The Phylogenetic Relationships of Lembocarpus and Goyazia (Gesneriaceae) Based on ndhF Sequences

9 Pages·2001·6.5 MB·

by  James F Smith

THE PHYLOGENETIC
James Smith 2
F.
OF
RELATIONSHIPS
AND
LEMBOCARPUS
GOYAZIA
BASED ON
(GESNERIACEAE)
SEQUENCES
ndhF
1
Abstract
Goy
known. Based on
sequences. Both genera are currently classified in the tribe Gloxinieae, but both are poorly its floral
may
Lembocarpus be placed Gloxinieae,
Goyazia Gloxinieae not controversial. in
morphology, the classification of in is
number
Kpiscieae, or Sinningieae. The acaulescent, tuberous nature of lembocarpus limits the of characters available
Phylo-
morphological analysis and has made classification and phylogenetic relationships difficult to resolve.
for a its
genetic analyses of ndhF sequences place both genera in Gloxinieae. Although the affinities within the tribe are
The Lembocarpus, and an additional
ambiguous for Goyazia, Lembocarpus is sister to Capanea. addition of Goyazia,
species of Capanea provide better resolution of relationships within Gloxinieae and Gesnerieae than had been obtained
previously from parsimony analysis. A maximum likelihood analysis is largely congruent with the parsimony tree.
Lembocarpus, ndhF.
Key words: cladistics, Gesneriaceae, Gloxinieae, (royazia,
Goy>
and phylogenetic analyses within
Classification
Gesneriaceae, particularly the neotropical subfam- inieae on the basis of its scaly rhizome, annular
much and shape.
Gesnerioideae, have received attention, nectary, corolla
ily
placement Le
numerous relationships within genera Goyazia, the of
revealing
& drawn
(Smith Sytsma, 1994a, b, c; Smith, 1994; Kvist within Gloxinieae (Wiehler, 1983) has criti-
& known, monotypic
Skog, 1988, 1989, 1993, 1996; Kvist, 1990), cism. Lembocarpus is a poorly
& & Amer-
within tribes (Smith Carroll, 1997; Smith At- genus that was once in cultivation in North
The an
kinson, 1998; Smith, 2000a, b), and among tribes ica, and apparently is no longer. plant is
& produces
Wiehler, 1995; Smith, 1996, 2000c; Smith acaulescent tuberous perennial that a sin-
(Burtt
and and inflorescence each season (Wiehler,
1997a, However, the classification gle leaf
et b).
al.,
phylogenetic relationships of numerous genera have 1983). The ovary is superior and the nectary is an-
(W
Among 1983). In his
remained unexamined. these genera are
U
treatment of the Gesneriaceae of the Guianas,
Lembocarpus be
Leeuwenberg considered to
Goyaz (1958)
Guianan endemic Rhoogeton
Goyazia similar both the
to
Le
or well known and the plant is not in cultivation in
American European gardens, the place- Gloxinieae).
North or
Lembocar-
ment genus Gloxinieae (Wiehler, 1983) Wiehler (1983) was the first to place
of this in
annular nec-
pus Gloxinieae on the basis of the
Goyazia in
Wiehler
Goias and Mato Grosso prov- tary and tuberous habit. Additionally,
Planalto of Brazil in
Le
The plant a creeping saxicolous perennial
inces. is
with slender stems, scaly rhizomes, and small or- ningia Nees (Gloxinieae sensu Wiehler, 1983) as
Lembocarpus Gloxinieae.
bicular-ovate leaves. Flowers are borne singly in additional support for in
and
the leaf axils and are structurally the same as those Several Sinningia species have tubers nearly
members which added further support for
Achimenes and other of Gloxi- superior ovaries,
of Pers.
was provided by NSF grant DEB-9317775, a grant from the American Gloxinia and
•Support project
for this
Research Associates Program. thank the following
Gesneriad Society, and a grant from Boise State University Faculty 1
L
Skog and Boggan of the Smithsonian Institution, K. Dunn. I). Turley, R. Stewart,
providing leaf material: E.
for J.
and M. Peixoto. also thank M. Peixoto and A. Chautems for assistance in Brazil, and two anonymous reviewers for
I
comments.
helpful
Idaho 83725, U.S.A.
1910 University Drive, Boise,
2 Department of Biology, Boise State University,
[email protected]
Gard. 135-143. 2001.
Ann. Missouri Bot. 88:
136
Annals
of the
Missouri Botanical Garden
Table Species sequenced GenBank
1. in this study with submission numbers and voucher specimens.
Letters
i
parentheses indicate herbaria where vouchers
are deposited.
GenBank
Species
Collection and voucher number
Achimenes
skinneri Lindl.
Cultivated (US) U62177
Anodiscus xanthophyllus (Poepp.) Mansf. Dunn
Peru. (SRP) AF040143
s.n.
onia
Bell sp. Dominican
Republic. Evans (SRP) AF040144
s.n.
Capanea
Fritsd
Colombia.
affinis Risaralda: Mpio. Santuario, Apr. AF206201
1
&
Amaya
1998, M. Smith 393 (COL)
C. grandiflora (Kunth) Decne. ex Planch
Venezuela. Merida: Sierra Nevada, Mucuy AF040145
re-
26
gion, Jan. 1989, Smith 1199 (WIS)
Diastema
racemiferum Benth.
Cultivated. Skog 7574 (US) U621S6
Eucodonia
andrieiixii (DC.) Wiehler Dunn
Cultivated. (SRP) AF040146
n.
s.
Gloxinia sylvatica (HBK) Kunth Cultivated. Dunn 9012051 (SRP) U62157
Goyazia
rupicola Taubert
Brazil. Goi^s: 17 Jan. 1998, Smith 3722 AF257485
et al.
(SRP)
Heppiella ulmifolia (Kunth) Hans!
Kcuador. Napo: near Baeza, 4 Apr. 1996. Smith AF040147
3427
(SRP)
Koellikeria erinoides (DC.) Mansf. Dunn
Brazil. (SRP) A FO 13709
s.n.
Kohleria spicata (Kunth) Oerst. Cultivated. Skog 7701 (US) U6218I
Ijembocarpus amoemis Leeuw.
French Guiana. Haut Oyapock, AF257486
Mt. Marcel,
St.
4478
21 Mar. 1976, Sastre (US)
Monopyle maerocarpa
Benth.
Cultivated (US)
U62197
Moussonia
strigosa (C. V. Morton) Cultivated. Dunn (SRP) AF040148
s.n.
Niphaea oblonga
Lindl. Mexico. Skog 7564
(US) U62160
Pea cea sp
r< Kcuador. Napo: near Baeza, 4 Apr. 1996, Smith AF040149
3425
(SRP)
P. hypocyrtiflora Regel South America. 3943
Cultivated. Smith (SRP) AF040150
Phinaea Kusby
albiflora
Cultivated (US) AF040151
&
Sanango
racemosum
(Ruiz Pav.) Barringer Kcuador. Wiehler (GRF)
U62144
Smithiantha cinnabarina (Linden) Kuntze
Cultivated (US) AF040152
&
Solenophora Denham
obliqua
D. L. D. N. Gib- Mexico. 71542
Breedlove (CAS) U 62202
sou
Gesnerieae
Gesneria
pedicellaris Alain Skog 7722
Cultivated. (US) U62192
Gesneria Urban
christii
Cultivated (US) U62191
Rytidophyilum tomentosum
(L.) Mart Cultivated. Skog 5364 (US) U62200
Rytidophyllum auriculatum Hook.
Cultivated (US) U62199
Episcieae
Alloplectus pan a mens Morton
is C. V. Panama. 28 Oct. 1993, Skog 7641 (US) A F0 3685
et al.
1
Allopleetus
sp.
Kcuador. Napo: San Rafael Falls, 4 Apr. 1996, A F0 13686
3418
Smith (SRP)
&
Alsobia Moore
dianthiflora (H. K. G. Wil- Cultivated. Skog 7969
\\. (US) A FO 13687
son) Wiehler
A. punctata (Lindl.) Wiehler
Mexico. Chiapas: Ocozocautla, winter 1991- A FO 3688
1
3600
1992, Smith (SRP)
Alsobia
sp.
Cultivated (US) A FO 13689
Alsobia
sp. 3599
Cultivated. Smith (SRP) A FO 13690
Ghrysothemis
friedrichsthaliana (Hanst.) H. K. Cultivated. Skog 7992
(US) A FO 13691
M
oore
Cobananthus
calochlamys
D. Sm.) Wiehl
(J. er Cultivated (US) A FO 13692
Codonanthe
elegans Wiehler
Belize. San Jose: Skog 5699 (US) U62178
Codonanthopsis peruviana Wiehler
Cultivated. Turley (SRP) A FO 13693
s.n.
Columnea ambigua
(Urban) Morley
Puerto Rico. Smith 3701 (SRP) A FO 13694
mira Morley
C. Panama. Smith 2450 (WIS) A FO 13695
Rushy
C. oblongifolia
Peru. Cuzco: Urubamba, Machu A
Prov. Picchu, 3 FO 13696
Apr. 1989, Smith 1721 (WIS)
137
Volume Number Smith
88,
1
Lembocarpus and Goyazia
2001 Relationships of
Table Continued.
1
.
Gen Bank
number
and voucher
Collection
Speeies
A
Smith 3369 (SKP) FO 13697
sanguined Hanst. Cultivated.
(Pers.)
C.
U62164
from Xalapa Mis-
Mexico. Vera Cruz: road to
schiedeana Sehlecht.
C.
Smith 288 (WIS)
santla,
A
Near Limon, 15 May 1994 no voucher FO 13698
Wiehler Ecuador.
Corytoplectus speciosus (Poepp.)
AF013699
Wiehler Ecuador. Napo: San Rafael Falls, 4 Apr. 1996,
D. urceolata
Smith 3416 (SKP)
AFO
13700
3947
Smith (SEP)
Cultivated.
Episc ia fimbria a Fritsch
t
AF013701
French Guiana. Mte. des Nouragues, 16 June
I^euw.
sphalera
E.
94-079
al (US)
1994, Feuillet
et
AF013702
NautUocalyx adenosiphon (Leeuw.) Wiehler Cultivated. Skog 7847 (US)
AF206197
Smith al 3726 (SRP)
Nematanthus allms Chautems, ined. Brazil. 17 Jan. 1998, et
AF206198
Mpo. 17 1998,
Hoehne Sao Paulo: Espiritu, Jan.
Brazil.
N. fritschii
3720
Smith (SRP)
et al.
AF013703
num Wiehler Cultivated. Smith 3944 (SRP)
Neomortonia miliaria (Hanst.)
AFO
13704
Wiehler Cultivated (US)
N. rosea
AF206199
Macho, 20
Panama. Bocas Cerro Pate
Wiehler del Toro:
Oerstedina cerricola
Hammel 5754
Nov. 1978, (US)
AFO
13705
Skog 7979 (US)
Paradrymonia aurea Wiehler Cultivated.
AF013706
R (SRP)
densa H. Wright) Wiehler Cultivated. Stewart s.n.
(C.
AF013707
fuquaiana Wiehler Ecuador. 6 Jan. 1990, Skog 7889 (US)
P.
AF2O620O
Kaieteur
Guyana. Potaro-Siparuni region, Falls,
Rhoogeton Lwbg.
viviparus
370
18 Oct. 1987, Kvist et al, (US)
AFO
708
Skog 7761 (US) 13
major Wiehler Cultivated.
Rufodorsia
Outgroups: Sinningieae
U62174
5399
Skog (US)
Paliavana prasinata (Ker-Gawl.) Fritsch Brazil.
& U62 75
Dunn 9104014 (SBP)
Wieh-
Sinningia brasiliensis (Regel Schmidt) Brazil. 1
er
1
U62201
7808
Skog (US)
Wiehler Cultivated.
Sinningia cooperi (Paxt.)
U62186
Cultivated (US)
Sinningia Clayb.
richii
U62203
Skog 7690 (US)
Cultivated.
Vanhouttea lanata Fritsch
many
based on charac-
the placement of Lembocarpus in Gloxinieae. Re- in Episcieae is largely the
hybrids were apparently premature; no hy- ters shares with Rhoogeton: superior ovary, tuber,
ports of it
acaulescent
between these genera have been documented calyx venation, inflorescence structure,
brids
and ovules only on outer placental surfaces
habit,
(Boggan, 1991).
Lemb (Leeuwenberg, 1958). The superior ovary and tu-
unknown
Gloxinieae, but are
based on seed berous habit are in
Gloxinieae sensu Fritsch (1894)
common annular nec-
characteristics, noting that the seed shape was sim- in Episcieae. In contrast, the
Gesneneae
some species of Episcia Mart, (tribe Epis- tary is known outside of Gloxinieae and
ilar to
cieae) but not like any species of Gloxinia. Thus only in Beslerieae, a tribe with several other defin-
would exclude the placement of
seed shape implies an affinity to Episcieae. Beau- ing characters that
among
more widespread
noted the shape of the Lembocarpus. Tubers are
fort-Murphy (1983) also that
was Smithiantha members of Sinningieae (sensu Smith et al., 1997b)
cells of the seed coat similar to
member Beaufort-Murphy than elsewhere in Gesnerioideae, implying that
Kuntze, a of Gloxinieae.
among
Lembocarpus Lembocarpus may best be placed these gen-
due Tubers also are known from members of Ep-
isolated position within Gesnerioideae to its era.
(Wiehler, 1983) but not other Gloxinieae
anomalous combination of seed characteristics. iscieae
Smith 1997b).
summary, morphological data support the (sensu et al.,
In
The Gloxinieae has received recent atten-
Lemb tramus either in Gloxinieae or tribe
phylogenetic relationships
The Gloxinieae based on tion with regard to the
Episcieae. similarity to is
&
However,
and genera (Smith Atkinson, 1998).
campanulate of
th e annular nectary, corolla, its
Lemb
placement Goyazia nor
shape of the cells of the seed coat. Its
138
Annals
of the
Missouri Botanical Garden
Monopyle
2(17)
Solenophora
15 Gl
Diastema
_
Sanango
1(13) Ge
1(12)
34
Phinea
Pearcea
55
Pearcea
(6)
Gloxinia
8(26)
Niphaea
um
Eucodonia
Gl
Achimenes
Kohleria
1(10)
Lemboca
rpus
11(27)
2J13)
33
2 Capanea
affinis
Heppiella
Moussonia
Anodiscus
,fil L _
2J9)
2 (15)
I Koellikeria
I
1
I Goyazia
3(8)
Gesneria
20
christii
(26)
8
5(15)
—
Gesneria
pedicellaris
Ge
29 —
Rytidophyllum
auriculatum
—
Rytidophyllum tomentosum
—
3 —
(12)
Bellonia
Gl
39
—
Smithiantha
^__
™
Alsobia
1(12) sp.
Alsobia
sp.
2(10 Alsobia dianthiflora
Cobananthus
calochlamys
(9)
Alsobia punctata
Episcia sphalera
'2(15)
51
Episcia fimbria
ta
Columnea
ambigua
(8)
Columnea
sanguinea
Columnea
mira
Alloplectus
sp.
1(4)
panamens
Alloplectus
is
17
Drymonia
1(11)
L(M
-
Neomortonia
rosea
Columnea
schiedeana
(8)
f
Columnea
oblongifolia
L(?}„
r
Neomortonia nummularia
i
Ep
Corytoplectus
1(11)
Rhoogeton
Nematanthus
albus
Nematanthus
fritschii
9(24)
Oerstedina
99
Rufodorsia
Paradrymonia
1 1 (24) fuquiana
100
(15)
1 Paradrymonia
aurea
i
L Capanea
grandiflora
Gl
Chrysothemis
2(12)
Nautilocalyx
Paradrymonia Ep
densa
Codonanthopsis
1 1 (30)
Codonanthe
elegans
73
_^__
1(23)
Paliavana
39
Vanhouttea
Si
1(24) Sinningia
richii
Sinningia
brasiliensis
Sinningia cooperi
139
Volume Number Smith
88,
1
Lembocarpus and Goyazia
2001 Relationships of
may
previous cladistic analyses of molecular data due carpus have affinities (Gloxinieae, Episcieae,
and Taxonomic choice derived from
the lack of leaf material. This study presents Sinningieae).
to
results regarding the phylogenetic position of these previous tribal analyses with Episcieae sister to
genera within the neotropical Gesneriaceae. Gloxinieae/Gesnerieae but Sinningieae sister to
The
these three tribes (Smith et al., 1997b). data
1.54%
Mkthods missing
Materials and matrix taxa contains cells
for all
based on sequence alignments.
total
and Genbank numbers
Voucher information for
sequences used analysis are presented
in this
all
PHYLOGENETIC ANALYSIS
DNA
was from
Table Goyazia isolated
in for sil-
1.
and Phylogenetic divergence was reconstructed using
ica gel dried material (Smith et al., 1992), the
PAUP maximum
ndhF gene was amplified in two overlapping sec- version 4.0d64 to implement par-
DNA
dons (positions 1-1350, and 972-2044). for simony (MP) (Farris, 1970; Farris et al., 1970;
&
maximum
Lembocarpus was obtained from herbarium speci- Swofford Maddison, 1987) and likeli-
mens (Savolainen 1995) using the DNEasy hood (MLE). In this study, trees were generated us-
et al.,
miniprep (Qiagen) following the manu- ing the general heuristic option. To search for is-
Plant kits
The ndhF gene Lembo- lands of equally parsimonious trees (Maddison,
facturer's instructions. for
Olmstead and Palmer
carpus was amplified two overlapping fragments 1991), the search strategy of
in
using primers 172-1350R and 972-2044R. The (1994) was implemented searching for 1000 trees
each subsequent analyses with the nearest
fragment smaller than those reported pre- in five
first is
viously (Smith et 1997b) since amplifying the neighbor interchange (NNI) search option in effect
al.,
DNA
from herbarium specimens required succes- and mulpars "off." Each of the results from the five
NNI
sive amplifications using internal primers. Initial searches was used as the starting tree(s) for a
DNA
(TBR) and
reconnection
followed procedures described search with tree bisection
amplifications
was used
elsewhere (Smith 1997b) using primers and mulpars "on." This search strategy for all
et al., 1
MP
2044R
1350R and 803 and the analyses.
the part for
for
first
MLE
second Subsequent amplifications required Because of the greater time involved in
part.
the use of 172 and 972 as forward primers, al- analysis, a smaller sampling was utilized. In this
though the same reverse primers resulted in sue- analysis, a full sampling of Gloxinieae was used
and
and one species each of Gesneria L. Rytido-
cessful amplifications.
Ep-
The was on Gloxinieae phyllum Mart. (Gesnerieae). Representatives of
focus of this analysis
since both Goyazia and Lembocarpus are currently iscieae were used as the outgroup with taxa select-
&
Lembocarpus
classified in this tribe (Burtt Wiehler, 1995). ed to include possible relatives of
MLE
However, since there considerable question re- (most notably Rhoogeton). trees were gener-
is
TBR
Lembocarpus, ated using the heuristic search option with
garding the tribal position of initial
MLE
Under Has-
analyses were conducted using representatives of and mulpars "on." the option, the
egawa model was used, which allows
tribes of the neotropical subfamily Gesnerioi- et al. (1985)
all
and
Members Old World Epithematae unequal nucleotide frequencies differential
deae. of the tribe for
The assumed
were used as the outgroup because some prelimi- rates for transitions and transversions.
nucleotide frequencies were estimated from the
nary results implied a potential relationship to this
= = G =
A C
0.27809, 0.15598, 0.17529,
tribe. Results from this preliminary analysis (not data:
MP
= MLE
T compared
more taxon sam- and 0.39063. trees were to
included) allowed for a restricted
&
Kishino-Hasegawa (Kishino
pling that would permit greater analytical flexibility trees using the test
MP
as well as minimizing homoplasy in the data set. Hasegawa, 1989). Additional trees were gen-
Subsequent analyses used only Gloxinieae, Ges- erated using the same search criteria described
MLE
same
nerieae, and Episcieae as well as Sinningieae as above with the taxa utilized in the anal-
The reduced
outgroup. choice of taxa for the anal- ysis.
where Lembo- Branch support analysis was performed to ex-
considered possible tribes
ysis all
= =
Figure One of 48 maximum parsimony trees of 3998 steps each, CI 0.38, RI 0.33. These trees were rooted
1.
using Sinningieae. Numbers above clades are deeay values; numbers in parentheses are branch lengths using the
aeetran option of PAUP. Numbers below clades indicate bootstrap values. Terminal branch lengths are not shown.
48 Abbre-
Branches shown with dashed lines indicate nodes that are not supported in a strict consensus of all trees.
= = = =
Ep Episcieae, Ge Gesnerieae, Gl Gloxinieae, Si Sinningieae.
viations for tribes are:
140
Annals
the
of
Missouri Botanical Garden
amine
trees that were one or two steps longer than Capanea
to as indicated by parsimony. Further,
the most-parsimonious trees (Bremer, 1988, 1994; Goyazia
is sister to a clade containing Moussonia
Donoghue
Clades
et al., 1992). that persisted in Kegel, Anodiscus Benth., and Koellikeria Regel
MP
consensus two beyond An
strict trees steps the most- same
(Fig. 2). analysis of the taxa used in
MLE
parsimonious were examined
trees using the con- the analysis resulted in two islands for a total
straints option search the 2076
to for shortest tree that of four trees of steps each (trees not shown).
did not contain that clade. Bootstrap analysis The Kishino-Hasegawa &
(Fel- (Kishino Hasegawa,
test
senstein, 1985) was performed using 100
replicates 1989) did not indicate a significant difference be-
TBR
with and mulpars "off and "on." Because tween MP and MLE
the
trees.
there substantial morphological and biogeograph-
is
ic data that imply a relationship between Lembo-
DISCUSSION
carpus and Rhoogeton, the constraints option was
GOYAZIA
to ass the most-parsimonious tree with
1
these genera as a monophyletic group. The
trees
The
results of this study support the placement
generated from the constraints anal.ys«is.we.re,com- „r r\ ' /r^^ o\^ wz-'W"
r»
°* Goyazi;a i• n Gloxi• ni• eae (Figs. i Wiehler
1, 2).
.
pared, to t,he unconstrained trees using& the Kishino- nQQQ\ ^«« *l^ £™» r u n\
' (19*5.5) was the hrst to pll ace Goyazi•a i• n hi• s Gloxi•-
&
Hasegawa .
(Kishino Hasegawa,
test 1989).
nieae, where has remained mod
in a revised
it
&
version of the classification system (Burtt Wieh-
Results
ler, 1995). Although Goyazia is always in a clade
with Moussonia, Anodiscus, and
Koellikeria, ex-
s
it
Ihe members
initial analysis utilized all of the
.l
*
^ ... ^ act reli a4t-ionship to these ot*iher genera • unresoli vedi
™r is
Gesnenoideae wi.th, t,he O/-lm d, Worli_di tr.i,be EpFith, e- n n w n
(n?r-ig. One ofr ftUhe t. rees ffrom 4t.he MP anal.ysi.s iJ?.
1). (Fig.
. . MP
matae as the outgroup (not sh, own). The analysi.s
Grr>
V
ed 5128
in 14 trees of steps each (trees not
*„ u *u o\ j h-i
n to #these th ree genera (/ir?i-g. and Koellikeri• a •
2), is
s,hown)x. tIn all ofr tihese trees bi otih Lr embiocarnus and
.,
. , .
Goy
Goy
and
azia Koellikeria are both found
in Brazil,
reduced data resulted 48 3998 M
set in trees of steps n
r '
whi ereas Moussoni•a • end,emi.c Cent.rmal, AAmerica,
is to
r ,. rr . .
each, from t,hree different i. slands most-parsimo- aa
of r
r „ an nd Anodiscus is native *to t*uhe Aandjes of Peru. cS-ince
=
nious trees, with consistency index
(CI) 0.38,
the majority of Gloxinieae are found west of the
=
and One
retention index (RI) 0.33. of these trees
Andes,
the most-parsimonious explanation
for this
is presented in figure with dashed lines
1 to in- u
j; -
dist, ributf ion wouli di ibe t. wo separat.e migrati. ons
to
. , u
j. 11 •
dicate clades that collapse the consensus
in strict
Goyazia
more
ot all trees. his tree is ,roesoilvoed*t,ha.n prior _
I *•
„ reli ationsihi• ps among tA ihese genera are accurate as
,
., , &
results from ndhY analyses (Smith Atkinson, u
F
• -
igure 2. However, weaki iboot*st* rap andi dje-
ooq\ tu u
1i 998) l i_ Gr*oyazia and\ Lie
cay index support herein for these relationships im-
bocarpus
within Gloxini although
the position of
i- .1 * more j l r
° / plies that dat* a are necessary before concliu-
.
Goyazia Lemb .
not resolved;
is
sions regarding the biogeography of these taxa can
Capanea
Decne. ex Planch. As demonstrated
in l
i
&
simulation (DeBry
studies Olmstead,
2000), the
bootstrap values with mulpars "on"
"off" or differed
LEMBOCARPUS
only by a few points (data not shown). The
val
on Figure are from the analysis with mulpars Goy
1
of analysis con-
s this
"off."
Lembocarpus
An
additional 14 steps beyond the most-parsi-
(Figs. This placement more
1, 2). is controversial
Lemb
since early descriptions of Lembocarpus noted
its
and Rhoogeton
into a monophyletic group
(not disparate calyx venation, inflorescence, acaulescent
Is
habit, outer ovule placentation, superior ovary, and
Rhoogeton
together are Nematanthus
sister to tubers (Leeuwenberg, 1958) as shared with Rhoog-
Schrader The
(Episcieae). Kishino-Hasegawa
test eton (Episcieae).
&
(Kishino Hasegawa,
1989) did not indicate a sig- Placement of Rhoogeton Episcieae
in is in itself
nificant difference between the constrained and un- controversial. Although morphological data and
constrained
trees.
2000b)
MLE
analysis resulted in a -In likelihood of for its placement there, ITS sequences imply
its
118.64716
Goyazia and Lembocarpus
(Fig.
2). position outside of Episcieae and
possibly within
both in Gloxinieae, and Lembocarpus
Gloxinieae
is sister (Smith, 2000b). Unfortunately, ITS
se-
5
Volume Number Smith 141
88,
1
Lembocarpus and Goyazia
2001 Relationships of
Monopyle
Solenophora
Gl
Diastema
_
Sanango
Ge
Phinea
Gloxinia
Niphaea
Eucodonia
Achimenes
Kohleria Gl
Lembocarpus
.
Capanea
afhnis
Heppiella
Pearcea
Pearcea
Moussonia
Anodiscus
Koellikeria
Goyazia
Gesneria
christii
Ge
Rytidophyllum auriculatum
Bellonia
Gl
Smithiantha
Nematanthus
fritschii
Codonanthe
elegans
Rhoogeton
Episcia fimbriata
Alsobia dianthiflora
Rufodorsia
Paradrymonia fuquiana
Columnea
schiedeana
=
Maximum 16118.M716. To minimize computer lime this
estimate likelihood
Figure likelihood tree, -In
2.
=
= Ge
Ep
used only representative taxa of Episcieae as outgroups. Abbreviations for tribes are: Episcieae,
analysis
=
Gesnerieae, Gl Gloxinieae.
142
Annals
of the
Garden
Missouri Botanical
DNA
quences could not be obtained using from
the unite these genera with the exception that they are
herbarium material used Lembocarpus among
for in this the few genera within Gloxinieae that lack
study. Despite the several morphological character scaly rhizomes. The placement Capanea
of Glox-
in
states shared between lembocarpus and Rhoogeton, inieae contradicts earlier cladistic analyses of ndhF
14 additional steps beyond the most-parsimonious sequences of Gloxinieae where was placed Ep-
in
it
&
trees are necessary to place Lembocarpus and iscieae (Smith Atkinson, The
1998). earlier anal-
Rhoogeton in a monophyletic group using ndhF se- ysis based on ndhF sequences only used Capanea
quence
Thus,
data. regardless of the tribal affinity grandiflora (Kunth) Decne. ex Planch., and Capa-
of Rhoogeton, it does not appear to be closely re- nea ajfinis is added to the present analysis. The
lem
b
source of discrepancy as yet unknown, but based
is
Lem
on results of ITS and ndhF sequences
(Smith,
bocarpus Gloxinieae comes from and
in the corolla 2000b), Capanea
clear that belongs
it is ajfinis in
seed. Although Beaufort-Murphy hypothe-
(1983) Gloxinieae.
may
sized that Lembocarpus have an isolated po-
sition within the Gesnerioideae due to a unique SUMMARY
combination
of characters, combination seed
this of
The ndhF
surface characters and superior ovary data presented here confirm the place-
best
is
ment
viewed as autapomorphic. The shape of both Goyazia and Lembocarpus within Glox-
of the cells of
Lembocarpus
the seed coat and campanulate corolla may be syn- inieae. is strongly supported by decay
Le index, moderate (relatively) bootstrap values, and
ae
.
MLE
Lem boc nnus analysis as sister to Capanea. Goyazia is
and Gloxinieae nectary- shape. Whereas the an- placed in a clade with Moussonia, Anodiscus, and
is
MP
MLE
nular nectary widespread Koellikeria in both and analyses, although
Gloxinieae,
is in also
it
^
is known from tribes Gesnerieae and Beslerieae. In e relationships among these four genera are not
MP
resolved Because ndhF seems
cladistic analyses of Gloxinieae and Gesnerieae falty in trees. to
^
based on ndhF sequences, 3e at e limits of resolving power within Glox-
the position of Gesner- its
'
&
ieae always creates a paraphyletic Gloxinieae inieae (also observed in tribe Episcieae; Smith
(Figs.
&
1, 2; Smith Atkinson, 1998). This implies that Carroll, 1997; Smith, 2000b), it will be essential in
if
not derived from within Gloxinieae, the Gesnerieae future investigations to add additional sequence
are closely related, and the annular nectary data to resolve more fully the intergeneric rclation-
like-
is
ly a synapomorphy for both tribes Gloxinieae and ships of this tribe.
The
Gesnerieae. Beslerieae are more distantly
re-
lated to Gloxinieae based on previous phylogenetic
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