Phylogeny of the Neotropical tribe Jacarandeae (Bignoniaceae)

Abstract and Figures

Premise: The tribe Jacarandeae includes Jacaranda (49 species) and Digomphia (3 species), two genera of trees and woody shrubs with Neotropical distribution. Jacarandeae is sister to the rest of the Bignoniaceae, but not much is known about interspecific and intergeneric relationships within this group. Methods: We reconstructed the phylogeny of Jacarandeae using chloroplast (ndhF, rpl32-trnL, trnL-F) and nuclear (ETS, PPR62) markers. Evolutionary relationships within Jacarandeae were inferred using Bayesian, Maximum Likelihood, and species tree approaches. The resulting phylogenetic framework was used as the basis to interpret the evolution of key morphological character states (i.e., stamen and calyx traits) and revise the infra-generic classification of the group. Results: Jacaranda and Digomphia belong to a well-supported clade, with Digomphia nested within Jacaranda. We propose the necessary taxonomic changes to recognize monophyletic taxa, including a broadly circumscribed Jacaranda divided into four sections: (1) Jacaranda sect. Nematopogon, species previously included in Digomphia and united by divided staminode apices and spathaceous calyces; (2) Jacaranda sect. Copaia, species with monothecal anthers and cupular calyces; (3) Jacaranda sect. Jacaranda, species with monothecal anthers and campanulate calyces; and (4) Jacaranda sect. Dilobos, species with dithecal anthers and cupular calyces, and including more than half of the species of the genus, all restricted to Brazil. Conclusions: As circumscribed here, Jacarandeae includes only a broadly defined Jacaranda divided into four sections. Each section is defined by a unique combination of anther and calyx morphologies.
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American Journal of Botany 106(12): 1589–1601, 2019; © 2019 Botanical Society of America 1589
Tribe Jacarandeae Fenzl (Bignoniaceae) traditionally consisted of
two genera (Jacaranda and Digomphia) and 50 species of trees and
shrubs that are distributed across the New World tropics (Fenzl,
1841; Gentry, 1992b). Jacaranda Juss. contains 47 species, the major-
ity of species in the tribe, and is distributed from southern Mexico
to northern Argentina (Gentry, 1992a; Lohmann and Ulloa Ulloa,
2006 onward; Da Silva-Castro, 2017; Farias-Singer, 2019 onward).
Members of this genus occupy a wide range of biomes and present
a variety of growth habits, ranging from rainforest canopy trees to
re-adapted xylopodial subshrubs. For example, Jacaranda copaia
(Aubl.) D. Don is a hyperdominant tree in lowland, wet tropical for-
ests from Belize to Bolivia (ter Steege etal., 2013), while Jacaranda
rugosa A.H. Gentry is endemic to the Caatingas of Pernambuco
State, Brazil (Gentry, 1992b; Milet-Pinheiro and Schlindwein, 2009).
Digomphia Benth., with only three species, consists of shrubs re-
stricted to lowland Amazonia and the Guiana shield (Gentry, 1992a;
Lohmann and Ulloa Ulloa, 2006 onward; Farias-Singer and Singer,
Jacaranda species are known to have an array of horticultural
and ethnobotanical uses (Gentry, 1992a). e species of greatest
horticultural importance is Jacaranda mimosifolia D. Don, perhaps
the most widely planted subtropical tree worldwide (Gentry, 1992a).
Described as a “long-lived pioneer species” (Scotti-Santaigne etal.,
2013), J. copaia is a good candidate for reforestation projects due
to its rapid growth in high-light environments and persistence
in mature forests. e Siona Indians of Amazonian Ecuador use
Phylogeny of the Neotropical tribe Jacarandeae
Audrey C. Ragsac1,5 , Rosana Farias-Singer2, Loreta B. Freitas3, Lúcia G. Lohmann4, and Richard G. Olmstead1
Manuscript received 21 May 2019; revision accepted 21 October 2019.
1 Department of Biology and Burke Museum,University of
Washington, Box 355325, Seattle, Washington 98195, USA
2 Fundação Zoobotânica do Rio Grande do Sul, Rua Dr. Salvador
França 1427, 90.690-000, Porto Alegre, Rio Grande do Sul, Brazil
3 Departamento de Genética,Instituto de Biociências,Universidade
Federal do Rio Grande do Sul, P.O. Box 15051, 91501-970, Porto
Alegre, Rio Grande do Sul, Brazil
4 Departamento de Botânica,Instituto de Biociências,Universidade de
São Paulo,Rua do Matão, 277, 05508-090, São Paulo, São Paulo, Brazil
5Author for correspondence (e-mail:
Citation: Ragsac, A. C., R. Farias-Singer, L. B. Freitas, L. G. Lohmann,
and R. G. Olmstead. 2019. Phylogeny of the Neotropical tribe
Jacarandeae (Bignoniaceae). American Journal of Botany 106(12):
PREMISE: The tribe Jacarandeae includes Jacaranda (49 species) and Digomphia (3 species),
two genera of trees and woody shrubs with Neotropical distribution. Jacarandeae is
sister to the rest of the Bignoniaceae, but not much is known about interspecic and
intergeneric relationships within this group.
METHODS: We reconstructed the phylogeny of Jacarandeae using chloroplast (ndhF,
rpl32-trnL, trnL-F) and nuclear (ETS, PPR62) markers. Evolutionary relationships within
Jacarandeae were inferred using Bayesian, Maximum Likelihood, and species tree
approaches. The resulting phylogenetic framework was used as the basis to interpret the
evolution of key morphological character states (i.e., stamen and calyx traits) and revise the
infra-generic classication of the group.
RESULTS: Jacaranda and Digomphia belong to a well-supported clade, with Digomphia
nested within Jacaranda. We propose the necessary taxonomic changes to recognize
monophyletic taxa, including a broadly circumscribed Jacaranda divided into four sections:
(1) Jacaranda sect. Nematopogon, species previously included in Digomphia and united
by divided staminode apices and spathaceous calyces; (2) Jacaranda sect. Copaia, species
with monothecal anthers and cupular calyces; (3) Jacaranda sect. Jacaranda, species with
monothecal anthers and campanulate calyces; and (4) Jacaranda sect. Dilobos, species with
dithecal anthers and cupular calyces, and including more than half of the species of the
genus, all restricted to Brazil.
CONCLUSIONS: As circumscribed here, Jacarandeae includes only a broadly dened
Jacaranda divided into four sections. Each section is dened by a unique combination of
anther and calyx morphologies.
KEY WORDS Bignoniaceae; Digomphia; Jacaranda; Jacarandeae; Neotropical ora;
phylogeny; systematics.
1590 American Journal of Botany
the capsular fruit of J. copaia as tools to shape pottery (Vickers
and Plowman, 1984; Gentry, 1992a). Ethnomedically, species in
Jacaranda have been used as treatment for venereal disease and
ailments of the skin, stomach, and intestines (Gentry, 1992a).
Jacaranone, a chemical constituent present in Jacaranda, has been
shown to have anticancer and antimicrobial properties and was pat-
ented for antitumor activity (Ogura etal., 1977; Gachet and Schuhly,
Two genera within the Bignoniaceae were originally recognized
based on fruit type (Linnaeus, 1753): Bignonia (including Bignonia
caerulea L.) with dehiscent fruit and Crescentia L. with indehiscent
fruit. Subsequent discoveries resulted in the splitting of these gen-
era and the proposal for several new genera, including Jacaranda,
which was described based on B. caerulea (Jussieu, 1789). De
Candolle (1838) accepted many of the newly proposed genera,
while recognizing Linnaeus’ groups as tribes Crescentieae and
Bignonieae, the latter containing two subgroups based on whether
the fruit dehiscence was loculicidal or septicidal.
e tribe Jacarandeae was rst circumscribed by Fenzl (1841),
to include Jacaranda, Distictis Mart. ex Meisn., and Platycarpum
Bonpl. Currently, Distictis is placed in tribe Bignonieae
(Lohmann, 2006), while Platycarpum is currently included in
Rubiaceae. Bentham (1846) initially described Digomphia as a
new genus of Scrophulariaceae, but later placed it in Jacarandeae
(Hooker, 1876). Hooker (1876) recognized four main tribes
in the Bignoniaceae: Bignonieae, Crescentieae, Jacarandeae,
and Tecomeae. He recircumscribed Jacarandeae to include
Jacaranda, Eccremocarpus Ruiz and Pav., Colea Bojer ex Meisn.,
and Parmentiera DC., the last three of which would later be
placed in tribes Eccremocarpeae (or Tourretieae), Coleeae, and
Crescentieae, respectively (Hooker, 1876; Olmstead etal., 2009).
Failing to recognize Jacarandeae, Bureau and Schumann (1896-
1897) returned Jacaranda and Digomphia to Tecomeae. Gentry
(1980, 1992b) adopted Schumanns classication, agreeing that
Hooker’s Jacarandeae was not a “natural group” and retained
Jacaranda and Digomphia in Tecomeae.
e rst molecular phylogeny of the Bignoniaceae recovered the
sole Jacaranda species sampled as sister to all other Bignoniaceae
(Spangler and Olmstead, 1999). A subsequent phylogeny of the
family based on increased sampling and including four species of
Jacaranda, corroborated the initial results (Olmstead etal., 2009).
Based upon both these results, Olmstead etal. (2009) resurrected
Jacarandeae, restricted to Jacaranda and Digomphia. More recent
phylogenetic work provided additional support for the mono-
phyly of the tribe, as well as to its sister-group relationship to the
rest of the family (AppendicesS1, S2; Farias-Singer etal., 2011).
Despite that, our understanding of phylogenetic relationships
among members of Jacarandeae remains poor.
Traits that distinguish Jacarandeae from the rest of Bignoniaceae
include the elongated and glandular staminode, typically bipin-
nately compound leaves (occasionally simply compound, or even
simple), and a broad, dehiscent, oblong to circular or elliptic fruit
attened perpendicular to the septum (Fig.1). Furthermore, most
taxa in Bignoniaceae have 2n = 40 chromosomes, but Jacaranda has
2n = 36 (Cordeiro etal., 2016).
Gentry (1992b) suggested that Digomphia should be recog-
nized as its own genus based on dierences in staminode apex
and leaf architecture. Digomphia has simple or simply pinnate
compound leaves, while most Jacaranda species have bipin-
nately compound leaves (Fig. 1; Gentry, 1992a, b). e apices
of Digomphia staminodia are bifurcating or trifurcating while
staminodia of Jacaranda are entire (Fig.1). e purpose of these
showy staminodia is uncertain, but they seem to be multifunc-
tional (Guimarães etal., 2008). In a pollination study of J. rugosa,
staminode presence was associated with signicantly higher pol-
linator visitation frequency compared to owers with staminodia
experimentally removed, suggesting that showy staminodia in
this tribe are important for pollinator attraction (Milet-Pinheiro
and Schlindwein, 2009). A pollination study of Jacaranda oxy-
phylla Cham. suggested that secretions from glandular trichomes
of staminodia attract pollinators, and that the physical location of
staminodia in owers play a role in increased pollinator contact
with reproductive organs (Guimarães etal., 2008). Most species
in Jacarandeae seem to be pollinated by euglossine bees (Gentry,
1974; Guimarães etal., 2008; Maues etal., 2008; Milet-Pinheiro
and Schlindwein, 2009).
Endlicher (1839) described three sections within Jacaranda
based on number of anther thecae and calyx morphology: (1) Copaia
with one anther sac and tubular, truncate calyces; (2) Hemilobos
with one anther sac and campanulate, ve-lobed calyces; and (3)
Dilobos with two anther sacs and campanulate calyces. Subsequent
classications (de Candolle, 1845; Gentry, 1992b) did not use ca-
lyx characters, recognizing sections only on the number of anther
thecae, Dilobos and Monolobos, with nearly equal numbers of spe-
cies in each group. Biogeographically, this classication segregates
Jacaranda into two main areas of the neotropics; section Monolobos
(hereaer sect. Jacaranda, as required by the ICN, Turland etal.,
2018) characterized by a single anther sac, and widespread through-
out Central America and Amazonia, and section Dilobos, charac-
terized by two anther sacs and found throughout South America,
with greatest diversity in eastern Brazil (Gentry, 1992b). e ab-
sence of one theca in Jacaranda is unique in Bignoniaceae and rare
in the greater core Lamiales (Gentry, 1980; Judd etal., 2015).
e dichotomy based on the number of anther thecae is sup-
ported by dierences in Jacaranda wood anatomy. Jacarandeae
are distinguished from the rest of the family by the narrow vessels,
paratracheal winged-aliform parenchyma, and non-storied rays
(Pace etal., 2015). Within Jacaranda, section Jacaranda has wood
anatomy marked by homocellular and uniseriate rays, while sect.
Dilobos has heterocellular and multiseriate rays (Dos Santos and
Miller, 1997). e only exception to this rule is found in Jacaranda
copaia. Even though J. copaia was assigned to sect. Jacaranda due
to the single anther sac, this species has a unique combination of
anatomical traits that are intermediate between sections (Gentry,
1992b, Dos Santos and Miller, 1997). Specically, J. copaia has mul-
tiseriate rays of homocellular composition, and the widest vessels of
any tree in Bignoniaceae (Pace etal., 2015).
Other clades in Bignoniaceae that have been studied extensively
include Bignonieae (e.g., Lohmann, 2006; Lohmann etal., 2013;
Lohmann and Taylor, 2014), Catalpeae (Li, 2008), Coleeae (Zjhra
etal., 2004; Callmander etal., 2016), Incarvillea (Chen etal., 2005),
and the Tabebuia alliance (Grose and Olmstead, 2007a, 2007b).
Molecular phylogenetics provides strong support for the circum-
scription and monophyly of these clades and provides a foundation
for the current study.
Understanding the evolutionary history of Jacarandeae re-
quires knowledge of relationships among species within this
tribe. e aims of this paper are to: (1) test the monophyly of
Jacarandeae; (2) test the monophyly of the two genera recog-
nized in the tribe, Jacaranda and Digomphia, as well as resolve
December 2019, Volume 106 Ragsac etal.—Phylogeny of Jacarandeae (Bignoniaceae) 1591
relationships among species therein; and (3) revise the classi-
cation of tribe Jacarandeae. To address these goals, we sampled
widely in Jacaranda and Digomphia and built a dataset compris-
ing three chloroplast (ndhF, rpl32-trnL, trnL-F) and two nuclear
(ETS, PPR62) markers, which was used to generate a phylogenetic
framework for the tribe using Bayesian, Maximum Likelihood,
and species tree approaches. Our phylogeny provides a founda-
tion for future investigation of the origin and maintenance of
species diversity within the group, and an improved understand-
ing of Bignoniaceae systematics.
We sampled 35 of the 50 species recognized in the most com-
prehensive treatment of Jacarandeae to date (Gentry, 1992a, b).
is sampling strategy included all three species of Digomphia,
and 32 out of 47 species of Jacaranda (including both subspe-
cies of J. copaia). Furthermore, ve species from related lineages
within Bignoniaceae (Bignonia capreolata L., Crescentia cujete L.,
FIGURE 1. Reproductive and vegetative morphology of Jacaranda and Digomphia. (A) Digomphia ceratophora with a divided staminode apex, spatha-
ceous calyx, and simple leaves. (B) D. ceratophora with round, dehiscent fruits. (C) Jacaranda mimosifolia with dehiscent fruit attened perpendicular
to the septum. (D) J. mimosifolia with a thyrse inorescence and bipinnately compound leaves. (E) Jacaranda ulei with a thyrse inorescence and
cupular calyx. (F) J. mimosifolia with an entire staminode, monothecal anthers, and style. (G) Jacaranda irwinii with simply pinnate compound leaves.
(H) Jacaranda copaia's crown of purple inorescences peeking through wet forest canopy near Inírida, Colombia. Photo credits: Mateo Fernandez (A),
Richard G. Olmstead (B, C, D, H), Rosana Farias-Singer (E, F), and Audrey C. Ragsac (G).
1592 American Journal of Botany
Chilopsis linearis (Cav.) Sweet, Eccremocarpus scaber Ruiz and
Pav., Tabebuia rigida Urb.), and ve species from closely related
lineages within the Lamiales (Paulownia tomentosa (unb.)
Steud., Schlegelia fuscata A.H. Gentry, Schlegelia parviflora
(Oerst.) Monach., Sesamum indicum L., Thunbergia alata Bojer
ex Sims) were also included as outgroups (Lohmann and Ulloa
Ulloa, 2006 onward; Olmstead etal., 2009; Refulio and Olmstead,
2014; Appendix1). Of the ingroup taxa, ve were collected in
the eld, six were obtained from plants in cultivation, and the
remaining samples were gathered from herbaria. Sampling of the
ingroup includes previously published sequences for two taxa
(Olmstead etal., 2009), with the remaining taxa representing new
DNA extraction, PCR amplication, and sequencing
Total genomic DNA was extracted from silica-dried material or her-
barium tissue using either the CTAB method (modied from Doyle
and Doyle, 1987) or DNeasy Kits (Qiagen, Valencia, California,
USA). ree chloroplast regions (ndhF, rpl32-trnL, trnL-F) were
amplied via polymerase chain reaction (PCR) using primers
developed in Olmstead and Sweere (1994), Taberlet etal. (1991),
and Zuntini et al. (2013), respectively. Two nuclear regions
(ETS and PPR62) were amplied using the 18s-IGS primer of
Baldwin and Markos (1998) with a custom primer used to amplify
ETS in the Lamiales (Beardsley and Olmstead, 2002) and PPR62
using primers developed in Yuan etal. (2009, 2010). PCR was per-
formed under the following general conditions: 94°C for 2 min,
followed by 35 cycles of 94°C for 30 sec, 50°C for 30 sec, 72°C for
1.5–2.5 min, followed by 72°C for 10 min. Amplied PCR products
were puried using 20% polyethylene glycol (PEG) precipitation
and 70% ethanol washes prior to sequencing. Cycle sequencing re-
actions were performed using BigDye 3.1 (Applied Biosystems Inc.,
Foster City, California, USA) using a standard Applied Biosystems
protocol. Products of sequencing reactions were puried by passing
through Sephadex G-50 columns (Sigma-Aldrich, St. Louis, MO)
or sodium acetate precipitation. An Applied Biosystems genetic an-
alyzer was used to generate raw sequence data.
Phylogenetic analyses
Sequence chromatogram editing and manual alignments were per-
formed in Geneious 10.2.3 (Kearse etal., 2012.). Data were analyzed
in both Maximum Likelihood and Bayesian frameworks in four
ways: (1) concatenated chloroplast loci (ndhF, rpl32-trnL, trnL-F);
(2) each nuclear locus separately; (3) concatenated nuclear loci
(ETS, PPR62); and (4) combined chloroplast-nuclear dataset par-
titioned. Chloroplast and nuclear datasets were analyzed separately
to detect any conicting phylogenetic signal.
Appropriate models of evolution for these data were deter-
mined using jModelTest 2.1.4 (Darriba et al., 2012) using the
Akaike information criterion (AIC). Phylogenetic reconstruc-
tions were performed in a Maximum Likelihood framework us-
ing RAxML 8.0.0 (Stamatakis, 2014), Bayesian framework using
MrBayes 3.1.2 (Ronquist etal., 2012), and Bayesian coalescent
framework using *BEAST 2.4.7 (Bouckaert et al., 2014). One
thousand bootstraps were performed with Maximum Likelihood.
All other parameters were kept at default values set in RAxML
8.0.0. Bayesian analyses used two replicate runs, each consisting
of four chains sampled every 1000 generations. Analyses were
assessed for convergence using Tracer 1.6 (Rambaut etal., 2014).
Analyses were run until convergence diagnostics indicated that
they had reached stationarity. When summarizing consensus
trees over all runs, the rst 25% of sampled trees were discarded
as burn-in.
In *BEAST, the chloroplast loci were treated as a single organ-
ellular (haploid) locus and nuclear loci were treated as autosomal.
Due to the degradation of herbarium specimen DNA, it was di-
cult to achieve complete sequencing coverage for all loci. erefore,
*BEAST was run twice—once including only taxa with full se-
quencing coverage for all loci and once including all taxa with ‘N’
inserted for missing data. e nal analysis used a GTR model
for all datasets, default speciation and clock models, and priors
for mean population size and birth rate were set to gamma distri-
butions with shape = 2. Runs were performed for 500 million gen-
erations, sampling every 5000 trees. TreeAnnotator 2.4.7 (Rambaut
and Drummond, 2017) was used to summarize the sampled trees
into single target trees, with the rst 25% of trees sampled discarded
as burn-in.
Morphological trait mapping
Stamen (monothecal or dithecal; staminodia entire or divided) and
calyx (cupular, campanulate, or spathaceous; persistent or not per-
sistent) morphology was assessed for each taxon (Gentry, 1992b;A.
Ragsac, personal observation). Ancestral state reconstructions were
performed in Mesquite 3.3 (Maddison and Maddison, 2017) on
the maximum clade-credibility tree generated from the *BEAST
tree using the “parsimony ancestral states” option.
Sequence data were derived from three cpDNA regions (ndhF, rpl32-
trnL, trnL-F) and two nDNA regions (ETS, PPR62). Sequences were
submitted to GenBank (Appendix1). Due to diculty in amplify-
ing and sequencing target DNA regions extracted from herbarium
specimens, about 20% of sequences from target loci were partial
or missing from the nal dataset.
For the nuclear dataset, new sequence data were generated for all
48 accessions included. Aer alignment, the total length of align-
ment for each dataset was 457 for ETS and 360 for PPR62, resulting
in a combined nuclear dataset of 817 positions. While the separate
ETS and PPR regions did not have the same sequencing coverage,
tree topologies were in agreement with regards to the major clades
Chloroplast sequences of ndhF and trnL-F for Jacaranda ar-
borea Urb., J. sparrei A.H. Gentry, and outgroups were obtained
from previous studies (Olmstead etal., 2009). Remaining sequences
for ndhF and trnL-F, as well as all rpl32-trnL sequences, were newly
generated for this study. Aer alignment, the total length of align-
ment for each dataset was 2001 for ndhF, 917 for trnL-F, and 1196
for rpl32-trnL, resulting in a combined chloroplast dataset of 4114
positions. When the nuclear regions were added to this dataset, the
resulting concatenated dataset contained 4931 positions.
Appropriate models of evolution were evaluated with JModelTest
2.1.4 and the Akaike information criterion (AIC), which deter-
mined that the GTR+I+G model was the most likely t for ETS
and all chloroplast loci, while GTR+I was the most likely t for the
PPR62 dataset (Darriba etal., 2012).
December 2019, Volume 106 Ragsac etal.—Phylogeny of Jacarandeae (Bignoniaceae) 1593
Phylogenetic trees generated from nuclear and chloroplast
datasets in both Bayesian and ML frameworks support a mono-
phyletic Jacarandeae. In the nuclear tree (Fig.2, AppendicesS3,
S4), four clades emerged, as follows: (1) a monophyletic
Digomphia (PP = 1, BS = 100%); (2) all species from Jacaranda
sect. Jacaranda (PP = 1, BS = 100%), except J. copaia; (3) J. co-
paia (including both subspecies) (PP = 1, BS = 100%); and (4)
a monophyletic Jacaranda sect. Dilobos (PP = 1, BS = 100%).
Jacaranda sect. Jacaranda appears as sister to Digomphia, al-
though poorly supported (PP = 0.63, BS = 72%), while Jacaranda
sect. Dilobos appears as sister to J. copaia and also is poorly sup-
ported (PP = 0.64, BS = 60%).
In contrast, dierences were found in the tree that resulted
from the analysis of the combined cpDNA dataset (Fig. 3,
AppendicesS5, S6). is topology includes six main clades: (1)
a monophyletic J. copaia (PP = 1, BS = 100%); (2) all Caribbean
species of Jacaranda sect. Jacaranda (PP = 0.98, BS = 85%); (3)
the remaining species of Jacaranda sect. Jacaranda (PP = 1,
BS = 71%); (4) Digomphia laurifolia Benth. and Digomphia den-
sicoma (Mart. ex DC.) Pilg. (PP = 1, BS = 73%); (5) Digomphia
ceratophora A.H. Gentry; and (6) Jacaranda sect. Dilobos (PP = 1,
BS = 98%). Neither Jacaranda sect. Jacaranda nor Digomphia ap-
pear as monophyletic, while Jacaranda sect. Dilobos and J. copaia
are monophyletic. e relationships among the six clades that
emerge in the chloroplast tree are largely unresolved, with weak
support for any nodes suggesting relationships among them in
the optimal tree. e previous results of Farias-Singer etal. (2011)
based on two chloroplast loci are consistent with the results ob-
tained here (AppendicesS1, S2).
Concatenated, partitioned analysis of the combined dataset ana-
lyzed in both Bayesian and ML frameworks (Fig.4, AppendicesS7,
S8) recovered the same four clades found in the tree that resulted
from the analyses of the nuclear dataset. Jacarandeae is again in-
ferred to be monophyletic. ere is low probability (PP = 0.7,
BS = 44%) of a clade comprising Jacaranda sect. Dilobos and J. co-
paia. e relationships among this clade, Jacaranda sect. Jacaranda
(except J. copaia), and J. copaia are unresolved (Fig.4).
To unravel incongruence among the results of the three previ-
ously described analyses, a coalescent species tree was generated
in *BEAST (Fig. 5). Like all previous analyses, the resulting tree
FIGURE 2. Bayesian majority rule consensus tree derived from the analysis of the concatenated nuclear dataset with both Bayesian posterior proba-
bility and ML bootstrap support values (* denotes a PP of 1 and bootstrap of 100%) indicated at nodes.
1594 American Journal of Botany
strongly supported a monophyletic Jacarandeae (PP = 1) contain-
ing four clades: (1) Digomphia (PP = 0.67); (2) J. copaia (PP = 1);
(3) Jacaranda sect. Jacaranda (PP = 0.2); and (4) Jacaranda sect.
Dilobos (PP = 1). Relationships among the four clades are unre-
solved. Unique to the *BEAST tree, J. copaia is weakly supported as
sister to the rest of the tribe (PP = 0.4).
Morphological trait mapping on the *BEAST maximum clade
credibility tree (Fig.5) suggested two equally parsimonious hy-
potheses for the evolution of anther thecae in Jacarandeae: (1)
a monothecal anther ancestral state led to the gain of a second
functional anther sac twice, i.e., once on the branch to Digomphia
and once on the branch to sect. Dilobos; and (2) a dithecal anther
ancestral state preceded the loss of a functional anther sac twice,
i.e., once on the branch to J. copaia and once on the branch to
sect. Jacaranda. Undivided staminodia were inferred to be the
ancestral condition in Jacarandeae, with divided staminodia
evolving in Digomphia. Spathaceous calyces of Digomphia and
campanulate calyxes of sect. Jacaranda are likely derived from
cupular calyces, and the persistent calyces of sect. Dilobos are de-
rived from deciduous calyces.
In this study, we used nuclear and chloroplast datasets to recon-
struct the phylogeny of Jacarandeae (Figs. 2‒5, Appendices S1,
S3-S8). e trees provide a consistent picture of the phylogeny of
the tribe, although dierences were detected. Below we summarize
phylogenetic relationships within the tribe and discuss the implica-
tion of those ndings for morphological evolution and the classi-
cation of the group.
Phylogenetic relationships of tribe Jacarandeae
Our primary goal was to resolve the main clades of Jacarandeae
via molecular phylogenetic methods and use them as a basis for
a revised classication. Using concatenated sequence and co-
alescent approaches, the results support that: (1) Jacarandeae
is monophyletic; (2) Digomphia is nested within Jacaranda; and
(3) Jacarandeae consists of four well-supported clades. However,
resolution of the relationships among clades is weak (Figs.2‒5).
Previous classications diered in their methods of grouping
FIGURE 3. Bayesian majority rule consensus tree derived from the analysis of the concatenated chloroplast dataset with both Bayesian posterior
probability and ML bootstrap support values (* denotes a PP of 1 and bootstrap of 100%) indicated at nodes.
December 2019, Volume 106 Ragsac etal.—Phylogeny of Jacarandeae (Bignoniaceae) 1595
taxa in Jacarandeae. While Endlicher (1839) recognized three
groups in Jacarandeae based on a combination of number of an-
ther sacs and calyx morphology, de Candolle (1845) and Gentry
(1992b) recognized two groups based only on number of anther
sacs. Digomphia was not recognized in Jacarandeae until Hooker
(1876), whose circumscription also included other Bignoniaceae
taxa. Olmstead etal. (2009) resurrected Jacarandeae to contain
only Jacaranda and Digomphia based on a hypothesis by Gentry
(1992b) that Digomphia is derived from within Jacaranda.
Our molecular phylogenetic results support the monophyly of
Digomphia and Jacaranda sect. Dilobos. However, our results
do not support the monophyly of Jacaranda sect. Jacaranda,
because J. copaia forms a distinct lineage from the rest of
sect. Jacaranda as reected in Endlicher’s (1839) classica-
tion. Jacarandeae’s monophyly, Digomphias placement within
Jacaranda, and paraphyly of Jacaranda sect. Jacaranda are con-
sistent with the results of Farias-Singer etal. (2011).
Our analyses recovered a monophyletic Digomphia nested
within Jacaranda, supporting the relationship posited by Gentry
(1992b) and the most recent circumscription of Jacarandeae
proposed by Olmstead etal. (2009). Digomphia has dithecal anthers,
as seen in sect. Dilobos. Despite that, Digomphia shows a number
of characters that are not found in any members of Jacaranda such
as the deeply divided staminode and large, foliaceous calyx. A sister
relationship between Digomphia and sect. Dilobos is weakly sup-
ported in trees generated from both *BEAST and the chloroplast
dataset (Figs.3, 5). In the tree that resulted from the analyses of
the nuclear dataset, Digomphia receives modest support as sister to
sect. Jacaranda. In the concatenated tree, the relationship between
Digomphia and other major clades is unresolved. Dos Santos and
Miller (1992) found the wide rays of Digomphia most similar to
those of J. copaia but these taxa do not appear as closely related
in our analyses. However, Dos Santos and Miller (1992) did not
nd any major anatomical dierences between Digomphia and
Jacaranda at the macroscopic or microscopic level, providing
additional evidence that the genus belongs in Jacaranda. Given
the morphological similarities between Jacaranda and Digomphia
and lack of clear evidence of a monophyletic Jacaranda excluding
Digomphia, the genus Jacaranda is best circumscribed broadly, in-
cluding Digomphia.
FIGURE 4. Bayesian majority rule consensus tree derived from the analysis of the concatenated nuclear and chloroplast datasets with both Bayesian
posterior probability and ML bootstrap support values (* denotes a PP of 1 and bootstrap of 100%) indicated at nodes.
1596 American Journal of Botany
We also found incongruence among gene trees and the species
tree with regards to the monophyly of Jacaranda sect. Jacaranda
(excluding J. copaia). is group receives low support in the species
tree and is not monophyletic in the tree that resulted from the analy-
ses of the chloroplast dataset, where it is split into two well- supported
clades, one in South America and the other in the Caribbean.
FIGURE 5. *BEAST maximum clade credibility species tree generated from chloroplast, ETS, and PPR62 loci. Weighted branches subtend clades re-
ceiving > 0.9 posterior probability; numbers indicate support for other nodes. Most parsimonious character histories for anther thecae, staminodia,
calyces, and calyx persistence are mapped onto branches, with character states for each taxon indicated at tips.
December 2019, Volume 106 Ragsac etal.—Phylogeny of Jacarandeae (Bignoniaceae) 1597
Anther thecae evolution
e number of anther thecae present in Jacaranda species has
been the traditional guide to classifying species within the genus.
e major clades generated in the coalescent tree (Fig.5) suggest
two possible directions of thecal evolution: (1) the loss of one
functional anther sac occurred twice from a typical dithecal an-
ther, once in J. copaia and once in sect. Jacaranda; or (2) a mono-
thecal ancestral state reversed to dithecal anthers on the branch
leading to Digomphia and sect. Dilobos. However, because mono-
thecal anthers are unique to Jacaranda in the Bignoniaceae, and
the gain of a functional trait once lost is relatively rare, scenario
one seems more likely. Alternatively, it is possible that our opti-
mal trees do not correctly reconstruct phylogenetic relationships
within the group and the loss of one anther sac was a unique
event without reversal.
Calyx evolution
As suggested by Endlicher (1839), calyx morphology can also
be used as a guide to distinguish Jacaranda species in each of
the four major clades uncovered. e major clades generated
in the coalescent tree (Fig. 5) suggest that campanulate caly-
ces in Jacaranda sect. Jacaranda and spathaceous calyces in
Digomphia each evolved once from a cupular calyx ancestral
state. Furthermore, calyx persistence evolves once on the branch
leading to Jacaranda sect. Dilobos from non-persistent calyces as
the ancestral state (Fig.5).
Taxonomic treatment
ese new molecular phylogenetic results warrant a new classi-
cation for the group. e tribe currently contains two genera,
Jacaranda and Digomphia. e results presented in this study sup-
port a single, well-supported clade including both genera, com-
prised of four well-supported subclades. We resurrect Jacaranda
sect. Nematopogon to house former Digomphia species and propose
that the three sections of Jacaranda designated by Endlicher (1839)
be resurrected. Names and diagnostic traits of the four sections are
provided below:
I. Jacaranda Jussieu, Gen. Pl. 138. 1789. Type: Jacaranda caeru-
lea (L.) Juss.
Leaves usually bipinnate, occasionally pinnate or simple.
Inorescence a few to many-owered terminal or axillary thyrse
or thyrsoid raceme. Flowers with short calyces; corolla blue or
blue-purple to magenta, tubular campanulate above a narrow basal
tube, the staminode elongate, exceeding the stamens. Fruit an ob-
long capsule attened perpendicularly to the septum. Seeds winged,
the wings membranaceous. Distribution: Belize, Guatemala, and the
Antilles to northern Argentina.
1. Jacaranda sect. Copaia Endlicher. Gen. Pl. 711. 1839. Type:
Jacaranda copaia (Aubl.) D. Don.
e clade containing the two subspecies of Jacaranda copaia
(J. copaia subsp. spectabilis (Mart. ex DC.) A.H. Gentry and J.
copaia subsp. copaia) receives strong support across all analyses.
Phylogenetic trees generated from nuclear and concatenated data-
sets suggest a position sister to Jacaranda sect. Dilobos, while the
species tree supports a position sister to the rest of the tribe.
e decision to recognize Jacaranda sect. Copaia is supported
by morphology and distribution that sets it apart from the rest of
the genus. Jacaranda copaia has monothecal anthers, but unlike all
other species in Jacaranda, species in this section have unlobed, cu-
pular calyces, a wood anatomy of multiseriate, homocellular rays
intermediate between sections Jacaranda and Dilobos, and the larg-
est wood vessels of any tree in the Bignoniaceae. Jacaranda copaia is
also the only rainforest canopy tree in Jacaranda that reaches 45 m
in height.
e decision to designate these two taxa as subspecies is
supported by previous monographic and phylogeographic
studies (Gentry, 1992b; Scotti-Saintagne et al., 2013). Based on
their mainly allopatric distribution (except for a small region
of range overlap in northern Brazil), Gentry (1992b) hypoth-
esized that it is likely that two species exist. Furthermore, mi-
crosatellite analyses of multiple populations of both subspecies
revealed evidence of two divergent chloroplast lineages within
J. copaia, corresponding to the ranges of the two subspecies.
Nuclear microsatellites, however, suggest that nuclear variation
between populations supports gene ow between these two di-
vergent chloroplast lineages (Scotti-Saintagne et al., 2013). In
this study, chloroplast analyses (Fig. 4) resulted in a paraphy-
letic Jacaranda copaia subsp. spectabilis, with our single acces-
sion of Jacaranda copaia subsp. copaia nested within the clade,
while nuclear analyses (Fig.2) placed the two accessions of J. co-
paia subsp. spectabilis together, sister to J. copaia subsp. copaia.
Combined analyses showed the same result as the nuclear tree.
ese dierent outcomes lend further support for the population
structuring described in Scotti-Saintagne etal. (2013), and our
decision to keep them all as one species.
Jacaranda copaia subsp. copaia is characterized by elliptic or ob-
long-elliptic leaets with distinct petioles, and capsule valves less
than 7 cm wide. Distribution: South American Guianas. Jacaranda
copaia subsp. spectabilis is characterized by rhomboid-elliptic, ses-
sile to subsessile leaets with acute to acuminate tips, and capsule
valves <6 cm. Distribution: widespread from Amazonia to Central
Ecologically, J. copaia is considered a “long-lived pioneer spe-
cies” in lowland tropical forests, experiencing fast growth in
light-rich environments, but persisting in mature forests (Scotti-
Saintagne et al., 2013). is species also tolerates drought and
can therefore inhabit seasonally dry forests. e broad ecological
tolerance of this species, particularly in its adaptations to xeric
environments, contributes to its being one of the most abundant
tree species in Amazonia (ter Steege etal., 2013), and could have
favored expansion from Amazonia to Central America through a
dry northern dispersal corridor along the Caribbean coast (Scotti-
Saintagne etal., 2013).
2. Jacaranda sect. Jacaranda. Type: Jacaranda caerulea (L.) Juss.
Trees derived from the analyses of the nuclear, concatenated, and
species tree datasets lend strong support for the monophyly of this
group, which contains most of the species formerly included in sect.
Monolobos. e Antillean species of Jacaranda form a clade within
section Jacaranda (Figs.2,4,5). e remaining species include trees
in sub-Amazonian dry areas and the xylopodial subshrub J. decur-
rens Cham.
e ca. 17 species in Jacaranda sect. Jacaranda are united
by the combination of monothecal anthers and a campanulate,
sometimes deeply lobed, calyx, as opposed to the cupular, mi-
nutely-toothed calyces of Jacaranda sect. Copaia. Distribution:
widespread, ranging from Central and South America to the
1598 American Journal of Botany
3. Jacaranda sect. Nematopogon A.P. de Candolle. Prodr. 9:
232. 1845. Type: Jacaranda densicoma Mart. ex DC. Basionym:
Nematopogon densicoma (Mart. ex DC.) Bureau and K. Schum.,
Fl. Bras. 8(2): 396. 1897. Synonym: Digomphia densicoma (Mart.
ex DC.) Pilg.
Jacaranda sect. Nematopogon consists of three species of the
former genus Digomphia. ere is strong support for the mono-
phyly of sect. Nematopogon in the trees that resulted from the
analyses of the nuclear and concatenated datasets, with modest
support in the species tree. is section is distinguished mor-
phologically from the rest of the tribe by the presence of deeply
divided staminode apices and spathaceous calyces. It shares dith-
ecal anthers with Jacaranda sect. Dilobos. ere is disagreement
among nuclear, chloroplast, and concatenated analyses regarding
the position of this clade within the tribe. is clade emerges as
sister to sect. Dilobos in both the chloroplast and species tree
analyses, sister to sect. Jacaranda in the nuclear gene tree, and
remains unresolved relative to the other sections in the combined
concatenated tree. e simple leaved species D. ceratophora is
sister to a clade that contains another simple leaved species (i.e.,
D. laurifolia), and a pinnately compound leaved species (i.e., D.
densicoma), suggesting a loss of compound leaves twice in the
a. Jacaranda ceratophora (A.H. Gentry) Ragsac, comb. nov.
Basionym: Digomphia ceratophora A.H. Gentry, Mem. N.Y.
Bot. Gard. 29: 270. 1978. Type: Venezuela. Amazonas: Cerro
Yapacana, 150 m, Maguire & Wurdack 34523 (holotype, NY; iso-
types, P, VEN).
Jacaranda ceratophora is characterized by 4-6 parted staminode
apices and narrowly elliptic, simple leaves. Distribution: Below
200 m elevation in the white sand savannas of Amazonas, Venezuela
and adjacent Colombia.
b. Jacaranda densicoma Mart. ex DC. Synonyms: Digomphia
densicoma (Mart. ex DC.) Pilg.; Nematopogon densicoma (Mart. ex
DC.) Bureau and K. Schum.
Jacaranda densicoma is the only species in Jacaranda sect.
Nematopogon that has pinnately compound leaves. Distribution:
Guayana highland region of northern Amazonian Brazil and south-
ern Venezuela and Colombia.
c. Jacaranda laurifolia (Benth.) Ragsac, comb. nov. Basionym:
Digomphia laurifolia Benth., Hooker’s London J. Bot. 5: 364. 1846;
Synonym: Nematopogon laurifolius (Benth.) Bureau and K. Schum.,
Fl. Bras. 8(2): 396. 1897. Type: Guyana. Roraima, Schomburgk 1049
Jacaranda laurifolia has bifurcate staminode apices, simple ellip-
tic leaves, and acute fruit apices. Distribution: Above 450 m in the
Guayana highlands of Venezuela, Guyana, and Brazil.
4. Jacaranda sect. Dilobos Endlicher. Gen. Pl. 712. 1839. Type:
Jacaranda jasminoides (unberg) Sandwith. Basionym: Bignonia
jasminoides unberg.
Jacaranda sect. Dilobos is the largest clade supported by all anal-
yses. It consists of ca. 30 species united by the presence of dithecal
anthers, undivided staminodia, and cupular, persistent calyces. e
group contains species with a wide variety of growth habits across
eastern South America, within which many species have endemic
distributions (e.g., J. rugosa in campo rupestre and J. jasminoides
in the Atlantic forest of Brazil). e lack of resolution within this
section suggests that it is undergoing a recent and rapid radiation.
Distribution: South America, concentrated in the Cerrado and
Atlantic forest.
e authors would like to thank L.H.M. Fonseca, and M. Beyer
for assistance in the eld. We also thank F, FTG, HUH, MO, NY,
and US herbaria for providing leaf tissue for DNA extraction. We
would also like to thank P.L. Irving, J. Chau, L. Frost, A.M. Bedoya,
W. Rahfeldt, R. Cramer, and P. Fabre for laboratory assistance. Two
anonymous reviewers provided helpful comments on this manu-
script. e work was supported by National Science Foundation
(NSF) grant DEB-1500905 to A.C.R. and R.G.O., NSF Systematics
grants DEB 1020369 to R.G.O., NSF GRFP and NSF GROW
Awards to A.C.R., Botanical Society of America and American
Society of Plant Taxonomists Graduate Research Awards to A.C.R.,
University of Washington Graduate Opportunities and Minority
Achievement Fellowship to A.C.R., and Achievement Rewards
for College Scientists Fellowship to A.C.R., a collaborative
FAPESP/NSF/NASA grant on the “Assembly and evolution of the
Amazonian biota and its environment” (FAPESP 2012/50260-6) to
L.G.L., and Conselho Nacional de Desenvolvimento Cientíco e
Tecnológico (CNPq) through Pq grants to L.G.L. (310871/2017-4),
L.B.F. (504981/2008-0), and R.F.S. (151978/2008-5).
Additional Supporting Information may be found online in the
supporting information tab for this article.
APPENDIX S1. Bayesian majority rule consensus tree derived
from the analysis of the concatenated chloroplast loci ndhF and
trnL-F (Farias-Singer etal., 2011). e clade highlighted in gray
is Jacarandeae. Bayesian posterior probability (PP) and ML boot-
strap support (BS) values are indicated at nodes (* denotes a PP of
1 and bootstrap of 100%). Outgroups are taxa from other tribes of
Bignoniaceae and families of Lamiales.
APPENDIX S2. Voucher information of samples used to generate
the tree in AppendixS1. See Index Herbariorum (http://sweet gum. ce/ih/) for herbarium information.
APPENDIX S3. Bayesian majority rule consensus tree derived
from the analysis of the concatenated nuclear dataset. Numbers on
nodes indicate posterior probabilities.
APPENDIX S4. Maximum likelihood tree derived from analysis of
the concatenated nuclear dataset. Numbers on nodes indicate boot-
strap values.
APPENDIX S5. Bayesian majority rule consensus tree derived
from the analysis of the concatenated chloroplast dataset. Numbers
on nodes indicate posterior probabilities.
APPENDIX S6. Maximum likelihood tree derived from analysis of
the concatenated chloroplast dataset. Numbers on nodes indicate
bootstrap values.
APPENDIX S7. Bayesian majority rule consensus tree derived
from analysis of the concatenated nuclear and chloroplast datasets.
Numbers on nodes indicate posterior probabilities.
APPENDIX S8. Maximum likelihood tree derived from analysis
of the concatenated nuclear and chloroplast datasets. Numbers on
nodes indicate bootstrap values.
December 2019, Volume 106 Ragsac etal.—Phylogeny of Jacarandeae (Bignoniaceae) 1599
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APPENDIX 1. Voucher information for sampled taxa. See Index Herbariorum (http://sweet ce/ih/) for herbarium information.
Taxa Locality
Collector and Collection
Bignonia capreolata L. RBG Kew, UK, living acc.
no voucher FJ887855.1 MN447857 FJ870021.1 MN445199 MN447816
Chilopsis linearis D. Don RSA Bot Gardens,
California, USA, in cult.
no voucher FJ887856.1 MN447858 FJ870024.1 MN445200 MN447817
Crescentia cujete L. Puerto Rico S.O. Grose PR 009 (WTU) MN416908 MN447859 MN445244 MN445201 MN447817
Digomphia ceratophora A.H.
Guainia, Inírida,
A.M. Bedoya 616 (WTU) MN416909 MN447864 MN445247 MN445209 MN447822
Digomphia densicoma (Mart.
ex DC.) Pilg.
Amazonas, Brazil P. Acevedo-Rodriguez
14674 (US)
MN416910 MN447865 MN445248 MN445207 MN447820
Digomphia laurifolia Benth. Bolivar, Venezuela O. Huber 11149 (NY) MN416911 MN447866 MN445249 MN445208 MN447821
Eccremocarpus scaber Ruiz
& Pav.
Seattle, Washington,
USA. escaped from
R.G. Olmstead 07-155
MN416912 MN447860 MN445245 MN445202 MN447819
Jacaranda acutifolia Bonpl. Santa Cruz, Bolivia M. Nee 50718 (NY) MN416913 MN447867 MN445250 MN445210 MN447823
Jacaranda arborea Urb. Jardin Botanico
Nacional, Havana,
Cuba, in cult.
R. G. Olmstead 1996-96
FJ887866.1 - FJ870039.1 MN445211 MN447824
Jacaranda bracteata Bureau &
K. Schum.
Rio de Janeiro, Brazil W. Bensen 39 (F) MN416914 MN447868 MN445251 MN445212 MN447825
Jacaranda brasiliana (Lam.)
Bahia, Brazil H. Irwin et al. 31597 (F) MN416915 MN447869 MN445252 MN445213 MN447826
Jacaranda caerulea (L.) Juss. Fairchild Tropical
Botanic Gardens,
Florida, USA, in cult.
T. Berman 2013-001 (FTG) MN416916 MN447870 MN445253 MN445214 MN447827
Jacaranda campinae A.H.
Gentry & Morawetz
Amazonas, Brazil C.A. Cid Ferreira 5816 (NY) MN416917 MN447871 MN445254 MN445215 MN447828
Jacaranda caroba (Vell.) DC. University of São Paulo,
São Paulo, Brazil, in
Living Acccession (SPF) MN416942 MN447872 MN445255 MN445242 MN447852
Jacaranda copaia ssp. copaia
(Aubl.) D. Don
Amapá, Brazil D.C. Daly 3937 (NY) MN416919 MN447874 MN445258 MN445217 MN447830
Jacaranda copaia ssp.
spectabilis (Mart. ex A.P. DC.)
A.H. Gentry
Roraima, Brazil M.J. Hopkins 620 (GH) MN416918 MN447873 MN445257 MN445216 MN447829
Jacaranda copaia ssp.
spectabilis (Mart. ex DC.)
A.H. Gentry
Pará, Brazil L.H.M. Fonseca 408B (SPF) MN416920 MN447875 MN445259 MN445218 MN447831
December 2019, Volume 106 Ragsac etal.—Phylogeny of Jacarandeae (Bignoniaceae) 1601
Taxa Locality
Collector and Collection
Jacaranda crassifolia
Rio de Janeiro, Brazil G. Gottsberger 11-12878
MN416921 MN447876 MN445256 MN445219 MN447832
Jacaranda cuspidifolia Mart.
ex DC.
Fairchild Tropical
Botanic Gardens,
Florida, USA, in cult.
T. Berman 2013-003 (FTG) MN416922 MN447877 MN445260 MN445220 MN447833
Jacaranda decurrens Cham. São Paulo, Brazil J.A. Lombardi 6445 (NY) MN416923 MN447878 MN445261 MN445221 MN447834
Jacaranda ekmanii Alain Dominican Republic Liogier (GH) MN416924 MN447879 MN445262 MN445222 MN447835
Jacaranda glabra (DC.)
Bureau & K. Schum.
Napo, Ecuador D. Irvine 832 (F) - - - MN445223
Jacaranda grandifoliolata A.H.
Bahia, Brazil L.H.M. Fonseca 389 (SPF) MN416943 MN447880 MN445263 MN445243 MN447853
Jacaranda irwinii A.H. Gentry Bahia, Brazil L.H.M. Fonseca 383 (SPF) MN416925 MN447881 MN445264 MN445224 MN447837
Jacaranda jasminoides
(Thunb.) Sandwith
Parnamirim, Brazil P. da C. Gadelha Neto 3440
MN416926 MN447882 MN445265 MN445225 MN447838
Jacaranda micrantha Cham. Brazil A. Kegler 383 (MO) MN416927 MN447883 MN445266 MN445226 MN447839
Jacaranda microcalyx A.H.
Bahia, Brazil A.H. Gentry 49997 (MO) MN416928 MN447884 MN445267 MN445227 MN447840
Jacaranda mimosifolia D. Don Bahia, Brazil, in cult. L.H.M. Fonseca 371 (SPF) MN416929 MN447885 MN445268 MN445228 MN447841
Jacaranda montana
São Paulo, Brazil K. Kubitzki 81-27 (NY) MN416930 MN447886 MN445269 MN445229 MN447842
Jacaranda mutabilis Hassl. Paraguay N. Soria 1785 (MO) - MN447887 MN445270 MN445230 MN447843
Jacaranda obovata Cham. Minas Gerais, Brazil R.M. Castro 610 (MO) MN416931 MN447888 MN445271 MN445231 -
Jacaranda obtusifolia Bonpl. Colombia L. Frost 13-013 (WTU) MN416932 MN447889 MN445272 MN445232 MN447844
Jacaranda oxyphylla Cham. Paraná, Brazil A. Krapovickas 40766 (GH) MN416933 MN447890 MN445273 MN445233 MN447845
Jacaranda paucifoliolata
Mart. ex DC.
Minas Gerais, Brazil V.C. Souza 10154 (MO) MN416934 MN447891 MN445274 MN445234 -
Jacaranda poitaei Urb. Peravia, Dominican
A.H. Gentry 50507 (MO) MN416935 - MN445275 MN445235 -
Jacaranda praetermissa
Brazil R.S. Albino 11474 (MO) MN416936 MN447892 MN445276 MN445236 MN447846
Jacaranda puberula Cham. Rio de Janeiro, Brazil V.F. Mansano 07-406 (MO) MN416937 MN447893 MN445277 MN445237 MN447847
Jacaranda racemosa Cham. Minas Gerais, Brazil V.C. Souza 8300 (MO) MN416938 MN447894 MN445278 -
Jacaranda rufa Silva Manso Vaca Diez, Beni, Bolivia: J.C. Solomon 7732 (MO) MN416939 MN447895 MN445279 MN445238 MN447848
Jacaranda simplicifolia K.
Schum. ex. Bureau & K.
Maranhão, Brazil R.S. Rosario 1073 (NY) MN416940 MN447896 - MN445239 MN447849
Jacaranda sparrei A.H. Gentry Waimea, Hawaii, USA,
in cult.
H. Descimon s.n. (MO) AF102631.1 MN447897 FJ870040.1 MN445240 MN447850
Jacaranda ulei Bureau & K.
Pará, Brazil L.H.M. Fonseca 406 (SPF) MN416941 MN447898 MN445280 MN445241 MN447851
Paulownia tomentosa Steud. University of
Arboretum, Seattle,
Washington, USA, in
R.G. Olmstead 88-008
L36406.1 - - MN445203 MN447854
Schlegelia fuscata A.H. Gentry Ibarra, Imbabura,
J.L. Clark 8578 (MO) HQ384828.1 MN447861 - MN445204 MN447855
Schlegelia parviflora (Oerst.)
Aragua, Venezuela A.H. Gentry 14221 (MO) L36410.1 - MN445246 MN445205 -
Sesamum indicum L. Côte d’Ivoire G. A. Ambe 48 (MO) L36413.1 MN447862 AF479010.1 MN445206 MN447856
Thunbergia alata Bojer x Sims Denver Botanical
Garden, Denver,
Colorado, USA, in cult.
no voucher - MN447863 KT075029.1 - -
APPENDIX 1. (Continued)
... No Brasil, está representada por 11 gêneros e 54 espécies, oito destas endêmicas (Lohmann et al. 2020). A Tribo Jacarandeae é constituída pelo gênero Jacaranda Juss., que inclui árvores e arbustos com flores caracterizadas pela presença de um estaminódio glandular alongado (Ragsac et al. 2019). No Brasil, ocorrem 39 espécies, das quais 34 são endêmicas (Lohmann et al. 2020). ...
... A corola é amarelo-intenso, com guias de néctar avermelhados. Jacaranda inclui 52 espécies distribuídas da Guatemala às Antilhas, alcançando o norte da Argentina (Ragsac et al. 2019). No Brasil, está representado por 36 espécies, 32 das delas endêmicas, sendo encontrado em todos os biomas (Lohmann et al. 2020). ...
... Este gênero caracteriza-se pelas folhas pinadas a bipinadas, raque alada, canaliculada ou subalada, corolas azul-arroxeadas ou lilás-magenta, presença de estaminódio desenvolvido e glandular, maior que os estames férteis, e fruto cápsula, comprimido perpendicularmente ao septo, oblongo-obovoide (Gentry 1992a ;Ragsac et al. 2019). (Lam.) ...
... In the past decades our knowledge of phylogenetic relationships within members of the Bignoniaceae has improved substantially thanks of phylogenetic reconstructions based on molecular data to the entire family (Spangler & Olmstead 1999, Olmstead et al. 2009), its main tribes (Zjhra et al. 2004, Lohmann 2006, Grose & Olmstead 2007a, Li 2008, Callmander et al. 2016, Ragsac et al. 2019, or key genera (Kaehler et al. 2012, 2019, Fonseca & Lohmann 2015, Medeiros & Lohmann 2015, Fonseca & Lohmann 2018, Thode et al. 2019, Carvalho-Francisco & Lohmann 2020. These phylogenetic reconstructions formed the basis for a series of new taxonomic treatments for the family (Grose & Olmstead 2007b, Lohmann & Taylor 2014. ...
... In this latter, it is remarkable that Jacaranda copaia (Aubl.) D.Don, the only species with anatomically intermediate wood, forms a separate lineage (Dos Santos & Miller 1997, Ragsac et al. 2019. There are also wood characters that consistently help delimit the clades most similar to Astianthus, i.e., Catalpeae and Tecomeae s.s. ...
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Background: Astianthus is a monospecific arborescent genus of Bignoniaceae that occur in the Pacific Coast of central Mexico and northern Central America, where it grows in dense populations along riversides. Its phylogenetic placement has remained controversial since Astianthus has unusual morphological characters such as a four-loculed ovary, and simple, pulvinate, verticillate leaves. Methods: Here we used three plastid markers ndhF, rbcL, and trnL-F, wood, and bark anatomical data to investigate the phylogenetic placement of Astianthus and assign it to one of Bignoniaceae’s main clades. Results: Our molecular phylogenetic analyses indicated that Astianthus belongs in tribe Tecomeae s.s., where other charismatic Neotropical Bignoniaceae genera such as Campsis and Tecoma are currently placed. Wood and bark anatomy support this placement, as Astianthus reunites a unique combination of features only known from members of Tecomeae s.s., such as storied axial parenchyma, the co-occurrence of homo- and heterocellular rays, septate fibers, and scattered phloem fibers in the bark. Conclusions: The placement of Astianthus within Tecomeae s.s. provides further support to previous proposals for the Neotropical origin of this Pantropical tribe.
... Jacaranda mimosifolia D. Don, belongs to the Bignoniaceae family, is a conspicuous ornamental tree feature of numerous metropolises in tropical and sub-tropical countries [81,82]. Although it is reported to be native to South America, similar to various ornamental trees, it was very extensively introduced throughout the centuries. ...
The progressive increase in the industrialisation, consequently the energy consumption and carbon footprint reacquaint the sustainable and environmental credentials of hydrated lime mortars and re-congregate the implications of such binders in masonry construction. The low water retaining characteristics of hydrated lime mortars that often results in insufficient bond development with the subsequent substrate, were manipulated through the incorporation of powdered Jacaranda seed pods and silica fume to establish mortar-masonry optimisation in practice. Although there are numerous studies addressing the agricultural waste incorporation in cement-based materials, the incorporation of Jacaranda seed pods on construction materials have not been reported in the literature. Characterisation study comprising the elemental composition and physical properties of Jacaranda seed pods were essential in assessing the compatibility of pods utilisation in hydrated lime mortars. The paper reports authentic experimental research results on the utilisation of Jacaranda seed pods as a binder substitute in hydrated lime mortars concerning the effect of dewatering on the long-term engineering properties and sustainability indices. The utilisation of powdered Jacaranda seed pods and silica fume demonstrated the eligibility of manipulating the strong water retaining characteristics of hydrated lime mortars to attain improved mortar-substrate optimisation in masonry construction. It is essentially shown in the paper that the degree of dewatering, governed by the incorporation of powdered Jacaranda seed pods and silica fume herein, could not be underestimated as this phenomenon yields approximately 18% increase in the compressive strength of hydrated lime mortars comprising both at short- and long-terms. Although the incorporation of both the powdered Jacaranda seed pods and silica fume decreased the water penetration depth and porosity of mortars by ~11% and 17% respectively, dewatering, an inevitable incident that occurs as soon as the freshly mixed mortars are placed on dry absorbent substrates, further yields a greater decrease in these physical measurements at 1 year. Incorporation of powdered Jacaranda seed pods and silica fume generated a crucial improvement on cost efficiency and eco-strength efficiency of mortars both at 91 days and 1 year. The results have shown that more than 30% reduction on the carbon footprint could be established through this practice. It is essentially established in the paper that the degree of dewatering, experienced at freshly-mixed stage, determines the authentic performance of hydrated lime mortars in masonry construction and governs the veritable sustainability analysis to be performed. The results reported in this paper does not only encourage the re-introduction of the hydrated lime binder in construction practice but also assert a cleaner alternative waste management route for the agricultural wastes.
... Bignoniaceae circumscription corroborated previous findings and included the tribe Jacarandeae (Fig. 3). The presence of the tribe Jacarandeae inside Bignoniaceae was previous reconstructed with modest support of bootstrap (Olmstead et al., 2009) or with maximum support of bootstrap and posterior probabilities in a study focusing on the tribe itself and not sampling the gene matK (Ragsac et al., 2019). Here a robust clade comprising Bignoniaceae was recovered and matK was the only incongruent region (Figs. ...
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Lamiales is one of the most intractable orders of flowering plants, with several changes in family composition, and circumscription throughout history. The order is worldwide distributed, occurring in tropical forests and frozen habitats. In this study, a comprehensive phylogeny of Lamiales was reconstructed using DNA sequences. The tree was used to infer dispersal patterns, focusing on the tropics and extratropics. Molecular and species geographic data available from public repositories were combined to address both objectives. A total of 6,910 species, and 842 genera of Lamiales were sampled using the Python tool PyPHLAWD. The tree was inferred using RAxML, and recovered a monophyletic Lamiales. All 26 families were recovered as monophyletic with high support. The families Bignoniaceae, and Plantaginaceae are remarkable examples. The first emerged as monophyletic and included tribe Jacarandeae, while the later emerged as monophyletic in its sensu lato and included both the tribes Angelonieae, and Gratioleae. Distribution points for all species were retrieved from GBIF. After filtering, 1,136,425 records were retained. Species were coded as present in extratropical or tropical environments. The in and out of the tropics dispersal patterns were inferred using a maximum likelihood approach that identifies hidden rate changes. The model recovered higher rates of transition from extratropics to tropics, estimating two rates of state transitions. When ancestral states are considered, more discrete transitions from extratropics to tropics were observed. The extratropical state was also inferred for the crown node of Lamiales and old nested nodes, revealing a rare pattern of transitions to the tropics throughout the upper Cretaceous and Tertiary. A significant phylogenetic signal was recovered for the in and out of the tropics dispersal patterns, showing that state transitions are not frequent enough to erase the effect of tree structure on the data.
... Jacaranda mimosifolia D. Don is a deciduous tree belonging to the Bignoniaceae family that originated in South America (Mostafa et al. 2014;Ragsac et al. 2019). It is now widely cultivated in the tropics and subtropics throughout the world (Gentry 1992). ...
Full-text available
Jacaranda mimosifolia D. Don is a deciduous tree widely cultivated in the tropics and subtropics of the world. It is famous for its beautiful blue flowers and pinnate compound leaves. In addition, this tree has great potential in environmental monitoring, soil quality improvement, and medicinal applications. However, a genome resource for J. mimosifolia has not been reported to date. In this study, we constructed a chromosome-level genome assembly of J. mimosifolia using PacBio sequencing, Illumina sequencing, and Hi-C technology. The final genome assembly was ~707.32 Mb in size, 688.76 Mb (97.36%) of which could be grouped into 18 pseudochromosomes, with contig and scaffold N50 values of 16.77 and 39.98 Mb, respectively. A total of 30,507 protein-coding genes were predicted, 95.17% of which could be functionally annotated. Phylogenetic analysis among 12 plant species confirmed the close genetic relationship between J. mimosifolia and Handroanthus impetiginosus. Gene family clustering revealed 481 unique, 103 significantly expanded, and 16 significantly contracted gene families in the J. mimosifolia genome. This chromosome-level genome assembly of J. mimosifolia will provide a valuable genomic resource for elucidating the genetic bases of the morphological characteristics, adaption evolution, and active compounds biosynthesis of J. mimosifolia.
... The impression on the leaves places the plant close to J. mimosifolia, the sister species of J. acutifolia (Ragsac et al. 2019), occurring in Peru exclusively as an introduced ornamental, currently extremely common in Lima. However, the smaller pinnae (Fig. 3) and the deeper blue colour of the flowers (Fig. 4) put our tree decisively within the variability of J. acutifolia. ...
Full-text available
Nine nomenclatural acts by Antonio Raimondi are assessed and commented. These include a new genus, six new species and two new combinations that are absent from or incorrectly cited in major databases. A new combination, Jacaranda acutifolia var. punctata is proposed for an endemic plant from central Peru. Lastly, Jacaranda punctata Raimondi and Puya raimondii Harms are neotypified and lectotypified, respectively.
... Although there is a fruitful history of population genetic and phylogenetic research on tropical forest trees using allozymes, microsatellite loci, and chloroplast loci (Hamrick and Murawski 1991;Lowe 2005;Lohmann 2006;Olmstead et al. 2009;Ragsac et al. 2019) modern genomic approaches have rarely been used to study these communities (c.f. Collevatti et al. 2019;Brousseau et al. 2020). ...
Full-text available
The lack of genomic resources for tropical canopy trees is impeding several research avenues in tropical forest biology. We present genome assemblies for two Neotropical hardwood species, Jacaranda copaia and Handroanthus (formerly Tabebuia) guayacan, that are model systems for research on tropical tree demography and flowering phenology. For each species, we combined Illumina short-read data with in vitro proximity-ligation (Chicago) libraries to generate an assembly. For J. copaia, we obtained 104X physical coverage and produced an assembly with N50/N90 scaffold lengths of 1.020 Mbp/0.277 Mbp. For H. guayacan, we obtained 129X coverage and produced an assembly with N50/N90 scaffold lengths of 0.795 Mbp/0.165 Mbp. J. copaia and H. guayacan assemblies contained 95.8% and 87.9% of benchmarking orthologs, although they constituted only 77.1% and 66.7% of the estimated genome sizes of 799 Mbp and 512 Mbp, respectively. These differences were potentially due to high repetitive sequence content (> 59.31% and 45.59%) and high heterozygosity (0.5% and 0.8%) in each species. Finally, we compared each new assembly to a previously sequenced genome for H. impetiginosus using whole-genome alignment. This analysis indicated extensive gene duplication in H. impetiginosus since its divergence from H. guayacan.
Full-text available
Most taxa in the Bignoniaceae have 2n = 40, but the basal clade Jacarandeae has 2n = 36, suggesting that x = 18 is the ancestral basic number for the family. Variations in heterochromatin band patterns in genera that are numerically stable, such as Jacaranda, could facilitate our understanding of the chromosomal and karyotypic evolution of the family. We characterized heterochromatin distributions in six Jacaranda species using chromomycin A3 (CMA) and 4’6-diamidino-2-phenylindole (DAPI). All of them had 2n = 36, including first counts for Jacaranda bracteata Bureau & K. Schum., Jacaranda irwinii A.H. Gentry, Jacaranda jasminoides (Thunb.) Sandwith, and Jacaranda rugosa A.H. Gentry. Their karyotypes had four to eight terminal CMA⁺/DAPI– bands per monoploid set. In the section Monolobos, Jacaranda brasiliana (Lam.) Pers. had eight terminal bands and Jacaranda mimosifolia D. Don had four; in the section Dilobos, J. bracteata had six bands per monoploid set, with the other species having five. While three species in the section Dilobos had the same number of terminal bands, J. irwinii had two additional pericentromeric bands and a proximal heterozygotic band, and J. bracteata had two distended CMA bands. The consistent records of 2n = 36 in Jacaranda may represent a plesiomorphic condition for the Bignoniaceae; therefore, the family originated from an ancestor with x = 18. However, 2n = 36 may represent a derived condition, and the family could have had an ancestral basic number of x = 20 that is still conserved in most representatives of the family.
Jacaranda heterophylla M.M.Silva-Castro, a new species of Jacaranda, is described and illustrated. This species is only known from a restricted area of Caatinga located between Lençóis and Palmeiras, at the Chapada Diamantina (Bahia, Brazil). It resembles J. grandifoliolata by the elliptic to obovate leaflets and J. jasminoides by the leaves that are pinnate at the base and bipinnate at the apex. However, J. heterophylla differs from J. grandifoliolata by the higher number of leaflets (17–19 vs. 3–5) and from J. jasminoides by the longer leaflets (6–15 cm vs. 1–4 cm). Information on its conservation status, distribution, and phenology are provided.
This study presents the most complete generic phylogenetic framework to date for the tribe Coleeae (Bignoniaceae), which is endemic to Madagascar and the other smaller islands in the western part of the Indian Ocean. The study is based on plastid and nuclear DNA regions and includes 47 species representing the five currently recognized genera (including all the species occurring in the western Indian ocean region). Bayesian and maximum likelihood analyses supported i) the monophyly of the tribe, ii) the monophyly of Phylloctenium, Phyllarthron and Rhodocolea and iii) the paraphyly of Colea due to the inclusion of species of Ophiocolea. The latter genus was also recovered paraphyletic due to the inclusion of two species of Colea (C. decora and C. labatii). The taxonomic implications of the mutual paraphyly of these two genera are discussed in light of morphological evidence, and it is concluded that the two genera should be merged, and the necessary new nomenclatural combinations are provided. The phylogenetic framework shows Phylloctenium, which is endemic to Madagascar and restricted to dry ecosystems, as basal and sister to the rest of the tribe, suggesting Madagascar to be the centre of origin of this clade. The remaining genera are diversified mostly in humid ecosystems, with evidence of multiple dispersals to the neighbouring islands, including at least two to the Comoros, one to Mauritius and one to the Seychelles. Finally, we hypothesize that the ecological success of this tribe might have been triggered by a shift of fruit-dispersal mode from wind to lemur.
Phylogenetics of Chilopsis and Catalpa (Bignoniaceae) was elucidated based on sequences of chloroplast ndhF and the nrDNA ITS region. In Bignoniaceae, Chilopsis and Catalpa are most closely related as sister genera. Our data supported section Macrocatalpa of the West Indies and section Catalpa of eastern Asian and North American continents. Within section Catalpa, Catalpa ovata of eastern Asia form a clade with North American species, C. speciosa and C. bignonioides, while the other eastern Asian species comprise a clade where C. duclouxii is sister to the clade of C. bungei and C. fargesii. The Caribbean species of Catalpa diverged early from the continental species. More studies are needed to test whether the phylogenetic pattern is common in eastern Asian-North American disjunct genera with species in the West Indies.