Table Of ContentPHYLOGENETIC ANALYSIS Mari and
Per Ola Karis,^ Kdllersjo,^
Kdre Bremer^
THE
OF CICHORIOIDEAE
WITH
(ASTERACEAE),
ON THE
EMPHASIS
MUTISIEAE^
Abstract
We
present a cladistic analysis of 53 taxa from Asteraceae using 72 characters of mainly morphological data.
monophyletic
Resuhs reveal subfamily Cichorioideae paraphyletic. The Mutisieae constitute a basal grade that contains
is
groups with Mutisia and Cardueae sensu The Arctotideae, Liabeae, Vernonieae, and Lactuceae form a clade
lato.
together with subfamily Asteroideae. Our results show some inconsistencies with those from cladistic analyses ol
molecular
data.
mor-
The noting strong
relationships within the Asteraceae, cladistic analysis of the family,
dis-
Bremer
dichotomy. s
cussed since Cassini and Lessing in the early nine- phological support for this
monophyly
teenth century, have been scrutinized using (1987) analysis further supported the
cla-
&
subfamily
methods (Bremer, 1987; Jansen Palmer, of the subfamily Asteroideae, and the
distic
assem-
Many emerged paraphyletic
1987, 1988; Jansen aL, 1990). Cichorioideae as a
et results
yieldea
The Jansen (1990)
of cladistic analyses were inconsistent with the blage. analyses of et al.
in-
monophyletic
The
only
trafamilial systems proposed by other authors quite different topologies.
(e.g.,
Wagner and
both
Carlquist, 1976; Cronquist, 1977, 1981; Jeffrey, groupings established in all trees,
& the
Asteroideae,
1978; Wagenitz, 1976). Jansen Palmer (1987) Dollo analyses, are the subfamily
an
Liabeae,
demonstrated that Mutisieae-Barnadesiinae (Ca- Vernonieae as the sister group to the
paraphyletic
brera, 1961, 1977) form a group the the Eupatorieae as a subclade within a
sister to rest
always sup-
of the family, a finding that was corroborated by Heliantheae sensu lato. Dollo analysis
Wag-
cpDNA but the
an apomorphic inversion the group, ported a monophyletic Cichorioideae,
in latter
Cichorioi-
Bremer came paraphyletic
(1987) also to this conclusion in his ner analysis resulted in a
1*
I*
bwe
This study was supported by a grant Compositae taxonomy and Asteraceae phylogeny from the i
for
^
We and o
analyses
Natural Science Research Council. thank Arne Anderberg for discussions of complex cladistic
Donog
We
letting us examine material that he had on loan from K. thank one anonymous reviewer, Michael J.
also
•
an
Stevens,
Vicki A. Funk, Larry Hufford, Christopher Humphries, Robert K. Jansen, Guy L. Nesom, Peter F.
J.
John comments
L. Strother for valuable on earlier versions of the manuscript.
,
StockholIm,
2 Department of Phanerogamic Botany, Swedish Museum of Natural History, P.O. Box 50007, S-104 05
Sweden.
Department Sweden.
3 of Systematic Botany, Uppsala University. P.O. Box 541, S-751 21 Uppsala,
Figure 1. One of the two final cladograms. Character distributions are given below. Note that 6^"^^*
l^^^^ch
an
node
shaded area are all assigned to Mutisieae-Gochnatiinae. All apomorphies/reversals are listed for each
Apomorphies
terminal taxon. with subsequent
reversals/parallelisms are without further labeling,
P^^^'^vJ^^tatc
— Muj
marked
=, marked brackets.
reversals are with the change in multistate characters indicated within
para e
characters have the state in question indicated within parentheses. Under the terminal taxa all characters are
unless otherwise indicated.
m
A 4'
Nod^^
Node 28(0- 37, 65; =33(0-1); Node Node 38(0-1), 42, 60;
1: 1), 2: 5, 58(0-1). 68; 3: 4. 30,
27(0-1), 29(0-1), 55; Node 5: -38(1-0); Node Node Node 8:
6: 50, 51; 7: 15, 42(1 ^0);
^^^^^^Jj.'
2\
29{\ -Oy, Node 9: -4, 55; Node 10: = 21, =48(0-3), =49(0-2); Node 11: -30; Node 12: =9, -IplJ
^^.
-15;
Node
48(3-1), 59; 13: =1, =45, =46(0-2). =49(2-4), =58(1-3); Node =66; Node 15: -^^
14:
^5^
^'"^
29(0-1), =58(3-0); Node 17: =23. =38(0-1); Node -50, -55; Node 19: =1;
5, 18:
f^:>^i),
4tHJ
27(1-3), -28(1-0). =42(0-2). 64; Node 21: =20; Node 22: =16(1-2); Node 23: =46(0-1),
8,
Ann. Missouri Bot. Card. 416-427.
79: 1992.
i
-
;
. L
Number
Volume 2
79, Karis et 417
al.
1992
Phylogenetic Analysis of Cichorioideae
L^
+
^
BARNADESIIN'AE,
outgroup
-
FrHfjF'j^*Lr+Bv_+Lr'^i_^Hl_j+P_HIl_BItJjP_xtIJB_'Lq-_i^1_-J_I-^ri'^^"FKTThl^JJH+Llrjr,liblTbLihjr4-x*Tj4rH''bF_^_Llb_-I^_J
_r_T_F_l_"_F T
"h I I F « h + q
WF'r".\V,\'---'--^'---*-'.^^-' -h n T IT brdBkrJrrbi b rh4Tnr¥BBH H¥^ab+^4*-^HF^b+H¥*B^_+B_F+r_'r¥_"*4FL'_L^b'4^¥BF"H^"4X~*B4"HbriiL~brX"n¥^*V-B"HJH^b"+H*BFbJHxIH*¥^B"¥^-B^^4-H~^«^L"*^h*^X'+^H"b^H^X'^h^-H+'d^~b+^JJ"H^'H-^4B."^¥*B"^*d^'HJ~^b¥*^+*X~^lflb^^cA^"L»F-b'J«*b~Fb»^*V+_'l"^*iX^~'XLb^.i^.bb"-hJ'L'"^^+x^~^^ABX-.vdr^i-H^4'~^.br^^4^J~^-_r*-Xr^r^^_+-~b^b."^^J+^^"d'^^~xX^t'^»X-~f*^_c-^L*"F.xX'^~¥b-+"-^X+*,"d.^^'"b+.^'B'i*b+^'*_.-X¥^"^bi<-^"L.h-"*^+.~^b---rJ"H-,"J*4LfXx-rJ-4xr~H_HV-~_-.f^rr^.'4i~H-_H^¥b.*IiHb^*.^wH"-v_**tBhb^¥^Fr"HB¥X"¥B1H"_bF^_¥^*J_"I_^J14*H_^*I_^4J_H'hF_J^L*H_^*IBF_^JBHJJJ^*_^_*^-f"-'-"*-*- T h4 + F b +ri_frH_H_h_ ' **4 ^^r^4m4 _» 44"¥F'" "'^Fl ^^riHbbVAdrHlri M wA mM r ._r. . 4X,-r_Hr^^i^-^F,^
:j^i '_+___'_'__' T_Ti_'_'_'_'_'_*_^_"_'_F^'_^F_'r1+ t'4_4¥-ITT-¥h1+¥44h"frr¥i_Tq¥T14-FrI-^H¥hI+H^I4¥4_4HrT"1HiqTH1 IFIHI^I rH¥HHIHI¥*»1HF¥¥FH11¥Tl¥r¥HI¥TFrWl»¥THTH'^T''¥TTT'FF"F»'""^™»^"+*»**^+**"*T"''*^" ^
:::>:::>: '^IVHP'Lb^driHrid
J-:-; 1 * ' 1 E^ h * q +F4TH_^^Eph hF4_¥ F '^^^^'^L.^BL' '-'-*-'-'-*r^-p'H'_VFV---VF^.---¥'-^--r'--x-'-';'-%'*hbnV"j--"f'c-'L-*"i*""-^i-'-x*iB-\x» 25^ Gerbera
h*¥4H^4-ihv
^vlirBHbdHL^^Bd_XbldbdbLJXBJX_HLdbL^^dd
v>-A-5^V:*V;.VV<>i-vV>t:>VaVixV-lx;V--V^"-.x^i-'j IJ'k'ri^H*^J Ak +^^-^T-^H-|r^_-b,^^--v_^|F-^T--nT^X4H,--r_^!rv1-HbFr^x^--.-'l^-^.^B-x_-rb-x_T11jnH^Tr Txvkrb+-Jrh«F h"»T\Vh^*,'4h"+~^~x-^^^'^"_bl'"^*b"^^^^*^^_If^ri'*¥__'L^._~'_Jv^~_1"r'I .J^L-'~±^~"b*-~^*^-~hJ^~'^^~-Ai~lv~r.i*'rL-."V^^-^M-^^rL'^-&^ 244 T- Trichocline
:-;::;-: ,"b^---4---'.'j---.'- h 4b-J-+H4b"hb-*A"-H--X"^d'rb-B-.H-.B--^',H--B--d"+A-r^-,A--r,-i-H"-b-.d'*H-b--d'-A"*L'A---A-^d"'d-"bh'J- H-- MUTISIINAE
¥ 1 * ^ '-\:-'-'-'F'-v^'.'HV-\'h",'+'4^-^-\"h",'-7h\vM,^t"hVv,;-"*"4"-
I Chaetanthera
f
JLulia
—
Vi*bV^j,-rh-,-. ^FI^riHBV b^^^Vn^X----b d^^-^rL X^,'-_*^_^^^_+^_'-T^v^^^^h^4F+'^^_,-**-r^-+,^-^^-_^-_ * __*-*_ - 'a^'- b*~**J M~K
mfc^j'i'rn^ ' ^mf^^ r' F ¥n¥ Pm ^^^^^Lxb -^ Mutisia
-hVT4t b
XJHBXLHbJ
*^ + - - - * -'
^+:^b:J^,^_-.]_-:*^.:H^_:-':'T'x_FTF:¥-¥m-__w:I::JI^JH^*I>l*^lI^^l^L'±;X"^d-':^-vH H h^^BLv'^rTLJ br"nrn4i-k.--ph_B'"Ar4_-Xv+"dq4-T--n¥-^r_-x'ri-i"H..-'.r.t-_i-H'.r_.¥i^-_-pH-_"1rrbb"Pri*+¥H_¥-b^fr'+.iJj'4*AT_'4.rdi**dH_^\r^_*',mV*_*m'*_i^r_*^v*-'vm_fvc'l*_4b"'_*r^-nxr^*a&-L'.&mi""^*x"^WA^-^H^r^fV--'^'j'.^'^-"*^d*- V-%¥^1 - H-^'l F^-I'^-H b-¥I ^I J¥ I4 ¥-I LT¥ r4%i ¥1^%¥T"^f'^f^^¥*", V b + h I -I - T_fi^._^LHJ -X+ Bl-.H.-J".*L+I_HHIHr-i_Hb_rH_ixJHtb4l+ ^ll_ PfcrirJ+^JVqfrc4v4r-TTHt-r_4hv4r*b44+v4r4HrFv+rhFrirTfrhJ rr ¥ I F_¥_F"_^_' 4_h 4 4 b + h_4_^ fc_* *_^__'^__"**-__^_.*_r+J*_.^_r^_r-i'+_^_r h'_' _'_'_f''4"_-'''_r
*'j ri h +^4 -I
'
h + H 4 q - F 4 - '_^
^'
bd b L
Lop
hopappus
4 , - . x_V'
-r-:--.' >>>^^^ r:¥v**L'.I^IJ-vhI' '-:>>: x>^r>'h T V Hi m , J- b^ bh ¥ brB+L Tl^qI4H^Fq-T¥-F+ -I Trixis
NASSAUVIINAE
^Leucheria
Jungia
?^;sjssgs^i:gis?s^^ V
^— --'-^^-^ ''m*r^w'^iWM\\'M*m^M'4''*l"" \'j'j'-"+'-\.^>^["]+">b"^'>hV^\ h%-*-4'!", JI
J ~L~.~^ M :j_-_- H-fc - * T t h TV-^-V." - - ¥^H__q F I ' ri~H~L^^b L^^^'ri b x'^'d'a IS r Nassauvia
.-:r;::I:X;'X-':d^>b:^-:v;f::4'v:.:::,;:,>>;v.^^^:'Mv::w-'>:-;^ q JIF 1HHJHbl4IJI^L^IFXLHL+b^Xnd-JidBbV-^JdVTb4+rdnxiL--Hrb4A'T-XLVdHbBBr^vKvxi '..
^ h 4^^Hh 4m4mT V h4*n4-vi h4+ ¥JH4 r 4 Macrachaenium
4 4 r r| . ri_X Fi'--'T1-
_ ^ '-^¥b^_I'¥_ hb q* "L"d"±"bj4rit~B+j"4fc*"db"*~4f_c"^b^^H_^'j_"fjc*_'_^_ -H.**.-_^.J>.^.
_^*:- i-H .^^x'.- '--"-'.'-'
^^bB
^^':j:^^vl'O--^^'.''--*^.^'-v:-4-.- r4 H J+^LnX L^-T-*-' _* *_ rBvnHB-V-^_4n^v_^B_T*_B+ nd4_a» p^E'- ' '-_•-_'_*+-__^ _ jWBWVi^JjH;r-a'-r'n-'H.!r.ap_iI.v&_T_^q.vqr.h —4.-1rF* ^_'_^4*.__-. + '.^h'_^H 4 F - ^_F_» *bX*.^.XbJ"bLX4riHbri ' --:---::':^->::x:x-:>-::'->:::-:^-:::>':::-;-:^;::^:::o:;-::->:-:>^;:^::>;;^::^-::^:;^^:-^:::^^^ -X XrdrikdIriJrL
+b4Xri^*~Lib.^'
l+4-'BhTHbLX4 V J J L J J I ^ F . F r-rf ^+ r# ¥ri ¥
J r IH_HL J b b I XJ b b blLriHa + t j -rh_4¥ +1 .I . bI rXv^^h:4"+.4vH,_^-+.-%nX,4'I-.,hX-H,L^*_^b+*-b_^L.x^_i'H.^.-\j-^_^'--^.x-v^--^.j,'.b*'^--V-^^T,,_--r^V-_^»^T'-.--V-*\"-¥b»"*-vbL''--.L-^j'J--'^'"^^-_^'L^dr-''"'*-i-^4"''F'4b-^-4-'lL^j*-"\-XX_^^-*b-''fm*-.cJ"'V^*-*XJ^b--*^-X*xx'\'_"I*j-^xJ_4-'-*bl-X-_**'^_4rL_-V"**i'-_dxt4-^*-b"_-4b_*^\l*''.r-_HH*^^*_I%lj~b.j1I~\^kxb*jl.I/kHJ.h^L^Lx*^J-J*"lHh^"jxI^I,I^j^HI^x-H'F^Wd'II*t'HI..fb__.F^Xl+LX¥¥'_JT^.'_4-4T¥rm4¥'i_m^*¥^._'*-^Fw_,Ir4**.b_Hr+,r^4_Ir0H--m^_T4m¥_¥-Tm_w4-¥3V 5rf Cirsium CARDUEAE incl. Echinops
I Carduus
.:>v'^.- v"p-^_ *
Echinops
I
* i*_-
Saussurea
LACtUCEAE
-^.*----^---1^'t'-i''-4V^+'d-,-p" _^_'_' _ ' ^..^. 'y« Berardia «&
^:->'-v-"-"^*-' v^ 4 4]
Carlina
.\-b^
: -^.^ :-:*:*-: '-*r* ¥ ' j""-T4» 41-4 r44^ Munnozia LIABEAE
-T M.' \\ '- x^ -^ '- '_ ^^*
I42J
- r ri *Liabuin
'--V-^
'^- #4t«*t***t4*fl#*»
* J_-
^^
^
•'•' m-i . . ^-M^M. ^ AA^-^ ^*--t^ii* -^T^'T^ rf^ ,TT *•
A fl ^ t
Pluchea
4
Inula
Helichrysum
^„
ASTEROIDEAE
^Senecio
Aster
,
J
52 Geissolepis
-58(l-0);Afode24; -60; Node =21, =45, =46(0-1);
-68; Node =2(0-2), 26:
18(0-1), -30; =58(1-3), 25:
^orfe 27; = =49(0-1), -68; Node
- -29(1 Node =38(0-1),
Node =12. --5588((11--00)),, =69; 29:
5, 28:
*0); 4, 12,
30: -28(1-0), Node =20, =49(1-5),
=48(0 -45, -46(1-0); 33:
Node -60; Node
31: 32: 19,
3); 1,
-55;
Node Node =32(0-1);
34: =3, =58(1-0); Node Node 36: -4, -30, =42(1-2); 37:
3355:: 33(1-3), 39;
/Vode38.=l, =
Node
34. 68; Node 39: = =26, =29(1 41, =48(3-2), 67; Node 40: 16(0-3); 41: 51.
15,
2),
-55;
Node 42: 4, =17, =18(0 Node 43: = 32(1-2), -33(1-0), =48(2-3), -60, -68; Node 44. 24,
1);
-37;
A^ode 45; 10, 13, -15, == 16(3-4), 25(0 =32(1-2), 43, 54, =71(0 Node 31, -33(1-0),
3).
=35-58(1-0),
-33(1- =48(2 47(0-2),
-68, =19,
Node Node 18(0-1), .1).
70; 47: = 55, 57; 48; 0).
-37, Node
'2; A^ode 49; 36, 44, =49(1-4). -50; Node 50: 58(1-0), -60. =63; Node 51: =17, 68; 52:
_,-
18(0-1).
28(1-2). =49(4-2), 53; Actinoseris 2(0-2). 25(0-1). 29(0-1). 45. 67, 69; Ainsliaea: 1.
_''l', 1,
44
1-0), -37, -38(1-0), 42(2-1), 49(1-5), 51, 52;
66; Arctotis: 10, 19, 21, 26, 27(1-2). 28(0-1),
^^ten
-36. 16(0-5),
-58(1 71(0-1); SerfcAeja;
50, 61; Berardia: -21,26,27(1-2), -0), 67, 3, 4,
2(0-2), 20,
18(0 -4, -27(1-0), 42(1-2),
_ 2), 24, 28(0-1), 31; Brachylaena: 26, 27(1-3). -28(1-0), 68; Carrfuui;
-50, 71(0-2); Cirsium: 36, 66. 67,
^^' ^2. 71(0-2); Car^iVia; 4 11, 14, 22, 23, 49(1-5), 66,
3,
V>L •
fiH
oo;
Dislephanus:
LAaefa/iiAera; -29(1-0), 42(0-1); Dicoma: 18(0-1). 69;
10, 43(0 -3); Cnicothamnus: 36.
-67;
29(1-2).
-38(1-0), 48(3 Echinops: 24, -38(1-0), 52; Eremothamnus: -1, 3, 22, 29(2-1), 31, 35,
1);
^Othrocephalum: 29(0-1), 46(0-1), 50, 67;
Gerbera:
14, 29(0-1), 45, 46(0 50, 58(1-2), 66, 67; 5.
1),
'
"^ochnatia: -5. 16(3-6), 46(1 61
21; Gongylolepis: 16(0-2), -46(1 Helichrysum: 4, 11. 2).
17,
6. 7, 0);
Hypochoeris
"esperomannia: -30, 51,58(1
32(0-1), 45, 48(0-1), 49(0 Hyaloseris: 10, 16(0 '4),
1);
-67.
Jungia:
49(4-3). 62; 14.
-54.
oSr2i' I *• 66; /«u/a; 17, -21. 25(0-2), -32(1-0). 47(2 -1_),.. 48(1 3).
,^
. , ..
r l
-12(0-. -41, Lophopappus:
55. 70;
1); Lacfuca; 9, -25(3-0). 63; Leucheria: 25(0-2), 38(0-2); Ltafrum.- 9, 22.
-
3. -15, -50. Macrachaenium: 36, 46(2
_ 5, luii^. 24. -37, -38(1-0), 39, 50, 61; 2(2), 32(j), 1 ).
18(1),
-55.
49(4-2); 3fu„„o2m; Mufma; 40, 45, -60, 66; Nassauvia: 3, 4, 38(2),
27(2), 58(0), 71(1); 1, 25(.1).,
.
66, Per/>a; 50;
67. -68; 62, 68; 48(4).
45.
Oldenburgia: Onoseris: 19. 24. 38<1). 39. 40. 42(1),
. 7; 18(1),
naz^: "
.
22. -15. -16(1-0). -17, -21. 23. 36. 49(2-3). 55. 68;
42(0-1),
45, 49(2-3); Pleiotaxis: 10.
^S
/-^Aea; Senecio:
Sco/jmiw; 14, 24. 37, 67(2y,
16(3-6). -32(1-0). -67; Saussurea: 56. 61. 70;
25(2).
^ -50.
42(1-0).
5. 9. 38(
46(l-2);So«cA«5;3 iffi
1
-p.- 2),
^; -50.
TruhocUn -37. Warionia: 24. 27(1-3). 40,
5 66. 70;
-I, 22.
39. 40. 48(3-4); Vernonia:
?••
55.62.
-
71(0-
Wunderlichia: 25(0
22.
1 ); ).
1
