Simon Et Al 2011 American Journal of Botany

American Journal of Botany 98(7): 1201–1221. 2011.
THE EVOLUTIONARY HISTORY OF MIMOSA (LEGUMINOSAE):

TOWARD A PHYLOGENY OF THE SENSITIVE PLANTS1
Marcelo F. Simon2,3,9, Rosaura Grether4, Luciano P. de Queiroz5,
Tiina E. Särkinen2, Valquíria F. Dutra6,7, and Colin E. Hughes2,8
2Department of Plant Sciences, University of Oxford, South Parks Rd, Oxford, OX1 3RB, U.K.; 3Embrapa Recursos Genéticos e
Biotecnologia, PqEB, Caixa Postal 02372 Brasília, DF 70770-917, Brazil; 4Departamento de Biología, Universidad Autónoma
Metropolitana-Iztapalapa, Apdo. Postal 55-535 09340 México, D.F. México; 5Departamento de Ciências Biológicas,
Universidade Estadual de Feira de Santana, km 03, BR 116, Campus, Feira de Santana, BA 44031-460, Brazil; 6Pós-graduação
em Botânica, Universidade Federal de Viçosa, Rua Dr. Milton Bandeira 336/34, Viçosa, MG 36570-000, Brazil; 7Departamento
de Ciências Biológicas, Universidade Federal do Espírito Santo, Av. Marechal Campos 1468, Vitória, ES 29043-900, Brazil; and
8Institute for Systematic Botany, University of Zurich, Zollikerstrasse 107, 8008 Zurich, Switzerland
• Premise of the study: Large genera provide remarkable opportunities to investigate patterns of morphological evolution and
historical biogeography in plants. A molecular phylogeny of the species-rich and morphologically and ecologically diverse
genus Mimosa was generated to evaluate its infrageneric classification, reconstruct the evolution of a set of morphological
characters, and establish the relationships of Old World species to the rest of the genus.
• Methods: We used trnD-trnT plastid sequences for 259 species of Mimosa (ca. 50% of the total) to reconstruct the phylogeny
of the genus. Six morphological characters (petiolar nectary, inflorescence type, number of stamens, number of petals, pollen
type, and seismonasty) were optimized onto the molecular tree.
• Key results: Mimosa was recovered as a monophyletic clade nested within the Piptadenia group and includes the former mem-
bers of Schrankia, corroborating transfer of that genus to Mimosa. Although we found good support for several infrageneric
groups, only one section (Mimadenia) was recovered as monophyletic. All but one of the morphological characters analyzed
showed high levels of homoplasy. High levels of geographic structure were found, with species from the same area tending to
group together in the phylogeny. Old World species of Mimosa form a monophyletic clade deeply nested within New World
groups, indicating recent (6–10 Ma) long-distance dispersal.
• Conclusions: Although based on a single plastid region, our results establish a preliminary phylogenetic framework for Mimosa
that can be used to infer patterns of morphological evolution and relationships and which provides pointers toward a revised
infrageneric classification.
Key words: biogeography; character evolution; dating analysis; Mimosoideae; molecular phylogeny; trnD-trnT; infrageneric
classification.
“The foliage of M. viva is said to form extensive mats in their patterns of morphological and geographic diversification,
low-lying pasture and, being instantly sensitive to shock, represents a formidable challenge for botanists (Frodin, 2004;
provides a canvas for ephemeral graffiti drawn upon it Rønsted et al., 2007). The fact that most infrageneric classifica-
with a stick or finger.” (Barneby, 1991) tions have been based on small numbers of conspicuous mor-
phological characters that often turn out to be homoplastic, and
The phenomenon of large genera and the fact that most large the frequent lack of sufficient morphological diagnosability that
angiosperm genera remain poorly known, especially in terms of coincides with robustly supported monophyletic clades have
meant that large and complex groups have been considered to
1 Manuscript
be “unclassifiable” (e.g., Carine and Scotland, 2002). However,
received 21 December 2010; revision accepted 13 April 2011.
these issues remain relatively unexplored for the majority of
The authors thank the following people and institutions that helped with
field collections or provision of plant material: E. James, the Millenium
species-rich genera. Large and geographically widespread gen-
Seed Bank Project-Kew Gardens, S. de Faria, M. T. Schifino-Wittmann, N. era can also reveal much more comprehensively than has been
Dahmer, R. Eastwood, J. Sutherland, G. Elliot, F. Reis, B. Boddey, E. possible to date the extent to which phylogenies are geographi-
Gross, G. Lewis (K), G. Salazar (MEXU), C. Proença (UB), S. Harris cally structured and whether or how that relates to ecological
(FHO), A. Martínez, S. Camargo (UAMIZ), M. Sousa, J. Nascimento, T. preferences (Schrire et al., 2009). Finally, such genera, if they
Pennington, M. Luckow, J. Wood, F. Inches (RBGE), M. Atahuachi, and C. are distributed across more than one continent, also provide op-
Fagg, and the authorities of Brazil and Mexico for permission to collect portunities for testing the hypothesis that many transcontinental
plant material (IBAMA authorization n. 02001.007621/2005), and also all disjunctions can be attributed to long-distance dispersal rather
managers for permission to collect in their properties. D. Spence and N. than vicariance (Renner, 2004).
Caithness, Oxford Univ. E-Science Centre for help running BEAST on the
National Computing Grid, the Clarendon Fund, Wolfson College, and the
Mimosa L. is one of the largest genera in the legume family
Systematics Association for support to M.F.S. during his stay at the with more than 500 species distributed mainly in the neotropics
University of Oxford, and the Royal Society for support to C.E.H. with ca. 40 species occurring in the Old World. Mimosa species
9 Author for correspondence (e-mail: msimon@cenargen.embrapa.br) grow in a diverse array of habitats including lowland tropical
rainforest, savanna, tropical and subtropical dry forest and thorn
doi:10.3732/ajb.1000520 scrub, mid-elevation subtropical forest, desert, grassland, and
American Journal of Botany 98(7): 1201–1221, 2011; http://www.amjbot.org/ © 2011 Botanical Society of America
1201
1202 American Journal of Botany [Vol. 98
wetland. The genus is particularly diverse in open vegetation caulon), the inclusion of the genus Schrankia Willd. within
and is rich in narrowly restricted endemics. Major concentra- Mimosa as series Quadrivalves, plus a major reordering into
tions of species diversity and endemism are located in Central series and subseries of the haplostemonous mimosas of Ben-
Brazil, subtropical South America (Paraguay, Argentina, Uru- tham’s section Eumimosa (= sections Mimosa and Calotham-
guay, and southern Brazil), and Mexico, with secondary clus- nos of Barneby). Because Old World species were not included
ters of species diversity in the Andes, the Caribbean, and in Barneby’s (1991) monograph, the placement of these species
Madagascar (Barneby, 1991; Grether et al., 1996; Simon and within the classificatory structure of the American species re-
Proença, 2000; Villiers, 2002). More than a quarter of the spe- main to be determined. A taxonomic account of the Malagasy
cies of Mimosa are restricted to the Cerrado, the savanna biome species did not attempt to relate them to Barneby’s classifica-
of central South America where highland endemics are abun- tion (Villiers, 2002). Despite the marked increment produced
dant (Simon and Proença, 2000). Mexico, with more than 100 by Barneby’s (1991) revision, where 130 new species and
species, is also remarkable for its high levels of Mimosa ende- around 200 new infraspecific taxa were added to the genus, the
mism (Grether et al., 1996). During the preparation of this pace at which new Mimosa taxa continue to be described is re-
study, a draft world checklist of Mimosa was assembled markable. Around 40 new taxa (35 new species) have been de-
(M. Simon, R. Grether, and C. Hughes, unpublished manuscript), scribed and a few nomenclatural updates published since
including ca. 536 described species. Most of them are distrib- Barneby’s revision (Villiers, 1990, 2002; Villareal, 1992; Barneby,
uted in the Americas (496) from the United States to Argentina, 1993, 1997; Turner, 1994a–c; Grether and Martínez-Bernal,
34 occur in Madagascar (32 endemics), and only a few in east 1996; Glazier and Mackinder, 1997; Fortunato and Palese, 1999;
Africa and south Asia (Appendix S1, see Supplemental Data Queiroz and Lewis, 1999; Grether, 2000; Silva and Secco, 2000;
with the online version of this article). Although it is clear that Izaguirre and Beyhaut, 2002, 2003; Atahuachi and Hughes,
most of the globally widespread species of Mimosa are recently 2006; Lefevre and Labat, 2006, Lewis et al., 2010; Simon et al.,
introduced neotropical invasives, it is possible that a few spe- 2010; Morales and Fortunato, 2010; Särkinen et al., 2011).
cies have native pantropical distributions (Barneby, 1991). Alongside his infrageneric classification, Barneby (1991)
Some mimosas have the reputation for being among the proposed a series of hypotheses for the evolution of a number
world’s most notorious invasive weeds, throughout much of of prominent morphological and anatomical characters includ-
tropical Africa, Asia, and Oceania. One species, M. pigra, is ing habit, armature, indumentum, foliage, pollen, and reproductive
among the 100 most invasive organisms on the planet (Lowe et structures (Table 1). Barneby’s set of evolutionary sequences
al., 2000). In contrast to this handful of widespread pantropical for Mimosa, in which primitive and derived character states were
weeds, the majority of species are globally rare, with numerous proposed is explicit, but these have never been subject to a
narrowly restricted endemics (e.g., Simon and Hay, 2003; comprehensive cladistic analysis. A few previous higher level
Särkinen et al., 2011). Mimosas thus span the complete spec- phylogenetic studies of mimosoid legumes using DNA se-
trum of modern geographic range sizes. Many species of Mi- quences have included small numbers of species of Mimosa
mosa are economically and socially important, being used in (Luckow et al., 2003, 2005). An analysis focused on the genus
reforestation (M. caesalpiniifolia), timber production (M. sca- Piptadenia s.l. (Jobson and Luckow, 2007) included nine spe-
brella), as medicinal plants (M. tenuiflora), and as a source of cies of Mimosa and provided strong support for the monophyly
firewood by subsistence farmers (Rachie, 1979; Camargo- of the genus and its placement within the Piptadenia group, as
Ricalde et al., 2001). Their ability to fix nitrogen (Reis et al., previously proposed (Lewis and Elias, 1981) and envisaged by
2010) and grow quickly make them important plants for soil Barneby (1991). A preliminary species-level molecular phylog-
conservation and enrichment, and some species have been used eny sampled 28 mainly southern South American species
in agroforestry systems and ecological restoration (Franco and (Bessega et al., 2008), but the very sparse taxon sampling limits
Faria, 1997; Dhillion and Camargo-Ricalde, 2005). Flowers of its utility to assess Barneby’s classification and hypotheses of
several Mimosa species are also a major source of nectar for character evolution. Here we present a densely sampled mo-
honey production (Villanueva, 1994; Fagundez and Caccavari, lecular phylogeny of Mimosa, including representatives of al-
2006; Sodre et al., 2007). Mimosa is also well known for its most all the major infrageneric groups and about half of the
seismonastic leaves, prompting specific epithets such as “sensi- described species. The evolution of selected morphological
tiva”, “pudica” (sold as an entertaining “pet plant”), “casta” and characters, including several key characters that are prominent
“viva” for species having some of the most striking examples of in defining the sectional classification, plus the variation in seis-
rapid movement of leaves and leaf parts in response to touch. monasty, are investigated. The main goals of this study were to
Thus far, the chemical and molecular bases for leaf movements (1) test the monophyly of the groups proposed in Barneby’s
have been investigated (Allen, 1969; Ueda and Nakamura, infrageneric classification, (2) analyze the relationships of the
2007), but little is known about how many times this trait Old World species to the rest of the genus, (3) provide a com-
evolved in Mimosa. parative framework to investigate the evolution of key morpho-
In his seminal taxonomic account of the legume subfamily logical characters previously used to differentiate major
Mimosoideae, Bentham (1875) divided Mimosa into sections infrageneric groups and test the evolutionary sequences pro-
Habbasia (diplostemonous species) and Eumimosa (haploste- posed by Barneby (1991), and (4) provide a time frame for the
monous species) and proposed a number of minor groups. An diversification of major clades.
authoritative taxonomic monograph for the New World species
built on Bentham’s system and extensively increased both the
infrageneric groups and the number of species (Barneby, 1991). MATERIALS AND METHODS
The main changes proposed by Barneby (1991) to Bentham’s
(1875) system were recognition of section Mimadenia, based Taxon sampling—The analysis presented here includes 284 accessions
on the presence of leaf nectaries, the division of section Hab- of Mimosa, representing taxa from all five sections and 37 of the 41 series
basia (sensu Bentham) into two sections (Habbasia and Bato- proposed by Barneby (1991), plus half of the Old World species and 13
July 2011] Simon et al.—Evolutionary history of MIMOSA 1203
outgroups. Three of the four series not represented here (Ephedroideae, Rondo- Phylogenetic analyses—Parsimony analysis was conduced using the pro-
nianae, and Pseudocymosae) are monospecific, while series Piresianae has just gram Nona (Goloboff, 1993), spawned from Winclada (Nixon, 2001). Tree
five species and could not be sampled due to lack of material. The 259 species searches employed 5000 replicates, holding 50 trees in each search, using the
sampled were chosen to cover the native geographic and ecological range of the tree bisection and reconnection (TBR) strategy and branch swapping, holding a
genus (Appendix S1). Most samples were collected in the field (silica-dried maximum of 100 000 most parsimonious trees. Additional searches for possible
leaves), but herbarium specimens and seedlings grown from seed collections in shorter trees were conducted using the parsimony ratchet (Nixon, 1999), with
the greenhouse were also used. For a few widely distributed species, multiple four independent runs of 200 iterations each, holding five trees per replicate and
accessions were included, either representing specimens from distant locations randomly reweighting 100 characters and constraining 10% of the nodes in
across the range or different infraspecific taxa. Accessions with taxon names, each iteration. Strict consensus bootstrap percentages were computed with
voucher specimens, locations, and GenBank accession numbers are listed in 1000 replicates and 10 tree-bisection reconnections, holding 10 trees per
Appendix 1. Most sequences used here come from previously published stud- replicate.
ies (Simon et al., 2009; Särkinen et al., 2011), while 17 new sequences are A Bayesian analysis for phylogenetic reconstruction and to estimate diver-
published for the first time (GenBank accessions JF694255–JF694271). No- gence times was conducted under the uncorrelated lognormal relaxed molecu-
menclature follows Barneby (1991), Villiers (2002) for the Malagasy species, lar clock approach implemented in the program BEAST version 1.4.8
Gamble (1920) for the Asian species, Brenan and Brummitt (1970) for the (Drummond and Rambaut, 2007). The GTR + G + I nucleotide substitution
African species, with relevant post-Barneby updates (Brummitt, 1993; Turner, model was chosen based on the Akaike information criterion as implemented in
1994a–c; Glazier and Mackinder, 1997; Grether, 2000) and additions (as noted the program Modeltest (Posada and Crandall, 1998). Divergence time estimates
earlier). for Mimosa were based on an initial higher level family-wide analysis of a large
matK data set containing 839 terminals (Simon et al., 2009). Age estimates
(mean and 95% credibility intervals) for the Mimosa crown node (24.0 Ma, CI
DNA extraction, amplification, and sequencing—Total DNA from silica- [18.0, 30.6]) and the section Mimadenia (clade A) crown node (16.6 Ma, CI
dried leaves or herbarium material was isolated using the DNeasy plant mini kit [10.1, 23.6]) obtained from the family-level analysis were used as calibration
(Qiagen, Crawley, UK). After initial screening of a set of 16 plastid DNA se- constraints using a normal distribution prior. Three runs of 107 generations each
quence loci (Appendix S2, see online Supplemental Data), the trnD-trnT inter- were performed, sampling every 104 generations. Results were assessed to have
genic spacer was selected as the most variable and informative plastid locus for reached stationarity and convergence using the program Tracer version 1.4
Mimosa. This region comprises the fragment located between a primer pair (Rambaut and Drummond, 2007), and data from multiple runs were combined
anchored within the trnDguc and trnTggu genes including a noncoding intergenic after exclusion of burn-in trees. All estimated parameters had sufficient effec-
spacer (ca. 85%) and the embedded trnYGUA and trnEUUC genes, and also two tive sample sizes (ESS), in most cases well above the minimum of 200 recom-
regions of mononucleotide repeats (poly A/T). For most taxa, the whole region mended in the BEAST manual.
was amplified in a single PCR reaction using primers trnD2 (designed for this
study) and trnTggu (Shaw et al., 2005). For degraded DNA templates derived
from herbarium material, amplifications were performed using internal primers Evolution of morphological characters—We selected a subset of five con-
trnEuuc and trnYgua (Shaw et al., 2005) in combination with the external primers, spicuous characters (Table 1) that play a central role in Barneby’s classification,
and in some cases another newly designed primer, trnT2. The primers used in to investigate the evolution of these characters as putative synapomorphies that
this study, in 5′ to 3′ orientation, are as follows: trnD2 (GTG TAC AGC ATG might provide diagnosability for well-supported infrageneric groups within Mi-
CAT ATT CTT ACG), trnYgua (CCG AGC TGG ATT TGA ACC A), trnEuuc mosa. In addition, we investigated the evolution of rapid seismonastic leaf
(AGG ACA TCT CTC TTT CAA GGA G), trnTggu (CTA CCA CTG AGT movement in the genus, which was treated here as a morphological character.
TAA AAG GG), and trnT2 (GAC GTA TCG CCG AGT AAT TCC).
Polymerase chain reactions (PCR) were conducted in a total volume of 25 Petiolar nectaries—Extrafloral nectaries are a common feature of many
µL, containing ~5–20 ng of DNA template, 1× Buffer, 0.5 mol/L of betaine, genera of Mimosoideae and Caesalpinioideae, and are often related to mutual-
1.5 mmol/L of MgCl2, 0.1 mmol/L of each dNTP, 0.5 µmol/L of each primer istic relationships between plants and ants, where the plant provides food to
and 0.6 U of Taq polymerase (Yorkshire Bioscience, UK). PCR conditions were ants, and in return, receives protection against herbivores and/or seed predators
94°C for 5 min; 30 cycles of 45 s at 94°C, 1 min at 55°C and 1 min at 72°C; (McKey, 1989). The presence of petiolar nectaries was a key character used by
followed by a final extension of 5 min at 72°C. PCR products were cleaned us- Barneby to distinguish section Mimadenia, which in his interpretation retained
ing exonuclease I and shrimp alkaline phosphatase (Exo/SAP) and sequenced this ancestral condition shared with closely related Piptadenioid sister groups
in four reactions using the two PCR primers and the two internal primers, fol- (Table 1).
lowing Big Dye v. 3.1 chemistry (Applied Biosystems, Warrington, UK). Prob-
lems in sequencing caused by poly A/T regions were overcome by performing
four sequencing reactions. Consensus sequences from the four sequence strands Type of inflorescence—The reduced flowers of Mimosa are grouped into
were assembled using Sequencher (v. 3.1; GeneCodes Corp., Ann Arbor, Mich- compact inflorescences that can be either a capitulum or a spike. Although in-
igan, USA), and then imported into BioEdit (Hall, 1999) for alignment using termediates (ellipsoidal capitula) are found in some species, in most cases, the
the program Clustal W (Thompson et al., 1994; spawned by BioEdit using de- inflorescences are clearly either globose or spicate, and the type of inflores-
fault parameters) and manual edition. Regions of problematic alignment, where cence has been an important character to define groups in Mimosa (e.g., series
homology could not be determined with confidence, were considered unalign- Dystachyae and Leiocarpae). Barneby (1991) argued that the spike is the ple-
able and excluded from the data set. Unambiguous indels were coded using the siomorphic condition in Mimosa (Table 1), but recognized many reversals from
program SeqState (Müller, 2005), following the simple gap coding method of capitulum to spike.
Simmons and Ochoterena (2000) and were used in the parsimony analysis. In-
dels associated with mononucleotide repeats, common features in our data set, Number of stamens—The number of flower parts, and especially the number
were not coded as gaps since these regions are particularly prone to length of stamens, has dictated the tribal classification of the mimosoids and was also
mutations. A full data set including coded indels is available in the database central in the infrageneric classification of Mimosa (Bentham, 1875; Lewis and
TreeBase (http://treebase.org, study accession number S11700). Elias, 1981; Barneby, 1991). Haplostemonous flowers in Mimosa are thought to
Table 1. Summary of evolutionary progressions for morphological traits in Mimosa as hypothesized by Barneby (1991).
Character Anterior / Primitive Posterior / Derived
Petiolar nectary Present, functional Reduced to nonfunctional spicule, obsolete

Inflorescence Axillary; flowers spicate, all hermaphroditic Paniculate, pseudoracemose, radical; flowers capitate, the lower staminate
Corolla 5-(6)-merous; the lobes membranous, 1-nerved 4-, then 3-merous; lobes either thickened or several-nerved
Androecium Diplostemonous, filaments free, connective ovate Haplostemonous, filaments monadelphous, connective orbicular
Pollen In large or moderate, compound tetrads In small, simple tetrads
Fig. 1. Phylogeny of Mimosa based on DNA sequences of the trnD-trnT noncoding plastid locus. The 50% majority-rule consensus tree from a Bayes-
ian analysis. Letters on nodes define well-supported clades discussed in the text, and colors represent geographic distribution. Numbers above and below
nodes are posterior probabilities and bootstrap percentages from the parsimony analysis (shown only for major clades), respectively. The asterisk represents
the only lettered clade not present in the parsimony strict consensus tree.
Fig. 1. Continued
have evolved via suppression of the antepetalous whorl of a putative primitive tive characters. Unalignable regions comprising 137 sites were
diplostemonous androecium, which is predominant in the Piptadenia group, excluded.
and are thus considered to be the derived state (Barneby, 1991; Table 1).
The levels of informative trnD-trnT variation in Mimosa are
high compared to other noncoding plastid regions, in line with
Number of petals—The corolla in Mimosa can have three, four, five, or
more rarely six petals, although some random variation within species has been
findings that the trnD-trnT spacer is one of the most variable
recorded (Barneby, 1991). Based on outgroup comparison with other members plastid regions (Shaw et al., 2007) and thus useful for phyloge-
of the Piptadenia group which all have five petals, Barneby (1991) suggested netic analyses at the species level. Furthermore, in Mimosa, the
that pentamerous flowers are plesiomorphic, with tetramerous and trimerous trnD-trnT locus is ca. 500 bp longer than the average of ca.
flowers representing pogressive apomorphic specializations (Table 1). 1000 bp reported by Shaw et al. (2005) and therefore provides
a larger number of variable sites. In fact, the trnD-trnT locus
Pollen type—Pollen in mimosoids is frequently arranged in polyads. In Mi- represents a promising option to resolve relationships across
mosa, polyads of four (tetrads), eight (bi-tetrads), 12, or 16 grains, have been
extensively characterized (Guinet, 1969; Caccavari, 1986, 1988, 1989, 2002;
the mimosoid subfamily as a whole, given that present mimo-
Lima et al., 2008). Caccavari (1988, 1989) hypothesized that pollen in Mimosa soid phylogenies suffer from lack of resolution and low substi-
evolved from simple small tetrads to large and more complex polyads, contrast- tution rates compared to other legume groups (Luckow et al.,
ing with Barneby’s (1991) idea in which pollen in tetrads occurs in the morpho- 2003, 2005; Lavin et al., 2005).
logically most derived and specialized groups in the genus (Table 1).
Leaf movement—The sensitivity of Mimosa leaves to touch (seismonasty)

Phylogenetic analysis and major Mimosa clades— The par-
varies widely across the genus. Some species close their leaflets almost in- simony analysis discovered the maximum set of 100 000 equally
stantly when touched, while others respond very slowly or are insensitive to any parsimonious trees of 1123 steps (CI = 0.56, RI = 0.92). A par-
stimulus. Despite the continuum in the degree of seismonasty across the genus, simony ratchet found an additional 971 trees of the same length,
we have chosen to recognize only two character states: absence of seismonasty which were combined to produce a strict consensus tree (online
including very slow leaf movements and rapid seismonastic leaf movements, in Appendix S4). The Bayesian analysis produced a very similar
which the leaflets can close in less than five seconds.
The morphological data set (online Appendix S3) is based primarily on pub- topology to the parsimony analysis, although somewhat better
lished studies (Gamble, 1920; Brenan and Brummitt, 1970; Barneby, 1991, resolved (Fig. 1). All the major clades are present in both analy-
1993; Grether and Martínez-Bernal, 1996; Fortunato and Palese, 1999; Grether, ses, except clade X, which is absent in the parsimony tree, but
2000; Villiers, 2002; Atahuachi and Hughes, 2006). Data on pollen type were this does not represent conflicting topology. Most nodes are
also compiled from the literature (Guinet, 1969; Caccavari, 1986, 1988, 1989, more strongly supported in the Bayesian tree than in the parsi-
2002; Grether and Martínez-Bernal, 1996; Flores-Cruz et al., 2006; Martínez-
Bernal, 2003; Lima et al., 2008; Buril et al., 2010) plus new data (R. Grether,
mony tree, even considering that bootstrap percentages are
unpublished data) and were available for 170 of the 297 terminals. Information more conservative than Bayesian posterior probabilities (Alfaro
on seismonastic leaf movements has been gathered mostly from our own field and Holder, 2006). The majority-rule Bayesian tree (Fig. 1)
experience and tests in the greenhouse with more than 30 species, because there serves as the basis for most of the subsequent analyses and dis-
are no systematic published data on this beyond a few records of leaf move- cussions in this paper.
ments reported by Barneby (1991). Although our knowledge about the extent of Mimosa was recovered as monophyletic with robust support in
seismonasty remains limited, the available data provide some preliminary evi-
dence about how many times seismonasty has evolved in Mimosa and how this
both parsimony and Bayesian analyses and includes the former
trait is distributed among groups. members of the genus Schrankia (clade G), which are deeply
Ancestral character state reconstruction analysis was used to investigate the nested within Mimosa in the molecular analysis, corroborating
evolution of these six characters in Mimosa, using trees derived from the trnD- their transfer to Mimosa by Barneby (1991). In addition, there is
trnT Bayesian analysis as a phylogenetic framework. All traits were coded as also strong support for the inclusion of the monospecific genus
discrete bistate or unordered multistate characters and optimized onto the 50% Schrankiastrum Hassler within Mimosa (Barneby, 1984). Al-
majority-rule consensus trees under a maximum parsimony criterion using the
package Mesquite version 2.01 (Maddison and Maddison, 2007). To account though we have not sampled S. insigne Hassler [=M. insignis
for topological uncertainty, the procedure “Trace over trees” was used to sum- (Hassler) Barneby], the position of M. dalyi, its putative sister
marize ancestral state reconstructions over a set of 1000 Bayesian trees sampled species based on morphological evidence, deeply nested within
at stationarity. This approach was used to identify unequivocal changes in char- clade N (Fig. 1), corroborates Barneby’s (1984) interpretation.
acter states across the tree and thus shed light on the morphological evolution The precise sister group relationships between Mimosa and
of Mimosa. In cases where there is variation within a species, for example, in a
species where flowers can have either four or five petals, the species was coded
other genera of the Piptadenia group are still not clear, espe-
as polymorphic. cially in the Bayesian tree. The parsimony strict consensus tree
(Appendix S4) suggests that the Eupiptadenia clade (Piptadenia
sensu Jobson and Luckow, 2007) could be the sister group of
RESULTS AND DISCUSSION Mimosa, although receiving no bootstrap support. In a study fo-
cused on the polyphyletic genus Piptadenia based on matK and
trnD-trnT data set— The aligned trnD-trnT matrix comprises trnL-trnF sequences, Jobson and Luckow (2007) also found
2262 bp after exclusion of regions of ambiguous alignment. A indications for the Eupiptadenia clade as sister to Mimosa, al-
total of 395 (17.5%) substitutions were parsimony-informative, though again with low support. This relationship received mod-
with 361 of these (16.0%) informative within Mimosa. Inter- erate (80%) bootstrap support in a parsimony analysis of three
genic spacers are susceptible to different types of structural mu- concatenated loci (matK, trnL-trnF, and trnD-trnT) for a small
tations such as insertions, deletions, and inversions (Kelchner, subset of 12 Piptadenia group taxa (M. Simon et al., unpub-
2000). Extreme examples in our data set are a 225-bp autapo- lished data). Overall, these results agree with the hypothesis of a
morphic inversion in M. detinens (excluded from the analysis) close relationship between Piptadenia s.l. and Mimosa based on
and a 300-bp deletion in M. dysocarpa. Unambiguous gaps, morphology (Barneby, 1991; Lewis and Elias, 1981) and also
many of them larger than 10 bp, were common across the whole with a higher level legume phylogeny (Simon et al., 2009).
alignment, and a total of 102 informative indels were coded and When we come to examine the new Mimosa phylogeny in
used in the parsimony analysis, making up 21% of all informa- relation to Barneby’s (1991) infrageneric classification, only
sections Mimadenia and Calothamnos (only recovered in the regated it from the other haplostemonous species. Indeed,
parsimony analysis) are monophyletic (Table 2). Section Mi- section Mimosa would be rendered monophyletic if Calotham-
madenia, which shares the plesiomorphy of extrafloral necta- nos is incorporated within it, which would correspond, with just
ries with the outgroup genera, is robustly supported as sister to a few exceptions (see Fig. 2d), to the group of haplostemonous
the rest of Mimosa in the trnD-trnT analysis (Fig. 1), in line mimosas envisaged by Bentham (1875) in his section Eumi-
with Barneby’s classification and hypothesized evolutionary mosa. Similar results, based on a smaller taxon sample, were
sequence. Sections Batocaulon and Habbasia form an exten- found by Bessega et al. (2008).
sive paraphyletic grade with members of the two sections inter- Despite these notable disagreements at sectional level be-
mingled (Fig. 2a). Section Calothamnos is nested within section tween the results presented here and Barneby’s (1991) classifi-
Mimosa and is only well-supported in the parsimony analysis cation, many of Barneby’s series and subseries within sections
due to a shared synapomorphic indel. There is no support for were recovered as monophyletic (Table 2). Insufficient taxon
the idea that section Calothamnos is derived from within sec- sampling or lack of resolution in some parts of the phylogeny
tion Batocaulon, as hypothesized by Barneby (1991), who seg- prevents the evaluation of all infrageneric groups, most notably
within series Mimosa, which was divided by Barneby into 37
subseries. Because of undersampling, these groups are not dis-
Table 2. Infrageneric classification of Mimosa (Barneby, 1991) with cussed further here. We focus our discussion on the 24 well-
the corresponding clades in the molecular phylogeny presented here
(Fig. 1) and the status of sections and series. The monophyly of some
supported clades A–X (Fig. 1) and review these groups in
groups could not be assessed because of lack of resolution in the tree relation to Barneby’s (1991) infrageneric classification (Appen-
or undersampling. * Series not sampled in this study. dix 2).
Neither taxon nor character sampling in this study were de-
Section / Series Corresponding clade Monophyletic signed to test species boundaries or within-species relation-
Mimadenia Barneby A yes
ships. However, there are indications that some species for
Myriadeniae Barneby A monotypic which multiple accessions were included do not form mono-
Glanduliferae Benth. A no phyletic clades. These include M. weberbaueri, M. montana
Revolutae Barneby A monotypic (clade C) (see Särkinen et al., 2011), M. gracilis (clade H), M.
Nothacaciae Barneby A yes setosa, M. claussenii (clade O), M. polycarpa (clades U and
Batocaulon DC. B,C,D,E,F,G,H,I,K,M,N no W), and M. fachinalensis (clade X). More detailed studies with
Distachyae Barneby B no rangewide sampling of multiple accessions and preferably in-
Andinae Barneby B(C) unresolved cluding a nuclear locus will be required to investigate the status
Acanthocarpae Benth. B unresolved of these species and their infraspecific variants.
Acantholobae Barneby B yes
Boreales Barneby B no
Leiocarpae Benth. B,D,N no Evolution of morphological characters— Optimization of a
Bimucronatae Barneby B(C),D,N no set of morphological characters onto the trnD-trnT phylogeny
Leucaenoideae Barneby B unresolved allowed the reconstruction of character states across the tree,
Rubicaules Benth. B,D(E),G no sheding light on the morphological evolution of Mimosa (Figs.
Fagaracanthae Barneby B undersampled 2, 3). Character optimizations were unequivocal for most nodes
Bahamenses Barneby B monotypic
Farinosae Barneby B(C),D no
in the Bayesian trees. The character optimizations across the
Ephedroideae Benth.* — — 1000 different Bayesian trees sampled at stationarity produced,
Echinocaulae Barneby H monotypic in almost all cases, the same result as the findings presented in
Paucifoliatae Benth. H no the majority-rule consensus tree. Levels of homoplasy, as mea-
Glandulosae (Benth.) Barneby L no sured by the consistency index (CI) and the retention index
Stipellares Benth. K yes (RI), varied widely between characters (Table 3), varing from
Auriculatae Barneby M monotypic
Caesalpiniifoliae Benth. D yes
no homoplasy (petiolar nectary) to highly homoplastic (inflo-
Ceratoniae Barneby D undersampled rescence type).
Cordistipulae Barneby I yes
Campicolae Barneby H undersampled Petiolar nectaries—For Mimosa, the presence of extrafloral
Filipedes Barneby H no nectaries is plesiomorphic and restricted to species of section
Quadrivalves Barneby G no Mimadenia (clade A), in line with Barneby’s (1991) hypothesis.
Plurijugae Karsten B,F no
Within Mimosa, the petiolar nectary provides a conspicuous di-
Calothamnos Barneby X unresolved agnostic character for that section. The ancestral character re-
Habbasia DC. J,L,M,O no construction suggests that section Mimadenia retained this
Setosae Barneby O no ancestral character state, which is present in all species of the
Pachycarpae Benth. O no Piptadenia group (outgroups), but absent from the rest of the
Habbasia DC. J yes
Bipinnatae DC. L yes genus Mimosa (Fig. 2b). This finding was consistent across all
Neptunioideae Barneby M no of the sampled Bayesian trees. There is no indication of reac-
Rondonianae Barneby* — — quisition of petiolar nectaries (reversal) in any other lineages
Rojasianae Barneby M undersampled within Mimosa, indicating lack of homoplasy in this character
Pseudocymosae Hassler* — — and its suitability as a diagnostic character (Table 3).
Piresianae Barneby* — — Marazzi et al. (2006) hypothesized that the presence of extra-
Mimosa L. P to X no floral nectaries in Senna (Caesalpinioideae) could represent a
Myriophyllae Benth. X unresolved key innovation in terms of plant defense strategies that pro-
Mimosa L. P,R,S,T,U,V,W,X no
Modestae Benth. Q,R no
moted large-scale diversification and colonization of a wide
range of habitats and climates on different continents. They
Fig. 2. Infrageneric classification and morphological evolution in Mimosa. (a) Mapping of Barneby’s (1991) five sections onto the trnD-trnT gene tree,
showing that most infrageneric sections are not monophyletic. (b–d) Character optimizations onto a 50% majority-rule Bayesian tree showing the evolution
of (b) petiolar nectaries (c) type of inflorescence, and (d) number of stamens.
found that the Senna clade characterized by the presence of ex- suggest additional independent switches to haplostemonous
trafloral nectaries has a much larger number of species than flowers.
clades which lack nectaries, and argued that presence of nectary
could provide a possible evolutionary innovation that prompted Number of petals—Number of petals is another highly ho-
this accelerated diversification (Marazzi and Sanderson, 2010). moplastic character with multiple unequivocal character state
Conversely, it is notable that in Mimosa the derived loss of peti- transitions hypothesized by the optimization analysis (Fig. 3a).
olar nectaries is associated with a much larger clade (515 spe- The ancestral condition in Mimosa was unequivocally assigned
cies) than the group that retained this characteristic (section as five petals, a character state retained from its mimosoid an-
Mimadenia, 15 species). Surveys of other groups are needed to cestors. This was also the unequivocal ancestral condition in
verify whether the presence of extrafloral nectaries and the oc- clade A, but not in clade B, where most reconstructions are
currence of ant mutualisms are related to higher rates of species equivocal, probably because of lack of resolution and due to the
diversification more widely in plants. frequent switches in petal number. The node subtending the
large group made up of clades D–X is assigned as tetramerous
Inflorescence type—Ancestral state reconstruction of this by 770 of the Bayesian trees, and this is the predominant state
character suggests high levels of homoplasy (Table 3), with fre- in Mimosa. Shifts from tetramerous to trimerous corollas were
quent shifts between capitula and spikes across the tree (Fig. observed in clade D (M. ceratonia, M. caesalpiniifolia, and M.
2c). Character optimization indicates an unequivocal change laticifera), in a subclade nested within clade H (series Campi-
from an ancestral spike (the most common state in the outgroup) colae and Filipedes), and in clade I (series Cordistipulae). A
to a capitulum at the base of the genus Mimosa. However, the reversal from four to five petals (supported by 943 trees) is hy-
predominant capitate inflorescences were reversed to spicate pothesized for clade F + G. A few species with six petals occur
several times (Fig. 2c) independently in different lineages: in clades A (M. colombiana) and G, but in both cases these spe-
clade A (M. myriadenia), clade B (several species), clade D (M. cies have a mixture of flowers with five and six petals. Overall,
caesalpiniifolia, M arenosa, M. ophthalmocentra and M. acus- the number of petals is variable within seven of the major clades
tistipula), clade F (M. invisa), clade I (M. xiquexiquensis), clade defined in this study (clades A–D, G, H, and N). Although nu-
N, and clade X. In clade B, spicate species are placed in a large merous switches in the number of petals are postulated to have
polytomy, shedding no light on the evolution of this character occurred in Mimosa, this character still can be a potentially use-
within this group. On the other hand, spikes probably evolved ful diagnostic character for clade I and a few subgroups within
only once in clade N (supported by 523 of the 1000 Bayesian major clades. It is important to note that some Mimosa species
trees), which suggests that the capitulum in M. lewisii repre- (19 sampled here) have occasional flowers with a different
sents a reversal. In clade X, M. dutrae evolved short spikes number of petals or sometimes similar proportions of two types
from a capitulum. It is expected that many more reversals may of flowers within an inflorescence. An extreme case of variation
have occurred in clade X, since many species of section Mi- in floral parts is found in series Fagaracanthae where flowers
mosa with spicate inflorescences were not sampled here. can consistently have from four to five petals (Barneby, 1991).
For the most part, Barneby (1991) viewed this as a conserva-
tive character only rarely putting species with different types of Pollen type—Polyads comprising eight pollen grains pre-
inflorescences together in the same group. For example, Barne- dominate across the outgroups, and this character state is un-
by’s placement of the sister species M. acutistipula and M. equivocally optimized as the ancestral state in Mimosa (Fig.
bimucronata (Fig. 1), which are virtually identical apart from 3b). An unequivocal change to polyads of 12 grains is hypoth-
their inflorescences in spikes and capitula (respectively), in dif- esized in a subclade in clade A, although M. revoluta, M.
ferent series, reflects this viewpoint. nothacacia, and M. townsendii retained the bi-tetrads. An ad-
ditional change to a 16-grain polyad is suggested for M. rufe-
Number of stamens—The ancestral diplostemonous condi- scens (the only one documented here). Pollen in bi-tetrads
tion predominates in Mimosa, with switches to haplostemonous characterizes clades B and D, which also unequivocally re-
flowers hypothesized to have occurred at least six times (Fig. tained the ancestral state. An unequivocal change from bi-tet-
2d, Table 3), with all these character state transitions unequivo- rads to tetrads, which is the predominant pollen type in Mimosa,
cal (i.e., present in all 1000 Bayesian trees). The most relevant is hypothesized to have occurred on the branch leading to the
transition in terms of classification is the one that segregates a large group comprising clades F–X. Only a single reversal to
monophyletic group containing the haplostemonous sections bi-tetrads is hypothesized to have occurred within this very
Mimosa and Calothamnos from the rest of the genus. Barneby large group (M. sinaloensis, clade G).
(1991) hypothesized that Bentham’s series Lepidotae evolved Previous attempts to correlate pollen morphology with the
haplostemony independently of the other haplostemonous mi- infrageneric classification in Mimosa have had little success
mosas, and on that basis established section Calothamnos, seg- (Caccavari, 1988), largely due to the nonmonophyly of many of
regating it from section Eumimosa. However, our results more the infrageneric groups. Here we show that pollen type is re-
closely support Bentham’s classification where all haploste- markably congruent with the plastid gene tree and that this
monous mimosas were placed in one large group. However, character presents relatively low levels of homoplasy (Table 3).
there are a few important exceptions where other haplostemon- Our results do not support the hypothesis of pollen evolution in
ous lineages are deeply nested within diplostemonous clades: which polyads with higher numbers of grains evolved from tet-
e.g., M. myriadenia in clade A, and series Plurijugae (clade F), rads (Caccavari, 1988, 1989), but agree with the evolutionary
which Bentham (1875) considered to belong within section Eu- progressions proposed by Barneby (1991), with large com-
mimosa. In contrast, Barneby (1991) correctly interpreted these pound polyads as plesiomorphic and tetrads as more derived
as independent reductions in stamen number and considered (Table 1). Other pollen traits not investigated here such as shape
these species unrelated to section Eumimosa. In addition, the and size of the polyads, presence of apertures, and structure of
placements of M. tejupilcana and M. leptocarpa (clade B) also the exine (Caccavari, 1988, 2002), may also be of taxonomic
Fig. 3. Morphological evolution in Mimosa. Character optimizations onto a 50% majority-rule Bayesian tree showing the evolution of (a) number of
petals, (b) pollen type, and (c) sensitive leaves. Missing data are indicated by slightly shorter terminal branches in (b) and (c).
significance and could be explored further. In a comprehensive estimates (Graybeal, 1998; Hillis, 1998), and for Mimosa, trnD-
palynological study of the northeastern Brazilian species of trnT provides enough informative data to construct a reason-
Mimosa, Lima et al. (2008) reported the occurrence of monads ably well-resolved and robustly supported gene tree. However,
and dyads in the genus. However, we view their findings with there can be many reasons why gene trees based on one or a few
caution. Visual inspection of their images suggests that monads loci fail to reflect the evolutionary relationships among species.
and dyads reported for M. ursina (figs. 42–46 in Lima et al., Issues related to horizontal gene transfer, hybridization, reticu-
2008) actually correspond to tetrahedric tetrads, in line with lation, incomplete lineage-sorting, paralogous genes, and
previous data for this species (Grether, 1997). Therefore, spe- pseudogenes, can all contribute to gene tree–species tree incon-
cies reported as containing monads or dyads by Lima et al. gruence (Doyle, 1997; Maddison, 1997). Therefore, sequencing
(2008) were instead coded here as having tetrads. of biparentally inherited nuclear genes will be needed to con-
firm the relationships presented here, especially given the exis-
Sensitive leaves—Seismonasty is hypothesized to have arisen tence of a significant number of known polyploids and putative
independently in eight lineages of Mimosa (Fig. 3c), including hybrids in the genus (Seijo and Fernandez, 2001; Morales et al.,
clades G, M, P, Q, R, T, and U. However, it is important to bear 2010; Dahmer et al., 2011).
in mind that this trait is present, albeit to a lesser degree, in
many other species in the genus. The biological significance of Biogeography and dating analysis— A general result found
seismonasty is not well known, but probably evolved as a de- here is the strong geographical structuring of the phylogeny, in
fense mechanism against herbivores or to avoid leaf damage in which species in any particular clade tend to occur in the same
regions with very wet climates, although this remains to be con- biogeographical region. Examples are clade B (USA, Mexico,
firmed experimentally. Detailed studies found different types of and Mesoamerica), clades C and W (Andes), clade E (Old
nyctinastic (night time) leaf movements within the mimosoid World), clade I (northeast Brazil), clade K (southern South
genera Desmanthus (Luckow, 1993) and Leucaena (Hughes, America), clades O and R (central Brazil), clade V (Mexico)
1998), in which the pinnae and petiole can move either down- and clade X (subtropical South America) (Fig. 1). In many in-
ward or upward, providing further evidence that patterns of stances, species previously regarded as taxonomically distantly
evolution of leaf movements are both complex and evolution- related were placed in the same biogeographically coherent
arily labile in mimosoid legumes. The preliminary analysis clade.
shown here suggests that rapid leaf movements evolved multi- Geography has been found to be an important predictor of
ple times in distinct lineages within Mimosa, suggesting that phylogenetic structure, particularly across the diverse season-
the physiological and morphological apparatus required for ally dry tropical forest plant groups that make up the so-called
seismonasty has been recruited recurrently during the evolution succulent biome (sensu Schrire et al., 2005). This structure has
of the genus. However, more detailed studies might identify been attributed to phylogenetic niche conservatism (Donoghue,
some coherent phylogenetic patterns and also ecological asso- 2008), dispersal limitation, and in situ speciation within indi-
ciations related to leaf movement. vidual geographical areas (Schrire et al., 2009; Pennington et
al., 2006, 2009). For example, previous studies have shown that
Toward a new infrageneric classification of Mimosa— Clas- phylogenies of typical phylogenetically niche conserved tropi-
sifications that reflect evolutionary relationships based on cal dry forest plant groups such as Coursetia, Poissonia (Legu-
monophyly combined with diagnosability are a widely accepted minosae), and Ruprechtia (Polygonaceae) have high levels of
goal of modern systematics (Carine and Scotland, 2002). Many phylogenetic geographic structure (Lavin, 2006; Pennington et
of the groups of Barneby’s (1991) infrageneric classification al., 2006; Queiroz and Lavin, 2011). A similar pattern of strong
are shown here to be nonmonophyletic (Table 2). However, geographic structure in Mimosa has been found in the predomi-
several of the 24 strongly supported clades highlighted here nantly seasonally tropical dry forest clades, including southern
lack any obvious diagnostic morphological apomorphies. This Mexico dry forests (clade V), Andean dry valleys (clades C and
is evident from the high degree of homoplasy found in some of W), and Caatinga in northeast Brazil (clade I) and Madagascar
the morphological characters investigated, such as number of (clade E), suggesting that endemic lineages confined to each of
petals and type of inflorescence (Figs. 2, 3; Table 3). However, these dry forest areas have persisted in relative isolation for
it is important to mention that a few clades received strong sup- several million years and that dispersal between highly disjunct
port from conspicuous morphological characters, such as sec- fragments of tropical dry forest has been limited (Pennington
tion Mimadenia (clade A; presence of petiolar nectaries) and et al., 2006).
series Cordistipulae (clade I; trimerous flowers). Unlike these predominantly dry forest groups, Mimosa is
A more detailed survey of other morphological traits not in- also diverse in other neotropical biomes, thereby offering the
vestigated here (such as armature, indumentum, calyx morphol- opportunity for more detailed investigations of the role of ecol-
ogy, and fruit type) may yet reveal reliable characters that define ogy/geography on the structuring of phylogenies and to provide
some of these groups and provide the diagnosability sought in insights into the timing of origin and evolution of different
modern classifications. However, it is not guaranteed that all biomes. For example, Mimosa lineages have been used as a
clades could be appropriately delimited by a robust set of mor- source of phylogenetic evidence to investigate the evolutionary
phological synapomorphies, given that many of these other history of several prominent biomes in the noetropics, such as
traits also seem to have evolved independently many times in the Cerrado (several clades; Simon et al., 2009), the Amazon
Mimosa. (clade A; Hoorn et al., 2010), and the Andes (clades A, C, and W;
The emphasis in this study is on taxon rather than character Pennington et al., 2010).
sampling to ensure adequate taxonomic and geographic cover- The time-calibrated phylogeny of Mimosa suggests a gradual
age for such a species-rich group. The trade-off for this is reli- accumulation of major lineages from 15 to 3 Ma (Fig. 4; online
ance on a single plastid locus. It has been shown that adding Appendix S5). Although not used in the dating analysis as con-
more taxa to phylogenies can improve accuracy of phylogenetic straints, putative Mimosa fossils described from the Oligocene
Table 3. Levels of homoplasy of six morphological characters mapped following an initial diversification in Africa or that possible
onto a molecular phylogeny of Mimosa, as measured by number of niches available for Mimosa on that continent were already oc-
estimated steps and consistency (CI) and retention (RI) indices. See cupied by other taxa, pre-empting a major diversification of Mi-
also Figs. 2 and 3. mosa in Africa.
Character Steps CI RI In contrast to the paucity of species in east Africa, Mimosa is
impressively diverse in Madagascar, with more than 30 species,
Petiolar nectary 1 1.00 1.00 the vast majority of them endemic (Villiers, 2002). Among the
Inflorescence type 24 0.04 0.55 native species, only M. latispinosa extends to Africa and other
Number of stamens 6 0.33 0.95
Number of petals 19 0.11 0.80
islands in the Indian Ocean, but some of these could be recent
Pollen type 7 0.42 0.94 introductions (Villiers, 2002). Because of lack of resolution
Sensitive leaves 8 0.12 0.50 among the African, Asian, and Malagasy clades, the present
Mimosa phylogeny sheds no light on possible biogeographical
or evolutionary scenarios for the diversification of the Old
World species and whether the Malagasy mimosas came from
of Mexico (Magallón-Puebla and Cevallos-Ferriz, 1994; Africa, which is the most frequent sister clade relationship for
Calvillo-Canadell and Cevallos-Ferriz, 2005) are compatible Malagasy groups (Yoder and Nowak, 2006), or from some-
with the time frame estimated for the evolution of genus where else. If this is the case in Mimosa, the imbalance in sister
(Mimosa stem node 28 Ma; Simon et al., 2009). The rate of clades between continental Africa and Madagascar in terms of
nucleotide substitution varied across the tree and deviated from species richness observed here, would be in line with what has
a constant rate (BEAST parameter ucld.stdev = 0.66 [0.54– been found for some other groups of plants (e.g., Janssen et al.,
0.79], mean and 95% credibility interval), indicating signifi- 2008).
cant branch rate heterogeneity among lineages. Particularly Legume groups across all three subfamilies, including the
high substitution rates were observed in M. strigillosa (clade J) Chapmannia–Diphysa–Ormocarpum group, the subtribe
(Fig. 4). Phaseolinae, the Dichrostachys/Calliandropsis clade, and the
Delonix–Lemuropisum/Conzattia–Heteroflorum, Parkinsonia,
Old world clade— Divergence time estimates for the lineage and Haematoxylon clades of the Peltophorum group, show sim-
that gave rise to the Old World species indicate that Mimosa ilar disjunct distributions and sister group relationships between
was established in that region during the late Miocene. The fact Madagascar/east Africa and the neotropics (Lavin and Luckow,
that the Old World clade is nested within a predominantly South 1993; Lavin et al., 2000, 2004; Thulin et al., 2004; Haston et al.,
American clade (Fig. 1) suggests that the ancestral area of the 2005; Schrire et al., 2005). Some of these distributions have
Old World mimosas could have been in South America. The been discussed in relation to the boreotropics hypothesis, which
ages estimated for the Old World clade (crown node 6.3 Ma, predicts that some plant lineages used to be more widespread in
stem node 10.0 Ma, Fig. 4) are much too young to support any the northern hemisphere spanning the north Atlantic and occu-
hypothesis involving continental vicariance between Africa and pying land connections during the Eocene or early Oligocene,
South America, since the initial split between these continents and consequently linking the floras of different continents
occurred ca. 100 Ma, with putative later connections, including (Wolfe, 1975; Lavin and Luckow, 1993; Schrire et al., 2003).
the Early Tertiary North Atlantic land bridge and the Walvis According to this hypothesis, the present peculiar disjunct dis-
Ridge, a possible Oligocene filter barrier linking Africa and tributions of these taxa could be explained by the extinction of
South America (Tiffney, 1985; Morley, 2003), also signifi- members of these lineages in most of the former range, which
cantly predating the late Miocene divergence time estimates for is supported by the occurrence of tropical fossil taxa in the Ter-
Old World Mimosa. This suggests that long-distance dispersal tiary of North America and Europe. However, the younger ages
provides the most plausible explanation for this remarkable dis- estimated for the disjunction between Old World and neotropi-
junction, in line with predominantly Miocene divergence time cal lineages of Mimosa and the possible sister group relation-
estimates for the majority of intracontinental and transcontinen- ship between South American elements and the Old World
tal crown groups in legumes (Lavin et al., 2004). Among the clade would not seem to support a boreotropical explanation in
South American members of clade D, there is an intriguing this case. Instead, long-distance oceanic dispersal, which has
suite of coastal species (e.g., M. bimucronata, which is abun- been proposed to explain several intercontinental disjunctions
dant in coastal Brazil), which reinforces the hypothesis of a in many legume and other plant groups (e.g., Lavin et al., 2004;
trans-Atlantic dispersal as the likely route for migration to the Renner, 2004), is more likely.
Old World. However, lack of resolution at the base of clades D
and E precludes more precise inference about the exact ances- Conclusions— This study presents a major step toward the
tral area of the Old World clade within the neotropics or about construction of a molecular phylogeny for the large and com-
where Mimosa first arrived in the Old World. plex legume genus Mimosa. One of the strengths of the present
One unresolved question is why Mimosa is so poorly repre- analysis is the extensive taxon sampling, which establishes an
sented in continental Africa compared to the extraordinary spe- overall phylogenetic framework for Mimosa in terms of mor-
cies diversity found in South America, considering that both phological evolution and relationships. Many of the informal
continents share similar climates and vegetation types. One clades identified here prompt taxonomic recognition, which
possible explanation is that there has not been enough time could eventually culminate in an updated infrageneric classifi-
since the arrival of Mimosa in the Old World for significant di- cation of the genus. However, it may not be entirely straightfor-
versification to have occurred in Africa. However, this seems ward to directly translate a phylogeny into a classification.
unlikely, as many younger Mimosa clades have diversified very Potential obstacles for a new classification for Mimosa are the
extensively in the New World within a short time span. Another lack of diagnostic morphological apomorphies for some well-
possibility could be widespread extinction of Mimosa lineages supported groups, the difficulties of implementing a strictly
Fig. 4. Chronogram (left) derived from the trnD-trnT Bayesian dating analysis using an uncorrelated relaxed molecular clock (maximum credibility
tree with averaged node ages across a set of 1000 Bayesian trees sampled at stationarity), and the corresponding phylogram (right). Letters are major clades
discussed in the text, and squares are calibration points used in the analysis (Mimosa crown node and section Mimadenia crown node). Scale in million
years (chronogram) and substitutions per site (phylogram).
rank-based classification following the Linnean system, the in- extensive morphological homoplasy, suggesting that particular
complete taxon sampling in the present analysis, which makes traits evolved independently several times in different groups.
placement of unsampled species tentative, and the limited data Another possibility, given that these results are based on a
set from a uniparentally inherited genome. maternally inherited plastid gene tree, is that hybridization (in-
The affinities of the Old World species, which were not for- cluding allopolyploidy) could have facilitated chlorolplast in-
mally assessed in the most recent revision of Mimosa (Barneby, trogression, thereby generating an exaggerated impression of
1991), are explicitly analyzed for the first time, suggesting a the extent of trans-specific geographic structure and potentially
late Miocene transoceanic dispersal from South America. The causing the erroneous inference of widespread homoplasy.
importance of geography and ecology, rather than morphology, Nevertheless, there are no indications of this type of conflict in
in predicting many of the major groups within Mimosa is one of our data set, since the known polyploids sampled here (27 spe-
the most striking findings. In many cases, morphologically het- cies) are placed within morphologically homogeneous clades.
erogeneous species grouped in robustly supported clades that In any case, the results presented here provide the essential
can only be understood when geography is taken into account. foundations for disentangling potentially reticulate from diver-
The presence of some morphologically disparate species in gent relationships, as well as a good interim working frame-
well-supported clades implies strong geographic structure and work for comparative studies across the genus as a whole.
Finally, this paper illustrates how evolutionary patterns can Dahmer, N., M. F. Simon, M. T. Schifino-Wittmann, C. E. Hughes, S. T. S.
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Appendix 1. Voucher information and GenBank accession numbers for taxa used in this study. Voucher specimens are deposited in the following herbaria: CEN =
Embrapa Recursos Genéticos e Biotecnologia; E = Royal Botanic Garden, Edinburgh; FHO = University of Oxford; HUEFS = Universidade Estadual de Feira
de Santana; ICN = Universidade Federal do Rio Grande do Sul; K = Royal Botanic Gardens, Kew; MEXU = Universidad Nacional Autónoma de México; MO
= Missouri Botanical Garden; NY = New York Botanical Garden; RB = Jardim Botânico do Rio de Janeiro; UAMIZ = Universidad Autónoma Metropolitana,
Iztapalapa; UB = Universidade de Brasília; VIC = Universidade Federal de Viçosa.
Section (given only for Mimosa species; Old World taxa were considered as belonging to section Batocaulon)—Taxon, Voucher specimen, Collection country,
herbarium, GenBank accessions.
OUTGROUPS
Anadenanthera colubrina (Vell.) Brenan, Hughes 2308, Bolivia, FHO, MEXU, FJ982052; M. domingensis Benth., Barneby 18276, Dominican
FJ981975; Microlobius foetidus (Jacq.) M.Sousa & G.Andrade, Hughes Republic, NY, FJ982055; M. dysocarpa Benth., Newman 296, USA, K,
2150, Mexico, FHO, FJ981976; Parapiptadenia excelsa (Griseb.) FJ982059; M. echinocaula Benth., Simon 679, Brazil, UB, FJ982060; M.
Burkart, Hughes 2425, Bolivia, FHO, FJ982235; Piptadenia adiantoides emoryana Benth., Grether 2842, Mexico, UAMIZ, FJ982061; M.
(Spreng.) Macbride, Simon 726, Brazil, FHO, FJ982236; P. buchtienii ervendbergii A.Gray, Martínez 35132, Mexico, MEXU, FJ982062; M.
Barneby, Hughes 2427, Bolivia, FHO, FJ982237; P. gonoacantha (Mart.) farinosa Griseb., Wood 21535, Bolivia, K, HM353086; M. filipes Mart.,
Macbride, Simon 735, Brazil, FHO, FJ982238; P. stipulacea (Benth.) Queiroz 10058, Brazil, HUEFS, FJ982065; M. galeottii Benth., Simon
Ducke, Simon 702, Brazil, FHO, FJ982239; P. trisperma (Vell.) Benth., 840, Mexico, MEXU, FJ982069; M. gatesiae Barneby, Simon 741, Brazil,
Armstrong 512, Brazil, FHO, FJ982240; P. viridiflora Benth., Hughes FHO, FJ982070; M. gemmulata Barneby var. gemmulata, Simon 690,
1681, Mexico, FHO, FJ982241; Pityrocarpa moniliformis Benth., Way Brazil, FHO, FJ982071; M. gracilis Benth. var. invisiformis Barneby,
SWM2449, Brazil, K, FJ982242; P. obliqua Macbride, Macqueen 439, Simon 762, Brazil, FHO, FJ982073; M. gracilis Benth. var. stipitata
Mexico, K, FJ982243; Stryphnodendron adstringens (Mart.) Coville, Barneby, Simon 745, Brazil, FHO, FJ982074; M. grandidieri Baill., Du
Simon 734, Brazil, FHO, FJ982244; S. obovatum Benth., Hughes 2397, Puy M56, Madagascar, K, FJ982075; M. guaranitica Chodat & Hassl.,
Bolivia, FHO, FJ982245. Nascimento 474, Brazil, HUEFS, FJ982076; M. guatemalensis (Hook. &
Batocaulon—Mimosa acantholoba (Willd.) Poir. var. acantholoba, Eastwood Arn.) Benth., Simon 831, Mexico, MEXU, FJ982077; M. hafomantsina
118, Peru, FHO, FJ981977; M. acantholoba Poir. var. eurycarpa Villiers, Lewis 2138, Madagascar, K, FJ982080; M. hamata Willd., Simon
(B.L.Rob.) Barneby, Montaño-Arias 28, Mexico, UAMIZ, FJ981978; M. 876, India, FHO, FJ982081; M. hexandra M.Micheli, Simon 711, Brazil,
aculeaticarpa Ortega, Simon 808, Mexico, MEXU, FJ981980; M. FHO, FJ982084; M. hexandra M.Micheli, Fabian-Martinez 128, Mexico,
acutistipula Benth. var. acutistipula, Simon 705, Brazil, FHO, FJ981981; MEXU, FJ982083; M. hondurana Britton, Simon 858, Mexico, MEXU,
M. adenantheroides (Martens & Galleotti) Benth., Martínez-Bernal 945, FJ982087; M. interrupta Benth., Queiroz 10485, Brazil, HUEFS,
Mexico, UAMIZ, FJ981982; M. adenantheroides (Martens & Galleotti) FJ982093; M. interrupta Benth., Queiroz 10584, Brazil, HUEFS,
Benth. var. hystricosa (Brandegee) R.Grether, ined., Tenorio 21201, JF694262; M. invisa Mart. ex Colla var. invisa, Simon 715, Brazil, FHO,
Mexico, MEXU, FJ981983; M. adenocarpa Benth., Simon 728, Brazil, FJ982094; M. jaenensis T.E.Särkinen, J.L.Marcelo-Peña & C.E.Hughes,
FHO, FJ981984; M. adenophylla Taub. ex Glaz., Simon 458, Brazil, UB, Särkinen 3070, Peru, FHO, HM353091; M. lacerata Rose, Hughes 2057,
FJ981985; M. adenophylla Taub. ex Glaz. var. mitis Barneby, Lima 184, Mexico, FHO, FJ982099; M. laticifera Rizzini & Mattos, Simon 599,
Brazil, HUEFS, FJ981986; M. aff. dalyi Barneby, Wood 24306, Bolivia, Brazil, UB, FJ982104; M. latispinosa Lam., Sutherland 206, Madagascar,
K, JF694256; M. aff. weberbaueri Harms, Pennington 17903, Peru, K, K, FJ982105; M. lepidota Herzog, Hughes 2469, Bolivia, FHO, FJ982107;
FJ981990; M. andina Benth., Lewis 2271, Ecuador, K, HM353071; M. M. leptantha Benth., Nascimento 471, Brazil, HUEFS, FJ982108; M.
antioquensis Killip ex Rudd var. isthmensis R.Grether, Simon 860, leptocarpa Rose, Rico 1014, Mexico, K, FJ982109; M. leucaenoides
Mexico, MEXU, FJ981995; M. apodocarpa Benth., Simon 635, Brazil, Benth., Montaño-Arias 8, Mexico, UAMIZ, FJ982110; M. levenensis
UB, FJ981997; M. arenosa (Willd.) Poir. var. leiocarpa (DC.) Barneby, Drake, Luckow 4453, Madagascar, FHO, FJ982111; M. lewisii Barneby,
Martínez-Bernal 923, Mexico, UAMIZ, FJ981998; M. artemisiana Simon 696, Brazil, FHO, FJ982112; M. luisana Brandegee, Simon 844,
Heringer & Paula, Faria 138, Brazil, RB, FJ981999; M. aspera M.E.Jones, Mexico, FHO, FJ982114; M. malacophylla A.Gray, Camargo-Ricalde
Simon 817, Mexico, MEXU, FJ982000; M. auriculata Benth., Hughes 530, Mexico, UAMIZ, FJ982115; M. martindelcampoi Medrano,
2405, Bolivia, FHO, FJ982002; M. bahamensis Benth., Way 132, Mexico, Camargo-Ricalde 527, Mexico, UAMIZ, FJ982117; M. menabeensis R.
K, FJ982003; M. benthamii Macbride, Simon 848, Mexico, MEXU, Vig. var. menabeensis, Sutherland 209, Madagascar, K, FJ982119; M.
FJ982004; M. bifurca Benth., Dahmer 4, Brazil, ICN, FJ982005; M. minarum Barneby, Nascimento 495, Brazil, HUEFS, FJ982120; M.
bimucronata Kuntze, Simon 301, Brazil, UB, FJ982006; M. biuncifera minutifolia B.L.Rob. & Greenm., Simon 810, Mexico, MEXU, FJ982121;
Benth., Simon 805, Mexico, MEXU, FJ982007; M. blanchetii Benth., M. misera Benth., Simon 703, Brazil, FHO, JF694264; M. mollis Benth.,
Simon 688, Brazil, FHO, FJ982008; M. borealis A.Gray, Simon 873, Simon 850, Mexico, MEXU, FJ982123; M. monancistra Benth., Simon
USA, FHO, FJ982010; M. busseana Harms, Clarke 26, Tanzania, K, 809, Mexico, MEXU, FJ982124; M. montana Kunth. var. montana,
FJ982013; M. caduca (Willd.) Poir., Lewis 3006, Ecuador, K, HM353080; Hughes 2225, Peru, FHO, FJ982125; M. montana Kunth. var. sandemanii
M. caesalpiniifolia Benth., Simon 756, Brazil, FHO, FJ982014; M. Barneby, Eastwood 125, Peru, FHO, HM353099; M. morroensis Barneby,
calcicola B.L.Rob., Simon 846, Mexico, MEXU, FJ982015; M. campicola Nascimento 244, Brazil, HUEFS, JF694266; M. mossambicensis Brenan,
Harms var. planipes Barneby, Simon 692, Brazil, FHO, FJ982018; M. Brummitt 8896, Malawi, K, FJ982126; M. myriocephala Baker, Rakoto
candollei R.Grether, Hughes 2394, Bolivia, FHO, FJ982020; M. ceratonia 329, Madagascar, K, FJ982128; M. nanchititlana R.Grether & Barneby,
L. var. ceratonia, Grimes 3223, Puerto Rico, NY, JF694259; M. ceratonia Grether 2938, Mexico, UAMIZ, FJ982129; M. nossibiensis Benth. var.
L. var. interior Barneby, Simon 727, Brazil, FHO, FJ982022; M. nossibiensis, Du Puy M350, Madagascar, K, FJ982131; M. nuttallii
cordistipula Benth., Simon 693, Brazil, FHO, FJ982029; M. coruscocaesia (DC.) B.L.Turner, Simon 875, USA, FHO, FJ982134; M. onilahensis
Barneby, Martins 469, Brazil, UB, FJ982030; M. corynadenia Britton & R.Vig., Du Puy M899, Madagascar, K, FJ982138; M. ophthalmocentra
Rose, Sousa 12896, Mexico, MEXU, FJ982031; M. costenya McVaugh, Mart. ex Benth., Way SWM2434, Brazil, K, FJ982139; M. orthocarpa
Simon 833, Mexico, MEXU, FJ982032; M. craspedisetosa R.H.Fortunato Spruce ex Benth., Simon 855, Mexico, MEXU, FJ982141; M. palmeri
& R.Palese, Wood 24488, Bolivia, K, JF694260; M. cruenta Benth., Rose, Simon 823, Mexico, MEXU, FJ982142; M. paucifolia Benth.,
Queiroz 12575, Brazil, HUEFS, FJ982033; M. crumenarioides Dutra 450, Brazil, VIC, JF694268; M. platycarpa Benth. var. platycarpa,
L.P.Queiroz & G.P.Lewis, Simon 722, Brazil, FHO, FJ982034; M. dalyi Simon 859, Mexico, MEXU, FJ982152; M. polyantha Benth., Simon 829,
Barneby, Wood 16487, Bolivia, K, FJ982039; M. delicatula Baill., Mexico, MEXU, FJ982153; M. polydidyma Barneby, Simon 719, Brazil,
Sutherland 262, Madagascar, K, FJ982043; M. depauperata Benth., FHO, FJ982157; M. prainiana Gamble, Maesen 3834, India, K, FJ982158;
Simon 801, Mexico, MEXU, FJ982045; M. detinens Benth., Sanchez 46, M. pseudosepiaria Harms, Simon 712, Brazil, FHO, FJ982161; M.
Bolivia, MO, FJ982046; M. diplotricha C.Wright ex Sauvalle var. psoralea Benth., Phillipson 3571, Madagascar, K, FJ982164; M.
diplotricha, Simon 877, Taiwan, FHO, FJ982050; M. diplotricha C. pteridifolia Benth., Simon 754, Brazil, FHO, FJ982165; M. purpusii
Wright ex Sauvalle var. diplotricha, Simon 600, Brazil, UB, FJ982049; Brandegee, Simon 841, Mexico, MEXU, FJ982167; M. quadrivalvis L.
M. distachya Cav. var. oligacantha (DC.) Barneby, Ku 365, Mexico, var. quadrivalvis, Camargo-Ricalde 532, Mexico, UAMIZ, FJ982169;
M. quitensis Benth., Lewis 2856, Ecuador, K, HM353116; M. rhodocarpa Taub., Simon 676, Brazil, UB, FJ982188; M. somnians Humb. & Bonpl.
(Britton & Rose) R.Grether, Hughes 2161, Mexico, FHO, FJ982175; M. ex Willd. var. lasiocarpa (Benth.) Barneby, Simon 736, Brazil, FHO,
robusta R.Grether, Simon 818, Mexico, MEXU, FJ982176; M. rubicaulis FJ982194; M. speciosissima Taub., Simon 753, Brazil, FHO, FJ982197;
Lam. subsp. himalayana (Gamble) H.Ohashi, Thomas 24/1, Nepal, K, M. splendida Barneby, Simon 739, Brazil, FHO, FJ982199; M. strigillosa
FJ982177; M. rupertiana B.L.Turner, Bye 12884, Mexico, MEXU, Torr. & A.Gray, Lievens 2666, USA, MEXU, FJ982201; M. stylosa
FJ982179; M. schomburgkii Benth., Hellin 15, Honduras, FHO, Barneby, Dutra 318, Brazil, VIC, FJ982203; M. ulei Taub. var. grallator
FJ982183; M. sericantha Benth., Simon 410, Brazil, UB, FJ982185; M. Barneby, Simon 777, Brazil, FHO, FJ982213; M. viperina M.F.Simon &
setuligera Harms, Simon 709, Brazil, FHO, FJ982189; M. similis Britton C.E.Hughes, Simon 461, Brazil, FHO, FJ982224; M. weddelliana Benth.,
& Rose, Simon 807, Mexico, MEXU, FJ982191; M. sinaloensis Britton Ritter 4604, Bolivia, MO, FJ982230.
& Rose, Simon 828, Mexico, MEXU, FJ982192; M. spirocarpa Rose,
Mimadenia—Mimosa colombiana Britton & Killip, Torres 21343, Colombia,
Simon 825, Mexico, MEXU, FJ982198; M. strobiliflora Burkart, Ribas
K, FJ982027; M. guilandinae (DC.) Barneby var. guilandinae, Prévost
3600, Brazil, HUEFS, FJ982202; M. tejupilcana R.Grether & A.Martínez-
3958, French Guiana, K, FJ982078; M. irrigua Barneby, Simon 694,
Bernal, Montaño-Arias 16, Mexico, UAMIZ, FJ982204; M. tenuiflora
Brazil, FHO, FJ982095; M. lepidophora Rizzini, Cardoso 1747, Brazil,
(Willd.) Poir., Simon 698, Brazil, FHO, FJ982205; M. texana Small var.
FHO, FJ982106; M. myriadenia (Benth.) Benth. var. punctulata (Benth.)
filipes (Britton & Rose) Barneby, Simon 845, Mexico, MEXU, FJ982207;
Barneby, Acevedó-Rdgz 7483, Ecuador, K, FJ982127; M. nothacacia
M. texana Small var. texana, Simon 803, Mexico, MEXU, FJ982208; M.
Barneby, Lewis 2353, Ecuador, K, FJ982132; M. pithecolobioides Benth.,
torresiae R.Grether, Torres-Colin 10040, Mexico, MEXU, FJ982209; M.
Dutra 317, Brazil, VIC, FJ982151; M. revoluta Benth., Hughes 2278,
ulbrichiana Harms, Simon 710, Brazil, FHO, FJ982212; M. uliginosa
Bolivia, FHO, FJ982174; M. rufescens Benth. var. rufescens, Ferreira
Chod. & Hassl., Queiroz 12608, Brazil, HUEFS, FJ982214; M. unipinnata
596, Brazil, K, FJ982178; M. townsendii Barneby, Lewis 3025, Ecuador,
B.D.Parfitt & Pinkava, Carranza 2355, Mexico, MEXU, FJ982215; M.
K, FJ982210; M. watsonii B.L.Rob., Simon 857, Mexico, MEXU,
uraguensis Hook. & Arn., Simon 862, cultivated, FHO, FJ982216; M.
FJ982228.
verrucosa Benth., Simon 706, Brazil, FHO, FJ982221; M. vilersii Drake,
Labat 3020, Madagascar, K, FJ982223; M. volubilis Villiers, Du Puy Mimosa—Mimosa acapulcensis B.L.Rob., Otero R2, Mexico, MEXU,
M739, Madagascar, K, FJ982226; M. waterlotii R.Vig., Schrire 2551, FJ981979; M. adamantina Barneby var. adamantina, Dutra 459,
Madagascar, K, FJ982227; M. weberbaueri Harms, Hughes 2043, Peru, Brazil, VIC, JF694255; M. aff. flagellaris Benth., Queiroz 12322, Brazil,
FHO, FJ982229; M. xavantinae Barneby, Farias 346, Brazil, UB, HUEFS, FJ981989; M. aff. incarum Barneby, Pennington 1715, Peru,
FJ982233; M. xiquexiquensis Barneby, Harley 54331, Brazil, HUEFS, E, HM353081; M. aff. polycarpa Kunth var. polycarpa, Eastwood
JF694271; M. zygophylla Benth., Camargo-Ricalde 525, Mexico, 89, Peru, FHO, HM353082; M. aff. xanthocentra Mart., Queiroz
UAMIZ, FJ982234. 10476, Brazil, HUEFS, FJ981991; M. affinis B.L.Rob., Simon 814,
Mexico, MEXU, FJ981992; M. albida Humb. & Bonpl. ex Willd. var.
Calothamnos—Mimosa aff. bathyrrhena Barneby, Simon 874, Brazil, FHO,
albida, Hughes 2083, Mexico, FHO, FJ981993; M. atlantica Barneby,
FJ981988; M. aurivillus Mart. var. aurivillus, Dutra 348, Brazil, VIC,
Ribas 4333, Brazil, HUEFS, FJ982001; M. boliviana Benth., Hughes
JF694257; M. aurivillus Mart. var. calothamnos (Benth.) Barneby,
2426, Bolivia, FHO, FJ982009; M. brevipetiolata Burkart var. hirtula
Dutra 347, Brazil, VIC, JF694258; M. daleoides Benth., Schinini 35683,
(Burkart) Barneby, Queiroz 12614, Brazil, HUEFS, FJ982012; M.
Argentina, MEXU, FJ982038; M. flocculosa Burkart, CNPF sn, Brazil,–,
callidryas Barneby, Cruz 94, Brazil, HUEFS, FJ982016; M. callithrix
FJ982067; M. incana Benth., Dahmer 2, Brazil, ICN, FJ982091; M.
Malme, Simon 684, Brazil, UB, FJ982017; M. casta L., Johnson 2189-80,
leprosa (Benth.) Macbride var. parviceps Barneby, Dutra 358, Brazil,
Panama, MEXU, FJ982021; M. chartostegia Barneby, Ribas 5085, Brazil,
VIC, JF694263; M. pilulifera Benth., Dahmer 3, Brazil, ICN, FJ982149;
HUEFS, FJ982023; M. coniflora Burkart, Ribas 3060, Brazil, HUEFS,
M. pilulifera Benth. var. pseudincana (Burkart) Barneby, Simon 878,
FJ982028; M. ctenodes Barneby, Hughes 2212, Peru, FHO, FJ982036;
Brazil, FHO, FJ982150; M. scabrella Benth., Lima 4055, Brazil, RB,
M. cyclophylla Taub., Simon 757, Brazil, FHO, FJ982037; M. deamii
FJ982181.
B.L.Rob., Martínez-Bernal 919, Mexico, UAMIZ, FJ982040; M. debilis
Habbasia—Mimosa adenotricha Benth., Dutra 332, Brazil, VIC, FJ981987; Humb. & Bonpl. ex Willd. var. debilis, Hughes 2393, Bolivia, FHO,
M. albolanata Taub. var. paucipinna (Benth.) Barneby, Simon 667, Brazil, FJ982041; M. dicerastes Barneby, Simon 448, Brazil, UB, FJ982047; M.
UB, FJ981994; M. antrorsa Benth., Fagg 1747, Brazil, UB, FJ981996; discobola Barneby, Simon 744, Brazil, FHO, FJ982051; M. dolens Vell.
M. brachycarpa Benth., Queiroz 10589, Brazil, HUEFS, FJ982011; M. var. dolens, Dutra 352, Brazi, VIC, JF694261; M. dolens Vell. var. rigida
camporum Benth., Faria 729, Brazil, RB, FJ982019; M. cisparanensis (Benth.) Barneby, Simon 879, Brazil, FHO, FJ982053; M. dryandroides
Barneby, Simon 568, Brazil, UB, FJ982024; M. claussenii Benth. var. Taub. ex Glaz., Ribas 3449, Brazil, HUEFS, FJ982057; M. dutrae Malme,
claviceps Barneby, Simon 766, Brazil, FHO, FJ982025; M. claussenii Dahmer 5, Brazil, ICN, FJ982058; M. fachinalensis Burkart, Dahmer
Benth. var. megistophylla Barneby, Simon 768, Brazil, FHO, FJ982026; 16, Brazil, ICN, FJ982063; M. fachinalensis Burkart, Dahmer 20, Brazil,
M. cryptothamnos Barneby, Simon 738, Brazil, FHO, FJ982035; M. ICN, FJ982064; M. flagellaris Benth., Queiroz 12545, Brazil, HUEFS,
decorticans Barneby, Simon 681, Brazil, UB, FJ982042; M. densa Benth. FJ982066; M. goldmanii B.L.Rob., Martínez-Bernal 921, Mexico,
var. densa, Simon 870, Brazil, FHO, FJ982044; M. diminuta M.F.Simon UAMIZ, FJ982072; M. gymnas Barneby, Silva 3541, Brazil, HUEFS,
& C.E.Hughes, Simon 866A, Brazil, FHO, FJ982048; M. dominarum FJ982079; M. hirsutissima Mart. var. barbigera (Benth.) Barneby, Simon
Barneby, Simon 776, Brazil, FHO, FJ982054; M. dormiens Humb. 765, Brazil, FHO, FJ982085; M. hirsutissima Mart. var. grossa Barneby,
& Bonpl. ex Willd., Guadarrama 6841, Mexico, MEXU, FJ982056; Queiroz 12854, Brazil, HUEFS, FJ982086; M. honesta Mart., Simon 720,
M. foliolosa Benth. var. pubescens Benth., Simon 733, Brazil, FHO, Brazil, FHO, FJ982088; M. hypoglauca Mart. var. hypoglauca, Simon
FJ982068; M. heringeri Barneby, Proença 2138, Brazil, UB, FJ982082; M. 723, Brazil, FHO, FJ982090; M. incarum Barneby, Hughes 2206, Peru,
humivagans Barneby, Simon 737, Brazil, FHO, FJ982089; M. josephina FHO, FJ982092; M. jacobita Barneby, Hughes 2400, Bolivia, FHO,
Barneby, Hughes 2398, Bolivia, FHO, FJ982097; M. kalunga M.F.Simon FJ982096; M. lactiflua Delile ex Benth., Hughes 2079, Mexico, FHO,
& C.E.Hughes, Simon 866, Brazil, FHO, FJ982098; M. laniceps Barneby, FJ982100; M. lamolina C.E.Hughes & G.P.Lewis, Hughes 2648, Peru,
Simon 773, Brazil, FHO, FJ982102; M. manidea Barneby, Simon 760, FHO, FJ982101; M. lanuginosa Glaz. ex Burkart var. lanuginosa,
Brazil, FHO, FJ982116; M. melanocarpa Benth., Simon 675, Brazil, UB, Simon 732, Brazil, FHO, FJ982103; M. loxensis Barneby, Lewis 2987,
FJ982118; M. neptuniodes Harms, Wood 22123, Bolivia, K, FJ982130; Ecuador, K, FJ982113; M. modesta Mart. var. modesta, Simon 708,
M. occidentalis Britton & Rose, Simon 821, Mexico, MEXU, FJ982136; Brazil, FHO, FJ982122; M. montis-carasae Barneby, Dutra 623, Brazil,
M. oligosperma Barneby, Simon 865, Brazil, FHO, FJ982137; M. pigra VIC, JF694265; M. nuda Benth. var. nuda, Hughes 2396, Bolivia, FHO,
L. var. berlandieri (A.Gray) B.L.Turner, Camargo-Ricalde 531, Mexico, FJ982133; M. oblonga Benth., Barbosa 463, Brazil, HUEFS, FJ982135;
UAMIZ, FJ982147; M. pigra L. var. dehiscens (Barneby) D.Glazier & M. orthacantha Benth., Barros sn, Brazil, K, FJ982140; M. papposa
Mackinder, Hughes 2414, Bolivia, FHO, FJ982148; M. poculata Barneby, Benth. var. papposa, Simon 601, Brazil, UB, FJ982143; M. parviceps
Queiroz 10160, Brazil, HUEFS, JF694269; M. pseudosetosa M.F.Simon Barneby, Dutra 437, Brazil, VIC, JF694267; M. pectinatipinna Burkart,
& C.E.Hughes, Simon 864, Brazil, FHO, FJ982162; M. regina Barneby, Hughes 2036, Peru, FHO, FJ982144; M. pedersenii Barneby, Queiroz
Simon 759, Brazil, FHO, FJ982173; M. setosa Benth. var. paludosa 12645, Brazil, HUEFS, FJ982145; M. per-dusenii Burkart, Ribas 4545,
(Benth.) Barneby, Simon 725, Brazil, FHO, FJ982186; M. setosa Benth. Brazil, HUEFS, FJ982146; Mimosa sp., Hughes 2642, Peru, FHO,
var. urbica Barneby, Simon 730, Brazil, FHO, FJ982187; M. setosissima HM353125; M. polycarpa Kunth var. subandina Barneby, Hughes 2462,
Bolivia, FHO, FJ982154; M. polycephala Benth. var. polycephala, Simon Simon 746, Brazil, FHO, FJ982193; M. sousae R.Grether, Martínez-
400, Brazil, UB, FJ982155; M. polydactyla Humb. & Bonpl. ex Willd., Bernal 918, Mexico, UAMIZ, FJ982195; Mimosa sp., Wood 23707,
Coradin 8682, Brazil, CEN, FJ982156; M. pseudocallosa Burkart, Ribas Bolivia, K, JF694270; M. sparsiformis Barneby, Dahmer 15, Brazil, ICN,
5845, Brazil, HUEFS, FJ982159; M. pseudoradula Glaz. ex Barneby var. FJ982196; M. sprengelii DC., Queiroz 12469, Brazil, HUEFS, FJ982200;
pseudoradula, Simon 664, Brazil, UB, FJ982160; M. psilocarpa B.L.Rob., M. tequilana S.Watson, Simon 813, Mexico, MEXU, FJ982206; M.
Martínez-Bernal 933, Mexico, UAMIZ, FJ982163; M. pudica L., Simon tricephala Cham. & Schltdl. var. nelsonii (B.L.Rob.) Chehaibar &
669, Brazil, UB, FJ982166; M. pyrenea Taub., Simon 678, Brazil, UB, R.Grether, Martínez-Bernal 920, Mexico, UAMIZ, FJ982211; M. ursina
FJ982168; M. radula Benth. var. imbricata (Benth.) Barneby, Simon Mart., Simon 704, Brazil, FHO, FJ982217; M. velloziana Mart. var.
731, Brazil, FHO, FJ982170; M. ramboi Burkart, Queiroz 12530, Brazil, velloziana, Simon 721, Brazil, FHO, FJ982218; M. venatorum Barneby,
HUEFS, FJ982171; M. ramulosa Benth., Queiroz 12340, Brazil, HUEFS, Simon 740, Brazil, FHO, FJ982219; M. verecunda Benth., Simon 749,
FJ982172; M. rusbyana Barneby & Fortunato, Särkinen 2071, Bolivia, Brazil, FHO, FJ982220; M. vestita Benth., Simon 769, Brazil, FHO,
FHO, FJ982180; M. schleidenii Herter, Queiroz 12348, Brazil, HUEFS, FJ982222; M. virgula Barneby, Silva 5134, Brazil, UB, FJ982225; M.
FJ982182; M. sensitiva L. var. sensitiva, Almeida 4, Brazil, HUEFS, woodii Atahuachi & C.E.Hughes, Hughes 2285, Bolivia, FHO, FJ982231;
FJ982184; M. sicyocarpa B.L.Rob., Calóncio 4936, Mexico, MEXU, M. xanthocentra Mart. var. subsericea (Benth.) Barneby, Hughes 2403,
FJ982190; M. skinneri Benth. var. desmodioides (Benth.) Barneby, Bolivia, FHO, FJ982232.
Appendix 2. Circumscriptions of 24 well-supported clades (A–X) found in the phylogenetic analysis (Fig. 1).
Clade A is a robustly supported early-diverging group that is equivalent to and Farinosae (M. farinosa), with the Old World clade E (see below)
Barneby’s (1991) section Mimadenia, a group of 15 species, of which nested within it. The well-supported subclade formed by M. arenosa, M.
11 were sampled here. This group retains the ancestral character state acutistipula, M. bimucronata, M. pseudosepiaria, M. ophthalmocentra,
of extrafloral nectaries shared with piptadenioid sister genera (Fig. 2b), and M. farinosa is somewhat heterogeneous in terms of flower morphology,
while across the rest of the genus the nectary has been reduced to a including species with flowers with 3 or 4 petals, and both spicate and
nonfunctional spicule or been lost entirely. In addition, species in section globose inflorescences. This morphological diversity is reflected in
Mimadenia generally have more (up to 16) pollen grains per polyad, Barneby’s (1991) classification, in which these species are assigned to
whereas the remaining species of Mimosa have polyads of eight or four three different series based on flower and inflorescence characteristics. The
grains (Fig. 3b). Clade A is further subdivided into two geographically nonmonophyletism of most series within clade D precludes an estimation
and ecologically distinct subclades, one comprising Andean elements of of the number of species in this group.
mid-elevation seasonally dry ecology (M. revoluta, M. townsendii and
M. nothacacia), and the other lowland species widespread across the Clade E includes all Old World Mimosa species sampled here (half of those
neotropics, including a number of rain forest lianas, semi-arid thorn scrub, currently known) nested within the neotropical clade D, suggesting a
and savanna species (Fig. 1). possible single dispersal event from the Americas to the Old World (see
main text for further discussion). The Asian (M. prainiana, M. rubicaulis,
Clade B is a rather large and morphologically diverse clade containing around 65 and M. hamata) and continental African species (M. mossambicensis and
species and brings together several series of section Batocaulon, including M. busseana) form well-supported subclades within clade E. However, the
series Dystachyae (16 species), Acanthocarpae (13), Acantholobae (2), relationships of these two groups to the Malagasy species, which make
Boreales (10), Leucaenoideae (3), Rubicaules (4 excluding Old World up the majority of the Old World species are uresolved and still unclear.
species, but see Bentham, 1875), Fagaracanthae (7), Bahamenses (1), Clade E is formed by a morphologically homogeneus group of species
and M. leptocarpa (series Plurijugae), plus clade C. The clade comprises with inflorescences in capitula, tetramerous flowers, diplostemony, and
morphologically disparate elements that include convergent haplostemony presence of recurved aculei, although many of these characters are also
in M. leptocarpa and M. tejupilcana (Fig. 2d), but is geographically well- shared with other members of clade D. The affinity of the Old World
defined with all species centered in the USA, Mexico, the Caribbean, and species to the South American members of clade D was unexpected, since
Central America, except for the predominantly South American clade C they had been previously tentatively placed within series Rubicaules with
nested within (see below). None of the series represented in clade B, except M. ervendbergii, M. malacophylla, and M. hondurana (clade B) and M.
Acantholobae, are monophyletic (Table 2). However, in some cases, lack sinaloensis (clade G) (Bentham, 1875; Barneby, 1991). The apparent
of resolution and/or sparse taxon sampling preclude assessment of the morphological similarity between the Old World species and neotropical
monophyly of infrageneric groups. For example, series Fagaracanthae, members of series Rubicaules appears to be an example of independent
an essentially Caribbean group rich in endemics, is represented by only evolution of similar morphological features, which is fairly common in
one species (M. domingensis), and more samples are needed to confirm Mimosa.
the affinity of this series to the Mexican species.
Clade F corresponds in part to series Plurijugae and potentially contains
Clade C, nested within clade B, is a taxonomically heterogeneous group of four species, two of which were sampled here. This clade represents
around 10 species that combines members of three different series of section one of the five hypothesized transitions from ancestral diplostemonous
Batocaulon including series Andinae (5), Bimucronatae (M. hexandra ), and to haplostemonous flowers in Mimosa (Fig. 2d). Mimosa leptocarpa
Farinosae (M. detinens). The last two series are shown to be polyphyletic and M. tejupilcana, previously assigned to series Plurijugae (Barneby,
since some of their species are nested within clade D (see below). Clade C 1991; Grether and Martínez-Bernal, 1996), grouped within clade B, and
is likely to also include M. ostenii, which is morphologically similar to M. therefore appear to have aquired haplostemony independently.
detinens in series Farinosae (Barneby, 1991) and also grows in lowlands in
southern South America. The same would apply to M. exalbescens, which Clade G supports the monophyly of the morphologically unified series
is the putative sibling of M. hexandra in series Bimucronatae where these Quadrivalves (section Batocaulon) with the inclusion of a single species
are the only species with trimerous flowers (Barneby, 1991). This group (M. sinaloensis) from series Rubicaules. This clade is defined by a
comprises dry lowland species distributed across the Paraguay basin, the herbaceous diffuse or humifuse growth habit, stem armature of serial
Chaco, northeast Brazil; dry-forests in Colombia, Venezuela, and Mexico; aculei, and diplostemonous flowers with 5–6 petals and has a center of
and members of the ecologically and geographically well-defined series species diversity in southern USA and Mexico, with only one species
Andinae, which includes a set of narrowly distributed endemics from low- extending to South America. The distinct, narrow, tetragonal, valvately
to mid-elevation Andean dry tropical forests (Särkinen et al., 2011). dehiscent pod, in which the replum is as wide as or wider than the valves,
is characteristic of some taxa and gave the name to this series. However,
Clade D is formed by species of section Batocaulon series Caesalpiniifoliae (M. this is not a synapomorphy for this clade, since similar pods have been
caesalpiniifolia and M. laticifera), Ceratoniae (M. ceratonia), Leiocarpae reported in other unrelated groups, and also because some taxa within
(in part), Bimucronatae (M. bimucronata and M. pseudosepiaria), series Quadrivalves, as well as M. sinaloensis, have pods that resemble
a conventional craspedium (Barneby, 1991). Barneby (1991) reduced the series were mentioned by Barneby (1991) and are further discussed in
former genus Schrankia (= series Quadrivalves) to a single species, M. Simon et al. (2010). All species grow in the Cerrado, and the vast majority
quadrivalvis, with 16 varieties. However, this delimitation has not been are highly specialized fire-adapted local endemics. The adaptive strategies
widely accepted, with subsequent recognition of several taxa placed within and morphological innovations to withstand fire are strikingly diverse and
M. quadrivalvis as distinct species (Grether, 2000; Turner, 1994a, b). include rosulate trees (with leaves crowded at the tips of the branches),
thickened terminal branches, persistent stipules that protect the trunk
Clade H comprises a taxonomically widespread set of species mostly endemic
from fire, and prostrate or wand-like, functionally herbaceous subshrubs
to the Cerrado biome in Brazil. The nine species sampled here represent
growing from stout xylopodia (Simon et al., 2009).
four series of section Batocaulon, including series Paucifoliatae (17
species), Campicolae (3), Filipedes (6), and Echinocaulae (1), making Clade P is a small group comprising M. pudica, M. polydactyla (series Mimosa,
a possible total of 27 species. Most of these species are functionally subseries Pudicae), and M. affinis (subseries Affines). It is difficult to
herbaceous fire-adapted Cerrado endemics that grow from a napiform estimate the potential number of species in this clade because both these
xylopodium, but a few extend to coastal and southern Brazil, Venezuela, subseries were found to be polyphyletic. All three species have a minute
and Belize. One species, M. diplotricha, is a pantropical invasive weed, calyx, highly sensitive leaves, grow preferentially on disturbed sites
which is represented here by accessions from Brazil and Taiwan. Species such as roadsides, and have a tendency to become weedy. The group is
with trimerous flowers corresponding to series Campicolae and Filipedes widespread from tropical South America to Mexico (M. affinis). Mimosa
form a well-supported subgroup within clade H (Fig. 3a). pudica is introduced and widely naturalized across the tropics.
Clade I corresponds to series Cordistipulae, a morphologically well-defined Clade Q comprises series Modestae, except for M. skinneri, which was placed
group among the diplostemonous species characterized by trimerous in clade R. Species of clade Q grow mainly in northeast Brazil. The only
flowers, absence of interpinnal spicules, and paraphyllidia (Barneby, species of series Modestae not sampled here is endemic to the Caribbean
1991). The clade is particularly diverse in northeast Brazil, with one (M. viva), and its placement within clade Q awaits confirmation. All
widespread species (M. guaranitica) occurring in southern South America species in this clade have seismonastic leaf movements (Fig. 3c).
and also disjunct in Mexico. Ten of the 13 known species were sampled
here. Clade R is a large clade (probably 34 species) formed mostly by Cerrado
endemics and includes members of Barneby’s (1991) subseries
Clade J is equivalent to series Habbasia of Barneby (1991), a group of Polycephalae (20 species), Dicerastes (1), Discobolae (1), Hirsutae
potentially 10 species (four sampled here) of mostly wet tropical and (11), and M. skinneri (Modestae). Mimosa hirsutissima and M. skinneri
warm temperate lowlands. One species (M. pigra) is a widespread weed extend their ranges beyond the Cerrado, while three other species grow
in tropical wetlands. in dry grasslands in Paraguay and Argentina. Like species of clade O, this
Clade K is congruent with series Stipellares, a group of 19 species that grow group also shows diverse fire adaptations, including erect virgate stems
in open grasslands in subtropical South America, although sampling arising each year from a xylopodium, and many of them are functionally
within this clade is rather sparse (six species). The clade is defined by the herbaceous subshrubs. The monotypic subseries Dicerastes and Discobolae
presence of a dilated paraphyllidium that simulates a minute leaflet at the are nested within the clade, confirming their affinities as envisaged by
base of the pinna (Barneby, 1991). Barneby (1991). Resolution within this clade is very poor. The placement
of subseries Polycephalae, Dicerastes, Discobolae, and Hirsutae in the
Clade L combines series Bipinnatae (12 species, only three sampled here) in same clade seems morphologically plausible given the overall similarities
section Batocaulon and M. adenocarpa from series Glandulosae section in flowers, indumentum and foliage. However, the inclusion of M. skinneri
Habbasia. The clade is widespread in the neotropics, mostly in wetlands, in this group is unexpected and intriguing given its morphological affinites
and comprises some weedy species.
with species of clade Q.
Clade M contains all four species of series Neptunioideae, series Rojasianae
Clade S includes M. xanthocentra and M. verecunda in subseries Pudicae and M.
(one of three species sampled), and the monotypic series Auriculatae,
jacobita, and probably its putative sibling species in subseries Bipennatulae,
making a total of eight species of herbs and shrubs that grow in open
M. bipennatula (not sampled here). All these species are restricted to the
lowlands across the Neotropics, with a few species endemic to the
Cerrados of Brazil and Bolivia, except M. xanthocentra, which has a
savannas of Brazil and Bolivia. The placement of M. auriculata as sister
broader geographic range across South and Central America. All species
to M. josephina is intriguing, since they are morphologically very distinct,
share a paleaceous-pappiform calyx and almost free filaments.
but geographically adjacent (sometimes sympatric) local endemics from
the Serrania de Chiquitos in eastern Bolivia. Barneby (1991) placed M. Clade T is a taxonomically heterogeneous group mostly comprising species of
auriculata in a monotypic series in section Batocaulon, since he found no subseries Mimosa (all except M. acapulcensis and M. sousae), but also
obvious close relative for this species. However, all the other species in this includes M. casta (subseries Castae) and M. tequilana (subseries Affines).
clade are from section Habbasia. The presence of broad striate stipules The estimated number of species in this clade is nine (seven sampled here),
provides one trait shared by all members of this clade. The position of M. and most of them are widespread across the neotropics, being very common
orthocarpa (series Glandulosae), as sister to this clade, is unexpected in in disturbed sites. The inclusion of M. tequilana within this clade is not
terms of the striking morphological dissimilarity. surprising given its reduced leaf morphology of only one pair of pinnae,
Clade N comprises most of series Leiocarpae (probably 21 species, 14 sampled each with two pairs of large leaflets, a character shared by most species
here) along with M. lewisii (series Bimucronatae). These species are sampled in this group. One exception is M. casta, which has far more leaflets
mainly shrubs and trees with spicate inflorescences (except M. lewisii) per pinna, but which in everything else resembles M. velloziana (subseries
and glands on the leaflets, except M. dalyi and M. lewisii, although the Mimosa). The Mexican species of subseries Mimosa, M. acapulcensis,
latter has a glandular indumentum. The remaining species of series and M. sousae, which are morphologically very similar to species in clade
Leiocarpae, which lack leaflet glands and have white flowers, apart from T, are placed in the mostly Mexican clade V (see below). This and other
M. dayli, are placed within clade D (see above). The molecular data does examples stress the apparent importance of geography in structuring the
not support the hypothesis of Grether (2000) that the species of series phylogeny of Mimosa. Although Barneby (1991) placed these species in
Dystachyae (Mexican centered group) with spicate inflorescences should different subseries, he had doubts on the matter and noted several possible
be incorporated into series Leiocarpae, which is mainly South American, inconsistencies in this group and anticipated the relationships between M.
since these two groups appear to be distantly related on the plastid gene casta, M. tequilana, and M. acapulcensis, as found here.
tree. It would be interesting to include M. puberula in the analysis, as Clade U combines species from subseries Pedunculosae with one of the two
it is the only Mexican species included by Barneby (1991) in series varieties of the polyphyletic M. polycarpa (subseries Polycarpae) sampled
Leiocarpae, to verify its placement in the phylogeny. here, but as sampling within this group remains very sparse (only two of 10
Clade O is a large clade, remarkably rich in life forms and taxa. It comprises species of subseries Pedunculosae), more species need to be sequenced to
Barneby’s series Setosae and Pachycarpae with ca. 50 species, of which properly ascertain the composition of this clade. A link between subseries
26 were sampled here. Four recently described species (Simon et al., Pedunculosae and Polycarpae was already predicted by Barneby (1991).
2010) are included here. The plastid topology prompts incorporation of The species of this clade grow in open habitats in Paraguay, Argentina,
series Setosae into series Pachycarpae.The similarities between these two Bolivia, and southern Brazil.
Clade V comprises subseries Lactifluae (excluding M. incarum) and also M. Clade X is a large clade that incorporates series Myriophyllae and many of
acapulcensis and M. sousae (subseries Mimosa). Of a total of 11 potential the subseries proposed by Barneby (1991) in section Mimosa and also
species in this clade, eight were sampled. The range of the group is mainly includes the whole of section Calothamnos. This clade is largely distributed
in southern Mexico, with one species extending to Central America. The in southern South America where subtropical climates predominate
peculiar leaf morphology of M. acapulcensis and M. sousae, of a single pair (Argentina, Uruguay, Paraguay, and southern Brazil) and also in highland
of pinnae bearing two large leaflets each, is typical of subseries Mimosa areas at lower latitudes. In spite of poor resolution and insufficient
(see clade T), suggesting that this characteristic evolved independently taxon sampling in clade X, there are indications that some of Barneby’s
in the genus. There is weak support for this essentially Mexican clade as groups may be monophyletic, such as subseries Obstrigosae (clade
sister to the Andean clade W, a geographical sister group relationship also formed by M. ramboi, M. ramulosa, and M. sprengellii). Furthermore,
found for clade C. the possibility that section Calothamnos is monophyletic cannot be
dismissed, since the relationships between elements of this group were
Clade W is taxonomically disparate and is better defined by its geographical unresolved in the Bayesian analysis. The fact that the parsimony analysis
distribution restricted to the Andes than any obvious morphological or (Appendix S4) suggests that this group is monophyletic is explained by
classificatory coherence. This clade combines members of subseries the inclusion of indels in the parsimony analysis that were not included
Bolivianae, Pectinatae, Polycarpae (M. woodii and M. loxensis), and in the Bayesian analysis. An analysis based on plastid and nuclear DNA
also M. incarum (subseries Lactifluae) and M. rusbyana (subseries sequences containing a better taxon sampling of Calothamnos supports
Castae). Many of them are local endemics, restricted to single inter- its monophyletism and provide some internal resolution within this group
Andean dry valleys. (Savassi-Coutinho, 2009).

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American Journal of Botany 98(7): 1201–1221. 2011.

THE EVOLUTIONARY HISTORY OF MIMOSA (LEGUMINOSAE):

Character Anterior / Primitive Posterior / Derived

Petiolar nectary Present, functional Reduced to nonfunctional spicule, obsolete

Leaf movement—The sensitivity of Mimosa leaves to touch (seismonasty)

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