

    P. c

    f res

    trict

    umC

    BS

    139

    152

    = D

    AO

    M 2

    4105

    8 =

    DTO

    182

    -E1

    = C

    V

    943

    Soi

    l, M

    alm

    esbu

    ry, S

    outh

    Afri

    can.

    a.JX

    1410

    63JX

    1574

    73K

    P06

    4611

    P. c

    f res

    trict

    umC

    BS

    139

    153

    = D

    AO

    M 2

    4105

    1 =

    DTO

    182

    -E3

    = C

    V

    948

    Soi

    l, M

    alm

    esbu

    ry, S

    outh

    Afri

    caK

    P01

    6818

    JX14

    1064

    JX15

    7474

    KP

    0646

    12

    P. c

    f res

    trict

    umC

    BS

    139

    170

    = D

    TO 1

    84-E

    4 =

    CV

    96

    Soi

    l, S

    telle

    nbos

    ch, S

    outh

    Afri

    can.

    a.JX

    1410

    56JX

    1574

    20K

    P06

    4644

    P. c

    f res

    trict

    umC

    BS

    139

    171

    = D

    TO 1

    84-E

    6 =

    CV

    101

    Soi

    l, S

    telle

    nbos

    ch, S

    outh

    Afri

    can.

    a.JX

    1410

    57JX

    1574

    21K

    P06

    4608

    P. c

    hala

    buda

    eC

    BS

    219

    .66

    = AT

    CC

    183

    22 =

    ATC

    C 1

    8329

    = F

    RR

    33

    93 =

    VK

    MF-

    1037

    Soi

    l, U

    krai

    ne; e

    x-ty

    peK

    P01

    6811

    KP

    0167

    48K

    P01

    6767

    KP

    0645

    72

    P. c

    iner

    asce

    nsN

    RR

    L 74

    8 =

    ATC

    C 4

    8693

    = B

    IOU

    RG

    E 9

    0 =

    FRR

    748

    =

    IMI 9

    2234

    = Q

    M 7

    555

    = Th

    om 4

    733.

    34U

    nkno

    wn;

    ex-

    type

    AF0

    3345

    5JX

    1410

    41JX

    1574

    05K

    P06

    4614

    P. c

    iner

    eoat

    rum

    CB

    S 2

    22.6

    6 =

    ATC

    C 2

    2350

    = F

    RR

    339

    0 =

    IJFM

    502

    4 =

    IMI 1

    1367

    6 =

    VK

    MF-

    856

    Fore

    st s

    oil,

    Kie

    v, U

    krai

    ne; e

    x-ty

    peK

    C41

    1700

    KJ8

    3444

    2K

    P12

    5335

    JN40

    6608

    P. c

    itreo

    nigr

    umC

    BS

    258

    .29

    = AT

    CC

    487

    36 =

    FR

    R 7

    61 =

    IMI 0

    9220

    9 =

    LSH

    BP

    20 =

    LS

    HB

    P 98

    = M

    UC

    L 28

    648

    = M

    UC

    L 29

    062

    = M

    UC

    L 29

    116

    = N

    RR

    L 76

    1

    Rot

    ting

    stem

    , Bel

    gium

    ; ex-

    type

    of P

    . citr

    eoni

    grum

    and

    P.

    subc

    inea

    rum

    AF0

    3345

    6E

    F198

    621

    EF1

    9862

    8JN

    1214

    74

    P. c

    itreo

    nigr

    umC

    BS

    308

    .48

    = AT

    CC

    104

    25 =

    FR

    R 2

    046

    = IM

    I 040

    575

    = N

    RR

    L 20

    46M

    ilita

    ry e

    quip

    men

    t, Fl

    orid

    a, U

    SA

    ; ex-

    type

    of P

    . citr

    eovi

    ride

    EF1

    9864

    7E

    F198

    623

    KP

    0167

    78K

    P06

    4616

    P. c

    itreo

    nigr

    umC

    BS

    321

    .59

    = AT

    CC

    183

    08 =

    FAT

    123

    = IF

    O 6

    224

    = IM

    I 06

    8225

    = N

    RR

    L 34

    04 =

    QM

    729

    0S

    oil,

    Japa

    n; e

    x-ty

    pe o

    f P. a

    eneu

    mK

    P01

    6812

    KP

    0167

    49K

    P01

    6769

    KP

    0645

    73

    P. c

    itreo

    nigr

    umN

    RR

    L 11

    87U

    nkno

    wn

    EF1

    9864

    6E

    F198

    622

    EF1

    9862

    6E

    F198

    501

  • Penicillium sect. ExilicaulisARTIC

    LE

    79v o l u m e 7 n o . 1

    tabl

    e 1.

    (Con

    tinue

    d).

    gen

    Ban

    k ac

    cess

    ion

    num

    bers

    Spec

    ies

    nam

    ec

    olle

    ctio

    n nu

    mbe

    rsSu

    bstr

    ate,

    loc

    atio

    nIt

    SB

    enA

    caM

    rPB

    2

    P. c

    itreo

    sulfu

    ratu

    mC

    BS

    139

    135

    = D

    TO 1

    80-G

    1 =

    CV

    11

    Air

    sam

    ple,

    Ste

    llenb

    osch

    , Sou

    th A

    frica

    JX14

    0937

    JX14

    1162

    JX15

    7415

    KP

    0646

    75

    P. c

    itreo

    sulfu

    ratu

    mC

    BS

    139

    140

    = D

    TO 1

    81-E

    1 =

    CV

    283

    Bra

    ct fr

    om P

    rote

    a re

    pens

    infru

    ctes

    cens

    , Ste

    llenb

    osch

    , Sou

    th A

    frica

    JX14

    0943

    JX14

    1163

    JX15

    7441

    KP

    0646

    76

    P. c

    itreo

    sulfu

    ratu

    mC

    BS

    139

    156

    = D

    TO 1

    82-G

    3 =

    CV

    100

    0S

    oil,

    Mal

    mes

    bury

    , Sou

    th A

    frica

    JX14

    0935

    JX14

    1164

    JX15

    7536

    KP

    0646

    77

    P. c

    itreo

    sulfu

    ratu

    mC

    BS

    139

    158

    = D

    TO 1

    83-B

    2 =

    CV

    145

    4B

    ract

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, M

    alm

    esbu

    ry, S

    outh

    Afri

    caJX

    1409

    39JX

    1411

    67JX

    1574

    85K

    P06

    4678

    P. c

    itreo

    sulfu

    ratu

    mC

    BS

    139

    162

    = D

    TO 1

    83-C

    5 =

    CV

    153

    2B

    ract

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, M

    alm

    esbu

    ry, S

    outh

    Afri

    caJX

    1409

    40JX

    1411

    68JX

    1574

    92K

    P06

    4679

    P. c

    itreo

    sulfu

    ratu

    mC

    BS

    139

    165

    = D

    TO 1

    83-G

    2 =

    CV

    201

    5B

    ract

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, S

    truis

    baai

    , Sou

    th A

    frica

    JX14

    0942

    JX14

    1169

    JX15

    7537

    KP

    0646

    80

    P. c

    itreo

    sulfu

    ratu

    mIM

    I 922

    28 =

    DTO

    290

    -I4U

    nkno

    wn;

    ex-

    type

    KP

    0168

    14K

    P01

    6753

    KP

    0167

    77K

    P06

    4615

    P. c

    itreo

    sulfu

    ratu

    mN

    RR

    L 31

    271

    Unk

    now

    n, S

    train

    of P

    . tox

    icar

    ium

    Miy

    ake

    fide

    Ser

    ra e

    t al.

    (200

    8)E

    F198

    660

    EF1

    9861

    4E

    F198

    641

    EF1

    9848

    6

    P. c

    itreo

    sulfu

    ratu

    mN

    RR

    L 61

    72U

    nkno

    wn,

    Stra

    in o

    f P. t

    oxic

    ariu

    m M

    iyak

    e fid

    e S

    erra

    et a

    l. (2

    008)

    EF1

    9865

    0E

    F198

    620

    EF1

    9863

    1E

    F198

    499

    P. c

    onso

    brin

    umC

    BS

    139

    141

    = D

    AO

    M 2

    4107

    1 =

    DTO

    181

    -F8

    = C

    V 4

    36M

    ite fr

    om P

    rote

    a re

    pens

    infru

    ctes

    cens

    , Ste

    llenb

    osch

    , Sou

    th A

    frica

    JX14

    0887

    JX14

    1134

    JX15

    7447

    KP

    0646

    18

    P. c

    onso

    brin

    umC

    BS

    139

    144

    = D

    AO

    M 2

    4107

    2 =

    DTO

    181

    -H9

    = C

    V

    547

    Soi

    l, S

    telle

    nbos

    ch, S

    outh

    Afri

    ca; e

    x-ty

    peJX

    1408

    88JX

    1411

    35JX

    1574

    53K

    P06

    4619

    P. c

    onso

    brin

    umC

    BS

    139

    147

    = D

    AO

    M 2

    4107

    3 =

    DTO

    182

    -B5

    = C

    V

    865

    Air

    sam

    ple,

    Mal

    mes

    bury

    , Sou

    th A

    frica

    JX14

    0889

    JX14

    1138

    JX15

    7463

    KP

    0646

    22

    P. c

    onso

    brin

    umC

    BS

    139

    151

    = D

    AO

    M 2

    4107

    4 =

    DTO

    182

    -C9

    = C

    V

    911

    Soi

    l, M

    alm

    esbu

    ry, S

    outh

    Afri

    caJX

    1408

    90JX

    1411

    41JX

    1574

    69K

    P06

    4625

    P. c

    onso

    brin

    umC

    BS

    139

    155

    = D

    AO

    M 2

    4107

    5 =

    DTO

    182

    -F5

    = C

    V 9

    77S

    oil,

    Mal

    mes

    bury

    , Sou

    th A

    frica

    JX14

    0891

    JX14

    1144

    JX15

    7477

    KP

    0646

    28

    P. c

    onso

    brin

    umD

    TO 1

    85-D

    4 =

    CV

    815

    Air

    sam

    ple,

    Mal

    mes

    bury

    , Sou

    th A

    frica

    n.a.

    JX14

    1136

    JX15

    7461

    KP

    0646

    20

    P. c

    onso

    brin

    umD

    TO 1

    85-D

    5 =

    CV

    817

    Air

    sam

    ple,

    Mal

    mes

    bury

    , Sou

    th A

    frica

    n.a.

    JX14

    1137

    JX15

    7462

    KP

    0646

    21

    P. c

    onso

    brin

    umD

    TO 1

    85-E

    2 =

    CV

    888

    Soi

    l, M

    alm

    esbu

    ry, S

    outh

    Afri

    can.

    a.JX

    1411

    39JX

    1574

    65K

    P06

    4623

    P. c

    onso

    brin

    umD

    TO 1

    85-E

    5 =

    CV

    894

    Soi

    l, M

    alm

    esbu

    ry, S

    outh

    Afri

    can.

    a.JX

    1411

    40JX

    1574

    66K

    P06

    4624

    P. c

    onso

    brin

    umD

    TO 1

    85-F

    5 =

    CV

    917

    Soi

    l, M

    alm

    esbu

    ry, S

    outh

    Afri

    can.

    a.JX

    1411

    42JX

    1574

    70K

    P06

    4626

    P. c

    onso

    brin

    umD

    TO 1

    85-G

    3 =

    CV

    941

    Soi

    l, M

    alm

    esbu

    ry, S

    outh

    Afri

    can.

    a.JX

    1411

    43JX

    1574

    72K

    P06

    4627

    P. c

    onso

    brin

    umD

    TO 1

    85-I4

    = C

    V 1

    095

    Mite

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, M

    alm

    esbu

    ry, S

    outh

    Afri

    can.

    a.JX

    1411

    45JX

    1574

    83K

    P06

    4629

    P. c

    onso

    brin

    umD

    TO 1

    86-A

    6 =

    CV

    145

    7B

    ract

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, M

    alm

    esbu

    ry, S

    outh

    Afri

    can.

    a.JX

    1411

    46JX

    1574

    86K

    P06

    4630

    P. c

    oryl

    ophi

    lum

    CB

    S 1

    2780

    7 =

    IMI 9

    2263

    = N

    RR

    L 79

    3 =

    FRR

    793

    Unk

    now

    n; e

    x-ty

    pe o

    f P. o

    bscu

    rum

    KP

    0168

    15K

    P01

    6761

    KP

    0167

    97K

    P06

    4654

    P. c

    oryl

    ophi

    lum

    CB

    S 1

    2780

    8 =

    NR

    RL

    799

    = IM

    I 922

    02U

    nkno

    wn;

    ex-

    type

    of P

    . chl

    orol

    euco

    nK

    P01

    6813

    KP

    0167

    52K

    P01

    6776

    KP

    0646

    13

    P. c

    oryl

    ophi

    lum

    CB

    S 2

    31.3

    8 =

    ATC

    C 1

    0452

    = IF

    O 7

    726

    = IM

    I 039

    817

    = N

    RR

    L 87

    2H

    umul

    us lu

    pulu

    s, W

    eihe

    nste

    phan

    , Ger

    man

    y; e

    x-ty

    pe o

    f P. h

    umul

    iJN

    6176

    96K

    P01

    6756

    KP

    0167

    87K

    P06

    4645

    P. c

    oryl

    ophi

    lum

    CB

    S 2

    54.3

    7 =

    IMI 0

    9220

    1 =

    LSH

    BP

    12 =

    MU

    CL

    2866

    9 =

    NR

    RL

    769

    Unk

    now

    n; e

    x-ty

    pe o

    f P. c

    andi

    dofu

    lvum

    KC

    4117

    12K

    P01

    6751

    KP

    0167

    73K

    P06

    4606

  • Visagie et al.ARTICLE

    80 i m a f U N G U S

    tabl

    e 1.

    (Con

    tinue

    d).

    gen

    Ban

    k ac

    cess

    ion

    num

    bers

    Spec

    ies

    nam

    ec

    olle

    ctio

    n nu

    mbe

    rsSu

    bstr

    ate,

    loc

    atio

    nIt

    SB

    enA

    caM

    rPB

    2

    P. c

    oryl

    ophi

    lum

    CB

    S 2

    59.6

    7 =

    ATC

    C 2

    2351

    = F

    RR

    340

    3 =

    IFO

    914

    6 =

    IMI 1

    0314

    8Li

    tter o

    f Bet

    ula,

    UK

    ; ex-

    type

    of P

    . coe

    rule

    oviri

    deK

    C41

    1717

    KP

    0167

    54K

    P01

    6779

    KP

    0646

    17

    P. c

    oryl

    ophi

    lum

    CB

    S 3

    12.4

    8 =

    TCC

    9784

    = A

    THU

    M28

    90 =

    CE

    CT

    2270

    =

    FRR

    802

    = IM

    I 039

    754

    = M

    UC

    L 28

    671

    = M

    UC

    L 29

    073

    = M

    UC

    L 29

    131

    = N

    RR

    L 80

    2 =

    QM

    751

    0

    Unk

    now

    n; e

    x-ty

    pe o

    f P. c

    oryl

    ophi

    lum

    and

    P. c

    itreo

    vire

    nsA

    F033

    450

    JX14

    1042

    KP

    0167

    80K

    P06

    4631

    P. c

    oryl

    ophi

    lum

    CB

    S 3

    30.7

    9 =

    IJFM

    514

    7A

    ir sa

    mpl

    e, B

    arce

    lona

    , Spa

    in; a

    uthe

    ntic

    P. c

    itreo

    vire

    ns s

    train

    GU

    9445

    57G

    U94

    4519

    GU

    9446

    07JN

    4065

    69

    P. c

    oryl

    ophi

    lum

    DTO

    180

    -E4

    = C

    V 2

    853

    Soi

    l, M

    alm

    esbu

    ry, S

    outh

    Afri

    caFJ

    2309

    97JX

    1410

    44JX

    1575

    23K

    P06

    4633

    P. c

    oryl

    ophi

    lum

    DTO

    180

    -E5

    = C

    V 2

    854

    Soi

    l, M

    alm

    esbu

    ry, S

    outh

    Afri

    caFJ

    2309

    96JX

    1410

    45JX

    1575

    22K

    P06

    4634

    P. c

    oryl

    ophi

    lum

    DTO

    184

    -C9

    = C

    V 2

    852

    Soi

    l, M

    alm

    esbu

    ry, S

    outh

    Afri

    caFJ

    2309

    98JX

    1410

    43JX

    1575

    21K

    P06

    4632

    P. c

    rave

    nian

    umC

    BS

    139

    138

    = D

    AO

    M 2

    4108

    2 =

    DTO

    180

    -I5 =

    CV

    92

    Soi

    l, S

    telle

    nbos

    ch, S

    outh

    Afri

    ca; e

    x-ty

    peJX

    1409

    00JX

    1410

    76JX

    1574

    18K

    P06

    4636

    P. d

    ecum

    bens

    CB

    S 2

    30.8

    1 =

    FRR

    741

    = IM

    I 190

    875

    = M

    UC

    L 29

    107

    = N

    RR

    L 74

    1U

    nkno

    wn

    sour

    ce, F

    lorid

    a, U

    SA

    ; ex-

    type

    AY15

    7490

    KJ8

    3444

    6K

    P01

    6782

    JN40

    6601

    P.

    dim

    orph

    ospo

    rum

    CB

    S 4

    56.7

    0 =

    NR

    RL

    5207

    = A

    TCC

    227

    83 =

    ATC

    C

    5250

    1 =

    FRR

    112

    0 =

    IMI 1

    4968

    0M

    angr

    ove

    swam

    p so

    il, T

    oora

    ddin

    , Aus

    tralia

    ; ex-

    type

    AF0

    8180

    4K

    J834

    448

    KP

    0167

    83JN

    1215

    17

    P. d

    ravu

    niB

    PI 8

    4424

    8G

    reat

    Ast

    rola

    be R

    eed

    from

    Dic

    tosp

    haer

    ia v

    ersl

    uyii,

    Dra

    vuni

    , Fi

    ji; e

    x-ty

    peAY

    4948

    56n.

    a.n.

    a.n.

    a.

    P. e

    rube

    scen

    sC

    BS

    318

    .67

    = AT

    CC

    185

    44 =

    CS

    IR 1

    040

    = FR

    R 8

    14 =

    IF

    O 3

    1734

    = IM

    I 136

    204

    = N

    RR

    L 62

    23N

    urse

    ry s

    oil,

    Pre

    toria

    , Sou

    th A

    frica

    ; ex-

    type

    AF0

    3346

    4H

    Q64

    6566

    EU

    4272

    81JN

    1214

    90

    P. fa

    giC

    BS

    689

    .77

    = C

    CM

    F-69

    6 =

    IJFM

    304

    9 =

    IMI 2

    5380

    6 =

    VK

    MF-

    2178

    Falle

    n le

    af (F

    agus

    syl

    vatic

    a) o

    n A

    ndos

    ol s

    oil,

    Spa

    in; e

    x-ty

    peA

    F481

    124

    KJ8

    3444

    9K

    P01

    6784

    JN40

    6540

    P. g

    riseo

    lum

    CB

    S 2

    77.5

    8 =

    ATC

    C 1

    8239

    = F

    RR

    267

    1 =

    IFO

    817

    5 =

    IMI 0

    7162

    6 =

    LSH

    BB

    323

    = N

    RR

    L 26

    71 =

    QM

    752

    3A

    cidi

    c du

    ne s

    and,

    Dor

    set,

    Eng

    land

    ; ex-

    type

    EF4

    2284

    8E

    F506

    213

    EF5

    0623

    2K

    P06

    4638

    P. g

    uttu

    losu

    mC

    BS

    141

    171

    = N

    RR

    L 90

    7 =

    ATC

    C 4

    8734

    = F

    RR

    907

    =

    Thom

    489

    4.16

    Soi

    l, U

    tah,

    US

    A; e

    x-ty

    peH

    Q64

    6592

    HQ

    6465

    76H

    Q64

    6587

    KP

    0646

    39

    P. h

    emitr

    achu

    mC

    BS

    139

    133

    = D

    AO

    M 2

    4109

    7 =

    DTO

    180

    -D7

    = C

    V

    2844

    Soi

    l, M

    alm

    esbu

    ry, S

    outh

    Afri

    caFJ

    2310

    02JX

    1410

    47JX

    1575

    25K

    P06

    4641

    P. h

    emitr

    achu

    mC

    BS

    139

    134

    = D

    AO

    M 2

    4109

    8 =

    DTO

    180

    -D8

    = C

    V

    2845

    Soi

    l, M

    alm

    esbu

    ry, S

    outh

    Afri

    ca; e

    x-ty

    peFJ

    2310

    03JX

    1410

    48JX

    1575

    26K

    P06

    4642

    P. h

    emitr

    achu

    mC

    BS

    139

    154

    = D

    AO

    M 2

    4109

    9 =

    DTO

    182

    -F1

    = C

    V 9

    64S

    oil,

    Mal

    mes

    bury

    , Sou

    th A

    frica

    JX14

    0916

    JX14

    1046

    JX15

    7475

    KP

    0646

    40

    P. h

    eter

    omor

    phum

    CB

    S 2

    26.8

    9S

    oil,

    She

    nnon

    gjia

    , Chi

    na; e

    x-ty

    peK

    C41

    1702

    KJ8

    3445

    5K

    P01

    6786

    JN40

    6605

    P. k

    atan

    gens

    eC

    BS

    247

    .67

    = AT

    CC

    183

    88 =

    IMI 1

    3620

    6 =

    NR

    RL

    5182

    Soi

    l, K

    atan

    ga, D

    R C

    ongo

    ; ex-

    type

    AF0

    3345

    8K

    P01

    6757

    KP

    0167

    88K

    P06

    4646

    P. k

    urss

    anov

    iiC

    BS

    625

    .67

    = AT

    CC

    183

    87 =

    FR

    R 3

    381

    = IJ

    FM 5

    045

    = IM

    I 129

    965

    = N

    RR

    L 33

    81 =

    VK

    MF-

    1244

    Mai

    ze-fi

    eld

    soil,

    Zp

    oro

    sk

    dist

    rict,

    Ukr

    aine

    ; ex-

    type

    EF4

    2284

    9K

    P01

    6758

    KP

    0167

    89K

    P06

    4647

    P. la

    eve

    CB

    S 1

    3666

    5 =

    KY

    1272

    7 =

    NB

    RC

    109

    724

    Fore

    st s

    oil,

    Thai

    land

    ; ex-

    type

    KF6

    6736

    9K

    F667

    365

    KF6

    6736

    7K

    F667

    371

  • Penicillium sect. ExilicaulisARTIC

    LE

    81v o l u m e 7 n o . 1

    tabl

    e 1.

    (Con

    tinue

    d).

    gen

    Ban

    k ac

    cess

    ion

    num

    bers

    Spec

    ies

    nam

    ec

    olle

    ctio

    n nu

    mbe

    rsSu

    bstr

    ate,

    loc

    atio

    nIt

    SB

    enA

    caM

    rPB

    2

    P. la

    pido

    sum

    CB

    S 3

    43.4

    8 =

    ATC

    C 1

    0462

    = C

    CT4

    477

    = IF

    O 6

    100

    = IM

    I 039

    743

    = N

    RR

    L 71

    8 =

    QM

    192

    8C

    anne

    d bl

    uebe

    rry,

    Was

    hing

    ton,

    US

    A; e

    x-ty

    peA

    F033

    409

    KJ8

    3446

    5FJ

    5309

    84JN

    1215

    00

    P. m

    acle

    nnan

    iae

    CB

    S 1

    98.8

    1 =

    DA

    R 3

    5238

    Rhi

    zopl

    ane

    of G

    ahni

    a ra

    dula

    , Vic

    toria

    , Aus

    tralia

    ; ex-

    type

    KC

    4116

    89K

    J834

    468

    KP

    0167

    91K

    P06

    4648

    P. m

    elin

    iiC

    BS

    139

    142

    = D

    AO

    M 2

    4110

    0 =

    DTO

    181

    -H6

    = C

    V

    535

    Soi

    l, S

    telle

    nbos

    ch, S

    outh

    Afri

    caJX

    1409

    19JX

    1410

    50JX

    1574

    51K

    P06

    4650

    P. m

    elin

    iiC

    BS

    139

    143

    = D

    AO

    M 2

    4110

    1 =

    DTO

    181

    -H8

    = C

    V

    542

    Soi

    l, S

    telle

    nbos

    ch, S

    outh

    Afri

    caJX

    1409

    20JX

    1410

    51JX

    1574

    52K

    P06

    4651

    P. m

    elin

    iiC

    BS

    139

    168

    = D

    AO

    M 2

    4110

    2 =

    DTO

    183

    -H8

    = C

    V

    2393

    Bra

    ct fr

    om P

    rote

    a re

    pens

    infru

    ctes

    cens

    , Stru

    isba

    ai, S

    outh

    Afri

    caJX

    1409

    17JX

    1410

    52JX

    1575

    27K

    P06

    4652

    P. m

    elin

    iiC

    BS

    139

    169

    = D

    AO

    M 2

    4110

    3 =

    DTO

    183

    -I1 =

    CV

    24

    04B

    ract

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, S

    truis

    baai

    , Sou

    th A

    frica

    JX14

    0918

    JX14

    1053

    JX15

    7528

    KP

    0646

    53

    P. m

    elin

    iiC

    BS

    218

    .30

    = AT

    CC

    104

    69 =

    FR

    R 2

    041

    = IF

    O 7

    675

    = IM

    I 040

    216

    = M

    UC

    L 29

    235

    = N

    RR

    L 20

    41 =

    QM

    759

    9Fo

    rest

    soi

    l, U

    SA

    ; ex-

    type

    AF0

    3344

    9K

    J834

    471

    KP

    0167

    92JN

    4066

    13

    P. m

    elin

    iiC

    BS

    280

    .58

    = AT

    CC

    183

    83 =

    IMI 0

    7162

    4 =

    LSH

    BB

    333

    = N

    RR

    L 26

    72 =

    QM

    752

    6C

    allu

    na h

    eath

    land

    soi

    l, E

    ngla

    nd; e

    x-ty

    pe o

    f P. r

    adul

    atum

    KC

    4117

    27K

    P01

    6763

    KP

    0168

    01K

    P06

    4658

    P. m

    elin

    iiC

    BS

    285

    .65

    Pin

    us b

    anks

    iana

    , Pet

    awaw

    a, C

    anad

    an.

    a.K

    P01

    6764

    KP

    0168

    02K

    P06

    4659

    P. m

    elin

    iiC

    BS

    340

    .61

    Soi

    l, P

    adov

    a, It

    aly

    n.a.

    KP

    0167

    60K

    P01

    6793

    KP

    0646

    49

    P. m

    enon

    orum

    NR

    RL

    5041

    0G

    arde

    n so

    il, C

    alifo

    rnia

    , US

    A; e

    x-ty

    peH

    Q64

    6591

    HQ

    6465

    73H

    Q64

    6584

    KF9

    0019

    4

    P. m

    erid

    ianu

    mC

    BS

    314

    .67

    = AT

    CC

    185

    45 =

    CS

    IR 1

    052

    = IM

    I 136

    209

    Gra

    ssla

    nd s

    oil,

    Pre

    toria

    , Sou

    th A

    frica

    ; ex-

    type

    AF0

    3345

    1K

    J834

    472

    KP

    0167

    94JN

    4065

    76

    P. m

    omoi

    iC

    BS

    139

    157

    = D

    AO

    M 2

    4107

    7 =

    DTO

    182

    -G4

    = C

    V

    1015

    Mite

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, M

    alm

    esbu

    ry, S

    outh

    A

    frica

    ; ex-

    type

    JX14

    0895

    JX14

    1073

    JX15

    7479

    KP

    0646

    73

    P. n

    amys

    low

    skii

    CB

    S 2

    02.8

    7 =

    IJFM

    782

    4S

    and

    dune

    soi

    l, U

    tah,

    US

    A; e

    x-ty

    pe o

    f P. fl

    avid

    ostip

    itatu

    mK

    C41

    1691

    KJ8

    3445

    1K

    P01

    6785

    KP

    0646

    37

    P. n

    amys

    low

    skii

    CB

    S 2

    04.8

    7S

    andy

    soi

    l, C

    hile

    ; ex-

    type

    of P

    . tur

    ris-p

    aine

    nse

    KC

    4116

    93K

    P01

    6766

    KP

    0168

    09K

    P06

    4681

    P. n

    amys

    low

    skii

    CB

    S 3

    53.4

    8 =

    ATC

    C 1

    1127

    = IM

    I 040

    033

    = M

    UC

    L 29

    226

    = N

    RR

    L 10

    70S

    oil u

    nder

    Pin

    us, B

    elar

    us, P

    olan

    d; e

    x-ty

    peA

    F033

    463

    JX14

    1067

    KP

    0167

    95JF

    4174

    30

    P. n

    epal

    ense

    CB

    S 2

    03.8

    4 =

    NH

    L 64

    82R

    ice

    soil,

    Kat

    hman

    du, N

    epal

    ; ex-

    type

    KC

    4116

    92K

    J834

    474

    KP

    0167

    96JN

    1214

    53

    P. o

    vatu

    mC

    BS

    136

    664

    = K

    Y 12

    726

    Fore

    st s

    oil u

    nder

    litte

    r of P

    inus

    car

    ibae

    a, K

    uala

    Lum

    pur,

    Mal

    ayas

    ia; e

    x-ty

    peK

    F667

    370

    KF6

    6736

    6K

    F667

    368

    KF6

    6737

    2

    P. p

    agul

    umC

    BS

    139

    166

    = D

    AO

    M 2

    4106

    9 =

    DTO

    183

    -H2

    = C

    V

    2224

    Bra

    ct fr

    om P

    rote

    a re

    pens

    infru

    ctes

    cens

    , Stru

    isba

    ai, S

    outh

    A

    frica

    ; ex-

    type

    JX14

    0898

    JX14

    1070

    JX15

    7519

    KP

    0646

    55

    P. p

    agul

    umC

    BS

    139

    167

    = D

    AO

    M 2

    4107

    0 =

    DTO

    183

    -H3

    = C

    V

    2236

    Bra

    ct fr

    om P

    rote

    a re

    pens

    infru

    ctes

    cens

    , Stru

    isba

    ai, S

    outh

    Afri

    caJX

    1408

    99JX

    1410

    71JX

    1575

    20K

    P06

    4656

    P. p

    arvu

    mC

    BS

    359

    .48

    = AT

    CC

    104

    79 =

    IFO

    773

    2 =

    IMI 0

    4058

    7 =

    NR

    RL

    2095

    = Q

    M 1

    878

    Soi

    l, N

    icar

    agua

    ; ex-

    type

    AF0

    3346

    0H

    Q64

    6568

    KF9

    0017

    3JN

    4065

    59

    P. p

    arvu

    mC

    BS

    570

    .73

    = AT

    CC

    280

    50 =

    ATC

    C 4

    8363

    = F

    RR

    15

    59 =

    NH

    L 64

    63S

    oil,

    Por

    t Mor

    esby

    , Pap

    ua N

    ew G

    uine

    a; e

    x-ty

    pe P

    . pap

    uanu

    mK

    C41

    1767

    KP

    0167

    62K

    P01

    6798

    KP

    0646

    57

  • Visagie et al.ARTICLE

    82 i m a f U N G U S

    tabl

    e 1.

    (Con

    tinue

    d).

    gen

    Ban

    k ac

    cess

    ion

    num

    bers

    Spec

    ies

    nam

    ec

    olle

    ctio

    n nu

    mbe

    rsSu

    bstr

    ate,

    loc

    atio

    nIt

    SB

    enA

    caM

    rPB

    2

    P. p

    hilip

    pine

    nse

    CB

    S 6

    23.7

    2 =

    FRR

    153

    2 =

    NH

    L 61

    30S

    oil f

    rom

    pin

    e fo

    rest

    , Luz

    on Is

    land

    , Phi

    lippi

    nes;

    ex-

    type

    KC

    4117

    70K

    J834

    482

    KP

    0167

    99JN

    4065

    43

    P. p

    imite

    ouie

    nse

    CB

    S 1

    0247

    9 =

    NR

    RL

    2554

    2K

    idne

    y ep

    ithel

    ial c

    ell c

    ultu

    re fl

    ask,

    Pre

    oria

    , US

    A; e

    x-ty

    peA

    F037

    431

    HQ

    6465

    69H

    Q64

    6580

    JN40

    6650

    P. ra

    cibo

    rski

    iC

    BS

    224

    .28

    = AT

    CC

    104

    88 =

    DS

    M24

    22 =

    FR

    R 2

    150

    = IF

    O 7

    676

    = IM

    I 040

    568

    = LS

    HB

    P 92

    = M

    UC

    L 29

    246

    = N

    RR

    L 21

    50

    Soi

    l und

    er c

    onife

    r, D

    uga

    Go

    lina,

    Pol

    and;

    ex-

    type

    AF0

    3344

    7JX

    1410

    69K

    P01

    6800

    JN40

    6607

    P. re

    pens

    icol

    aC

    BS

    139

    159

    = D

    AO

    M 2

    4107

    9 =

    DTO

    183

    -B5

    = C

    V

    1486

    Bra

    ct fr

    om P

    rote

    a re

    pens

    infru

    ctes

    cens

    , Mal

    mes

    bury

    , Sou

    th A

    frica

    JX14

    0892

    JX14

    1149

    JX15

    7489

    KP

    0646

    66

    P. re

    pens

    icol

    aC

    BS

    139

    160

    = D

    AO

    M 2

    4108

    0 =

    DTO

    183

    -B8

    = C

    V

    1495

    Bra

    ct fr

    om P

    rote

    a re

    pens

    infru

    ctes

    cens

    , Mal

    mes

    bury

    , Sou

    th

    Afri

    ca; e

    x-ty

    peJX

    1408

    93JX

    1411

    50JX

    1574

    90K

    P06

    4660

    P. re

    pens

    icol

    aC

    BS

    139

    161

    = D

    AO

    M 2

    4108

    1 =

    DTO

    183

    -C2

    = C

    V

    1514

    Bra

    ct fr

    om P

    rote

    a re

    pens

    infru

    ctes

    cens

    , Mal

    mes

    bury

    , Sou

    th A

    frica

    JX14

    0894

    JX14

    1151

    JX15

    7491

    KP

    0646

    67

    P. re

    pens

    icol

    aC

    BS

    139

    175

    = D

    TO 1

    86-A

    8 =

    CV

    147

    9B

    ract

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, M

    alm

    esbu

    ry, S

    outh

    Afri

    caK

    P01

    6820

    JX14

    1147

    JX15

    7487

    KP

    0646

    64

    P. re

    pens

    icol

    aC

    BS

    139

    176

    = D

    TO 1

    86-A

    9 =

    CV

    148

    4B

    ract

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, M

    alm

    esbu

    ry, S

    outh

    Afri

    can.

    aJX

    1411

    48JX

    1574

    88K

    P06

    4665

    P. re

    pens

    icol

    aD

    TO 1

    86-B

    5 =

    CV

    154

    6B

    ract

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, M

    alm

    esbu

    ry, S

    outh

    Afri

    can.

    a.JX

    1411

    52JX

    1574

    93K

    P06

    4668

    P. re

    stric

    tum

    CB

    S 3

    67.4

    8 =

    ATC

    C 1

    1257

    = F

    RR

    174

    8 =

    IMI 0

    4022

    8 =

    NR

    RL

    1748

    = Q

    M 1

    962

    Soi

    l, H

    ondu

    ras;

    ex-

    type

    AF0

    3345

    7K

    J834

    486

    KP

    0168

    03JN

    1215

    06

    P. ru

    befa

    cien

    sC

    BS

    139

    145

    = D

    AO

    M 2

    4107

    6 =

    DTO

    181

    -I5 =

    CV

    597

    Mite

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, S

    telle

    nbos

    ch, S

    outh

    Afri

    caJX

    1408

    96JX

    1410

    72JX

    1574

    54K

    P06

    4662

    P. ru

    befa

    cien

    sC

    BS

    145

    .83

    = C

    EC

    T 27

    52S

    andy

    soi

    l und

    er P

    inus

    sp.

    , Val

    lado

    lid, S

    pain

    ; ex-

    type

    KC

    4116

    77K

    J834

    487

    KP

    0168

    04JN

    4066

    27

    P. ru

    bidu

    rum

    CB

    S 6

    09.7

    3 =

    NR

    RL

    6033

    = A

    TCC

    280

    51 =

    ATC

    C

    4823

    8 =

    FRR

    155

    8 =

    IMI 2

    2855

    1 =

    NH

    L 64

    60S

    oil,

    Wew

    ak, P

    apua

    New

    Gui

    nea;

    ex-

    type

    AF0

    3346

    2H

    Q64

    6574

    HQ

    6465

    85JN

    4065

    45

    P. s

    mith

    iiC

    BS

    256

    .87

    = FR

    R 2

    663

    = IM

    I 288

    724

    Drie

    d fis

    h (D

    ecap

    teru

    s sp

    .), In

    done

    sia;

    ex-

    type

    of P

    . cor

    ynep

    horu

    mK

    C41

    1716

    KP

    0167

    55K

    P01

    6781

    KP

    0646

    35

    P. s

    mith

    iiC

    BS

    261

    .87

    = FR

    R 2

    743

    Pas

    teur

    ised

    frui

    t jui

    ce, S

    ydne

    y, A

    ustra

    lia; e

    x-ty

    pe o

    f P. s

    abul

    osum

    KC

    4117

    19K

    P01

    6765

    KP

    0168

    05K

    P06

    4672

    P. s

    mith

    iiC

    BS

    276

    .83

    = C

    EC

    T 27

    44 =

    IMI 2

    5969

    3S

    ecal

    e ce

    real

    e, Z

    amor

    a, S

    pain

    ; ex-

    type

    KC

    4117

    23K

    J834

    492

    KP

    0168

    06JN

    4065

    89

    P. s

    triat

    ispo

    rum

    CB

    S 7

    05.6

    8 =

    ATC

    C 2

    2052

    = C

    CR

    C 3

    1679

    = F

    RR

    827

    =

    IMI 1

    5174

    9 =

    MU

    CL

    3120

    2Le

    af li

    tter o

    f Aca

    cia

    karr

    oo, P

    otch

    efst

    room

    , Sou

    th A

    frica

    ; ex-

    type

    AF0

    3893

    8JX

    1411

    56K

    P01

    6807

    JN40

    6538

    P. s

    ubtu

    rcos

    eum

    CB

    S 1

    3912

    9 =

    DA

    OM

    241

    093

    = D

    TO 1

    80-C

    5 =

    CV

    28

    17S

    oil,

    Mal

    mes

    bury

    , Sou

    th A

    frica

    FJ23

    1004

    JX14

    1158

    JX15

    7533

    KP

    0646

    69

    P. s

    ubtu

    rcos

    eum

    CB

    S 1

    3913

    0 =

    DA

    OM

    241

    094

    = D

    TO 1

    80-C

    6 =

    CV

    28

    20S

    oil,

    Mal

    mes

    bury

    , Sou

    th A

    frica

    FJ23

    1005

    JX14

    1159

    JX15

    7534

    KP

    0646

    70

    P. s

    ubtu

    rcos

    eum

    CB

    S 1

    3913

    1 =

    DA

    OM

    241

    095

    = D

    TO 1

    80-C

    7 =

    CV

    28

    26S

    oil,

    Mal

    mes

    bury

    , Sou

    th A

    frica

    FJ23

    1007

    JX14

    1160

    JX15

    7535

    KP

    0646

    71

    P. s

    ubtu

    rcos

    eum

    CB

    S 1

    3913

    2 =

    DA

    OM

    241

    096

    = D

    TO 1

    80-C

    9 =

    CV

    28

    35S

    oil,

    Mal

    mes

    bury

    , Sou

    th A

    frica

    ; ex-

    type

    FJ23

    1006

    JX14

    1161

    JX15

    7532

    KP

    0646

    74

    P. s

    ubtu

    rcos

    eum

    CB

    S 1

    3914

    6 =

    DA

    OM

    241

    078

    = D

    TO 1

    82-A

    7 =

    CV

    79

    5A

    ir sa

    mpl

    e, M

    alm

    esbu

    ry, S

    outh

    Afri

    caJX

    1409

    34JX

    1411

    57JX

    1574

    59K

    P06

    4663

    P. te

    rren

    umC

    BS

    254

    .87

    = FR

    R 2

    660

    Drie

    d fis

    h (D

    ecap

    teru

    s sp

    .), In

    done

    sia;

    ex-

    type

    of P

    . cha

    lybe

    umK

    C41

    1713

    KJ8

    3444

    0K

    P01

    6775

    JN40

    6596

  • Penicillium sect. ExilicaulisARTIC

    LE

    83v o l u m e 7 n o . 1

    tabl

    e 1.

    (Con

    tinue

    d).

    gen

    Ban

    k ac

    cess

    ion

    num

    bers

    Spec

    ies

    nam

    ec

    olle

    ctio

    n nu

    mbe

    rsSu

    bstr

    ate,

    loc

    atio

    nIt

    SB

    enA

    caM

    rPB

    2

    P. te

    rren

    umC

    BS

    313

    .67

    = AT

    CC

    185

    47 =

    CS

    IR 1

    022

    = IM

    I 136

    208

    Soi

    l, S

    outh

    Afri

    ca; e

    x-ty

    peA

    M99

    2111

    KJ8

    3449

    6K

    P01

    6808

    JN40

    6577

    P. te

    rren

    umC

    BS

    622

    .72

    = D

    SM

    2418

    = N

    HL

    6128

    = N

    RR

    L 58

    24Fo

    rest

    soi

    l und

    er P

    inus

    sp.

    , Luz

    on Is

    land

    , Phi

    lippi

    nes;

    ex-

    type

    of P

    . lu

    zoni

    acum

    AF0

    3344

    6K

    P01

    6759

    KP

    0167

    90JN

    4065

    44

    P. v

    elut

    inum

    CB

    S 2

    50.3

    2 =

    ATC

    C 1

    0510

    = C

    EC

    T 23

    18 =

    IJFM

    510

    8 =

    IMI 0

    4057

    1 =

    NR

    RL

    2069

    = Q

    M 7

    686

    = V

    KM

    F-37

    9S

    putu

    m fr

    om m

    an, A

    ppel

    door

    n, N

    ethe

    rland

    s; e

    x-ty

    peA

    F033

    448

    JX14

    1170

    KP

    0168

    10K

    P06

    4682

    P. v

    inac

    eum

    CB

    S 3

    89.4

    8 =

    ATC

    C 1

    0514

    = F

    RR

    739

    = IM

    I 029

    189

    = N

    RR

    L 73

    9 =

    QM

    674

    6S

    oil,

    Uta

    h, U

    SA

    ; ex-

    type

    AF0

    3346

    1H

    Q64

    6575

    HQ

    6465

    86JN

    4065

    55

    P. x

    anth

    omel

    inii

    CB

    S 1

    3916

    3 =

    DA

    OM

    241

    104

    = D

    TO 1

    83-C

    7 =

    CV

    16

    77S

    oil,

    Stru

    isba

    ai, S

    outh

    Afri

    ca; e

    x-ty

    peJX

    1409

    21JX

    1411

    20JX

    1574

    95K

    P06

    4683

    P. x

    anth

    omel

    inii

    CB

    S 1

    3917

    9 =

    DTO

    186

    -F4

    = C

    V 1

    923

    Mite

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, S

    truis

    baai

    , Sou

    th A

    frica

    n.a.

    JX14

    1126

    JX15

    7509

    KP

    0646

    89

    P. x

    anth

    omel

    inii

    CB

    S 1

    3918

    0 =

    DTO

    186

    -F9

    = C

    V 1

    942

    Mite

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, S

    truis

    baai

    , Sou

    th A

    frica

    n.a.

    JX14

    1128

    JX15

    7510

    KP

    0646

    90

    P. x

    anth

    omel

    inii

    CB

    S 1

    3918

    1 =

    DTO

    186

    -G4

    = C

    V 1

    969

    Mite

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, S

    truis

    baai

    , Sou

    th A

    frica

    n.a.

    JX14

    1129

    JX15

    7511

    KP

    0646

    91

    P. x

    anth

    omel

    inii

    DA

    OM

    241

    105

    = D

    TO 1

    83-F

    6 =

    CV

    190

    5B

    ract

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, S

    truis

    baai

    , Sou

    th A

    frica

    JX14

    0924

    JX14

    1125

    JX15

    7508

    KP

    0646

    88

    P. x

    anth

    omel

    inii

    DTO

    183

    -D3

    = C

    V 1

    745

    Mite

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, S

    truis

    baai

    , Sou

    th A

    frica

    JX14

    0922

    JX14

    1121

    JX15

    7499

    KP

    0646

    84

    P. x

    anth

    omel

    inii

    DTO

    183

    -E7

    = C

    V 1

    844

    Mite

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, S

    truis

    baai

    , Sou

    th A

    frica

    JX14

    0923

    JX14

    1122

    JX15

    7504

    KP

    0646

    85

    P. x

    anth

    omel

    inii

    DTO

    186

    -E5

    = C

    V 1

    871

    Bra

    ct fr

    om P

    rote

    a re

    pens

    infru

    ctes

    cens

    , Stru

    isba

    ai, S

    outh

    Afri

    can.

    a.JX

    1411

    23JX

    1575

    06K

    P06

    4686

    P. x

    anth

    omel

    inii

    DTO

    186

    -E8

    = C

    V 1

    886

    Bra

    ct fr

    om P

    rote

    a re

    pens

    infru

    ctes

    cens

    , Stru

    isba

    ai, S

    outh

    Afri

    can.

    a.JX

    1411

    24JX

    1575

    07K

    P06

    4687

    P. x

    anth

    omel

    inii

    DTO

    186

    -I7 =

    CV

    231

    2B

    ract

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, S

    truis

    baai

    , Sou

    th A

    frica

    n.a.

    JX14

    1130

    JX15

    7514

    KP

    0646

    92

    P. x

    anth

    omel

    inii

    DTO

    186

    -I9 =

    CV

    232

    9B

    ract

    from

    Pro

    tea

    repe

    ns in

    fruct

    esce

    ns, S

    truis

    baai

    , Sou

    th A

    frica

    n.a.

    JX14

    1131

    JX15

    7515

    KP

    0646

    93

  • Visagie et al.ARTICLE

    84 i m a f U N G U S

    P. atrolazulinum (321 total, six each from air and soil) in Protea repens infructescences and from their mite communities was especially apparent.

    PhylogenyAn ITS phylogeny (Fig. 1) was used to compare all ex-type strains from section Exilicaulis species and sort them into smaller clades for the BenA, CaM, RPB2 and combined phylogenies (Figs 26). The aligned ITS barcode data set was 526 bp long with K2+G+I and GTR+G+I the most suitable substitution models for ML and BI respectively. Table 2 summarises the most suitable substitution models used for the remaining phylogenies. The ITS phylogeny distributed strains into six main clades, similar to the results of Houbraken & Samson (2011). Species with biverticillate conidiophores were resolved in a well-supported clade distinct from monoverticillate species. We also divided the biverticillate clade into subclades, referred to here as the P. melinii- and P. corylophilum-clades, to present the data concisely. The monoverticillate species resolved in four clades, referred to here as the P. restrictum-, P. citreonigrum-, P. decumbens-, and P. parvum-clades. ITS performs poorly as a DNA barcode marker in the P. melinii-, P. corylophilum-, and P. restrictum-clades, although it still can be used to identify the important P. corylophilum. On the other hand, ITS can successfully identify all species from the P. citreonigrum-, P. decumbens-, and P. parvum-clades.

    Penicillium melinii-clade (Fig. 2) Phylogenetic data divided the fynbos strains into three clades, two representing new species described below as P. hemitrachum and P. xanthomelinii. The remaining strains were placed in a clade with P. melinii and P. radulatum. Although there is sequence variation within the clade, phylogenies were in conflict and following GCPSR we identified our strains as P. melinii and consider P. radulatum a synonym. Ex-type strains of P. terrenum, P. chalybeum, and P. luzoniacum were included

    in a coherent clade and we consider them to represent one species with P. terrenum the oldest name. Similarly, we consider P. flavidostipitatum and P. turris-painense synonyms of P. namyslowskii, while P. corynephorum and P. sabulosum are synonyms of P. smithii.

    Penicillium corylophilum-clade (Figs 34) The largest group of fynbos isolates belong to this clade, with nine species identified. GCPSR revealed seven coherent clades distinct from known species, which are introduced as new species below. Penicillium repensicola and P. subturcoseum strains were consistently resolved in coherent clades as close relatives. Penicillium momoii and P. cravenianum were represented by single isolates that shifted phylogenetic positions depending on the gene used. However, the combined phylogeny places P. cravenianum as a close relative to P. pagulum, while P. momoii seems most similar to P. rubefaciens. Sequence variation was observed in the clades described as P. atrolazulinum, P. consobrinum and P. pagulum. However, no coherent clades were observed within strains of P. atrolazulinum and P. consobrinum. For P. pagulum, only two strains were isolated and although they exhibited sequence variation, they were morphologically identical. Remaining fynbos isolates were identified as P. corylophilum and P. rubefaciens. Ex-type strains of P. candidofulvum, P. coeruleoviride, P. corylophilum, P. chloroleucon, P. citreovirens, P. humuli, and P. obscurum clustered together. Again, this clade displays sequence variation, but the only consistent branch is the one encompassing the entire clade. As a result, we accept P. corylophilum as the correct name for the clade and consider the remaining species synonyms.

    Penicillium citreonigrum-clade (Fig. 5) Based on the multigene analyses, we accept three species in this clade, namely P. citreonigrum, P. cinerascens and P. citreosulfuratum. Penicillium toxicarium (CBS 351.51) was recently shown to be synonymous with P. trzebinskii (Houbraken et al. 2014). On the other hand, Serra et al. (2008) pointed out that P.

    table 2. Overiview and details used for phylogenetic analyses.dataset

    P. citreonigrum clade P. corylophilum clade P. melinii clade P. parvum clade

    BenA dataset

    Length (bp) 449 420 454 427

    Substitution model (BI) K80+G SYM+G HKY+G SYM+G

    Substitution model (ML) K2+I K2+G K2+G K2+G

    CaM dataset

    Length (bp) 495 504 490 530

    Substitution model (BI) K80+I SYM+I GTR+G GTR+I+G

    Substitution model (ML) K2+G K2+G K2+G K2+G

    RPB2 dataset

    Length (bp) 877 695 838 888

    Substitution model (BI) GTR+I+G SYM+I+G SYM+G GTR+I+G

    Substitution model (ML) K2+G K2+G TN93+G K2+G

    Concatenated dataset

    Length (bp) 1821 1619 1782 1845

    Substitution model (BI) GTR+G SYM+I+G GTR+I+G GTR+I+G

    Substitution model (ML) K2+I TN93+G K2+G T92+I+G

  • Penicillium sect. ExilicaulisARTIC

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    P. corylophilum-clade

    P. restrictum-clade

    P. citreonigrum-clade

    P. decumbens-cladeP. parvum-clade

    P. melinii-cladeAF033449 P. melinii CBS 218.30TKC411727 P. melinii CBS 280.58 (P. radulatumT)AF033409 P. lapidosum CBS 343.48TFJ231003 P. hemitrachum CBS 139134TJN617706 P. atrosanguineum CBS 380.75TKC411723 P. smithii CBS 276.83T

    KC411716 P. smithii CBS 256.87 (P. corynephorumT)KC411719 P. smithii CBS 261.87 (P. sabulosumT)

    KC411736 P. burgense CBS 325.89TKC411713 P. terrenum CBS 254.87 (P. chalybeumT)KC411691 P. namyslowskii CBS 202.87(P. flavidostipitatumT)KC411689 P. maclennaniae CBS 198.81TAF033463 P. namyslowskii CBS 353.48TAF033447 P. raciborskii CBS 224.28TKC411693 P. namyslowskii CBS 204.87 (P. turris-painenseT)AF033448 P. velutinum CBS 250.32TAF033446 P. terrenum CBS 622.72 (P. luzoniacumT)AM992111 P. terrenum CBS 313.67T

    JX140921 P. xanthomelinii CBS 139163TJX140913 P. atrolazulinum CBS 139136T

    AF481124 P. fagi CBS 689.77TKC411677 P. rubefaciens CBS 145.83T

    JX140895 P. momoii CBS 139157TJX140893 P. repensicola CBS 139160TJX140898 P. pagulum CBS 139166TJX140900 P. cravenianum CBS 139138TJX140888 P. consobrinum CBS 139144TFJ231006 P. subturcoseum CBS 139132T

    GU944557 P. corylophilum CBS 330.79 (P. citreovirensAuthentic)KC411712 P. corylophilum CBS 254.37 (P. candidofulvumT)KP016813 P. corylophilum CBS 127808 (P. chloroleuconT)KC411717 P. corylophilum CBS 259.67 (P. coeruleovirideT)AF033450 P. corylophilum CBS 312.48TJN617696 P. corylophilum CBS 231.38 (P. humuliT)KP016815 P. corylophilum CBS 127807 (P. obscurumT)

    EF422849 P. kurssanovii CBS 625.67TAF033457 P. restrictum CBS 367.48T

    KC411700 P. cinereoatrum CBS 222.66TAF033458 P. katangense CBS 247.67T

    AF033451 P. meridianum CBS 314.67TKC411702 P. heteromorphum CBS 226.89T

    KC411770 P. philippinense CBS 623.72TKP016811 P. chalabudae CBS 219.66T

    KC411758 P. arabicum CBS 414.69TAF033455 P. cinerascens NRRL 748T

    EF198647 P. citreonigrum CBS 308.48 (P. citreovirideT)AF033456 P. citreonigrum CBS 258.29TKP016812 P. citreonigrum CBS 321.59 (P. aneumT)

    KP016814 P. citreosulfuratum IMI 92228TEF198660 P. citreosulfuratum NRRL 31271 (P. toxicarium fide Serra et al. 2008)

    AF033454 P. alutaceum CBS 317.67TAY157490 P. decumbens CBS 230.81T

    AY494856 P. dravuni F01V25TKF667369 P. laeve CBS 136665T

    KF667370 P. ovatum CBS 136664TEF422848 P. griseolum CBS 277.58T

    AF081804 P. dimorphosporum CBS 456.70TKJ511291 P. canis NRRL 62798T

    HQ646591 P. menonorum NRRL 50410TAF033464 P. erubescens CBS 318.67T

    AF038938 P. striatisporum CBS 705.68TKC411692 P. nepalense CBS 203.84T

    AF033462 P. rubidurum CBS 609.73TKC411767 P. parvum CBS 570.73 (P. papuanumT)AF033460 P. parvum CBS 359.48T

    KC411754 P. catenatum CBS 352.67TAF033461 P. vinaceum CBS 389.48T

    HQ646592 P. guttulosum NRRL 907TAF037431 P. pimiteouiense CBS 102479T

    KM189449 P. trzebinskii CBS 351.51 (P. toxicarium)

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    Fig. 1. Phylogenetic tree of Penicillium sect. Exilicaulis species ex-type strains using the ITS region. Penicillium toxicarium (CBS 351.51) was chosen as outgroup. Posterior probabilities (pp) and/or bootstrap values (bs) higher than 0.95 and 80 respectively, are given above thickened branches. Names in bold indicate fynbos strains, names in blue indicate new species strains, T = ex-type strain.

  • Visagie et al.ARTICLE

    86 i m a f U N G U S

    P. xanthomelinii CV 2312 P. xanthomelinii CV 2329 P. xanthomelinii CBS 139181 P. xanthomelinii CBS 139179P. xanthomelinii CV 1886 P. xanthomelinii CV 1871 P. xanthomelinii CV 1745 P. xanthomelinii CBS 139163TP. xanthomelinii CV 1844 P. xanthomelinii DAOM 241105 P. xanthomelinii CBS 139180

    P. melinii CBS 139142P. melinii CBS 139143

    P. melinii CBS 218.30TP. melinii CBS 139168P. melinii CBS 139169P. melinii CBS 340.61 P. melinii CBS 280.58 (P. radulatumT)P. melinii CBS 285.65

    P. smithii CBS 256.87 (P. corynephorumT)P. smithii CBS 261.87 (P. sabulosumT)

    P. smithii CBS 276.83TP. maclennaniae CBS 198.81T

    P. velutinum CBS 250.32TP. hemitrachum CBS 139154 P. hemitrachum CBS 139133P. hemitrachum CBS 139134T

    P. atrosanguineum CBS 380.75TP. raciborskii CBS 224.28T

    P. namyslowskii CBS 202.87 (P. flavidostipitatumT)P. namyslowskii CBS 353.48TP. namyslowskii CBS 204.87 (P. turris-painenseT)

    P. lapidosum CBS 343.48TP. burgense CBS 325.89TP. terrenum CBS 254.87 (P. chalybeumT)

    P. terrenum CBS 313.67TP. terrenum CBS 622.72 (P. luzoniacumT)

    P. restrictum CBS 367.48T

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    P. xanthomelinii CBS 139181P. xanthomelinii CV 2312P. xanthomelinii CBS 139180P. xanthomelinii CBS 139179P. xanthomelinii DAOM 241105P. xanthomelinii CV 1886P. xanthomelinii CV 1871P. xanthomelinii CV 1844P. xanthomelinii CV 1745P. xanthomelinii CBS 139163TP. xanthomelinii CV 2329P. melinii CBS 218.30TP. melinii CBS 139168

    P. melinii CBS 139169P. melinii CBS 139143

    P. melinii CBS 280.58 (P. radulatumT)P. melinii CBS 285.65P. melinii CBS 340.61P. melinii CBS 139142

    P. hemitrachum CBS 139154P. hemitrachum CBS 139134TP. hemitrachum CBS 139133

    P. maclennaniae CBS 198.81TP. smithii CBS 276.83T

    P. smithii CBS 256.87 (P. corynephorumT)P. smithii CBS 261.87 (P. sabulosumT)

    P. velutinum CBS 250.32TP. atrosanguineum CBS 380.75T

    P. raciborskii CBS 224.28TP. namyslowskii CBS 202.87 (P. flavidostipitatumT)P. namyslowskii CBS 204.87 (P. turris-painenseT)P. namyslowskii CBS 353.48T

    P. lapidosum CBS 343.48TP. burgense CBS 325.89T

    P. terrenum CBS 254.87 (P. chalybeumT)P. terrenum CBS 313.67T

    P. terrenum CBS 622.72 (P. luzoniacumT)P. restrictum CBS 367.48T

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    P. xanthomelinii CV 2312P. xanthomelinii CV 1871P. xanthomelinii CBS 139179P. xanthomelinii CBS 139181P. xanthomelinii CV 1844P. xanthomelinii CBS 139180P. xanthomelinii CBS 139163TP. xanthomelinii DAOM 241105P. xanthomelinii CV 1886P. xanthomelinii CV 1745P. xanthomelinii CV 2329

    P. melinii CBS 139142P. melinii CBS 139143

    P. melinii CBS 218.30TP. melinii CBS 139168P. melinii CBS 139169P. melinii CBS 340.61P. melinii CBS 280.58 (P. radulatumT)P. melinii CBS 285.65

    P. smithii CBS 256.87 (P. corynephorumT)P. smithii CBS 261.87 (P. sabulosumT)

    P. smithii CBS 276.83TP. maclennaniae CBS 198.81T

    P. velutinum CBS 250.32TP. hemitrachum CBS 139134TP. hemitrachum CBS 139154P. hemitrachum CBS 139133

    P. atrosanguineum CBS 380.75TP. raciborskii CBS 224.28T

    P. namyslowskii CBS 353.48TP. namyslowskii CBS 202.87 (P. flavidostipitatumT)P. namyslowskii CBS 204.87 (P. turris-painenseT)

    P. lapidosum CBS 343.48TP. burgense CBS 325.89T

    P. terrenum CBS 254.87 (P. chalybeumT)P. terrenum CBS 622.72 (P. luzoniacumT)

    P. terrenum CBS 313.67TP. restrictum CBS 367.48T

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    P. xanthomelinii CV 1871P. xanthomelinii CBS 139163TP. xanthomelinii CBS 139180P. xanthomelinii CV 2312P. xanthomelinii CV 1844P. xanthomelinii CBS 139179P. xanthomelinii CBS 139181P. xanthomelinii CV 1886P. xanthomelinii CV 1745P. xanthomelinii CV 2329

    P. xanthomelinii DAOM 241105P. melinii CBS 139142P. melinii CBS 139143

    P. melinii CBS 218.30TP. melinii CBS 139168P. melinii CBS 139169

    P. melinii CBS 285.65P. melinii CBS 340.61P. melinii CBS 280.58 (P. radulatumT)

    P. smithii CBS 256.87 (P. corynephorumT)P. smithii CBS 261.87 (P. sabulosumT)

    P. smithii CBS 276.83TP. maclennaniae CBS 198.81T

    P. velutinum CBS 250.32TP. hemitrachum CBS 139134TP. hemitrachum CBS 139154P. hemitrachum CBS 139133

    P. namyslowskii CBS 353.48TP. namyslowskii CBS 204.87 (P. turris-painenseT)P. namyslowskii CBS 202.87 (P. flavidostipitatumT)

    P. lapidosum CBS 343.48TP. burgense CBS 325.89T

    P. terrenum CBS 254.87 (P. chalybeumT)P. terrenum CBS 622.72 (P. luzoniacumT)

    P. terrenum CBS 313.67TP. atrosanguineum CBS 380.75T

    P. raciborskii CBS 224.28TP. restrictum CBS 367.48T

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    RPB2 BenA|CaM|RPB2

    CaM

    Fig. 2. Phylogenetic tree of Penicillium se