American Journal of Botany 88(3): 503–533. 2001.
THE
DALBERGIOID LEGUMES
(FABACEAE):
DELIMITATION OF A PANTROPICAL
MONOPHYLETIC CLADE1
MATT LAVIN,2,3 R. TOBY PENNINGTON,4 BENTE B. KLITGAARD,5
JANET I. SPRENT,6 HAROLDO CAVALCANTE DE LIMA,7 AND
PETER E. GASSON5
3
Department of Plant Sciences, Montana State University, Bozeman, Montana 59717 USA;
Tropical Biology Group, Royal Botanic Garden Edinburgh, 20a Inverleith Row, Edinburgh EH3 5LR, UK;
5
Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK;
6
Department of Biological Sciences, University of Dundee, Dundee DD1 4HN, UK; and
7
Jardim Botânico do Rio de Janeiro, Rua Pacheco Leão No. 915, Gavea 22.460 Rio de Janeiro—RJ, Brazil
4
A monophyletic pantropical group of papilionoid legumes, here referred to as the ‘‘dalbergioid’’ legumes, is circumscribed to include
all genera previously referred to the tribes Aeschynomeneae and Adesmieae, the subtribe Bryinae of the Desmodieae, and tribe
Dalbergieae except Andira, Hymenolobium, Vatairea, and Vataireopsis. This previously undetected group was discovered with phylogenetic analysis of DNA sequences from the chloroplast trnK (including matK) and trnL introns, and the nuclear ribosomal 5.8S
and flanking internal transcribed spacers 1 and 2. All dalbergioids belong to one of three well-supported subclades, the Adesmia,
Dalbergia, and Pterocarpus clades. The dalbergioid clade and its three main subclades are cryptic in the sense that they are genetically
distinct but poorly, if at all, distinguished by nonmolecular data. Traditionally important taxonomic characters, such as arborescent
habit, free stamens, and lomented pods, do not provide support for the major clades identified by the molecular analysis. Short shoots,
glandular-based trichomes, bilabiate calyces, and aeschynomenoid root nodules, in contrast, are better indicators of relationship at this
hierarchical level. The discovery of the dalbergioid clade prompted a re-analysis of root nodule structure and the subsequent finding
that the aeschynomenoid root nodule is synapomorphic for the dalbergioids.
Key words:
aeschynomenoid nodule; dalbergioid legumes; Fabaceae; papilionoid legumes; root nodule.
The ‘‘dalbergioid’’ legumes are a previously unrecognized
monophyletic group of papilionoid legumes in spite of the extensive taxonomic history of its four constituents: tribes Adesmieae, Aeschynomeneae, Dalbergieae, and Desmodieae subtribe Bryinae. The formal recognition of this group represents
a major rearrangement of papilionoid legumes. It combines
elements conventionally considered disparate and characterized as either ‘‘primitive’’ or having varying levels of ‘‘advancement’’ (Fig. 1).
The Dalbergieae originally included tropical trees with
fused floral parts and indehiscent pods (Bentham, 1860). Three
subtribes were recognized: Pterocarpeae with samaroid pods,
Lonchocarpeae marked by pods having at most small marginal
wings, and Geoffroyeae having drupaceous fruits. Polhill
(1971, 1981d, 1994) revised this classification by combining
morphological evidence with that of seed chemistry and wood
anatomy. This new Dalbergieae included 19 tropical woody
genera mainly from Bentham’s Pterocarpeae and Geoffroyeae.
Lonchocarpinae were relegated to a closer relationship with
other legumes that accumulated nonprotein amino acids in
seed (e.g., Evans, Fellows, and Bell, 1985). The revised Dalbergieae were diagnosed by supposedly plesiomorphic flower
morphologies (i.e., free keel petals, staminal filaments partly
fused and without basal fenestrae), pods with specialized seed
chambers, and seeds that accumulated alkaloids or other than
nonprotein amino acids. Geesink (1981, 1984) accepted Polhill’s circumscription with slight modification, whereas Sousa
and de Sousa (1981) proposed a classification similar to Bentham’s because Dalbergieae (sensu Polhill, 1981d) supposedly
shared a determinate inflorescence with the Lonchocarpinae.
The Aeschynomeneae (Rudd, 1981a) are one of five tribes
traditionally characterized by lomented pods (Polhill, 1981a).
Although some Aeschynomeneae lack such pods (e.g., Arachis, Ormocarpopsis, Diphysa spp., Ormocarpum spp., Pictetia
spp.), none of the members of this tribe have ever been confused or classified with the genera of Dalbergieae. Adesmieae
(Polhill, 1981f) have a notable history independent of the other
dalbergioid legumes. This is because this tribe combines a presumed plesiomorphic trait, free staminal filaments, with a supposedly very derived one, lomented pods. This combination
has suggested either a taxonomically isolated position or a
relationship with other papilionoids also with free stamens
(e.g., Burkart, 1952). Bryinae, with lomented pods, possess
other traits confirming its placement in the tribe Desmodieae
(e.g., explosive secondary pollen presentation; Ohashi, Polhill,
and Schubert, 1981). However, Bryinae have seeds that do not
accumulate nonprotein amino acids and lack a structural mu-
Manuscript received 11 January 2000; revision accepted 2 June 2000.
The authors thank Angela Beyra-M., Alfonso Delgado, Colin Hughes, JeanNoel Labat, Gwilym Lewis, Darien Prado, Mats Thulin, and Martin Wojciechowski for kindly providing seed or leaf material of many of the species
analyzed during this study, Alfonso Delgado, Martin Wojciechowski, and an
anonymous reviewer for providing comments that greatly improved the manuscript, Mats Thulin for making available his observations on the nectary disk
in Ormocarpum and close relatives, William Anderson for loaning copies of
the figures taken from Flora Novo-Galiciana, Sergio Faria for providing unpublished information on root nodule morphology, Karin Douthit, Shona
McInroy, and Maureen Warwick for illustrating the figures, and Tom Turley
for technical laboratory assistance. This study was supported by a grant from
the United States National Science Foundation (DEB-9615203), the Leverhulme Trust, and the Royal Botanic Garden Edinburgh Molecular Phylogenetic project.
2
Author for reprint requests (e-mail: mlavin@montana.edu).
1
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Fig. 1. Putative relationships among tribes of the subfamily Papilionoideae according to Polhill (1981a). Tribes underscored include genera that are now
known to be members of the dalbergioid clade (e.g., Desmodieae then included subtribe Bryinae, and Robinieae the genus Diphysa). Accumulation of nonprotein
amino acids and fusion of floral parts occur frequently in Tephrosieae and all tribes positioned above it. The absence of such traits is traditionally viewed as
primitive and is most frequent in tribes positioned below Tephrosieae.
tation in the chloroplast rpl2 locus (Bailey et al., 1997). Both
are atypical of the rest of Desmodieae.
In spite of a taxonomic history of Dalbergieae that has been
separate from those of Aeschynomeneae, Adesmieae, and
Bryinae, we present evidence that they collectively form a
monophyletic group. The focus on these putatively disparate
taxa was motivated by the taxonomic distribution of the distinctive aeschynomenoid root nodule (Corby, 1981; Faria et
al., 1994) and four cladistic analyses: three involving nonmolecular data (Lavin, 1987; Chappill, 1995; Beyra-M. and
Lavin, 1999), and one with rbcL sequence data (Doyle et al.,
1997). We have expanded on these previous analyses by sampling exhaustively to reveal the exact constituents of the dalbergioid clade and enumerate the nonmolecular characters that
have been used in the conventional tribal classification of these
legumes. As such, we demonstrate where molecular and nonmolecular data are taxonomically concordant. We also show
that many traditionally important taxonomic characters in this
group are more homoplasious than previously considered. Because taxon sampling has focused on just the putative members of the dalbergioid clade, a point to be briefly addressed
here but more thoroughly developed elsewhere is the higher
level relationships of this newly recognized clade (Hu et al.,
2000; Pennington et al., in press; M. Wojciechowski et al.,
unpublished data).
MATERIALS AND METHODS
DNA sequence data—DNA isolations, polymerase chain reaction (PCR)
amplifications, and template purifications were performed with Qiagen Kits
(i.e., DNeasy Plant Mini Kit, Taq PCR Core Kit, QIAquick PCR Purification
Kit; Qiagen, Santa Clarita, California, USA). DNA sequences analyzed were
the nuclear ribosomal 5.8S and flanking internal transcribed spacers (ITS1
and ITS2), the chloroplast trnK intron, including matK, and the trnL intron.
PCR and sequencing primers for ITS and 5.8S sequences are described in
Beyra-M. and Lavin (1999) and Delgado-Salinas et al. (1999). Primers for
matK and flanking trnK intron sequences are described in Lavin et al. (2000).
Primers for the trnL intron are described by Taberlet et al. (1991). DNA
sequencing was performed on an automated sequencer at the Iowa State University DNA Sequencing Facility (Ames, Iowa, USA) and Davis Sequencing
(Davis, California, USA).
DNA sequences were aligned manually with Se-Al (Rambaut, 1996). Bias
introduced by the manual alignment was evaluated with a sensitivity analysis
(cf. Whiting et al., 1997; Beyra-M. and Lavin, 1999; Delgado-Salinas et al.,
1999). Alignment-variable regions were variably aligned or excluded, a step
matrix (cf. Cunningham, 1997) was invoked or not, and gaps were treated as
missing data, a fifth state, or as separate characters. Each of the different
sensitivity analyses were subjected to the same heuristic search options. Missing data included 12.9% of the matK/trnK data set, 5.4% of the trnL data set,
1.5% of the ITS/5.8S data set, and 7.6% of the nonmolecular data set.
Maximum parsimony analyses were performed with PAUP* (Swofford,
2000). Heuristic search options included 100 random-addition replicates, treebisection-reconnection branch swapping, and steepest descent. A maximum
of 10 000 trees was allowed to accumulate, which is sufficient to capture all
topological variation (cf. Sanderson and Doyle, 1993). Clade stability tests
involved bootstrap resampling (Felsenstein, 1985; Sanderson, 1995), where
each of the 10 000 bootstrap replicates was subjected to heuristic search options that included one random-addition sequence per replicate, swapping with
tree-bisection-reconnection, and invoking neither steepest descent nor mulpars.
Taxon sampling—Sampling of molecular and nonmolecular data was as
exhaustive as possible at the generic level in order to determine membership
in the dalbergioid clade, as well as the principal phylogenetic structure within
this clade. Molecular and nonmolecular data were obtained for at least one
species from every genus ever placed in the Dalbergieae (Burkart, 1952; Polhill, 1981d), Aeschynomeneae (Rudd, 1981a), Adesmieae (Polhill, 1981f), or
Bryinae (Ohashi, Polhill, and Schubert, 1981). The only exception is the presumably extinct genus Peltiera (Labat and Du Puy, 1997), where no successful PCR amplifications were obtained from the few available DNAs. In addition to the advantages of being able to detail the taxonomic implications,
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exhaustive sampling for molecular data increases the probability of subdividing long branches (e.g., Hillis, 1998).
Our original intent was to sample the same DNA accessions for each of
the data sets. This proved impossible for DNA sequences because of inconsistencies in DNA quality and quantity and PCR amplification. We consequently had to resort to multiple methods of sampling. The DNA sequence
data were sampled using the exemplar approach. Multiple species per terminal
taxon were sampled where possible (Appendix A). Because nonmolecular data
are generally open to visual inspection across all species of a particular terminal taxon, the ‘‘democratic’’ method of sampling (Bininda-Emonds, Bryant,
and Russell, 1998) was used for nonmolecular data. In this approach, we
included all possible character states represented by any one terminal, which
was usually a traditionally recognized genus (i.e., multistate terminal taxa
were coded). The reasoning is that in the evaluation of traditionally important
taxonomic characters, the degree of polymorphisms within terminals should
be explicitly enumerated. For those few terminals in which species-level phylogenetic analysis has been completed (e.g., Andira and Pictetia), we employed the ancestral method of sampling nonmolecular data (Bininda-Emonds,
Bryant, and Russell, 1998). The justification for ultimately combining data
that have been sampled differently is that a combined analysis should still
allow us to best estimate where the traditionally important taxonomic characters lie on the continuum from strongly phylogenetically constrained to
maximally homoplasious.
The genera Bergeronia, Dalbergiella, Lonchocarpus, and Muellera have
been placed in the tribe Dalbergieae (e.g., Burkart, 1952; Geesink, 1981) and
Pongamiopsis has been synonymized with the genus Aeschynomene (Hutchinson, 1964). However, they were not included in this analysis because other
phylogenetic analyses (Lavin et al., 1998; Hu et al., 2000) have shown these
genera to be closely related to Millettia and relatives, all of which accumulate
nonprotein amino acids in seed. Similarly, Poecilanthe and Cyclolobium
should be allied with more basal Papilionoideae that accumulate alkaloids in
seed (Greinwald et al., 1995; Lavin et al., 1998; Hu et al., 2000). This is the
reason that Poecilanthe is retained as a designated outgroup.
Outgroups were sampled extensively as part of large-scale molecular phylogenetic studies of the subfamily Papilionoideae (Hu et al., 2000; Pennington
et al., in press; M. Wojciechowski et al., unpublished data). Sampling outgroups was guided by phylogenetic studies involving nonmolecular data (e.g.,
Chappill, 1995; Herendeen, 1995; Beyra-M. and Lavin, 1999). For example,
all outgroups chosen have leaves with punctate glands, a trait common to
dalbergioids. In the end, the outgroups retained in this analysis included Acosmium and Myrospermum (tribe Sophoreae; Polhill, 1981b), Dipteryx and Pterodon (Dipterygeae; Polhill, 1981c), Poecilanthe (variously classified; see Lavin and Sousa, 1995), and Apoplanesia, Amorpha, Eysenhardtia, and Marina
(tribe Amorpheae; Barneby, 1977; Polhill, 1981e). This sampling was considered sufficient to demonstrate membership in the dalbergioid clade. The findings reported here did not change with a more extensive sampling of outgroups.
Sampling for the molecular data was re-evaluated as aligned DNA sequences accumulated. It became obvious that the matK/trnK sequences were by far
the most informative at higher taxonomic levels, as seen in increased resolution in the strict consensus and higher bootstrap values. The primary effort
then changed to sample as exhaustively as possible matK/trnK sequences and,
secondarily, the ITS/5.8S and trnL intron sequences. Thus, the data analysis
of this study centers on the matK/trnK data set. Sampling of ITS/5.8S sequences was guided by species level analyses of certain dalbergioid genera
(e.g., Beyra-M. and Lavin, 1999; Lavin et al., 2000). Sampling of the trnL
intron data was guided by a phylogenetic analysis of putatively basal Papilionoideae (Pennington et al., in press). Unevenness in sampling was exacerbated by inconsistencies in PCR amplifications (mentioned above). A combined molecular analysis was not attempted because unevenness in sampling
would result in a combined data set not exhaustively sampled at the genus
level. Thus, consensus among the data sets was evaluated by congruence of
the major clades resolved with high bootstrap values (cf. Huelsenbeck, Bull,
and Cunningham, 1996).
Nonmolecular character analysis—A nonmolecular data set was devel-
505
oped from that in Beyra-M. and Lavin (1999) and is presented in Appendix
B. Characters that have been considered traditionally important in the taxonomy of Dalbergieae, Aeschynomeneae, Adesmieae, and Bryinae (e.g., Burkart, 1952; Ohashi, Polhill, and Schubert, 1981; Polhill, 1981d; Rudd, 1981a;
Sousa and de Sousa, 1981) were targeted for analysis. As discussed above,
multistate taxa were coded as polymorphic (cf. Weins, 1995; Weins and Servedio, 1997), in spite of the recommendation of Nixon and Davis (1991).
Although this can underestimate the degree of homoplasy (see individual character discussions in Appendix B), splitting polymorphic terminals into two or
more monomorphic ones does not change our findings (e.g., as evaluated in
the fashion of a sensitivity analysis). This is because the focus is strictly at
wide-scale relationships of groups of genera, and the potentially problematic
polymorphisms are at a different level, within genera. Polymorphisms are
discussed in the presentation of characters or ingroup terminal taxa (Appendices B and C). Inapplicable character states in certain terminals (e.g., leaf
traits of Ramorinoa, a genus that doesn’t produce leaves) were variously treated as a missing state, an uncertain state, or an extra state (as in a sensitivity
analysis). The nonmolecular data were gathered primarily from field observations or herbarium specimens. Literature reports were usually verified by
observations of the plants.
RESULTS
Parsimony analysis of the 1266 informative sites from the
95 taxa by 2966 sites matK/trnK data set produced 10 000 trees
(the set maximum) each with a minimal length of 4352, a
consistency index of 0.570 and a retention index of 0.830. The
monophyly of the dalbergioid clade, including all genera of
Aeschynomeneae, Adesmieae, Bryinae, and most Dalbergieae,
was very well supported by bootstrap analysis (Fig. 2). Four
members of tribe Dalbergieae (Andira, Hymenolobium, Vatairea, and Vataireopsis) and two sampled genera of Dipterygeae
(Dipteryx and Pterodon) were not included. Indeed, the sister
group to the dalbergioid clade includes genera sampled from
the tribe Amorpheae (Apoplanesia and Amorpha). Within the
dalbergioid clade, there are three well-supported subclades
marked as the Adesmia, Dalbergia, and Pterocarpus clades
(Fig. 2). The earliest branching Adesmia clade includes the
genus Adesmia (sole member of the tribe Adesmieae) and
mostly herbaceous to subshrubby genera of the tribe Aeschynomeneae (Poiretia, Amicia, Zornia, Chaetocalyx, and Nissolia). The remaining two subclades each include members of
the Aeschynomeneae and Dalbergieae. The Pterocarpus clade
additionally includes two genera, Brya and Cranocarpus, of
Desmodieae (subtribe Bryinae).
For the 481 informative sites from the 118 taxa by 719 sites
ITS/5.8S data set, 120 trees were generated each with a minimal length of 5009, a consistency index of 0.259, and a retention index of 0.714. The same higher level relationships
described for the matK/trnK analysis were resolved in this
analysis, though with less bootstrap support (Fig. 3). Although
the Pterocarpus clade was resolved in the strict consensus of
the parsimony analysis, it was resolved in less than 50% of
the analyses of the bootstrap replicates. In no case (majorityrule bootstrap consensus or strict consensus of minimal length
trees) was the sister-group relationship of the Amorpheae samples resolved.
Analysis of the 293 informative sites from the 93 taxa by
737 sites trnL intron data set generated 10 000 trees each with
a minimal length of 1102, a consistency index of 0.603, and
a retention index of 0.804. Although the dalbergioid clade is
well resolved by bootstrap analysis, only the Adesmia clade
is further resolved (Fig. 4). Not in any case was the Dalbergia
or Pterocarpus clades resolved as monophyletic. Regardless,
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Fig. 2. Bootstrap majority rule (50%) consensus from the analysis of matK/trnK sequences. The dalbergioid clade and its three constituent subclades are
indicated.
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507
Fig. 3. Bootstrap majority rule (50%) consensus from the analysis of ITS/5.8S sequences. The dalbergioid clade and two of its three constituent subclades
are indicated. The clade marked by a closed circle was also detected in the analysis of matK/trnK and trnL intron sequences.
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Fig. 4. Bootstrap majority rule (50%) consensus from the analysis of trnL intron sequences. The dalbergioid clade and the Adesmia subclade are indicated.
Clades marked by a closed circle were also detected in the analysis of matK/trnK and ITS/5.8S sequences.
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the relationships resolved by majority-rule bootstrap consensus
did not conflict with those similarly resolved in either the
matK/trnK and ITS/5.8S analyses.
Analysis of the 55 nonmolecular characters (Appendix B)
yielded poorly resolved and supported relationships, such that
the majority-rule bootstrap consensus was largely unresolved
above the genus level. Resolved intergeneric relationships include a clade with Aeschynomene, Cyclocarpa, Bryaspis, Geissaspis, Humularia, Kotschya, Smithia, and Soemeringia (60%
bootstrap support), one with Chapmannia, Arachis, and Stylosanthes (65%), Brya and Cranocarpus (67%), Chaetocalyx
and Nissolia (100%), Amicia, Poiretia, and Zornia (67%), and
Ormocarpopsis and Peltiera (93%). Because Peltiera is not
represented by DNA sequence data, this nonmolecular data
provide the only evidence for its relationships (the relationships of Peltiera are a focus of another study; M. Thulin and
M. Lavin, unpublished data). The only well-supported clade
that was resolved during this analysis and that was not seen
during the previous molecular analyses was one with Etaballia
and Inocarpus (80%), apomorphically diagnosed as having
nearly regular flowers (characters 22–23 in Appendix B).
Because of the poorly resolved relationships obtained from
analysis of the nonmolecular data set, it was combined with
the matK/trnK data set in order to explore the evolution of the
traditionally important taxonomic characters. Integration with
just the matK/trnK is justified by how well this data set can
resolve relationships (discussed in MATERIALS AND
METHODS) and because of noncompatibility of molecular
data sets with respect to sampling. Parsimony analysis of the
1319 informative characters of the combined matK/trnK and
nonmolecular data set (95 taxa by 3021 characters) produced
2340 trees with a minimal length of 4664, each with a consistency index of 0.551 and a retention index of 0.821. The
resulting relationships are essentially those described previously for the analysis of just the matK/trnK data set (Fig. 5).
Sensitivity analysis—Making different assumptions about
the molecular data sets, deleting characters with many missing
entries (e.g., nonmolecular characters 50–54), splitting polymorphic terminals into two or more monomorphic ones, or
recoding inapplicable nonmolecular characters to uncertain
states, missing data, or as an extra state, did not affect the
results described above (Figs. 2–5). The monophyly of the
dalbergioid legumes was consistently resolved, as generally
was the monophyly of the three constituent subclades. There
were no cases of clades with bootstrap values over 70% that
conflicted among the molecular data sets. Also, clades with
high bootstrap values (i.e., .90%) in individual analyses of
the matK/trnK, ITS/5.8S, trnL intron, or combined nonmolecular and matK/trnK data sets were consistently resolved regardless of the assumptions made about any one of the particular data sets. This is exemplified by analysis of just the matK
coding region (i.e., excluding the flanking noncoding portion
of the trnK intron), where some accessions in the data matrix
were missing either the 59 or 39 half of this locus (for a total
of 12.1% missing entries). The strict consensus of the parsimony analysis of the matK locus was essentially identical to
that of the analysis of the matK/trnK data set. Bootstrap analysis resulted in values that were sometimes lower than in the
analysis of the entire matK/trnK data set: 80% for the Amorpheae 1 dalbergioid clade, 100% for the dalbergioid clade,
100% for the Adesmia clade, 94% for the Dalbergia clade, and
71% for the Pterocarpus clade.
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DISCUSSION
As now circumscribed, the dalbergioids comprise 44 genera
(Appendix C) and ;1100 species of trees, shrubs, and perennial to annual herbs. Included are economically important
hardwoods (e.g., Dalbergia and Pterocarpus spp.), forage legumes (Stylosanthes spp.), and crops (e.g., Arachis spp.). Like
most pantropical legume taxa, the dalbergioids are concentrated in the neotropics and subSaharan Africa. Although the
position of the dalbergioid clade within the Fabaceae is not
fully developed here, its sister group is the tribe Amorpheae,
which contains eight New World genera confined mostly to
warm temperate and tropical North America. What is generally
certain of higher level relationships is that the dalbergioids are
distantly related to papilionoids that accumulate nonprotein
amino acids in seed. This most notably includes Lonchocarpus, Derris, Millettia, and Hologalegina (e.g., tribes Robinieae, Galegeae, etc.; Wojciechowski, Sanderson, and Hu,
1999), which at times have been taxonomically confused with
various elements now included in the dalbergioid clade.
Implications for traditional classifications—The classification of certain genera into tribes and subtribes of Papilionoideae (e.g., Rudd, 1981a; Ohashi, Polhill, and Schubert,
1981; Ohashi, 1999; Polhill, 1981a, d) needs to be greatly
modified in light of the evidence presented here. The genera
Brya and Cranocarpus (subtribe Bryinae of tribe Desmodieae)
share many unusual synapomorphies, such as periporate pollen
and glochidiate trichomes, that have served to obscure higher
level relationships. The explosive pollen presentation mechanism that Brya shares in common with Desmodieae is shown
to have evolved independently. So have the lomented pods that
Brya and Cranocarpus share with Desmodieae.
Four of the five subtribes of Aeschynomeneae are either
monotypic (e.g., Discolobiinae) or are polyphyletic. Aeschynomeneae subtribe Ormocarpinae includes three different elements: Diphysa, Ormocarpum, Ormocarpopsis (and Peltiera), and Pictetia form one lineage in the Dalbergia clade,
Fiebrigiella is in the Pterocarpus clade, and Chaetocalyx and
Nissolia are part of the Adesmia clade. The pod valves with
distinctive parallel venation that previously allied all of these
genera now are considered to have evolved on three separate
occasions. Indeed, this derived pod trait is homologous among
Fiebrigiella, Chapmannia, Arachis, and Stylosanthes.
Aeschynomeneae subtribe Poiretiinae includes two different
elements. Amicia, Poiretia, and Zornia form a monophyletic
group within the Adesmia clade, and Weberbauerella is phylogenetically isolated within the Dalbergia clade. The marked
pustular glands of Weberbauerella are no longer considered
homologous to those of Amicia, Poiretia, and Zornia. In the
recent classification of Japanese legumes (Ohashi, 1999), Poiretia and Zornia are classified as the sole members of the tribe
Poiretieae, a taxonomy that finds no support in this analysis.
Aeschynomeneae subtribe Aeschynomeninae includes eight
genera (Aeschynomene, Cyclocarpa, Soemmeringia, Kotschya,
Smithia, Humularia, Bryaspis, and Geissaspis) that form a
very well-supported monophyletic group. A nonmolecular
character supporting this relationship is the medifixed stipule,
although it is not universal in this clade and has evolved independently in Zornia. An extrapolation from our small sample, however, suggests that species of Aeschynomene having
basifixed stipules (e.g., A. fascicularis and A. purpusii) are
more closely related to Machaerium and Dalbergia than they
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Fig. 5. Bootstrap majority rule (50%) consensus from the analysis of combined nonmolecular and matK/trnK sequence data. The dalbergioid clade and its
three constituent subclades are indicated.
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are to the species of Aeschynomene with medifixed stipules.
Thus, the subtribe Aeschynomeninae includes two disparate
elements.
Only Aeschynomeneae subtribe Stylosanthinae, with Arachis, Stylosanthes, and Chapmannia (and the segregates Pachecoa and Arthrocarpum), has been long recognized as a distinct
taxonomic group and is also revealed as monophyletic in this
analysis. The well-known nonmolecular character supporting
the monophyly of this clade is a sessile papilionoid flower with
a long hypanthium. However, these three genera are very
closely related to Fiebrigiella and Fissicalyx and together all
of these genera are set apart from other members of the Pterocarpus clade by large genetic distances. Notably, nonmolecular characters do not support most of the relationships in this
clade that are so well supported by independent molecular
data. For example, there are no known nonmolecular data that
support the monophyly of the genus Chapmannia (Thulin,
2000) or the relationship of Fissicalyx and Fiebrigiella.
The tribe Dalbergieae also is not monophyletic. Excluded
from the dalbergioid clade are Andira, with 30 species largely
confined to the neotropics and with one species distributed in
the neotropics and tropical Africa (Lima, 1990; Pennington,
1996; Pennington, Aymard, and Cuello, 1997), Hymenolobium
with 10–15 species in tropical South America and one species
in Central America (Polhill, 1981d; Lima, 1982a, 1990), Vatairea with seven species from Mexico to Brazil (Lima, 1982b,
1990), and Vataireopsis with three species in Brazil and the
Guianas (Polhill, 1981d; Lima, 1990). The distinction of these
four genera from others traditionally included in the tribe Dalbergieae has been noted with wood anatomy (Baretta-Kuipers,
1981) and estimates of overall similarity (Lima, 1990). For
example, the wood of Andira, Hymenolobium, Vatairea, and
Vataireopsis lacks the storied structure and uniseriate rays that
are characteristic of dalbergioid wood and is generally of less
commercial value.
The remaining genera of the tribe Dalbergieae belong to
either the Dalbergia or Pterocarpus clades. Only Dalbergia and
Machaerium are part of the Dalbergia clade, where they are
most closely related to Aeschynomene species that have basifixed stipules. The rest of the genera previously classified in
the tribe Dalbergieae form the bulk of the Pterocarpus clade
along with some genera previously classified in the tribe Aeschynomeneae (e.g., Fiebrigiella, Chapmannia, Arachis, Stylosanthes, and Discolobium) and subtribe Bryinae of Desmodieae.
The genera of Dipterygeae (Taralea, Dipteryx, and Pterodon; Polhill, 1981c) are not part of the dalbergioid clade. Burkart (1952) originally included Dipteryx (then Coumarouna)
in the tribe Dalbergieae, and a phylogenetic analysis of nonmolecular data by Beyra-M. and Lavin (1999) suggested Dipterygeae was part of the dalbergioid clade. Even the combination of paripinnate leaves bearing glandular punctae is
known only from Dipterygeae and the dalbergioid legumes.
However, this analysis strongly suggests that the punctate
glands are plesiomorphic because they are found in all genera
included in this analysis. Paripinnate leaves evolved independently among Dipterygeae and various elements in the dalbergioid clade.
Phylogenetic information among the various nonmolecular
characters—While the matK/trnK phylogeny was not greatly
influenced by the addition of the 55 nonmolecular characters
(compare Figs. 2 and 5), there is some phylogenetic infor-
511
mation in the nonmolecular characters, as evinced by high retention indices (Table 1). The consistency (CI) and retention
(RI) indices for each of the 55 nonmolecular characters (Appendix B) in the combined analysis were compared to the
same values obtained when each of the nonmolecular characters was mapped onto the matK/trnK phylogeny. In the combined analysis, the average CI and RI were 0.427 and 0.672,
respectively. When mapped onto the matK/trnK trees, the average CI and RI were 0.390 and 0.627, respectively. Regardless of the small but significant differences (for RI, two-tailed
t test, t 5 2.94, P 5 0.005, df 5 52), no character had a
higher consistency or retention index when mapped onto the
matK/trnK phylogeny as when combined with the matK/trnK
sequence data during parsimony analysis. This suggests that
mapping a few selected nonmolecular characters onto a molecular phylogeny may involve a bias of excess levels of homoplasy.
Different classes of characters (e.g., vegetative, floral, and
fruiting) were equally as prone to having homoplasy overestimated when mapped onto a molecular phylogeny. These include, for example, an asymmetric leaflet base (character 9 in
Appendix B), persistent floral bracts (character 16), and a long
pod stipe (character 33). The states of the leaflet base had an
average retention index of 1.000 in the combined analysis and
0.500 when mapped to the matK/trnK trees (Table 1). The
corresponding values were 0.667 and 0.167 for the states of
the floral bracts, and 0.647 and 0.559 for the pod stipe (Table
1). Also, no particular class of characters (e.g., vegetative, floral, and fruiting) was more informative than another. For vegetative characters (1–13, 44–45, 50–55), the average retention
index is 0.705. For floral characters (14–30, 46–49), it is
0.604. For fruiting characters (31–43), the average retention
index is 0.726. These differences are not significant (singlefactor ANOVA, F 5 1.174, P 5 0.317, df 5 52). The lack
of a difference in behavior among the various classes of characters, as also generally found by Bateman and Simpson
(1998) for vascular plants, weakens the suggestion of Tucker
and Douglas (1994) that floral characters necessarily provide
the best taxonomic information in Leguminosae. These findings also weaken the implication that pod morphology is prone
to higher rates of convergent evolution than other types of
characters (e.g., Geesink, 1984; Hu et al., 2000).
Conventional taxonomic evidence—Some traditionally important taxonomic characters are determined in this analysis to
be more homoplasious than previously considered. This is especially true of the character states of growth habit, staminal
fusion, and pod segmentation. Herbaceous and woody relatives
generally are separated into different taxonomic groups when
a temperate vs. tropical distinction correlates with habit (Judd,
Sanders, and Donoghue, 1994). This is especially true of papilionoid legumes where tribes have been categorized by habit
(e.g., temperate herbaceous vs. tropical woody tribal division
in Polhill, 1981a, 1994). An herbaceous habit (number 1 in
Appendix B) has evolved at least three times in monomorphic
condition but more times than this in polymorphic condition
(Table 1). The Adesmia clade contains mostly herbaceous species, although some species of Adesmia and Poiretia are
shrubs. That an herbaceous growth form maps as the ancestral
state in the Adesmia clade stands in contrast to the conventional wisdom that woody taxa form basal clades in tropical
Papilionoideae (e.g., Polhill, 1981a; Tucker and Douglas,
1994).
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[Vol. 88
TABLE 1. Average lengths (L) and consistency (CI) and retention (RI) indices for each of the 55 nonmolecular characters. These are compared
for the combined analysis and when each of the 55 is mapped onto the matK/trnK phylogeny. An ‘‘5’’ indicates that the CI and RI of the
combined and mapped character are equal. A ‘‘.’’ signifies a higher CI and RI value for a character in the combined analysis compared to
when mapped. The reverse situation did not occur.
Combined analysis
Mapped
Number
L
CI
RI
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
Mean
SD
5.5
1.0
6.0
2.0
8.0
5.0
4.0
8.0
1.0
2.0
9.0
7.0
2.0
2.0
7.0
2.0
10.0
10.0
6.0
4.0
4.0
5.0
2.0
8.0
5.0
15.0
2.0
10.0
5.0
1.0
9.0
6.0
13.0
2.5
1.0
17.0
2.0
5.0
1.0
2.0
3.0
11.0
4.0
5.0
7.0
1.0
14.5
3.0
6.0
2.0
4.0
7.0
3.0
1.0
7.0
5.4
3.9
0.417
1.000
0.333
0.500
0.125
0.200
0.500
0.125
1.000
0.500
0.111
0.143
1.000
0.500
0.429
0.500
0.200
0.200
0.833
0.500
0.250
0.200
0.500
0.125
0.200
0.267
0.500
0.200
0.200
1.000
0.250
0.167
0.077
0.417
1.000
0.235
0.500
0.400
1.000
1.000
0.667
0.182
0.250
0.200
0.143
1.000
0.069
0.333
0.500
0.500
0.250
0.143
0.333
1.000
0.286
0.427
0.301
0.850
1.000
0.667
0.833
0.562
0.789
0.500
0.767
1.000
0.833
0.724
0.625
1.000
0.667
0.778
0.667
0.778
0.778
0.955
0.833
0.812
0.200
0.000
0.364
0.429
0.633
0.857
0.800
0.000
0.825
0.583
0.647
0.912
0.409
0.889
0.625
1.000
1.000
0.750
0.500
0.571
0.600
0.500
1.000
0.625
0.333
0.571
0.750
0.400
0.739
0.000
1.000
0.667
0.672
0.254
Genera containing both woody and herbaceous species also
occur in the clade containing Aeschynomene sect. Aeschynomene, Kotschya, Humularia, and Geissaspis. The same is true
for the clade including Fiebrigiella, Chapmannia, Stylosan-
.
5
5
5
5
5
5
5
.
5
.
5
5
5
.
.
.
.
.
5
5
5
5
5
5
5
5
5
5
5
.
.
.
.
5
.
5
5
5
5
5
.
5
.
5
5
.
5
.
5
5
5
5
.
5
L
CI
RI
6.0
1.0
6.0
2.0
8.0
5.0
4.0
8.0
2.5
2.0
10.0
7.0
2.0
2.0
9.0
3.5
11.5
12.0
7.0
4.0
4.0
5.0
2.0
8.0
5.0
15.0
2.0
10.0
5.0
1.0
10.0
7.0
16.0
3.0
1.0
17.5
2.0
5.0
1.0
2.0
3.0
12.0
4.0
6.0
7.0
1.0
15.0
3.0
8.0
2.0
4.0
7.0
3.0
2.0
7.0
5.8
4.2
0.333
1.000
0.333
0.500
0.125
0.200
0.500
0.125
0.417
0.500
0.100
0.143
1.000
0.500
0.333
0.292
0.175
0.167
0.714
0.500
0.250
0.200
0.500
0.125
0.200
0.267
0.500
0.200
0.200
1.000
0.200
0.143
0.062
0.333
1.000
0.229
0.500
0.400
1.000
1.000
0.667
0.167
0.250
0.167
0.143
1.000
0.067
0.333
0.375
0.500
0.250
0.143
0.333
0.500
0.286
0.390
0.279
0.800
1.000
0.667
0.833
0.562
0.789
0.500
0.767
0.500
0.833
0.690
0.625
1.000
0.667
0.667
0.167
0.736
0.722
0.909
0.833
0.812
0.200
0.000
0.364
0.429
0.633
0.857
0.800
0.000
0.800
0.500
0.559
0.882
0.387
0.889
0.625
1.000
1.000
0.750
0.444
0.571
0.500
0.500
1.000
0.611
0.333
0.286
0.750
0.400
0.739
0.000
0.667
0.667
0.627
0.258
thes, and Arachis. Fissicalyx and some species of Chapmannia
are woody in a clade dominated by herbaceous to subshrubby
species. Representing yet two other clades, species of Machaerium, Dalbergia, Brya, and Cranocarpus vary from trees
March 2001]
LAVIN
ET AL.—DALBERGIOID LEGUMES
513
Figs. 6–8. Selected nonmolecular characters (scale bar 5 1 cm for all figures). 6. Aeschynomenoid root nodule associated with lateral root (character number
55, Appendix B). 7. Short shoots of Ormocarpum (character number 2). 8. Pseudopetiole of Arachis (character number 4).
or shrubs to weak subshrubs. Clearly, there is no evidence
from this analysis that the ability to produce a strongly woody
growth habit is a good indicator of relationship.
The staminal character number 26 (Appendix B) includes
five states that provide an average length of 15.0 to the most
parsimonious trees. The consistency index of 0.267 and the
retention index of 0.633 demonstrate that this character is
homoplasious. Even state zero, free staminal filaments, added
a length of two because this state occurs ancestrally in some
of the outgroup genera and represents a reversion in the genus
Adesmia. That a legume group with free stamens can evolve
this condition secondarily from a fused condition (e.g., 9 1 1
diadelphous) is not surprising. Four species of Pictetia have
nearly free staminal filaments in a clade otherwise represented
by species with fused filaments (Beyra-M. and Lavin, 1999).
Also, Käss and Wink (1995, 1997) have implicitly shown in
an unrelated papilionoid group that the evolution of staminal
morphology does not necessarily involve a unique transformation from free filaments into the fused condition. Perhaps
related to this issue, Klitgaard (1999a) showed that order of
initiation and loss of stamens are more variable among the
dalbergioids than previously appreciated. No doubt, the a
priori view that free staminal filaments represent necessarily a
plesiomorphic condition among papilionoid legumes will have
to be abandoned.
All papilionoid legumes with lomented pods were at one
time classified together, although more recently five tribes
(Adesmieae, Aeschynomeneae, Coronilleae, Desmodieae, and
Hedysareae) were thought to have gained this pod type independently (Polhill, 1981a). We scored three states pertaining
to articulation of pod segments (number 31 in Appendix B),
which added an average length of 9.0 to the most parsimonious trees. The consistency index of 0.250 and a retention index
of 0.825 suggest that, although homoplasious, this character
provided phylogenetic resolution towards the tips of the tree.
The Adesmia clade is uniform for lomented pods, but the Dalbergia and Pterocarpus clades are variable, with a minimum
of three separate origins of this pod type in each of these
clades. What was thought to be two separate origins of lomented pods in Adesmieae and Aeschynomeneae is now considered at least six origins combined with at least two reversals, and not counting polymorphisms.
New taxonomic evidence—In contrast to the above, a few
previously overlooked characters are shown by analysis of
combined molecular and nonmolecular data to be taxonomically informative. Short shoots (character 2 in Appendix B)
evolved only once in the clade containing Pictetia, Ormocarpum, and Ormocarpopsis (also Peltiera). However, the support
for this clade is moderate (Fig. 5), both in this analysis, and
in those of Beyra-M. and Lavin (1999) and Lavin et al. (2000).
Bilabiate calyx lobes (state 2 of character 19 in Appendix B)
mark the monophyly of the clade containing Aeschynomene
sect. Aeschynomene, Smithia, Kotschya, Humularia, Cyclocarpa, Soemmeringia, Bryaspis, and Geissaspis. In contrast to
short shoots, this calyx morphology marks a very well-supported clade (Fig. 5). The other nonmolecular characters with
a high retention index (Table 1), however, either mark small
clades (e.g., characters 13 and 46 and the clade with Brya and
Cranocarpus), or have homoplasy that was underestimated because of scoring polymorphic taxa (e.g., see characters 39 and
40 in Appendix B).
The aeschynomenoid root nodule (Fig. 6, character 55 in
Appendix B) is the most notable nonmolecular character in
that it is inferred to be a synapomorphy for the dalbergioid
clade. The idea that nodule morphology could be a useful
character in legume taxonomy was pioneered by Corby
(1981). He described a number of shapes, named according to
the genus from which he had most observations. The aeschynomenoid type has as its main feature a small oblate nodule
(transverse diameter greater than axial) with determinate
growth. Corby noted that aeschynomenoid nodules are often
associated with fine rootlets, but his otherwise excellent drawings omitted these ‘‘for clarity.’’ Such nodules were found pri-
514
AMERICAN JOURNAL
marily in the tribes Adesmieae, and Aeschynomeneae, but also
in some members of the Abreae, Dalbergieae, Phaseoleae, and
Robinieae (Corby, 1988). On his retirement, Corby kindly
gave the Sprent laboratory his collection of preserved nodules.
These were used, together with new material, for more detailed
structural studies. As a result, the definition of an aeschynomenoid nodule has been adapted to include additional features.
In particular, this nodule is always associated with a lateral or
(in the case of stem nodules) adventitious root. The central
infected tissue contains few or no uninfected cells. Differentiated infection threads are not involved in the process of infection, which (where studied in detail) takes place at the lateral root junction (Sprent, Sutherland, and Faria, 1989). All
nodules of the tribe Aeschynomeneae that have been examined
conform to this description, together with ten genera of the
Dalbergieae: Centrolobium, Dalbergia, Etaballia, Geoffroea,
Machaerium, Platymiscium, Platypodium, Riedeliella, Tipuana, and Pterocarpus (two Brazilian species, P. rohrii and P.
santalinoides are not known to nodulate). The evidence for
Adesmia, Brya, and Cranocarpus, although slightly less detailed, is entirely consistent with the revised description of
aeschynomenoid nodules.
Members of the Dalbergieae that have been omitted from
the revised clade on morphological and molecular grounds
would also be omitted on grounds of nodule structure (Andira
and Hymenolobium) or absence of nodules (Vatairea and Vataireopsis; Sprent, Sutherland, and Faria, 1989). Two genera
of the dalbergioid clade that do not nodulate are Chaetocalyx
and Nissolia (Faria and Lima, 1998). Both of these are lianes.
Notably, a group of species in Acacia with a semiscandent
habitat cannot nodulate (Harrier et al., 1997). These acacias
have retained some of the characters associated with nodulation, such as some of the nod genes, and the ability to stimulate rhizobial attachment to roots. It was thus suggested that
they may have lost the ability to nodulate because, living on
the forest margins, they were not nitrogen limited (Harrier,
1995). It would be interesting to carry out similar tests on
Chaetocalyx and Nissolia as one of their principal habitats is
forest margins.
It is now generally agreed that nodulation in legumes may
have evolved more than once (Sprent, 1994; Soltis et al.,
1995). One of these nodulation events involved an infection
process through a wound, such as where a lateral or an adventitious root emerges. Compared with the more familiar root
hair infection pathway (see Sprent and Sprent, 1990 for details), this pathway is simpler, involving less complex recognition systems. Apart from some species of the mimosoid genus Neptunia (James et al., 1992), this wound infection pathway is associated with only aeschynomenoid nodules. In Neptunia, however, nodule processes subsequent to infection
involve production of infection threads and development of an
indeterminate nodule.
Our phylogenetic results are in agreement with molecular
and biochemical evidence that nodule structure and infection
site are largely plant determined (e.g., Gualtieri and Bisseling,
2000). Given a phylogenetic lineage, nodule morphology and
infection processes are generally the same regardless of which
species or genus of rhizobia is involved (six genera of bacteria
nodulating legumes are now recognized, and they are collectively known as rhizobia). Another general inference is derived
from the observation that all species of the genus Aeschynomene that have stem nodules are nodulated by photosynthetic
rhizobia (Molouba et al., 1999). Given that the aeschynome-
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noid root nodule has an unelaborated morphology and infection mode, the ancestral rhizobial form could have been photosynthetic. As legumes moved into drier areas, nodules developed on roots and lost photosynthetic ability (Sprent, 1994).
A phylogenetic classification—The dalbergioid legumes are
similar to a group of Papilionoideae that includes also Amorpheae and Dipterygieae. They share a distinctive combination
of a base chromosome number of x 5 10 (Goldblatt, 1981),
wood with uniseriate stored rays, vegetative growth with glandular punctae, flowers with fused keel petals or staminal filaments, and seeds that do not accumulate nonprotein amino
acids (derived from Beyra-M. and Lavin, 1999). The dalbergioids differ and are apomorphically defined (sensu de Queiroz
and Gauthier, 1994) as having glandular-based trichomes on
vegetative or floral organs, a well-developed abaxial calyx
lobe, and the ‘‘aeschynomenoid’’ root nodule. All of these
traits have been secondarily transformed in some constituents
of the dalbergioid clade (see characters 11, 19, and 55 in Appendix B; also Table 1).
The dalbergioid clade is distinguished more by molecular
than nonmolecular data. It is another legume example of a
cryptic clade, like ‘‘Neo-Astragalus’’ (Wojciechowski et al.,
1993) and the ‘‘temperate herbaceous clade’’ (Sanderson and
Wojciechowski, 1996). Regardless, it is informally recognized
here as a distinctive taxonomic group. Furthermore, the three
major constituent subclades are informally recognized and Appendix C enumerates the 44 current dalbergioid genera accordingly. The three subclades of dalbergioids are:
The Adesmia clade—This includes the genera Adesmia (of
tribe Adesmieae; Polhill, 1981f) and Poiretia, Amicia, Zornia,
Chaetocalyx, and Nissolia of the tribe Aeschynomeneae. This
clade is apomorphically defined as having an herbaceous
growth habit (modified in some descendants—character 1),
leaves with few opposite leaflets (evolved in parallel in Arachis and close relatives—character 8), and pedicels confluent
with the calyx (modified only in a few species of Nissolia—
character 17). A node-based definition (sensu de Queiroz and
Gauthier, 1994) includes all descendants from the common
ancestor of Adesmia and Amicia.
The Dalbergia clade—This includes Dalbergia and Machaerium (of tribe Dalbergieae; de Candolle, 1825; Polhill,
1981d), and the following genera of Aeschynomeneae (sensu
Rudd, 1981a): Aeschynomene (all infrageneric taxa), Soemmeringia, Cyclocarpa, Kotschya, Smithia, Humularia, Bryaspis, Geissaspis, Weberbauerella, Diphysa, Pictetia, Ormocarpum, Ormocarpopsis, and Peltiera. This clade is apomorphically defined as having diadelphous staminal filaments splitting readily or tardily into two flanges, usually in a 5 1 5
arrangement (polymorphic with a 9 1 1 diadelphous condition
in many species and occasionally monodelphous in Machaerium—character 26), and a persistent staminal flange that in
some cases reflexes upward above the developing fruit (character 28). A node-based definition includes all descendants
from the common ancestor of Dalbergia and Cyclocarpa.
The Pterocarpus clade—This includes Pterocarpus, Tipuana, Platypodium, Reideliella, Centrolobium, Grazielodendron, Paramachaerium, Ramorinoa, Inocarpus, Etaballia,
Platymiscium, Cascaronia, Fissicalyx, Geoffroea from Dalbergieae; Brya and Cranocarpus from Desmodieae; and Fie-
March 2001]
LAVIN
ET AL.—DALBERGIOID LEGUMES
brigiella, Chapmannia, Stylosanthes, Arachis, and Discolobium from Aeschynomeneae. This clade is apomorphically defined as having commonly caducous bracteoles (character 18)
and seedlings producing a simplified eophyll (secondarily
transformed in Arachis and close relatives—character 45). A
node-based definition includes all descendants from the common ancestor of Pterocarpus and Riedeliella.
Although data from matK/trnK, trnL, and ITS/5.8S were not
combined in a single analysis, results from individual analyses
showed significant consensus combined with no significant
conflict. The combined matK/trnK and nonmolecular analysis
yielded very robust results to support the conclusions outlined
above. This study demonstrates that matK/trnK sequences provide excellent resolution at the broadest phylogenetic levels
dealt with in this study. This same locus, along with ITS/5.8S,
gives excellent resolution to within and among closely related
genera. In contrast, trnL provides the least resolution. Ultimately, this study provides a framework for future studies that
deal taxonomically with individual dalbergioid genera. There
is now sufficient data from which to guide the choice of potential sister groups or outgroups in such studies.
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GenBank accession nos.a
Voucherb
Species
Locality
Acosmium panamense (Benth.) Yakovlev
Adesmia boronoides Hook. f.
Adesmia corymbosa Clos
Adesmia grandiflora Gillies ex Hook. &
Arn.
Adesmia lanata Hook. f.
Adesmia lanata 1
Adesmia lotoides Hook. f.
Adesmia muricata (Jacq.) DC.
Adesmia pinnifolia Hook. & Arn.
Adesmia retrofracta Hook. & Arn.
Adesmia schneideri Phil.
Adesmia villosa Hook. f.
Adesmia volckmannii Phil.
Adesmia volckmannii 1
Aeschynomene americana L.
México. Oaxaca. Temascal
Argentina. Neuquén. Las Lajas
Argentina. Chubut. Paso de Indios
Argentina. Mendoza. Lujan de Cuyo
Hughes 1308 (FHO)
Lavin 8252 (MONT)
Lavin 8283 (MONT)
Lavin 8232b (MONT)
Argentina. Neuquén. Las Lajas
Argentina. Chubut. Tecka
Argentina. Chubut. Paso de Indios
Argentina, Mendoza, Luján de Cuyo
Argentina. Mendoza. Penitentes
Argentina. Neuquén. Chos Malal
Argentina. Chubut. Paso de Indios
Argentina. Chubut. Tecka
Argentina. Mendoza. Las Leñas
Argentina. Neuquén. Catan Lil
Cuba. Camagüey
Lavin 8256 (MONT)
Lavin 8276 (MONT)
Lavin 8279 (MONT)
Lavin 8233 (MONT)
Lavin 8235 (MONT)
Lavin 8249 (MONT)
Lavin 8281 (MONT)
Lavin 8274 (MONT)
Lavin 8245 (MONT)
Lavin 8258 (MONT)
Beyra-M. 554 (MONT)
Venezuela: Mérida
Lavin 5730 (MONT)
GBAN-AF189025
U.S.A. Louisiana. Allen
Thomas & Allen NLU3 (LSU)
GBAN-U59892
U.S.A. North Carolina. Hyde Co.
Zimbabwe. National Botanic Gardens
México. Oaxaca
U.S.A. Florida. Lake Co.
U.S.A., Louisiana. Beaufort
Carulli 58 (LSU)
Lavin s.n. (MONT)
Lavin 5833 (MONT)
Fairbrothers et al. 82 (LSU)
Carulli 42 (LSU)
GBAN-AF068141
GBAN-AF189026
Ecuador. Loja
Argentina. Tucumán. Amaicha
Mexico. Puebla. Nauzontla
U.S.A. Wyoming
México. Veracruz. Lago Catemaco
Guatemala. El Progreso. El Raucho
Bolivia. Santa Cruz. Rio Parapetı́
R. T. Pennington 654 (E)
Lavin 5773 (MONT)
Delgado s.n. (MEXU)
Lavin 6221 (BH)
Lavin 8214 (MEXU)
Hughes 254 (FHO)
CIAT 22250 (MONT)
GBAN-AF183502
GBAN-AF183501
GBAN-AF269174
GBAN-U59890
GBAN-U59889
GBAN-AF187093
GBAN-AF203553
Argentina. Corrientes. Utuzaingo
Brazil. Mato Grosso do Sul. Coxim
CIAT 22249 (MONT)
CIAT 22227 (MONT)
GBAN-AF203554
GBAN-AF203556
trnL intron
GBAN-AF208891
GBAN-AF208900
GBAN-AF270863
GBAN-AF208901
GBAN-AF183494
GBAN-AF183495
GBAN-AF187097
GBAN-AF187098
GBAN-AF183493
GBAN-AF183497
GBAN-AF183498
GBAN-AF183496
GBAN-AF183499
GBAN-AF142690
GBAN-U59893
GBAN-AF272085
GBAN-AF272086
GBAN-AF272087
GBAN-AF142695
GBAN-AF208929
GBAN-AF272083
GBAN-AF272084
GBAN-AF208927
GBAN-AF203587
GBAN-AF270870
GBAN-AF270869
GBAN-AF203586
GBAN-AF208930
GBAN-AF208928
GBAN-AF203583
GBAN-AF208902
GBAN-AF270861
GBAN-AF142681
GBAN-AF270860
GBAN-AF208899
GBAN-AF208893
GBAN-AF208898
GBAN-AF156675
GBAN-AF203555
GBAN-AF203552
GBAN-AF208948
GBAN-AF203597
GBAN-AF208947
GBAN-AF203596
GBAN-AF203599
GBAN-AF203607
GBAN-AF208946
GBAN-AF270876
GBAN-AF208950
GBAN-AF203557
GBAN-AF203551
GBAN-AF204233
GBAN-AF203558
GBAN-AF068142
GBAN-AF203560
519
A. Bhagwat, T. G. Krishna, and R. K. unknown
Mitra (unpublished)
Arachis magna Krapov., W. C. Greg. & Bolivia. Santa Cruz. San Ignacio
CIAT 22248 (MONT)
Simpson
Arachis major Krapov. & W. C. Greg.
Brazil. Mato Grosso do Sul. AquiCIAT 22225 (MONT)
dauana
Arachis palustris Krapov., W. C. Greg. Brazil. Sâo Paulo. Miracema
CIAT 22245 (MONT)
& Valls
Arachis pintoi Krapov. & W. C. Greg.
Brazil. Minas Gerais. Unai
CIAT 22237 (MONT)
Arachis triseminata Krapov. & W. C.
Brazil
CIAT 22224 (MONT)
Greg
Arachis villosa Benth.
Uruguay
CIAT 22254 (MONT)
Brya ebenus (L.) DC.
Cuba. Camagüey. Sabanas de Cromo Beyra-M. 523 (MONT)
Brya hirsuta Borhidi
Cuba. Sierra Nipe
Lavin 7110 (MONT)
matK/trnK
GBAN-AF142684
ET AL.—DALBERGIOID LEGUMES
Aeschynomene indica 1
Aeschynomene pfundii Taub.
Aeschynomene purpusii Brandegee
Aeschynomene rudis Benth.
Aeschynomene virginica Britton, Stern
& Poggenb.
Amicia glandulosa H. B. & K.
Amicia medicaginea Griseb.
Amicia zygomeris DC.
Amorpha fruticosa L.
Andira galeottiana Standl.
Apoplanesia paniculata Presl.
Arachis batizocoi Krapov. & W. C.
Greg.
Arachis correntina (Burkart) Krapov.
Arachis hermannii Krapov. & W. C.
Greg
Arachis hypogaea L.
GBAN-AF187084
LAVIN
Aeschynomene fascicularis Cham. &
Schlect.
Aeschynomene indica L.
ITS/5.8S
March 2001]
APPENDIX A.
520
APPENDIX A.
Continued.
GenBank accession nos.a
Species
Voucherb
Locality
Sierra Leone. Waterloo
Argentina
Brazil. Rio de Janeiro. Botanic Gardens
blanchetiana (Benth.) Rudd Brazil. Bahia, Rio Lajendo
brasiliensis (Vogel) Benth. México. Veracruz
brasiliensis 1
Argentina. Misiones. Rowing Club
glaziovii Taub.
Brazil. Bahia
klugii Rudd
Brazil. Acre
latisiliqua (Poir.) Benth. ex Costa Rica. Puntarenas. Corcovado
ITS/5.8S
Dawe 424 (K)
Klitgaard 100 (K)
H. C. Lima s.n. (E)
GBAN-AF204234
GBAN-AF204235
Chaetocalyx
Chaetocalyx
Chaetocalyx
Chaetocalyx
Chaetocalyx
Chaetocalyx
Hemsl.
Chaetocalyx nigricans Burkart
Chaetocalyx scandens (L.) Urb.
Chapmannia floridana Torr. & Gray
Chapmannia floridana 1
Hatschbach 56922 (F)
Ventura 14977 (MO)
Prado s.n. (MONT)
Coradin et al. 5741 (NY)
Daly 6778 (NY)
Kernan 122 (MO)
GBAN-AF183505
GBAN-AF183507
GBAN-AF183506
GBAN-AF203566
Vanni 2955 (F)
CIAT 20536 (MONT)
Gunter 121 (FLAS)
Judd 3162 (FLAS)
GBAN-AF183508
GBAN-AF183509
GBAN-AF203543
GBAN-AF203562
Miller & Alexander 14039 (E)
Miller & Alexander 14016 (E)
Nuñez 11153 (MO)
GBAN-AF203545
GBAN-AF203546
GBAN-AF203542
Thulin & Gifri 8829 (UPS)
GBAN-AF204232
Thulin & Gifri 8807 (UPS)
Miller & Alexander 14241 (E)
Kuchar 15450 (UPS)
GBAN-AF203548
GBAN-AF203544
Saynes V. 1286 (MEXU)
Torres C. 997 (MEXU)
GBAN-AF068168
GBAN-AF068169
Diphysa racemosa Rose
Diphysa racemosa 1
Sousa S. 7070 (MO)
Tenorio 4950 (MO)
GBAN-AF068163
GBAN-AF189030
GBAN-AF068140
GBAN-AF189022
GBAN-AF208905
GBAN-AF203585
GBAN-AF270865
GBAN-AF203600
GBAN-AF203605
GBAN-AF203592
GBAN-AF203593
GBAN-AF203598
GBAN-AF203606
GBAN-AF203591
GBAN-AF270875
GBAN-AF272067
GBAN-AF142696
GBAN-AF203581
GBAN-AF208942
GBAN-AF208940
GBAN-AF208941
GBAN-AF260644
GBAN-AF208943
GBAN-AF208951
GBAN-AF208924
GBAN-AF208921
Harris 1924
GBAN-AF208919
GBAN-AF189023
GBAN-AF189024
GBAN-AF203582
GBAN-AF203580
GBAN-AF203580
GBAN-AF068160
GBAN-AF068166
GBAN-U59891
GBAN-AF068161
GBAN-AF068162
GBAN-AF189029
GBAN-AF068167
GBAN-AF203574
GBAN-AF208920
GBAN-AF208923
GBAN-AF208922
GBAN-AF203575
GBAN-AF203601
GBAN-AF203608
GBAN-AF208912
GBAN-AF208913
[Vol. 88
México. Oaxaca. El Puente
México. Puebla. El Coro
GBAN-AF203547
GBAN-AF189060
GBAN-AF272070
GBAN-AF270866
BOTANY
Kirkpatrick 1759 (E)
Ghafoor & Goodman 4435 (E)
Polhill 5308 (K)
Hughes 1237 (FHO)
R. T. Pennington 668 (E)
Lavin 5801 (MONT)
Macqueen 309 (MONT)
Lavin 5801a (MONT)
Sousa S. 10616 (MO)
Haber 1322 (MO)
Sousa 9115 (MO)
Lavin 5823 (MEXU)
Thulin & Gifri 8829 (UPS)
Kallunki et al. 540 (MO)
Maxwell 89-1487 (E)
Hughes 1253 (FHO)
R. T. Pennington 859 (E)
Polhill 5309 (K)
trnL intron
GBAN-AF208932
GBAN-AF208958
GBAN-AF208966
OF
Argentina. Misiones. Iguazú
Brazil. Roraima. Pacaraima
U.S.A. Florida. Lake Co.
U.S.A. Florida: Highlands Co. Lake
Wales Ridge
Chapmannia gracilis (Balf. f.) Thulin
Yemen. Socotra
Chapmannia gracilis 1
Yemen. Socotra
Chapmannia prismatica (Sessé & Moci- México. Michoacán. Palos Marı́as
ño) Thulin
Chapmannia reghidensis Thulin & Mc- Yemen. Socotra
Kean
Chapmannia reghidensis
Yemen. Socotra
Chapmannia sericea Thulin & McKean Yemen. Socotra
Chapmannia somalensis (Hillcoat &
Somalia
Gillett) Thulin
Chapmannia tinireana Thulin
Yemen. Socotra
Cranocarpus martii Benth.
Brazil. Bahia. Itacaré
Cyclocarpa stellaris Baker
Thailand
Dalbergia congestiflora Pittier
El Salvador. Santa Ana. Metapan
Dalbergia foliolosa Benth.
Bolivia. Yungas
Dalbergia melanoxylon Guillemin &
Africa
Perrottet
Dalbergia pachycarpa (De Wild. & T.
Africa
Durand) De Wild.
Dalbergia sissoo Roxb.
India. Himachal Pradesh
Dalbergia sissoo 1
Pakistan. Baluchistan
Dalbergia vaccinifolia Vatke
Africa
Dalbergia sp.
El Salvador. Sonsonate
Dalbergia sp. 1
Ecuador. Zamora
Diphysa americana (Miller) M. Sousa
México. Puebla. Santiago Nopala
Diphysa americana 1
México. Chiapas. San Fernando
Diphysa americana 2
México. Puebla. Santiago Nopala
Diphysa floribunda Benth. & Oerst.
México. Oaxaca. Putla
Diphysa humilis Oerst.
Costa Rica. Puntarenas. Santa Elena
Diphysa macrophylla Lundell
México. Oaxaca. Salina Cruz
Diphysa ormocarpoides (Rudd) M. Sou- México. Oaxaca. San José
sa & R. Antonio
Diphysa ormocarpoides 1
México. Oaxaca. San Pedro Totalapan
Diphysa ormocarpoides 2
México. Oaxaca. Tehuantepec
matK/trnK
GBAN-AF272068
GBAN-AF272072
GBAN-AF270883
AMERICAN JOURNAL
Bryaspis lupulina (Benth.) Duvign.
Cascaronia astragalina Griseb.
Centrolobium sp.
Continued.
GenBank accession nos.a
Species
Diphysa racemosa 2
Diphysa sennoides Benth.
Diphysa spinosa Rydberg
Diphysa spinosa 1
Diphysa suberosa S. Watson
gentryi Rudd
hirsuta DC.
leiogyne Sandwith
shottii A. Gray
trnL intron
GBAN-AF068164
GBAN-AF189031
Cabrera 3024 (MO)
GBAN-AF189032
Sousa 6264 (MO)
GBAN-AF189033
Lavin 5814 (MO)
Kearns et al. s.n. (MONT)
GBAN-AF189034
GBAN-AF068165
Brazil. Goiás
Costa Rica. Heredia, Sarapiqui
R. T. Pennington 494 (E)
R. T. Pennington 613 (E)
GBAN-AF187090
GBAN-AF183492
GBAN-AF272092
GBAN-AF208896
Argentina. Formosa. Formosa
Bolivia. Santa Cruz. Chiquitos
Guyana. Kanuku Mts.
Cristobal & Krapovickas 2167 (MO) GBAN-AF189058
Frey et al. 531 (MO)
GBAN-AF189059
Janson-Jacobs 107 (K)
GBAN-AF270874
GBAN-AF270873
GBAN-AF272073
GBAN-AF272074
GBAN-AF208964
GBAN-AF208963
GBAN-AF208960
México. Zacatecas. Jalapa
Bolivia. Tarija
Ecuador. Loja
Bolivia. Santa Cruz. Villagrande
Venezuela. Bolivar
Malawi
Lavin 5052 (MONT)
Ehrich 65 (NY)
Lewis et al. 3823 (K)
Nee 36224a (NY)
Wurdack 34436 (K)
Hilliard & Burtt 4305 (E)
GBAN-AF187096
GBAN-AF203541
GBAN-AF203561
U.S.A. Arizona (seed source)
Lavin 750 (MONT)
GBAN-AF189057
Chile.
Ecuador. Loja
Brazil. Rio de Janeiro. Botanic Garden
Congo. Katanga
Gardner & Knees 5823 (E)
R. T. Pennington 659 (E)
Lima s.n. (E)
GBAN-AF270879
GBAN-AF270862
GBAN-AF208952
Symoens 14132 (K)
GBAN-AF272069
GBAN-AF208936
Costa Rica. La Selva Biological Station
U.K. Royal Botanica Gardens, Kew
Malawi
R. T. Pennington 614 (E)
R. T. Pennington s.n. (E)
Salubeni 3060 (E)
French Guinea
Armour 8400 (E)
Brazil. Goiás
Brazil. Roraima
Colombia. Tolima. Ibagué
México. Oaxaca. Chazumba
Nicaragua. Boaco. Ojo de Agua
U.S.A. Texas. Austin. cultivated
R. T. Pennington 487 (E)
Lewis 1598 (E)
R. T. Pennington 703 (E)
Lavin 5341 (MONT)
Hughes 424 (FHO)
Turner s.n. (MONT)
Kew Living Collection Accession
1990-901
Mexico. Sonora. Guayabo
Mexico. Mexico. Ixtapan
Mexico. Jalisco. Tapalapa
Mexico. Sonora
no voucher
VanDevender 93-189 (NY)
Roe 1904 (F)
Magallanes 2902 (F)
Joyal 2094 (NY)
GBAN-AF208911
GBAN-AF189061
GBAN-AF260645
GBAN-AF203576
GBAN-AF203589
GBAN-AF203590
GBAN-AF272063
GBAN-AF272064
GBAN-AF208939
GBAN-AF208938
GBAN-AF208931
GBAN-AF270880
GBAN-AF208962
GBAN-AF187087
GBAN-AF272079
GBAN-AF272080
GBAN-AF270878
GBAN-AF272065
GBAN-AF208965
GBAN-AF208934
GBAN-AF208935
GBAN-AF208926
GBAN-AF203564
GBAN-AF187095
GBAN-AF187085
GBAN-AF187086
GBAN-AF142692
GBAN-AF208925
GBAN-AF142679
GBAN-AF208892
GBAN-AF265559
GBAN-AF203563
GBAN-AF183510
GBAN-AF270868
GBAN-AF208906
GBAN-AF208908
GBAN-AF270867
GBAN-AF208907
521
Nissolia
Nissolia
Nissolia
Nissolia
matK/trnK
McVaugh 23043 (MO)
Garcı́a M. 484 (MO)
Nelson 7754 (MO)
ET AL.—DALBERGIOID LEGUMES
Eysenhardtia sp.
Fiebrigiella gracilis Harms
Fiebrigiella gracilis 1
Fiebrigiella gracilis 2
Fissicalyx fendleri Benth.
Geissaspis descampsii De Wild. & T.
Durand
Geoffroea decorticans (Gillies ex Hook.
& Arn.) Burkart
Geoffroea decorticans 1
Geoffroea spinosa Jacq.
Grazielodendron rio-docensis H. C.
Lima
Humularia corbisieri (De Wild.) Duvign.
Hymenolobium mesoamericana H. C.
Lima
Inocarpus fagifer (Parkinson) Fosberg
Kotschya aeschynomenoides (Baker) J.
Dewit & Duvign.
Kotschya ochreata (Taub.) J. Dewit &
Duvign.
Machaerium acutifolium Vogel
Machaerium inundatum Ducke
Machaerium sp.
Marina sp.
Myrospermum frutescens Jacq.
Myrospermum sousanum A. Delgado &
M.C. Johnst.
Myrospermum sousanum 1
México. Jalisco. La Huerta
México. Oaxaca. Teposcolula
Honduras. Francisco Morazán. Tegucigalpa
México. Chiapas. Amatenango de
Valle
México. Oaxaca. Santa Cruz Mixtepec
México. Oaxaca. Juchetango
México. Sonora. Alamos
ITS/5.8S
LAVIN
Diphysa suberosa 1
Diphysa thurberi (A. Gray) Rydberg ex
Standl.
Dipteryx alata Vogel
Dipteryx panamensis (Pittier) Record &
Mell.
Discolobium psoraleifolium Benth.
Discolobium pulchellum Benth.
Etaballia guianensis Benth.
Voucherb
Locality
March 2001]
APPENDIX A.
522
APPENDIX A.
Continued.
GenBank accession nos.a
Species
Ormocarpopsis aspera R. Vig.
Ormocarpopsis calcicola R. Vig.
Locality
ITS/5.8S
matK/trnK
GBAN-AF068148
GBAN-AF068145
GBAN-AF203568
DuPuy 2363 (K)
GBAN-AF068149
GBAN-AF203567
Phillipson 2924 (K)
GBAN-AF068147
Phillipson 3508 (K)
GBAN-AF068143
Du Puy et al. M132 (K)
Keraudren 1369 (K)
GBAN-AF068144
GBAN-AF068146
Labat et al. 2882 (P)
GBAN-AF189035
GBAN-AF203570
Du Puy et al. M716 (P)
Thulin & Gifri 8781 (UPS)
Thulin et al. 9746 (UPS)
GBAN-AF189036
GBAN-AF189037
GBAN-AF189040
GBAN-AF203572
Capuron 24625-SF (P)
Rakotozafy 986 (P)
Greenway 14054 (MO)
Wieland 4681 (UPS)
Wieland 4357 (MO)
Faden 74/958 (MO)
Chapman 8492 (MO)
Balsinhas 2842 (MO)
Torre 9458 (MO)
J. D. Manning 747 (MO)
Amshoff 1299 (MO)
Gilbert 1309 (MO)
DeWilde 5498 (MO)
Roach s.n. (QLD)
GBAN-AF189038
GBAN-AF189039
GBAN-AF189041
GBAN-AF189042
GBAN-AF189043
GBAN-AF068155
GBAN-AF068150
GBAN-AF068151
GBAN-AF068152
GBAN-AF189044
GBAN-AF068154
GBAN-AF068156
GBAN-AF068157
GBAN-AF068159
Aubreville s.n. (P)
Thulin et al. 6891 (UPS)
Schlieben 5766 (P)
CFRP 1043 (MO)
Thulin & Warfa 5818 (UPS)
Amshoff 9887 (MO)
GBAN-AF189045
GBAN-AF189046
GBAN-AF189047
GBAN-AF068153
GBAN-AF189048
GBAN-AF189049
Stephens 820 (MO)
GBAN-AF068158
Jungner s.n. (UPS)
Thulin et al. 9267 (UPS)
Janson-Jacobs 97 (K)
GBAN-AF189050
GBAN-AF189051
GBAN-AF204237
Axelrod 4788 (NY)
Axelrod 2877 (NY)
Acevedo et al. 2046 (NY)
Atha & Zanoni 730 (NY)
Beyra-M. s.n. (MONT)
Lavin 7108 (MONT)
Beyra-M. s.n. (MONT)
GBAN-AF068175
GBAN-AF068174
GBAN-AF208918
GBAN-AF208914
GBAN-AF260646
GBAN-AF260647
GBAN-AF203602
GBAN-AF203571
GBAN-AF208917
OF
GBAN-AF203603
GBAN-AF203569
GBAN-AF208916
GBAN-AF260648
GBAN-AF260649
GBAN-AF208915
GBAN-AF203573
GBAN-AF272062
GBAN-AF203577
GBAN-AF203604
GBAN-AF203579
GBAN-AF203578
GBAN-AF260650
GBAN-AF208959
GBAN-AF260906
GBAN-AF208910
[Vol. 88
GBAN-AF068171
GBAN-AF068176
GBAN-AF068177
trnL intron
BOTANY
Peltier 4416 (MO)
Capuron 24240-SF (K)
AMERICAN JOURNAL
Madagascar. Mohobo
Madagascar. Ambongo. Antsakoamanera
Ormocarpopsis itremoensis Du Puy &
Madagascar. Fianarantsoa; AmbatofiLabat
nandrahana
Ormocarpopsis mandrarensis Dumaz-le- Madagascar. Toliara; Andohahela ReGrande
serve
Ormocarpopsis parvifolia Dumaz-leMadagascar. Toliara; Tsiombe
Grande
Ormocarpopsis parvifolia 1
Madagascar. Toliara; Beloha
Ormocarpopsis tulearensis Du Puy &
Madagascar. Toliara
Labat
Ormocarpum bernierianum (Baill.) Du Madagascar. Antsiranana
Puy & Labat
Ormocarpum bernierianum 1
Madagascar. Antsiranana
Ormocarpum coeruleum Balf. f.
Yemen. Socotra
Ormocarpum dhofarense Hillcoat & Gil- Yemen
lett
Ormocarpum drakei R. Vig.
Madagascar. Menabe. Antsalova
Ormocarpum drakei 1
Madagascar. Bekopaka
Ormocarpum flavum Gillett
Tanzania. Iringa Distr.
Ormocarpum gillettii Thulin
Somalia
Ormocarpum gillettii 1
Somalia. Hobyo Dist.
Ormocarpum keniense Gillett
Kenya. Meru
Ormocarpum kirkii S. Moore
Malawi. Southern Region
Ormocarpum kirkii 1
South Africa. Kloof Forest
Ormocarpum kirkii 2
Mozambique. Momba
Ormocarpum klainei Tisser.
Cameroon. Southwest. Lake Barombi
Ormocarpum megalophyllum Harms
Cameroon. Forest Preserve
Ormocarpum muricatum Chiov.
Kenya. Mandera
Ormocarpum muricatum 1
Ethiopia. Harrar
Ormocarpum orientale (Spreng.) Merr. Australia. Queensland, North Kennedy
Ormocarpum pubescens (Hochst.) Cufod Afrique occidentale
Ormocarpum rectangulare Thulin
Somalia
Ormocarpum schliebenii Harms
Tanzania. Lindi Distr.
Ormocarpum sennoides (Willd.) DC.
Tanzania. Zaraninje Forest
Ormocarpum somalense Gillett
Somalia
Ormocarpum trachycarpum (Taub.)
Ethiopia. Harar
Harms
Ormocarpum trichocarpum (Taub.) Burtt Natal. Gunamanini Pan
Davy
Ormocarpum verrucosum P. Beauv.
Cameroon
Ormocarpum yemenense Gillett
Yemen
Paramachaerium schomburgkii (Benth.) Guyana. Kanuku Mts.
Ducke
Pictetia aculeata (Vahl) Urb.
Puerto Rico. Cabo Rojo
Pictetia aculeata 1
Puerto Rico. Guanica
Pictetia aculeata 2
St. John. Coral Bay
Pictetia aculeata 3
Puerto Rico. Guanica
Pictetia angustifolia Griseb.
Cuba. Camagüey. Loma de la Coca
Pictetia marginata Sauv.
Cuba. Hoguı́n. Sierra Nipe
Pictetia marginata 1
Cuba. Las Villas-Camagüey
Voucherb
Continued.
GenBank accession nos.a
Species
Pictetia mucronata (Griseb.) Beyra-M.
& Lavin
Pictetia mucronata 1
Pictetia nipensis (Urb.) Beyra-M. &
Lavin
Pictetia obcordata DC.
Voucherb
Locality
ITS/5.8S
GBAN-AF068172
Cuba. Camagüey. Tagarro
Cuba. Sierra Nipe
Beyra-M. s.n. (MONT)
Ekman 10001 (NY)
GBAN-AF068173
GBAN-AF189052
Dominican Republic. Cambita Garabita
Cuba. Santiago de Cuba
Zanoni 40488 (NY)
GBAN-AF068170
Ekman 8403 (NY)
GBAN-AF203565
Dominican Republic. Baoruco
Garcı́a et al. 623 (NY)
GBAN-AF068178
Klitgaard 35 (K)
R. T. Pennington
R. T. Pennington
R. T. Pennington
R. T. Pennington
Lima s.n. (RB)
GBAN-AF189053
GBAN-AF189054
Poiretia angustifolia Vogel
Poiretia punctata (Willd.) Desv.
Brazil
Ecuador. Napo. Jatun Sacha
Colombia. Antioquia. Rio Claro
Colombia. Antioquia. Rio Claro
Brazil. Goiás. Chapada dos Veadeiros
Brazil. Rio de Janeiro. Botanic Garden
Brazil. Goiás. Niquelândia
Ecuador. Guayas
Pterocarpus acapulcensis Rose
Pterocarpus indicus Willd.
Pterocarpus indicus 1
Pterocarpus macrocarpus Kurz
Pterocarpus sp.
Pterodon pubescens Benth.
Ramorinoa girolae Speg.
Pictetia spinosa (A. Rich.) Beyra-M. &
Lavin
Pictetia sulcata (P. Beauv.) Beyra-M. &
Lavin
Platymiscium filipes Benth.
Platymiscium stipulare Benth.
Platymiscium sp.
Platypodium elegans Vogel
Platypodium elegans 1
Poecilanthe parviflora Benth.
649
692
688
488
(E)
(E)
(E)
(E)
matK/trnK
GBAN-AF270872
GBAN-AF270871
GBAN-AF270877
GBAN-AF187089
GBAN-AF142687
Fonseca et al. 1419 (MO)
Madsen 63491 (MO)
GBAN-AF183503
GBAN-AF183504
GBAN-AF270864
GBAN-AF272081
GBAN-AF272082
Mexico. Oaxaca. Santiago Astata
Philippines. Luzon. Mt. Izarog
unknown locality
Puerto Rico. Fajardo. cultivated
Colombia. Antioquia. Rio Claro
Brazil. Goiás
Lavin 5325 (MONT)
Pennington 718 (E)
Henderson s.n. (NY)
Lavin 721 (MONT)
R. T. Pennington 690 (E)
R. T. Pennington 476 (E)
GBAN-AF269175
GBAN-AF269177
GBAN-AF269176
GBAN-AF142691
GBAN-AF203588
GBAN-AF187091
no voucher
GBAN-AF204236
Riedeliella graciliflora Harms
U.K. Royal Botanic Gardens, Kew.
cultivated
Brazil. Mato Grosso do Sul
GBAN-AF272094
GBAN-AF272095
GBAN-AF270881
Smithia ciliata Royle
Soemmeringia semperflorens Mart.
Nepal
Brazil. Roraima, Ilha da Maracá
Stainton et al. 4048 (E)
Lewis 1600 (E)
Stylosanthes angustifolia Vogel
Stylosanthes calcicola Small
Stylosanthes capitata Vogel
Stylosanthes hamata (L.) Taub.
Stylosanthes humilis Kunth
Stylosanthes fruticosa (Retz.) Alston
Stylosanthes gracilis H. B. & K. 1
Stylosanthes gracilis 2
Stylosanthes grandiflora M. B. Ferreira
& S. Costa 1
Stylosanthes grandiflora 2
Stylosanthes guianensis (Aubl.) Swartz 1
Stylosanthes guianensis 2
Stylosanthes guianensis 3
Stylosanthes guianensis 4
Stylosanthes guianensis 5
Stylosanthes guianensis 6
Stylosanthes guianensis 7
(Vander Stappen et al., 1998)
(Vander Stappen et al., 1998)
Brazil. Mato Grosso. Rondonopolis
Cuba. Santiago de Cuba
(Vander Stappen et al., 1998)
(Vander Stappen et al., 1998)
(Vander Stappen et al., 1998)
(Vander Stappen et al., 1998)
(Vander Stappen et al., 1998)
Stappen
Stappen
Stappen
Stappen
Stappen
Stappen
Stappen
Stappen
et
et
et
et
et
et
et
et
al.,
al.,
al.,
al.,
al.,
al.,
al.,
al.,
1998)
1998)
1998)
1998)
1998)
1998)
1998)
1998)
GBAN-AF208961
GBAN-AF208897
GBAN-AF208904
GBAN-AF189027
GBAN-AF203549
GBAN-AF203550
GBAN-AF272090
GBAN-AF272091
GBAN-AF272066
GBAN-AF272088
GBAN-AF272089
GBAN-AF203595
GBAN-AF203594
GBAN-AF208954
GBAN-AF208895
GBAN-AF208957
GBAN-AF208949
GBAN-AF208933
GBAN-AF208937
GBAN-AJ230726
GBAN-AJ230728
GBAN-AF208945
GBAN-AF208944
GBAN-AJ230738
GBAN-AJ230731
GBAN-Y13483
GBAN-Y13488
GBAN-Y13486
CIAT 136
CIAT 1283
Schofield DNA
CPI 34906
CPI 18750
CIAT 1283
CIAT 184
GBAN-Y13487
GBAN-Y13480
GBAN-Y13481
GBAN-Y13482
GBAN-Y13485
GBAN-Y13489
GBAN-Y13490
GBAN-Y13491
523
(Vander
(Vander
(Vander
(Vander
(Vander
(Vander
(Vander
(Vander
GBAN-AF208955
GBAN-AF208953
Ratter et al. 7494 (E)
CIAT 1693 (MONT)
Beyra-M. 595 (MONT)
trnL intron
ET AL.—DALBERGIOID LEGUMES
Beyra-M. s.n. (MONT)
LAVIN
Cuba. Camagüey. La Mina
March 2001]
APPENDIX A.
524
APPENDIX A.
Continued.
GenBank accession nos.a
Species
Stylosanthes
Stylosanthes
Stylosanthes
Stylosanthes
Stylosanthes
guianensis 8
hamata (L.) Taub.
hippocampoides Mohlenbr.
humilis H. B. & K.
humilis 1
Locality
Voucherb
Lavin 5796 (MONT)
R. T. Pennington s.n. (E)
R. T. Pennington 495 (E)
R. T. Pennington 689 (E)
R. T. Pennington 587 (E)
GBAN-AF183491
GBAN-AF187088
Vataireopsis surinamensis H. C. Lima
Weberbauerella brongniartioides Ulbr.
(Vander Stappen et al., 1998)
Cuba. Santiago de Cuba
(Vander Stappen et al., 1998)
(Vander Stappen et al., 1998)
(Vander Stappen et al., unpublished
data)
(Vander Stappen et al., unpublished
data)
(Vander Stappen et al., unpublished
data)
(Vander Stappen et al., unpublished
data)
(Vander Stappen et al., unpublished
data)
(Vander Stappen et al., unpublished
data)
Argentina. Salta. La Cruz
Spain. Barcelona. Cultivated
Brazil. Goiás. Chapada dos Veadeiros
Colombia. Antioquia. Rio Claro
Costa Rica. Puntarenas, Penı́nsula de
Osa
Guyana. Iwokrama Reserve Area
Perú. Arequipa. Lomas de Mollendo
R. T. Pennington 385 (E)
Dillon 3909 (F)
GBAN-AF189028
Weberbauerella brongniartioides 1
Weberbauerella brongniartioides 2
Perú. Arequipa. Lomas de Mejı́a
Perú. Arequipa. Lomas de Mejı́a
Dillon 3742 (F)
Dillon 4818 (F)
Zornia sp.
Zornia sp. 1
México. Zacatecas. Fresnillo
(Vander Stappen et al., unpublished
data)
Lavin 5039 (MONT)
Stylosanthes humilis 2
Stylosanthes humilis 4
Stylosanthes humilis 5
Stylosanthes humilis 6
GBAN-AF203594
trnL intron
GBAN-AJ230733
GBAN-AF208944
GBAN-AJ230738
GBAN-AJ101325
GBAN-AJ101326
GBAN-AJ101327
GBAN-AJ101328
GBAN-AJ101329
GBAN-AJ010330
GBAN-AF189056
GBAN-AF270882
GBAN-AF208956
GBAN-AF265560
GBAN-AF142680
GBAN-AF272075
GBAN-AF272076
GBAN-AF272071
GBAN-AF272077
GBAN-AF272078
GBAN-AF203584
GBAN-AF208894
GBAN-AF208909
BOTANY
GBAN-AF183500
GBAN-AF270859
OF
Tipuana tipu (Benth.) Kuntze
Tipuana tipu 1
Vatairea macrocarpa (Benth.) Ducke
Vatairea sp.
Vatairea sp. nov.
GBAN-AF203550
GBAN-Y13484
matK/trnK
AMERICAN JOURNAL
Stylosanthes humilis 3
cultivar SG 220
Beyra M. 595 (MONT)
ITS/5.8S
GBAN-AF208903
GBAN-AJ230748
Note: Nexus files of nonmolecular and molecular data are located at http://gemini.oscs. montana.edu/;mlavin/data/dalbdat.htm.
a The prefix GBAN- has been added to all GenBank accession numbers to link the online version of American Journal of Botany to GenBank but is not part of the actual accession
number.
b CIAT accessions were grown from seed, and a herbarium specimen was prepared from the greenhouse plant and deposited at MONT.
[Vol. 88
March 2001]
LAVIN
ET AL.—DALBERGIOID LEGUMES
APPENDIX B
Nonmolecular characters and character states. All references to clades are
those derived from the combined matK/trnK and nonmolecular analysis (Fig.
5). References to ancestral states were inferred with the reconstruct tree option
in PAUP (Swofford, 2000) and the trace option in MacClade (Maddison and
Maddison, 1999).
Vegetative characters
1. Habit: 0) woody (trees to shrubs), 1) herbaceous (subshrubs to herbs),
2) twining and herbaceous, 3) twining and woody. Predominantly herbaceous
genera sometimes include subshrubby species, whereas woody genera usually
do not, thus explaining the coding for state number 1. A herbaceous habit
arose independently in the following clades: one represented by Fiebrigiella
and Arachis, another by Chaetocalyx and Poiretia, and one by Weberbauerella and Kotschya. The twining herbaceous habit is restricted to the Adesmia
clade where it is known from some species of Poiretia (Rudd, 1972c) and all
Chaetocalyx (Rudd, 1958) and Nissolia (Rudd, 1956). A twining woody habit
occurs in polymorphic condition in the clade with Dalbergia and Machaerium.
2. Short shoots: 0) absent or not regularly present and then not covered by
persistent stipules, 1) regularly present and covered by distichously arranged
persistent stipules from the axils of which are born the inflorescence (Fig. 7).
The short shoot condition is restricted to the clade including all descendants
of the most recent common ancestor of Pictetia and Ormocarpopsis. Very
similar short shoots were described for Poitea (tribe Robinieae; Lavin, 1993),
which is also from the Greater Antilles.
3. Stipule modifications: 0) attached to stem at base (basifixed) and foliaceous, 1) attached to stem in the middle and foliaceous (peltate or medifixed),
2) basifixed and lignescent. Medifixed stipules are referred to as appendiculate
(e.g., Rudd, 1981a) and are evolved independently in a clade including Aeschynomene sect. Aeschynomene, Cyclocarpa, Humularia, Geissaspis, Smithia,
and another including just Zornia. Lignescent stipules evolved independently
in polymorphic condition in the liana-forming species of Machaerium, in most
species of Brya, and in all species of Pictetia. In Brya, the leaves of the long
shoot are entirely transformed into a single spine.
4. Pseudopetiole: 0) absent, 1) present (Fig. 8). A pseudopetiole is traditionally defined as a petiole with stipules attached. It is here described as a
pulvinus (leaf base) that is projected away from the main axis of the stem.
The stipules are attached to this projected portion of the stem, and they superficially appear as if they are adnate to the petiole. The pseudopetiole
evolved independently in a clade including just Adesmia, and another including Arachis and Stylosanthes.
5. Leaf rachis in cross section: 0) terete, 1) with a single continuous groove
(canaliculate). A terete leaf rachis is recorded from Discolobium, Dalbergia,
Machaerium, and Ormocarpopsis, Peltiera, Platymiscium, Centrolobium,
Grazielodendron, Etaballia, Fissicalyx, Peltiera, and Pterocarpus, and in
polymorphic condition from Ormocarpum, Aeschynomene (all subgroups) and
closely related genera (Cyclocarpa, etc.). Grooved leaf rachises occur in the
rest of the genera, except where the leaves are uniformly sessile, as in Brya
and Inocarpus, and this trait is then scored as inapplicable. Otherwise, leaf
rachises vary continuously between narrowly grooved and distinctly canaliculate. The motivation for using this trait is that terete leaf rachises are shown
to be derived (but in polymorphic condition) in two clades: that including all
descendents but Pictetia of the most recent common ancestor of Dalbergia
and Ormocarpopsis, and that including most descendants of the recent common ancestor of Platymiscium and Pterocarpus.
6. Distal end of leaf rachis: 0) terminated by a leaflet, 1) not terminated by
a leaflet (a mucro is often present). A leaf rachis not terminated by a leaflet
is found in the large clade including Aeschynomene sect. Aeschynomene, Cyclocarpa, Humularia, Soemmeringia, Kotschya, Smithia, Geissaspis, and
Bryaspis. This type of leaf also has evolved independently in the outgroup
samples of Dipterygeae (Dipteryx and Pterodon), the clade including Amicia,
Zornia, Adesmia, Arachis, and Poiretia, the clade including just Aeschynomene sect. Ochopodium, and the clade including Stylosanthes and Arachis.
7. Number of leaflets per leaf: 0) leaves unifoliolate/simple, 1) leaves trito 20-foliolate, 2) leaves more than 20-foliolate. State zero occurs uniformly
in Etaballia, Inocarpus, and Brya, and in polymorphic condition in Cranocarpus. State two is restricted to just the Dalbergia clade where it occurs
uniformly in Weberbauerella, and predominantly so (i.e., polymorphic) in
Machaerium, Dalbergia, and all the sections and series of Aeschynomene
(Aeschynomene, Viscidulae, Pleuronerviae, and Scopariae). This state is capturing ‘‘fern-like’’ leaves where the leaflets abut laterally, are narrowly ellip-
525
tic, and have parallel lateral margins. Simple leaves are scattered throughout
but with most occurrences (usually in polymorphic condition) in the Pterocarpus clade (Discolobium, Etaballia, Inocarpus, Platypodium, Byra, and
Cranocarpus).
8. Leaflet arrangement: 0) alternate, 1) opposite. Two large clades have
evolved opposite leaflets independently. One includes Adesmia, Chaetocalyx,
Nissolia, Poiretia, Amicia, Zornia, and the other includes Fissicalyx, Fiebrigiella, Chapmannia, Stylosanthes, and Arachis. Opposite leaflets have evolved
sporadically mostly within the Pterocarpus clade (Grazielodendron, Riedeliella, Cranocarpus, Paramachaerium), and rarely in the Dalbergia clade
(Smithia). The genera with uniformly simple or unifoliolate leaves (e.g., Etaballia, Inocarpus, Brya, and Ramorinoa) were marked inapplicable. The species of Cranocarpus with imparipinnate leaves have opposite leaflets, and this
condition is used to represent the genus. A terminal taxon is scored for opposite leaflets if all constituent species predominate with this condition. A
terminal taxon is scored for a polymorphic condition only if some constituent
species have uniformly opposite leaflets and others have uniformly alternate
leaflets.
9. Leaflet base: 0) symmetric, 1) asymmetric. The asymmetric state is restricted to the Dalbergia clade, where it has evolved independently and polymorphically in Pictetia and Aeschynomene (all subgroups except Scoparia)
and uniformly in Humularia, Bryaspis, Geissaspis, Kotschya, and Smithia. An
asymmetric base of the leaflet is correlated with an eccentric midrib and probably related to a nyctinastic leaflet movement that involves a forward twisting
and folding of each leaflet. This ‘‘forward-folding’’ type is very similar to
the leaflet movements in legume subfamilies Mimosoideae and Caesalpinioideae, as well as the papilionoid genus Sesbania, and it has been observed in
species of Aeschynomene, Arachis, Diphysa, Dalbergia, and Machaerium.
10. Tannin deposits on the abaxial surface of dried leaflets: 0) absent, 1)
present. Tanniniferous patches on dried leaflets have evolved independently
in the clade including Arachis, Stylosanthes (polymorphic in these first two),
and Chapmannia (this is the subtribe Stylosanthinae of Rudd, 1981a) and in
two genera endemic to Madagascar, Ormocarpopsis and Peltiera (Labat and
Du Puy, 1996, 1997). Reddish tannin deposits usually occur in reticulate patterns demarcating individual epidermal cells. In Ormocarpopsis and Peltiera,
they can be concentrated along the leaflet midrib.
11. Glandular-based trichomes: 0) absent, 1) present (Figs. 9, 10). This type
of trichome is a synapomorphy for the dalbergioid group, where it is found
on the stems, leaves, inflorescence, or ovary. Although synapomorphic, the
glandular-based trichomes have been secondarily lost several times in each of
the Adesmia, Dalbergia, and Pterocarpus clades. In addition to most genera
of the formally recognized tribe Aeschynomeneae, glandular-based trichomes
are found in Centrolobium, Grazielodendron, Ramorinoa, Etaballia, Riedeliella, Fissicalyx, Paramachaerium, Peltiera, and polymorphic in Brya, Cranocarpus, Dalbergia, and Machaerium.
12. Pustular glands: 0) absent, 1) present (Fig. 11). The latter condition is
thought to be a derivation of the general dalbergioid trait of punctate glands
(all members of the ingroup and outgroup possess punctate glands on the
leaflets). There has been further development in the size and color of the
common punctate gland such that they protrude outward from the plane of
the leaflet, calyx, or ovary and are brownish red to blackish in color. Pustular
glands are known from genera outside the dalbergioid clade (Acosmium, Myrospermum, Amorpha, Apoplanesia, Dipteryx, and Pterodon), and have
evolved in four separate instances within the dalbergioid clade (Geoffroea and
Cascaronia; Poiretia, Amicia, and Zornia; Weberbauerella; and Centrolobium).
13. Stipitate glands: 0) absent, 1) present from non-glochidiate trichomes,
2) present from microscopically glochidiate trichomes (Fig. 12). Such glandular trichomes are usually present on stems or leaves, but can also occur on
ovaries and pods. Stipitate glands have evolved in the clade including Brya,
Cranocarpus, and Grazielodendron (uniquely from glochidiate trichomes in
the first two genera), and in polymorphic condition in the clade including
Adesmia, Chapmannia, and Stylosanthes. In Brya and Cranocarpus, stipitate
glands are found, in addition to the foliage, on the ovary where they persist
with the mature fruit. The high tree scores for this character (Table 1) do not
account for the optimizations of polymorphic codings where Chapmannia,
Stylosanthes, and Adesmia were assigned state zero during parsimony analysis.
Inflorescence characters
14. Inflorescence position: 0) axillary, 1) terminal. The first state corresponds to leafy flowering branches that are indeterminate with vegetative
growth from the apical meristem. The second refers to leafy flowering branch-
526
AMERICAN JOURNAL
OF
BOTANY
[Vol. 88
Figs. 9–12. Selected nonmolecular characters. 9. Glandular-based trichome of dalbergioid legumes (character number 11; scale bar 5 200 mm). 10. Base
of trichome where glandular exudate is secreted (scale bar 5 20 mm). 11. Pustular glands on leaflet of Centrolobium (character number 12; scale bar 5 200
mm). 12. Glochidiate trichomes on leaf of Brya (character number 13; scale bar 5 20 mm).
es whose growth is terminated by the inflorescence. The relatively high scores
(Table 1) reflect the uniform occurrence of state one in the clade including
Apoplanesia and Amorpha, and in the two species of Geoffroea. Other cases
of independent evolution but in polymorphic condition include Reideliella,
most outgroup genera, and sporadically throughout the Dalbergia clade (Aeschynomene subgroups, Kotschya, and Smithia).
15. Inflorescence type: 0) racemose, 1) axillary subumbel, 2) solitary axillary flowers, 3) helicoid cymes. Helicoid cymes have evolved several times
but in all cases within the Dalbergia clade (Dalbergia, Machaerium, Aeschynomene, Kotschya, and Smithia). They appear to arise readily from any inflorescence condition (i.e., note the polymorphic codings for most of these genera). In the axillary subumbel, the internodes of the rachis are telescoped
down almost completely, as in Chaetocalyx and Nissolia. Solitary flowers
have evolved independently and uniformly in Brya and the clade with Arachis
and Stylosanthes. Notably, polymorphic codings for this character are highly
localized to the genera in the clade that includes the most recent common
ancestor of Dalbergia and Ormocarpopsis.
16. Floral bracts: 0) smaller than the flower or fruit, 1) larger than the
flower or fruit. Large floral bracts have evolved independently in Zornia, and
the clade including Bryaspis, Geissaspis, and Humularia. These two states
are markedly discontinuous where the smaller bract is barely visible.
Floral characters
17. Pedicels: 0) articulated with the calyx, 1) confluent with calyx, 2) absent, flowers sessile. The Adesmia clade is marked by pedicels confluent with
the calyx, with the exception of a very few species of Nissolia. State zero is
most common among the rest of the dalbergioid clade and could be the ancestral condition to the Dalbergia and Pterocarpus clades. If so, then a transition to pedicels confluent with the calyx has occurred many times independently (Reideliella, Ramorinoa, Centrolobium, Brya, Cranocarpus, Weberbauerella, and Geissaspis, Bryaspis, and Humularia). Sessile flowers have
evolved three times, once in Chapmannia, Arachis, and Stylosanthes (the subtribe Stylosanthinae of Rudd, 1981a), and again in Etaballia and Inocarpus.
18. Bracteoles: 0) persistent, 1) caducous, 2) not or irregularly produced.
Bracteoles persisting paired at the end of the pedicel after abscission of the
flower or with the developing or mature fruit are common to the dalbergioids.
Caducous signifies that the bracteoles fall before the flower aborts or before
the pod begins to form. Caducous bracteoles are highly localized in the clade
that includes all descendants of the most recent common ancestor of Pterocarpus and Platymiscium. Such bracteoles have also evolved independently
in Geoffroea, Riedeliella, and several of the outgroups. Bracteoles occur irregularly (i.e., mostly singly and variously along the pedicel) or not at all in
four separate clades: one including Weberbauerella, another with Humularia,
Geissaspis, yet another with Amicia, Poiretia, Zornia, Chaetocalyx, and Nissolia, and finally in Cascaronia.
19. Calyx lobe fusion: 0) five more or less equally spaced lobes, 1) five
separate lobes but with the abaxial one (lower or carinal) the largest and
separate from laterals, 2) a two-lipped calyx with the abaxial lobe fused completely or nearly so to the two lateral lobes, and the upper two lobes com-
March 2001]
LAVIN
ET AL.—DALBERGIOID LEGUMES
527
Figs. 13–16. Petal characters (scale bar 5 5 mm for all figures). Figs. 13–15. Petals differentiated into blade and claw in Geoffroea (character number 23).
13. Standard. 14. Wing. 15. Keel. 16. Petals not differentiated into a blade and claw in Inocarpus.
pletely fused, 3) Dipteryx type, 4) Fissicalyx type, 5) Inocarpus type. Character state one is synapomorphic for the dalbergioid clade, and it is most
distinctive developmentally with the abaxial sepal initiating with the larger
size and faster growth rate relative to the other sepals (Klitgaard, 1999a). The
most notable derivation from this condition within the dalbergioids is state
two, which occurs in the clade with Aeschynomene sect. Aeschynomene, Cyclocarpa, Soemmeringia, Kotschya, Smithia, Geissaspis, Bryaspis, and Humularia (Aeschynomeneae subtribe Aeschynomeninae of Rudd, 1981a). The
Dipteryx type occurs in Dipteryx, Pterodon, and Amicia zygomeris, where the
upper two calyx lobes are greatly enlarged, contrasting with the diminutive
lower three lobes. The Fissicalyx type evolved only in Fissicalyx, where all
the calyx lobes occur as an upper lip (spathaceous). The Inocarpus type has
three lips, the lower formed by the carinal lobe, and two lateral lips formed
by one lateral and one vexillar lobe. Also, the Socotran species of Chapmannia have yet another type where the upper lip of a bilabiate calyx comprises
the two upper and two lateral calyx lobes, and the lower lip comprises just
the carinal lobe. Chapmannia, however, was scored for state one because it
represents the ancestral state for the genus (see the Chapmannia phylogeny
in Lavin et al., 2000). Similarly, the ancestral state for Pictetia is state one
(Beyra-M. and Lavin, 1999).
20. Hypanthium: 0) not well developed, petals and stamens arising at the
base of the ovary, 1) short-tubular, petals and stamens arising from a rim
positioned above the ovary base but not above the ovary itself, 2) long-tubular,
where petals and stamens arise from a rim located above the ovary. The
calyces of Acosmium, Apoplanesia, Amorpha, and Etaballia have a poorly
developed hypanthium (the last genus represents the only reversion to a loss
of the hypanthium among dalbergioid legumes). Among the dalbergioids, state
one predominates and is ancestral. State two is confined to the clade including
Chapmannia, Arachis, and Stylosanthes.
21. Petal coloration: 0) predominantly whitish to reddish or violet, 1) predominantly yellow. The large majority of dalbergioid legumes have yellow
petals, and this is inferred to be ancestral. Notable exceptions include the
clade with Dalbergia and Machaerium (polymorphic), as well as Grazielodendron, Paramachaerium, Ormocarpum (polymorphic), and Adesmia (the
species with solitary axillary flowers) where whitish to violet petals are common.
22. Corolla symmetry: 0) bilateral (papilionoid), 1) nearly radial. State zero
predominates among dalbergioids and indeed most papilionoids. Notably, a
nearly radially symmetric flower has evolved independently four times: once
each in Inocarpus, Etaballia, Reideliella, and the clade with the samples of
Amorpheae (Apoplanesia and Amorpha). Nearly radially symmetric flowers
have also evolved independently in the four species of Pictetia where state
zero is considered ancestral (Beyra-M. and Lavin, 1999).
23. Petal morphology: 0) petals abruptly differentiated into a blade and
claw (Figs. 13–15), 1) petals ligulate, the claw and blade not distinguishable
(Fig. 16). State one evolved separately in Etaballia and Inocarpus. This character is not dependent on character number 22 because Amorpheae, Reideliella, and four species of Pictetia have a nearly radial flower symmetry with
petals differentiated into a blade and claw.
24. Keel petals: 0) free, 1) connate, at least along the carinal margin if not
to near the tip. Free keel petals in papilionoid legumes have been the hallmark
of the tribes Swartzieae and Sophoreae. Acosmium and Myrospermum, traditionally placed in the tribe Sophoreae, have free keel petals. Among the
dalbergioid legumes, fused keel petals represent the ancestral condition that
has reverted back to the free condition only in Etaballia, Geoffroea, Riedeliella, Platypodium, and Tipuana (all confined to the Pterocarpus clade).
25. Wing petals: 0) smooth, 1) crimped. Crimped wing petals are distinctly
much broader than the adjacent keel petals. The evolution of such wing petals
has occurred in the clade including Paramachaerium, Pterocarpus, Ramorinoa, Paramachaerium, Tipuana, and Platypodium, and separately in that including Geoffroea.
Androecial characters
26. Staminal filaments: 0) all free from the base, 1) diadelphous 9 1 1, 2)
open monodelphous, 3) closed monodelphous, 4) diadelphous [5 1 5, 5 1 4
1 1, or 4 1 1 1 4 1 1] with at least two phalanges of fused filaments. State
two is the inferred ancestral condition of the dalbergioid legumes. State four
is synapomorphic for the Dalbergia clade (though species of Ormocarpum,
Pictetia, Diphysa, Machaerium, and Dalbergia are polymorphic). State four,
however, has evolved independently in Platypodium and Discolobium (both
of the Pterocarpus clade). The free stamens of Adesmia are inferred to represent a reversion from an open monodelphous condition. State one is uncommon among dalbergioids and is monomorphic only among some members the
Pterocarpus clade (Grazielodendron, Geoffroea, and Ramorinoa). The diadelphous 9 1 1 condition is associated with weakly developed basal fenestrae (Klitgaard, 1999a).
27. Anther size and attachment: 0) monomorphic, basi- to dorsi-fixed, 1)
dimorphic, the smaller anthers usually dorsifixed, the larger basifixed. Character state one evolved independently in the clade with Amicia (polymorphic),
Poiretia, and Zornia, and again in that with Arachis and Stylosanthes. Dimorphic anthers also have other sporadic occurrences, such as in Aeschynomene genistioides (Aeschynomene sect. Ochopodium; Rudd, 1967, 1972a), the
monotypic Aeschynomene subgen. Bakerophyton (Verdcourt, 1971), and in a
Mesoamerican clade of Platymiscium species (B. Klitgaard, unpublished data).
28. Staminal flange and filaments post-anthesis: 0) readily caducous, not
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persisting with the maturing fruit, 1) persistent on the abaxial side of fruit, 2)
persistent on adaxial side of fruit. This trait is only partially conditional upon
character number 27 (see Beyra-M. and Lavin, 1999). Persistent stamens are
diagnostic of the Dalbergia clade where the predominant condition is state
one. State two occurs in a clade with Aeschynomene sect. Aeschynomene,
Kotschya, etc. Persistent stamens evolved separately in Brya, Cascaronia, and
several outgroup genera.
Gynoecial characters
29. Locule: 0) encompassing nearly the entire length of the ovary, 1) confined to the basal end of the ovary. The locule is situated just above the stipe
in state one, and a large portion of the distal end is solid. All five occurrences
of this state are inferred to be cases of independent evolution (Vatairea, Vataireopsis, Tipuana, Centrolobium, and Paramachaerium).
30. Nectary disk: 0) absent; 1) present. A nectary disk surrounding the base
of the ovary is known from Paramachaerium (Rudd, 1981a), most species of
Ormocarpum (M. Thulin and M. Lavin, unpublished data), and occasionally
in Machaerium (Klitgaard, 1999a). The retention index is undefined in this
character (Table 1) because Machaerium and Ormocarpum were assigned an
ancestral state of zero.
Fruit and seed characters
31. Pod valves: 0) loments present during early stages of fruit development,
1) loments present by late stages, 2) valves continuous. Articulations forming
late during pod development occur in Ormocarpum, Pictetia, Diphysa, Chaetocalyx, and Nissolia. Some species in the first of these three genera form
inarticulate pods. Only in the Adesmia clade is the lomented condition uniformly present. Both of the Dalbergia and Pterocarpus clades combine genera
with articulate and inarticulate pods.
32. Pod margins: 0) straight, with no marginal constrictions between seeds
1) constricted between seeds. State one has evolved numerous times independently in Discolobium, Fiebrigiella, Brya and Cranocarpus (polymorphic),
Amicia, and Giessaspis, Bryaspis, Humularia, Kotschya, and Smithia. Aeschynomene, Ormocarpum, and Diphysa are distinctively polymorphic for this
character.
33. Stipe of mature pod: 0) absent to less than half the length of the calyx
tube, 1) surpassing the length of the calyx tube. State one has evolved most
uniformly in the clade with Dalbergia, Machaerium, Diphysa, Ormocarpum,
Ormocarpopsis, Peltiera, and Pictetia. Among dalbergioids, state zero predominates only in the Adesmia clade. Otherwise, both states have been gained
and lost on many separate occasions, particularly in the Pterocarpus clade.
34. Nervation of the mature pod valve in the region of the seed chamber:
0) primarily reticulate, 1) primarily longitudinally parallel. This trait has
evolved once in the clade with Arachis, Stylosanthes, Chapmannia, and Fiebrigiella, (not Fissicalyx, however), and again in the clade with Chaetocalyx
and Nissolia. Some species of Diphysa, Ormocarpum, and Pictetia have pods
with strong longitudinal nerves, but these genera were optimized during analysis as having state zero.
35. Replum: 0) placental margin disarticulating with the pod valves or
articles, 1) the valves or articles disarticulating separately from the persistent
placental margin. The last state is gained independently in Adesmia sect. Muricatae, Cyclocarpa, and in species of Aeschynomene sect. Aeschynomene (e.g.,
A. villosa). This character had an undefined retention index (Table 1) because
Adesmia and Aeschynomene were optimized for state zero.
36. Development of pod wings: 0) not winged, 1) wing from expansion of
the ovary wall, 2) wing from expansion of the ovary sutures, 3) wing from
attenuation of the distal end of the ovary (i.e., the style), 4) winged from
attenuation of the proximal end of the ovary (i.e., the stipe). This character
is derived from Lima’s (1990) developmental work on samaroid fruits of tribe
Dalbergieae. He distinguished wings whose area of origin was the ovary walls
(state one); wings with an origin of the ovary sutures (state two); and wings
with an origin from the distal end of the ovary (state three). Lima considered
Vatairea and Vataireopsis to have a separate state, with wings derived from
the solid distal portion of the ovary. This reflects the different morphology of
the gynoecium in these two genera (see state one of character 29). We consider that the origin of the wing in these taxa is merely from the distal end
of the ovary, and thus they are coded with state three. A further modification
is that we consider the basal wing of Platypodium to be derived from an
expansion of the stipe (state four). Developmental anatomical work could
confirm these distinctions. State one has evolved independently in both the
Dalbergia (Dalbergia and Weberbauerella) and Pterocarpus clades (Platymiscium, Cranocarpus, Grazielodendron, Ramorinoa, Pterocarpus, Fissicalyx,
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and Riedeliella). State three has evolved in every occurrence separately (Vatairea, Vataireopsis, Nissolia, Machaerium (polymorphic), Tipuana, Centrolobium, and Paramachaerium).
37. Inner epidermis and endocarp: 0) lignescent, 1) spongy and adhering
to the mature seeds. State one evolved in the clade with Chaetocalyx and
Nissolia, and again in that with Pictetia, and perhaps again in Peltiera (the
sister genus of Ormocarpopsis—see Labat and Du Puy, 1997).
38. Mesocarp: 0) lignescent, 1) spongy, 2) fleshy. State one arose independently and uniformly in Pictetia and Fiebrigiella, and sporadically in Ormocarpum, Chapmannia, Nissolia, and Chaetocalyx. State two occurs in the
fleshy vertebrate dispersed fruits of Dipteryx, Andira and Geoffroea, all instances of independent evolution. Polymorphic terminals were optimized for
state zero, effectively underestimating the actual levels of homoplasy.
39. Exocarp: 0) adnate to the mesocarp, 1) loosely attached to mesocarp,
2) separate from the mesocarp by the formation of a distinct air chamber.
This last trait occurs in many species of Diphysa and a few species of Nissolia
(e.g., N. leiogyne). State one is characteristic of Ormocarpopsis. The high tree
scores for this character (Table 1) resulted from state zero being assigned to
the polymorphic terminals Diphysa and Nissolia during parsimony analysis.
40. Pod coiling: 0) coiled in a forward directed manner, 1) not coiled, 2)
coiled in a laterally directed manner. The forward coil of the pod is confined
to Discolobium, whereas the lateral coil has evolved in Cyclocarpa and various species of Aeschynomene sect. Aeschynomene and Ormocarpum. Although the lateral coil is restricted to members of the Dalbergia clade, state
one was assigned to the polymorphic terminals Aeschynomene and Ormocarpum during parsimony analysis, resulting in high tree scores (Table 1).
41. Pod valve ornamentation: 0) not present, 1) multiseriate trichomes, 2)
crests and bumps. Multiseriate trichomes persisting on the mature pod valve
have evolved in many separate occasions throughout the dalbergioid legumes,
as in Adesmia (polymorphic), Ormocarpum (polymorphic), Brya, and Centrolobium (where they become spinose). Pod valves with crests or bumps have
evolved once in the clade with Poiretia, Amicia, and Zornia, and again in
polymorphic condition among various species of Aeschynomene sect. Aeschynomene. The polymorphic terminals Adesmia, Ormocarpum, and Aeschynomene were assigned state zero during parsimony analysis, thus resulting in
relatively high tree scores (Table 1).
42. Seed shape: 0) lenticular to spherical with a centrally placed hilum, 1)
reniform with a central recessed hilum, 2) longitudinally elongate with the
hilum placed toward the end toward the style. The last condition is characteristic of most dalbergioids and indeed ancestral to that clade. However, state
two has evolved independently in Dipterygeae (Dipteryx and Pterodon), Hymenolobium, Vatairea (polymorphic), and Vataireopsis. Among dalbergioids,
Aeschynomene, Kotschya, Smithia, and Platymiscium have reniform seeds
(three separate origins), and Ormocarpopsis and Peltiera have spherical seeds.
43. Orientation of the seed in the fruit: 0) longitudinal, 1) oblique to transverse. Oblique to transverse orientation of seeds is confined to a subclade
including Platymiscium, Centrolobium, Paramachaerium, Pterocarpus, Ramorinoa, and Tipuana (Lima, 1990). Such seeds have also evolved independently in Hymenolobium (polymorphic).
Seedling characters
44. Position of the eophylls: 0) alternate, 1) opposite. Among dalbergioids,
opposite eophylls are confined to the Pterocarpus clade where they are known
from Platymiscium, Grazielodendron, Ramorinoa, Centrolobium, and Geoffroea.
45. Number of leaflets in the first eophyll: 0) one, 1) more than one. Dalbergioids commonly have eophylls that are not strongly differentiated from
the adult leaves (i.e., multifoliolate). The Pterocarpus clade is exceptional in
having all known instances where the eophylls are unifoliolate. Data for characters 44 and 45 for Ramorinoa came from Burkart (1952, p. 238).
Pollen characters
46. Aperture type: 0) tricolporate, 1) periporate. Tricolporate apertures are
the general and most common type in legumes. Among the dalbergioids, periporate pollen is known from only Brya and Cranocarpus.
47. Pollen pore: 0) without an operculum, 1) with an operculum. An operculum is a distinctly delimited ectexinous structure that covers the ectoaperture, which in the case of all dalbergioids means the colpus. State one has
been gained independently many times throughout all three principal clades
of the dalbergioid legumes.
48. Colpi (the polar region): 0) colpi short, not anastomosing at the poles
of the pollen grain, the polar region entire, 1) colpi longer, the ends of colpi
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anastomosing, forming syncolpi. State one evolved independently and uniformly in Humularia and Vataireopsis. This state is polymorphic for Aeschynomene, Kotschya, and Smithia. Thus, state one is confined to the Dalbergia
clade among the dalbergioid legumes.
49. Wall stratification (Guinet and Ferguson, 1989): 0) well-developed endexine and footlayer, 1) thickening of the endexine at least at the apertures
combined with a reduction in the foot layer, 2) reduction of the endexine
combined with a thickening of the foot layer, 3) reduction of the endexine
and foot layer combined with an elongation of the columellae. State one is
most common among the dalbergioids and is inferred to be ancestral in this
clade. State two evolved independently in Adesmia and the outgroup Myrospermum. State three evolved once in the clade with Geissaspis and Bryaspis.
Wood characters
50. Ray size: 0) three or more cells wide, and taller than 20 cells high, 1)
1–2 cells wide and less than 15–20 cells high. Many of the dalbergioid genera
have narrow short rays (state one). Good examples include Platymiscium,
Fissicalyx, and the nondalbergioid Dipteryx. Outgroup genera have storied
rays that are larger than this. Wood characters were scored from examination
of slides in the collections at the Jodrell Laboratory, Kew, and at the Forest
Products Laboratory, Madison, Wisconsin, USA, and by reference to descriptions and photographs in Baretta-Kuipers (1981), Gasson (1994, 1999), Miles
(1978), and Détienne and Jacquet (1983). Wheeler, Bass, and Gasson (1989)
provide thorough definitions for all of the wood characters used in this analysis. Details on the wood anatomy of Chaetocalyx, Nissolia, Poiretia, Amicia,
Zornia, Chapmannia, Arachis, Stylosanthes, Soemmeringia, Smithia and Geissaspis come from Cumbie (1960). Unfortunately, the information is presented
in such a way that only a few character states can be coded. Amorpha fruticosa has been illustrated and described by Schweingruber (1990), Adesmia
horrida by Roig (1986), Discolobium by Cozzo (1949, 1950), and Paramachaerium by Brizicky (1960). No information on the wood anatomy is available for Riedeliella, Cranocarpus, Fiebrigiella, Cyclocarpa, Kotschya, Bryaspis, Humularia, Weberbauerella, Ormocarpum, Ormocarpopsis, and Peltiera.
51. Ray arrangement: 0) not storied, 1) storied. In legumes, storied rays,
axial parenchyma, and adjacent vessels are common and can be observed in
tangential longitudinal section. Although considered a very useful anatomical
character, both diagnostically and cladistically, there are many legume genera
with storied rays that are irregular, or obvious in short rays and less so in
taller rays which may be axially fused. Storied rays are particularly strongly
developed in Dipteryx, Pterocarpus, Platymiscium, Grazielodendron, Etaballia, Inocarpus, Dalbergia, Machaerium, and Aeschynomene, all of which have
short rays. The taxa with larger rays often do not exhibit such regular storied
arrangement, and axial fusion is often the cause, as in species of Acosmium.
52. Composition of cells in rays: 0) homocellular, 1) heterocellular. This
feature is observed in radial longitudinal section. Homocellular rays are composed entirely of procumbent ray cells. Heterocellular rays in legumes are
composed mainly of procumbent cells, but there are also some square or
upright cells, usually in a row or rows at the ray margins (i.e., at the top or
bottom of a ray). These two character states are not mutually exclusive. Juvenile wood often tends to be more heterocellular than mature wood, and it
is not always apparent where exactly a wood sample was taken from if the
pith in the stem is not included.
53. Crystals in ray cells: 0) absent, 1) present in some ray cells. Prismatic
crystals of calcium oxalate are found in many, if not most legumes. They are
almost ubiquitous in chambered axial parenchyma strands, but in a few genera
can also be found in ray cells. The main difficulty with this character is that
if the crystals are rare they can be overlooked. They are searched for in radial
longitudinal section, because they are even more difficult to find in tangential
longitudinal section.
54. Axial parenchyma: 0) not abundantly aliform and confluent, 1) abundantly aliform and confluent. Axial parenchyma patterns in legumes are very
difficult to code. All the legumes in this study have predominantly paratracheal parenchyma, with the addition of some apotracheal diffuse parenchyma
in particularly Dalbergia and Platymiscium. This ranges continuously from
scanty paratracheal, vasicentric, aliform, to confluent. In the opinion of one
of us (Gasson), these all constitute one character. They could each be coded
as character states, but virtually all wood samples in the legumes exhibit more
than one condition. Unilaterally paratracheal parenchyma is found in some of
the taxa, and probably forms part of this continuum. Banded parenchyma,
which could be treated separately, may be an extreme form of confluent parenchyma, particularly if the bands are several cells wide. Some taxa in the
study group do have narrow bands, but they are not distinguished here. The
choice of the two character states above serves to separate four of the genera
529
very well, but does not distinguish all the other complicated variations on the
paratracheal theme exhibited by the taxa coded as zero. Aeschynomene is very
different, in that it has such abundant parenchyma, that the fibers exhibit a
winged-aliform appearance.
Nitrogen fixation character
55. Root nodule: 0) none produced, 1) produced as a non-aeschynomenoid
nodule, 2) produced as an aeschynomenoid nodule (Fig. 6; Corby, 1981; see
discussion). State two is synapomorphic for the dalbergioid clade, although
some genera in this clade are known not to produce nodules at all (i.e., Chaetocalyx and Nissolia), as is the case for some nondalbergioids (e.g., Myrospermum, Dipteryx, Pterodon, Vatairea, and Vataireopsis). Some outgroups
produce nodules of a type other than aeschynomenoid (e.g., Acosmium, Poecilanthe, Andira, Hymenolobium, and Amorpha). Although lost within the
dalbergioid clade (Table 1), the aeschynomenoid root nodule is not encountered elsewhere in the legume family. The stem (but not root) nodules of
Sesbania rostrata (tribe Robinieae) are superficially similar, but they differ
from the aeschynomenoid type in having an apical meristem, albeit ephemeral
(J. Sprent, unpublished data). Three dalbergioid genera, Cyclocarpum, Geissaspis, and Paramachaerium, are known to produce nodules, but the exact
type is unknown. These genera were variously coded as having missing data
or state one. Such alternative coding did not affect how these genera were
related with respect to the three major subclades of the dalbergioid clade.
Future studies incorporating nodule morphology in a phylogenetic analysis
will do well to recognize specific morphologies independent of nodule categories. Specifically, these would include characters such as the apical meristem (absent vs. present), infection site (associated with emergent rootlet or
not), infection threads (absent vs. present), and central tissue (uniformly infected vs. uninfected). The aeschynomenoid type is defined as having the first
state of each of these four characters. Regardless, this coding strategy would
not change our findings because the dalbergioids would be nearly uniform in
occurrence for the states of these four characters.
APPENDIX C
Enumeration of the constituent genera of the dalbergioid clade. The emphasis in the discussion of each of the dalbergioid genera is on the diagnostic
traits that are presumably autapomorphic.
The Adesmia clade
Adesmia DC. is diagnosed by stipules attached a pseudopetiole. Although
also found in Stylosanthes and Arachis, the projected portions of the nodes
of these two genera are nearly as long as the petiole. In Adesmia, the nodal
projections extend to much less than half the length of the petiole. In addition,
Adesmia uniquely combines free staminal filaments and lomented pods (Polhill, 1981f). Adesmia comprises about 230 species centered in Chile and Argentina (Burkart, 1949, 1954, 1960, 1962, 1964, 1966, 1967; Ulibarri, 1978,
1980, 1982a, b, 1984, 1987, 1990). The genus contains two distinct monophyletic subgroups, according to ITS/5.8S sequence analysis (Fig. 3). One is
marked by inflorescences of usually solitary axillary flowers with pedicels
confluent with the calyx, stipules (or at least scars) that are connate around
the stem, and pods that lack glandular-based trichomes, multiseriate trichomes,
or the raised pericarp reticulations (e.g., Adesmia lanata and A. villosa). The
second clade is characterized by inflorescences of terminal racemes, subumbels, or panicles, flowers articulated with the pedicel, stipules (or scars) that
are not connate around the stem, and pod loments that commonly bear some
type of ornamentation, for example, large glandular-based trichomes, long
multiseriate plumose trichomes, or very prominent reticulate venation (e.g.,
Adesmia muricata and A. volckmannii). Phylogenetic analysis of ITS/5.8S
sequence data strongly supports the monophyly of Adesmia, as does matK/
trnK.
Chaetocalyx DC. (Figs. 17–23) is paraphyletic with respect to Nissolia, an
issue that is the focus of another study (M. Lavin and D. Prado, unpublished
data). It possesses no autapomorphic traits and is characterized like Nissolia
(with twining herbaceous stems and ebracteolate flowers) but lacking the sterile (usually samaroid) terminal loment of the mature pod. The glandular-based
trichomes on the calyx of most species of Chaetocalyx are not diagnostic and
the species of Chaetocalyx form a rather homogeneous assemblage. The supposedly obvious division between species with laterally flattened or winged
fruits vs. those with terete fruits (as coded in Beyra-M. and Lavin, 1999) is
not resolved with 5.8S/ITS sequence analysis. Chaetocalyx includes ;13 neotropical species centered in dry forests of South America (Rudd, 1958, 1972b,
1996).
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Figs. 17–28. Representative species of the Adesmia clade (scale bar 5 1 cm for all figures). Figs. 17–23. Chaetocalyx brasiliensis. 17. Habit. 18. Calyx.
19. Gynoecium. 20. Androecium. 21. Keel petal. 22. Wing petal. 23. Standard. Figs. 24–27. Nissolia wislizenii. 24. Habit. 25. Cauline leaf. 26. Flower. 27.
Fruits. 28. Nissolia microptera, leafy stem with fruits. Reproduced from Volume 5 of Flora Novo-Galiciana by Rogers McVaugh.
Nissolia Jacq. is derived from within Chaetocalyx (Figs. 2, 3, and 5) and
characterized by the autapomorphy of pods with a sterile (usually samaroid)
terminal loment (Figs. 24–28). This genus contains ;13 species centered in
tropical dry forests of Mexico and Central America (Rudd, 1956, 1970a,
1975b). Chaetocalyx and Nissolia lack floral bracteoles, otherwise occurring
among dalbergioids in Poiretia, Amicia, Zornia, Cyclocarpa, Humularia,
Geissaspis, and Bryaspis.
Amicia Kunth occurs in Mexico, Ecuador, Peru, Bolivia, and Argentina
(Rudd, 1981a). This genus is closely related to Poiretia and Zornia. All three
have legumes with crests or bristles on each pod article and leaves that are
usually paripinnate (a few species of Poiretia have imparipinnate leaves).
Amicia differs from Poiretia and Zornia in having blunt keel petals, a staminal
sheath that is split open above, and anthers that are mostly uniform. A recent
attempt to segregate Poiretia and Zornia from Amicia (Ohashi, 1999) is not
supported by this analysis.
Poiretia Vent. is confined to the Neotropics but with most species from
Brazil to northern Argentina (Rudd, 1972c). The genus is similar to Zornia,
but differs in its usually twining habit and racemose inflorescences with small,
single flower bracts at each node.
Zornia J. F. Gmel. occurs in southeastern United States, the Neotropics with
a center of diversity in Brazil, and throughout sub-Saharan Africa (Mohlenbrock, 1961, 1962). It is marked by medifixed stipules (independently evolved
in Aeschynomene sect. Aeschynomene and relatives), leaves with digitately
arranged few leaflets, and sessile flowers in axils of large paired bracts.
The Pterocarpus clade
Discolobium Benth. is readily diagnosed by its pod that coils in a forward
direction with each of three turns compressed together into a single disc. Only
the middle loment is fertile. Its 4 1 1 1 4 1 1 diadelphous staminal column
is not unique and is found sporadically among the dalbergioids. Discolobium
comprises eight species distributed from northern Argentina to adjacent Brazil
and Paraguay (Rudd, 1981a).
Riedeliella Harms comprises three species endemic to southeastern Brazil
and Paraguay (Lima and Studart da Fanseca Vaz, 1984). Like Inocarpus,
Etaballia, and some species of Pictetia, the flowers of Riedeliella are nearly
radially symmetric. Lima and Studart da Fanseca Vaz (1984) propose the close
relationship of Etaballia and Riedeliella in the tribe Acosmieae (Yakovlev,
1972), a group also with essentially radially symmetric flowers, although with
free staminal filaments. Riedeliella differs from Etaballia and Inocarpus in
having paripinnate leaves and a long exerted style, and in this analysis it is
suggested to be not most closely related to Etaballia, but rather to Discolobium.
Brya P. Br. is recorded to have explosive pollen release (León and Alain,
1951, p. 315, fig. 131), a form of pollen presentation that is unique among
dalbergioid legumes. Also, Brya is characterized by its leaves from the long
shoots being transformed into spines. Brya is sister to Cranocarpus, as evidenced by the shared occurrence of leaves, stems, inflorescences, and pods
bearing capitate glandular trichomes that are microscopically glochidiate, and
by periporate pollen (Ferguson and Skvarla, 1981). Brya includes four species
endemic to the Greater Antilles (Ohashi, Polhill, and Schubert, 1981; Lewis,
1988).
Cranocarpus Bentham comprises three species endemic to Brazil (Harley,
1978; Ohashi, Polhill, and Schubert, 1981). In all respects Cranocarpus is
like Brya but the leaves from the long shoots are not transformed into spines.
The yellow petals, base chromosome number of x 5 10, storied wood structure (Record, 1919), and simple axillary racemes or solitary flowers of Brya
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531
Figs. 29–47. Representative species of the Pterocarpus clade (scale bar 5 1 cm for all figures). Figs. 29–36. Platymiscium trifoliolatum. 29. Flowering
branch. 30. Branch of fruiting inflorescences with wall of one fruit cut away to show seed. 31. Calyx. 32. Androecium. 33–34. Wing petals. 35. Keel petals.
36. Standard. Figs. 37–47. Pterocarpus orbiculatus. 37. Detached leaf. 38. Inflorescence. 39. Mature fruits. 40. Immature fruits. 41. Calyx. 42. Androecium.
43. Gynoecium. 44. Keel petals. 45–46. Wing petals. 47. Standard. Reproduced from Volume 5 of Flora Novo-Galiciana by Rogers McVaugh.
and Cranocarpus are traits strongly suggestive of a relationship with the dalbergioid legumes.
Platymiscium Vogel (Figs. 29–36) comprises 18 neotropical species centered in Mexico and northeastern Brazil. The genus is unique in having opposite leaves with interpetiolar stipules (Lima, 1990; Klitgaard, 1999a, b).
Centrolobium Mart. ex Benth. comprises six tropical species from Panama
to Colombia, Ecuador, Venezuela, Brazil, and Bolivia (Rudd, 1954; Lima,
1985). The genus is well marked by its orange peltate glands covering the
leaves and inflorescences, and winged pods in which the seed-bearing portion
is covered with spines.
Grazielodendron Lima is a monotypic genus endemic to Brazil (Lima,
1983, 1990). The laterally compressed pod of Grazielodendron is distinguished by having an additional wing-like extension of the dorsal margin. The
winged dorsal margin is markedly delineated from the main winged body of
the pod.
Pterocarpus Jacq. (Figs. 37–47) comprises 20 species distributed pantropically (Rojo, 1972). Of the genera with wide, crimped wing petals (i.e., Paramachaerium, Geoffroea, Pterocarpus, Ramorinoa, Paramachaerium, Tipuana, and Platypodium), Pterocarpus is diagnosed by its pod that is winged
from an attenuation of the pod body all around the seed chamber (Polhill,
1981d; Lima, 1990). The pods are variable in this genus with some winged
and bristly (e.g., P. angolensis), others winged and not bristly (e.g., P. indicus), and rarely not winged and not bristly (i.e., P. amazonum).
Tipuana (Benth.) Benth. is a monotypic genus of subtropical forests in
Bolivia and northwestern Argentina (Rudd, 1974). Of the genera with wide,
crimped wing petals (see description of Pterocarpus), Tipuana is diagnosed
by its pod that is winged from the style, the seed chambers being proximal
to the wing (Lima, 1990; Polhill, 1981d).
Platypodium Vogel includes one or two species in Panama, Guatemala,
Venezuela, Colombia, Bolivia, Brazil, and Paraguay. Of the genera with wide,
crimped wing petals (see description of Pterocarpus), Platypodium is diagnosed by its pod that is winged from the stipe, the seed chambers being distal
to the wing (Polhill, 1981d; Lima, 1990).
Paramachaerium Ducke includes five species from Panama, Guyana, Peru,
and Brazil (Rudd, 1981a, b; Lima, 1990). Of the dalbergioid tree genera with
laterally broadened and crimped wing petals, Paramachaerium has reddish to
violet petals rather than the typical yellow pigment. This genus is unusual in
its nectariferous disk surrounding the base of the ovary, a trait independently
evolved in certain species of Machaerium and Ormocarpum.
Ramorinoa Speg. is a monotypic genus from west-central Argentina. The
genus is very well marked by its leafless pungent branches (Burkart, 1952;
Polhill, 1981d; Lima, 1990). As remarked by Burkart (1952), the genus is so
highly modified vegetatively that morphology provides few clues to its closest
relationships.
Inocarpus J. R. & G. Forster is very distinctive in having all five ligulate
petals fused at base, and with the ten stamens fused by their filaments to the
corolla tube (similar to some genera of Amorpheae). Inocarpus comprises one
to three species and is geographically distinctive in being restricted to Malaysia and adjacent Pacific islands (Polhill, 1981d). The only other dalbergioid
genus restricted to Asia is Geissaspis.
Etaballia Benth. is very similar to Inocarpus, except that its leaves are
unifoliolate rather than simple, and the staminal filaments are monodelphous
with no split along the adaxial side. Etaballia is monotypic and unlike Inocarpus is neotropical, occurring in Guyana, Venezuela, and Brazil (Rudd,
1970b).
Geoffroea Jacq. comprises two species from Colombia and Venezuela south
to Chubut, Argentina, and also on the Galapagos Islands possibly due to
cultivation (Ireland and Pennington, 1999). Of the genera with wide, crimped
wing petals (see description of Pterocarpus), Geoffroea is diagnosed by its
sessile ovary that develops into a fleshy drupe (Polhill, 1981d; Lima, 1990).
532
AMERICAN JOURNAL
OF
BOTANY
[Vol. 88
Figs. 48–69. Representative species of the Dalbergia clade (scale bar 5 1 cm for all figures except where noted). Figs. 48–58. Dalbergia congestiflora. 48.
Leafy branch. 49. Flowering branch. 50. Fruits. 51. Seed. 52. Androecium. 53. Anther (scale bar 5 1 mm). 54. Calyx. 55. Keel petals. 56–57. Wing petals. 58.
Standard. Figs. 59–69. Machaerium kegelii. 59. Flowering branch. 60–61. Nodes with stipular spines. 62. Androecium. 63. Gynoecium. 64. Calyx. 65. Keel
petals. 66. Wing petal. 67. Standard. 68. Flower. 69. Fruit. Reproduced from Volume 5 of Flora Novo-Galiciana by Rogers McVaugh.
Cascaronia Griseb. is not readily diagnosed, but the combination of its
leaves and pods with dark pustular glands, pods with strong longitudinal
nerves, arborescent habit, inflorescences of axillary racemes, and small yellow
petals is unique. Cascaronia is a monotypic genus from northern Argentina
and adjacent Paraguay and Bolivia (Polhill, 1981d).
Fissicalyx Bentham is a monotypic genus from Venezuela and Guyana, and
marked by its spathaceous calyx (all five lobes are on the adaxial lip), porate
anthers, and pods with a fusiform seed chamber bearing a closely veined
membranous wing on both margins (Polhill, 1981d; Lima, 1990). There is no
morphological evidence to suggest that this genus is closely related to Fiebrigiella, as revealed by DNA sequence analysis.
Fiebrigiella Harms is a monotypic genus from Bolivia and Ecuador (Burkart and Vilchez, 1971). The pods of Fiebrigiella have prominent continuous
parallel venation on the lateral walls, once suggesting an affinity to Chaetocalyx and Nissolia (Rudd, 1981a), but now considered homologous to such
pods of the genera Chapmannia, Stylosanthes, and Arachis.
Chapmannia Torr. & Gray is recently expanded from monotypic (Gunn,
Norman, and Lassetter, 1980) to include seven species of seasonally dry vegetation, two New World (Florida and Mesoamerica), and five Old World (Somalia and the Yemeni island Socotra; Thulin, 2000). Arthrocarpum Balf. f.
(Gillett, 1966) and Pachecoa Standl. & Steyerm. (Burkart, 1957) are synonymized. The genus is diagnosed by its dried (herbarium preserved) leaflets
with uniformly reddish reticulate tannin deposits on the abaxial surface. Chapmannia is sister to Arachis, and Stylosanthes; the species of these two latter
genera do not consistently show the reddish tannin reticulations. Chapmannia,
Arachis, and Stylosanthes form a monophyletic group marked in part by their
sessile flowers with long hypanthia. Within this group, Chapmannia maintains
the plesiomorphic spicate inflorescence, whereas Arachis and Stylosanthes
have inflorescences of solitary axillary flowers.
Stylosanthes Swartz and Arachis share the synapomorphy of stipules united
to nodal projections, which in turn are superficially continuous with the petiole
(a trait known also from Adesmia). Stylosanthes is distinguished from Arachis
by having lomented, nongeocarpic pods, which are presumably plesiomorphic,
as well as ovaries that are uniformly covered by uniseriate trichomes, an
autapomorphy. The ;25 species of Stylosanthes are distributed in warm temperate to tropical regions of the world, but with a center of diversity in the
neotropics (Mohlenbrock, 1957, 1960, 1963; Rudd, 1981a).
Arachis L. is distinguished from Stylosanthes by its flowers with a very
long and narrow hypanthium, a gynophore (Moctezuma and Feldman, 1998)
that renders the pods geocarpic, nonlomented glabrous pods, and mostly four
leaflets per leaf. The 69 species of Arachis originate in South America from
a region including Brazil south to northern Argentina (Krapovickas and Gregory, 1994).
The Dalbergia clade
Dalbergia L. f. (Figs. 48–58) is diagnosed by small ovate to obovate anthers with short transverse slits at dehiscence. The genus includes over 100
species distributed pantropically, but with centers of diversity in Amazonia
and Indo-Asia (Prain, 1904; Pittier, 1922; Polhill, 1981d; Lima, 1990; de Carvalho, 1997).
Machaerium Pers. (Figs. 59–69) includes ;120 neotropical species, although M. lunatum (L. f.) Ducke also occurs in western Africa. Machaerium
is related to Dalbergia (Polhill, 1981d; Doyle et al., 1997) and Aeschynomene
March 2001]
LAVIN
ET AL.—DALBERGIOID LEGUMES
sect. Ochopodium, as evinced in part by inflorescences of helicoid cymes (but
polymorphic in all three taxa). Machaerium differs in its spinescent recurved
stipules (on at least the climbing species) and pods that are usually distally
winged, or at least have the seed chamber toward the base (Rudd, 1973, 1977,
1986, 1987; Polhill, 1981d; de N. Carmo-Bastos, 1987; Lima, 1990).
Aeschynomene L. sect. Ochopodium. Aeschynomene sensu lato includes
species that do not fit the diagnosis of the other dalbergioid genera. It is for
this reason that the genus is treated with two terminal taxa, sects. Ochopodium
(with basifixed stipules) and Aeschynomene (with medifixed stipules). Section
Ochopodium, according to DNA sequence analysis, is more closely related to
Machaerium than to sect. Aeschynomene (e.g., section Ochopodium is represented by Aeschynomene purpusii and A. fasicularis in Fig. 5). Regardless,
it is not certain if either of these two sections are monophyletic, a topic that
will have to be taken up elsewhere given their large taxonomic size. As such,
sect. Ochopodium includes ;101 species distributed pantropically (Rudd,
1955, 1967, 1975a).
Aeschynomene sect. Aeschynomene is diagnosed by medifixed stipules,
which are also characteristic of the closely related Smithia and Geissaspis.
Thus, this taxon (represented by Aeschynomene americana, A. indica, A. pfundii, A. rudis, and A. virginica in Figs. 2 and 5), potentially lacking any obvious
morphological apomorphy, could be paraphyletic with respect to at least some
of the genera listed immediately below. It is beyond the scope of this analysis
to address this potential problem. As such, sect. Aeschynomene comprises
;50 species with a pantropical distribution (Léonard, 1954; Rudd, 1955,
1959, 1972a; Verdcourt, 1971; Fernandes, 1996).
Soemmeringia Mart. is characterized by a scarious standard petal that persists with the mature pod, which is independently evolved in some species of
Ormocarpum. Soemmeringia is a monotypic, neotropical genus from Brazil,
Bolivia, and Venezuela (Rudd, 1981a). Soemmeringia, along with Cyclocarpa,
Kotschya, Smithia, Geissaspis, Bryaspis, and Humularia (below), are all
closely related to sect. Aeschynomene because of their paripinnate leaves,
usually alternate leaflets, and bilabiate calyces (Rudd, 1981a).
Cyclocarpa Afz. ex Bak. is diagnosed by pods that have one lateral spiral,
the pod articles of which disarticulate from a persistent placental margin or
replum. This monotypic genus is locally common across tropical Africa, and
in southeast Asia (Laos and Borneo) and northern Australia (Hepper, 1958).
Kotschya Endl. and Smithia (below) are characterized by an inflorescence
of a dense strobilate helicoid cyme, a pod enclosed by the calyx and in which
the articles are folded against each other. Kotschya differs in having alternate
leaflets that each bear 2–7 basal nerves, as well as basifixed stipules. Kotschya
comprises 31 species restricted to tropical Africa and Madagascar (Gillett,
Polhill, and Verdcourt, 1971; Verdcourt, 1974; Rudd, 1981a).
Smithia Ait. differs from Kotschya by its medifixed stipules and opposite
leaflets each bearing one main nerve. Smithia comprises ;30 species mainly
in Asia and Madagascar (Gillett, Polhill, and Verdcourt, 1971; Verdcourt,
1974; Rudd, 1981a).
Geissaspis Wight & Arn. together with Bryaspis and Humularia (below)
are characterized by large inflorescence bracts that completely envelop the
subtending flower and fruit (independently evolved in Zornia). Geissaspis and
Bryaspis differ by ebracteolate flowers, and Geissaspis differs from Bryaspis
by its medifixed stipules. Geissaspis comprises three species confined to tropical and subtropical central and southeast Asia, but not crossing Wallace’s line
(Gledhill, 1968; Rudd, 1981a)
533
Bryaspis Duvign. includes two species from tropical west Africa (Gledhill,
1968; Hepper, 1958; Gillett, Polhill, and Verdcourt, 1971; Rudd, 1981a). Unlike Geissaspis, the inflorescence bracts of Bryaspis are markedly imbricate.
Humularia Duvign. differs from Geissaspis and Bryaspis by emarginate
inflorescence bracts and panduriform standard petals. Humularia comprises
;40 species confined to central Africa (Gledhill, 1968; Gillett, Polhill, and
Verdcourt, 1971; Verdcourt, 1974; Rudd, 1981a).
Weberbauerella Ulbrich is diagnosed by the combination of its herbaceous
habit, pustular glands densely covering the stems, leaves, and inflorescences
(including petals), and leaves with well over 40 leaflets. Similar pustular
glands on the petals are known from Poiretia, but this genus is marked by
leaves with four leaflets, and a sometimes climbing habit. Weberbauerella
contains two species confined to sand in southern coastal Peru (Ferreyra,
1951; Rudd, 1981a).
Pictetia DC. is characterized by spiny stipules, short shoots bearing distichously arranged stipules (shared with Ormocarpum, Ormocarpopsis, and
Peltiera), coriaceous leaflets that in all but two species have spinescent mucros, and pods with two-ribbed placental margins. Pictetia includes eight species confined to Cuba, Hispaniola, Puerto Rico, and the Virgin Islands excluding St. Croix (Beyra-M. and Lavin, 1999).
Diphysa Jacq. has been characterized by its mature pods that have an exocarp distinctly inflated and separated from the mesocarp. However, Diphysa
ormocarpoides and D. spinosa have laterally flattened lomented pods very
similar to species of Ormocarpum and Pictetia (Antonio and Sousa, 1991).
The monophyly of this genus is strongly supported, however, by phylogenetic
analysis of molecular data (see also Beyra-M. and Lavin, 1999; Lavin et al.,
2000). The genus includes about ten species centered in Mexico and Central
America (M. Lavin, unpublished data).
Ormocarpum P. Beauv. is diagnosed by most species forming a cylindrical
nectary disk surrounding the base of the ovary (M. Thulin and M. Lavin,
unpublished data). This trait otherwise is known in a few species of Machaerium and Paramachaerium. This genus of ;20 species is primarily African. Three species occur on the southern Arabian Peninsula in Yemen (including Socotra) and Oman, and one to two species occur in tropical Asia
and Australia (Gillett, 1966; Rudd, 1981a; Thulin, 1990). According to ITS/
5.8S sequence data (see also Lavin et al., 2000), Ormocarpum comprises two
lineages (one with and one without the intrastaminal disk) that are collectively
paraphyletic with respect to Ormocarpopsis (and Peltiera). This issue is being
addressed in a separate study (M. Thulin and M. Lavin, unpublished data).
Ormocarpopsis R. Viguier has short shoots with persistent distichously arranged stipules shared with Ormocarpum, Peltiera, and Pictetia. In this context, its non-lomented pod with a smooth exocarp (no evidence of prominent
parallel nervation on the pod valves) and tannin patches on the abaxial surface
of dried leaflets are diagnostic. Ormocarpopsis comprises six species endemic
to Madagascar (Labat and Du Puy, 1996).
Peltiera Labat & Du Puy includes two endemic Madagascan species that
are sister to Ormocarpopsis (Labat and Du Puy, 1997). These two genera
share a distinctive tannin patterning on the abaxial surface of herbarium-dried
leaflets where tannin deposits are concentrated along the midrib. Like Ormocarpopsis, the flowers of Peltiera lack a nectary disk (M. Thulin and M.
Lavin, unpublished data), and the pods, though lomented and with all but one
loment aborting, contain spherical seeds. The pod valves in the seed-bearing
article are dehiscent. Unfortunately, both species of Peltiera are probably
extinct due to the clearing of forests from which they were known.