ARTICLE 59 et et al. et al species et al .//7 ( 9 et al .//. F Versicolores " Key words: Aspergillus amoenus Aspergillus austroafricanus Aspergillus creber Aspergillus cvjetkovicii Aspergillus fructus Aspergillus jensenii Aspergillus protuberus Aspergillus puulaauensis Aspergillus subversicolor Aspergillus sydowii Aspergillus tabacinus Aspergillus tennesseensis Aspergillus venenatus Aspergillus versicolor concordance analysis phylogeny systematics et al ./// 0 .//G
is the most widely reported and et al. ./66 # IMA FUNGUS · VOLUME 3 · NO 1: 59–79 NO 1: 3 · · VOLUME FUNGUS IMA et al .//. % et al. 67Q/ # J et al .//7 % and A. venenatus . PCR primer pairs K .//. ; 9 ! 67VR & 9 E , Aspergillus versicolor % # ? 9 F 67QV & 6 ( 9 P .//7 K =( al .//G ? .//Q studied species in section Versicolores < # = 67Q/ B # 9 J ./6/ 67.V B 9 P 67QR N0F= et al. ./6/ F > .//R .//G ( N et al. .//6 J = N P .//7 ? 9 F ./6/ " < \ < G Aspergillus austroafricanus, A. creber, A. A. creber, Aspergillus austroafricanus, A. sydowii, A. tabacinus and A. A. sydowii, versicolor
K Versicolores Versicolores & versicolor. L P . Using scanning electron
: nine new species and multilocus DNA DNA and multilocus new species : nine Versicolores was originally erected as the Aspergillus ' protein Tsr1 processing , and pre-rRNA in sick building syndrome studies have a positive reaction for all of the newly in sick building syndrome studies have a positive Mcm7 A. subversicolor & 9 E 67.V section section Versicolores Versicolores
67QR < J L ( 6 section 6Q A. versicolor != = N0F ;0F . N0F
Gams et al. . 0 ( ; + F ; J # M J N F ( [ < R/7 N @F G7Q2.=/R/7 %?J+ F " .// ; 6G/ 0 E 0> /Q/WW ! ( ? ; M 0 E F M ; F ; characteristics and accepted the seven species previously # K 67Q7 ( .//Q 677G # surface ornamentation of most species of the section and removed seven species from section sequence species in the section based on multilocus DNA 62 J%? K 67Q7 < Article info: J 1 6W > ./6.X F 1 W > ./6.X ( 1 .6 > ./6. A. amoenus, A. protuberus, are described as new species and A. amoenus, of molecular and phenotypic differences are accepted as distinct species on the basis used to detect A. versicolor < numerous isolates belonging to numerous isolates belonging cvjetkovicii, A. fructus, A. jensenii, A. puulaauensis, A. subversicolor, A. tennesseensis A. subversicolor, A. puulaauensis, A. jensenii, A. fructus, cvjetkovicii, replication licensing factor replication licensing factor ; 9 ! R 67VR Aspergillus Abstract: © 2012 International Mycological Association © 2012 International Mycological conditions: distribute and transmit the work, under the following are free to share - to copy, You Attribution: INTRODUCTION Aspergillus Aspergillus versicolor G 6 . Aspergillus Aspergillus sequence based phylogeny sequence T > # Non-commercial: No derivative works: Any of the above conditions can be waived if you get For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Nothing in this license impairs or restricts the author’s moral rights. permission from the copyright holder. ' # & 9 ; 672R group J # M J N F 6Q6R 0 M # J ( " V6V/2 MJFX 1 J ; 9 ! Aspergillus 67VR # Z # VOLUME 3 · NO. 1 16/ RR7Q3 ./6. /G /6 /W Jurjevic, Peterson & Horn
# ' N0F by Peterson et al ./6/ J V 7R E . N et al .//G Versicolores GR 7V E G/ X R6 E V/ X W. E The species described and accepted are monophyletic. V/ X < W. E R [
ARTICLE a higher annealing temperature was used to obtain single MATERIALS AND METHODS N0F ' #G 6 Fungal isolates F " GWG/ ' F & # < 133 3 ; ' = & 6 # J ' 133 F ; J # E E 0;;+ 3 E ' ( " 133 # E+MJ&F+L & et al. 6772 J ' @ Culture methods >0QRGW7QM>0QR26G6 %!VR.6WV %!VR.6WQ %!VR.6QRM E E < E F R %!VR.6QW %!VR.67V %!VR../G %!VR../7M%!VR..66 E .R E GW E < ?%F E ./J %!VR..62M%!VR..6V %!VR...V %!VR..V2 %!VR..VV ?2/ ?V/ .R E 6/ ( 67Q/ %!VR..WGM%!VR..WR %!VR..Q2 %!VR..76 %!VR..7WM K .//. ?2/ . ] < / R ] %!VR..77 %!VR.G/.M%!VR.G/2 %!VR.G62 %!VR.GR. < 2/ ] X ?V/ . ] < %!VR.GR2 %!VR.GV6M%!VR.GVG %!VR.GW. %!VR.GW7 / R ] < V/ ] E %!VR.GQRM%!VR.GQW %!VR.G7/M%!VR.G7. %!VR.2/. and appearance were recorded and photographs were made %!VR.22/ %!VR.22. %!VR.227M%!VR.2R6 %!VR.2V/ 6/= .R E E %!VR.2VW %!VR.2WGM%!VR.2WR %!VR.2WQM%!VR.2Q/ ; 676. %!VR.27/ >G/6QQ7M>G/6Q7V ;2R ( (FM( 2 / 6/ J .//G ! = Microscopy R/// ? < 00" / 6 ] & \=6// < R/// & < N"! was used as the starting point for parsimony analysis with F O P & ; gently pressing a ca ./ & R O P & ; W/ ] 6/// F + N? .R// ? G 6. P 9 ; ' .//6 ; ' 9 N"! # & P .//G & Mcm7, Tsr1 and RPB2 data sets J included only protein-coding sequences and each data set was F 0 6 . G & BT2 and CF J+; NW/ loci included protein-coding and intron regions and the data ( ? < F @&; ( ( .//G F ( = # # = DNA methods ? # ? E ?E?E E 6.R= R & 6/V # + % \ .R + < E < E .// .MG 9 J 677R .R E # concordance phylogenetic species recognition concepts of = N Taylor et al. ./// E E&F # 6 < X = centrifugation and DNA was precipitated from the aqueous . ' # & acids were collected by centrifugation, the pellet was rinsed N et W/ ] # al. .//G J ÉW/ ] 6// )+ É/ 7R N et al. .//G DNA was diluted ca 616// & A. versicolor in a PCR != BT2 CF N et al .//R "&J = N0F ID ;0F . RPB2 ' N0F Mcm7 = ;0F < < Tsr1 N et al .//R
60 IMA FUNGUS Aspergillus section Versicolores
Table 1. ( # ARTICLE NRRL number Provenance Aspergillus amoenus ? ? .R/VR2 ..V MJF1 < 676G .GV @ 1 ? < Berberis , 67G/ M. Roberg VRQ MK1 < 67.7 G. A. Ledingham 2QGQ %' # 0;;+ .GV # E # J 67V. <= GRV// MJF1 P K ( < Gandoderma australe .//R, D.T. Wicklow F=.G..Q " 1 K < 67WQ B. Muthappa Aspergillus asperescens J ? ? .7.QGR 2WW/ %<= = Aspergillus austroafricanus sp. nov., ? ? Q//R7W .GG J F 1 E 67.. V. A. Putterill <= Aspergillus creber sp. nov., ? ? Q//R7Q .G6 J F 1 E 67.. V. A. Putterill VR22 F [ 1 < \ 67W7 A. Wellman .RV.W > 1 " < 677V T. Goto RQRQG MJF1 ( # < .//Q Z. Jurjevic RQRQ2 MJF1 E < .//Q Z. Jurjevic RQRQW MJF1 E < .//Q Z. Jurjevic RQR7. MJF1 E < .//Q Z. Jurjevic <= RQR7W MJF1 0 > < .//Q Z. Jurjevic RQV/6 MJF1 0 > 1 .//7 Z. Jurjevic RQV/V MJF1 ( # < .//7 Z. Jurjevic RQV/W MJF1 ( # < .//7 Z. Jurjevic RQV6. MJF1 0 > < .//7 Z. Jurjevic RQVW/ MJF1 0 > < .//7 Z. Jurjevic RQVW. MJF1 @ < .//7 Z. Jurjevic RQVWG MJF1 @ < .//7 Z. Jurjevic RQVWR MJF1 [ < .//7 Z. Jurjevic Aspergillus cvjetkovicii sp. nov., ? ? Q//R77 ..W MJF1 0 > < 676R G.W. Wilson <= .G/ E 1 < 676W Round 2V2. M 1 0;;+ 67V7 D. I. Fennell L 2V2. RQR7G MJF1 E < .//Q Z. Jurjevic Aspergillus fructus sp. nov., ? ? Q//V// .G7 MJF1 E < 67G7 Bliss <= .26 M 1 < 676V L. McCulloch Aspergillus jensenii sp. nov., ? ? Q//V/6 ..R MK1 676G E & Dade .GR MK1 + < 67G/ H. Raistrick .2/ MJF1 0 " < 6766 C. N. Jensen, E & L type strain of A. globosus
RQRQ. MJF1 ? < .//Q Z. Jurjevic RQV// MJF1 ? < .//Q Z. Jurjevic <= RQVW6 MJF1 ( # < .//7 Z. Jurjevic RQVW2 MJF1 [ < .//7 Z. Jurjevic Aspergillus multicolor J ? ? .7.Q27 2WWR %<= =
VOLUME 3 · NO. 1 61 Jurjevic, Peterson & Horn
Table 1. E NRRL number Provenance Aspergillus protuberus ? ? G.VVR/ VV6 MK1 < 67.7 G. A. Ledingham GR/R # < 67VQ <= ARTICLE RQV6G MJF1 0 > < .//7 Z. Jurjevic RQW2W MJF1 0 > < .//7 Z. Jurjevic RQW2Q MJF1 0 > < .//7 Z. Jurjevic RQ77/ MJF1 E < .//7 Z. Jurjevic RQ776 MJF1 E < .//7 Z. Jurjevic Aspergillus puulaauensis sp. nov., ? ? Q//V/. GRV26 MJF1 P ( R R P .// < .//G D. T. Wicklow <= RQV/. MJF1 L B < .//7 Z. Jurjevic V.6.2 MJF1 P < Inonotus .//G D. T. Wicklow V.R6V E 1 F < ca 677/ S. P. Abbot, ' # MF?P WVR6 Aspergillus subversicolor sp. nov., ? ? Q//V/G RQ777 " 1 K < 67W/ B. Muthappa <= Aspergillus sydowii ? ? .W7VGV .R/ M 1 67G/ E & M. Swift. .R2 MJF1 @ L 672/ M. M. Harris 2WVQ MJF1 E < 67V7 V.2R/ & 1 < 67WW E. G. Simmons Aspergillus tabacinus ? ? RG7R22 VR7 MK1 < 67.7 G. A. Ledingham 2W76 M 1 < 67G2 Y. Nakazawa <= R/G6 Unknown: type isolate of A. versicolor # magnus J # "![ 67V. V.2Q6 0 1 K < 67WW Aspergillus tennesseensis sp. nov., ? ? Q//V/2 ..7 M 1 E & 676W R. Thaxter .G2 MJF1 ? # < 67.W C. Thom 6G6R/ MJF1 & < < 67Q2 B. W. Horn <= 6G6R. MJF1 & < < 67Q2 B. W. Horn Aspergillus venenatus sp. nov., ? ? Q//V/R 6G62W MJF1 & 1 < 67Q2 B. W. Horn <= 6G62Q MJF1 & < < 67Q2 B. W. Horn 6G627 MJF1 & < < 67Q2 B. W. Horn V.2RW MJF1 ? < 67Q7 D. T. Wicklow Aspergillus versicolor ? ? 6W.6R7 .GQ MJF1 < 67GR V. K. Charles <= R.67 J F 1 ( # 67W/ J. P. van der Walt. 6G622 MJF1 & < < 67Q2 B. W. Horn 6G62R MJF1 & < < 67Q2 B. W. Horn 6G62V MJF1 & < < 67Q2 B. W. Horn Aspergillus species, undescribed RG/ % " 1 < 67GQ Shumard 6G6R6 MJF1 & 1 < < 67Q2 B. W. Horn
RESULTS N et al. .//G & ( .//Q A. versicolor, A. Phylogenic analysis of sequence data tabacinus, A. amoenus, A. protuberus and A. sydowii each J < # ! 6 !
62 IMA FUNGUS Aspergillus section Versicolores ARTICLE
Fig. 1. ( N0F ' & Versicolores contains three subclades, the A. versicolor subclade, the A. sydowii subclade and the A. subversicolor & É7/ ] É/ 7/ " 0;;+ 6G6R6 A. tennesseensis but may represent a distinct " 0;;+ RG/ A. amoenus
& = isolate lineages, A. subversicolor and A. austroafricanus, The section Versicolores clade contained three & ! 6 1 A. sydowii subclade containing = 0;;+ 6G6R6 0;;+ RG/ A. sydowii, A. creber, A. venenatus, A. tennesseensis, A.
VOLUME 3 · NO. 1 63 Jurjevic, Peterson & Horn ARTICLE
Fig. 2. Aspergillus amoenus 0;;+ 2QGQ 7 .R E 6/ E F E F # ?%F ?%F # J # 56/ 3 @ = 56/ 3
cvjetkovicii, A. jensenii and A. puulaauensisX A. versicolor ;2W < subclade containing A. versicolor, A. tabacinus, A. fructus, present in some isolates, faint reddish soluble pigment present A. protuberus, A. amoenus and A. austroafricanusX in some isolates, reverse mostly reddish brown hues, with some A. subversicolor subclade containing the single species E 6/ ?%F .R E ! A. subversicolor J = A. sydowii in . M .GMGG # the A. sydowii subclade, in the A. versicolor subclade or with shallow sulcations, colony center often with funicular hyphal in a distinct clade containing only A. sydowii ! J6MJR aggregates, sporulation in blue-green to gray-green shades, no J " < 0;;+ ..V # & Mcm7 locus from A. sydowii < # despite numerous attempts and thus A. sydowii does not " W E F R E ! JG J " " W E F GW E & ! 6 A. sydowii as a V member of the A. sydowii subclade with strong statistical Stipes ! . " R GRM 6//MV// M66// & . RM 2MW MQ É7/ ] ) vesicles 2M WM6W M.6 ) É/ 7/ conidial heads biseriate, metulae # 63G # GMV MQ & . RM2 / MR R ) phialides RM VMQ M 66 & .MG ) TAXONOMY conidia ! . . RMG R MR ) Previously described species 0;;+ GRV// 6.M.. 3 ?2/ Aspergillus amoenus ? ; Hedwigia 7016GQ 67G6 ? ? .R/VR2 Aspergillus protuberus ? =E# # 4 ! . M Mikrobiologija 51 667 67VQ ? ? G.VVR/ Type: Germany1 ? < Berberis 67G/ M. ! G M Roberg 0;;+ 2QGQN < Synonym: Aspergillus versicolor # protuberus ? = E# # 4 K Mycol Pap 1611 6G7 67Q7 Description: Colonies 6/ E F .R E ! . M .RM2/ Type: Yugoslavia: < ca GM2 0;;+ ..V 67VQ 0;;+ GR/RN < sporulating moderately to well, conidial heads in grayish green
64 IMA FUNGUS Aspergillus section Versicolores ARTICLE
Fig. 3 Aspergillus protuberus 0;;+ GR/R 7 .R E 6/ E F E F # ?%F ?%F # J # # 56/ 3 ; 56/ 3 @ 56/ 3 @ 56/ 3
Description: Colonies 6/ E F .R E ! Type: Sine loc 1 E & 67G/ M. Swift G M .QMG2 0;;+ .R/N < .M2 hyphae give a mealy appearance in some areas of some Description: Colonies 6/ E F .R E ! isolates, sporulation moderate with conidial heads often 2 M .WMGW # creamy white but sometimes patches of yellow-green conidia sporulating well, conidial heads deep bluish gray-green ;2W ;6 < ;2. < # = ;2/ to reddish brown, reddish-brown soluble pigment, reverse soluble pigment present, reverse brownish red or orange # ;G7 ;.7 ;./ ;6 E E 6/ ?%F .R E ! 2 M 6/ ?%F .R E ! G M .WMG. GWM2Q # \ 2MR shallow sulcations, sporulating in dark grayish blue-green < # color, funicular hyphal aggregates often seen centrally, " W E F R E GW E < # 0;;+ 2WVQ " W E F Stipes ! G M R E " 6./M G//M GW E 6/M6W 6/ Q// M6.R/ & 2M6/ ) Stipes ! 2 6//MR// 3 & 2MW vesicles, vesicles pyriform to spatulate, rarely subspherical, 3 vesicles RM6/ M6R 3 conidial VM 6/M.2 M.W ) conidial heads biseriate, metulae heads biseriate, metulae # # VMW # # GM 2MW MQ & . RM2 R MR R & .MG 3 phialides WM6/ & . /M. R 3 ) phialides 2M RMQ M66 & .MG MG R ) ! 2 conidia ! 2 . RMG / MR 3 present, conidia ! G . /M . RMG R MR ) hülle cells ! G Aspergillus tabacinus 0 et al J. Agr. Chem. Soc. Japan 101 6WW 67G2 ? ? RG7R22 Aspergillus sydowii 9 J & 9 E ! R M Aspergilli162W 67.V Synonym: Aspergillus versicolor # magnus J J. Fac. ? ? .W7VGV Agric. Hokkaido Univ. 491 622 67R/ ! 2 M Basionym: Sterigmatocystis sydowi 9 J Ann. Type: Sine loc.1 < 67G2 Y. Nakazawa 0;;+ Mycol 111 .R 676G 2W76N <
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Fig. 4. Aspergillus sydowi 0;;+ .R/ 7 .R E 6/ E F E F # ?%F ?%F # J # 56/ 3 ( 56/ 3 J 56/ 3
Fig. 5. Aspergillus tabacinus 0;;+ 2W76 7 .R E 6/ E F E F # ?%F ?%F # J # # 56/ 3 @ = 56/ 3
Description: Colonies 6/ E F .R E ! ?%F .R E ! R M 6WMG/ R M G/MG. 0;;+ 2W76 # # .MG # 0;;+ R/G6 0;;+ V.2Q6 \ ;2W = < < # # when present, no soluble pigment, reverse uncolored in " W E F R E GW E 0;;+ R/G6 E 6/
66 IMA FUNGUS Aspergillus section Versicolores ARTICLE
Fig. 6. Aspergillus versicolor 0;;+ .GQ 7 .R E 6/ E F E F # ?%F ?%F # 5R/ 3 J # 56/ 3 @ 56/ 3
Stipes ! R " W E F R E W/M G//MW// M7// & 2MQ M7 " W E F ) vesicles RM QM6R M.. ) GW E Q conidial heads biseriate, metulae covering half to entire Stipes ! V M # GMQ M7 & . RM2 R MR R ) phialides RMQ M66 2RM .//MWR/ M & .MG MG R ) 6/R/ & 2M RMQ M6. ) vesicles pyriform to spatulate, conidia ! R VM 7M6W M./ ) conidial heads biseriate, metulae . RM GM2 MW ) # # GMV M7 & . RM2 R ) phialides 2M RMW M66 & .MG ) conidia ! Aspergillus versicolor B & Annali Bot 71 7 V .M 67/Q . RMG R MV R ) hülle cells globose, ? ? 6W.6R7 0;;+ R.67 ?2/ ! V M Basionym: Sterigmatocystis versicolor B in ? Thèse de médicine 0 2716R 67/G Observations: & <= 0;;+ .GQ 67GR ' Type: Sine loc.: 67GR V. K. Charles 0;;+ .GQN < of dark red soluble pigment and scarlet colony reverse on CYA, from the more recent isolates that were placed in the F;J E E 67W/ 67Q2 & Description: Colonies 6/ E F .R E ! V M 0;;+ R.67 0;;+ 6G622 0;;+ 6G62R .QMGV 2MR 0;;+ 6G62V ' sporulating well, conidial heads pale grayish green near F ;2W # A. versicolor based < N0F ' # # soluble pigment, reverse vinaceous or brown or scarlet New species 0;;+ .GQ E 6/ ?%F .R E ! V M .6MG6 Aspergillus austroafricanus > # J L ( aggregates, sometimes dominating colony appearance, 9 L P sp. nov. sporulating in pale to dark bluish green to gray green color, ? ? Q//R7W < ! W M not present in others, reverse pale yellow, yellow orange Etymology: " J F
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Fig. 7. Aspergillus austroafricanus 0;;+ .GG 7 .R E 6/ E F E F # ?%F ?%F # J # 56/ 3 @ = 56/ 3
Fig. 8. Aspergillus creber 0;;+ RQRQG 7 .R E 6/ E F E F # ?%F ?%F # J # 56/ 3 @ 56/ 3
Type: South Africa1 E E & 67.. V. overgrowth by clumped hyphae making surface appear A. Putterill (" QQ/762 M mealy, sporulating well, conidial heads near sage green 0;;+ .GG W .R E E F ?%F ;2W < # E 6/ ?%F .R E Diagnosis: E = GW E ! W M .W # E F < # " W Description: Colonies 6/ E F .R E ! W M E F R E GW E .GM.2
68 IMA FUNGUS Aspergillus section Versicolores ARTICLE
Fig. 9. Aspergillus cvjetkovicii 0;;+ 2V2. 7 .R E 6/ E F E F # ?%F ?%F # 0 5R/ 3 J # 56/ 3 @ 56/ 3 @ 56/ 3
Stipes ! W ;2W ca 6 0;;+ .G6 2/M 6//MGR/ MR// 3 & GMR MV ) vesicles pyriform to with vivid brown soluble pigment, other isolates no soluble 2M VM6. M6R ) conidial heads biseriate, < # # metulae # 63G # GMW M7 & . RM2 R " W E F R E ) phialides 2M RMW M7 & .M . RMG M2 ) GW E Stipes ! Q 6/M W/M2R/ MVR/ < GM present, conidia ! W . RMG R 2MW MQ ) vesicles pyriform to spatulate and occasionally M2 R ) 2M WM6W M.R ) conidial heads biseriate, metulae GM 2MV MQ < . RM2 R MR ) phialides 2M RMQ M Aspergillus creber > # J L ( 9 L 6/ < .MG M2 ) conidia ! Q spherical to subspherical, P sp. nov. . RM GM2 M7 ) ? ? Q//R7Q ! Q M Etymology1 ! + creber meaning numerous Aspergillus cvjetkovicii > # J L ( 9 ' L P sp. nov. ? ? Q//R77 Type: USA: California1 < 0 # .//Q Z. ! 7 M Jurjevic (" Q//76. M X Etymology: 0 E# # 4 M # 0;;+ RQR7. W .R E E F ?%F T X ËÌ = [=ÌÌ# = Ë
Diagnosis: ( = GW Type: USA: New Jersey: < 676R W. Wilson (" E E F ?%F QQ/7/7 M 0;;+ ..W W E F ?%F .R E E F ?%F
Description: Colonies 6/ E F .R E ! Diagnosis: ( GW E Q M 6QM.V GMR < mm centrally, peripheral areas white or yellow, central area E F sporulating well, conidial heads pea green to artemisia green ;2W < Description: Colonies 6/ E F .R E ! 7 M pigment, reverse clay colored to cinnamon or reddish brown .2M.7 ;.7 E 6/ ?%F .R E ! Q M .MG 6QM.. 6M. ;2W 0;;+ overgrown centrally with hyphae aggregated into funicles, RQR7G < = cinnamon, reddish brown soluble pigment, reverse yellowish
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Fig. 10. Aspergillus fructus 0;;+ .G7 7 .R E 6/ E F E F # ?%F ?%F # J # 56/ 3 @ 56/ 3 @ 56/ 3
;.7 # Description: Colonies 6/ E F .R E ! ;G7 E 6/ ?%F .R E ! 7 M 6/ M .7MG7 6WMGV 2MR 0;;+ RQR7G ;2W 0;;+ ..W 0;;+ .G/ < 0;;+ 2V2. RQR7G 0;;+ .G7 # produce soluble pigment, reverse brownish orange or pale = ;. E " W E F R E GW E 6/ ?%F .R E ! 6/ M ..MG. diam, slightly sulcate, centrally covered by hyphal tufts, Stipes ! 7 M sporulation in yellow-green hues near artemisia green 2/M .//MW// MQR/ & GM 2MW MQ ) vesicles pyriform ;2W < # RM 7M6Q M.G ) 0;;+ .26 \ ?%F " conidial heads biseriate, metulae covering half to entire W E F R E # GMV MQ & . RM2 R ) phialides RMQ M6/ & .MG M " W E F GW E 2 ) G. ) 2 Stipes ! 6/ R/M occasionally present, conidia ! 7 6R/M2// MR// & 2MW 3 vesicles pyriform to spatulate, .M . RMG R MR ) VM 7M6W M.6 ) conidial heads biseriate, metulae spinulose, hülle cells ! 7 # # .M GMW M7 & . RM2 R MW ) phialides RM VMQ M66 & .MG M2 ) Aspergillus fructus > # J L ( 9 L conidia ! P sp. nov. 6/ .M ? ? Q//V// . RMG R M2 R ) hülle cells ! 6/ ! 6/ M Etymology: ! Aspergillus jensenii > # J L ( 9 L Type: USA: California1 < 67G7 Bliss (" P sp. nov. QQ/76R M 0;;+ .G7 W ? ? Q//V/6 .R E E F ?%F ! 66 M Etymology: 0 E 0 > Diagnosis: Resembling A. versicolor GW E reported this species as Aspergillus globosus > 6R/M2// 3 # .//MWR/ homonym of A. globosus + 3 A. versicolor.
70 IMA FUNGUS Aspergillus section Versicolores ARTICLE
Fig. 11. Aspergillus jensenii 0;;+ RQVW6 7 .R E 6/ E F E F # ?%F ?%F # J # 56/ 3 ( 56/ 3 @ 56/ 3
Type: USA: Montana1 < [ .//Q Z. G. ) 2 R ) Jurjevic (" QQ/76/ M 0;;+ RQV// W .R E E F ?%F ! 66 conidia ! 66 spherical to . RM GM Synonym: Aspergillus globosus > E M # 2 R MW ) 6RM./ F %< J G6R1 2Q. 676. X 3 0;;+ RQRQ. + 6Q/7 Aspergillus puulaauensis > # J L ( 9 Diagnosis1 E GW E L P sp. nov. conidial color near celandine, tawny olive to dark umber ? ? Q//V/. # E F ! 6. M Etymology " ( R R P P X Description: Colonies 6/ E F .R E ! Ë M=M= =M= = Ë 66 M ./M.W raised or sunken, with clumped hyphal aggregates common Type: USA: Hawaii1 < .//G in some isolates, sporulating moderately well, conidial heads D.T. Wicklow (" QQ/766 M ;2W 0;;+ GRV26 W .R E E F ?%F < soluble pigment faint or intense yellow brown, in one case Diagnosis: " # # ;G7 ?2/ GW E ;G E 6/ ?%F .R E ! 66 M 6WMG/ Description: Colonies 6/ E F .R E ! 6. M have funicular tufts of aerial hyphae centrally, sporulating well ..M.R RMV = < with funicular hyphal clumps, sporulation light, conidial heads pigment either light brown or reddish brown, brownish orange ;2W < in one isolate, reverse pale yellow or orange or brownish soluble pigment when present brown, reverse yellowish to clay " W E F R E GW E ;G7 ;.7 E 6/ ?%F .R E ! 6. M .6M.R Stipes ! 66 # ;2W 2RM .//MW// M6/// < # & MG 2MW MQ ) vesicles pyriform to spatulate, rarely " W E F R E RM WM6V M.. ) conidial heads GW E biseriate, metulae # 63G # GMQ & . RM Stipes ! 6. GRM 2 MR ) phialides 2M RMQ M66 & .MG ) 6//MR// MW// & GM 2MW ) vesicles pyriform to spatulate,
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Fig. 12. Aspergillus puulaauensis 0;;+ GRV26 7 .R E 6/ E F E F # ?%F ?%F # J # 56/ 3 ? 5R/ 3 P 56/ 3 @ 56/ 3
RM QM6Q M.6 ) conidial Aspergillus subversicolor > # J L ( 9 heads biseriate, metulae # # GM 2M L P sp. nov. W M7 & . RM2 ) phialides RMW M6/ & .MG ) ? ? Q//V/G ! 6G M present, conidia ! 6. . RM GM Etymology1 Aspergillus versicolor 2 MR R ) hülle cells ! 6. M 66M67 3 Type: India: Karnataka1 < 67W/ ?2/ B. Muthappa (" QQ/76Q M 0;;+ RQ777 W .R E E F ?%F
Fig. 13. Aspergillus subversicolor 0;;+ RQ777 7 .R E 6/ E F E F # ?%F ?%F # J # 56/ 3 J 56/ 3
72 IMA FUNGUS Aspergillus section Versicolores ARTICLE
Fig. 14. Aspergillus tennesseensis 0;;+ 6G6R/ 7 .R E 6/ E F E F # ?%F ?%F # J # 56/ 3 @ 56/ 3
Diagnosis1 E = GW E Diagnosis1 ( = GW slowly on all media, producing yellow soluble pigment on E ?%F E F < Description: Colonies 6/ E F .R E ! 62 M Description: Colonies 6/ E F .R E ! ..MG/ 6G M 6QM./ RMV mat, radially sulcate, centrally raised or sunken, overgrown centrally, wrinkled, sporulating sparsely, conidial heads by clumps of aerial hyphae in some isolates, sporulating well ;2W < ;2W # ;6R E < # 6/ ?%F .R E ! 6G M 6.M62 in brownish orange shades near honey yellow or chamois diam, low, plane, velutinous, sporulating in bluish green color ;G/ E 6/ ?%F .R E ! 62 M ;2W < # ./M2V # " W E F R E GW E ;2W < no soluble pigment, reverse uncolored, pale lemon yellow, Stipes ! 6G " W E F R E GW E V/M .R/M2R/ MRR/ & 2MW M6/ ) vesicles VM 6/M6W M.. ) conidial Stipes ! 62 heads biseriate, metulae # 63G GRM 6//MG// M2// & 2MW ) vesicles # GM 2MW M7 & .M . RM2 ) .MG WM 6/M6V M6Q ) conidial heads biseriate, phialides, RMQ M6/ & .MG ) metulae # # 2MV MQ & . RM2 ) conidia ! phialides RMQ M66 & .MG ) 6G conidia ! . RM GM2 MW ) 62 . RM GM2 MQ ) Aspergillus tennesseensis > # J L ( 9 L P sp. nov. Aspergillus venenatus > # J L ( 9 ? ? Q//V/2 L P sp. nov. ! 62 M ? ? Q//V/R Etymology1 " & ! 6R M Etymology1 ( < Type: USA: Tennessee1 < < 67Q2 B.W. Horn (" QQ/76W M 0;;+ Type: USA: Tennessee1 < < 67Q2 B.W. 6G6R/ W .R E E F ?%F Horn (" QQ/76V M 0;;+ 6G62W W .R E E F ?%F
VOLUME 3 · NO. 1 73 Jurjevic, Peterson & Horn ARTICLE
Fig. 15. Aspergillus venenatus 0;;+ 6G62W 7 .R E 6/ E F E F # ?%F ?%F # J # 56/ 3 ( 56/ 3 @ 56/ 3 @ 56/ 3
Diagnosis1 ( GW E Aspergillus cvjetkovicii, A. sydowii and A. venenatus < E F ?%F A. sydowii isolates grow GW E A. cvjetkovicii and A. venenatus isolates do Description: Colonies 6/ E F .R E ! A. cvjetkovicii < 6R M ..MG6 soluble pigment on CYA, while A. venenatus isolates produce ;2W = < ;2. < # Aspergillus amoenus, A. austroafricanus and A. tabacinus # ;6R E 6/ = [ A. amoenus ?%F .R E ! 6R M 6WM.2 GW E A. tabacinus isolates produce no lightly sulcate, low, central tufted funicular aggregates of soluble pigment and A. austroafricanus produces reddish aerial hyphae, sporulating well in deep green color near slate E F ;2W < # The remaining eight species produce conidia with # " W noticeably roughened walls, but the ornamentation is not E F R E GW E & eight species, A. versicolor and A. fructus, have roughened Stipes ! 6R GW E & # ./M 6//M2// MR// & 2MW 3 vesicles # 6R/M2// VM 7M6W M.6 ) conidial 3 A. fructus # .//MWR/ 3 A. versicolor L heads biseriate, metulae # # GM < A. fructus # A. versicolor 2MW M7 & . RM2 MR ) phialides RM VMQ M66 & .MG G R isolates and while separation of these species using ) phenotype on standard media appears possible, until more ! 6R hülle cells ! 6R present in some isolates, conidia ! 6R ' GM2 MV ) Genealogical concordance species recognition clearly ! 6 J GW E Phenotypic species recognition are A. protuberus, A. creber, A. jensenii, A. puulaauensis, A. Growth rates of species on different media are presented in subversicolor and A. tennesseensis Aspergillus protuberus & . E F < Phenotypic recognition of species in section Versicolores ;6 # ?%F is based on smooth, roughened or spinulose conidia, conidial \ A. jensenii isolates produce brown < E F ?%F CYA colony reverse colors from tawny olive to dark umber, GW E ;2W F A. puulaauensis
74 IMA FUNGUS Aspergillus section Versicolores
Table 2. E Versicolores # W " .R E < ARTICLE Species CYA MEA CY20 M40Y M60Y CYA at 37 °C A. amoenus ./M.7 66M6R 66M./ 62M.. 6RM.2 V A. austroafricanus 6QM67 6VM6W .2M.R 6QM67 6WM6Q - A. creber 6WM.G 66M6R 6RM.. 6QM.G 67M.2 - A. cvjetkovicii 6RM.6 62M6W 6WM./ 6VM.2 6RM.R - A. fructus 6GM./ 6/M6V 7M6W 6/M.G 6/M./ 2 A. jensenii 6VM./ 7M6G 6RM./ .6M.V ..M.Q - A. protuberus 6WM.R 66M6Q 62M.. .6M.2 ./M.2 - A. puulaauensis 6QM.6 66M6. 6WM67 67M.. 6QM.6 - A. subversicolor 6GM62 VMW 6/M66 6RM6V 6VM6Q - A. sydowii ./M.R ./M.R .6M.V .GM.V .GM.W Q A. tabacinus .6M.V 6.M6V .6M.V QM.G QM.G - A. tennesseensis ./M.. 6.M62 6WM67 67M.. 67M.. - A. venenatus 62M6W QM6/ 6RM6V 67M.G 6WM.6 - A. versicolor ./M.V 6/M6Q 67M.. .6M.V ..M.R Q
Table 3. ( "&J "&J Versicolores "&J ITS genotype Predicted species A A. amoenus, A. fructus, A. protuberus, A. tabacinus, A. versicolor A. subversicolor C A. austroafricanus D A. cvjetkovicii, A. jensenii, A. tennesseensis, A. venenatus % A. sydowii ! A. sydowii G A. amoenus P A. tabacinus " A. creber, A. versicolor > A. puulaauensis K A. creber + A. creber ? A. jensenii N A. creber
?2/ # F [ A. versicolor isolates, several appear to have mutated and consequently and A. amoenus GW E A. produce colonies that have a wet appearance when grown on jensenii ?2/ A. subversicolor CYA or produce only moist aerial aggregates of hyphae with # ?%F ?2/ " & . E F N0F ' N0F ' A. tennesseensis ?%F # is the most reliable means for identifying species within this ;2W section. = Aspergillus creber "&J Versicolores E F ?%F & G J and conidial color on either medium is pea green to sage @ F # ;2W different species and genotype D is present in four different There is considerable variation in colony appearance " A. within species and considerable overlap in colony appearance creber " K + 0 "&J between species, making species separation within section Versicolores " included in this study were propogated in vitro for several
VOLUME 3 · NO. 1 75 Jurjevic, Peterson & Horn
DISCUSSION et al. .//7 # ( .//Q " < Aspergillus section Versicolores could easily be dropped Versicolores isolates was made using CYA cultures grown for from Aspergillus < & W .R E K 9 ( 67QQ & # A. versicolor sensu lato isolates suggests that section
ARTICLE J ' Versicolores should be retained as a monophyletic and useful 6/ .R E E F # < Aspergillus versicolor is the most reported fungal species ; 9 ! 67VR in section Versicolores from damp indoor environments 6/M62 L 6/ > .//G ; et al. .//R Versicolores J J J J J 9 [ # J .//2 . L ! 6 & "&J using A. versicolor- N et al .//RX was not included because it contained few informative # < nucleotides and because its veracity as a phylogenetic for A. subversicolor and A. sydowiiX ' @ et al. .//R & "&J " A. creber and A. jensenii some themselves however may be of interest for bar-coding studies isolates did not amplify even though the genotypes were & ' identical with isolates that did amplify, suggesting degradation Versicolores are of the “two intron” type and probably have ' N0F a different evolutionary origin than the “three intron” type of & = Versicolores isolates in this study were = ( .//Q T> none comprised A. versicolor sensu & 6 [ " # A. versicolor < N0F ' = < MJF MJF P 67VV ' J F & < L geographically, but was not commonly encountered among characters in section Versicolores Aspergillus creber was the # P R most frequently isolated species from indoor air samples in ' F N0F ' MJF 6G < A. protuberus & et al ./// N et al. # A. jensenii .//G Aspergillus versicolor sensu stricto may not be Phylogenetic recognition of species occasionally makes it & A. cvjetkovicii and ( et al ./66 A. puulaauensis & because the phenotypic characters of the species overlap other newly described species, A. fructus, A. austroafricanus, < A. subversicolor, A. tennesseensis and A. venenatus, were ! 0;;+ RG/ & 0;;+ 6G6R6 6 F et al. ./6/ # # the temperate regions of the world and are much less diverse ' & ! A. versicolor and A. fructus the limited number MJF # # from many regions of the world may represent much of the diversity present in section Versicolores species, but the phylogenetic data are unequivocal and so A. & "&J ' fructus = = Prior to this publication A. versicolor was a species with et al. ./6/ J et al documented genetic and phenotypic variation that did not ./6. " Versicolores "&J # ! # are now known in section Versicolores "&J & G # G ] "&J ' 2 ] # Petromyces this section, use of multiple loci is the most reliable means Aspergillus Flavi \ and P. N0F ' = Versicolores nomius 62 # B et al ( ./6. .//7 " Petromyces clade, one species may possess Viable propagules of A. versicolor have been recovered # K et N J K =( et al .//G al. 67QW ( et al. .//6 J et al. ./6. L # # the validity of some species in the Petromyces clade have & F;J E E # A. ' B et al. .//7 versicolor MK # # ( .//Q B Upon sequence and phenotypic analysis, these isolates were
76 IMA FUNGUS Aspergillus section Versicolores
A. amoenus, A. tabacinus and A. REFERENCES ARTICLE protuberus, all of which occur in the A. versicolor subclade ! 6 F A. creber 0;;+ VR22 A. F FJ J KF J ; &N ./6/ " sydowii \ composition is geographically patterned and more diverse in F [ P < Proceedings of the National species in section Versicolores Aspergillus versicolor has Academy of Sciences MJF 1071 6GW2QM6GWRG " F ! # >E J " ; " + +J J J B et al .//Q " E F ./66 F = ?%F E ./J ?2/ Applied and Environmental Microbiology 77: ?V/ ./ 2/ V/ ] # 26Q/M26QQ F & . >L ? KF F N; L &> K EF # ?V/ K N( " >" JF ( (@ ? JF .//7 J Versicolores have a remarkably broad Patterns of susceptibility of Aspergillus isolates recovered from # patients enrolled in the transplant-associated infection surveillance Aspergillus versicolor \ < Journal of Clinical Microbiology 47: G.W6MG.WR precursor sterigmatocystin, a compound that is mutagenic NF J K+ 677R @ # B # 9 J ./6/ F " 1 Experimental and Molecular Approaches to Plant infested with three morphotypes of A. versicolor, all of which Biosystematics (E P 9 F@ J 1 .Q7MG/G J produce sterigmatocystin, have been implicated in dairy + ? 1 ? @ < N 0 P M ? ? L ./6/ E < B 9 P 67QR & # N0F A. versicolor, A. Applied Microbiology and Biotechnology 871 77M tennesseensis and A. venenatus, and as these species 6/Q occur in the two main subclades of section Versicolores N &; ; N ? ? .//R F ! 6 < & Versicolores # # Journal of species in agricultural commodities and their role in Microbiological Methods 611 7M6V < ' " N >; > N> & % ( F & >L .//G to sterigmatocystin, recent studies have revealed numerous Reproductive isolation and phylogenetic divergence in # ! et al. ./66 + Neurospora: comparing methods of species recognition in a et al. ./66 A. versicolor sensu lato > et al. .//W Evolution 571 .W.6M.W26 discovered novel nucleotide analogs from A. puulaauensis Domsch KP @ L F &P 67Q/ Compendium of Soil which was reported under the name A. versicolor Fungi B 6 F ( MK Aspergillus versicolor has been implicated as the % NF EJ &+ J NE K B( ! # T >0 .//7 " < et al. ./6. International Journal of Molecular Sciences 101 R2W6=R2Q2 et al .//7 % J + F J N J P + F ! P"B ( P %< ? .//. [ < Aspergillus versicolor et al. .//R L Versicolores clinical isolates and sterigmatocystin in carpet dust from damp indoor 0;;+ .R2 A. sydowii # Applied and Environmental Microbiology 68: 0;;+ ..V A. versicolor, is here GQQVMGQ7/ A. amoenus # ! >? J NP & J L ;? ./66 of fungal species to fungal antibiotics, a more detailed study J 1 of A. versicolor clinical isolates might be of value to guide V= < < # % ( et al. ./66 > Journal of Organic Chemistry 76: R7R2MR7RQ @ >% E # J% E E ? +=> L >; ACKNOWLEDGEMENTS J + J=" ? J # EE E > K # B > > J E ! ? > ( J L F % ? @ # # ( P J @P N [ J NR% E % # + N # + P E @ @P =( N @ => J N # & >P > B K ( F ! ? J %M F N N & L MJNF ( "+ + J M # ( # ?F [ ; ? ? & & K & F F % F P E K ? ? 0 LE N NL E ? P ? ? ( ? ! ; ? N ( J ?J [ equipment does not constitute a guarantee or warranty by the United JF L .//R J ' Aspergillus nidulans and J N F # comparative analysis with A. fumigatus and A. oryzae. Nature < MJNF 4381 66/RM666R ' # @ L E ? [ FPJ ( >" J ;F 67QR " < Aspergillus " 1 Advances in Penicillium and
VOLUME 3 · NO. 1 77 Jurjevic, Peterson & Horn
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78 IMA FUNGUS Aspergillus section Versicolores
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