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A new generic circumscription of Hydrochorea
(Leguminosae, Caesalpinioideae, mimosoid clade)
with an amphi-Atlantic distribution
Marcos Vinicius Batista Soares1,2*, Erik Jozef Mathieu Koenen3*,
João Ricardo Vieira Iganci1,4, Marli Pires Morim5
1Universidade Federal do Rio Grande do Sul, Programa de Pós-Graduação em Botânica, Av. Bento Gonçalves
9500, Bloco IV, Prédio 43433, 91501-970, Porto Alegre, Rio Grande do Sul, Brazil 2Rua dos Bandeirantes
1020, Caranazal, 68040-329, Santarém, Pará, Brazil 3Evolutionary Biology & Ecology, Université Libre
de Bruxelles, Faculté des Sciences, Campus du Solbosch - CP 160/12, Avenue F.D. Roosevelt 50, 1050 Brux-
elles, Belgium 4Instituto de Biologia, Universidade Federal de Pelotas, Campus Universitário Capão do Leão,
Travessa Andre Dreyfus s/n, 96010-900, Capão do Leão, Rio Grande do Sul, Brazil 5Instituto de Pesquisas
Jardim Botânico do Rio de Janeiro, Av. Pacheco Leão 915, 22460-030, Rio de Janeiro, Brazil
Corresponding author: Erik Jozef Mathieu Koenen (erikk_botany@gmx.com)
Academic editor: Gwilym P. Lewis|Received 25 February 2022|Accepted 23 June 2022|Published 22 August 2022
Citation: Soares MVB, Koenen EJM, Iganci JRV, Morim MP (2022) A new generic circumscription of Hydrochorea
(Leguminosae, Caesalpinioideae, mimosoid clade) with an amphi-Atlantic distribution. In: Hughes CE, de Queiroz
LP, Lewis GP (Eds) Advances in Legume Systematics 14. Classication of Caesalpinioideae Part 1: New generic
delimitations. PhytoKeys 205: 401–437. https://doi.org/10.3897/phytokeys.205.82775
Abstract
Hydrochorea and Balizia were established to accommodate four and three species, respectively, that were
previously included in dierent ingoid genera, based primarily on dierences in fruit morphology. Both
genera have Amazonia as their centre of diversity, extending to Central America and the Brazilian Atlantic
Rainforest. Previous phylogenetic evidence showed Balizia to be paraphyletic with respect to Hydrocho-
rea, and species of Hydrochorea and Balizia were placed in a large unresolved polytomy with species of
Jupunba. Here we present a new phylogenomic analysis based on 560 exons, from which 686 orthologous
alignments were derived for gene tree inference. is analysis conrms a paraphyletic Balizia in relation to
Hydrochorea, together with two African species formerly placed in Albizia nested within the clade. Jupunba
macradenia was resolved as sister to the clade combining those taxa. However, quartet support is low for
several of the branches at the base of the clade combining the genera Jupunba, Balizia and Hydrochorea,
suggesting that rapid initial divergence in this group led to extensive incomplete lineage sorting and con-
sequently poor phylogenetic resolution. Because of these phylogenomic complexities, we decided to use
morphology as the main guide to consider Hydrochorea as a distinct genus from Jupunba, and Balizia as
* ese authors contributed equally to this work.
Copyright Marcos Vinicius Batista Soares et al. This is an open access article distributed under the terms of the Creative Commons Attribution
License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source
are credited.
PhytoKeys 205: 401–437 (2022)
doi: 10.3897/phytokeys.205.82775
https://phytokeys.pensoft.net
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Marcos Vinicius Batista Soares et al. / PhytoKeys 205: 401–437 (2022)
402
a new synonym for Hydrochorea. e taxonomic treatment includes the study of collections from various
herbaria and eldwork expeditions. We present a re-circumscribed Hydrochorea accommodating a total
of 10 species, including six new combinations, ve new synonyms, one new taxonomic status, two cor-
rections of nomenclature category for lectotypes, and a second step lectotype and three new lectotypes. A
new species from the Brazilian Amazon is described and illustrated. An identication key for all species of
Hydrochorea is presented, together with comments and illustrations.
Keywords
Albizia, Balizia, Cathormion, Fabaceae, nomenclature, taxonomy
Introduction
Rupert C. Barneby and James W. Grimes established a new generic system for most
of the ingoid mimosoids of the Americas in a landmark monographic series (Barneby
and Grimes 1996, 1997; Barneby 1998). In it they created seven new genera, including
Hydrochorea Barneby & J.W. Grimes and Balizia Barneby & J.W. Grimes, which were
established to accommodate four and three species, respectively, that were previously
included (as many ingoid species have been) in several dierent genera such as Albizia
Durazz., Arthrosamanea Britton & Rose, Cathormion Hassk. and Pithecellobium Mart.,
among others (Lewis and Rico Arce 2005; Brown 2008). While Barneby and Grimes
(1996: p. 35) stated that Hydrochorea and Balizia “have arisen from common ancestry”,
these were nonetheless treated as separate genera based on dierences in fruit morphol-
ogy that were ascribed to adaptation to dierent habitats and seed dispersal strategies:
Hydrochorea was dened based on lomentiform fruits adapted to water-borne dispersal
in seasonally inundated habitats, while Balizia was described as a genus of “terra rme”
forest (even though at least two of its three species were mentioned to also often occur
on riverbanks), and recognized mainly based on having indehiscent or follicular fruits,
with a septate endocarp but not lomentiform. Barneby and Grimes (1996) recognized
Balizia, Hydrochorea, and Abarema Pittier s.l. as closely related genera, distinguished by
fruit morphology (Iganci and Morim 2009, 2012). However, Abarema was shown to be
polyphyletic (Iganci et al. 2016) and the type species of the genus, Abarema cochliacarpos
(Gomes) Barneby & J.W. Grimes is placed in the Inga clade, together with the recently
described A. diamantina E. Guerra, M.P. Morim & Iganci (Guerra et al. 2016, 2019).
e other species of Abarema s.l. were segregated in the two genera Jupunba Britton &
Rose and Punjuba Britton & Rose (Soares et al. 2021). ese ndings question all the
former classications of those taxa, which were mostly based on fruit morphology, and
call for further studies aiming to better understand fruit and seed morphology in the
context of the evolution of dispersal strategies and ecological adaptations.
Recent phylogenetic evidence (Iganci et al. 2016; Koenen et al. 2020b; Soares et al.
2021; Ringelberg et al. 2022; and a new analysis presented here) has shown Balizia to
be paraphyletic with respect to Hydrochorea. Furthermore, two African species that were
formerly placed in several genera including Albizia Durazz. and Cathormion Hassk.
Recircumscription of Hydrochorea 403
were also shown to be most closely related to Hydrochorea (Koenen et al. 2020b), and
their general morphological features and ecology are virtually indistinguishable from
Neotropical species of Hydrochorea.
Besides the advances in phylogenetic and phylogenomic methods, recent eldwork
collecting programmes have greatly contributed to herbarium collections of Amazo-
nian taxa (Milliken et al. 2010; Cardoso et al. 2017; Ulloa et al. 2017; BFG 2021), and
furthermore, the Reora Program has led to the online availability of nearly all Brazil-
ian plant collections (Pearce et al. 2020; BFG 2021), providing excellent opportunities
for synoptic taxonomic revisions.
Here we present a taxonomic update including a new generic circumscription of
Hydrochorea based on phylogenomic and morphological evidence, along with a no-
menclatural review presented as a synopsis of the genus, which includes new combina-
tions, new synonyms, and the description of a newly discovered species from the Up-
per Rio Negro. We include an identication key, illustrations, and distribution maps
for the 10 species now accommodated in Hydrochorea.
Materials and methods
Taxonomy
Standard herbarium taxonomy practices were used for analysis of all species studied in the
present work. e collections (including digital images) of the following herbaria were
analysed: A, BM, BR, CTBS, E, F, G, GH, HUEFS, IAN, INPA, K, MG, MO, NY, P,
PEL, R, RB, SP, US and Z (iers 2022). Fieldwork was carried out especially in the Up-
per Rio Negro region of Amazonian Brazil, where we collected four species of Balizia and
Hydrochorea, including a species new to science. All the new collections were incorporated
into the RB herbarium, in the Rio de Janeiro Botanical Garden, and duplicates were sent
to partner institutions. Fresh leaf samples were stored in silica gel for total DNA extraction.
We also visited the collections in NY to study the specimens that Barneby and Grimes
(1996) worked with for their taxonomic account. Combined with studying the large num-
ber of new collections that have been made in the past 25 years since publication of the
Barneby and Grimes (1996) taxonomic account, we were able to review their taxonomic
decisions based on the relatively limited herbarium material available to them at the time.
Online databases were used to view digital images of specimens including types, es-
pecially the Reora Virtual Herbarium (REFLORA -Herbário Virtual 2022), National
Institute of Science and Technology (INCT-Splink 2022), and JSTOR (2022). Geo-
graphic distributions of each species were inferred based on specimen labels and litera-
ture (Barneby and Grimes 1996; Soares 2015). e morphological characters were de-
scribed following Beentje (2016) and Iganci and Morim (2009, 2012), for habit, leaves,
inorescences, owers, pods, and seeds. Original descriptions of all taxa were analysed,
and nomenclature was revised according to the International Code of Nomenclature
of algae, fungi and plants (the Shenzen code; Turland et al. 2018). Except for the new
Marcos Vinicius Batista Soares et al. / PhytoKeys 205: 401–437 (2022)
404
species described here, and the two African species now placed in Hydrochorea, all other
species have been described earlier in detail by specialists (Barneby and Grimes 1996).
us, for those species, here we only present new combinations, taxonomic notes and
a reference to the literature where the complete description is available. Synonyms are
accepted following Barneby and Grimes (1996) and are only listed here when either
lectotypication or nomenclatural correction is needed.
Exon selection, matrix assembly and phylogenomic analysis
To better evaluate the evidence for monophyly of the studied genera, or lack thereof, we
have performed new analyses based on a selection of exons with anking non-coding re-
gions derived from the sequencing data of Koenen et al. (2020b) and Ringelberg et al.
(2022), including network analyses and quantication of supporting bipartitions across
gene trees for alternative topologies. e sequencing methods are described in those pub-
lications and here we only briey describe our methods when they dier from Koenen et
al. (2020b) and Ringelberg et al. (2022). In the original Mimobaits probe design (Koenen
et al. 2020b), exons were predicted, and anking untranslated regions (UTRs) were also
(partially) included. From this reference exon set, we selected all of those that are longer
than 500bp, which for initial or terminal exons includes the UTR. Read data of Koenen
et al. (2020b) and Ringelberg et al. (2022) for the accessions of the Jupunba clade plus six
outgroup accessions were mapped against these exons and non-matching reads discarded.
Read quality ltering and de novo assembly methods followed Koenen et al. (2020b), and
after clustering the assembled contigs to the reference sequences of the exons, initial align-
ments and gene trees were inferred using MAFFT (Katoh et al. 2005) and RAxML (Stama-
takis 2014), respectively. en, mono- and paraphyletic groups per species were collapsed
to select a single allele in case multiple alleles were reconstructed (Yang and Smith 2014),
followed by cutting long internal branches to splice potential paralogs into separate align-
ments using the cut_long_branches.py script of Yang and Smith (2014). e resulting
clusters were then realigned with MAFFT and used to infer multilabeled gene trees using
RAxML. Finally, gene trees with a single tip per species were extracted from these gene trees
using the maximum inclusion (MI) method of Yang and Smith (2014) and used in species
tree analysis in ASTRAL-III (Zhang et al. 2018) with default settings and nodes with less
than 10% bootstrap support collapsed as suggested by the authors of the software. A l-
tered supernetwork was constructed in SplitsTree4 (Huson 1998) from the same set of gene
trees but with nodes with less than 50% bootstrap support collapsed and with the mintrees
parameter set to 100 (25% of the total number of gene trees). Quantication of supporting
bipartitions across gene trees for alternative topologies followed the same methodology as
Koenen et al. (2020a).
Results
Our herbarium taxonomic work has resulted in the synopsis presented below. is
includes a total of six new combinations, including a new status for a species that had
Recircumscription of Hydrochorea 405
been treated at varietal rank by Barneby and Grimes (1996), as well as the description
of a new species. Balizia and its sections are placed in the synonymy of Hydrochorea
and two new heterotypic synonyms, one at species and one at varietal rank, are pro-
posed, as well as two lectotype corrections. A second step lectotype and three new
lectotypes are designated.
Phylogenomic analysis
A total of 560 exons were selected for gene tree inference, from which 398 MI gene
trees were extracted after clustering and ltering (Suppl. materials 1, 2). e species
tree based on these gene trees (Fig. 1A, Suppl. material 3) does not provide qualita-
tively dierent results from those of Ringelberg et al. (2022), showing a paraphyletic
Balizia relative to Hydrochorea together with the two African Cathormion/Albizia spe-
cies. All currently recognised species of Balizia and Hydrochorea, including the type
species of both genera, were sampled, with the exception of H. marginata, although
this species was included in the studies of Iganci et al. (2016) and Soares et al. (2021).
Also, Jupunba macradenia (Pittier) M.V.B. Soares, M.P. Morim & Iganci was found as
sister of the clade combining these genera. However, quartet support is low for several
of the branches at the base of the clade combining the genera Jupunba, Balizia and Hy-
drochorea, suggesting that rapid initial divergence in this group led to extensive incom-
plete lineage sorting and consequently poor phylogenetic resolution. is is further
reinforced by the ltered supernetwork (Fig. 1B), which also clearly shows the para-
phyly of Balizia, and shows that Hydrochorea and Balizia together form a group that is
separate from Jupunba, but with a complex network structure indicative of incomplete
lineage sorting. When the monophyly of these genera and their sister-group relation-
ships are evaluated based on the number of compatible bipartitions across gene trees,
it is clear that Albizia sect. Arthrosamanea (Britton & Rose) Barneby & J.W. Grimes,
Punjuba and Hydrochorea (including the two African Cathormion/Albizia species) are
each monophyletic, while Balizia and Jupunba are only supported to be monophyletic
by a small number of gene trees (Fig. 1C). Support for the sister-group relationship
of Jupunba macradenia with Balizia + Hydrochorea is supported by c. 10% of the gene
trees. is is higher than what is found for some other species of Jupunba that are sister
to Balizia + Hydrochorea in some gene trees, but nonetheless the separation between
these genera is not very clear likely due to signicant incomplete lineage sorting.
Discussion
In this study, we have made an in-depth investigation of the generic delimitation issues
surrounding the genera Balizia, Cathormion and Hydrochorea, to reconcile morphological
characters of the group with phylogenetic relationships and to propose a revised classica-
tion. While the uncovered phylogenomic complexity adds further diculty to the goal
of achieving a stable classication for these taxa, we conclude that the taxa with either
indehiscent, follicular or lomentiform fruits, that are septate between the seeds at least in
Marcos Vinicius Batista Soares et al. / PhytoKeys 205: 401–437 (2022)
406
Figure 1. Phylogenomics of the Jupunba clade A ASTRAL-III species tree based on 398 gene trees, pos-
terior probability values are shown only for those nodes for which support is lower than 1.0, and pie charts
on several crucial nodes indicate alternative quartet support B ltered super-network of the same gene
tree set with the genus Hydrochorea as circumscribed in this study indicated by a grey ellipse C bar graphs
indicating numbers of compatible bipartitions across the same gene tree set in the maximum-likelihood es-
timate (ML) and when only taking into account bipartitions that receive at least 50 or 80% bootstrap sup-
port. e abbreviations that are used are Bal = Balizia, Hyd = Hydrochorea, and Jup = Jupunba. Note that
the taxonomy of Albizia sect. Arthrosamanea is updated in this volume by Aviles Peraza et al. (2022), where
new binomials in the genus Pseudalbizzia are presented for the majority of the species of this section.
Recircumscription of Hydrochorea 407
the endocarp, are preferably all classied within a recircumscribed Hydrochorea, separate
from the genus Jupunba which is characterized by dehiscent fruits that are never septate
between the seeds. Extensive incomplete lineage sorting surrounding the early evolution
of these genera means that they are phylogenomically not well separated (Fig. 1), but in
order to ensure both diagnosability and stability of names, we believe keeping these as
separate genera is justied (i.e., transferring some species to Hydrochorea to account for
the non-monophyly of Balizia is preferable to moving all taxa to a morphologically het-
erogeneous Jupunba).
One of the most interesting aspects of Hydrochorea is the evolution of its fruit mor-
phology and dehiscence in adaptation to water-borne seed dispersal, which presumably
led to its distribution in riparian, swamp and periodically inundated forests on both
sides of the Atlantic, as trans-oceanic dispersal is presumed to be relatively likely in
hydrochorous plants. Much attention was traditionally given to pod morphology in
mimosoids, in attempts to classify the ingoid genera, as one of the most easily observ-
able characters to visually distinguish the taxa (Barneby and Grimes 1996; Lewis and
Rico-Arce 2005). Lomentiform pods are typical of most species of Hydrochorea, but are
also found in Albizia s.s. (Albizia dolichadena I.C. Nielsen, Albizia moniliformis (DC.)
F. Muell., Albizia rosulata (Kosterm.) I.C. Nielsen and Albizia umbellata (Vahl) E.J.M.
Koenen) and Albizia sect. Arthrosamanea (Barneby and Grimes 1996; Aviles Peraza
et al. 2022). e craspedia, as found in Mimosa L., Adenopodia C. Presl and Entada
Adans., are somewhat similar, but in those genera there is a replum (a persistent frame-
work formed by the upper and lower suture of the craspedium) that stays attached to
the infructescence after the 1-seeded articles have been shed. e fruit of Cathormion
altissimum (Hook. f.) Hutch. & Dandy (which is transferred to a new genus by Koenen
2022) is also similar, but the fruit of that species diers in containing aerenchymous
tissue on the seminiferous nuclei to promote oating. ese various lomentiform in-
goid fruits also dier in whether they break up while still attached to the tree as is the
case in most species of Hydrochorea and some species of Albizia sect. Arthrosamanea, or
whether the fruit falls from the tree entire and only tardily breaks up into 1-multiple
seeded articles afterwards, as appears to be the case in most of the other genera. Phylo-
genetic evidence clearly indicates that these fruits have all evolved independently from
one another (Ringelberg et al. 2022), presumably in response to adaptation to riparian
and periodically inundated habitats. e homoplasious nature of these similar fruits
has led to many species having been moved around between dierent genera, based on
their fruit morphology (Nielsen 1981). We note that the legume fruit appears to be
amenable to the evolution of lomentiform fragmentation, given the similar craspedia
that were independently derived at least twice and, moreover, lomentiform fruits are
also present in several lineages of subfamily Papilionoideae.
Recent advances in molecular systematics of ingoid legumes also demonstrated the
pod morphology to be less informative than previously thought (Souza et al. 2013;
Iganci et al. 2016; Koenen et al. 2020b; Soares et al. 2021; Souza et al. 2022). Barneby
and Grimes (1996) stated that Balizia and Hydrochorea closely resemble Albizia but are
distinguished by indeterminate inorescence-axes and vegetative branches arising from
Marcos Vinicius Batista Soares et al. / PhytoKeys 205: 401–437 (2022)
408
sylleptic and proleptic buds, pinnate leaet venation, and truncate ovaries. e authors
also highlighted the strong similarities in ower morphology shared between Abarema
s.l., Balizia and Hydrochorea.
Iganci et al. (2016) found the species of Hydrochorea and Balizia in a large un-
resolved polytomy together with Jupunba species. e Bayesian results of Soares et
al. (2021) are in line with the generic delimitation proposed here, with 1.0 posterior
probability supporting Jupunba, Hydrochorea and Balizia as monophyletic considering
matK sequences only, and a monophyletic Jupunba and paraphyletic Hydrochorea in
relation to Balizia when considering ETS sequences only. However, there was no boot-
strap support for a monophyletic Jupunba and neither for the clade uniting Hydrocho-
rea and Balizia. e results of Ringelberg et al. (2022) show that J. macradenia is more
closely related to Hydrochorea and Balizia than to other species of Jupunba, in contrast
to the phylogenetic position of the same accession in Soares et al. (2021), but in ac-
cordance with the phylogenomic results presented here. Notably, Soares et al. (2021)
included three accessions of J. macradenia, which were rmly nested in a monophyletic
Jupunba, suggesting that further phylogenomic analyses with more accessions included
will need to be carried out to further test the monophyly of Jupunba.
Phylogenetic evidence (Iganci et al. 2016; Koenen et al. 2020b; Soares et al. 2021;
Ringelberg et al. 2022; and a new analysis presented here) that shows Balizia to be para-
phyletic with respect to Hydrochorea, as well as the eld discovery of a new species that is
morphologically intermediate between the two genera, with crypto-lomentiform pods
that resemble more the follicle of Balizia pedicellaris (DC.) Barneby & J.W. Grimes
than the lomentiform pods of Hydrochorea, but a species of seasonally inundated forest
with hydrochorous seed dispersal, prompted us to decide that the two genera are best
combined. Furthermore, two African species that were formerly placed in several gen-
era, including Albizia and Cathormion, were shown to be closely related to Hydrochorea
based on phylogenomic evidence (Koenen et al. 2020b). ese species are therefore best
accommodated within Hydrochorea, as their general morphological features and ecol-
ogy are virtually indistinguishable from Neotropical species of Hydrochorea.
Koenen et al. (2020b) and Ringelberg et al. (2022) also show that Albizia sect.
Arthrosamanea is placed in the Jupunba clade, being more closely related to Jupunba,
Punjuba, Balizia and Hydrochorea than to Albizia s.s. ese results are reinforced by
Aviles Peraza et al. (2022) who proposed nomenclatural updates to solve this situation
based on more extensive sampling of Albizia sect. Arthrosamanea. In our study (Fig. 1),
we have still referred to these species as Albizia sect. Arthrosamanea but we refer the
reader to Aviles Peraza et al. (2022) for presentation of a new taxonomy of the group.
Based on parsimony analysis of morphological characters only, Barneby and Grimes
(1996) recognized pollen polyads comprising 16 grains as a synapomorphy for the Ab-
arema alliance, while other alliances that they recognized (e.g., the Samanea, Chloroleu-
con and Inga alliances) presented a polymorphic polyad number. Indeed, Guinet and
Grimes (1997) studied all Neotropical species of Hydrochorea and Balizia for their poly-
ads and we observed that the nested African species Cathormion obliquifoliolatum (De
Wild.) G.C.C. Gilbert & Boutique (Deighton 3618, K) and Cathormion rhombifolium
Recircumscription of Hydrochorea 409
(Benth.) Keay (Germain 87, K) also have 16-celled polyads. Polyad morphology was also
highlighted as diagnostic for the recognition of Afrocalliandra E.R. Souza & L.P. Quei-
roz, a genus segregated from Calliandra Benth. (Souza et al. 2013), and more attention
should be given to this character in future studies. Furthermore, the taxa in the Jupunba
clade share the simultaneous presence of vegetative and reproductive branches (sylleptic),
a character considered by Grimes (1999) as uncommon amongst the ingoid legumes.
us, possessing polyads with 16 grains and sylleptic branches could circumscribe ei-
ther the clade comprised of Hydrochorea, Jupunba and Punjuba, or a more conservative
Jupunba s.l. Forthcoming studies, including more samples of unstable taxa in current
molecular analyses, new eld collections and advances in phylogenomic analysis, will
hopefully resolve this question.
We did not include Hydrochorea acreana (J.F. Macbr.) Barneby & J.W. Grimes in our
synopsis and the name is here considered as incertae sedis. Pods from this species were not
known to Barneby and Grimes (1996), and herbarium specimens are dicult to iden-
tify when owers and especially fruits are unavailable. e type specimen (Kruko 5631,
NY334624) includes owers arranged in a large terminal panicle composed of umbel-
liform pseudoracemes of capitula, diering from the axillary to terminal inorescences
in Hydrochorea and Jupunba that are not paniculate, and always have sylleptic branches
present. Citing fruiting Central American collections, Hydrochorea acreana was combined
into Abarema s.l. by Rico-Arce (1999) as Abarema acreana (J.F. Macbr.) L. Rico. However,
the Central American collections cited by Rico-Arce (1999) were identied by Barneby
and Grimes as either Abarema macradenia (Pittier) Barneby & J.W. Grimes or Abarema ad-
enophora (Ducke) Barneby & J.W. Grimes, with which we agree. Interestingly, a specimen
from Acre which was collected in 1995 (Oliveira 691; NY00662831) and was presumably
not seen by Barneby and Grimes (1996), but was identied as A. acreana by L. Rico-Arce,
does include unripe pods and this material is likely conspecic with the type material of
H. acreana. However, as discussed before, fruit morphology often has been shown to be
rather misleading in mimosoid taxonomy due to homoplasy, and we point out the dif-
ferences in inorescence structure from Jupunba as discussed above. Soares et al. (2021)
resolved H. acreana as sister to Albizia subdimidiata (Splitg.) Barneby & J.W. Grimes, and
not closely related to Hydrochorea nor Jupunba, but these analyses were based on ETS se-
quences only and morphologically the material does not bear much resemblance to species
of Albizia sect. Arthrosamanea, in which Albizia subdimidiata is placed. Given the taxon’s
unusual combination of morphological characters, it may well represent an isolated lineage
that merits recognition as a distinct genus; this decision is pending further study.
Taxonomic treatment
Hydrochorea Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 23. 1996.
Figs 2, 3
Balizia Barneby & J.W. Grimes, syn. nov., Mem. New York Bot. Gard. 34(1). 23.
1996. Type: Balizia pedicellaris (DC.) Barneby & J.W. Grimes.
Marcos Vinicius Batista Soares et al. / PhytoKeys 205: 401–437 (2022)
410
Balizia sect. Leucosamanea Barneby & J.W. Grimes, syn. nov., Mem. New York Bot.
Gard. 34(1). 36. 1996. Type: Balizia leucocalyx (Britton & Rose) Barneby & J.W.
Grimes.
Balizia Barneby & J.W. Grimes sect. Balizia syn. nov., Mem. New York Bot. Gard. 34
(1). 37. 1996. Type: Balizia pedicellaris (DC.) Barneby & J.W. Grimes.
Type. Hydrochorea corymbosa (Rich.) Barneby & J.W. Grimes.
Description. Shrubs and trees, unarmed; branches grey to brown pilosulous
to glabrescent, cylindrical; stipules persistent or caducous. Leaves bipinnate, with
1–15 pairs of pinnae; petiole canaliculate or cylindrical, grey to brown pilosulous
or glabrous; nectaries sessile to stipitate, orbicular, patelliform, or cupuliform, the
rst either near mid-petiole or between the rst pinnae pair, and often along the leaf
rachis, between the leaet pairs; leaets 2–33 pairs per pinna, petiolate to subses-
sile, rhombic-ovate, rhombic-lanceolate, rhombic-oblong, rhombic-obovate, ovate,
elliptic, oblong, lanceolate or oblanceolate, grey to brown pilosulous, ciliate or gla-
brous, concolorous or more often discolorous, venation pinnate. Inorescence con-
sisting of umbelliform capitula or corymbiform racemes, arising singly or fasciculate
from the axils of coeval or hysteranthous leaves, bracts generally caducous; bracteoles
persistent or caducous. Flowers heteromorphic, pedicellate in peripheral owers,
mostly pentamerous, and sessile in the larger terminal ower, 5–8-merous; calyx
green, gamosepalous, campanulate, or tubular, pubescent, ciliate or glabrous; corolla
pinkish to reddish, yellowish or whitish, gamopetalous, infundibuliform, campanu-
late, or tubular, glabrous, puberulent, ciliate or pilose at the apex; androecium with
(10–)12–60(–75) stamens; laments white, greenish or roseate, fused into a tube,
included in peripheral owers or exserted beyond the corolla in the terminal ower;
stemonozone present, anthers dorsixed; ovary superior, sessile, truncate at the apex,
usually pubescent or sometimes glabrous. Fruits sessile or shortly stipitate, straight
or slightly recurved, either lomentiform, the seeds released in one-seeded articles, or
woody and indehiscent, the exocarp with transverse bres and the endocarp hard
and septate, or follicular, with similar exocarp but the septate endocarp papyraceous
and shed along with the seeds, or crypto-lomentiform with follicular dehiscence, the
exocarp smooth and the endocarp remaining attached to the seeds forming 1-seeded
articles. Seeds with a hard testa, with pleurogram complete or narrowly U-shaped.
Distribution and habitat. North America (Mexico), Central America (Belize, Costa
Rica, Guatemala, Honduras and Nicaragua), South America (Brazil, Bolivia, Colombia,
Ecuador, French Guiana, Guyana, Peru, Suriname and Venezuela) and Africa (Congo Ba-
sin and West Africa) (Fig. 2A). Hydrochorea species occur in riparian habitats, inundated
and non-inundated wet tropical forests of the Orinoco and Amazon basins, pre-Andean
Amazonia along the Nor-Yungas and Pando in Bolivia, Vaupés in Colombia and Huá-
nuco in Peru, Central Brazilian Savanna, the Atlantic Rainforest of Brazil and extending
to northern South America in Venezuela and the Guianas and the Gulf-Caribbean low-
lands until Mexico, and one species in coastal tidal swamp forests in Upper Guinea (West
Africa) and one species in riparian and seasonally inundated forests in the Congo Basin.
Recircumscription of Hydrochorea 411
Note. Since the names Hydrochorea and Balizia were published in the
same publication (Barneby and Grimes 1996), neither has priority, although
Hydrochorea was treated as genus 1 and Balizia as genus 2, Hydrochorea thus ap-
pearing rst in the publication. e name Hydrochorea is here chosen to represent
the recircumscribed genus, especially since the name is appropriate for most of its
species, and most Balizia species are also thought to frequently use water-borne
seed dispersal, and all but one species (B. elegans) are reported to often occur along
river-banks. e name Balizia, being an anagram of Albizia, is less appropriate
Figure 2. e genus Hydrochorea Barneby & J.W. Grimes A e amphi-atlantic geographic distribution
of Hydrochorea B Hydrochorea pedicellaris (DC.) M.V.B. Soares, Iganci & M.P. Morim foliage and fruits
C Hydrochorea corymbosa (Rich.) Barneby & J.W. Grimes foliage and fruits D Hydrochorea panurensis
(Spruce ex Benth.) M.V.B. Soares, M.P. Morim & Iganci foliage and fruits E Hydrochorea uaupensis M.P.
Morim, Iganci & E.J.M. Koenen in habitat, with foliage and fruits F Flowers of H. uaupensis after rain
G mature fruits of H. uaupensis. B, C from M.V.B Soares D from D. Cardoso E–G from J.R.V. Iganci.
Marcos Vinicius Batista Soares et al. / PhytoKeys 205: 401–437 (2022)
412
Figure 3. e genus Hydrochorea Barneby & J.W. Grimes (continued). Species from the Americas
A owering branch of Hydrochorea corymbosa (Rich.) Barneby & J.W. Grimes B close-up of inorescence
of H. corymbosa C discolorous leaves of H. corymbosa D close-up of inorescence of Hydrochorea panu-
rensis (Spruce ex Benth.) M.V.B. Soares, M.P. Morim & Iganci E unripe lomentiform pod of H. panu-
rensis F close-up of inorescence of Hydrochorea pedicellaris (DC.) M.V.B. Soares, Iganci & M.P. Morim,
with a few peripheral owers removed to expose sessile terminal owers G unripe pods of H. pedicellaris
H dehisced follicular pods of H. pedicellaris showing papery septate endocarp I detail of primary rachis
Recircumscription of Hydrochorea 413
given that several of its species have previously been placed in Albizia and therefore
the name may suggest close kinship, while actually being most closely related to
the genus Jupunba.
Identification key to the species of Hydrochorea
1a Species from Congo Basin and West Africa ................................................. 2
2a Adaxial leaet surface shiny, abaxial leaet surface glabrous, apart from the
ciliate midrib or with few scattered short hairs especially on and near the mid-
rib; calyx and corolla green to greenish white, corolla lobes glabrous or with a
few short white hairs around the apex, Congo Basin (Democratic Republic of
Congo, Central African Republic and Gabon) ............ 6. H. obliquifoliolata
2b Adaxial leaet surface dull, abaxial leaet surface pilose with varying density
of hairs (rarely nearly glabrous); calyx and corolla white, upper half of corolla
lobes rusty pilose to villous, West Africa (Senegal, Guinea-Bissau, Guinea,
and Sierra Leone) ............................................................. 9. H. rhombifolia
1b Species from North, Central and South America ........................................3
3a Pinnae 1–jugate on every leaf (seldom 2-jugate and then the true petiole very
short) .......................................................................................................... 4
4a Calyx covering the corolla in bud; owers glabrous, terminal ower with tu-
bular calyx ......................................................................... 7. H. panurensis
4b Calyx not covering the corolla in bud; owers puberulous, terminal ower
with campanulate calyx .......................................................5. H. marginata
3b Pinnae 2– or more jugate (seldom 1–jugate on some leaves of the same indi-
vidual) ........................................................................................................5
5a Leaets up to 10 pairs per pinna ................................................................. 6
6a Pinnae 1–2 jugate, leaets ovate to rhombic-ovate, corolla of peripheral owers
up to 1.5 mm long; follicular crypto-lomentiform fruit ....... 10. H. uaupensis
6b Pinnae (2–)3–6-jugate; leaets rhombic-oblong, rhombic-ovate or rhombic-
lanceolate; corolla of peripheral owers with more 1.5 mm long; fruit inde-
hiscent or lomentiform ...............................................................................7
7a Leaets rhombic-oblong; corolla of peripheral owers more than 7 mm long
........................................................................................... 4. H. leucocalyx
7b Leaets rhombic-ovate to rhombic-lanceolate; corolla of peripheral owers
up to 6 mm long ..................................................................1. H.corymbosa
of H. pedicellaris showing interpinnal extra-oral nectaries J inorescence of Hydrochorea uaupensis M.P.
Morim, Iganci & E.J.M. Koenen showing large sessile central ower and pedicellate peripheral owers
K unripe crypto-lomentiform pod and seed enveloped by septate endocarp of H. uaupensis; African spe-
cies L inorescence of Hydrochorea obliquifoliolata (De Wild.) E.J.M. Koenen M pinnae of Hydrochorea
rhombifolia (Benth.) E.J.M. Koenen showing rhombic leaets. A–E, J, K Erik Koenen F-I Colin Hughes
L Jan Wieringa M William Hawthorne. Vouchers A–C J.R.V. Iganci 862 D, E M.P. Morim 563 F–I L.P.
Queiroz 15529 J, K M.P. Morim 577 L J.J. Wieringa 6519 M unvouchered.
Marcos Vinicius Batista Soares et al. / PhytoKeys 205: 401–437 (2022)
414
5b Leaets in more than 10 pairs per pinna ..................................................... 8
8a Corolla of peripheral owers 8–10 mm long, fruit indehiscent, not lomenti-
form ........................................................................................ 2. H. elegans
8b Corolla of peripheral owers up to 7.5 mm long, fruit lomentiform or fol-
licular .........................................................................................................9
9a Pinnae 3–5-jugate; fruit lomentiform ................................ 3. H. gonggrijpii
9b Pinnae 6–17-jugate; fruit follicular, with septate endocarp and transverse b-
ers in the exocarp .............................................................. 8. H. pedicellaris
1. Hydrochorea corymbosa (Rich.) Barneby & J.W. Grimes, New York Bot. Gard.
74(1): 27. 1996.
Fig. 2A, C, 3A-C, 4
Pithecellobium subcorymbosum Hoehne [as Pithecolobium], Comiss. Linhas Telegr. Es-
tratég., Mato Grosso-Amazonas, Bot. 8: 18, Ic. 133. 1919. Type: B, Mato
Grosso, São Luiz de Cáceres, nas margens do rio Paraguai, perto da Campina, Hoeh-
ne 4582 (lectotype, designated here from amongst the syntypes: R! [R000003169];
isolectotype: SP).
Basionym. Mimosa corymbosa Rich., Actes Soc. d’Hist. Nat. Paris 1: 113. 1792.
Type material. French Guiana, frequens in sylvis ripariis uvii Kourou, Louis
Claude Richard s.n. (lectotype, designated by Barneby and Grimes 1996, p. 27, as
holotype, here corrected: P [P02142909] digital image!).
Distribution and habitat. Bolivia, Brazil, Colombia, Ecuador, French Guiana,
Guyana, Paraguay, Peru, Venezuela. Hydrochorea corymbosa occurs in periodically
or permanently inundated riparian forest, gallery forest, and open vegetation, up to
480m elevation (Barneby and Grimes 1996).
Notes. Hydrochorea corymbosa is morphologically similar to H. gonggrijpii by
its leaves with (2–)3–6 pairs of pinnae (3–5 pairs of pinnae in H. gonggrijpii),
but diers by presenting (4–)5–11(–14) leaet pairs per pinnae (vs. (12–)14–35
in H. gonggrijpii). Hydrochorea corymbosa has a wide distribution in the Brazilian
Amazon, and displays wide morphological plasticity. Barneby and Grimes (1996)
recognised the specimen Louis Claude Richard s.n. (P02142909) as holotype, al-
though the species protologue did not present a type specimen. e specimen
does represent original material that the author associated with the taxon, being a
specimen collected by the author and annotated as Mimosa corymbosa. us, Louis
Claude Richard s.n. (P02142909) is here corrected to lectotype (Art. 9.3, 9.4, 9.8
and 9.10; Turland et al. 2018).
Selected specimens examined. B, Amazonas: São Gabriel da Cachoeira,
entre Assunção do Içana e Camarão, mato de Igapó, margem do rio, 10 July 2012,
J.R.V. Iganci 862 (RB). B, Pando: Federico Roman, bordo del Río Abuna, 18
November 2006, S. Altamirano & H. Ramos 4293 (K). C, Vaupés: Mitú
Recircumscription of Hydrochorea 415
Figure 4. Hydrochorea corymbosa (Rich.) Barneby & J.W. Grimes A branch with inorescences
B–D extra-oral nectaries E peripheral ower bud F peripheral ower G terminal ower bud H terminal
ower I fruit J seed. A, E–H from M.V.B. Soares 75 B–D from M.V.B. Soares 180 I–J from M.V.B. Soares
174. Illustration by Alex Pinheiro.
Marcos Vinicius Batista Soares et al. / PhytoKeys 205: 401–437 (2022)
416
and Vicinity, lower rio Kubiyú, 26 September 1976, Zarucchi 2147 (INPA). E-
, Francisco de Orellana: Estación Cientíca Yasuní, Río Tiputini, este de la Car-
retera Repsol-YPF, km 7 desvío hacia el pozo Tivacuno, Laguna Herradura, 20 April
1999, G. Villa 177 (K). G: Potaro-Siparuni, riparian zone lower Kuribrong,
April 2010, Zartman et al. 8002 (INPA). P, Loreto: Jenaro Herrena, Cano Su-
pay, ooded forest along cano, 23 May 2002, T.D. Pennington et al. 17430 (K).
V, Amazonas: Departamento Rio Negro, middle part of the Río Baria, 21
July 1984, G. Davidse 27570 (K).
2. Hydrochorea elegans (Ducke) M.V.B. Soares, Iganci & M.P. Morim, comb. nov.
urn:lsid:ipni.org:names:77303827-1
Fig. 2A
Balizia elegans (Ducke) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 40
1996. Albizia elegans (Ducke) L. Rico, Novon, 9(4): 556. 1999.
Albizia duckeana L. Rico, syn. nov., Kew Bull. 55(2): 404. 2000. Type: based on Pithe-
cellobium elegans Ducke.
Basionym. Pithecellobium elegans Ducke [as Pithecolobium], Arch. Jard. Bot. Rio de
Janeiro 3: 64. 1922.
Type material. B, in silvis non inundatis, prope Alcobaca (Tocantins),
A. Ducke 16271 (lectotype, designated by Barneby and Grimes 1996, p. 40: MG
[MG00016271], digital image!; isolectotypes: G [G00359898] digital image!, MG,
P [P03093819] digital image!, R [R000002384] digital image!, RB [RB10177]!, US
[US1040853] digital image!, US [US00000336] digital image!, US [US00610722]
digital image!).
Distribution and habitat. Bolivia, Brazil, Costa Rica, Ecuador, Honduras, Nica-
ragua, Peru. Hydrochorea elegans occurs in primary rain forest, up to 350 m elevation
(Barneby and Grimes 1996).
Notes. Hydrochorea elegans has a morphological anity with H. pedicellaris, as al-
ready pointed out by Ducke (1922) and by Barneby and Grimes (1996). However, the
corolla of peripheral owers is larger (8–10 mm long) in H. elegans than in H. pedicel-
laris (up to 7.5 mm long). Ducke (1922) and Barneby and Grimes (1996) also com-
mented on the similarity between the fruit of both species, but the fruits of H. elegans
are indehiscent (vs. follicular dehiscence in H. pedicellaris). Hydrochorea elegans has a
disjunct distribution between hylaean Brazil and Costa Rica and Nicaragua.
Selected specimens examined. B, Rondônia: Porto Velho, área do Res-
ervatório da Usina Hidrelétrica de Samuel, 15 June 1986, C.A.C. Ferreira 7458 (K).
C R, Limón: Talamanca, Fila Carbon, Finca de Pedro Bolivar, 25 May 1999,
O. Valerde 1175 (K).
Recircumscription of Hydrochorea 417
Figure 5. Hydrochorea gonggrijpii (Kleinhoonte) Barneby & J.W. Grimes A branch with inorescences
B, C extra-oral nectaries D peripheral ower bud E peripheral ower F terminal ower bud G terminal
ower H leaet. A–H from Fróes 28045. Illustration by Alex Pinheiro.
Marcos Vinicius Batista Soares et al. / PhytoKeys 205: 401–437 (2022)
418
3. Hydrochorea gonggrijpii (Kleinhoonte) Barneby & J.W. Grimes, Mem. New
York Bot. Gard. 74 (1): 25. 1996.
Figs 2A, 5
Basionym. Pithecellobium [as Pithecolobium] gonggrijpii Kleinhoonte Recueil Trav.
Bot. Néerl. 22: 414. 1926.
Type material. S, im Reservat der Zanderij I, die nummerierten Baume
n. 102 (Herb. [Acad.Rhenotraiect.J n. 1529, im Dez. 1915, und n. 4350bl. im Juli
1919) und n. 141 (Herb. n. 4357, bl. Im Juli 1919.)” 141, 10/VII/1919”, Forest Bu-
reau 4357 (lectotype, designated here from amongst the syntypes: IAN [IAN49436]!;
isolectotypes: A [A00064017] digital image!, BR [BR0000005170067] digital im-
age!, K [K000527996]!, K [K000527995]!, MO [MO954361] digital image!, NY
[NY00334660] digital image!, NY [NY00334661] digital image!, NY [NY00334662]
digital image!, P [P01818508] digital image!, U [U U0003385] digital image!, U
[U0003384] digital image!, US [US00629380] digital image!).
Distribution and habitat. Brazil, Colombia, Guyana, French Guiana, Suriname,
Venezuela. Hydrochorea gonggrijpii occurs along riverbanks, gallery forest margins, and
low-lying swamp forests, at 40–1400 m elevation (Barneby and Grimes 1996).
Notes. In the nomenclatural treatment of H. gonggrijpii Barneby and Grimes
(1996: p. 25) maintained the specimens “Surinam: im Reservat der Zanderij I, die
nummerierten Baume n. 102 (Herb. [Acad. Rhenotraiect. J n. 1529, im Dec. 1915,
und n. 4350 bl. im Juli 1919) und n. 141 (Herb. n. 4357, bl. im Juli 1919.)” as syn-
types. In the present work, the specimen Forest Bureau 4357 (IAN49436) is designated
as lectotype (Art. 9.3, Turland et al. 2018).
Selected specimens examined. B, Amazonas: Presidente Figueiredo, Cachoeira
do boto, 21 September 2007, Carvalho-Sobrinho et al 1632 (RB). C, Vaupés:
Mitú and vicinity, lower Río Kubiyú, along river, 26 September 1976, J.L. Zarucchi s.n.
(K). S. Plantas de Tafelberg (Table Mountain), 10 August 1944, Maguire 24273
(RB). V, Bolivar: Distrito Piar, gallery forest bordering savana, vicinity of Guad-
equen (Buadequen), Río Acanán (auent of Río Carrao), Cerros Los Hermanos, 20 May
1986, Lat 5°26'N, Long 62°17'W, alt 470 meters, J.A. Steyermark et al. 131865 (NY).
4. Hydrochorea leucocalyx (Britton & Rose) Iganci, M.V.B. Soares & M.P. Morim,
comb. nov.
urn:lsid:ipni.org:names:77303828-1
Fig. 2A
Balizia leucocalyx (Britton & Rose) Barneby & J.W. Grimes, in Mem. New York Bot.
Gard. 74(1): 85. 1996.
Basionym. Samanea leucocalyx Britton & Rose, N. Amer. Fl. 23: 34. 1928.
Recircumscription of Hydrochorea 419
Type material. M. Tabasco, El Limon, J. N. Rovirosa 976 (lectotype, designat-
ed by Barneby and Grimes 1996, p. 36, as holotype, here corrected: US [US13198371]
digital image!, clastotypus (fragm. + photo): NY [NY00003824] digital image!).
Distribution and habitat. Belize, Guatemala, Honduras, Mexico. Hydrochorea
leucocalyx occurs in wet tropical forests, often along riverbanks, seldom in anthropo-
genic pastures, up to 400 m elevation (Barneby and Grimes 1996).
Notes. Amongst the species of Hydrochorea, H. leucocalyx is one of the few that does
not occur in Amazonia. It has anities with the new species described in this treat-
ment (see H. uaupensis) and is mainly distinguished by the lomentiform indehiscent
fruit (vs. the cryptoloment in H. uaupensis). Barneby and Grimes (1996) recognised
the specimen J. N. Rovirosa 976 as holotype, but in the species protologue (Britton and
Rose 1928), the authors did not indicate the herbarium where the type specimen was
deposited. us, following Art. 9.10 of the International Code of Botanical Nomen-
clature (Turland et al. 2018), the specimen J. N. Rovirosa 976 (US13198371) is here
corrected to lectotype.
Selected specimens examined. H: 7 September 1932, W.S. Schipp 1024
(K). M, Chiapas: km 12 carretera Pénjamo-Chancalá, 8 June 1960, J.P. Chavelas
et al. s.n. (K).
5. Hydrochorea marginata (Spruce ex Benth.) Barneby & J.W. Grimes, Mem. New
York Bot. Gard. 74(1): 29. 1996.
Figs 2A, D, 6
Basionym. Pithecellobium [as Pithecolobium] marginatum Spruce ex Benth., Trans.
Linn. Soc. London 30: 586. 1875.
Type material. B, Barra, by a stream [Prov. Rio Negro], Spruce 1658 (lecto-
type, designated by Barneby and Grimes 1996, p. 31: K [K000528011]!; isolectotypes:
E [E00313848] digital image!, F [V0058733F] digital image!], G [G275450] digital
image!, P [P03094432] digital image!, P [P03094430] digital image!).
Distribution and habitat. Brazil and Venezuela. Hydrochorea marginata occurs in
Amazonia, in ooded areas and along riverbanks and lake shores.
Notes. Barneby and Grimes (1996) considered H. marginata to comprise three
varieties, H. marginata var. panurensis (Benth.) Barneby & J.W. Grimes, H. mar-
ginata var. scheryi Barneby & J.W. Grimes, and Hydrochorea marginata var. mar-
ginata. Hydrochorea marginata var. panurensis is recognized at the species level in
this treatment, and H. marginata var. scheryi is placed as a synonym of H. panu-
rensis because we identied no morphological diagnostic characters that support
them as independent taxa. Both these taxonomic decisions are discussed under
H. panurenis.
Selected specimens examined. B: Amazonas, Rio Negro between Moreira
and Rio Arirahá, 13 October 1971, G.T. Prance 15206 (NY).
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420
6. Hydrochorea obliquifoliolata (De Wild.) E.J.M. Koenen, comb.nov.
urn:lsid:ipni.org:names:77303829-1
Figs 2A, 3L
Pithecellobium obliquifoliolatum (De Wild.) J. Léonard, in Compt. Rend. Sem. Agric.
Yangambi Comm. No. 67, 868 (1947).
Pithecellobium obliquifoliolatum (De Wild.) Aubrév., Fl. Forest. Soudano-Guin. 290
(1950), in obs., Aubrev. in Not. Syst., ed. Humbert, xiv. 57 (1950) nom. illeg.;
Arthrosamanea obliquifoliolata (De Wild.) G.C.C. Gilbert & Boutique, Fl. Congo
Belge & Ruanda-Urundi iii. 194 (1952).
Cathormion obliquifoliolatum (De Wild.) G.C.C. Gilbert & Boutique, Bull. Soc. Roy.
Bot. Belgique 90: 309 (1958).
Basionym. Albizia obliquifoliolata De Wild., Bull. Jard. Bot. État Bruxelles 7: 253 (1920).
Type material. D R C, Congo Belge, Eala, Lau-
rent 1823 (lectotype, designated here: BR [BR0000008916334]!; isolectotype: BR
[BR0000008916662]!).
Description. Trees up to 30 m in height and up to 1m DBH, the bark with both small
scattered and long transverse linear lenticels, the indumentum consisting of a dense rusty
to golden-brown pubescence covering the young twigs, petiole and primary rachis, with
more sparse pubescence on peduncles and pinna-rachises except for dense rows of hairs at
the margins of the otherwise glabrous canaliculate adaxial side of the pinna rachises, often
also the canaliculate primary rachis of the leaf sparsely pubescent to glabrous adaxially.
Stipules linear deltoid to falcate, 2–3 mm long, adaxially glabrous except at apex, densely
pubescent, caducous. Leaves with (1–)2(–3) pairs of pinnae, petiole pulvinate and slightly
attened at base, (1.5–)2–3.5(–4.5) cm long, with a sessile concave circular to triangular
nectary at the apex, c. 0.8–1.5 mm in diameter, rachis usually canaliculate adaxially, (0–
)1.5–3(–6) cm long, if the leaf 3-jugate then usually with an inter-pinnal nectary, similar
to the petiolar one, in between the middle pair of pinnae, apical nectary usually lacking,
pinnae distinctly pulvinate, and usually with minute paraphyllidia at the apex of the pul-
vini, pinna-rachises canaliculate adaxially, the groove glabrous, c. (3–)4–11(–15) cm long,
with short stipitate circular to elliptical cupular or trumpet-shaped nectaries c. (0.2–)0.5–
1mm in diameter. Leaets in (3–)5–7(–8) pairs per pinna, subsessile on a c. 0.5 mm long
pulvinule, widely spaced so that the margins do not overlap, bicolorous, dark green and
shiny above, pale dull green beneath, rhomboid and often distinctly curved towards pinna
apices, base asymmetrically obtuse or slightly oblique and the apex rounded or shallowly
emarginate, sometimes mucronate, (1.4–)2.2–3.5(–4.7) × (0.6–)1.1–1.8(–2.2)cm, except
the apical pair that is asymmetrically elliptic with oblique base and emarginate apex, (1.8–
)2.7–4.8(–5.5) × (1.0–)1.5–2.5 cm, venation pinnate with (6–)12–16(–18) secondary
veins brochidodromous, prominent on both surfaces or prominulous to slightly sunken on
upper surface, and reticulate tertiary venation, often prominent on upper surface, obscure
beneath, margins and midrib ciliate on both surfaces, lamina glabrous but for a few short
scattered appressed hairs. Inorescences (10–)15–20 owered umbelliform capitula, on
long slender peduncles arising 1–2 from axillary buds of coeval or caducous leaves, held
Recircumscription of Hydrochorea 421
Figure 6. Hydrochorea marginata (Spruce ex Benth.) Barneby & J.W. Grimes A branch with inores-
cences B–D extra-oral nectaries E peripheral ower bud F peripheral ower G terminal ower bud
H terminal ower I leaet. A– D, F, H, I from A. Carlos et al. 066 E, G from C. Ferreira et al. 7260. Il-
lustration by Alex Pinheiro.
Marcos Vinicius Batista Soares et al. / PhytoKeys 205: 401–437 (2022)
422
above the foliage, the axillary meristems usually not continuous beyond the peduncles and
aborted prior to fruit set, dimorphic with a single enlarged terminal ower and often one
dispositioned peripheral ower c. 0.5 cm below the others, on peduncles 4–8(–12) cm
long. Bracts linear to spatulate, sometimes bilobed at apex, c. 2–3.5 × 0.5mm, pubescent
with longer hairs at apex. Peripheral owers on pedicels 2–6 mm long, calyx pentamerous,
green to greenish white, c. 3–4 mm long, the deltoid lobes c. 0.5 × 0.5 mm, glabrous,
corolla pentamerous, green to greenish white, c. 5–6 mm long, the lobes c. 2–3 × 2 mm,
glabrous or with short white hairs around the apices of the lobes, androecium consisting
of c. 20–30 stamens, c. 2.1–2.5 cm long, the laments white to pale green at apex, fused
into a tube c. 3 mm long, with dorsixed pale yellowish green anthers, pollen in 16-celled
plano-compressed disc-shaped polyads, pistil c. 2.5–2.8 cm long, ovary c. 3 mm long,
pubescent in upper half, the pale green to white style emerging from it at an angle of c.
45°, with a green funnel-shaped stigma, extending beyond the stamens. Terminal ower
sessile to subsessile, similar to peripheral owers but broadly campanulate and larger, calyx
c. 3.5–5 mm and corolla c. 7.5–9 mm long, the laments thicker and staminal tube c.
8–10mm long, exserted well beyond the corolla tube. Pods falcate and weakly articu-
lated, base often tapering into a c. 5 mm long stipe, (3–)6–12 seeded with a thin papery
fruit wall and slightly thickened rim, dark brown to black outside when ripe, light brown
inside, (3.7–)5.5–9.6 × 1.2–1.4cm, breaking up into 1-seeded articles 0.4–0.7(–1.0) cm
long, the basal and apical articles up to 1 cm long, seed c. 6.5 × 4.5 × 0.5 mm, the testa
hard, light brown with a darker brown closed elliptic pleurogram, c. 4 × 2 mm.
Distribution and habitat. Gabon, Central African Republic, Congo-Brazzaville,
Democratic Republic of the Congo. Hydrochorea obliquifoliolata occurs in the Congo
Basin, and is a species of swamp forests, seasonally inundated forests and riverbanks.
Notes. e similarities to Cathormion rhombifolium, the other African species that
is here transferred to Hydrochorea, are discussed below.
Selected specimens examined. G: Ogooué-Lolo, road Okondja to Bambidie
and Lastoursville, 21 km SW of Okondja, 7 February 2008, J.J. Wieringa 6519 (BR, K).
D R C: Yafunda, rive guache, près d’Isangi, 8 September
1938, J. Louis 11175 (BR). Boendu, August 1938, Du Bois 904 (BR), G. Couteaux 55
(BR). Bolomba, 7 November 1957, C. Évrard 2746 (BR). Bongoy, 4 January 1958, C.
Évrard 3191 (BR). Botsima, route station-village, 28 January 1991, J.B.M.M. Dhetch-
uvi 321 (BR). Yangambi, île Tutuku en face du plateau de l’Isalowe, 3 January 1940,
R.G.A. Germain 87 (BR). Bokondji, 28 September 1959, De Wanckel 162 (BR).
7. Hydrochorea panurensis (Spruce ex Benth.) M.V.B. Soares, M.P. Morim & Iganci,
comb. nov.
urn:lsid:ipni.org:names:77303830-1
Figs 2A, 7
Hydrochorea marginata var. panurensis (Benth.) Barneby & J.W. Grimes, Mem. New York Bot.
Gard. 74: 32. 1996. Type: based on Pithecellobium panurense Spruce ex Benth., syn. nov.
Recircumscription of Hydrochorea 423
Figure 7. Hydrochorea panurensis (Spruce ex Benth.) M.V.B. Soares, M.P. Morim & Iganci A branch
with inorescences B, C extra-oral nectaries D peripheral ower E peripheral ower bud F terminal
ower bud G terminal ower H terminal ower longitudinal section I leaet. A–I from Wurdack & Ad-
derley 43618. Illustration by Alex Pinheiro.
Marcos Vinicius Batista Soares et al. / PhytoKeys 205: 401–437 (2022)
424
Hydrochorea marginata var. scheryi Barneby & J.W. Grimes, Mem. New York Bot.
Gard. 74: 32. 1996. Type: V, at Sanariapo, Territorio Federal Amazonas,
Llewellyn Williams 15953 (lectotype rst step, designated by Barneby and Grimes
1996, p. 32, as holotype, here corrected: F; lectotype second step, here designat-
ed: F [V0058706F], digital image!; isotypes: F [V0058707F] digital image!), G
[G00365761] digital image!, GH [GH00060404] digital image! K [K0005279]
digital image!, MO [MO954357] digital image!, MO [MO954358] digital im-
age!), NY [NY00334620] digital image! US [US00000401] digital image!, US
[US00385625] digital image!, VEN [VEN2237] digital image!, syn. nov.
Basionym. Pithecellobium [as Pithecolobium] panurense Spruce ex Benth., Trans. Linn.
Soc. London 30: 586. 1875.
Type material. B, in silvis ‘Gapó’ ad umen Uaupés prope Panuré, prov. do
Alto Amazonas, Spruce 2425 (lectotype rst step, designated by Barneby and Grimes
1996, p. 32, as holotype, here corrected: K; lectotype second step, here designated:
K [K000528008]!; isolectotypes: E [E00313845] digital image!, F, [V0058739F]
digital image!, G [G00365356] digital image!, G [G00365687] digital image!, GH
[GH00063965] digital image!, K [K000528007]!, NY [NY334690], P [P03094382]
digital image!, P [P03094383] digital image!, RB [RB00708599]!).
Distribution and habitat. Brazil, Venezuela. Hydrochorea panurensis occurs in season-
ally ooded Amazonian sites along rocky stream banks and ecotone with gallery forests.
Notes. Ducke (1949) considered Pithecellobium panurense as a form of P. marginatum,
considering the vegetative characters to be more important taxonomically than the dier-
ences between the owers of those taxa. Barneby and Grimes (1996) moved P. panurense to
Hydrochorea as H. marginata var. panurensis. However, Hydrochorea panurense has diagnos-
tic oral characters which distinguish it from H. marginata. e oral buds of H. panurense
have the corolla covered by the calyx; the owers are glabrous and the terminal ower has
a tubular calyx (vs. calyx not covering the corolla in the oral buds, pubescent owers, and
calyx of the terminal ower campanulate in H. marginata). Barneby and Grimes (1996)
also described H. marginata var. scheryi, distinguishing it from H. marginata var. panuren-
sis only by the pedicel of the peripheral owers, this 4–5 mm long. (vs. 6.5–13 mm long in
H. marginata var. panurensis). Barneby and Grimes (1996) emphasized, however, that this
is the only character to distinguish between the two taxa. In the present study we consider
H. marginata var. scheryi to be a synonym of H. panurensis.
A second step lectotypication is designated here for both H. panurensis and
H. marginata var. scheryi, since the material that Barneby and Grimes (1996) recog-
nised as holotypes consists of two sheets in both cases and they did not select lectotypes
from amongst these sheets (Art. 9.10,9.17; Turland et al. 2018).
Selected specimens examined. B, Amazonas: Barcelos, Serra do Araçá, Rio
Araçá à 13 h de Barcelos, 28 July 1985, Silva 389 (INPA); São Gabriel da Cachoeira.
Margem do Rio Içana em direção a comunidade Camarão, 0°48'35.8"N, 67°32'10"W,
19 July 2012, Morim, M.P., Iganci, J.R.V, Bonadeu F & Koenen, E. 563 (RB, Z). V-
, Amazonas: Rio Casiquiare, 11 November 1959, Wurdack & Adderley 43407 (IAN).
Recircumscription of Hydrochorea 425
8. Hydrochorea pedicellaris (DC.) M.V.B. Soares, Iganci & M.P. Morim, comb.
nov.
urn:lsid:ipni.org:names:77303831-1
Figs 2A, B, 3F-I
Balizia pedicellaris (DC.) Barneby & J.W. Grimes, in Mem. New York Bot. Gard.
74(1): 85. 1996.
Albizia pedicellaris (DC.) L. Rico, in Novon 9(4): 555. 1999.
Basionym. Inga pedicellaris DC., Prodr. 2: 441. 1825.
Type material. F G, “...in Cayenna” (lectotype, designated by
Barneby and Grimes 1996, p. 36, as holotype, here corrected: G-DC = F Neg. 6972,
digital image!).
Distribution and habitat. Bolivia, Brazil, Colombia, Ecuador, French Guiana,
Guyana, Peru, Suriname, Venezuela. Hydrochorea pedicellaris occurs in non-inundated
primary rainforest in Amazonia, in the lowlands of the Atlantic Rainforest, and in gal-
lery forests, up to 200 m elevation, and occasionally at 700–800 m elevation in Bolivia,
Ecuador and eastern Brazil (Barneby and Grimes 1996).
Notes. Hydrochorea pedicellaris is the only species of the genus that occurs in a
range of environments including areas of the Brazilian Atlantic Forest. It has an an-
ity with H. elegans (see comment under that species), but when it is in fruit it is easily
recognized by its follicular dehiscence, and an exocarp with deep, transversal ssures.
Barneby and Grimes (1996) recognised a specimen in the G-DC herbarium as holo-
type, but in the species protologue (De Candolle 1825), the author did not indicate
a type specimen. us, the specimen at G-DC should be considered a lectotype, here
corrected (Art. 9.10; Turland et al. 2018).
Selected specimens examined. B, La Paz: Province of Larecaja, Tuiri, 12
September 1989, B. Kruko 10886 (K). B, Amazonas: São Gabriel da Cachoeira,
Rio Içana, na comunidade Camarão, terra rme, 0°37'23"N, 67°26'57"W, 20 July
2012, Iganci, J.R.V, Morim, M.P., Bonadeu F., Koenen, E. 870 (RB); Espírito Santo:
Linhares Fragmento em frente a casa do Reis, Sítio Santo Domingo, Restinga arbórea
de cordões arenosos, 19°21'6"S, 39°43'31"W, 13 March 2007, R.D. Ribeiro et al. 812
(RB). G: Territorio Federal Delta Amacuro, 29 May 1964, L.M. Berti 225 (K).
P: Palcazú, Pasco Oxapampa, localidad Mayro, 20 May 2010, R. Vásquez et al.
36546 (K). S: Zenderij, November 1944, M. Koeleroe 237 (RB). V:
Altiplanicie de Nuria, 15 July 1960, J.A. Steyermark 86335 (K).
9. Hydrochorea rhombifolia (Benth.) E.J.M. Koenen, comb. nov.
urn:lsid:ipni.org:names:77303832-1
Figs 1A, 3M
Feuilleea rhombifolia (Benth.) Kuntze, Revis. Gen. Pl. 1: 189 (1891).
Marcos Vinicius Batista Soares et al. / PhytoKeys 205: 401–437 (2022)
426
Cathormion rhombifolium (Benth.) Keay, Kew Bull. 8(4): 489 (1953).
Basionym. Albizia rhombifolia Benth., London J. Bot. 3: 87 (1844).
Type material. G, Conakry, Heudelot 735 (lectotype designated here
from amongst the syntypes: K [K000043955]!; isolectotypes: K [K000043954]!,
K [K000043949]!, P [P00418271] digital image!, P [P00418272] digital image!, P
[P00418270] digital image!).
Description. Trees or shrubs up to 12 m tall, the young stems, all leaf-axes and
peduncles puberulent-tomentulose with rusty brown hairs. Stipules deltoid, c. 1 mm
long, puberulent-tomentulose, caducous. Leaves with 2–3 pairs of pinnae, petiole
pulvinate, ventrally attened above pulvinule and with central groove in upper half,
2–3.5(–8.5) cm long, rachis ventrally grooved, 1.5–4(–12.5) cm long, pinna rachises
pulvinate, ventrally grooved, (3.2–)4–6(–12) cm long. Nectaries present at the petiole
apex just below the rst pair of pinnae as well as just below each further pair of pinnae,
sessile or shortly stipitate on stipe to 0.5 mm, cupular or sometimes concave, circular
and 0.8–2.2 mm in diameter, and between the upper 2–3 pairs of leaets, trumpet-
shaped and then on a short stipe 0.5 mm or cupular and (sub)sessile, the lower ones
circular and the upper ones elliptical, 0.8–1.5 × 0.8–1.1 mm. Minute paraphyllidia
sometimes present at the apex of the pinna-pulvinus. Leaets in 4–6 pairs per pinna,
closely spaced, bicoloured leaets often with partly overlapping margins, bright green
above and pale green beneath, dull on both surfaces, rhomboid with a pulvinate sessile
oblique base and rounded to slightly emarginate apex, increasing in size towards pinna
apex, (1.1–)1.7–3.5(–5.1) × (0.5–)1.2–1.8(–2.3) cm, except for the apical pair which
has a less oblique to nearly acute base, (2.1–)2.5–4.5(–5.7) × (1.1–)1.5–2.5(–3.2) cm;
venation pinnate with 8–12(–18) secondary veins brochidodromous, tertiary vena-
tion reticulate, prominulous on both surfaces, midribs ciliate on both sides, the lower
leaet surface pilose with a variable density of brownish to white hairs, rarely almost
glabrous, sometimes villose particularly near the midrib giving a rusty orange-brown
appearance. Inorescences umbelliform capitula, axillary to co-eval leaves on pedun-
cles (4.5–)5–9.5 cm long, dimorphic with 6–16 peripheral owers and 1–2 terminal
ower(s) with elongated exserted staminal tubes. Bracts spatulate, c. 1.8 mm long, pu-
berulent with minute rusty hairs, caducous. Peripheral owers on pedicels of 1–4 mm,
calyx pentamerous, white, 3–3.5 mm long, fused, the deltoid lobes 1–1.3 mm long,
glabrous or with few minute hairs, corolla pentamerous, white, 6–8 mm long, fused
in the lower half, glabrous, pilose to villose in the upper half, androecium 1.6–2.3 cm
long, consisting of 20–28 stamens with white laments fused at the base into a short
tube of c. 2 mm, anthers dorsixed, pollen in 16-celled plano-compressed disc-shaped
polyads, gynoecium with a c. 2 mm long ovary, pubescent on the upper half, the 1.6–
2.5 cm long white style emerging from it at an angle of c. 45°, with a funnel-shaped
stigma, extending beyond the stamens. Terminal ower(s) similar but larger and more
robust in appearance, calyx c. 4.5 mm long with c. 1.5 mm long lobes, corolla c. 9 mm
long, androecium with 30–36 stamens that are thicker and fused into a tube 7–10 mm
long, exserted well beyond the corolla tube, and with a sunken nectariferous disk be-
Recircumscription of Hydrochorea 427
low the base of the ovary, gynoecium otherwise similar to that of the peripheral ow-
ers. Pods straight to falcate, 6–12-seeded with a thin papery fruit wall and thickened
rim, dark brown outside when ripe, whitish grey inside, (4.5–)7–12.5 × 1.4–1.9 cm,
breaking up into 1-seeded articles 0.6–1.1 cm long, seed c. 7 × 4.5 × 2 mm, the testa
hard, light brown with a wide lighter brown closed pleurogram.
Distribution and habitat. Known from the tidal riverine systems near the coast from
Senegal to Sierra Leone. Hydrochorea rhombifolia occurs often abundantly, in permanent
or tidal swamp forest, including on the edge of mangrove swamps, and in gallery forests.
Notes. Bentham (1844) described Albizia rhombifolia, before designation of holo-
types was required by the International Code of Botanical Nomenclature. Keay (1953)
made the new combination Cathormion rhombifolium and cited the holotype as being
at Kew. However, there are three specimens of Heudelot 735 at K, the type that was
cited by Bentham, leaving it ambiguous as to which one of these represents the holo-
type. erefore, the specimen from Herbarium Benthamianum (the oldest deposited
specimen dating to 1854) is here designated as a lectotype: it has leaves and owers,
and is more richly annotated than the other two specimens.
Hydrochorea obliquifoliolata and H. rhombifolia are morphologically very similar
and have sometimes been confused in herbaria, despite their clearly dierent geo-
graphical distributions. e species are readily separated by the darker appearance of
the leaets of H. obliquifoliolata, which have a distinct shine on the upper surface and
the lower surface usually (sub-)glabrous (vs. a usually rusty pilose lower leaet surface
in H. rhombifolia). e leaets of H. rhombifolia are also more closely spaced than
those of H. obliquifoliolata, the latter not having overlapping margins. Furthermore,
the ower colour of the two species is clearly dierent (as per the key), a characteristic
which remains apparent when comparing dried owering specimens in the herbarium,
and the corolla lobes of H. obliquifoliolata are glabrous or with a few short apical hairs
(vs. pilose to villous on the upper half in H. rhombifolia).
Selected specimens examined. S L: Mange, 7 February 1939, F.C.
Deighton 3618 (K), Rokupr, 25 May 1953, F.C. Deighton 5925 (K), Kasanko (Ma-
fore), 3 December 1950, T.S. Jones 52 (K), near Tassin and Kukum, 17 January 1892,
G.F. Scott Elliot 4418 (K); G-B: Gabu, Ponte do rio Colufe, 10 June 1949,
Espirito Santo 2500 (K).
10. Hydrochorea uaupensis M.P. Morim, Iganci & E.J.M. Koenen, sp. nov.
urn:lsid:ipni.org:names:77303833-1
Figs 2A, E-G, 3J, K, 8
Diagnosis. Hydrochorea uaupensis is morphologically similar in appearance to H. leu-
cocalyx (Britton & Rose) Iganci, M.V.B. Soares & M.P. Morim by its leaets and in-
orescence, however it diers by having a red or green calyx, pink corolla, 1–2 pairs
of pinnae, and crypto-lomentiform fruits (vs. white calyx and corolla, 3–5(–6) pairs of
pinnae and indehiscent fruits in H. leucocalyx).
Marcos Vinicius Batista Soares et al. / PhytoKeys 205: 401–437 (2022)
428
Type material. B, Amazonas, São Gabriel da Cachoeira. Igarapé Tibuari,
auente do Vaupés 0°05'5"N, 67°23’16"W, 23 July 2012, . and fr., M.P. Morim,
J.R.V. Iganci, F. Bonadeu, E. Koenen 577 (holotype: RB [RB00728413]!; isotypes:
HUEFS!, INPA!, K!, MBM!, MG!, MO!, NY!, PEL!, S!, US!, Z!).
Description. Trees 2–6 m tall, trunk not observed, partially underwater during sea-
sonal inundation. Branches, leaf axes and peduncles sparsely pubescent to glabrescent.
Stipules linear, to 1.2 cm long, densely pubescent on outer surface. Leaves with 1–2 pairs
of pinnae; petiole including pulvinus 1.5–4.5 cm, cylindrical; rachis 0 or 3–4(–9) cm,
glabrous, canaliculate; extraoral nectaries borne between rst or both pairs of pinnae, ses-
sile, patelliform and smaller nectaries usually present between the leaet pairs; pinnae 3–5
jugate; leaets subsessile on pulvinules, chartaceous, c. 2–4(–6) × 1.5–2(–3) cm, rhomboid
or ovate, apex emarginate or obtuse, sometimes with a minute mucro, base asymmetri-
cally oblique to acute; adaxial and abaxial surfaces glabrous, discolorous, adaxial surface
sometimes lustrous; venation pinnate with c. 11–17 secondary veins, tertiary venation
reticulate and prominent on both surfaces when leaets dry. Inorescences dimorphic,
umbelliform with 7–10 peripheral owers and an enlarged sessile terminal ower, pedun-
cle 4–6(–8)cm. Bracts and bracteoles not seen. Flowers with a reddish or green calyx, pink
corolla and white laments, the ower buds oblong, ca. 8 mm long, with the corolla con-
cealed by the calyx prior to anthesis, peripheral owers on pedicels 0.7–1.5 cm, calyx cam-
panulate, c. 9 mm long, 5-angulate due to prominently raised veins, sparsely puberulent or
ciliate at the apex of the lobes, corolla tubular, with a prominulous midvein on the lobes,
c. 1.5 cm long, sparsely puberulent on the upper half of the lobes, stamens c. 50–60, the
laments white, c. 3 cm long, exserted from the corolla ca. 2 cm; ovary glabrous, 3–4mm,
sub-truncate to truncate at the apex, style 3.5–4 cm, stigma funnel-shaped; terminal ower
similar to peripheral owers but more robust and c. 5 mm wide at base, calyx c. 1.2 cm
long, corolla 1.6 cm long, stamens ca. 75, ca. 3.5 cm long. Pods typically 1–3 per in-
fructescence, crypto-lomentiform, up to 15-seeded, oblong, slightly curved, lignescent, c.
9.5 × 2.5 cm excluding a ca. 5 mm long mucro, dehiscence follicular, the smooth exocarp
and transversely brous mesocarp continuous, the endocarp septate, enveloping the seeds
which are released in monospermous articles. Seeds not seen in mature state, oblong, c.
1.6 × 0.4–0.7 cm, pleurogram extending from apex to base, c. 1.3 × 0.3–0.4 cm, closed.
Distribution and habitat. Brazil. Known only from the Upper Rio Negro region in
the Brazilian Amazon (Amazonas state), in seasonally inundated “campinarana” vegetation.
Phenology. Flowering and fruiting in July.
Etymology. e specic epithet refers to the type locality, near the River Uaupés,
in the state of Amazonas, Brazil. e indigenous people living in this area (e.g., the
Tucanos) were known as Uaupés, and later the river took the same name.
Notes. Hydrochorea uaupensis is only known from Amazonas state, Brazil, where it
was collected at “Igarapé Tibuari”, in the municipality of São Gabriel da Cachoeira, dur-
ing eldwork in July 2012. e species grows in open vegetation on white sand, known
in Brazil as campinarana in the Amazon Domain. During times of ood, only the tree-
tops are exposed above the water line. A second herbarium collection from close to the
type locality (Rio Tourí, a. do Rio Negro, igapó; R.L. Fróes 28691, IAN [IAN78279]),
Recircumscription of Hydrochorea 429
Figure 8. Hydrochorea uaupensis M.P. Morim, Iganci & E.J.M. Koenen A branch with inorescences
B branch with fruits C ovary D terminal ower E peripheral ower F dehisced fruit G detail of fruit
endocarp forming 1-seeded articles H, I monospermous articles. A–I from M.P. Morim et al. 577. Illustra-
tion by João Augusto Castor Silva.
Marcos Vinicius Batista Soares et al. / PhytoKeys 205: 401–437 (2022)
430
of which we have only seen an image, is here tentatively included under H. uaupensis be-
cause the fruit and leaves match the type material and the owers are described as pink
on the specimen label. Since these two occurrence records are close to the borders with
Colombia and Venezuela, the species is to be expected in those two countries.
e phylogenetic position of H. uaupensis, as the sister lineage of the clade composed
of Hydrochorea sensu Barneby and Grimes (1996) and the African Hydrochorea spp.,
provides ample support for this as a distinct taxon and a species new to science, as it
does not form a sister pair with any other known species. Furthermore, this phylogenetic
position is in line with the fruit morphology of the species being intermediate between
Balizia and Hydrochorea, adding further support, along with the paraphyly of Balizia, for
not maintaining these as distinct genera.
Conservation status. Data decient. e species is known only from two adjacent
localities in the Upper Rio Negro region of Amazonas state, Brazil. More collections
are needed to assess the species’ conservation status.
Conclusion
Our results provide signicant advances in the generic delimitation of Hydrochorea and
related taxa, as well as broadening our understanding of ongoing diversication in these
taxa. e uncertain phylogenomic position of Jupunba macradenia, and other species of
Jupunba, sharing a relatively large number of incompletely sorted genes with Hydrochorea,
leads to further diculties in our ability to delimit genera in a group where classication
has been notoriously unstable. Nevertheless, given the complex evolutionary patterns
across the genome presented by the Jupunba clade taxa, we decided to use morphology
as our main guide for taxonomic decisions, re-circumscribing Hydrochorea to include
ten species to account for the paraphyly of Balizia, while incomplete lineage sorting sur-
rounding the divergence between Hydrochorea and Jupunba does not falsify these two
genera as natural groups. Furthermore, not transferring all these species to Jupunba, al-
though a cautious decision, avoids the publication of more new names while safeguarding
morphological diagnosability. e species treated here as Hydrochorea form a morpho-
logically homogeneous group in terms of vegetative and oral characters, although the
fruits are variable as observed in other mimosoid genera (e.g., Aviles Peraza et al. 2022).
Nevertheless, fruit type is useful for the identication of some species of Hydrochorea.
Quantitative characters, such as the number of pinnae per leaf and number of leaets per
pinna, also can be useful for identication of some species, but we commonly observed
overlap in these characters between species and even variation on the same individual.
Acknowledgements
e authors are grateful to the curators of all herbaria consulted, with special thanks to
RB, NY and K. We thank Francismeire Bonadeu MSc., for her contributions during
Recircumscription of Hydrochorea 431
eldwork and for preparing herbarium labels; the National Indian Foundation (FU-
NAI) for providing permits for research inside indigenous areas in Amazonia; the in-
digenous community Baniwa, for receiving us during eldwork along the Içana River,
in the Brazilian Amazon; Eimear NicLughada, for sending high resolution specimen
images from K; Alex Pinheiro and João Augusto Castor Silva for the illustrations.
JRVI thanks the Brazilian National Council for Scientic and Technological Devel-
opment (CNPq-310075/2020-3; 311847/2021-8) for the research grants received.
We thank Domingos Cardoso and Marcelo Simon for sharing eldwork photographs
and herbarium specimen images. MVBS thanks the Federal University of Amazonas
(UFRA), the Museu Paraense Emílio Goeldi (MPEG), and the Federal University
of Rio Grande do Sul (UFRGS), for infrastructure support of this work; the Co-
ordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES - Oce for the
Advancement of Higher Education; Grant no. PNADB 922/2010) for a grant sup-
porting an MSc. scholarship and travel to consult herbaria; the Brazilian National
Council for Scientic and Technological Development (CNPq-141414/2016-2) for a
PhD. scholarship. EJMK wishes to thank Gwilym Lewis for acting as SYNTHESYS+
host at K. EJMK was supported by the Swiss National Science Foundation (Early.
Postdoc.Mobility fellowship P2ZHP3_199693) and received further support from
the SYNTHESYS+ project http://www.synthesys.info/ which is nanced by European
Community Research Infrastructure Action under the H2020 Integrating Activities
Programme, Project number 823827. We would like to thank Gwilym Lewis and
Marcelo Simon for their reviews that provided very useful comments that have im-
proved the manuscript.
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Appendix 1
Vouchers and European Nucleotide Archive (ENA, https://www.ebi.ac.uk/ena/) ac-
cession numbers for molecular phylogenetic data. Note that the taxonomy of Albizia
Recircumscription of Hydrochorea 435
sect. Arthrosamanea is updated in this volume by Aviles Peraza et al. (2022), where new
binomials in the genus Pseudalbizzia Britton & Rose are presented for the majority of
the species of this section.
Acacia rostellifera Benth., Murphy 466 (MEL): ERS11697109; Afrocalliandra redac-
ta (J.H. Ross) E.R. Souza & L.P. Queiroz, Germishuizen 5585 (PRE): ERS11697117;
Albizia adinocephala Britton & Rose, Hughes 1070 (FHO): ERS11697120; Albizia
burkartiana Barneby & J.W. Grimes, Stival-Santos 678 (RB): ERS4812854 ; Albi-
zia coripatensis (Rusby) Schery, Hughes 2433 (FHO): ERS11697123; Albizia decandra
(Ducke) Barneby & J.W. Grimes, Vilhena 231 (NY): ERS11697124; Albizia edwallii
(Hoehne) Barneby & J.W. Grimes, Dalmaso 272 (RB): ERS4812856 ; Albizia glabrip-
etala (H.S. Irwin) G.P. Lewis & P.E. Owen, Lewis 1652 (K): ERS11697125; Albizia
inundata (Mart.) Barneby & J.W. Grimes, Wood 26530 (K): ERS4812859; Albizia
multiora (Kunth) Barneby & J.W. Grimes, Hughes 3090 (FHO): ERS11697127; Al-
bizia niopoides (Spruce ex Benth.) Burkart, Simon 1601 (CEN): ERS11697128; Albi-
zia sinaloensis Britton & Rose, Hughes 1576 (K): ERS11697130; Albizia subdimidiata
var. minor Barneby & J.W. Grimes, Gorts 341 (K): ERS11697131; Albizia subdimidia-
ta (Splitg.) Barneby & J.W. Grimes var. subdimidiata, Ferreira 210 (K): ERS11697129;
Albizia tomentella Miq., Leach & Dunlop 3801 (L): ERR9867596; Albizia tomentosa
Standl., Hughes 648 (K): ERS11697132; Albizia xerophytica J.Linares, Hughes 1435
(K): ERS11697133; Archidendron clypearia (Jack) I.C. Nielsen, Wieringa 1849 (WAG):
ERS11697136; Calliandra bella Benth., Queiroz 15696 (HUEFS): ERS11697156;
Enterolobium cyclocarpum (Jacq.) Griseb., Macqueen & Styles 75 (K): ERS11697204;
Hydrochorea corymbosa (Rich.) Barneby & J.W. Grimes (1), Bonadeu 655 (RB):
ERS4812902 ; Hydrochorea corymbosa (Rich.) Barneby & J.W. Grimes (2), Iganci
862 (RB): ERS4812903 ; Hydrochorea elegans (Ducke) Iganci, M.V.B. Soares & M.P.
Morim, Iganci 870 (RB): ERS11697146; Hydrochorea gonggrijpii (Kleinh.) Barneby
& J.W. Grimes, Tillett 45696 (K): ERS11697223; Hydrochorea leucocalyx (Britton
& Rose) Iganci, M.V.B. Soares & M.P. Morim, Aguilar 1939 (NY): ERS11697147;
Hydrochorea panurensis (Spruce ex Benth.) M.V.B. Soares, M.P. Morim & Iganci,
Morim 563 (NY): ERS11697224; Hydrochorea obliquifoliolata (De Wild.) E.J.M.
Koenen, Wieringa 6519 (WAG): ERS4812863 ; Hydrochorea pedicellaris (DC.) Iganci,
M.V.B. Soares & M.P. Morim, Queiroz 15529 (HUEFS): ERS4812877 ; Hydrochorea
rhombifolia (Benth.) E.J.M. Koenen, Deighton 3618 (K): ERS11697168; Hydrochorea
uaupensis M.P. Morim, Iganci & E.J.M. Koenen, Morim 577 (RB): ERS4812878 ;
Jupunba abbottii Britton & Rose, Zanoni 21220 (NY): ERS11697071; Jupunba as-
plenifolia Britton & Rose, Ekman 6383 (NY): ERS11697072; Jupunba barbouriana
(Standl.) M.V.B. Soares, M.P. Morim & Iganci, Iganci 847 (RB): ERS11697073; Ju-
punba brachystachya (DC.) M.V.B. Soares, M.P. Morim & Iganci, Lima 7438 (RB):
ERS11697074; Jupunba cochleata (Willd.) M.V.B. Soares, M.P. Morim & Iganci, Bon-
adeu 673 (RB): ERS11697076; Jupunba commutata (Barneby & J.W. Grimes) M.V.B.
Soares, M.P. Morim & Iganci, Maguire 46145 (NY): ERS11697077; Jupunba lamen-
tosa, (Benth.) M.V.B. Soares, M.P. Morim & Iganci Lima 7487 (RB): ERS11697079;
Jupunba oribunda (Spruce ex Benth.) M.V.B. Soares, M.P. Morim & Iganci, Iganci
Marcos Vinicius Batista Soares et al. / PhytoKeys 205: 401–437 (2022)
436
883 (RB): ERS11697080; Jupunba idiopoda (S.F. Blake) M.V.B. Soares, M.P. Morim
& Iganci, Quesada 1718 (NY): ERS11697081; Jupunba laeta (Benth.) M.V.B. Soares,
M.P. Morim & Iganci, Mori 25147 (NY): ERS11697083; Jupunba langsdori (Benth.)
M.V.B. Soares, M.P. Morim & Iganci, Ribeiro 728 (RB): ERS11697084; Jupunba leu-
cophylla (Spruce ex Benth.) M.V.B. Soares, M.P. Morim & Iganci, Iganci 839 (RB):
ERS11697086; Jupunba longipedunculata (H.S. Irwin) M.V.B. Soares, M.P. Morim
& Iganci, Cardona 2682 (NY): ERS11697088; Jupunba macradenia (Pittier) M.V.B.
Soares, M.P. Morim & Iganci, Lourteig 3021 (NY): ERS11697089; Jupunba mi-
crocalyx (Spruce ex Benth.) M.V.B. Soares, M.P. Morim & Iganci, Iganci 855 (RB):
ERS11697090; Jupunba nipensis Britton & Rose, Mayo 19662 (NY): ERS11697091;
Jupunba oppositifolia Britton & Rose, Liegier 16014 (NY): ERS11697092; Jupunba
oxyphyllidia (Barneby & J.W. Grimes) M.V.B. Soares, M.P. Morim & Iganci, Yonker
6157 (NY): ERS11697093; Jupunba rhombea (Benth.) M.V.B. Soares, M.P. Morim &
Iganci, Iganci 261 (RB): ERS11697087; Jupunba trapezifolia Moldenke, Simon 1600
(CEN): ERS4812839 ; Jupunba villosa (Iganci & M.P. Lima) M.V.B. Soares, M.P.
Morim & Iganci, Borges 423 (RB): ERS11697095; Punjuba callejasii (Barneby & J.W.
Grimes) M.V.B. Soares, M.P. Morim & Iganci, Daly 5935 (NY): ERS11697075; Pun-
juba killipii Britton & Rose, Palodorios 6252 (NY): ERS11697082; Punjuba lehman-
nii Britton & Rose ex Britton & Killip, Escobar 7465 (NY): ERS11697085; Punjuba
racemiora Britton & Rose, Jimenez & Soares 3626 (USJ): ERS11697094.
Supplementary material 1
Supplementary data le S1
Authors: Marcos Vinicius Batista Soares, Erik Jozef Mathieu Koenen3, João Ricardo
Vieira Iganci, Marli Pires Morim
Data type: zip. archiv.
Explanation note: Alignments of 560 nuclear loci in FASTA format (zipped).
Copyright notice: is dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Link: https://doi.org/10.3897/phytokeys.205.82775.suppl1
Recircumscription of Hydrochorea 437
Supplementary material 2
Supplementary data le S2
Authors: Marcos Vinicius Batista Soares, Erik Jozef Mathieu Koenen3, João Ricardo
Vieira Iganci, Marli Pires Morim
Data type: zip. archiv.
Explanation note: Gene trees of 560 nuclear loci in Newick format (zipped).
Copyright notice: is dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Link: https://doi.org/10.3897/phytokeys.205.82775.suppl2
Supplementary material 3
Supplementary data le S3
Authors: Marcos Vinicius Batista Soares, Erik Jozef Mathieu Koenen3, João Ricardo
Vieira Iganci, Marli Pires Morim
Data type: zip. archiv.
Explanation note: MI gene trees (one tip per accession) of 398 nuclear loci with nodes
with < 10% bootstrap support collapsed, in Newick format.
Copyright notice: is dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Link: https://doi.org/10.3897/phytokeys.205.82775.suppl3
Available via license: CC BY 4.0
Content may be subject to copyright.
