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Research Article
Misleading fruits: The non-monophyly of Pseudopiptadenia and Pityrocarpa supports generic re-circumscriptions and a new genus within mimosoid legumes
expand article infoLeonardo M. Borges‡, Peter W. Inglis§, Marcelo F. Simon§, Pétala Gomes Ribeiro|, Luciano P. de Queiroz|
‡ Universidade Federal de São Carlos, São Carlos, Brazil
§ Embrapa Recursos Genéticos e Biotecnologia, Parque Estação Biológica, Brasília, Brazil
| Universidade Estadual de Feira de Santana, Feira de Santana, Brazil

Corresponding author: Leonardo M. Borges ( aquitemcaqui@gmail.com )

Academic editor: Colin E. Hughes
© 2022 Leonardo M. Borges, Peter W. Inglis, Marcelo F. Simon, Pétala Gomes Ribeiro, Luciano P. de Queiroz.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Borges LM, Inglis PW, Simon MF, Ribeiro PG, de Queiroz LP (2022) Misleading fruits: The non-monophyly of Pseudopiptadenia and Pityrocarpa supports generic re-circumscriptions and a new genus within mimosoid legumes. In: Hughes CE, de Queiroz LP, Lewis GP (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae Part 1: New generic delimitations. PhytoKeys 205: 239-259. https://doi.org/10.3897/phytokeys.205.82275
Open Access

Abstract

Generic delimitation in Piptadenia and allies (mimosoid legumes) has been in a state of flux, particularly caused by over-reliance on fruit and seed morphology to segregate species out of Piptadenia into the genera Parapiptadenia, Pityrocarpa and Pseudopiptadenia. Although supporting their segregation from Piptadenia, previous phylogenetic analyses suggested that some of these segregated genera are not monophyletic. Here, we test the monophyly of Parapiptadenia, Pityrocarpa and Pseudopiptadenia with dense taxon sampling across these genera, including the type species of each genus. Our analysis recovers Parapitadenia as monophyletic, but places Pseudopiptadenia species in two distinct lineages, one of which includes all three species of Pityrocarpa. Given that the type species of both Pseudopiptadenia and Pityrocarpa are nested in the same clade, we subsume Pseudopiptadenia under the older name Pityrocarpa. The remaining Pseudopiptadenia species are assigned to the new genus Marlimorimia. Alongside high molecular phylogenetic support, recognition of Parapiptadenia, Pityrocarpa and Marlimorimia as distinct genera is also supported by combinations of morphological traits, several of which were previously overlooked.

Keywords

Caesalpinioideae, Fabaceae, Leguminosae, Parapiptadenia, Stryphnodendron clade, tropical America

Introduction

Generic delimitation in the mimosoid legumes is being continually revised, notably across the informal Piptadenia group sensu Lewis and Elias (1981), which included Anadenanthera Speg., Microlobius C. Presl, Mimosa L., Parapiptadenia Brenan, Piptadenia Benth., Pityrocarpa (Benth.) Britton & Rose, Pseudopiptadenia Rauschert and Stryphnodendron Mart. Most of the proposed generic re-circumscriptions within the Piptadenia group have involved segregating species out of Piptadenia, which was morphologically poorly-defined (Brenan 1955) and is known to be polyphyletic (Luckow et al. 2003; Jobson and Luckow 2007; Simon et al. 2016; Ribeiro et al. 2018). While previous phylogenetic and phylogenomic analyses confirm the segregation of Parapiptadenia, Pityrocarpa and Pseudopiptadenia and place them together with Stryphnodendron and Microlobius in the Stryphnodendron clade sensu Koenen et al. (2020), the monophyly of these three genera is still uncertain because of incomplete taxon sampling in previous analyses (Simon et al. 2016; Koenen et al. 2020; Ringelberg et al. 2022).

Species of Parapiptadenia, Pityrocarpa and Pseudopiptadenia are trees inhabiting Neotropical rain forests and seasonally dry tropical forests and woodlands (SDTFWs sensu Queiroz et al. 2017), with the majority of species in South America and just two taxa in North America (Pi. obliqua (Pers.) Brenan var. obliqua and Ps. psilostachya (DC.) G.P. Lewis & M.P. Lima) (Brenan 1955, 1963; Rauschert 1982; Lima and Lima 1984; Lewis and Lima 1991; Queiroz 2009). Their bipinnate leaves vary widely in the number of pinnae, as well as leaflet number, size and shape. Flowers are pentamerous, dialipetalous or gamopetalous and arranged in elongated spikes. The diverse fruits and seeds have been the most prominent traits used to define each genus (Brenan 1955; Lewis and Elias 1981). Parapiptadenia includes six species with plano-compressed fruits opening along both sutures (typical legumes) and flat, compressed, narrowly-winged seeds lacking a pleurogram. Eleven species with similar seeds, but with follicles (fruits splitting along the upper suture only) were placed in Pseudopiptadenia (Rauschert 1982; Lewis and Lima 1991). The three species in Pityrocarpa, which was first proposed as a section of Piptadenia (Bentham 1842), differ from the other two genera by their regularly constricted moniliform legumes and lentiform whitish seeds with an U-shaped pleurogram (Jobson and Luckow 2007).

The first phylogenetic analysis including these three genera recovered each as monophyletic, with Pseudopiptadenia contorta (DC.) G.P. Lewis & M.P. Lima and Ps. psilostachya forming a clade sister to Pityrocarpa (three species sampled), while the relationship of Parapiptadenia (three species sampled) to other genera was uncertain (Fig. 1; Jobson and Luckow 2007). The relationships amongst these genera and the putative monophyly of Pseudopiptadenia were later questioned by analyses with larger DNA sequence datasets and increased taxon sampling (Simon et al. 2016; Ribeiro et al. 2018). In these analyses, Parapiptadenia (four species sampled) emerged as sister to a clade including all sampled species of Pseudopiptadenia (five species, including Ps. contorta and Ps. psilostachya), except Ps. brenanii G.P. Lewis & M.P. Lima, which was sister to Pityrocarpa (Fig. 1). This latter clade appeared more closely related to Stryphnodendron and Microlobius than to the group formed by Parapiptadenia and Pseudopiptadenia. Phylogenomic analyses with sparse taxonomic sampling recovered slightly different relationships between these three genera (Fig. 1), but reinforced the non-monophyly of Pseudopiptadenia (Lima et al. 2022; Ringelberg et al. 2022).

Figure 1. 

Topological differences amongst phylogenetic analyses of the Stryphnodendron clade.

While it is clear that the non-monophyly of Pseudopiptadenia means that taxonomic adjustments are needed, the type species of the genus, Ps. leptostachya, has not been included in any previous phylogenetic analyses, raising doubts about its placement and, hence, about which generic name should be applied to the clade containing that species. In this study, we infer the phylogenetic relationships between Parapiptadenia, Pityrocarpa and Pseudopiptadenia using near-complete taxon sampling, including the type species of all three genera, and re-evaluate the circumscriptions of these genera, based on the resulting phylogenetic hypothesis.

Materials and methods

Phylogenetic inference

To further test the polyphyly of Pseudopiptadenia indicated by previous studies (Simon et al. 2016; Ribeiro et al. 2018; Ringelberg et al. 2022) and further investigate sister group relationships across the Stryphnodendron clade, we carried out phylogenetic analyses including near-complete sampling of species of Parapiptadenia, Pityrocarpa, Pseudopiptadenia and allies. Phylogenetic analyses were based on the nuclear ribosomal 5.8S subunit and internal transcribed spacer region (nrITS) and plastid regions matK and trnD-trnT. We generated 60 new sequences (21 nrITS, 23 matK, 16 trnD-trnT), including two accessions of Ps. leptostachya, the type species of Pseudopiptadenia, sampled here for the first time. Published sequences of other members of the Stryphnodendron clade and other genera were obtained from GenBank (Hughes et al. 2003; Simon et al. 2009; Simon et al. 2016; LPWG 2017; Ribeiro et al. 2018). Sampling comprised 60 accessions, including nine species (18 accessions) of Pseudopiptadenia (only the poorly known Ps. colombiana and Ps. pittieri were not sampled), all three species of Pityrocarpa (six accessions), all six known species of Parapipitadenia (11 accessions), plus representatives of the allied genera Microlobius (monospecific; two accessions) and Stryphnodendron (14 accessions, including members of the three major lineages of this non-monophyletic genus; see Lima et al. 2022). A selection of mimosoid lineages closely related to the Stryphnodendron clade (Jobson and Luckow 2007; Simon et al. 2016; Ribeiro et al. 2018; Ringelberg et al. 2022) were included as outgroups. Voucher details and GenBank accession numbers are provided in Table 1 and in the Suppl. material 1.

Table 1.
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Voucher information and GenBank accession numbers for taxa used in this study. Newly-generated sequences are in bold. See the Suppl. material 1 for a digital version.

Taxon Voucher Herbarium nrITS matK trnD-trnT
Inga edulis Mart. Queiroz 13797; Pennington 13282 HUEFS; K JX870764 AF523078 JQ417383
Lachesiodendron viridiflorum (Kunth) P.G. Ribeiro, L.P. Queiroz & Luckow Queiroz 13090 HUEFS MG001274 MG001286 MG001305
Microlobius foetidus (Jacq.) M. Sousa & G. Andrade Hughes 2150 FHO KT364047 KT364172 FJ981976
Microlobius foetidus (Jacq.) M. Sousa & G. Andrade Macqueen 432 FHO AF458783 AF523095 (No data)
Mimosa palmeri Rose Simon 823 FHO KT364059 KT364212 FJ982142
Mimosa pigra L. Hughes 2414 FHO KT364060 KT364213 FJ982148
Mimosa ursina Mart. Simon 704 CEN KT364061 KT364210 FJ982217
Parapiptadenia blanchetii (Benth.) Vaz & M.P. Lima Queiroz 15358 HUEFS OM575100 ON409904 ON409927
Parapiptadenia blanchetii (Benth.) Vaz & M.P. Lima Thomas 12372 NY OM575099 ON409905 (No data)
Parapiptadenia excelsa (Griseb.) Burkart Hughes 2425 FHO KT364062 KT364160 FJ982235
Parapiptadenia ilheusana G.P. Lewis Neves 1659 RB OM575101 KY046081 ON409928
Parapiptadenia pterosperma (Benth.) Brenan Cardoso 2359 HUEFS OM575102 ON409906 ON409929
Parapiptadenia pterosperma (Benth.) Brenan Ribeiro 902 HUEFS MG001260 ON409910 MG001292
Parapiptadenia rigida (Benth.) Brenan Marestoni 26 HUEFS MG001261 ON409909 (No data)
Parapiptadenia zehntneri (Harms) M.P. Lima & H.C. Lima Cotarelli 2029 HUEFS OM575104 ON409907 (No data)
Parapiptadenia zehntneri (Harms) M.P. Lima & H.C. Lima Pereira-Silva 3102 CEN KT364063 KT364063 KT364108
Parapiptadenia zehntneri (Harms) M.P. Lima & H.C. Lima Queiroz 10974 HUEFS OM575105 ON409908 (No data)
Parapiptadenia zehntneri (Harms) M.P. Lima & H.C. Lima Queiroz 15692 HUEFS OM575106 KX302341 (No data)
Piptadenia gonoacantha (Mart.) J.F. Macbr. Simon 735 FHO KT364065 DQ790620 FJ982238
Piptadenia stipulacea (Benth.) Ducke Simon 702; Queiroz 3115 CEN; HUEFS KT386296 DQ790634 FJ982239
Pityrocarpa leucoxylon (Barneby & J.W. Grimes) Luckow & R.W. Jobson Fernandez 2909 NY (No data) DQ790622 (No data)
Pityrocarpa moniliformis (Benth.) Luckow & R.W. Jobson Melo 7518 HUEFS (No data) ON409911 ON409936
Pityrocarpa moniliformis (Benth.) Luckow & R.W. Jobson Queiroz 9084 HUEFS ON191501 ON409912 (No data)
Pityrocarpa moniliformis (Benth.) Luckow & R.W. Jobson Way 2449 K KT364067 KT364162 FJ982242
Pityrocarpa obliqua (Pers.) Brenan subsp. brasiliensis (G.P. Lewis) Luckow & R.W. Jobson Queiroz 12903 HUEFS ON191500 ON409920 (No data)
Pityrocarpa obliqua (Pers.) Brenan subsp. obliqua Macqueen 439 FHO KT364068 KT364206 FJ982243
Pseudopiptadenia bahiana G.P. Lewis & M.P. Lima Melo 138 HUEFS OM575115 ON409916 ON409930
Pseudopiptadenia bahiana G.P. Lewis & M.P. Lima Queiroz 15381 HUEFS MG001277 MG001290 ON409931
Pseudopiptadenia bahiana G.P. Lewis & M.P. Lima Queiroz 15504 HUEFS OM575114 ON409917 ON409932
Pseudopiptadenia brenanii G.P. Lewis & M.P. Lima Borges 680 SPF KT364069 (No data) KT364111
Pseudopiptadenia brenanii G.P. Lewis & M.P. Lima Cardoso 2807 HUEFS OM575108 ON409914 ON409937
Pseudopiptadenia brenanii G.P. Lewis & M.P. Lima Harley 56005 HUEFS OM575109 ON409915 (No data)
Pseudopiptadenia brenanii G.P. Lewis & M.P. Lima Queiroz 15585 HUEFS MG001278 ON409913 ON409938
Pseudopiptadenia contorta (DC.) G.P. Lewis & M.P. Lima Queiroz 15507 HUEFS (No data) KT364155 KT364113
Pseudopiptadenia contorta (DC.) G.P. Lewis & M.P. Lima Queiroz 15582 HUEFS MG001279 KX302348 MG001308
Pseudopiptadenia inaequalis (Benth.) Rauschert Lima 7790 RB OM575111 ON409921 ON409939
Pseudopiptadenia leptostachya (Benth.) Rauschert Lima 8231 RB OM575113 ON409922 ON409940
Pseudopiptadenia leptostachya (Benth.) Rauschert Lima 8326 RB OM575112 ON409923 ON409941
Pseudopiptadenia psilostachya (DC.) G.P. Lewis & M.P. Lima Simon 1245 CEN KT364070 KT364170 KT364114
Pseudopiptadenia schumanniana (Taub.) G.P. Lewis & M.P. Lima Lima 7938 RB OM575110 ON409924 ON409942
Pseudopiptadenia sp. Neves 1675 RB OM575116 ON409918 ON409933
Pseudopiptadenia sp. Ribeiro 351 HUEFS OM575117 ON409919 (No data)
Pseudopiptadenia suaveolens (Miq.) J.W. Grimes = P. psilostachya Moacir & Clovis sn IAN OM575119 ON409925 ON409934
Pseudopiptadenia warmingii (Benth.) G.P. Lewis & M.P. Lima Queiroz 12761 HUEFS OM575118 ON409926 ON409935
Senegalia macrostachya (Rchb. ex DC.) Kyal. & Boatwr. Miller 1322 CANB KY688790 KY688920 (No data)
Senegalia nigrescens (Oliv.) P.J.H. Hurter Maurin 255 JRAL JQ265858 GQ872237 (No data)
Stryphnodendron adstringens (Mart.) Coville Souza 29702 ESA KT364072 KT364198 KT364116
Stryphnodendron coriaceum Benth. Scalon 718 ESA (No data) KT364200 KT364120
Stryphnodendron duckeanum Occhioni Simon 1343 CEN KT364076 (No data) KT364122
Stryphnodendron fissuratum E.M.O. Martins Ivanauskas sn ESA KT364077 KT364175 KT364124
Stryphnodendron foreroi E.M.O. Martins Assis 1143 SPF KT364079 KT364201 KT364126
Stryphnodendron gracile Rizzini & Heringer Scalon 458 ESA KT364080 KT364177 KT364127
Stryphnodendron obovatum Benth. Scalon 712 ESA KT364081 KT364182 KT364130
Stryphnodendron occhionianum E.M.O. Martins Simon 1597 CEN KT364083 (No data) KT364132
Stryphnodendron paniculatum Poepp. Simon 1058 CEN KT364084 (No data) KT364133
Stryphnodendron polyphyllum Mart. Mello-Silva 2659 SPF KT364086 KT364184 KT364136
Stryphnodendron pulcherrimum (Willd.) Hochr. Simon 980 CEN KT364087 (No data) KT364137
Stryphnodendron roseiflorum (Ducke) Ducke Scalon 728 ESA KT364090 KT364193 KT364143
Stryphnodendron rotundifolium Mart. Scalon 715 ESA KT364094 KT364194 KT364147
Stryphnodendron velutinum Scalon Scalon 719 ESA KT364101 KT364187 KT364153

Total DNA was extracted from about 20 mg of silica gel-dried leaf material using a modified CTAB-based protocol (Inglis et al. 2018a). We checked DNA quality and integrity using agarose gel electrophoresis and DNA quantity and purity estimated by Nanodrop spectrophotometry (Thermo Scientific). Laboratory procedures, primer sequences and amplification protocols followed Inglis et al. (2018b) for nrITS and Simon et al. (2016) for matK and trnD-trnT. PCR products were prepared for direct Sanger sequencing using ExoSAP (ThermoFisher) and both DNA strands were sequenced using the Big Dye v.3.1 kit (Applied Biosystems), using the amplification primers. We obtained further sequences included in the analysis from GenBank (Table 1).

We assembled contigs using Geneious Prime 2021 (https://www.geneious.com) and aligned matrices with MAFFT v.7 (Katoh and Standley 2013). Maximum Likelihood (ML) phylogenetic analysis was performed using IQ-TREE (Nguyen et al. 2015), using 1000 ultrafast bootstrap replicates to estimate branch support (Hoang et al. 2018) and models estimated with ModelFinder (Kalyaanamoorthy et al. 2017). Trees were drawn with FigTree (http://tree.bio.ed.ac.uk/software/figtree/) and rooted using Lachesiodendron viridiflorum (Kunth) P.G. Ribeiro, L.P. Queiroz & Luckow, following Ringelberg et al. (2022). Analyses of individual loci produced similar topologies, although the plastid trees were substantially less well-resolved compared to the nrITS phylogeny. In the absence of major incongruence between individual gene trees, we inferred phylogenetic relationships with a concatenated dataset (nrITS, matK, trnD-trnT) containing 3280 bp and 13% of missing data and used it as the basis for proposing taxonomic rearrangements.

Data Resources

The data underpinning the analysis reported in this paper are deposited in GitHub at https://doi.org/10.5281/zenodo.6611789

Results and discussion

Our densely sampled phylogenetic analysis recovers Parapitadenia as monophyletic, reinforces the non-monophyly of Pseudopiptadenia and shows that Pityrocarpa is also non-monophyletic (Fig. 2). Although the backbone of the phylogeny remains weakly-supported, the three main clades relevant to the delimitation of genera and the taxonomic decisions proposed here have full (100%) bootstrap support.

Figure 2. 

Phylogeny of the Stryphnodendron clade, based on Maximum Likelihood analysis of the concatenated nrITS, matK and trnD-trnT data. Significant ultrafast bootstrap values (> 90%) are given above branches. The tree was rooted using Lachesiodendron viridiflorum. Scale bar: expected number of changes per site; dotted branches not to scale.

The first clade, hereafter referred to as Pseudopiptadenia pro parte, includes Ps. bahiana G.P. Lewis & M.P. Lima, Ps. contorta, Ps. psilostachya, Ps. warmingii (Benth.) G.P. Lewis & M.P. Lima and a putative new species yet to be described. The second clade, hereafter referred to as the Pityrocarpa clade, encompasses the remaining Pseudopiptadenia species, including the type species of the genus, Ps. leptostachya, intermixed with accessions of the three species of Pityrocarpa, including Pi. moniliformis (Benth.) Luckow & R.W. Jobson, the type species of Pityrocarpa.

The placement of Parapiptadenia, Pityrocarpa, and Pseudopiptadenia species in three distinct lineages and the robustly supported monophyly of Parapiptadenia agree with previous phylogenetic analyses (Fig. 1; Simon et al. 2016; Ribeiro et al. 2018; Lima et al. 2022; Ringelberg et al. 2022). However, the relationships amongst these three clades and other members of the Stryphnodendron clade remain unclear, because of the lack of support across the backbone of the clade (Figs 1 and 2) and disagreement with previous analyses. For example, although analyses of nuclear and plastid data (Simon et al. 2016; Ribeiro et al. 2018) also placed Pseudopiptadenia p.p. and Parapiptadenia in the same clade, this group could be sister to the remainder of the Stryphnodendron clade (Simon et al. 2016) or sister to the clade comprising Stryphnodendron and Microlobius (Ribeiro et al. 2018). Phylogenomic analyses based on 997 nuclear genes (Lima et al. 2022; Ringelberg et al. 2022) placed Pseudopiptadenia p.p. as sister to a group including Stryphnodendron duckeanum Occhioni f. plus a clade formed by Parapiptadenia and the Pityrocarpa clade. Furthermore, these nodes across the backbone of the Stryphnodendron clade show high gene tree conflict (Ringelberg et al. 2022) coinciding with very short branches and weak support in both conventional and phylogenomic analyses, highlighting the difficulties of inferring relationships across this part of the mimosoid phylogeny.

Despite uncertainties regarding generic relationships, our results provide an additional example of how over-reliance on particular traits, in this case fruits and seeds (Brenan 1955; 1963; Lewis and Elias 1981), may lead to unnatural taxonomies. Presence of follicles and of flat and winged seeds, which were used to diagnose Pseudopiptadenia, are respectively shared by most lineages within the Stryphnodendron clade or homoplastic between Pseudopiptadenia p.p. and members of the Pityrocarpa clade. All this is not to say that fruits have no taxonomic significance, as the vast majority of Parapiptadenia species have distinctive legumes with valves plicate above the seeds, not seen in any other member of the Stryphnodendron clade. Nonetheless, most species in the Pityrocarpa clade, even though variable in seed morphology (flat and winged vs. lentiform and wingless), share a number of similarities, including the position of the extrafloral nectaries between or just below the first pair of pinnae; few pinnae pairs; inflorescence spikes in general solitary and axillary to coeval leaves; and bifoliolate seedlings (Fig. 3). These features are not shared with most Pseudopiptadenia p.p. species, which have extrafloral nectaries on the lower half of the petiole; many pairs of pinnae; inflorescence spikes arranged in complex efoliate synflorescences; and pinnate or bipinnate seedlings (see Table 2). Although fairly homogeneous within the Pityrocarpa clade and Pseudopiptadenia p.p., the characters highlighted above sometimes vary amongst and within species, particularly in a context including Parapiptadenia. For example, solitary inflorescences occur in species of both Parapiptadenia and the Pityrocarpa clade, while Pseudopiptadenia p.p. species sometimes do not have spikes arranged in complex synflorescences (e.g. particular specimens of Ps. bahiana and Ps. contorta). Nonetheless, taken together, the traits highlighted here provide better recognition of these lineages as distinct genera than fruit morphology alone.

Table 2.
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Morphological comparison amongst Parapiptadenia, Pityrocarpa and Marlimorimia. Traits in bold highlight diagnostic features separating Pityrocarpa and Marlimorimia. EFN - Extrafloral nectary.

Parapiptadenia Pityrocarpa Marlimorimia
Pinnae number 1–8 1–4 (rarely to 5 in Pi. brenanii) 5–10 or more (2–5 in M. bahiana and M. colombiana)
Petiolar EFN position Variable across species Between or just below the first pair of pinnae Between the base and the middle of the petiole
Spike arrangement Solitary, axillary or supra-axillary to coeval leaves Solitary (very rarely up to 2 in Pi. moniliformis), axillary to coeval leaves 2–many fasciculate and further arranged in efoliate terminal pseudoracemes or on efoliate nodes below mature leaves
Petals Reddish (yellowish in Pa. excelsa and Pa. rigida); united at the base (free in Pa. rigida) White to yellowish or greenish; free and glabrous (united in Pi. leucoxylon) White to yellowish or greenish; united and pubescent
Fruit type (dehiscence) Legume Follicle Follicle
Fruit shape Flat compressed, valves plicate above the seeds (except in Pa. excelsa) Flat compressed, valves not plicate above the seeds Flat compressed, valves not plicate above the seeds
Fruit margins Straight to shallowly sinuous Deeply constricted (moniliform) (sinuous in P. brenanii). Straight, (shallowly and irregularly sinuous in M. bahiana, M. colombiana and M. warmingii), sometimes constricted where seeds abort
Valve consistency and indumentum Thin, chartaceous, glabrous Thick, coriaceous, mostly pubescent Thin, coriaceous (thicker and harder in M. warmingii), glabrous
Embryo plumule Developed and multifid Rudimentary (developed and multifid in P. brenanii) Developed and multifid
Seedling eophylls Pinnate Bifoliolate Pinnate or bipinnate
Figure 3. 

Inflorescences of Parapiptadenia, Pityrocarpa and Marlimorimia A Parapiptadenia pterosperma (Benth.) Brenan showing reddish inflorescences in the axils of coeval leaves B Pa. rigida (Benth.) Brenan showing yellowish inflorescences and fruits with valves plicate above the seeds C Pityrocarpa brenanii showing whitish, solitary spikes in the axils of coeval leaves D Pi. moniliformis showing yellowish, solitary spikes in the axil of a coeval leaf E Marlimorimia bahiana (G.P. Lewis & M.P. Lima) L.P. Queiroz & L.M. Borges, showing whitish spikes clustered in efoliate terminal pseudoracemes F Marlimorimiasp. showing yellowish spikes clustered in efoliate terminal pseudoracemes (Photos: A PG Ribeiro; B RT de Queiroz C–E LP Queiroz; F G Siqueira).

These results from phylogenetic and morphological analyses provide robust support for re-circumscription of Pseudopiptadenia as it was traditionally conceived and also Pityrocarpa. Given that the type species of these two genera are nested in the same clade and that no morphological traits support the recognition of a narrow circumscription of Pseudopiptadenia, we subsume the name Pseudopiptadenia under Pityrocarpa, the oldest validly published generic name (Britton and Rose 1928; Lewis and Lima 1991; Turland et al. 2018). We assign the remaining Pseudopiptadenia species to the new genus Marlimorimia.

Key to the genera Parapiptadenia, Pityrocarpa, and Marlimorimia. See Lima et al. (2022) for a key to all genera of the Stryphnodendron clade

1 Petals reddish (yellowish in Pa. excelsa and Pa. rigida); fruit a legume (dehiscing along both sutures), the valves plicate above the seeds (except in Pa. excelsa) Parapiptadenia
– Petals white to yellowish or greenish; fruit a follicle (splitting along one suture only), the valves not plicate above the seeds 2
2 Petiolar nectary just below or between the first pair of pinnae; spikes solitary in axils of coevally developing leaves; petals free (united in Pi. leucoxylon) and glabrous; fruit margins deeply constricted (sinuous in Pi. brenanii) Pityrocarpa
– Petiolar nectary between the base and the middle of the petiole; spikes 2-many-fasciculate, the fascicles usually arranged in efoliate terminal pseudoracemes or on efoliate nodes below the leaves; petals united and pubescent; fruit margins straight or shallowly and irregularly sinuous, sometimes constricted where seeds abort Marlimorimia

Taxonomy

Pityrocarpa (Benth.) Britton & Rose, N. Amer. Fl. 23(3): 190. 1928.

Monoschisma Brenan, Kew Bull. 10(2): 179. 1955, nom. inval., non Monoschisma Duby, Mém. Soc. Phys. Genève 19: 294. 1868. Type. Monoschisma leptostachyum (Benth.) Brenan, syn. nov.

Pseudopiptadenia Rauschert, Taxon 31(3): 559. 1982. Type. Pseudopiptadenia leptostachya (Benth.) Rauschert, syn. nov.

Basionym

Piptadenia sect. Pityrocarpa Benth., J. Bot. (Hooker) 4: 339. 1842.

Type

Pityrocarpa moniliformis (Benth.) Luckow & R.W. Jobson [≡ Piptadenia moniliformis Benth., designated by Britton and Rose 1928].

Description

Unarmed trees or shrubs. Leaves bipinnate; petiole with an extrafloral nectary between or shortly below the first pair of pinnae; pinnae 1–4 (5) pairs, exceptionally to 10 pairs in Pi. leptostachya; leaflets 1–10 pairs per pinna, rarely to 20 pairs (Pi. brenanii and Pi. leptostachya), mostly rhomboid sometimes also asymmetrically elliptical or lanceolate. Inflorescences spikes, solitary in the axils of coeval leaves, commonly pendulous. Flowers pentamerous; petals free (except possibly Pi. leucoxylon), glabrous; stamens 10, anther gland present; ovary shortly stipitate and included within or exserted from the corolla. Fruit a follicle, dehiscing along the lower suture, flat compressed, mostly moniliform, the margins deeply and regularly constricted, rarely sinuous margins and shallowly constricted (Pi. brenanii and occasionally in Pi. leucoxylon); valves stiffly coriaceous. Seeds mostly flat compressed with a coriaceous testa and a narrow marginal wing, lacking a pleurogram or, less frequently, ovoid or discoid with a hard, whitish testa, wingless and with a ‘U’-shaped pleurogram (Pi. leucoxylon, Pi. moniliformis and Pi. obliqua); embryo with a rudimentary plumule (except Pi. brenanii). Seedlings with bifoliolate eophylls.

Distribution

Pityrocarpa is distributed in tropical America, from Mexico to southern Brazil and Paraguay. Most species occur in the Brazilian Atlantic rainforests (Pi. inaequalis, Pi. leptostachya, Pi. schumanniana), in the northern Amazonian rainforests (Pi. leucoxylon), in seasonally dry tropical forests and woodlands in the north-eastern Brazilian Caatinga (Pi. brenanii, Pi. moniliformis, Pi. obliqua subsp. brasiliensis), western Mexico (Pi. obliqua subsp. obliqua) or in Venezuelan savannas and Paraguayan Chaco (Pi. moniliformis).

Notes

As circumscribed here, Pityrocarpa includes seven species, all with a moniliform fruit, with the margins deeply constricted between the seeds (Fig. 4). This trait is shared by species formerly included in Pityrocarpa (sensu Jobson and Luckow 2007) and some species previously placed in the genus Pseudopiptadenia (sensu Lewis and Lima 1991). These two genera had been separated based on seed morphology, Pityrocarpa characterised by ovoid or discoid seeds with a hard, whitish seed coat and a ‘U’-shaped pleurogram, while Pseudopiptadenia included species with flat compressed and narrowly winged seeds with a coriaceous testa lacking a pleurogram. Pityrocarpa brenanii and Pi. leucoxylon have fruits with only shallowly sinuous margins, more similar to species of the genus Marlimorimia.

Figure 4. 

Fruits of Pityrocarpa species A Pi. brenanii (from Lewis et al. 1899, NY) B Pi. inaequalis (from Moreira et al. 3, F) C Pi. leptostachya (from Baez et al. 1174, NY) D Pi. leucoxylon (from de Bruijn 1750, NY) E Pi. moniliformis (from Nunes 597, HUEFS) F Pi. obliqua subsp. brasiliensis (from Mori 11837, NY) G Pi. schumanniana (from Lima 2994, RB).

Besides sharing these fruit traits, Pityrocarpa species also have leaves with few pinnae (1 to 4 [5] pairs, rarely up to 10 pairs in Pi. leptostachya) and relatively large rhomboid leaflets compared to species of Marlimorimia. One exception are the leaves of Pi. brenanii, which are similar to those of M. bahiana. All species of Pityrocarpa present an extrafloral nectary between or shortly below the first pair of pinnae, in contrast to species of Marlimorimia that have the nectary below mid-petiole, frequently close to the pulvinus.

Floral traits, although previously disregarded as being generically diagnostic in the group, provide further evidence for the distinction between Pityrocarpa and Marlimorimia. The solitary inflorescence spikes in the axils of coevally developing leaves in Pityrocarpa contrast with the more complex synflorescences of Marlimorimia (Fig. 3; see notes under Marlimorimia). All species of Pityrocarpa have free and glabrous petals, except for Pi. leucoxylon, in which the petals are connate for a little over 1 mm (Barneby and Grimes 1984).

Lima (1985) and Lewis and Lima (1991) provided additional information on embryos and seedlings that are potentially useful for distinguishing Pityrocarpa from Marlimorimia. Embryos of Pityrocarpa species have a rudimentary plumule, while in Marlimorimia, the plumule is developed and multifid. This seems to be correlated with seedling morphology as the studied species of Pityrocarpa have bifoliolate eophylls and those of Marlimorimia species have pinnate or bipinnate eophylls (Lewis and Lima 1991). Pityrocarpa brenanii, however, has embryo morphology more similar to that reported for species of Marlimorimia (Lewis and Lima 1991).

Pityrocarpa brenanii (G.P. Lewis & M.P. Lima) L.P. Queiroz & L.M. Borges, comb. nov.

urn:lsid:ipni.org:names:77303780-1

Basionym

Pseudopiptadenia brenanii G.P. Lewis & M.P. Lima, Arch. Jard. Bot. Rio de Janeiro 30: 50–51. 1991.

Type

Brasil, Bahia, Harley et al. 21346 (holotype CEPEC; isotypes BR, K, M, MBM, MEXU, NY, RB, US).

Pityrocarpa inaequalis (Benth.) L.P. Queiroz & Marc.F. Simon, comb. nov.

urn:lsid:ipni.org:names:77303781-1

Monoschisma inaequale (Benth.) Brenan, Kew Bull. 10(2): 179. 1955.

Pseudopiptadenia inaequalis (Benth.) Rauschert, Taxon 31(3): 559. 1982.

Basionym

Piptadenia inaequalis Benth., J. Bot. (Hooker) 4: 339. 1842.

Type

Brazil, Rio de Janeiro, Pohl 1386 (lectotype K 000504704, designated here; isolectotype K 000504706).

Pityrocarpa leptostachya (Benth.) L.P. Queiroz & P.G. Ribeiro, comb. nov.

urn:lsid:ipni.org:names:77303782-1

Monoschisma leptostachyum (Benth.) Brenan, Kew Bull. 10(2): 179. 1955.

Pseudopiptadenia leptostachya (Benth.) Rauschert, Taxon 31(3): 559. 1982.

Basionym

Piptadenia leptostachya Benth., J. Bot. (Hooker) 4: 339. 1842.

Type

Brasil, Sellow s.n. (Lectotype K 000504709, designated here; isolectoypes F 0360957F [fragment], K 000504710, TUB 009699).

Note

Lewis and Lima (1991) unintentionally lectotypified this name by indicating the holotype to be at B and the isotype to be at K. However, the B specimen was destroyed and, hence, cannot serve as a lectotype. Moreover, K holds two duplicates of an un-numbered Sellow collection. Here, we chose the one previously belonging to Bentham’s herbarium as the lectotype.

Pityrocarpa leucoxylon (Barneby & J.W. Grimes) Luckow & R.W. Jobson, Syst. Bot. 32(3): 573. 2007.

Basionym

Piptadenia leucoxylon Barneby & J.W. Grimes, Brittonia 36(3): 236–238. 1984.

Type

Venezuela: Bolivar, de Bruijn 1750 (holotype NY; isotypes MO, VEN, US).

Pityrocarpa moniliformis (Benth.) Luckow & R.W. Jobson, Syst. Bot. 32(3): 573. 2007.

Stryphnodendron piptadenioides E.M.O. Martins, Leandra 5(6): 90. 1975. Type. Brazil. Pernambuco, “lectum in silva pluviali ad S. José Belmonte”, Mata da Mina, 29 Oct 1971, Ramalho 52 (holotype RFA 17173) .

Stryphnodendron consimile E.M.O. Martins, Leandra 5(6): 92. 1975. Type. Brazil. Piauí, “habitat in caatinga ad Paulistana”, Fazenda Altamira, 04 Nov 1974, Lima 1330 (holotype RFA 17172).

Basionym

Piptadenia moniliformis Benth., J. Bot. (Hooker) 4: 339. 1842.

Type

Brazil, Bahia, Serra de Jacobina, Blanchet 2701 (lectotype K 000090193, designated here; isolectotypes F, K 000205897, MO, NY 00003233).

Pityrocarpa obliqua (Pers.) Brenan, Kew Bull. 10(2): 176. 1955.

Acacia thibaudiana DC., Prodr. 2: 456. 1825.

Piptadenia obliqua (Pers.) J.F. Macbr., Contr. Gray Herb. 59: 17. 1919.

Basionym

Sophora obliqua Pers., Syn. Pl. 1: 452. 1805.

Type

“Amer. australi?”, Herb. D. Thibaud. (not located).

1.6.1. Pityrocarpa obliqua subsp. obliqua

Pityrocarpa obliqua subsp. brasiliensis (G.P. Lewis) Luckow & R.W. Jobson, Syst. Bot. 32(3): 573. 2007.

Basionym

Piptadenia obliqua subsp. brasiliensis G.P. Lewis, Kew Bull. 46(1): 160–162. 1991.

Type

Brazil, Bahia, Mori et al. 9519 (holotype CEPEC; isotypes HUEFS, K, NY).

Pityrocarpa schumanniana (Taub.) L.P. Queiroz & L.M. Borges, comb. nov.

urn:lsid:ipni.org:names:77303784-1

Pseudopiptadenia schumanniana (Taub.) G.P. Lewis & M.P. Lima, Arch. Jard. Bot. Rio de Janeiro 30: 53. 1991.

Basionym

Piptadenia schumanniana Taub., Flora 75: 75. 1892.

Type

Brazil, “Brasilia austro-orientale”, Rio de Janeiro, Glaziou 13774 (lectotype R 00008369, designated here; isolectotypes A 00064056, F 0058675F, K 000504703, MPU 016109, NY 00003244, NY 00003245, US 00001018, US 00997081).

Marlimorimia L.P. Queiroz, L.M. Borges, Marc.F. Simon & P.G. Ribeiro, gen. nov.

urn:lsid:ipni.org:names:77303785-1

Newtonia sect. Neonewtonia Burkart, Fl. Il. Catarin. fasc. LEGU: 285. 1979, syn. nov. Type. Newtonia nitida (Benth.) Brenan (= Marlimorimia contorta (DC.) L.P. Queiroz & P.G. Ribeiro).

Diagnosis

Marlimorimia shares with Pityrocarpa the follicle, a fruit dehiscing along the lower suture only, and flat, compressed winged seeds, which lack a pleurogram. It can be differentiated from Pityrocarpa by the position of the extrafloral nectary on the petiole (from the base to the mid-petiole in Marlimorimia vs. between or just below the first pair of pinnae in Pityrocarpa); inflorescence spikes clustered in terminal pseudoracemes or in fascicles at efoliate nodes, surpassed by mature leaves (vs. solitary spikes in the axils of coeval leaves); petals united and joined into a gamopetalous corolla (vs. petals free and glabrous); and fruits with margins straight to shallowly sinuous (vs. margins deeply constricted).

Type

Marlimorimia contorta (DC.) L.P. Queiroz & P.G. Ribeiro

Description

Unarmed trees. Leaves bipinnate; petiole with an extrafloral nectary well below the first pair of pinnae, close to the pulvinus, always below mid-petiole; pinnae 5–10 to many pairs per leaf (2–3 pairs in M. colombiana and 3–5 in M. bahiana); leaflets mostly > 10 pairs per pinna, (6–8 in M. colombiana), mostly oblong to linear from an asymmetrical base, rarely rhomboid (M. bahiana). Inflorescences spikes, grouped in fascicles, these being arranged in terminal pseudoracemes or forming clusters below the coeval leaves. Flowers pentamerous; petals united into a gamopetalous corolla, pubescent; stamens 10, anther gland present; ovary shortly stipitate and included or exserted from the corolla. Fruit a follicle, dehiscing along the lower suture, flat compressed, straight, curved or longitudinally twisted, the margins usually straight, rarely irregularly sinuous and only becoming constricted where the seeds fail to develop (M. bahiana and M. warmingii), valves coriaceous, thin or thick. Seeds flat compressed with a coriaceous testa, presenting a narrow or somewhat wider marginal wing, pleurogram lacking; embryo with a developed, multifid plumule (unknown in M. colombiana and M. pittieri). Seedlings with pinnate or bipinnate eophylls (unknown in M. bahiana, M. colombiana and M. pittieri).

Distribution

Marlimorimia comprises six species with a bicentric distribution in the two main areas of tropical humid forests in South America. Three species occur in eastern Brazil, two of which are restricted to the Atlantic wet forests (Marlimorima bahiana and M. warmingii) and M. contorta, which extends to inland semi-deciduous forests. The three other species are distributed in northern South America. Marlimorimia psilostachya is widely distributed across Amazonia, sparsely extending to Central America (Costa Rica) and M. colombiana and M. pittieri have restricted ranges in Colombia and Venezuela, respectively.

Etymology

The genus Marlimorimia is named in honour of Dr. Marli Pires Morim, taxonomist at the Rio de Janeiro Botanical Garden, for her outstanding contribution to our knowledge of the diversity and taxonomy of Brazilian mimosoid legumes.

Notes

The new genus Marlimorimia is proposed to accommodate a monophyletic group of species, previously classified in Pseudopiptadenia (sensu Lewis and Lima 1991; Luckow 2005), but which could not retain the genus name, because its type species is now included in Pityrocarpa.

Besides the molecular phylogenetic evidence, morphology also supports recognition of Marlimorimia as distinct from Pityrocarpa. Marlimorimia brings together most of the species formerly placed in Pseudopiptadenia which have multipinnate leaves, small oblong to linear leaflets and fruits with straight (or shallowly sinuous) margins. Marlimorimia bahiana and M. colombiana, however, have leaves with few pinnae and rhomboid leaflets.

Species of Marlimorimia have more complex inflorescences than those of Pityrocarpa. While the spikes of Pityrocarpa are solitary in the axils of coevally developing leaves, Marlimorimia species have spikes in fascicles of 2–3, which are arranged in terminal efoliate pseudoracemes or clustered on nodes below mature leaves (Fig. 3). Sometimes, as leaves expand, Marlimorimia synflorescences may resemble those of Pityrocarpa and Parapiptadenia (e.g. particular specimens of M. contorta such as Hatschbach 50149 [NY]). Nonetheless, flowers of Marlimorimia have pubescent petals united into a gamopetalous corolla (vs. free glabrous petals in the majority of Pityrocarpa species).

Two types of fruits are found in Marlimorimia (Fig. 5). Some species have long linear fruits, frequently curved or longitudinally twisted with straight margins (M. colombiana, M. contorta, M. pittieri and M. psilostachya), while M. bahiana and M. warmingii have oblong fruits with shallowly sinuous margins. The valves of the fruits are woody, although usually thin, becoming thicker and harder in M. warmingii.

Figure 5. 

Fruits of Marlimorimia species A Marlimorimia bahiana (from Amorim 1009, NY) B Marlimorimia colombiana (from Killip 16268, NY) C Marlimorimia contorta (from Gomes 257, NY) D Marlimorimia pittieri (from Guevara 1264, F) E Marlimorimia psilostachya (from Rabelo 2753, NY) F Marlimorimia warmingii (from Nunes et al. 2, NY).

The seeds of Marlimorimia, although superficially similar to those of most species of Pityrocarpa, have embryos with multifid plumules that result in seedlings with pinnate or bipinnate eophylls (Lima 1985; Lewis and Lima 1991).

Marlimorimia bahiana (G.P. Lewis & M.P. Lima) L.P. Queiroz & L.M. Borges, comb. nov.

urn:lsid:ipni.org:names:77303786-1

Basionym

Pseudopiptadenia bahiana G.P. Lewis & M.P. Lima, Arch. Jard. Bot. Rio de Janeiro 30: 54–55. 1991.

Type

Brasil, Bahia, Mori & King 12223 (holotype CEPEC; isotypes HUEFS, K, NY, RB).

Marlimorimia colombiana (Britton & Killip) L.P. Queiroz & Marc.F. Simon, comb. nov.

urn:lsid:ipni.org:names:77303787-1

Pseudopiptadenia colombiana (Britton & Killip) G.P. Lewis

Basionym

Stryphnodendron colombianum Britton & Killip, Ann. New York Acad. Sci. 35(3): 155. 1936.

Type

Colombia, Santander, Killip & Smith 16268 (holotype NY 00003356; isotypes A, GH, US).

Notes

In the absence of phylogenetic evidence, the petiolar extrafloral nectaries located at mid-petiole and fruits with straight to shallowly sinuous margins support the transfer of Pseudopiptadenia colombiana to Marlimorimia.

Marlimorimia contorta (DC.) L.P. Queiroz & P.G. Ribeiro, comb. nov.

urn:lsid:ipni.org:names:77303788-1

Piptadenia nitida Benth., J. Bot. (Hooker) 4: 336. 1842.

Piptadenia contorta (DC.) Benth., Trans. Linn. Soc. Lond. 30: 368. 1875.

Newtonia nitida (Benth.) Brenan, Kew. Bull. 10 (2): 182. 1955.

Newtonia contorta (DC.) Burkart, Fl. Il. Catarin. fasc. LEGU: 289. 1979.

Pseudopiptadenia contorta (DC.) G.P. Lewis & M.P. Lima, Arch. Jard. Bot. Rio de Janeiro 30: 57. 1991.

Basionym

Acacia contorta DC., Prodr. 2: 470. 1825.

Type

Brasil, Rio de Janeiro, Raddi s.n. (lectotype FI, designated here).

Marlimorimia pittieri (Harms) L.P. Queiroz & L.M. Borges, comb. nov.

urn:lsid:ipni.org:names:77303789-1

Piptadenia similis Britton & Killip, Ann. New York Acad. Sci. 35(3): 156. 1936. Holotype Colombia, Barranquilla, Elias 263 (US).

Pseudopiptadenia pittieri (Harms) G.P. Lewis, Kew Bull. 46(1): 118. 1991.

Basionym

Piptadenia pittieri Harms, Notizbl. Bot. Gart. Berlin-Dahlem 8(71): 51–52. 1921.

Type

Venezuela, Carabobo, Pittier 8859 (lectotype US 00001013, designated here; isolectotypes GH 00064052, NY 00003236).

Notes

Although Pseudopiptadenia pittieri was not included in the phylogenetic analyses, the presence of extrafloral nectaries at the base of the petiole, spikes arranged in pseudoracemes and fruits with straight margins support its transfer to Marlimorimia.

Marlimorimia psilostachya (DC.) L.P. Queiroz & Marc.F. Simon, comb. nov.

urn:lsid:ipni.org:names:77303790-1

Piptadenia psilostachya (DC.) Benth., J. Bot. (Hooker) 4: 336. 1842.

Piptadenia suaveolens Miq., Linnaea 18: 589–590. 1845. Type Surinam, Bergendaal, Focke 936 (holotype U).

Newtonia psilostachya (DC.) Brenan, Kew. Bull. 10 (2): 182. 1955.

Newtonia suaveolens (Miq.) Brenan, Kew. Bull. 10 (2): 182. 1955.

Pseudopiptadenia psilostachya (DC.) G.P. Lewis & M.P. Lima, Arch. Jard. Bot. Rio de Janeiro 30: 55. 1991.

Pseudopiptadenia suaveolens (Miq.) J.W. Grimes, Brittonia 45(1): 27. 1993.

Basionym

Acacia psilostachya DC., Prodr. 2: 457. 1825.

Type

French Guiana, Cayenne, Martin 2 (lectotype K 000504699, designated by Lewis & Lima 1991; isolectotype P 02930999).

Notes

Contrary to Grimes (1993), who recognised Pseudopiptadenia psilostachya and Ps. suaveolens as distinct species, we agree with Lewis and Lima (1991) on the synonymisation of Ps. suaveolens under M. psilostachya. These plants grow sympatrically and the traits used by Grimes (1993) to support recognition of two species are too variable to be diagnostic.

Marlimorimia warmingii (Benth.) L.P. Queiroz & P.G. Ribeiro, comb. nov.

urn:lsid:ipni.org:names:77303791-1

Piptadenia glaziovii Harms, Repert. Spec. Nov. Regni Veg. 17: 203. 1921. Type. Brasil, Rio de Janeiro, Serra da Estrela, Glaziou 8440 (lectotype K, designated by Lewis and Lima 1991).

Newtonia glaziovii (Harms) Burkart ex Barth & Yoneshigue, Mem. Inst. Oswaldo Cruz 64: 102. 1966.

Newtonia warmingii (Benth.) G.P. Lewis, Legumes of Bahia p. 111. 1987.

Pseudopiptadenia warmingii (Benth.) G.P. Lewis & M.P. Lima, Arch. Jard. Bot. Rio de Janeiro 30: 54. 1991.

Basionym

Mimosa warmingii Benth., Trans. Linn. Soc. London 30(3): 413. 1875.

Type

Brasil, Minas Gerais, Lagoa Santa, Warming s.n. (lectotype K 000504702, designated by Lewis and Lima 1991).

Acknowledgements

We thank fellow plant collectors, particularly Haroldo de Lima, Danilo Neves and Domingos Cardoso, for providing silica-dried leaf samples used in our phylogenetic analysis. We also thank Marli Morim for her valuable inputs to the manuscript, Thais Cury de Barros for a discussion on anther gland morphology and Gwilym Lewis, an anonymous reviewer and particularly Colin Hughes for their suggestions. L.P. Queiroz acknowledges support from CNPq (processes 303585/2016-1 and 440487/2015-3) and FAPESB (processes APP 096/2016 and PTX0004/2016). P.G. Ribeiro acknowledges a Ph.D. grant from Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES; process 5414130) and the GM and REFLORA project grants from CNPq (processes 130515/2010-8 and 563533/2010-2). M.F. Simon acknowledges support from CNPq (305570/2021-8). The use of DNA from the Brazilian species is authorised by SISGEN.

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  • Simon MF, Pastore JFB, Souza AF, Borges LM, Scalon VR, Ribeiro PG, Santos-Silva J, Souza VC, Queiroz LP (2016) Molecular phylogeny of Stryphnodendron (Mimosoideae, Leguminosae) and generic delimitations in the Piptadenia group. International Journal of Plant Sciences 177(1): 44–59. https://doi.org/10.1086/684077
  • Turland NJ, Wiersema JH, Barrie FR, Greuter W, Hawksworth DL, Herendeen PS, Knapp S, Kusber W-H, Li D-Z, Marhold K et al. (2018) International Code of Nomenclature for Algae, Fungi, and Plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Koeltz Botanical Books. https://doi.org/10.12705/Code.2018

Supplementary material

Supplementary material 1 

Voucher information and GenBank accession numbers (internal transcribed spacer-ITS sequences) for taxa used in this study

Leonardo M. Borges, Peter W. Inglis, Marcelo F. Simon, Pétala Gomes Ribeiro, Luciano P. de Queiroz

Data type: excel file

Explanation note: Voucher information and GenBank accession numbers for taxa used in this study.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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