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A phylogeny of the Areae (Araceae) implies that Typhonium, Sauromatum, and the Australian species of Typhonium are distinct clades

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Abstract and Figures

With in excess of 70 species, the Southeast Asian/Australian genus Typhonium is the largest genus of the Areae, a tribe that includes up to nine smaller genera of which Sauromatum and Lazarum have recently been reduced to the synonymy of Typhonium. To test the circumscription and relationships of Typhonium to the other Areae, we used chloroplast and nuclear DNA sequences (4319 aligned nucleotides) for 86 of the total 153 species, including representatives of all relevant genera. In the resulting phylogeny, Typhonium species fall into three well-supported clades: the first comprises most Typhonium species, including the type, T. trilobatum ; the second consists of the type of Sauromatum, S. guttatum, and other species formerly placed in that genus; the third includes only Australian endemics. Each of the remaining six genera of Areae is monophyletic. Sauromatum and Typhonium are not sister groups, requiring the recognition of Sauromatum. The Australian clade also needs to be ranked as a genus to achieve similar levels of morphological, geographic, and genetic distinctness among the genera of Areae. However, since only 10 of the 16 described Australian endemics currently placed in Typhonium have so far been sequenced, not including the type of the name of the Australian genus Lazarum, we refrain from applying this name to the Australian clade. Among the nomenclatural and taxonomic results of this study are a key to the nine species of Sauromatum, and five new combinations. We also report two new chromosome counts and discuss the implications of the molecular phylogeny for the evolution of Sauromatum karyotypes.
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INTRODUCTION
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A p h y l o g e n y o f t h e A r e a e ( A r a c e a e ) i m p l i e s t h a t Typhonium,
Sauromatum, and the Australian species of Typh onium are
distinct clades
Natalie Cusimano,1 Matthew D. Barrett,2,3 Wilbert L.A. Hetterscheid4 & Susanne S. Renner1
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Abstract
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Keywords
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MATERIALS AND METHODS
Taxon sampling and sequencing. —
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+N,+,-. %/0 # &,)7?,7?67?&?*@?K,AF,@M+,+7,@>7??,)BB?&&'+*&/,5&,
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7+6.)&@,6>F.+D?*F,+N,W?**?7,-,X>)*D,2388!4/,=>'&,@+@).&,@+,ZZ,
&6?B'?&G,7?67?&?*@?K,AF,#Z,)BB?&&'+*&/, 5..,&6?B'?&,M'@>,@>?'7,
&+%7B?&,)*K,>?7A)7'%(,C+%B>?7&,2M>?7?,)66.'B)A.?4,)7?,.'&@?K,
'*,@>?,566?*K'U/
=+,K?K%B?,6>F.+D?*?@'B,7?.)@'+*&>'6&G,M?,7?.'?K,+*,),*%B.?)7,
.+B%&G,@>?, 6>F@+B>7+(?,$, D?*?, 2I.,N4G,)*K,@M+,B>.+7+6.)&@,
.+B'G,@>?,$-9HD1$-8SH('*@?7D?*'B,&6)B?7,)*K,@>?,@WQ5),8(D?*?,
2'$/Y4G,M>'B>,B+*@)'*&,),D7+%6,],'*@7+*,@>)@,?*B+K?&,@>?,()@%J
7)&?,^,2&"'Y4,+6?*,7?)K'*D,N7)(?/,=+@).,SQ5,N7+(,&'.'B)JK7'?K,
.?)C?&,M)&,?U@7)B@?K,M'@>,@>?,Q%B.?+P6'*,6.)*@,R'@,)BB+7K'*D,
@+,@>?,()*%N)B@%7?7b&,67+@+B+.,2I)B>?7?FJQ)D?.G,Sh7?*G,g?7J
()*F4/,P?_%?*B'*D,+N,@>?, B)/, 3"88, *%B.?+@'K?, 2*@4J.+*D,'$/Y(
()7R?7G,)(6.'N'?K,'*,+*?,6'?B?,M'@>,@>?,67'(?7,6)'7,@7*^J"#9!V,
2K'B+@4;@7*^J9TW,2`+>*&+*, -,P+.@'&G,9##!4G,M)&,67+A.?()@'B/,
$+*&?_%?*@.FG,M?, K?&'D*?K,*?M, '*@?7*)., 67'(?7&, )*K, )(6.'J
N'?K,@>?,&?B@'+*,'*,@M+,6'?B?&:,@7*^J"#9!V;@7*^JWI,)*K,@7*^J
VI;,@7*^J9TW/,=>?,*?M,67'(?7,&?_%?*B?&,)7?,)&,N+..+M&:,@7*^,
JWI, 1ƍJ55g5=g==g5=$g=555=55g5ggJ"ƍ,)*K, @7*^,
JVI,1ƍJg====g$=g=$5==5=gg555==$$J"ƍ/,I.,N(M)&,
).&+,)(6.'N'?K,'*, @M+,6'?B?&,M'@>,@>?, *?M.F,K?&'D*?K, 67'(J
?7&:,538V;<18W,)*K,!"8V;5W:,538V:,1ƍJ$5$=$55=$$=5
$55 5$=gg$J"ƍG,<18W:,1ƍJ5g5=$$5=55$5===gg=g5
==g=J"ƍG,!"8V:,1ƍJ$=$g=g5=g=$=g=$5$55=55gJ"ƍ,)*K,
5W:,1ƍJg55=5g$5=$$5===$55$5=$J"ƍ/,=>?,$-9HD1$-8SH(
'*@?7D?*'B,&6)B?7,M)&,)(6.'N'?K,%&'*D,@>?,67'(?7&,)*K,i$W,
B+*K'@'+*&,K?&B7'A?K,'*,W?**?7,-,X>)*D,2388!4/
i+.F(?7)&?,B>)'*, 7?)B@'+*&,2i$W4,M?7?, 6?7N+7(?K, M'@>,
98,I,67'(?7&,'*,31,.,7?)B@'+*&G,%&'*D,L'+=>?7(,SQ5,6+.FJ
(?7)&?,2g?*?B7)N@G,[hK'*D>)%&?*G,g?7()*F4/,=>?,'*'@').,&@?6,
+N,1,('*,)@,#1j$,M)&,N+..+M?K,AF,"1,BFB.?&,+N,#1j$,N+7,"8,&,
N+7,SQ5,K?*)@%7)@'+*G,T8j$,N+7,T8,&,N+7,67'(?7,)**?).'*DG,)*K,
<3j$,N+7,3, ('*, )*K,!8,&, N+7,67'(?7,?U@?*&'+*/,i$W,67+K%B@&,
M?7?, B+*@7+..?K,AF, ?.?B@7+6>+7?&'&, +*,)*, ?@>'K'%(,A7+('K?J
&@)'*?K, 9k,)D)7+&?,D?.,M'@>,@>?, [)(AK),SQ5,&'d?,()7R?7/,
i$W,67+K%B@&,M?7?,6%7'N'?K,%&'*D, ?'@>?7, i7+(?D), Y'd)7K
l
,
Pm,g?.,)*K,i$W,$.?)*Jn6,PF&@?(,+7,5D?*B+%7@,5Ii%7?l,
i$W,6%7'N 'B)@'+*,R'@,)*K,_%)*@'N'?K,?.?B@7+6>+7?@'B)..FG,%&'*D,
[)(AK),SQ5,)&,&@)*K)7K/,]N,(%.@'6.?,A)*K&,M?7?,K?@?B@?KG,)*,
)KK'@'+*).,?.?B@7+6>+7?&'&,M)&,6?7N+7(?K,@+,?UB'&?,)*K,)*).Fd?,
@>?(,&?6)7)@?.F/,P?_%?*B'*D,7?.'?K,+*,L'D,SF?,=?7('*)@+7,R'@&,
!!9
$%&'()*+,-,)./,0,Phylogenetics of Typhonium and SauromatumTA XO N,1#,234,0,567'.,3898:,!"#;!!<
2566.'?K,L'+&F&@?(&G,Y)7 7'*D@+*G,n/^/4,)*K,@>?,)(6.'N'B) @'+*,
67'(?7&/,$FB.?,&?_%?*B'*D,67+K%B@&,M?7?,B.?)*?K,AF,P?6>)K?U,
gJ18,P%6?7N'*?,D?.,N'.@7)@'+*,25(?7&>)(G,n66&).),PM?K?*4,+*,
I%.@'PB7??*, =IJEm,(?(A7)*?, 6.)@?&,2I'..'6+7?G,L?KN+7KG,
n/P/5/4,)BB+7K'*D,@+,@>?, ()*%N)B@%7?7&b,67+@+B+.&,@+,7?(+C?,
%*'*B+76+7)@?K,*%B.?+@'K?&/,V7)D(?*@&,M?7?,&?6)7)@?K,+*,)*,
5L],"988,5C)*@,B)6'..)7F,&?_%?*B?7G,)&&?(A.?K,)*K,?K'@?K,%&J
'*D,@>?,&+N@M)7?,P?_%?*B>?7,2g?*?,$+K?&G,5**,57A+7G,I'B>'J
D)*G,n/P/5/4G,)*K,L[5P=J&?)7B>?K,'*,g?*L)*R/,P?_%?*B?&,)7?,
K?6+&'@?K,'*,g?*L)*R,2N+7,)BB?&&'+*,*%(A?7&,&??,566?*K'U4/
Alignments and phylogenetic analyses.
,5.'D*(?*@,,&,
2=)A.?,94,M?7?,D?*?7)@?K,'*,I)B$.)K?,2I)KK'&+*,-, I)KK'J
&+*G,9##34,)*K,B+*@'*%+%&.F,)Ko%&@?K,()*%)..F/,=>?,@>7??,K)@),
6)7@'@'+*&,M?7?,N'7&@,)*).Fd?K,&?6)7)@?.FG,)*K,'*,@>?,)A&?*B?,+N,
&@)@'&@'B)..F,&%66+7@?K,@+6+.+D'B).,B+*@7)K'B@'+*&,2ZZ8k4G,@>?F,
M?7?,@>?*,B+(A'*?K/
i>F.+D?*?@'B,'*N?7?*B?,+N,@>?,B+(A'*?K,K)@),2!"9#,).'D*?K,
*%B.?+@'K?&4,7?.'?K, +*, ()U'(%(,.'R?.'>++K,2I[4,)&,'(6.?J
(?*@?K,'*, @>?, W5UI[,L.)BRL+U,2P@)()@)R'&, -, )./G,388ZG,
>@@6:cc6>F.+A?*B>/C'@).J'@/B>c7)U(.JAAc4,)*K,+*,L)F?&')*,)*).FJ
&'&,)&,'(6.?(?*@?K,'*,I7L)F?&,C/"/9/3,2E%?.&?*A?BR,-,W+*J
_%'&@G,3889H,W+*_%'&@,-,E%?.&?*A?BRG,388"4/,L++@&@7)66'*D,
%*K?7, I[, %&?K,9888,7?6.'B)@?&, 6?7N+7(?K, '*,W 5UI[/, 5..,
&?)7B>?&,7?.'?K,+*,@>?,g=W,p,g,(+K?.G,M'@>,(+K?.,6)7)(?@?7&,
?&@'()@?K,K%7'*D,7%*&/,2W5UI[,%&?&,@>?,g=W$5=,)667+U'()J
@'+*,+N,@>?,g=W,p,g,(+K?.G,M'@>,@>?,D)((),&>)6?,6)7)(?@?7,
>)C'*D,31,7)@?,B)@?D+7'?&/4,
L)F?&')*, 7 %*&, M?7?, &@) 7@?K, N7+( , '*K?6?*K?*@, 7 )*K+(,
&@)7@'*D, @7??&,)*K,7?6?)@?K, N+%7, @'(?&/,I)7R+C,B>)'*,I+*@?,
$)7.+,7%*&,?U@?*K?K,N+7,3,('..'+*,D?*?7)@'+*&G,M'@>,@7??&,&)(J
6.?K,?C?7F,3888@>(D?*?7)@'+*,27?&%.@'*D,'*,9889,@7??&,N+7,?)B>,
7%*4/,Y?,%&?K,),N .)@,S'7'B>.?@,67'+7,N+7,@>?,7?.)@'C?,*%B.?+@'K?,
N7?_%?*B'?&, )*K,7)@?,6)7)(?@?7&G,),K'&B7?@?,%*'N+7(,67'+7,N+7,
@+6+.+D'?&G,)*K,)*, ?U6+*?*@').,K'&@7'A%@'+*,2(?)*,984,N+7,@>?,
ȖJ&>)6?,6)7)(?@?7,)*K,)..,A7)*B>,.?*D@>&/,$+*C?7D?*B?,M)&,
)&&?&&?K,AF,B>?BR'*D,@>)@,294,N'*).,.'R?.'>++K&,)*K,()o+7'@F,
7%.?,@+6+.+D'?&,)(+*D,7%*&,M?7?,&'('.)7H,234,@>?,&@)*K)7K,K?J
C')@'+*&,2PS4,+N,&6.'@,N7?_%?*B'?&,M?7?,[8/89H,2"4,@>?,B+*C?7J
D?*B?,K')D*+&@'B,2@>?,6+@?*@').,&B).?,7?K%B@'+*,N)B@+7,D'C?*,AF,
I7L)F?&4,)667+)B>?K,9H,)*K,2!4,AF,?U)('*'*D,@>?,6.+@,67+C'K?K,
AF,I7L)F?&,+N,@>?,D?*?7)@'+*,*%(A?7,C?7&%&,@>?,.+D,67+A)A'.'@F,
+N,@>?,K)@)/,=W5$OW,2W)(A)%@,-,S7%((+*KG,388<4,M)&,%&?K,
@+,)&&?&&,M>?@>?7,7%*&,>)K,7?)B>?K,B+*C?7D?*B?/,=7??&,&)C?K,
67'+7,@+,B+*C?7D?*B?,M?7?,K'&B)7K?K,)&,A%7*J'*,2988,@7??&4,)*K,
),B+*&?*&%&,@7??,B+*&@7%B@?K,N7+(,@>?,7?()'*'*D,"T8!,@7??&/
Chromosom e counts. —
,$>7+(+&+(?,*%(A?7&,M?7?,+AJ
@)'*?K,N+7,!4('3/'":#9"'#&(2.'C'*D,6.)*@,*%(A?7,E/5W/8!34,)*K,
!4(.0$8#'#&(2E/5W/8"T4,N7+(,'*K'C'K%).&,B%.@'C)@?K,AF,@>?,@>'7K,
)%@>+7,'*,@>?,A+@)*'B).,D)7K?*, +N,Y)D?*'*D?*/, \NN&67'*D,+N,
@>?&?,)BB?&&'+*&,)7?,*+M,2388#4,'*,B%.@'C)@'+*,)@,@>?,I%*'B>,
L+@)*'B).,g)7K?*/,O)B>,B+%*@,'&,A)&?K,+*,98,@+,91,*%B.?',N7+(,
+*?,'*K'C'K%)./,=+,+A@)'*,D++KJ_%).'@F,B>7+(+&+().,&67?)K&G,
7++@,@'6,(?7'&@?(&,M?7?,B+..?B@?K,'*,@>?,(+7*'*DG,67?@7?)@?K,N+7,
";1,>+%7&,M'@>,B+.B>'B'*?,)@,!j$G,N'U?K,M'@>,?@>)*+.J)B?@'B,)B'K,
2",:௘௘94G,)*K,& @ +7?K ,) @,;38 j$,%*@' . ,% & ?/,V+7,A) &'B ,R ) 7 F +@F6?,)&& ? &&J
(?*@G,>FK7+.Fd?K,(?7'&@?(&,M?7?,&@)'*?K,M'@>,PB>'NNb&,7?)D?*@G,
&_%)&>?K,%*K?7,),B+C?7,&.'6G,)*).Fd?K,%*K?7,),.'D>@,('B7+&B+6?G,
)*K,K+B%(?*@?K,%&'*D,),K'D'@).,'()D?,B)6@%7?,&F&@?(/
RESULTS
Phylogeny. —
(=>?,B+(A'*?,,,K,&?_%?*B?,()@7'U,2!"9#,B>)7J
)B@?7&G,#Z,)BB?&&'+*&H,=)A.?,94,F'?.K?K,),M?..J&%66+7@?K,57?)?,
B.)K?,'*, M>'B>, ).., D?*?7),M'@>, (+7?, @>)*,+*?,&6?B'?&, M?7?,
(+*+6>F.?@'BG,?UB?6@, N+7, +,-. % /0# &G,M>'B>,&6.'@,'*@+, @>7??,
M?..J&%66+7@?K,B.)K?&,2V'D/,94/,=>?, .)7D?&@,+N,@>?&?G,&'&@?7,@+,
@>?,7?()'*'*D,D?*?7)G,B+*@)'*&,"9,+N,@>?,13,&)(6.?K, &6?B'?&,
+N, +,-.% /0# &G,'*B.%K'*D, @>?, @F6?,+4 ( '$ 09 %=" ' #&/, =>?,&?B+*K,
a+,-.% /0# &b,B.)K?,'*B.%K?&,+*.F,5%&@7).')*,?*K?('B&,2V'D/,94H,
M?,7?N?7,@+,@>'&,B.)K?,)&,@>?,5%&@ 7).')*,B.)K?/,Y'@>'*,@>?,5%&@7)J
.')*,B.)K?G,@>?7?,)7?,@M+,&@)@'&@'B)..F,&%66+7@?K,&%AB.)K?&G,+*?,
B+*@)'*'*D,+4("908&0;%90#&G(+4(O09=3$'00,)*K,+4("/5#8'09%=0#&,
2@>?,.)@@?7,*+@,(+*+6>F.?@'B4G,)*K,+*?,B+*@)'*'*D,+4(/#60=":1
:"'#&G(+4(-$"3'3$&088#&,)*K, @>?, %*K?&B7'A?K, &6?B'?&,+4, &6/,
^%*%*%77)G,+4,&6/,i7'*B?,W?D?*@,)*K,+4,&6/,I+7D)*,W'C?7/,=>?,
6>F.+D?*?@'B,6+&'@'+*&,+N,+@>?7,5%&@7).')*,@)U), )7?,*+@,M?..,
7?&+.C?K/,=>?,*?U@JA7)*B>'*D,B.)K?,'&,+.3$0%-.%/#&G(N+..+M?K,
AF,@>?, @>'7K(a+,-.% /0#&b(B.)K?G,M>'B>,'*B.%K?&,)..,&6?B'?&,)@,
+*?,@'(?,6.)B?K,'*G(+7,(+76>+.+D'B)..F,&'('.)7, @+G,!"#$%&"1
'#&,2V'D/,94/,Y'@>'*,!"#$%&"'#&G,@>?7?,)7?,@>7??,&@)@'&@'B)..F,
&%66+7@?K,B.)K?&G,+*?,+N,!4(.0$8#'#&()*K(!4('3/'":#9"'#&G(+*?,
+N,!4( =$370-38()*K,!4(73/%8#&G()*K,+*?,B+(67'&'*D(!4(6073$1
80;%90#&G, !4(5"%905%/53/83G()*K,!4(.%$8;039600/,=>?,&'&@?7,7?J
.)@'+*&>'6,A?@M??*,@>?,.)@@?7,@M+,'&,).&+, M?..,&%66+7@?K/,=>?,
?*@'7?,!"#$%&"'#&(B.)K?,'&,&'&@?7,@+,),M?..J&%66+7@?K,B.)K?,+N,
@>?,7?()'*'*D,N'C?,D?*?7),+N,57?)?G,M>'B>,)7?,B?*@?7?K,'*,@>?,
I?K'@?77)*?)*,7?D'+*/
Chromosome numbers.
,V+7 ,!"#$%&"(('#&G(M?,+A@)'*?K,
*?M,B+%*@&, +N,H/,q,3T,N+7,!4(.0$8#'#&()*K,!4('3/'":#9"'#&/,
=+D?@>?7,M'@>,@>?,+@>?7,B+%*@&,)C)'.)A.?,N+7,@>?,D?*%&,2V'D/,3,
)*K,S'&B%&&'+*4G,),A)&?,*%(A?7,+N,<(\,9",B)* ,A?,' *N?77? K G,%* K?7,
@>?,)&&%(6@'+*,@>)@,>'D>?7,*%(A?7&,'*,@>?,D?*%&,)7?,@?@7)6.+'K,
+7,+B@+6.+'K/
Table .
,P'd?&,+N,@>?,'*K'C'K%).,)*K,B+(A'*?K,B>.+7+6.)&@,)*K,*%B.?)7,K)@),()@7'B?&/
5.'D*?K,*%B.?+@'K?& 5BB?&&'+*&,
SQ5,.+B%& =+@)., OUB.%K?K ]*B.%K?K =+@). ]*D7+%6 \%@D7+%6
'$/Y 3<9# 9T# 2550 94 #3 3
$-9HD1$-8SH , Z<3 933 750 58 1T 3
I.,N 99#3 9<" 1019 51 !# 3
$+(A'*?K,K)@) 4319 98 #T 3
!!3
TA XO N,1#,234,0,567'.,3898:,!"#;!!<$%&'()*+,-,)./,0,Phylogenetics of Typhonium and Sauromatum
Fig. .
,I)U'(%(,.'R?.'>++K,
6>F.+D?*F,N+7,#T,)BB?&&'+*&,
7?67?&?*@'*D,ZT,&6?B'?&,+N,@>?,
*'*?,D?*?7),+N,57?)?,A)&?K,
+*,*%B.?)7,)*K,B>.+7+6.)&@,
&?_%?*B?&,2!"9#,).'D*?K,*%B.?+J
@'K?&4/,m).%?&,)A+C?,A7)*B>?&,
7?N?7,@+,6+&@?7'+7,67+A)A'.'@'?&,
N7+(,L)F?&')*,'*N?7?*B?,23,('.J
.'+*,D?*?7)@'+*&4G,@>+&?,A?.+M,
A7)*B>?&,@+,A++@&@7)6,&%66+7@,
26?7B?*@)D?&,+N,9888,7?6.'B)@?&4,
%*K?7,()U'(%(,.'R?.'>++K/
Dracunculus canariensis
Dracunculus vulgaris
Arum korolkowii
Arum sp. nov. 26940
Arum italicum
Arum nigrum
Arum purpureospathum
Arum sp. nov. I 21 05 M.N.
Arum balansanum
Arum cylindracaeum
Arum maculatum
Arum cyrenaicum
Arum concinnatum
Arum rupicola
Arum dioscoridis
Arum orientale
Arum creticum
Arum megobrebi
Arum hygrophilum
Arum pictum
Biarum pyramii
Biarum davisii
Biarum ditschianum
Biarum straussii
Biarum bovei
Biarum carduchorum
Biarum tenuifolium
Biarum dispar SBL 564
Biarum kotschyi
Helicodiceros muscivorus
Eminium jaegeri
Eminium spiculatum
Sauromatum diversifolium
Sauromatum gaoligongense
Sauromatum horsfieldii
Sauromatum venosum
Sauromatum brevipes
Sauromatum giganteum
Sauromatum tentaculatum
Sauromatum hirsutum
Theriophonum infaustum
Theriophonum dalzellii
Typhonium eliosurum
Typhonium brownii H.AR.043
Typhonium aff. brownii AF4782
Typhonium aff. brownii BG9276
T peltandroides
Typhonium aff. liliifolium
Typhonium russell-smithii
Typhonium sp. PrinceRegent MDB1033
Typhonium sp. MorganRiver
Typhonium sp. PrinceRegent RLB1716
Typhonium nudibaccatum lobed
Typhonium nudibaccatum ovate
Typhonium praetermissum
Typhonium sp. Kununurra RLB3069
Typhonium sp. Kununurra MDB2264
Typhonium alismifolium
Typhonium angustilobium BG9277
Typhonium wilbertii
Typhonium wilbertii AF2544
Typhonium angustilobium HS2007/32
Typhonium flagelliforme Baume 1
Typhonium flagelliforme
Typhonium flagelliforme 174
Typhonium digitatum
Typhonium sp. nov. H.AR.532
Typhonium violifolium 2
Typhonium cordifolium
Typhonium sp. nov. H.AR.555
Typhonium glaucum
Typhonium corrugatum sp. nov.
Typhonium huense
Typhonium lineare
Typhonium gallowayi
Typhonium griseum
Typhonium orbifolium
Typhonium reflexum
Typhonium pedunculatum
Typhonium varians
Typhonium tubispathum
Ty ph o n iu m ec h i nu l a tu m
Typhonium adnatum
Typhonium circinnatum
Typhonium roxburghii
Typhonium aff. saraburiense
Typhonium trilobatum
Typhonium gracile
Typhonium sp. nov. H.AR.543
Typhonium albidinervum
Typhonium baoshanense
Typhonium jingpingense
Typhonium blumei
Typhonium subglobosum
Typhonium filiforme 1
Typhonium bachmaense
Arisaema tortuosum
Arisaema speciosum
100
100
100
98
100
96
100 100
100
100
100
98
100
99
99
100
96
100
100 98
99
96
100
100
1
1
1
1
1
1
0.99
1
1
0.98
1
1
0.99
1
11
1
0.99
1
11
1
1
1
1
1
1
10.99
1
11
1
1
1
1
1
1
1
1
0.98
0.98
1
1
1
Australian clade
Mediterranean clade
Sauromatum
Typhonium
!!"
$%&'()*+,-,)./,0,Phylogenetics of Typhonium and SauromatumTA XO N,1#,234,0,567'.,3898:,!"#;!!<
DISCUSSION
Phylogeny. —
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K?67?&&?KJD.+A+&?,B+7(&,)&,N+%*K,'*,(+&@,+@>?7,5%&@7).')*,@)U)/
Leaf shape
Spathe tube
Types of staminodes
Chromosome numbers
26
26
52
26, 52, 104
26
26
52
Distribution
Thailand
Thailand
Afghanistan to China
Afghanistan to China,
tropical Africa
S China
Indonesia, Thailand,
S China, Myanmar
Himalaya to SE Asia
N Thailand, Yunnan (China)
Nepal
pedatifid/
pedatisect
simple
connate
convolute
2 types
1 type
upper sterile part rugose
or with minute processes
96
Fig. .
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>)&,A??*,.+&@,@M'B?,2'*,!4(505"/'3#&()*K(!4(6073$80;%90#&4/
Chromosome numbers in Typhoni um, Sauromatum, and
the Australian clade.
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'(6+7@)*@,7+.?,'*,@>?,?C+.%@'+*,+N,+,- .%/0 #&G,.?)K'*D,@+,@>?,?UJ
'&@'*D,C)7'?@F,+N,B>7+(+&+(?,*%(A?7&/,=>?,K7)&@'B,7?K%B@'+*,+N,
B>7+(+&+(?,*%(A?7,B+%.K,>)C?,>)66?*?K,@>7+%D>,B>7+(+&+(?,
N%&'+*G,@7)*&.+B)@'+*&,+N,B>7+(+&+(?,6)7@&,)*Kc+7,.+&&,+N,SQ5/
V+7,@>?,5%&@7).')*,B.)K?G,+*.F,@M+,B>7+(+&+(?,*%(A?7&,
>)C?,A??*, 7?6+7@?KG, A+@>,?U@7?(?.F,>'D>:,+4(390%8#$#&( 23/, t,
9884()*K,+4(=$%O/00,23/,q,9T8H,L7'DD&,'*,OC)*&G,9#T94/,5.+*D,
M'@>,)*,%*K?&B7'A?K,&6?B'?&,N7+(,Q?M,P+%@>,Y).?&G,@>?&?,
&6?B'?&,()F,A?.+*D,@+,),6+.F6.+'K,B+(6.?U/
TAXONOMIC CONCLUSIONS
\%7,7?&%.@&,&>+M,@>),,@,@>?,A7+)K.F,B'7B%(&B7'A?K,+,-.%/ 0#&,
+N,E?@@?7&B>?'K,-,L+FB?,238884,'&,6+.F6>F.?@'B,)*K,B+(67'&?&,
@>7??,K'&@'*B@,B.)K?&G,M>'B>,&>+%.K,A?,7?B+D*'d?K,)@,D?*?7'B,7)*R,
@+,)B>'?C?,),A).)*B?K,B.)&&'N'B)@'+*,+N,@>?,57?)?/,=>?,5&')*,)*K,
I).?&')*,&6?B'?&,N+7(,),B.)K?,2+,-. % /0# &,&/&@7/4,@>)@,'&,&'&@?7,@+,
@>?,+@>?7,D?*?7),+N,57?)?/,=>?,5%&@7).')*,&6?B'?&,+N,+,-.% /0#& ,
&+,N)7,&?_%?*B?K, N+7(, ), K'&@'*B@,B.)K?/,+.3$0%-.%/#&G,N7+(,
&+%@>?7*,]*K')G, K'C?7D?&,*?U@G,N+..+M?K,AF,!"#$%&"'#&G,*+M,
B+(6+&?K,+N,*'*?,&6?B'?&G,)*K,&'&@?7,@+,), B.)K?,'*B.%K'*D,@>?,
I?K'@?77)*?)*,D?*?7),+N,57?)?,22$#&G(?0"$#&G(>$":#/:#9#8G(
W&0/0#&G(X390:%60:3$%84/,=>?, D?*%&, !"#$%&"'#&,B)*,A?,B'7J
B%(&B7'A?K, *+@,+*.F,D?*?@'B)..FG,A%@,).&+, (+76>+.+D'B)..F,)&,
M?,>)C?,&>+M*,>?7?/,566.'B)@'+*,+N,@>?,*)(?,)"*"$#&,N+7,@>?,
5%&@7).')*,B.)K?,2V'D/,94,)M)'@&,@>?,&?_%?*B'*D,+N,@>?,@F6?,+N,
@>'&,D?*?7'B,*)(?/,5@,@>'&,&@)D?G,@>?7?N+7?G,+,-. %/0# &,&/&@7/,)*K,
@>?,5%&@7).')*,B.)K?,)7?,*+@,K?N'*?K,(+76>+.+D'B)..FG,A%@,+*,+%7,
(+.?B%.)7,?C'K?*B?,)7?,B.?)7.F,K'&@'*B@/
]*,@>?,N+..+M'*DG,M?,7?&%77?B@,@>?,D?*%&,!"#$%&"'#&G(()R?,
@>?,*?B?&&)7F,N'C?,*?M,B+(A'*)@'+*&G,)*K,67?&?*@,),R?F,@+,@>?,
*'*?,&6?B'?&,+N,!"#$%&"'#&/,S?@)'.?K,K?&B7'6@'+*&,+N,)..,&6?B'?&,
)*K,%6J@+JK)@?,'*N+7()@'+*,+*,@>?'7,D?+D7)6>'B,K'&@7'A%@'+*,)7?,
)C)'.)A.?,?.&?M>?7?,2P7'A++*(),-,)./G,9##!H,Y)*D,-,['G,9###H,
E?@@?7&B>?'K,-,L+FB?G,3888H,E?@@?7&B>?'K,-,)./G,38894/
!"#$%&"'#&(PB>+@@,'*,PB>+@@,-,O*K.'B>?7G,I?.?@/,L+@/:,9</,9Z"3,
;,[?B@+@F6?,2K?&'D*)@?K,'*,=)U+*,9T:,19Z/,9#T<4:,!4(5#''"'#&(
PB>+@@,22$#&(5#''"'#&(Y). . 'B>,9Z"9G,*+*,P). '&A % 7 F,9<#T 4/
!"#$%&"'#&(=$370-38(2E++R/,N/4,Q/O/,L7+M*,'*,g)7K/,$>7+*/G,
&?7/,"G,"!234:,#"/,9#8",Ł(+,-.%/ 0#& (=$3 70-38(E++R/,N/,'*,V./,
L7'@/,]*K'),T:,199/,9Z#",;,PF*@F6?&:,S)7o??.'*DG,<188,N@/G,
N9"$J3(HE_DC(2^4H,`+7?,i+R7'G,<T88,N@/G,G"&&03(84/4,2^4/
!"#$%&"'#&(5"%905%/53/83(X/[/,Y)*D,-,E/,[','*,5B@),L+@/,
s%**)*/G,&%66./,99:,T9/,9###,Ł(+,-.%/0#&(5"%905%/53/83(
2X/[/, Y)*D,-,E/, ['4, E?@@/, -, i/$/,L+FB?,'*, 57+'K?)*),
3":,19/,3888,;,E+.+@F6?:,$>'*)G,s%**)*,67+C/G,L)+&>)*,
u')*(G,)0(X3/5(`(G4(a#:J3$'(SSFDP2,2^nQ4/
!"#$%&"'#&(.%$8;039600( I'_/,'*,V./,Q?K/, ]*K/, ":,9#T/,9Z11,Ł(
+,- .% /0 #& ( .% $8 ;0 39 600 ( 2I'_/4,P@??*'&,'*,L%../,`)7K/,L+@/,
L%'@?*d+7DG,&?7/,"G,9<:,!8"/,9#!Z,;,E+.+@F6?:,`)C)G,\?*)D)J
7)*G,X%$83;0396(84/4,2^4/,vV+7,N%..,&F*+*F(F,&??,P7'A++*(),
-,)./G,9##!/w
!"#$%&"'#&(73/%8#&(2S7F)*K/,?U,5'@/4,^%*@>,'*,O*%(/,i./,":,
3Z/,9Z!9,Ł(2$ #&(73/ %8# &(S7F)*K/,?U,5'@/,'*,E+7@/,^?M/,":,
"91/,9<Z#,Ł(>38&380"(73/%8#&(2S7F)*K/,?U,5'@/4,W)N/,'*,
V./,=?..%7/,":,T"/,9Z"<,Ł(!"#$%& "'# &,5#''"'#&(25'@/4,PB>+@@,
C)7/,73/%8#&(25'@/4,O*D./,'*,iN.)*d?*7/,]m,3"V,2E?N@,<"4:,
931/,9#38,Ł(+,-.%/0#&(73/%8#&(2S7F)*K/,?U,5'@/4,E?@@/,
-,i/$/,L+FB?,'*,57+'K?)*),3":,19/,3888,;,E+.+@F6?:,i.)*@,
+N,%*R*+M*,+7'D'*,'*@7+K%B?K,'*@+,B%.@'C)@'+*,)@,^?M,AF,
Y'..')(,I).B+.(,'*,9<<!,2LI4/,vV+7,N%..,&F*+*F(F,&??,
E?@@?7&B>?'K,)*K,L+FB?G,3888/w
,Sauromatum brevipilosum(2E?@@/,-,I/,P'd?(+7?4,$%&'()*+,
-,E?@@/G,comb. nov.,Ł(+,-.%/ 0#& (=$3 70-09%8# &(E?@@/,-,I/,
P'd?(+7?,'*,57+'K?)*),3":,13/,3888,;,E+.+@F6?:,]*K+*?J
&')G,P%()@?7)G,Y?&@,P%()@?7)G,*?)7,5?R&)>G,X3''3$8:.306(
X42a4DP_1+,2+7'D/,B+../,!0*3&%$3(84/44,N.+M?7?K,'*,B%.@/,'*,
[?'K?*,L+@/,g)7K/G,3#,Q+C,9###,2[G,&6'7'@,B+../4/
,Sauromatum diversifolium(2Y)../,?U, PB>+@@4,$%&'()*+, -,
E?@@/G,comb. nov.,Ł(+,-.%/0#&(6073$80;%90#&(Y)../, v'*,
Y)..'B>b&,Q%(?7/,['&@,*/,Z#""/,9#!#G,*+(/,*%K/w,?U,PB>+@@,
'*,57+'K?)?,9":,38/,9Z11,Ł(X3'3 $%8'" 908(60 73$80 ;% 90 "(PB>+@@,
'*,\?&@?77/,L+@/,Y+B>?*A./,<:,3T</,9Z1<,;,E+.+@F6?:,Q?6).G,
Y)..'B>b&,Q%(?7/, ['&@/, *+/,Z#""),'*, 9Z39,2^4/,vV+7,N%..,
&F*+*F(F,&??,P7'A++*(),-,)./G,9##!/w
,Sauromatum giganteum(2O*D./4,$%&'()*+,-,E?@@/,comb. nov/,
Ł(+,-.%/0#&(505"/'3#&(O*D./,'*,L+@/,`)>7A/,PF&@/,!:,TT/,
9ZZ ", ; , E+. +@ F6 ?: , $> '* )G , L? 'o' *D G, !J"'8:.J%O(84/4,2[O4/,
vV+7,N%..,&F*+*F(F,&??,P7'A++*(),-,)./G,9##!/w
,Sauromatum hirsutum(2P/s/, E%4, $ %&' () *+, - , E?@ @/G ,comb.
nov/,Ł(2$08"3&"(.0$8#'#&(P/s/,E%,'*, S)*&R,L+@/,57R/,
3"2!4:, !1!/, 9#TZ, Ł(+,-.%/0#&(.0$8#'#&(2P/s/, E%4, `/,
I%7)@),-,I)F+,'*,^?M,L%../,!T294:,93#/,9##9,;,E+.+@F6?:,
=>)'.)*KG,I","-(FPFP(2$4/
,Sauromatum tentaculatum(2E?@@/4,$%&'()*+,-,E?@@/G,comb.
nov.,Ł,+,-.% /0# &('3/' ":#9 "'#& (E?@@/,'*,57+'K?)*),3!:,!#/,
3889,;,E+.+@F6?:,=>)'.)*K,PY,"<G,^)*B>)*)A%7'G,P)*DR.)J
A%7',S'&@7/G,[)',Y+,P%AK'&@7/G,=++*D,s)',Y'.K.'N?,W?&?7C?G,
L)*,P)*?>,i)M*DG,Y?&@,&'K?,+N,i)*?>,.'(?&@+*?,(+%*@)'*G,
"88, (,X3''3$8:.306(X42a4DQH1+,2+7'D/,B+../,b4K4(B"<O399(
PF1EQ_ 4G,N .+M?7?K,'*,B%.@/,'*,@>?,[?'K?*,L+@/,g)7K/,#,567,
9##<,2L^ VG,&6'7 '@,B +../4/
!!T
TA XO N,1#,234,0,567'.,3898:,!"#;!!<$%&'()*+,-,)./,0,Phylogenetics of Typhonium and Sauromatum
Bogner, J. & Boyce, P.C. 388Z/,W&0/0#&(]"353$0(257 )B?)?4G,),*? M,&6 ?J
B'?&,N7+(,*+7 @>M?&@?7 *,]7)*/,L09963/%O0","Z:,9!#;91"/
Bogner, J. & Petersen, G. 388</,=>?,B>7+(&+(?,*%(A?7&,+N,@>?,)7+'K,
D?*?7)/,2$%063"/","8:,Z3;#8/
Key to Sauromatum species
9, =M+,@F6?&,+N,&@)('*+K?&G,%66?7,B.?)7.F,+N,K'NN?7?*@,&>)6?,
+7,7?K%B?K,B+(6)7?K,@+,@>?,.+M?7,+*?&,,/,/,/,/,/,/,/,/,/,/,/,/,/, 3
9, \*.F,+*?,@F6?,+N,&@)('*+K?&G,%66?7,&@?7'.?,6)7@,.+*D'@%K'J
*)..F,D7++C?K,)*K,7+%D>?*?K,M'@>,67+o?B@'+*&,+7,67+B?&&?&G,
+7,*)R?K/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/, 1
3, P6)@>?,@%A?,N%&?KH, ";1,A%.A'.&,'*, @>?,.+M?7, 6)7@, +N, @>?,
6?@'+.?,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/ S. gaoligongense
3, P6)@>?,@%A?,B+*C+.%@?/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,"
", [+M?7,&@)('*+K?&,B.?)7.F,B.)C)@?/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,!
", [+M?7, &@)('*+K?&,B.)C)@?c&6)@>%.)@?G, B.)C)@?,6)7@,)*C'.,
&>)6?K, +7, K+7&+JC?*@7)..F,N.)@@?*?KG,K)7R, 6%76.?H,%66?7,
&@)('*+K?&,*)77+M.F,&6'*K.?,&>)6?KG,M>'@'&>,,/,/,/,/,/,/,/,/,/, ,
/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/ S. tentaculatum
!, [?)C?&,&'(6.?G,+C)@?G,B+7K)@?,@+,>)&@)@?H,%66?7,&@)('*+K?&,
&()..?7,@>)*,@>?,.+M?7,+*?&G,&%A%.)@?,+7,+BB)&'+*)..F,)AJ
&?*@,,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,S. giganteum
", [?)C?&,6?K)@'&?B@H,%66?7,&@)('*+K?&,N'.'N+7(G,D7)K%)..F,
&>+7@?7,@+M) 7K&,@>?,().?,d+*?,,/,/,/,/,/,/,/,/,/,/, S. horsfieldii
1, P6)@>?,@%A?,N7??H,.?)N,A.)K?,&'(6.?,)*K,+C)@?J.)*B?+.)@?G,
B%*?)@?, @+,>)&@)@?,";1J.+A?K,+7, 1;#JN+.'+.)@?,6?K)@'&?B@H,
%66?7,&@?7'.?,6)7@,+N,@>?,&6)K'U, *)R?K, +7, M'@>, )6'B%.)@?,
67+o?B@'+*&/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/ S. diversifolium
1, P6)@>?,@%A?,N%&?KG,)@,.?)&@,A)&).,6)7@ /,/,/,/,/,/,/,/,/,/,/,/,/,/,/,T
T, i?@'+.?,)*K,.?)N,A.)K?,M'@>,>)'7&H,%66?7,&@?7'.?,6)7@,+N,@>?,
&6)K'U,)U'&,B+(6.?@?.F,*)R?K/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,<
T, i?@'+.?,)*K,.?)N,A.)K?,M'@>+%@,>)'7&G,%66?7,&@?7'.?,6)7@,+N,
@>?,&6)K'U,)U'&,7+%D>G, &>)DDF,+7,M'@>,('*%@?, 67+B?&&?&,
2>'D>.F,7?K%B?K,&@)('*+K?&4,,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,Z
<, E)'7&,K'&@'*B@,)*K,.+*DH,).&+,+%@&'K?,&%7N)B?,+N,&6)@>?,B+CJ
?7?K,M'@>,>)' 7&,,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/, S. hirsutum
<, E)'7&, &>+7@, 2B), 8/1, ((4H, +%@&'K?, &%7N)B?, +N, &6)@>?,
&(++@>/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,S. brevipilosum
Z, [?)N.?@&,+A.+*DJ.)*B?+.)@?G,)6?U,)B%('*)@?H,'*N.+7?&B?*B?,
%6,@+,!8,B(,.+*DH,&6)@>?,@%A?,K)7R,6%76.?,'*&'K?G,&6)@>?,
A.)K?,'*&'K?,()B%.)@?,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/S. venosum
Z, [?)N.?@&, .'*?)7J.)*B?+.)@?G, )6?U, .+*D, )B%('*)@?H,.+M?7,
B.)C)@?,&@)('*+K?&,M>'@?G,%66?7,('*%@?,67+B?&&?&,6%76.?H,
'*N .+7?&B?*B?,()U/,</1,B(,.+*DH,&6)@>?,@%A?,'*&'K?,D7??*J
'&>,@+,M>'@?G,&6)@>?,A.)K?,'*&'K?,*+@,()B%.)@?G,K%..,6%76.?,
A)&)..F,)*K,6'*R,)A+C? /,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,/,S. brevipes
ACKN OWLEDG EMENTS
V+7, 6.) *@, () @?7').G,M?,@ >)*R,`+ &?N,L+D* ?7G,[',E? *DG ,$.)7 ?,E?7J
&B+C'@B>G, Y+.N7)(, [+A'*,)*K, 5..)*, g)..+M)F/,Y?,).&+,@>)*R, `+&?N,
L+D*?7,)*K,@M+,)*+*F(+%&,7?C'?M?7&,N+7,@>?'7,B+((?*@&/,=>'&,&@%KF,
M)&,&%66+7 @?K,AF,SVg,D7)*@,WO,T8"c<J9/
LITERATURE CITED
Boyce, P.C. 9##"/,+.3(53/#8(57%(/,^?M:,=>?,L?*@>)(JI+U+*,=7%&@G,
W+F).,L+@)*'B,g)7K?*&G,^?M/
Boyce, P.C. 388T/,2$#&:,5,K?B)K?,+N,B>)*D?/,2$%063"/",3#:,9"3;9"</
Boyce, P.C. 388Z/, 5, (+*+D7)6>, +N, @>?, D?*%&, ?0"$#&c&6?B')., 6)7@/,
N#$'08d8(?%'4(B"54(31:,9;99#/
Brown, N.E. 9#8"/,Q?M,+7,*+@?M+7@>F,6.)*@&,;,!"#$%&"'#&(=$370-384,
G"$64(N.$%/G,&?7/,"G,"!:,#";#!/
Dao, Z.L., Chen, S.-T., Ji, Y.-H. & Li, H. 388</,+,-.%/0#&(="%8."1
/3/83(X/[/,S)+,-,E/,[':,5,*?M,&6?B'?&,+N,57)B?)?,N7+(,M?&@?7*,
s%**)*G,$>'*)/,2:'"(I.,'%'"<4(!0//,!1:,3"!;3"Z/
Evans, O. 9#T9/,57 )B ?) ?/,N%/'$4(U3O(!%#'.(L"938(U"'94(X3$=4@(K94(!3$4(
39c33:,T;9"/
French, J.C., Chung, M.G. & Hur, Y.K. 9##1/,$ >.+7+6.)&@,SQ5,6>FJ
.+D?*F,+N,@>?,57'N .+7)?/, i6/, 311;3<1,'*:,W%K)..G, i/`/G,$7'AAG, i/G,
$%@.?7G,S/V/,-,E%(6>7'?&G,$/`/,2?K&/4G(B%/%: %' ,936% /8e(!,8'3 &" '1
0:8("/6(37%9#'0%//,^?M:,W+F).,L+@)*'B,g)7K?*&/
Hay, A. 9##3/,5,*?M,5%&@7).')*,D?*%&,+N,57)B?)?G,M'@>, *+@?&,+*,D?J
*?7'B,.'('@&,)*K,A'+D?+D7)6>F,+N,@>?,57?)?/,?%'4(b4()0//4(!%:/,98#:,
!3<;!"!/
Hay, A. 9##"/,=>?,D?* %&,+,-.%/0#&(257)B?)?;57?)?4,'*,5%&@7).)&')/,
?9#&3","<:,"!1;"<T/
Hay, A. 9##</,=M+,*?M,&6?B'?&,)*K, ),*?M, B+(A'*)@'+*,'*, 5%&@7).')*,
+,-.%/0#&(257)B?)?,@7'A?,57?)?4/,W60/=#$5.(b4(?%'/,1!:,"3#;""T/
Hay, A. & Taylor, S.M. 9## T/, 5, * ?M, & 6? B'? &, +N , +,-.% /0# &, PB>+@@,
257)B?)? ;57?)?4,N7+(,@>?,Q+7@>?7 *,=?77'@+7FG,M'@>,*+@?&,+*,@>?,
B+*&?7C)@'+*,&@)@%&,+N,@M+,57?)?, ?*K?('B,@+, @>?,='M',]&.)*K&/,
+39 %-3 ",T:,1T";1TZ/
Hetterscheid, W.L.A. & Boyce, P.C. 3888/,5,7?B.)&&'N'B)@'+*,+N,!"#1
$%&"'#&(PB>+@@,)*K,*?M,&6?B'?&,+N,+,-.%/0#&(PB>+@@,257)B?)?4/,
2$%063"/",3":,!Z;11/
Hetterscheid, W.L.A. & Galloway, A. 388T/,Q?M,+,-. %/0 #& ,257)J
B?)?4,&6?B'?&,N7+(,=>)'.)*K/,2$%063"/",3#:,Z8;Z1/
Hetterscheid, W.L.A. & Nguyen, V.D. 3889/,=>7??,*?M,&6?B'?&, +N,
+,-.%/0#&(257)B?)?4,N7+(,m'?@*)(/,2$%063"/",3!:,3!;3#/
Hetterscheid, W.L.A., Sookchaloem, D. & Murata, J. 3889/,+,-. % 1
/0#&(257)B?)?4,+N, =>)'.)*K:,Q?M, &6?B'?&,)*K, ),7?C'&?K,R?F/,2$1
%063"/",3!:,"8;11/
Hooker, J.D. 9Z#"/,K9%$"(%;(?$0'08.(f/60"G(C+./,TG(g$:.063"3('%(N,-3$"1
:3"3/,[+*K+*:,W??C?/
Hu, S.Y. 9#T Z/, 5 7) B? )?/ , P@ %K '?& , '*, @ >? , V.+ 7), +N , => )' .)* K, !9/, >"/8((J(
?%'4(2$J4,3":,!99;!1</
Huelsenbeck, J.P. & Ronquist, F. 3889/,I7L)F?&:,L)F?&')*,'*N?7?*B?,
+N,6>F.+D?*?@'B,@7??&/,?0%0/;%$&"'0:8,9<:,<1!;<11/
Johnson, L.A. & Soltis, D.E. 9##!/,B"'Y(SQ5,&?_%?*B?&,)*K,6>F.+D?J
*?@'B,7?B+*&@7%B@'+*,'*,P)U'N7)D)B?)?,&/&@7/,!,8'4(?%'/,9#:,9!";91T/
Kunth, C.S. 9Z!9/,W/#&3$"'0%(-9"/'"$#&G,C+./,"/,P@%@@D)7 @:,$+@@)?/
Lobin, W., Neumann, M., Bogner, J. & Boyce, P.C. 388</,5, *?M,
2$#&( &6?B'?&,257?)?G, 57)B?)?4, N7+(, QO,=%7R?F, )*K, g?+7D')/,
L09963/%O0","<:,!!1;!!#/
Maddison, W.P. & Maddison, D.R. 9##3/,B":N9"63e(2/"9,808(%;(-.,1
9%53/,("/6( :."$":'3$(37%9#'0%/G(C?7&'+*, "/8/, P%*K?7.)*KG, I)&J
&)B>%&?@@&:,P'*)%?7/
Mayo, S.J., Bogner, J. & Boyce, P.C. 9##</,+.3(53/3$"(%;( 2$":3"3/,
^?M:,=>?,=7%&@??&G,W+F).,L+@)*'B,g)7K?*&/
Miquel, F.A.W. 9Z11/,K9%$"( 7"/( U363$9"/68:.(f/603G,C+./,"/,5(&@?7J
K)(:,$/g/,C)*,K?7,i+&@/
Miquel, F.A.W. 9ZT!/,I)*@'&&),57).')B?)7%(G,O7'B)B?)7%(G,$%6%.'NJ
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Ronquist, F. & Huelsenbeck, J.P. 388"/,I7L)F?&,":,L)F?&')*,6>F.+D?J
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*%(A?7,'*,57)B?)?/,U%7%/,93:,3ZT;3Z#/
Appendix.
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... ex Schott) Cusimano & Hett. (Cusimano et al. 2010), and one recently published species S. meghalayense D.K. Roy, A.D. Talukdar, B.K. Sinha & M. Dutta Choudhury (Roy et al. 2014). ...
... These morphological characters showed that collected plant is a species Sauromatum Schott. But, the morphological characters of its tuber, leaf blade, petiole, spathe tube, spadix, structure of staminodes, male and female zone, stipitate appendix, and bifid pointed stigma made it is distinct from the so far known species of the genus (Hooker 1894;Cusimano et al. 2010). ...
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Sauromatum nangkarense A. Nangkar & H. Tag, a new species of Sauromatum Schott, (Araceae) from Kimin, Papum Pare District of Arunachal Pradesh, India, is described and illustrated with photographs of living specimens. The new species resembles Sauromatum brevipes and S. hirusutum with their staminodes short and curved downward and the spathe-tube is convulated. The new species differs from these in having subcylindric tubers, pinnate leaves, with male zone blonde, cream, sterile flowers clavate, pink, stipitate and curved downward covering female zone partially. Detailed description and a range of photographs have been provided.
... ex Schott) Cusimano & Hett. (Cusimano et al. 2010), and one recently published species S. meghalayense D.K. Roy, A.D. Talukdar, B.K. Sinha & M. Dutta Choudhury (Roy et al. 2014). ...
... These morphological characters showed that collected plant is a species Sauromatum Schott. But, the morphological characters of its tuber, leaf blade, petiole, spathe tube, spadix, structure of staminodes, male and female zone, stipitate appendix, and bifid pointed stigma made it is distinct from the so far known species of the genus (Hooker 1894;Cusimano et al. 2010). ...
... IntroductionCusimano et al., 2010). During field exploration in South Garo Hills district of Meghalaya, Northeast India, one of us (DKR) made some collections (only one mature) of an aroid plant. ...
... As the mature individual was collected only in fruiting stage, so could not observe the flower characters. Even though, the morphological characters of its tuber, leaf blade, petiole, spathe tube, fruiting zone of spadix, berries, etc. so distinct from its all species within the genus Sauromatum known to science (Hooker, 1894; Cusimano et al., 2010). Therefore, the plant is described here as a new species. ...
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Sauromatum meghalayense D.K. Roy, A.D. Talukdar, B.K. Sinha & M. Dutta Choudhury (Araceae), a new species is described and illustrated from the Indian state of Meghalaya based on live plant.
... In the family Araceae, Typhonium is a medium-sized genus with an estimated number of species ranging from 37 (Mayo et al. 1997), 69 including Sauromatum (Govaerts et al. 2002), to about 100 (Hetterscheid and Sookchaloem 2012). They are distributed from East Himalaya throughout tropical and subtropical Asia as far east as New Guinea and Australia (Mayo et al. 1997, Govaerts et al. 2002, Hetterscheid 2013, eMonocot Team 2021, though molecular studies indicate that nearly all the hitherto recognised New Guinea and Australian Typhonium species are in a distinct clade for which the generic name Lazarum A. Hay is available (Cusimano et al. 2010). Recently, since 2000 to now, 36 new species have been described from Indochina including Thailand (Hetterscheid and Boyce 2000, Hetterscheid and Nguyen 2001, Nguyen 2008, Nguyen and Croat 2010, Galloway 2012, 2015, Hetterscheid and Sookchaloem 2012, Hetterscheid 2013, Luu et al. 2017, Sookchaloem and Maneeanakekul 2017, Van et al. 2017, Nguyen et al. 2021). ...
Article
Typhonium kbangense is described as a new species from Vietnam. It is one of eleven species recently found in central and southern Vietnam. It belongs to the pedate leaf blade group of species and is closest to T. bachmaense and T. dongnaiense, but is differentiated from the two latter by having much longer sterile flowers, not dense as in T. dongnaiense, and without being clavate apically as in T. bachmaense. A table of morphological characters for the three species, the ecological characteristics, specific habitat and conservation status of the new species are estimated and provided.
... There can, however, be significant variation in floral traits between Typhonium species which may reflect selection by different pollinator assemblages (Sayers et al., 2020). In this study, we focus on two closely related Typhonium species, T. angustilobum and T wilbertii, which form a single monophyletic group and have similar geographical distributions in tropical Far North Queensland (FNQ) (Cusimano et al., 2010;Hay, 2011), and were observed to be pollinated by different insect orders (Coleoptera and Diptera). We set out to (1) identify their pollinators; (2) compare the mechanism of thermogenesis and the genes involved; and (3) examine the floral traits, both sensory (i.e. ...
Article
Background Flowers which imitate insect oviposition sites likely represent the most widespread form of floral mimicry, exhibit the most diverse floral signals and are visited by two of the most speciose and advanced taxa of insect – beetles and flies. Detailed comparative studies on brood-site mimics pollinated exclusively by each of these insect orders are lacking, limiting our understanding of floral trait adaptation to different pollinator groups in these deceptive systems. Methods Two closely related and apparent brood-site mimics Typhonium angustilobum and T. wilbertii (Araceae) observed to trap these distinct beetle and fly pollinator groups were used to investigate potential divergence in floral signals and traits most likely under pollinator-mediated selection. Trapped pollinators were identified and their relative abundances enumerated, and thermogenic, visual, and chemical signals and morphological traits were examined using thermocouples and qRT-PCR, reflectance, gas chromatography-mass spectrometry, floral measurements and microscopy. Key Results T. angustilobum and T. wilbertii were functionally specialised to trap saprophagous Coleoptera and Diptera, respectively. Both species shared similar colour and thermogenic traits and contained two highly homologous AOX genes (AOX1a and AOX1b) most expressed in the thermogenic tissue and stage (unlike pUCP). Scent during the pistillate stage differed markedly - T. angustilobum emitted a complex blend of sesquiterpenes, and T. wilbertii, a dung mimic, high relative amounts of skatole, p-cresol, and irregular terpenes. Species differed significantly in floral morphology related to trapping mechanisms. Conclusions Functional specialisation and pollinator divergence were not associated with differences in anthesis rhythm and floral thermogenic or visual signals between species, but with significant differences in floral scent and morphological features, suggesting these floral traits are critical for the attraction and filtering of beetle or fly pollinators in these two brood-site mimics.
... Anggota puak Areae terdiri dari 14 marga dengan Typhonium merupakan marga terbesar dengan jumlah mencapai 68 jenis (TPL 2013). Marga ini tersebar luas di kawasan Asia Selatan, Asia Tenggara, Asia Timur, Malesia, dan Australia (Hay 1993;Cusimano et al. 2010;NGPS 2018). Selain itu, marga ini juga dapat ditemukan di kawasan Afrika dan Amerika melalui proses introduksi (Nicolson dan Sivadasan 1981). ...
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The study on the dynamics of Typhonium was carried out through the Bogor Botanic Gardens collections catalog and direct observations in the garden. The results showed that there were 8 species of Typhonium which had once been listed as garden's collection and planted in Bogor Botanic Gardens. There was a change in the species composition of Typhonium starting from 1844 to 1985. In 2018, two non-collection species of Typhonium were found growing abundantly in the Bogor Botanic Gardens. The updated data of Typhonium record in Bogor Botanic Gardens catalog is also presented.
... Secondary centres of species abundance and diversity associated with geophytism occur in the monsoonal tropics of Africa, Madagascar and IndoMalaya (e.g. Bogner 1972;Ittenbach 2003;Cusimano et al. 2010;Bogner and Nusbaumer 2012;Boyce et al. 2012;Hetterscheid and Claudel 2014), and in warm temperate Asia (e.g. Li et al. 2010). ...
Article
Phylogenetic relationships within Araceae tribe Schismatoglottideae are elucidated based on nuclear ITS and plastid matK. Thirty genera are recognised, composed of 13 pre-existing genera (Apoballis, Aridarum, Bakoa, Bucephalandra, Fenestratarum, Galantharum, Ooia, Phymatarum, Pichinia, Piptospatha, Schismatoglottis, Schottariella and Schottarum), five resurrected genera (Colobogynium, Gamogyne, Heteroaridarum, Hottarum and Rhynchopyle) and 11 new genera: Bakoaella, Bidayuha, Burttianthus, Gosong, Hera, Kiewia, Nabalu, Pursegloveia, Naiadia, Tawaia and Toga. Significant recircumscription is proposed for the pre-existing genera Aridarum, Bakoa and Piptospatha. Schismatoglottis remains imperfectly delineated despite resolution of a well-supported corpus defined by hapaxanthic stems and containing the nomenclature type. An updated taxonomic treatment is presented and all necessary nomenclatural changes are made including a new generic name, Vesta to replace Hestia, which has been discovered to be preoccupied by Hestia, a fossil lycopsid. All genera, including representatives of each clade pertinent to Schismatoglottis sens. str. are illustrated from living plants.
... Steenis was accepted as a synonym of Sauromatum horsfieldii Miq. (Cusimano et al., 2010). Therefore, prior to this paper, 15 species of Typhonium have been known for the country. ...
Article
Typhonium thatsonense Luu & H. T. Van (Araceae) is described and illustrated as a new species from the That Son Mountains, Mekong Delta, Vietnam. It most closely resembles T. supraneeae A. Galloway, Petra Schmidt & Sinhab. in its trifoliolate leaves and general appearance of the inflorescence but differs in many leaf and floral characters.
... Banerji's (1947) account of T. trilobatum is the only developmental study of another Typhonium species of which we are aware. It should be noted here that, in the view ofCusimano et al. (2010), who studied genetic evidence of the phylogeny of the genus, Australian Typhonium spp. and those from Asia belong to different clades and the Australian species may warrant generic recognition. ...
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Typhonium sp. Kununurra is an undescribed, perennial, cormous herb belonging to the Family Araceae. It produces leaves and flowers only during the northern Australian wet season. Its known distribution is restricted to grey vertisols ('black soils') on the ancestral flood plain of the lower Ord River in the east Kimberley, Western Australia and adjacent areas of the Northern Territory, northern Australia. It is Declared Rare Flora under Western Australian legislation (but its conservation status has not yet been assessed in the NT). Although several subpopulations are known, few are on land with conservation-secure tenure and no previous studies have distinguished immature from mature plants. This study had two objectives; to locate additional subpopulations on conservation-secure land and to identify development stages in the life history of the species so that immature plants can be identified and excluded from future population-size estimates. Because IUCN Red List guidelines for the estimation of population sizes require that only reproductively mature individuals are included (IUCN 2001, 2014), it is critically important that future subpopulation surveys for this species distinguish immature from mature plants and take into account possible hydrological/climatic conditions that might affect survey results. Following exceptionally dry, hot conditions, many known subpopulations were still dormant at the time of the study (early March 2013) and, perhaps because of that, no new subpopulations were located. However, there had been sufficient, localised rainfall for some, isolated emergence. Three developmental stages are described, newly germinated seedlings, immature plants and reproductively mature plants, all of which were recognisable in the field. The overwhelming proportion of all subpopulations observed in this study consisted of immature plants and the proportion of immature to mature plants suggests a high mortality rate during the 3+ years plants can take to mature; however, an alternative explanation involving sustained dormancy of mature corms in 'dry' years is canvassed.
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Three new species of Typhonium (T. bachmaense, T. lineare and T. penicillatum) are described from Vietnam.
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The Araceae genus Sauromatum is put into synonymy of the genus Typhonium. Several new nomenclatural combinations are presented.
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Typhonium baoshanense Z. L. Dao & H. Li (Araceae: Areae), a new species from western Yunnan, China, is described and illustrated. This species differs mainly from other species of Typhonium by the presence of one to two rings of sterile male flowers above the fertile male flowers, and from T. blumei Nicolson & Sivadasan by a conic female part of the spadix, the appendix not being truncate at its base, and by the angustate limb of the spathe. The chromosome number of Typhonium blumei, which occurs in same area, is 2n=52, while the chromosome number of the new species is 2n=10. The chromosome number of the new species is the lowest in the family Araceae. Key words Typhonium, Typhonium baoshanense Z. L. Dao & H. Li, Araceae, new species, Yunnan, China. Typhonium Schott (Araceae: Areae) is a genus containing about 60 (44) species (Schott, 1832; Sriboonma et al., 1994; Hay, 1997; Mayo et al., 1997; Hetterscheid & Boyce, 2000; Hetterscheid & Nguyen, 2001; Hetterscheid et al., 2001; Wang et al., 2002), excluding Sauromatum Schott species which were moved into Typhonium by Hetterscheid and Boyce (2000). The genus Typhonium is generally distributed in south, southeast and east Asia, and extends throughout the Pacific area to Australasia. Some species are weeds and have become naturalized in various parts of the world outside Asia. There are 16 species reported in northern and southern provinces of China except Jilin and Xinjiang (Li, 1979; Li & Liu, 1983; Li & Long, 1998; Wang et al., 2002). In the summer of 2003, during an expedition in the Baihualing, Baoshan, Yunnan Province, we encountered an unusual population of Typhonium. After having carefully examined the relevant herbarium specimens, reviewed the relevant literatures, and investigated the chromosome number, we provide here the morphological and karyological evidence and conclude that these plants are sufficiently distinct to be recognized as a new species.
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Pertelaan polong dan pohon tipe Ormosia incerta Koord. (yang direduksi ke O. penangensis Ridl.) dari Jawa diberikan. Trifidacanthus Merr. direduksi menjadi Desmodium dan suatu kombinasi baru diusulkan. Platyspermation Guillaumin, semula dimasukkan dalam Myrtaeeae, dipindah ke Saxifragaceae. Juncus bufonius L. direkam sebagai pendatang di G. Kinabalu. Dua Fimbristylis direkam dari Australia Utara dan Oreobolus kiikenthalii Steen. direkam dari G. Mulu (Sarawak).
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Comparative DNA sequencing of matK, a maturase coding gene located within the intron of the chloroplast gene trnK, was evaluated for phylogenetic utility using genera of Saxifragaceae s. str. The entire matK gene was sequenced for two members of the family, Sullivantia sullivantii and Saxifraga integrifolia. Comparison of base substitution rates between these two species indicated that matK evolves approximately three-fold faster than rbcL. Comparative sequencing of 754 base pairs within matK was subsequently conducted using 25 genera in Saxifragaceae s. str. and two outgroup taxa. Summed over the 31 taxa sequenced for this 754 base pair region, 40% of the base positions were variable and 15.6% were potentially informative. Five insertion/deletion events of three or six base pairs were also detected. Skewness and randomization tests both suggest that significant non-random structure is present in the matK data set. Parsimony analyses provided 72 most parsimonious trees of 223 steps (excluding autapomorphies) with a consistency index of 0.565. Several well-supported groups of genera are highly concordant with relationships suggested by two other chloroplast DNA data sets: chloroplast DNA restriction sites and rbcL sequences.
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Theriophonum Bl., most closely related to Typhonium, is endemic to South and Central India and Sri Lanka. Five quite distinct species (four have been treated as genera) are accepted: T. dalzellii, T. fischeri, T. infaustum, T. minutum (the most variable species), and T. sivaganganum.
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A new species Typhonium watanabei J.Murata, Sookchaloem & W.Hett. is described from Thailand.
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Arisaema hirsutum S. Y. Hu, described from Thailand and based on a fruiting specimen, is transferred to Typhonium. A revised description supplemented by information from a flowering specimen is provided.