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# The Taxonomic Status of Genera within the Fabeae
## (Vicieae), with a Special Focus on Pisum

T. Ellis, Petr Smýkal, Nigel Maxted, Clarice Coyne, Claire Domoney, Judith

Burstin, Yanis Bouchenak-Khelladi, Noam Chayut

► To cite this version:

T. Ellis, Petr Smýkal, Nigel Maxted, Clarice Coyne, Claire Domoney, et al. The Taxonomic Status of Genera within the Fabeae (Vicieae), with a Special Focus on Pisum. Diversity, 2024, 16 (7), pp.365. 10.3390/d16070365. hal-05269690

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# The Taxonomic Status of Genera within the Fabeae (Vicieae), with a Special Focus on *Pisum*

T. H. Noel Ellis <fnref n="1" />,* , Petr Smýkal <fnref n="2" />, , Nigel Maxted <fnref n="3" />, Clarice J. Coyne <fnref n="4" />, Claire Domoney <fnref n="1" />, , Judith Burstin <fnref n="5" />, , Yanis Bouchenak-Khelladi <fnref n="5" /> and Noam Chayut <fnref n="1" />

**Abstract:** The taxonomy of the tribe Fabeae (Vicieae) has long been problematic, but an analysis by Schaefer et al. in 2012 gave an exceptionally clear view of the tribe and noted the possibility that some nomenclatural adjustments may be required at some future date. These authors suggested several options, expressing some preferences. However, there has been a recent change to formally accepted names, implementing one of these possibilities, but without any additional relevant information. This change seems unjustified and unhelpful. We therefore present an argument for the retention, or re-instatement, of the genera *Pisum*, *Vavilovia*, and *Lens* until such time as new data support this requirement and there is no nomenclatural solution that is both accurate and convenient.

**Keywords:** Leguminosae; systematics; phylogenetics; *Pisum sativum*; *Lathyrus oleraceus*; nomenclature

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**Citation:** Ellis, T.H.N.; Smýkal, P.; Maxted, N.; Coyne, C.J.; Domoney, C.; Burstin, J.; Bouchenak-Khelladi, Y.; Chayut, N. The Taxonomic Status of Genera within the Fabeae (Vicieae), with a Special Focus on *Pisum*. *Diversity* **2024**, *16*, 365. https://doi.org/10.3390/d16070365

Academic Editor: Michael Wink

Received: 3 June 2024

Revised: 21 June 2024

Accepted: 22 June 2024

Published: 26 June 2024

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**Copyright:** © 2024 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
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Attribution (CC BY) license (https://
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4.0/).

# 1. Introduction

Legumes (Leguminosae or Fabaceae) are a large and diverse family of angiosperms [1,2] recently expanded and reorganized to include six subfamilies [2]. Most agriculturally important legumes are found within the Papilionoideae subfamily, and within this subfamily, an informally named monophyletic group of genera, the Inverted Repeat Lacking Clade (IRLC) [3,4], includes the majority of temperate legume crop and pasture species.

The IRLC, which includes, among others, the tribes Cicereae, Wisterieae, Trifolieae, and Fabeae (Vicieae), is defined as papilionoid legume species lacking one of the two inverted repeats of the plastid genome commonly found in land plants and freshwater green algae (the Viridiplantae). As is the case for the other group of land plants where this type of plastid genome is found (twice in the Pinophyta), this structural rearrangement affects the rate of evolution of plastid genes usually located within the inverted repeat [5]. Analysis of the phylogenetics of *Medicago* (Trifolieae) presented evidence for a gain of an IR structure in *M. minima* [6]. Taken together, these observations suggest that phylogenies within the IRLC that depend on plastid genome divergence should be treated with caution.

Fabeae (Vicieae) sensu lato includes the genera *Vicia*, *Lens*, *Lathyrus*, *Pisum*, and *Vavilovia* [7], and a comprehensive phylogenetic analysis of this group was presented by Schaefer et al. [8], who concluded that these genera were not monophyletic, noting that *Pisum* and *Vavilovia* were nested within *Lathyrus*, and that *Lens* was nested within *Vicia* and furthermore, that a group of *Vicia* species, belonging to sections *Ervum* and *Ervilia*, were early-diverging lineages of the entire tribe. Schaefer et al. [8] discussed the taxonomic

<footnote n="1">¹ John Innes Centre, Norwich Research Park, Colney Lane, Norwich NR4 7UH, UK;
claire.domoney@jic.ac.uk (C.D.); noam.chayut@jic.ac.uk (N.C.)</footnote>

<footnote n="2">² Department of Botany, Faculty of Science, Palacky University, Slechtitelu 27, 77900 Olomouc, Czech Republic;
petr.smykal@upol.cz</footnote>

<footnote n="3">³ School of Biosciences, University of Birmingham, Edgbaston, Birmingham B15 2TT, UK;
n.maxted@bham.ac.uk</footnote>

<footnote n="4">⁴ Department of Horticulture, Washington State University, Pullman, WA 99164, USA; coynec@wsu.edu</footnote>

<footnote n="5">⁵ Agroécologie, Institut Agro Dijon, INRAE, Univ. Bourgogne, Univ. Bourgogne Franche-Comté,
F-21000 Dijon, France; judith.burstin@inrae.fr (J.B.); yanis.bouchenak-khelladi@inrae.fr (Y.B.-K.)</footnote>

<footnote>* Correspondence: noel.ellis2@jic.ac.uk; Tel.: +44-1603-450000</footnote>
implications of this phylogeny and suggested two broad possibilities; the first was to generate a multiplicity of new genera, and the second was to amalgamate all into a single genus, *Vicia*. One alternative was to merge *Pisum* and *Vavilovia* with *Lathyrus*, and *Lens* with *Vicia*, creating three genera in the tribe, the basal genus comprised of *Vicia* sections *Ervum* and *Ervilia*, *Vicia* (excluding sections *Ervum* and *Ervilia* but including *Lens*), and *Lathyrus*, broadened to include *Pisum* and *Vavilovia*.

This three-genus scheme has been implemented, in part [9]; here, we discuss the basis and implications for this taxonomic restructuring and argue for delaying any change in the taxonomy of this tribe until more robust information is available.

## 2. The Fabeae

The general features of the phylogenetic tree presented by Schaefer et al. [8] are summarized in Figure 1; the data and interpretations of Schaefer et al. [8] remain the best available description of relationships within the Fabeae.

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**Figure 1.** (A) Summary of the consensus Maximum Likelihood phylogeny of the tribe Fabeae from Additional File 12, Figure S10, of Schaefer et al. [8]. Bootstrap values are to the right, and for reference, nodes are indicated by a letter. *Lathyrus-1* = most *Lathyrus*; *Lathyrus-2* = *L. neurolobus*; *Lathyrus-3* = *L. articulatus*, *L. clymenum*, and *L. ochrus*; *Vicia-1* = *Vicia pubescens*, *V. tenuissima*, and *V. tetrasperma*; *Vicia-2* = most *Vicia*; *Vicia-3* = *V. malosana*, *V. claessensii*, *V. paucifolia*, *V. iranica*, and *V. subvilloosa*; *Vicia-4* = *V. crocea*; *Vicia-5* = *V. gigantea*, *V. nigricans*, and *V. menziesii*; *Vicia-6* = *V. cypria*, *V. lunata*, *V. sylvatica*, *V. ervilia*, *V. articulata*, *V. monantha*, and *V. hirsuta*. (B) Summary Chronogram of the Fabeae for relevant
nodes, from Additional File 11, Figure S9, of Schaefer et al. [8]. Numbers are age estimates in Myr. The grey boxes indicate the 95% confidence intervals of the age estimates. In part B, nodes indicated by letters with a prime refer to the equivalent node with the same letter, as shown in part A, and as a letter, when there is no corresponding bifurcation in the two source data files. Names in grey are the suggested new genus names [8].

## 2.1. *The Consensus Maximum Likelihood (ML) Phylogeny*

The basal bifurcation in the consensus ML phylogeny separates *Vicia*-6 (*V. cypria*, *V. lunata*, *V. sylvatica*, *V. ervilia*, *V. articulata*, *V. monantha*, and *V. hirsuta*) from all other species, with a high degree of certainty (93% bootstrap support). It is this bifurcation and the paraphyly of *Vicia* that argues for the entire tribe being considered to be the monophyletic genus, *Vicia*. While this interpretation is well supported, it adds no information regarding the internal relationships of the Fabeae as a whole, which is provided by the recognition of additional genera within the Fabeae.

The reinstatement of the genus *Ervilia*, corresponding to this group of species, branching at node "i" (Figure 1), has the potential to enable the fragmentation of the tribe into more than one genus. These species have not been renamed; for example, the name *Vicia ervilia* (L.) Willd. is accepted in The World Checklist of Vascular Plants (WCVP) [9]. In addition, the remaining *Vicia* species are paraphyletic, and the other genera are nested within *Vicia*.

The creation of another genus *Ervium*, branching at node "e" (Figure 1), has also been proposed [8]. This genus would comprise *Vicia pubescens*, *V. tenuissima*, and *V. tetrasperma* (*Vicia*-1 in Figure 1). However, this retains a paraphyletic *Vicia*, including *Lens*. The genus *Ervium* is not recognized by WCVP [9], but *Lens culinaris* is no longer accepted and has been replaced by *Vicia lens* (L.) Coss. & Germ. Strangely, *Lens culinaris* subsp. *odemensis* (Ladiz.) and *Lens culinaris* subsp. *tomentosus* (Ladiz.) remain [10].

The *Ervium* proposal creates a monophyletic group at node "d" (Figure 1) in the consensus ML phylogeny, and this group includes *Pisum* and *Vavilovia* nested within three *Lathyrus* lineages. However, this group has only 53% bootstrap support. *Pisum* and *Vavilovia* are distinct, each with 100% bootstrap support, even though the monospecific genus *Vavilovia formosa* was formerly *Pisum formosum* [9]. The combined *Pisum* plus *Vavilovia* is monophyletic, with 99% bootstrap support, and is thus distinct from (or within) *Lathyrus*.

An important issue to resolve is the status of the nodes labeled "b", "c" and "d" (Figure 1) in the ML phylogeny. These do not have strong bootstrap support, raising the possibility that the three groups of *Lathyrus* species may be a monophyletic clade, rather than a paraphyletic grade, as in this figure. Supposing that *Lathyrus* and *Pisum* plus *Vavilovia* are two monophyletic clades derived at node "d" (Figure 1, condensing the nodes "b" and "c", with less than 50% support, and "d", which musters 53% support), then, in that case, there is no requirement to re-assign any generic status. On the other hand, if we accept that there are two monophyletic lineages, designated *Lathyrus*-1 and *Lathyrus*-1&2 in the chronogram of Figure 1B, then it would be possible either to elevate the smaller subgroup of *Lathyrus* to a new genus, or to combine *Lathyrus* with *Pisum* plus *Vavilovia*. The former option has four genera, and the latter has only one. Furthermore, it should be noted that the diversity within *Pisum* and *Vavilovia* is not fully represented in this tree, both with respect to the number of *Pisum* subspecies represented [11–13] and with respect to the molecular data available. Indeed, whereas *Lathyrus* accessions were described both by ITS and plastid DNA sequences, most *Pisum* accessions where only characterized by ITS sequences. A total of 15 out of 18 *Pisum* accessions had either one or no chloroplast sequence, which may have significant effects on the robustness of the phylogenetic relationships inferred, especially if the sampling of chloroplast data for *Lathyrus* is higher.

The majority of *Lathyrus* species are represented by *Lathyrus*-1 in Figure 1 and form a monophyletic group, but with 61% bootstrap support; in some trees, some of these species are placed differently, presumably in association with *L. neurolobus*. The third group of *Lathyrus* (*L. articulatus*, *L. clymenum*, and *L. ochrus*) is itself distinct and monophyletic, with
100% bootstrap support in the ML tree. The arrangement of these three *Lathyrus* groups and the *Pisum* plus *Vavilovia* group is uncertain, with the highest bootstrap support of 53% for node “d” and less than 50% support for nodes “b” and “c”, suggesting that these dichotomies may collapse when adding more data and/or taxa for further phylogenetic inferences (Figure 1).

## 2.2. *The Chronogram*

There are many useful analyses in the work of Schaefer et al. [8], one of which is a chronogram that attempted to date divergences within the Fabeae. The overall branching structure that relates to the present discussion is given in Figure 1B. There are several striking features of this structure. The first is that the proposed *Ervilia* group is basally branched, but the age estimate of this divergence overlaps slightly with the divergence of *Vicia* plus *Lens* from *Lathyrus*, *Pisum*, and *Vavilovia*. All *Vicia*, including the proposed *Ervum*, are a monophyletic lineage, but are distinct from *Lens* in this analysis [8]. This is consistent with retaining *Lens* as a separate genus.

In the chronogram, the two minor *Lathyrus* lineages are distinct from the remaining (and the majority of) *Lathyrus* species, but the age estimate for this divergence overlaps substantially with the estimated age of the divergence of *Pisum* plus *Vavilovia* from all *Lathyrus*. The estimate of the *Pisum*/*Vavilovia* divergence time overlaps, to some degree, with their divergence from the majority of *Lathyrus*, presumably reflected in the 99% bootstrap support for this clade in the ML tree.

## 3. Discussion

The phylogenetic analysis of the Fabeae presented by Schaefer et al. [8] is a thorough analysis of this tribe, and remains the best information we have regarding species and genus relationships in the Fabeae. These authors suggested several possible nomenclatural changes within the tribe that might better reflect the new understanding of species relationships which they uncovered, but did not present a firm and comprehensive set of new species and genus names. No new data have yet become available to lend weight to any of the possible new systems of species and genus names, yet a new system has been partially implemented by WCVP [9] and other authors [14–16]. This seems premature and precipitate.

Kenicer and Parsons [10] argue, “A major change is that *Pisum* and *Vavilovia* are now included in *Lathyrus*. If an alternative approach had been taken with *Pisum* and *Vavilovia*, maintaining them as separate genera, then *Lathyrus* would have to lose *L. clymenum*, *L. neurolobus* and *L. ochrus*. On balance the inclusion of *Pisum* and *Vavilovia* in *Lathyrus* entails fewer name changes and more accurately reflects the morphology of the group, meaning the inwardly hairy style is a defining character of the genus”. Whether this, in fact, entails fewer name changes depends on the number of species within *Pisum* plus *Vavilovia*. In fact, this change demands at least four name changes (*Pisum sativum*, *P. elatius*, and *P. fulvum*, plus *Vavilovia formosa*) compared to renaming four *Lathyrus* species. This numerical point of view is not the only consideration; a search in Web of Science [17] with the term “*Pisum sativum*” returned 41,916 results for this species, while a search with the term “*Lathyrus*” returned 4834 results for this genus, reflecting the focus of interest in one of the species to be renamed.

We have focused attention on the work of Schaefer et al. [8] because this is the most complete study in terms of species sampling. Nevertheless, additional studies have been published which are relevant to the issue of the taxonomic position of *Pisum* with respect to *Lathyrus*. Robledillo et al. [18] present a phylogeny, based on centromeric repeated sequences (which are unusual within *Pisum* and *Lathyrus*), which positions *Pisum* as basal to a monophyletic group which includes *Lathyrus*, but distinct from a monophyletic *Vicia*. Zhao et al. [19], based on an analysis of 15,000 nuclear genes in the Fabaceae, shows the same relationship. However, two plastid gene-based phylogenies [20,21] place *Pisum* within a monophyletic *Lathyrus* clade. In these analyses, the position of *L. clymenum* (and *L. ochrus*
in [20]) is critical to the interpretation of the relationship of *Pisum* (and *Vavilovia*) to the remainder of *Lathyrus*, but the support for a particular branching structure is low. The branching pattern of these phylogenies may depend on the species included or excluded from the analysis and also on whether the plastid or nuclear genome is being analyzed. Similarly, the nature of the nuclear DNA sequences may again have an impact, reflecting patterns of gene duplication [19] or perhaps their function [18].

We recognize that taxonomic names should reflect our understanding of species relationships; nomenclature should above all be useful for distinguishing Organismal Taxonomical Units under study. When there are substantial changes to our understanding, a new nomenclatural system is appropriate, and this should recognize due priority, as well the value of nomenclatural stability. *Pisum* has been used as a name since antiquity. However, we see no good evidence that *Pisum* and *Vavilovia* (and *Lens*) need to be renamed. This may eventually become desirable, given new knowledge which is not yet available. If or when this situation changes, we think it would be wise to implement changes that will be minimally disruptive to the scientific literature. If the creation of a small number of new genera enables the preservation of *Vicia*, *Lens*, *Lathyrus*, *Pisum*, and *Vavilovia*, then that would seem a sensible change, as indeed Schaefer et al. [8] proposed for *Ervum* and *Ervilia*. Similarly, Kenicer and Parsons [14] presented a diagram of the relationships within the Fabeae, with eight taxonomic groups combined in a way that creates four genera.

If future phylogenetic analysis does confirm that there is a monophyletic clade in which *Pisum* and *Vavilovia* are indeed embedded within *Lathyrus*, then the creation of a new genus from within *Lathyrus* might enable the retention of *Pisum* and *Vavilovia*; if so, this would seem a desirable change.

**Author Contributions:** T.H.N.E. prepared the first draft of the manuscript. P.S., N.M., J.B., and Y.B.-K. contributed to the taxonomic interpretations, as presented and edited the text. C.J.C., C.D., and N.C. edited the manuscript and contributed to the discussion. All authors have read and agreed to the published version of the manuscript.

**Funding:** T.H.N.E. and N.C. acknowledge the receipt of long-term investments from the UK Research Infrastructure Biotechnology and Biological Sciences Research Council (UKRI-BBSRC) through the National Bioscience Research Infrastructure grant (BBS/E/JI/23NB0001) and the UK Department for Environment, Food, and Rural Affairs (Defra) through the Provision and Maintenance of the Pea Genebank to Facilitate R&D Need grant (C5515). C.D. acknowledges funding support from UKRI BBSRC (BBS/E/J/000PR9799). P.S. acknowledges funding from the Grant Agency of Palacky University, PrF-2024-001.

**Acknowledgments:** We thank Julie Hofer, Andrey Sinjushin, Valery Malecot, and Tom Warkentin for useful discussions.

**Conflicts of Interest:** The authors declare no conflicts of interest.

## References

1. Lewis, G.P.; Schrire, B.; Mackinder, B.; Lock, M. *Legumes of the World*; Kew Publishing: Kew, UK, 2005.

2. Azani, N.; Babineau, M.; Bailey, C.D.; Banks, H.; Barbosa, A.R.; Pinto, R.B.; Boatwright, J.S.; Borges, L.M.; Brown, G.K.; Bruneau, A.; et al. A new subfamily classification of the Leguminosae based on a taxonomically comprehensive phylogeny: The Legume Phylogeny Working Group (LPWG). *Taxon* **2017**, *66*, 44–77. [CrossRef]

3. Wojciechowski, M.F.; Sanderson, M.J.; Hu, J.M. Evidence on the monophyly of Astragalus (Fabaceae) and its major subgroups based on nuclear ribosomal DNA ITS and chloroplast DNA trnL intron data. *Syst. Bot.* **1999**, *24*, 409–437. [CrossRef]

4. Duan, L.; Li, S.-J.; Su, C.; Sirichamorn, Y.; Han, L.-N.; Ye, W.; Lôc, P.K.; Wen, J.; Compton, J.A.; Schrire, B.; et al. Phylogenomic framework of the IRLC legumes (Leguminosae subfamily Papilionoideae) and intercontinental biogeography of tribe Wisterieae. *Mol. Phylogenet. Evol.* **2021**, *163*, 107235. [CrossRef] [PubMed]

5. Zhu, A.; Guo, W.; Gupta, S.; Fan, W.; Mower, J.P. Evolutionary dynamics of the plastid inverted repeat: The effects of expansion, contraction, and loss on substitution rates. *New Phytol.* **2016**, *209*, 1747–1756. [CrossRef] [PubMed]

6. Choi, I.-S.; Jansen, R.; Ruhlman, T. Lost and Found: Return of the Inverted Repeat in the Legume Clade Defined by Its Absence. *Genome Biol. Evol.* **2019**, *11*, 1321–1333. [CrossRef] [PubMed]

7. Kupicha, F.K. Vicieae. In *Advances in Legume Systematics*; Polhill, R.M., Raven, P.M., Eds.; Royal Botanic Gardens: Kew, UK, 1981; pp. 377–381.
8. Schaefer, H.; Hechenleitner, P.; Santos-Guerra, A.; Menezes de Sequeira, M.; Pennington, R.T.; Kenicer, G.; Carine, M.A. Systematics, biogeography, and character evolution of the legume tribe Fabeae with special focus on the middle-Atlantic island lineages. *BMC Evol. Biol.* **2012**, *12*, 250. [CrossRef]

9. Royal Botanic Gardens, Kew. The World Checklist of Vascular Plants (WCVP): Fabaceae. Checklist Dataset. 2023. Available online: https://www.gbif.org/dataset/f7053f73-74fb-4c9f-ab63-de28c61140c2 (accessed on 9 May 2024).

10. Ferguson, M.E.; Maxted, N.; van Slageren, M.V.; Robertson, L.D. A re-assessment of the taxonomy of *Lens* Mill. (Leguminosae, Papilionoideae, Vicieae). *Bot. J. Linn. Soc.* **2000**, *133*, 41–59. [CrossRef]

11. Hellwig, T.; Abbo, S.; Sherman, A.; Coyne, C.; Saranga, Y.; Lev-Yadun, S.; Main, D.; Zheng, P.; Ophir, R. Limited divergent adaptation despite a substantial environmental cline in wild pea. *Mol. Ecol.* **2020**, *29*, 4322–4336. [CrossRef] [PubMed]

12. Hellwig, T.; Abbo, S.; Sherman, A.; Ophir, R. Prospects for the natural distribution of crop wild-relatives with limited adaptability: The case of the wild pea *Pisum fulvum*. *Plant Sci.* **2021**, *310*, 110957. [CrossRef] [PubMed]

13. Hellwig, T.; Abbo, S.; Ophir, R. Phylogeny and disparate selection signatures suggest two genetically independent domestication events in pea (Pisum L.). *Plant J.* **2022**, *110*, 419–439. [CrossRef] [PubMed]

14. Kenicer, G.; Parsons, R. *Lathyrus the Complete Guide*; Royal Horticultural Society: Peterborough, UK, 2021.

15. Rix, M.; Nesbitt, M.; King, C. 1063. Lathyrus oleraceus Lam. *Curtis's Bot. Mag.* **2023**, *40*, 197–205. [CrossRef]

16. Kosterin, O.E. Abyssinian pea (*Lathyrus schaeferi* Kosterin nom. Nov. pro *Pisum abyssinicum* A. Br.) is a problematic taxon. *Vavilovskii Zhurnal Genet. I Sel. Vavilov J. Genet. Breed.* **2017**, *21*, 158–169. [CrossRef]

17. Web of Science. Available online: https://www.webofscience.com/wos/alldb/basic-search (accessed on 19 May 2024).

18. Ávila Robledillo, L.; Neumann, P.; Koblížková, A.; Novák, P.; Vrbová, I.; Macas, J. Extraordinary Sequence Diversity and Promiscuity of Centromeric Satellites in the Legume Tribe Fabeae. *Mol. Biol. Evol.* **2020**, *37*, 2341–2356. [CrossRef] [PubMed]

19. Zhao, Y.; Zhang, R.; Jiang, K.-W.; Qi, J.; Hu, Y.; Guo, J.; Zhu, R.; Zhang, T.; Egan, A.N.; Yi, T.-S.; et al. Nuclear phylotranscriptomics and phylogenomics support numerous polyploidization events and hypotheses for the evolution of rhizobial nitrogen-fixing symbiosis in Fabaceae. *Mol. Plant.* **2021**, *14*, 748–773. [CrossRef] [PubMed]

20. Moghaddam, M.; Kazempour-Osaloo, S. Extensive survey of the ycf4 plastid gene throughout the IRLC legumes: Robust evidence of its locus and lineage specific accelerated rate of evolution, pseudogenization and gene loss in the tribe Fabeae. *PLoS ONE* **2020**, *15*, e0229846. [CrossRef] [PubMed]

21. da Silva, F.G.; de Moraes Filho, R.M.; Martins, L.S.; da Silva Ramos, R.; Silva, G.C. Plastid marker-based phylogeny reveals insights into relationships among Papilionoideae species. *Genet. Resour. Crop. Evol.* **2024**, *71*, 439–454. [CrossRef]

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