IMA FUNGUS VOLUME 3 NO 1: Aspergillus section Versicolores: nine new species and multilocus DNA sequence based phylogeny ARTICLE Abstract: repli

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(1)IMA FUNGUS · VOLUME 3 · NO 1: 59–79. 16/RR7Q3 ./6./G/6/W. T > # 6J L( . LP G. ART I CLE. Aspergillus section Versicolores: nine new species and multilocus DNA sequence based phylogeny. 6. %?J+F ".//; 6G/0 E 0>/Q/WW !

(2) ( ? ; M 0 E F M ; F ; J # MJ N F 6Q6R 0 M # J ( " V6V/2 MJFX . 1 J Z # .. G. 0 ( ; +

(3) F ; J # MJN F ([<R/7N @FG7Q2.=/R/7. Abstract: Ã=

(4)

(5) = N0F;0F .N0F replication licensing factor Mcm7, and pre-rRNA processing protein Tsr1 ' numerous isolates belonging to Aspergillus versicolor.&

(6)

(7) Aspergillus austroafricanus, A. creber, A. cvjetkovicii, A. fructus, A. jensenii, A. puulaauensis, A. subversicolor, A. tennesseensis and A. venenatus are described as new species and A. amoenus, A. protuberus, A. sydowii, A. tabacinus and A. versicolor are accepted as distinct species on the basis of molecular and phenotypic differences. PCR primer pairs used to detect A. versicolor in sick building syndrome studies have a positive reaction for all of the newly

(8) < A. subversicolor. 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. Article info: J

(9) 16W> ./6.XF 1W> ./6.X(

(10) 1.6> ./6.. INTRODUCTION Aspergillus section Versicolores was originally erected as the Aspergillus versicolor

(11) &9E 67.V

(12) ' #

(13) &9; 672R ; 9 ! 67VR # Aspergillus 6Q A. versicolor group. Gams et al.67QR < ; 9! O67VR Using scanning electron J%? K 67Q7 < surface ornamentation of most species of the section and removed seven species from section Versicolores K 677G #

(14) characteristics and accepted the seven species previously. #

(15) K 67Q7 Versicolores ( .//Q species in the section based on multilocus DNA sequence 62 Aspergillus. Aspergillus versicolor is the most widely reported and studied species in section Versicolores"

(16) Net al.67Q/ # J et al..//6J et al.//.% et al.//.F et al../6/F et al../66# ( 9 P .//7 K =( et al .//G ?

(17) .//Q > .//G (. et al .//R et al .//7 % et al .//7 ( 9 P .//7 ? 9F ./6/ " < $ < 6? 9F

(18) 67QV& et al.///0 .//G B È # 9J ./6/ %# Versicolores species <

(19) # =

(20) = N et al. 67Q/, K .//. ; 9 ! 67VR & 9 E 67.V B 9 P 67QR N0F=

(21) . © 2012 International Mycological Association You are free to share - to copy, distribute and transmit the work, under the following conditions: Attribution:

(22)

(23)

(24) Non-commercial: No derivative works:

(25) 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. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights.. VOLUME 3 · NO. 1. 59.

(26) ART I CLE. Jurjevic, Peterson & Horn. # ' N0F V

(27) N et al.//G Versicolores The species described and accepted are monophyletic.. MATERIALS AND METHODS Fungal isolates & # < &

(28) 6 #

(29) F ; J # E E 0;;+ ( " 133. #. Culture methods E E < EF R É.RÉ GWÉ < ?%FE./J ?2/ ?V/ .RÉ 6/ ( 67Q/ K .//.?2/ .] < /R] < 2/] X?V/ .] < /R ] < V/ ] E and appearance were recorded and photographs were made 6/=

(30) .RÉE ; 676. .

(31) ;2R. Microscopy ? <

(32) /6 ] & \=6// <

(33) N"! F

(34) gently pressing a ca ./ R W/] F+ N?.R//

(35) N"! # & J F 0 J+; NW/ ( ? ( ( .//G F

(36) (. DNA methods E 6.R= +% $ .R+ <

(37) E .// .8G .RÉ

(38) # =

(39) N E&F

(40) < X

(41) centrifugation and DNA was precipitated from the aqueous ' # & acids were collected by centrifugation, the pellet was rinsed W/ ] # 6//+ DNA was diluted ca 616// Ã=

(42) BT2 CF "&J = N0F ID ;0F . RPB2 N0F Mcm7 = ;0F Tsr1 . 60. by Peterson et al ./6/ J

(43) 7RÉ .

(44) GR 7V É G/ X R6 É V/ X W. É V/ X < W.É R [

(45)

(46) a higher annealing temperature was used to obtain single

(47) N0F '

(48) #G6 F"GWG/ '

(49) F 133

(50) 3 ; '

(51) = . J ' 133 3 E. ' E+MJ&F+L & et al. 6772J ' @ >0QRGW7Q8>0QR26G6 %!VR.6WV %!VR.6WQ %!VR.6QR8 %!VR.6QW %!VR.67V %!VR../G %!VR../78%!VR..66 %!VR..628%!VR..6V %!VR...V %!VR..V2 %!VR..VV %!VR..WG8%!VR..WR %!VR..Q2 %!VR..76 %!VR..7W8 %!VR..77 %!VR.G/.8%!VR.G/2 %!VR.G62 %!VR.GR. %!VR.GR2 %!VR.GV68%!VR.GVG %!VR.GW. %!VR.GW7 %!VR.GQR8%!VR.GQW %!VR.G7/8%!VR.G7. %!VR.2/. %!VR.22/ %!VR.22. %!VR.2278%!VR.2R6 %!VR.2V/ %!VR.2VW %!VR.2WG8%!VR.2WR %!VR.2WQ8%!VR.2Q/ %!VR.27/ >G/6QQ78>G/6Q7V ( (FM(2/

(52) 6/ J .//G! = R///

(53) 00"

(54) < R///& was used as the starting point for parsimony analysis with ) * &;

(55) ) * &;

(56) 6///

(57)

(58) ? G6. P

(59) 9 ;' .//6 ;' 9 P

(60) .//G & Mcm7, Tsr1 and RPB2 data sets included only protein-coding sequences and each data set was 6. G& BT2 and CF loci included protein-coding and intron regions and the data < F @&; = #

(61) # =

(62) ? # ? E ?E?E R6/V # E

(63) < 9 J 677R concordance phylogenetic species recognition concepts of Taylor et al..///E # 6 = .

(64)

(65)

(66) N et al..//GJ ÉW/]

(67) É/7R

(68)

(69) N et al..//G & A. versicolor in a PCR N et al.//R '

(70) < < N et al.//R IMA FUNGUS.

(71) Aspergillus section Versicolores. Table 1.( # Provenance. ..V. MJF1 < 676G. .GV. @ 1? < Berberis ,67G/M. Roberg. VRQ. MK1 <

(72) 67.7G. A. Ledingham. 2QGQ GRV// F=.G..Q. ART I CLE. NRRL number. Aspergillus amoenus? ?.R/VR2. %' # 0;;+.GV # E

(73) # J 67V. <= MJF1P K ( <

(74) Gandoderma australe.//R, D.T. Wicklow " 1K <

(75) . 67WQB. Muthappa. Aspergillus asperescensJ? ?.7.QGR 2WW/. %<= = . Aspergillus austroafricanus sp. nov.,? ?Q//R7W .GG. JF 1E 67..

(76) V. A. Putterill <= . Aspergillus creber sp. nov.,? ?Q//R7Q .G6 VR22. JF 1E 67..

(77) V. A. Putterill F [ 1 < $

(78) 67W7A. Wellman. .RV.W. > 1"

(79) < 677VT. Goto. RQRQG. MJF1( # < .//QZ. Jurjevic. RQRQ2. MJF1E < .//QZ. Jurjevic. RQRQW. MJF1E < .//QZ. Jurjevic. RQR7.. MJF1E < .//QZ. Jurjevic <= . RQR7W. MJF10 > < .//QZ. Jurjevic. RQV/6. MJF10 > 1 .//7Z. Jurjevic. RQV/V. MJF1( # < .//7Z. Jurjevic. RQV/W. MJF1( # < .//7Z. Jurjevic. RQV6.. MJF10 > < .//7Z. Jurjevic. RQVW/. MJF10 > < .//7Z. Jurjevic. RQVW.. MJF1@ < .//7Z. Jurjevic. RQVWG. MJF1@ < .//7Z. Jurjevic. RQVWR. MJF1[ < .//7Z. Jurjevic. Aspergillus cvjetkovicii sp. nov., ? ?Q//R77 ..W. MJF10 > < 676RG.W. Wilson <= . .G/. E 1 < 676WRound. 2V2. RQR7G. M1 0;;+67V7D. I. Fennell L2V2. MJF1E < .//QZ. Jurjevic. Aspergillus fructus sp. nov., ? ?Q//V// .G7. MJF1E < 67G7Bliss <= . .26. M1 < 676VL. McCulloch. Aspergillus jensenii sp. nov., ? ?Q//V/6 ..R. MK1676G E&

(80) Dade. .GR. MK1+ < 67G/H. Raistrick. .2/. MJF10 " < 6766C. N. Jensen, E&

(81) L type strain of A. globosus. RQRQ.. MJF1? < .//QZ. Jurjevic. RQV//. MJF1? < .//QZ. Jurjevic <= . RQVW6. MJF1( # < .//7Z. Jurjevic. RQVW2. MJF1[ < .//7Z. Jurjevic. Aspergillus multicolorJ ? ?.7.Q27 2WWR. VOLUME 3 · NO. 1. %<= = . 61.

(82) Jurjevic, Peterson & Horn. ART I CLE. Table 1.E NRRL number. Provenance. Aspergillus protuberus? ?G.VVR/ VV6 GR/R. MK1 <

(83) 67.7G. A. Ledingham # <

(84)

(85)

(86) 67VQ <= . RQV6G. MJF10 > < .//7Z. Jurjevic. RQW2W. MJF10 > < .//7Z. Jurjevic. RQW2Q. MJF10 > < .//7Z. Jurjevic. RQ77/. MJF1E < .//7Z. Jurjevic. RQ776. MJF1E < .//7Z. Jurjevic. Aspergillus puulaauensis sp. nov., ? ?Q//V/. GRV26. MJF1P (O O P .// <

(87) .//GD. T. Wicklow <= . RQV/.. MJF1L B < .//7Z. Jurjevic. V.6.2. MJF1P <

(88) Inonotus.//GD. T. Wicklow. V.R6V. E 1F

(89) <

(90) ca677/S. P. Abbot, ' # MF?PWVR6. Aspergillus subversicolor sp. nov., ? ?Q//V/G RQ777. " 1K <

(91) . 67W/B. Muthappa <= . Aspergillus sydowii? ?.W7VGV .R/ .R2 2WVQ V.2R/. M1 67G/ E&

(92) M. Swift. MJF1@ L 672/M. M. Harris MJF1E < 67V7 & 1 < 67WWE. G. Simmons. Aspergillus tabacinus? ?RG7R22 VR7. MK1 <

(93) 67.7G. A. Ledingham. 2W76. M1 <

(94) 67G2Y. Nakazawa <= . R/G6. Unknown: type isolate of A. versicolor# magnusJ # "![67V.. V.2Q6. 0 1K < 67WW. Aspergillus tennesseensis sp. nov., ? ?Q//V/2 ..7. M1 E&676W

(95) R. Thaxter. .G2. MJF1? # < 67.WC. Thom. 6G6R/. MJF1& << 67Q2B. W. Horn <= . 6G6R.. MJF1& << 67Q2B. W. Horn. Aspergillus venenatus sp. nov., ? ?Q//V/R 6G62W. MJF1& 1 < 67Q2B. W. Horn <= . 6G62Q. MJF1& << 67Q2B. W. Horn. 6G627. MJF1& << 67Q2B. W. Horn. V.2RW. MJF1? < 67Q7D. T. Wicklow. Aspergillus versicolor? ?6W.6R7 .GQ. MJF1 <

(96) 67GRV. K. Charles <= . R.67. JF 1( # 67W/ J. P. van der Walt.. 6G622. MJF1& << 67Q2B. W. Horn. 6G62R. MJF1& << 67Q2B. W. Horn. 6G62V. MJF1& << 67Q2B. W. Horn. Aspergillus species, undescribed RG/ 6G6R6. % " 1 <

(97)

(98) 67GQShumard MJF1& 1 << 67Q2B. W. Horn. RESULTS Phylogenic analysis of sequence data J < # . 62.

(99) N et al. .//G & ( .//Q A. versicolor, A. tabacinus, A. amoenus, A. protuberus and A. sydowii each ! 6 ! IMA FUNGUS.

(100) Aspergillus section Versicolores. ART I CLE Fig. 1.( N0F ' & Versicolores contains three subclades, the A. versicolor subclade, the A. sydowii subclade and the A. subversicolor

(101) &

(102) É7/]

(103) É/7/

(104)

(105) " 0;;+6G6R6 A. tennesseensis but may represent a distinct " 0;;+RG/ A. amoenus

(106) . & = isolate lineages, A. subversicolor and A. austroafricanus,

(107) & = 0;;+ 6G6R6 0;;+ RG/ VOLUME 3 · NO. 1.

(108) The section Versicolores clade contained three

(109) ! 61 A. sydowii subclade containing A. sydowii, A. creber, A. venenatus, A. tennesseensis, A.. 63.

(110) ART I CLE. Jurjevic, Peterson & Horn. Fig. 2. Aspergillus amoenus0;;+2QGQ 7 .RÉ 6/ EF

(111) EF. # ?%F ?%F # J

(112)

(113) #

(114) 56/@

(115) =

(116) 56/. cvjetkovicii, A. jensenii and A. puulaauensisX A. versicolor subclade containing A. versicolor, A. tabacinus, A. fructus, A. protuberus, A. amoenus and A. austroafricanusX A. subversicolor subclade containing the single species A. subversicolor J = A. sydowii in the A. sydowii subclade, in the A. versicolor subclade or in a distinct clade containing only A. sydowii ! J68JR J " # & Mcm7 locus from A. sydowii despite numerous attempts and thus A. sydowii does not ! JGJ " &

(117) ! 6 A. sydowii as a member of the A. sydowii subclade with strong statistical "

(118) O

(119) É7/]

(120) É/7/

(121)

(122) . TAXONOMY Previously described species Aspergillus amoenus ? ;

(123) Hedwigia 7016GQ 67G6 ? ?.R/VR2 ! . 8 Type: Germany1? <Berberis 67G/ M. Roberg0;;+2QGQ½ < Description: Colonies 6/ EF .R É ! . 8

(124) .R82/ G82 0;;+ ..V sporulating moderately to well, conidial heads in grayish green. 64. ;2W < present in some isolates, faint reddish soluble pigment present in some isolates, reverse mostly reddish brown hues, with some E 6/?%F .RÉ! .8 .G8GG # with shallow sulcations, colony center often with funicular hyphal aggregates, sporulation in blue-green to gray-green shades, no

(125) < 0;;+ ..V

(126) < # . "

(127) W EF R É "

(128) WEF GWÉ V Stipes ! .

(129) GR86//8V//866// .R828W8Q vesicles 28W86W8.6 conidial heads biseriate, metulae # 63G # G8V8Q .R82/8RR phialides R8V8Q8 66 .8G

(130)

(131) conidia ! .

(132) .R8GR8R 0;;+ GRV//

(133) 6.8.. ?2/ . Aspergillus protuberus Mikrobiologija 5166767VQ ? ?G.VVR/ ! G 8. ? Ê =E# # ¯. Synonym: Aspergillus versicolor# protuberus? Ê = E# # ¯K MycolPap16116G767Q7 Type: Yugoslavia: <

(134)

(135)

(136) ca 67VQ0;;+GR/R½ < . IMA FUNGUS.

(137) Aspergillus section Versicolores. ART I CLE Fig. 3Aspergillus protuberus0;;+GR/R 7 .RÉ 6/ EF

(138) EF. # ?%F ?%F # J

(139)

(140) # #

(141) 56/;

(142) 56/@

(143)

(144) 56/@

(145)

(146) 56/. Type: Sine loc1 E & 67G/ M. Swift 0;;+.R/½ < . Description: Colonies 6/ EF .R É ! G 8

(147) .Q8G2 .82 hyphae give a mealy appearance in some areas of some isolates, sporulation moderate with conidial heads often creamy white but sometimes patches of yellow-green conidia ;2W ;6 <

(148) # = ;2/ soluble pigment present, reverse brownish red or orange ;./

(149) ;6E 6/?%F .RÉ! G8 .W8G. $ 28R <

(150) # "

(151) WEF RÉ GWÉ Stipes ! G 8

(152)

(153) 6./8G//8 Q//86.R/ 286/ vesicles, vesicles pyriform to spatulate, rarely subspherical, V86/8.28.W conidial heads biseriate, metulae # # G828W8Q.R82R8RR phialides 28R8Q866 .8G8GR

(154) present, conidia ! G

(155) ./8.R8GR8R hülle cells ! G

(156) . Description: Colonies 6/ EF .R É ! 2 8

(157) .W8GW # sporulating well, conidial heads deep bluish gray-green ;2. <

(158) to reddish brown, reddish-brown soluble pigment, reverse # ;G7 ;.7 E 6/ ?%F .R É ! 28 GW82Q # shallow sulcations, sporulating in dark grayish blue-green color, funicular hyphal aggregates often seen centrally, <

(159) #

(160) 0;;+ 2WVQ "

(161) W EF RÉ "

(162) GWÉ 6/86W 6/ Stipes ! 2 6//8R// 28W vesicles

(163)

(164) R86/ 86R conidial heads biseriate, metulae# # V8W .8GphialidesW86/./8.R ! 2

(165)

(166) conidia ! 2

(167)

(168)

(169) .R8G/ 8R . Aspergillus sydowii 9 J & 9 E Aspergilli162W67.V ? ?.W7VGV ! 2 8. Synonym: Aspergillus versicolor# magnusJ J. Fac. Agric. Hokkaido Univ. 49162267R/. Basionym: Sterigmatocystis sydowi 9 J Ann. Mycol111.R676G VOLUME 3 · NO. 1. Aspergillus tabacinus 0 et al J. Agr. Chem. Soc. Japan 1016WW67G2 ? ?RG7R22 ! R 8. Type: Sine loc.1 <

(170) 67G2Y. Nakazawa0;;+ 2W76½ < . 65.

(171) ART I CLE. Jurjevic, Peterson & Horn. Fig. 4. Aspergillus sydowi0;;+.R/ 7 .RÉ 6/ EF

(172) EF # ?%F ?%F # J

(173)

(174) #

(175) 56/(

(176) 56/J

(177)

(178)

(179) 56/. Fig. 5. Aspergillus tabacinus0;;+2W76 7 .RÉ 6/ EF

(180) EF. # ?%F ?%F # J # #

(181) 56/@

(182) =

(183) 56/. Description: Colonies 6/ EF .R É ! R 8

(184) G/8G. .8G #

(185) ;2W

(186) < when present, no soluble pigment, reverse uncolored in 0;;+ R/G6

(187) E 6/. 66. ?%F .R É ! R8 6W8G/ 0;;+2W76 # # 0;;+ R/G6 0;;+ V.2Q6 $

(188) = <

(189) # # "

(190) W EF R É GW É IMA FUNGUS.

(191) Aspergillus section Versicolores. ART I CLE Fig. 6. Aspergillus versicolor0;;+.GQ 7 .RÉ 6/ EF

(192) EF # ?%F ?%F #

(193) 5R/J

(194)

(195) #

(196) 56/@

(197)

(198) 56/. Stipes ! R

(199) W/8G//8W//87// 28Q87 vesicles R8Q86R8.. conidial heads biseriate, metulae covering half to entire # G8Q87 .R82R8RR phialides R8Q866 .8G8GR

(200)

(201) conidia ! R

(202) .R8G828W . Aspergillus versicolor B &

(203) Annali Bot717 67/Q ? ?6W.6R7 ! V 8 Basionym: Sterigmatocystis versicolorB in? Thèse de médicine0 2716R67/G Type: Sine loc.:67GRV. K. Charles0;;+.GQ½ < Description: Colonies 6/EF .RÉ! V 8

(204) .Q8GV 28R sporulating well, conidial heads pale grayish green near ;2W <

(205) #

(206) # soluble pigment, reverse vinaceous or brown or scarlet 0;;+ .GQ E 6/ ?%F .R É ! V8 .68G6 aggregates, sometimes dominating colony appearance, sporulating in pale to dark bluish green to gray green color, <

(207) not present in others, reverse pale yellow, yellow orange VOLUME 3 · NO. 1. "

(208) W EF R É "

(209) WEF GWÉ Q Stipes ! V 8

(210)

(211) 2R8.//8WR/8 6/R/ 28R8Q86. vesicles pyriform to spatulate, V8786W8./ conidial heads biseriate, metulae # # G8V87.R82R phialides 28R8W866 .8G

(212) conidia ! V

(213) .8 .R8GR8VR hülle cells globose,

(214) 0;;+R.67 ?2/

(215) Observations: & <= 0;;+.GQ 67GR ' of dark red soluble pigment and scarlet colony reverse on CYA, from the more recent isolates that were placed in the F;JE E

(216) 67W/ 67Q2& . 0;;+R.670;;+6G6220;;+6G62R 0;;+ 6G62V '

(217) F # A. versicolor based N0F ' . New species Aspergillus austroafricanus > # JL( 9LP sp. nov. ? ?Q//R7W ! W 8 Etymology: " JF . 67.

(218) ART I CLE. Jurjevic, Peterson & Horn. Fig. 7. Aspergillus austroafricanus0;;+.GG 7 .RÉ 6/ EF

(219) EF. # ?%F ?%F # J

(220)

(221) #

(222) 56/ @

(223) =

(224) 56/. Fig. 8. Aspergillus creber0;;+RQRQG 7 .RÉ 6/ EF

(225) EF # ?%F ?%F # J

(226)

(227) #

(228) 56/@

(229)

(230) 56/. Type: South Africa1 E E & 67.. V. A. Putterill (" QQ/762 8 0;;+.GG W .RÉEF ?%F Diagnosis: E = GW É

(231)

(232) EF Description: Colonies 6/EF .RÉ! W 8

(233) .G8.2 . 68. overgrowth by clumped hyphae making surface appear mealy, sporulating well, conidial heads near sage green ;2W <

(234)

(235) . #

(236) E 6/?%F .RÉ ! W8 .W #

(237) <

(238) # "

(239) W EF R É GW É IMA FUNGUS.

(240) Aspergillus section Versicolores. ART I CLE Fig. 9. Aspergillus cvjetkovicii0;;+2V2. 7 .RÉ 6/ EF

(241) EF. # ?%F ?%F # 0

(242)

(243) 5R/J

(244)

(245) #

(246) 56/@

(247)

(248) 56/@

(249)

(250) 56/. Stipes ! W 2/86//8GR/8R//G8R8Vvesicles pyriform to 28V86.86R conidial heads biseriate, metulae# 63G # G8W87.R82R phialides28R8W87.8.R8G82

(251) present, conidia ! W

(252) .R8GR 82R . Aspergillus creber > # J L( 9 L P sp. nov. ? ?Q//R7Q ! Q 8 Etymology1 ! + creber meaning numerous ' Type: USA: California1 < 0# .//Q Z. Jurjevic (" Q//76. 8 X 0;;+RQR7. W .RÉEF ?%F. ;2W ca 6

(253) 0;;+ .G6 with vivid brown soluble pigment, other isolates no soluble < # #

(254)

(255) "

(256) WEF RÉ GWÉ Stipes! Q 6/8W/82R/8VR/<G8 28W8Qvesicles pyriform to spatulate and occasionally

(257)

(258) 28W86W8.R conidial heads biseriate, metulae G828V8Q < .R82R8R phialides 28R8Q8 6/<.8G82conidia! Q spherical to subspherical, .R8G8287 . . Aspergillus cvjetkovicii > # JL( 9 LP sp. nov. ? ?Q//R77 ! 7 8 Etymology: 0 E# # ¯M # T

(259) X ËÌ =[=ÌÌ# = Ë. Diagnosis: ( = GW É

(260) EF ?%F EF ?%F. Type: USA: New Jersey: < 676RW. Wilson(" QQ/7/78 0;;+..W W .RÉEF ?%F. Description: Colonies 6/ EF .R É ! Q 8

(261) 6Q8.V G8R mm centrally, peripheral areas white or yellow, central area sporulating well, conidial heads pea green to artemisia green ;2W <

(262) pigment, reverse clay colored to cinnamon or reddish brown ;.7 E 6/ ?%F .R É ! Q8 6Q8.. 68. overgrown centrally with hyphae aggregated into funicles, = . Diagnosis: ( GWÉ <

(263) EF. VOLUME 3 · NO. 1. Description: Colonies 6/EF .RÉ! 7 8

(264) .28.7 .8G ;2W 0;;+ RQR7G <

(265)

(266) cinnamon, reddish brown soluble pigment, reverse yellowish. 69.

(267) ART I CLE. Jurjevic, Peterson & Horn. Fig. 10. Aspergillus fructus0;;+.G7 7 .RÉ 6/ EF

(268) EF # ?%F ?%F # J #

(269) 56/@

(270)

(271) 56/@

(272)

(273) 56/. ;.7 # ;G7 E 6/ ?%F .R É ! 78 6W8GV 0;;+ RQR7G . 0;;+ ..W 0;;+ .G/

(274)

(275) 0;;+2V2. RQR7G produce soluble pigment, reverse brownish orange or pale "

(276) W EF R É GW É Stipes ! 7 8 2/8.//8W//8QR/ G828W8Q vesicles pyriform .

(277) R8786Q8.G conidial heads biseriate, metulae covering half to entire # G8V8Q .R82R phialides R8Q86/ .8G8 2 G.

(278) occasionally present, conidia ! 7

(279) .8.R8GR8R spinulose, hülle cells! 7

(280) . Aspergillus fructus > # JL( 9L P sp. nov. ? ?Q//V// ! 6/ 8. Description: Colonies 6/ EF .R É ! 6/ 8

(281) .78G7 28R

(282) ;2W <

(283)

(284) 0;;+.G7 # = ;. E 6/ ?%F .R É ! 6/8 ..8G. diam, slightly sulcate, centrally covered by hyphal tufts, sporulation in yellow-green hues near artemisia green ;2W <

(285) #

(286) 0;;+.26 $ ?%F"

(287) WEF RÉ "

(288) W EF GW É 2 Stipes! 6/ R/8 6R/82//8R// 28W vesicles pyriform to spatulate, V8786W8.6 conidial heads biseriate, metulae # # .8G8W87.R82R8W phialides R8V8Q866 .8G82 .

(289)

(290) conidia ! 6/

(291) .8 .R8GR82R hülle cells! 6/

(292) . Etymology:! Type: USA: California1 < 67G7 Bliss (" QQ/76R8 0;;+.G7 W .RÉEF ?%F Diagnosis: Resembling A. versicolor GW É

(293) 6R/82//# .//8WR/ A. versicolor.. 70. Aspergillus jensenii > # JL( 9L P sp. nov. ? ?Q//V/6 ! 66 8 Etymology: 0 E 0 > reported this species as Aspergillus globosus> homonym of A. globosus+ . IMA FUNGUS.

(294) Aspergillus section Versicolores. ART I CLE Fig. 11. Aspergillus jensenii0;;+RQVW6 7 .RÉ 6/ EF

(295) EF # ?%F ?%F # J

(296)

(297) #

(298) 56/(

(299) 56/@

(300)

(301) 56/. Type: USA: Montana1 < [ .//Q Z. Jurjevic("QQ/76/8 0;;+ RQV// W .RÉEF ?%F Synonym: Aspergillus globosus > E M # F %< J G6R12Q.676.X + 6Q/7 Diagnosis1 E GW É conidial color near celandine, tawny olive to dark umber # EF Description: Colonies 6/ EF .R É ! 66 8

(302) ./8.W raised or sunken, with clumped hyphal aggregates common in some isolates, sporulating moderately well, conidial heads ;2W <

(303)

(304) soluble pigment faint or intense yellow brown, in one case.

(305) # # ;G7

(306)

(307) ;GE 6/?%F .RÉ! 668 6W8G/ have funicular tufts of aerial hyphae centrally, sporulating well

(308) = <

(309) pigment either light brown or reddish brown, brownish orange in one isolate, reverse pale yellow or orange or brownish. "

(310) WEF RÉ GWÉ Stipes ! 66

(311) 2R8.//8W//86/// 8G28W8Q vesicles pyriform to spatulate, rarely

(312) R8W86V8.. conidial heads biseriate, metulae# 63G # G8Q.R8 28R phialides 28R8Q866 .8G . VOLUME 3 · NO. 1. G. 2R

(313) ! 66 conidia! 66 spherical to

(314) .R8G8 2R8W

(315) 6R8./

(316) 0;;+RQRQ.

(317) . Aspergillus puulaauensis > # JL( 9 LP sp. nov. ? ?Q//V/. ! 6. 8 Etymology" (O O P P X ËM=M= =M= = Ë Type: USA: Hawaii1 <

(318) .//G D.T. Wicklow("QQ/7668 0;;+GRV26 W .RÉEF ?%F Diagnosis: "

(319) ?2/ GWÉ Description: Colonies 6/EF .RÉ! 6. 8

(320) ..8.R R8V with funicular hyphal clumps, sporulation light, conidial heads. ;2W < soluble pigment when present brown, reverse yellowish to clay ;G7 ;.7E 6/?%F .RÉ! 6.8 .68.R # ;2W <

(321) # "

(322) WEF RÉ GWÉ Stipes! 6. GR8 6//8R//8W//G828Wvesicles pyriform to spatulate,. 71.

(323) ART I CLE. Jurjevic, Peterson & Horn. Fig. 12. Aspergillus puulaauensis0;;+GRV26 7 .RÉ 6/ EF

(324) EF. # ?%F ?%F # J #

(325) 56/?

(326) 5R/P

(327) 56/@

(328)

(329) 56/.

(330) R8Q86Q8.6 conidial heads biseriate, metulae# # G828 W87.R82phialidesR8W86/.8G

(331) present, conidia ! 6. .R8G8 28RR hülle cells ! 6.8 66867 ?2/ . Aspergillus subversicolor > # JL( 9 LP sp. nov. ? ?Q//V/G ! 6G 8 Etymology1 Aspergillus versicolor Type: India: Karnataka1 <

(332) . 67W/ B. Muthappa("QQ/76Q8 0;;+RQ777 W .RÉEF ?%F. Fig. 13. Aspergillus subversicolor0;;+RQ777 7 .RÉ 6/ EF

(333) EF. # ?%F ?%F # J

(334)

(335) #

(336) 56/J

(337)

(338)

(339) 56/. 72. IMA FUNGUS.

(340) Aspergillus section Versicolores. ART I CLE Fig. 14. Aspergillus tennesseensis0;;+6G6R/ 7 .RÉ 6/ EF

(341) EF # ?%F ?%F # J #

(342) 56/@

(343)

(344) 56/. Diagnosis1E = GWÉ slowly on all media, producing yellow soluble pigment on EF

(345) < Description: Colonies 6/ EF .R É ! 6G 8

(346) 6Q8./ R8V centrally, wrinkled, sporulating sparsely, conidial heads. ;2W <

(347) # ;6R E 6/?%F .RÉ! 6G8 6.862 diam, low, plane, velutinous, sporulating in bluish green color ;2W <

(348) #

(349) "

(350) WEF RÉ GWÉ Stipes ! 6G

(351) V/8 .R/82R/ 8RR/ 28W86/ vesicles

(352)

(353) V86/86W8.. conidial heads biseriate, metulae# . 63G # G828W87.8.R82

(354) .8G phialides,R8Q86/.8G

(355) conidia ! 6G

(356) .R8G828W . Aspergillus tennesseensis > # J L ( 9LP sp. nov. ? ?Q//V/2 ! 62 8 Etymology1" & . Diagnosis1( = GW É ?%F Description: Colonies 6/EF .RÉ! 62 8

(357) ..8G/ mat, radially sulcate, centrally raised or sunken, overgrown by clumps of aerial hyphae in some isolates, sporulating well ;2W <

(358)

(359) # in brownish orange shades near honey yellow or chamois ;G/ E 6/ ?%F .R É ! 628 ./82V # ;2W < no soluble pigment, reverse uncolored, pale lemon yellow,

(360) "

(361) W EF R É GW É Stipes! 62

(362) GR86//8G//82// 28W vesicles W86/86V86Q conidial heads biseriate, metulae# # 28V8Q.R82 phialidesR8Q866.8G

(363) conidia ! 62

(364) .R8G828Q . Aspergillus venenatus > # JL( 9 LP sp. nov. ? ?Q//V/R ! 6R 8 Etymology1( < . Type: USA: Tennessee1 << 67Q2B.W. Horn("QQ/76W8 0;;+ 6G6R/ W .RÉEF ?%F. VOLUME 3 · NO. 1. Type: USA: Tennessee1 << 67Q2 B.W. Horn("QQ/76V8 0;;+ 6G62W W .RÉEF ?%F. 73.

(365) ART I CLE. Jurjevic, Peterson & Horn. Fig. 15. Aspergillus venenatus0;;+6G62W 7 .RÉ 6/ EF

(366) EF. # ?%F ?%F # J #

(367) 56/(

(368) 56/@

(369)

(370) 56/@

(371)

(372) 56/. Diagnosis1( GWÉ <

(373) EF ?%F Description: Colonies 6/ EF .R É ! 6R 8

(374) ..8G6 ;2W

(375) = ;2. <

(376) # #

(377)

(378) ;6RE 6/ ?%F .R É ! 6R8 6W8.2 lightly sulcate, low, central tufted funicular aggregates of aerial hyphae, sporulating well in deep green color near slate ;2W <

(379) # #

(380) "

(381) W EF RÉ GWÉ Stipes ! 6R

(382) ./86//82//8R// 28W vesicles V8786W8.6 conidial heads biseriate, metulae# # G8 28W87 .R828R phialides R8V8Q866 .8GGR

(383) ! 6R hülle cells! 6R present in some isolates, conidia ! 6R

(384) G828V . Phenotypic species recognition Growth rates of species on different media are presented in &

(385) . Phenotypic recognition of species in section Versicolores is based on smooth, roughened or spinulose conidia, conidial <

(386) EF ?%F

(387) GWÉ . 74. Aspergillus cvjetkovicii, A. sydowii and A. venenatus A. sydowii isolates grow GWÉ A. cvjetkovicii and A. venenatus isolates do A. cvjetkovicii < soluble pigment on CYA, while A. venenatus isolates produce <

(388) Aspergillus amoenus, A. austroafricanus and A. tabacinus = [ A. amoenus GW É A. tabacinus isolates produce no soluble pigment and A. austroafricanus produces reddish

(389)

(390) EF The remaining eight species produce conidia with noticeably roughened walls, but the ornamentation is not

(391) & eight species, A. versicolor and A. fructus, have roughened GWÉ& #

(392) # 6R/82// A. fructus # .//8WR/ A. versicolor L < A. fructus # A. versicolor isolates and while separation of these species using phenotype on standard media appears possible, until more

(393) '

(394)

(395) Genealogical concordance species recognition clearly

(396) ! 6 J GWÉ are A. protuberus, A. creber, A. jensenii, A. puulaauensis, A. subversicolor and A. tennesseensis Aspergillus protuberus EF < ;6 #

(397) ?%F $ A. jensenii isolates produce brown CYA colony reverse colors from tawny olive to dark umber, ;2W F A. puulaauensis IMA FUNGUS.

(398) Aspergillus section Versicolores. Table 2.E Versicolores # W"

(399) .RÉ < CYA. MEA. CY20. M40Y. M60Y. CYA at 37 °C. A. amoenus. ./8.7. 6686R. 668./. 628... 6R8.2. V. A. austroafricanus. 6Q867. 6V86W. .28.R. 6Q867. 6W86Q. -. A. creber. 6W8.G. 6686R. 6R8... 6Q8.G. 678.2. -. A. cvjetkovicii. 6R8.6. 6286W. 6W8./. 6V8.2. 6R8.R. -. A. fructus. 6G8./. 6/86V. 786W. 6/8.G. 6/8./. 2. A. jensenii. 6V8./. 786G. 6R8./. .68.V. ..8.Q. -. A. protuberus. 6W8.R. 6686Q. 628... .68.2. ./8.2. -. A. puulaauensis. 6Q8.6. 6686.. 6W867. 678... 6Q8.6. -. A. subversicolor. 6G862. V8W. 6/866. 6R86V. 6V86Q. -. A. sydowii. ./8.R. ./8.R. .68.V. .G8.V. .G8.W. Q. A. tabacinus. .68.V. 6.86V. .68.V. Q8.G. Q8.G. -. A. tennesseensis. ./8... 6.862. 6W867. 678... 678... -. A. venenatus. 6286W. Q86/. 6R86V. 678.G. 6W8.6. -. A. versicolor. ./8.V. 6/86Q. 678... .68.V. ..8.R. Q. ART I CLE. Species. Table 3.(

(400) "&J . "&J Versicolores"&J

(401) .

(402) 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/

(403) [ A. versicolor and A. amoenus

(404) GWÉ A. jensenii ?2/A. subversicolor # ?%F ?2/ &

(405) .

(406) EF A. tennesseensis ?%F # ;2W

(407) = Aspergillus creber

(408) EF ?%F and conidial color on either medium is pea green to sage ;2W There is considerable variation in colony appearance within species and considerable overlap in colony appearance between species, making species separation within section Versicolores " included in this study were propogated in vitro for several VOLUME 3 · NO. 1. #

(409) F isolates, several appear to have mutated and consequently produce colonies that have a wet appearance when grown on CYA or produce only moist aerial aggregates of hyphae with " . N0F ' N0F ' is the most reliable means for identifying species within this section. "&J Versicolores. &

(410) GJ

(411) @ F # different species and genotype D is present in four different " A. creber "K+0 "&J . 75.