The Dalbergioid Legumes (Fabaceae): Delimitation of a Pantropical Monophyletic Clade

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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

3Department of Plant Sciences, Montana State University, Bozeman, Montana 59717 USA;4Tropical Biology Group, Royal Botanic Garden Edinburgh, 20a Inverleith Row, Edinburgh EH3 5LR, UK;

5Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK;6Department of Biological Sciences, University of Dundee, Dundee DD1 4HN, UK; and

7Jardim Botanico do Rio de Janeiro, Rua Pacheco Leao No. 915, Gavea 22.460 Rio de Janeiro—RJ, Brazil

A monophyletic pantropical group of papilionoid legumes, here referred to as the ‘‘dalbergioid’’ legumes, is circumscribed to includeall genera previously referred to the tribes Aeschynomeneae and Adesmieae, the subtribe Bryinae of the Desmodieae, and tribeDalbergieae except Andira, Hymenolobium, Vatairea, and Vataireopsis. This previously undetected group was discovered with phy-logenetic analysis of DNA sequences from the chloroplast trnK (including matK) and trnL introns, and the nuclear ribosomal 5.8Sand 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 geneticallydistinct but poorly, if at all, distinguished by nonmolecular data. Traditionally important taxonomic characters, such as arborescenthabit, 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 thishierarchical level. The discovery of the dalbergioid clade prompted a re-analysis of root nodule structure and the subsequent findingthat 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 unrecognizedmonophyletic group of papilionoid legumes in spite of the ex-tensive taxonomic history of its four constituents: tribes Ades-mieae, Aeschynomeneae, Dalbergieae, and Desmodieae sub-tribe Bryinae. The formal recognition of this group representsa major rearrangement of papilionoid legumes. It combineselements conventionally considered disparate and character-ized as either ‘‘primitive’’ or having varying levels of ‘‘ad-vancement’’ (Fig. 1).

The Dalbergieae originally included tropical trees withfused floral parts and indehiscent pods (Bentham, 1860). Threesubtribes were recognized: Pterocarpeae with samaroid pods,Lonchocarpeae marked by pods having at most small marginalwings, and Geoffroyeae having drupaceous fruits. Polhill(1971, 1981d, 1994) revised this classification by combiningmorphological evidence with that of seed chemistry and wood

1 Manuscript received 11 January 2000; revision accepted 2 June 2000.The authors thank Angela Beyra-M., Alfonso Delgado, Colin Hughes, Jean-

Noel Labat, Gwilym Lewis, Darien Prado, Mats Thulin, and Martin Wojcie-chowski for kindly providing seed or leaf material of many of the speciesanalyzed during this study, Alfonso Delgado, Martin Wojciechowski, and ananonymous reviewer for providing comments that greatly improved the man-uscript, Mats Thulin for making available his observations on the nectary diskin Ormocarpum and close relatives, William Anderson for loaning copies ofthe figures taken from Flora Novo-Galiciana, Sergio Faria for providing un-published information on root nodule morphology, Karin Douthit, ShonaMcInroy, and Maureen Warwick for illustrating the figures, and Tom Turleyfor technical laboratory assistance. This study was supported by a grant fromthe United States National Science Foundation (DEB-9615203), the Lever-hulme Trust, and the Royal Botanic Garden Edinburgh Molecular Phyloge-netic project.

2 Author for reprint requests (e-mail: [email protected]).

anatomy. This new Dalbergieae included 19 tropical woodygenera mainly from Bentham’s Pterocarpeae and Geoffroyeae.Lonchocarpinae were relegated to a closer relationship withother legumes that accumulated nonprotein amino acids inseed (e.g., Evans, Fellows, and Bell, 1985). The revised Dal-bergieae were diagnosed by supposedly plesiomorphic flowermorphologies (i.e., free keel petals, staminal filaments partlyfused and without basal fenestrae), pods with specialized seedchambers, and seeds that accumulated alkaloids or other thannonprotein amino acids. Geesink (1981, 1984) accepted Pol-hill’s circumscription with slight modification, whereas Sousaand de Sousa (1981) proposed a classification similar to Ben-tham’s because Dalbergieae (sensu Polhill, 1981d) supposedlyshared a determinate inflorescence with the Lonchocarpinae.

The Aeschynomeneae (Rudd, 1981a) are one of five tribestraditionally characterized by lomented pods (Polhill, 1981a).Although some Aeschynomeneae lack such pods (e.g., Arach-is, Ormocarpopsis, Diphysa spp., Ormocarpum spp., Pictetiaspp.), none of the members of this tribe have ever been con-fused or classified with the genera of Dalbergieae. Adesmieae(Polhill, 1981f) have a notable history independent of the otherdalbergioid legumes. This is because this tribe combines a pre-sumed plesiomorphic trait, free staminal filaments, with a sup-posedly very derived one, lomented pods. This combinationhas suggested either a taxonomically isolated position or arelationship with other papilionoids also with free stamens(e.g., Burkart, 1952). Bryinae, with lomented pods, possessother 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 notaccumulate nonprotein amino acids and lack a structural mu-

504 [Vol. 88AMERICAN JOURNAL OF BOTANY

Fig. 1. Putative relationships among tribes of the subfamily Papilionoideae according to Polhill (1981a). Tribes underscored include genera that are nowknown to be members of the dalbergioid clade (e.g., Desmodieae then included subtribe Bryinae, and Robinieae the genus Diphysa). Accumulation of nonproteinamino acids and fusion of floral parts occur frequently in Tephrosieae and all tribes positioned above it. The absence of such traits is traditionally viewed asprimitive and is most frequent in tribes positioned below Tephrosieae.

tation in the chloroplast rpl2 locus (Bailey et al., 1997). Bothare atypical of the rest of Desmodieae.

In spite of a taxonomic history of Dalbergieae that has beenseparate from those of Aeschynomeneae, Adesmieae, andBryinae, we present evidence that they collectively form amonophyletic group. The focus on these putatively disparatetaxa was motivated by the taxonomic distribution of the dis-tinctive aeschynomenoid root nodule (Corby, 1981; Faria etal., 1994) and four cladistic analyses: three involving non-molecular data (Lavin, 1987; Chappill, 1995; Beyra-M. andLavin, 1999), and one with rbcL sequence data (Doyle et al.,1997). We have expanded on these previous analyses by sam-pling exhaustively to reveal the exact constituents of the dal-bergioid clade and enumerate the nonmolecular characters thathave been used in the conventional tribal classification of theselegumes. As such, we demonstrate where molecular and non-molecular data are taxonomically concordant. We also showthat many traditionally important taxonomic characters in thisgroup are more homoplasious than previously considered. Be-cause taxon sampling has focused on just the putative mem-bers of the dalbergioid clade, a point to be briefly addressedhere but more thoroughly developed elsewhere is the higherlevel 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 PurificationKit; Qiagen, Santa Clarita, California, USA). DNA sequences analyzed werethe nuclear ribosomal 5.8S and flanking internal transcribed spacers (ITS1and 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 formatK and flanking trnK intron sequences are described in Lavin et al. (2000).Primers for the trnL intron are described by Taberlet et al. (1991). DNAsequencing was performed on an automated sequencer at the Iowa State Uni-versity DNA Sequencing Facility (Ames, Iowa, USA) and Davis Sequencing(Davis, California, USA).

DNA sequences were aligned manually with Se-Al (Rambaut, 1996). Biasintroduced 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 stepmatrix (cf. Cunningham, 1997) was invoked or not, and gaps were treated asmissing data, a fifth state, or as separate characters. Each of the differentsensitivity analyses were subjected to the same heuristic search options. Miss-ing 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, tree-bisection-reconnection branch swapping, and steepest descent. A maximumof 10 000 trees was allowed to accumulate, which is sufficient to capture alltopological variation (cf. Sanderson and Doyle, 1993). Clade stability testsinvolved bootstrap resampling (Felsenstein, 1985; Sanderson, 1995), whereeach of the 10 000 bootstrap replicates was subjected to heuristic search op-tions that included one random-addition sequence per replicate, swapping withtree-bisection-reconnection, and invoking neither steepest descent nor mul-pars.

Taxon sampling—Sampling of molecular and nonmolecular data was asexhaustive as possible at the generic level in order to determine membershipin the dalbergioid clade, as well as the principal phylogenetic structure withinthis clade. Molecular and nonmolecular data were obtained for at least onespecies from every genus ever placed in the Dalbergieae (Burkart, 1952; Pol-hill, 1981d), Aeschynomeneae (Rudd, 1981a), Adesmieae (Polhill, 1981f), orBryinae (Ohashi, Polhill, and Schubert, 1981). The only exception is the pre-sumably extinct genus Peltiera (Labat and Du Puy, 1997), where no success-ful PCR amplifications were obtained from the few available DNAs. In ad-dition to the advantages of being able to detail the taxonomic implications,

March 2001] 505LAVIN ET AL.—DALBERGIOID LEGUMES

exhaustive sampling for molecular data increases the probability of subdivid-ing long branches (e.g., Hillis, 1998).

Our original intent was to sample the same DNA accessions for each ofthe data sets. This proved impossible for DNA sequences because of incon-sistencies in DNA quality and quantity and PCR amplification. We conse-quently had to resort to multiple methods of sampling. The DNA sequencedata were sampled using the exemplar approach. Multiple species per terminaltaxon were sampled where possible (Appendix A). Because nonmolecular dataare generally open to visual inspection across all species of a particular ter-minal taxon, the ‘‘democratic’’ method of sampling (Bininda-Emonds, Bryant,and Russell, 1998) was used for nonmolecular data. In this approach, weincluded all possible character states represented by any one terminal, whichwas usually a traditionally recognized genus (i.e., multistate terminal taxawere coded). The reasoning is that in the evaluation of traditionally importanttaxonomic characters, the degree of polymorphisms within terminals shouldbe explicitly enumerated. For those few terminals in which species-level phy-logenetic analysis has been completed (e.g., Andira and Pictetia), we em-ployed the ancestral method of sampling nonmolecular data (Bininda-Emonds,Bryant, and Russell, 1998). The justification for ultimately combining datathat have been sampled differently is that a combined analysis should stillallow us to best estimate where the traditionally important taxonomic char-acters lie on the continuum from strongly phylogenetically constrained tomaximally homoplasious.

The genera Bergeronia, Dalbergiella, Lonchocarpus, and Muellera havebeen placed in the tribe Dalbergieae (e.g., Burkart, 1952; Geesink, 1981) andPongamiopsis has been synonymized with the genus Aeschynomene (Hutch-inson, 1964). However, they were not included in this analysis because otherphylogenetic analyses (Lavin et al., 1998; Hu et al., 2000) have shown thesegenera to be closely related to Millettia and relatives, all of which accumulatenonprotein amino acids in seed. Similarly, Poecilanthe and Cyclolobiumshould be allied with more basal Papilionoideae that accumulate alkaloids inseed (Greinwald et al., 1995; Lavin et al., 1998; Hu et al., 2000). This is thereason that Poecilanthe is retained as a designated outgroup.

Outgroups were sampled extensively as part of large-scale molecular phy-logenetic studies of the subfamily Papilionoideae (Hu et al., 2000; Penningtonet al., in press; M. Wojciechowski et al., unpublished data). Sampling out-groups 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 todalbergioids. In the end, the outgroups retained in this analysis included Acos-mium and Myrospermum (tribe Sophoreae; Polhill, 1981b), Dipteryx and Pter-odon (Dipterygeae; Polhill, 1981c), Poecilanthe (variously classified; see Lav-in and Sousa, 1995), and Apoplanesia, Amorpha, Eysenhardtia, and Marina(tribe Amorpheae; Barneby, 1977; Polhill, 1981e). This sampling was consid-ered sufficient to demonstrate membership in the dalbergioid clade. The find-ings reported here did not change with a more extensive sampling of out-groups.

Sampling for the molecular data was re-evaluated as aligned DNA sequenc-es accumulated. It became obvious that the matK/trnK sequences were by farthe most informative at higher taxonomic levels, as seen in increased reso-lution in the strict consensus and higher bootstrap values. The primary effortthen changed to sample as exhaustively as possible matK/trnK sequences and,secondarily, the ITS/5.8S and trnL intron sequences. Thus, the data analysisof this study centers on the matK/trnK data set. Sampling of ITS/5.8S se-quences was guided by species level analyses of certain dalbergioid genera(e.g., Beyra-M. and Lavin, 1999; Lavin et al., 2000). Sampling of the trnLintron data was guided by a phylogenetic analysis of putatively basal Papi-lionoideae (Pennington et al., in press). Unevenness in sampling was exac-erbated by inconsistencies in PCR amplifications (mentioned above). A com-bined molecular analysis was not attempted because unevenness in samplingwould result in a combined data set not exhaustively sampled at the genuslevel. Thus, consensus among the data sets was evaluated by congruence ofthe major clades resolved with high bootstrap values (cf. Huelsenbeck, Bull,and Cunningham, 1996).

Nonmolecular character analysis—A nonmolecular data set was devel-

oped from that in Beyra-M. and Lavin (1999) and is presented in AppendixB. Characters that have been considered traditionally important in the tax-onomy of Dalbergieae, Aeschynomeneae, Adesmieae, and Bryinae (e.g., Bur-kart, 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 Ser-vedio, 1997), in spite of the recommendation of Nixon and Davis (1991).Although this can underestimate the degree of homoplasy (see individual char-acter discussions in Appendix B), splitting polymorphic terminals into two ormore monomorphic ones does not change our findings (e.g., as evaluated inthe fashion of a sensitivity analysis). This is because the focus is strictly atwide-scale relationships of groups of genera, and the potentially problematicpolymorphisms are at a different level, within genera. Polymorphisms arediscussed in the presentation of characters or ingroup terminal taxa (Appen-dices B and C). Inapplicable character states in certain terminals (e.g., leaftraits of Ramorinoa, a genus that doesn’t produce leaves) were variously treat-ed as a missing state, an uncertain state, or an extra state (as in a sensitivityanalysis). The nonmolecular data were gathered primarily from field obser-vations or herbarium specimens. Literature reports were usually verified byobservations of the plants.

RESULTS

Parsimony analysis of the 1266 informative sites from the95 taxa by 2966 sites matK/trnK data set produced 10 000 trees(the set maximum) each with a minimal length of 4352, aconsistency index of 0.570 and a retention index of 0.830. Themonophyly of the dalbergioid clade, including all genera ofAeschynomeneae, Adesmieae, Bryinae, and most Dalbergieae,was very well supported by bootstrap analysis (Fig. 2). Fourmembers of tribe Dalbergieae (Andira, Hymenolobium, Vatai-rea, and Vataireopsis) and two sampled genera of Dipterygeae(Dipteryx and Pterodon) were not included. Indeed, the sistergroup to the dalbergioid clade includes genera sampled fromthe tribe Amorpheae (Apoplanesia and Amorpha). Within thedalbergioid clade, there are three well-supported subcladesmarked as the Adesmia, Dalbergia, and Pterocarpus clades(Fig. 2). The earliest branching Adesmia clade includes thegenus Adesmia (sole member of the tribe Adesmieae) andmostly herbaceous to subshrubby genera of the tribe Aeschy-nomeneae (Poiretia, Amicia, Zornia, Chaetocalyx, and Nis-solia). The remaining two subclades each include members ofthe Aeschynomeneae and Dalbergieae. The Pterocarpus cladeadditionally includes two genera, Brya and Cranocarpus, ofDesmodieae (subtribe Bryinae).

For the 481 informative sites from the 118 taxa by 719 sitesITS/5.8S data set, 120 trees were generated each with a min-imal length of 5009, a consistency index of 0.259, and a re-tention index of 0.714. The same higher level relationshipsdescribed for the matK/trnK analysis were resolved in thisanalysis, though with less bootstrap support (Fig. 3). Althoughthe Pterocarpus clade was resolved in the strict consensus ofthe parsimony analysis, it was resolved in less than 50% ofthe analyses of the bootstrap replicates. In no case (majority-rule bootstrap consensus or strict consensus of minimal lengthtrees) was the sister-group relationship of the Amorpheae sam-ples resolved.

Analysis of the 293 informative sites from the 93 taxa by737 sites trnL intron data set generated 10 000 trees each witha minimal length of 1102, a consistency index of 0.603, anda retention index of 0.804. Although the dalbergioid clade iswell resolved by bootstrap analysis, only the Adesmia cladeis further resolved (Fig. 4). Not in any case was the Dalbergiaor Pterocarpus clades resolved as monophyletic. Regardless,


Fig. 2. Bootstrap majority rule (50%) consensus from the analysis of matK/trnK sequences. The dalbergioid clade and its three constituent subclades areindicated.


Fig. 3. Bootstrap majority rule (50%) consensus from the analysis of ITS/5.8S sequences. The dalbergioid clade and two of its three constituent subcladesare indicated. The clade marked by a closed circle was also detected in the analysis of matK/trnK and trnL intron sequences.


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.


the relationships resolved by majority-rule bootstrap consensusdid not conflict with those similarly resolved in either thematK/trnK and ITS/5.8S analyses.

Analysis of the 55 nonmolecular characters (Appendix B)yielded poorly resolved and supported relationships, such thatthe majority-rule bootstrap consensus was largely unresolvedabove the genus level. Resolved intergeneric relationships in-clude a clade with Aeschynomene, Cyclocarpa, Bryaspis, Geis-saspis, Humularia, Kotschya, Smithia, and Soemeringia (60%bootstrap support), one with Chapmannia, Arachis, and Sty-losanthes (65%), Brya and Cranocarpus (67%), Chaetocalyxand Nissolia (100%), Amicia, Poiretia, and Zornia (67%), andOrmocarpopsis and Peltiera (93%). Because Peltiera is notrepresented by DNA sequence data, this nonmolecular dataprovide the only evidence for its relationships (the relation-ships of Peltiera are a focus of another study; M. Thulin andM. Lavin, unpublished data). The only well-supported cladethat was resolved during this analysis and that was not seenduring the previous molecular analyses was one with Etaballiaand Inocarpus (80%), apomorphically diagnosed as havingnearly regular flowers (characters 22–23 in Appendix B).

Because of the poorly resolved relationships obtained fromanalysis of the nonmolecular data set, it was combined withthe matK/trnK data set in order to explore the evolution of thetraditionally important taxonomic characters. Integration withjust the matK/trnK is justified by how well this data set canresolve relationships (discussed in MATERIALS ANDMETHODS) and because of noncompatibility of moleculardata sets with respect to sampling. Parsimony analysis of the1319 informative characters of the combined matK/trnK andnonmolecular data set (95 taxa by 3021 characters) produced2340 trees with a minimal length of 4664, each with a con-sistency index of 0.551 and a retention index of 0.821. Theresulting relationships are essentially those described previ-ously for the analysis of just the matK/trnK data set (Fig. 5).

Sensitivity analysis—Making different assumptions aboutthe molecular data sets, deleting characters with many missingentries (e.g., nonmolecular characters 50–54), splitting poly-morphic terminals into two or more monomorphic ones, orrecoding inapplicable nonmolecular characters to uncertainstates, missing data, or as an extra state, did not affect theresults described above (Figs. 2–5). The monophyly of thedalbergioid legumes was consistently resolved, as generallywas the monophyly of the three constituent subclades. Therewere no cases of clades with bootstrap values over 70% thatconflicted among the molecular data sets. Also, clades withhigh bootstrap values (i.e., .90%) in individual analyses ofthe matK/trnK, ITS/5.8S, trnL intron, or combined nonmolec-ular and matK/trnK data sets were consistently resolved re-gardless of the assumptions made about any one of the partic-ular data sets. This is exemplified by analysis of just the matKcoding region (i.e., excluding the flanking noncoding portionof the trnK intron), where some accessions in the data matrixwere missing either the 59 or 39 half of this locus (for a totalof 12.1% missing entries). The strict consensus of the parsi-mony analysis of the matK locus was essentially identical tothat of the analysis of the matK/trnK data set. Bootstrap anal-ysis resulted in values that were sometimes lower than in theanalysis of the entire matK/trnK data set: 80% for the Amor-pheae 1 dalbergioid clade, 100% for the dalbergioid clade,100% for the Adesmia clade, 94% for the Dalbergia clade, and71% for the Pterocarpus clade.

DISCUSSION

As now circumscribed, the dalbergioids comprise 44 genera(Appendix C) and ;1100 species of trees, shrubs, and peren-nial to annual herbs. Included are economically importanthardwoods (e.g., Dalbergia and Pterocarpus spp.), forage le-gumes (Stylosanthes spp.), and crops (e.g., Arachis spp.). Likemost pantropical legume taxa, the dalbergioids are concen-trated in the neotropics and subSaharan Africa. Although theposition of the dalbergioid clade within the Fabaceae is notfully developed here, its sister group is the tribe Amorpheae,which contains eight New World genera confined mostly towarm temperate and tropical North America. What is generallycertain of higher level relationships is that the dalbergioids aredistantly related to papilionoids that accumulate nonproteinamino acids in seed. This most notably includes Lonchocar-pus, Derris, Millettia, and Hologalegina (e.g., tribes Robi-nieae, Galegeae, etc.; Wojciechowski, Sanderson, and Hu,1999), which at times have been taxonomically confused withvarious elements now included in the dalbergioid clade.

Implications for traditional classifications—The classifi-cation of certain genera into tribes and subtribes of Papilion-oideae (e.g., Rudd, 1981a; Ohashi, Polhill, and Schubert,1981; Ohashi, 1999; Polhill, 1981a, d) needs to be greatlymodified in light of the evidence presented here. The generaBrya and Cranocarpus (subtribe Bryinae of tribe Desmodieae)share many unusual synapomorphies, such as periporate pollenand glochidiate trichomes, that have served to obscure higherlevel relationships. The explosive pollen presentation mecha-nism that Brya shares in common with Desmodieae is shownto have evolved independently. So have the lomented pods thatBrya and Cranocarpus share with Desmodieae.

Four of the five subtribes of Aeschynomeneae are eithermonotypic (e.g., Discolobiinae) or are polyphyletic. Aeschy-nomeneae subtribe Ormocarpinae includes three different el-ements: Diphysa, Ormocarpum, Ormocarpopsis (and Pelti-era), and Pictetia form one lineage in the Dalbergia clade,Fiebrigiella is in the Pterocarpus clade, and Chaetocalyx andNissolia are part of the Adesmia clade. The pod valves withdistinctive parallel venation that previously allied all of thesegenera now are considered to have evolved on three separateoccasions. Indeed, this derived pod trait is homologous amongFiebrigiella, Chapmannia, Arachis, and Stylosanthes.

Aeschynomeneae subtribe Poiretiinae includes two differentelements. Amicia, Poiretia, and Zornia form a monophyleticgroup within the Adesmia clade, and Weberbauerella is phy-logenetically isolated within the Dalbergia clade. The markedpustular glands of Weberbauerella are no longer consideredhomologous to those of Amicia, Poiretia, and Zornia. In therecent classification of Japanese legumes (Ohashi, 1999), Poir-etia and Zornia are classified as the sole members of the tribePoiretieae, a taxonomy that finds no support in this analysis.

Aeschynomeneae subtribe Aeschynomeninae includes eightgenera (Aeschynomene, Cyclocarpa, Soemmeringia, Kotschya,Smithia, Humularia, Bryaspis, and Geissaspis) that form avery well-supported monophyletic group. A nonmolecularcharacter supporting this relationship is the medifixed stipule,although it is not universal in this clade and has evolved in-dependently in Zornia. An extrapolation from our small sam-ple, however, suggests that species of Aeschynomene havingbasifixed stipules (e.g., A. fascicularis and A. purpusii) aremore closely related to Machaerium and Dalbergia than they


Fig. 5. Bootstrap majority rule (50%) consensus from the analysis of combined nonmolecular and matK/trnK sequence data. The dalbergioid clade and itsthree constituent subclades are indicated.


are to the species of Aeschynomene with medifixed stipules.Thus, the subtribe Aeschynomeninae includes two disparateelements.

Only Aeschynomeneae subtribe Stylosanthinae, with Arach-is, Stylosanthes, and Chapmannia (and the segregates Pache-coa and Arthrocarpum), has been long recognized as a distincttaxonomic group and is also revealed as monophyletic in thisanalysis. The well-known nonmolecular character supportingthe monophyly of this clade is a sessile papilionoid flower witha long hypanthium. However, these three genera are veryclosely related to Fiebrigiella and Fissicalyx and together allof these genera are set apart from other members of the Pter-ocarpus clade by large genetic distances. Notably, nonmolec-ular characters do not support most of the relationships in thisclade that are so well supported by independent moleculardata. For example, there are no known nonmolecular data thatsupport the monophyly of the genus Chapmannia (Thulin,2000) or the relationship of Fissicalyx and Fiebrigiella.

The tribe Dalbergieae also is not monophyletic. Excludedfrom the dalbergioid clade are Andira, with 30 species largelyconfined to the neotropics and with one species distributed inthe neotropics and tropical Africa (Lima, 1990; Pennington,1996; Pennington, Aymard, and Cuello, 1997), Hymenolobiumwith 10–15 species in tropical South America and one speciesin Central America (Polhill, 1981d; Lima, 1982a, 1990), Va-tairea with seven species from Mexico to Brazil (Lima, 1982b,1990), and Vataireopsis with three species in Brazil and theGuianas (Polhill, 1981d; Lima, 1990). The distinction of thesefour genera from others traditionally included in the tribe Dal-bergieae has been noted with wood anatomy (Baretta-Kuipers,1981) and estimates of overall similarity (Lima, 1990). Forexample, the wood of Andira, Hymenolobium, Vatairea, andVataireopsis lacks the storied structure and uniseriate rays thatare characteristic of dalbergioid wood and is generally of lesscommercial value.

The remaining genera of the tribe Dalbergieae belong toeither the Dalbergia or Pterocarpus clades. Only Dalbergia andMachaerium are part of the Dalbergia clade, where they aremost closely related to Aeschynomene species that have basi-fixed stipules. The rest of the genera previously classified inthe tribe Dalbergieae form the bulk of the Pterocarpus cladealong with some genera previously classified in the tribe Aes-chynomeneae (e.g., Fiebrigiella, Chapmannia, Arachis, Sty-losanthes, and Discolobium) and subtribe Bryinae of Desmo-dieae.

The genera of Dipterygeae (Taralea, Dipteryx, and Ptero-don; Polhill, 1981c) are not part of the dalbergioid clade. Bur-kart (1952) originally included Dipteryx (then Coumarouna)in the tribe Dalbergieae, and a phylogenetic analysis of non-molecular data by Beyra-M. and Lavin (1999) suggested Dip-terygeae was part of the dalbergioid clade. Even the combi-nation of paripinnate leaves bearing glandular punctae isknown only from Dipterygeae and the dalbergioid legumes.However, this analysis strongly suggests that the punctateglands are plesiomorphic because they are found in all generaincluded in this analysis. Paripinnate leaves evolved indepen-dently among Dipterygeae and various elements in the dal-bergioid clade.

Phylogenetic information among the various nonmolecularcharacters—While the matK/trnK phylogeny was not greatlyinfluenced by the addition of the 55 nonmolecular characters(compare Figs. 2 and 5), there is some phylogenetic infor-

mation in the nonmolecular characters, as evinced by high re-tention indices (Table 1). The consistency (CI) and retention(RI) indices for each of the 55 nonmolecular characters (Ap-pendix B) in the combined analysis were compared to thesame values obtained when each of the nonmolecular char-acters was mapped onto the matK/trnK phylogeny. In the com-bined analysis, the average CI and RI were 0.427 and 0.672,respectively. When mapped onto the matK/trnK trees, the av-erage CI and RI were 0.390 and 0.627, respectively. Regard-less of the small but significant differences (for RI, two-tailedt test, t 5 2.94, P 5 0.005, df 5 52), no character had ahigher consistency or retention index when mapped onto thematK/trnK phylogeny as when combined with the matK/trnKsequence data during parsimony analysis. This suggests thatmapping a few selected nonmolecular characters onto a mo-lecular phylogeny may involve a bias of excess levels of ho-moplasy.

Different classes of characters (e.g., vegetative, floral, andfruiting) were equally as prone to having homoplasy overes-timated when mapped onto a molecular phylogeny. These in-clude, for example, an asymmetric leaflet base (character 9 inAppendix B), persistent floral bracts (character 16), and a longpod stipe (character 33). The states of the leaflet base had anaverage retention index of 1.000 in the combined analysis and0.500 when mapped to the matK/trnK trees (Table 1). Thecorresponding values were 0.667 and 0.167 for the states ofthe floral bracts, and 0.647 and 0.559 for the pod stipe (Table1). Also, no particular class of characters (e.g., vegetative, flo-ral, and fruiting) was more informative than another. For veg-etative characters (1–13, 44–45, 50–55), the average retentionindex is 0.705. For floral characters (14–30, 46–49), it is0.604. For fruiting characters (31–43), the average retentionindex is 0.726. These differences are not significant (single-factor ANOVA, F 5 1.174, P 5 0.317, df 5 52). The lackof a difference in behavior among the various classes of char-acters, as also generally found by Bateman and Simpson(1998) for vascular plants, weakens the suggestion of Tuckerand Douglas (1994) that floral characters necessarily providethe best taxonomic information in Leguminosae. These find-ings also weaken the implication that pod morphology is proneto higher rates of convergent evolution than other types ofcharacters (e.g., Geesink, 1984; Hu et al., 2000).

Conventional taxonomic evidence—Some traditionally im-portant taxonomic characters are determined in this analysis tobe more homoplasious than previously considered. This is es-pecially true of the character states of growth habit, staminalfusion, and pod segmentation. Herbaceous and woody relativesgenerally are separated into different taxonomic groups whena temperate vs. tropical distinction correlates with habit (Judd,Sanders, and Donoghue, 1994). This is especially true of pap-ilionoid legumes where tribes have been categorized by habit(e.g., temperate herbaceous vs. tropical woody tribal divisionin Polhill, 1981a, 1994). An herbaceous habit (number 1 inAppendix B) has evolved at least three times in monomorphiccondition but more times than this in polymorphic condition(Table 1). The Adesmia clade contains mostly herbaceous spe-cies, although some species of Adesmia and Poiretia areshrubs. That an herbaceous growth form maps as the ancestralstate in the Adesmia clade stands in contrast to the conven-tional wisdom that woody taxa form basal clades in tropicalPapilionoideae (e.g., Polhill, 1981a; Tucker and Douglas,1994).


TABLE 1. Average lengths (L) and consistency (CI) and retention (RI) indices for each of the 55 nonmolecular characters. These are comparedfor 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 thecombined and mapped character are equal. A ‘‘.’’ signifies a higher CI and RI value for a character in the combined analysis compared towhen mapped. The reverse situation did not occur.

Number

Combined analysis

L CI RI

Mapped

L CI RI

12345

5.51.06.02.08.0

0.4171.0000.3330.5000.125

0.8501.0000.6670.8330.562

.5555

6.01.06.02.08.0

0.3331.0000.3330.5000.125

0.8001.0000.6670.8330.562

6789

10

5.04.08.01.02.0

0.2000.5000.1251.0000.500

0.7890.5000.7671.0000.833

555.5

5.04.08.02.52.0

0.2000.5000.1250.4170.500

0.7890.5000.7670.5000.833

1112131415

9.07.02.02.07.0

0.1110.1431.0000.5000.429

0.7240.6251.0000.6670.778

.555.

10.07.02.02.09.0

0.1000.1431.0000.5000.333

0.6900.6251.0000.6670.667

1617181920

2.010.010.06.04.0

0.5000.2000.2000.8330.500

0.6670.7780.7780.9550.833

.

.

.

.5

3.511.512.0

7.04.0

0.2920.1750.1670.7140.500

0.1670.7360.7220.9090.833

2122232425

4.05.02.08.05.0

0.2500.2000.5000.1250.200

0.8120.2000.0000.3640.429

55555

4.05.02.08.05.0

0.2500.2000.5000.1250.200

0.8120.2000.0000.3640.429

262728

15.02.0

10.0

0.2670.5000.200

0.6330.8570.800

555

15.02.0

10.0

0.2670.5000.200

0.6330.8570.800

2930313233

5.01.09.06.0

13.0

0.2001.0000.2500.1670.077

0.000

0.8250.5830.647

55...

5.01.0

10.07.0

16.0

0.2001.0000.2000.1430.062

0.000

0.8000.5000.559

3435363738

2.51.0

17.02.05.0

0.4171.0000.2350.5000.400

0.912

0.4090.8890.625

.5.55

3.01.0

17.52.05.0

0.3331.0000.2290.5000.400

0.882

0.3870.8890.625

3940414243

1.02.03.0

11.04.0

1.0001.0000.6670.1820.250

1.0001.0000.7500.5000.571

555.5

1.02.03.0

12.04.0

1.0001.0000.6670.1670.250

1.0001.0000.7500.4440.571

4445464748

5.07.01.0

14.53.0

0.2000.1431.0000.0690.333

0.6000.5001.0000.6250.333

.55.5

6.07.01.0

15.03.0

0.1670.1431.0000.0670.333

0.5000.5001.0000.6110.333

4950515253

6.02.04.07.03.0

0.5000.5000.2500.1430.333

0.5710.7500.4000.7390.000

.5555

8.02.04.07.03.0

0.3750.5000.2500.1430.333

0.2860.7500.4000.7390.000

5455MeanSD

1.07.05.43.9

1.0000.2860.4270.301

1.0000.6670.6720.254

.5

2.07.05.84.2

0.5000.2860.3900.279

0.6670.6670.6270.258

Genera containing both woody and herbaceous species alsooccur in the clade containing Aeschynomene sect. Aeschyno-mene, Kotschya, Humularia, and Geissaspis. The same is truefor the clade including Fiebrigiella, Chapmannia, Stylosan-

thes, and Arachis. Fissicalyx and some species of Chapmanniaare woody in a clade dominated by herbaceous to subshrubbyspecies. Representing yet two other clades, species of Ma-chaerium, Dalbergia, Brya, and Cranocarpus vary from trees


Figs. 6–8. Selected nonmolecular characters (scale bar 5 1 cm for all figures). 6. Aeschynomenoid root nodule associated with lateral root (character number55, 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 evidencefrom this analysis that the ability to produce a strongly woodygrowth habit is a good indicator of relationship.

The staminal character number 26 (Appendix B) includesfive states that provide an average length of 15.0 to the mostparsimonious trees. The consistency index of 0.267 and theretention index of 0.633 demonstrate that this character ishomoplasious. Even state zero, free staminal filaments, addeda length of two because this state occurs ancestrally in someof the outgroup genera and represents a reversion in the genusAdesmia. That a legume group with free stamens can evolvethis condition secondarily from a fused condition (e.g., 9 1 1diadelphous) is not surprising. Four species of Pictetia havenearly free staminal filaments in a clade otherwise representedby species with fused filaments (Beyra-M. and Lavin, 1999).Also, Kass and Wink (1995, 1997) have implicitly shown inan unrelated papilionoid group that the evolution of staminalmorphology does not necessarily involve a unique transfor-mation from free filaments into the fused condition. Perhapsrelated to this issue, Klitgaard (1999a) showed that order ofinitiation and loss of stamens are more variable among thedalbergioids than previously appreciated. No doubt, the apriori view that free staminal filaments represent necessarily aplesiomorphic condition among papilionoid legumes will haveto be abandoned.

All papilionoid legumes with lomented pods were at onetime classified together, although more recently five tribes(Adesmieae, Aeschynomeneae, Coronilleae, Desmodieae, andHedysareae) were thought to have gained this pod type inde-pendently (Polhill, 1981a). We scored three states pertainingto articulation of pod segments (number 31 in Appendix B),which added an average length of 9.0 to the most parsimoni-ous trees. The consistency index of 0.250 and a retention indexof 0.825 suggest that, although homoplasious, this characterprovided phylogenetic resolution towards the tips of the tree.The Adesmia clade is uniform for lomented pods, but the Dal-bergia and Pterocarpus clades are variable, with a minimum

of three separate origins of this pod type in each of theseclades. What was thought to be two separate origins of lo-mented pods in Adesmieae and Aeschynomeneae is now con-sidered at least six origins combined with at least two rever-sals, and not counting polymorphisms.

New taxonomic evidence—In contrast to the above, a fewpreviously overlooked characters are shown by analysis ofcombined molecular and nonmolecular data to be taxonomi-cally informative. Short shoots (character 2 in Appendix B)evolved only once in the clade containing Pictetia, Ormocar-pum, and Ormocarpopsis (also Peltiera). However, the supportfor this clade is moderate (Fig. 5), both in this analysis, andin 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 Aeschynomenesect. Aeschynomene, Smithia, Kotschya, Humularia, Cyclocar-pa, Soemmeringia, Bryaspis, and Geissaspis. In contrast toshort shoots, this calyx morphology marks a very well-sup-ported clade (Fig. 5). The other nonmolecular characters witha high retention index (Table 1), however, either mark smallclades (e.g., characters 13 and 46 and the clade with Brya andCranocarpus), or have homoplasy that was underestimated be-cause of scoring polymorphic taxa (e.g., see characters 39 and40 in Appendix B).

The aeschynomenoid root nodule (Fig. 6, character 55 inAppendix B) is the most notable nonmolecular character inthat it is inferred to be a synapomorphy for the dalbergioidclade. The idea that nodule morphology could be a usefulcharacter in legume taxonomy was pioneered by Corby(1981). He described a number of shapes, named according tothe genus from which he had most observations. The aeschy-nomenoid type has as its main feature a small oblate nodule(transverse diameter greater than axial) with determinategrowth. Corby noted that aeschynomenoid nodules are oftenassociated with fine rootlets, but his otherwise excellent draw-ings omitted these ‘‘for clarity.’’ Such nodules were found pri-


marily in the tribes Adesmieae, and Aeschynomeneae, but alsoin some members of the Abreae, Dalbergieae, Phaseoleae, andRobinieae (Corby, 1988). On his retirement, Corby kindlygave the Sprent laboratory his collection of preserved nodules.These were used, together with new material, for more detailedstructural studies. As a result, the definition of an aeschyno-menoid 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 centralinfected tissue contains few or no uninfected cells. Differen-tiated infection threads are not involved in the process of in-fection, which (where studied in detail) takes place at the lat-eral root junction (Sprent, Sutherland, and Faria, 1989). Allnodules of the tribe Aeschynomeneae that have been examinedconform to this description, together with ten genera of theDalbergieae: Centrolobium, Dalbergia, Etaballia, Geoffroea,Machaerium, Platymiscium, Platypodium, Riedeliella, Tipu-ana, and Pterocarpus (two Brazilian species, P. rohrii and P.santalinoides are not known to nodulate). The evidence forAdesmia, Brya, and Cranocarpus, although slightly less de-tailed, is entirely consistent with the revised description ofaeschynomenoid nodules.

Members of the Dalbergieae that have been omitted fromthe revised clade on morphological and molecular groundswould also be omitted on grounds of nodule structure (Andiraand Hymenolobium) or absence of nodules (Vatairea and Va-taireopsis; Sprent, Sutherland, and Faria, 1989). Two generaof the dalbergioid clade that do not nodulate are Chaetocalyxand Nissolia (Faria and Lima, 1998). Both of these are lianes.Notably, a group of species in Acacia with a semiscandenthabitat cannot nodulate (Harrier et al., 1997). These acaciashave retained some of the characters associated with nodula-tion, such as some of the nod genes, and the ability to stim-ulate rhizobial attachment to roots. It was thus suggested thatthey may have lost the ability to nodulate because, living onthe forest margins, they were not nitrogen limited (Harrier,1995). It would be interesting to carry out similar tests onChaetocalyx and Nissolia as one of their principal habitats isforest margins.

It is now generally agreed that nodulation in legumes mayhave evolved more than once (Sprent, 1994; Soltis et al.,1995). One of these nodulation events involved an infectionprocess through a wound, such as where a lateral or an ad-ventitious root emerges. Compared with the more familiar roothair infection pathway (see Sprent and Sprent, 1990 for de-tails), this pathway is simpler, involving less complex recog-nition systems. Apart from some species of the mimosoid ge-nus Neptunia (James et al., 1992), this wound infection path-way is associated with only aeschynomenoid nodules. In Nep-tunia, however, nodule processes subsequent to infectioninvolve production of infection threads and development of anindeterminate nodule.

Our phylogenetic results are in agreement with molecularand biochemical evidence that nodule structure and infectionsite are largely plant determined (e.g., Gualtieri and Bisseling,2000). Given a phylogenetic lineage, nodule morphology andinfection processes are generally the same regardless of whichspecies or genus of rhizobia is involved (six genera of bacterianodulating legumes are now recognized, and they are collec-tively known as rhizobia). Another general inference is derivedfrom the observation that all species of the genus Aeschyno-mene that have stem nodules are nodulated by photosyntheticrhizobia (Molouba et al., 1999). Given that the aeschynome-

noid root nodule has an unelaborated morphology and infec-tion mode, the ancestral rhizobial form could have been pho-tosynthetic. As legumes moved into drier areas, nodules de-veloped on roots and lost photosynthetic ability (Sprent, 1994).

A phylogenetic classification—The dalbergioid legumes aresimilar to a group of Papilionoideae that includes also Amor-pheae and Dipterygieae. They share a distinctive combinationof a base chromosome number of x 5 10 (Goldblatt, 1981),wood with uniseriate stored rays, vegetative growth with glan-dular punctae, flowers with fused keel petals or staminal fila-ments, and seeds that do not accumulate nonprotein aminoacids (derived from Beyra-M. and Lavin, 1999). The dalber-gioids differ and are apomorphically defined (sensu de Queirozand Gauthier, 1994) as having glandular-based trichomes onvegetative or floral organs, a well-developed abaxial calyxlobe, and the ‘‘aeschynomenoid’’ root nodule. All of thesetraits have been secondarily transformed in some constituentsof the dalbergioid clade (see characters 11, 19, and 55 in Ap-pendix B; also Table 1).

The dalbergioid clade is distinguished more by molecularthan nonmolecular data. It is another legume example of acryptic clade, like ‘‘Neo-Astragalus’’ (Wojciechowski et al.,1993) and the ‘‘temperate herbaceous clade’’ (Sanderson andWojciechowski, 1996). Regardless, it is informally recognizedhere as a distinctive taxonomic group. Furthermore, the threemajor constituent subclades are informally recognized and Ap-pendix C enumerates the 44 current dalbergioid genera ac-cordingly. The three subclades of dalbergioids are:

The Adesmia clade—This includes the genera Adesmia (oftribe Adesmieae; Polhill, 1981f) and Poiretia, Amicia, Zornia,Chaetocalyx, and Nissolia of the tribe Aeschynomeneae. Thisclade is apomorphically defined as having an herbaceousgrowth habit (modified in some descendants—character 1),leaves with few opposite leaflets (evolved in parallel in Ar-achis and close relatives—character 8), and pedicels confluentwith the calyx (modified only in a few species of Nissolia—character 17). A node-based definition (sensu de Queiroz andGauthier, 1994) includes all descendants from the commonancestor of Adesmia and Amicia.

The Dalbergia clade—This includes Dalbergia and Ma-chaerium (of tribe Dalbergieae; de Candolle, 1825; Polhill,1981d), and the following genera of Aeschynomeneae (sensuRudd, 1981a): Aeschynomene (all infrageneric taxa), Soem-meringia, Cyclocarpa, Kotschya, Smithia, Humularia, Bryas-pis, Geissaspis, Weberbauerella, Diphysa, Pictetia, Ormocar-pum, Ormocarpopsis, and Peltiera. This clade is apomorphi-cally defined as having diadelphous staminal filaments split-ting readily or tardily into two flanges, usually in a 5 1 5arrangement (polymorphic with a 9 1 1 diadelphous conditionin many species and occasionally monodelphous in Machaer-ium—character 26), and a persistent staminal flange that insome cases reflexes upward above the developing fruit (char-acter 28). A node-based definition includes all descendantsfrom the common ancestor of Dalbergia and Cyclocarpa.

The Pterocarpus clade—This includes Pterocarpus, Tipu-ana, Platypodium, Reideliella, Centrolobium, Grazieloden-dron, Paramachaerium, Ramorinoa, Inocarpus, Etaballia,Platymiscium, Cascaronia, Fissicalyx, Geoffroea from Dal-bergieae; Brya and Cranocarpus from Desmodieae; and Fie-


brigiella, Chapmannia, Stylosanthes, Arachis, and Discolob-ium from Aeschynomeneae. This clade is apomorphically de-fined as having commonly caducous bracteoles (character 18)and seedlings producing a simplified eophyll (secondarilytransformed in Arachis and close relatives—character 45). Anode-based definition includes all descendants from the com-mon ancestor of Pterocarpus and Riedeliella.

Although data from matK/trnK, trnL, and ITS/5.8S were notcombined in a single analysis, results from individual analysesshowed significant consensus combined with no significantconflict. The combined matK/trnK and nonmolecular analysisyielded very robust results to support the conclusions outlinedabove. This study demonstrates that matK/trnK sequences pro-vide excellent resolution at the broadest phylogenetic levelsdealt with in this study. This same locus, along with ITS/5.8S,gives excellent resolution to within and among closely relatedgenera. In contrast, trnL provides the least resolution. Ulti-mately, this study provides a framework for future studies thatdeal taxonomically with individual dalbergioid genera. Thereis now sufficient data from which to guide the choice of po-tential sister groups or outgroups in such studies.

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APPENDIX B

Nonmolecular characters and character states. All references to clades arethose derived from the combined matK/trnK and nonmolecular analysis (Fig.5). References to ancestral states were inferred with the reconstruct tree optionin PAUP (Swofford, 2000) and the trace option in MacClade (Maddison andMaddison, 1999).

Vegetative characters

1. Habit: 0) woody (trees to shrubs), 1) herbaceous (subshrubs to herbs),2) twining and herbaceous, 3) twining and woody. Predominantly herbaceousgenera sometimes include subshrubby species, whereas woody genera usuallydo not, thus explaining the coding for state number 1. A herbaceous habitarose independently in the following clades: one represented by Fiebrigiellaand Arachis, another by Chaetocalyx and Poiretia, and one by Weberbauer-ella and Kotschya. The twining herbaceous habit is restricted to the Adesmiaclade where it is known from some species of Poiretia (Rudd, 1972c) and allChaetocalyx (Rudd, 1958) and Nissolia (Rudd, 1956). A twining woody habitoccurs in polymorphic condition in the clade with Dalbergia and Machaer-ium.

2. Short shoots: 0) absent or not regularly present and then not covered bypersistent stipules, 1) regularly present and covered by distichously arrangedpersistent stipules from the axils of which are born the inflorescence (Fig. 7).The short shoot condition is restricted to the clade including all descendantsof the most recent common ancestor of Pictetia and Ormocarpopsis. Verysimilar 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 folia-ceous, 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 Aes-chynomene sect. Aeschynomene, Cyclocarpa, Humularia, Geissaspis, Smithia,and another including just Zornia. Lignescent stipules evolved independentlyin polymorphic condition in the liana-forming species of Machaerium, in mostspecies of Brya, and in all species of Pictetia. In Brya, the leaves of the longshoot are entirely transformed into a single spine.

4. Pseudopetiole: 0) absent, 1) present (Fig. 8). A pseudopetiole is tradi-tionally defined as a petiole with stipules attached. It is here described as apulvinus (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 su-perficially appear as if they are adnate to the petiole. The pseudopetioleevolved independently in a clade including just Adesmia, and another includ-ing 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 inpolymorphic condition from Ormocarpum, Aeschynomene (all subgroups) andclosely related genera (Cyclocarpa, etc.). Grooved leaf rachises occur in therest of the genera, except where the leaves are uniformly sessile, as in Bryaand Inocarpus, and this trait is then scored as inapplicable. Otherwise, leafrachises vary continuously between narrowly grooved and distinctly canalic-ulate. The motivation for using this trait is that terete leaf rachises are shownto be derived (but in polymorphic condition) in two clades: that including alldescendents but Pictetia of the most recent common ancestor of Dalbergiaand Ormocarpopsis, and that including most descendants of the recent com-mon ancestor of Platymiscium and Pterocarpus.

6. Distal end of leaf rachis: 0) terminated by a leaflet, 1) not terminated bya leaflet (a mucro is often present). A leaf rachis not terminated by a leafletis found in the large clade including Aeschynomene sect. Aeschynomene, Cy-clocarpa, Humularia, Soemmeringia, Kotschya, Smithia, Geissaspis, andBryaspis. This type of leaf also has evolved independently in the outgroupsamples of Dipterygeae (Dipteryx and Pterodon), the clade including Amicia,Zornia, Adesmia, Arachis, and Poiretia, the clade including just Aeschyno-mene sect. Ochopodium, and the clade including Stylosanthes and Arachis.

7. Number of leaflets per leaf: 0) leaves unifoliolate/simple, 1) leaves tri-to 20-foliolate, 2) leaves more than 20-foliolate. State zero occurs uniformlyin Etaballia, Inocarpus, and Brya, and in polymorphic condition in Crano-carpus. State two is restricted to just the Dalbergia clade where it occursuniformly in Weberbauerella, and predominantly so (i.e., polymorphic) inMachaerium, Dalbergia, and all the sections and series of Aeschynomene(Aeschynomene, Viscidulae, Pleuronerviae, and Scopariae). This state is cap-turing ‘‘fern-like’’ leaves where the leaflets abut laterally, are narrowly ellip-

tic, and have parallel lateral margins. Simple leaves are scattered throughoutbut with most occurrences (usually in polymorphic condition) in the Ptero-carpus clade (Discolobium, Etaballia, Inocarpus, Platypodium, Byra, andCranocarpus).

8. Leaflet arrangement: 0) alternate, 1) opposite. Two large clades haveevolved opposite leaflets independently. One includes Adesmia, Chaetocalyx,Nissolia, Poiretia, Amicia, Zornia, and the other includes Fissicalyx, Fiebri-giella, Chapmannia, Stylosanthes, and Arachis. Opposite leaflets have evolvedsporadically mostly within the Pterocarpus clade (Grazielodendron, Riede-liella, Cranocarpus, Paramachaerium), and rarely in the Dalbergia clade(Smithia). The genera with uniformly simple or unifoliolate leaves (e.g., Eta-ballia, Inocarpus, Brya, and Ramorinoa) were marked inapplicable. The spe-cies of Cranocarpus with imparipinnate leaves have opposite leaflets, and thiscondition is used to represent the genus. A terminal taxon is scored for op-posite leaflets if all constituent species predominate with this condition. Aterminal taxon is scored for a polymorphic condition only if some constituentspecies have uniformly opposite leaflets and others have uniformly alternateleaflets.

9. Leaflet base: 0) symmetric, 1) asymmetric. The asymmetric state is re-stricted to the Dalbergia clade, where it has evolved independently and poly-morphically in Pictetia and Aeschynomene (all subgroups except Scoparia)and uniformly in Humularia, Bryaspis, Geissaspis, Kotschya, and Smithia. Anasymmetric base of the leaflet is correlated with an eccentric midrib and prob-ably related to a nyctinastic leaflet movement that involves a forward twistingand folding of each leaflet. This ‘‘forward-folding’’ type is very similar tothe leaflet movements in legume subfamilies Mimosoideae and Caesalpinioi-deae, as well as the papilionoid genus Sesbania, and it has been observed inspecies 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 independentlyin the clade including Arachis, Stylosanthes (polymorphic in these first two),and Chapmannia (this is the subtribe Stylosanthinae of Rudd, 1981a) and intwo genera endemic to Madagascar, Ormocarpopsis and Peltiera (Labat andDu Puy, 1996, 1997). Reddish tannin deposits usually occur in reticulate pat-terns 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 typeof trichome is a synapomorphy for the dalbergioid group, where it is foundon the stems, leaves, inflorescence, or ovary. Although synapomorphic, theglandular-based trichomes have been secondarily lost several times in each ofthe Adesmia, Dalbergia, and Pterocarpus clades. In addition to most generaof the formally recognized tribe Aeschynomeneae, glandular-based trichomesare found in Centrolobium, Grazielodendron, Ramorinoa, Etaballia, Riede-liella, Fissicalyx, Paramachaerium, Peltiera, and polymorphic in Brya, Cran-ocarpus, Dalbergia, and Machaerium.

12. Pustular glands: 0) absent, 1) present (Fig. 11). The latter condition isthought to be a derivation of the general dalbergioid trait of punctate glands(all members of the ingroup and outgroup possess punctate glands on theleaflets). There has been further development in the size and color of thecommon punctate gland such that they protrude outward from the plane ofthe leaflet, calyx, or ovary and are brownish red to blackish in color. Pustularglands are known from genera outside the dalbergioid clade (Acosmium, My-rospermum, Amorpha, Apoplanesia, Dipteryx, and Pterodon), and haveevolved in four separate instances within the dalbergioid clade (Geoffroea andCascaronia; Poiretia, Amicia, and Zornia; Weberbauerella; and Centrolob-ium).

13. Stipitate glands: 0) absent, 1) present from non-glochidiate trichomes,2) present from microscopically glochidiate trichomes (Fig. 12). Such glan-dular trichomes are usually present on stems or leaves, but can also occur onovaries and pods. Stipitate glands have evolved in the clade including Brya,Cranocarpus, and Grazielodendron (uniquely from glochidiate trichomes inthe first two genera), and in polymorphic condition in the clade includingAdesmia, Chapmannia, and Stylosanthes. In Brya and Cranocarpus, stipitateglands are found, in addition to the foliage, on the ovary where they persistwith the mature fruit. The high tree scores for this character (Table 1) do notaccount for the optimizations of polymorphic codings where Chapmannia,Stylosanthes, and Adesmia were assigned state zero during parsimony analy-sis.

Inflorescence characters

14. Inflorescence position: 0) axillary, 1) terminal. The first state corre-sponds to leafy flowering branches that are indeterminate with vegetativegrowth from the apical meristem. The second refers to leafy flowering branch-


Figs. 9–12. Selected nonmolecular characters. 9. Glandular-based trichome of dalbergioid legumes (character number 11; scale bar 5 200 mm). 10. Baseof trichome where glandular exudate is secreted (scale bar 5 20 mm). 11. Pustular glands on leaflet of Centrolobium (character number 12; scale bar 5 200mm). 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 includingApoplanesia and Amorpha, and in the two species of Geoffroea. Other casesof independent evolution but in polymorphic condition include Reideliella,most outgroup genera, and sporadically throughout the Dalbergia clade (Aes-chynomene subgroups, Kotschya, and Smithia).

15. Inflorescence type: 0) racemose, 1) axillary subumbel, 2) solitary ax-illary flowers, 3) helicoid cymes. Helicoid cymes have evolved several timesbut in all cases within the Dalbergia clade (Dalbergia, Machaerium, Aeschy-nomene, Kotschya, and Smithia). They appear to arise readily from any inflo-rescence condition (i.e., note the polymorphic codings for most of these gen-era). In the axillary subumbel, the internodes of the rachis are telescopeddown almost completely, as in Chaetocalyx and Nissolia. Solitary flowershave evolved independently and uniformly in Brya and the clade with Arachisand Stylosanthes. Notably, polymorphic codings for this character are highlylocalized to the genera in the clade that includes the most recent commonancestor of Dalbergia and Ormocarpopsis.

16. Floral bracts: 0) smaller than the flower or fruit, 1) larger than theflower or fruit. Large floral bracts have evolved independently in Zornia, andthe clade including Bryaspis, Geissaspis, and Humularia. These two statesare markedly discontinuous where the smaller bract is barely visible.

Floral characters17. Pedicels: 0) articulated with the calyx, 1) confluent with calyx, 2) ab-

sent, 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 ismost common among the rest of the dalbergioid clade and could be the an-cestral condition to the Dalbergia and Pterocarpus clades. If so, then a tran-sition to pedicels confluent with the calyx has occurred many times indepen-dently (Reideliella, Ramorinoa, Centrolobium, Brya, Cranocarpus, Weber-bauerella, and Geissaspis, Bryaspis, and Humularia). Sessile flowers haveevolved three times, once in Chapmannia, Arachis, and Stylosanthes (the sub-tribe 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 theflower or with the developing or mature fruit are common to the dalbergioids.Caducous signifies that the bracteoles fall before the flower aborts or beforethe pod begins to form. Caducous bracteoles are highly localized in the cladethat includes all descendants of the most recent common ancestor of Ptero-carpus and Platymiscium. Such bracteoles have also evolved independentlyin Geoffroea, Riedeliella, and several of the outgroups. Bracteoles occur ir-regularly (i.e., mostly singly and variously along the pedicel) or not at all infour separate clades: one including Weberbauerella, another with Humularia,Geissaspis, yet another with Amicia, Poiretia, Zornia, Chaetocalyx, and Nis-solia, and finally in Cascaronia.

19. Calyx lobe fusion: 0) five more or less equally spaced lobes, 1) fiveseparate lobes but with the abaxial one (lower or carinal) the largest andseparate from laterals, 2) a two-lipped calyx with the abaxial lobe fused com-pletely or nearly so to the two lateral lobes, and the upper two lobes com-


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. Char-acter state one is synapomorphic for the dalbergioid clade, and it is mostdistinctive developmentally with the abaxial sepal initiating with the largersize and faster growth rate relative to the other sepals (Klitgaard, 1999a). Themost notable derivation from this condition within the dalbergioids is statetwo, which occurs in the clade with Aeschynomene sect. Aeschynomene, Cy-clocarpa, Soemmeringia, Kotschya, Smithia, Geissaspis, Bryaspis, and Hu-mularia (Aeschynomeneae subtribe Aeschynomeninae of Rudd, 1981a). TheDipteryx type occurs in Dipteryx, Pterodon, and Amicia zygomeris, where theupper two calyx lobes are greatly enlarged, contrasting with the diminutivelower three lobes. The Fissicalyx type evolved only in Fissicalyx, where allthe calyx lobes occur as an upper lip (spathaceous). The Inocarpus type hasthree lips, the lower formed by the carinal lobe, and two lateral lips formedby one lateral and one vexillar lobe. Also, the Socotran species of Chapman-nia have yet another type where the upper lip of a bilabiate calyx comprisesthe two upper and two lateral calyx lobes, and the lower lip comprises justthe carinal lobe. Chapmannia, however, was scored for state one because itrepresents the ancestral state for the genus (see the Chapmannia phylogenyin 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 thebase of the ovary, 1) short-tubular, petals and stamens arising from a rimpositioned 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. Thecalyces of Acosmium, Apoplanesia, Amorpha, and Etaballia have a poorlydeveloped hypanthium (the last genus represents the only reversion to a lossof the hypanthium among dalbergioid legumes). Among the dalbergioids, stateone predominates and is ancestral. State two is confined to the clade includingChapmannia, Arachis, and Stylosanthes.

21. Petal coloration: 0) predominantly whitish to reddish or violet, 1) pre-dominantly yellow. The large majority of dalbergioid legumes have yellowpetals, and this is inferred to be ancestral. Notable exceptions include theclade with Dalbergia and Machaerium (polymorphic), as well as Grazielo-dendron, Paramachaerium, Ormocarpum (polymorphic), and Adesmia (thespecies with solitary axillary flowers) where whitish to violet petals are com-mon.

22. Corolla symmetry: 0) bilateral (papilionoid), 1) nearly radial. State zeropredominates among dalbergioids and indeed most papilionoids. Notably, anearly radially symmetric flower has evolved independently four times: onceeach in Inocarpus, Etaballia, Reideliella, and the clade with the samples ofAmorpheae (Apoplanesia and Amorpha). Nearly radially symmetric flowershave also evolved independently in the four species of Pictetia where statezero is considered ancestral (Beyra-M. and Lavin, 1999).

23. Petal morphology: 0) petals abruptly differentiated into a blade andclaw (Figs. 13–15), 1) petals ligulate, the claw and blade not distinguishable(Fig. 16). State one evolved separately in Etaballia and Inocarpus. This char-acter is not dependent on character number 22 because Amorpheae, Reide-liella, and four species of Pictetia have a nearly radial flower symmetry withpetals differentiated into a blade and claw.

24. Keel petals: 0) free, 1) connate, at least along the carinal margin if notto near the tip. Free keel petals in papilionoid legumes have been the hallmarkof the tribes Swartzieae and Sophoreae. Acosmium and Myrospermum, tra-ditionally placed in the tribe Sophoreae, have free keel petals. Among thedalbergioid legumes, fused keel petals represent the ancestral condition thathas reverted back to the free condition only in Etaballia, Geoffroea, Riede-liella, Platypodium, and Tipuana (all confined to the Pterocarpus clade).

25. Wing petals: 0) smooth, 1) crimped. Crimped wing petals are distinctlymuch broader than the adjacent keel petals. The evolution of such wing petalshas occurred in the clade including Paramachaerium, Pterocarpus, Ramori-noa, Paramachaerium, Tipuana, and Platypodium, and separately in that in-cluding 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 41 1, or 4 1 1 1 4 1 1] with at least two phalanges of fused filaments. Statetwo is the inferred ancestral condition of the dalbergioid legumes. State fouris 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 (bothof the Pterocarpus clade). The free stamens of Adesmia are inferred to rep-resent a reversion from an open monodelphous condition. State one is uncom-mon among dalbergioids and is monomorphic only among some members thePterocarpus clade (Grazielodendron, Geoffroea, and Ramorinoa). The di-adelphous 9 1 1 condition is associated with weakly developed basal fenes-trae (Klitgaard, 1999a).

27. Anther size and attachment: 0) monomorphic, basi- to dorsi-fixed, 1)dimorphic, the smaller anthers usually dorsifixed, the larger basifixed. Char-acter state one evolved independently in the clade with Amicia (polymorphic),Poiretia, and Zornia, and again in that with Arachis and Stylosanthes. Di-morphic anthers also have other sporadic occurrences, such as in Aeschyno-mene genistioides (Aeschynomene sect. Ochopodium; Rudd, 1967, 1972a), themonotypic Aeschynomene subgen. Bakerophyton (Verdcourt, 1971), and in aMesoamerican clade of Platymiscium species (B. Klitgaard, unpublished data).

28. Staminal flange and filaments post-anthesis: 0) readily caducous, not


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 uponcharacter number 27 (see Beyra-M. and Lavin, 1999). Persistent stamens arediagnostic of the Dalbergia clade where the predominant condition is stateone. State two occurs in a clade with Aeschynomene sect. Aeschynomene,Kotschya, etc. Persistent stamens evolved separately in Brya, Cascaronia, andseveral outgroup genera.

Gynoecial characters

29. Locule: 0) encompassing nearly the entire length of the ovary, 1) con-fined to the basal end of the ovary. The locule is situated just above the stipein state one, and a large portion of the distal end is solid. All five occurrencesof this state are inferred to be cases of independent evolution (Vatairea, Va-taireopsis, Tipuana, Centrolobium, and Paramachaerium).

30. Nectary disk: 0) absent; 1) present. A nectary disk surrounding the baseof the ovary is known from Paramachaerium (Rudd, 1981a), most species ofOrmocarpum (M. Thulin and M. Lavin, unpublished data), and occasionallyin Machaerium (Klitgaard, 1999a). The retention index is undefined in thischaracter (Table 1) because Machaerium and Ormocarpum were assigned anancestral 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 forminglate during pod development occur in Ormocarpum, Pictetia, Diphysa, Chae-tocalyx, and Nissolia. Some species in the first of these three genera forminarticulate pods. Only in the Adesmia clade is the lomented condition uni-formly present. Both of the Dalbergia and Pterocarpus clades combine generawith articulate and inarticulate pods.

32. Pod margins: 0) straight, with no marginal constrictions between seeds1) constricted between seeds. State one has evolved numerous times indepen-dently in Discolobium, Fiebrigiella, Brya and Cranocarpus (polymorphic),Amicia, and Giessaspis, Bryaspis, Humularia, Kotschya, and Smithia. Aes-chynomene, Ormocarpum, and Diphysa are distinctively polymorphic for thischaracter.

33. Stipe of mature pod: 0) absent to less than half the length of the calyxtube, 1) surpassing the length of the calyx tube. State one has evolved mostuniformly in the clade with Dalbergia, Machaerium, Diphysa, Ormocarpum,Ormocarpopsis, Peltiera, and Pictetia. Among dalbergioids, state zero pre-dominates only in the Adesmia clade. Otherwise, both states have been gainedand 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 hasevolved once in the clade with Arachis, Stylosanthes, Chapmannia, and Fie-brigiella, (not Fissicalyx, however), and again in the clade with Chaetocalyxand Nissolia. Some species of Diphysa, Ormocarpum, and Pictetia have podswith strong longitudinal nerves, but these genera were optimized during anal-ysis as having state zero.

35. Replum: 0) placental margin disarticulating with the pod valves orarticles, 1) the valves or articles disarticulating separately from the persistentplacental margin. The last state is gained independently in Adesmia sect. Mur-icatae, Cyclocarpa, and in species of Aeschynomene sect. Aeschynomene (e.g.,A. villosa). This character had an undefined retention index (Table 1) becauseAdesmia and Aeschynomene were optimized for state zero.

36. Development of pod wings: 0) not winged, 1) wing from expansion ofthe ovary wall, 2) wing from expansion of the ovary sutures, 3) wing fromattenuation of the distal end of the ovary (i.e., the style), 4) winged fromattenuation of the proximal end of the ovary (i.e., the stipe). This characteris derived from Lima’s (1990) developmental work on samaroid fruits of tribeDalbergieae. He distinguished wings whose area of origin was the ovary walls(state one); wings with an origin of the ovary sutures (state two); and wingswith an origin from the distal end of the ovary (state three). Lima consideredVatairea and Vataireopsis to have a separate state, with wings derived fromthe solid distal portion of the ovary. This reflects the different morphology ofthe gynoecium in these two genera (see state one of character 29). We con-sider that the origin of the wing in these taxa is merely from the distal endof the ovary, and thus they are coded with state three. A further modificationis that we consider the basal wing of Platypodium to be derived from anexpansion of the stipe (state four). Developmental anatomical work couldconfirm these distinctions. State one has evolved independently in both theDalbergia (Dalbergia and Weberbauerella) and Pterocarpus clades (Platym-iscium, Cranocarpus, Grazielodendron, Ramorinoa, Pterocarpus, Fissicalyx,

and Riedeliella). State three has evolved in every occurrence separately (Va-tairea, Vataireopsis, Nissolia, Machaerium (polymorphic), Tipuana, Centro-lobium, and Paramachaerium).

37. Inner epidermis and endocarp: 0) lignescent, 1) spongy and adheringto the mature seeds. State one evolved in the clade with Chaetocalyx andNissolia, and again in that with Pictetia, and perhaps again in Peltiera (thesister genus of Ormocarpopsis—see Labat and Du Puy, 1997).

38. Mesocarp: 0) lignescent, 1) spongy, 2) fleshy. State one arose indepen-dently and uniformly in Pictetia and Fiebrigiella, and sporadically in Or-mocarpum, Chapmannia, Nissolia, and Chaetocalyx. State two occurs in thefleshy vertebrate dispersed fruits of Dipteryx, Andira and Geoffroea, all in-stances of independent evolution. Polymorphic terminals were optimized forstate 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 treescores for this character (Table 1) resulted from state zero being assigned tothe 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 confinedto Discolobium, whereas the lateral coil has evolved in Cyclocarpa and var-ious species of Aeschynomene sect. Aeschynomene and Ormocarpum. Al-though the lateral coil is restricted to members of the Dalbergia clade, stateone was assigned to the polymorphic terminals Aeschynomene and Ormocar-pum 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 valvehave evolved in many separate occasions throughout the dalbergioid legumes,as in Adesmia (polymorphic), Ormocarpum (polymorphic), Brya, and Centro-lobium (where they become spinose). Pod valves with crests or bumps haveevolved once in the clade with Poiretia, Amicia, and Zornia, and again inpolymorphic condition among various species of Aeschynomene sect. Aeschy-nomene. The polymorphic terminals Adesmia, Ormocarpum, and Aeschyno-mene were assigned state zero during parsimony analysis, thus resulting inrelatively 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 thehilum placed toward the end toward the style. The last condition is charac-teristic of most dalbergioids and indeed ancestral to that clade. However, statetwo has evolved independently in Dipterygeae (Dipteryx and Pterodon), Hy-menolobium, 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 trans-verse. Oblique to transverse orientation of seeds is confined to a subcladeincluding Platymiscium, Centrolobium, Paramachaerium, Pterocarpus, Ra-morinoa, and Tipuana (Lima, 1990). Such seeds have also evolved indepen-dently 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 knownfrom Platymiscium, Grazielodendron, Ramorinoa, Centrolobium, and Geof-froea.

45. Number of leaflets in the first eophyll: 0) one, 1) more than one. Dal-bergioids commonly have eophylls that are not strongly differentiated fromthe adult leaves (i.e., multifoliolate). The Pterocarpus clade is exceptional inhaving all known instances where the eophylls are unifoliolate. Data for char-acters 44 and 45 for Ramorinoa came from Burkart (1952, p. 238).

Pollen characters

46. Aperture type: 0) tricolporate, 1) periporate. Tricolporate apertures arethe general and most common type in legumes. Among the dalbergioids, per-iporate pollen is known from only Brya and Cranocarpus.

47. Pollen pore: 0) without an operculum, 1) with an operculum. An oper-culum is a distinctly delimited ectexinous structure that covers the ectoaper-ture, which in the case of all dalbergioids means the colpus. State one hasbeen gained independently many times throughout all three principal cladesof the dalbergioid legumes.

48. Colpi (the polar region): 0) colpi short, not anastomosing at the polesof the pollen grain, the polar region entire, 1) colpi longer, the ends of colpi


anastomosing, forming syncolpi. State one evolved independently and uni-formly in Humularia and Vataireopsis. This state is polymorphic for Aeschy-nomene, Kotschya, and Smithia. Thus, state one is confined to the Dalbergiaclade among the dalbergioid legumes.

49. Wall stratification (Guinet and Ferguson, 1989): 0) well-developed end-exine and footlayer, 1) thickening of the endexine at least at the aperturescombined with a reduction in the foot layer, 2) reduction of the endexinecombined with a thickening of the foot layer, 3) reduction of the endexineand foot layer combined with an elongation of the columellae. State one ismost common among the dalbergioids and is inferred to be ancestral in thisclade. State two evolved independently in Adesmia and the outgroup Myros-permum. 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 generahave narrow short rays (state one). Good examples include Platymiscium,Fissicalyx, and the nondalbergioid Dipteryx. Outgroup genera have storiedrays that are larger than this. Wood characters were scored from examinationof slides in the collections at the Jodrell Laboratory, Kew, and at the ForestProducts Laboratory, Madison, Wisconsin, USA, and by reference to descrip-tions and photographs in Baretta-Kuipers (1981), Gasson (1994, 1999), Miles(1978), and Detienne and Jacquet (1983). Wheeler, Bass, and Gasson (1989)provide thorough definitions for all of the wood characters used in this anal-ysis. Details on the wood anatomy of Chaetocalyx, Nissolia, Poiretia, Amicia,Zornia, Chapmannia, Arachis, Stylosanthes, Soemmeringia, Smithia and Geis-saspis come from Cumbie (1960). Unfortunately, the information is presentedin such a way that only a few character states can be coded. Amorpha fruti-cosa has been illustrated and described by Schweingruber (1990), Adesmiahorrida by Roig (1986), Discolobium by Cozzo (1949, 1950), and Parama-chaerium by Brizicky (1960). No information on the wood anatomy is avail-able for Riedeliella, Cranocarpus, Fiebrigiella, Cyclocarpa, Kotschya, Bryas-pis, 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 intangential longitudinal section. Although considered a very useful anatomicalcharacter, both diagnostically and cladistically, there are many legume generawith storied rays that are irregular, or obvious in short rays and less so intaller rays which may be axially fused. Storied rays are particularly stronglydeveloped in Dipteryx, Pterocarpus, Platymiscium, Grazielodendron, Etabal-lia, Inocarpus, Dalbergia, Machaerium, and Aeschynomene, all of which haveshort rays. The taxa with larger rays often do not exhibit such regular storiedarrangement, and axial fusion is often the cause, as in species of Acosmium.

52. Composition of cells in rays: 0) homocellular, 1) heterocellular. Thisfeature is observed in radial longitudinal section. Homocellular rays are com-posed entirely of procumbent ray cells. Heterocellular rays in legumes arecomposed mainly of procumbent cells, but there are also some square orupright cells, usually in a row or rows at the ray margins (i.e., at the top orbottom of a ray). These two character states are not mutually exclusive. Ju-venile wood often tends to be more heterocellular than mature wood, and itis not always apparent where exactly a wood sample was taken from if thepith in the stem is not included.

53. Crystals in ray cells: 0) absent, 1) present in some ray cells. Prismaticcrystals of calcium oxalate are found in many, if not most legumes. They arealmost ubiquitous in chambered axial parenchyma strands, but in a few generacan also be found in ray cells. The main difficulty with this character is thatif the crystals are rare they can be overlooked. They are searched for in radiallongitudinal section, because they are even more difficult to find in tangentiallongitudinal section.

54. Axial parenchyma: 0) not abundantly aliform and confluent, 1) abun-dantly aliform and confluent. Axial parenchyma patterns in legumes are verydifficult to code. All the legumes in this study have predominantly paratra-cheal parenchyma, with the addition of some apotracheal diffuse parenchymain particularly Dalbergia and Platymiscium. This ranges continuously fromscanty paratracheal, vasicentric, aliform, to confluent. In the opinion of oneof us (Gasson), these all constitute one character. They could each be codedas character states, but virtually all wood samples in the legumes exhibit morethan one condition. Unilaterally paratracheal parenchyma is found in some ofthe taxa, and probably forms part of this continuum. Banded parenchyma,which could be treated separately, may be an extreme form of confluent pa-renchyma, particularly if the bands are several cells wide. Some taxa in thestudy group do have narrow bands, but they are not distinguished here. Thechoice of the two character states above serves to separate four of the genera

very well, but does not distinguish all the other complicated variations on theparatracheal theme exhibited by the taxa coded as zero. Aeschynomene is verydifferent, in that it has such abundant parenchyma, that the fibers exhibit awinged-aliform appearance.

Nitrogen fixation character

55. Root nodule: 0) none produced, 1) produced as a non-aeschynomenoidnodule, 2) produced as an aeschynomenoid nodule (Fig. 6; Corby, 1981; seediscussion). State two is synapomorphic for the dalbergioid clade, althoughsome genera in this clade are known not to produce nodules at all (i.e., Chae-tocalyx and Nissolia), as is the case for some nondalbergioids (e.g., Myros-permum, Dipteryx, Pterodon, Vatairea, and Vataireopsis). Some outgroupsproduce nodules of a type other than aeschynomenoid (e.g., Acosmium, Poe-cilanthe, Andira, Hymenolobium, and Amorpha). Although lost within thedalbergioid clade (Table 1), the aeschynomenoid root nodule is not encoun-tered elsewhere in the legume family. The stem (but not root) nodules ofSesbania rostrata (tribe Robinieae) are superficially similar, but they differfrom the aeschynomenoid type in having an apical meristem, albeit ephemeral(J. Sprent, unpublished data). Three dalbergioid genera, Cyclocarpum, Geis-saspis, and Paramachaerium, are known to produce nodules, but the exacttype is unknown. These genera were variously coded as having missing dataor state one. Such alternative coding did not affect how these genera wererelated with respect to the three major subclades of the dalbergioid clade.Future studies incorporating nodule morphology in a phylogenetic analysiswill do well to recognize specific morphologies independent of nodule cate-gories. Specifically, these would include characters such as the apical meri-stem (absent vs. present), infection site (associated with emergent rootlet ornot), infection threads (absent vs. present), and central tissue (uniformly in-fected vs. uninfected). The aeschynomenoid type is defined as having the firststate of each of these four characters. Regardless, this coding strategy wouldnot change our findings because the dalbergioids would be nearly uniform inoccurrence for the states of these four characters.

APPENDIX C

Enumeration of the constituent genera of the dalbergioid clade. The em-phasis in the discussion of each of the dalbergioid genera is on the diagnostictraits that are presumably autapomorphic.

The Adesmia clade

Adesmia DC. is diagnosed by stipules attached a pseudopetiole. Althoughalso found in Stylosanthes and Arachis, the projected portions of the nodesof these two genera are nearly as long as the petiole. In Adesmia, the nodalprojections extend to much less than half the length of the petiole. In addition,Adesmia uniquely combines free staminal filaments and lomented pods (Pol-hill, 1981f). Adesmia comprises about 230 species centered in Chile and Ar-gentina (Burkart, 1949, 1954, 1960, 1962, 1964, 1966, 1967; Ulibarri, 1978,1980, 1982a, b, 1984, 1987, 1990). The genus contains two distinct mono-phyletic subgroups, according to ITS/5.8S sequence analysis (Fig. 3). One ismarked by inflorescences of usually solitary axillary flowers with pedicelsconfluent with the calyx, stipules (or at least scars) that are connate aroundthe stem, and pods that lack glandular-based trichomes, multiseriate trichomes,or the raised pericarp reticulations (e.g., Adesmia lanata and A. villosa). Thesecond clade is characterized by inflorescences of terminal racemes, subum-bels, or panicles, flowers articulated with the pedicel, stipules (or scars) thatare not connate around the stem, and pod loments that commonly bear sometype of ornamentation, for example, large glandular-based trichomes, longmultiseriate plumose trichomes, or very prominent reticulate venation (e.g.,Adesmia muricata and A. volckmannii). Phylogenetic analysis of ITS/5.8Ssequence data strongly supports the monophyly of Adesmia, as does matK/trnK.

Chaetocalyx DC. (Figs. 17–23) is paraphyletic with respect to Nissolia, anissue that is the focus of another study (M. Lavin and D. Prado, unpublisheddata). It possesses no autapomorphic traits and is characterized like Nissolia(with twining herbaceous stems and ebracteolate flowers) but lacking the ster-ile (usually samaroid) terminal loment of the mature pod. The glandular-basedtrichomes on the calyx of most species of Chaetocalyx are not diagnostic andthe species of Chaetocalyx form a rather homogeneous assemblage. The sup-posedly obvious division between species with laterally flattened or wingedfruits vs. those with terete fruits (as coded in Beyra-M. and Lavin, 1999) isnot resolved with 5.8S/ITS sequence analysis. Chaetocalyx includes ;13 neo-tropical species centered in dry forests of South America (Rudd, 1958, 1972b,1996).


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) andcharacterized by the autapomorphy of pods with a sterile (usually samaroid)terminal loment (Figs. 24–28). This genus contains ;13 species centered intropical dry forests of Mexico and Central America (Rudd, 1956, 1970a,1975b). Chaetocalyx and Nissolia lack floral bracteoles, otherwise occurringamong 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 threehave legumes with crests or bristles on each pod article and leaves that areusually paripinnate (a few species of Poiretia have imparipinnate leaves).Amicia differs from Poiretia and Zornia in having blunt keel petals, a staminalsheath that is split open above, and anthers that are mostly uniform. A recentattempt to segregate Poiretia and Zornia from Amicia (Ohashi, 1999) is notsupported by this analysis.

Poiretia Vent. is confined to the Neotropics but with most species fromBrazil 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 witha center of diversity in Brazil, and throughout sub-Saharan Africa (Mohlen-brock, 1961, 1962). It is marked by medifixed stipules (independently evolvedin Aeschynomene sect. Aeschynomene and relatives), leaves with digitatelyarranged 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 forwarddirection 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 columnis not unique and is found sporadically among the dalbergioids. Discolobiumcomprises eight species distributed from northern Argentina to adjacent Braziland Paraguay (Rudd, 1981a).

Riedeliella Harms comprises three species endemic to southeastern Braziland Paraguay (Lima and Studart da Fanseca Vaz, 1984). Like Inocarpus,Etaballia, and some species of Pictetia, the flowers of Riedeliella are nearlyradially symmetric. Lima and Studart da Fanseca Vaz (1984) propose the closerelationship of Etaballia and Riedeliella in the tribe Acosmieae (Yakovlev,1972), a group also with essentially radially symmetric flowers, although withfree staminal filaments. Riedeliella differs from Etaballia and Inocarpus inhaving paripinnate leaves and a long exerted style, and in this analysis it issuggested to be not most closely related to Etaballia, but rather to Discolob-ium.

Brya P. Br. is recorded to have explosive pollen release (Leon and Alain,1951, p. 315, fig. 131), a form of pollen presentation that is unique amongdalbergioid legumes. Also, Brya is characterized by its leaves from the longshoots being transformed into spines. Brya is sister to Cranocarpus, as evi-denced by the shared occurrence of leaves, stems, inflorescences, and podsbearing capitate glandular trichomes that are microscopically glochidiate, andby periporate pollen (Ferguson and Skvarla, 1981). Brya includes four speciesendemic 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 islike 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 struc-ture (Record, 1919), and simple axillary racemes or solitary flowers of Brya


Figs. 29–47. Representative species of the Pterocarpus clade (scale bar 5 1 cm for all figures). Figs. 29–36. Platymiscium trifoliolatum. 29. Floweringbranch. 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 dal-bergioid legumes.

Platymiscium Vogel (Figs. 29–36) comprises 18 neotropical species cen-tered in Mexico and northeastern Brazil. The genus is unique in having op-posite leaves with interpetiolar stipules (Lima, 1990; Klitgaard, 1999a, b).

Centrolobium Mart. ex Benth. comprises six tropical species from Panamato Colombia, Ecuador, Venezuela, Brazil, and Bolivia (Rudd, 1954; Lima,1985). The genus is well marked by its orange peltate glands covering theleaves and inflorescences, and winged pods in which the seed-bearing portionis covered with spines.

Grazielodendron Lima is a monotypic genus endemic to Brazil (Lima,1983, 1990). The laterally compressed pod of Grazielodendron is distin-guished by having an additional wing-like extension of the dorsal margin. Thewinged dorsal margin is markedly delineated from the main winged body ofthe pod.

Pterocarpus Jacq. (Figs. 37–47) comprises 20 species distributed pantrop-ically (Rojo, 1972). Of the genera with wide, crimped wing petals (i.e., Par-amachaerium, Geoffroea, Pterocarpus, Ramorinoa, Paramachaerium, Tipu-ana, and Platypodium), Pterocarpus is diagnosed by its pod that is wingedfrom an attenuation of the pod body all around the seed chamber (Polhill,1981d; Lima, 1990). The pods are variable in this genus with some wingedand bristly (e.g., P. angolensis), others winged and not bristly (e.g., P. indi-cus), and rarely not winged and not bristly (i.e., P. amazonum).

Tipuana (Benth.) Benth. is a monotypic genus of subtropical forests inBolivia and northwestern Argentina (Rudd, 1974). Of the genera with wide,crimped wing petals (see description of Pterocarpus), Tipuana is diagnosedby its pod that is winged from the style, the seed chambers being proximalto 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 diag-nosed by its pod that is winged from the stipe, the seed chambers being distalto 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 withlaterally broadened and crimped wing petals, Paramachaerium has reddish toviolet petals rather than the typical yellow pigment. This genus is unusual inits nectariferous disk surrounding the base of the ovary, a trait independentlyevolved in certain species of Machaerium and Ormocarpum.

Ramorinoa Speg. is a monotypic genus from west-central Argentina. Thegenus is very well marked by its leafless pungent branches (Burkart, 1952;Polhill, 1981d; Lima, 1990). As remarked by Burkart (1952), the genus is sohighly modified vegetatively that morphology provides few clues to its closestrelationships.

Inocarpus J. R. & G. Forster is very distinctive in having all five ligulatepetals fused at base, and with the ten stamens fused by their filaments to thecorolla tube (similar to some genera of Amorpheae). Inocarpus comprises oneto three species and is geographically distinctive in being restricted to Ma-laysia and adjacent Pacific islands (Polhill, 1981d). The only other dalbergioidgenus restricted to Asia is Geissaspis.

Etaballia Benth. is very similar to Inocarpus, except that its leaves areunifoliolate rather than simple, and the staminal filaments are monodelphouswith no split along the adaxial side. Etaballia is monotypic and unlike Ino-carpus is neotropical, occurring in Guyana, Venezuela, and Brazil (Rudd,1970b).

Geoffroea Jacq. comprises two species from Colombia and Venezuela southto Chubut, Argentina, and also on the Galapagos Islands possibly due tocultivation (Ireland and Pennington, 1999). Of the genera with wide, crimpedwing petals (see description of Pterocarpus), Geoffroea is diagnosed by itssessile ovary that develops into a fleshy drupe (Polhill, 1981d; Lima, 1990).


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. Keelpetals. 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 itsleaves and pods with dark pustular glands, pods with strong longitudinalnerves, arborescent habit, inflorescences of axillary racemes, and small yellowpetals is unique. Cascaronia is a monotypic genus from northern Argentinaand adjacent Paraguay and Bolivia (Polhill, 1981d).

Fissicalyx Bentham is a monotypic genus from Venezuela and Guyana, andmarked by its spathaceous calyx (all five lobes are on the adaxial lip), porateanthers, and pods with a fusiform seed chamber bearing a closely veinedmembranous wing on both margins (Polhill, 1981d; Lima, 1990). There is nomorphological evidence to suggest that this genus is closely related to Fie-brigiella, as revealed by DNA sequence analysis.

Fiebrigiella Harms is a monotypic genus from Bolivia and Ecuador (Bur-kart and Vilchez, 1971). The pods of Fiebrigiella have prominent continuousparallel venation on the lateral walls, once suggesting an affinity to Chaeto-calyx and Nissolia (Rudd, 1981a), but now considered homologous to suchpods 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 veg-etation, two New World (Florida and Mesoamerica), and five Old World (So-malia and the Yemeni island Socotra; Thulin, 2000). Arthrocarpum Balf. f.(Gillett, 1966) and Pachecoa Standl. & Steyerm. (Burkart, 1957) are synon-ymized. The genus is diagnosed by its dried (herbarium preserved) leafletswith uniformly reddish reticulate tannin deposits on the abaxial surface. Chap-mannia is sister to Arachis, and Stylosanthes; the species of these two lattergenera do not consistently show the reddish tannin reticulations. Chapmannia,Arachis, and Stylosanthes form a monophyletic group marked in part by theirsessile flowers with long hypanthia. Within this group, Chapmannia maintains

the plesiomorphic spicate inflorescence, whereas Arachis and Stylosantheshave inflorescences of solitary axillary flowers.

Stylosanthes Swartz and Arachis share the synapomorphy of stipules unitedto nodal projections, which in turn are superficially continuous with the petiole(a trait known also from Adesmia). Stylosanthes is distinguished from Arachisby having lomented, nongeocarpic pods, which are presumably plesiomorphic,as well as ovaries that are uniformly covered by uniseriate trichomes, anautapomorphy. The ;25 species of Stylosanthes are distributed in warm tem-perate to tropical regions of the world, but with a center of diversity in theneotropics (Mohlenbrock, 1957, 1960, 1963; Rudd, 1981a).

Arachis L. is distinguished from Stylosanthes by its flowers with a verylong and narrow hypanthium, a gynophore (Moctezuma and Feldman, 1998)that renders the pods geocarpic, nonlomented glabrous pods, and mostly fourleaflets per leaf. The 69 species of Arachis originate in South America froma region including Brazil south to northern Argentina (Krapovickas and Greg-ory, 1994).

The Dalbergia clade

Dalbergia L. f. (Figs. 48–58) is diagnosed by small ovate to obovate an-thers with short transverse slits at dehiscence. The genus includes over 100species distributed pantropically, but with centers of diversity in Amazoniaand Indo-Asia (Prain, 1904; Pittier, 1922; Polhill, 1981d; Lima, 1990; de Car-valho, 1997).

Machaerium Pers. (Figs. 59–69) includes ;120 neotropical species, al-though M. lunatum (L. f.) Ducke also occurs in western Africa. Machaeriumis related to Dalbergia (Polhill, 1981d; Doyle et al., 1997) and Aeschynomene


sect. Ochopodium, as evinced in part by inflorescences of helicoid cymes (butpolymorphic in all three taxa). Machaerium differs in its spinescent recurvedstipules (on at least the climbing species) and pods that are usually distallywinged, 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 includesspecies that do not fit the diagnosis of the other dalbergioid genera. It is forthis reason that the genus is treated with two terminal taxa, sects. Ochopodium(with basifixed stipules) and Aeschynomene (with medifixed stipules). SectionOchopodium, according to DNA sequence analysis, is more closely related toMachaerium than to sect. Aeschynomene (e.g., section Ochopodium is rep-resented by Aeschynomene purpusii and A. fasicularis in Fig. 5). Regardless,it is not certain if either of these two sections are monophyletic, a topic thatwill 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. pfun-dii, A. rudis, and A. virginica in Figs. 2 and 5), potentially lacking any obviousmorphological apomorphy, could be paraphyletic with respect to at least someof the genera listed immediately below. It is beyond the scope of this analysisto address this potential problem. As such, sect. Aeschynomene comprises;50 species with a pantropical distribution (Leonard, 1954; Rudd, 1955,1959, 1972a; Verdcourt, 1971; Fernandes, 1996).

Soemmeringia Mart. is characterized by a scarious standard petal that per-sists with the mature pod, which is independently evolved in some species ofOrmocarpum. 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 allclosely 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 orreplum. This monotypic genus is locally common across tropical Africa, andin southeast Asia (Laos and Borneo) and northern Australia (Hepper, 1958).

Kotschya Endl. and Smithia (below) are characterized by an inflorescenceof a dense strobilate helicoid cyme, a pod enclosed by the calyx and in whichthe articles are folded against each other. Kotschya differs in having alternateleaflets that each bear 2–7 basal nerves, as well as basifixed stipules. Kotschyacomprises 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 oppositeleaflets each bearing one main nerve. Smithia comprises ;30 species mainlyin 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 thesubtending flower and fruit (independently evolved in Zornia). Geissaspis andBryaspis differ by ebracteolate flowers, and Geissaspis differs from Bryaspisby its medifixed stipules. Geissaspis comprises three species confined to trop-ical and subtropical central and southeast Asia, but not crossing Wallace’s line(Gledhill, 1968; Rudd, 1981a)

Bryaspis Duvign. includes two species from tropical west Africa (Gledhill,1968; Hepper, 1958; Gillett, Polhill, and Verdcourt, 1971; Rudd, 1981a). Un-like Geissaspis, the inflorescence bracts of Bryaspis are markedly imbricate.

Humularia Duvign. differs from Geissaspis and Bryaspis by emarginateinflorescence bracts and panduriform standard petals. Humularia comprises;40 species confined to central Africa (Gledhill, 1968; Gillett, Polhill, andVerdcourt, 1971; Verdcourt, 1974; Rudd, 1981a).

Weberbauerella Ulbrich is diagnosed by the combination of its herbaceoushabit, pustular glands densely covering the stems, leaves, and inflorescences(including petals), and leaves with well over 40 leaflets. Similar pustularglands on the petals are known from Poiretia, but this genus is marked byleaves with four leaflets, and a sometimes climbing habit. Weberbauerellacontains two species confined to sand in southern coastal Peru (Ferreyra,1951; Rudd, 1981a).

Pictetia DC. is characterized by spiny stipules, short shoots bearing disti-chously arranged stipules (shared with Ormocarpum, Ormocarpopsis, andPeltiera), coriaceous leaflets that in all but two species have spinescent mu-cros, and pods with two-ribbed placental margins. Pictetia includes eight spe-cies confined to Cuba, Hispaniola, Puerto Rico, and the Virgin Islands ex-cluding St. Croix (Beyra-M. and Lavin, 1999).

Diphysa Jacq. has been characterized by its mature pods that have an exo-carp distinctly inflated and separated from the mesocarp. However, Diphysaormocarpoides and D. spinosa have laterally flattened lomented pods verysimilar to species of Ormocarpum and Pictetia (Antonio and Sousa, 1991).The monophyly of this genus is strongly supported, however, by phylogeneticanalysis of molecular data (see also Beyra-M. and Lavin, 1999; Lavin et al.,2000). The genus includes about ten species centered in Mexico and CentralAmerica (M. Lavin, unpublished data).

Ormocarpum P. Beauv. is diagnosed by most species forming a cylindricalnectary disk surrounding the base of the ovary (M. Thulin and M. Lavin,unpublished data). This trait otherwise is known in a few species of Ma-chaerium and Paramachaerium. This genus of ;20 species is primarily Af-rican. Three species occur on the southern Arabian Peninsula in Yemen (in-cluding Socotra) and Oman, and one to two species occur in tropical Asiaand Australia (Gillett, 1966; Rudd, 1981a; Thulin, 1990). According to ITS/5.8S sequence data (see also Lavin et al., 2000), Ormocarpum comprises twolineages (one with and one without the intrastaminal disk) that are collectivelyparaphyletic with respect to Ormocarpopsis (and Peltiera). This issue is beingaddressed in a separate study (M. Thulin and M. Lavin, unpublished data).

Ormocarpopsis R. Viguier has short shoots with persistent distichously ar-ranged stipules shared with Ormocarpum, Peltiera, and Pictetia. In this con-text, its non-lomented pod with a smooth exocarp (no evidence of prominentparallel nervation on the pod valves) and tannin patches on the abaxial surfaceof dried leaflets are diagnostic. Ormocarpopsis comprises six species endemicto Madagascar (Labat and Du Puy, 1996).

Peltiera Labat & Du Puy includes two endemic Madagascan species thatare sister to Ormocarpopsis (Labat and Du Puy, 1997). These two generashare a distinctive tannin patterning on the abaxial surface of herbarium-driedleaflets where tannin deposits are concentrated along the midrib. Like Or-mocarpopsis, the flowers of Peltiera lack a nectary disk (M. Thulin and M.Lavin, unpublished data), and the pods, though lomented and with all but oneloment aborting, contain spherical seeds. The pod valves in the seed-bearingarticle are dehiscent. Unfortunately, both species of Peltiera are probablyextinct due to the clearing of forests from which they were known.

The Dalbergioid Legumes (Fabaceae): Delimitation of a Pantropical Monophyletic Clade

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