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Monophyly and Phylogenetic Relationships in Lymania (Bromeliaceae: Bromelioideae) Based on
Morphology and Chloroplast DNA Sequences
Author(s): Leandro de Oliveira Furtado de Sousa, Tânia Wendt, Gregory K. Brown, Dorothy E.
Tuthill and Timothy M. Evans
Source: Systematic Botany, Vol. 32, No. 2 (Apr. - Jun., 2007), pp. 264-270
Published by: American Society of Plant Taxonomists
Stable URL: http://www.jstor.org/stable/25064246
Accessed: 19-01-2016 20:21 UTC
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Systematic Botany (2007), 32(2): pp. 264-270
? Copyright 2007 by theAmerican Society of Plant Taxonomists

in Lymania
and Phylogenetic
(Bromeliaceae:
Relationships
Monophyly
on
DNA Sequences
Based
and
Bromelioideae)
Chloroplast
Morphology
de Sousa,1 T?nia Wendt,1-4 Gregory
de Oliveira
Furtado
E. Tuthill,2
and Timothy M. Evans3
Dorothy

Leandro

Federal do Rio de Janeiro, IB,CCS, Ilha do Fund?o, 21941-590,

de Bot?nica, Universidade

departamento

Rio

2Department of Botany, University
Department,

3Biology

de

82071 and Marie

Laramie, Wyoming
34236

Florida

Sarasota,

35 East
College,
for correspondence

Hope
4Author

Brazil;

Janeiro-RJ,

of Wyoming,

Gardens,

K. Brown,2

12th street,

Selby Botanical

U.S.A.;

Holland,

49423-9000

Michigan,

U.S.A.

(twendt@biologia.ufrj.br)

Communicating Editor:Matt Lavin
conducted
Abstract.
A cladistic
and sequences
from three
of Lymania was
using morphology
analysis
and trnh-trnF
The
the matK coding
DNA
spacers.
region and the psbA-trnH
intergenic
regions:
chloroplast
related genera were
examined.
Of the nine
of the genus and the phylogenetic
among
relationships
monophyly
in Bromeliaceae
to southern Bahia, Brazil. Lymania is the first genus
Lymania species, eight are endemic
subfamily
and molecular
to be subjected
to a combined morphological
Bromelioideae
analysis. The genera of Bromelioideae
and
to classify
and there has been disagreement
about their interrelationships
difficult
have been particularly
test
than molecular
data alone. The partition homogeneity
data show better resolution
Morphological
monophyly.
a single most parsimonious
a combined
tree. In the combined
of the two data sets, yielding
analysis
supported
and the genus
is closely related to species of Aechmea
of Lymania is moderately
supported,
analysis, monophyly
distinctiveness
low molecular
and subg. subg. Ortigiesia.
The morphological
coupled with
subg. Lamprococcus
indicates relatively recent and rapid speciation within
divergence
a framework
and molecular
data as done in this study provides
that could foster better taxonomic
rearrangements.
Keywords:

Brazil,

is a well-defined

Bromeliaceae

monophyly

combined

cladistics,

matK,

analysis,

monocot

its

family,

analyses
Bromeliaceae

and Brown
(Gilmartin
et al.
et al. 1993; Duvall

characters
1992; Clark

trnL-trnF.

psbA-trnH,

of

data

such

even

than

robust

data.

and molecular

morphological

being supported by several phylogenetic

and morphological
et al.
1987; Gaut

Lymania. The combined
analysis of morphological
for future research on other Bromelioideae
genera

not

have

been
have

though
they
of molecular

analyses

or

Combined
in

undertaken
more

proven

morphological
1992; Nandi

1993; Givnish

et al. 1999, 2004). Three subfamilies of

data

Bromeliaceae

have

et al. 1998; Chase et al. 2000; Sytsma and Pires 2001).

been

traditionally
recognized,
and Tillandsioideae

Pitcairnioideae,

Bromelioideae,

(Mez 1934-1935; Smith and Downs
Several
the
al.

phylogenetic

relationships
1990;

Terry

studies

have

the subfamilies
among
et al. 1997a; Horres
et al.

(Ranker
2000),

et

had

and

et

al.

2005).

All

these

works

reveal

genera

within

Bromeliaceae

often

Sanderson

nine

to delimit natural groups (Brown and Terry 1992;
Grant 1993), and the generic limits are frequently
change (e.g., Read 1984; Smith and
undergoing
Brown and Lerne 2005). The absence
1992;
Spencer
delimited monophyletic
of convincingly
groups,
iswell known
especially within the Bromelioideae,
(Duval et al. 2003; Faria et al. 2004).
of natural groups will be
The circumscription
studies using
aided
by phylogenetic
greatly

and

Araeococcus,

some

fail

from

added
Forzza
vided

a

limited

winged-ovaries
Aechmea
subgenus
New
Ronnbergia.

(1984)

all

to accommodate

species
that were
Lamprococcus,
were
species

to Lymania (Lerne 1987, 2006; Lerne and
2001), and recently De Sousa (2004) pro
the

revision.

first

diagnosed
most
genera

with

questionable.
examine

the

relatives,

which

a

distrib

narrowly
from

known

(De Sousa 2004). Read

Lymania
or

furrowed-

segregated

conflict within the generic concepts presented in the
of the family by
last comprehensive monograph
Smith and Downs (1974, 1977, 1979)
characters traditionally used to
Morphological
circumscribe

described

with

genera (Gilmartin and Brown 1986; Varadarajan and
Gilmartin 1988; Terry et al. 1997b; Faria et al. 2004;
Barfuss

(Donoghue

number of collections

on

focused

and

Read
comprises
Lymania
are
which
uted
species,

1974,1977,1979).
been

alone

only
in Bromelioideae,

relationships
not
have

is
genus
as with
is

the monophyly

study
among
been
Our

framework.

phylogenetic

and

characters,

current

The

the

However,

a few

to

is designed
and

Lymania
evaluated
specific

its

within
aims

are

to: 1) evaluate the utility of a combined phyloge
and molecular data
netic analysis of morphological
on

a

group

of

closely

related

Bromelioideae

of Lymania using
species, 2) assess the monophyly
three chloroplast
and
from
sequences
morphology
DNA

regions

{matK,

trnL-trnF,

and

264
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All use subject to JSTOR Terms and Conditions

psbA-trnH),

and

DE SOUSA ET AL.: PHYLOGENETIC

2007]

3)

examine

and the distribution
within

among

relationships

this

Lymania

of specific morphological

species,

traits

genus.

and

Materials

Methods

Taxon

23 species
Our data matrix
Sampling.
comprises
five genera
of subfamily
Bromelioideae
representing
(Ap
all recognized
of Lymania were
1). Almost
pendix
species
L.
B.
included
for
marantoides
(L.
Sm.) Read and L.
except
to obtain material.
l?nguida Lerne, for which we were unable
of the genera Aechmea
(10 species), Araeococcus
(2),
Samples
and Ronnbergia
some Lymania
chosen
because
(2) were
were
included
in these genera.
Two
species
previously
as outgroups
based
species of Cryptanthus were
designated
on Terry et al. (1997a) and Faria et al. (2004). Almost
all plant
material was collected
from naturally
occurring
populations
two years of field studies
in the eastern
conducted
during
Brazilian Atlantic
Rain Forest of Rio de Janeiro, Espirito
of Aechmea pedicellata and
Santo, and Bahia States. Material
Araeococcus goeldianus were
obtained
from The Marie
Selby
Botanical Gardens
(SEL).
Data.
We scored 60 morphological
char
Morphological
acters for this study (Appendix
annotation
2, 3). Meticulous
the habit and color of the plant structures, as well
concerning
as photo documentation,
were made
in the field or blooming
ex-situ. Flowers and fruits in different periods
of maturation
were
in 70% ethanol prior to examination.
The
preserved
characters were
determined
after preparing
the
remaining
herbarium
vouchers.
Continuous
variables
leaf
(e.g., mature
to discrete
on
variables
converted
based
length) were
as it was possible,
observed
gaps between
species. As much
or clones were used for the
the same individual
morpholog
ical and molecular
DNA

data

(Appendix

1).
and

The
Amplification
Sequencing.
were
extraction
obtained
from living
samples
collected
in the field or in cultivation
and quickly
plants
dried in silica gel. Total DNA was
isolated using a modifica
tion of the 2X CTAB buffer method
1987;
(Doyle and Doyle
Smith et al. 1991) followed by purification
with
the QIAamp
DNA Mini Kit (Qiagen). The protocols
for PCR amplification
and sequencing
of all three cpDNA
regions follow Johnson
and Soltis
of matK, we used
(1995). For PCR amplification
two and five from Crayn et al. (2000). psbA-trnH and
primers
were
trnL-trnF primer
taken from Sang et al.
sequences
all DNA
(1997). Almost
sequences were originally
produced
Extraction,
for DNA

for this study, except for Cryptanthus
the
beuckeri, for which
matK sequence
earlier as
(GenBank AF 539965) was deposited
1). For the total combined
part of another study (Appendix
data matrix
3.7% of the cells
plus molecular),
(morphology
were
scored as ''missing/'
Data Analysis.
All matrices
for this study were deposited
in TreeBASE
were
(study number SI 754). Cladistic
analyses
maximum
version
(PAUP
performed
using
parsimony
Swofford
conducted
searches were
4.0b.l0,
2003). Heuristic

OF LYMANIA

265

search were conducted
bootstrap
replicates with full heuristic
to evaluate
for each clade.
support
For parsimony
the morphological
and cpDNA
analysis,
data sets were
each analyzed
and in combina
individually
tion. Before combining
the separate
data sets, congruence
between
them was examined
the incongruence
using
length
differences
is referred to
(ILD) test (Farris et al. 1995), which
as the partition
test in PAUP. Cunningham
homogeneity
that combining
the data improved or did not
(1997) indicated
if a P value of the ILD test
reduce the phylogenetic
accuracy
was greater than 0.01.
For Bayesian
of the combined
analyses
morphological/
molecular
data matrix,
of evolution were
appropriate models
selected
for the matK region (HKY + G) and the non-coding
ratio test as
cpDNA
regions (F81 +1 + G) using a hierarchical
in Modeltest
and Crandall
(Posada
1998).
implemented
ver. 3.1.2 (Huelsenbeck
Bayesian
analyses
using MrBayes
and Ronquist
and Huelsenbeck
2001; Ronquist
2003) con
sisted of 5,000,000
with
taken every
generations,
samples
Trees from the first 1.2 million
10,000 generations.
genera
tions were
discarded
(i.e. the first 25% of the generations
were
the "burn-in"
stage).

RESULTS
Morphological
dataset

The morphological

Analysis.
nine

produced

most

trees

parsimonious

of

219 steps (CI = 0.45, RI = 0.59; Table 1). The strict
consensus

tree

from

data

morphological

showed

that the ingroup iswell supported as monophyletic
with a bootstrap value of 100%, but there is little
resolution

to

sister

the

clade

containing
and
subg.
was
found

support

bootstrap
Molecular

matrix

45

Ortgiesia,
for this

no

though
relationship.
molecular

aligned

informative.
data

molecular
trees

parsimonious

83

of

1782
and

uninformative,

parsimony

parsimony
of

analysis

that is

a total of 1858 characters with

had

31 being

Lymania
of Aechmea

species

The

Analysis.

constant,

being

shown).

forms a polytomy

and

Lamprococcus

subg.

(tree not

genera

among

is monophyletic,

yielded
(CI

steps

cladistic

The
=

214

most

0.86,

RI

=

0.79; Table 1). The strict consensus (not shown) is
largely unresolved, with a large basal polytomy
and

four main

is not

clades.

The

of

monophyly

Lymania

supported.

Combined Morphological
and Molecular Analy
sis. The partition homogeneity
test produced a P
value of 0.15, indicating combinability for the two
data

sets.

data

resulted

The

parsimony
in a
single

of combined
analysis
most
tree
parsimonious

on,
using the TBR branch swapping
algorithm, MULTREES
and 10000 random addition
replicates. All characters were
and assigned
unordered
Relative
levels of
equal weight.
and
in the data
sets were
homoplasy
synapomorphy
calculated
the consistency
index
(CI), the retention
using
index
index
(RI), and the rescaled
(RC) as
consistency
in PAUP* 4.0M0. The cpDNA
implemented
sequences were
and combined
into a single molecular
aligned manually
matrix.
A matrix
of morphological
characters was
created
the computer
MacCLADE
version
4.0
program
using
and Maddison
character
(Maddison
2000). Morphological
were
state changes
and AC
mapped
using MacClade

in overall topology to that produced by parsimony,
but with slightly poorer resolution and a paraphy
letic (instead of polyphyletic)
Ronnbergia (Fig. 2).
The monophyly
of Lymania is moderately
sup

CTRANS

and

character

state

optimization.

One

thousand

=
=
0.56, RI
0.61; Table 1).
(Fig. 1) of 310 steps (CI
Bayesian

ported
is sister
and

resulted

analysis

in these

analyses.

to Aechmea

subg.

sister

Ortgiesia.

in a tree

In each

from
species
Araeococcus

to the clade

that was

analysis,
subg.
is

formed

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All use subject to JSTOR Terms and Conditions

similar

Lymania

Lamprococcus
monophyletic

by

species

of

266

SYSTEMATIC

Table
related

1. Summary
genera.

for parsimony

statistics

trnL-trnF

psbA-trnH
inmatrix
of characters
of constant characters
of uninformative

No.
No.
No.
No.

characters
16
of informative

No.

characters
of trees

No.

of steps

-

of morphological,

analyses

602
577

matK

389
375

Total Molecular

9 6
-

1858

31
1

83

219

RC

310

0.67

-

2

Figure

and

Lymania
subg.

Aechmea

and

Lamprococcus

(subg.

Ortgiesia).

of this genus has long been
polyphyly
and Downs
1974, 1979). While
(Smith
recognized

generic

delimitation

supported

of

in previous

present

nized

Integration
of Morphological
Data
in Lymania
and Related
Aechmea

and Molecular
The
Species.
has

not

been

(Faria et al. 2004)

studies

rit
?

necessary
genera.

Scotland et al. (2003) noted that morphological
data are currently utilized
less for phylogeny
reconstructions

Aechmea lingulata

This

Araeococcus goeldianus

seven
includes
of nine
recog
more
extensive
of
species,
sampling
other
will
be
Bromelioideae
genera
to clarify
the two
between
relationships
study

Lymania
and
Aechmea

Aechmea turbinocalyx

Ronnbergiabrasiliensis

0.450.56
0.59 0.61
0.27
0.34
31

and
the

Discussion

Morphology + Molecular

5687
9

CI 0.86
RI
0.79

-

than

are molecular

is true for bromeliad

(Ranker

et al.

2000; Horres

1990;

Terry

phylogenetic

et al.

et al. 2000; Duval

1997a,

Araeococcus parviflorus

Ronnbergiabrasiliensis

Ronnbergia neoregelioides
Araeococcus goeldianus

Lymaniaglobosa

Araeococcus parviflorus

Lymaniabrachycaulis

Lymaniaalvimii

Lymaniacorallina

Lymaniaspiculata

Aechmea pedicellata

Lymaniaazurea

Aechmea gamosepala

Lymaniasmithii

Aechmea gracilis

Lymaniaglobosa

Aechmea capixabae

Lymaniabrachycaulis

Aechmea warasii

Lymaniacorallina

Aechmea racinae

Aechmea pedicellata

Aechmea miniata

Aechmea gamosepala

Aechmea carvalhoi

Aechmea gracilis

Ronnbergianeoregelioides

FiG. 1. Single most
parsimonious
neous analysis of morphological
and
data of Lymania and related
sequence
deae with Cryptanthus as the outgroup.
than 50% are given
above
greater

et al.

Aechmea lingulata

Lymaniasmithii

c

b; Crayn

Aechmea turbinocalyx

Lymaniaazurea

hy^L

studies

et al. 2003; Barfuss

Lymaniaspiculata

C

data.

sequence

Lymaniaalvimii

100
1 56

and

49 4

45

214

sets for Lymania

60 1918
0
1782

1782

16
-

data

Morphology

21

8

and combined

molecular,

867
830

...
-

[Volume 32

BOTANY

Aechmea capixabae

Cryptanthusbeuckeri

Aechmea warasii

Cryptanthusbromelioides

Aechmea miniata
tree from

the simulta

DNA
three chloroplast
species of Bromelioi
Bootstrap percentages
the branches.
Selected
onto the parsimonious
tree are indicated
characters mapped
in the list of
with black bars. Numbers
refer to characters
and states discussed
characters.
Characters
morphological
in text.

Aechmea racinae
Aechmea carvalhoi
Cryptanthusbeuckeri

d

Cryptanthusbromelioides

tree from Bayesian
of mor
Consensus
analysis
and cpDNA
sequence data. Posterior probabilities
phological
greater than 50% are given above the branches.
FiG. 2.

This content downloaded from 132.174.255.116 on Tue, 19 Jan 2016 20:21:16 UTC
All use subject to JSTOR Terms and Conditions

DE SOUSA ET AL.: PHYLOGENETIC

2007]

et

al.

lower
support

show higher

of

levels of
and

resolution,
and

(e.g.,
bootstrap
than molecular

values)
An
increased

probability
et al. 2003).
an

levels

that

assumption

general

data should

morphological
homoplasy,
levels
of

a

is

There

2005).

additional

posterior

(Scotland
in
characters

of

and this is
increases
accuracy,
generally
of the primary
of molecular
advantages
However
DNA
studies
undertaken
for

analysis
seen as one

data.

and

Bromeliaceae,

low

specifically
suffered
from

have

melioideae,

level of phylogenetic
low

unusually
our

study,

a more highly
data

subfamily

an

rate

to an

due
et al.

(Terry

alone.

data

morphological

The

combined

resolution
complete
and higher
support

1997a,

in

resulted
within

species
across

the

Lymania
tree

(Figs.

1

and 2). This is the first study in Bromeliaceae
in
which morphological
and molecular
data have
been

combined

a

for

phylogenetic

and

analysis,

this approach appears to hold great promise for
systematic studies in the family. While combining
data

from

sets

data

different

still

some

remains

what controversial (see reviews by de De Queiroz
et al. 1995; Huelsenbeck
et al. 1996), there are
in

reasonable
of
ysis
exhibits

data

favor

arguments
sets.
In Bromeliaceae,

of

combined

anal

a

that
family
and
convergent
parallel
a low nu
evolution
with
morphological
coupled
cleotide
substitution
of multiple
rate,
integration
data

sets

yielding
cation.

a

provides
a stable,

of

the

perhaps

combined

The morphological

sets

data

each

for a monophyletic
genus,
character
unambiguously
of a furrowed
presence
been

to define

used

low
relatively
there is support

no

single
unites

or

the mono

support

phyly of Lymania, albeit with
support values (Figs. 1, 2). While

has

for
hope
classifi

greatest

phylogenetically-based

of Lymania.

Monophyly
and

deal

great

winged

the genus,

morphological
the
clade.

The

which
ovary,
in
is also present

Aechmea carvalhoi (Fig. 1). Likewise, a bottleform
rosette is found in all Lymania species, but it has
been

in Araeococcus

derived

(Fig. 1). This high
evolution
parallel
sistently
generic

and

systematics

attempts

hampered

has

to establish

well

we

have not yet found

morphological
data
support

character
maintenance

for

Lymania

of A.

that incorporate
could

Lamprococcus

subg.
of
the

of

circumscription

among

Relationships
bined

Lymania

Lymania

with

species,

falling

clades

1, 2). One
(Figs.
and L. smithii)

resolution

complete
members

of

that

L.
(L. aivimii,
spiculata,
is united
the
of
presence
by
and a furrowed
disintegrate,

soon

to winged)

ovary

clade

the

(L. brachycaulis,

within

genus

(Fig. 1). The second

and L. globosa) is united by flower
from

2-3.5

sepals
these

characters

L. corallina,

length (ranging

acute
to attenuate,
apex
sepal
and ovary
While
each
of
winged.
within
is homoplasious
Brome

cm),

car?nate,

a whole,

as

the

supported

clade

(as opposed

lioideae

The com

into two relatively well

genus

bracts

in

resulted

analyses

and

Species

Traits.

of Morphological

are

they

unambiguous

within

synapomorphies
While
material

of

Lymania.
L. marantoides

and

the newly

described L. l?nguida (Lerne 2006) was not available
for this study, the distribution of characters within
some insight
the Lymania clade may provide
the

regarding

has

of

placement

two

these

It

species.

that each of these two
hypothesized
species is morphologically
closely related to L.
corallina (Lerne 2006; De Sousa 2004). The presence

of

been

the alate-carinate
clade,
ovary
close
affinity

winged
tively

characters,

logical
within

that

the winged
apex.
having
bract

clade
ovary

unite
ovary may
which
would
to L.

corallina.

make
however,
more
problematic.
clade
also
share

them

with

the

a rela
support
Other
morpho
the placement
Members
an

acute

of
sepal
as

is described
however,
Lymania
l?nguida,
an obtuse
the scape
Likewise,
apex.
sepal
as soon
in L.
is described
disinte
l?nguida

a character
grating,
the furrowed
ovary
acteristics
from each

state
clade.
of

the

that
The
two

unites

members

presence
clades may

of

of
char

suggest

a relatively basal position for L. l?nguida within the
Lymania clade or hybrid origin. It is not possible to
hypothesize
because
ships

more

about

information
characters

is not

L. marantoides
regarding
available.

relation
several

mor

and
that

phological

con

We
thank
Acknowledgements.
the Brazilian
Forestry
Service
(IBAMA) and the Brazilian Research Council
(CNPq)
for the field collection permits;
the numerous
colleagues who
us
with
assisted
plant collection; H. Luther, who provided
some of the plant material; A. P. G. Faria for assistance with
data analysis and for constructive
comments; Matt Lavin and
on the
three anonymous
reviewers
for helpful
comments

solid

circumscriptions.

Although
agnostic
current

of convergent
degree
a difficulty
illustrates

Bromelioideae

pervades

as

parviflorus

be
relationships
remain
unclear.

that future studies

Distribution

scape
produced

analyses

among
values

subgenus

the

Lymania.

L. azurea,

than molecular

resolved phylogeny

result

and
and

species
in modification

among

unexpectedly

resolution

substitution

Bro

et al. 2000; Barfuss et al. 2005).

b; Horres
In

for

267

however,

Lamprococcus,
tween
this

It is possible

lower

data

number

OF LYMANIA

a single di
the
Lymania,
of
generic

status, as advocated by De Sousa (2004). Our study
included only six (of ca. 14) species from A. subg.

the United
States National
Science Foundation
manuscript;
the Brazilian
for funding;
(DEB-0129446,
DEB-0129414)
Council
for Graduate
Studies
(CAPES) for a research grant

This content downloaded from 132.174.255.116 on Tue, 19 Jan 2016 20:21:16 UTC
All use subject to JSTOR Terms and Conditions

268

systematic

to L. O. F. De Sousa; and CNPq for a productivity
grant to T.
Wendt.
This paper is part of a Master's
at
thesis undertaken
in Botany of the Universidade
the Post-Graduate
Program
Federal do Rio de Janeiro by the first author.
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in the
systematics
frontier. Taxon 50:

(Mart, ex Schult. & Schult.f.) Lindm.: Brazil, Bahia, Sousa 453
EF110647.
EF110671,
(RFA), EF110624,
Cryptanthus Otto &
A.Dietr.
C. beuckeri E. Morren:
Brazil, Bahia, Wendt 489
Otto &
EF110672, EF110648. C. bromelioides
(RFA), AF53965,
: Brazil, Rio de Janeiro, Ro?as s.n. (RFA), EF110637,
A.Dietr
L. alvimii
EF110684, EF110661. Lymania Read
(L.B.Sm. &
s.n.
Read
Read:
Bahia,
Brazil,
Read)
(RFA), EF110625,
EF110649.
L. azurea Lerne: Brazil, Bahia, Sousa
EF110673,
L. brachycaulis
EF110660.
EF110683,
(RFA), EF110636,
(Baker) L.F. Sousa: Brazil, Bahia, Sousa 443 (RFA), EF110639,
ex Beer)
no sequence obtained, EF110663. L. corallina
(Brong.
Read: Brazil, Bahia, Sousa 458 (RFA), EF110640,
EF110686,
EF110664. L. globosa
Lerne: Brazil, Bahia, Lerne 2989 (RFA),
EF110626, EF110674, EF110650. L. smithii Read: Brazil, Bahia,
Sousa 407 (RFA), EF110627, EF110675, EF110651. L. spiculata
Sousa 416 & 417 (RFA),
Lerne & Forzza:
Brazil,
Bahia,
EF110628, EF110676, EF110652. Ronnbergia E.Morren & Andr?
?
E. Pereira & I. A. Penna: Brazil, Bahia, Lerne
R. brasiliensis
3017 (RFA), EF110644, EF110690, EF110668. R. neoregelioides
Lerne: Brazil, Bahia, Lerne 4368 (SEL), EF110632,
EF110679,
EF110656.
2. Morphological
characters
of Lymania (Bromeliaceae).

Appendix
Olmstead.
1997a.
in Bromeliaceae
locus ndhF.
plastid

and-.
1997b. Phylogenetic
relationships
in subfamily
Tillandsioideae
(Bromeliaceae)
using ndhF
sequences.
Systematic Botany 22: 333-345.
1988. Phylogenetic
G. S. and A. J. Gilmartin.
Varadarajan,
of groups of genera within
the subfamily
relationships
13:
Pitcairnioideae
(Bromeliaceae).
Botany
Systematic
-,-,

283-293.
1. Taxonomic
for combined morpho
Appendix
sampling
and molecular
of Lymania and
analysis
phylogenetic
as the
with
of Bromelioideae
Cryptanthus
species
to those
correspond
outgroup.
Subgenera
assignments
and Downs
1979).
(i.e., Smith
traditionally
recognized
are listed
in the following
details
Voucher
sequences:
and herbaria
collection
locality, collector name and number,
logical
related

at which
accession
trnH.

269

406

Press.
York: Hafner
-.
and
1977. Tillandsioideae
in Flora Neotropica. Mon.
Pp. 663-1492
York: Hafner Press.
-.
and
1979. Bromelioideae

New
-

OF LYMANIA

is deposited
GenBank
the voucher
(parentheses).
are in the order matK, trnL-trnF, psbA
numbers

A. lingulata
Aechmea
(L.)
subg. Aechmea Ruiz & Pav.
Baker: Brazil,
Santo, F aria 81 (RFA), EF110643,
Espirito
EF110689, EF110667. Aechmea subg. Lamprococcus (Beer) Baker
A. capixabae L.B.Sm.: Brazil, Espirito Santo, Faria 90 (RFA),
E. Pereira &
EF110657. A. carvalhoi
EF110633,
EF110680,
Lerne: Brazil, Bahia, Lerne 579 (RFA), EF110641,
EF110687,
ex Baker: Brazil, Bahia,
EF110665. A. miniata
(Beer) Hort,
EF110662. A. pedicel
Sousa 436 (RFA), EF110638, EF110685,
in Selby
lata Lerne & H. Luther: Brazil, Espirito
Santo,
Bello s.n. (SEL, RFA), EF110629, EF110677,
Garden
collection,
A. racinae L.B.Sm.: Brazil, Espirito Santo, Faria 80
E. Pereira:
(RFA), EF110635, EF110682, EF110659. A. warasii
EF110681,
Santo, Faria 76 (RFA), EF110634,
Brazil, Espirito
A.
EF110658. Aechmea subg. Macrochordion
(deVriese) Baker
turbinocalyx Mez: Brazil, Bahia, Wendt 461 (RFA), EF110622,
EF110645. Aechmea
EF110669,
subg. Ortgiesia
(Regel) Mez
Wittm:
A. gamosepala
Brazil, Rio de Janeiro, Wendt 351
(RFA), EF110631, no sequence obtained, EF110655. A. gracilis
EF110653.

Lindm.: Brazil, Rio de Janeiro, Sousa 464 (RFA), EF110642,
EF110688, EF110666. Araeococcus
subg. Araeococcus Brongn.
in
A. goeldianus
L.B.Sm.:
Garden
French Guiana,
Selby
s.n. (SEL), EF110623, EF110670, EF110646.
collection, Moonen
A. parviflorus
Araeococcus
Mez
subg. Pseudaraeococcus

analyses

used

in the cladistic

1 =
0 = broadly
infundibuliform;
shape:
=
= flattened.
2.
2
3
infundibuliform;
narrowly
lageniform;
Number
of leaves per rosette: 0 = < 10; 1 = 10 to 15; 2 = >
a water
15. 3. Rosette
tank: 0 = yes; 1
impounding
forming
= no. 4. Habit:
1 = primarily
0 = primarily
epiphytic;
5. Stolon
2 = facultative.
formation:
terrestrial or saxicolous;
=
0 = primarily
from the
from between
leaves; 1
primarily
base of the rosette. 6. Form of clonal growth:
0 = elliptical
=
=
=
linear cluster; 2
cluster; 1
stolon; 3
climbing
pendent
leaf length (include blade and sheath, cm):
stolon. 7. Mature
=
=
=
<
>
>
of leaf blade
0
45 to 65; 2
65. 8. Width
45; 1
= <
= > 2.5 to
= > 5. 9. Leaf blade base
(cm): 0
2.5; 1
5; 2
=
not or slightly differentiated;
1 = distinctly
and sheath: 0
but not petiolate;
2 = petiolate
blade
differentiated
between
10. Leaf blade
1 =
0 = absent;
and sheath.
channeling:
1
11. Shape of the leaf apex: 0 = acute or attenuate;
present.
= rounded or obtuse. 12. Leaf
armament:
0 = none
margin
=
= serrate. 13.
and entire, to obscurely
serrate; 1
Scape: 0
erect and hidden
in the center of the rosette; 1 = erect or
and
the rosette; 2 = pendulous
slightly curved and exceeding
rosette.
the
14.
0 = yes,
Scape bracts present:
exceeding
soon disintegrating;
15.
1 = yes,
2 = no.
persistent;
of scape
Distribution
bracts:
0 = erect and
imbricate,
1 = erect, not
the scape
the scape;
imbricate,
covering
=
2
16. Scape bract
the scape exposed.
exposed;
spreading,
=
=
0
armed. 17. Scape bract color: 0 =
entire; 1
margin:
or pale (tan); 1 = rose or red. 18. Scape bract apex:
greenish
1 =
to attenuate;
19.
to obtuse.
0 = acute
rounded
= <
= > 10 to
= >
Inflorescence
(cm): 0
10; 1
20; 2
length
in relation
to the
20. 20. Inflorescence
and scape position
1 =
not exceeding
the leaf-sheaths;
leaves: 0 = nidular,
but not exceeding
the leaf-sheaths,
the leaf-blades;
exceeding
2 = completely
21. Inflorescence
the leaf-blades.
exceeding
1 = simple or
0 = none, simple, never branched;
branching:
at the base; 2 = bipinnate;
3 = tripinate or
rarely branched
=
on rachis or
22. Flowers
more; 4
congested
corymbiform.
0 = no, rachis visible; 1 = yes, rachis
inflorescence
branches:
23. Flower
not visible.
the rachis or
arrangement
along
0 = polystichous;
1 = distichous;
2
inflorescence
branches:
=
= in
on main axis, and distichous
fascicles; 3
polystichous
=
=
on the branches.
24. Primary bracts: 0
absent; 1
present.
1 = entire. 26.
bract margins:
0 = armed;
25. Primary
=
=
rose or red. 27.
Primary bract color: 0
greenish or pale; 1
=
1 = membranaceous
coriaceous;
Primary bract texture: 0
1. Rosette

This content downloaded from 132.174.255.116 on Tue, 19 Jan 2016 20:21:16 UTC
All use subject to JSTOR Terms and Conditions

270

systematic

or chartaceous.
28. Primary
bract apex: 0 = acute to long
1 = rounded
to obtuse.
29. Floral bracts:
0 =
attenuate;
or inconspicuous.
1 = absent
30. Floral
bract
present;
0 = entire; 1 = armed. 31. Floral bract color: 0 =
margins:
=
= rose or
blue, lilac or purple.
red; 2
greenish, pale or tan; 1
=
32. Floral bract apex, excluding
the terminal
spine: 0
to obtuse; 1 = attenuate
rounded
to acute. 33. Terminal
spine
1 =
of floral bract apex: 0 = very short and inconspicuous;
=
or
2
but
shorter
than
the
bract;
conspicuous,
equaling
(length/width
longer than the bract. 34. Floral bract shape
=
ovate (1:1); 1 = narrow ovate (3:1); 2 =
ratio): 0
broadly
=
=
ovate
no; 1
(2:1). 35. Floral bracts car?nate: 0
yes, one
= <
keel. 36. Flower
0
cm):
0.5; 1
pedicel,
length (excluding
= > 0.5 to
= > 2 to
= > 3.5 cm to
= > 5. 37.
3.5; 3
2; 2
5; 4
38. Sepal color: 0 = white,
Pedicel:
0 = absent; 1 = present.
=
= red or
=
or
or
1
rose; 3
tan;
blue,
lilac,
greenish
purple; 2
=
or
4
39.
bicolored.
orange;
yellow
Sepal apex,
distinctly
= rounded
terminal
to obtuse; 1 = acute
excluding
spine: 0
=
or
to attenuate.
40. Sepal
terminal
0
absent
spine:
1 = present, but shorter than the sepal; 2 =
inconspicuous;
or
the sepal. 41. Sepal
0 =
symmetry:
equaling
exceeding
or
1 = strongly asymmetric.
42.
symmetric
slight asymmetric;
car?nate: 0 = yes; 1 = no. 43. Sepal connation:
0 =
Sepals
= connate
free or mostly
free; 1
by 1/3 or more of its length.

botany

[Volume 32

cream or greenish;
color: 0 = white,
1 = blue, lilac or
2 = yellow or orange. 45. Petal apex: 0 = rounded
to
1 = acute to attenuate.
46. Petal shape: 0 = linear,
or nearly so; 1 = spathulate, with distinct
margins
parallel
blade and claw. 47. Petal appendages:
0 = absent; 1 = pouch
or sac-like; 2 = ligulate with conspicuous
3
fringed margins;
=
48. Petal with
lateral folds on
ligulate with entire margins.
=
=
the adaxial
0
1
surface:
49. Petal
absent;
present.
at anthesis:
orientation
0 = erect or cucullate;
1 = spreading
or recurved.
50. Petal connation:
0 = free; 1 = connate.
51.
= <
= > 0.5 to <
= > 1 to
Filament
0.5; 1
1; 2
length (cm): 0
=
=
^ 2. 52. Filament
< 2; 3
1 =
shape: 0
compl?nate;
=
filiform. 53. Filament
fusion: 0 = none,
filaments
free; 1
to the petals;
filaments
adnate
2 = yes,
yes, antepetalous
a tube. 54. Anther
filaments
(cm):
connate,
length
forming
0 = < 0.2; 1 = > 0.2 to < 0.5; 2 = > 0.5. 55. Ovary
shape in
cross-section:
0 = circular; 1 = trigonous.
56. Ovary wall
ornamentation:
0 = none, smooth and regular; 1 = furrowed;
2 = winged;
4 = pilose. 57. Ovary
3 = verrucose;
color: 0 =
or green; 1 = yellow
or orange; 2 = rose or red. 58.
white
in relation
to anthers
at anthesis:
0 =
Stigma
position
=
or
equaling
slightly shorter than the anthers; 1
exceeding
the anthers. 59. Ovules
1 =
0 = yes, caudate;
appendaged:
= >
= < 5.
no. 60. Ovule
1
number
locule:
0
5;
per
44. Petal

purple;
obtuse;

Appendix
3. Matrix
character states (from Appendix
of morphological
2) used in a phylogenetic
related species of Bromelioideae
with Cryptanthus as the outgroup.
characters or missing
Inapplicable
a dash.
Character number
Aechmea

capixabae
Aechmea carvalhoi
Aechmea gamosepala
Aechmea gracilis
Aechmea
lingulata
Aechmea miniata
Aechmea pedicellata
Aechmea racinae
Aechmea turbinocalyx
Aechmea warasii
Araeococcus
goeldianus
Araeococcus
parviflorus
beuckeri
Cryptanthus
Cryptanthus
bromelioides
Lymania
Lymania
Lymania
Lymania

alvimii
azurea
brachycaulis
corallina

Lymania globosa
Lymania smithii
Lymania

spiculata

Ronnbergia
brasiliensis
Ronnbergia
neoregelioides

analysis of Lymania and
data are indicated with

10

20

1102111111
1000120111

0110000012
1110201012
1010100012
1110000011

2030-1

-10300

1111012000

203111101
0100-00

-10000
0111010202

1101011000
1111002100
1111012100

1110010022

2001101000

1110100002

12021-0100

1102100110
0202102200
1002111111
110-100011
1002110111

30

2001111000

40

1112010202
0121010002
-10400
-11300
-21200

50

60

1011031000
0011010001
10110020-0

1011002010

1110110110

0000000000

1101012000
1100103000
1102002000

1001032000
1001041100

1100110110
1101002000

0011000000

0102101010

0110100001
0120000002
1010100001

01001-1111

1120000002

201111101
0000-1
0000-1
2001101000
1000-1

1002100110

0110000012

3001101001

0012000001

0100000000

2000130110

0010100012

3001111000

0102-00

2121000000

0111001001
0112130010

0100010010

3011000220

0010100111

0000000110
3121100110

3111000010
2000130111
2000130111
2000130111
2000130111
2000130111
2000130111
2000130111

0102-00

4121001000
2001101000

0111020011
0000010000

2101100111
1011010000
1011010100
1011022100
1011022100
1011020100
0011010100
1011010000

1202101011

1000130010

0102010001
-21100

1011032000
1012032000
-000000-10

2001101000

0000010000

0010000010
1110010111
1110010110
1010010100
1010010100
1010010111
1110010111
1110010111

0010001002

2001111000

2110010101

1101100010

1011001010

0000000000

0100-00

0111020001

1110110111

2011000110

1111100012
1111100012
0100001100
0010001101
0000100011
0011100012
1111100012

200110101

2101111100

-10000

1101020210

200111111

-20210

200110101
200110101

-20010
-10000

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