Neod 131 1

A Journal of Neotropical Biodiversity ISSN 2358-2847 (online)
ARTICLE 02 Sep 2020 DOI: 10.13102/neod.131.1
A molecular phylogeny of the pantropical papilionoid

legume Aeschynomene supports reinstating the
ecologically and morphologically coherent genus Ctenodon
Domingos B. O. S. Cardoso1 | Cilene M. J. Mattos2 | Fabiana Filardi2 |
Alfonso Delgado-Salinas3 | Matt Lavin4 | Pedro L. R. de Moraes5 |
Fernando Tapia-Pastrana6 | Haroldo C. de Lima2
ABSTRACT
1
Instituto de Biologia, Universidade Federal
da Bahia, Rua Barão de Jeremoabo, s.n.,
Ondina, 40170-115, Salvador, Bahia, Brazil.
The pantropical and morphologically diverse genus Aeschynomene
2
Instituto de Pesquisas Jardim Botânico do
Rio de Janeiro, Rua Pacheco Leão, 915, (Leguminosae, Papilionoideae) comprises ca. 180 species, but its
22460-030, Rio de Janeiro, RJ, Brazil.
monophyly has never been comprehensively addressed in a molecular
3
Departamento de Botánica, Instituto de phylogenetic context of the dalbergioid clade. Here we have assembled the
Biología, Universidad Nacional Autónoma de
México, Apartado postal 70-233, 04510, most complete DNA sequence data (nuclear ribosomal ITS/5.8S and the
CDMX, México.
plastid genes trnL intron and protein-coding matK), representatively
4
Plant Sciences and Plant Pathology, sampled both in Aeschynomene and the closely related Dalbergia and
Montana State University, Bozeman,
Montana 59717, USA. Machaerium, as well all currently accepted genera from the dalbergioids.
The Bayesian and maximum likelihood phylogenetic analyses
5
Universidade Estadual Paulista “Júlio de
Mesquita Filho,” Instituto de Biociências, Rio unequivocally confirm the non-monophyly of Aeschynomene. The
Claro, São Paulo, Brazil.
pantropically distributed Aeschynomene sect. Aeschynomene, often
6
Facultad de Estudios Superiores Zaragoza,
inhabiting riparian and wetland habitats, is paraphyletic with respect to
UNAM, Laboratorio de Genecología, Batalla 5
de Mayo s/n esquina Fuerte de Loreto, Col. the mostly African genera Bryaspis, Cyclocarpa, Geissaspis, Humularia,
Ejército de Oriente, Iztapalapa, C.P. 09230,
Ciudad de México, Mexico. Kotschya, and Smithia, as well as the South American monotypic genus
1 Soemmeringia. The strongly supported and morphologically coherent
Correspondence: cardosobot@gmail.com
clade comprising the American species traditionally classified within
Aeschynomene sect. Ochopodium, with a greater ecological predilection for
neotropical fire-prone savannas and the seasonally dry tropical forests and
woodlands, is here elevated to generic rank. As such, this lineage is re-
established as the previously monotypic genus Ctenodon. A complete
description of Ctenodon and 78 new combinations for the American
species and associated infraspecific taxa are provided, as well as 24
lectotypifications.
1 | www.neodiversity.org Neodiversity 13: 1–38. 2020

Cardoso et al. – Molecular phylogeny of Aeschynomene
1 | INTRODUCTION
Molecular systematics have largely enhanced our understanding of evolutionary relationships
in the Leguminosae in the last decades, providing the basis for new phylogenetic realignments (e.g.
Wojciechowski et al. 2004; Bruneau et al. 2008; Cardoso et al. 2012a, 2012b; Gagnon et al. 2013,
2016; Koenen et al. 2020), placing phylogenetically enigmatic genera (e.g. Cardoso et al. 2015,
2017; Queiroz et al. 2017), and reorganizing subfamily and tribal classifications (e.g. Queiroz et al.
2015; LPWG 2017; De-la-Estrella et al. 2018). While there are several monotypic or species-poor
genera still lacking any molecular data for assessing their phylogenetic placement, comprehensively
addressing the monophyly of morphologically-heterogenous and species-rich genera is another
important step to advance the legume systematics (Cardoso et al. 2013; LPWG 2013). This is
particularly the case of Aeschynomene L. (Papilionoideae), a pantropically distributed genus of about
180 species (Klitgaard & Lavin 2005), of which at least 80 occur predominantly in the fire-prone
savannas and seasonally dry tropical forests and woodlands, but also in Oak forests, pine-oak
forests as well in pine woodlands.
The American species of Aeschynomene were the focus of a taxonomic revision by Rudd (1955),
and since many new species have been described thus far (Rudd 1975, 1989; Lewis 1985, 1992,
1993; Queiroz & Cardoso 2008; Delgado-Salinas & Sotuyo 2012; Silva & Antunes 2014; Antunes &
Silva 2017, 2019; Cardoso et al. 2019). Some have added striking new morphological features that
were previously unknown in the genus. For example, Aeschynomene was often described as having
pinnately-compound leaves, but aphyllous and unifoliolate species have been discovered (Lewis
1992; Cardoso et al. 2019). In addition to the heterogeneity in terms of leaf morphology, there is a
clear morphological separation in the traditional concept of Aeschynomene that is reflected in its
infrageneric classification (Vogel 1838; Rudd 1955; Fernandes 1996). Accordingly, the species with
peltate or medifixed stipules, loment fruits with articles separated by septa, and a bilabiate calyx
were classified in Aeschynomene sect. Aeschynomene, whilst Aeschynomene sect. Ochopodium
comprised the species with basifixed stipules, loment articles separated by an isthmus, and
campanulate calyx.
Many species-rich legume genera as traditionally circumscribed have already been thoroughly
investigated in molecular phylogenies, such as the mimosoids Inga Mill. (Nicholls et al. 2015) and
Mimosa L. (Simon et al. 2011), the caesalpinioid Caesalpinia Plum. ex L. sensu lato (Gagnon et al.
2013, 2016) and Chamaecrista Moench (Conceição et al. 2009; Rando et al. 2016), the cercidoid
Bauhinia Plum. ex L. s.l. (Sinou et al. 2009), and the papilionoids Astragalus L. (Wojciechowski et al.
1999; Wojciechowski 2005), Dalbergia L.f. (Vatanparast et al. 2013), Indigofera L. (Schrire et al.
2009), Lupinus L. (Drummond et al. 2012), Swartzia (Torke & Schaal 2008), and Vigna Savi s.l.
(Delgado-Salinas et al. 2011). However, the systematics of Aeschynomene has been evaluated at
most by only a handful DNA sequences (Lavin et al. 2001; Ribeiro et al. 2007; Cardoso et al. 2013,
2019; LPWG 2017). Nevertheless, all molecular phylogenetic analyses have indicated that the
traditionally circumscribed Aeschynomene could represent an intermingled collection of distinct

evolutionary lineages. Lavin et al. (2001) were the first to recognize the non-monophyly of
Aeschynomene using three molecular (ITS, matK-trnK, and trnL) and a morphological data set, which
supported A. sect. Ochopodium apart from Aeschynomene sensu stricto. Later, Ribeiro et al. (2007)
showed the most consistent evidence that A. sect. Ochopodium could be more closely related to
Dalbergia and Machaerium Pers. than to A. sect. Aeschynomene. Despite their relatively good
sampling within the latter genera, Aeschynomene was represented by just nine species in a combined
analysis of nuclear ribosomal ITS/5.8S and plastid trnL intron DNA sequence data. Although focused
on understanding the evolution of nodulation in Aeschynomene, the studies of Chaintreuil et al.
(2013) and Brottier et al. (2018) also provided additional evidence of the non-monophyly of
Aeschynomene, after including an improved taxon sampling of 31 and 42 species of the genus,
respectively.
Here we assemble the most comprehensive molecular dataset to address the monophyly of
Aeschynomene s.l. in a phylogenetic analysis, not just with denser taxon sampling but also
combining variation from ITS/5.8S, trnL intron and the protein-coding gene matK. We also explore
whether the contrasting ecology and evolutionary history of A. sect. Aeschynomene and A. sect.
Ochopodium associated to distinct biomes can be expressed in terms of differences between climatic
niche variation among extant species in the different clades.
2 | MATERIALS AND METHODS

Taxon sampling and DNA sequence data—The monophyly of Aeschynomene s.l. was
evaluated using broad taxon sampling of nuclear ribosomal ITS/5.8S and the plastid genes trnL
intron and protein-coding matK. These genes have often been successfully used in phylogenetic
studies at several taxonomic levels in legumes (e.g. Bruneau et al. 2008; Cardoso et al. 2012a, 2013,
2015, 2017; Queiroz et al. 2015, 2017; LPWG 2017; De-la-Estrella et al. 2018), particularly in the
dalbergioids (Lavin et al. 2001; Ribeiro et al. 2007; Cardoso et al. 2012b; Fortuna-Peres et al. 2018).
We assembled a matK dataset for an individual analysis including all currently accepted genera
of the dalbergioid legume clade (Lavin et al. 2001; Cardoso et al. 2013), but maximizing sampling as
much as possible across the taxonomic diversity within Aeschynomene (53 spp. sampled), Dalbergia
(36 spp. sampled), and Machaerium (51 spp. sampled). The early branching amorphoid genera were
used as outgroups, so that the total sampling of the individual matK dataset included 198 species.
We also built an individual ITS/5.8S dataset and a concatenated dataset (ITS/5.8S, matK, and trnL
intron) focusing on just the Dalbergia clade, and using the two genera of the Pterocarpus clade as
outgroups. A total of 54 species of Aeschynomene, 17 of Dalbergia, and 15 of Machaerium were
sampled for the ITS/5.8S dataset, while the concatenated dataset included 67 species of
Aeschynomene, 41 of Dalbergia, and 54 of Machaerium. These non-overlapping numbers in taxon
coverage for the concatenated dataset reflects the fact that we wanted to maximize taxon sampling
even if some terminals had incomplete data. Combined phylogenetic analyses of empirical data and
computational simulations have largely demonstrated that missing data do not negatively affect the

results (Wiens 2003). Rather, analyses including taxa with less than 50% of data can even help
breaking up long branches and improving phylogenetic accuracy (Wiens 2005; Wiens & Tiu 2012).
Most sequences used here were already published in our previous molecular systematic studies
(Lavin et al. 2001; Cardoso et al. 2012b, 2019; LPWG 2017), but our sampling was also largely
increased by recent evolutionary-focused studies in Aeschynomene nodulation (Chaintreuil et al.
2013; Brottier et al. 2018), all of which are taxonomically curated, vouchered, and have sequences
available in GenBank (www.ncbi.nlm.nih.gov). GenBank accession numbers follow the terminal taxon
labels in the presented phylogenetic trees. For the 15 newly generated sequences of ITS/5.8S, DNA
isolation, polymerase chain reaction (PCR), amplifications, and sequencing were performed as
described in our previous study (e.g. Lavin et al. 2001). All these new DNA sequences and associated
voucher information are deposited in GenBank, and they are readily identified in the ITS
phylogenetic tree by the accessions beginning with “MT”.
All DNA datasets were aligned manually with AliView version 1.17.1 (Larsson 2014). Accurate
homologies among the nucleotide sites of the protein-coding matK gene were assessed by inserting
gaps corresponding to complete codons in amino acid translated sequences (Wojciechowski et al.
2004). The individual DNA alignments were combined using a newly developed R package
(D.Cardoso, unpubl. data) with functions that automatically build the concatenated dataset with
missing data (sensu Wiens 2003).
Estimation of phylogenetic relationships—We used maximum likelihood and Bayesian
inference as the phylogenetic reconstruction approaches to estimate evolutionary relationships
based on the individual ITS/5.8S and matK datasets of and the concatenated dataset of ITS/5.8S,
matK, and trnL intron sequences. The maximum likelihood tree was inferred with RAxML v8.2.812
(Stamatakis 2014), using the evolutionary model GTRGAMMA, where the invariant sites and gamma
distribution were estimated during the run. Branch supports (BS) (Felsenstein 1985) were estimated
by letting RAxML halt bootstrapping automatically (Stamatakis et al. 2008). The Bayesian analyses
were run in MrBayes version 3.2.6 (Ronquist et al. 2012), using a best-fit nucleotide substitution
model (GTR + I + G) selected via the Akaike information criterion (AIC) in jmodeltest2 (Darriba et al.
2012). In two separate runs of a Metropolis-coupled Markov Chain Monte Carlo (MCMC) permutation
of parameters, eight simultaneous chains run for 15 million generations, sampling one tree at each
10,000th generation. A Bayesian majority-rule consensus tree at 50% was produced after a burn-in
of 25%, where the group frequencies represent posterior probabilities (PP) (Huelsenbeck et al.
2002). We used FigTree version 1.4.0 for visualization of the Bayesian tree (Rambaut 2012) and
Adobe Illustrator for final editing. All phylogenetic analyses were run in the CIPRES Science Gateway
v. 3.3 on-line portal (www.phylo.org) (Miller et al. 2010).
Climate data—Because of conspicuous visual habitat differences between Aeschynomene sect.
Ochopodium, centered in the savanna and seasonally dry tropical forests and woodlands (Rudd
1955; Queiroz & Cardoso 2008; Silva & Antunes 2014; Cardoso et al. 2019), and Aeschynomene sect.
Aeschynomene, most abundant and diverse in low elevation riparian and wetland settings (Rudd
1955; Antunes & Silva 2017, 2019), we analyzed climate data associated with collection information

accessed from Global Biodiversity Information Facility (GBIF, www.gbif.org). We used the R package
‘rgbif’ (Chamberlain et al. 2020) to retrieve the ~36,000 herbarium records for all American species
of A. sect. Ochopodium and all species of A. sect. Aeschynomene and the phylogenetically related
mostly African genera sampled in our combined molecular phylogeny. Data cleaning involved first
standardizing the collector name (tagged in GBIF as “recordedBy”) and associated number
(“recordNumber”). We used a newly developed package of R functions (D.Cardoso, unpubl. data)
which automatically clean the “recordedBy” column in the database by keeping just a uniformly
formatted name of the principal collector. By doing so, we could exclude the duplicate specimens
among the different herbaria more confidently. We then excluded records without any geographical
coordinates or collector names, as well as any georeferenced records collected before 1930, so as to
exclude the mostly erroneously georeferenced records of old, historical collections. Then we
excluded duplicates by identifying those records with the same principal collector and associated
number. After that we used the R package ‘CoordinateCleaner’ for further cleaning the database
with respect to coordinates at sea, as country and province centroids, country capitals, urban areas,
as well as around biodiversity institutions, which often comes from cultivated individuals (Zizka et al.
2019).
Because we also excluded all rampant, widespread species (e.g. A. americana, A. rudis, A. histrix,
etc.) with an estimated range size greater than 800,000 Km 2, as calculated with the R package ‘letsR’
(Vilela & Villalobos 2015), our database for the analysis of climatic niche variation ended up with
just ~3,500 records from among 48 species of A. sect. Aeschynomene and related genera, and 51
American species of A. sect. Ochopodium. The reasoning here is that the signature of a constraining
ecology would be left only on those relatively confined, small-range-sized species, since the more
widespread, mostly weedy herbs usually break ecological constraints and have a too large climatic
niche breadth, erasing any signature of a putatively phylogenetically inherited ecology. Using the R
package ‘raster’ (Hijmans et al. 2020), we extracted WorldClim data (Fick & Hijmans 2017) and 250
m spatial resolution elevation data (http://srtm.csi.cgiar.org/srtmdata/) from the final set of
geolocated Aeschynomene collections as well as for the mostly African related genera. Climate
variables with greatest variance, identified using a standard biplot of the principal components
analysis (PCA), were then analyzed for any significant differences between the Aeschynomene
sections and related genera.
Taxonomic treatment—This work is based on standard revisionary studies, including a
comprehensive literature review and analysis of specimens deposited in herbaria (acronyms
according to Thiers 2020 [continuously updated]). The taxonomy adopted here at species and
infraspecific level follows mostly Rudd’s (1955, 1959, 1967, 1972, 1975, 1989) revisionary studies
on Aeschynomene s.l., but with subsequent updates (Lewis 1985, 1992, 1993; Fernandes 1996;
Queiroz & Cardoso 2008; Queiroz 2009; Delgado-Salinas & Sotuyo 2012; Silva & Antunes 2014;
Vanni 2016; Antunes & Silva 2019; Cardoso et al. 2019). Additional information, including detailed
data and images for the specimens and names examined, has been consulted in the following
databases: Flora do Brasil 2020, GBIF, IPNI, JSTOR Global Plants (https://plants.jstor.org), The Plant

List, and Tropicos®. Locations of known but unseen duplicates of specimens are included. Barcode
numbers are cited consistently whenever available. Homotypic synonyms are indicated by the
symbol “≡”, whereas heterotypic synonyms are indicated by “=”.
3 | RESULTS AND DISCUSSION
The Bayesian and maximum likelihood phylogenetic single region analyses of matK and
ITS/5.8S datasets strongly resolve several key nodes among Aeschynomene s.l. and related genera
(Figs. 1-2). These individual analyses essentially provide independent tests for the non-monophyly
of Aeschynomene s.l. which collectively falls in two main clades. Such results are also largely
congruent with the more strongly resolved combined analysis (Fig. 3).
The phylogenetic analysis of matK and combined data strongly support the Aeschynomene
sect. Aeschynomene as paraphyletic with respect to the mostly African genera Bryaspis P.A.Duvign.,
Cyclocarpa Afzel. ex Urb., Geissaspis Wight & Arn., Humularia P.A.Duvign., Kotschya Endl., and
Smithia Aiton, as well as the South American monotypic genus Soemmeringia Mart. (Figs. 1, 3).
Interestingly, species in this clade, particularly those in A. sect. Aeschynomene, are often hydrophytes
growing in swamps, temporary or permanent ponds, and along streams and riverbanks (Rudd 1955;
Antunes & Silva 2017; Chaintreuil et al. 2013), an ecological predilection that perhaps explains why
stem nodulation ability have evolved several times in this clade (Chaintreuil et al. 2013; Brottier et
al. 2018). Whether the whole clade will be circumscribed into an expanded concept of
Aeschynomene s.str. amalgamated with the remaining genera should be the focus of a more resolved
phylogeny and detailed comparative morphological studies. We believe that further nomenclatural
changes here would be precipitated because of the poor taxon sampling across the morphologically
and geographically diverse related genera from Africa and Asia. Also, our individual ITS Bayesian
analysis recovered A. sect. Aeschynomene as a weakly supported sister clade of one comprising the
abovementioned mostly African genera. Some phylogenetic studies of legumes involving next-
generation phylogenomic approaches (Nicholls et al. 2015; Ojeda et al. 2019; Duan et al. 2020;
Koenen et al. 2020; Zhang et al. 2020) suggest improved resolution could be attained by applying
them to A. sect. Aeschynomene.
All our analyses are also congruent in strongly resolving the sister relationship between
Aeschynomene sect. Ochopodium and Machaerium, and then this clade as sister to Dalbergia (Figs. 1-
3). These relationships have been shown in previous studies with lower taxon sampling focused
primarily on the broader dalbergioid legumes (Lavin et al. 2001; Cardoso et al. 2012, 2013) or just
on Aeschynomene s.l. and related genera (Ribeiro et al. 2007; Chaintreuil et al. 2013; Vatanparast et
al. 2013; Brottier et al. 2018). Interestingly, species of A. sect. Ochopodium are morphologically
similar to Machaerium and Dalbergia by sharing the 5-lobed, campanulate calyx and often having
pubescent outer surface of the standard petal.

Psorodendron spinosum AY391822

Psorothamnus scoparius AY391821
Dalea lanata AY391790
Marina scopa AY391811
1.0/100 Amorpha fruticosa AY391785
Parryella filifolia AY391812
Apoplanesia paniculata AF270860
Errazurizia megacarpa AY391804
Eysenhardtia orthocarpa AY386909
Adesmia volckmanii AF142690
Chaetocalyx brasiliensis AF270866
1.0/100 Nissolia hirsuta AF270868
Poiretia angustifolia AF270864
Amicia glandulosa AF203583
Zornia orbiculata KX595215
Discolobium psoraleifolium AF270874
Riedeliella graciliflora KY045894
Platymiscium stipulare AF270872
1.0/90 Brya ebenus AF270876
Cranocarpus martii AF270875
Cascaronia astragalina AF272072
Geoffroea spinosa AF270879
Fiebrigiella gracilis AF203590
Fissicalyx fendleri AF272063
Chapmannia gracilis AF203592
Arachis pintoi AF203596
Stylosanthes capitata AF203595
Acosmium lentiscifolium JX124417
Grazielodendron riodocense AF270862
Platypodium elegans AF270877
Etaballia dubia AF272074
Paramachaerium ormosioides KP177927
1.0/100 Pterocarpus monophyllus KP177928
Maraniona lavinii AY247263
Centrolobium microchaete EU401409
Ramorinoa girolae AF270881
Inocarpus fagifer AF270878
Tipuana tipu AF270882
Weberbauerella brongniartioides AF272075
Diphysa americana AF203574
Zygocarpum yemenense AF203573
Pictetia aculeata AF203577
Ormocarpum kirkii AF20357
0.6/46 Ormocarpopsis calcicola AF203568
Peltiera nitida GU951672
Aeschynomene fluminensis KX687571
Aeschynomene parviflora MF281589
Aeschynomene villosa KX687572
Aeschynomene americana MF281585
Aeschynomene unijuga MF281587
Aeschynomene montevidensis MF281590
Aeschynomene paraguayensis MF281591
Aeschynomene tambacoundensis MF281597
1.0/98 1.0/100 Aeschynomene filosa KX932464
Aeschynomene rostrata MF281595
Aeschynomene deamii MF281594
Aeschynomene pratensis KX687576
Aeschynomene pratensis MF281611
Aeschynomene selloi MF281610
Aeschynomene sensitiva MF281609
Aeschynomene
Aeschynomene ciliata KX687578
Aeschynomene denticulata MF281599
Aeschynomene magna MF281600
Aeschynomene indica AF272084
Aeschynomene scabra KX687579
Aeschynomene evenia KX687580
Aeschynomene pluriarticulata MF281603
Aeschynomene rudis AF270869
Aeschynomene virginica AF203586
Aeschynomene patula MF281592
prob Soemmeringia semperflorens MF281614
1 Geissaspis cristata MF281620
Geissaspis tenella MF281621
Cyclocarpa stellaris MF281615
Smithia abyssinica MF281616
Smithia sensitiva MF281617
Kotschya strigosa MF281631
Kotschya africana MF281630
Kotschya lutea MF281629
Aeschynomene pulchella MF281627
Aeschynomene bella MF281624
Aeschynomene pygmaea MF281626
Humularia descampsii MF281632
Aeschynomene heurckeana MF281625
Humularia kapiriensis MF281633
Aeschynomene abyssinica MF281623
Aeschynomene elaphroxylon KT821189
1.0/100 Aeschynomene uniflora KT821213
Bryaspis lupulina MF281618
Bryaspis humularioides MF281619
Aeschynomene crassicaulis KT821185
Aeschynomene pfundii KT821195
Aeschynomene afraspera KT821180
Aeschynomene aspera KT821184
0.5 Aeschynomene fluitans KT821192
Aeschynomene nilotica KT821194
Aeschynomene deightonii MF281612
Aeschynomene mediocris MF281613
Aeschynomene cristata KT821186
Aeschynomene schimperi KX687574
Dalbergia ovalis KY046042
Dalbergia miscolobium KY200512
Dalbergia spruceana KY045870
0.01 Dalbergia inundata KY045978
Dalbergia elegans KY046039
Dalbergia villosa KY118273
Dalbergia acuta KY118272
Dalbergia foliolosa KX816356
1.0/99 Dalbergia densiflora KY046038
Dalbergia glaucescens KY046041
Dalbergia
Dalbergia arbutifolia LC194935
Dalbergia retusa JQ587584
Dalbergia melanoxylon LC194942
Dalbergia boehmii LC194934
Dalbergia cochinchinensis FR854102
Dalbergia latifolia LC194940
Dalbergia lanceolaria JX506655
Dalbergia nigrescens FR854107
Dalbergia oliveri LC194937
Dalbergia polyadelpha LC194929
Dalbergia nigra KX816360
Dalbergia bracteolata LC194933
Dalbergia glabra JQ587581
Dalbergia sisso LC194943
Dalbergia rimosa HM049527
Dalbergia tonkinensis FR854117
Dalbergia maritinii LC194941
Dalbergia humbertii LC194930
Dalbergia lemurica LC194939
Dalbergia armata JF270735
1.0/100 Dalbergia congestiflora AF142696
Dalbergia decipularis KY045969
Dalbergia hortensis KY045970
Dalbergia riparia KY045992
Dalbergia brasiliensis LC194932
Dalbergia frutescens KX816358
Aeschynomene mollicula KX687560
Aeschynomene fascicularis AF142695
Aeschynomene purpusii AF270870
Aeschynomene brasiliana KX687568
Aeschynomene chicocesariana KX932467
1.0/100
Aeschynomene riedeliana KX687562
Aeschynomene scoparia KX687561 Ctenodon
Aeschynomene viscidula KX687567
Aeschynomene brevipes KX687563
Aeschynomene paniculata KX687564
Aeschynomene elegans KX932465
Aeschynomene falcata KX932466
Machaerium kegelii JQ587754
Machaerium quinata KX898297
Machaerium pedicellatum KX898294
1.0/99 Machaerium ruddianum KY045988
Machaerium firmum KX898272
Machaerium fulvovenosum KX898274
Machaerium incorruptibile KX898280
Machaerium leucopterum KX898284
Machaerium scleroxylon KX898300
Machaerium jobimianum KX898282
Machaerium obovatum KX898291
matK Machaerium floridum KY045954
Machaerium macaense KY045995
Machaerium nyctitans KX898289
1.0/96
Machaerium
Machaerium ovalifolium KX898292
Machaerium tortipes KY045877
Machaerium caratinganum KX898268
Machaerium aculeatum KX898258
Machaerium condensatum KX898269
Machaerium ferox KY045905
Machaerium isadelphum KX898281
Machaerium amplum KX898260
Machaerium biovulatum KX898261
Machaerium riparium KJ593923
Machaerium milleflorum KJ593919
Machaerium salvadorense KJ593925
Machaerium eriocarpum KY046057
Machaerium declinatum KX898271
Machaerium uncinatum KX898304
Machaerium cobanense JQ589879
Machaerium hirtum KY045906
Machaerium lunatum KM521425
Machaerium pittieri JQ587760
Machaerium seemannii JQ587764
Machaerium aureum KX898259
Machaerium oblongifolium KX898290
Machaerium floribundum KY046058
Machaerium punctatum KX898296
Machaerium glabrum KX898275
Machaerium millei KX898285
Machaerium nigrum KX898287
Machaerium stipitatum KX898301
Machaerium lanceolatum KX898283
Machaerium ternatum KX898303
Machaerium brasiliense KX898263
Machaerium salzmannii KX898298
Machaerium acutifolium KX898256
Machaerium cantarellianum KX898267
Machaerium opacum KY045997
Machaerium paraguariense KY046059
Machaerium villosum KX898305
Figure 1. A matK Bayesian majority-rule consensus phylogram comprehensively-sampled across all lineages of the
dalbergioid legumes (199 terminals by 1617 aligned sites). Aeschynomene (sensu Rudd 1955) appears as non-
monophyletic, where Aeschynomene sect. Ochopodium, herein as the reinstated genus Ctenodon, is more closely
related to Machaerium. Posterior probabilities from 0.5 to 1.0 are shown as color gradient from red to black on the
branches. Numbers on some key nodes are posterior probabilities and maximum likelihood bootstrap values.
GenBank accession numbers follow the terminal taxon labels.

Pterocarpus indicus JN083482

0.99/79 Centrolobium sclerophyllum EU401440
Diphysa ormocarpoides AF068169
Pictetia aculeata AF068175
Zygocarpum yemenense AF189051
Ormocarpopsis itremoensis AF068149
Ormocarpum keniense AF068155
Smithia abyssinica KC560763
Kotschya africana AB828601
Kotschya lutea KC560762
0.74/- Soemmeringia semperflorens AF189027
Geissaspis cristata AB828602
Bryaspis lupulina AF204234
Bryaspis humularioides AB828600
Aeschynomene montevidensis KC540621
Aeschynomene
Aeschynomene filosa EF451063
1.0/99 Aeschynomene americana FM242591
Aeschynomene villosa FM242618
Aeschynomene fluminensis FM242598
Aeschynomene parviflora KC540620
0.99/79 Aeschynomene abyssinica KC540629
Aeschynomene bella KC540628
0.79/- Aeschynomene crassicaulis FM242594
Aeschynomene pfundii AF189026
Aeschynomene elaphroxylon KC540624
Aeschynomene uniflora FM242635
Aeschynomene cristata FM242625
Aeschynomene schimperi FM242633
Aeschynomene aspera FM242623
0.72/- Aeschynomene afraspera FM242585
Aeschynomene fluitans KC540622
Aeschynomene nilotica KC560767
Aeschynomene deamii KC540625
Aeschynomene tambacoundensis FM242634
Aeschynomene pratensis KC899209
0.88/58 Aeschynomene sensitiva FM242614
Aeschynomene ciliata FM242624
Aeschynomene denticulata FM242626
Aeschynomene evenia KC163296
Aeschynomene indica KC560765
Aeschynomene scabra FM242605
Aeschynomene rudis FM242609
Aeschynomene hispida FM242592
Aeschynomene virginica KC560766
Weberbauerella brongniartioides AF189028
Dalbergia ecastaphyllum EF451071
Dalbergia ovalis EF451073
Dalbergia foliolosa AF189022
prob Dalbergia miscolobium EF451070
1 1.0/96 Dalbergia elegans EF451066
Dalbergia cuiabensis EF451065
Dalbergia villosa EF451068
0.71/-
Dalbergia cochinchinensis KM521367
Dalbergia oliveri KM521397
Dalbergia nigrescens KM521386
Dalbergia
Dalbergia assamica KM521356
Dalbergia lanceolaria KM521379
Dalbergia melanoxylon KM276150
Dalbergia brasiliensis EF451076
Dalbergia congestiflora AF068140
Dalbergia rimosa KM521403
Dalbergia sissoo AF189023
Aeschynomene viscidula MT902881
Aeschynomene brevifolia KC540626
1.0/93
Aeschynomene falcata FM242597
Aeschynomene elegans MT902883
1.0/100 Aeschynomene egena MT902865
Aeschynomene tumbezensis MT902862
Aeschynomene chicocesariana MK881625
Aeschynomene vogelii EF451089
0.5 Aeschynomene riedeliana MT902888
Aeschynomene brasiliana KC540627
Aeschynomene petraea MT902870
Aeschynomene simulans MT902869
Ctenodon
Aeschynomene hintonii MT902868
Aeschynomene lyonnetii MT902867
Aeschynomene palmeri MT902866
Aeschynomene histrix FM242599
1.0/66 Aeschynomene paniculata EF451086
0.03 Aeschynomene brevipes MT902882
Aeschynomene martii EF451088
Aeschynomene mollicula MT902871
Aeschynomene fascicularis AF189025
Aeschynomene nivea MT902872
Aeschynomene purpusii MT902876
Aeschynomene compacta MT902873
Machaerium nictitans EF451083
Machaerium scleroxylon EF451084
Machaerium aculeatum EF451080
Machaerium hirtum EF451081
Machaerium lunatum KM521421
ITS/5.8S
0.98/81 Machaerium salvadorense KM521422
Machaerium gracile EF451099
Machaerium
Machaerium millei KM521424
Machaerium inundatum AF203564
Machaerium oblongifolium EF451096
Machaerium glabrum EF451095
Machaerium stipitatum EF451092
Machaerium villosum EF451094
Machaerium acutifolium EF451090
Machaerium opacum EF451098
Figure 2. An ITS/5.8S Bayesian majority-rule consensus phylogram (101 terminals by 743 aligned sites) showing the
non-monophyly of Aeschynomene (sensu Rudd 1955), where A. sect. Ochopodium, herein as the reinstated genus
Ctenodon, is strongly supported as more closely related to Machaerium. Posterior probabilities from 0.5 to 1.0 are
shown as color gradient from red to black on the branches. Numbers on some key nodes are posterior probabilities
and maximum likelihood bootstrap values. GenBank accession numbers follow the terminal taxon labels.
Despite being phylogenetically closely related, A. sect. Ochopodium differs from Machaerium
by the mostly multi-seeded loment fruits and mostly herbaceous or shrubby habit (vs. mostly trees
or woody vines and fruits often a single-seeded samara with basal seed chamber and a terminal
wing, or a wingless nucoid legume in Machaerium). Despite the limited taxon sampling, patterns of
chromosomal and flower development variation as well as seedling morphology add further support
on the distinctiveness of A. sect. Ochopodium from the remaining Aeschynomene species (Sampaio et
al. 2013; Tapia-Pastrana et al. 2020; Rodrigues et al. 2019). Species of A. sect. Ochopodium have

Centrolobium microchaete
1.0/100 Pterocarpus monophyllus
Weberbauerella brongniartioides
0.64/46 Diphysa americana
Pictetia aculeata
1.0/99 Diphysa ormocarpoides
Zygocarpum yemenense
Ormocarpum kirkii
Ormocarpopsis calcicola
Peltiera nitida
Aeschynomene fluminensis
Aeschynomene parviflora
Aeschynomene americana
Aeschynomene unijuga
Aeschynomene villosa
Aeschynomene montevidensis
Aeschynomene paraguayensis
Aeschynomene tambacoundensis
1.0/100 Aeschynomene filosa
Aeschynomene rostrata
Aeschynomene deamii
Aeschynomene pratensis
Aeschynomene selloi
Aeschynomene sensitiva
Aeschynomene evenia Aeschynomene
Aeschynomene indica
Aeschynomene ciliata
Aeschynomene denticulata
Aeschynomene magna
Aeschynomene scabra
Aeschynomene pluriarticulata
Aeschynomene rudis
Aeschynomene hispida
Aeschynomene virginica
Aeschynomene patula
Aeschynomene crassicaulis
Aeschynomene pfundii
Aeschynomene elaphroxylon
Aeschynomene uniflora
Aeschynomene aspera
Aeschynomene afraspera
Aeschynomene fluitans
1.0/100 Aeschynomene nilotica
Aeschynomene deightonii
Aeschynomene mediocris
Aeschynomene cristata
Aeschynomene schimperi
Aeschynomene pulchella
Aeschynomene bella
Aeschynomene pygmaea
Aeschynomene abyssinica
Humularia kapiriensis
Aeschynomene heurckeana
Humularia descampsii
Soemmeringia semperflorens
prob Bryaspis humularioides
1 Bryaspis lupulina
Geissaspis cristata
Geissaspis tenella
Cyclocarpa stellaris
Smithia ciliata
Smithia abyssinica
Smithia sensitiva
Kotschya strigosa
Kotschya africana
Kotschya lutea
Kotschya ochreata
Dalbergia pachycarpa
Dalbergia ecastaphyllum
Dalbergia ovalis
Dalbergia inundata
Dalbergia elegans
Dalbergia cuiabensis
Dalbergia villosa
Dalbergia
Dalbergia miscolobium
1.0/100 Dalbergia spruceana
Dalbergia acuta
Dalbergia foliolosa
Dalbergia densiflora
0.5 Dalbergia glaucescens
Dalbergia arbutifolia
Dalbergia melanoxylon
Dalbergia retusa
Dalbergia boehmii
Dalbergia cochinchinensis
Dalbergia latifolia
Dalbergia oliveri
Dalbergia polyadelpha
0.02 Dalbergia nigrescens
Dalbergia assamica
Dalbergia lanceolaria
Dalbergia nigra
Dalbergia bracteolata
Dalbergia glabra
1 Dalbergia armata
Dalbergia congestiflora
Dalbergia decipularis
Dalbergia hortensis
Dalbergia riparia
Dalbergia brasiliensis
Dalbergia frutescens
Dalbergia sisso
1.0/100 Dalbergia rimosa
Dalbergia tonkinensis
Dalbergia maritinii
Dalbergia sissoo
Dalbergia humbertii
Dalbergia lemurica
Aeschynomene mollicula
Aeschynomene fascicularis
Aeschynomene nivea
Aeschynomene compacta
Aeschynomene purpusii
1.0/99 Aeschynomene martii
Ctenodon
Aeschynomene histrix
Aeschynomene scoparia
Aeschynomene brevipes
Aeschynomene paniculata
Aeschynomene chicocesariana
Aeschynomene riedeliana
Aeschynomene vogelii
Aeschynomene egena
Aeschynomene tumbezensis
Aeschynomene viscidula
Aeschynomene brevifolia
Aeschynomene elegans
Aeschynomene falcata
Aeschynomene brasiliana
Aeschynomene petraea
Aeschynomene simulans
1.0/99 Aeschynomene hintonii
Aeschynomene lyonnetii
Aeschynomene palmeri
Machaerium kegelii
Machaerium quinata
Machaerium pedicellatum
Machaerium ruddianum
Machaerium firmum
Machaerium fulvovenosum
Machaerium incorruptibile
Machaerium leucopterum
Machaerium scleroxylon
Machaerium nictitans
Machaerium jobimianum Machaerium
Machaerium obovatum
Machaerium floridum
Machaerium macaense
1.0/99 Machaerium nyctitans
Machaerium ovalifolium
Machaerium tortipes
Machaerium caratinganum
Machaerium condensatum
Machaerium lunatum
Machaerium amplum
Machaerium biovulatum
ITS
Machaerium aculeatum
Machaerium ferox
Machaerium isadelphum
matK
Machaerium riparium
Machaerium milleflorum
Machaerium salvadorense
trnL
Machaerium eriocarpum
Machaerium cobanense
Machaerium hirtum
Machaerium declinatum
Machaerium uncinatum
Machaerium pittieri
Machaerium seemannii
Machaerium gracile
Machaerium millei
Machaerium nigrum
Machaerium glabrum
Machaerium stipitatum
Machaerium lanceolatum
Machaerium ternatum
Machaerium brasiliense
Machaerium inundatum
Machaerium aureum
Machaerium oblongifolium
Machaerium floribundum
Machaerium punctatum
Machaerium salzmannii
Machaerium cantarellianum
Machaerium paraguariense
Machaerium villosum
Machaerium acutifolium
Machaerium opacum
Figure 3. A Bayesian majority-rule consensus phylogram of combined ITS/5.8S, matK, and trnL intron sequence data
(187 terminals by 2976 aligned sites), densely-sampled within Aeschynomene and the closely related species rich
genera Dalbergia and Machaerium. The non-monophyly of Aeschynomene (sensu Rudd 1955) is evinced, where A.
sect. Ochopodium, herein as the reinstated genus Ctenodon, is strongly supported as more closely related to
Machaerium. Posterior probabilities from 0.5 to 1.0 are shown as color gradient from red to black on the branches.
Numbers on specific key nodes are posterior probabilities and maximum likelihood bootstrap values. The
photographs by Domingos Cardoso illustrate the general floral architecture of the reinstated genus Ctenodon and its
closely related Dalbergia and Machaerium.

Figure 4. Geographical distribution of the reinstated genus Ctenodon in the American continent. Yellow points are
records for species that were sampled in our molecular phylogenies, whereas the gray points are from any species
that no accession were sampled in our phylogenies. The photograph by Domingos Cardoso shows the floral
morphology of Ctenodon sabulicolus, an endemic species of the Caatinga seasonally dry woodlands along the São
Francisco River sand dunes of Northeastern Brazil.
always been taxonomically differentiated from A. sect. Aeschynomene by possessing non-peltate,

basifixed stipules, campanulate calyx with five subequal lobes, and loments with articles distinctly
joined by an isthmus (Vogel 1838; Rudd 1955; Fernandes 1996). Also, they are terrestrial subshrubs
or shrubs, sometimes treelets or a small tree reaching 8 m tall, that are more ecologically confined
in fire-prone savannas and the seasonally dry tropical forests and woodlands, but also in oak forests,
pine-oak forests as well in pine woodlands.
Although elevation and most climate variables differed significantly between A. sect.
Ochopodium and Aeschynomene s.str. lineage, the actual differences were slight (Fig. 5). For example,
the average elevation for the Aeschynomene s.str. lineage was 669 m and that of the A. sect.
Ochopodium lineage 601 m, a difference of only 68 meters, which was significant given that the

95% CI of differences between the two clades ranged 30-107 m. The one notable climate difference
was annual precipitation (Fig. 5), which averaged 1543 mm for Aeschynomene s.str. lineage and
1059 mm for A. sect. Ochopodium lineage. The 95% CI of differences between the two clades for
annual precipitation ranged 442-527 mm. Notably, these differences span the 1200-1300 mm
threshold that distinguishes the lowland tropical seasonally dry biomes from wetter biomes (e.g.
Oliveira-Filho et al. 2013; Ringelberg et al. 2020). This finding reflects the observation that A. sect.
Ochopodium lineage is centered in seasonally dry neotropical forests and woodlands as well as
savanna, whereas Aeschynomene s.str. lineage is centered in wetter lowland tropical and subtropical
biomes. Inspection of the species distribution maps in Rudd (1955) reveals that species of A. sect.
Ochopodium have more limited or narrower geographical distributions than species of A. sect.
Aeschynomene. This pattern reflects the finding that dry lowland tropical biomes, such as seasonally
dry tropical forests and woodlands, are more dispersal limited than wet lowland tropical biomes, at
least with respect to woody plants (e.g. Pennington et al. 2009; Pennington & Lavin 2016).
Here we argue that this ecologically, morphologically, and phylogenetically coherent clade of
A. sect. Ochopodium species should to be recognized at generic rank. Therefore we decided to
reinstate the name Ctenodon Baill. to include all species of A. sect. Ochopodium (sensu Vogel 1838;
Rudd 1955). Ctenodon was originally described as a monotypic genus by Baillon (1870) based on the
Brazilian endemic, savanna-inhabiting species Ctenodon weddellianus Baill. [= Ctenodon oroboides
(Benth.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima], which was later transferred to Aeschynomene by
Rudd (1955). Although we have not sampled the type species of Ctenodon in our phylogenetic
analyses, it is most likely related to the A. sect. Ochopodium lineage, because it bears the same
morphological features traditionally used to define the section, and that are consistently found in
all species of this clade (Rudd 1955). Ctenodon oroboides is somewhat distinctive by just the ovate-
falcate leaflets. All other characters used once to classify it at genus level (e.g. pubescent fruits) are
found variously in A. sect. Ochopodium (Rudd 1955).
The infrageneric name Ochopodium published earlier by Vogel (1838) cannot be raised to
generic rank. Ctenodon is the legitimate generic name according to the Code Art. 11.2 (Turland et al.
2018) because there is no priority for a name outside the rank at which it was published. The generic
names Balisaea Taub. and Secula Small, originally published for species later transferred to A. sect.
Ochopodium (Rudd 1955, 1972), are here treated as synonyms of Ctenodon. All other generic names
(Herminiera Guill. & Perr., 1832; Macromiscus Turcz., 1846; Rueppelia A.Rich., 1847; Aedemone
Kotschy, 1858) ever associated under synonymy of Aeschynomene s.l. (Rudd 1955) and older than
Ctenodon were based on type species that are shown here phylogenetically nested within the
Aeschynomene s.str. lineage (Fig. 3). Noteworthy is the Old World species Aeschynomene
elaphroxylon (Guill. & Perr.) Taub. that is strongly resolved within Aeschynomene s.str. lineage (Fig. 3),
but has been suggested by Taubert (1894), when synonymizing the genus Herminiera, as
morphologically close to A. sect. Ochopodium.

Ctenodon as herein newly circumscribed comprises 66 species in America but the genus could
include as many as ca. 120 species. However, this will depend on a detailed taxonomic revision with
the pantropical and African species, as well as the complex of infraspecific variation in American
species. Because the taxonomic diversity of the African species of A. sect. Ochopodium is
underrepresented in our molecular phylogenies, the nomenclatural proposals include only the
American species. In the Americas, Ctenodon is particularly diverse in Mexico and Brazil, but there are
also a few endemic species in the Andes (Fig. 4; Rudd 1955; Delgado-Salinas & Sotuyo 2012; Flora
do Brasil 2020 under construction 2020). Species of the genus are poorly known with respect to
floral and pollination biology, but the recent study by Carleial et al. (2015) described a striking
mechanism of reflexed flowers in Ctenodon mucronulatus (= Aeschynomene amorphoides) that is rare
in papilionoid legumes, but previously observed only in some species of the phylogenetically related
genus Machaerium. The unique architecture of the papilionate flowers in C. mucronulatus involves
the wing petals and keel highly curved against the standard petal that lies in obliquely horizontal
position. The entire flower then serves as landing platform for pollinators and, more unusually, the
distinctly-colored keel tip functions as color marking (maculae), which is otherwise a predominant
feature of the standard petal in papilionoid legumes (Carleial et al. 2015).
30
3000
25
Annual Mean Temperature (C)
Elevation (m)
2000
20
15
1000
10
0
ochopodium$clade ochopodium$clade
35
6000
30
Temperature Annual Range
Annual Precipitation (mm)
25
4000
20
2000
15
10
0
Aeschynomene clade Ctenodon Aeschynomene clade Ctenodon
Figure 5. Elevation and climate differences between species of the Aeschynomene clade (Aeschynomene sensu stricto
and the mostly African phylogenetically related genera) and the clade comprising Ctenodon species. For elevation,
means for the two clades, respectively, are 669 versus 601 m (95% CI for the differences in these means range 30 –
107 m). For annual mean temperature, means for the two clades, respectively, are 22.8 versus 23.2 C (95% CI for the
differences in these means range -0.5–-0.11 C). For annual mean precipitation, means for the two clades,
respectively, are 1543 versus 1059 mm (95% CI for the differences in these means range 442–527 mm). For annual
mean temperature range (a measure of seasonality), means for the two clades, respectively, are 17.2 versus 17.8 C
(95% CI for the differences in these means range -0.88–-0.30 mm).

4 | TAXONOMY
Ctenodon Baill., Adansonia 9: 236. 1870. Type: Ctenodon weddellianus Baill., as “weddellianum” [=
Ctenodon oroboides (Benth.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima].
= Balisaea Taub., Bot. Jahrb. Syst. 21: 436. 1895. Type: Balisaea genistoides Taub. [≡ Ctenodon
genistoides (Taub.) D.B.O.S.Cardoso, Filardi & H.C.Lima].
= Secula Small, Fl. Miami [Small] 90: 200. 1913. Type: Secula viscidula (Michx.) Small [≡ Ctenodon
viscidulus (Michx.) D.B.O.S.Cardoso & A.Delgado].
= Aeschynomene sect. Ochopodium Vogel, Linnaea 12: 86. 1838 ≡ Aeschynomene subgen.
Ochopodium (Vogel) J.Léonard, Bull. Jard. Bot. État. Brux. 24: 84. 1954. Lectotype (designated by
Rudd 1955): Aeschynomene falcata (Poir.) DC. [≡ Ctenodon falcatus (Poir.) D.B.O.S.Cardoso,
P.L.R.Moraes & H.C.Lima].
Herbs, pyrophytic or fire-sensitive subshrubs, shrubs, treelets or a small tree reaching up to 8 m tall;
erect or prostrate; the stems covered by glandular hairs or simple pubescence, or the plants
sometimes glabrous. Stipules striate, subcordate or triangular to lanceolate, non-peltate, basifixed.
Leaves pinnately compound, 5–80-foliolate or more unifoliolate, or scale-like and then giving plant
an aphyllous appearance; petiole often including a swollen pulvinus at the base; leaflets
membranaceous to coriaceous, symmetrical or oblique, oblong to orbiculate, elliptic to ovate or
obovate, or more rarely needle-like, glabrous to densely pubescent, with or without marginal
glandular hairs. Inflorescence axillary or sometimes terminal, 2–3-flowered racemes to densely-
flowered compound racemes or panicle-like inflorescences; pedicel bracteolate. Flowers bilaterally
symmetrical, papilionate; calyx campanulate, lobes 5; petals bright to light yellow, red, pale pink or
red purplish, clawed, the standard often pubescent externally, distinctly clawed; wing and keel
glabrous; stamens 10, the filaments half the length fused into a sheath, monadelphous, sometimes
diadelphous, glabrous, the anthers uniform, dorsifixed; gynoecium with the ovary stipitate, laterally
compressed. Fruit a (1–)2–18-articulate loment, distinctly stipitate, the articles distinctly joined by
an isthmus, glabrous or pubescent; seeds reniform, light brown to black, smooth.
Ctenodon acapulcensis (Rose) A.Delgado, comb. nov.

Basionym: Aeschynomene acapulcensis Rose, Contr. U.S. Natl. Herb. 5: 191. 1899. Type:
MEXICO, Guerrero, Acapulco, Oct 1894-Mar 1895, E. Palmer 126 (holotype: US00001870;
isotypes: BM000931568, V0058790F, V0058791F, V0058792F, V0058804F, GH00058419,
GH00058420, K000297149, MICH1107075, MO-126284, NY00004974, PH00000045,
UC80616, US00930889, US00930890).
= Aeschynomene picachensis Brandegee, Univ. Calif. Publ. Bot. 6: 181. 1915. Type: MEXICO, Oaxaca,
Cerro de Pichaco, Jul 1914, C.A. Purpus 7162 (holotype: UC175043; isotypes: BM000931569,
V0058799F, GH00058437, MO-128347, NY00004987, US00001919).
Distribution—Mexico.

Ctenodon afranianus D.B.O.S.Cardoso, Filardi & H.C.Lima, nom. nov.

Basionym: Aeschynomene venulosa Afr.Fern., O Táxon Aeschynomene no Brasil: 111. 1996,
nom. illeg., hom. post. (non Verdc. 1970). Type: BRAZIL, Alagoas, Maceió, Faz. Pau-Brasil, Jul
1977, A. Fernandes & Matos s.n. (holotype: EAC0003363).
Distribution—Brazil.
Ctenodon benthamii (Rudd) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Basionym: Aeschynomene mollicula var. benthamii Rudd, Contr. U.S. Natl. Herb. 32: 131. 1955
≡ Aeschynomene platycarpa Benth., Fl. Bras. 15(1): 63. 1859, nom. illeg., hom. post. (non Michx.
1803). Type: BRAZIL, Minas Gerais, “in saxosis umbrosis prope fluv. Jiquitinonha
[Jequitinhonha]”, Dec 1824, L. Riedel 591 (lectotype designated by Rudd 1955: K000222449;
isolectotype: LE00002482).
≡ Aeschynomene benthamii (Rudd) Afr.Fern., O Táxon Aeschynomene no Brasil: 113. 1996,
nom. inval.
Ctenodon bradei (Rudd) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Basionym: Aeschynomene bradei Rudd, J. Wash. Acad. Sci. 49: 51. 1959. Type: BRAZIL, Rio de
Janeiro, Santa Maria Madalena, Pedra Dubois, Feb 1935, J. Santos Lima & A.C. Brade 14220
(holotype: RB00540115, RB00168508; isotype: US00001864).
Ctenodon brasilianus (Poir.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.

Basionym: Hedysarum brasilianum Poir., Encycl. [J. Lamarck & al.] 6(2): 448. 1805. Type:
BRAZIL, Rio de Janeiro, circa Rio de Janeiro, Jul 1767, P. Commerson s.n. (holotype: P-JU [Catal.
No. 15624] – P00678649; isotypes: V0059317F, P01167838, P02913671, P02913680 [F neg.
36628]).
≡ Aeschynomene brasiliana (Poir.) DC., Prodr. [A. P. de Candolle] 2: 322. 1825.
= Cassia biflora Mill., Gard. Dict., ed. 8. n. 14. 1768, nom. illeg., hom. post. (non L. 1753) ≡ Cassia
houstoniana Collad., Hist. Nat. Méd. Casses 132. 1816 ≡ Aeschynomene biflora Fawc. & Rendle, Fl.
Jamaica 4: 27. 1920. Type: JAMAICA, 1730, W. Houstoun s.n. (holotype: BM000931561).
= Aeschynomene paucijuga DC., Prodr. [A. P. de Candolle] 2: 321. 1825. Type: FRENCH GUIANA,
Cayenne, 1819, G.S. Perrottet s.n. (holotype: G-DC [F neg. 6959]; possible isotype: P02913696).
= Aeschynomene paucijuga var. subscabra DC., Prodr. [A. P. de Candolle] 2: 321. 1825. Type: “In
America merid.”, Anonymous s.n. (holotype: G-DC n.v.).
= Hedysarum hirtum Vell., Fl. Flumin.: 319. 1829; idem in Fl. Flumin. Icon. 7: t. 151. 1831, nom. illeg.,
hom. post. (non L. 1753). Type: BRAZIL, “Habitat campis paulo supra relatis”, Oct. Lectotype
(designated here): [icon] original plate on parchment of “Florae Fluminensis” (digital object:
mss1198656_155) in the Manuscript Section of Fundação Biblioteca Nacional, Rio de Janeiro

(Coleção Flora Fluminense; fonte: Conceição Vellozo, 1790, Florae Fluminensis Icones
fundamentales. t. VII. p. [prancha] 151; I-17,04,001 – Manuscritos).
= Aeschynomene trisperma Klotzsch, Reis. Br.-Guiana [Ri. Schomburgk] 3: 1103. 1849. Type:
GUYANA, “auf trockner Savanne in der Umgebung des Rué-imeru”, Sep-Oct, Ri. Schomburgk s.n.
(unlocated).
= Aeschynomene guaricana Pittier, Bol. Tecn. Minist. Agric. 5: 41. 1944. Type: VENEZUELA, Aragua,
entre San Juan de los Morros y Uberito, Dec 1923, H. Pittier 11324 (holotype: VEN3782).
Distribution—Widespread in tropical America.
Ctenodon brevipes (Benth.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.

Basionym: Aeschynomene brevipes Benth., Fl. Bras. (Martius) 15(1): 66. 1859. Type: BRAZIL,
Piauí, rocky places near Oeiras, May 1839, G. Gardner 2097 (lectotype designated here:
K000092347; isolectotypes: BM000931738, GH n.v., L0018459). Other syntype: BRAZIL, Piauí,
in campis agrestibus, May 1819, C.F.P. von Martius s.n. (M0233719 [F neg. 6269]).
= Aeschynomene brevipes var. uliginosa Benth., Fl. Bras. (Martius) 15(1): 66. 1859. Type: BRAZIL,
Maranhão, Jun 1841, G. Gardner 5997 (holotype: K000222439; isotypes: BM000931740,
K000222438).
Ctenodon carvalhoi (G.P.Lewis) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Basionym: Aeschynomene carvalhoi G.P.Lewis, Kew Bull. 40(3): 599. 1985. Type: BRAZIL, Bahia,
Palmeiras, Morro do Pai Inácio, Km 224 da Rodovia BR 242, alt. 1000 m, Dec 1981, G.P. Lewis
et al. 962 (holotype: CEPEC00030212; isotypes: HUEFS [99842], K000092398, M0233720,
NY00004997, RB00540116, SPF00040115, US00091042).
Ctenodon carichanicus (Rudd) D.B.O.S.Cardoso & H.C.Lima, comb. nov.

Basionym: Aeschynomene brasiliana var. carichanica Rudd, J. Wash. Acad. Sci. 49: 48. 1959.
Type: VENEZUELA, Bolívar, north end of Cerro Gavilan (Cerro Carichana), 100-300 m, Dec
1955, J.J. Wurdack & J.V. Monachino 40885 (holotype: US00001886; isotypes: F0058807F,
NY00004995, VEN40998).
≡ Aeschynomene carichanica (Rudd) G.B.Rodr. & G.Agostini, Acta Bot. Venez. 16(1): 123. 1991.
Distribution—Venezuela.
Ctenodon chicocesarianus (D.B.O.S.Cardoso & G.Ramos) D.B.O.S.Cardoso, comb. nov.

Basionym: Aeschynomene chicocesariana D.B.O.S.Cardoso & G.Ramos, Syst. Bot. 44(4): 811.
2019. Type: BRAZIL, Bahia, Ibicoara, Gerais do Machombongo, 13 o13’58”S, 41o17’16”W, 1182
m, Jan 2015, D. Cardoso et al. 3640 (holotype: HUEFS0239018; isotypes: ALCB, CEPEC, K, MO,
NY, RB).

Ctenodon compactus (Rose) A.Delgado, comb. nov.

Basionym: Aeschynomene compacta Rose, Contr. U.S. Natl. Herb. 5(4): 191. 1899. Type:
MEXICO, Oaxaca, Tomellin Canyon, Oct 1894, C.G. Pringle 5645 (lectotype designated by Rudd
1955: US00001904; isolectotypes: GH00058427, MEXU01169220, UC80604, UVMVT024909).
Other syntypes: MEXICO, Oaxaca, from Juchitan to Chivela, Jun 1895, E.W. Nelson 2630
(US01108278). MEXICO, [Oaxaca], Cuesta de Ejulta, Naialtepec [Manialtepec], Jun 1895, L.C.
Smith 452 (GH n.v., US02044797).
= Aeschynomene oaxacana Brandegee, Univ. Calif. Publ. Bot. 6: 181. 1915. Type: MEXICO, Oaxaca,
Cerro de Picacho, Jul 1914, C.A. Purpus 7179 (holotype: UC175011; isotypes: BM000931570,
V0058798F, GH00058433, MO-126286, NY00004984, US00001916).
Distribution—Costa Rica, Guatemala, and Mexico.
Ctenodon egenus (J.F.Macbr.) D.B.O.S.Cardoso & H.C.Lima, comb. nov.

Basionym: Aeschynomene mollicula var. egena J.F.Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser.
13: 443. 1943. Type: PERU, Cajamarca, Prov. Jaen, mountains between the rivers Tabaconas
and Marañón, 1000-1200 m, Apr 1912, A. Weberbauer 6176 (holotype: F0043450F; isotypes:
F0043443F, GH00058461, NY00005007, US00001888).
≡ Aeschynomene egena (J.F.Macbr.) Rudd, Contr. U.S. Natl. Herb. 32: 122. 1955.
Distribution—Peru.
Ctenodon elegans (Schltdl. & Cham.) D.B.O.S.Cardoso & A.Delgado, comb. nov.
Basionym: Aeschynomene elegans Schltdl. & Cham., Linnaea 5(4): 583. 1830. Type: MEXICO,
Veracruz, Hacienda de la Laguna, Sep 1828, C.J.W. Schiede & F. Deppe s.n. [231] (lectotype
designated here: HAL0098429; isolectotype: G [F neg. 27926]). Other syntype: MEXICO,
Veracruz, “in collibus apricis prope Jalapam”, Aug 1828, C.J.W. Schiede & F. Deppe s.n. [231]
(HAL0098428). Possible original material: Hacienda de la Laguna, Aug 1829, C.J.W. Schiede &
F. Deppe s.n. (BM000931567).
≡ Aeschynomene falcata (Poir.) DC. var. elegans (Schltdl. & Cham.) Kuntze, Revis. Gen. Pl. 1:
158. 1891.
Ctenodon elegans var. elegans

= Aeschynomene tecta Vogel, Linnaea 12: 87. 1838 ≡ Aeschynomene falcata var. plurijuga Benth., Fl.
Bras. (Martius) 15(1): 68. 1859, p.p. Type: BRAZIL, “inter Campos-Victoria”, F. Sellow s.n. (holotype:
B†; probable isotype: K000222428).
= Aeschynomene falcata (Poir.) DC. var. elegans (Schltdl. & Cham.) Kuntze f. glabrior Kuntze, Revis.
Gen. Pl. 1: 158. 1891. Type: VENEZUELA, Caracas, 1874, C.E.O. Kuntze s.n. (holotype:
NY00005001).

= Aeschynomene arenicola Brandegee, Univ. Calif. Publ. Bot. 10: 408. 1924. Type: MEXICO, Chiapas,
on sandy soil in pine forests near Hacienda Monserrate, Sep 1923, C.A. Purpus 9148 (holotype:
UC220377; isotypes: GH n.v., MO-126288, NY00004977, US00001867).
Ctenodon elegans var. robustior (Rudd) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.
Basionym: Aeschynomene elegans var. robustior Rudd, J. Wash. Acad. Sci. 49: 50. 1959. Type:
BRAZIL, Goiás, “Brejon prope Santa Cruz”, Jun 1820, J.E. Pohl s.n. [2560] (holotype:
K000222429; isotypes: M0233731, W0067211).
Ctenodon falcatus (Poir.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.

Basionym: Hedysarum falcatum Poir., Encycl. [J. Lamarck & al.] 6(2): 448. 1805. Type: BRAZIL,
Rio de Janeiro, “le long de la rivière de Rio-Janeiro”, P. Commerson s.n. (holotype: P-JU [Catal.
No. 15623] – P00678648; isotypes: F0059318F, P02297636, P02297637 [F neg. 36629],
P00296593, P02913700).
≡ Aeschynomene falcata (Poir.) DC., Prodr. [A. P. de Candolle] 2: 322. 1825.
Distribution—Argentina, Bolivia, Brazil, Chile, Colombia, and Paraguay.
Ctenodon falcatus var. falcatus

= Hedysarum diffusum Vell., Fl. Flumin.: 320. 1829; idem in Fl. Flumin. Icon. 7: t. 153. 1831, nom.
illeg., hom. post. (non Willd. 1802). Type: BRAZIL, “Habitat campis maritimis. Offendi ad urbis
collem, quo inædificatum est Divi Antonii Cænobium”, Nov. Lectotype (designated here): [icon]
original plate on parchment of “Florae Fluminensis” (digital object: mss1198656_157) in the
Manuscript Section of Fundação Biblioteca Nacional, Rio de Janeiro (Coleção Flora Fluminense;
fonte: Conceição Vellozo, 1790, Florae Fluminensis Icones fundamentales. t. VII. p. [prancha] 153;
I-17,04,001 – Manuscritos).
= Aeschynomene falcata var. paucijuga Benth., Fl. Bras. (Martius) 15(1): 67, tab. 14. 1859, includes
type of species.
= Aeschynomene apoloana Rusby, Bull. New York Bot. Gard. 6: 511. 1910. Type: BOLIVIA, Apolo,
grassy prairie, Mar 1902, R.S. Williams 25 (holotype: NY00004994; isotypes: BM000931553,
K000222453, US00001868).
Ctenodon falcatus var. hassleri (Rudd) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.
Basionym: Aeschynomene falcata var. hassleri Rudd, J. Wash. Acad. Sci. 49: 52. 1959. Type:
PARAGUAY, near Concepción, Sep 1901, E. Hassler 7461 (holotype: K000222433; isotypes:
BM000538162, G n.v., NY00005000, P02297638, P02913580, S13-12015).
Ctenodon falcatus var. minor (N.F.Mattos) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.
Basionym: Aeschynomene falcata var. minor N.F.Mattos, Loefgrenia 53: 2. 1971. Type: BRAZIL,
Mato Grosso do Sul, 3 léguas ao sul de Campo Grande, Fazenda Ibirussu, Jan 1933, J. Otero
173 (holotype: SP000945).

Ctenodon fascicularis (Schltdl. & Cham.) A.Delgado, comb. nov.

Basionym: Aeschynomene fascicularis Schltdl. & Cham., Linnaea 5: 584. 1830. Type: MEXICO,
Veracruz, “inter la Laguna Verde et Actopan”, Mar 1829, C.J.W. Schiede & F. Deppe s.n.
(holotype: B†; lectotype designated here: HAL0098430).
= Adesmia mimosoides G.Don, Gen. Hist. 2: 282. 1832. Type: MEXICO, ex Herb. Lambert (not found).
= Aeschynomene fruticosa Sessé & Moc., Pl. Nov. Hisp.: 122. 1890. Type: MEXICO, Michoacán,
Apatzingam [Apatzingán], Sep, M. de Sessé & J. M. Mociño s.n. (Possible original material: [1940] F,
MA601244, MA601245; [1941] F, MA601242; [1942] F, MA601243).
= Aeschynomene oligantha Micheli, Mém. Soc. Phys. Genève 34: 256. 1903, nom. illeg., hom. post.
(non Welw. ex Baker 1871). Type: MEXICO, Guerrero, La Mata San Dios, 200 m, Oct 1898, É.
Langlassé 431 (holotype: G; isotypes: GH00058434, K000297148, MEXU00051016, P02914003,
P02914004, US00001915).
Distribution—Colombia, Costa Rica, El Salvador, Guatemala, Honduras, Mexico, Nicaragua, and
Venezuela.
Ctenodon foliolosus (Rudd) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Basionym: Aeschynomene foliolosa Rudd, Contr. U.S. Natl. Herb. 32: 91. 1955. Type:
COLOMBIA, Guaviare [Vaupés], San José del Guaviare, 270 m, Nov 1939, J. Cuatrecasas 7697
(holotype: US00001897; isotypes: COL000001730, V0058810F).
Distribution—Bolivia, Brazil, and Colombia.
Ctenodon genistoides (Taub.) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Basionym: Balisaea genistoides Taub., Bot. Jahrb. Syst. 21: 437. 1896. Type: BRAZIL, Goiás, “in
campis montium Serra de Balisa dictorum”, Sep 1892, E.H.G. Ule 2837 [14] (holotype: B†;
isotypes: HBG506552, P03383092, P03383093, P03383094, R000065277).
≡ Aeschynomene genistoides (Taub.) Rudd, Phytologia 23: 321. 1972.
Ctenodon genistoides var. genistoides

= Aeschynomene irwinii Rudd, Phytologia 15: 117, fig. 1. 1967. Type: BRAZIL, Goiás, Chapada dos
Veadeiros, ca. 13 km N.W. of Veadeiros, 1200 m, Oct 1965, H.S. Irwin et al. 9367 (holotype:
US00001908; isotype: NY00005002).
Ctenodon genistoides var. latifoliolus (G.P.Lewis) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.
Basionym: Aeschynomene genistoides var. latifoliola G.P.Lewis, Kew Bull. 49(1): 95. 1994. Type:
BRAZIL, Goiás, Alto Paraíso, Córrego Veredas, Nov 1991, G. Hatschbach et al. 55944 (holotype:
MBM146604; isotypes: C10011415, CEPEC [62941], G00364569, HBG520675, INPA [201071],
K000222450).

Ctenodon gracilis (Vogel) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.

Basionym: Aeschynomene gracilis Vogel, Linnaea 12: 89. 1838. Type: BRAZIL, loco haud
indicato, “inter Campos-Victoria”, s.d., F. Sellow s.n. (lectotype designated here: K000189448;
probable isolectotype: K000189535).
= Aeschynomene portoricensis Urb., Symb. Antill. (Urban) 1(2): 325. 1899. Type: PUERTO RICO,
Arecibo, Laguna Tortuguero, near Manatí, Apr 1887, P.E.E. Sintenis 6657 (lectotype designated
here: US00001921; isolectotypes: BM000931560, V0058805F, GH, HBG520670, K000222411,
L0018471, M0233746, MO-022345, NY00004992, P02912509). Other syntype: PUERTO RICO,
Maricao, Aguadilla, Sierra de Naguabo ad Rio Blanco, Nov 1886, P.E.E. Sintenis 4889 (US).
Distribution—Brazil, Colombia, and Puerto Rico.
Ctenodon graminoides (G.P.Lewis) D.B.O.S.Cardoso & H.C.Lima, comb. nov.

Basionym: Aeschynomene graminoides G.P.Lewis, Kew Bull. 47(1): 143. 1992. Type: BRAZIL,
Goiás, Santo Antônio do Descoberto, divisa do Distrito Federal, Jan 1976, E.P. Heringer 15338
(holotype: IBGE; isotypes: RB00306230, RFA [17839], UB, UEC051896).
Ctenodon hintonii (Sandwith) A.Delgado, comb. nov.

Basionym: Aeschynomene hintonii Sandwith, Hooker's Icon. Pl. 35(2): tab. 3448. 1943. Type:
MEXICO, Guerrero, Coyuca, Placeres [Placeres de Oro], Mar 1934, G.B. Hinton et al. 6104
(lectotype designated here: K000118279; isolectotypes: K000118280, MICH1107078,
NY00004983, RSA0002726, UC1098467, US00001895).
Ctenodon histrix (Poir.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.

Basionym: Aeschynomene histrix Poir., Encyc. [J. Lamarck & al.], Suppl. 4: 77. 1816. Type:
FRENCH GUIANA, “île de Cayenne”, s.d., J. Martin s.n. (lectotype designated here: P02912637;
isolectotype: P02912675 ex Herb. Poiret in Herb. Moquin-Tandon).
≡ Secula histrix (Poir.) Small, Man. S.E. Fl.: 728. 1933.
Ctenodon histrix var. histrix

= Aeschynomene cassioides Desv. ex Ham., Prodr. Pl. Ind. Occid.: 51. 1825. Type: FRENCH GUIANA,
Cayenne, ex Herb. Desvaux (holotype: P02912656).
= Aeschynomene echinus Vogel, Linnaea 12: 92. 1838. Type: BRAZIL, loco haud indicato, s.d., F. Sellow
s.n. (holotype: B†).
= Aeschynomene conferta Benth., Ann. Nat. Hist. 3: 433. 1839. Type: GUYANA, 1837, R.H.
Schomburgk 187 (lectotype designated here: K000222418; isolectotypes: BM000931556,

BR0000005119516, F0058808F, GH00058449, L1970334, MICH1107076, P02913646,

P02913648, TCD0004354, US00001903).
= Aeschynomene mucronulata Benth., J. Bot. (Hooker) 2: 56. 1840 ≡ Aeschynomene histrix var.
mucronulata Benth., Fl. Bras. (Martius) 15(1): 69. 1859. Type: GUYANA, dry savannahs, 1839, R.H.
Schomburgk 822 (lectotype designated here: K000222419; isolectotypes: BM000931555, G [F
neg. 27929]).
= Aeschynomene pineticola Standl. & L.O.Williams, Ceiba 1: 79. 1950. Type: HONDURAS: Morazán,
rocky pine forest, along road from El Zamorano toward San Antonio de Oriente, 825-950 m, Sep-
Nov 1948, P.C. Standley 12657 (holotype: F; isotype: EAP15549).
Ctenodon histrix var. apanus (Rudd) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.
Basionym: Aeschynomene histrix var. apana Rudd, J. Wash. Acad. Sci. 49: 50. 1959. Type:
PARAGUAY, Centurión, “Zwischen Rio Apa und Rio Aquidaban, Dec 1908, K. Fiebrig 4387
(holotype: US00001894; isotypes: BM000538153, E00296480, E00296481, GH,
K000222425).
Ctenodon histrix var. densiflorus (Benth.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.
Basionym: Aeschynomene densiflora Benth. J. Bot. (Hooker) 2: 56. 1840. Type: GUYANA, R.H.
Schomburgk 846 (lectotype designated here: K000222423; isolectotypes: BM000931554,
F0360740F, G00364570, L0018467, P02912697, P02913643, P02913644, TCD0004351,
US00001901).
≡ Aeschynomene histrix var. densiflora (Benth.) Rudd, Contr. U.S. Natl. Herb. 32: 84. 1955.
Ctenodon histrix var. incanus (Benth.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.
Basionym: Basionym: Aeschynomene histrix var. incana Benth., Fl. Bras. (Martius) 15(1): 69.
1859. Type: URUGUAY, Montevideo, F. Sellow s.n. (holotype: B†; lectotype designated here:
K000222421; isolectotypes: V0058811F, K000222420, L1970338, M0233737, P02912696
[Herbier Impérial du Brésil no. 1486], W0067216).
≡ Aeschynomene incana Vogel, Linnaea 12: 90. 1838, nom. illeg., hom. post. (non G.Mey. 1818).
= Aeschynomene puberula DC., Prodr. [A. P. de Candolle] 2: 321. 1825. Type: “America meridionale”,
s.d., Anonymous s.n. (holotype: G-DC [F neg. 6960]).
Ctenodon histrix var. multijugus (Chodat & Hassl.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.
Basionym: Aeschynomene brasiliana f. multijuga Chodat & Hassl., Bull. Herb. Boissier (ser. 2) 4:
882. 1904. Type: PARAGUAY, Departamento Canindeyú, “in campo pr. flumen Carimbatay”, Dec
1898-1899, E. Hassler 5814 (lectotype designated by Vanni 2016: NY00004996; isolectotypes:
BM000538155, G00381609, G00381610, G00381611, G00381612, GH00058453,
K000222426, MO-022336, P02913685, P02913994, S13-12024).
≡ Aeschynomene histrix var. multijuga (Chodat & Hassl.) Rudd, J. Wash. Acad. Sci. 49: 49. 1959.

Ctenodon interruptus (Benth.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.

Basionym: Aeschynomene interrupta Benth., J. Bot. (Hooker) 2(9): 56. 1840. Type: GUYANA, Rio
Branco, 1840, R.H. Schomburgk 803 (lectotype designated here: K000222445; isotypes: B † [F
neg 2147], BM000931551, F, G, GH00058454, IAN [230], K000222446, L1970772, MO-
1282367, P02913986, TCD0004350, US00001893).
Distribution—Brazil and Guyana.
Ctenodon lacabuendianus (Brandão) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Basionym: Aeschynomene laca-buendiana Brandão, Daphne 1(3): 5. 1991. Type: BRAZIL, Minas
Gerais, Mocambinho, km 18 após o trevo de Janaúba-Mocambinho, J.P. Laca-Buendia 1500
(holotype: RB; isotype: PAMG).
Ctenodon langlassei (Micheli ex Rudd) A.Delgado, comb. nov.

Basionym: Aeschynomene langlassei Micheli ex Rudd, Contr. U.S. Natl. Herb. 32: 117. 1955.
Type: MEXICO, Guerrero, Sierra Madre, 1200 m, Feb 1899, E. Langlassé 847 (holotype:
US00001892; isotypes: G, GH00058429, K000297140, MPU022831, P02913969,
P02913982).
Ctenodon leptostachyus (Benth.) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Basionym: Aeschynomene leptostachya Benth., Fl. Bras. (Martius) 15(1): 65 (1859). Type:
BRAZIL, Goiás, in paludosis, prope Salinas, 1844, H.A. Weddell 2113 (holotype: K000222437
[Photo: P02913991]; isotype: P02914026).
Ctenodon lewisianus (Afr.Fern.) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Basionym: Aeschynomene lewisiana Afr.Fern., O Táxon Aeschynomene no Brasil: 78. 1996. Type:
BRAZIL, Bahia, Lençóis, Rio Mucugezinho, próximo a BR-242, próximo ao Morro do Pai Inácio,
ca. 1000 m, Dec 1984, G.P. Lewis et al. s.n. (holotype: EAC0016790).
Ctenodon lorentzianus (Bacigalupo & Vanni) D.B.O.S.Cardoso & H.C.Lima, comb. nov.
Basionym: Aeschynomene lorentziana Bacigalupo & Vanni, Hickenia 2(6): 33, f. 1. 1984. Type:
ARGENTINA, Entre Rios, Depto. Federación, Cantera de Chaviyú, Nov 1976, N.L. Troncoso et al.
1363 (holotype: SI001487; isotype: CTES0000458).
Distribution—Argentina.

Ctenodon lyonnetii (Rudd) A.Delgado, comb. nov.

Basionym: Aeschynomene lyonnetii Rudd, Acta Bot. Mex. 8: 31. 1989. Type: MEXICO, Guerrero,
Iguala, Sep 1937, E. Lyonnet 1717 (holotype: MEXU00278344; isotypes: CAS0000118,
MEXU00278345, US00288959).
Ctenodon marginatus (Benth.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.

Basionym: Aeschynomene marginata Benth., Fl. Bras. (Martius) 15(1): 66. 1859. Type: BRAZIL,
Piauí, in sandy woods near Oeiras, Apr 1839, G. Gardner 2098 (lectotype designated here:
K000092312; isolectotypes: B† [F neg. 2149], BM000931741, F0058812F, GH00058458,
GH00058459, L0018474, NY00005005, NY00005006, P02913972, P02913975, S-R-9501,
W0067220). Other syntype: BRAZIL, Ceará, sandy hills Aracaty, Jul 1838, G. Gardner 1543
(K000092313).
Distribution—Bolivia and Brazil.
Ctenodon marginatus var. marginatus
Ctenodon marginatus var. grandiflorus (Benth.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.
Basionym: Aeschynomene marginata var. grandiflora Benth., Fl. Bras. (Martius) 15(1): 67. 1859.
Type: BRAZIL, Piauí, dry rocky hills near the city of Oeiras, May 1839, G. Gardner 2099
(lectotype designated here: K000092315; isolectotypes: BM000931739, GH00058456,
GH00058457, L0018475, NY00005004, NY00005022, P02913971, P02913973, P02913976,
S-R-9500, W0067221, W18890044776).
Ctenodon martii (Benth.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.

Basionym: Aeschynomene martii Benth., Fl. Bras. (Martius) 15(1): 62, t. 13. 1859. Type: BRAZIL,
Bahia, “in sylvis Catingas ad fluv. Salitre prope Joazeiro”, Apr 1819, C.F.P. von Martius Observ.
no. 2322 (lectotype designated here: M0233740). Other syntype: BRAZIL, Minas Gerais, “in
campis et sepibus Serro Frio”, Jul 1818, C.F.P. von Martius s.n. (M0233741). Other original
material: BRAZIL, Bahia et Minas Gerais, s.d., C.F.P. von Martius s.n. (M0233742); idem, s.d.,
C.F.P. von Martius s.n. (M0233739).
= Aeschynomene arbuscula Rizzini, Leandra 4-5: 15, t. 13. 1974, nom. illeg., hom. post. (non Baker f.
1932) ≡ Aeschynomene rizzinii Schoenberg & Ferreira, Bol. Mus. Bot. Munic. 32: 1. 1977. Type:
BRAZIL, Bahia, Campo Formoso, Fazenda Lagoa do Porco, Apr 1973, D.P. Lima 13109 (holotype:
RB00540113; isotypes: HUCPE [219], HUEFS [93748], IPA [75500], PEUFR [14597]).

Ctenodon matosii (Afr.Fern.) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Basionym: Aeschynomene matosii Afr.Fern., O Táxon Aeschynomene no Brasil: 97. 1996. Type:
BRAZIL, Piauí, Sete Cidades, Piracuruca, Jul 1991, A. Fernandes & Matos s.n. (holotype:
EAC0017998 – 2 sheets).
Ctenodon molliculus (Kunth) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Basionym: Aeschynomene mollicula Kunth, Nov. Gen. Sp. [quarto ed.] 6: 532. 1823. Type: PERU,
“inter Paramo de Yamoca et villam Colasey... (Provincia Jaen de Bracamoros) ”, Aug, A.J.A.
Bonpland & F.W.H.A. von Humboldt s.n. [3586] (holotype: P00679170; isotypes: B-W 13729-01
0 [F neg. 2241], F0058813F, P02913965).
Distribution—Brazil, Colombia, Ecuador, and Peru.
Ctenodon molliculus var. molliculus

= Aeschynomene pauciflora Vogel, Linnaea 12: 93. 1838. Type: BRAZIL, Minas Gerais, Santo Antonio
do Monte, Nov 1818, F. Sellow s.n. (holotype: B†).
Ctenodon molliculus var. breviflorus (Rudd) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.
Basionym: Aeschynomene mollicula var. breviflora Rudd, Contr. U.S. Natl. Herb. 32: 130. 1955.
Type: COLOMBIA, Norte de Santander, between Chinácota and La Esmeralda, Mar 1927, E.P.
Killip & A.C. Smith 20921 (holotype: US00001889; isotypes: GH00058460, NY00005021).
Ctenodon monteiroi (Afr.Fern. & P.Bezerra) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.
Basionym: Aeschynomene monteiroi Afr.Fern. & P.Bezerra, Bradea 2(50): 332. 1979. Type:
BRAZIL, Ceará, Independência, encosta da Serra, Apr 1978, A. Fernandes & Matos s.n.
(holotype: EAC0003772 – 2 sheets).
= Aeschynomene monteiroi var. psilantha G.P.Lewis, Kew Bull. 40(3): 603. 1985. Type: BRAZIL, Piauí,
São Raimundo Nonato, Fundação Ruralista (Sede), 320 m, Jan 1982, G.P. Lewis & H.P.N. Pearson
1135 (holotype: CEPEC00029563; isotypes: K000092359, RB00540117).
Ctenodon mucronulatus (Benth. ex Hemsl.) A.Delgado, comb. nov.

Basionym: Drepanocarpus mucronulatus Benth. ex Hemsl., Diagn. Pl. Nov. Mexic. 1: 8. 1878.
Type: MEXICO, Jalisco, Bolaños, Guadalaxara [Guadalajara], s.d., T. Coulter s.n. (holotype:
K000297144; isotype: TCD).
= Brya amorphoides S.Watson, Proc. Amer. Acad. Arts 22: 406. 1887 ≡ Aeschynomene amorphoides
(S.Watson) Rose ex B.L.Rob., Proc. Amer. Acad. Arts 29: 315. 1894. Type: MEXICO, Jalisco, Tequila,
in deep barranca, Aug-Sep 1886, E. Palmer 414 (holotype: GH00058424; isotypes: K000297147,
NY00006284, US00001874, US00930891).

= Hedysarum arborescens Sessé & Moc., Fl. Mexic., ed. 2. 170. 1894. Type: MEXICO, Jalisco, “in
Prædio S. Tomæ, prope Hostotipaquillum” [Santo Tomás near Hostotipaquillo], Jul, M. de Sessé &
J.M. Mociño 2009 (lectotype designated by Nelson Sutherland 1997: MA601239; isolectotype: F).
Other original material: M. de Sessé & J.M. Mociño 1938 (F, MA601240); M. de Sessé & J.M. Mociño
1948ter (F, MA601241); [icon] Torner Collection of Sessé and Mociño Biological Illustration No.
1890: drawing by Echeverría, 1790-1791, "Hedissarum arborescence" (Hunt Institute for Botanical
Documentation: Art Accession Number: 6331.1890).
= Aeschynomene bracteolaris L.Riley, Bull. Misc. Inform. Kew 1923(3): 115. 1923. Type: MEXICO,
Sinaloa, San Ignacio, El Coacoyal, May 1921, J. González Ortega 586 (holotype: K000297145;
isotype: K000297146).
= Aeschynomene guadalajarana M.E.Jones, Contr. W. Bot. 18: 46. 1933. Type: MEXICO, Jalisco, La
Barranca, Guadalajara, Nov 1930, M.E. Jones 27212 (Syntype: RSA0002730); idem, La Barranca,
Nayarit, Feb 1927, M.E. Jones 23052 (Syntype: F, RSA0023934).
Ctenodon nanus (Glaz. ex Rudd) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.
Basionym: Aeschynomene nana Glaz. ex Rudd, Phytologia 15: 118. 1967. Type: BRAZIL, Goiás,
entre As Brancas et Hector près du Rio Roncador, Dec 1894, A.F.M. Glaziou 20921 (holotype
fide Rudd 1967: P; lectotype designated here: P02914072; isolectotypes: BR0000005189861,
C10011416, F n.v., G [F neg. 27930], IAN [23706], K000222442, MG, NY00005011,
P02914067, P02914071, RB00158237, S-R-9502).
Ctenodon nicaraguensis (Oerst.) A.Delgado, comb. nov.

Basionym: Brya nicaraguensis Oerst., Vidensk. Meddel. Naturhist. Foren. Kjøbenhavn 1853: 13.
1853. Type: NICARAGUA, “paa Vulkanen El Viejo ... og i Provindsen Segovia”, Nov 1847, A.S.
Ørsted 4701 (holotype: C10011419; isotypes: F0058966F, K000297138, S13-12057).
≡ Aeschynomene nicaraguensis (Oerst.) Standl., Trop. Woods 34: 41. 1933.
= Aeschynomene calderoniana Standl., J. Washington Acad. Sci. 14: 93. 1924. Type: EL SALVADOR,
Santa Ana, vicinity of Santa Ana, Jan 1930, P.C. Standley 20367 (holotype: US00001905; isotypes:
GH00058426, NY00004979).
Distribution—El Salvador, Guatemala, Honduras, Mexico, and Nicaragua.
Ctenodon niveus (Brandegee) A.Delgado, comb. nov.

Basionym: Aeschynomene nivea Brandegee, Proc. Calif. Acad. Sci., ser. 2, 2: 150. 1889. Type:
MEXICO, Baja California Sur, La Purísima, Feb 1889, T.S. Brandegee s.n. (holotype: UC80594;
isotypes: GH00058431, NY n.v., PH00021835, US00001909).

Ctenodon oroboides (Benth.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.

Basionym: Aeschynomene oroboides Benth., Fl. Bras. (Martius) 15(1): 64. 1859. Type: BRAZIL,
Mato Grosso do Sul, in campis siccis secus Rio Pardo, Sep 1826, L. Riedel 502 (lectotype
designated here: K000222436; isolectotypes: B † [F neg. 2151], F0058815F, LE, P02914077).
Other syntypes: BRAZIL, loco haud indicato, s.d., J. Schücht s.n. [45] (K000222434,
W0027269). BRAZIL, Goiás, “Sertao d’Amaroleité” [Sertão de Amaro Leite], Sep 1844, H.A.
Weddell 2771 (K000920313, P02914076).
= Ctenodon weddellianus Baill., Adansonia 9: 237. 1870, as “Weddellianum”. Type: BRAZIL, Goiás,
“Sertao d’Amaroleité” [Sertão de Amaro Leite], Sep 1844, H.A. Weddell 2771 (lectotype
designated here: P02914076; isolectotype: K000920313). Other syntype: BRAZIL, Mato Grosso,
s.d., Herbier Impérial du Brésil no. 214 (P02914079, P02914080).
Ctenodon palmeri (Rose) A.Delgado, comb. nov.

Basionym: Aeschynomene palmeri Rose, Contr. U.S. Natl. Herb. 5: 192. 1899. Type: MEXICO,
Guerrero, near Acapulco, 1894-1895, E. Palmer 106a (holotype: US00001913; isotypes: GH
n.v., K000297141).
Ctenodon paniculatus (Willd. ex Vogel) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.
Basionym: Aeschynomene paniculata Willd. ex Vogel, Linnaea 12(1): 95. 1838. Type: BRAZIL,
loco haud indicato, s.d., F.W. Sieber s.n. [J. C. von Hoffmannsegg] (lectotype designated here:
B-W 13740-00 0, B-W 13740-01 0 [F neg. 2242]]; possible isolectotype: P02914020). Other
syntypes: BRAZIL, Bahia, F. Sellow s.n. (B† [F0058816F – F fragm.]). BRAZIL, Bahia, Cruz de
Casma, Jul 1835, B. Luschnath 223 (B†, LE, W18890119766). BRAZIL, loco haud indicato, A.I.
Gomes s.n. (B†).
= Aeschynomene hedysaroides M.Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 10(2): 181. 1843.
Type: MEXICO, Oaxaca, “Cordillera”, savannes de Zacatepeque, Sep 1840, H.G. Galeotti 3184
(holotype: BR0000005119554; isotypes: G [F neg. 27928], P02912794).
= Aeschynomene laevis M.Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 10(2): 181. 1843, nom.
illeg., hom. post. (non Noronha 1790). Type: MEXICO, Veracruz, les savanes de Zacuapan, Oct 1840,
H.G. Galeotti 3261 (holotype: BR0000005119882; isotype: P02913984).
= Aeschynomene gracilis Miq., Linnaea 18: 566. 1844, nom. illeg., hom. post. (non Vogel 1838). Type:
SURINAME, in savana arenosa ad plant. Berlyn, Sep 1844, H.C. Focke 966 (holotype: U0003457;
isotypes: BR0000013054298, GH00058452).
Ctenodon paucifolius (Vogel) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Basionym: Aeschynomene paucifolia Vogel, Linnaea 12(1): 94. 1838. Type: BRAZIL, Minas
Gerais, Santo Antonio do Monte, Nov 1818, F. Sellow s.n. (holotype: B†; probable isotype:

K000222440). Possible original material: BRAZIL, Minas Gerais, Serra do Galheiro, Oct 1818, F.
Sellow 1004 (B† [F neg. 2152]).
Ctenodon paucifoliolatus (Micheli) A.Delgado, comb. nov.

Basionym: Aeschynomene paucifoliolata Micheli, Mém. Soc. Phys. Genève 34: 256, t. 9. 1903.
Type: MEXICO, Guerrero, El Calabazal, 200 m, Oct 1898, E. Langlassé 474 (holotype: G;
isotypes: GH00058435, K000297143, MEXU00051044, MPU022827, P02912304, P02912305,
US00001912).
Ctenodon petraeus (B.L.Rob.) A.Delgado, comb. nov.

Basionym: Aeschynomene petraea B.L.Rob., Proc. Amer. Acad. Arts 27: 166. 1893. Type:
MEXICO, Jalisco, rocky hills near Guadalajara, May 1891, C.G. Pringle 5147 (holotype:
GH00058436; isotypes: K000297136, MEXU01169216, US00001911, UVMVT024910).
Ctenodon petraeus var. petraeus
Ctenodon petraeus var. grandiflorus (Rudd) A.Delgado, comb. nov.

Basionym: Aeschynomene petraea var. grandiflora Rudd, Contr. U.S. Natl. Herb. 32: 122. 1955.
Type: MEXICO, Durango, near Chacala, Feb 1899, E.A. Goldman 329 (holotype: US00001910;
isotype: NY00004986).
= Aeschynomene glomerata M.E.Jones, Contr. W. Bot. 15: 139. 1929. Type: MEXICO, Nayarit,
Acaponeta, at El Tigre mine, Mar 1927, M.E. Jones [23057] 23025 (holotype: RSA0002731;
isotypes: F0058797F, GH n.v., NY00004982).
Ctenodon petraeus var. madrensis (Micheli) A.Delgado, comb. nov.

Basionym: Aeschynomene petraea var. madrensis (Micheli) Rudd, Contr. U.S. Natl. Herb. 32: 121.
1955 ≡ Aeschynomene madrensis Micheli, Mém. Soc. Phys. Genève 34: 255, t. 8. 1903. Type:
MEXICO, Guerrero, “Sierra Madre, versant oriental”, [vicinity of Río Coyuquilla], 1500 m, Feb
1899, E. Langlassé 865 (holotype: G; isotypes: GH00058430, K000297134, P02913980,
P02913981, US00001891).
= Aeschynomene pringlei Rose, Contr. U.S. Natl. Herb. 8: 312. 1905. Type: MEXICO, Morelos,
limestone hills near Jojutla, Aug-Oct 1902, C.G. Pringle 8709 (lectotype designated here:
US00001923; isolectotypes: ARIZ-BOT-0004085, BM000931565, C10011422, CAS0000119,
CM0945, DAO000424204, E00394280, ENCB003331, V0058801F, GH00058439, GOET004619,
HBG520669, K000297135, L0018468, L0018469, L0018470, LL00371217, M0233747,
MEXU01169217, MEXU01169218, MICH1107080, MIN1000048, MO-128346, MSC0129843,

NY00004978, P02912480, P02912502, P02912503, P02912504, PH00000047, RSA0002727, S-

G-8844, UC80589, US01107508, UVMVT024911, UVMVT053376).
Ctenodon pinetorus (Brandegee) A.Delgado, comb. nov.

Basionym: Aeschynomene pinetorum Brandegee, Univ. Calif. Publ. Bot. 10: 408. 1924. Type:
MEXICO, Chiapas, in pine forests near Hacienda Monserrate, Sep 1923, C.A. Purpus 9064
(holotype: UC220371; isotypes: F0058800F, GH00058438, MO-128349, NY00004988,
US00001920).
= Aeschynomene chiapensis Brandegee, Univ. Calif. Publ. Bot. 10: 407. 1924. Type: MEXICO, Chiapas,
near Hacienda Monserrate, Sep 1923, C.A. Purpus 9147 (holotype: UC220376; isotypes:
F0058794F, GH00058425, MEXU00053652, MO-128348, NY00004980, US01108256).
Ctenodon pleuronervius (DC.) D.B.O.S.Cardoso & H.C.Lima, comb. nov.

Basionym: Aeschynomene pleuronervia DC., Prodr. [A. P. de Candolle] 2: 321. 1825. Smithia
domingensis Balb. ex DC., Prodr. [A. P. de Candolle] 2: 321. 1825, nomen. Type: DOMINICAN
REPUBLIC, Santo Domingo, s.d., C.L.G. Bertero s.n. (holotype: G-DC [F neg. 33436]; isotypes:
MO-022343, MO-022344, P02912467).
= Aeschynomene aurea Leonard, J. Washington Acad. Sci. 17: 67. 1927. Type: HAITI, Artibonite,
vicinity of Ennery, Puilboreau road, 800 m, Jan 1926, E.C. Leonard 8818 (holotype: US00001866;
isotype: GH00058444).
Distribution—Haiti and Dominican Republic.
Ctenodon podocarpus (Vogel) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Basionym: Aeschynomene podocarpa Vogel, Linnaea 12(1): 89. 1838 ≡ Aeschynomene falcata
var. plurijuga Benth., Fl. Bras. (Martius) 15(1): 68. 1859, p.p. Type: BRAZIL, “Brasil. merid.”, loco
haud indicato, s.d., F. Sellow s.n. (holotype: B†).
Ctenodon priscoanus (Afr.Fern.) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Basionym: Aeschynomene priscoana Afr.Fern., O Táxon Aeschynomene no Brasil: 86. 1996. Type:
BRAZIL, Piauí, Ribeiro Gonçalves, Estação Ecológica de Uruçuí-Una, Apr 1981, A. Fernandes s.n.
(holotype: EAC0010005).
Ctenodon purpusii (Brandegee) A.Delgado, comb. nov.

Basionym: Aeschynomene purpusii Brandegee, Zoë 5(11): 247. 1908. Type: MEXICO, Veracruz,
Zacuapán, 1906, C.A. Purpus 1904 (holotype: UC83413; isotypes: F0058802F, GH00058440,
MO-128350, NY00004989, P02912496, UC84877, US00001924, US00001925).

Ctenodon racemosus (Vogel) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Basionym: Aeschynomene racemosa Vogel, Linnaea 12(1): 92. 1838. Type: BRAZIL, Minas
Gerais, Santo Antonio do Monte, Nov 1818, F. Sellow s.n. (holotype: B†).
Ctenodon riedelianus (Taub.) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Basionym: Aeschynomene riedeliana Taub., Flora 72: 424. 1889. Type: BRAZIL, Minas Gerais, in
glareosis pr. Diamantina, Dec 1824, L. Riedel 1223 (holotype: LE n.v.; isotypes: B † [F neg. 2154],
K000222448, P02913159).
Ctenodon rosei (C.V.Morton) A.Delgado, comb. nov.

Basionym: Aeschynomene rosei C.V.Morton, Contr. U.S. Natl. Herb. 29: 84. 1944 ≡
Aeschynomene fruticosa Rose, Contr. U.S. Natl. Herb. 5(4): 192. 1899, nom. illeg., hom. post.
(non Sessé & Moc. 1889). Type: MEXICO, Sinaloa, on stony hills near Topolobampo, Sep 1897,
E. Palmer 204 (holotype: US00001896; isotypes: ARIZ-BOT-0004084, CAS0000117, FI010920,
MICH1107081, P02297639, RSA0002732, S-G-8843, UC770356).
Ctenodon sabulicolus (L.P.Queiroz & D.B.O.S.Cardoso) D.B.O.S.Cardoso, comb. nov.

Basionym: Aeschynomene sabulicola L.P.Queiroz & D.B.O.S.Cardoso, Bot. J. Linn. Soc. 157(4):
750. 2008. Type: BRAZIL, Bahia, Casa Nova, Fazenda Santarém, 9 o31’47”S, 41o21’34”W, Oct
2004, L.P. Queiroz et al. 9640 (holotype: HUEFS0089017; isotypes: K, MBM336389, MO-
2196354, NY00990979, SPF00181181).
Ctenodon scoparius (Kunth) D.B.O.S.Cardoso & H.C.Lima, comb. nov.

Basionym: Aeschynomene scoparia Kunth, Nov. Gen. Sp. [quarto ed.] 6: 531. 1823. Type: PERU,
Piura, “prope Guancabambam Peruvionorum” [Huancabamba], Aug, A.J.A. Bonpland & F.W.H.A.
von Humboldt s.n. [3543] (holotype: P00679169; isotypes: B-W 13741-01 0) [F neg. 2243], F
n.v., P02912477).
Distribution—Ecuador, Peru, and Suriname.
Ctenodon simplicifolius (G.P.Lewis) D.B.O.S.Cardoso & H.C.Lima, comb. nov.

Basionym: Aeschynomene simplicifolia G.P.Lewis, Kew Bull. 47(1): 141. 1992. Type: BRAZIL,
Goiás, Chapada dos Veadeiros, about 22 km of Alto Paraíso, 1200 m, Aug 1971, D.R. Gifford &
S.G. da Fonseca G241 (holotype: E00296478; isotype: UB [43048]).

Ctenodon simulans (Rose) A.Delgado, comb. nov.

Basionym: Aeschynomene simulans Rose, Contr. U.S. Natl. Herb. 5: 192. 1899. Type: MEXICO,
Sinaloa, between Rosario and Colomas, Jul 1897, J.N. Rose 1616 (lectotype designated by
Rudd 1955: US00001929; isolectotypes: GH00058441, K000297137). Other syntype: MEXICO,
Tepic, Nayarit, Acaponeta, Jun 1897, J.N. Rose 1487 (US01108255).
Ctenodon soniae (G.P.Lewis) D.B.O.S.Cardoso & H.C.Lima, comb. nov.

Basionym: Aeschynomene soniae G.P.Lewis, Kew Bull. 49(1): 93. 1994. Type: BRAZIL, Bahia, 27
km S of Livramento do Brumado on road to Brumado near two hamlets of Fazendinha and
Itapicuru, Apr 1991, G.P. Lewis & S.M.M. de Andrade 2006 (holotype: CEPEC00050387;
isotypes: BR0000005119943, E00296477, G00364571, HBG520662, HBG520663, HUEFS
[99848, 99849], K000092403, K000092404, M0233754, MBM221652, MEXU00607347, MO-
022339, NY00022635, SI001488, SPF00092629, SFV, TEX00371218, US00432869).
Ctenodon sousae (Rudd ex A.Delgado & Sotuyo) A.Delgado, comb. nov.

Basionym: Aeschynomene sousae Rudd ex A.Delgado & Sotuyo, Revista Mex. Biodivers. 83(2):
330. 2012. Type: MEXICO, Oaxaca, Tehuantepec District, municipio Santo Domingo
Tehuantepec, a 15 km sobre la desviación Buenos Aires a San Miguel Tenango, 16 o18’48.2”N,
95o30’10.7”W, 700 m, Oct 2010, A. Delgado-Salinas & S. Sotuyo 2732 (holotype: MEXU;
isotypes: GH, K, MO, NY, TEX, US).
Ctenodon standleyi (Ant.Molina) A.Delgado, comb. nov.

Basionym: Aeschynomene standleyi Ant.Molina, Ceiba 3(2): 92. 1952. Type: HONDURAS,
Department of Morazán, drainage of the Rio Yeguare, 900 m, Nov. 1948, A. Molina Rosito
1708a (holotype: US00001930; isotype: F n.v.).
Distribution—Honduras, Nicaragua, Costa Rica, and Mexico.
Ctenodon tenuis (Griseb.) D.B.O.S.Cardoso & H.C.Lima, comb. nov.

Basionym: Aeschynomene tenuis Griseb., Cat. Pl. Cub. [Grisebach]: 72. 1866. Type: CUBA,
Vueldabajo, Jul [1860-1864], C. Wright 2306 (Isotypes: BM000931559, GH00058447,
K000222412, MO-022347, NY00004993, P02913163, S13-12035, S-R-9506, US00001931,
YU001421).
Distribution—Cuba.

Ctenodon tumbezensis (J.F.Macbr.) D.B.O.S.Cardoso & H.C.Lima, comb. nov.

Basionym: Aeschynomene tumbezensis J.F.Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 8: 101.
1930. Type: PERU, Tumbez, Mountains east of Hacienda Chicama, 400-500 m, Feb 1927, A.
Weberbauer 7669 (holotype: F0043417F; isotypes: GH00058465, NY00005015, US00001955).
Distribution—Ecuador and Peru.
Ctenodon veadeiranus (M.J.Silva, & L.L.C.Antunes) D.B.O.S.Cardoso & H.C.Lima, comb. nov.
Basionym: Aeschynomene veadeirana M.J.Silva, & L.L.C.Antunes, Phytotaxa 184(1): 32. 2014.
Type: BRAZIL, Goiás, Teresina de Goiás, 13 o54’39.5”S, 47o22’48”W, 1278 m, Jul 2014, M.J.
Silva et al. 6127 (holotype: UFG; isotypes: CEN00100963, RB01340731, SP003689,
UB0040052).
Ctenodon venezolanus (Rudd) D.B.O.S.Cardoso & H.C.Lima, comb. nov.

Basionym: Aeschynomene brasiliana var. venezolana Rudd, Contr. U.S. Natl. Herb. 32: 80. 1955.
Type: VENEZUELA, Miranda, La Cortada, road to Guatire, near Petare, Nov 1923, H. Pittier
11233a (holotype: VEN3775; isotype: US00001906).
≡ Aeschynomene venezolana (Rudd) G.B.Rodr. & G.Agostini, Acta Bot. Venez. 16(1): 123. 1991.
Distribution—Venezuela.
Ctenodon vigil (Brandegee) A.Delgado, comb. nov.

Basionym: Aeschynomene vigil Brandegee, Proc. Calif. Acad. Sci. ser. 2, 3: 128. 1891. Type:
MEXICO, Baja California Sur, San José del Cabo, Oct 1891, T.S. Brandegee 142 (holotype:
UC80586; isotypes: GH00058443, PH00000049).
Ctenodon viscidulus (Michx.) D.B.O.S.Cardoso & A.Delgado, comb. nov.

Basionym: Aeschynomene viscidula Michx., Fl. Bor.-Amer. (Michaux) 2: 74. 1803. Type: UNITED
STATES OF AMERICA, “in arenosis insulæ Cumberland et Floridæ”, A. Michaux s.n. (holotype: P-
MICH ?). Possible original material: “Amériq.[ue] Sept.[entrionale]”, s.d., A. Michaux s.n.
(P02913175).
≡ Aeschynomene prostrata Poir., Encyc. [J. Lamarck & al.] Suppl. 4: 76. 1816.
≡ Secula viscidula (Michx.) Small, Fl. Miami: 90. 1913.
= Aeschynomene eriocarpa Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 9. 1943. Type:
GUATEMALA, Zacapa, rocky hills in vicinity of Santa Rosalia, Oct 1939, J.A. Steyermark 29313
(holotype: F0058796F).

= Aeschynomene gilbertoi Brandão, Daphne 1(2): 5. 1991. Type: BRAZIL, Minas Gerais, Mocambinho,
Nov 1989, J.F. Macedo 567 (holotype: RB; isotype: PAMG).
Distribution—USA, Mexico, Central America, Bolivia, Brazil, and Venezuela.
Ctenodon vogelii (Rudd) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Basionym: Aeschynomene vogelii Rudd, J. Wash. Acad. Sci. 49: 48. 1959. Type: BRAZIL, Minas
Gerais, Serra do Itambé, Oct 1818, F. Sellow s.n. (1382) (lectotype designated by Rudd 1959:
K000222432; isolectotypes: BM000931742, V0058817F, NY00005009).
≡ Aeschynomene falcata (Poir.) DC. var. multijuga Benth., Fl. Bras. 15(1): 68. 1859.
Ctenodon warmingii (Micheli) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Basionym: Aeschynomene warmingii Micheli, Vidensk. Meddel. Dansk Naturhist. Foren.
Kjøbenhavn: 68. 1875. Type: BRAZIL, Minas Gerais, Lagoa Santa, Dec 1865, J.E.B. Warming
[3015/2] (lectotype designated here: C10011424; isolectotypes: B [F neg. 2155], C10011423,
F, P02913199).
Ctenodon weberbaueri (Ulbr.) D.B.O.S.Cardoso & H.C.Lima, comb. nov.

Basionym: Aeschynomene weberbaueri Ulbr., Bot. Jahrb. Syst. 37(5): 554. 1906. Type: PERU,
Amazonas, Chachapoyas, “supra Balsas, in declivibus vallis Marañon fluminis loco stepposo
2000-2100 m s. m.”, Jun 1904, A. Weberbauer 4274 (holotype: B† [F neg. 2156]; isotypes:
V0043416F, USM n.v.).
Distribution—Ecuador and Peru.
5 | ACKNOWLEDGEMENTS
We gratefully acknowledge the two reviewers for critically reading an early version of the
manuscript. DBOSC’s research in plant biodiversity is also supported by grants from CNPq (Research
Productivity Fellowship, grant no. 308244/2018-4; Edital Universal grant no. 422325/2018-0),
FAPESB (grant no. APP0037/2016), and The Royal Society (Newton Advanced Fellowship no. NAF\
R1\180331). FF would like to thank the Senior Postdoctoral grant from FAPERJ (E-
26/201.562/2018 - process 236717), and the Reflora Program/JBRJ.

6 | LITERATURE CITED
Antunes, L.L.C. & Silva, M.J. 2017. New amphibious species of Aeschynomene (Leguminosae,
Papilionoideae, Dalbergieae) from the North Region of Brazil. Systematic Botany 42: 823–829.
Antunes, L.L.C. & Silva, M.J. 2019. Aeschynomene eridesii (Leguminosae, Papilionoideae, Dalbergiae),
a new species from seasonally flooded environments of Brazil. Phytotaxa 424: 123–129.
Baillon, H.E. 1870. Du nouveau genre Ctenodon. Adansonia 9: 236–237.
Brottier, L., Chaintreuil, C., Simion, P., Scornavacca, C., Rivallan, R., Mournet, P., Moulin, L., Lewis, G.P.,
Fardoux, J., Brown, S.C., Gomez-Pacheco, M., Bourges, M., Hervouet, C., Gueye, M., Duponnois,
R., Ramanankierana, H., Randriambanona, H., Vandrot, H., Zabaleta, M., DasGupta, M., D’Hont,
A., Giraud, E. & Arrighi, J-F. 2018. A phylogenetic framework of the legume genus
Aeschynomene for comparative genetic analysis of the Nod-dependent and Nod-independent
symbioses. BMC Plant Biology 18: 333.
Bruneau, A., Mercure, M., Lewis, G.P. & Herendeen, P.S. 2008. Phylogenetic patterns and
diversification in the caesalpinioid legumes. Botany 86: 697–718.
Cardoso, D., Queiroz, L.P., Pennington, R.T., Lima, H.C., Fonty, E., Wojciechowski, M.F. & Lavin, M.
2012a. Revisiting the phylogeny of papilionoid legumes: New insights from comprehensively
sampled early-branching lineages. American Journal of Botany 99: 1991–2013.
Cardoso, D., Lima, H.C., Rodrigues, R.S., Queiroz, L.P., Pennington, R.T. & Lavin, M. 2012b. The
realignment of Acosmium sensu stricto with the Dalbergioid clade (Leguminosae:
Papilionoideae) reveals a proneness for independent evolution of radial floral symmetry
among early-branching papilionoid legumes. Taxon 61: 1057–1073.
Cardoso, D., Pennington, R.T., Queiroz, L.P., Boatwright, J.S., Van Wyk, B.E., Wojciechowski, M.F. &
Lavin, M. 2013. Reconstructing the deep-branching relationships of the papilionoid legumes.
South African Journal of Botany 89: 58–75.
Cardoso, D., São-Mateus, W.M.B., Cruz, D.T., Zartman, C.E., Komura, D.L., Kite, G., Prenner, G.,
Wieringa, J.J., Clark, A., Lewis, G., Pennington, R.T. & Queiroz, L.P. 2015. Filling in the gaps of
the papilionoid legume phylogeny: The enigmatic Amazonian genus Petaladenium is a new
branch of the early-diverging Amburaneae clade. Molecular Phylogenetics and Evolution 84:
112–124.
Cardoso, D., Harris, D.J., Wieringa, J.J, São-Mateus, W.M.B., Batalha-Filho, H., Torke, B.M. Prenner, G.
& Queiroz, L.P. 2017. A molecular-dated phylogeny and biogeography of the monotypic
legume genus Haplormosia, a missing African branch of the otherwise American-Australian
Brongniartieae clade. Molecular Phylogenetics and Evolution 107: 431–442.
Cardoso, D.B.O.S., Ramos, G., São-Mateus, W.M.B. & Queiroz, L.P. 2019. Aeschynomene
chicocesariana, a striking new unifoliolate legume species from the Brazilian Chapada
Diamantina and its phylogenetic placement in the Dalbergioid clade. Systematic Botany 44:
810–817.

Carleial, S., Delgado-Salinas, A., Domínguez, C.A. & Terrazas, T. 2015. Reflexed flowers in
Aeschynomene amorphoides (Fabaceae: Faboideae): a mechanism promoting pollination
specialization? Botanical Journal of the Linnean Society 177: 657–666.
Chaintreuil, C., Arrighi, J.-F., Giraud, E., Miché, L., Moulin, L., Dreyfus, B., Munive-Hernández, J.A.,
Villegas-Hernandez, M.C. & Béna, G. 2013. Evolution of symbiosis in the legume genus
Aeschynomene. New Phytologist 200: 1247–1259.
Chamberlain, S., Barve, V., Mcglinn, D., Oldoni, D., Desmet, P., Geffert, L. & Ram, K. 2020. rgbif:
Interface to the Global Biodiversity Information Facility API. R package version 2.2.0, <URL:
https://CRAN.R-project.org/package=rgbif>.
Conceição, A.S., Queiroz, L.P., Lewis, G.P., Andrade, M.J.G., Almeida, P.R.M., Schnadelbach, A.S. & van
den Berg, C. 2009. Phylogeny of Chamaecrista Moench (Leguminosae-Caesalpinioideae) based
on nuclear and chloroplast DNA regions. Taxon 58: 1168–1180.
Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. 2012. jModelTest 2: More models, new heuristics
and parallel computing. Nature Methods 9: 772.
De-la-Estrella, M., Forest, F., Klitgård, B., Lewis, G.P., Mackinder, B.A., Queiroz, L.P., Wieringa, J.J. &
Bruneau, A. 2018. A new phylogeny-based tribal classification of subfamily Detarioideae, an
early branching clade of florally diverse tropical arborescent legumes. Scientific Reports 8:
6884.
Delgado-Salinas, A. & Sotuyo, S. 2012. A new species of Aeschynomene (Papilionoideae: Dalbergieae)
from Oaxaca, Mexico. Revista Mexicana de Biodiversidad 83: 329–333.
Delgado-Salinas, A., Thulin, M., Pasquet, R., Weeden, N. & Lavin, M. 2011. Vigna (Leguminosae)
sensu lato: The names and identities of the American segregate genera. American Journal of
Botany 98: 1694–1715.
Drummond, C.S., Eastwood, R.J., Miotto, S.T.S. & Hughes, C.E. 2012. Multiple continental radiations
and correlates of diversification in Lupinus (Leguminosae): Testing for key innovation with
incomplete taxon sampling. Systematic Biology 61: 443–460.
Duan L, Harris, A.J., Su, C., Zhang, Z.-R., Arslan, E., Ertugrul, K., Loc, P.K., Hayashi, H., Wen, J. & Chen,
H.-F. 2020. Chloroplast phylogenomics reveals the intercontinental biogeographic history of
the liquorice genus (Leguminosae: Glycyrrhiza). Frontiers in Plant Science 11: 793.
Felsenstein, J. 1985. Confidence limits on phylogenies: An approach using the bootstrap. Evolution
39: 783–791.
Fernandes, A. 1996. O táxon Aeschynomene no Brasil. Fortaleza: EUFC.
Fick, S.E. & Hijmans, R.J. 2017. WorldClim 2: New 1km spatial resolution climate surfaces for global
land areas. International Journal of Climatology 37: 4302–4315.
Flora do Brasil 2020 under construction. Jardim Botânico do Rio de Janeiro. Available at:
<http://floradobrasil.jbrj.gov.br/>. Accessed on: 07 Aug. 2020.

Fortuna-Perez, A.P., Silva, M.J., Queiroz, L.P., Lewis, G.P., Simões, A.O., Tozzi, A.M.G.A., Sarkinen, T. &
Souza, A.P. 2018. Phylogeny and biogeography of the genus Zornia (Leguminosae:
Papilionoideae: Dalbergieae). Taxon 62: 723–732.
Gagnon, E., Lewis, G.P., Sotuyo, J.S., Hughes, C.E. & Bruneau, A. 2013. A molecular phylogeny of
Caesalpinia sensu lato: Increased sampling reveals new insights and more genera than
expected. South African Journal of Botany 89: 111–127.
Gagnon, E., Bruneau, A., Hughes, C.E., Queiroz, L.P. & Lewis, G.P. 2016. A new generic system for the
pantropical Caesalpinia group (Leguminosae). PhytoKeys 71: 1–160.
Hijmans, R, and others. 2020. raster: Geographic data analysis and modeling. https://cran.r-
project.org/web/packages/raster/index.html
Huelsenbeck, J.P., Larget, B., Miller, R.E. & Ronquist, F. 2002. Potential applications and pitfalls of
Bayesian inference in phylogeny. Systematic Biology 51: 673–688.
Klitgaard, B.B. & Lavin, M. 2005. Tribe Dalbergieae sens. lat. In: Lewis, G.P., Schrire, B.D., MacKinder,
B.A. & Lock, M. (eds.). Legumes of the World. Royal Botanic Gardens, Kew, London, pp. 307–
335.
Koenen, E.J.M., Ojeda, D.I., Steeves, R., Migliore, J., Bakker, F.T., Wieringa, J.J., Kidner, C., Hardy, O.J.,
Pennington, R.T., Bruneau, A. & Hughes, C.E. 2020. Large-scale genomic sequence data resolve
the deepest divergences in the legume phylogeny and support a near-simultaneous
evolutionary origin of all six subfamilies. New Phytologist 225: 1355–1369.
Larsson, A. 2014. AliView: A fast and lightweight alignment viewer and editor for large datasets.
Bioinformatics 30: 3276–3278.
Lavin, M., Pennington, R.T., Klitgaard, B.B., Sprent, J.I., Lima, H.C. & Gasson, P.E. 2001. The
dalbergioid legumes (Fabaceae): Delimitation of a pantropical monophyletic clade. American
Journal of Botany 88: 503–533.
Lewis, G.P. 1985. Two new taxa and one little-known species of Aeschynomene from Brazil. Kew
Bulletin 40: 599–605.
Lewis, G.P. 1992. Two new species of Aeschynomene (Leguminosae: Papilionoideae) from Brazil. Kew
Bulletin 47: 141–145.
Lewis, G.P. 1993. Two new taxa of Aeschynomene (Leguminosae: Papilionoideae) from Brazil. Kew
Bulletin 49: 93–97.
LPWG [The Legume Phylogeny Working Group], 2013. Legume phylogeny and classification in the
21st century: Progress, prospects and lessons for other species-rich clades. Taxon 62: 217–248.
LPWG [The Legume Phylogeny Working Group]. 2017. A new subfamily classification of the
Leguminosae based on a taxonomically comprehensive phylogeny. Taxon 66: 44–77.
Miller, M.A., Pfeiffer, W. & Schwartz, T. 2010. Creating the CIPRES Science Gateway for inference of
large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop
(GCE), New Orleans, Louisiana.

Nelson-Sutherland, C. 1997. Material tipo de la colección de Sessé y Mociño en el Real Jardín

Botánico de Madrid. Anales del Jardín Botánico de Madrid 55(2): 375–418.
Nicholls, J.A., Pennington, R.T., Koenen, E.J.M., Hughes, C.E., Hearn, J., Bunnefeld, L., Dexter, K.G.,
Stone, G.N. & Kidner, C.A. 2015. Using targeted enrichment of nuclear genes to increase
phylogenetic resolution in the neotropical rain forest genus Inga (Leguminosae: Mimosoideae).
Frontiers in Plant Science 6: 710.
Ojeda, D.I., Koenen, E., Cervantes, S., De-la-Estrella, M., Banguera-Hinestroza, E., Janssens, S.B.,
Migliore, J., Demenou, B.B., Bruneau, A., Forest, F. & Hardy, O.J. 2019. Phylogenomic analyses
reveal an exceptionally high number of evolutionary shifts in a florally diverse clade of African
legumes. Molecular Phylogenetics and Evolution 137: 156–167.
Oliveira-Filho, A.T., Cardoso, D., Schrire, B.D., Lewis, G.P., Pennington, R.T., Brummer, T. J., Rotella, J.
& Lavin, M. 2013. Stability structures tropical woody plant diversity more than seasonality:
Insights into the ecology of high legume-succulent-plant biodiversity. South African Journal of
Botany 89: 42–57.
Pennington, R.T., Lavin, M. & Oliveira-Filho, A. 2009. Woody plant diversity, evolution and ecology in
the tropics: perspectives from seasonally dry tropical forests. Annual Review of Ecology,
Evolution, and Systematics 40: 437–457.
Pennington, R.T. & Lavin, M. 2016. The contrasting nature of woody plant species in different
neotropical forest biomes reflects differences in ecological stability. New Phytologist 210: 25–
37.
Queiroz, L.P. 2009. Leguminosas da Caatinga. Universidade Estadual de Feira de Santana, Feira de
Santana.
Queiroz, L.P. & Cardoso, D.B.O.S. 2008. A new species of Aeschynomene L. (Leguminosae,
Papilionoideae) from a continental sand dune area in north-eastern Brazil. Botanical Journal of
the Linnean Society 157: 749–753.
Queiroz, L.P., Pastore, J.F.B., Cardoso, D., Snak, C., Lima, A.L.C., Gagnon, E., Vatanparast, M., Holland,
A.E. & Egan, A.N. 2015. A multilocus phylogenetic analysis reveals the monophyly of a
recircumscribed papilionoid legume tribe Diocleae with well-supported generic relationships.
Molecular Phylogenetics and Evolution 90: 1–19.
Queiroz, L.P., São-Mateus, W., Delgado-Salinas, A., Torke, B.M., Lewis, G.P., Dorado, O., Ardley, J.K.,
Wojciechowski, M.F. & Cardoso, D. 2017. A molecular phylogeny reveals the Cuban enigmatic
genus Behaimia as a new piece in the Brongniartieae puzzle of papilionoid legumes. Molecular
Phylogenetics and Evolution 109: 191–202.
Rambaut, A. 2012. FigTree v1.4.0. University of Oxford, Oxford, UK. Website
<http://tree.bio.ed.ac.uk/software/figtree/>.
Rando, J.G., Zuntini, A.R., Conceição, A.S., van den Berg, C., Pirani, J.R. & Queiroz, L.P. 2016.
Phylogeny of Chamaecrista ser. Coriaceae (Leguminosae) unveils a lineage recently diversified
in Brazilian Campo Rupestre vegetation. International Journal of Plant Sciences 177: 3–17.

Ribeiro, R.A., Lavin, M., Lemos-Filho, J.P., Mendonça-Filho, C.A.V., Santos, F.R. & Lovato, M.B. 2007.
The genus Machaerium (Leguminosae) is more closely related to Aeschynomene sect.
Ochopodium than to Dalbergia: Inferences from combined sequence data. Systematic Botany
32: 762–771.
Ringelberg, J.J., Zimmermann, N.E., Weeks, A., Lavin, M. & Hughes, C.E. 2020. Biomes as
evolutionary arenas: Convergence and conservatism in the trans‐continental succulent biome.
Global Ecology and Biogeography 29: 1100–1113.
Rodrigues, R.S., Hartmann, L.S. & Flores, A.S. 2019. Seedling morphology of some Brazilian taxa of
Aeschynomene (Leguminosae) and its systematic relevance. Flora 255: 69–79.
Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L.,
Suchard, M.A. & Huelsenbeck, J.P. 2012. MrBayes 3.2: Efficient Bayesian phylogenetic inference
and model choice across a large model space. Systematic Biology 61: 539–542.
Rudd, V.E. 1955. The American species of Aeschynomene. Contributions from the United States
National Herbarium 32: 1–172.
Rudd, V.E. 1959. Supplementary studies in Aeschynomene I: Series Viscidulae, including a new species
and five new varieties. Journal of the Washington Academy of Sciences 49: 45–52.
Rudd, V.E. 1967. Supplementary studies in Aeschynomene II: Series Pleuronerviae. Phytologia 15:
114–119.
Rudd, V.E. 1972. Reduction of Balisaea to Aeschynomene. Phytologia 23: 321–322.
Rudd, V.E. 1975. Supplementary studies in Aeschynomene III: Series Scopariae in Mexico and Central
America. Phytologia 31: 431–434.
Rudd, V.E. 1989. A new species and reconsiderations in Aeschynomene series Scopariae
(Leguminosae, Papilionoideae) in Mexico. Acta Botanica Mexicana 8: 31–33.
Sampaio, D.S., Moço, M.C.C. & Mariath, J.E.A. 2013. Floral ontogeny of Aeschynomene falcata and A.
sensitiva (Leguminosae: Papilionoideae) supports molecular phylogenetic data. Plant
Systematics and Evolution 299: 499–513.
Schrire, B.D., Lavin, M., Barker, N.P. & Forest, F. 2009. Phylogeny of the tribe Indigofereae
(Leguminosae–Papilionoideae): geographically structured more in succulent-rich and
temperate settings than in grass-rich environments. American Journal of Botany 96: 816–852.
Silva, M.J. & Antunes, L.L.C. 2014. An update of the Brazilian species of Aeschynomene sect.
Ochopodium ser. Viscidulae including a new species and a new synonym. Phytotaxa 184: 31–
38.
Simon, M.F., Grether, R., Queiroz, L.P., Särkinen, T.E., Dutra, V.F. & Hughes, C.E. 2011. The
evolutionary history of Mimosa (Leguminosae): Toward a phylogeny of the sensitive plants.
American Journal Botany 98: 1201–1221.
Sinou, C., Forest, F., Lewis, G.P. & Bruneau, A. 2009. The genus Bauhinia s.l. (Leguminosae): A
phylogeny based on the plastid trnL-trnF region. Botany 87: 947–960.

Stamatakis, A. 2014. RAxML version 8: A tool for phylogenetic analysis and post-analysis of large
phylogenies. Bioinformatics 30: 1312–1313.
Stamatakis, A., Hoover, P. & Rougemont, J. 2008. A rapid bootstrap algorithm for the RAxML web
servers. Systematic Biology 57: 758–771.
Tapia-Pastrana, F., Delgado-Salinas, A. & Caballero, J. 2020. Patterns of chromosomal variation in
Mexican species of Aeschynomene (Fabaceae, Papilionoideae) and their evolutionary and
taxonomic implications. Comparative Cytogenetics 14: 157–182.
Taubert, P. 1894. Leguminosae. In: Engler, A. & Prantl, K. (eds.). Die natürlichen Pflanzenfamilien.
Wilhelm Englemann, Leipzig, Germany, pp. 70–396.
Thiers, B. 2020 [constinuously updated]. Index Herbariorum: A global directory of public herbaria and
associated staff. New York Botanical Garden’s Virtual Herbarium. Available from:
http://sweetgum.nybg.org/science/ih/ (accessed 15 Aug 2020).
Torke, B.M. & Schaal, B.A., 2008. Molecular phylogenetics of the species-rich neotropical genus
Swartzia (Leguminosae, Papilionoideae) and related genera of the swartzioid clade. American
Journal of Botany 95: 215–228.
Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, W., Hawksworth, D.L., Herendeen, P.S., Knapp, S.,
Kusber, W.-H., Li, D.-Z., Marhold, K., May, T.W., McNeill, J., Monro, A.M., Prado, J., Price, M.J. &
Smith, G.F. (eds.) 2018. International Code of Nomenclature for algae, fungi, and plants
(Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China,
July 2017. Regnum Vegetabile 159. Koeltz Botanical Books, Glashütten, Germany.
Vanni, R.O. 2016. El género Aeschynomene (Leguminosae-Dalbergieae) en Argentina y Paraguay.
Boletín de la Sociedad Argentina de Botánica 51(4): 705–725.
Vatanparast, M., Klitgård, B.B., Adema, F.A.C.B., Pennington, R.T., Yahara, T. & Kajita, T. 2013. First
molecular phylogeny of the pantropical genus Dalbergia: Implications for infrageneric
circumscription and biogeography. South African Journal of Botany 89: 143–149.
Vilela, B. & Villalobos, F. 2015. letsR: A new R package for data handling and analysis in
macroecology. Methods in Ecology and Evolution 6: 1229–1234.
Vogel, J.R.T. 1838. De Hedysareis Brasiliae. Linnaea 12: 51–111.
Wiens, J.J. 2003. Missing data, incomplete taxa, and phylogenetic accuracy. Systematic Biology 52:
528–538.
Wiens, J.J. 2005. Can incomplete taxa rescue phylogenetic analyses from long branch attraction?
Systematic Biology 54: 731–742.
Wiens, J.J. & Tiu, J. 2012. Highly incomplete taxa can rescue phylogenetic analyses from the
negative impacts of limited taxon sampling. PLoS One 7: e42925.
Wojciechowski, M.F. 2005. Astragalus (Fabaceae): A molecular phylogenetic perspective. Brittonia 57:
382–396.

Wojciechowski, M.F., Sanderson, M.J. & Hu, J.M. 1999. Evidence on the monophyly of Astragalus
(Fabaceae) and its major subgroups based on nuclear ribosomal DNA ITS and chloroplast DNA
trnL intron data. Systematic Botany 24: 409–437.
Wojciechowski, M.F., Lavin, M. & Sanderson, M.J. 2004. A phylogeny of legumes (Leguminosae)
based on analysis of the plastid matK gene resolves many well-supported subclades within the
family. American Journal of Botany 91: 1846–1862.
Zhang, R., Wang, Y.-H., Jin, J.-J., Stull, G.W., Bruneau, A., Cardoso, D., Queiroz, L.P., Moore, M.J.,
Zhang, S.-D., Chen, S.-Y., Wang, J., Li, D.-Z. & Yi, T.-S. 2020. Exploration of plastid
phylogenomic conflict yields new insights into the deep relationships of Leguminosae.
Systematic Biology 69: 613–622.
Zizka, A., Silvestro, D., Andermann, T., Azevedo, J., Ritter, C.D., Edler, D., Farooq, H., Herdean, A.,
Ariza, M., Scharn, R., Svanteson S., Wengstrom, N., Zizka, V. & Antonelli, A. 2019.
CoordinateCleaner: Standardized cleaning of occurrence records from biological collection
databases. Methods in Ecology and Evolution 10: 744–751.
Accepted by: Cássio van den Berg on 20th Aug 2020

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A Journal of Neotropical Biodiversity ISSN 2358-2847 (online)

ARTICLE 02 Sep 2020 DOI: 10.13102/neod.131.1

A molecular phylogeny of the pantropical papilionoid

1 | www.neodiversity.org Neodiversity 13: 1–38. 2020

2 | www.neodiversity.org Neodiversity 13: 1–38. 2020

2 | MATERIALS AND METHODS

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4 | www.neodiversity.org Neodiversity 13: 1–38. 2020

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3 | RESULTS AND DISCUSSION

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Psorodendron spinosum AY391822

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Pterocarpus indicus JN083482

8 | www.neodiversity.org Neodiversity 13: 1–38. 2020

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always been taxonomically differentiated from A. sect. Aeschynomene by possessing non-peltate,

10 | www.neodiversity.org Neodiversity 13: 1–38. 2020

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Aeschynomene clade Ctenodon Aeschynomene clade Ctenodon

12 | www.neodiversity.org Neodiversity 13: 1–38. 2020

Ctenodon acapulcensis (Rose) A.Delgado, comb. nov.

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Ctenodon afranianus D.B.O.S.Cardoso, Filardi & H.C.Lima, nom. nov.

Ctenodon benthamii (Rudd) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Ctenodon bradei (Rudd) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Ctenodon brasilianus (Poir.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.

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Ctenodon brevipes (Benth.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.

Ctenodon carvalhoi (G.P.Lewis) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Ctenodon carichanicus (Rudd) D.B.O.S.Cardoso & H.C.Lima, comb. nov.

Ctenodon chicocesarianus (D.B.O.S.Cardoso & G.Ramos) D.B.O.S.Cardoso, comb. nov.

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Ctenodon compactus (Rose) A.Delgado, comb. nov.

Ctenodon egenus (J.F.Macbr.) D.B.O.S.Cardoso & H.C.Lima, comb. nov.

Ctenodon elegans var. elegans

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Ctenodon falcatus (Poir.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.

Ctenodon falcatus var. falcatus

17 | www.neodiversity.org Neodiversity 13: 1–38. 2020

Ctenodon fascicularis (Schltdl. & Cham.) A.Delgado, comb. nov.

Ctenodon foliolosus (Rudd) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Ctenodon genistoides (Taub.) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Ctenodon genistoides var. genistoides

18 | www.neodiversity.org Neodiversity 13: 1–38. 2020

Ctenodon gracilis (Vogel) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.

Ctenodon graminoides (G.P.Lewis) D.B.O.S.Cardoso & H.C.Lima, comb. nov.

Ctenodon hintonii (Sandwith) A.Delgado, comb. nov.

Ctenodon histrix (Poir.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.

Ctenodon histrix var. histrix

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BR0000005119516, F0058808F, GH00058449, L1970334, MICH1107076, P02913646,

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Ctenodon interruptus (Benth.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.

Ctenodon lacabuendianus (Brandão) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Ctenodon langlassei (Micheli ex Rudd) A.Delgado, comb. nov.

Ctenodon leptostachyus (Benth.) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

Ctenodon lewisianus (Afr.Fern.) D.B.O.S.Cardoso, Filardi & H.C.Lima, comb. nov.

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Ctenodon lyonnetii (Rudd) A.Delgado, comb. nov.

Ctenodon marginatus (Benth.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.

Ctenodon marginatus var. marginatus

Ctenodon martii (Benth.) D.B.O.S.Cardoso, P.L.R.Moraes & H.C.Lima, comb. nov.