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Bolanical Journal o f the Linnean Sociely (1993), 112: 293-309. With 6 figures
A cladistic analysis of Nassauvia Comm. ex
Juss. (Asteraceae, Mutisieae) and related
genera
SUSANA E. FREIRE, JORGE V. CRISCI AND LILIANA KATINAS
Departamento Cient$co de Plantas Vasculares. Museo de La Plata, Paseo del Bosque
sln., 1900 La Plata, Argentina
zyxwvuts
Received April 1992, accepted for publication December 1992
FREIRE, S. E., CRISCI, J. V. & KATINAS, L., 1993. A cladistic analysis of Nassauvia
Comm. ex Juss. (Asteraceae, Mutisieae) and related genera. Nassauuia and the most closely
related genera Calopappus and Triptilion from the southern Andes and Patagonia of South America,
form a rnonophyletic group diagnosed by the following synapomorphies: cypsela trichomes single
two-celled, cypsela testa with strengthened cells, pollen grains spheroidal to spheroidal-oblate, colpi
membrane with sexine processes, pappus bristles two to six, and pappus deciduous. Furthermore,
“Wassauuia, Triptilion, and Calopappus form a group with two other Andean genera, Moscharia and
Polyachyrus, diagnosed by occurrence of pseudocephalia and a reduction in the number of flowers to
five, three or one. A cladistic analysis of the group was undertaken using 35 characters from
morphology, anatomy, and palynology. The monophyletic terminal taxa were the 38 species of
Nassauuia, the genus Triptilion, the monotypic genus Calopappus, the genus Polyachyrus; and the genus
Moscharia. Character polarity was based on outgroup comparison using Cephalopappus. The analysis
resulted in 223 equally parsimonious cladograms, each with 70 steps and a consistency index of0.57.
A successive weighting procedure was applied, resulting in 15 cladograms with a consistency index
of 0.82. Results of the cladistic analysis support most of the current systematic classification of
Nassauuia, with three exceptions: ( 1) Nassauuia (excluding Calopappus) is paraphyletic; ( 2 ) section
Masligophorus appears to be a polyphyletic group ( N ,pygmaea does not cluster with the remaining
species of the section); (3) section Panargyrum (without N. lagascae = section Caloptilium) appears to
be a paraphyletic group. The capitula arranged in cymose conflorescences in Triptilion are regarded
as a primitive condition which gave rise to all stages present in Nassauuia (conflorescence spicate,
pseudocephalium, capitula solitary), The capitula arranged in pseudocephalia in Moscharia and
Polyac/yrus are regarded as a parallel development to the pseudocephalium found in Nassauuia.
Nassauuia, subgenus Strongyloma appears as the most primitive taxon, with its spicate conflorescence,
whereas section Mastigophorus with its solitary capitulum is thought-derived. These results
correspond well with cytological data where species of the subgenus Strongyloma have n = 1 I and the
species of section Masfigophorus are tetraploids ( n = 22).
ADDITIONAL KEY WORDS:-Cytology
~
evolution of heads
~
phylogeny.
CONTENTS
Introduction .
. . . . . .
Material and methods.
. . . .
Results . . . . . . . .
Discussion
. . . . . . .
Placement of Calopappus . . .
Evolution of flowering heads . .
Pappus
. . . . . . .
Cytology . . . . . . .
Geographic distribution and ecology
0024-4074/93/080293
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0 1993 The Linnean Society of London
294
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S. E. FREIRE E T AL.
Acknowledgements
References
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INTRODUCTION
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308
308
zy
Nassauvia Comm. ex Juss. is found in the southern Andes and Patagonia of
South America. T h e 38 species of Nassauvia (Arroyo & Marticorena, 1988;
Cabrera, 1982) have been divided into two subgenera and four sections based on
plant growth habit, cypsela morphology, pappus, and leaf shape (Cabrera, 1.c.).
Although some hypotheses of relationships between species have been postulated
(Cabrera, 1982; Hunziker et al., 1991; Stuessy et al., 1988), their genealogy is still
unclear.
Nassauvia, with Triptilion, Calopappus, Moscharia and Polyachyrus, represent an
interesting group within the subtribe Nassauviinae (Crisci, 1974a, 1980) because
of the several disjunct distribution patterns of closely related species and
advanced characters within the family, such as the occurrence of secondary
capitula or pseudocephalia in several species. Nevertheless, Nassauvia, Triptilion
and Calopappus form a monophyletic group based on several synapomorphies
(cypsela trichomes single two-celled, cypsela testa with strengthened cells, pollen
grains spheroidal to spheroidal-oblate, colpi membrane with sexine processes,
pappus bristles two to six, pappus deciduous).
This paper presents a cladistic analysis of Nassauvia using a Wagner parsimony
algorithm (Farris et al., 1970). To test the monophyly of Nassauvia, we have
included the most closely related genera Triptilion, Calopappus, Polyachyrus, and
Moscharia as part of the ingroup. All of them have been recently monographed
(Nassauvia: Cabrera, 1982; Calopappus: Crisci & Freire, 1986; Triptilion: Katinas et
al., 1992; Polyachyrus: Ricardi & Weldt, 1974; Moscharia: Crisci, 1974b).
MATERIAL AND METHODS
Nassauvia, Triptilion, Calopappus, Moscharia and Polyachyrus belong to the
subtribe Nassauviinae (Crisci, 1974a), which is delimited from the rest of the
Mutisieae by the following three characters: disc florets bilabiate and style
branches truncate a t the apex with a crown of collector hairs.
The monophyletic terminal taxa of this analysis are: (1) the genus Polyachyrus
(seven species), which is characterized as a monophyletic group by the presence
of pseudocephalia composed of up to ten heads disposed at different levels;
(2) the genus Moscharia (two species), which can be also delimited from all other
genera of the subtribe by the pseudocephalia composed of eight to ten heads
attached a t the same level; (3) the genus Triptilion (seven species) which is
characterized as a monophyletic group by a pappus formed of four to five paleae
plicate along the median line, and expanded and laciniate in the upper part;
(4)the monotypic genus Calopappus; (5) the 38 species of Nassauvia. Table 1
shows their geographical distribution.
Data from 35 characters were derived from plant habit, leaf morphology,
growth characteristics, capitula arrangement, capitula morphology, cypsela
morphology, pappus morphology and pollen grain type (Cabrera, 1982; Grau,
1980; Parra & Marticorena, 1972). Table 2 shows the characters and character
states used in the cladistic analysis.
z
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CLADISTICS OF NASSAUVIA
295
TABLE
1. Taxa studied, acronyms, and geographical distributions
Taxa
Acronym
Geographical distribution (Cabrera, 1982; Crisci, 1974a)
Calopappus
Cephalopappus
Macrachaenium
Moscharia
CAL
CEP
MAC
MOS
central Chile
northeastern of Brazil (Bahia)
Nothofagus forest
central Chile
Nassauuia
Subgen. Jvassauuia
Sect . Caloptilium
N . lagascae
LAG
Argentina (S Mendoza to Santa Cruz), Chile (Aconcagua to
Cautin, and Magallanes)
Sect . MastigophoruJ
N . ameghinoi
N . gaudichaudii
N. hilii
N. juniperina
N . pentacaenoides
N. pygmaea
AME
GAU
HIL
JUN
PEN
PYG
Argentina (S Chubut and Santa Cruz)
Argentina (Falkland Islands = Mas Malvinas)
Argentina (Neuqutn)
Argentina (Chubut, Santa Cruz)
Argentina (Santa Cruz)
Argentina (Neuqutn to Tierra del Fuego, Isla de 10s Estados),
Chile (Ruble to Magallanes)
Sect. Nassauuia
N. argentea
N. argvrophylla
N. coronipappa
N. chubutensis
N. cumingii
AGE
AGY
COR
CHU
CUM
Argentina (Neuqutn), central Chile
Argentina (Neuqutn to Chubut)
Chile (Ultima Esperanza)
Argentina (Chubut)
Argentina (S San Juan to Mendoza), Chile (Coquimbo to
Santiago)
Argentina (Neuqutn, Rio Negro, Chubut), Chile (Llanquihue,
Magallanes)
Argentina (Neuqutn), Chile ( T a k a to Bio-Bio)
Argentina (Rio Negro to S Chubut)
Argentina (S Tierra del Fuego, Isla de 10s Estados)
Argentina (Santa Cruz, Tierra del Fuego)
Argentina (Mendoza, Neuqukn), Chile (Santiago to Curicb)
Argentina (S Mendoza, N Neuquin)
Argentina (S Neuqutn, NW Rio Negro)
Argentina (Mendoza, N Neuqukn), Chile (Santiago to Bio-Bio)
central Chile (Linares)
Argentina (Mendoza to Santa Cruz), Chile (Aconcagua to
Magallanes)
Argentina (Mendoza)
Argentina (Santa Cruz)
Argentina (Falkland Islands = Mas Malvinas)
Chile (Curic6 to Malleco), Argentina
Argentina (S Mendoza, N Neuqukn)
N. dentata
DEN
N. digitata
N . dusmii
N . latissima
N . magellanica
N . pinnigera
N . planifolia
N . pulcherrina
N. pyramidalis
N . ramosissima
N. revoluta
DIG
DUS
LAT
MAG
PIN
PLA
PUL
PYR
RAM
REV
N. ruizii
N. sceptrum
N. serpens
N. sprengeiioides
N . sublobata
RUI
SCE
SER
SPR
SUB
Sect. Panargyrurn
N . aculeata
ACU
zyx
N . darwinii
N . glomerata
N. looseri
N. unijlora
DAR
GLT
LOO
UNI
Argentina (Mendoza to Tierra del Fuego), Chile (Aconcagua to
Magallanes and Aisen)
Argentina (Neuqutn to Tierra del Fuego), Chile
Argentina (N Mendoza, N Neuqukn), Chile (Santiago to T a k a )
Chile (Santiago)
Argentina (S San Juan, N Mendoza), Chile (Aconcagua)
Subgen. Stronuloma
N . axillaris
N. fuegiana
N . glomerulosa
N . maeuiae
N . uticina
AX1
FUE
GLS
MAE
ULI
S Bolivia to central Chile and Argentina (N Chubut)
Argentina (S Mendoza to N Tierra del Fuego)
Argentina (S Mendoza to Santa Cruz)
Argentina (S Santa Cruz)
Argentina (Rio Negro to Santa Cruz)
Polyachyru
POL
S Peru, central Chile
Triptilion
TRI
central Chile, Argentina (Neuqutn to Santa Cruz)
296
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S. E. FREIRE E T A L .
TABLE
2. Characters and character states used in cladistic analysis of Nussauviu and allied genera
Characters
Character states
1 Habit
2 Life cycle
3 Limb of leaves multicostate
4 Brachyblast
5 Leaves
6 Shape of leaf
7 Shape of leaf base
8 Auricles
9 Cross section of leaf
10 Basal leaves in rosettes
11 Phyllotaxy
12 Type of inflorescence
0. Herbs; I . Shrubs
0. Perennial; 1. Annual
0. Absent; 1. Present
0. Absent; I . Present
0. Entire; 1. Partite
0. Obovate, ovate; 1. Linear-subulate
0. Non-vaginate; 1. Vaginate
0. Absent; 1. Present
0. Aplanate; 1. Circular
0. Present; 1. Absent
0. Spreading; 1. Crowded
0. Conflorescence cymose (paniculate, spicate or corymbose);
I . Pseudocephalium; 2. Solitary
0. Absent; 1. Present
0. Conical; 1. Convex
0. Monomorphic; 1. Dimorphic
0. In branched cyme; 1. Solitary
0. 2; 1. I ; 2. 3
0. Outer bracts longer than inner; 1. All the same length
0. More than 5; 1. Less than 5
0. Less than 15 mm; 1. More than 17 mm
0. 5 (rarely 4); 1. 1-2 (rarely 3)
0. White; 1. Violet or pink
0. Present; 1 . Absent
0. Double trichomes ‘Zwillingshaare’; 1. Single trichomes twocelled
0. External layer of the testa with epidermal cells not strengthened; 1. Lateral and basal walls of the testa epidermis
strengthened
0. Subprolate to prolate-spheroidal; 1. Spheroidal to
spheroidal-oblate
0. Smooth; 1. With sexine processes
0. Infratectum thickness; 1. Tectum and infratectum equal
thickness
0. Plumose; 1. Entire or ciliate
0. Completely plumose; I . Plumose on the distal half
0. 10 or more; I . 2-6
0. Flat; 1. Plicate
0. Hairy (less than 0.1 mm wide); 1. Chaffy (0.2 to 1 mm
wide); 2. Paleaceous (more than 1.5 mm)
0. Persistent; 1. Deciduous
0. Pappus and corollas (upper lip) equal; 1. Pappus shorter
than corollas (upper lip); 2. Pappus reduced or absent
13 Pseudoinvolucre
14 Receptacle of the pseudocephalium
15 Capitula of the pseudocephalium
16 Type of pseudocephalia
17 Number of series of involucre
18 Length of involucral bracts
19 Number of bracts of involucre
20 Height of involucre
21 Number of flowers per capitulum
22 Colour of corollas
23 Cypselas pubescent
24 Cypsela hairs
25 Cross-section of cypsela testa
26 Shape of pollen grains
27 Colpus membrane
28 ’rectum and infratectum
29 Margin of pappus bristles
30 Pappus plumose
3 1 Number of pappus bristles
32 Surface of pappus bristles
33 Type of pappus
34 Pappus duration
35 Length of bristles pappus
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Character polarity was determined by outgroup comparison (Humphries &
Funk, 1984; Maddison et al., 1984; Watrous & Wheeler, 1981) with Cephalopappus
as the outgroup which, according to Crisci (1980), appears at the base of the tree
branch of Nassauvia, Triptilion, Calopappus, Moscharia and Polyachyrus. Nassauvia,
related genera and Cephalopappus form a monophyletic group diagnosed by a
reduction in the size of capitula and the reduction in the size or number of
pappus bristles. Because the pappus is absent in Cephalopappus, we used
Macrachaenium as a second outgroup (Karis et al., 1992). However, identical
results were obtained by using ‘question marks’ for these characters involving the
pappus in Cephalopappus.
Table 3 contains the data matrix used in this analysis. Four of the 35
characters (12, 17, 33 and 3 5 ) had more than two character states. Characters
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CLADISTICS OF NASSAUVIA
297
TABLE
3. Data matrix
Taxa
1 2 3 4 5 6 7 8 9 10 I 1
1 OUT
2 CAL
3 MOS
4 POL
5 TRI
6 ACU
7 AGE
8 AGY
9 AME
10 AX1
1 1 COR
12 CUM
13 CHU
14 DAR
15 DEN
16 DIG
17 DUS
18 FUE
19 GAU
20 GLS
21 GLT
22 HIL
23 JUN
24 LAG
25 LAT
26 LOO
27 MAE
28 MAG
29 PEN
30 PIN
31 PLA
32 PUL
33 PYG
34 PYR
35 RAM
36 REV
37 RUI
38 SCE
39 SER
40 SPR
41 SUB
42 ULI
43 UNI
12 13 1415 1617 18 19202122232425262728293031 3233 3435
0 0 0 0 0 0 0 0 0 0 0 O ? ? ? ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
1 0 0 0 0 1 1 0 0 1 1 2 ? ? ? ?2 0 0 1 0 0 I ? 1 1 0 1 1 2 0 0 1 0 0
0 1 0 0 1 ? 0 1 0 0 0 1 1 1 1 0 l ? 1 0 1 1 0 0 0 0 1 0 0 0 0 0 1 0 2
0 1 0 0 1 ? 0 1 0 0 0 1 0 0 0 0 1 ? 0 0 1 1 0 0 0 0 1 0 0 0 0 0 1 0 1
0 1 0 0 1 ? 0 0 0 0 0 0 ? ? ? ?0 1 0 0 0 1 0 1 1 1 1 1 0 1 I 1 1 1 1
0 0 0 0 0 1 0 0 0 1 1 0 ? ? ? ? 0 0 0 0 0 0 1 ? 1 I 1 1 0 0 0 0 0 0 0
0010000001 1 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 1 ? 1 0 1 I 1
0010000001 1 1 0 0 0 1 0 0 0 0 0 0 I ? 1 1 1 1 1 ? 1 0 1 1 1
0000011011 1 2 ? ? ? ? 0 0 0 0 0 0 1 ? 1 1 1 1 1 ? 1 0 1 1 0
100101100 1 1 O ? ? ? ? 0 0 0 0 0 0 0 1 1 1 1 1 1 ? 1 0 1 1 1
0 0 ? 0 0 0 0 0 ? 1 1 1 0 0 0 1 0 0 0 0 1 0 1 ? 1 1 1 1 l ? 1 1 2 0 1
0 0 ? 0 0 0 0 0 0 I 1 I 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 1 ? 1 0 1 1 1
0010100001 ? O ? ? ? ? 0 0 0 0 0 0 1 ? I 1 1 1 1 ? 1 0 1 1 1
000001000 1 ? 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 0 0 0 0 0 0 1
0010000001 1 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 1 ? 1 0 1 1 1
0 0 1 0 1 ? 0 0 0 1 1 1 0 0 0 1 0 0 0 0 0 0 l ? 1 1 1 1 1 ? 1 0 1 1 1
001000000 I 1 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 1 ? 1 0 1 1 1
100101100 1 I O ? ? ? ? 0 0 0 0 0 0 0 1 1 1 1 1 1 ? 1 0 1 1 1
000001101 I I Z ? ? ? ? 0 0 0 0 0 0 1 ? 1 1 1 1 I ? 0 0 1 1 0
100101100 I I O ? ? ? ? 0 0 0 0 0 0 0 1 1 1 1 1 1 2 1 0 1 1 1
0 0 ? 0 1 ? 0 0 ?I I 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 I ? 0 0 0 0 0
000001101 1 I 2 ? ? ? ? 0 0 0 0 0 0 1 ? 1 1 I I I ? 1 0 1 1 0
000001 101 1 I 2 ? ? ? ? 0 0 0 0 0 0 1 ? 1 1 1 1 1 ? 1 0 I I 0
001000000 1 1 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 0 0 0 0 0 0 0
0010000001 1 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 1 2 1 0 1 1 1
00?00100?1 1 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 1 ? 0 0 0 O ?
1001011001 l o ? ? ? ? 0 0 0 0 0 0 0 1 1 1 1 1 1 ? 1 0 1 1 1
0010000001 1 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 1 ? 1 0 1 1 1
000001101 1 1 Z ? ? ? ?0 0 0 0 0 0 1 ? 1 1 1 1 1 ? 0 1 2 1 0
001000000 1 1 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 1 ? 1 0 1 1 1
001000000 1 1 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 1 ? 1 0 1 1 1
001000000 1 1 1 0 0 0 1 0 0 0 0 0 0 I ? 1 1 1 1 1 ? 1 0 I 1 ?
000000100 1 1 1 0 0 0 1 0 0 0 0 0 0 I ? 1 1 1 I 1 ? 1 0 1 1 0
001000000 1 1 1 0 0 0 1 0 0 0 0 0 0 I ? 1 1 1 1 1 ? 1 0 1 1 1
001000000 1 1 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 I 1 1 ? 1 0 1 1 ?
001000000 1 1 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 1 ? 1 0 1 1 1
001000000 1 1 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 1 ? 1 0 I 1 1
001000?00 1 1 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 1 ? 1 0 1 1 1
001000000 I 1 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 l ? 1 0 1 1 1
001000000 I 1 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 1 ? 1 0 1 1 1
00101?0001 I 1 0 0 0 1 0 0 0 0 0 0 1 ? 1 1 1 1 I ? I 0 1 1 1
1001011001 I O ? ? ? ? 0 0 0 0 0 0 0 1 1 1 1 1 I ? I 0 1 1 1
0 0 0 0 1 ? 0 0 0 I I 2 ? ? ? ? 0 0 0 O ? 0 1 ? 1 1 I I I ? 0 0 0 0 0
zyxwvutsrqpo
17, 33 and 35 were treated as non-additive (equivalent to ‘unordered’ in other
analyses). Character 12 (type of inflorescence) was treated as additive. Despite
the fact that for some authors (e.g. Hauser & Presch, 1991) the hypothesis of
order in character changes is unnecessary, we believe that character 12 should be
treated as ordered because as was pointed out by Cabrera (1982) the changes
from cymose conflorescence to solitary capitula seem to be more parsimonious
when it has a transformation series through the pseudocephalium. Characters 6,
13-16, 24, 30 are related to characters 5, 12, 23 and 29, respectively. Therefore
we have coded those taxa with partite leaves (character 5, state l ) , without a
pseudocephalium (character 12, state 0 or 2), with glabrous cypselas (character
23, state l ) , with entire or ciliate pappus bristles (character 29, state 1) as ‘?’ in
characters 6, 13-16, 24 and 30.
298
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S. E. FREIRE ET AL.
The data were analysed using Farris’s phylogenetic program, Hennig86
version 1.5 (Farris, 1988), applying the options ‘mhennig’ for constructing
multiple initial cladograms, followed by ‘bb*’ (branch breaker), an heuristic for
determining the shortest tree. We also used the successive approximations
weighting procedure in Hennig86, which calculates weights from the last fits to
the prior round of most parsimonious trees. The weights are based on the
rescaled consistency index ( r c ) , which is the product of character consistency (c)
and character retention index ( r ) . The value is rescaled to lie within a range of
0-1 0. T h e weighting procedure is repeated on successively produced trees until
the trees no longer change (Farris, 1989). When the analysis yielded more than
one tree, we constructed a strict consensus tree using the Nelsen option of
Hennig86.
zyxw
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RESULTS
Two hundred and twenty three equally parsimonious cladograms were
generated from our data matrix. All have 70 steps and a consistency index of
0.57. If we exclude the autapomorphies for the genera Moscharia (characters 13,
14, 15, 19), Triptilion (characters 18, 30), and Calopappus (character 20), the
length is 63 steps and the consistency index is 0.50. The strict consensus tree of
the 223 cladograms (Fig. 1) shows that six monophyletic groups appear in all of
them: ( 1) MOS-POL (annual, leaves partite, auricles, pseudocephalium,
involucre one-seriate, flowers one to three, corollas violet); (2) all taxa except
MOS and POL (cypsela hairs single two-celled, testa epidermis strengthened,
pollen grains spheroidal to spheroidal oblate, colpi membrane with sexine
processes, pappus bristles two to six, pappus deciduous); (3) all taxa except
MOS-POL and T R I (basal leaves not in rosette, leaves crowded, pappus bristles
entire or ciliate); (4) UNI-GLT-LOO (LAG-DAR-ACU) (leaves non-vaginate,
hairy pappus); (5) LAG-DAR-ACU (pappus bristles plumose); ( 6 ) AXI-FUEGLS-MAE-ULI (shrubs, brachyblasts, leaves linear-subulate, leaves vaginate) .
When the successive approximations weighting procedure was applied, 15
minimum length trees were obtained after the third round of weighting, with a
length of 292 steps (the high value for the length is a function of the weight being
scaled up to a value of lo), and the consistency improved to 0.82. The values of
the range and number of steps and consistency indices (c), retention indices ( r ) ,
and weights (rc x 100) for each character are listed in Table 4. After the
procedure, the maximum weight of 10 was assigned to characters 4,8-11, 13-20,
and 23-26; a weight of 4 to characters 3, 29 and 35; a weight of 3 to character
33; a weight of 2 to characters 2, 6-7, 12, 21-22, 31 and 34; a weight of 0 to
characters 1, 5, 27 and 32. The strict consensus tree of the 15 cladograms (Fig. 2)
shows seven additional monophyletic groups not included in the strict consensus
tree of the 223 cladograms, listed with their synapomorphies: (1) all taxa except
MOS-POL, T R I and the species of the subgenus Strongyloma (pseudocephalium
solitary, cypselas glabrous); (2) PYG ( ( (HIL-JUN-AME) GAU-PEN)
(CAL(UNI(GLT-LOO(LAG-DAR-ACU)))))
(leaves vaginate, pappus and
corolla of equal long); (3) ( ( (HIL-JUN-AME)GAU-PEN(CAL(UNI(GLT-LOO
(LAG-DAR-ACU) ) ) ) ) (leaves linear-subulate, capitula solitary and bristles
pappus 10 or more); (4)(HIL-JUN-AME)GAU-PEN (cross section of leaf
circular); ( 5 )HIL-JUN-AME (pappus bristles two to six); (6) CAL(UN1-GLO-
z
OUT
MOS
POL
TRI
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z
7
zyxwvutsrqponmlk
AGY
COR
CUM
DEN
DIG
DUS
LAT
I
-
MAG
PIN
I
PUL
PYR
RAM
REV
RUI
SCE
SER
SPR
SUB
CHU
PVG
HIL
JUN
AM€
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MAG
PIN
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PUL
I
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MAE
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ACU
Figure 1. Strict consensus cladogram from the 223 most parsimonious
cladograms. Numbers 1-6 show six monophyletic groups that appear
in all of the 223 cladograms. Refer to Table 1 for acronyms.
Figure 2. Strict consensus cladogram from 15 most parsimonious cladograms after
applying the successive weighting procedure. Numbers 1-7 show seven additional
monophyletic groups not included in Fig. 1. Refer to Table 1 for acronyms.
zyxwvutsrq
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S. E. FREIRE E'T AL.
300
TABLE
4. Character consistencies (c) and retention indices ( r ) as the best fits on the 15 equally
parsimonious trees from Hennig86, used to calculate weights. Final weights were obtained after the
third round of the successive weighting procedure in Hennig86. Weights were scaled to integers
Character
~
Range of
step
Number of
step
Consistency
index (c)
Retention
index ( r )
Weight
(rc x 10)
Final
weight
~
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
I
1
1
1
1
1
1
1
1
1
I
2
1
1
1
1
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
2
I
2
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2
2
2
1
0.50
0.50
0.50
7
0.14
0.33
0.33
1
1
3
3
1
1
1
1
6
1
I
1
I
2
1
1
1
2
2
1
1
1
1
2
1
2
1
3
3
4
3
4
1
I
I
0.33
1
1
1
1
1
1
1
1
0.50
0.50
1
1
1
1
0.50
1
0.50
1
0.33
0.33
0.50
0.33
0.50
0
0.50
0.92
1
0.14
0.77
0.71
I
1
1
1
0.60
1
1
1
1
1
1
1
1
0.50
0.50
1
1
1
1
0
1
0.83
1
0.81
0
0
2
4
10
2
4
4
10
10
10
10
2
10
10
10
10
10
10
10
10
2
2
10
10
10
10
0
10
4
10
4
0
0.71
3
0.80
0.8 I
2
4
0
2
4
10
0
2
2
10
10
10
10
2
10
10
10
10
10
10
10
10
2
2
10
10
10
10
0
10
4
10
2
0
3
2
4
LOO(LAG-DAR-ACU))) (pappus persistent); ( 7 ) GLT-LOO(LAG-DARACU) (pseudocephalium) .
The 15 trees obtained after successive weighting can be further simplified to
ten topologies without loss of parsimony. The reason for this is that the Hennig86
regards trees as distinct if there are any different character interpretations to
distinguish them and producing extra branches in some cladograms (Carr et al.,
1990). In five of our 15 cladograms appear simultaneously two groups: (GAUPEN) and (HIL-JUN-AME), which cannot be supported at the same time in
the same cladogram.
Figure 3 illustrates one of ten equally parsimonious cladograms. Figure 4
shows the ten different configurations for the positions of N . coronipappa,
N. chubutensis and HIL-JUN-AME-GAU-PEN for the middle of the cladogram.
The differences in the positions of N . coronipappa and N . chubutensis (Fig. 3)
result from changes in character 3 (presence or absence of leaves multicostate)
and character 12 (presence of pseudocephalium). Character 3 is missing for
N . coronipappa (i.e. '?'), so the presence of this character is assumed in Figure
4a, c, e, i, whereas absence is assumed in Figure 4b, d, f-h, j. We have no
zy
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Figure 3. O n e of the ten most parsimonious cladograms found after applying the successive weighting procedure. Consistency index = 0.82. Character state changes are
superimposed onto the cladograms; single lines = synapomorphies; double lines = homoplasies (parallel or convergent evolution), X = reversals. Refer to Table 1 for acronyms.
5
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characters 3, 12 and 31.
JV. coronipapfa (COR), N . chubutensis (CHU), JV. gaudichaudii (GAU), JV. pentacaenoides (PEN),
N . hiffii (HIL),N.juniperina UUN), and N . ameghinoi (AME), resulting from the changes in
Figure 4. Variation in the topologies of the middle of the ten most parsimonious trees found after
applying the successive weighting procedure. The ten configurations show the different positions of
I -
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CLADISTICS OF NASSAUVIA
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303
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After
Character
12
number
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coded
o + 1 + 2
0
cladograrn
construct ion
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17
31
32
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33
34
36
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Figure 5. Character transformation for selected characters as coded and after cladogram
construction.
304
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S. E. FREIRE E T A L .
reason to choose between the alternative patterns found in character 3. I n
character 12, the hypothesis that the pseudocephalium is lost (Fig. 4a-d, f-g)
requires a developmentally more complex pattern of changes than the
alternative of two parallel presences (Fig. 4e, h-j). We have chosen the last case
where N . chubutensis, is isolated from the remaining species of the section
Nassauvia, because it is the only species that inhabits the Patagonian steppe,
whereas the other species of section Nassauvia are present in the High Andean
steppe.
The different positions of HIL-JUN-AME-GAU-PEN complex are the result
of changes in character 31. Character 31 (number of pappus bristles) show two
patterns: one reversal to ten or more pappus bristles with one reduction to two to
six pappus bristles (Fig. 4c, f-g, i-j) and independent reversals to ten or more
pappus bristles (Fig. 4a-b, d-e, h). We have chosen to illustrate in the reversal
followed by reduction in Figure 3.
The cladogram chosen (Fig. 3 ) shows that 18 characters remain as originally
coded (characters 4, 8-1 1, 13-16, 18-20, 23-26, 28, and 30). Of the remaining
17 characters (Fig. 5), seven show parallelisms (characters 1-3, 5, 21, 22 and
32), five show parallelisms and reversals (characters 6, 7, 12, 33 and 34) and four
show reversals (characters 27, 29, 31 and 35).
DISCUSSION
Our analysis suggests that Nassauvia, is a paraphyletic group. However, if
Calopappus is included within the genus, Nassauvia is likely to be monophyletic.
The monophyletic groups found in the cladograms of our analysis correspond
to those of the current infrageneric classification of Nassauvia (Cabrera, 1982)
with two exceptions: (1) section Mastigophorus is polyphyletic because N . pygmaea
does not cluster with the remaining species; and (2) section Panargyrum is
paraphyletic because N. lagascae ( = section Caloptilium) is not included within
the section Panargyrum.
The major section of the subgenus Nassauvia, section Nassauvia, is a
monophyletic group diagnosed by multicostate leaves. Within section Nassauvia,
there is no resolution among the species in this analysis. The other monophyletic
group is subgenus Strongyloma defined by four synapomorphies: shrubs with
brachyblasts, leaves linear-subulate and vaginate.
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Placement of Calopappus
In two studies of the subtribe Nassauviinae (Crisci, 1974a, 1980) Calopappus is
considered as a valid genus separated from Nassauvia. According to Crisci (L.c.),
Triptilion and Nassauvia form the sister group of Calopappus, and these three
genera together form the sister group of Polyachyrus and Moscharia. Calopappus was
recognized as a distinct genus (Crisci & Freire, 1986) based on capitula length,
17-25 mm long, pollen morphology (‘type Calopappus’, Crisci, 1974a) and three
rows of involucral bracts. However, in light of the results obtained here,
Nassauvia constitutes a monophyletic genus only when Calopappus is included in
it, diagnosed by the presence of entire or ciliate pappus bristles, crowded and
basal cauline leaves not occurring in a rosette. The distinctive appearance of
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CLADISTICS OF NASSAUVZA
305
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Calopappus is due to the possession of several derived characters (autapomorphies)
mentioned above.
Within Nassauuia, Calopappus is related to species of subgenus Nassauuia by its
glabrous cypselas. Within this large subgenus, Calopappus combines the species of
section Mastigophorus (excluding N. pygmaea) and the species of section Panargyrum
(including section Caloptilium). These three groups have linear-subulate leaves,
solitary capitula, and ten or more pappus bristles.
Calopappus was previously included in subgenus Mastigophorus (Weddell, 1855)
and compared with N. pentacaenoides of section Mastigophorus by Cabrera (1982).
Nassauuia pentacaenoides possesses pollen grains of intermediate size as compared
with Calopappus and other species of Nassauuia. On the other hand, Calopappus
appears to be related to members of section Panargyrum by its persistent pappus.
Evolution o f Jowering heads
Evolution of capitula in Asteraceae has been the subject of several
investigations during the 1960s and 70s (Burtt, 1961, 1977; Leppik, 1960, 1970,
1977; Zohary, 1950). Often observed is a reduction in the number of flowers in
the capitula and the aggregation of capitula into capitate secondary
inflorescences (pseudocephalium, Troll, 1928). Approximately 70 genera (7% of
the genera in the family) have pseudocephalia, and occur in nine tribes (Crisci,
1974b; Good, 1931; Stuessy, 1978). Within the Nassauviinae, Nassauuia and
related genera show a trend towards reduction of the number of flowers in the
capituIa and the aggregation of capitula into capitate secondary inflorescences,
as pseudocephalia (Crisci, 1974a). I n Nassauuza all stages are found between a
cymose conflorescence and a solitary capitulum or pseudocephalium.
In our cladistic analysis (Fig. 6) the corymbose conflorescence in Trz$tilion is
regarded as primitive, from which three different degrees of head condensation
were derived: ( 1 ) spicate conflorescence (subgenus Strongyloma, and
N. chubutensis); (2) heads arranged in pseudocephalium in cymose or racemose
clusters (Moscharia and Polyachyrus); (3) solitary pseudocephalium (section
Nassauuia except N . chubutensis, JV. pygmaea, N. lagascae, section Panargyrum except
N . unijora and N . aculeata). Within the context of the cladogram presented, the
solitary capitula ( N . uniJora, Calopappus, section Mastigophorus except N. pygmaea)
are regarded as derived, originating by a reduction in the total number of the
capitula from a cymose conflorescence. By the same context, the occurrence of
secondary heads in Moscharia and Polyachyrus is regarded as a parallel
development from the pseudocephalia found in Nassauuia.
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Pappus
From the cladogram variation in the pappus can be examined as an
evolutionary scenario (Fig. 6). From the condition in Macrachaenium of plumose
hairy and numerous pappus bristles, there is an increase in the width of pappus
bristles (Moscharia, Polyachyrus), followed by a reduction in size (in Moscharia) or
number of pappus bristles, becoming deciduous (disfunctional) in the latter case.
From the chaffy pappus composed of two to six pappus bristles, two evolutionary
lines evolved: one (culminating in Triptilion) where the pappus bristles are
plumose only in the upper part, and a second major line that begins with ciliate
306
zyxwvuts
zyxwvut
zyxwv
zyx
zy
S. E. FREIRE E l AL.
d
Y
m
a
zyxwvu
zyxwvu
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CLADISTICS OF NASSAUVZA
307
or entire pappus bristles (subgenus Strongyloma, section Nassauvia, N . pygmaea)
and culminates in the plesiomorphic condition (as reversals, i.e. hairy and
numerous pappus bristles) becoming again persistent (section Panargyrum
including Caloptilium) . T h e paleaceous pappus occurs twice on the cladogram in
Nassauvia, in N. coronipappa and N . pentacaenoides and presumably evolved from a
chaffy pappus. Within Nassauvia, Cabrera (1982) considered the hairy plumose
pappus as the most primitive type and the chaffy or paleaceous pappus as the
result of the union of many hairs. Accordingly, he suggested section Panargyrum
as the basal complex of the genus Nassauvia which has given rise to sections
Mastigophorus, Caloptilium and Nassauvia. I n contrast, the results of our analysis
shows the section Panargyrum (including Caloptilium) as the most derived group.
Cytology
Within the tribe Mutisieae only about 10% of the c. 1000 species have been
examined cytologically (Crisci, 1976). Both Crisci (1976) and Cabrera (1977)
reported that counts are available for 26 of the 89 genera in the tribe. I n the
subtribe Nassauviinae, chromosome numbers have been published for 37 of the
300 species (12%) and ten for the other 22 genera (Crisci, 1976).
On the basis of our results we can assess some cytological data within Nassauvia
and related genera (Fig. 6). In this group chromosome numbers are known for
Moscharia ( n = 20, Crisci, 1974a), and 1 1 species of Nassauvia ( N . darwinii n = 11,
Moore in Crisci, 1974a; N . gaudichaudii n = 22, Moore, 1967; N . magellanica
n = 11, Moore in Crisci, 1974a; N . serpens n = 11, Moore, 1967; N . axillaris
n = 11, Hunziker et al., 1991; N. chubutensis n = 11, Hunziker et al., 1991;
N. glomerulosa n = 1 1 , Hunziker et al., 1991; N . lagascae n = 11, Hunziker et al.,
1991; N . revoluta n = 11, Hunziker et al., 1991; N . uniJEora n = 11, Hunziker et al.,
1991; N . aculeata n = 44, Hunziker et al., 1991).
The basic number in Nassauvia is x = 11; species are reported at three ploidy
levels: 2x, 4x and 8x (Hunziker et al., 1991). Our results correspond well with this
interpretation. If the trend of evolution within Nassauvia has been from an
ancestral type such as Cephalopappus (with paniculate conflorescence), then
subgenus Strongyloma (with its spicate conflorescence), would presumably be
closer in morphology to the ancestral stock than any other extant group within
the genus. The presence of diploid chromosome number of n = 11 in N . axillaris
and N . glomerulosa supports the hypothesis that subgenus is primitive. According
to its solitary capitula the tetraploid N . gaudichaudii ( n = 22) belongs to an
advanced section of Nassauvia. Nassauvia aculeata ( n = 44) is placed as the most
derived species on the cladogram.
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Geographic distribution and ecology
The distribution of the taxa (Table 1, Fig. 6) shows that four genera,
Moscharia, Polyachyrus, Triptilion and Calopappus, are restricted to central Chile
(The ‘Chilean Province’ mentioned in Cabrera and Willink, 1973). However,
Pobachyrus encroaches northern Chile and extreme southern Peru (The ‘Desert
Province’ of Cabrera and Willink, 1.c.) and Triptilion is found also in the western
part of Patagonia (The ‘Subantarctic Province’ of Cabrera and Willink, 1.c.).
Excluding Nassauvia, they do not form a monophyletic group in any of our
308
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S . E. FREIRE E T A L .
cladograms. These four genera present several apomorphic character states, such
as the annual life cycle ( Triptilion, Moscharia, Polyachyrus) , compacted head
clusters or pseudocephalia (Moscharia, Polyachyrus) or monocephalous flowering
stems (Calopappus), basally branched scrub shrub (Calopappus), linear-subulate
shaped leaves (Calopappus), and succulent leaves (Polyachyrus).
Nassauuia is found in two biogeographic provinces (Cabrera, 1982),
Patagonian and Altoandean provinces, with different ecological requirements.
The Patagonian steppe species of Nassauuia (subgenus Strongyloma; N . chubutensis;
section Mastigophorus excluding N . pygmaea and N . gaudichaudii; N . darwinii and
N . aculeata of section Panargyrum) grow under dry conditions and most of them
possess linear-subulate leaves. Within the Patagonian steppe species there are
two species groups, subgenus Strongyloma which is relatively primitive because of
its spicate conflorescence and pubescent cypselas, and section Mastigophorus with
advanced characters, such as solitary capitula. The Altoandean steppe species of
Nassauuia (N.pygmaea; N . lagascae; section Nassauvia except N . chubutensis and
N . serpens; section Panargyrum except N . darwinii and N . aculeata) grow under
more humid conditions. They have aggregations of capitula into secondary
clusters or pseudocephalia (Fig. 6). This head condensation is probably
correlated with an increase in elevation and shortening of flowering season
(Stuessy et al., 1988).
Only two species of Nassauuia grow in the Insular Province. According to the
cladogram, one of them ( N . serpens) is related to the Altoandean species, and the
other ( N . gaudichaudii) is closely related to the Patagonian species. The
complexity of the biota inhabiting the Faikland Islands was pointed out by
Morrone (1992), who identified them as a panbiogeographic node (sensu Croizat,
1958).
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ACKNOWLEDGEMENTS
We thank Paul Berry, Angel L. Cabrera, Peter Hoch and Juan J. Morrone for
critical review and discussion of the manuscript. We thank Nelly Vittet and
Piero Marchionni for technical assistance and Iona Nieva and Hugo Calvetti for
help with the illustrations. Finally we are grateful for the valuable review of
Christopher Humphries. This research was partially supported by grant number
4662-91 from the National Geographic Society. The continued support of the
Consejo Nacional de Investigaciones Cientificas y TCcnicas (CONICET),
Argentina to which the authors belong is gratefully acknowledged.
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a