!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
TAXON
2,0'+30)4"/.$0,5'6$",-"&.'"!"#$"%&'"($)*"#+,"'+-"%/'+*'%"+,(/0'1"
78)(/)630)+')'9"*6-'44':"-4,("+.0,4,/0)%&")6:"63+0')4"6,6;+,:*65"
%'<3'6+'%")6:"*&%"&)1,6,(*+"*(/0*+)&*,6%
1
5
2
2
3
2
4"
9,2"'&6"+):'-2&2/2()"'$);"#$('<)="6/+25),>)9&,+,?5<)@'&A(#-&25),>)9(+?#"$(<)B"C,A-C")DE<)FFGGG)9(+?#"$(<)H(#I&")
FGGGG)J"?#(I<)!#,"2&"
Abstract
!"%."'/+").5#"%&$"+&)
)
1
>+,#" )
.-I412#'* ).-IJ12#'>K )2#';12#'H
)
Keywords !"%."'/+"
Supplementary Material
4-5'(/%"
INTRODUCTION
4'$#,-"6()
#
#
#
*(+&,-.(#%")
23/-)
7',-%"
8&%,'&/%
!"%."'/+"
4#(2&")
#
#
#
!"#$"%&'()
"
!"%."'/+")
#
0$#"&"'1
!"%."'/+"
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
#
!"!
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
TAXON
#
#
#
'
'
'
!"%."'/+"
#
'
#
'
#
#
!"%."'/+").5#"%&$"+&-
'
#
'
!"%."'/+")
#
#
!"%."'/+"
#
!"%."'/+"
#
!"%."'/+"
!"%."'/+"
#
#
#
#
#
!"$
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
TAXON
1
$"+&-
=+,#")0/#,."(")
K($1!3(6C+&-2)
.-I412#'* ).-IJ12#'>K )2#';12#'H
)
MATERIALS AND METHODS
Taxon sampling. —
#
#
-&6,+,#)
)%,'2('(?#&'"
)+&%('-&-
#
Fig. .
!"%."'/+").5#"%&$"+&-)
)
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
1
%"2C"()
#
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
TAXON
#
!"%."'/+"
#
#
DNA extraction, amplification, and sequencing. —
#
#
#
GUG
CAU
UCC
GCU
#
#
#
#
#
#
Phylogenetic analyses. —
#
#
#
#
Topological incongruence and alternative hypothesis
testing. —
#
#
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
#
#
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
TAXON
#
#
#
#
#
RESULTS
DNA regions and alignments. —
#
Chloroplast genealogies. —
#
#
#
.-I41
2#'*
1
$"+&-
!"%."'/+").5#"%&$"+&-
Table .
.-I412#'*
.-IJ12#'>K
2#';12#'H
a
$"
b
$!
c
8
=
=
=
a
b
c
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
&$"+&$&>+,#"
TAXON
1
#
1
#
#
#
#
#
#
#
$"+&-
#
1
#
#
#
#
#
Phylogenetic analyses of combined data. —
#
#
#
Nuclear genealogy. —
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
TAXON
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
.-I412#'* .-IJ1
#
Fig. .
2#'>K 2#';12#'H
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
TAXON
Fig. .
#
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
TAXON
DISCUSSION
#
#
#
#
=+,#") 0/#,."("
K($1!3(6C+&-2
"'$)9&I+&,?#".35),>)!"%."'/+"6("(
#
L,#+$)!3(6C+&-2)
"
-&6,+,#
."'/+").5#"%&$"+&-
1
-&6,+,#)
)
1
#
!"%1
#
#
#
$"+&-
#
1
#
Species delimitation within C. pyramidalis. — !"%."'/+")
.5#"%&$"+&-
#
!"%."'/+").5#"%&$"+&-
Table .
>?>>@
>?>AB
#
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
Fig. ."
2#'* .-IJ12#'>K 2#';12#'H
."'/+"
TAXON
.-I41
!"%1
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
TAXON
M&.(#")"%%,$52(-)
#
#
)
!"%1
!"%1
."'/+").5#"%&$"+&."'/+")"/-2#,"$#&"2&6"
#
!"#$"%&'()%"#&2&%"
!"#$"%&'()%"#&2&%"
"'23/-)2('/&>,+&/-
1
2#,"$#&"2&6"
0$#"&1
#
#
%(+&-(++('-&-
#
Campanula pyramidalis s.str. (P.). —
0$#"&"'23/-)-(#.5++&>,+&/-
&$"+&-
#
1
!"%."'/+").5#"%&$"+&-
Table .
#
#
#
#
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
#
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
TAXON
!"%."'/+").5#"%&$"+&-
#
#
#
!"%."'/+").5#"%&$"+&-
Campanula austroadriatica sp. nov. (P.). —
#
#
#
#
#
Fig. ."
!"%."'/+").5#"%&$"+&B, )
C,
A,
#
#&"2&6")
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
1
"
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
TAXON
#
Taxa not accepted within the C. pyramidalis complex. —
#
#
Campanula pyramidalis s.l. (P.). —
#
#
1
$"+&-
-"+"
1
%"2C"(
)
6&'"
%&$"+&-
"+I"
-,'&&)
6,%."62"
6"+51
1
25%,'-&&
1
#
1
>+,#")
TAXONOMIC TREATMENT
!"#$"%&'"( $)*"#+,"'+-
"C'+&,&$/' :'%*56)&':".'4'
#
#
Campanula versicolor. —
!"%."'/+").5#"%&$"+&-
#
#
.5#"%&$"+&-
!
#
1
2"#/%)
#
#
#
)
K(+I,/#'()!,$()
!"%."'/+").5#"%&$"+&!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
Fig. . !"%."'/+")"/-2#,"$#&"2&6")
F,
G,
H,
I,"
TAXON
A,
B,
J,
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
C, D,
E,
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
TAXON
!"#$"%&'"("&-.*/",*+".+0"
%/?"6,D?
#
)
"
#&'(%
C')D'%"
ACKNOWLEDGEMENTS
;
+'6+'
8)0$1
8,4,00)
#&)('6%
#
#&$0'
ED)4$
"
8)/%30'
#
#
#
"#'':%
LITERATURE CITED
#
!"%."'/+"
#
#
!"%1
#
."'/+")"#?"("
#
:+)N/"2(#'"#&,
#
#
#
#
#
#
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
TAXON
#
#
!"%."'/+")#,2/'$&>,+&"
=+,#"),#&('2"+&#
"
K,',?#".3&()$(-)!"%."'/+O(!"%."'/+")+/-&2"'&6"
"
!"%."'/+")$#"I&>,+&"
3,2-.,2
!"%."'/+"
+,6/+"#(-
#
!"%1
."'/+"
"
N/&'Q/(1
1
2&6&."2&,'P)9,,C),>)4I-2#"62-
#
;(,+,?5
!"%."'/+"
#
;(,+,?5
=+,#")0/#,."("
#
#
#
0A,+/2&,'
*(+&,-.(#%")
!"%."'/+"
:+)N/"2(#'"#&,
#I68)
!"%."'/+"
"
#
4-5'(/%")6,%,-&>,#%(
!"%."'/+")(+"2&'(-
!"%."'/+"-P)B3(&#)6/+2&A"2&,')"'$)6+"--&>&6"2&,'
#
!%"
!"%."'/+"
L('2&"
!"%."'/+"
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
TAXON
!"%."'/+"
K($1!3(6C+&-2P)4)6#&2&6"+)&'A('2,#5),>)
A"-6/+"#).+"'2-),>)23()6&#6/%1K($&2(##"'("')6,/'2#&(-
#
K($1!3(6C+&-2
1
R&'"#&$#
#
#
#
#
#
!"%."'/+"
#
!"%."'/+"
!"%."'/+")#,2/'$&>,+&"
!"%."'/+"
K,/'2"&')>+,#"),>);#((6(
R&'1
#
"#,%5-)I,?$"',A&
"
#
!"#$"%&'()%"#&2&%"
!"#$"%&'()%"#&2&%")
!"#$"%&'()
."'6&6&&)
0$#"&"'23/- +"C/-&6&&)
#
*(#($&25
!"2"+,?/-) =+,#"() 4/-2#&"(
H5%.(2"+"(
#
('6(
25.(-
!"%."'/+").5#"%&$"+&:'2(#'"2&,'"+)!,'>(#('6()
1
"
7#$(#),/2),>)63",-P)8&''"("').+"'2)'"%(-)"'$)23(&#)
!"%."'/+"
#
M(#?+(&63('$()!3,#,+,?&()$(#)
L,#+$)63(6C+&-2)"'$)I&I+&,?#".35),>)!"%."'/+1
"6("(
!"%."'/+"-P)4)?"#$('(#S-)?/&$(
!"%."'/+"
)
&
2(#'"2&,'"+)."#2&6&."2&,'P)9,,C),>)4I-2#"62-
!"%."'/+"
H.(6&(-).+"'2"#/%
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
1
1
!"%."'/+"
7',-%"
#
#
#
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
R&'"#,+"6(#2")%,-,#('-&-
TAXON
#
#
!#"2"(?/-
K(-.&+/-
!"%."'/+"
=+,#")$S:2"+&"
#
"!"%."'/+"
#
2)4+.)01(
#
9&,&'>,#%"2&6:'2(#'"2&,'"+)!,$(),>)
4$,.2($)I5)23()0&?32(('23):'2(#'"2&,'"+)9,2"'&6"+)!,'?#(--)K(+1
#
#
L"3+('I(#?&"
"
#
L"3+('1
I(#?&"
!+"$&-2&6-)
M&,+")-/"A&#
"7':?9"
)=+,#"
M(#?+(&63('$()!3,#,+,?&()$(#)
"
1
!"%."'/+")
#
9&,&'>,#%"2&6-
"
#
"
"
!,#$&>,+&"()
!,'-.(62/-)>+,#"()(/#,."("(
H5%.35"'$#&>,#%(-)
!"%."'/+" !"%."'/+"
#
!"%."'/+"
H&65,6,$,'
"
N/&'Q/(+,6/+"#(!"%."'/+"
#
!"%."'/+")
9,2
"
4'$#,-"6()C,%,A('-&-
"
!"#$"%&'()"6#&-
#
9&,&'>,#%"2&6-
H6&('6(
!%%
"
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
TAXON
#
K(-,2#&2,')"+.(-2#&-
9"+C"')I&,$&A(#-&251."22(#')"'$).#,6(--)&')
"
M&.(#")"%%,$52(-
!"%."'/+")2#&63,6"+56&'"1.&63+(#&
4-5'(/%"
"
0$#"&"'23/-)
"6("(
#
!"%."'/+"
!52,+,?&")
"
!"%."'/+1
0$#"&"'23/0$#"&"'23/- -(#.5++&>,+&/- .&+,-/+/-
#
0$#"&"'23/-)
-(#.5++&>,+&/-
!"%."'/+").5#"%&$"+&-)
!"%."'/+").5#"%&$"+&-)
)!"%."'/+").5#"%&$"+&-)
-('-63">2
#
)!"%."'/+").5#"%&$"+&-)
)!"%."'/+").5#"%&$"+&-)
1
?#".3&6"+)-2/$5
#
1
1
-63">2
.5#"%&$"+&-)
Appendix .
#
.-I412#'* ).-IJ12#'>K )2#';12#'H
!"#$"%&'"($)*"#+,"'+-"+,(/0'1
!2($)*"#+,"'+-
!2($)*"#+,"'+)
)
H/#&'")FEGD
H/#&'")FEGE
)
)
)
)
)
)
)
)
)
)
)
H/#&'")
)
)
)
EGTEE
)
)
)
AFF
)
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
)
)
#
)
H/#&'")
!"#$%&'()&*+%",-"&.'"!"#$"%&'"($)*"#+,"'+-(+,(/0'1
TAXON
Appendix .
)
)
)
!2("&-.*/",*+".+0"
)
)
-2%2
)
U"-.#&6"
)
)
)
#
8&I(#)EGTED
#
!2(-30&%,+4'/*"
)%,'2('(?#&'"
GHI
)
)%,'2('(?#&'"
!2(-30&%,+4'/*"
"#
!2(-30&%,+4'/*"
G>>"
!2(53*-+0/'/*)
)
)+&%('-&-2%2
)
J
)
K@>"
K&C"6)EGTDG
K&C"6)EGTDF
)
)
)
)
)
)
)
HHB
)
)
)
)
%"2C"()
#
)
H@I"
)
)
E3&54,3/%J"
)
)
-2%2
)
)
H/#&'"
!"#$%&'(&)(*"+&#,(-%,"'.%+/0(.&(1#%'.(2"#$%&'34
"-2%2
H/#&'"
-2%2