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Reassignment of Species of Paraphyletic Junellia s. l. to the New Genus Mulguraea (Verbenaceae) and New Circumscription of Genus Junellia: Molecular and Morphological Congruence

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  • Museo de Ciencias Antropológicas y Naturales (UNLAR) - Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET)

Abstract and Figures

Verbenaceae tribe Verbeneae includes three major genera, Verbena, Glandularia, and Junellia, which form a recently diversified group. Junellia is a South American genus, whereas Verbena and Glandularia are distributed in temperate regions of both South and North America. Seven noncoding chloroplast regions were sequenced including intergenic spacers and/or introns in trnD-trnT, trnS-trnG, trnS-trnfM, trnT-trnL, trnG, trnL, and trnL-trnF; the nuclear ITS and ETS regions were also sequenced. Together with previous studies, these results suggest that Junellia, as traditionally conceived, is a paraphyletic group of two separate clades. Junellia should be restricted to the clade containing the type, J. micrantha, which also includes Glandularia subgenus Paraglandularia and the genus Urbania. Consequently Urbania and Glandularia subgenus Paraglandularia are reduced to synonymy under Junellia, and eight new combinations in Junellia are proposed: Junellia ballsii, J. crithmifolia, J. fasciculata, J. hookeriana, J. lucanensis, J. occulta, J. pappigera and J. origenes. The remaining Junellia species form a monophyletic group here designated as the new genus Mulguraea, with 13 new combinations: Mulguraea arequipense, M. asparagoides, M. aspera var. aspera, M. aspera var. longidentata, M. cedroides, M. cinerascens, M. echegarayi, M. hystrix, M. ligustrina var. ligustrina, M. ligustrina var. lorentzii, M. scoparia, M. tetragonocalyx and M. tridens. Verbena and Glandularia s. s. are both monophyletic based on ETS/ITS data, but neither is monophyletic based on cpDNA data. Relationships within each genus are still not wholly resolved, nevertheless there is evidence that South and North American Verbena might both be monophyletic. Verbena and Glandularia are sister groups, and together they are sister to the reconstituted Junellia. Mulguraea is sister to the group comprising all the three genera, Verbena, Glandularia, and Junellia.
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Systematic Botany (2009), 34(4): pp. 777–786
© Copyright 2009 by the American Society of Plant Taxonomists
777
In the most recent classification of Verbenaceae ( Atkins 2004 )
tribe Verbeneae Schauer comprises five genera, Verbena L.
with 44 species ( O’Leary et al. 2007a ), Glandularia J. F. Gmel.
with 84 species ( Peralta 2009 ), Junellia Moldenke with 39 spe-
cies ( Peralta et al. 2008 ), and the monotypic Urbania Phil. and
Hierobotana Briq. These genera, often combined under Verbena
in early treatments ( Schauer 1847 ; Briquet 1895 ; Perry 1933 ;
Troncoso 1974 ; Table 1 ), constitute a recent and rapidly diver-
sifying group, referred to as the “ Verbena complex” ( Yuan and
Olmstead 2008a , b ). All Verbeneae share a dry schizocarpic
fruit, separating at maturity into four 1-seeded mericarps.
Most species of this tribe are found in temperate North or
South America.
Junellia is distinguished by its shrubby habit, woody root-
stock, and reduced, sometimes spiny, leaves, and a base chro-
mosome number of x = 10, rarely x = 9 ( Botta and Brandham
1993 ). Glandularia and Verbena are mostly leafy herbs, ocas-
sionally subshrubs, without woody rootstocks or spines.
Verbena differs from Glandularia principally by its base chro-
mosome number of x = 7 vs. x = 5 ( Schnack and Covas 1944 ),
as well as several morphological and anatomical characters
( Schnack and Covas 1946 ; Schnack 1964 ; Umber 1979 ; Botta
1989 ; O’Leary et al. 2007b ).
Junellia is a South American genus, distributed along the
arid Andean region, from Perú, Bolivia, Chile, and Argentina,
with the greatest species richness in the Andean-Patagonian
domain ( Cabrera and Willink 1973 ). Verbena and Glandularia
are distributed in temperate regions of both South and North
America, with several species naturalized in the Old World.
Yuan and Olmstead (2008a , b ), using cpDNA and the nuclear
genes waxy , PHOT1, and PHOT2 sequence data, showed that
both Glandularia (excluding subgenus Paraglandularia Schnack
& Covas) and Verbena are monophyletic, with base chromo-
some numbers of x = 5 and x = 7 as putative synapomorphies,
respectively. These studies also provided initial evidence that
Junellia , as currently circumscribed ( Botta 1989 ; Peralta et al.
2008 ), is not monophyletic. The main objective of this study is
to extend sampling of Junellia to provide a solid phylogenetic
framework for generic realignments.
Materials and Methods
Taxon Sampling— For phylogenetic analyses, 70 taxa of the Verbena
complex were included; all but 20 of these taxa had been used in a previ-
ous study ( Yuan and Olmstead 2008a ), where detailed voucher informa-
tion can be found. Information on the 20 new taxa is in Appendix 1 . For the
cpDNA analyses, trees were rooted using Lippia salsa and Aloysia virgata as
outgroups; both belong to tribe Lantaneae, sister group of tribe Verbeneae
(Marx and Olmstead, unpubl. data). There is too much ETS/ITS sequence
divergence between Aloysia/Lippia and the Verbena complex to permit an
unambiguous alignment, and thus, the use of Aloysia/Lippia as outgroups.
Subsequently, the ETS/ITS trees were rooted using the “ Junellia II” group
( Yuan and Olmstead 2008b ) as a functional outgroup, based on previous
studies on the Verbena complex ( Yuan and Olmstead 2008a , b ) and ongo-
ing work towards resolving the phylogeny of the Verbenaceae family
based on cpDNA (Marx and Olmstead, unpubl. data) and five nuclear
pentatricopeptide (PPR) genes (Yuan et al. 2009). This “ Junellia II” group is
treated as a new genus, Mulguraea , in this study (see below).
DNA Extraction, Amplification and Sequencing— The material used
for DNA extraction was taken from plants collected from wild populations
and dried in silica gel or from herbarium specimens (taxa with asterisk in
Appendix 1 ). For leaves dried in silica gel, DNA extraction was performed
using a modified CTAB protocol by Doyle and Doyle (1987 ), adapted for
small amounts of plant material. For herbarium material, DNA extrac-
tion was conducted using the DNeasy plant mini kit (Qiagen, Hilden,
Germany). Seven noncoding chloroplast regions were amplified. The
trnS-trnG spacer and trnG intron were generally amplified together as the
trnS-G fragment, using primers trnS ( Shaw et al. 2005 ) and trnG(V) ( Yuan
and Olmstead 2008a ). When this was not possible internal primers trnG-S
F(V) and trnG-S R(V), designed by Yuan and Olmstead (2008a ), were used
to amplify the region in two fragments. The trnT-trnL spacer, trnL intron,
and trnL-trnF spacer were amplified together as the trnT-F fragment using
primers “a” and “f” ( Taberlet et al. 1991 ). When this was not possible inter-
nal primers “b” and “c” ( Taberlet et al. 1991 ) were used to amplify the
region in two fragments. The trnS-trnfM region was amplified using prim-
ers trnS and trnfM ( Shaw et al. 2005 ). The trnD-trnT region was amplified
Reassignment of Species of Paraphyletic Junellia s. l. to the New Genus
Mulguraea (Verbenaceae) and New Circumscription of Genus Junellia :
Molecular and Morphological Congruence
Nataly O’Leary, 1
,
4 Yao-Wu Yuan, 2
,
3 Amelia Chemisquy, 1 and Richard G. Olmstead 2
1 Instituto de Botánica Darwinion, Labardén 200, San Isidro, Argentina
2 Department of Biology, University of Washington, Box 355325 Seattle, Washington 98195 U.S.A.
3 Present address: Department of Plant Biology, University of Georgia, Athens, Georgia 30602 U.S.A.
4 Author for correspondence ( noleary@darwin.edu.ar )
Communicating Editor: Mark P. Simmons
Abstract— Verbenaceae tribe Verbeneae includes three major genera, Verbena , Glandularia, and Junellia, which form a recently diversified
group. Junellia is a South American genus, whereas Verbena and Glandularia are distributed in temperate regions of both South and North
America. Seven noncoding chloroplast regions were sequenced including intergenic spacers and/or introns in trnD-trnT, trnS-trnG, trnS-trnfM,
trnT-trnL, trnG, trnL, and trnL-trnF ; the nuclear ITS and ETS regions were also sequenced. Together with previous studies, these results sug-
gest that Junellia, as traditionally conceived, is a paraphyletic group of two separate clades. Junellia should be restricted to the clade containing
the type, J. micrantha , which also includes Glandularia subgenus Paraglandularia and the genus Urbania . Consequently Urbania and Glandularia
subgenus Paraglandularia are reduced to synonymy under Junellia, and eight new combinations in Junellia are proposed: Junellia ballsii ,
J. crithmifolia, J. fasciculata, J. hookeriana, J. lucanensis, J. occulta, J. pappigera and J. origenes . The remaining Junellia species form a mono-
phyletic group here designated as the new genus Mulguraea , with 13 new combinations: Mulguraea arequipense , M. asparagoides , M. aspera
var. aspera , M. aspera var. longidentata, M. cedroides , M. cinerascens , M. echegarayi , M. hystrix, M. ligustrina var. ligustrina, M. ligustrina
var. lorentzii, M. scoparia, M. tetragonocalyx and M. tridens. Verbena and Glandularia s. s. are both monophyletic based on ETS/ITS data, but
neither is monophyletic based on cpDNA data. Relationships within each genus are still not wholly resolved, nevertheless there is evidence
that South and North American Verbena might both be monophyletic. Verbena and Glandularia are sister groups, and together they are sister to
the reconstituted Junellia . Mulguraea is sister to the group comprising all the three genera, Verbena, Glandularia, and Junellia .
Keywords— Glandularia, new combinations, Paraglandularia, phylogeny, taxonomy, Verbena, Verbeneae.
778 SYSTEMATIC BOTANY [Volume 34
using primers trnDF and trnTR ( Shaw et al. 2005 ). When this was not pos-
sible internal primers designed by Yuan and Olmstead (2008a ), trnD2T(V)
and trnT2D(V), were used to amplify the region in two fragments. The
internal and external transcribed spacer regions of nuclear DNA (ITS and
ETS) were amplified using primers ITS-4 and ITS-5 ( White at al. 1990 ),
and primers 18S-IGS ( Baldwin and Markos 1998 ) and ETS-B ( Beardsley
et al. 2003 ), respectively. Polymerase chain reaction and sequencing pro-
tocols followed those of Yuan and Olmstead (2008a ) for their chloroplast
DNA data. Some sequencing reactions were also performed by Macrogen
Inc. using ABI PRISM BigDyeTM Terminator Cycle Sequencing Kits with
AmpliTaq DNA polymerase (Applied Biosystems, Seul, Korea).
The initially generated ITS/ETS sequences display > 1% intraindivid-
ual polymorphic sites in one Junellia species and a few Glandularia species.
For these taxa, the primer pair ITS-4 and ETS-B was used to amplify a
fragment including ETS and the entire 18S, ITS1, 5.8S, and ITS2 regions,
totaling ca. 3 kb. Polymerase chain reaction reactions were performed
using PfuUltra
TM II Fusion HS DNA Polymerase (Stratagene, La Jolla,
California), following the protocol used by Yuan and Olmstead (2008a ) to
generate their nuclear DNA data. After purification, PCR products were
cloned using the TOPO TA Cloning kit (Invitrogen, Carlsbad, California)
and six to eight positive clones were sequenced for the ETS and ITS region
but not the 18S DNA. The cloned ETS and ITS sequences were identified
by clone numbers. All newly generated sequences from this study have
been deposited in GenBank ( Appendix 1 ).
Phylogenetic Analysis— Sequences were aligned manually using the
program Se-Al version 2.0a11 ( Rambaut 1996 ) based on the similarity cri-
terion ( Simmons 2004 ). Alignment matrices were submitted to TreeBASE
study number S2349. Two poly-A/C regions in the trnS-trnfM fragment
(positions 626–643, 686–693) were excluded from subsequent analyses
due to uncertainty of homology assessment. Parsimony-informative gaps
were coded as binary characters using the simple gap coding approach
( Simmons and Ochoterena 2000 ; Graham et al. 2000 ) and were included
in both parsimony and Bayesian analyses. Phylogenetic analyses were
conducted on the combined cpDNA dataset and the combined ITS/ETS
dataset separately. Parsimony analyses were performed using PAUP*
v.4.0b10 ( Swofford 2002 ). Heuristic searches were performed with 500
random stepwise addition replicates and TBR branch swapping with the
MULTREES option in effect. Clade support was estimated by bootstrap
analyses ( Felsenstein 1985 ) using 500 pseudo-replicates. Bayesian MCMC
analyses ( Yang and Rannala 1997 ) were conducted using MrBayes v.3.1.2
( Ronquist and Huelsenbeck 2003 ). A mixed-model approach ( Ronquist and
Huelsenbeck 2003 ) was employed to integrate the parsimony-informative
gaps as binary characters with nucleotide data. Each final dataset (cpDNA,
ITS/ETS) was divided into two partitions, the “DNA” partition and
“gap” partition. The Akaike information criterion (AIC; Akaike 1974 ) as
implemented in Modeltest v.3.7 ( Posada and Crandall 1998 ) was used to
determine the model of sequence evolution that best fit the DNA partition
(TVM + G and GTR + I + G for the cpDNA and ITS/ETS dataset, respec-
tively). The restriction site (binary) model in MrBayes v.3.1.2 ( Ronquist
and Huelsenbeck 2003 ) was used for the “gap” partition, with ascertain-
ment bias for gap characters incorporated (lset coding = informative). We
carried out two independent runs of 1,000,000 generations from a ran-
dom starting tree using the default priors and four Markov chains (one
cold and three heated chains), sampling one tree every 100 generations.
The program Tracer v.1.4 ( Rambaut and Drummond 2007 ) was used to
examine the output parameters from the Bayesian MCMC analyses.
Stationarity was assumed when the estimated value of –lnL and all other
parameters had stabilized. Trees prior to reaching stationarity were dis-
carded as burn-in and the remaining trees from the two independent runs
were pooled to calculate the majority rule consensus tree.
Results and Discussion
Phylogenetic Analyses— Figure 1 represents one of the 12
maximum parsimonious (MP) trees inferred from cpDNA
data and Fig. 2 represents one of the 7,137 MP trees inferred
from ITS/ETS data, respectively. Bayesian analyses gave
very similar results, and none of the few differences was
supported by > 50% bootstrap value (BS) or > 0.95 posterior
probability (PP). With more extensive sampling of Junellia
and Glandularia , the cpDNA tree topology is consistent with
previous results based on cpDNA data ( Yuan and Olmstead
2008a ). Junellia is paraphyletic and can be divided into two
major clades ( Fig. 1 ). The three species from Glandularia
subgenus Paraglandularia are well nested within one of the
Junellia clades. Neither Verbena nor Glandularia (excluding
Paraglandularia ) is monophyletic in the cpDNA tree, a con-
sequence of two potential intergeneric chloroplast transfers
( Yuan and Olmstead 2008a ). Within Verbena , both the South
and North American group are monophyletic ( Fig. 1 ). The
ITS/ETS tree topology is largely consistent with previous
results from nuclear gene data ( Yuan and Olmstead 2008a , b )
in the monophyly of both Verbena and Glandularia (exclud-
ing subgenus Paraglandularia ), and is also consistent with the
cpDNA phylogeny in the paraphyly of Junellia and the posi-
tion of Glandularia subgenus Paraglandularia . In addition, like
the cpDNA data, the ITS/ETS data suggest that both South
and North American Verbena are monophyletic.
Mulguraea a New Genus Segregated from Junellia— A
recent taxonomic treatment of Junellia ( Peralta et al. 2008 )
recognized two subgenera: Junellia and Thryothamnus , each
composed of three sections. Subgenus Junellia is character-
ized by an enlarged style base covering the mericarp apex,
whereas the style base in subgenus Thryothamnus is some-
what inserted between the four mericarps, as is the case in the
rest of Verbeneae. Subgenus Thryothamnus was initially recog-
nized as a genus by Philippi (1895 ) and is distinguished by its
deeply 5-sected calyx and its cylindrical stems.
Junellia as currently circumscribed ( Botta 1989 ; Peralta et al.
2008 ) is not monophyletic based on previous studies using
chloroplast DNA and nuclear waxy , PHOT1 and PHOT2 genes
Table 1. Comparison of historical treatments of Verbena , Glandularia, and Junellia.
Schauer ( 1847 ), Briquet ( 1895 ), Perry ( 1933 ) Troncoso ( 1974 ) Botta ( 1989 ), Atkins ( 2004 ), O’Leary et al. ( 2007a ), Peralta et al. ( 2008 ), (2009)
Verbena L.
sect. Verbenaca Schauer
Verbena Verbena
ser. Verbena
ser. Leptostachyae Schauer sect. Verbena ser. Pachystachyae Schauer
ser. Pachystachyae Schauer
ser. Pungentes Schauer
Junellia Moldenke
subg. Junellia
sect. Junellia
sect. Junelliopsis (Botta) Peralta & Múlgura
ser. Pauciflorae Briquet sect. Guedesia Botta
ser. Acerosae Briquet
ser. Junciformes Briquet
sect. Junellia (Mold.) Tronc. subg. Thryothamnus Botta
sect. Thryothamnus Botta
ser. Verticilliflorae Schauer sect. Verticiflora (Schauer) Botta
sect. Dentium Peralta & Múlgura
ser. Nobiles Schauer Glandularia J. F. Gmel. Glandularia
subg. Glandularia
sect. Glandularia (J. F. Gmel.) Schauer subg. Paraglandularia Schnack & Covas
2009] O’LEARY ET AL.: REDEFINITION OF JUNELLIA AND A NEW GENUS MULGURAEA 779
Fig. 1. One of the 12 maximun parsimony (MP) trees from the combined cpDNA data. The topology is similar to the Bayesian consensus tree.
Bootstrap values (BS) and Bayesian posterior probabilities (PP) supporting the corresponding branches are shown when BS 50% or PP 0.95 (BS/PP).
The asterisks indicate that BS < 50% when PP 0.95 or PP < 0.95 when BS 50% of the same branch. Numbers after J. juniperina designate different voucher
accesions. The arrow shows the position of Urbania pappigera . Glandularia species formerly placed in subgenus Paraglandularia are in bold. The asterisks
indicate species from Junellia sugbenus Thryothamnus sect. Thryothamnus ; Junellia subgenus Junellia species are shaded.
780 SYSTEMATIC BOTANY [Volume 34
Fig. 2. One of the 7,137 maximun parsimony (MP) trees from the combined ETS/ITS data. The topology is similar to the Bayesian consensus tree.
Node support (BS/PP) is shown as in Fig. 1 . Clone numbers are designated after species names when the individual sampled display >1% polymorphic
sites at the ETS/ITS region. Numbers after J. juniperina designate voucher accessions. The arrow shows the position of Urbania pappigera . Glandularia
species formerly placed in subgenus Paraglandularia are in bold. The asterisks indicate species from Junellia sugbenus Thryothamnus sect. Thryothamnus ;
Junellia subgenus Junellia species are shaded. Base chromosome numbers are represented along the branch leading to each of the four genera.
2009] O’LEARY ET AL.: REDEFINITION OF JUNELLIA AND A NEW GENUS MULGURAEA 781
( Yuan and Olmstead 2008a , b ). These results are here con-
firmed by our expanded chloroplast DNA data and the newly
generated ETS/ITS data (see Figs. 1 2 ), which suggest that
species from Junellia subgenus Thryothamnus sect. Dentium and
sect. Verticiflora form a clade distinct from the rest of Junellia .
Even though we sampled only six of the 13 taxa that belong
to these two sections due to difficulty in obtaining material,
we assume that the rest will fall within this clade based on
the shared morphological synapomorphy of anther connec-
tive surpassing the theca. This character state is not found in
any other Junellia , Glandularia, or Verbena species, all of which
have anther connectives not surpassing the theca (except four
Verbena species from ser. Pachystachyae ). In addition, a MITE
(miniature inverted-repeat transposable element) insertion in
waxy gene intron 8 and a MITE insertion in PHOT1 intron 13
were found in all Verbena , Glandularia , and Junellia s. s. species,
but not in Junellia subgenus Thryothamnus sect. Dentium and
sect. Verticiflora ( Yuan and Olmstead 2008a , b ). These MITE
insertions were suggested to be a molecular signature defin-
ing the clade of Verbena + Glandularia + Junellia s. s. ( Yuan and
Olmstead 2008a , b ), further corroborating the evidence that
Junellia is not monophyletic as currently circumscribed.
Given such strong evidence from both molecular and
morphological characters, species of Junellia subgenus
Thryothamnus sects. Dentium and Verticiflora are here circum-
scribed as the new genus Mulguraea.
Taxonomic Position of Glandularia subgenus Paraglan-
dularia— Schnack and Covas (1978 ) established Glandularia
subgenus Paraglandularia to include three South American
species, G. crithmifolia (Gillies & Hook. ex Hook.) Schnack &
Covas, G. hookeriana Covas & Schnack, and G. origenes (Phil.)
Schnack & Covas, which differed from the rest of the spe-
cies because of their subshrubby habit, winged or subwinged
mericarps and chromosome number 2 n = 20. This chromo-
some number is uncommon in Glandularia , given that all
South American Glandularia were believed to be diploids with
2 n = 10. Botta (1992 ) accepted subg. Paraglandularia and added
some defining morphological character states for this group,
including glabrous corolla throat, mericarps narrowed at the
base, and a fruiting calyx longer than the mericarps with non-
contorted teeth. Botta (1992 ) also suggested that G. ballsii was
better placed in subgenus Paraglandularia given the presence
of these character states in this species. All species in this sec-
tion are interpreted to have unbranched inflorescences, or
if branched then the lateral branches briefly pedunculated
(P. Peralta, pers. comm.), appearing as compressed inflo-
rescences because of the brief basal internodes ( Martínez et
al. 1996 ). In some species internodes are so brief and lateral
branches are so many, that it looks as if the flowers are arranged
in heads (e.g. G. ballsii and G. origenes ). G. fasciculata (Benth.)
P. Jörg. shares this type of inflorescence as well as characters
related to mericarp morphology, based on which it was also
grouped into subgenus Paraglandularia ( Peralta 2009 ).
All but one of the morphological characters that define sub-
genus Paraglandularia are also found in all species of Junellia
subgenus Thryothamnus sect. Thryothamnus , the exception
being the winged mericarps, which are found in only some
species. The close similarity between these two groups had
previously been noted by Botta (1989 ). Our phylogenetic anal-
yses indicate that G. crithmifolia, G. hookerina , and G. fasciculata
form a clade with species belonging to sect. Thryothamnus , all
forming a subclade within Junellia ( Fig. 2 ) or all part of a basal
grade within Junellia ( Fig. 1 ). This is also supported by pre-
vious studies using nuclear genes waxy, PHOT1 and PHOT2
( Yuan and Olmstead 2008a , b ), where G. crithmifolia always
is nested within Junellia . Unfortunately, we were unable to
obtain any living material from the remaining members of
Glandularia subgenus Paraglandularia ( G. ballsii, G. lucanensis ,
G. occulta, and G. origenes ) mainly because they are not wide-
spread. Two of these species are endemic to Perú ( G. lucan-
ensis and G. occulta ) and one ( G. ballsii ) is from the northern
Argentine province of Jujuy; herbarium specimens were not
of sufficient quality for DNA extractions. However, we pre-
dict that these four other species will group with Junellia ,
given that they share the character states that differentiate this
group from subgenus Glandularia and unite them with Junellia
sect. Thryothamnus . Consequently seven new combinations
are made. The inclusion of these species in Junellia means the
chromosome number 2 n = 20 is not a tetraploid number of
x = 5, but a diploid number of x = 10, as Junellia and Mulguraea
species have ( Botta and Brandham 1993 : 147–148).
Taxonomic Position of the genus Urbania— Urbania Philippi
(1891 ) is a monotypic genus, confined to the high Andean puna
of Argentina and Chile. Its taxonomic position has been con-
troversial, principally due to its close morphological similar-
ity to Junellia , from which it has been differentiated ( Troncoso
1974 ; Botta 1988 ) only by its hairy calyx with long hygroscopic
hairs. Our results show that Urbania is nested within the
Junellia clade ( Figs. 1 2 ), so recognition of Urbania as a differ-
ent genus, based only on the presence of a hairy calyx, renders
Junellia paraphyletic. Besides, Urbania and Junellia , especially
species from sect. Guedesia , share many morphological char-
acter states, including the cushion habit and homomorphic
leaves that are imbricate, spineless, and somewhat fleshy. The
transfer of Urbania to Junellia makes the presence of a hairy
calyx an autapomorphy for this new species of Junellia, and
does not justify its segregation as a different genus.
Redefinition of Junellia Boundaries— Our results show
that Junellia s. s. is restricted to four of the six sections classi-
fied by Peralta et al. (2008 ). These are sects. Junellia , Guedesia ,
Junelliopsis, and Thryothamnus . Species from the remaining two
sections, sects. Dentium and Verticiflora , have been transferred
to the new genus Mulguraea . However, Junellia has gained
seven species from Glandularia subg. Paraglandularia and one
species from the monotypic genus Urbania . As a consequence,
the recircumscribed Junellia comprises 36 species. The type
species of Junellia is Junellia serpyllifolia (Speg.) Moldenke
(1940 ), which is a synonym of J. micrantha (Phil.) Moldenke.
This species was included in our analyses, and the results
confirm the association of the name Junellia with this clade.
Species from subg. Paraglandularia , which have the style
base inserted between the mericarps, group with species of
sect. Thryothamnus ( Figs. 1 2 ), which have an enlarged base
style covering the apex of the mericarps. This renders the tra-
ditional division of Junellia into two subgenera, based upon
the presence or absence of an enlarged style base, no longer
correct. To further investigate if the sections within Junellia s. s.
constitute natural groups we should sample more taxa and
obtain more DNA sequence data.
Glandularia and Verbena— Our nuclear ETS/ITS data
indicate that both Glandularia and Verbena are monophyletic,
in agreement with the nuclear data in Yuan and Olmstead
(2008a ; 2008b ). However, neither Verbena nor
Glandularia are
monophyletic based on chloroplast data. These results sug-
gest two intergeneric chloroplast transfers from Verbena to
Glandularia ( Yuan and Olmstead 2008a ).
782 SYSTEMATIC BOTANY [Volume 34
Within Verbena , the North American species, treated by
O’Leary et al. (unpublished results) as series Verbena , were
recovered as monophyletic in this study ( Figs. 1 2 ), consistent
with results from the nuclear waxy and PHOT1 genes but incon-
gruent with the PHOT2 gene ( Yuan and Olmstead 2008a , b ).
Species of series Verbena are all diploids, with cortical chloro-
phyllous parenchyma as a continuous band and filiform or
much elongated cylindric inflorescences. South American spe-
cies of Verbena were also suggested to be monophyletic by the
chloroplast and nuclear ITS/ETS data ( Figs. 1 2 ), consistent
with PHOT1 gene data but not with PHOT2 or waxy sequence
data ( Yuan and Olmstead 2008a , b ). These species are treated
by O’Leary et al. (2007a ) in series Pachystachyae being mostly
polyploids with compressed cylindric inflorescences.
Within Glandularia , relationships resolved by the chloro-
plast DNA ( Fig. 1 ), nuclear ITS/ETS ( Fig. 2 ), waxy ( Yuan and
Olmstead 2008a ), PHOT1 and PHOT2 ( Yuan and Olmstead
2008b ), were largely incongruent. The incongruence is most
likely due to incomplete lineage sorting or/and extensive
recent gene flow. Potential gene flow among Glandularia spe-
cies was also implied by the intraindividual heterogeneity
(see Fig. 2 ) in the ribosomal ITS/ETS region of many dip-
loid species. The ribosomal ITS/ETS region is expected to be
homogenized rapidly in the absence of extensive gene flow,
presumably by highly efficient unequal crossing over or gene
conversion ( Eickbush and Eickbush 2007 ).
Relationship Among Genera of Tribe Verbeneae: Verbena,
Glandularia, Junellia and Mulguraea— This study corrobo-
rates previous inferences ( Yuan and Olmstead 2008a , b ) that
Verbena and Glandularia are sister groups, and Junellia as recir-
cumscribed here is sister to the Verbena + Glandularia clade.
The new genus Mulguraea is sister to this more inclusive clade.
Although Junellia and Mulguraea share many morphologi-
cal symplesiomorphies, such as a base chromosome number
x = 10, shrubby habit, unbranched inflorescences, and narrow
mericarp bases, these results suggest that Junellia s. s. is more
closely related to Verbena and Glandularia than to Mulguraea.
Given that both Mulguraea and Junellia are restricted to South
America, the origin of the Verbena complex is inferred to be
in South America. This also indicates that the chromosome
number x = 10 might be ancestral, and that x = 7 and x = 5
are derived from x = 10, as suggested by Yuan and Olmstead
(2008a ), in contrast with the view of Botta and Brandham
(1993 ).
Taxonomic Treatment
Mulguraea N. O’Leary & P. Peralta, gen. nov.—TYPE: Verbena
asparagoides Gillies & Hook. ex Hook., Bot. Misc. 1: 165.
1829. [ Mulguraea asparagoides (Gillies & Hook. ex Hook.)
N. O’Leary & P. Peralta].
Frutices caulibus cylindricus vel subcylindricus. Foliis
dimorphis, raro homomorphis, macroblastorum integris aut
trisectis, apice pungentibus, brachyblastorum integris, car-
nosis, inermis. Corolla faux glabra. Connectivum staminum
quam techae longius. Inflorescentiae capitula globosa vel
maturitate elongata efficientes, solitaria.
Subshrubs with cylindric or subcylindric stems, generally
divided in macroblast and brachyblasts. Leaves dimorphic,
sometimes homomorphic, macroblast leaves entire or tri-
sected, spinescent; brachyblast leaves entire, sometimes fleshy,
never spinescent. Many flowered, unbranched inflorescences,
forming cylindric spikes or racemes, rachis elongated during
maturity or not. Flowers slightly zygomorphic, calyx tubular,
4 or 5 teethed, never contorted at maturity. Corolla hypocra-
teriform, interior with retrorse hairs, throat pubescent or gla-
brous. Stamens 4, didynamous, attached towards the upper
middle of corolla tube, generally inserted, anthers with con-
nective tissue surpassing the theca, not appendaged. Ovary
bicarpellate, 4 locular, 4 ovular, with a filiform style more than
three times the length of the ovary, style base inserted between
mericarps, bilobed stigmatic tissue. Mericarps 4, narrowed at
the base, winged or not. Basic chromosome number x = 10.
Distribution— Eleven species and two varieties inhabiting
arid zones in southern South America.
Notes— The new genus is named in honor of Prof. M. E.
Múlgura (Instituto de Botánica Darwinion), who has been
working on Verbenaceae for more than 20 yr and is a well-
known specialist on Junellia . Because the species Mulguraea
asparagoides has typical characters states that define this
genus, it is designated as the type species. Mulguraea is sis-
ter to the clade Verbena + Glandularia + Junellia s. s. ( Figs. 1 – 2 .)
Sections are not recognized within Mulguraea until more taxa
have been analyzed.
See Peralta et al. (2008 ) for synonymies, complete descrip-
tion, distribution data, additional type information, and obser-
vations referred to each newly combined taxon. For a key to
the species of Mulguraea , see key to Junellia sect. Dentium and
sect. Verticiflora in Peralta et al. (2008 ).
The genus Dipyrena Hook. (tribe Priveae Briq.) has a sub-
drupaceous fruit separating into two mericarps and alternate
leaves; Junellia has a dry fruit separating into four mericarps
and opposite leaves. Ravenna (2008 ) transfered several Junellia
species to Dipyrena , but Múlgura and Peralta (unpublished
results) reject these new combinations on the basis of these
morphological differences between the two genera. Three of
these species, J. arequipensis, J. cinerascens, and J. scoparia are
here transfered to Mulguraea .
Mulguraea arequipense (Botta) N. O’Leary & P. Peralta, comb.
nov. Verbena arequipense Botta, Darwiniana 28: 237. [1987]
1988. Junellia arequipense (Botta) Botta, Darwiniana 29:
392. 1989. Dipyrena arequipensis (Botta) Rav., Onira11(15):
43. 2008.—TYPE: PERÚ. Arequipa: encima de baños
de Jesús, 23 Apr. 1961, 2,600–2,700 m, A. Ferreyra 14261
(holotype: SI!).
Mulguraea asparagoides (Gillies & Hook. ex Hook.)
N. O’Leary & P. Peralta, comb. nov. Verbena asparagoides
Gillies & Hook. ex Hook., Bot. Misc. 1: 165. 1829. Junellia
asparagoides (Gillies & Hook. ex Hook.) Moldenke, Lilloa
5: 395. 1940.—TYPE: ARGENTINA. Mendoza: “near
the upper … from Portezuelo a Uspallata to the valley
of Uspallata”, s. f., J. Gillies s.n. (lectotype, designated
by Peralta et al. 2008 : 371: K photo SI!; isotype: K, photo
SI!).
Mulguraea aspera (Gillies & Hook. ex Hook.) N. O’Leary &
P. Peralta, comb. nov. Verbena aspera Gillies & Hook. ex
Hook., Bot. Misc. 1: 163. 1829. Junellia aspera var. aspera
(Gillies & Hook. ex Hook.) Moldenke, Lilloa 5: 393.
1940.—TYPE: ARGENTINA. Mendoza: Paramillo de
Uspallata “near Los Hornillos, eastern descent from
Paramillo de Uspallata”, Nov. 1822, J. Gillies s.n. (lecto-
type, designated by Botta, 1989 : 392: K, photo SI!; isotype:
BM, photo SI!).
2009] O’LEARY ET AL.: REDEFINITION OF JUNELLIA AND A NEW GENUS MULGURAEA 783
Mulguraea aspera var. longidentata (Moldenke) N. O’Leary &
P. Peralta, comb. nov. Junellia longidentata Moldenke,
Known Geogr. Distr. Verb. Avicen.: 77. 1942. Verbena aspera
var. longidentata (Moldenke) Botta, Darwiniana 25: 338.
1984. Junellia aspera var. longidentata (Moldenke) Múlgura
& P. Peralta, Ann. Missouri Bot. Gard. 95(2): 338–390.
2008.—TYPE: ARGENTINA. Tucumán. Amaicha, 2090 m,
29 Dec. 1912, L. Castillón 2460 (holotype: LIL 31753, photo
SI!).
Mulguraea cedroides (Sandwith) N. O’Leary & P. Peralta,
comb. nov. Verbena cedroides Sandwith, Kew Bull. 1927:
184. 1927. Junellia cedroides (Sandwith) Moldenke, Lilloa
5: 394. 1940.—TYPE: ARGENTINA. Neuquén. Zapala:
Zapala, 27 Nov. 1925, H. F. Comber 189 (holotype: K, photo
SI!; isotypes: BM, K, LP!, TEX).
Mulguraea cinerascens (Schauer) N. O’Leary & P. Peralta,
comb. nov. Verbena cinerascens Schauer, in DC Prodr. 11:
545. 1847. Diostea cinerascens (Schauer) Moldenke, Revista
Sudameric. Bot. 5: 1. 1937. Junellia cinerascens (Schauer)
Botta, Darwiniana 29: 392. 1989. Dipyrena cinerascens
(Schauer) Rav., Onira 11(15): 44. 2008.—TYPE: CHILE. IV
Región. Coquimbo, 1829, C. Gay s. n. (lectotype, desig-
nated by Botta, 1989 : 392: G, photo FM 7855!; isotypes:
F, G, K, photo SI!).
Mulguraea echegarayi (Hieron.) N. O’Leary & P. Peralta,
comb. nov. Verbena echegarayi Hieron., Bol. Acad. Nac. Ci.
4: 66. 1881. Junellia echegarayi (Hieron.) Moldenke, Lilloa
5: 395. 1940.—TYPE: ARGENTINA. San Juan. Leoncito,
Jan. 1876, S. Echegaray s. n. (holotype: CORD!).
Mulguraea hystrix (Phil.) N. O’Leary & P. Peralta, comb. nov.
Verbena hystrix Phil., Anales Univ. Chile 90: 610. 1895.
Junellia hystrix (Phil.) Moldenke, Known Geogr. Dist.
Verben. Avicen.: 77. 1942.—TYPE: CHILE. Atacama:
Acerillos, desierto de Atacama, Nov. 1844, Villanueva s. n.,
SGO 42514 (lectotype, designated by Acevedo de Vargas
1951: 55: SGO; isotype: SGO, photo SI!).
Mulguraea ligustrina (Lag.) N. O’Leary & P. Peralta, comb.
nov. Verbena ligustrina Lag., Gen. Sp. Pl.: 18. 1816. Lippia
ligustrina (Lag.) Britton, Trans. New York Acad. Sci. 9:
181. 1890. Aloysia ligustrina (Lag.) Small, Flora SE United
States: 1013. 1903. Junellia ligustrina var. ligustrina
(Lag.) Moldenke, Phytologia 2(11): 466. 1948.—TYPE:
ARGENTINA. Santa Cruz. Deseado: a 107 km al S del
límite con Chubut, ruta Nac. 3, 25 Feb. 1990, M. N. Correa
et al. 10256 (neotype, designated by Peralta et al. 2008 :
378: SI!).
Mulguraea ligustrina var. lorentzii (Niederl. ex Hieron.)
N. O’Leary & P. Peralta, comb. nov. Verbena lorentzii
Niederl. ex Hieron., Bol. Acad. Nac. Ci. Córdoba 3(4): 370.
1880. Junellia lorentzii (Niederl. ex Hieron.) Moldenke,
Lilloa 5: 397. 1940. Junellia ligustrina var. lorentzii (Niederl.
ex Hieron.) Moldenke, Phytologia 47: 222. 1980.—TYPE:
ARGENTINA. Río Negro. Sep.-Nov., 1874, C. Berg
100 (lectotype, designated by Peralta et al. 2008 : 379:
CORD).
Mulguraea scoparia (Gillies & Hook. ex Hook.) N. O’Leary &
P. Peralta, comb. nov. Verbena scoparia Gillies & Hook. ex
Hook., Bot. Misc. 1: 161. 1829. Diostea scoparia (Gillies &
Hook. ex Hook.) Miers, Trans. Linn. Soc. London 27: 104.
1869. Junellia scoparia (Gillies & Hook. ex Hook.) Botta,
Darwiniana 29: 392. 1989. Dipyrena scoparia (Gillies &
Hook. ex Hook.) Rav., Onira 11(15): 44. 2008.—TYPE:
ARGENTINA. Mendoza: Las Heras, shrub valley near
Villavicencio, J. Gillies s. n. (lectotype, designated by
Botta, 1989 : 392: K, photo SI!; isotype: K, photo SI!).
Mulguraea tetragonocalyx (Tronc.) N. O’Leary & P. Peralta,
comb. nov. Verbena tetragonocalyx Tronc., Darwiniana
8: 481. 1949. Junellia tetragonocalyx (Tronc.) Moldenke,
Phytologia 3: 167. 1949.—TYPE: ARGENTINA. Chubut.
Paso de Indios: 23 km al E de El Sombrero, 6 Jan. 1948,
Soriano 2839 (holotype: SI!; isotype: K, photo SI!).
Mulguraea tridens (Lag.) N. O’Leary & P. Peralta, comb. nov.
Verbena tridens Lag., Gen. Sp. Pl. 19. 1816. Junellia tridens
(Lag.) Moldenke, Lilloa 5: 402. 1940.—TYPE: Argentina.
Chubut. Florentino Ameghino: a 7 km del cruce de Ruta
Provincial 28 con Ruta Nacional 3, 24 Feb. 1990, N. Correa
10227 (neotype, designated by Peralta et al. 2008 : 373:
SI!).
New transfers to Junellia— Eight new combinations under
Junellia have resulted from this study. See Botta (1988 ) for syn-
onymies, complete descriptions, distribution data, additional
type information, and observations referred to each newly
combined taxa of Junellia previously placed under Glandularia
subgenus Paraglandularia or under the monotypic Urbania ,
respectively. Given that Urbania ( Philippi 1891 ) has priority
over Junellia ( Moldenke 1940 ), a formal proposal to conserve
the name Junellia over Urbania was made to the International
Committee on Nomenclature and has been recently accepted
( O’Leary et al. 2009 ).
Junellia ballsii (Moldenke) N. O’Leary & P. Peralta, nov.
comb. Verbena ballsii Moldenke, Phytologia 1: 477. 1940.
Glandularia ballsii (Moldenke) Botta, Hickenia 2: 61.
1992.—TYPE: ARGENTINA. Jujuy, Chorru Valley, near
Tilcara, 12,800 ft, 13 Feb 1939, E. K. Balls 6036 (holotype:
US; isotypes: K, US, W).
Junellia crithmifolia (Gillies & Hook. ex Hook.) N. O’Leary &
P. Peralta, nov. comb. Verbena crithmifolia Gillies & Hook.
ex Hook. Bot. Misc. 1: 169. 1829. Glandularia crithmifolia
(Gillies & Hook. ex Hook.) Schnack & Covas, Darwiniana
6: 475. 1944.—TYPE: ARGENTINA. “along the foot of
mountains near Mendoza and San Luis de la Punta”,
J. Gillies s. n. (lectotype, designated by Botta, 1993 : 18, K;
isolectotype: K).
Junellia fasciculata (Benth.) N. O’Leary & P. Peralta, nov.
comb. Verbena fasciculata Benth., Bot. Voy. Sulphur.: 153.
1844. Glandularia fasciculata (Benth.) P. Jörg., Mem. New
York Bot. Gard. 92: 463. 2005.—TYPE: PERÚ. Lima.
“Huamantango”, Hinds 707 (holotype: K, isotypes: K,
SI).
Junellia hookeriana (Covas & Schnack) N. O’Leary &
P. Peralta, nov. comb. Glandularia hookeriana Covas &
Schnack, Revista Argent. Agron. 12: 57. 1945. Verbena
hookeriana (Covas & Schnack) Moldenke, Phytologia 2:
149. 1946.—TYPE: ARGENTINA. Mendoza. Dpto. La
Paz: entre La Paz y Desaguadero, 15 Dec 1943, G. Covas
2106 (holotype: SI).
Junellia lucanensis (Moldenke) N. O’Leary & P. Peralta,
nov. comb. Verbena lucanensis Moldenke, Phytologia 3:
784 SYSTEMATIC BOTANY [Volume 34
279. 1950. Glandularia lucanensis (Moldenke) Botta,
Hickenia 2: 128. 1995.—TYPE: PERÚ. Ayacucho, Luca-
nas, between Nazca and Puquio, 1,500–2,000 msm,
19 Mar 1949, R. Ferreira 5493 (holotype: NY; isotypes:
MO, US).
Junellia occulta (Moldenke) N. O’Leary & P. Peralta, nov.
comb. Verbena occulta Moldenke, Phytologia 3: 280. 1950.
Glandularia occulta (Moldenke) P. Jörg., Mem. New York
Bot. Gard. 92: 463. 2005.—TYPE: PERÚ. La Libertad, Prov.
Bolívar, cerca al Nevado Cajamarquilla, 12 Sep 1946,
R. Ferreira 1298 (holotype: NY; isotypes: NY, SI)
Junellia origenes (Phil.) N. O’Leary & P. Peralta, nov. comb.
Verbena origenes Phil., Linnaea 29: 20. 1857–58. Glandularia
origenes (Phil.) Schnack and Covas, Darwiniana 6: 475.
1944.—TYPE: CHILE. Andium Coquimbensium, monte
del Toro, Nov 1836, C. Gay 1906 (holotype: SGO 54769;
isotype: SGO 54769).
Junellia pappigera (Phil.) N. O’Leary & P. Peralta, nov. comb.
Urbania pappigera Phil., An. Mus. Nac. Chile (Verz. Antof.
Pfl.), 2° sect. Bot. 8: 60. 1891.—TYPE: CHILE. “ad. ped.
montis Miñique, I-1885”, s/leg. (lectotype, designated by
Botta 1988 : 480: SGO 42497).
Key to the Genera of Tribe Verbeneae
1. Subshrubs or cushion plants, mericarp base narrowed, inflorescences generally unbranched (sometimes with simple branching),
base chromosome number x = 9 or x = 10 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Anther connective never surpassing the theca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Junellia
2. Anther connective surpassing the theca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mulguraea
1. Herbs, mericarp base thickened, not narrowed, branched inflorescences (at times condensed and appearing simple), base
chromosome number x = 5 or x = 7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Style generally more than 3 × longer than ovary, mature calyx usually longer than fruit and with teeth generally contorted,
chromosome number x = 5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glandularia
3. Style never more than 3 × longer than ovary, mature calyx shorter than fruit and with teeth not contorted, chromosome
number x = 7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Verbena
Acknowledgments. This study was possible thanks to grants to
M. E. Múlgura and N. O’Leary (PIP 5262, CONICET) and grants to
R. G. Olmstead (NSF: DEB-0542493, DEB-0710026), a postdoctoral fellow-
ship to N. O’Leary from CONICET, and a Plant Molecular Systematics
Graduate Fellowship to Y. W. Yuan from the University of Washington,
Department of Biology. The authors are grateful to Mark Simmons and
two anonymous reviewers for helpful comments on an earlier version of
this manuscript.
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Appendix 1. Accession data for plants included in the molecular phy-
logenetic analyses. Voucher information is only given for sequences gen-
erated in this study, GenBank numbers from this study are in bold, the rest
were submitted prior to this study in connection with a previous paper
( Yuan and Olmstead 2008a ). For voucher information for species marked
with ‡ see Yuan and Olmstead (2008a ). GenBank accesion numbers are in
the following sequence: trnS-G / trnD-T / trnS-fM / trnT-F / ETS / ITS (—
indicates sequence not obtained). The number in parentheses represents
different clones. The taxa marked by an asterisk represent DNA extrac-
tions from herbarium specimens.
Aloysia Palau — A. virgata (Ruiz and Pav.) Juss. ‡, EF583119 / EF583172 /
EF576717 / EF571570 / FJ867545 / FJ867398
Glandularia J. F. Gmel. — G. araucana (Phil.) Botta‡, EF583104 /
EF583157 / EF576702 / EF571555 / (c2)FJ867576, (c4)FJ867577 / (c2)
FJ867429, (c4)FJ867430 . G. aristigera (S. Moore) Tronc. ‡, EF583106 /
EF583159 / EF576704 / EF571557 / (c3)FJ867570, (c6)FJ867571 / (c3)
FJ867423, (c6)FJ867424 . G. aurantiaca (Speg.) Botta‡, EF583103 / EF583156 /
EF576701 / EF571554 / (c3)FJ867572, (c4)FJ867573, (c6)FJ867574 / (c3)
FJ867425, (c4)FJ867426, (c6)FJ867427 . G. bajacalifornica (Moldenke) Umber,
EF583097 / EF583150 / EF576695 / EF571548 / — / —. G. balansae
(Briq.) N. O’Leary, Paraguay: Canindeyú, Múlgura 3576 (CTES, FCQ, SI),
FJ867511 / FJ867476 / FJ867489 / FJ867532 / FJ867596 / —. G. bipinnatifida
Nutt. ‡, EF583084 / EF583137 / EF576682 / EF571535 / (TX2)FJ867588,
(TX1)FJ867589 / (TX2)FJ867440, (TX1)FJ867441 . G. canadensis (L.) Nutt.,
EF583093 / EF583146 / EF576691 / EF571544/ — / —. G. cheitmaniana
(Moldenke) Botta and Poggio‡, Argentina, Jujuy, Marazzi 260 (SI), FJ867508 /
FJ867474 / FJ867493 / FJ867530 / FJ867595 / FJ867444 . Glandularia chirica-
hensis Umber‡, EF583095 / EF583148 / EF576693 / EF571546 / FJ867584 /
FJ867436 . G. crithmifolia (Gillies and Hook. ex Hook.) Schnack and Covas‡,
EF583107, EF586160, EF576705, EF571558, FJ867593 / FJ867443 . G. delticola
(Small) Umber, EF583090 / EF583143 / EF576688 / EF571541 / — / —.
G. dissecta (Spreng.) Schnack and Covas‡, EF583085 / EF583138 / EF576683 /
EF571536 / (c1)FJ867565, (c5)FJ867566 / (c1)FJ867418, (c5)FJ867419 .
G. fasciculata Benth.*, Perú: Ancash, Cano 1717 (MO), FJ867514 / FJ867472 /
FJ867491 / FJ867528 / FJ867597 / FJ867447 . G. flava (Gillies and Hook. ex
Hook.) Schnack and Covas‡, EF583102 / EF583155 / EF576700 / EF571553 /
FJ867575 / FJ867428 . G goodingii var. goodingii (Briq.) Solbrig‡, EF583096 /
EF583149 / EF576694 / EF571547 / FJ867585 / FJ867437 . G. goodingii var.
neptifolia Tidestr. ‡, EF583089 / EF583142 / EF576687 / EF571540 / (c4)
FJ867586, (c5)FJ867587 / (c4)FJ867438, (c5)FJ867439 . G. guaranitica Tronc.
‡, EF583087 / EF583140 / EF576685 / EF571538 / (c5)FJ867581, (c6)
FJ867582, (c8)FJ867583 / (c5)FJ867433, (c6)FJ867434, (c8)FJ867435 . G. hook-
eriana Covas and Schnack, Argentina: Catamarca, RGO 2007-080 (MERL,
SI, WTU), FJ867510 / FJ867471 / FJ867495 / FJ867527 / FJ867590 / FJ867448 .
G. incisa (Hook.) Tronc. ‡, EF583086 / EF583139 / EF576684 / EF571537 /
FJ867567 / FJ867420 . G. mendocina (Phil.) Covas and Schnack‡, EF583100
/ EF583153 / EF576698 / EF571551 / (c5)FJ867568, (c8)FJ867569 /
(c5)FJ867421, (c8)FJ867422 . G. microphylla (Kunth) Cabrera‡, EF583101 /
EF583154 / EF576699 / EF571552 / FJ867579 / FJ867432 . G. parodii Covas
and Schnack, EF583099 / EF583152 / EF576697 / EF571550 / — / —.
G. peruviana (L.) Small, Argentina: Jujuy, RGO 2007-066 (MERL, SI, WTU),
FJ867512 / FJ867477 / FJ867490 / FJ867533 / FJ867591 / FJ867445 . G. pumila
(Rydb.) Umber, EF583094 / EF583147 / EF576692 / EF571545 / — / —.
G. santiaguensis Covas and Schnack, Argentina, Salta, RGO 2007-005 (MERL,
SI, WTU), FJ867509 / FJ867473 / FJ867494 / FJ867529 / FJ867594 / —. G. scrobic-
ulata (Griseb.) Tronc., Argentina, Jujuy, Marazzi 259 (SI), FJ867513 / FJ867475
/ FJ867492 / FJ867531 / FJ867592 / FJ867446 . G. subincana Tronc., Argentina:
Corrientes, RGO 2004-117 (WTU), EF583088 / EF583141 / EF576686 /
EF571539 FJ867580 / FJ867442 . G. tampensis Small, EF583091 / EF583144
/ EF576689 / EF571542 / — / —. G. tenera (Spreng.) Cabrera‡, EF583105 /
EF583158 / EF576703 / EF571556 / FJ867578 / FJ867431 . G. teucriifolia (Mart.
and Galeotti) Umber, EF583098 / EF583151 / EF576696 / EF571549 / — / —.
G. verecunda Umber, EF583092 / EF583145 / EF576690 / EF571543 / — / —
Junellia http://www.tropicos.org/name/40033922 Moldenke
J. asparagoides (Gillies and Hook. ex Hook.) Moldenke‡, EF583116 /
EF583169 / EF576714 / EF571567 / FJ867616 / FJ867458 . J. aspera (Gillies
and Hook. ex Hook.) Moldenke‡, EF583117 / EF583170 / EF576715 /
EF571568 / FJ867617 / FJ867459. J. aspera var. longidentata (Moldenke)
Múlgura and P. Peralta, Argentina: Jujuy, Soza 1824 (WTU), FJ867523 /
FJ867479 / FJ867502 / FJ867539 / FJ867618 / FJ867460 . J. caespitosa (Gillies
and Hook. ex Hook.) Moldenke, Argentina: San Juan, Meglioli 174 (SI),
FJ867520 / FJ867481 / FJ867500 / FJ867536 / FJ867606 / FJ867466 . J. con-
natibracteata (Kuntze) Moldenke‡, EF583108 / EF583161 / EF576706 /
EF571559 / FJ867613 / FJ867453 . J. juniperina 1 (Lag.) Moldenke, EF583112 /
EF583165 / EF576710 / EF571563 / — / —. J. juniperina 2 (Lag.) Moldenke‡,
EF583113 / EF583166 / EF576711 / EF571564 / FJ867612 / FJ867455 . J. ligus-
trina var. lorentzii (Niederl. ex Hieron.) Moldenke, Argentina: Neuquén,
RGO 2007-092 (MERL, SI, WTU), FJ867522 / FJ867486 / FJ867504 / FJ867542
/ FJ867603 / FJ867468 . J. micrantha (Phil.) Moldenke, Argentina: Mendoza,
RGO 2004-198 (MERL, SI), FJ867524 / FJ867482 / FJ867498 / FJ867537 /
FJ867602 / FJ867462 . J. minima (Meyen) Moldenke‡, EF583109 / EF583162
/ EF576707 / EF571560 / FJ867611 / FJ867454 . J. scoparia (Gillies and Hook.
ex Hook.) Botta‡, EF583115 / EF583168 / EF576713 / EF571566 / FJ867615
/ FJ867457 . J. selaginoides (Walp.) Moldenke*, Chile: Elqui, Lammers et al.
7597 (US), FJ867518 / FJ867485 / FJ867505 / FJ867541 / FJ867604 / FJ867463 .
J. seriphioides (Gillies and Hook. ex Hook.) Moldenke‡, EF583110 /
EF583163 / EF576708 / EF571561 / FJ867610 / FJ867452 . J. spathulata
(Gillies and Hook. ex Hook.) Moldenke‡, EF583114 / EF583167 / EF576712
/ EF571565 / FJ867614 / FJ867456 . J. succulentifolia (Kuntze) Moldenke*,
Argentina, Bonifacino 418 (US), FJ867517 / — / FJ867503 / FJ867540 /
FJ867599 / FJ867461 . J. tonini var. mulinoides (Speg.) P. Peralta and Múlgura*,
Argentina: Neuquén, Kiesling 10081c (SI), FJ867521 / FJ867484 / FJ867499
/ FJ867538 / FJ867601 / FJ867467 . J. tonini var. tonini (Kuntze) Moldenke,
Argentina: Neuquén, Morrone 5756 (SI), FJ867525 / FJ867483 / FJ867506 /
FJ867534 / FJ867600 / FJ867464 . J. tridens (Lag.) Moldenke*, Argentina:
Santa Cruz, Bonifacino 500 (US), FJ867516 / FJ867487 / FJ867501 / FJ867543 /
FJ867605 / FJ867465 . J. ulicina (Phil.) Moldenke, Argentina: Mendoza, RGO
2004-201 (MERL, SI), FJ867519 / FJ867480 / FJ867497 / FJ867535 / FJ867607 /
FJ867469 . J. uniflora (Phil.) Moldenke‡, EF583111 / EF583164 / EF576709 /
EF571562 / (c2)FJ867608, (c7)FJ867609 / (c2)FJ867450, (c7)FJ867451 .
Lippia L. L. salsa Griseb. ‡, EF583118 / EF583117 / EF576716 /
EF571569 / FJ867546 / FJ867399 .
Urbania Phil. U . pappigera Phil., Argentina: Salta, RGO 2007-036
(MERL, SI, WTU), FJ867515 / FJ867478 / FJ867496 / FJ867544 / FJ867598
/ FJ867449.
Verbena L. — V. bonariensis L. ‡, EF583067 / EF583120 / EF576665 /
EF571518 / FJ867548 / FJ867401 . V. bracteata Cav. ex Lag. and Rodr. ‡,
EF583075 / EF583128 / EF576673 / EF 571526 / FJ867555 / FJ867408 .
786 SYSTEMATIC BOTANY [Volume 34
V. canescens Kunth‡, EF583082 / EF583135 / EF576680 / EF571533 /
FJ867559 / FJ867412 . V. halei Small‡, EF583083 / EF583136 / EF576681 /
EF571534 / FJ867556 / FJ867409 . V. hastata L. ‡, EF583080 / EF583133 /
EF576678 / EF571531 / FJ867562 / FJ867415 . V. hispida Ruiz and Pav.,
Argentina: Jujuy, Soza 1826 (WTU), — / — / — / — / FJ867549 / FJ867402 ;
Argentina: Misiones, Zuloaga 8266 (SI), FJ867507 / FJ867470 / FJ867488 /
FJ867526 / — / —. V. intermedia Gillies and Hook. ex Hook. ‡, EF583071 /
EF583124 / EF576669 / EF571522 / FJ867547 / FJ867400 . V. lasiostachys
Link‡, EF583072 / EF583125 / EF576670 / EF571523 / FJ867564 / FJ867417 .
V. litoralis Kunth‡, EF583069 / EF583122 / EF576667 / EF571520 /
FJ867551 / FJ867404 . V. macdougalii Heller‡, EF583081 / EF583134 /
EF576679 / EF571532 / FJ867554 / FJ867407 . V. menthifolia Benth. ‡,
EF583076 / EF583129 / EF576674 / EF571527 / FJ867560 / FJ867413 .
V. montevidensis Spreng. ‡, EF583070 / EF583123 / EF576668 / EF571521 /
FJ867552 / FJ867405 . V. neomexicana var. hirtella L. M. Perry‡, EF583078 /
EF583131 / EF576676 / EF571529 / FJ867557 / FJ867410 . V. officinalis L. ‡,
EF583074 / EF583127 / EF576672 / EF571525 / FJ867561 / FJ867414 .
V. orcuttiana L. M. Perry‡, EF583079 / EF583132 / EF576677 / EF571530 /
FJ867553 / FJ867406 . V. perennis Wooton‡, EF583077 / EF583130 / EF576675 /
EF571528 / FJ867558 / FJ867411 . V. rigida Spreng. ‡, EF583068 / EF583121
/ EF576666 / EF571519 / FJ867550 / FJ867403 . V. urticifolia L. ‡, EF583073 /
EF583126 / EF576671 / EF571524 / FJ867563 / FJ867416 .
... However, several biogeographic patterns are possible for the North American lineages, including (1) long-distance dispersal from South American arid zones, (2) a migration route connecting South American and North American arid zones, or (3) de novo evolution of plants in North American arid zones from wet tropical forest ancestors. We review previous evidence from Lantaneae and Verbeneae ( Yuan and Olmstead, 2008a , b ;O'Leary et al., 2009 ;Lu-Irving et al., 2014 ) and present new evidence from Verbena and Citharexylum to improve our understanding of the origin of North American arid-zone Verbenaceae. ...
... Because this study integrates previously published results ( Aloysia : Lu-Irving et al., 2014 ), a previously studied group for which additional sampling was obtained (Verbeneae: Yuan and Olmstead, 2008a ;O'Leary et al., 2009 ), and a group ( Citharexylum ) not previously subject to phylogenetic analysis, sampling and DNA regions vary among the diff erent case studies. For a table of vouchers, herbaria housing voucher specimens, and GenBank accession numbers accompanying sequence data used in this study, see Appendix S1 in the Supplemental Data with this article. ...
... Polymerase chain reaction protocols for the two chloroplast regions, ndhF and the trnT-F fragment, the latter of which includes the trnT-trnL spacer, trnL intron, and trnL-trnF spacer, followed those of Marx et al. (2010) and Yuan and Olmstead (2008a) for ndhF and trnT-F , respectively. Internal (ITS) and external transcribed spacer (ETS) regions of nuclear 18S/26S rDNA were amplifi ed as done by O'Leary et al. (2009) . Th e PPR11 gene was amplifi ed as done by Yuan et al. (2010) . ...
Article
Premise of the study: Verbenaceae originated and initially diversified in South America in wet forest habitats. They have diversified extensively in arid habitats in both South and North America. This study aims to understand the origin of the North American arid-land members of Verbenaceae. Methods: A phylogenetic approach is used to examine four genera (Aloysia, Citharexylum, Glandularia, Verbena) in three distinct clades with representatives in North American deserts and disjunct South and North American distributions. Phylogenetic analyses were conducted using maximum likelihood and Bayesian approaches. Analyses included both plastid and nuclear DNA regions and include the first study of Citharexylum and an expanded sampling of tribe Verbeneae (Glandularia and Verbena). Ancestral areas were reconstructed for each group. Key results: North American desert species of Aloysia and Glandularia were likely derived from ancestors in arid temperate South America, perhaps by long-distance dispersal. The pattern for Verbena was less clear, with evidence from plastid DNA implicating an Andean dispersal route to the North American clade, whereas nuclear data suggest that the Andean and North American species resulted from independent dispersals from southern South America. A previously unrecognized clade of Andean Verbeneae was discovered, raising the possibility of an Andean origin of Verbena or Verbena and Glandularia. North American desert species of Citharexylum represent multiple, independent origins from mesic habitat ancestors in Mesoamerica. Conclusions: North American arid-zone Verbenaceae are derived from South and Central American ancestors via multiple avenues, including long-distance, amphitropical dispersal, Andean migration corridors, and in situ evolution of desert-adapted species.
... Verbenaceae is flowering plant family in the large asteroid order Lamiales (Refulio & Olmstead, 2014). The Verbenaceae included 35 genera and 830 species (Atkins, 2004;O'Leary et al., 2009O'Leary et al., , 2012Thode et al., 2013). In Brazil Verbenaceae is represented by 300 species in 16 genera (Salimena & Mulgura, 2015). ...
... Verbena species depicted close phylogenetic association based on plastid markers ndhF and trnL-trnF by Thode et al., (2013). O'Leary et al., (2009) also studied the relationship of Verbena species on the basis of chloroplast trnD-trnT, trnS-trnG, trnS-trnfM, trnT-trnL, trnG, trnL, trnL-trnF and the nuclear ITS and ETS regions. ...
... Además, estudios filogenéticos morfológicos en Glandularia (Peralta, 2010) y estudios filogenéticos moleculares en la tribu Verbeneae (Yuan & Olmstead, 2008a;2008b) sugieren que los taxones de Glandularia subgénero Paraglandularia deberían agruparse bajo el género Junellia. O'Leary et al. (2009a), ampliando el número de taxones estudiados e incluyendo nuevos marcadores moleculares, demuestran que Glandularia subgénero Paraglandularia y el género monotípico Urbania pappigera Phil. forman parte del mismo clado que las especies de Junellia, y proponen ocho nuevas combinaciones bajo éste último género. ...
... Urbania), Botta (1988: 480) menciona que el tejido conectivo de las anteras supera la longitud de las tecas. Sin embargo, el análisis del material de herbario evidencia lo contrario, corroborando los estudios moleculares (O'Leary et al., 2009a) que posicionan a este taxón dentro de Junellia y no dentro de Mulguraea. ...
Article
Full-text available
Asinopsis of genus Junellia is here presented, following the recent re-circumscription of the genus based on morphologically founded molecular phylogenetic studies. A key to the 37 Junellia species recognized by the latest delimitation of the genus is here proposed, as well as an actualized description of the genus and its differentiation from related genera from tribe Verbeneae. Nine species and one variety of Junellia, never taxonomically treated before, are here described and/or illustrated, or their description is here amended. Two new combinations: Junellia hookeriana var. catamarcensis and Junellia trifida, and eight new synonyms are here proposed.
... Targeted loci were selected for their phylogenetic utility and use in previous studies of Verbenaceae (O'Leary et al., 2009;Yuan et al., 2008Yuan et al., , 2009aYuan et al., , 2009bMarx et al., 2010;Lu-Irving and Olmstead, 2013;Lu-Irving et al. 2014;Frost et al., 2017). Seven plastid regions (cpDNA: matK, ndhF, rbcL, rpl32, rpoC2, trnT-trnL, and trnL-trnF), two spacer regions (ETS and ITS) of nuclear 18 S/26 S rDNA (nrDNA), and five lowcopy nuclear genes (waxy, PPR 24, PPR 62, PPR 70, and PPR 123) were included in this study. ...
Article
Premise: As a family of Neotropical origin and primarily Neotropical distribution, the Verbenaceae are a good but understudied system with which to understand Neotropical evolution. Tribe Citharexyleae comprises three genera: Baillonia, Citharexylum-one of the largest genera in Verbenaceae-and Rehdera. A molecular phylogenetic approach was taken to resolve intergeneric relationships in Citharexyleae and infrageneric relationships in Citharexylum. The phylogeny is used to elucidate character evolution in a widespread, morphologically diverse Neotropical genus. Methods: Seven plastid regions, two nuclear ribosomal spacers, and six low-copy nuclear loci were analyzed for 64 species of Citharexyleae. Phylogenetic analyses were conducted using maximum likelihood, Bayesian inference, and multispecies coalescent approaches. Habit, presence or absence of thorns, inflorescence architecture, flower color, fruit color, and geography were examined to identify diagnostic character states for clades within Citharexylum. Results: Rehdera is resolved as sister to Citharexylum, and Baillonia nested within Citharexylum. Two species, C. oleinum and C. tetramerum, are not closely related to tribe Citharexyleae, but may be related to members of tribe Duranteae instead. Seven clades within Citharexylum are inferred, each characterized by a combination of geography, fruit color and/or maturation, and inflorescence architecture. There is evidence of correlated evolution between habit, axillary inflorescences, and flower number per inflorescence. Shrubs with reduced inflorescences have evolved repeatedly. Conclusions: A subgeneric classification for Citharexylum is proposed. Although suites of associated traits are found, character morphology has been labile throughout Citharexylum's evolutionary history. Morphological diversity may be related to adaptation to differing mesic and xeric habitats.
... The genus Junellia, belonging to the Verbenaceae family, is represented by around 40 species distributed from Peru and Bolivia to Argentina and Chile, with mainly Andean-Patagonian distribution demonstrating strong adaptation to adverse environmental conditions (O'Leary et al., 2009(O'Leary et al., , 2011;Peralta et al., 2008). In Chile, 22 species inhabit (Rodríguez et al., 2018), five of which are endemic, distributed mainly in the high Andean areas of the arid regions of the north, a few species in the central Andean mountain range and others in the extreme south (Rodríguez et al., 2018) In the Chilean highlands, they are used medicinally to treat digestive disorders, colds, cough, fever and bladder ailments (Trivelli and Huerta, 2014;Ortiz et al., 2019). ...
Article
Ethnopharmacological relevance Chilean population relies on medicinal plants for treating a wide range of illnesses, especially those of the gastrointestinal system. Junellia spathulata (Gillies & Hook.) Moldenke var. spathulata (Verbenaceae), called as “verbena-azul-de-cordilleira”, is a medicinal plant native to Argentina and Chile traditionally used for treating digestive disorders. Although the species of the genus are important as therapeutic resources for the Andean population, the plants are very scarcely studied. Aims of the study The purpose of the present study was to find out the main constituents and investigate the protective effect of J. spathulata against oxidative stress induced by the potent oxidant 2,2′-azobis (2-amidinopropane) dihydrochloride (AAPH) in human hepatoblastoma cells. Materials and methods The crude methanol extract of J. spathulata and an iridoid obtained by chromatographic processes were tested to access the hepatoprotective effect and cytotoxicity in HepG2 cell. In addition, the reducing power of the samples and their ability to scavenge free radicals were evaluated using FRAP and ORAC assay systems. Results The iridoid asperuloside, the main compound of the crude methanol extract of J. spathulata, was isolated and identified by means of NMR analysis. The crude methanol extract of J. spathulata and asperuloside protected HepG2 cells against oxidative damage triggered by AAPH-derived free radicals. This effect can be credited to the ability of the extract and asperuloside to protect the liver cells from chemical-induced injury, which might be correlated to their free radical scavenging potential. Conclusions This study experimentally evidenced the ethnopharmacological usefulness of J. spathulata as a treatment of digestive disorders. Our result could stimulate further investigations of hepatoprotective agents in other Chilean Junellia species.
... El ciclo vital es desconocido, pero es posible suponer que sea holocíclico, como en A. matilei. El género Mulguraea se ha constituido recientemente con 13 especies sudamericanas (O'Leary et al., 2009). M. aspera se encuentra distribuida entre Río Negro al sur y Catamarca al norte (Instituto de Botánica Darwinion, 2017). ...
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Full-text available
Se describen tres especies del género Aphis Linnaeus (Hemiptera, Aphididae, Aphidinae) a partir de hembras vivíparas ápteras y aladas recogidas sobre especies de Junellia, Lippia y Mulguraea (Verbenaceae) en localidades de la Argentina, respectivamente: Aphis junelliae González Rodríguez & Nieto Nafría sp. n., Aphis lippiae Ortego & Nieto Nafría sp. n., Aphis mulguraeae Nieto Nafría & Mier Durante sp. n. Se exponen las características que permiten diferenciar las nuevas especies entre sí y de las restantes especies de Aphis conocidas en América del Sur. urn:lsid:zoobank.org:pub:A365C410-F84A-40DE-8C5B-13C2FA768EFC
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Premise of the study Tribe Citharexyleae comprises three genera: Baillonia, Citharexylum , and Rehdera . While there is good support for these genera as a clade, relationships between genera remain unresolved due to low sampling of the largest genus, Citharexylum . A molecular phylogenetic approach was taken to resolve intergeneric relationships in Citharexyleae and infrageneric relationships in Citharexylum . Methods Seven chloroplast regions, two nuclear ribosomal spacers, and six low-copy nuclear loci were analyzed for 64 species of Citharexyleae. Phylogenetic analyses were conducted using maximum likelihood, Bayesian inference, and Bayesian multi-species coalescent approaches. Habit, presence/absence of thorns, inflorescence architecture, flower color, fruit color, and geography were examined to identify diagnostic characters for clades within Citharexylum . Key results Intergeneric relationships resolved Rehdera as sister to Citharexylum and Baillonia nested within Citharexylum . Two species, C. oleinum and C. tetramerum , fell outside of Citharexyleae close to tribe Duranteae. There is strong support for seven clades within Citharexylum , each characterized by a unique combination of geography, fruit color/maturation, and inflorescence architecture. Conclusions Baillonia is included in Citharexylum; Rehdera is retained as a distinct genus. A subgeneric classification for Citharexylum is proposed.
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— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.