Morphology and character evolution of seedlings in Leptolobieae (Leguminosae, Papilionoideae)

Morphology and character evolution of seedlings in Leptolobieae (Leguminosae, Papilionoideae) | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Morphology and character evolution of seedlings in Leptolobieae (Leguminosae, Papilionoideae) JACIARA CERQUEIRA DA SILVA, Ely Simone Cajueiro Gurgel, André Olmos Simões This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7973373/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Molecular phylogenetic evidence demonstrate the monophyly of five genera ( Bowdichia, Diplotropis, Guianodendr on, Leptolobium and Staminodianthus ) into a recircumscribed Leptolobieae. Even though recognition of the tribe is highly supported by molecular data, morphological characterization of its taxa is still challenging. In this context, our study aimed to study the seedling morphology of eleven species from all genera of Leptolobieae. The seedling morphology of Staminodianthus racemosus is here described for the first time. The observed characters were scored into a morphological matrix and mapped onto a phylogeny in order to detect potential synapomorphies for major clades. Three morphological types of seedlings were observed and described: phanero-epigeo-foliaceous seedlings (PEF) in Bowdichia virgilioides and Leptolobium spp., crypto-hipogeo-reserve seedlings (CHR) in Guianodendron praeclarum and Diplotropis martiusii, and phanero-hipogeo-reserve seedlings (PHR) in S. racemosu s. PEF morphotype was reconstructed as the mostly likely state in the most recent common ancestor (MRCA) of Leptolobieae, with subsequent transitions to the CHR morphotype in Diplotropus and Guianodendron , and to the PHR morphotype in S. racemosus . Another five seedling characters were reconstructed as potential synapomorphies for clades in Leptolobieae: hypocotyl shape and length, epicotyl length, presence of glands at the eophyl axyl and the number of leaflets at the seedling first node. For Leptolobium , the phyllotaxy, number of eophylls from first to third node, leaflet shape, apex and base form, and the presence of stipels are diagnostic characters within species and could be used for taxonomic purposes. ancestral character state reconstruction Leptolobieae seedling Staminodianthus Figures Figure 1 Figure 2 Introduction Papilionoideae is the largest subfamily in Leguminosae, encompassing about 14,000 species from 503 genera and 28 tribes (LPWG, 2017). Within Papilionoideae, tribe Leptolobieae sensu Cardoso et al. (2012) is a monophyletic group that comprises 29 species from five genera: Bowdichia Kunth (2 spp.), Diplotropis Benth. (10 spp.), Leptolobium Vogel (13 spp.), Guianodendron Sch. Rodr. & A.M.G. Azevedo (1 sp.) and Staminodianthus D.B.O.S. Cardoso, H.C. Lima & L.P. Queiroz (3 spp.). The tribe is defined by a number of morphological synapomorphies from its flowers and fruits: subequal, symetrical lower petals (wings and keel), smooth lateral petals (wings), staminal filaments free at base, compressed samaroid fruits, and bracts persistent at fruiting stages (Cardoso et al., 2012a). A number of phylogenetic studies focused on the Genistoid clade published in the last two decades (Wojciechowski et al., 2004; Cardoso et al., 2012; Rodrigues & Tozzi, 2007) have caused significant changes in the traditional circumscription of Leptolobieae. The Genistoid clade is a large group comprising a number of papilionoid genera, such as Baptisia Vent., Crotalaria L., Genista L., Lupinus L., Sophora L. and Thermopsis R. Br., mainly found in the Southern hemisphere and that has been consistently found as monophyletic in several studies. One of the most significant changes in Leptolobieae was the recognition of two neotropical genera, Bowdichia Kunth and Diplotropis Benth, previously ascribed to tribe Sophoreae (Wojciechowski et al. (2004), as a highly supported clade within Genistoids and sister to representatives of Leptolobieae (Cardoso et al., 2013b). Bowdichia and Diplotropis are morphologically alike in their floral morphology, sharing the following features: zygomorphic flowers and corolla with unequally sized, spathulate to narrowly obovate wings and keels. The two genera can be distinguished by a number of reproductive features of flowers and seeds. Bowdichia has a suborbicular standard, wings and keel inflexed, superimposed and without auricles, and seeds slightly compressed seeds with a hard testa and oval hilum. Diplotropis , on the other hand, have a sagitate, reflex standard, auricles strongly inflexed and not superimposed, strongly compressed seeds with a chartaceous testa and inconspicuous elliptic hilum. (Kirkbride et al, 2003; Cardoso et al., 2013b; Lima & Cardoso, 2015). The phylogenetic proximity of Bowdichia and Leptolobium is somehow intriguing. Even though the two genera have distinct floral morphology, their fruits, seeds and seedlings have a number of similar features that may represent putative synapomorphies: PEF seedlings (“Phanero-Epigeous-Foliaceous) with intercotyledonary glands and quadrangular hypocotil (Rodrigues & Tozzi, 2007b; Rodrigues & Tozzi 2008a), samaroid fruits and compressed seeds. In traditional classifications, Leptolobium has been included into the synonymy of Acosmium , with its taxa distributed in two sections, Acosmium sect. Leptolobium (Vogel) Yakovlev and Acosmium sect. Mesitis (Vogel) Yakovlev. Phylogenetic evidence, however, has supported the paraphyly of Acosmium (Rodrigues & Tozzi, 2008b), with Leptolobium as one of its segregates that should be restablished to the generic level. In its current circumscription, Leptolobium comprises 13 species, but no infrageneric division has been proposed due to the lack of morphological diagnostic features. Another segregate of Acosmium is Acosmium sect. praeclara Yakovlev. Morphological and molecular evidence has shown that this monospecific section is a distinct lineage within Leptolobieae, from which a new genus, Guianodendron , has been proposed by Rodrigues & Tozzi in 2006. Staminodianthus is another new genus that has been recently recognized in Leptolobieae (Cardoso et al. 2012a, Cardoso et al., 2013a). It is formed by three species that has been traditionally recognized as a section within Diplotropis , D . sect. racemosae H. Lima. Its placement as sister to Guianodendron preclarum within the Bowdichia clade, and not with the other sampled species of Diplotropis , was reported by Cardoso et al. 2013a and resulted in its recognition as a distinct genus. Staminodianthus share a number of morphological features with Diplotropis , such as zygomorphic flowers, fringed petals, fleshy auricles and non-differentiated lateral and lower petals, but can be distinguished from the latter by the presence of five staminodes plus five fertile stamens (vs. ten fertile stamens), leaflets shorter than 4 cm (vs. leaflets bigger than 4 cm) and inflorescences in axyllary or terminal racemes (vs. terminal panicles) (Cardoso et al., 2013b). The presence of five fertile stamens is a putative synapomorphy of the Staminodianthus + Guianodendron clade, with a subsequent loss of the staminodes in Guianodendron (Rodrigues & Tozzi 2006, Cardoso et al. 2012a; Cardoso et al., 2013b). Even though recognition of Leptolobieae and its genera is highly supported by phylogenetic evidence (Cardoso et al., 2012, Cardoso et al., 2013a, Cardoso et al., 2013b; Wojciechowski, et al., 2004), morphological characterization f these lineages is difficult due to the reduced number of potential synapomorphic features from vegetative and reproductive organs. Incorporation of data from other sources may provide useful information for taxonomic characterization and evolutionary studies. This seems to be the case in Leptolobieae, in which the seedling morphology has been described by Rodrigues and Tozzi (2007b; 2008a) for species belonging to all genera but Staminodianthus . In 2012, Cardoso et al. incorporated morphological data from seedlings on a phylogenetic study in Leguminosae, including data from Rodrigues and Tozzi (2007b; 2008a). Their results confirmed that seedling morphology provide a number of characters congruent with the obtained phylogenies, thus suggesting that these features may be conservative in Leguminosae lineages. In this context, our study aim to describe the seedling morphology from representatives of Leptobieae and related tribes, based on new data and by revising all information from the literature, and to analyse it on a phylogenetic framework. The seedling morphology of Staminodianthus is here described for the first time. Material and methods Bibliographic revision Information concerning taxonomy, morphology and anatomy of seedlings, vegetative and reproductive organs of species and genera from Leptolobieae and related tribes was obtained online by using the following search websites: Google Scholar ( https://scholar.google.com.br ), Clarivate, Scopus, Periódicos Capes ( Portal .periodicos. CAPES - Portal .periodicos. CAPES ) and Scielo ( SciELO.org ). The following combinations of keywords were used in our searches: " Seedlings + Morfology + Leptolobieae”, “ Systematic + review + Exostyleae or Leptolobieae”, and “ Phylogeny + Leguminosae + Exostyleae or Leptolobieae”. We have also performed searches with taxonomic names used in previous classifications, such as “ Bowdichia + Clade + Phylogeny” and Lecointea + clade + systematics”. For our study, we have selected the literature in which seedling morphology has been described. Taxon sampling The taxa selected for our study have been previously cited in Cardoso et al. (2013b) and LPWP (2013), and their seedling morphology have been previously described in Rocas (2004) and Rodrigues & Tozzi (2007b e 2008a). The only exception was Stamonidianthus racemosus , from which seedling morphology is here described for the first time (Fig. 1 ). In order to get information about available collections, area of occurence, phenology and potential collecting sites, we have examined the IAN and MG collections and made online searches at the following websites: Reflora ( http://reflora.jbrj.gov.br/ ) , JABOT ( http://hrb.jbrj.gov.br/v2/consulta.php ) , SpeciesLink ( http://splink.cria.org.br/ ) , The Plant List ( http://www.theplantlist.org/1.1/browse/A/ ) and Global Biodiversity Information Facility ( https://www.gbif.org/ ). Fruits of Staminodianthus racemosus were collected in two populations found at the Amazon forest in Mato Grosso state on the municipalities of Nova Canaã do Norte and Itaúba, and vouchers were deposited in MG. The collected propagules were then packed and sent to the Nova Esperança forest nursery on the municipality of Ipixuna do Pará, in Pará State, where they were processed and sown. Fruit processing and propagule generation Fruit processing and propagule generation After collection and transportation to the germination sites, fruits were initially kept in room temperature for 24 hours and processed until dispersal units were completely cleaned. Fruits were then washed and umidified to facilitate seed removal. Seeds were manually extracted by fruit maceration and planted after complete removal. Some fruits with enclosed seeds were also planted to enhance the probability of obtaining seedlings. To make the best possible processing, fruit and seed characteristics were taken into consideration for each species, removing fruit and seeds with conspicuous malformations, mechanical injuries or with evidences of predation. Dispersal units were then sterilized with 5% sodium hypochlorite and stored in labeled plastic bags. At this stage, seeds with injuries and/or evidence of predation were discarded. Germination, seedling formation and morphological analysis Seeds were planted at 0,5cm depth in plastic trays with 80 x 40 x 20 cm. Substrate for sowing was composed of sand, sawdust and vermiculite (1:1), and irrigated every day to keep moisture. The plastic trays were kept in wood benches inside a greenhouse (forest nursery). After germination, seedlings were transfered to poliethylene bags and kept them in controled conditions. We have considered post-seminal development the period from seed swelling to complete eophyll expansion. Morphological description of structures present in all stages of post-seminal development were made in plants with primary root, hipocotyl and normal cotyledons. Daily observations wer made on the developing plants, with special attention to morphological features that were potentially diagnostic and useful in species identification. A developing plant was considered a seedling when eophyls were completely developed ( sensu Duke & Polhill, 1981; Oliveira, 2001). Only seedlings with primary roots, hipocotyl, cotyledons, epicotyl and well-developed eophylls were selected for morphological analysis and characterization. All vegetative features described and ilustrated were present in both post-seminal and seedling stages and are presented in details in Table 3. Measures from epicotyl and hipocotyl were coded as short, medium and long,with intervals defined by the mean values obtained for each analysed specimen. Diagnostic morphological features of fruits, seeds, post-seminal developmental stages and seedlings were illustraded by photographs and schematic drawings, the Procreate program was used to create the illustrations. An identification key for genera of Leptolobieae based on seedling characters was made on the Xper3 platform ( http://www.xper3.fr/ ). Xper3 is an analytical tool that allows storage and edition of different sources of character, which can be used to build an interactive key for a given study group. Phylogenetic inference and character state reconstruction Phylogenetic trees were generated with sequences from three molecular markers, two from the cpDNA ( trn L intron and mat K gene) and one from the nDNA (ITS/5.8S). These markers have been traditionally used in several phylogenetic studies in Leguminosae and has generated well-resolved phylogenies for both Leptolobieae and Genistoid clades (e.g., Cardoso et al. 2012, 2013 a, b, Wojciechowski et al., 2004). Both internal and external groups previously deposited in GenBank ( https://ncbi.nlm.nih.gov/genbank/ ). Most sequences used in our study were generated by Cardoso et al. (2013b). Sophora tomentosa L. and Poecilanthe parviflora Benth. were selected as outgroups to represent the core Genistoid clade, following previous studies that suggested its sister position to the Leptolobieae clade (Moraes, 2007; Oliveira, 2001; Rodrigues & Tozzi, 2007; Valadares & Môro, 2009). A list of the sampled species with Genbank accession numbers is provided in Appendix 1. The obtained sequences were aligned in the software MAFFT v7.0 (Katho & Standley, 2013) with the standard configuration and organized in individual matrices for each molecular marker. Matrices were analyzed separately and, as no incongruence has been detected, concatenated on a total evidence matrix. Phylogenetic analyses were made from both Maximum Likelihood (ML) and Bayesian Inference (BI) approaches for individual and concatenated matrices. Substitution models were inferred for each individual matrix in software jModeltest v2.1.6, (Darriba et al., 2012), as implemented in the CIPRES Science Gateway v3.3 platform (MILLER et al., 2010). ML analyses were performed in RaxML-HPC v8.2.10 (Stamatakis, 2014), also in Cipres Platform. The GTRGAMMA model was selected for bootstrap analysis, with 1000 replicates. Bootstrap (BS) values were scored as follows: 84% as high support. BI analyses was performed in MrBayes v3.2.6 (Ronquist et al., 2012), in Cipres platform. A total of 100 million generations were obtained, with one cold and three MCMC chains, with trees sampled at each 2,000 generations. Two independent MCMC chains were executed to assure data confiability. MCMC chain convergence was verified in Tracer v1.7.1 (Rambaut et al., 2018), with an effective sample size (ESS) of 200 or higher considered as significant. Thje first 25% of sampled posterior-probability trees from each independent run were discharged as burnin, and a clade credibility tree was built with the remaining 75% trees in TreeAnnotator v2.5.1 (Bouckaert et al., 2014). Posterior probability (PP) values were scored as follows: 0.94 as strong support (Rabosky & Glor, 2010). In order to identify synapomorphies that are congruent with major clades and genera in Leptolobieae, 32 morphological characters from seedlings and adult plants were selected. The same taxa sampled for the phylogenetic analysis were selected for ancestral character state reconstruction of morphological characters. Character coding was derived mainly from direct observation of plants in the field, herbarium specimens and from information available in the literature (Cardoso et al., 2012a; Cardoso et al., 2013a; Rodrigues & Tozzi, 2007b, Rodrigues & Tozzi, 2008a; Oliveira, 2001; Mansano et al., 2004a; Felippi et al., 2014). The character and character states are given in Table 1 , and the coded morphological matrix in Table 2 . The matrix contains polymorphic codings whenever a character is variable for a given taxon. Character evolution was reconstructed onto the maximum credibility tree generated in the Bayesian analysis from the total evidence matrix by using the Maximum Likelihood State Reconstruction package for Mesquite v3.81 under Mk1 model (Maddison & Maddison, 2023). States were reconstructed as proportional likelihoods for each node, and a summary of the most relevant results is shown in Fig. 2 . Table 1 List of character and character states from seedlings and adult plants selected for this study. nº Character States 1 Seedling morphotype 0 = CHR 1 = PEF 2 = PHR 2 Pulvinus 0 = absent 1 = present 3 Eophyl nyctinasty 0 = absent 1 = present 4 Lenticells 0 = absent 1 = present 5 Hypocotyl shape 0 = cylindric 1- quadrangular 6 Hypocotyl length 0 = short (1–2 mm) 1 = long (12–62 mm) 7 Epycotyl length 0 = short (3–40 mm) 1 = long (70–130 mm) 8 Leaflet shape 0 = ovate 1 = oblong 2 = elliptic 3 = cordate 9 Leaflet base shape 0 = cuneate 1 = rounded 2 = obtuse 3 = cordate 10 Leaflet apex shape 0 = obtuse 1 = acute 2 = emarginate 3 = acuminate 4 = rounded 11 Number of leaflets at first node 0 = 1 1 = 3 2 = 12 12 Glands at eophyl axyl 0 = absent 1 = present 13 Phyllotaxy of first node 0 = opposite 1 = alternate 2 = subopposite 14 Stipels 0 = absent 1 = present 15 Stipel shape 0 = absent 1 = linear 2 = linear-lanceolate 3 = ovate 16 Stipels 0 = absent 1 = present 17 Leaflet symmetry 0 = assymetric 1 = symmetric 18 Canaliculate rachis and petiole 0 = absent 1 = present 19 Cataphylls 0 = absent 1 = present 20 Curvature of calyx + hypanth 0 = straight 1 = curved 21 Corolla symmetry 0 = zygomorphic 1 = actinomorphic 22 Petals color 0 = blue to lilac-purplish 1 = pinkish 2 = yellow 3 = withish 23 Fringed petals 0 = absent 1 = present 24 Fertile stamens 0 = ten 1 = five 25 Staminodes 0 = absent 1 = present 26 Fruit type 0 = regular legume 1 = nut-like legume 2 = compressed samaroid legume 27 Bracts 0 = caducifolious 1 = persistent in fruit 28 Aril 0 = absent 1 = present 29 Endosperm 0 = present 1 = absent 30 Radicle shape 0 = linear 1 = bulbose 31 Inflorescence type 0 = raceme 1 = congested panicle 2 = branched, lax panicle 32 Root nodules in seedlings 0 = absent 1 = present Table 2 Morphological data matrix used for the ancestral reconstruction of character evolution with coded states. ? = missing data; polymorphic states were coded with all possible states for a given taxa separated by a dash (/). Espécies 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 Bowdichia virgilioides 1 0 0 0 1 1 0 0 1/2 0/1 0,1 1 1 1 2 1 1 1 0 1 0 0 1 0 0 2 1 0 0 1 2 0 Diplotropis matiusii 0 0 0 1 0 0 1 2 1/2 3 0 0 0 1 1 1 0 1 0 1 0 1 1 0 0 1 1 1 1 1 2 0 Guianodendron praeclarum 0 0 0 0 0 0 1 2 0 3 1 0 0 1 3 1 1 0 0 0 1 3 0 1 0 2 1 1 1 1 2 0 Leptolobium brachystachyum 1 0 0 0 1 1 0 1 2 0 0/1 1 1 1 2 0 1 0 0 0 1 3 0 0 0 2 0 0 0 1 1 0 Leptolobium bijugum 1 0 0 0 1 1 0 0 2 0 0 1 1 1 2 0 1 0 0 0 1 3 0 0 0 2 0 0 0 1 1 0 Leptolobium dasycarpum 1 0 0 0 1 1 0 0 2 1 0 1 0 1 2 0 1 0 0 0 1 3 0 0 0 2 0 0 0 1 2 0 Leptolobium elegans 1 0 0 0 1 1 0 3 3 0/2 0 1 0/2 1 2 0 1 0 0 0 1 3 0 0 0 2 0 0 0 1 2 0 Leptolobium nitens 1 0 0 0 1 1 0 0 2 1 0/1 1 0 1 2 1 1 0 0 0 1 3 0 0 0 1 0 0 0 1 2 0 Poecilanthe parviflora 0 0 1 1 0 0 1 0/2 0/2 0/3 2 1 1 1 2 1 1 1 0 ? 0 3 ? 0 0 0 0 0 1 ? 0 0 Sophora tomentosa 0 1 0 0 0 0 1 0 1 4 0 0 1 0 0 0 1 0 1 ? 0 2 ? 0 0 0 0 0 ? ? 0 1 Staminodianthus racemosus 2 1 0 0 0 0 1 2 2 0,4 1 0 1 1 1 0 1 1 1 1 0 1 1 1 1 1 1 1 1 1 0 0 Results Seedling morphology We recognized three morphological types of seedlings in the studied species of Leptolobieae. Bowdichia virgilioides and Leptolobium spp. have phanero-epigeo-foliaceous seedlings (PEF), whereas G. praeclarum and D. martiusii have crypto-hipogeo-reserve seedlings (CHR) and S. racemosu s have phanero-hipogeo-reserve seedlings (PHR) (Rodrigues e Tozzi, 2007b; 2008). The species of Leptolobieae studied here can be distinguished by the shape and length of the hypocotyl. In terms of elevation of the cotyledons, they can be divided in two groups: seedlings with quadrangular hypocotyl ( Bowdichia virgilioides and Leptolobium spp.), and seedlings with cylindrical hypocotyl ( G. praeclarum, D. martiusii and S. racemosus). The size of the hypocotyl is also a diagnostic feature, with Bowdichia virgilioides and Leptolobium spp. having a medium to long-sized hypocotyl (12–62 mm) and species of G. praeclarum, D. martiusii and S. racemosus with much smaller hypocotyls (1–2 mm). The epicotyl is cylindrical in all studied species of Leptolobieae, but its size is variable. In Bowdichia virgilioides and Leptolobium spp., the epicotyl is short, but conspicuously longer in G. praeclarum, D. martiusii and S. racemosus. Cotyledons can either function as photosynthetic sites or storage organs to assure an ideal growth of seedlings during initial stages of development (Garwood, 1995). Seedlings of B. virgilioides and Leptolobium spp. have leafy cotyledons, whereas S. racemosus has thick, storage-rich cotyledons. The insertion point between hypocotyl and epycotyl, a feature associated to hypocotyl elongation, is another diagnostic character in Leptolobieae. In B. virgilioides and Leptolobium spp., the insertion point is located above the surface, but in S. racemosus it is found at the ground level. The seedlings of G. praeclarum and D. martiusii have cotyledons hidden into the seminal remains. The endosperm stored at these cotyledons function as a nutritive tissue for vertical development of the seedling. Cotyledon shape in phanerocotiledonary seedlings are elliptic in B. virgilioides and Leptolobium spp. and ovate in S. racemosus . The base is obtuse in the three genera, but it can also be slightly oblique or cordate in B. virgilioides and Leptolobium spp. Nictinasty was not observed in the cotyledons from all analyzed taxa, but was present in the eophyls of species of B. virgilioides , S. racemosus and Leptolobium spp. (e xcept L. dasycarpum ), In that taxa, the adaxial face of the leaflets are exposed by the descendent orientation of the eophyls. In D. martiusii , the eophyl morphology differs significantly during plant development. Seedlings have a single leaflet, but young plants have compound leaves with several, imparipinated leaflets. The number of eophyls from the first to third nodes is highly variable, being constant only in D. martiusii, L. elegans, L. dasycarpum and L. bijugum , with one eophyl per node. In all taxa, eophyls have broquidodromous venation. Eophyls/leaflets were elliptic in D. martiusii, G. praeclarum and S. racemosus , ovate in B. virgilioides, L. panamense, L. bijugum, L. dasycarpum and L. nitens , cordate in L. elegans and oblong in L. brachystachyum . The apex was acute in L. nitens, L. dasycarpum e L. panamense , obtuse in L. bijugum, L. brachystachyum, L. elegans and acuminate in D. martiusii and G. praeclarum. In B. virgilioides and S. racemosus , eophyls/leaflets were mainly obtuse, but acute and rounded apices were ocasionally observed in the former and latter, respectively. The base was always symetrical, obtuse to rounded in B. virgilioides and D. martiusii , obtuse in L. nitens, L. dasycarpum, L. brachystachyum, L. bijugum and S. racemosus , cuneate in G. praeclarum , and cordate in L. elegans. Canaliculate petiole and rachis were observed in B. virgilioides, D. martiusii and S. racemosus. Intercotyledonary glands were observed in B. virgilioides and Leptolobium spp., but absent in the remaining taxa. Stipels were observed in all taxa, being linear in D. martiusii and S. racemosus , linear-lanceolate in B. virgilioides and Leptolobium spp., and ovate in G. praeclarum. Stipels were observed in B. Virgilioides, D. martiusii, G. praeclarum and Leptolobium nitens , but were absent in the remaining taxa. Leaves with opposite phylotax y at the first node is a shared feature among B. Virgilioides, L. nitens, L. brachystachyum, L. bijugum and S. racemosum , Morphological description of fruits, seeds and seedlings of Staminodianthus racemosus Fruit samaroid, one-seeded, greenish when young and brown when mature and dry. oblong, apex caudate, base cuneate, venation reticulated and conspicuous on both faces. Seeds green, glabrous, rounded and flattened, 11–15 x 9–12 mm (Fig. 1 ). Seedling of phanero-hypogeo-reserve type (PHR), germinating at the 15th day after sowing; cotyledons green, glabrous, sessile, 11 x 9 x 2 mm, nictinasty absent. Hipocotyl cylindric, short, about 1 mm length and 2,5 mm width, glabrous; epicotyl cylindric, light green, about 66 mm length and 2 mm width, sparsely pubescent; cataphylls 2, opposite, 5 x 1,5 mm, caducous, slightly canaliculate, located at the intermediary region between the hypocotyl and first node. Eophyls alternate, 3-foliolate at the first node, 5-foliolate at the second node and 57-foliolate at the third node, leaflets 9–18 x 6–10 mm, elliptic, apex obtuse to rounded, base obtuse, rachis and petiole canaliculate; stiple linear, 2mm, stipels absent, venation brochidodromous, nictinasty present, descendant orientation (Fig. 1 ). The seedling morphological type, cotyledon shape, number of leaflets in the three nodes of the eophyl, the absence of stipels and presence of cataphylls are diagnostic features of the genus that allows its recognition within Leptolobieae. Phylogenetic analysis Leptolobieae was recovered as monophyletic, with all sampled taxa strongly supported (BS = 1.0) on a single clade (Fig. 2 ). The five sampled taxa of Leptolobium are nested into a strongly supported clade (BS = 0.96) sister to Bowdichia virgilioides. A sister relationship of Guianodendron and Staminodianthus was also recovered (BS = 1.0), as well as the sister relationship of this clade to Diplotropis (BS = 1.0). Similar results were observed in both individual and combined datasets. Morphological matrix and character state reconstruction Of the 32 coded characters, 20 are related to the seedling morphology and 12 to inflorescence/flower morphology (Tables 2 , 3). Six characters from seedlings were reconstructed as potential synapomorphies for clades within Leptolobieae. The PEF plantlet morphotype (char. 1, state 1) is the mostly likely state in the most recent common ancestor (MRCA) of Leptolobieae. Transition from PEF to CHR (char. 1, state 0) may have occurred once in the MRCA common ancestor of the Diplotropus + Guianodendron + Staminodianthus clade, or, alternatively, twice in this clade, once in Diplotropus and again in Guianodendron . The PHR morphotype (char. 1, state 2) is an autapomorphy of Staminodianthus , but our reconstruction is equivocal whether it have evolved from PEF or CHR types. Quadrangular (character 5, state 1) and long (character 6, state 1) hypocotyls were reconstructed as the most likely states at the MRCA of Bowdichia + Leptolobium , probably evolving from cylindric, shorter hypocotyls. Epicotyls with a length of 70 mm or longer (character 7, state 1) is the most likely condition in the MRCA of Leptolobieae, from which epicotyls shorter than 40 mm (character 7, state 0), has unambiguously evolved in the MRCA of Bowdichia + Leptolobium . The presence of glands at the eophyl axyl (character 12, state 1) is a novelty for Bowdichia + Leptolobium and may have evolved once in the MRCA of this clade, as this feature is not present in any other genera of Leptolobieae. The presence of three leaflets at the seedling first node (char. 11, state 1), instead of one, four or 12, is an autapomorphy of Guianodendron . A number of morphological features from adult were unambiguously reconstructed as synapomorphies for genera in Leptolobieae: presence of pulvin (char. 2, state 1), cataphylls (char. 19, state 1), staminodes (char. 25, state 1), and racemose inflorescences (char. 31, state 0) for Staminodianthus ; flowers lilac to blue (char. 22, state 0) in Bowdichia ; flowers actinomorphic (char. 21, state 1) and early senescent bracts (char. 27, state 0) in Leptolobium ; presence of lenticells (char. 4, state 1), linear stipels (15, state 0) and assymetric basal leaflets (char. 17, state 0) in Diplotropis ; and stipels ovate (char. 15, state 2) and flowers actinomorphic (char. 21, state 1) in Guianodendron . Other features reconstructed as synapomorphies for major clades are the presence of only five fertile stamens (char. 24, state 1) in Guianodendron + Staminodianthus , seeds with arils (char. 28, state 1) in Diplotropis + Guianodendron + Staminodianthus , and samaroid fruits (char. 26, state 2) and seeds with endosperm (char. 29, state 1) in Bowdichia + Leptolobium. Discussion Seedlings have a number of morphological features that are variable among taxa and can provide useful information for taxonomic and evolutionary studies. In Leguminosae, seedlings have been extensively studied in several genera and provide useful characters for delimitation of groups at different taxonomic levels (e.g., Ye, 1983; Lima, 1990; Oliveira, 2001; Rodriguez & Tozzi 2007). In Leptolobieae and related tribes, Rodriguez & Tozzi (2008) used seedling characters from species of Acosmium, Leptolobium and Guianodendron to evaluate the systematic value of these characters. Later on, Cardoso et al. (2012, 2013a) has used seedling characters on a phylogenetic context to detect potential synapomorphies and evolutionary trends in Leptolobieae. Our study follow that approach by adding new information from seedling morphology to a simplified phylogeny of Leptolobieae, in which all genera from the tribe were sampled. Our results support to the recognition of Staminodianthus as a distinct genus from Diplotropis and its placement as sister to Guianodendron. Elevation of Diplotropis sect. racemosae to the generic rank was proposed by Cardoso et al. (2013a), based on molecular and morphological evidence. The authors reported six apomorphies unique to Staminodianthus within the Genistoid legumes, all related to the flower architecture: actinomorphic symmetry calyx with a curved hypanthium, well-differentiated standard petal, lateral and lower petals without auricles, and androecium with five fertile stamens and five staminodes. At the time of this publication, seedlings of Staminodianthus were unkown to science, so that there was no information available in the literature to compare with seedlings from other genera in the tribe. Our observation and description of seedlings of S. racemosus has provided additional evidence supporting recognition of the genus. Its morphotype, phanero-hypogeo-reserve type (PHR), have thick, expanded cotyledons at the ground level with energetic reserve, an unusual condition exclusive of this genus in Leptolobieae. In addition, seedlings of S. racemosus share six features with seedlings from the sampled species of Diplotropis and Guianodendron , reinforcing the phylogenetic affinities of the three genera. A close relationship of Bowdichia and Leptolobium is also supported by our results. Despite having distinct floral features, which has placed them in different tribes in traditional classifications of Leguminosae, the phylogenetic proximity of these two genera is strongly supported in studies focused on the Genistoid and Leptolobieae clades (e.g., Cardoso et al. 2012, 2013). These studies also reported two features from seedlings (presence of intercotiledonary glands and quadrangular hypocotyls) shared by the two genera that represent potential synapomorphies. Our study corroborated these results and reconstructed two additional character states from seedlings as unequivocal synapomorphies for the Bowdichia + Leptolobium clade: epicotyls shorter than 40 mm and hypocotyls longer than 11 mm. Within Leptolobium , the phyllotaxy, number of eophylls from first to third node, leaflet shape, apex and base form, and the presence of stipels were variable among the five sampled taxa and thus potentially useful for species delimitation. For that reason, we provide here an identification key based solely on seedling characters. Our sampling, however, is lower than 50% of the valid taxa in Leptolobium (13 species), so that the taxonomic value of these features could not be properly tested. A larger sampling within the genus is required to address this question in future studies and to build a more complete identification key. Key to the species of Leptolobium 1 Stipels present, hipocotyl 58-62mm length, epicotyl 35-40mm length .............. L. nitens. 1’ Stipels absent, hipocótilo 17-37mm length, epicotyl 8-28mm length ...............2 2 Phylotaxy of the eophyl from the first node opposite ......................................................................3 3 Nictinasty of leaflets present, hipocotyl 35-37mm length, blade ovate, apex obtuse, base cordate ....................... L. elegans . 3’ Nictinasty of leaflets absent, hipocotyl 22-33mm length, blade cordate, apex acute, base obtuse ...................... L. dasycarpum . 2’ Phylotaxy of the eophyl from the first node alternate..................................................4 4 Cotyledons 5-10mm width, secondary veins with angles of 60–70º to the primary vein, blade ovate ............................................. L. bijugum . 4’. Cotyledons 11-12mm width, secondary veins forming angles of 70–85º to the primary vein, blade oblong ............................................L. brachystachyum . Our study has reinforced the systematic value of seedling features in Leguminosae, specially in tribe Leptolobieae. Morphological description of seedlings of Staminodianthus racemosus has provided additional evidence of the validation of its generic status and close relationship to Guianodendron and Diplotropis . Observation and description of seedlings from five species of Leptolobium demonstrates that several seedling features are constant at the specific level but variable within species and therefore can be used for systematic purposes. Future studies including more species from both Leptolobieae and Genistoid clade, with description of their seedling morphology and further character mapping on a phylogenetic approach, are greatly desired to improve our knowledge on the systematics and evolution of these lineages of Leguminosae. Declarations Author Information Authors and Affiliations Universidade Federal Rural da Amazônia / Museu Paraense Emílio Goeldi, Botany Department, Belém, Pará, Brazil. https://orcid.org/0000-0002-9715-158X Jaciara Cerqueira da Silva Museu Paraense Emílio Goeldi, Ministry of Science and Technology, Botany Department, Belém, Pará, Brazil. https://orcid.org/0000-0002-9488-7532 Ely Simone Cajueiro Gurgel Universidade Estadual de Campinas, Instituto de Biologia, Campinas, São Paulo, Brazil. https://orcid.org/0000-0001-6555-8759 André Olmos Simões Corresponding Author Correspondence to: Jaciara Cerqueira da Silva – [email protected] Funding Scholarship granted by the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), process no. 132296/2020-9. This research is part of the master’s dissertation of the first author, conducted within the Graduate Program in Biological Sciences of the Universidade Federal Rural da Amazônia (UFRA) / Museu Paraense Emílio Goeldi (MPEG), Belém, Pará, Brazil. Declarations and Ethical Statements Each author made a substantial contribution to the conception and design of the study, as well as to data acquisition, analysis, and interpretation. All authors have approved the final version of the manuscript for publication. The authors declare that they have no conflicts of interest, financial or otherwise, that could be perceived as influencing the content of this article. Conflict of interest Each author made a substantial contribution to the conception and design of the study, as well as to data acquisition, analysis, and interpretation. All authors have approved the final version of the manuscript for publication. The authors declare that they have no conflicts of interest, financial or otherwise, that could be perceived as influencing the content of this article. Author Contributions Collection and monitoring of seedling development, manuscript structuring, and writing (JCS). Morphological conceptualization of the group, data validation, and manuscript review (ESCG). Critical analysis of the text regarding phylogenetic systematics, data validation, english writing and final formatting (AOS). Acknowledgments We thank Dr. Mathias Engels for collecting botanical material of Staminodianthus racemosus. We are also grateful to Dr. Felipe Martins for preparing the illustrations. This study was supported by the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), which provided a research scholarship. References Azani N, Babineau M, Bailey CD, Banks H, Barbosa AR, Pinto RB et al. 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1","display":"","copyAsset":false,"role":"figure","size":952459,"visible":true,"origin":"","legend":"\u003cp\u003eFruits, seeds, and seedlings of \u003cem\u003eS. racemosus\u003c/em\u003e (a), fruit; (b), seed; (c), seedling emergence; (d), seedling with fully expanded eophylls; (e), nyctinastic movement of leaflets with downward orientation; (f), detail of brochidodromous venation; (g), metaphylls of the young plant. Legend: co = cotyledons; c = cataphylls; e = eophyll; ep = epicotyl\u003c/p\u003e","description":"","filename":"Fig.1.png","url":"https://assets-eu.researchsquare.com/files/rs-7973373/v1/3b6a4f731b253f0807e2bfea.png"},{"id":97673376,"identity":"f844a399-3a55-455d-8a72-7d5e1a3989d6","added_by":"auto","created_at":"2025-12-08 09:40:00","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":224970,"visible":true,"origin":"","legend":"\u003cp\u003eClade credibility tree based on the Bayesian analysis of the combined molecular dataset with mapping of selected seedling characters. Posterior probabilities are indicated at the right side of the corresponding node. White circles denote unambiguous reconstructions for characters 1, 5, 6, 7, 11 and 12. Blue and green circles denote two alternative reconstructions for states 0 and 2 in character 1. Numbers above the circles correspond to the character numbers, and below the circles to the state numbers. Character numbers and states are as follows: 1, Seedling morphotype. 0: CHR; 1: PEF; 2: PHR. 5, Hypocotyl shape. 0: cylindric; 2: quadrangular. 6, Hypocotyl length. 0: short (1–2mm); 1: long (12–62mm). 7, Epicotyl length. 0: 0: short (3–40mm); 1: long (70–130mm). 11, Number of leaflets at first node. 0: 1; 1: 3; 2: 12. 12, Glands at eophyl axyl. 0: absent; 1: present.\u003c/p\u003e","description":"","filename":"Fig.2.png","url":"https://assets-eu.researchsquare.com/files/rs-7973373/v1/42bedfc7b2c16ac0589e74ab.png"},{"id":100547727,"identity":"aabc3302-5a44-41fc-8dcb-32586895aafe","added_by":"auto","created_at":"2026-01-19 08:16:29","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2198397,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7973373/v1/f541d12a-6189-44c5-b5c2-e3b18954cf45.pdf"},{"id":97673403,"identity":"07ea118d-424e-4572-b877-0447503b8eb7","added_by":"auto","created_at":"2025-12-08 09:40:04","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":6476,"visible":true,"origin":"","legend":"","description":"","filename":"ArtigoBrazilianJournalofBotanyAppendix1.docx","url":"https://assets-eu.researchsquare.com/files/rs-7973373/v1/3823c03771f469fac0b47f81.docx"}],"financialInterests":"","formattedTitle":"Morphology and character evolution of seedlings in Leptolobieae (Leguminosae, Papilionoideae)","fulltext":[{"header":"Introduction","content":"\u003cp\u003ePapilionoideae is the largest subfamily in Leguminosae, encompassing about 14,000 species from 503 genera and 28 tribes (LPWG, 2017). Within Papilionoideae, tribe Leptolobieae sensu Cardoso et al. (2012) is a monophyletic group that comprises 29 species from five genera: \u003cem\u003eBowdichia\u003c/em\u003e Kunth (2 spp.), \u003cem\u003eDiplotropis\u003c/em\u003e Benth. (10 spp.), \u003cem\u003eLeptolobium\u003c/em\u003e Vogel (13 spp.), \u003cem\u003eGuianodendron\u003c/em\u003e Sch. Rodr. \u0026amp; A.M.G. Azevedo (1 sp.) and \u003cem\u003eStaminodianthus\u003c/em\u003e D.B.O.S. Cardoso, H.C. Lima \u0026amp; L.P. Queiroz (3 spp.). The tribe is defined by a number of morphological synapomorphies from its flowers and fruits: subequal, symetrical lower petals (wings and keel), smooth lateral petals (wings), staminal filaments free at base, compressed samaroid fruits, and bracts persistent at fruiting stages (Cardoso et al., 2012a).\u003c/p\u003e\u003cp\u003eA number of phylogenetic studies focused on the Genistoid clade published in the last two decades (Wojciechowski et al., 2004; Cardoso et al., 2012; Rodrigues \u0026amp; Tozzi, 2007) have caused significant changes in the traditional circumscription of Leptolobieae. The Genistoid clade is a large group comprising a number of papilionoid genera, such as \u003cem\u003eBaptisia\u003c/em\u003e Vent., \u003cem\u003eCrotalaria\u003c/em\u003e L., \u003cem\u003eGenista\u003c/em\u003e L., \u003cem\u003eLupinus\u003c/em\u003e L., \u003cem\u003eSophora\u003c/em\u003e L. and \u003cem\u003eThermopsis\u003c/em\u003e R. Br., mainly found in the Southern hemisphere and that has been consistently found as monophyletic in several studies. One of the most significant changes in Leptolobieae was the recognition of two neotropical genera, \u003cem\u003eBowdichia\u003c/em\u003e Kunth and \u003cem\u003eDiplotropis\u003c/em\u003e Benth, previously ascribed to tribe Sophoreae (Wojciechowski et al. (2004), as a highly supported clade within Genistoids and sister to representatives of Leptolobieae (Cardoso et al., 2013b). \u003cem\u003eBowdichia\u003c/em\u003e and \u003cem\u003eDiplotropis\u003c/em\u003e are morphologically alike in their floral morphology, sharing the following features: zygomorphic flowers and corolla with unequally sized, spathulate to narrowly obovate wings and keels. The two genera can be distinguished by a number of reproductive features of flowers and seeds. \u003cem\u003eBowdichia\u003c/em\u003e has a suborbicular standard, wings and keel inflexed, superimposed and without auricles, and seeds slightly compressed seeds with a hard testa and oval hilum. \u003cem\u003eDiplotropis\u003c/em\u003e, on the other hand, have a sagitate, reflex standard, auricles strongly inflexed and not superimposed, strongly compressed seeds with a chartaceous testa and inconspicuous elliptic hilum. (Kirkbride et al, 2003; Cardoso et al., 2013b; Lima \u0026amp; Cardoso, 2015). The phylogenetic proximity of \u003cem\u003eBowdichia\u003c/em\u003e and \u003cem\u003eLeptolobium\u003c/em\u003e is somehow intriguing. Even though the two genera have distinct floral morphology, their fruits, seeds and seedlings have a number of similar features that may represent putative synapomorphies: PEF seedlings (\u0026ldquo;Phanero-Epigeous-Foliaceous) with intercotyledonary glands and quadrangular hypocotil (Rodrigues \u0026amp; Tozzi, 2007b; Rodrigues \u0026amp; Tozzi 2008a), samaroid fruits and compressed seeds.\u003c/p\u003e\u003cp\u003eIn traditional classifications, \u003cem\u003eLeptolobium\u003c/em\u003e has been included into the synonymy of \u003cem\u003eAcosmium\u003c/em\u003e, with its taxa distributed in two sections, \u003cem\u003eAcosmium\u003c/em\u003e sect. \u003cem\u003eLeptolobium\u003c/em\u003e (Vogel) Yakovlev and \u003cem\u003eAcosmium\u003c/em\u003e sect. \u003cem\u003eMesitis\u003c/em\u003e (Vogel) Yakovlev. Phylogenetic evidence, however, has supported the paraphyly of \u003cem\u003eAcosmium\u003c/em\u003e (Rodrigues \u0026amp; Tozzi, 2008b), with \u003cem\u003eLeptolobium\u003c/em\u003e as one of its segregates that should be restablished to the generic level. In its current circumscription, \u003cem\u003eLeptolobium\u003c/em\u003e comprises 13 species, but no infrageneric division has been proposed due to the lack of morphological diagnostic features. Another segregate of \u003cem\u003eAcosmium\u003c/em\u003e is \u003cem\u003eAcosmium\u003c/em\u003e sect. \u003cem\u003epraeclara\u003c/em\u003e Yakovlev. Morphological and molecular evidence has shown that this monospecific section is a distinct lineage within Leptolobieae, from which a new genus, \u003cem\u003eGuianodendron\u003c/em\u003e, has been proposed by Rodrigues \u0026amp; Tozzi in 2006.\u003c/p\u003e\u003cp\u003e\u003cem\u003eStaminodianthus\u003c/em\u003e is another new genus that has been recently recognized in Leptolobieae (Cardoso et al. 2012a, Cardoso et al., 2013a). It is formed by three species that has been traditionally recognized as a section within \u003cem\u003eDiplotropis\u003c/em\u003e, \u003cem\u003eD\u003c/em\u003e. sect. \u003cem\u003eracemosae\u003c/em\u003e H. Lima. Its placement as sister to \u003cem\u003eGuianodendron preclarum\u003c/em\u003e within the Bowdichia clade, and not with the other sampled species of \u003cem\u003eDiplotropis\u003c/em\u003e, was reported by Cardoso et al. 2013a and resulted in its recognition as a distinct genus. \u003cem\u003eStaminodianthus\u003c/em\u003e share a number of morphological features with \u003cem\u003eDiplotropis\u003c/em\u003e, such as zygomorphic flowers, fringed petals, fleshy auricles and non-differentiated lateral and lower petals, but can be distinguished from the latter by the presence of five staminodes plus five fertile stamens (vs. ten fertile stamens), leaflets shorter than 4 cm (vs. leaflets bigger than 4 cm) and inflorescences in axyllary or terminal racemes (vs. terminal panicles) (Cardoso et al., 2013b). The presence of five fertile stamens is a putative synapomorphy of the \u003cem\u003eStaminodianthus\u003c/em\u003e\u0026thinsp;+\u0026thinsp;\u003cem\u003eGuianodendron\u003c/em\u003e clade, with a subsequent loss of the staminodes in \u003cem\u003eGuianodendron\u003c/em\u003e (Rodrigues \u0026amp; Tozzi 2006, Cardoso et al. 2012a; Cardoso et al., 2013b).\u003c/p\u003e\u003cp\u003eEven though recognition of Leptolobieae and its genera is highly supported by phylogenetic evidence (Cardoso et al., 2012, Cardoso et al., 2013a, Cardoso et al., 2013b; Wojciechowski, et al., 2004), morphological characterization f these lineages is difficult due to the reduced number of potential synapomorphic features from vegetative and reproductive organs. Incorporation of data from other sources may provide useful information for taxonomic characterization and evolutionary studies. This seems to be the case in Leptolobieae, in which the seedling morphology has been described by Rodrigues and Tozzi (2007b; 2008a) for species belonging to all genera but \u003cem\u003eStaminodianthus\u003c/em\u003e. In 2012, Cardoso et al. incorporated morphological data from seedlings on a phylogenetic study in Leguminosae, including data from Rodrigues and Tozzi (2007b; 2008a). Their results confirmed that seedling morphology provide a number of characters congruent with the obtained phylogenies, thus suggesting that these features may be conservative in Leguminosae lineages. In this context, our study aim to describe the seedling morphology from representatives of Leptobieae and related tribes, based on new data and by revising all information from the literature, and to analyse it on a phylogenetic framework. The seedling morphology of \u003cem\u003eStaminodianthus\u003c/em\u003e is here described for the first time.\u003c/p\u003e"},{"header":"Material and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003eBibliographic revision\u003c/h2\u003e\u003cp\u003eInformation concerning taxonomy, morphology and anatomy of seedlings, vegetative and reproductive organs of species and genera from Leptolobieae and related tribes was obtained online by using the following search websites: Google Scholar (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://scholar.google.com.br\u003c/span\u003e\u003cspan address=\"https://scholar.google.com.br\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e), Clarivate, Scopus, Peri\u0026oacute;dicos Capes (\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003ePortal .periodicos. CAPES - Portal .periodicos. CAPES\u003c/span\u003e) and Scielo (\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003eSciELO.org\u003c/span\u003e). The following combinations of keywords were used in our searches: \" Seedlings\u0026thinsp;+\u0026thinsp;Morfology\u0026thinsp;+\u0026thinsp;Leptolobieae\u0026rdquo;, \u0026ldquo; Systematic\u0026thinsp;+\u0026thinsp;review\u0026thinsp;+\u0026thinsp;Exostyleae or Leptolobieae\u0026rdquo;, and \u0026ldquo; Phylogeny\u0026thinsp;+\u0026thinsp;Leguminosae\u0026thinsp;+\u0026thinsp;Exostyleae or Leptolobieae\u0026rdquo;. We have also performed searches with taxonomic names used in previous classifications, such as \u0026ldquo;\u003cem\u003eBowdichia\u003c/em\u003e\u0026thinsp;+\u0026thinsp;Clade\u0026thinsp;+\u0026thinsp;Phylogeny\u0026rdquo; and \u003cem\u003eLecointea\u003c/em\u003e\u0026thinsp;+\u0026thinsp;clade\u0026thinsp;+\u0026thinsp;systematics\u0026rdquo;. For our study, we have selected the literature in which seedling morphology has been described.\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eTaxon sampling\u003c/h3\u003e\n\u003cp\u003eThe taxa selected for our study have been previously cited in Cardoso et al. (2013b) and LPWP (2013), and their seedling morphology have been previously described in Rocas (2004) and Rodrigues \u0026amp; Tozzi (2007b e 2008a). The only exception was \u003cem\u003eStamonidianthus racemosus\u003c/em\u003e, from which seedling morphology is here described for the first time (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eIn order to get information about available collections, area of occurence, phenology and potential collecting sites, we have examined the IAN and MG collections and made online searches at the following websites: Reflora (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://reflora.jbrj.gov.br/\u003c/span\u003e\u003cspan address=\"http://reflora.jbrj.gov.br/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e)\u003c/span\u003e, JABOT (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://hrb.jbrj.gov.br/v2/consulta.php\u003c/span\u003e\u003cspan address=\"http://hrb.jbrj.gov.br/v2/consulta.php\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e)\u003c/span\u003e, SpeciesLink (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://splink.cria.org.br/\u003c/span\u003e\u003cspan address=\"http://splink.cria.org.br/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e)\u003c/span\u003e, The Plant List (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://www.theplantlist.org/1.1/browse/A/\u003c/span\u003e\u003cspan address=\"http://www.theplantlist.org/1.1/browse/A/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e)\u003c/span\u003e and Global Biodiversity Information Facility (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.gbif.org/\u003c/span\u003e\u003cspan address=\"https://www.gbif.org/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e).\u003c/span\u003e\u003c/p\u003e\u003cp\u003eFruits of \u003cem\u003eStaminodianthus racemosus\u003c/em\u003e were collected in two populations found at the Amazon forest in Mato Grosso state on the municipalities of Nova Cana\u0026atilde; do Norte and Ita\u0026uacute;ba, and vouchers were deposited in MG. The collected propagules were then packed and sent to the Nova Esperan\u0026ccedil;a forest nursery on the municipality of Ipixuna do Par\u0026aacute;, in Par\u0026aacute; State, where they were processed and sown.\u003c/p\u003e\n\u003ch3\u003eFruit processing and propagule generation\u003c/h3\u003e\n\u003cdiv class=\"Heading\"\u003eFruit processing and propagule generation\u003c/div\u003e\u003cp\u003eAfter collection and transportation to the germination sites, fruits were initially kept in room temperature for 24 hours and processed until dispersal units were completely cleaned. Fruits were then washed and umidified to facilitate seed removal. Seeds were manually extracted by fruit maceration and planted after complete removal. Some fruits with enclosed seeds were also planted to enhance the probability of obtaining seedlings.\u003c/p\u003e\u003cp\u003eTo make the best possible processing, fruit and seed characteristics were taken into consideration for each species, removing fruit and seeds with conspicuous malformations, mechanical injuries or with evidences of predation.\u003c/p\u003e\u003cp\u003eDispersal units were then sterilized with 5% sodium hypochlorite and stored in labeled plastic bags. At this stage, seeds with injuries and/or evidence of predation were discarded.\u003c/p\u003e\n\u003ch3\u003eGermination, seedling formation and morphological analysis\u003c/h3\u003e\n\u003cp\u003eSeeds were planted at 0,5cm depth in plastic trays with 80 x 40 x 20 cm. Substrate for sowing was composed of sand, sawdust and vermiculite (1:1), and irrigated every day to keep moisture. The plastic trays were kept in wood benches inside a greenhouse (forest nursery). After germination, seedlings were transfered to poliethylene bags and kept them in controled conditions. We have considered post-seminal development the period from seed swelling to complete eophyll expansion. Morphological description of structures present in all stages of post-seminal development were made in plants with primary root, hipocotyl and normal cotyledons. Daily observations wer made on the developing plants, with special attention to morphological features that were potentially diagnostic and useful in species identification.\u003c/p\u003e\u003cp\u003eA developing plant was considered a seedling when eophyls were completely developed (\u003cem\u003esensu\u003c/em\u003e Duke \u0026amp; Polhill, 1981; Oliveira, 2001). Only seedlings with primary roots, hipocotyl, cotyledons, epicotyl and well-developed eophylls were selected for morphological analysis and characterization. All vegetative features described and ilustrated were present in both post-seminal and seedling stages and are presented in details in Table\u0026nbsp;3. Measures from epicotyl and hipocotyl were coded as short, medium and long,with intervals defined by the mean values obtained for each analysed specimen.\u003c/p\u003e\u003cp\u003eDiagnostic morphological features of fruits, seeds, post-seminal developmental stages and seedlings were illustraded by photographs and schematic drawings, the Procreate program was used to create the illustrations. An identification key for genera of Leptolobieae based on seedling characters was made on the Xper3 platform (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://www.xper3.fr/\u003c/span\u003e\u003cspan address=\"http://www.xper3.fr/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e).\u003c/span\u003e Xper3 is an analytical tool that allows storage and edition of different sources of character, which can be used to build an interactive key for a given study group.\u003c/p\u003e\n\u003ch3\u003ePhylogenetic inference and character state reconstruction\u003c/h3\u003e\n\u003cp\u003ePhylogenetic trees were generated with sequences from three molecular markers, two from the cpDNA (\u003cem\u003etrn\u003c/em\u003eL intron and \u003cem\u003emat\u003c/em\u003eK gene) and one from the nDNA (ITS/5.8S). These markers have been traditionally used in several phylogenetic studies in Leguminosae and has generated well-resolved phylogenies for both Leptolobieae and Genistoid clades (e.g., Cardoso et al. 2012, 2013 a, b, Wojciechowski et al., 2004). Both internal and external groups previously deposited in GenBank (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://ncbi.nlm.nih.gov/genbank/\u003c/span\u003e\u003cspan address=\"https://ncbi.nlm.nih.gov/genbank/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e).\u003c/span\u003e Most sequences used in our study were generated by Cardoso et al. (2013b). \u003cem\u003eSophora tomentosa\u003c/em\u003e L. and \u003cem\u003ePoecilanthe parviflora\u003c/em\u003e Benth. were selected as outgroups to represent the core Genistoid clade, following previous studies that suggested its sister position to the Leptolobieae clade (Moraes, 2007; Oliveira, 2001; Rodrigues \u0026amp; Tozzi, 2007; Valadares \u0026amp; M\u0026ocirc;ro, 2009). A list of the sampled species with Genbank accession numbers is provided in Appendix 1.\u003c/p\u003e\u003cp\u003eThe obtained sequences were aligned in the software MAFFT v7.0 (Katho \u0026amp; Standley, 2013) with the standard configuration and organized in individual matrices for each molecular marker. Matrices were analyzed separately and, as no incongruence has been detected, concatenated on a total evidence matrix.\u003c/p\u003e\u003cp\u003ePhylogenetic analyses were made from both Maximum Likelihood (ML) and Bayesian Inference (BI) approaches for individual and concatenated matrices. Substitution models were inferred for each individual matrix in software jModeltest v2.1.6, (Darriba et al., 2012), as implemented in the CIPRES Science Gateway v3.3 platform (MILLER et al., 2010). ML analyses were performed in RaxML-HPC v8.2.10 (Stamatakis, 2014), also in Cipres Platform. The GTRGAMMA model was selected for bootstrap analysis, with 1000 replicates. Bootstrap (BS) values were scored as follows: \u0026lt;75% as weak support, 75%-84% as moderate support, and \u0026gt;\u0026thinsp;84% as high support. BI analyses was performed in MrBayes v3.2.6 (Ronquist et al., 2012), in Cipres platform. A total of 100\u0026nbsp;million generations were obtained, with one cold and three MCMC chains, with trees sampled at each 2,000 generations. Two independent MCMC chains were executed to assure data confiability. MCMC chain convergence was verified in Tracer v1.7.1 (Rambaut et al., 2018), with an effective sample size (ESS) of 200 or higher considered as significant. Thje first 25% of sampled posterior-probability trees from each independent run were discharged as burnin, and a clade credibility tree was built with the remaining 75% trees in TreeAnnotator v2.5.1 (Bouckaert et al., 2014). Posterior probability (PP) values were scored as follows: \u0026lt;0.7 as weak support, 0.7\u0026ndash;0.94 as moderate support, and \u0026gt;\u0026thinsp;0.94 as strong support (Rabosky \u0026amp; Glor, 2010).\u003c/p\u003e\u003cp\u003eIn order to identify synapomorphies that are congruent with major clades and genera in Leptolobieae, 32 morphological characters from seedlings and adult plants were selected. The same taxa sampled for the phylogenetic analysis were selected for ancestral character state reconstruction of morphological characters. Character coding was derived mainly from direct observation of plants in the field, herbarium specimens and from information available in the literature (Cardoso et al., 2012a; Cardoso et al., 2013a; Rodrigues \u0026amp; Tozzi, 2007b, Rodrigues \u0026amp; Tozzi, 2008a; Oliveira, 2001; Mansano et al., 2004a; Felippi et al., 2014). The character and character states are given in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, and the coded morphological matrix in Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e. The matrix contains polymorphic codings whenever a character is variable for a given taxon. Character evolution was reconstructed onto the maximum credibility tree generated in the Bayesian analysis from the total evidence matrix by using the Maximum Likelihood State Reconstruction package for Mesquite v3.81 under Mk1 model (Maddison \u0026amp; Maddison, 2023). States were reconstructed as proportional likelihoods for each node, and a summary of the most relevant results is shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eList of character and character states from seedlings and adult plants selected for this study.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"3\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003en\u0026ordm;\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCharacter\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eStates\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eSeedling morphotype\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;CHR\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;PEF\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u0026thinsp;=\u0026thinsp;PHR\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003ePulvinus\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;absent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;present\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eEophyl nyctinasty\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;absent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;present\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eLenticells\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;absent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;present\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eHypocotyl shape\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;cylindric\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1- quadrangular\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eHypocotyl length\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;short (1\u0026ndash;2 mm)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;long (12\u0026ndash;62 mm)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eEpycotyl length\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;short (3\u0026ndash;40 mm)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;long (70\u0026ndash;130 mm)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"3\" rowspan=\"4\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"3\" rowspan=\"4\"\u003e\u003cp\u003eLeaflet shape\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;ovate\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;oblong\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u0026thinsp;=\u0026thinsp;elliptic\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e3\u0026thinsp;=\u0026thinsp;cordate\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"3\" rowspan=\"4\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"3\" rowspan=\"4\"\u003e\u003cp\u003eLeaflet base shape\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;cuneate\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;rounded\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u0026thinsp;=\u0026thinsp;obtuse\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e3\u0026thinsp;=\u0026thinsp;cordate\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"4\" rowspan=\"5\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"4\" rowspan=\"5\"\u003e\u003cp\u003eLeaflet apex shape\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;obtuse\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;acute\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u0026thinsp;=\u0026thinsp;emarginate\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e3\u0026thinsp;=\u0026thinsp;acuminate\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e4\u0026thinsp;=\u0026thinsp;rounded\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003e11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eNumber of leaflets at first node\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u0026thinsp;=\u0026thinsp;12\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eGlands at eophyl axyl\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;absent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;present\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003e13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003ePhyllotaxy of first node\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;opposite\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;alternate\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u0026thinsp;=\u0026thinsp;subopposite\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e14\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eStipels\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;absent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;present\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"3\" rowspan=\"4\"\u003e\u003cp\u003e15\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"3\" rowspan=\"4\"\u003e\u003cp\u003eStipel shape\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;absent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;linear\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u0026thinsp;=\u0026thinsp;linear-lanceolate\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e3\u0026thinsp;=\u0026thinsp;ovate\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e16\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eStipels\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;absent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;present\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e17\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eLeaflet symmetry\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;assymetric\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;symmetric\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eCanaliculate rachis and petiole\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;absent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;present\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e19\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eCataphylls\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;absent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;present\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e20\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eCurvature of calyx\u0026thinsp;+\u0026thinsp;hypanth\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;straight\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;curved\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e21\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eCorolla symmetry\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;zygomorphic\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;actinomorphic\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"3\" rowspan=\"4\"\u003e\u003cp\u003e22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"3\" rowspan=\"4\"\u003e\u003cp\u003ePetals color\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;blue to lilac-purplish\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;pinkish\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u0026thinsp;=\u0026thinsp;yellow\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e3\u0026thinsp;=\u0026thinsp;withish\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e23\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eFringed petals\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;absent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;present\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e24\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eFertile stamens\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;ten\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;five\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e25\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eStaminodes\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;absent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;present\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003e26\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eFruit type\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;regular legume\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;nut-like legume\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u0026thinsp;=\u0026thinsp;compressed samaroid legume\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e27\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eBracts\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;caducifolious\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;persistent in fruit\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e28\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eAril\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;absent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;present\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e29\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eEndosperm\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;present\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;absent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eRadicle shape\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;linear\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;bulbose\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003e31\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eInflorescence type\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;raceme\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;congested panicle\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u0026thinsp;=\u0026thinsp;branched, lax panicle\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e32\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eRoot nodules in seedlings\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u0026thinsp;=\u0026thinsp;absent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026thinsp;=\u0026thinsp;present\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eMorphological data matrix used for the ancestral reconstruction of character evolution with coded states. ? = missing data; polymorphic states were coded with all possible states for a given taxa separated by a dash (/).\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"33\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c13\" colnum=\"13\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c14\" colnum=\"14\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c15\" colnum=\"15\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c16\" colnum=\"16\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c17\" colnum=\"17\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c18\" colnum=\"18\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c19\" colnum=\"19\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c20\" colnum=\"20\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c21\" colnum=\"21\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c22\" colnum=\"22\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c23\" colnum=\"23\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c24\" colnum=\"24\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c25\" colnum=\"25\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c26\" colnum=\"26\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c27\" colnum=\"27\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c28\" colnum=\"28\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c29\" colnum=\"29\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c30\" colnum=\"30\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c31\" colnum=\"31\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c32\" colnum=\"32\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c33\" colnum=\"33\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEsp\u0026eacute;cies\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c9\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c10\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c11\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c12\"\u003e\u003cp\u003e11\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c13\"\u003e\u003cp\u003e12\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c14\"\u003e\u003cp\u003e13\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c15\"\u003e\u003cp\u003e14\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c16\"\u003e\u003cp\u003e15\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c17\"\u003e\u003cp\u003e16\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c18\"\u003e\u003cp\u003e17\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c19\"\u003e\u003cp\u003e18\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c20\"\u003e\u003cp\u003e19\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c21\"\u003e\u003cp\u003e20\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c22\"\u003e\u003cp\u003e21\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c23\"\u003e\u003cp\u003e22\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c24\"\u003e\u003cp\u003e23\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c25\"\u003e\u003cp\u003e24\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c26\"\u003e\u003cp\u003e25\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c27\"\u003e\u003cp\u003e26\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c28\"\u003e\u003cp\u003e27\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c29\"\u003e\u003cp\u003e28\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c30\"\u003e\u003cp\u003e29\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c31\"\u003e\u003cp\u003e30\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c32\"\u003e\u003cp\u003e31\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c33\"\u003e\u003cp\u003e32\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eBowdichia virgilioides\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e1/2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e0/1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0,1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c19\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c20\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c21\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c22\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c23\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c24\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c25\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c26\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c27\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c28\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c29\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c30\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c31\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c32\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c33\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDiplotropis matiusii\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e1/2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c19\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c20\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c21\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c22\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c23\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c24\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c25\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c26\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c27\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c28\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c29\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c30\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c31\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c32\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c33\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eGuianodendron praeclarum\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c19\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c20\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c21\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c22\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c23\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c24\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c25\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c26\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c27\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c28\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c29\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c30\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c31\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c32\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c33\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eLeptolobium brachystachyum\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0/1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c19\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c20\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c21\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c22\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c23\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c24\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c25\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c26\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c27\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c28\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c29\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c30\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c31\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c32\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c33\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eLeptolobium bijugum\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c19\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c20\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c21\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c22\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c23\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c24\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c25\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c26\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c27\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c28\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c29\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c30\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c31\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c32\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c33\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eLeptolobium dasycarpum\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c19\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c20\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c21\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c22\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c23\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c24\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c25\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c26\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c27\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c28\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c29\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c30\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c31\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c32\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c33\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eLeptolobium elegans\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e0/2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0/2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c19\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c20\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c21\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c22\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c23\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c24\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c25\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c26\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c27\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c28\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c29\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c30\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c31\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c32\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c33\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eLeptolobium nitens\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0/1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c19\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c20\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c21\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c22\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c23\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c24\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c25\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c26\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c27\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c28\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c29\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c30\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c31\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c32\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c33\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003ePoecilanthe parviflora\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0/2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0/2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e0/3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c19\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c20\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c21\"\u003e\u003cp\u003e?\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c22\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c23\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c24\"\u003e\u003cp\u003e?\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c25\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c26\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c27\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c28\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c29\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c30\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c31\"\u003e\u003cp\u003e?\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c32\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c33\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eSophora tomentosa\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c19\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c20\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c21\"\u003e\u003cp\u003e?\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c22\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c23\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c24\"\u003e\u003cp\u003e?\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c25\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c26\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c27\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c28\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c29\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c30\"\u003e\u003cp\u003e?\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c31\"\u003e\u003cp\u003e?\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c32\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c33\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eStaminodianthus racemosus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e0,4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c19\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c20\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c21\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c22\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c23\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c24\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c25\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c26\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c27\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c28\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c29\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c30\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c31\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c32\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c33\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e\u003ch2\u003eSeedling morphology\u003c/h2\u003e\u003cp\u003eWe recognized three morphological types of seedlings in the studied species of Leptolobieae. \u003cem\u003eBowdichia virgilioides\u003c/em\u003e and \u003cem\u003eLeptolobium\u003c/em\u003e spp. have phanero-epigeo-foliaceous seedlings (PEF), whereas \u003cem\u003eG. praeclarum\u003c/em\u003e and \u003cem\u003eD. martiusii\u003c/em\u003e have crypto-hipogeo-reserve seedlings (CHR) and \u003cem\u003eS. racemosu\u003c/em\u003es have phanero-hipogeo-reserve seedlings (PHR) (Rodrigues e Tozzi, 2007b; 2008).\u003c/p\u003e\u003cp\u003eThe species of Leptolobieae studied here can be distinguished by the shape and length of the hypocotyl. In terms of elevation of the cotyledons, they can be divided in two groups: seedlings with quadrangular hypocotyl (\u003cem\u003eBowdichia virgilioides\u003c/em\u003e and \u003cem\u003eLeptolobium\u003c/em\u003e spp.), and seedlings with cylindrical hypocotyl ( \u003cem\u003eG. praeclarum, D. martiusii and S. racemosus).\u003c/em\u003e The size of the hypocotyl is also a diagnostic feature, with \u003cem\u003eBowdichia virgilioides\u003c/em\u003e and \u003cem\u003eLeptolobium\u003c/em\u003e spp. having a medium to long-sized hypocotyl (12\u0026ndash;62 mm) and species of \u003cem\u003eG. praeclarum, D. martiusii and S. racemosus\u003c/em\u003e with much smaller hypocotyls (1\u0026ndash;2 mm).\u003c/p\u003e\u003cp\u003eThe epicotyl is cylindrical in all studied species of Leptolobieae, but its size is variable. In \u003cem\u003eBowdichia virgilioides\u003c/em\u003e and \u003cem\u003eLeptolobium\u003c/em\u003e spp., the epicotyl is short, but conspicuously longer in \u003cem\u003eG. praeclarum, D. martiusii and S. racemosus.\u003c/em\u003e\u003c/p\u003e\u003cp\u003eCotyledons can either function as photosynthetic sites or storage organs to assure an ideal growth of seedlings during initial stages of development (Garwood, 1995). Seedlings of \u003cem\u003eB. virgilioides\u003c/em\u003e and \u003cem\u003eLeptolobium\u003c/em\u003e spp. have leafy cotyledons, whereas \u003cem\u003eS. racemosus\u003c/em\u003e has thick, storage-rich cotyledons. The insertion point between hypocotyl and epycotyl, a feature associated to hypocotyl elongation, is another diagnostic character in Leptolobieae. In \u003cem\u003eB. virgilioides\u003c/em\u003e and \u003cem\u003eLeptolobium\u003c/em\u003e spp., the insertion point is located above the surface, but in \u003cem\u003eS. racemosus\u003c/em\u003e it is found at the ground level.\u003c/p\u003e\u003cp\u003eThe seedlings of \u003cem\u003eG. praeclarum\u003c/em\u003e and \u003cem\u003eD. martiusii\u003c/em\u003e have cotyledons hidden into the seminal remains. The endosperm stored at these cotyledons function as a nutritive tissue for vertical development of the seedling.\u003c/p\u003e\u003cp\u003eCotyledon shape in phanerocotiledonary seedlings are elliptic in \u003cem\u003eB. virgilioides\u003c/em\u003e and \u003cem\u003eLeptolobium\u003c/em\u003e spp. and ovate in \u003cem\u003eS. racemosus\u003c/em\u003e. The base is obtuse in the three genera, but it can also be slightly oblique or cordate in \u003cem\u003eB. virgilioides\u003c/em\u003e and \u003cem\u003eLeptolobium\u003c/em\u003e spp.\u003c/p\u003e\u003cp\u003eNictinasty was not observed in the cotyledons from all analyzed taxa, but was present in the eophyls of species of \u003cem\u003eB. virgilioides\u003c/em\u003e, \u003cem\u003eS. racemosus\u003c/em\u003e and \u003cem\u003eLeptolobium\u003c/em\u003e spp. \u003cem\u003e(e\u003c/em\u003except \u003cem\u003eL. dasycarpum\u003c/em\u003e), In that taxa, the adaxial face of the leaflets are exposed by the descendent orientation of the eophyls.\u003c/p\u003e\u003cp\u003eIn \u003cem\u003eD. martiusii\u003c/em\u003e, the eophyl morphology differs significantly during plant development. Seedlings have a single leaflet, but young plants have compound leaves with several, imparipinated leaflets. The number of eophyls from the first to third nodes is highly variable, being constant only in \u003cem\u003eD. martiusii, L. elegans, L. dasycarpum\u003c/em\u003e and \u003cem\u003eL. bijugum\u003c/em\u003e, with one eophyl per node. In all taxa, eophyls have broquidodromous venation.\u003c/p\u003e\u003cp\u003eEophyls/leaflets were elliptic in \u003cem\u003eD. martiusii, G. praeclarum\u003c/em\u003e and \u003cem\u003eS. racemosus\u003c/em\u003e, ovate in \u003cem\u003eB. virgilioides, L. panamense, L. bijugum, L. dasycarpum\u003c/em\u003e and \u003cem\u003eL. nitens\u003c/em\u003e, cordate in \u003cem\u003eL. elegans\u003c/em\u003e and oblong in \u003cem\u003eL. brachystachyum\u003c/em\u003e. The apex was acute in \u003cem\u003eL. nitens, L. dasycarpum\u003c/em\u003e e \u003cem\u003eL. panamense\u003c/em\u003e, obtuse in \u003cem\u003eL. bijugum, L. brachystachyum, L. elegans\u003c/em\u003e and acuminate in \u003cem\u003eD. martiusii\u003c/em\u003e and \u003cem\u003eG. praeclarum.\u003c/em\u003e In \u003cem\u003eB. virgilioides\u003c/em\u003e and \u003cem\u003eS. racemosus\u003c/em\u003e, eophyls/leaflets were mainly obtuse, but acute and rounded apices were ocasionally observed in the former and latter, respectively. The base was always symetrical, obtuse to rounded in \u003cem\u003eB. virgilioides\u003c/em\u003e and \u003cem\u003eD. martiusii\u003c/em\u003e, obtuse in \u003cem\u003eL. nitens, L. dasycarpum, L. brachystachyum, L. bijugum\u003c/em\u003e and \u003cem\u003eS. racemosus\u003c/em\u003e, cuneate in \u003cem\u003eG. praeclarum\u003c/em\u003e, and cordate in \u003cem\u003eL. elegans.\u003c/em\u003e\u003c/p\u003e\u003cp\u003eCanaliculate petiole and rachis were observed in \u003cem\u003eB. virgilioides, D. martiusii\u003c/em\u003e and \u003cem\u003eS. racemosus.\u003c/em\u003e Intercotyledonary glands were observed in \u003cem\u003eB. virgilioides\u003c/em\u003e and \u003cem\u003eLeptolobium\u003c/em\u003e spp., but absent in the remaining taxa. Stipels were observed in all taxa, being linear in \u003cem\u003eD. martiusii\u003c/em\u003e and \u003cem\u003eS. racemosus\u003c/em\u003e, linear-lanceolate in \u003cem\u003eB. virgilioides\u003c/em\u003e and \u003cem\u003eLeptolobium\u003c/em\u003e spp., and ovate in \u003cem\u003eG. praeclarum.\u003c/em\u003e Stipels were observed in \u003cem\u003eB. Virgilioides, D. martiusii, G. praeclarum\u003c/em\u003e and \u003cem\u003eLeptolobium nitens\u003c/em\u003e, but were absent in the remaining taxa.\u003c/p\u003e\u003cp\u003eLeaves with opposite phylotax\u003cem\u003ey\u003c/em\u003e at the first node is a shared feature among \u003cem\u003eB. Virgilioides, L. nitens, L. brachystachyum, L. bijugum\u003c/em\u003e and \u003cem\u003eS. racemosum\u003c/em\u003e,\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eMorphological description of fruits, seeds and seedlings of Staminodianthus racemosus\u003c/h3\u003e\n\u003cp\u003eFruit samaroid, one-seeded, greenish when young and brown when mature and dry. oblong, apex caudate, base cuneate, venation reticulated and conspicuous on both faces. Seeds green, glabrous, rounded and flattened, 11\u0026ndash;15 x 9\u0026ndash;12 mm (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Seedling of phanero-hypogeo-reserve type (PHR), germinating at the 15th day after sowing; cotyledons green, glabrous, sessile, 11 x 9 x 2 mm, nictinasty absent. Hipocotyl cylindric, short, about 1 mm length and 2,5 mm width, glabrous; epicotyl cylindric, light green, about 66 mm length and 2 mm width, sparsely pubescent; cataphylls 2, opposite, 5 x 1,5 mm, caducous, slightly canaliculate, located at the intermediary region between the hypocotyl and first node. Eophyls alternate, 3-foliolate at the first node, 5-foliolate at the second node and 57-foliolate at the third node, leaflets 9\u0026ndash;18 x 6\u0026ndash;10 mm, elliptic, apex obtuse to rounded, base obtuse, rachis and petiole canaliculate; stiple linear, 2mm, stipels absent, venation brochidodromous, nictinasty present, descendant orientation (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eThe seedling morphological type, cotyledon shape, number of leaflets in the three nodes of the eophyl, the absence of stipels and presence of cataphylls are diagnostic features of the genus that allows its recognition within Leptolobieae.\u003c/p\u003e\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e\u003ch2\u003ePhylogenetic analysis\u003c/h2\u003e\u003cp\u003eLeptolobieae was recovered as monophyletic, with all sampled taxa strongly supported (BS\u0026thinsp;=\u0026thinsp;1.0) on a single clade (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The five sampled taxa of \u003cem\u003eLeptolobium\u003c/em\u003e are nested into a strongly supported clade (BS\u0026thinsp;=\u0026thinsp;0.96) sister to \u003cem\u003eBowdichia virgilioides.\u003c/em\u003e A sister relationship of \u003cem\u003eGuianodendron\u003c/em\u003e and \u003cem\u003eStaminodianthus\u003c/em\u003e was also recovered (BS\u0026thinsp;=\u0026thinsp;1.0), as well as the sister relationship of this clade to \u003cem\u003eDiplotropis\u003c/em\u003e (BS\u0026thinsp;=\u0026thinsp;1.0). Similar results were observed in both individual and combined datasets.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec12\" class=\"Section2\"\u003e\u003ch2\u003eMorphological matrix and character state reconstruction\u003c/h2\u003e\u003cp\u003eOf the 32 coded characters, 20 are related to the seedling morphology and 12 to inflorescence/flower morphology (Tables\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e, 3). Six characters from seedlings were reconstructed as potential synapomorphies for clades within Leptolobieae. The PEF plantlet morphotype (char. 1, state 1) is the mostly likely state in the most recent common ancestor (MRCA) of Leptolobieae. Transition from PEF to CHR (char. 1, state 0) may have occurred once in the MRCA common ancestor of the \u003cem\u003eDiplotropus\u003c/em\u003e\u0026thinsp;+\u0026thinsp;\u003cem\u003eGuianodendron\u003c/em\u003e\u0026thinsp;+\u0026thinsp;\u003cem\u003eStaminodianthus\u003c/em\u003e clade, or, alternatively, twice in this clade, once in \u003cem\u003eDiplotropus\u003c/em\u003e and again in \u003cem\u003eGuianodendron\u003c/em\u003e. The PHR morphotype (char. 1, state 2) is an autapomorphy of \u003cem\u003eStaminodianthus\u003c/em\u003e, but our reconstruction is equivocal whether it have evolved from PEF or CHR types. Quadrangular (character 5, state 1) and long (character 6, state 1) hypocotyls were reconstructed as the most likely states at the MRCA of \u003cem\u003eBowdichia\u003c/em\u003e\u0026thinsp;+\u0026thinsp;\u003cem\u003eLeptolobium\u003c/em\u003e, probably evolving from cylindric, shorter hypocotyls. Epicotyls with a length of 70 mm or longer (character 7, state 1) is the most likely condition in the MRCA of Leptolobieae, from which epicotyls shorter than 40 mm (character 7, state 0), has unambiguously evolved in the MRCA of \u003cem\u003eBowdichia\u0026thinsp;+\u0026thinsp;Leptolobium\u003c/em\u003e. The presence of glands at the eophyl axyl (character 12, state 1) is a novelty for \u003cem\u003eBowdichia\u0026thinsp;+\u0026thinsp;Leptolobium\u003c/em\u003e and may have evolved once in the MRCA of this clade, as this feature is not present in any other genera of Leptolobieae. The presence of three leaflets at the seedling first node (char. 11, state 1), instead of one, four or 12, is an autapomorphy of \u003cem\u003eGuianodendron\u003c/em\u003e.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eA number of morphological features from adult were unambiguously reconstructed as synapomorphies for genera in Leptolobieae: presence of pulvin (char. 2, state 1), cataphylls (char. 19, state 1), staminodes (char. 25, state 1), and racemose inflorescences (char. 31, state 0) for \u003cem\u003eStaminodianthus\u003c/em\u003e; flowers lilac to blue (char. 22, state 0) in \u003cem\u003eBowdichia\u003c/em\u003e; flowers actinomorphic (char. 21, state 1) and early senescent bracts (char. 27, state 0) in \u003cem\u003eLeptolobium\u003c/em\u003e; presence of lenticells (char. 4, state 1), linear stipels (15, state 0) and assymetric basal leaflets (char. 17, state 0) in \u003cem\u003eDiplotropis\u003c/em\u003e; and stipels ovate (char. 15, state 2) and flowers actinomorphic (char. 21, state 1) in \u003cem\u003eGuianodendron\u003c/em\u003e. Other features reconstructed as synapomorphies for major clades are the presence of only five fertile stamens (char. 24, state 1) in \u003cem\u003eGuianodendron\u003c/em\u003e\u0026thinsp;+\u0026thinsp;\u003cem\u003eStaminodianthus\u003c/em\u003e, seeds with arils (char. 28, state 1) in \u003cem\u003eDiplotropis\u003c/em\u003e\u0026thinsp;+\u0026thinsp;\u003cem\u003eGuianodendron\u003c/em\u003e\u0026thinsp;+\u0026thinsp;\u003cem\u003eStaminodianthus\u003c/em\u003e, and samaroid fruits (char. 26, state 2) and seeds with endosperm (char. 29, state 1) in \u003cem\u003eBowdichia\u0026thinsp;+\u0026thinsp;Leptolobium.\u003c/em\u003e\u003c/p\u003e\u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eSeedlings have a number of morphological features that are variable among taxa and can provide useful information for taxonomic and evolutionary studies. In Leguminosae, seedlings have been extensively studied in several genera and provide useful characters for delimitation of groups at different taxonomic levels (e.g., Ye, 1983; Lima, 1990; Oliveira, 2001; Rodriguez \u0026amp; Tozzi 2007). In Leptolobieae and related tribes, Rodriguez \u0026amp; Tozzi (2008) used seedling characters from species of \u003cem\u003eAcosmium, Leptolobium\u003c/em\u003e and \u003cem\u003eGuianodendron\u003c/em\u003e to evaluate the systematic value of these characters. Later on, Cardoso et al. (2012, 2013a) has used seedling characters on a phylogenetic context to detect potential synapomorphies and evolutionary trends in Leptolobieae. Our study follow that approach by adding new information from seedling morphology to a simplified phylogeny of Leptolobieae, in which all genera from the tribe were sampled.\u003c/p\u003e\u003cp\u003eOur results support to the recognition of \u003cem\u003eStaminodianthus\u003c/em\u003e as a distinct genus from \u003cem\u003eDiplotropis\u003c/em\u003e and its placement as sister to \u003cem\u003eGuianodendron.\u003c/em\u003e Elevation of \u003cem\u003eDiplotropis\u003c/em\u003e sect. \u003cem\u003eracemosae\u003c/em\u003e to the generic rank was proposed by Cardoso et al. (2013a), based on molecular and morphological evidence. The authors reported six apomorphies unique to \u003cem\u003eStaminodianthus\u003c/em\u003e within the Genistoid legumes, all related to the flower architecture: actinomorphic symmetry calyx with a curved hypanthium, well-differentiated standard petal, lateral and lower petals without auricles, and androecium with five fertile stamens and five staminodes. At the time of this publication, seedlings of \u003cem\u003eStaminodianthus\u003c/em\u003e were unkown to science, so that there was no information available in the literature to compare with seedlings from other genera in the tribe. Our observation and description of seedlings of \u003cem\u003eS. racemosus\u003c/em\u003e has provided additional evidence supporting recognition of the genus. Its morphotype, phanero-hypogeo-reserve type (PHR), have thick, expanded cotyledons at the ground level with energetic reserve, an unusual condition exclusive of this genus in Leptolobieae. In addition, seedlings of \u003cem\u003eS. racemosus\u003c/em\u003e share six features with seedlings from the sampled species of \u003cem\u003eDiplotropis\u003c/em\u003e and \u003cem\u003eGuianodendron\u003c/em\u003e, reinforcing the phylogenetic affinities of the three genera.\u003c/p\u003e\u003cp\u003eA close relationship of \u003cem\u003eBowdichia\u003c/em\u003e and \u003cem\u003eLeptolobium\u003c/em\u003e is also supported by our results. Despite having distinct floral features, which has placed them in different tribes in traditional classifications of Leguminosae, the phylogenetic proximity of these two genera is strongly supported in studies focused on the Genistoid and Leptolobieae clades (e.g., Cardoso et al. 2012, 2013). These studies also reported two features from seedlings (presence of intercotiledonary glands and quadrangular hypocotyls) shared by the two genera that represent potential synapomorphies. Our study corroborated these results and reconstructed two additional character states from seedlings as unequivocal synapomorphies for the \u003cem\u003eBowdichia\u0026thinsp;+\u0026thinsp;Leptolobium\u003c/em\u003e clade: epicotyls shorter than 40 mm and hypocotyls longer than 11 mm.\u003c/p\u003e\u003cp\u003eWithin \u003cem\u003eLeptolobium\u003c/em\u003e, the phyllotaxy, number of eophylls from first to third node, leaflet shape, apex and base form, and the presence of stipels were variable among the five sampled taxa and thus potentially useful for species delimitation. For that reason, we provide here an identification key based solely on seedling characters. Our sampling, however, is lower than 50% of the valid taxa in \u003cem\u003eLeptolobium\u003c/em\u003e (13 species), so that the taxonomic value of these features could not be properly tested. A larger sampling within the genus is required to address this question in future studies and to build a more complete identification key.\u003c/p\u003e\u003cdiv id=\"Sec14\" class=\"Section2\"\u003e\u003ch2\u003eKey to the species of Leptolobium\u003c/h2\u003e\u003cp\u003e1 Stipels present, hipocotyl 58-62mm length, epicotyl 35-40mm length .............. \u003cem\u003eL. nitens.\u003c/em\u003e\u003c/p\u003e\u003cp\u003e1\u0026rsquo; Stipels absent, hipoc\u0026oacute;tilo 17-37mm length, epicotyl 8-28mm length ...............2\u003c/p\u003e\u003cp\u003e2 Phylotaxy of the eophyl from the first node opposite ......................................................................3\u003c/p\u003e\u003cp\u003e3 Nictinasty of leaflets present, hipocotyl 35-37mm length, blade ovate, apex obtuse, base cordate ....................... \u003cem\u003eL. elegans\u003c/em\u003e.\u003c/p\u003e\u003cp\u003e3\u0026rsquo; Nictinasty of leaflets absent, hipocotyl 22-33mm length, blade cordate, apex acute, base obtuse ...................... \u003cem\u003eL. dasycarpum\u003c/em\u003e.\u003c/p\u003e\u003cp\u003e2\u0026rsquo; Phylotaxy of the eophyl from the first node alternate..................................................4\u003c/p\u003e\u003cp\u003e4 Cotyledons 5-10mm width, secondary veins with angles of 60\u0026ndash;70\u0026ordm; to the primary vein, blade ovate .............................................\u003cem\u003eL. bijugum\u003c/em\u003e.\u003c/p\u003e\u003cp\u003e4\u0026rsquo;. Cotyledons 11-12mm width, secondary veins forming angles of 70\u0026ndash;85\u0026ordm; to the primary vein, blade oblong ............................................L. \u003cem\u003ebrachystachyum\u003c/em\u003e.\u003c/p\u003e\u003cp\u003eOur study has reinforced the systematic value of seedling features in Leguminosae, specially in tribe Leptolobieae. Morphological description of seedlings of \u003cem\u003eStaminodianthus racemosus\u003c/em\u003e has provided additional evidence of the validation of its generic status and close relationship to \u003cem\u003eGuianodendron\u003c/em\u003e and \u003cem\u003eDiplotropis\u003c/em\u003e. Observation and description of seedlings from five species of \u003cem\u003eLeptolobium\u003c/em\u003e demonstrates that several seedling features are constant at the specific level but variable within species and therefore can be used for systematic purposes. Future studies including more species from both Leptolobieae and Genistoid clade, with description of their seedling morphology and further character mapping on a phylogenetic approach, are greatly desired to improve our knowledge on the systematics and evolution of these lineages of Leguminosae.\u003c/p\u003e\u003c/div\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAuthor Information\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthors and Affiliations\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eUniversidade Federal Rural da Amaz\u0026ocirc;nia / Museu Paraense Em\u0026iacute;lio Goeldi, Botany Department, Bel\u0026eacute;m, Par\u0026aacute;, Brazil.\u003c/p\u003e\n\u003cp\u003ehttps://orcid.org/0000-0002-9715-158X\u003c/p\u003e\n\u003cp\u003eJaciara Cerqueira da Silva\u003c/p\u003e\n\u003cp\u003eMuseu Paraense Em\u0026iacute;lio Goeldi, Ministry of Science and Technology, Botany Department, Bel\u0026eacute;m, Par\u0026aacute;, Brazil.\u003c/p\u003e\n\u003cp\u003ehttps://orcid.org/0000-0002-9488-7532\u003c/p\u003e\n\u003cp\u003eEly Simone Cajueiro Gurgel\u003c/p\u003e\n\u003cp\u003eUniversidade Estadual de Campinas, Instituto de Biologia, Campinas, S\u0026atilde;o Paulo, Brazil.\u003c/p\u003e\n\u003cp\u003ehttps://orcid.org/0000-0001-6555-8759\u003c/p\u003e\n\u003cp\u003eAndr\u0026eacute; Olmos Sim\u0026otilde;es\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCorresponding Author\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eCorrespondence to: Jaciara Cerqueira da Silva \u0026ndash; [email protected]\u003c/p\u003e\n\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eScholarship granted by the Conselho Nacional de Desenvolvimento Cient\u0026iacute;fico e Tecnol\u0026oacute;gico (CNPq), process no. 132296/2020-9.\u003c/p\u003e\n\u003cp\u003eThis research is part of the master\u0026rsquo;s dissertation of the first author, conducted within the Graduate Program in Biological Sciences of the Universidade Federal Rural da Amaz\u0026ocirc;nia (UFRA) / Museu Paraense Em\u0026iacute;lio Goeldi (MPEG), Bel\u0026eacute;m, Par\u0026aacute;, Brazil.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDeclarations and Ethical Statements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eEach author made a substantial contribution to the conception and design of the study, as well as to data acquisition, analysis, and interpretation. All authors have approved the final version of the manuscript for publication. The authors declare that they have no conflicts of interest, financial or otherwise, that could be perceived as influencing the content of this article.\u003c/p\u003e\u003cp\u003e\u003cstrong\u003eConflict of interest\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eEach author made a substantial contribution to the conception and design of the study, as well as to data acquisition, analysis, and interpretation. All authors have approved the final version of the manuscript for publication. The authors declare that they have no conflicts of interest, financial or otherwise, that could be perceived as influencing the content of this article.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor Contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eCollection and monitoring of seedling development, manuscript structuring, and writing (JCS). Morphological conceptualization of the group, data validation, and manuscript review (ESCG). Critical analysis of the text regarding phylogenetic systematics, data validation, english writing and final formatting (AOS).\u003c/p\u003e\u003cp\u003e\u003cstrong\u003eAcknowledgments\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe thank Dr. Mathias Engels for collecting botanical material of \u003cem\u003eStaminodianthus racemosus.\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eWe are also grateful to Dr. Felipe Martins for preparing the illustrations.\u003c/p\u003e\n\u003cp\u003eThis study was supported by the Conselho Nacional de Desenvolvimento Cient\u0026iacute;fico e Tecnol\u0026oacute;gico (CNPq), which provided a research scholarship.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAzani N, Babineau M, Bailey CD, Banks H, Barbosa AR, Pinto RB et al. 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American journal of botany 91: 1846\u0026ndash;1862. http://dx.doi.org/10.3732/ajb.91.11.1846.\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eYahara T, Javadi F, Onoda Y, De Queiroz LP, Faith DP, Prado DE (2013) Global legume diversity assessment: Concepts, key indicators, and strategies. Taxon 62: 249\u0026ndash;266. https://doi.org/10.12705/622.12.\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eYakovlev GP (1976). Survey of genera \u003cem\u003eZollernia\u003c/em\u003e Wied-Neuw. \u0026amp; Nees and \u003cem\u003eLecointea\u003c/em\u003e Ducke. Botanicheskii Zhurna 61: 1304\u0026ndash;1308.\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eYe, N. 1983. Descriptions of various seedlings of leguminous plants. Phytologia 54: 190\u0026ndash;218.\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"ancestral character state reconstruction, Leptolobieae, seedling, Staminodianthus","lastPublishedDoi":"10.21203/rs.3.rs-7973373/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7973373/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eMolecular phylogenetic evidence demonstrate the monophyly of five genera (\u003cem\u003eBowdichia, Diplotropis, Guianodendr on, Leptolobium\u003c/em\u003e and \u003cem\u003eStaminodianthus\u003c/em\u003e) into a recircumscribed Leptolobieae. Even though recognition of the tribe is highly supported by molecular data, morphological characterization of its taxa is still challenging. In this context, our study aimed to study the seedling morphology of eleven species from all genera of Leptolobieae. The seedling morphology of \u003cem\u003eStaminodianthus racemosus\u003c/em\u003e is here described for the first time. The observed characters were scored into a morphological matrix and mapped onto a phylogeny in order to detect potential synapomorphies for major clades. Three morphological types of seedlings were observed and described: phanero-epigeo-foliaceous seedlings (PEF) in \u003cem\u003eBowdichia virgilioides\u003c/em\u003e and \u003cem\u003eLeptolobium\u003c/em\u003e spp., crypto-hipogeo-reserve seedlings (CHR) in \u003cem\u003eGuianodendron praeclarum\u003c/em\u003e and \u003cem\u003eDiplotropis martiusii, and\u003c/em\u003e phanero-hipogeo-reserve seedlings (PHR) in \u003cem\u003eS. racemosu\u003c/em\u003es. PEF morphotype was reconstructed as the mostly likely state in the most recent common ancestor (MRCA) of Leptolobieae, with subsequent transitions to the CHR morphotype in \u003cem\u003eDiplotropus\u003c/em\u003e and \u003cem\u003eGuianodendron\u003c/em\u003e, and to the PHR morphotype in \u003cem\u003eS. racemosus\u003c/em\u003e. Another five seedling characters were reconstructed as potential synapomorphies for clades in Leptolobieae: hypocotyl shape and length, epicotyl length, presence of glands at the eophyl axyl and the number of leaflets at the seedling first node. For \u003cem\u003eLeptolobium\u003c/em\u003e, the phyllotaxy, number of eophylls from first to third node, leaflet shape, apex and base form, and the presence of stipels are diagnostic characters within species and could be used for taxonomic purposes.\u003c/p\u003e","manuscriptTitle":"Morphology and character evolution of seedlings in Leptolobieae (Leguminosae, Papilionoideae)","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-12-06 00:15:36","doi":"10.21203/rs.3.rs-7973373/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"ffcc6c9d-3b3c-40f8-913a-98e07ef8def7","owner":[],"postedDate":"December 6th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2026-01-16T18:38:30+00:00","versionOfRecord":[],"versionCreatedAt":"2025-12-06 00:15:36","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-7973373","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7973373","identity":"rs-7973373","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}