The Phylogeny of the Alstroemeriaceae, Based on Morphology, rps16 Intron, and

rbcL Sequence Data

Author(s): Lone Aagesen and A. Mariel Sanso
Source: Systematic Botany, 28(1):47-69.
Published By: The American Society of Plant Taxonomists
URL: http://www.bioone.org/doi/full/10.1043/0363-6445-28.1.47

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Systematic Botany (2003), 28(1): pp. 47–69
q Copyright 2003 by the American Society of Plant Taxonomists

The Phylogeny of the Alstroemeriaceae, Based on Morphology, rps16

Intron, and rbcL Sequence Data

LONE AAGESEN1 and A. MARIEL SANSO2

1
Botanical Institute, Gothersgade 140, DK-1123 Copenhagen K, Denmark.
Present address: Instituto de Botánica Darwinion, C. C. 22, 1642 San Isidro, Argentina
Author for correspondence (laagesen@darwin.edu.ar)
2
Instituto de Botánica Darwinion, C. C. 22, 1642 San Isidro, Argentina—Dpto. de Ciencias Biológicas,
Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Argentina

Communicating Editor: James F. Smith

ABSTRACT. Separate and combined analyses of morphology, rps16, and rbcL data were conducted on a total of 27 Al-
stroemeriaceae species including 23 Alstroemeria, three Bomarea, and Leontochir. We wished to examine the monophyly of
Alstroemeria and Bomarea as well as explore the phylogenetic position of Leontochir and two controversial monotypic genera
segregated from Alstroemeria: Schickendantzia and Taltalia. Monophyly of the Chilean and Brazilian Alstroemeria species were
explored. The results support monophyly of the family. Alstroemeria and Bomarea are both monophyletic with the former
genus being more strongly supported. Bomarea and the monotypic genus Leontochir form a clade distinguished from Alstroe-
meria by a set of morphological characters. Bomarea could not be defined by morphological characters without including
Leontochir. Resolution within Alstroemeria is only weakly supported and formal subdivisions should not be made until the
Brazilian members are revised and/or more appropriate phylogenetic markers at the species level have been found. Alstroe-
meria graminea or A. pygmaea never appear in a position that makes their segregation from Alstroemeria possible (as Taltalia
and Schickendantzia, respectively), without making Alstroemeria paraphyletic. The resolution best supported in this study
divides Alstroemeria into subclades roughly corresponding to geography: a clade from northern Chile, a clade from central
Chile, and a clade containing the Brazilian species. Most of the trans-andean species could not be unambigously placed by
this study.

Alstroemeriaceae Dumort. are mainly South Amer-
ican. They are erect or twining herbs living in a wide
range of habitats from cloud forests to swamps and
desert areas. The family includes two major genera:
Alstroemeria L. (;50 species), which has become a very
popular cut flower (the Inca lily), and Bomarea Mirb.
(;100 species), as well as the monotypic genus, Leon-
tochir Phil., and two controversial monotypic genera
segregated from Alstroemeria, Schickendantzia Pax and
Taltalia Ehr. Bayer.
The monophyly of the family has never been dis-
puted. Resupinate leaves are characteristic for nearly
all members of the family. The leaves are twisted near
the base or along the lamina, hence the abaxial leaf
surface becomes the functionally adaxial surface. The
flowers are epigynous and are nearly always zygo-
morphic as the angles between the inner lower tepal
and the two inner upper tepals exceeds 1208. The zy-
gomorphy is reinforced by the position of the fila-
ments, which are located in the lower part of the flower
during anthesis. Furthermore, all members of the fam-
ily have pseudo-basifixed anthers and thickened roots
modified to store nutrients and water (the only excep-
tions are Alstroemeria graminea Phil. and A. hygrophila
Meerow, Tombolato & F. K. Meyer).
The family Alstroemeriaceae was recognised by
Dumortier in 1829 but the genera were placed in the
Amaryllidaceae by various other authors (e.g., Herbert
1837; Baker 1888; Pax 1888; Pax and Hoffmann 1930),
due to the epigynous flowers and the umbel-like inflo-
47
rescence in most species. Buxbaum (1954) argued that
the flowers are pseudo-epigynous as the epigyny is
formed by the union of the basal part of tepals and
filaments with the ovary. Huber (1969), who studied
the seed morphology, and Dahlgren et al. (1985) both
grouped the genera within Liliales (sensu Dahlgren et
al. 1985). Dahlgren et al. (1985) recognized the Alstroe-
meriaceae as a separate family, and mentioned char-
acters such as perigonal nectaries (and lack of septal
nectaries) and variegated pattern on the tepals as pos-
sible synapormorphies for the order Liliales. Hutchin-
son (1959) grouped the Alstroemeriaceae together with
Petermanniaceae and Philesiaceae in the order Alstroe-
meriales.
In recent analyses based on DNA sequence data
Alstroemeriaceae form a clade with Luzuriaga Ruiz &
Pav., Uvulariaceae p.p., and Colchicaceae (sensu Dahl-
gren et al. 1985). This clade was found by Chase et al.
(1995a) based on data from the chloroplast gene rbcL,
by Chase et al. (1995b) based on data from a combined
analysis of rbcL and morphological data, and by Rudall
et al. (2000) based on a combined analysis of data from
rbcL, trnL-F, and morphology. These analyses show no
support for a close relationship between the Orchida-
ceae and the Alstroemeriaceae as hypothesised by
Dahlgren et al. (1985).
The genus Alstroemeria is strictly South American,
ranging from Venezuela (38 North) to Tierra del Fuego,
Argentina (538 South). It is primarily found in the
southern Andes and the Brazilian highlands, with the
majority of the taxa in Chile and Brazil.
48 SYSTEMATIC BOTANY
The Chilean Alstroemeria species were reviewed by
Bayer (1987), who recognised 31 species. Sanso (1996)
more recently reviewed the Argentinean Alstroemeria
species and recognized one endemic species. The Bra-
zilian Alstroemeria species are currently under revision
by Marta Camargo (University of Saõ Paulo).
Few attempts have been made to make an infrage-
neric classification of Alstroemeria. Baker (1888) divided
the genus into Chilean and Brazilian species using
mostly quantitative characters from the leaf and um-
bel. The Brazilian species were further divided accord-
ing to the leaf structure: thin leaves or rigid, strongly
nerved leaves, but no formal divisions were made and,
Bayer (1987), who revised the Chilean species, noted
that a subdivision of the genus is premature until the
Brazilian species can be included. However, Ravenna
(2000) proposed the section Andesine based on floral
characters, including species from the Andean region
and Patagonia.
Bomarea is found from Central Mexico and the An-
tilles (248 North) to Chile (408 South). They are mainly
vines from tropical and cloud forests, but erect, often
high Andean taxa, are also found. Only the Argenti-
nean Bomarea species have been reviewed recently
(Sanso and Xifreda 1995). Herbert (1837) divided what
is now Bomarea into three different genera mainly
based on habit. Baker (1888) retained this division at
a subgeneric level: Bomarea, Wichuraea (M. Roemer)
Baker, and Sphaerine (Herb.) Baker. Using floral and in-
florescence characters Killip (1936) later divided one of
the subgenera, Bomarea, into sections. Later Neuendorf
(1977) added a new section based on pollen characters
within the same subgenus. No recent revision of the
genus is available.
The monotypic genus Leontochir is found only in the
Atacama desert of Chile. Leontochir differs from the
other members of the family by its habit, the presence
of Fritillaria-like nectaries (Wilkin 1997), and a uni-loc-
ular ovary with parietal placentation. The monotypic
genus Schickendantzia includes small, high Andean
herbs segregated from Alstroemeria by Pax (1890) on
the basis of the nearly actinomorphic flowers and the
presence of a unilocular ovary. In contrast, Sanso
(1996) and Sanso and Xifreda (1999) studied the ovary
and found that it is trilocular with axile placentation
as in the other Alstroemeria species. They further noted
that nearly actinomorphic flowers are also present in
other Alstroemeria species. Hence the species was in-
cluded in Alstroemeria as A. pygmaea Herb.
Recently Bayer (1998a, 1998b) established the mono-
typic genus Taltalia based on the Chilean species Al-
stroemeria graminea, the only known annual Alstroe-
meriaceae species.
Hunziker (1973) challenged the validity of Bomarea,
Leontochir, and Schickendantzia due to the lack of clear
cut characters separating the genera. However, Sanso
[Volume 28
and Xifreda (1995, 2001) showed that despite confu-
sion in the literature, Alstroemeria and Bomarea differ in
morphology of the storage roots, morphology and de-
hiscence of the capsule, lack or presence of a sarcotes-
ta, ornamentation of the exine, and chromosome num-
ber.
Various authors have discussed the relative degree
of specialization among genera of the family. All dis-
cussions have been based on comparative studies in
the complete absence of a suitable phylogenetic frame-
work. Whyte (1929) studied the chromosome comple-
ment of Alstroemeria and Bomarea and suggested that
Bomarea has arisen from Alstroemeria. This was disput-
ed by Hunziker and Xifreda (1990) and Sanso and
Hunziker (1998) who also studied the chromosomes
but found it more likely that Alstroemeria had risen
from Bomarea, than vice versa. Furthermore, based on
morphology, Bayer (1988) considered the characters of
Alstroemeria as more derived than those of Bomarea. Ko-
senko (1994), who studied pollen morphology, agreed
that characters from Alstroemeria (and Schickendantzia)
seemed to be more derived than those of Bomarea and
Leontochir.
It is the goal of the present study to investigate the
monophyly of Alstroemeria and Bomarea and to examine
the phylogenetic position of the three monotypic gen-
era to determine whether they can be retained without
rendering either Alstroemeria or Bomarea paraphyletic.
We furthermore wish to explore whether the Brazilian
and the Chilean Alstroemeria species form monophy-
letic groups within the genus.
MATERIALS AND METHODS
Taxon Sampling. In this analysis twenty-three Alstroemeria
species, including A. graminea and A. pygmaea (Schickendantzia 1
Taltalia), have been used. Fifteen species belong to the Chilean
species group and seven species belong to the Brazilian species
group. The species have been chosen to represent most of the mor-
phological variation found within the genus. Furthermore, Leon-
tochir ovallei Phil. and three Bomarea species, one from each sub-
genus, are included. The complete lack of an integrated revision
of Bomarea, combined with the large number of species and the
wide distribution of the genus makes it very hazardous to do a
sub-sample within the genus for the purpose of a cladistic anal-
ysis. We have no preconceived opinion about monophyly of the
subgenera. The included Bomarea species represent the three dis-
tinct growth habits found within the genus and do not cover the
entire morphological diversity within the genus. We do, however,
consider the chosen sub-sample sufficient for a cursory test of the
monophyly of the genus.
Three species from Luzuriaga have been included as outgroups
together with Uvularia grandiflora Sm. and Disporum viridescens
(Maxim.) Nakai to represent the clade found in the phylogenetic
analyses based on DNA sequence data (Chase et al. 1995a, b, Ru-
dall et al. 2000). Finally, Dioscorea dodecaneura Vell. has been in-
cluded for the purpose of rooting (Table 1). All characters were
scored according to the states found in the included species. Pos-
sible polymorphisms within the outgroup genera are therefore not
represented.
Scoring of Morphological Characters. The scoring of morpho-
logical data was done mainly on fresh or FAA-fixed material, either
in the field or in cultivation, supplemented by a few characters
2003] AAGESEN & SANSO: PHYLOGENY OF ALSTROEMERIACEAE
taken from the literature (mainly Wilbur 1963; Bayer 1987; Rod-
rı́guez and Marticorena 1987; Arroyo and Leuenberger 1988). For
Alstroemeria andina, A. apertiflora, A. graminea, A. insignis, A. longis-
tyla, A. sellowiana, Leontochir ovallei, and Luzuriaga polyphylla only
herbarium material was available. As no material was available for
A. caryophyllaea, all characters were scored from its original de-
scription (Jacquin 1798) and Curtiss (1790). Specimens from the
following herbaria were examined: BA, BAA, BAB, BAF, BHCB,
CEN, CTES, ESA, F, G, HUFU, IBGE, K, LIL, LP, MBM, MCNS,
MERL, SGO, SI, SP, SPF, UEC, and UPCB. For the outgroup taxa,
only the states of the included species have been scored. Voucher
information not included in Sanso and Xifreda (1995) and Sanso
(1996) is listed in Appendix 3.
The characters, including discussions of their scoring, are listed
in Appendix 1. The morphological matrix is found in Appendix
2. Unless otherwise stated in the text, the character is coded as
non-additive. When polymorphism was detected all observed
states were scored for the corresponding species.
The Alstroemeria species are mainly distinguished by quantita-
tive characters of doubtful use in cladistics (Pimentel and Riggins
1987). Qualitative characters have been difficult to find, and those
included in the present analysis may not be distinct if more Al-
stroemeria species are included. For example, the extremes of the
variation found in some characters often seem phylogenetically
informative, e.g., height of plants. Some species (A. graminea, A.
patagonica, and A. pygmaea) are very small whereas others attain a
considerable height, e.g., A. angustifolia and A. aurea. However,
nearly all intermediate heights are found in other species and the
apparent phylogenetic information of this character depends on
sample choice. As quantitative characters cannot be reliably coded,
many characters that make up the difference between the Alstroe-
meria species were left out. Some quantitative characters have, nev-
ertheless, been included when only a few species had intermediate
states. These include the characters size relation among inner te-
pals (char. 37) and shape of apex of the inner tepals (char. 38).
The multistate non-additive characters present a special prob-
lem when some of the states are considered nested. One example
comes from character 6 (leaves resupinate) and character 7 (resu-
pination of leaves complete or partial, twisting only some leaves).
Obviously, all species with resupinate leaves are bound to be
scored as having either partially or completely resupinate leaves
in character 7. If it was known which of the states is the plesiom-
orphic one, the two characters could be combined into a single
additive character. However, if this character were left as unor-
dered the homology statement between the two alternative states
of resupinate leaves would be lost. Each state could appear indi-
vidually without the cost of extra steps. When the ordering of a
multistate character is avoided, but one wishes to maintain the
homology statement between the alternative states, problems arise
during the coding of the character. One commonly used approach
is to code the unit (in this case whether the leaves are resupinate
or not) as a present/absent character, and the nested states in a
separate character where species lacking the unit are coded as
inapplicable (Pleijel 1995). This allows either state to be the nested
one, while the homology statement between the two states is main-
tained. The approach does, however, suffer from an unreasonable
optimization of the missing data (Maddison 1993). In the present
case, the missing data will be assigned as partly or completely
resupinate leaves according to the most parsimonious resolution.
This, however, does not make any sense. It is simply not possible
to have partly or completely resupinate leaves in character 7 with-
out having resupinate leaves in character 6. A possible solution to
the problem is to recode the two characters into a single character
using a step matrix as discussed by Maddison (1993). Then the
gain or loss of resupination could be coded as more costly than
the change between having either completely or partially resupi-
nate leaves. The use of step matrices for recoding complex char-
acters also has been applied in the context of DNA sequence anal-
yses (Wheeler 1999; Simmons and Ochoterena 2000). However,
apart from the problem of assigning justifiable relative weights of
the transformations within the step matrix (Maddison 1993), a sec-
ond drawback with this differential weighting within a character
49
is the relatively high or low weights that individual transforma-
tions receive compared to the weight of transformations in other
characters of the data set (Lee and Bryant 1999). In case of char-
acter conflict, the result would be biased towards the transfor-
mations of high weights.
In this study we apply the more commonly used method of
scoring the nested states as inapplicable in the species lacking the
feature. Although this may cause impossible state assignments at
the internal nodes the method was chosen because all analyses of
the matrices are done using both equal weights of the characters
as well as their implied weights. When analyzing under implied
weights, the program (PeeWee) used in this study does not allow
optimization of step matrices. Furthermore, Strong and Lipscomb
(1999), who reviewed various methods for coding inapplicable
data, found drawbacks in all of them. The authors argued that the
‘reductive coding’—used here—analyzed with NONA is the cod-
ing method which leads to tree topologies that best reflect our
observations.
DNA Extraction. Total genomic DNA was extracted from fresh
or silica-dried plant material using the DNeasyy Plant Mini Kit
(Qiagen) according to the instructions provided by the manufac-
turer. DNA from Alstroemeria caryophyllaea was donated by J. I.
Davis, Cornell University. DNA was available for all species except
Alstroemeria insignis, A. longistyla, A. sellowiana, and Luzuriaga po-
lyphylla. Voucher information is listed in Table 1.
rps16. The rps16 (ribosomal protein S16) intron was amplified
from total genomic DNA using the primers rpsF (59-GTGGTA-
GAAAGCAACGTGCGACTT-39) and rpsR2 (59-TCGGGATCGAA-
CATCAATTGCAAC-39; Oxelman et al. 1997). PCR amplification
was performed under standard conditions, and the double strand-
ed products were purified using the QIAquickTM PCR Purifica-
tion Kit (QIAGEN*) according to manufacturer9s instructions. Pu-
rified PCR products were sequenced using the ABI PRISMy Dye
Terminator Cycle Sequencing Ready Reaction Kit (Perkin-Elmer).
The PCR amplification primers were used also as sequencing
primers. In addition, two internal sequencing primers were used
in order to produce complete sequences: rps16-551F (59-CCGAAG-
TAATGTCTAAACCCAA-39) and rps16-680R (59-CTCATAACTCA-
AGTTGGGTAA-39; Asmussen et al., 2000). Cycle-sequencing prod-
ucts were analyzed on an Applied Biosystems 377 automated
DNA sequencer. The obtained sequences were checked and assem-
bled using the program SequencherTM ver. 3.0 (Gene Codes Corp,
Ann Arbor, MI).
rbcL. rbcL sequences were produced from 13 Alstroemeriaceae
species selected to represent the family. For amplification and se-
quencing of rbcL, the same method was followed as described
above. For PCR amplification the following primers were used:
RH-1s and Z1204RS; the same primers were used as sequencing
primers in addition to Z427s and Z674RS (all primers from G.
Zurawski, DNAX Research Institute, Palo Alto, California, but
slightly reduced in length).
The choice of including rbcL, which is mostly used for resolving
phylogenetic relationship at higher taxonomic levels (Soltis and
Soltis 1998), was based on preliminary results obtained from an-
alyzing morphology and rps16 data. In the preliminary results
Bomarea appeared as polyphyletic or paraphyletic when analyzing
rps16 on its own or combined with morphology. To settle the re-
lationships of the Bomarea species, rbcL was considered appropri-
ate. However, as rbcL is often found to be almost invariable at the
species level there was no justification for sequencing all species
included in this study. The choice of the 13 species used for am-
plifying rbcL was based on the preliminary cladograms. We se-
lected species that were likely to represent different clades of the
trees.
When only a subset of the species are represented in one of the
data sets a large number of missing data is bound to appear when
combining the data sets unless the combined matrix is pruned.
Missing data are known to have undesirable effects on phyloge-
netic analysis (Nixon and Davis 1991; Platnick et al. 1991; Mad-
dison 1993). The missing data discussed here differ from the in-
applicable data discussed under the coding of the morphological
characters. When optimising inapplicable data combinations of
50 SYSTEMATIC BOTANY
TABLE 1. Taxa used in the morphological and molecular study
listed alphabetically. Vouchers are indicated for the species from
which DNA sequences have been obtained. Those from plants cul-
tivated at the Copenhagen Botanical Garden are deposited at the
Herbarium of the Botanical Institute of Copenhagen (Denmark).
C-numbers correspond to purified-DNA samples deposited at the
Botanical Institute of Copenhagen. References are given to voucher
specimens and herbaria as designated by acronym in Holmgren
et al. (1990). Accession numbers and references to already pub-
lished sequences extracted from GenBank archives also are indi-
cated. 1 sequence from B. hirtella (H.B.K.) Herb. 2 sequence from D.
sessile D. Don. Distribution follows the taxon name, in parentheses;
abbreviations: AR: Argentina, BO: Bolivia, BR: Brazil, CH, Chile,
CHI: China, CO: Colombia, PA: Paraguay, PE: Perú, UR: Uruguay.
Alstroemeria andina Phil. subsp. venustula (Phil.) Ehr. Bayer
(AR-CH). Kiesling et al. 9216 (SI), C648, AR. Prov. San
Juan. Dto. Iglesias. Cordillera de la Ortiga. rps16
AY120370, rbcL AY120366.
A. angustifolia Herb. subsp. angustifolia (CH). Cult. at Copen-
hagen Botanical Garden (P1995-5003), C435, CH. Valparaı́-
so. Valparaı́so. rps16 AY120371, rbcL AY120358.
A. apertiflora Baker (AR-BR-PA). Ribas 687 (BHCB), C92, BR.
Mun. São José dos Pinhais. rps16 AY120372.
A. aurea Graham (AR-CH). Aagesen w/n (SI), C81, AR.
Prov. Neuquén. Dto. Minas. Lags. Epulauquen. rps16
AY120373. Cult. at Copenhagen Botanical Garden (P1995-
5043), C441, CH. Araucaria, rbcL AY120359.
A. caryophyllaea Jacq. (BR). From J. I. Davis, Cornell Universi-
ty, C4. rps16 AY120374.
A. graminea Phil. (CH). von Bohlen 1275 (SGO), C85, CH.
Antofagasta. Taltal. rps16 AY120375, rbcL AY120360.
A. hookeri Schultes subsp. cummingiana (Herb.) Ehr. Bayer
(CH). Cult. at Copenhagen Botanical Garden (P1995-5010),
C448, CH. Coquimbo. Pan Am. rps16 AY120376.
A. insignis Hérincq (BR).
A. isabellana Herb. (AR-BR-PA-UR). Rua (BAA 24543), C445,
AR. Prov. Misiones. Posadas. Pantano del Zaimán. rps16
AY120377, rbcL AY120361.
A. ligtu L. subsp. ligtu (CH).
A. longistyla Schenk (BR).
A. magnifica Herb. subsp. magnifica (CH). Cult. at Copenha-
gen Botanical Garden (P1995-5031), C449, CH. Valparaı́so.
rps16 AY120378.
A. pallida Graham (CH). Cult. at Copenhagen Botanical Gar-
den (P1995-5035), C444, CH. Santiago. Lagunillas. rps16
AY120380.
A. patagonica Phil. (AR-CH). Aagesen w/n (SI), C82, AR.
Prov. Neuquén. Dto. Catán-Lil. Entre Catán-Lil y La Ne-
gra. rps16 AY120381, rbcL AY120362.
A. pelegrina L. (CH). Cult. at Copenhagen Botanical Garden
(P1995-5037), C437, CH. Coquimbo. Cerca de Totoralillo.
rps16 AY120382, rbcL AY120363.
A. presliana Herb. (AR-CH). Aagesen w/n (SI), C80, AR.
Prov. Neuquén. Dto. Minas. Camino Lags. de Epulauquen.
rps16 AY120383.
A. pseudospathulata Ehr. Bayer (AR-CH). Xifreda and Sanso
2004 (SI), C89a, AR. Prov. Neuquén. Dto. Chos-Malal. En-
tre Chos-Malal y Andacollo. rps16 AY120384.
A. psittacina Lehm. (AR-BR-UR). Cult. at Facultad de Agro-
nomı́a, UBA (Argentina), C91a, AR. Cdad. de Buenos Ai-
res. rps16 AY120385, rbcL AY120364.
A. pygmaea Herb. (AR-BO-PE). Aagesen w/n (SI), C79b, AR.
Prov. Tucumán. Dto. Trancas. Huanlinchay. rps16
AY120386, rbcL AY120365.
A. revoluta Ruiz & Pav. (CH). Cult. at Copenhagen Botanical
Garden (P1995-5050), C434, CH. Del Maule. Pte. Longo-
milla. rps16 AY120379.
[Volume 28
TABLE 1. Continued.
A. sellowiana Seub. (BR).
A. versicolor Ruiz & Pav. (CH). Cult. at Copenhagen Botani-
cal Garden (P1995-5053), C447, CH. Gen. O’ Higgins. Ter-
mas de Cauquenes. rps16 AY120387.
A. werdermannii Ehr. Bayer (CH). Cult. at Copenhagen Botan-
ical Garden (P1995-5054), C446, CH. Atacama. Huasco.
rps16 AY120388.
Bomarea boliviensis Baker (AR-BO). Aagesen w/n (SI), C75b,
AR. Prov. Jujuy. Dto. Tumbaya. Lag. de Volcán. rps16
AY120389, rbcL AY120368.
B. edulis (Tuss.) Herb. (AR-BO-BR-CO-PA, Central America).
Aagesen w/n (SI), C76a, AR. Prov. Jujuy. Dto. Gral. M.
Belgrano. Lags. de Yala. rps16 AY120390. Rudall et al., un-
publ. rbcL Z772551.
B. macrocephala Pax (AR). Aagesen w/n (SI), C78b, AR. Prov.
Tucumán. Dto. Tafı́. Entre El Infiernillo y Tafı́ del Valle.
rps16 AY120391, rbcL AY120367.
Dioscorea dodecaneura Vell. (AR-BO-BR-PA-PE).
Disporum viridescens (Maxim.) Nakai (CHI). Cult. at Copen-
hagen Botanical Garden (S1939-1565), C422, rps16
AY120394. Shinwari et al. (1994), rbcL D173762.
Leontochir ovallei Phil. (CH). Cult. at Copenhagen Botanical
Garden, C676, CH. Atacama. Carrizal Bajo. rps16
AY120392, rbcL AY120369.
Luzuriaga marginata (J. Gaertn.) Benth. (AR-CH). Aagesen
w/n (SI), C74, AR. Prov. Rı́o Negro. Dto. Bariloche. Pto.
Blest. rps16 AY120393.
L. polyphylla (Hook.) J. F. MacBr. (CH).
L. radicans Ruiz & Pav. (AR-CH). Rudall et al., unpubl. rbcL
Z77300.
Uvularia grandiflora Sm. (U.S.A.). Cult. at Copenhagen Botan-
ical Garden (1080-4), C857, rps16 AY120395. Kawano and
Kato, unpubl, rbcL AB09950.
states that are not logical occur at the internal nodes. When miss-
ing data come from states that simply were not scored the opti-
mization at the internal nodes is legitimate (Maddison 1993). How-
ever, missing data can result in a high number of equally optimal
trees and a poorly resolved consensus tree, but they can not alter
the topology obtained from the observed data (Kitching et al.
1998). Therefore, although there is a high number of missing data
in some of the matrices included in this study this should not have
a misleading effect on the conclusions as long as these are drawn
from the strict consensus trees.
Sequence Alignment. The sequences were aligned by hand us-
ing the software BioEdit ver. 4.8.10 (Hall 1999). All sequences and
alignments are deposited in GenBank under accession numbers
AY120358–AY120395.
Data Matrices. Eight matrices were constructed. Three matri-
ces included the individual data sets (morphology, rps16 intron,
and rbcL). Furthermore, five combined matrices were constructed:
morphology plus rps16 or rbcL, and one including rbcL sequence
data and all species for which rps16 sequence data was available.
An additional rbcL plus rps16 matrix was constructed including
only species sequenced for both molecular data sets. Finally a
combined matrix was constructed including all three data sets con-
taining all species where rps16 intron sequence data were avail-
able. All matrices can be obtained from the first author. Number
of terminals and informative characters for each matrix are shown
in Table 2.
Data Analyses. Analyses using equal weights were performed
with NONA ver. 2 (Goloboff 1998a). In contrast to PAUP ver. 3.1.1
(Swofford 1993) and Hennig86 ver. 1.5 (Farris 1988), the default
setting in NONA (amb-) calculates trees retaining a branch only
when unambiguous support is available (collapsing a branch if its
minimum length is zero) while Hennig86 and PAUP retain a
2003] AAGESEN & SANSO: PHYLOGENY OF ALSTROEMERIACEAE
TABLE 2. Attributes of the eight data sets. MPT 5 most parsimonious trees; CI 5 consistency index; RI 5 retention index; * 5
significantly incongruent (P , 0.05) using the incongruence length difference test (Farris et al. 1994).
nr. informative nr.
characters terminals
Morphology 57 33
rps16 81 26
rcL 55 16
Morphology 1 rps16* 138 26
Morphology 1 rbcL 112 26
rps16 1 rbcL complete 136 26
rps16 1 rbc reduced* 132 16
Morphology 1 rps16 1 rbcL 193 26
branch if its length in at least one of the possible optimizations
differs from zero (Coddington and Scharff 1994). All analyses
were run using the default settings amb- and poly5 (polytomies
allowed). The following search strategy was applied to all data
sets: five hundred subsearches were performed, each constructing
a Wagner tree using a random addition sequence, swapping the
initial tree with TBR (tree bisection and reconnection) and retain-
ing a maximum of two trees in each replicate (hold/2; mult*500).
The resulting trees were swapped using TBR (max*). These two
commands were repeated until no new trees were found. To ob-
tain different results in each mult replication the random seed set-
ting was changed to zero (rseed 0), using the time as seed for the
random number generator that calculates the input order of the
taxa during the mult replicates (Goloboff 1998a).
Bremer support values (BS, Bremer 1994) were calculated find-
ing suboptimal trees up to five steps longer than the shortest
tree(s) holding a maximum of 2,000 trees. This can be done in
NONA using a loop finding suboptimal trees of a given length by
swapping previous trees held in memory and calculating BS val-
ues in each round. As starting trees for the first round of the loop
all most parsimonious trees held in memory were swapped until
the treebuffer was filled (syntax: hold 2000; loop 1 5; sub#; find*;
bs; k N; stop;), where N refers to the number of optimal trees held
in memory. BS values higher than 5 were calculated finding 5,000
suboptimal trees 10, 20, 30 etc. steps longer than the shortest
tree(s) (e.g., hold 5000; sub10; find*; bs;). BS values higher than 5
were checked searching 5,000 suboptimal trees one step shorter
than the support under test, while support values higher than 20
were checked searching 20,000 suboptimal trees one step shorter
than the support under test. Note that when using a limited tree-
buffer and a heuristic search method the BS values are estimated
and not calculated exactly. The support may be the one indicated
or less. This is because the limited number of trees used for cal-
culating the BS values at each step represents only a fraction of
the actual number of trees with a given number of extra steps.
High BS values are more likely to be overestimated than low BS
values are, simply because there may be many more trees with a
high number of extra steps. The highest BS values reported here
should not be taken as absolute values but rather as indicators for
highly supported groups. Jackknife frequency values (Farris et al.
1996) based on 10,000 Wagner trees were calculated using the
available instruction file for NONA, the jak.run file.
The incongruence length difference test, ILD (Farris et al. 1994)
was performed for each combination of the individual data sets
using the program Winclada. ver. 0.9.9 (Nixon 1999). The matrices
were pruned to include only common terminals and informative
characters before the test. The test was done on the basis of 1,000
iterations, each finding a Wagner tree submitted to TBR swapping
holding a maximum of 10 trees for each submatrix.
We did not apply the ILD test to decide whether to combine the
data sets or not, because we agree with Nixon and Carpenter
(1996) that the ILD is the best measure of incongruence and regard
the test as a means of determining whether the amount of incon-
gruence is large or small. In case of significant incongruence, we
use the test to explore the source of this incongruence. If the in-
51
Jackknife
nr. MPT/ frequency
% resolved nodes CI RI index
3/87.1 0.40 0.68 0.14
4/47.8 0.87 0.90 0.17
6/61.5 0.81 0.89 0.54
4/75 0.56 0.69 0.21
2/87 0.52 0.70 0.17
970/47.8 0.82 0.88 0.14
2/92.3 0.83 0.88 0.58
6/82.6 0.60 0.72 0.20
congruence is caused by a particular taxon (as is the case in this
study), we still consider that combining the unmodified matrices
is the best approach for placing this conflicting taxon.
Additional analyses using implied weights (Goloboff 1993) were
run in PeeWee ver. 3 (Goloboff 1998b) using the same search strat-
egies as when analysing with equal weights. In cases of character
conflicts, PeeWee downweights homoplastic characters in propor-
tion to their amount of extra steps (homoplasy). The strength with
which a homoplastic character is downweighted by PeeWee de-
pends on the shape of the weighting function. Six different con-
cavities are available in the program, concavity six being the mild-
est and concavity one the strongest weighting function (Goloboff
1998b). All six concavities were explored and will hereafter be
referred to as conc one to six.
There are strong arguments for and against the use of equal
weighting versus differential weighting of the characters (e.g., Go-
loboff 1993; Källersjö et al. 1999). We use implied weights as a
means for exploring the sensitivity of the matrices to different
weighting schemes of the characters. The robustness of a group to
changes in the analytic parameters has been used in DNA anal-
yses as a means for obtaining firm conclusions of monophyly
(Wheeler 1995). The implicit assumption is that the appropriate
analytical parameter is unknown. When a group appears under
all analytical parameters we may conclude that the group is mono-
phyletic and this conclusion will be independent of the analytical
parameters. In the present context a choice between the use of
equal weights and implied weights can be made on the basis of
theoretical assumptions, but when using implied weights it is not
clear by which strength a homoplastic character should be down-
weighted. We consider the use of equal weights and implied
weights, using different concavities, as a gradual change of the
analytical parameters. If a group is resistant to variation in the
weighting scheme few or no characters contradict the monophyly
and the group may also be found when new evidence is added.
RESULTS
Morphology. The morphological data set comprises
57 informative characters with 6.6% of the cells coded
as missing. When analyzing the data matrix with equal
weights two trees were obtained (212 steps, CI50.39,
RI50.68). The strict consensus tree is shown in Fig. 1A.
Both Bomarea and Alstroemeria are monophyletic and
sister groups to one another. The Brazilian Alstroemeria
species fall in two clades that branch off as sequential
sister-groups to the remaining Alstroemeria species,
with both informal groups of Baker (1888) being
monophyletic and the Chilean/Argentinean species
complex forming a clade.
When analyzed using implied weights nearly all
52 SYSTEMATIC BOTANY [Volume 28

FIG. 1. Strict consensus trees from separate and combined analyses. Numbers below branches refer to Bremer support
values, numbers above branches denote jackknife frequencies. Group a1 refers to the Brazilian Alstroemeria species with rigid
leaves. Group a2 refers to the Brazilian Alstroemeria species with herbaceous leaves. Group b refers to the Chilean/Argentinean
species complex. Group c refers to the central Chilean species group. A. Strict consensus tree from two optimal trees resulting
from morphology analyzed under equal weights. B. Strict consensus tree from four optimal trees resulting from rps16 analyzed
under equal weights. C. Strict consensus tree from six optimal trees resulting from rbcL analyzed under equal weights. D.
Strict consensus tree from four optimal trees resulting from morphology and rps16 analyzed under equal weights.
2003] AAGESEN & SANSO: PHYLOGENY OF ALSTROEMERIACEAE
conc values resulted in different trees that again dif-
fered from the topology found when using equal
weights. A strict consensus tree between the trees
found when using conc four to six only includes the
monophyletic Alstroemeriaceae wherein B. boliviensis
and B. edulis are sister groups and Alstroemeria is
monophyletic. Within Alstroemeria the group including
A. angustifolia, A. hookeri, A. revoluta and A. versicolor,
also found when using equal weights, is retained to-
gether with a clade consisting of A. patagonica and A.
pygmaea. This latter clade is lost when results from
conc three are added to the strict consensus tree. Trees
obtained when using conc two and one do not modify
this result any further.
rps16. Among Alstroemeriaceae the length of the
rps16 intron varied from 824 bp (A. hookeri) to 882 bp
(A. apertiflora). We were not able to amplify the com-
plete rps16 intron in A. pelegrina and Bomarea edulis,
consequently only partial sequences are included for
these in the matrix. In Alstroemeria pelegrina, A. presli-
ana, Disporum viridescens, and Luzuriaga marginata, the
amplified sequence fragments lack aprox. 20 bp near
the middle of the sequences which could not be am-
plified. Several gaps, varying from one bp to 25 bp in
length were introduced in the alignment. In cases of
ambiguous alignment, gaps were introduced to make
the positions non-informative. Two hypervariable are-
as were excluded from the analysis. Gaps were treated
as missing characters, while a total of eight informative
and non-problematic indels were coded as binary char-
acters and added to the end of the alignment. The final
matrix includes 923 characters, of which 81 are poten-
tially phylogenetically informative, and 5.32% of the
data matrix cells scored as missing data.
Four trees were found (164 steps, CI50.87, RI50.90)
by analyzing the rps16 matrix with equal weights. The
strict consensus tree is shown in Fig. 1B. Bomarea ma-
crocephala is embedded within Alstroemeria rendering
Alstroemeria paraphyletic and Bomarea polyphyletic.
The remaining Bomarea species are a sister group to
Leontochir. The resolution within Alstroemeria is mod-
erate. Bomarea macrocephala is sister species to A. psit-
tacina and the clade groups with A. patagonica. Two
minor clades appear, one including A. graminea and A.
werdermannii the other A. presliana and A. revoluta,
while six of the purely Chilean species are gathered in
a clade. Analyzing under implied weights all six conc
values yielded the same tree as found using equal
weights.
rbcL. In the final matrix nucleotide positions
57,648 to 58,803 of the rbcL gene were used (numbers
corresponding to the Nicotiana tabacum L. chloroplast
genome sequence [Shinozaki et al 1986; GenBank ac-
cession number Z00044]). The matrix includes 1156
characters of which 55 are potentially phylogenetically
53
informative with 0.11% of the data matrix cells scored
as missing data.
When analyzing the rbcL matrix under equal
weights, six trees were found (121 steps, CI50.81,
RI50.89). The strict consensus tree is shown in Fig. 1C.
Alstroemeriaceae is monophyletic and monophyly of
both Alstroemeria and Bomarea is supported. Leontochir
and Bomarea form a clade which is the sister group of
Alstroemeria. Within Alstroemeria little resolution is ob-
tained. The two species included from the Brazilian
species complex (A. isabellana and A. psittacina) are sis-
ter species within a clade that also contains A. aurea
and A. patagonica 1 A. pygmaea. When analyzed under
implied weights all six conc values yielded the same
tree as found using equal weights.
Morphology and rps16. Combining the morpholog-
ical data set with rps16 intron data resulted in a matrix
including 984 characters of which 138 are potentially
phylogenetically informative and with 5.21% of the
cells scored as missing. The two matrices are signifi-
cantly incongruent (P50.050) according to the ILD test
(Farris et al. 1994). The conflict is apparently caused
by Bomarea macrocephala. When this species was ex-
cluded from the test there was no significant conflict
between the two data sets (P50.343). When analyzing
the matrix under equal weights four trees are found
(steps5355, CI50.56, RI50.69). The strict consensus
tree is shown in Fig. 1D. As when analyzing the rps16
intron data alone, Bomarea is not monophyletic. Bomarea
macrocephala is the sister group to Alstroemeria. Leonto-
chir is sister group to the remaining two species of
Bomarea. Among the Alstroemeria species resolution is
moderate. The two Brazilian species with herbaceous
leaves form a clade which is sister group to the re-
mainder of the genus. In this second monophyletic
group A. pelegrina is the sister group to the rest of the
Alstroemeria species. Further resolution within Alstroe-
meria is restricted to a clade consisting of four Chilean
species, a clade containing A. andina and A. werder-
mannii, and a clade wherein A. pseudospathulata is the
sister group to a clade containing the Brazilian species
with rigid leaves and the clade A. patagonica and A.
pygmaea.
When applying weights conc three to six yielded
trees nearly identical to those obtained when analyz-
ing under equal weights but adding resolution to the
collapsed part of Alstroemeria (see Fig. 2A). As op-
posed to the results shown in Fig. 2A (strict consensus
tree from conc six), when applying stronger weights
against homoplasy (conc three to five) A. presliana and
A. revoluta are placed as sister groups. When using
even stronger weights (conc one and two) all Brazilian
species are placed basal within Alstroemeria (though
not forming a clade) and the positions of A. aurea and
A. andina are altered.
Morphology and rbcL. Combining the morpholog-
54 SYSTEMATIC BOTANY [Volume 28

FIG. 2. Strict consensus trees from separate and combined analyses. Numbers below branches refer to Bremer support
values, numbers above branches denote jackknife frequencies. Group a1 refers to the Brazilian Alstroemeria species with rigid
leaves. Group a2 refers to the Brazilian Alstroemeria species with herbaceous leaves. Group b refers to the Chilean/Argentinean
species complex. Group c refers to the central Chilean species group. A. Strict consensus tree from four optimal trees resulting
from morphology and rps16 analyzed with implied weights, conc six. B Strict consensus tree from two optimal trees resulting
from morphology and rbcL analyzed under equal weights. C. Strict consensus tree from 970 optimal trees resulting from the
complete rps16 and rbcL matrix analyzed under equal weights. D. Strict consensus tree from two optimal trees resulting from
the reduced rps16 and rbcL matrix analyzed under equal weights.
2003] AAGESEN & SANSO: PHYLOGENY OF ALSTROEMERIACEAE
ical data set with rbcL data gives a matrix of 1218 char-
acters of which 112 are potentially phylogenetically in-
formative and with 21.57% of the cells scored as miss-
ing. According to the ILD test (Farris et al. 1994) the
two matrices are not significantly incongruent (P 5
0.143).
When analyzed under equal weights two trees were
obtained (steps 5 308, CI 5 0.52, RI 5 0.70); the strict
consensus tree is shown in Fig. 2B. Both Alstroemeria
and Bomarea are monophyletic and Leontochir forms a
clade with Bomarea. Within Alstroemeria, the Brazilian
species with herbaceous leaves form a clade which is
sister group to the rest of the genus. The two remain-
ing Brazilian species are embedded in a clade contain-
ing the Chilean/Argentinean species.
When analyzed under implied weights several re-
arrangements within Alstroemeria occur. Using conc
six, five and three a single tree is obtained placing the
Brazilian species in a clade embedded among Chilean
species. Among other clades found in the equally
weighted analyses only the sister group relationship
between A. patagonica and A. pygmaea and between A.
andina and A. werdermannii reappeared in this tree,
while various rearrangements among the remaining
Alstroemeria species occurred. Using conc four and two
a second tree appeared in addition to the first. This
tree corresponds to the tree previously found but lacks
part of the resolution. Using conc one only this second
tree appeared (trees not shown).
rps16 and rbcL. The complete rbcL and rps16 ma-
trix including 26 taxa includes 2079 characters of which
136 are potentially phylogenetic informative, and
18.81% of the data matrix cells are scored as missing.
When analysing the complete rbcL and rps16 matrix
under equal weights 970 trees were obtained (290
steps, CI 5 0.82, RI 5 0.88). The strict consensus tree
is shown in Fig. 2C. Both Alstroemeria and Bomarea are
monophyletic. As in the rbcL tree Leontochir forms a
clade with Bomarea, and this clade is the sister group
to Alstroemeria. Within Alstroemeria, A. andina, A. gra-
minea, and A. werdermannii form a clade, which is the
sister group to the rest of the Alstroemeria species. Fur-
ther resolution within the genus is poor, restricted to
six Chilean species grouped in a clade, and the sister
group relationship between A. presliana and A. revoluta.
Analyzing under implied weights yielded the same
970 trees under all six conc values.
When including only species sequenced for both mo-
lecular data sets the reduced rbcL and rps16 matrix
includes 2079 characters of which 132 are potentially
phylogenetic informative, and 9.3% of the data matrix
cells are scored as missing. The two matrices are sig-
nificantly incongruent according to the ILD test (Farris
et al. 1994), with P 5 0.006. A considerable part of the
conflict is, however, caused by Bomarea macrocephala.
Excluding this species from the analysis removes suf-
55
ficient character conflict to obtain a nonsignificant re-
sult in the ILD test (P 5 0.410).
When analyzing the reduced rbcL and rps16 matrix
under equal weights two trees were found (282 steps,
CI50.83, RI50.88). The strict consensus tree is shown
in Fig. 2D. The tree includes the same clades as the
strict consensus tree found when analyzing the unre-
duced data set, but the resolution within Alstroemeria
is more detailed. This added resolution concurs with
the topology found when analyzing the rbcL sequence
data alone. When analyzing under implied weights all
six conc values yielded the same tree as found when
using equal weights.
Morphology and rps16 and rbcL. Combining the
three data sets yields a matrix including 2140 charac-
ters of which 193 are potential phylogenetically infor-
mative and with 14.75 % of the cells scored as missing.
When analyzed under equal weights six trees were ob-
tained (steps 5 485, CI 5 0.60, RI 5 0.72); the strict
consensus tree is shown in Fig. 3A. Both Alstroemeria
and Bomarea are monophyletic and Leontochir forms a
clade with Bomarea. Within Alstroemeria, the Brazilian
species form a clade, which is sister group to the rest
of the genus. Among the Chilean/Argentinean species
A. patagonica and A. pygmaea is the sister group to the
remaining species. All other minor species (A. pseudos-
pathulata, A. graminea, A. andina, and A. werdermannii)
are gathered in a clade which is the sister group of a
clade consisting of purely Chilean species.
When analyzed under implied weights, all six conc
functions concur on the same pattern shown in Fig.
3B. Using conc six and five a single tree was found,
while when using conc four, four trees were obtained,
one identical to the one found when using conc six and
five and three trees identical to the trees found when
using conc three. Conc two and one both yielded the
two same trees, different from those found when using
higher conc values but not changing the consensus.
The results differ from those obtained when using
equal weights mainly by placing A. andina, A. graminea
and A. werdermannii as sister group to the rest of the
genus while the Brazilian species are embedded with-
in the Argentinean/Chilean clade, still forming a
monophyletic group.
DISCUSSION
Attributes of the Data Set. Three individual and
five combined data matrices were analyzed. Table 2
shows different attributes of the data sets. Generally
the CI is a good indicator of stability: in data sets hav-
ing CI higher than 0.80 in the equally weighted anal-
ysis there were no changes when homoplastic charac-
ters were downweighted using implied weights (Go-
loboff 1993). The direct relationship between level of
homoplasy in the data matrix and the effect of using
implied weights is no surprise since the concavity
56 SYSTEMATIC BOTANY [Volume 28

FIG. 3. Strict consensus trees from separate and combined analyses. Numbers below branches refer to Bremer support
values, numbers above branches denote jackknife frequencies. Group a refers to the Brazilian Alstroemeria species. Group b
refers to the Chilean/Argentinean species complex. Group c refers to the central Chilean species group. A. Strict consensus
tree from six optimal trees found when combining all data sets analyzed under equal weights. B. Strict consensus tree from
seven optimal trees found when combining all data sets analyzed under implied weights using conc one to six.

function used to downweight a homoplastic character
is a modification of the CI (Goloboff 1993). In data sets
having CI between 0.55 and 0.60, the effect of applying
weights was moderate, restricted to rearrangement of
a few clades or polytomies found during the equally
weighted analyses were resolved. Generally high con-
cordance was found between the results obtained us-
ing distinct conc values. Only in the data sets with
lowest CI (0.40 and 0.52) were major changes found
when applying weights to the characters.
Of the individual data sets the rps16 data matrix is
least homoplastic, but no more than 48% of the nodes
were resolved. Furthermore, the low jackknife frequen-
cy index suggests low support for the resolution. The
rbcL data is only slightly more homoplastic but more
nodes are resolved (61%) and the higher jackknife fre-
quency index indicates better support for the obtained
resolution. The morphological data showed the highest
level of homoplasy. Although 87% of the nodes were
resolved, the jackknife index was low, indicating poor
support for the topology. Among the individual data
sets only the morphological one was sensitive to
changes in weighting schemes. Although well resolved
topologies appeared when analyzing this matrix, these
were not robust to changes in weighting schemes.
When implied weights were used nearly all conc val-
ues resulted in different resolutions that again differed
from the resolution obtained when analyzing the ma-
trix using equal weights. Furthermore, when the ma-
trix was combined with the DNA sequence data the
resulting matrices were sensitive to changes in weight-
ing schemes, while when analyzed on their own or
combined, the rps16 and the rbcL matrices were insen-
sitive to different weighting schemes. The high amount
of character conflict suggests that within Alstroemeri-
aceae various parallelisms and reversals have occurred
among the morphological characters. Although some
information is gained when adding morphology to the
combined matrix (compare Fig. 2C and Fig. 3B), the
most obvious influence of adding the morphological
characters is to destabilize the results due to character
conflict among the morphological characters (compare
Fig. 3 A and B).
Among the combined analyses the rps16 and rbcL
data included the smallest amount of homoplasy. In
the strict consensus tree derived from running the
complete matrix less than half of the potential nodes
are resolved and the jackknife index was low, indicat-
ing poor support for the topology. When the matrix
was reduced to include only taxa in common between
2003] AAGESEN & SANSO: PHYLOGENY OF ALSTROEMERIACEAE
the two data sets the result is a highly resolved tree
with a reasonably high jackknife index. This matrix
showed overall the best scores among all the combined
data sets. According to the ILD test (Farris et al. 1994)
the two matrices are incongruent. The conflict is prob-
ably due to the different rps16 sequence of Bomarea ma-
crocephala. Excluding this species from the analysis re-
moves sufficient character conflict to obtain a nonsig-
nificant result in the ILD test. However, despite the
character conflict both combined matrices are resistant
to changes in weighting schemes.
When morphology is combined with either the rps16
or the rbcL data the resulting matrices include mod-
erate to high levels of homoplasy. The resulting strict
consensus trees are well resolved but with low support
for the topology. There is no significant conflict be-
tween morphology and the rbcL data, but morphology
conflicts with the rps16 data. Again the conflict is ap-
parently caused by Bomarea macrocephala. When this
species was excluded from the test there was no sig-
nificant conflict between the two data sets. According
to both morphology and rbcL data, Bomarea macroce-
phala clearly belongs in Bomarea. Sampling within Bom-
area is very limited and we feel that the presence of an
Alstroemeria-like rps16 intron sequence in this species
should be confirmed in other individuals and/or other
Bomarea species (especially subgenus Wichuraea) before
a hypothesis concerning the divergent nature of the
sequence can be constructed. The sequence differs at
several positions from all other rps16 sequences used
in this study. Although a simple contamination of the
DNA sample from B. macrocephala can not be ruled out,
this would additionally require various modifications
of this sequence during the amplification.
When all data sets are combined the level of ho-
moplasy is moderate and a well resolved strict consen-
sus tree is found in the analysis using equal weights.
However, the support for the topology is low as judged
by the jackknife index. The final resolution receives
considerably higher support if morphology is left out,
and the topology only partly coincides with the par-
ticularly well supported one found in the reduced rbcL
and rps16 data sets. This condition changes if weights
are applied to the fully combined data set. Using im-
plied weights, the topology found in the reduced rbcL
and rps16 data set reappears though somewhat col-
lapsed.
We mainly base our discussion on the trees obtained
under implied weights combining all data sets (Fig.
3B). This topology closely resembles the particularly
well supported one found when combining rbcL and
rps16, and it is stable under all weighting schemes ex-
cept when using equal weights. However, the topology
found under equal weights (Fig. 3A), including a basal
dichotomy between Brazilian and Argentinean/Chi-
lean species is likewise considered.
57
Sister Group Relationship of the Alstroemeriaceae. The
Alstroemeriaceae are monophyletic in all analyses. When
combining all data sets monophyly is supported with a
Bremer support of 35, which is higher than the additive
value for the individual data sets, and a jackknife fre-
quency of 100.
The sister group relationships have not been clearly
resolved in previous analyses (Chase et al. 1995a, b;
Rudall et al. 2000). In these analyses Alstroemeriaceae
were invariably included in a clade also containing Lu-
zuriaga and a monophyletic group containing species
from Uvulariaceae p.p. and Colchicaceae. Only in the
combined analysis of Rudall et al. (2000) based on trnL-
F and rbcL sequences and morphology does this clade
receive solid support with a bootstrap value of 100. In
the analysis of Chase et al (1995a) based on rbcL, the
relationships among the three groups were not re-
solved. When rbcL was combined with morphology
(Chase et al. 1995b) Alstroemeriaceae and Uvulari-
aceae p.p. 1 Colchicaceae are sister groups but the
clade has a Bremer support of only 2. In Rudall et al.
(2000) the Alstroemeriaceae also appear as sister to the
Uvulariaceae p.p. 1 Colchicaceae clade but again the
support is low with a bootstrap value of only 54.
Our analysis only superficially explored the sister
group relationships of the Alstroemeriaceae. When an-
alyzing morphology on its own, and rooting the tree
with Dioscorea, a clade is formed with Luzuriaga, Dis-
porum, and Uvularia. We conducted an informal anal-
ysis adding rbcL data to the morphological matrix in-
cluding Dioscorea. In the results, Alstroemeriaceae is
the sister group to the Disporum-Uvularia clade as in
previous analyses but with a low Bremer support of 1
and no jackknife support.
Apparently there is only weak evidence for a sister
group relationship between Alstroemeriaceae and the
Uvulariaceae p.p.-Colchicaceae clade (Colchicaceae
sensu Rudall et al. [2000]). These results are, however,
not convincing and Rudall et al. (2000) also considered
a sister group relationship between Luzuriaga and Al-
stroemeriaceae as likely.
Circumscription of the Genera of the Alstroemeriaceae.
Both Bomarea and Alstroemeria are monophyletic in all
analysis, except when the rps16 data are analyzed
alone or combined with morphology. The rps16 data
places Bomarea macrocephala within Alstroemeria due to
two base substitutions shared with Alstroemeria and a
six bp indel shared with A. psittacina. Forcing Alstroe-
meria to be monophyletic in the rps16 data set adds one
step and places B. macrocephala as sister group to Al-
stroemeria, which is the position of B. macrocephala in
the combined morphology and rps16 analyses. Forcing
Bomarea to be monophyletic in the rps16 data set adds
four steps to the tree.
Alstroemeria. Only when the rps16 data are ana-
lyzed alone is monophyly of Alstroemeria not main-
58 SYSTEMATIC BOTANY
tained. Both morphology and rbcL data support mono-
phyly of the genus with a Bremer support of 4 and 5,
respectively, and a jackknife frequency of 81 and 90.
The moderately high Bremer support and the high
jackknife frequency suggest low conflict concerning
the monophyly of Alstroemeria in these two data sets.
When the rps16 data set is combined with morphology
or rbcL a monophyletic Alstroemeria has a Bremer sup-
port that is slightly lower to somewhat higher than the
support found in the separate data sets. Although the
rps16 data places Bomarea macrocephala within Alstroe-
meria, this signal is not sufficiently strong to lower the
support for monophyly of the latter when combined
with other data sets. If all data sets are combined, the
Alstroemeria clade has a Bremer support of 11 and a
jackknife frequency of 74.
When all data sets are combined the Alstroemeria
clade is defined by 11 morphological characters, five
of which are free of homoplasy (two characters related
to the veins of the outer tepals, explosive dehiscence
of the fruit, a capsule which gradually tapers towards
the apex, and globose seeds). These five characters also
characterize Alstroemeria in the equally weighted anal-
ysis. Striate pollen exine and the shape of the storage
roots are potential synapomorphies, but these charac-
ters are ambiguously optimized at the family level
(e.g., there are three different hypotheses of pollen
sculpture evolution that are equally parsimonious: stri-
ate pollen plesiomorphic within the Alstroemeriaceae,
altered to reticulate pollen in the Bomarea-Leontochir
clade, reticulate pollen plesiomorphic within the family
and striate pollen a synapormorphy for Alstroemeria, or
a third, different, pollen sculpturing could be plesiom-
orphic within the family and both striate and reticulate
pollen sculpturing are synapormorphies for Alstroe-
meria and the Bomarea-Leontochir clade, respectively).
Chromosome number does not appear as a synapor-
morphy for Alstroemeria, as the same number occurs in
Disporum (other chromosome numbers are found in
other Disporum species and homology between the
chromosome numbers found in Alstroemeria and in
Disporum is dubious). Furthermore, monophyly of Al-
stroemeria is supported by base substitutions within
both molecular data sets.
Taltalia and Schickendantzia. Neither Alstroemeria
graminea nor A. pygmaea ever occur in a position that
make their segregation from Alstroemeria possible. El-
evating either species to genus level as proposed by
Bayer (1998a) and Pax (1890) and recently again by
Bayer (1998b) makes Alstroemeria paraphyletic. To re-
tain Alstroemeria, both species must remain within the
genus, as previously proposed for A. pygmaea by Sanso
and Xifreda (1999).
Bomarea. Monophyly of Bomarea is not supported
by the rps16 data and the genus does not appear as
monophyletic when these data are combined with
[Volume 28
morphology. Morphology and rbcL data support
monophyly of the genus, but Bremer support values
and jackknife frequencies are low (1/64 and 2/65 re-
spectively). When all data sets are combined Bomarea
has a Bremer support of 3, but the clade does not ap-
pear when the data set is jackknifed.
In both the equally weighted analysis and in the
analysis using implied weights monophyly of Bomarea
is supported by one homoplastic morphological char-
acter related to indumentum of the leaves, apart from
base substitutions in rbcL. Within Bomarea, B. macroce-
phala, belonging to the subgenus Wichuraea, is placed
as sister group of B. edulis and B. boliviensis, which be-
long to subgenera Bomarea and Sphaerine, respectively.
The B. edulis-B. boliviensis clade is defined by two ho-
moplastic characters (papillae at the inner tepals; chars.
40 and 41) and a single substitution in rps16.
Bomarea is sister group to Leontochir in all analyses,
except when morphology is analyzed alone. Alstroe-
meria and Bomarea share five morphological characters
not found in Leontochir (zygomorphic flowers, multi-
colored outer and inner tepals, presence of a dotted-
striated color pattern on the tepals, and filaments as
long as upper inner tepals). The zygomorphic flowers
and the tepal pattern show no reversals within the
clade. When DNA data are added Leontochir is sister
group to Bomarea (or Bomarea p.p. when analyzing
rps16) and all the above mentioned characters are am-
biguously optimized at the family level, e.g., either
having evolved in parallel in Alstroemeria and Bomarea
or showing reversals in Leontochir.
Leontochir. The monotypic Leontochir is sister
group to Bomarea. The genus Leontochir is characterised
by its prostrate habit, the presence of Fritillaria-like
nectaries and an unilocular ovary (Wilkin 1997).
Bomarea-Leontochir. A morphologically based di-
agnosis of Bomarea is somewhat difficult. All characters
mentioned by Sanso and Xifreda (1995) as separating
Alstroemeria and Bomarea at the genus level (gradual
dehiscence of fruit, presence of a sarcotesta, reticulate
pollen sculpturing, and chromosome number) are
shared with Leontochir, and hence, diagnostic for the
clade Leontochir-Bomarea. The only morphological char-
acter that defines the clade in our study is the presence
of hyaline hairs at the lower surface of the leaves (char.
12). However, considering that our analysis only in-
cluded three of the ;100 taxa in Bomarea, a diagnosis
based only on presence of hairs at the lower surface of
the leaves seems premature. Monophyly of Bomarea is,
however, also supported by rbcL data. Retaining both
Leontochir and Bomarea as genera could be supported
by our results, but merging Leontochir within Bomarea
makes a morphological diagnosis possible.
The Bomarea-Leontochir clade is supported by rbcL
data and all combined analyses including this data set.
When all data sets are combined, the clade has mod-
2003] AAGESEN & SANSO: PHYLOGENY OF ALSTROEMERIACEAE
erate Bremer support (4) and no jackknife support. The
clade is supported by nine character state changes. All
changes are non-homoplastic, including three morpho-
logical characters (apex of inner tepals truncate, pres-
ence of sarcotesta, and chromosome number). The
gradual dehiscence of the fruit is shared with the out-
group species while pollen sculpturing and the shape
of the storage roots are ambiguously optimized at the
family level (see discussion above).
Alstroemeria versus Bomarea-Leontochir. Hun-
ziker (1973) disputed the validity of maintaining Al-
stroemeria and Bomarea as two separate genera, and
Bayer (1998) considered the genera to be morphologi-
cally very similar, but clearly distinguished by chro-
mosome numbers. We consider the monophyly of Al-
stroemeria to be particularly well supported, and the
genus is easily distinguishable from the Bomarea-Leon-
tochir clade by fruit, seed, root, and pollen characters
as pointed out by Sanso and Xifreda (2001). Our re-
sults do not support the notion that either of these
genera is nested within the other, as discussed by var-
ious authors (Whyte 1929; Hunziker and Xifreda 1990;
Sanso and Hunziker 1998; Bayer 1988; Kosenko 1994).
A discussion concerning derived characters within the
family must be postponed until the sister group rela-
tionship of the family has been resolved and the ple-
siomorphic character states of the family can be estab-
lished with more precision.
Relationships within Alstroemeria. The infrage-
neric divisions proposed by Ravenna (2000) do not
agree with the provisions of the ICBN (Greuter et al.
2000). It is, however, beyond the scope of the present
paper to make the necessary taxonomic and nomen-
clature changes. Bayer (1987) considered a subdivision
of Alstroemeria premature until the Brazilian species
have been reviewed.
The overall resolution within Alstroemeria is high
when the three data sets are combined and the matrix
analyzed under equal weights. Support for the branch-
es is low, however, with Bremer supports of 1, rarely
2 and only few clades reappearing when the matrix is
jackknifed. Five groups do, however, remain stable
when applying weights to the characters (Fig. 3 A and
B). Hence, although the support of the branches is low,
character conflict concerning monophyly of these
groups is limited.
ALSTROEMERIA APERTIFLORA , A. ISABELLANA, A. CARY-
OPHYLLAEA, AND A. PSITTACINA . This clade corre-
sponds to the Brazilian species complex. Based on
AFLP data, Han et al. (2000) analyzed the genetic dis-
tances by cluster analysis among 22 Alstroemeria spe-
cies available in breeding programs. In the analysis
both the Brazilian and Chilean species groups were
found, with the Brazilian group being best supported
based on bootstrap values. The authors were aware
that their results might be biased by the limited num-
59
ber of Brazilian species included (only three species),
one of which, A. inodora, was synonymized with A.
psittacina by Sanso (1996).
Monophyly of the Brazilian species is supported in
all analyses where the three data sets are combined.
The sampling of the Brazilian species is limited, how-
ever, and more thorough sampling is needed to con-
firm monophyly of this species group.
When analyzed under equal weights, the Brazilian
species are placed near the root of the genus (Fig. 3A).
The flowers of some Brazilian Alstroemeria species su-
perficially resemble Bomarea flowers more than the
flowers of some Chilean species do (i.e. smaller and
more tubular flowers with the same color pattern on
all inner tepals) and there is some support for the Bra-
zilian species being placed near the root of the genus
when all data sets are combined as a result of the in-
fluence of the morphological characters (Fig. 1A). How-
ever, when implied weights are used the Brazilian spe-
cies are placed among the Argentinean/Chilean spe-
cies (Fig. 3B), which is the position favored by rbcL (see
Fig. 1C). At present, monophyly of the Brazilian spe-
cies seems supported, but morphology and DNA se-
quence data disagree about the position of the clade.
The monophyly of the informal subgroups needs con-
firmation.
ALSTROEMERIA ANDINA , A. GRAMINEA AND A.
WERDERMANNII. When implied weights are used Al-
stroemeria andina, A. graminea, and A. werdermannii form
a clade that is sister group to the remaining species
(Fig. 3B).
Within the clade Alstroemeria graminea and A. wer-
dermannii are sister species and defined by a substi-
tution and a 25 bp indel in the rps16 intron; both
changes are non-homoplastic. Both species are found
on the coast of northern Chile between 258 S and 298
S. Alstroemeria andina grows slightly south of this area
but generally at higher elevations than 3,000 m (Sanso
1996). Other species occupying the same area were not
included in the analysis.
When equal weights are used the group is placed
deeper within Alstroemeria together with A. pseudospa-
thulata. The group appears as sister group to a purely
Chilean clade (see below) based on substitutions in
rbcL.
ALSTROEMERIA ANGUSTIFOLIA , A. HOOKERI, A. MAGNIFI-
CA, A. PALLIDA, A. PELEGRINA, AND A. VERSICOLOR. This
clade contains central Chilean species (298 S–378 S
[Bayer 1987]) and is defined by two substitutions in
rps16. A close relationship between A. angustifolia, A.
hookeri, and A. pallida was predicted by Bayer (1987).
Although the clade is only weakly supported, it per-
sists under all weighting schemes.
ALSTROEMERIA PATAGONICA AND A. PYGMAEA. The
two species included in this clade are small-statured
(no more than 6–15 cm high). Alstroemeria patagonica is
60 SYSTEMATIC BOTANY
a trans-Andean species from Argentina and Chile oc-
cupying dry areas from sea level to 1,000 m. In con-
trast, A. pygmaea is a high-Andean species (3,500–4,400
m) from Argentina, Bolivia, and Peru (Sanso 1996).
The clade is resistant to different weighting schemes,
and is reasonably well supported when equal weights
are used, with a Bremer support of 2, and a jackknife
frequency of 78.
The position of the clade is not unequivocally re-
solved in the present analysis. When analyzed under
equal weights Alstroemeria patagonica and A. pygmaea is
the sister group to the remaining Argentinean/Chi-
lean Alstroemeria species. When analyzed under im-
plied weights the position of Alstroemeria patagonica
and A. pygmaea is not resolved. The clade is either
grouped with A. aurea and A. pseudospathulata or with
the Brazilian Alstroemeria species.
ALSTROEMERIA AUREA , A. PRESLIANA, A. PSEUDOSPATHU-
LATA , AND A. REVOLUTA . No clear placement of these
four species could be established as their position var-
ied with each weighting scheme. With the exception of
Alstroemeria aurea, rbcL data is lacking for the species.
Han et al. (2000) considered it likely that the Chilean
and Brazilian Alstroemeria species originated as two in-
dependent groups from the widely spread A. aurea. In
our analysis there is no support for A. aurea being the
sister group of the remaining species. However, based
on rbcL data (Fig. 1C), A. aurea, among other trans-
Andean species, may have shared a common ancestor
with the Brazilian species.
ACKNOWLEDGEMENTS. The authors wish to thank Conny As-
mussen who provided us with the rps16 primers and assistance
in the laboratory. We are grateful to Gitte Petersen and Louise
Pedersen who helped us at various point of this work including
sequencing of the rbcL DNA data. We also wish to thank Charlotte
Hansen for technical assistance. We also thank Cecilia Xifreda who
supplied us with the Dioscorea data and for helpful comments, and
Marta Camargo for receiving LAA at Saõ Paulo University, and
for allowing us to access to the Brazilian Alstroemeria species col-
lection. Ole Seberg, Gitte Petersen, Juan Héctor Hunziker, Jim
Smith, Dennis Stevenson, and three anonymous reviewers sup-
ported valuable comments and help improving this manuscript.
We thank the Argentina National Parks for permission to conduct
fieldwork and the curators of the Herbaria for facilities. The Uni-
versity of Copenhagen supported Lone Aagesen as well as all lab-
oratory work (Ph.D. grant). The research was partially supported
by a grant from the Consejo Nacional de Investigaciones Cientı́-
ficas y Técnicas of Argentina (CONICET): project PEI No. 0339/
98.
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APPENDIX 1
Characters used in the morphological analyses.
0. Persistence of aerial parts: persistent (0); dying after
each growth season (1). All species included in this
study are rhizomatous and perennial, the only ex-
ception being Alstroemeria graminea, which is annual
(Bayer 1998a, b). In all Alstroemeriaceae, Disporum
viridescens (pers. observation), and Uvularia (Wilbur
1963) the aerial part withers after each growth sea-
son. Luzuriaga species are perennial with somewhat
lignified stems (Rodrı́guez and Marticorena 1987)
and the aerial parts are persistent.
[Volume 28
1. Habit: erect to reclining (0); prostrate (1); twining (2);
climbing (3).
2. Storage roots: absent (0); present (1). In Alstroemeria
and Bomarea some of the roots are thickened and
modified to store nutrients and water (Uphof 1952;
Sanso and Xifreda 2001). The only known excep-
tions are Alstroemeria graminea, which is an annual
herb without storage roots (Bayer 1998a), and A.
hygrophila (Meerow et al. 1999). Wilbur (1963) men-
tions that three species of Uvularia possess thick-
ened roots while fibrous roots are found in the re-
maining two species. No morphological or ana-
tomical studies on Uvularia roots have been carried
out. We considered the storage roots present in Al-
stroemeriaceae and the thickened roots in Uvularia
as non-homologous. In Luzuriaga and Disporum
thickened storage roots have not been reported. In
Dioscorea a storage organ is present. However, this
is not homologous with the roots present in Al-
stroemeriaceae. According to Huber (1998) the
storage organ of Dioscorea has secondary growth
and is derived from the hypocotyl.
3. Shape of storage roots: fusiform (0); ovoid to roundish
(1).
4. Phyllotaxy: spiral (0); distichous (1).
5. Leaves anatomy inverted: no (0); yes (1). In all Al-
stroemeriaceae species the anatomy of the leaves
is inverted, with the stomata (and indumentum
when present) located at the morphological adaxial
surface. This pattern is recognizable even in spe-
cies having stomata on both surfaces, as the inver-
sion is still identifiable by the protuberance of the
main nerves or the hooded apex of the leaf, both
characters noted at the adaxial surface. The invert-
ed anatomy is present regardless of wether or not
the leaves are resupinate.
6. Leaves resupinate: no (0); yes (1). In nearly all spe-
cies belonging to Alstroemeriaceae the leaves are
resupinate making the adaxial surface (occupied
by stomata, indumentum, etc., see char. 5) as the
functional lower surface. A few Alstroemeriaceae
species included in this study lack resupination.
7. Resupination of leaves: complete (0); partial, twisting
only some leaves (1). Although nearly all Alstroe-
meriaceae species have more or less resupinate
leaves, variation in the degree of resupination oc-
curs among the species. In species with completely
resupinate leaves the twisting of the leaves starts
at the base, turning the leaf 1808 within a short
distance, with all leaves repeating this pattern. In
other species the resupination may be more or less
complete, with the twisting occurring along the en-
tire leaf, this being twisted 1808 or less while other
leaves lack any sign of resupination. The Luzuriaga
species have been scored as having completely re-
supinate leaves as all leaves are placed with the
2003] AAGESEN & SANSO: PHYLOGENY OF ALSTROEMERIACEAE
adaxial surface towards the ground although the
twisting does not exceed 908.
8. Leaf texture: herbaceous (0); crassus (1); coriaceous
(2). Most Alstroemeria species have thin leaves, but
species with somewhat thicker leaves occur mostly
among the Chilean species.
9. Proximal part of leaves: petiole-like (0); not constrict-
ed (1).
10. Form of leaves: lanceolate to ovate (0); linear (1).
11. Surface of the functional abaxial side of the leaf:
glabrous (0); with papillae (1); with hyaline hairs (2).
All Alstroemeria species included in this study have
glabrous leaves (papillae are found in leaves of A.
andina subsp. venustula and A. insignis), while the
included Bomarea species have hyaline hairs on the
functional abaxial side. Some species may be poly-
morphic for the character (Sanso and Xifreda
1995). Hyaline hairs were found in some Uvularia
species, but not in Disporum or Luzuriaga. In the
latter genus, papillate cells are present, either on
the entire surface or in the intercostal areas only
(Arroyo and Leuenberger 1988). The character has
been coded as additive in the analyses as we con-
sider the hyaline hairs to be prolonged papillae,
hence state 1 and 2 are homologous and should be
coded as internested states (Lipscomb 1992).
12. Papillae at the apex of the leaves: absent (0); present
(1). In the Brazilian Alstroemeria species with stiff
rigid leaves, inconspicuous papillae are found at
the leaf apex. The character has also been coded
as present in species having papillae covering the
whole surface (see char. 11).
13. Epicuticular waxes on the functional abaxial sur-
face of the leaves: Convallaria-type (0); wax stalks (1);
absent (2); non-oriented crystalloids (3). All studied
Alstroemeria and Bomarea species as well as Leon-
tochir have parallel wax scales (Convallaria-type) on
the functional abaxial leaf surface (Sanso and Xif-
reda 1998). In Uvularia epicuticular waxes of the
Convallaria-type are found on both surfaces while
in Luzuriaga the functional abaxial surface is cov-
ered with wax stalks (Frölich and Barthlott 1988).
In Dioscorea non-oriented crystalloids are found
(Frölich and Barthlott 1988). Information concern-
ing waxes is lacking in Disporum.
14. Stomata present on both the adaxial and abaxial
surface of the leaves: absent from abaxial surface (0);
present on both surfaces (1).
15. Leaf margin: entire (0); minutely serrate to crenate (1);
with papillae (2). In Alstroemeriaceae the margins
of the leaves are composed of elongate, hyaline
cells. In some species these cells are straight and
the leaf margin entire, while in others small pa-
pillae or teeth are found, making the margin mi-
nutely crenate to serrate. In a few species these are
prolonged into papillae. The character has been
63
coded as additive in the analyses (see discussion
for char. 11).
16. Sclerenchyma in bundle sheaths: no (0); yes (1).
Sclerenchyma is found in the bundle sheaths in
Leontochir and all Bomarea species included in this
study, as well as in the Brazilian Alstroemeria spe-
cies corresponding to Baker’s (1888) informal
group with rigid leaves. The condition is also
found in Luzuriaga (Arroyo and Leuenberger 1988).
17. Sclerenchyma in bundle sheaths extended as girders
from the vascular bundles towards both leaf surfac-
es: no (0), yes (1). The presence of conspicuous areas
of sclerenchyma extending as girders from the vas-
cular bundles towards both leaf surfaces is charac-
teristic of the Brazilian species with rigid, strongly
nerved leaves (Sanso and Xifreda 1998).
18. Leaves of flowering shoots: abundant (resembling
those of the vegetative shoot) (0); sparse (reduced in size)
(1). Fertile shoots having only few, reduced leaves
are found mainly in one of the Brazilian species
groups. In these the vegetative shoots are leafy
with leaves bigger than those found at the fertile
shoots. In Leontochir and Alstroemeria graminea veg-
etative shoots are not found.
19. Synflorescences: including 2 or more paracladia (0); in-
cluding only a single paracladium (1). The synfloresc-
ences in species belonging to Alstroemeriaceae are
composed of one to several paracladia, each con-
sisting of a cyme with one or more flowers. Most
species produce 2 or more paracladia, but in a few
species, mainly belonging to the Chilean species
group, only a single paracladium is formed.
20. Pherophylls of axillary flowers of first order: simi-
lar to upper leaves (0); smaller than upper leaves, leaf-
like or reduced to bracts (1); bigger than upper leaves
(2). In Alstroemeria the pherophylls (subtending
leaves) of the axillary flowers of first order are sim-
ilar to the upper leaves of the flowering shoot in
most species. In some species the pherophylls are
strongly reduced, often into bracts. In Bomarea, the
pherophylls are either bract-like or leaf-like. In the
latter case, they are either smaller or larger than
the upper leaves but never of the same size. In
Uvularia grandiflora and Disporum viridescens no dif-
ference in size between pherophylls and other
leaves was seen. In Luzuriaga radicans the synflo-
rescences are axillary brachyblasts, each composed
of several lateral flowers, while the apical meristem
seems to abort. Pherophylls that subtend the
brachyblast have the same size and shape as the
assimilating leaves. In L. marginata and L. polyphylla
uniflorous synflorescences, preceded by bracts, are
found at the apex of a seasonal growth unit and
apparently ending the growth of this shoot. It has
not been possible to determine whether the flower
is terminal or axillary, but in both cases the cor-
64 SYSTEMATIC BOTANY
responding pherophyll is similar to the assimilat-
ing leaves.
21. Position of anthetic flowers: vertical (0); horizontal,
angle between peduncle and ovary ; 908 (1); pendent 32.
(2). In nearly all Alstroemeriaceae species the flow-
ers are horizontally oriented during anthesis with 33.
the angle between peduncle and ovary being ;908.
During development of the fruit this angle be-
comes more open and the mature fruit is eventu-
ally vertically oriented. The only exceptions to this
pattern are seen in A. pygmaea and Leontochir,
where the anthetic flowers are vertically oriented.
22. Perianth position: hypogynous (0); epigynous (1). 34.
23. Position of inner tepals in anthetic flowers: actino-
morphic (0); zygomorphic, angle between upper tepals 35.
and median tepal strongly exceeding 1208 (1). In all
members of the Alstroemeriaceae the flowers are
more or less zygomorphic due to the position of the
inner tepals. In only two of the included species are
actinomorphic flowers found to be mixed with flow-
ers showing some degree of zygomorphy.
24. Background color of outer tepals: homogeneous (0);
composed of various colors (1).
25. Color of outer tepals: white (0); yellow to orange (1);
rose to red (2); greenish (3).
26. Striated to dotted color pattern on the tepal: absent
(0); present (1).
27. Color pattern of outer tepals: intense, always present 36.
(0); weak, often lacking (1). In many Alstroemeria spe-
cies color pattern is absent from the outer tepals 37.
though specimens having a few spots or lines on
the outer tepals may always be found. In other spe-
cies, e.g. A. psittacina, the outer tepals are always
provided with spots or lines forming a pattern as
intense as the one seen on the inner tepals. 38.
28. Color pattern of inner tepals: the inner median tepal
with the same pattern as the lateral inner tepals (0); the 39.
inner median tepal with weak pattern or often lacking
(1). The pattern of the inner tepals can be similar
on all inner tepals or the lower tepal may be pro-
vided with only a weak pattern, or in some spec-
imens completely lacking.
29. Margin of outer tepals: entire (0); finely serrate to 40.
crenate (1). 41.
30. Outer tepals with wings: absent (0); present (1). In
most Alstroemeria species the outer tepals are com-
posed of a central part with parallel nerves anas-
tomosing at the apex, and a more or less wing-like
extension having nerves arising at the base or cen-
ter of the tepal and terminating at the margin.
Among the outgroup species the nervation of the
outer tepals is parallel to curved.
31. Apex of outer tepals: glabrous (0); with short papillae
(1). The apex of the tepals may be covered with
papillae, often shorter and denser than the ones
found at the basal part of the inner tepals. In Lu- 42.
[Volume 28
zuriaga marginata and L. radicans the apices of the
tepals are papillate though very inconspicuously in
the latter species.
Central vein of outer tepals: not prominent (0); prom-
inent (1).
First pair of lateral veins of outer tepals: not prom-
inent (0); prominent (1). In most Alstroemeria species
the first lateral veins of the outer tepals are prom-
inent from apex to base. At the level of insertion
of the tepals the lateral and central nerve can be
seen at the ovary as three ribs, which soon become
confluent.
Second pair of lateral veins of outer tepals: promi-
nent (0); not prominent (1).
Anastomosing point of main veins of outer tepals:
forming a flat apex of the tepal (0); forming a hooded
apex of the tepal (1); overtopped by the tepal (2). In
Alstroemeria (except A. versicolor) the apices of the
outer tepals are hood-formed at the point where
the central nerves anastomose (Sanso 1996: fig. 10
D). In Bomarea and Leontochir the same pattern is
found, but the anastomosing point is overtopped
by lateral tissue of the tepals (Neuendorf 1977: fig.
5 C right). The character is coded additive because
in species with overtopped main nerves the anas-
tomosing point of these is hooded, hence state 2 is
nested within state 1.
Backgroun color of inner tepals: homogeneous (0);
composed of various colors (1).
Inner median tepal differing in size from lateral
ones: no (0); yes (1). In nearly all Alstroemeria spe-
cies the inner median tepal is slightly (to conspic-
uously) shorter and often a little wider than the
two lateral ones.
Apex of inner tepals: attenuate (0); obtuse (1); trun-
cate (2).
Inner tepals base: smooth (0); papillate (1). The basal
part of the inner tepals in most species of Alstroe-
meriaceae are covered with long dense papillae.
The papillae are most pronounced at the margins
of the tepals but in some species the adaxial sur-
face also is covered.
Inner tepals apex: smooth (0); papillate (1).
Intercostal area between the central and the first lat-
eral nerves: without channels (0); with channels (1).
Due to the prominence of the central and first lateral
nerves the intercostal area is channelled (see Bayer
1987; abb. 64- lateral view) in all Alstroemeriaceae
species included in this study, except A. andina. In
some species the structure is somewhat less clear as
in A. cf. sellowiana. As the character can only be
scored on live material, the character has been
scored as unknown when only herbarium material
was available. In Luzuriaga radicans the same pattern
is found though it is not as conspicuous.
Base of the inner tepals: without nectar groove (0); with
2003] AAGESEN & SANSO: PHYLOGENY OF ALSTROEMERIACEAE
nectar groove (1). In Leontochir the base of all inner
tepals is closed to form a pocket that contains nectar
(Wilkin 1997). In all other Alstroemeriaceae species,
the proximal part of the two inner upper tepals is
channelled and contain nectar at anthesis. In Luzur-
iaga nectar is secreted at the base of all tepals before
anthesis, with the area of nectar secretion not being
histologically recognizable (Daumann 1970). Nectar
grooves are also found in Uvularia and Disporum
(Dahlgren et al. 1985; Wilbur 1963).
43. Length of style and filaments: nearly as long as the
lateral inner tepals (0); clearly shorter than the tepals (1).
44. Form of filament: straight (0); curved (1). In Alstroe-
meriaceae the filaments are more or less curved,
occupying the lower part of the flower. Straight fil-
aments are found only in Alstroemeria pygmaea.
45. Filaments forming a loop after dehiscence of an-
thers: no (0); yes (1). In all Alstroemeriaceae species
the anthers apparently move downward and out-
ward after dehiscence (see Hunziker [1991] and
Aizen and Basilio [1995] for a description of sta-
men and style movement in Alstroemeria psittacina
and A. aurea, respectively). In addition, some Al-
stroemeria species have very long filaments that
form a loop after dehiscence of the anthers.
46. Proximal part of filaments: glabrous (0); papillate (1).
47. Color of anthers: yellow to orange (0); purple (1); green
(2), blue (3); brown (4). The color of the anthers varies
within the Alstroemeriaceae. The color scored is of
the mature anther just before dehiscence.
48. Anther attachment: dorsifixed (0); basifixed (1); pseu-
do-basifixed (2). The anthers of Alstroemeriaceae are
pseudo-basifixed, with the pollen sacs elongated
beyond the point of insertion of the filaments.
Among the outgroups the anthers are either dor-
sifixed or basifixed.
49. Anther dehiscence: introrse (0); extrorse (1); latrorse
(2). According to Huber (1969), Dahlgren et al.
(1985), and Sanso (1996), anther dehiscence in Al-
stroemeria is introrse. Sanso & Xifreda (1995) noted
the same condition in Bomarea. Goldblatt (1995)
disputed the interpretation of the anthers as in-
trorse, arguing that at least during anthesis they
are latrorse, with no hint of an introrse condition
in the buds. We agree with Goldblatt’s interpreta-
tion that the anthers are best interpreted as la-
trorse. There is a weak sign of the anthers being
introrse before they open. At that stage all four
thecae can be seen from the adaxial side, while
from the abaxial side only the two dorsal thecae
are seen. When opening the longitudinal slits are
oriented laterally with the two pollen presenting
surfaces eventually becoming completely flat, still
laterally oriented. While dehiscence takes place,
the curvature of the filaments brings the open an-
ther into a central position with regards to the
65
opening of the flower while the anther turns 908,
bringing the pollen presenting surfaces into an in-
trorse/extrorse position. Among Alstroemeriaceae
included, only Bomarea boliviensis seems to differ
from this pattern. In this species the pollen pre-
senting surfaces never become flat and the anthers
are introrse. In Luzuriaga radicans and L. marginata
the dehiscence is introrse, while in L. polyphylla the
pollen sacs open by pores and the character has
not been applied to this species.
50. Pollen sculpturing: striate-reticulate (0); reticulate
(1); foveolate (2). In all examined species of Alstroe-
meria the sculpturing of the exine is striate-reticu-
late, but reticulate in Bomarea and Leontochir (Schul-
ze 1978; Kosenko 1994; Sanso and Xifreda 2001).
In Disporum viridescens the exine is reticulate too
(Rudall et al. 2000), while in Luzuriaga and Uvu-
laria it is finely foveolate (Heusser 1971; Zavada
1983). In Dioscorea dodecaneura the exine pattern is
reticulate (Xifreda, pers. comm.).
51. Style development: completed before anther dehiscence
(0); completed after anther dehiscence (1). In nearly all
Alstroemeriaceae species examined the style is
very short during anther dehiscence, with prolon-
gation of the style and spreading of the stylodia
taking place after pollen presentation is completed.
52. Base of style: terete to three-lobed (0); three-edged (1).
In most Alstroemeriaceae the style is three-edged
particularly towards the base, while in Alstroemeria
isabellana and in Bomarea macrocephala, as well as in
the outgroup, the style is three-lobed to terete in
cross section.
53. Ovary winged: no (0); yes (1). In Alstroemeriaceae
the three angles of the base of the style continue
as ridges on the ovary uniting with the ridges
formed by the central nerve of the outer tepals.
54. Stigma divided into stylodia: no (0); yes (1). In Al-
stroemeriaceae, Uvularia and Disporum the style is
divided into 3 stylodia, while in Luzuriaga the style
is 3-lobed but not divided into stylodia.
55. Fruit type: capsule (0); berry (1).
56. Fruit dehiscence: gradual (0); explosive (1). A com-
plete analysis of fruit morphology and dehiscence
is presented in Sanso and Xifreda (2001). In Alstroe-
meria dehiscence is loculicidal and explosive, split-
ting from the base (into three parts). The loculicidal
capsule in Bomarea and Leontochir opens gradually,
beginning at the upper region (Sanso and Xifreda
2001). According to Wilbur (1963) the capsules
found in Uvularia open tardily and loculicidally.
57. Apex of capsule: gradually tapering towards the apex (0);
truncate (1). The fruit of Alstroemeria is rounded, more
or less oblong, and slightly or notably umbonate to-
wards the apex. In Bomarea and Leontochir it is semi-
spherical, turbinate and more or less truncate at the
apex (Sanso and Xifreda 2001). In Uvularia grandiflora
66 SYSTEMATIC BOTANY [Volume 28

the capsules are more or less truncate, but variation
is found within the genus (Wilbur 1963).
58. Sarcotesta: absent (0); present (1). Sarcotesta are found
in all species of Bomarea so far studied (Sanso and
Xifreda 2001) and in Leontochir (Wilkin 1997). A sar-
cotesta is found in isolated species belonging to the
Liliaceae sensu lato (Danilova et al. 1995). Recently
Norderstam (1998) reported the presence of a fleshy
aril or swollen strophiole in species of Colchicaceae
and Uvulariaceae, among them in Uvularia. Al-
though the aril found in Uvularia may be homolo-
gous to the sarcotesta found in Bomarea and Leon-
tochir, the character is coded as absent in Uvularia.
The lack of information concerning sarcotesta in
Uvularia makes us doubt its presence.
59. Form of seed: globose (0); rounded-ellipsoidal (1); an-
gular-subangular (2); compressed and winged (3). In Al-
stroemeria the seeds are globose while those found
in Bomarea and Leontochir are rounded as are the
seeds of Disporum (Don 1941) and Uvularia (Nor-
derstam 1998). In Luzuriaga the seeds are angular-
subangular (Rodriguez and Maticorena 1987).
60. Chromosome number (2n): 14 (0), 16 (1), 18 (2), 20
(3), 40 (4). All Alstroemeria species so far studied
are diploid 2n52x516, except an unidentified tet-
raploid collection of Goodspeed (1940). The Bom-
area species studied are diploid 2n52x518 except
B. hirtella (HBK) that is reported as tetraploid,
(Tsuchiya and Hang 1989; Hunziker and Xifreda
1990; Buitendijk and Ramanna 1996; Sanso (2002);
Sanso and Hunziker 1998). Leontochir has the same
number as Bomarea, 2n 5 18 (Bayer 1998). The
chromosome number found in representatives of
Uvularia is 2n514 (Love and Love 1981; Therman
and Denniston 1984; Utech 1980). Most species of
Disporum have 2n 5 6 but other numbers (2n 5
12, 14, 18, 22, 30, 32) have been reported (Kuro-
sawa 1983; Lee 1979; Love and Love 1981; Sen
1974; Tamura et al. 1992; Utech 1980). The chro-
mosome number reported in Luzuriaga parviflora is
2n 5 20 (Benzenberg and Hair 1963) and in Dios-
corea dodecaneura, 2n 5 40 (in this latter genus, the
base numbers are x 5 9, 10 and 12 [Huber 1998]).

APPENDIX 2. Morphological data matrix.

0 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

Als. andina 1 0 1 0 0 1 1 1 1 1 0 1 1 0 1 0 0 — 0 1 0 1 1 1 0 [02] 1 1 1 0 1 0

Als. angustifolia 1 0 1 0 0 1 1 0 0 1 1 0 0 ? 1 1 0 — — 0 0 1 1 1 0 2 1 1 [01] 1 1 0
Als. apertiflora 1 0 1 0 0 1 0 — 0 1 0 0 1 0 1 0 1 1 1 0 1 1 1 1 0 [12] 1 [01] 0 0 1 0
Als. aurea 1 0 1 0 0 1 1 0 0 0 0 0 0 0 0 1 0 — 0 0 0 1 1 1 0 1 1 1 1 1 1 0
Als. caryophyllaea 1 0 1 0 0 1 1 0 0 0 0 0 ? ? ? 0 0 — 1 0 1 1 1 1 1 [02] 1 0 0 ? 1 ?
Als. graminea 1 0 0 — 0 1 1 1 0 1 0 0 0 0 1 0 0 — — 0 0 1 1 1 1 0 1 1 1 1 1 0
Als. hookeri 1 0 1 0 0 ? 1 0 0 1 1 0 0 ? 1 1 0 — 0 0 0 1 1 1 0 2 1 1 1 1 1 0
Als. insignis 1 0 1 0 0 1 1 0 0 0 0 1 1 ? 0 0 ? ? 0 0 0 1 1 1 0 2 1 0 0 1 1 1
Als. isabellana 1 0 1 0 0 1 1 1 0 1 0 0 1 0 1 0 1 1 1 0 1 1 1 1 1 [12] 1 1 0 0 0 1
Als. ligtu 1 0 1 0 0 1 1 0 0 0 0 0 0 ? 0 2 0 — 0 0 0 1 1 1 0 2 1 1 1 1 1 [01]
Als. longistyla 1 0 1 0 0 1 0 — 0 1 0 0 1 ? 1 0 ? ? 1 0 1 1 1 1 1 [12] 1 1 0 0 1 0
Als. magnifica 1 0 1 0 0 1 1 0 0 0 0 0 0 ? 0 2 0 — 1 0 0 1 1 1 0 2 1 1 1 1 1 0
Als. pallida 1 0 1 0 0 1 1 0 0 1 0 0 0 ? 1 1 0 — 0 0 0 1 1 1 1 [02] 1 1 1 1 1 0
Als. patagonica 1 0 1 0 0 1 1 1 1 1 [01] 0 0 0 1 1 0 — [01] 1 0 1 1 1 0 1 1 1 1 [01] 1 0
Als. pelegrina 1 0 1 0 0 1 1 0 0 0 0 0 0 0 0 0 0 — 0 0 1 1 1 1 1 [02] 1 1 1 1 1 0
Als. presliana 1 0 1 0 0 1 1 0 0 0 0 0 0 0 1 1 0 — 0 0 0 1 1 1 0 2 1 1 1 1 1 0
Als. pseudospathulata 1 0 1 0 0 1 1 1 1 [01] 0 0 0 0 1 0 0 — 0 0 0 1 1 1 0 1 1 1 1 1 1 0
Als. psittacina 1 0 1 0 0 1 1 0 0 0 0 0 0 0 0 0 0 — 1 0 0 1 1 1 1 2 1 0 0 0 1 1
Als. pygmaea 1 0 1 0 0 1 1 1 0 1 0 0 0 0 1 1 0 — 0 1 0 0 1 [01] 0 1 1 1 [01] 0 1 0
Als. revoluta 1 0 1 0 0 1 1 0 0 1 1 0 0 ? 1 1 0 — 0 0 0 1 1 1 0 2 1 1 1 0 1 0
Als. sellowiana 1 0 1 0 0 1 0 — 0 1 0 0 1 ? 0 0 1 1 1 0 0 1 1 1 0 1 1 1 0 0 1 0
Als. versicolor 1 0 1 0 0 1 1 0 0 1 1 0 0 ? 1 1 0 — 0 0 0 1 1 1 1 2 1 0 0 0 1 0
Als. werdermannii 1 0 1 0 0 1 1 1 1 0 0 0 0 ? 1 1 0 — 0 0 0 1 1 1 0 2 1 1 1 0 0 0
Bol. boliviensis 1 0 1 1 0 1 1 0 0 1 0 2 0 0 0 0 1 0 0 0 1 1 1 1 0 1 1 1 [01] 0 0 0
Bol. edulis 1 2 1 1 0 1 1 0 0 0 0 2 0 0 0 0 1 0 0 0 1 1 1 [01] 1 2 1 1 0 0 [01] [01]
Bol. macrocephala 1 0 1 1 0 1 1 0 0 1 1 2 0 0 0 1 1 0 0 0 2 1 1 1 1 2 1 0 0 0 0 0
Dioscorea 1 2 0 — 0 0 0 — 0 1 0 0 0 3 0 0 0 — 1 0 1 1 1 0 0 3 0 — — 0 0 0
Dis. viridescens 1 0 0 — 1 0 0 — 0 0 0 0 0 2 0 1 ? ? 0 0 0 2 0 0 0 0 0 — — 0 0 0
Leo. ovallei 1 1 1 1 0 1 1 0 0 0 0 0 0 0 0 0 1 0 — 0 1 0 1 0 0 2 0 — — 0 1 0
Luz. marginata 0 3 0 — 1 1 1 0 2 0 0 1 1 1 0 0 1 0 1 1 1 2 0 0 0 0 0 — — 0 0 1
Luz. polyphylla 0 3 0 — 1 1 1 0 0 0 0 1 1 1 0 0 1 0 1 1 1 2 0 0 0 0 0 — — 0 0 0
Luz. radicans 0 3 0 — 1 1 1 0 0 0 0 1 1 1 0 1 1 0 1 0 0 2 0 0 0 0 0 — — 0 0 1
Uvu. grandiflora 1 0 0 — 1 0 0 — 0 1 0 2 0 0 0 1 ? ? 0 0 0 2 0 0 0 1 1 0 0 0 0 0
2003] AAGESEN & SANSO: PHYLOGENY OF ALSTROEMERIACEAE 67

APPENDIX 3. Parque Nacional Los Alerces, Cusato 4740 (BAA).

Voucher specimens used in the morphological anal-
yses not included in Sanso and Xifreda (1995) and San-
so (1996).
Alstroemeria andina. ARGENTINA. San Juan: Dpto. Ig-
lesias, Rio de las Taguas, Cordillera de la Ortiga,
3800 msm, 30 Jan 1999, Kiesling et al. 9216 (SI).
CHILE. Región IV: Prov. Coquimbo, Illapel, Car-
én, 18 Dic 1971, Beckett et al. 4612 (SGO).
A. angustifolia. DENMARK. Copenhagen: Cult. Hort.
Bot. Garden, University of Copenhagen P1995–
5003 (C).
A. apertiflora. BRAZIL. Mato Grosso: Mun. Rio Brilhan-
te, Rod. BR-267, 22 Oct 1970, Hatschbach 25088
(MBM). Paraná: Mun. Contenda, Rodovia do Xis-
to, 22 Oct 1967, Hatschbach 17552 (F). Mun. Curi-
tiba, Boqueirão, 8 Oct 1987, Silva 335 (MBM,
UPCB). Mun. Piraquara, São Roque, 9 Dec 1986,
Silva 264 (ESA). Mun. São José dos Pinhais, Rio
Pequeño, 20 Oct 1994, Ribas 687 (BHCB). Mun. Ti-
jucas do Sul, Rincão, 21 Oct 1977, Hatschbach
40440 (CTES).
A. aurea. ARGENTINA. Chubut: Dpto. Futaleufu,
Neuquen: Dpto. Catán-Lil, Sierra del Chachil, Ca-
jón Chico, 28 Jan 1965, Rúgolo and Agrasar 393
(BAA). Dpto. Minas, Reserva Prov. Lagunas de
Epulauquen, en bosque de la laguna superior, 20
Jan 1998, Aagesen s.n. (BAA). Dpto. Huiliches, 20
km desde Junin de los Andes hacia Lago Lolog,
22 Jan 1998, Aagesen s.n. (BAA).
A. graminea. CHILE. Región II: Prov. de Antofagasta,
Dpto. Taltal, hills southeast of Taltal, 25 Nov 1925,
Johnston 5099 (BA). Taltal, 7 Oct 1983, Elgueta 15
(SGO). Quebrada Paposo, 650 msm, ladera exp.
sur, 5 Sept 1991, von Bohlen 1275 (SGO). Paposo,
Quebrada Los Peralitos, 18 Oct 1991, Brinck w/n
(SGO 122848). Prov. de Atacama: Dpto. Chañaral,
vicinity of Puerto Chañaral, hills back of El Bar-
quito, 28–29 Oct 1925, Johnston 4811 (BA).
A. hookeri. DENMARK. Copenhagen: Cult. Hort. Bot.
Garden, University of Copenhagen P1995–5010,
5011 (C).
A. cf. insignis. BRAZIL. São Paulo: Mun. São Bernardo
do Campo, Parque Caminhos do Mar, Calçada de
Lorena, trecho A, 23 Jan 1991, Ferreira s.n. (SP).
Mun. Ubatuba, Beira de estrada, serra que liga

APPENDIX 2. Extended.

32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60

1 1 0 1 1 1 0 1 0 0 0 0 1 1 1 2 2 2 0 1 1 1 1 0 1 0 0 0 1
1 1 1 1 1 1 0 1 0 1 0 1 1 1 1 1 2 2 0 1 1 1 1 0 1 0 0 0 1
1 1 1 1 0 1 0 1 1 1 0 0 1 1 0 0 2 2 0 1 1 1 1 0 1 0 0 0 ?
1 1 1 1 [01] 1 0 1 0 1 0 0 1 1 1 0 2 2 0 1 1 1 1 0 1 0 0 0 1
1 ? ? 1 1 1 0 ? ? ? 0 0 1 0 ? 1 ? ? ? ? ? 1 1 ? ? ? ? ? 1
1 1 1 1 1 1 0 1 0 ? 0 1 1 0 [01] 0 2 2 0 ? 1 1 1 0 1 0 0 0 1
1 1 1 1 1 1 0 1 0 1 0 0 1 1 1 4 2 2 0 1 1 1 1 0 1 0 0 0 1
1 1 1 1 0 1 0 1 1 1 0 0 1 0 1 1 2 2 ? 1 1 1 1 0 1 0 0 ? ?
1 1 1 1 1 1 0 1 0 1 0 0 1 0 0 2 2 2 0 1 0 1 1 0 1 0 0 0 1
1 1 1 1 1 1 0 1 1 1 0 0 1 1 1 1 2 2 0 1 1 1 1 0 1 0 0 0 1
1 1 1 1 1 0 0 1 0 1 0 0 1 0 1 0 2 2 ? 1 1 1 1 0 1 ? 0 ? ?
1 1 1 1 1 1 [01] 1 0 1 0 1 1 1 1 2 2 2 0 1 1 1 1 0 1 0 0 0 1
1 1 1 1 1 1 0 1 0 1 0 0 1 1 1 [024] 2 2 0 1 1 1 1 0 1 0 0 0 1
1 1 1 1 0 0 0 1 0 1 0 0 1 1 0 0 2 2 0 1 1 1 1 0 1 0 0 0 1
1 1 1 1 1 1 0 1 0 1 0 1 1 0 1 2 2 2 0 1 1 1 1 0 1 0 0 0 1
1 1 1 1 1 1 0 1 0 1 0 0 1 1 1 0 2 2 0 1 1 1 1 0 1 0 0 0 ?
1 1 1 1 0 1 0 1 0 1 0 0 1 1 1 0 2 2 0 1 1 1 1 0 1 0 0 0 1
1 1 0 1 1 1 0 1 1 1 0 0 1 0 1 2 2 2 0 1 1 1 1 0 1 0 0 0 1
1 1 1 1 0 0 0 0 0 1 0 1 0 0 0 0 2 2 0 1 1 1 1 0 1 0 0 0 1
1 1 0 1 1 1 0 1 1 1 0 0 1 0 1 4 2 2 0 1 1 1 1 0 1 0 0 0 ?
1 1 1 1 0 1 0 1 0 1 0 0 1 0 0 ? 2 2 ? 1 1 1 1 0 1 0 ? ? ?
1 1 1 0 1 1 0 1 0 1 0 0 1 1 1 4 2 2 0 1 1 1 1 0 1 0 0 0 1
1 1 0 1 1 1 0 1 0 1 0 0 1 1 1 1 2 2 0 1 1 1 1 ? ? ? ? ? ?
0 0 0 2 0 1 [12] 1 1 1 0 0 1 0 1 0 2 0 1 1 1 1 1 0 0 1 1 1 2
0 0 0 2 1 0 1 1 1 1 0 0 1 0 [01] [023] 2 2 1 1 0 1 1 0 0 1 1 1 2
0 0 0 0 1 0 2 0 0 1 0 0 1 0 0 2 2 2 1 1 1 1 1 0 0 1 1 1 2
1 0 0 — 0 0 0 0 0 0 0 1 1 0 0 0 0 0 1 — 0 0 0 1 — — 0 3 4
0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 1 1 0 0 0 1 1 — — 0 ? 1
0 0 0 2 0 0 2 0 0 1 1 1 1 0 0 1 2 2 1 ? 1 1 1 0 0 1 1 1 2
0 0 0 1 0 0 0 0 1 0 0 1 0 0 0 0 0 0 2 0 0 0 0 1 — — 0 2 3
1 1 0 0 0 0 0 0 0 ? 0 1 0 0 0 ? 1 — 2 0 0 0 0 1 — — 0 2 3
0 0 0 1 0 0 0 0 1 1 0 0 0 0 0 0 1 0 2 0 0 0 0 1 — — 0 2 3
0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 1 2 0 0 0 1 0 0 1 0 1 0
68 SYSTEMATIC BOTANY
Ubatuba a Taubaté, a 13 km de Ubatuba, 22 Dec
1996, Assis and Skorupa 340 (SPF). Mun. Salesó-
polis, margem do Rio Claro, 30 Nov 1966, Kuhl-
mann 2763 (SP). Salesopolis, Casa Grande, Estação
Biologica de Boraceia, 8 Dec 1981, Filho 718 (UEC).
A. isabellana. ARGENTINA. Corrientes: Dpto. Ituzain-
go, Playadito, 7 km W de Colonia Liebig, 20 Jan
1993, Schinini 27574 (CTES). Rincón de Playadito,
5 km N de San Alonso, 25 Jan 1995, Tressens and
Barret 5180 (CTES). Misiones: Dpto. Capital, Po-
sadas, Pantano del Zaimán, 26 Dec 1997, Rua s.n.
(BAA).
A. ligtu. DENMARK. Copenhagen: Cult. Hort. Bot.
Garden, University of Copenhagen P1995–5015,
5016, 5028 (C).
A. cf. longistyla. BRAZIL. Distrito Federal: Mun. Brasi-
lia, 27 Dec 1978, Heringer 18392 (IBGE). Zoológico,
18 Oct 1976, Heringer 16212 (IBGE). Entrada BSB/
Planaltina, 6 Oct 1980, Heringer Salles 217 (IBGE).
Reserva Ecológica do IBGE, 14 Oct 1986, Mendon-
ça & Alvarenga 756 (IBGE). Reserva ecologica do
Guara, 26 Aug 1993, Pereira da Silva 1746 (CEN).
Mun. Planaltina, Gallery forest and adjacent cer-
rado slopes, Córrego Moneiro, vicinity of Planal-
tina, elevation 950 m, 29 Sep 1965, Irwin et al. 8774
(SPF). Goiás: Mun. Alto Paraı́so, Fda. Mato Fundo,
Oct 1990, Hatschbach 54670 (MBM). Mun. Formo-
sa, near Lagoa Feia, ca. 7 km S.E. of Formosa, 11
Oct 1965, Irwin et al. 9156 (SPF). Mun. Rio Verde,
BR-060, próximo ao Rio Doce, 19 Sep 1974, Hatsch-
bach 34976 (MBM). Mun. Pouso Alto, Chapada
dos Veadeiros, brejo a 1800 m.sm., 21 Dec 1968,
Graziela 601 (UB). Minas Gerais: Mun. Gouveia,
Serra do Espinaco, 13 Nov 1971, Hatschbach 27866
(MBM). Mun. Uberlândia, Estação Ecológica do
Panga, 12 Oct 1993, Romero et al. 513 (HUFU). Pa-
raná: Mun. Palmas, Estrada Ponte Serrada, 5 Dec
1971, Hatschbach 28252 (MBM).
A. magnifica. DENMARK. Copenhagen: Cult. Hort.
Bot. Garden, University of Copenhagen P1995–
5030, 5031 (C).
A. pallida. DENMARK. Copenhagen: Cult. Hort. Bot.
Garden, University of Copenhagen P1995–5035 (C).
A. patagonica. ARGENTINA. Neuquén: Dpto. Catán-
Lil, RN 40, 14 km desde Catán-Lil hacia La Negra,
22 Jan 1998, Aagesen s.n. (BAA).
A. pelegrina. DENMARK. Copenhagen: Cult. Hort. Bot.
Garden, University of Copenhagen P1995–5037
(C).
A. presliana. ARGENTINA. Neuquén: Dpto. Minas,
camino hacia la Reserva Prov. Lagunas de Epu-
lauquen, ca. 2 km despues de la confluencia de los
rı́os Buraleo y Nahueve, 20 Jan 1998, Aagesen s.n.
(BAA). DENMARK. Copenhagen: Cult. Hort. Bot.
Garden, University of Copenhagen P1995–5040,
5041 (C).
[Volume 28
A. psittacina. ARGENTINA. Buenos Aires: Capital Fed-
eral, cult. Hort. Bot. Fac. Agronomia, 5 Dec 1967,
Quesada s.n. (BA).
A. pulchra. DENMARK. Copenhagen: Cult. Hort. Bot.
Garden, University of Copenhagen P1995–5047
(C).
A. pygmaea. ARGENTINA. Tucumán: Dpto. Trancas,
Hualinchay, Rodeo Largo, 15 Jan 1998, Aagesen s.n.
(BAA).
A. revoluta. DENMARK. Copenhagen: Cult. Hort. Bot.
Garden, University of Copenhagen P1995–5050
(C).
A. cf. sellowiana. BRAZIL. Paraná: Mun. Campina
Grande do Sul, Pico Caratuva, 20 May 1967,
Hatschbach 16455 (MBM). Mun. Guaratuba, Serra
do Araçatuba, 21 Jan 1994, Kummrow 3369, 3387
(ESA, MBM). Mun. Quatro Barras, Morro Mãe Ca-
tira, 9 Apr 1986, Cordeiro 281 (MBM).
A. versicolor. DENMARK. Copenhagen: Cult. Hort. Bot.
Garden, University of Copenhagen P1995–5053
(C).
A. wedermannii. DENMARK. Copenhagen: Cult. Hort.
Bot. Garden, University of Copenhagen P1995–
5003 (C).
Bomarea boliviensis. ARGENTINA. Jujuy: Dpto. Tum-
baya, Laguna de Volcán, laderas occidentales, 8
Jan 1998, Aagesen s.n. (BAA). Salta: Dpto. Chi-
coana, RP 33, 41 km desde el desvı́o de El Carril,
13 Jan 1998, Aagesen s.n. (BAA).
B. edulis. ARGENTINA. Jujuy: Dpto. Capital, camino
hacia Laguna de Yala, 4 km desde RN 9, 12 Jan
1998, Aagesen s.n. (BAA). Tunal, 8 Apr 1988, Rot-
man 669 (LIL). Dpto. El Carmen, camino entre
Ruta 9 y Dique Las Maderas, 4 Mar 1987, Mar-
chessi et al. s.n. (BAA). Misiones: Dpto. Iguazú,
Parque Nacional Iguazú, 1 Dec 1993, Vanni et al.
3125 (CTES).
B. macrocephala. ARGENTINA. Catamarca: Dpto. An-
dalgalá, El Candado, Feb 1916, Jörgensen s.n. (BA).
Salta: Dpto. Chicoana, RP 33, km 50, 12 Dec 1995,
Hilger et al. s.n. (BAA). Tucuman: Dpto. Tafi, cam-
ino entre El Infernillo y Tafı́ del Valle, 14 Jan 1998,
Aagesen s.n. (BAA).
Leontochir ovallei. CHILE. Región II: Prov. de Atacama,
Carrizal Bajo, Oct 1886, Heisse s.n. (SGO 72917),
17 Sept 1957, Ricardi s.n. (SGO 73065), 20 Nov
1963, Bolı́var Rojas s.n. (SGO 75234).
Luzuriaga marginata. ARGENTINA. Rı́o Negro: Dpto.
Bariloche, Parque Nacional Nahuel Huapi, Puerto
Bles, senda a Lago los Clavos, 19 Dec 1998, Aagesen
s.n. (BAA). Laguna Frı́a, 15 Feb 1948, Dawson and
Schwabe 2763 (BAA). Tierra del Fuego: Dpto. Us-
huaia, Ushuaia, 5 Mar 1962, Robertson 63 (BAA).
L. radicans. ARGENTINA. Rı́o Negro: Dpto. Bariloche,
Parque Nacional Nahuel Huapi, Puerto Bles, cam-
ino a Las Cantaras, sobre árboles o en el suelo, 25
2003] AAGESEN & SANSO: PHYLOGENY OF ALSTROEMERIACEAE 69

Jan 1998; 18 Dec 1998, Aagesen s.n. (BAA). Parque
Nacional Nahuel Huapi, Puerto Bles, Laguna
Frias, May 1980, Cusato 2138 (BAA). Parque Na-
cional Nahuel Huapi, Valle del Rı́o Frias, 5 Feb
1946, Boelcke 2064 (BAA). CHILE. Región VIII:
Concepción, Reserva Universitaria Hualpén, 8 Jan
1964, Boelcke 10634 (BAA). Región IX: Temuco, ca-
mion de Lago Villarica a balsa sobe Rı́o Tolben, 6
Nov 1955, Boelcke 7348 (BAA). Región X: Pangui-
pulli, Fundo Trafún, 13 Jan 1962, Boelcke 8950
(BAA). Fundo Trafún, en piso selva y en troncos,
14 Jan 1962, Boelcke 8957 (BAA).