The dalbergioid legumes (Fabaceae): delimitation of a pantropical monophyletic clade

Matt Lavin; R. Toby Pennington; Bente B. Klitgaard; Janet I. Sprent; Haroldo Cavalcante de Lima; Peter E. Gasson

The classification of the legume family proposed here addresses the long-known non-monophyly of the traditionally recognised subfamily Caesalpinioideae, by recognising six robustly supported monophyletic subfamilies. This new classification uses as its framework the most comprehensive phylogenetic analyses of legumes to date, based on plastid matK gene sequences, and including near-complete sampling of genera (698 of the currently recognised 765 genera) and ca. 20% (3696) of known species. The matK gene region has been the most widely sequenced across the legumes, and in most legume lineages, this gene region is sufficiently variable to yield well-supported clades. This analysis resolves the same major clades as in other phylogenies of whole plastid and nuclear gene sets (with much sparser taxon sampling). Our analysis improves upon previous studies that have used large phylogenies of the Leguminosae for addressing evolutionary questions, because it maximises generic sampling and provides a phylogenetic tree that is based on a fully curated set of sequences that are vouchered and taxonomically validated. The phylogenetic trees obtained and the underlying data are available to browse and download, facilitating subsequent analyses that require evolutionary trees. Here we propose a new community-endorsed classification of the family that reflects the phylogenetic structure that is consistently resolved and recognises six subfamilies in Leguminosae: a recircumscribed Caesalpinioideae DC., Cercidoideae Legume Phylogeny Working Group (stat. nov.), Detarioideae Burmeist., Dialioideae Legume Phylogeny Working Group (stat. nov.), Duparquetioideae Legume Phylogeny Working Group (stat. nov.), and Papilionoideae DC. The traditionally recognised subfamily Mimosoideae is a distinct clade nested within the recircumscribed Caesalpinioideae and is referred to informally as the mimosoid clade pending a forthcoming formal tribal and/or cladebased classification of the new Caesalpinioideae. We provide a key for subfamily identification, descriptions with diagnostic charactertistics for the subfamilies, figures illustrating their floral and fruit diversity, and lists of genera by subfamily. This new classification of Leguminosae represents a consensus view of the international legume systematics community; it invokes both compromise and practicality of use.

Using as a starting point Doyle (1996), we compiled a morphological cladistic matrix of 54 coded taxa (31 wholly extinct, and 23 at least partly extant) and 102 informative characters in order to explore relationships among gymnosperms in general and pteridosperms in particular. Our core analysis omitted six supplementary fossil taxa and yielded 21 most-parsimonious trees that generated two polytomies in the strict consensus tree, both among pteridosperms; the first affected several hydraspermans, and the second affected the three peltasperm/ corystosperm taxa analyzed. The resulting topology broadly resembled topologies generated during previous morphological cladistic analyses that combined substantial numbers of extant and extinct higher taxa. The five groups that include extant taxa were relatively well resolved as monophyletic and yielded the familiar Anthophyte topology (cycads (Ginkgo (conifers (Gnetales, angiosperms))))), strongly contradicting most recent DNA-based studies that placed Gnetales as sister to, or within, conifers. These five extant groups were embedded in the derived half of a morphologically diverse spectrum of extinct taxa that strongly influenced tree topology and elucidated patterns of acquisition of morphological character-states, demonstrating that pteridosperms and other, more derived “stem-group” gymnosperms are critical for understanding seed-plant relationships. Collapses in strict consensus trees usually reflected either combinations of data-poor taxa or “wildcard” taxa that combine character states indicating strongly contradictory placements within the broader topology. Including three progymnosperms and identifying the aneurophyte progymnosperm as outgroup proved crucial to topological stability. An alternative progymnosperm rooting allowed angiosperms to diverge below cycads as the basalmost of the extant groups, a morphologically unintuitive position but one that angiosperms have occupied in several recent molecular studies. We therefore believe that such topologies reflect inadequate rooting, which is inevitable in analyses of seed plants that use only extant taxa where the outgroups can only be drawn from ferns and/or lycopsids, groups that are separated from extant seed-plants by a vast phylogenetic discontinuity that is bridged only by wholly fossil groups. Given the rooting problem, and the poverty of the hypotheses of relationship that can be addressed using only extant taxa, morphology-based trees should be treated as the initial phylogenetic framework, to subsequently be tested using molecular tools and employing not only molecular systematics but also evolutionary-developmental genetics to test ambiguous homologies. Among several possible circumscriptions of pteridosperms, we prefer those that imply paraphyly rather than polyphyly and exclude only one monophyletic group, providing one cogent argument for the inclusion of extant cycads in pteridosperms. Although pteridosperms cannot realistically be delimited as a monophyletic group, they remain a valuable informal category for the plexus of gymnosperms from which arose several more readily defined monophyletic groups of seed-plants. The ideal solution of recognizing several monophyletic groups, each of which incorporates a “crown-group” and one or more pteridosperms, is not yet feasible, due to uncertainties of relationship and difficulties to satisfactorily delimiting the resulting groups using reliable apomorphies. Exploration of the matrix demonstrated that coding all of the organs of a plant (extinct or extant) and dividing significantly polymorphic coded taxa are highly desirable, thereby justifying the substantial investment of time required to reconstruct individual conceptual whole plants from disarticulated fossil organs.

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). 1 503 504 AMERICAN JOURNAL OF BOTANY [Vol. 88 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, March 2001] LAVIN ET AL.—DALBERGIOID LEGUMES 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, 506 AMERICAN JOURNAL OF BOTANY [Vol. 88 Fig. 2. Bootstrap majority rule (50%) consensus from the analysis of matK/trnK sequences. The dalbergioid clade and its three constituent subclades are indicated. March 2001] LAVIN ET AL.—DALBERGIOID LEGUMES 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. 508 AMERICAN JOURNAL OF BOTANY [Vol. 88 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. March 2001] LAVIN ET AL.—DALBERGIOID LEGUMES 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. 509 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 510 AMERICAN JOURNAL OF BOTANY [Vol. 88 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. March 2001] LAVIN ET AL.—DALBERGIOID LEGUMES 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). 512 AMERICAN JOURNAL OF BOTANY [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- OF BOTANY [Vol. 88 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. 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A contribution to the system of the order Fabales Nakai (Leguminales Jones). Botanical Journal 57: 585–594 (in Russian—English translation at Royal Botanic Gardens, Kew, Richmond, Surrey, UK). Voucher specimen information for the molecular data. 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 528 AMERICAN JOURNAL 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, OF BOTANY [Vol. 88 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 March 2001] LAVIN ET AL.—DALBERGIOID LEGUMES 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). 530 AMERICAN JOURNAL OF BOTANY [Vol. 88 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 March 2001] LAVIN ET AL.—DALBERGIOID LEGUMES 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.

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