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PhytoKeys 127: 1–37 (2019)
The rise of Cynometra and the fall of Maniltoa: a generic re-circumscription and...
doi: 10.3897/phytokeys.127.29817
RESEARCH ARTICLE
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The rise of Cynometra (Leguminosae)
and the fall of Maniltoa: a generic re-circumscription
and the addition of 4 new species
Aleksandar Radosavljevic1,2,3
1 Chicago Botanic Garden, 1000 Lake Cook Road, Glencoe, Illinois 60022, USA 2 National Museum of
Natural History, Smithsonian Institution, 10th St and Constitution Ave NW, Washington, D. C. 20560, USA
3 Northwestern University, 633 Clark St, Evanston, Illinois 60202, USA
Corresponding author: Aleksandar Radosavljevic (aleks.rado@u.northwestern.edu)
Academic editor: Clifford Morden | Received 28 September 2018 | Accepted 31 May 2019 | Published 19 July 2019
Citation: Radosavljevic A (2019) The rise of Cynometra (Leguminosae) and the fall of Maniltoa: a generic re-
circumscription and the addition of 4 new species. PhytoKeys 127: 1–37. https://doi.org/10.3897/phytokeys.127.29817
Abstract
Cynometra L. is a genus of ca. 85 species of shrubs to large trees. It is amongst the largest genera in the
legume subfamily Detarioideae and one of the few with a pantropical distribution. Perhaps due to this
wide distribution and high diversity, systematists and taxonomists have struggled with the classification of
Cynometra and its close ally, the genus Maniltoa Scheff. Recent phylogenetic studies have shown that many
of the African species are more closely related to other genera and that the genus Maniltoa is nested within
a clade of Indo-Pacific Cynometra. Here, I present an emended circumscription of Cynometra that excludes
the African species defined by jointed pedicels and dehiscent fruits and includes the species formerly
recognised in Maniltoa. New combinations in Cynometra are also provided for those species that require
them. Additionally, four new species of Neotropical Cynometra are described and illustrated: Cynometra
cerebriformis sp. nov. from the lower Rio Trombetas in Brazil; Cynometra dwyerii sp. nov. from the Darien
gap region of Panama; C. tumbesiana sp. nov. from the dry tropical forests of Ecuador and Peru; and C.
steyermarkii sp. nov. from the foothills of the western Cordillera de la Costa in Venezuela.
Keywords
Cynometra L, Detaroideae, Leguminosae, Maniltoa Scheff, new species, taxonomy
Copyright Aleksandar Radosavljevic. 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.
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Aleksandar Radosavljevic / PhytoKeys 127: 1–37 (2019)
Introduction
The genus Cynometra L. (Leguminosae) has a broad tropical distribution, is relatively species rich and many of its species are poorly represented in herbaria. The
approximately 85 species of trees and (some) shrubs in the genus are spread somewhat evenly amongst four regions (Figure 1): the Neotropics (from southern Mexico
and the Caribbean to northern Argentina; Dwyer 1958, Sprada Tavares and da Silva
1992), mainland tropical Africa (equatorial forest belt; Léonard 1951), Madagascar
and the Comoros Islands (Du Puy et al. 2002) and the Indo-Pacific (extending from
the Western Ghats eastward to Fiji; Knaap-van Meeuwen 1970, Smith 1985). Perhaps
unsurprisingly, the genus has long troubled taxonomists and systematists who have
struggled with its diversity in several attempts to revise and classify regional groupings
of the species (Léonard 1951, Dwyer 1958, Knaap-van Meeuwen 1970). Further, its
relationship to the much smaller Pacific genus Maniltoa Scheff. has remained equivocal. Several important phylogenetic studies of the Caesalpinioideae sensu lato suggested that Cynometra was not monophyletic (Bruneau et al. 2001, 2008). More recent
studies, with greater taxonomic sampling, have provided additional evidence that the
genus is polyphyletic and, furthermore, that one of the lineages is paraphyletic with
respect to Maniltoa (Mackinder et al. 2013, Radosavljevic et al. 2017, de la Estrella et
al. 2018). In order for the classification of Cynometra to reflect a monophyletic taxon,
a new genus circumscription must be provided and Maniltoa should be subsumed
into Cynometra. In addition, during the course of herbarium study and fieldwork,
several undescribed species were revealed. Those species are here described. This paper
is divided into three parts: I. A new circumscription of the genus Cynometra; II. The
transfer of all species of Maniltoa into Cynometra; and III. Description of four new
species of Cynometra.
Materials and methods
The following herbaria allowed the author to study their collections in person or
provided loans: A, ATH, BM, BRI, CANB, F, IAN, INPA, K, MG, MO, NY, P,
RB, SING and US. Morphological characters were assessed and measured from
herbarium material. Flowers and immature fruit were rehydrated in warm Pohl’s
solution (Pohl 1965) prior to dissection and measurement. Other characters were
measured directly from the herbarium specimens. For the new species from Ecuador,
flower colour, habit and habitat information were taken from label data and field
observations by Gwilym Lewis and Bente Klitgaard (pers. comm.); for the other
three species, these features were described from label data. Reproductive status is
indicated with the following abbreviations (alone or in combination): fl, flowering; fr,
fruiting; im, immature; st, sterile. Illustrations were made at the National Museum of
Natural History, Washington DC, USA by Mattias S. Lanas (Cynometra steyermarkii),
Katherine Rudebusch (C. tumbesiana) and Alice Tangerini (C. cerebriformis and
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The rise of Cynometra and the fall of Maniltoa: a generic re-circumscription and...
b.
c.
Cynometra L. emend. Rados.
a.
Cynometra bauhiniifolia 1
Cynometra bauhiniifolia 2
Cynometra portoricensis 1
Cynometra portoricensis 2
Cynometra vogelii
Cynometra spruceana
Cynometra spruceana
var. phaselocarpa
Cynometra sp 3
Cynometra stenopetala
Cynometra bauhiniifolia 3
Cynometra longifolia 1
Cynometra longifolia 2
Cynometra spruceana
var procera
Cynometra marginata 2
Cynometra marginata 1
Cynometra longicuspis 2
Cynometra longicuspis 1
Cynometra oaxacana 2
Cynometra oaxacana 1
Cynometra retusa 3
Cynometra retusa 2
Cynometra retusa 1
Maniltoa megalocephala
Maniltoa gemmipara
Maniltoa sp
Maniltoa schefferi
Cynometra sp 2
Cynometra ramiflora 1
Cynometra cebuensis
Cynometra sp 4
Maniltoa browneoides
Cynometra ramiflora 2
Cynometra minutiflora
Cynometra malaccensis
Cynometra aff ramiflora
Cynometra cauliflora
Cynometra roseiflora
Cynometra iripa
Cynometra floretii
Cynometra madagascarensis
Cynometra dauphinensis
Cynometra abrahamii
Scorodolpheus
Clade
Zenkerella citrina
Normandiodendron bequaertii
Taxa excluded
from Cynometra
d.
Cynometra filifera
Cynometra greenwayi
Cynometra gilmanii
Cynometra suaheliensis 2
Cynometra longipedicellata
Cynometra webberi
Cynometra sp 5
Cynometra sp 1
Cynometra brachyrrhachis
Cynometra suaheliensis 1
Dicymbe altsonii
Dicymbe aff. hymenaea
Polystemonanthus dinklagei
Hymenostegia
Clade
Leonardoxa africana
Talbotiella gentii
Loesenera kalantha
Tamarindus indica
Paramacrolobium coeruleum
Cryptosepalum staudtii
Humboldtia vahliana
Figure 1. A Map showing global distribution of Cynometra and Maniltoa (white hatching) B ML majority
rules consensus tree from Radosavljevic et al. (2017) with branches coloured by species distributions. Star
indicates type species of Cynometra C indehiscent fruits typical of Cynometra as circumscribed here; D dehiscent fruits typical of those species formerly included in Cynometra, but excluded by the new circumscription.
C. dwyerii) from herbarium specimens. Two of the new species are described based
on a single collection: in both cases, the taxa are morphologically and geographically
distinct and, after extensive herbarium study, the author is confident that the available
material, while limited, is sufficient to distinguish two new entities at the species level.
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Part I: A new circumscription of the genus
Linnaeus established the genus Cynometra and included two species from southeast Asia in the first edition of Species Plantarum, Cynometra cauliflora L. and C.
ramiflora L. (Linnaeus, 1753). This original concept of Cynometra was of a genus
distinguished by compound leaves, four reflexed sepals, five equal petals and by single-seeded, indehiscent pods with thick and tuberculate valves (Linnaeus’ description does not explicitly mention ‘indehiscent pods’, but it is a character present in
both of his original species). For nearly the next century, this circumscription was
generally followed by taxonomists around the world. The limits of the genus were
first expanded by the publication of two new species from the Indo-Pacific region:
Cynometra polyandra Roxb., described in 1820 from Indian material and Cynometra
grandiflora A.Gray, described in 1854 from Fijian specimens. These species differed
most notably from the traditional understanding of Cynometra by having more numerous stamens (35–40 in C. grandiflora and approximately 50 in C. polyandra;
Roxburgh 1820, Gray 1854). Bentham (1865a) later described the African species C.
floribunda Benth. and C. laxiflora Benth., species with large, petaloid bracteoles and
only two showy petals which he placed in a new subgenus, Hymenostegia (the fruit
of the two species also differs from Cynometra, but these were unknown at the time).
Scheffer (1876) established the segregate genus Maniltoa after viewing collections
from New Guinea; believing the material to be conspecific, he included a single species, Maniltoa grandiflora (A.Gray) Scheff. (based on Cynometra grandiflora A.Gray).
While the most apparent difference between the new genus Maniltoa and typical
Cynometra was the number of stamens, Scheffer (1876) argued that the genus was
distinct based on the size of the flowers, the number of seeds, caducous filiform stipules, the presence of a tendril terminating the leaf rachis and bracts subtending each
pedicel in the inflorescence. He does not include or reference C. polyandra, another
polyandrous species. Maniltoa was not universally accepted at first and Taubert’s
(1894) treatment of the Leguminosae for Engler’s Die Natürlichen Pflanzenfamilien
treated Maniltoa as a section of Cynometra (along with the sections Eucynometra,
Hymenostegia and Pseudocynometra). A few years later, however, Harms (1897) broke
apart this broad concept of Cynometra. He elevated Hymenostegia to the rank of genus and restored Maniltoa by combining it with section Pseudocynometra (consisting
of a single species, C. polyandra) on the basis of greater stamen number and the presence of prominent buds enclosed in imbricate scarious bract scales (Harms 1897,
1902, 1919). This understanding of Cynometra and Maniltoa in the Indo-Pacific
was further refined by Knaap-Van Meeuwen (1970) who sought to better define the
limits between the two genera – she again placed greatest significance on the stamen
number, placing those with 8–13 stamens in Cynometra and those with 15–80 in
Maniltoa. While many authors have followed this delimitation (Verdcourt 1979;
Hou et al 1996), some have continued to question the distinctionof the two genera
(Soerianegara 1993; Pan et al. 2010).
The rise of Cynometra and the fall of Maniltoa: a generic re-circumscription and...
5
In Africa, misunderstanding of the delimitation of Cynometra persisted, even after Harm’s (1897) treatment and the turn of the century marked the beginning of
a period of rapid taxonomic expansion in both the number of species and, consequently, the characters describing the generic limits of Cynometra. As explained (in
great detail) by Léonard (1951), the first few species described in Africa generally
matched the Linnaean concept (e.g. Cynometra vogelii Hook.f. in 1849 from Nigeria;
C. mannii Oliv. in 1871 from Cameroon). However, with the description of Cynometra sessiliflora Harms, Harms (1899) expanded the understanding of Cynometra
to include species with five erect sepals (instead of four reflexed sepals) and, most
importantly for the continued confusion regarding the genus, dehiscent pods with
thin, flat valves. Not long after, the publication of Cynometra alexandri C.H.Wright,
C. gilletii De Wild., C. laurentii De Wild., C. lujae De Wild., C. oddonii De Wild.
and C. pedicellata De Wild. added more species with five sepals and dehiscent fruits
and further expanded the genus to include species with an intrastaminal disc and
numerous ovules (Wright 1902; De Wildeman 1904, 1905, 1906, 1907). Harms
(1907) also continued to expand the generic conception of Cynometra in Africa to
include species with alternate leaflets, distichous flowers and a stipe fused with the
receptacle wall when he described C. leptantha Harms, C. multijuga Harms and C.
pierreana Harms. Cynometra brachyura Harms, expanded the genus to again include
species with only two petals (Harms 1913). As a result, species with all manner of
combinations of these characters were placed in Cynometra (see Baker 1930 and Pellegrin 1949). While many taxonomists, working in Africa, noted the heterogeneity of
the genus, it was not until Léonard undertook a thorough and careful revision of the
African species that a coherent circumscription of Cynometra began to emerge (Baker
1930; Lebrun 1933; Aubréville 1936; Léonard 1951, 1957).
Léonard’s work (1951, 1957) was a major advancement in revising the boundaries
of Cynometra and its allies in Africa, which had become so broad as to make genera
nearly indistinguishable. Léonard (1951) removed from Cynometra any taxa with the
stipe of the ovary adnate to the hypanthium or eccentrically inserted (species with a
free, central stipe but alternate leaflets with translucent dots were moved to Gilletiodendron Vermoesen). To accommodate his revised concept of the genus, Léonard (1951,
1957) described a new genus, Lebruniodendron J.Léonard, resurrected the genera Gilletiodendron Vermoesen and Zenkerella Taub. and transferred numerous species out of
Cynometra and into the genera Hymenostegia (Benth.) Harms, Plagiosiphon Harms,
Schotia Jacq. and Scorodophloeus Harms. He placed the remaining Cynometra into three
informal groups; they share a centrally inserted stipe, opposite leaflets, five equal petals
and imbricate sepals, but differ notably in characters such as inflorescence structure,
presence or absence of a staminal disc, presence or absence of foliar glands and fruit
shape and dehiscence. Léonard (1951) acknowledged that the genus remained heterogenous – the species of ‘group one’ resembled ‘typical Cynometra’, but the others were
possibly misplaced. Later, he further assessed the genera of African Amherstieae and
Cynometreae, revising the tribal and generic limits using characters from germination
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mode, seedling architecture and wood structure, amongst others. (Léonard 1957). A
thorough examination of the genus Cynometra, unfortunately, was not included in
that otherwise exhaustive study. Léonard and others recognised that, as contemporarily
circumscribed, the genus was likely polyphyletic and would require further revisionary
studies at a global scale (Aubréville 1968, 1970; Breteler 1996; Léonard 1996).
An early morphological cladistic analysis by Temu (1990) provided the first phylogenetic evidence for the non-monophyly of African Cynometra. The results showed two
distinct clades of Cynometra in tropical Africa, one of which included Maniltoa. Later
morphological and molecular phylogenetic analyses, albeit with limited sampling of
Cynometra and Maniltoa, also found no evidence to support monophyly of Cynometra
and suggested that it may be paraphyletic with respect to Maniltoa (Bruneau et al.
2001, 2008). More recent studies which included a greater taxon sampling from Cynometra and Maniltoa have found support for two clades of Cynometra, one consisting
of exclusively East African species and the other containing species from the rest of the
genus plus Maniltoa (Mackinder et al. 2013; Radosavljevic et al. 2017; de la Estrella
et al. 2018). Radosavljevic et al. (2017), which featured 47 accessions from 36 species
of Cynometra and Maniltoa, recovered two strongly supported clades of Cynometra
sensu lato, which are each diagnosed by several morphological characters (Figure 1).
One clade (Cynometra clade A of Radosavljevic et al. 2017) is comprised of exclusively
east African species and is grouped as sister to a clade composed of Dicymbe Spruce
ex Benth. and Polystemonanthus Harms with strong support. The other clade, which
includes the type species Cynometra cauliflora L., contains Cynometra species from the
Neotropics, the remaining Afrotropical species and the Indo-Pacific species (Cynometra clade B of Radosavljevic et al. 2017). Maniltoa is nested within clade B and is also
recovered as non-monophyletic. This clade of Cynometra and Maniltoa is placed in
a clade with Zenkerella Taub., Normandiodendron J. Léonard and the Scorodophloeus
clade, again with strong support. The species of Cynometra clade A are characterised
by the absence of foliar extra-floral nectaries, paniculate inflorescences (with two exceptions), articulated pedicels, a well-developed intrastaminal disc, 2–4 ovules and
dehiscent fruit. Cynometra clade B is characterised by the presence of foliar extra-floral
nectaries (in most species), racemose inflorescences, simple pedicels, the absence of an
intrastaminal disc, 1–2 ovules and indehiscent fruit. The species of Cynometra clade A
correspond to Léonard’s (1951) groups 2 and 3 while Cynometra clade B, which contains the generitype Cynometra cauliflora L., corresponds to group 1. Léonard (1951,
1996) predicted as much; he referred to group 1 as ‘typical Cynometra’ and believed
that Cynometra alexandri and C. hankei Harms were likely misplaced. De la Estrella et
al. (2018), who sampled broadly across Detarioideae, placed Cynometra and Maniltoa
in a re-circumscribed Amherstieae.
To reconcile the discrepancy between the current classification of Cynometra and
Maniltoa and our understanding of their evolutionary relationships, I present here an
emended generic circumscription of Cynometra that excludes the species of Cynometra
clade A and includes the species formerly placed in Maniltoa. A treatment that proposes a new genus for the species of Cynometra clade A is in preparation.
The rise of Cynometra and the fall of Maniltoa: a generic re-circumscription and...
7
Taxonomic treatment
Cynometra L. Sp. Pl. 1:382. 1753.
Syn. Iripa Adans., Fam. 2: 508. 1763. Cynomora R.Hedw. Gen. Pl. [R. Hedwig]. 1806.
Type. Cynometra cauliflora L. (lectotype, designated by A.S. Hitchcock, in Hicthcock and Green 1959, pg. 152)
Metrocynia Thouars, Gen. Nov. Madagasc. 22. 1806. Type. Metrocynia commersoniana DC.
Maniltoa Scheff., Ann. Jard. Bot. Buitenzorg 1: 20. 1876. Type. Cynometra grandiflora
A.Gray.
Schizosiphon K.Schum., Fl. Kais. Wilh. Land 101. 1889. Schizoscyphus K.Schum ex Taubert, nom. superfl. Bot. Centralbl. 41: 265. 1890. Type. Schizosiphon rosea K. Schum.
Pseudocynometra Kuntze in Post & Kuntze, Lex. Gen. Phan. Phan. 464. 1903. Type.
Cynometra polyandra Roxb.
Trees or shrubs (infrequent), evergreen (rarely deciduous), buttressed or not, growth
flush-wise, new growth flaccid and whitish or reddish at emergence, becoming green
with maturity, vegetative buds covered in a series of imbricate scales. Stipules lateral,
free, linear or filamentous, early caducous, scars typically not visible on mature growth.
Leaves pulvinate, petiolate, rachis (if present) terete or caniculate, often terminating in
a filiform outgrowth, axes glabrous or pubescent, paripinnate with 1–16 pairs of opposite leaflets (rarely unifoliolate); leaflets petiolulate but sometimes appearing sessile
because of decurrent lamina, blade lanceolate, ovate, elliptic, oblong, obovate, oblanceolate or trapeziform, symmetrical to strongly asymmetrical, glabrous or sparsely pubescent abaxially, glabrous adaxially (rarely with sparse pubescence along midvein),
margins entire, apex broadly obtuse to acuminate, retuse or emarginate, mucronate,
base oblique, distal margin decurrent to petiolule; laminar nectaries usually present,
abaxial, submarginal, embedded in laminar surface, shallow, without noticeably raised
edges. Inflorescences axillary or ramiflorous (rarely cauliflorous), racemose, 1–2(–3),
per axil, buds enclosed by imbricate bracts, appearing conical, ovoid or cigar-shaped
in silhouette; pedicels simple (not articulated), filamentous in anthesis, accrescent and
lignified in fruit; bracts enclosing inflorescence during development prior to emergence, imbricate, distichous, deciduous or persistent; bracteoles not enclosing buds,
paired, inserted along proximal half of pedicel, opposite or subopposite, caducous.
Flowers bisexual, actinomorphic; hypanthium present (sometimes indistinct), either
turbinate-campanulate and short or tubular and partially to completely enveloping
ovary; disc absent; sepals (3–)4(–5), reflexed at anthesis, unequal, deltoid, ovate, elliptic or oblong; petals 5, crumpled or smooth, incurved, erect or horizontal, equal,
linear, oblong, or oblanceolate, with rounded or acute apices; stamens (8–)10–80,
filaments free or briefly connate basally; anthers dorsifixed, versatile, longitudinally
dehiscent; ovary inserted +/- centrally, occasionally eccentric, free, short-stipitate or
subsessile, pubescence varied but rarely glabrous, light green or pinkish, often turning red post anthesis, ovules 1 (2); style eccentric, glabrous; stigma capitate. Legumes
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indehiscent, splitting along suture after germination due to action of emerging shoot,
orbicular to oblong, often laterally compressed, smooth to deeply fissured, often apiculate (in some taxa only when immature), valves 0.5–4.0 mm thick, cork-like in several
species, remaining attached to cotyledons during and after germination. Seeds 1–2,
enclosed in fruit until germination. Seedling germination epigeal.
The emended description above reflects the merger of Maniltoa and Cynometra s.s.
and the exclusion of the east African taxa with dehiscent pods, paniculate inflorescences
and articulated pedicels (Tables 1, 2). While many taxa in the former Maniltoa are no
doubt distinctive and, in many cases, striking when compared to the more unassuming Cynometra, a close examination of their morphological characteristics reveals a strong
similarity. The number of stamens has been perhaps the most distinctive and most often
cited character separating the two genera. Cynometra has long been known to have only
10 stamens, while Maniltoa has been traditionally treated as having anywhere from 15–80
stamens. Indeed, this proliferation in stamen number is rare in the Detarioideae. However, a closer look at the Indo-Pacific Cynometra reveals several examples of species which
regularly have as few as eight stamens (Cynometra glomerulata Gagnep.) or as many as 12
(Cynometra katikii Verdc.) and some with variation across the range within a species (8–10
stamens in Cynometra cauliflora L. and 10–15 stamens in Cynometra ramiflora L.; Knaapvan Meeuwen 1970, Verdcourt 1979). So, while the majority of species, placed within
Cynometra, do have 10 stamens, there is lability amongst this trait in the taxa of the region.
Several other characteristics of Maniltoa that supposedly distinguish it from Cynometra are also incorrect. The first is the presence of conspicuous ‘bract covered’ vegetative and
reproductive buds (Scheffer 1876, Harms 1902, 1919). In some species of Maniltoa, these
buds can be many centimetres long and several centimetres in diameter. The scales themselves are often tan or brown, but can be whitish or pinkish and are often covered with
parallel striations running longitudinally along the surface. These types of buds are also
present in Cynometra, although they are much smaller, in some cases only 4 or 5 mm long.
Likewise, the scales are similar in shape, vestiture and surface texture. Another general
characteristic used to separate the two genera has been the sturdiness of the inflorescence
rachis in Maniltoa (Knaap-van Meeuwen 1970). However, this is likely related to the
overall general difference in size of the flowers and inflorescence between the two genera:
Maniltoa tend to have larger buds, bracts, inflorescences and individual flowers. The larger
inflorescences of the Maniltoa species may be an adaptation to mammal pollination as
there are reports of marsupials and bats visiting the flowers of Maniltoa species in Australia
and the Pacific Islands (Marshall 1985, Endress 1994). Scheffer (1876) cited the presence of a tendril terminating the leaf rachis as a distinguishing character of Maniltoa, but
Dwyer (1958) notes that this trait is present in several Cynometra. Finally, differences in
wood anatomy given by Knaap-van Meeuwen (1970) have been found to be insufficiently
distinct from one another and overlapping in range (Soerianegara 1993; Pan et al. 2010).
The most clearly distinguishing feature of the newly emended Cynometra with
respect to the taxa here excluded (Cynometra clade A) is the indehiscent pod. Indeed,
if one examines the taxa misplaced in Cynometra over the years, they nearly all share
the characteristic of dehiscent pods. Unfortunately, Linnaeus makes no mention of the
The rise of Cynometra and the fall of Maniltoa: a generic re-circumscription and...
9
Table 1. Species excluded from Cynometra. Distributions correspond to regions given in text. Species
may occupy only a part of the overall region. MTA = mainland tropical Africa.
Species
Cynometra alexandri C.H. Wright
Cynometra ananta Hutch. & Dalziel
Cynometra brachyrrhachis Harms
Cynometra engleri Harms
Cynometra filifera Harms
Cynometra fischeri Baker f.
Cynometra gilletii De Wild.
Cynometra gillmanii J. Léonard
Cynometra greenwayi Brenan
Cynometra hankei Harms
Cynometra leonensis Hutch. & Dalziel
Cynometra longipedicellata Harms
Cynometra lujae De Wild.
Cynometra nyangensis Pellegr.
Cynometra oddonii De Wild.
Cynometra palustris J. Léonard
Cynometra pedicellata De Wild.
Cynometra sessiliflora Harms
Cynometra suaheliensis (Taub.) Baker f.
Cynometra ulugurensis Harms
Cynometra webberi Baker f.
Distribution
MTA
MTA
MTA
MTA
MTA
MTA
MTA
MTA
MTA
MTA
MTA
MTA
MTA
MTA
MTA
MTA
MTA
MTA
MTA
MTA
MTA
nature of the pods’ dehiscence. Likewise, subsequent treatments were vague. Bentham
(1865b), Taubert (1894) and Harms (1897) all described the pods as ‘two-valved’ without further elaboration. Léonard (1951) and Aubréville (1968, 1970) cite the pods of
‘typical Cynometra’ as dehiscent or indehiscent, likely owing to the fruit of Cynometra
mannii, the valves of which are not dehiscent but, upon pressing and drying, often
split and rupture in several places along axes roughly perpendicular to the sutures.
A similar occurrence is observed in the Neotropical species Cynometra bauhiniifolia
Benth. When Harms (1899) described C. sessiliflora, he set a precedent regarding the
pod characters that contributed to the imprecise delimitation of Cynometra and subsequently led to the taxonomic confusion surrounding Cynometra in Africa.
With this new circumscription, the two genera are united and share radially symmetric flowers, bract covered buds (with bracts persisting on the inflorescence), flowers with early caducous bracteoles not enveloping the flowers and indehiscent fruits
(which may play a role in dispersal as many taxa are associated with alluvial habitats
or appear as drift ‘fruit’ in coastal environments; Ridley 1930, Léonard 1951, Clarke
et al. 2001, Tomlinson 2016). The new classification changes the overall distribution of the taxa. Prior to this, Cynometra was a genus with its diversity distributed
somewhat evenly across the American, African and Asian tropics. However, with the
exclusion of many of the African species and the inclusion of the former Maniltoa,
Cynometra is now a genus whose primary centre of diversity is the Indo-Pacific region, with a secondary centre in the Neotropics and the majority of Afrotropic taxa
restricted to Madagascar.
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Aleksandar Radosavljevic / PhytoKeys 127: 1–37 (2019)
Table 2. Accepted species of Cynometra. Taxonomy follows: Knaap Van-Meeuwen 1970; Léonard 1951;
Dwyer 1958. New combinations and new species presented in text are not included. Distributions correspond to regions given above. Species may occupy only a part of the overall region. COM = Comoros
Islands; INP = Indo-Pacfic; MAD = Madagascar; MTA = mainland tropical Africa; NEO = Neotropics.
Species
Cynometra abrahamii Du Puy & R.Rabev.
Cynometra americana Vogel
Cynometra ankaranensis Dupuy & R.Rabev.
Cynometra aurita R.Vig.
Cynometra bauhiniifolia Benth.
Cynometra beddomei Prain
Cynometra bourdillonii Gamble
Cynometra brachymischa Harms
Cynometra capuronii Du Puy & R.Rabev.
Cynometra cauliflora L.
Cynometra cloiselii Drake
Cynometra commersoniana Baill.
Cynometra congensis De Wild.
Cynometra copelandii (Elmer) Elmer
Cynometra craibii Gagnep.
Cynometra crassifolia Benth.
Cynometra cubensis A.Rich.
Cynometra cuneata Tul.
Cynometra dauphinensis Dupuy & R.Rabev.
Cynometra dongnaiensis Pierre
Cynometra duckei Dwyer
Cynometra elmeri Merr.
Cynometra falcata A.Gray
Cynometra fissicuspis (Pittier) Pittier
Cynometra floretii Labat & O.Pascal
Cynometra glomerulata Gagnep.
Cynometra grandiflora A.Gray
Cynometra hemitomophylla (Donn.Sm.) Rose
Cynometra hondurensis Dwyer
Cynometra hostmanniana Tul.
Cynometra humboldtiana Stergios
Cynometra insularis A.C.Sm.
Cynometra iripa Kostel.
Cynometra katikii Verdc.
Cynometra letestui (Pellegr.) J.Léonard
Cynometra longicuspis Ducke
Distribution
MAD
NEO
MAD
MAD
NEO
INP
INP
INP
MAD
INP
MAD
MAD
MTA
INP
INP
NEO
NEO
NEO
MAD
INP
NEO
INP
INP
NEO
COM
INP
INP
NEO
NEO
NEO
NEO
INP
INP
INP
MTA
NEO
Species
Cynometra longifolia Huber
Cynometra lukei Beentje
Cynometra lyallii Baker
Cynometra macrocarpa A.S.Tav.
Cynometra madagascariensis Baill.
Cynometra malaccensis Meeuwen
Cynometra mannii Oliv.
Cynometra marginata Benth.
Cynometra marleneae A.S.Tav.
Cynometra mayottensis Labat & O.Pascal
Cynometra megalophylla Harms
Cynometra microflora R.S.Cowan
Cynometra minutiflora F.Muell.
Cynometra mirabilis Meeuwen
Cynometra novoguineensis Merr. & L.M.Perry
Cynometra oaxacana Brandegee
Cynometra parvifolia Tul.
Cynometra pervilleana Baill.
Cynometra polyandra Roxb.
Cynometra portoricensis Krug & Urb.
Cynometra ramiflora L.
Cynometra retusa Britton & Rose
Cynometra sakalava Du Puy & R.Rabev.
Cynometra sanagaensis Aubrev.
Cynometra schlechteri Harms
Cynometra schottiana Hochr.
Cynometra simplicifolia Harms
Cynometra spruceana Benth.
Cynometra stenopetala Dwyer
Cynometra travancorica Bedd.
Cynometra trinitensis Oliv.
Cynometra vogelii Hook.f.
Cynometra warburgii Harms
Cynometra whitfordii Elmer
Cynometra yokotai Kaneh.
Cynometra zeylanica Kosterm.
Distribution
NEO
MTA
MAD
NEO
MAD
INP
MTA
NEO
NEO
COM
MTA
NEO
INP
INP
INP
NEO
NEO
MAD
INP
NEO
INP
NEO
MAD
MTA
MTA
NEO
INP
NEO
NEO
INP
NEO
MTA
INP
INP
INP
INP
Part II: Reduction of Maniltoa into Cynometra
The generic rearrangements require new combinations to reflect the merging of Maniltoa into Cynometra. For Maniltoa, in most cases, the taxonomy proposed by Knaap-van
Meeuwen (1970) has been followed, but for some species limits, the interpretations of
Verdcourt (1979) have been followed; such deviations are noted. Species, for which a
combination already exists in Cynometra, are not listed (Knaap-van Meeuwen 1970). In
total, 18 new combinations and one new name are proposed. A lectotype is designated for
Maniltoa megalocephala Harms and a neotype is designated for Maniltoa peekelii Harms.
The rise of Cynometra and the fall of Maniltoa: a generic re-circumscription and...
11
1. Cynometra basifoliola (Verdc.) Rados., comb. nov.
urn:lsid:ipni.org:names:77199243-1
Basionym. Maniltoa basifoliola Verdc., Kew Bull. 37: 129. 1982. Type. PAPUA NEW
GUINEA. Madang province: Madang subprovince [district], cleared area utilized as a
gravel pit on north side of Madang Usino Highway on banks of Gogol River, 145 37
E, 05 15 S, 100 m alt., 25 April 1979, S.H. Somer & P. Katik LAE 75185 (holotype:
K; isotypes: BM, L, M).
Notes. The arrangement of leaflets for which this species is named (i.e. the basalmost pair of leaflets inserted just above petiole and separated from terminal pair of
leaflets by a relatively long rachis) is unusual amongst the species that were formerly
included in Maniltoa; however some species of Cynometra have a similar arrangement
(e.g. Cynometra sakalava Du Puy & R.Rabev from Madagascar).
2. Cynometra brassii (Merr. & L.M.Perry) Rados., comb. nov.
urn:lsid:ipni.org:names:77199244-1
Basionym. Maniltoa brassii Merr. & L.M.Perry, J. Arnold Arbor. 23: 398. 1942. Type.
BRITISH NEW GUINEA [PAPUA NEW GUINEA]. Central Division: U-uma River, 14 May 1926, [fl., imm. fr.], L. J. Brass 1428, (holotype: A; isotype: K).
Notes. The protologue lists the collection year as 1928, however 1926 appears
to be the correct date. While the label affixed to the holotype does list the collection
date as “14 May 1928”, this appears to be a transcription error. The handwritten
slip attached to the holotype, presumably filled out by the collector Brass, gives
the date as “14/5/26”. Additionally, the printed label is titled “Arnold Arboretum
Expedition, 1925–1926”. The label on the isotype also has the collection date as 14
May 1926.
3. Cynometra browneoides (Harms) Rados., comb. nov.
urn:lsid:ipni.org:names:77199245-1
Basionym. Maniltoa browneoides Harms, Notizbl. Königl. Bot. Gart. Berlin 3:
190. 1902. Synonym. Pseudocynometra browneoides (Harms) Kuntze, Deutsche
Bot. Monatsschr. 21: 173. 1903. Type. [INDONESIA]. Java. [West Java: Bogor
Botanic Gardens], 1880–1882, [fl], H. O. Forbes 1204a (holotype: B, destroyed;
isotype: BM).
Synonym. Maniltoa gemmipara Scheff. ex Backer, Voorl. Schoolfl. Java 104: in clavi. 1908. Type. [INDONESIA]. Cultivated at Weltevreden (Djakarta), 1908, Backer
s.n. (holotype: L, fide Knaap-van Meeuwen 1970).
Notes. According to Knaap-van Meeeuwen (1970, p. 42), the type material for
Maniltoa browneoides came from a cultivated specimen grown at Bogor Botanical Gar-
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Aleksandar Radosavljevic / PhytoKeys 127: 1–37 (2019)
dens from seeds or seedlings collected by Forbes on New Guinea, however no citation
is given as the source for this information. The holotype for M. gemmipara is listed as
being at Leiden by Knaap-van Meeuwen, however there are no specimens there annotated as such. There are two specimens, however, from Java collected in 1908 that seem
to match the details in the protologue. These are filed as M. gemmipara and have the
registration numbers L.3886447 and L.3886448.
4. Cynometra cynometroides (Merr. & L.M.Perry) Rados., comb. nov.
urn:lsid:ipni.org:names:77199246-1
Basionym. Maniltoa cynometroides Merr. & L.M.Perry, J. Arnold Arbor. 23: 398.
1942. Type. BRITISH NEW GUINEA [PAPUA NEW GUINEA]. Palmer River, 2
mi. below junction Black River, 100m alt., Jun 1936, [fr], L. J. Brass 6903 (holotype:
A; isotypes: BM, BO, BRI, L).
5. Cynometra fortuna-tironis (Verdc.) Rados., comb. nov.
urn:lsid:ipni.org:names:77199247-1
Basionym. Maniltoa fortuna-tironis Verdc., Kew Bull. 32 (1): 243. 1977. Type. PAPUA NEW GUINEA. Central District [Central Province]: Rouna Falls area, forest by
tributary of R. Laloki, 300 m alt., 2 Jan. 1976, B. Verdcourt, C.R. Huxley, & Dodd 4899
(holotype: K; isotypes: LAE, UPNG).
6. Cynometra lenticellata (C.T.White) Rados., comb. nov.
urn:lsid:ipni.org:names:77199248-1
Basionym. Maniltoa lenticellata C.T.White, J. Arnold Arbor. 8: 130. 1927. Type.
PAPUA [PAPUA NEW GUINEA]. Northern Division: Sageri, July 1922, [fl], C. E.
Lane-Poole 203 (holotype: BRI; isotype: A).
7. Cynometra lenticellata var. villosa (Verdc.) Rados., comb. nov.
urn:lsid:ipni.org:names:77199249-1
Basionym. Maniltoa lenticellata var. villosa Verdc., Kew Bull. 32 (1): 241. 1977. Type.
TERRITORY OF NEW GUINEA [PAPUA NEW GUINEA]. Morobe district [Morobe province]: Lae subdistrict [Lae district], Kassam Pass, 6°20'S, 146°00'E, 3500
ft alt., [fl], J.S. Womersley & J. Vandenberg NGF 37192 (holotype: LAE; isotypes: A,
BISH, BO, BRI, CANB, K, L, NSW, PNH, SING, UPNG, US)
The rise of Cynometra and the fall of Maniltoa: a generic re-circumscription and...
13
8. Cynometra mariettae (van Meeuwen) Rados., comb. nov.
urn:lsid:ipni.org:names:77199250-1
Basionym. Maniltoa mariettae van Meeuwen, Blumea 18 (1): 37. 1970. Type. TERRITORY OF NEW GUINEA [PAPUA NEW GUINEA]. Morobe District [Morobe
Province]: Yalu, 6°36'S, 146°52'E, 50 ft alt., Jul 1944, [fl], C.T. White, H.E. Dadswell,
& L.S. Smith, NGF 1661 (holotype: BRI; isotype: CANB).
9. Cynometra megalocephala (Harms) Rados., comb. nov.
urn:lsid:ipni.org:names:77199251-1
Basionym. Maniltoa megalocephala Harms, Bot. Jahrb. Syst. 55 (1): 52. 1917. Type.
NORDÖSTLICH NEU-GUINEA [PAPUA NEW GUINEA]. Sepik area, 1912–1913,
C. L. Ledermann 7857, 7895, 10616 (syntypes: B, destroyed). PAPUA NEW GUINEA.
Sepik area, Harms, Bot. Jahrb. Syst. 55 (1): 53, tab. 2. 1917. (lectotype, here designated)
Notes. All three syntypes cited in Harms’ original description were destroyed and
no isotypes have been located. Therefore plate 2 (tab. 2) from Harms 1917 is designated as the lectotype (Figure 2).
10. Cynometra minor (A.C.Sm.) Rados., comb. nov.
urn:lsid:ipni.org:names:77199252-1
Basionym. Maniltoa minor A.C.Sm., Sargentia 1: 37. 1942. Type. FIJI. Lau Province:
Moala Island, near Maloku, 20–24 Mar 1934, [fl], A.C. Smith 1333 (holotype: GH;
isotype: BISH, K, NY, S, WIS, US).
11. Cynometra plurijuga (Merr. & L.M.Perry) Rados., comb. nov.
urn:lsid:ipni.org:names:77199253-1
Basionym. Maniltoa plurijuga Merr. & L.M.Perry, J. Arnold Arbor. 23: 398. 1942.
Type. NETHERLANDS NEW GUINEA [INDONESIA]. [Papua:] 2 km southwest
of Bernhard Camp, Idenburg River, 650 m alt., 3 April 1939, L.J. Brass & C. Versteegh
13539 (holotype: A; isotype: BO, BRI, L).
12. Cynometra psilogyne (Harms) Rados., comb. nov.
urn:lsid:ipni.org:names:77199254-1
Basionym. Maniltoa psilogyne Harms, Bot. Jahrb. Syst. 55 (1): 50. 1917. Type. KAISER-WILHELMSLAND [Papua New Guinea]. At Kaulo, ca. 250 m alt., 1 December
1907, R. Schlechter 16934 (holotype: B; isotypes: A, E, G, K, L, S, Z).
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Aleksandar Radosavljevic / PhytoKeys 127: 1–37 (2019)
13. Cynometra rosea (K.Schum.) Rados., comb. nov.
urn:lsid:ipni.org:names:77199255-1
Basionym. Schizosiphon roseus K.Schum., Fl. Kais. Wilh. Land 101. 1889. Syn.
Schizoscyphus roseus (K.Schum.) Warb., nom superfl., Bot. Jahrb. Syst. 13: 331. 1891.
Maniltoa rosea (K.Schum.) Meeuwen, Blumea 18: 35. 1970. Type. KAISER-WILHELMSLAND [PAPUA NEW GUINEA]. Astrolabebay, Im Hochwalde von Constantinhafen, July [1886–1887], Hollrung 492 (holotype: B, destroyed; isotypes not
found). PAPUA NEW GUINEA. Madang Province: Madang District, Naikum, Josephstaal, 04°45'30"S, 145°00'30"E, 1 Sept. 1958, K.J. White NGF 10226 (neotype,
designated by Knaap-van Meeuwen 1970, p. 35: K; isoneotypes: BRI, CANB).
Maniltoa urophylla Harms, Bot. Jahrb. 55: 51. 1917. Type. NORDÖSTLICH
NEU-GUINEA [PAPUA NEW GUINEA]. Kameelsrücken, lager G, 6–900 m, Sept.
1912, C. L. Ledermann 8848 (holotype: B, destroyed; isotypes: WRSL, LE, not located).
Notes. I agree with Verdcourt (1979) that this synonymy is questionable. However, with the original type material destroyed and to avoid creating an unnecessary
new combination, I have elected to follow Knaap-van Meeuwen’s treatment here.
14. Cynometra schefferi (K.Sch.) Rados., comb. nov.
urn:lsid:ipni.org:names:77199256-1
Basionym. Maniltoa schefferi K.Sch., Fl. Kais. Wilh. Land 10. 1889. Syn. Pseudocynometra schefferi Kuntze, Deutsche Bot. Monatsschr. 21: 173. 1903. Type. PAPUA
NEW GUINEA. Dore, Teysmann s.n. (lectotype, designated by Knaap-van Meeuwen
1970, p. 45: L).
Maniltoa hollrungii Harms, Notizbl. Königl. Bot. Gart. Berlin 3: 189. 1902. Syn.
Pseudocynometra hollrungii Kuntze, Deutsche Bot. Monatsschr. 21: 173. 1903. Type.
KAISER-WILHELMSLAND [PAPUA NEW GUINEA]. Augusta-Station, Aug.
1887, Hollrung 689 (holotype: B, destroyed; isotypes not found).
15. Cynometra schefferi var. peekelii (Harms) Rados., comb. nov.
urn:lsid:ipni.org:names:77199257-1
Basionym. Maniltoa peekelii Harms, Bot. Jahrb. Syst. 55 (1): 50. 1917. Syn. Maniltoa
schefferi var. peekelii (Harms) Verdc. Type. [PAPUA NEW GUINEA]. Neu-Mecklenburg (New Ireland): Lemakot, Garamate, Strandebene, July 1912, G. Peekel 841 (holotype: B, destroyed). PAPUA NEW GUINEA. New Ireland District [Province]: [Island
of New Ireland], Kavieng Sub-District [District], ca. 26 miles from Kavieng, inland
from Lavongai, 2°46'S, 151°2'E, 0 m alt., 24 Jan. 1967, M.J.E. Coode, T. Cropley, &
P. Katik NGF 29604 (neotype, here designated: K; isoneotypes: A, CANB, L, LAE).
Note. When Verdcourt (1977) published the above name at a new rank, he did
not name a neotype. He did cite several representative specimens and specifically men-
The rise of Cynometra and the fall of Maniltoa: a generic re-circumscription and...
15
Figure 2. Illustration of Maniltoa megalocephala Harms, reproduced from protologue and designated
here as the lectotype.
16
Aleksandar Radosavljevic / PhytoKeys 127: 1–37 (2019)
tioned Coode et al NGF 29604 as a specimen he was certain belonged to this species.
Given that it is well represented in herbaria and was collected near the original type
locality, this specimen has been chosen as the neotype.
16. Cynometra steenisii (van Meeuwen) Rados., comb. nov.
urn:lsid:ipni.org:names:77199258-1
Basionym. Maniltoa steenisii van Meeuwen, Blumea 18 (1): 40. 1970. Type. TERRITORY OF PAPUA [PAPUA NEW GUINEA]. Northern District [Northern Province]: Tufi subdistrict, near Budi Barracks, 9°32'S, 148°58'E, 75 m alt., 26 August
1954, R.D. Hoogland 4581 (holotype: CANB).
Notes. Knaap-van Meeuwen incorrectly cites the holotype as being housed in BRI.
17. Cynometra steenisii var. rodneyensis (Verdc.) Rados., comb. nov.
urn:lsid:ipni.org:names:77199266-1
Basionym. Maniltoa steensisii var. rodneyensis Verdc., Kew Bull. 32 (1): 241. 1977.
Type. TERRITORY OF PAPUA [PAPUA NEW GUINEA]. Central District [Central
Province]: Abau Subdistrict [Abau District], Cape Rodney, Mori River, 200 ft alt., 20
June 1968, E. E. Henty NGF 38561 (holotype: LAE; isotypes: A, BISH, BO, BRI,
CANB, K, L, NSW, SING).
18. Cynometra vestita (A.C.Sm.) Rados., comb. nov.
urn:lsid:ipni.org:names:77199267-1
Basionym. Maniltoa vestita A.C.Sm., J. Arnold Arbor. 31: 170. 1950. Type. FIJI. Vanua Levu. Mathuata [Macuata] province: east of Lambasa [Labasa], on the southern
slopes of Mt Numbuiloa, 3 Nov 1947, [fl], A.C. Smith 6442 (holotype: A; isotypes:
BISH, BRI, K, L, LE, NY, P, S, US).
19. Cynometra vitiensis Rados., nom. nov.
urn:lsid:ipni.org:names:77199269-1
Basionym. Maniltoa floribunda A.C.Sm., J. Arnold Arbor. 31: 169. 1950, non Cynometra floribunda Benth, Trans Linn. Soc. London 25: 318. 1865. Type. FIJI. Viti
Levu. Nandronga-Navosa Province: southern slopes of Nausori Highlands, in drainage
of Namosi Creek above Tumbenasolo, 300–400 m alt., 29 May 1947, A.C. Smith 4588
(holotype: A; isotypes: BISH, BRI, K, L, LE, NY, P, S, US).
Notes. This species is named after the nation of Fiji, where it is endemic, but somewhat common and widespread, occurring on at least six islands (Smith 1985).
The rise of Cynometra and the fall of Maniltoa: a generic re-circumscription and...
17
Part III: Four new species of Cynometra
A monograph of the Neotropical species of Cynometra was published by Dwyer (1958),
but it was limited by the lack of fertile material. In the intervening decades, however,
many new collections have been made, particularly in the Amazon basin, the centre of
diversity of the genus in the New World. This has made revisionary work more feasible
and uncovered previously undocumented diversity. In the context of the ongoing taxonomic and phylogenetic studies, several new species have been discovered in existing
herbarium collections. Four new species are described below: Cynometra cerebriformis
sp. nov. from the lower Rio Trombetas in Para State, Brazil; Cynometra dwyerii sp. nov.
from the Darien gap region of Panama; C. tumbesiana sp. nov. from the dry tropical
forests of Ecuador and Peru; and C. steyermarkii sp. nov. from the foothills of the western Cordillera de la Costa in Venezuela (Figure 3).
1. Cynometra cerebriformis Rados., sp. nov.
urn:lsid:ipni.org:names:77199270-1
Figures 4, 5
Type. BRAZIL. Pará: [Mun. Oriximiná] Rio Trombetas, near Cachoeira Porteira, 24
May 1974, [fl.], D. G. Campbell et al., P22338 (holotype US; isotype F, INPA, MO, NY).
Description. Tree to 20 m tall; bark not seen; branchlets rough, lenticels prominent, bud scale scars partially encircling stems near base, new growth with short
scattered pubescence, glabrous or nearly so at maturity. Stipules not seen. Leaves bifoliolate, pulvinate, axes glabrous or with sparse pubescence concentrated on adaxial
surfaces; petioles 3.0–4.0 mm long, 1.0 mm wide, transversely corrugated; petiolules
0.5–1.0 mm long, 0.5 mm wide, transversely corrugated; leaflets appearing sessile,
coriaceous, obelliptic to obovate, asymmetric, primary vein eccentric, proximal side
2.0–2.7 times wider than distal, 3.1–4.5 cm long, 1.1–1.9 cm wide, abaxial surface
with scattered raised areas both surfaces glabrous, primary venation pinnate, secondary venation brochidodromous-eucamptodromous, 3–4 basal acrodromous veins,
decurrent to primary vein, prominent abaxially, their course barely visible adaxially,
tertiary venation difficult to discern on either surface even under magnification,
margins entire, apex acute, usually short acuminate (acumen to 4.0 mm), retuse,
mucronate, base oblique, acute, distal side narrowly cuneate with margin nearly parallel to midvein for 8.0–10.0 mm, proximal side convex, decurrent to petiolule,
laminar glands absent. Inflorescence an axillary raceme, (1–)2 per axil, bracteate,
axes ferrugino-pilose; peduncle 1.0–2.0 mm, rachis 2.0–5.0 mm long, flowers spirally arranged, 2–10 per raceme; pedicels 7.0–10.0 mm and filamentous in anthesis,
to 18.0 mm and accrescent in fruit; bracts subtending individual flowers, scale-like,
quickly deciduous, brown, broadly elliptical to deltoid, strongly convex 1.5–2.5 mm
long, 1.0–1.5 mm wide, striate, abaxial surface pubescent, pubescence denser at
base and along margins, glabrous adaxially; bracteoles not seen. Flowers bisexual,
radially symmetric, pentamerous, delicate; hypanthium cupular, 0.7–1.0 mm deep,
18
Aleksandar Radosavljevic / PhytoKeys 127: 1–37 (2019)
Figure 3. Distribution of Cynometra cerebriformis, C. dwyerii, C. steyermarkii and C. tumbesiana.
surrounding basal portion of ovary, fleshy, abaxial surface pubescent, adaxial surface
glabrous; sepals 4, imbricate, reflexed, abaxial and adaxial sepals larger than lateral,
greenish-white, petaloid, oblong to elliptic, apices acute to rounded, 3.0–4.0 mm
long, 1.0–2.0 mm wide, pubescence on abaxial surface near apex, with faint parallel
venation; petals 5, equal, white, crumpled texture, curving inwards, oblanceolate,
4.0–5.0 mm long, 1.0–1.5 mm wide, adaxial surface with minute appressed hairs,
pinnate venation; stamens 10, filaments free, subequal, 7.0–8.5 mm long, anthers
dorsifixed, versatile, longitudinal dehiscence, ellipsoid, to 1.0 mm long, glabrous;
ovary centrally inserted, free, stipitate, obliquely elliptical, 2.5–3.0 mm long, 1.5–
2.0 mm wide, tomentose, stipe 0.5 mm, style apical, 2.0–2.5 mm long, glabrous,
eccentric, curving downwards, stigma capitate. Legume indehiscent, oblate, rugose,
apiculate when immature, 14.2–17.5 mm long, 7.1–9.2 mm wide, 12.1–13.9 mm
thick, valves pubescent, wall of pericarp up to 2.5 mm thick, brown. Seeds 1 per
pod, filling locule, dark brown.
Distribution and ecology. Currently, this species appears restricted to the Trombetas River Basin of Para State, Brazil. However, it is quite possible that the range is
more extensive given that the species occurs in seasonally flooded forests and the genus
is known for water dispersed fruits (Ridley 1930, Clarke et al. 2001, Tomlinson 2016).
Little else is known about this taxon.
Phenology. Flowering specimens have been collected in May. Fruiting specimens
have been collected in June and August.
The rise of Cynometra and the fall of Maniltoa: a generic re-circumscription and...
19
Figure 4. Illustration of Cynometra cerebriformis. A Habit B bud with imbricate bracts C leaf base showing corrugated petiole and petiolule D inflorescence rachis showing bract scars on main axis and remnants
of bracteoles on pedicels E flower F infructescence with single pod G bract H sepal I petal J longitudinal
section of hypanthium and receptacle; sepals, petals and stamens removed. A–E, G–J Campbell et al.
P22338, US F G. Martinelli 7016, US.
20
Aleksandar Radosavljevic / PhytoKeys 127: 1–37 (2019)
Figure 5. Photograph of the holotype of Cynometra cerebriformis (Campbell et al. P22338, US).
Etymology. Cynometra cerebriformis is named after the brain-like appearance of
the mature fruit.
Additional specimens examined. BRAZIL. Pará: Municipio Oriximiná: Rio
Trombetas shore, Porteira cemetery, 3 Jun 1974 [im fr], D. G. Campbell et al. P22510
The rise of Cynometra and the fall of Maniltoa: a generic re-circumscription and...
21
(INPA, NY, US); Margem direita do Rio Mapuera, entre as Cach[oeira] Paraiso
Grande e Maracajá, Área do reservatório da 2a etapa, 00°58'S, 57°35'W, 12 Aug 1986
[im fr], C.A. Cid Ferreira et al. 7659 (INPA, NY); Rio Trombetas, river banks downriver from Cachoeira Porteira, N to NE bank (Between C.P. and IBDF Reverval – Lago
do Jacaré), 17 Jun 1980 [fr], C. Davidson & G. Martinelli 10324 (INPA, NY, US); Rio
Trombetas, margem esq. entre o Lago Jacaré e Cachoeira Porteira, 70 m alt., 17 Jun
1980 [fr], G. Martinelli 7016 (INPA, NY, RB).
Notes. This taxon has been collected in the areas around Santarem and Oriximiná
in Para, Brazil. Many of the specimens have been annotated by Adalea Sprada Tavares
as Cynometra duckei ssp. trombetensis, but I can find no record of publication and several
of the specimens thus annotated are assigned to different taxa in her unpublished thesis
(Sprada Tavares 1987). The available material of C. duckei is limited and only a few fruiting specimens exist. While a case can be made that C. duckei shares certain vegetative
traits with C. cerebriformis (prominent lenticels, the smoothness of the adaxial surface of
the leaflets), it also shares characteristics with several other taxa, including C. spruceana
var. spruceana (long pedicels, leaflet shape) and C. marginata var. laevis (smooth leaflet
surface, nearly sessile leaflets). In light of the characters separating C. cerebriformis from
other taxa (see below), the author has opted to describe this taxon at the species level.
Cynometra cerebriformis differs from C. duckei primarily in the shape of the leaflets. Cynometra cerebriformis differs from C. spruceana var. spruceana in several ways.
The leaflets of C. cerebriformis are generally smaller than those of C. spruceana var.
spruceana and the surface is nearly smooth, while the secondary veins are quite obvious
in C. spruceana. Cynometra cerebriformis also lacks the basal laminar gland present in C.
spruceana and many other Cynometra taxa. Finally, the fruit of C. spruceana is approximately 1.5×–2.5× larger than the fruit of C. cerebriformis and the valves are smooth to
slightly rugulose, lacking the strongly rugose surface of C. cerebriformis.
Cynometra cerebriformis differs from C. marginata var. laevis by having an acute
leaflet base, short acumen and oblate rugose fruit; C. marginata var. laevis has an obtuse
leaflet base, long acumen and a globose fruit with a prominent raised suture ridge.
2. Cynometra dwyerii Rados., sp. nov.
urn:lsid:ipni.org:names:77199272-1
Figures 6, 7
Type. PANAMA. Darién [now Comarca Emberá-Wounaan]: vicinity of Campamento Buena Vista, Río Chucunaque above confluence with Río Tuquesa, [08°23'N,
77°47'W] 5 July 1959, [fr.], W. L. Stern 941 (holotype US; isotype MO).
Description. Tree to approximately 20 m tall; bark not seen; branchlets lenticelate, pubescent when young, becoming glabrous with age. Stipules not seen. Leaves
bifoliolate, axes pubescent, transversely corrugated; petioles 4.5–5.5 mm long; petiolules 1.0–1.5 mm long, inconspicuous, leaflets appearing sessile; leaflets narrowly
obovate to obovate, occasionally sub-trapeziform, strongly asymmetric, primary vein
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Aleksandar Radosavljevic / PhytoKeys 127: 1–37 (2019)
Figure 6. Illustration of Cynometra dwyerii. A Leaf base (adaxial surface) showing corrugated petiole
and pubescence along midrib B habit C leaf base (abaxial surface) showing basal acrodromous veins arising from leaflet pulvinus and laminar glands D leaflet apex (abaxial surface) E Dissected fruit, proximal
surface F dissected fruit, distal surface G longitudinal section of fruit with single seed H reconstruction
of fruit. A–H Stern et al. 941, US.
The rise of Cynometra and the fall of Maniltoa: a generic re-circumscription and...
Figure 7. Photograph of the holotype of Cynometra dwyerii (Stern et al. 941, US).
23
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Aleksandar Radosavljevic / PhytoKeys 127: 1–37 (2019)
eccentric, proximal side 2.8–3.7 times wider than distal, 3.1–3.9 cm long, 1.4–1.9 cm
wide, thin, abaxial surface sparsely pubescent, more so on midvein and major secondaries, adaxial surface with pubescence restricted primarily to midvein, occasional hairs
scattered on lamina, primary venation pinnate, secondary venation brochidodromouseucamptodromous, 2(–3) basal acrodromous veins, decurrent to primary, prominent
abaxially, only slightly less so adaxially, tertiary venation visible on both surfaces at
10× magnification, margins entire, apex acute, weakly acuminate (to 2.0 mm), retuse,
mucronate, base oblique, acute, distal side narrowly cuneate, proximal side slightly
concave to cuneate, decurrent to petiolule, laminar glands present, 3–6 per leaflet,
arranged in a row approximately halfway between margin and midvein, restricted to
distal portion of lamina, typically adjacent to tertiary veins, crateriform, less than 1.0
mm in diameter. Inflorescences not seen, position inferred as axillary from remnant
of peduncle. Flowers not seen. Legume indehiscent, roughly globose, to 4.7 cm in
diameter, surface of valves rugulose, wall of pericarp up to 3.0 mm thick, deep brown
colour at maturity. Seeds 1 per pod, filling entire cavity, dark brown.
Distribution and ecology. Known only from the type locality in the Darien Gap
region of Panama; the area where the type was collected is primarily lowland moist
tropical forest.
Phenology. The type was collected with mature fruit in July.
Etymology. The specific epithet honours the contributions of Dr. John Dwyer,
who published the first monograph of the Neotropical species of Cynometra.
Note. This species is known only from the type collection, however the combination of vegetative characters and fruit morphology make it clearly distinct from other
Cynometra species. Superficially, this species resembles C. bauhiniifolia, given its small
leaflets with prominent secondary venation. However, the pubescence and arrangement of laminar glands clearly distinguish it from other Neotropical species of Cynometra, which usually have just a single, basal laminar gland. In fact, the combination
of bifoliolate leaves with small leaflets and several submarginal laminar glands is unique
across the entire genus. When these characters are combined with the large, globose
fruit, it is clear that this is a distinct species.
Additional specimens examined. None.
3. Cynometra steyermarkii Rados., sp. nov.
urn:lsid:ipni.org:names:77199276-1
Figures 8–10
Type. VENEZUELA. Miranda: Distrito Brión, Selva siempre verde a lo largo de la
quebrada afluente del río Aricagua, 3.9 km oeste del Pueblo Seco, 1.6 km oeste de Aricagua, 75 m alt., 24–25 March 1973, [fl, fr], J. A. Steyermark & V. Carreño Espinoza,
106937 (holotype: US; isotypes: F, VEN n.v.).
Description. Tree to 25 m tall; bark not seen, sapwood reddish; branchlets glabrous, lenticelate. Stipules not seen. Leaves bifoliolate, axes glabrous; petioles 6.5–
The rise of Cynometra and the fall of Maniltoa: a generic re-circumscription and...
25
9.0 mm long, 2.0 mm wide, transversely corrugated; petiolules 2.0–3.0 mm long,
1.0 mm wide, transversely corrugated; leaflets, coriaceous, elliptic to slightly obovate,
asymmetric, primary vein eccentric, proximal side 2.3–3.1 times wider than distal,
5.7–7.1 cm long, 2.7–3.6 cm wide, discolorous, abaxial and adaxial surface glabrous,
primary venation pinnate, secondary venation brochidodromous-eucamptodromous,
2(–3) basal acrodromous veins, decurrent to primary vein, prominent abaxially, slightly raised adaxially, tertiary venation visible on abaxial surface without magnification,
margins entire, apex obtuse, usually rounded but occasionally acuminate (to 3.0 mm),
retuse, mucronate, base oblique, acute, distal side strongly cuneate, proximal concave
to convex, decurrent to petiolule, single laminar gland present on some leaflets, abaxial, near basal margin of proximal lamina and insertion point of petiolule, typically
adjacent to tertiary veins, crateriform, 0.5 mm in diameter. Inflorescence an axillary
raceme, bracteate, axes densely ferrugino-puberulent; peduncle together with rachis
to 6.0 mm long, flowers spirally arranged, 15 per raceme; pedicels 5.5–6.0 mm, pubescent, accrescent in fruit; bracts subtending individual flowers, scale-like, deciduous, lustrous, brown, broadly elliptical, strongly convex 1.0–2.0 mm long, 1.5–2.5
mm wide, striate, abaxial surface with appressed pubescence along medial surface and
along margins, glabrous adaxially; bracteoles paired, inserted near base of pedicel,
subopposite, linear to oblanceolate, 2.5–3.0 mm long, 0.75 mm wide, acute at apex,
densely pubescent on abaxial surface, less so adaxially. Flowers bisexual, radially symmetric, pentamerous, delicate; hypanthium much reduced, less than 0.5 mm deep,
with a few scattered hairs on abaxial and adaxial surface; sepals 4, imbricate, reflexed,
slightly unequal, white, petaloid, oblong to obovate, 2.5–3.5 mm long, 1.0–1.5 mm
wide, faintly striate, appressed pubescence on abaxial surface near insertion point and
along central axis; petals 5, equal, white, oblanceolate, 3.5–4.5 mm long, 1.0–1.5 mm
wide, pinnate venation; stamens 10, filaments appearing free but possibly connate
for approximately 0.25 mm at base, subequal, 4.0–5.5 mm long, anthers dorsifixed,
versatile, longitudinal dehiscence, ellipsoid, to 0.75 mm long, glabrous; ovary centrally inserted, free, sessile, obliquely elliptical, 2.5–4.0 mm long, 1.5–3.0 mm wide,
pubescent at base and along suture, becoming glabrous with age, style apical, 1.5–2.5
mm long, glabrous, eccentric, stigma capitate. Legume indehiscent, globose, shortly
apiculate (to 0.75 mm), 1.5 cm in diameter, surface of granulose, with very short pubescence, wall of pericarp up to 2.0 mm thick, brown. Seeds 1 per pod (1 ovule per
ovary based on dissections), dark brown.
Distribution and ecology. Known from a single collection made in Miranda
State, Venezuela. The species was collected growing along a small stream in the foothills of the Cordillera de la Costa at approximately 75 m above sea level. While much
of this region is covered in xeric shrublands and thorn forests, this site corresponds to
a low gallery forest growing along a tributary of the Río Aricagua.
Phenology. The type material was collected at the end of March with late flowers
and nearly mature fruit.
Etymology. The specific epithet honours Dr. Julian A. Steyermark, the collector of
the type material and a prolific collector of neotropical plants. Over the course of his
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Aleksandar Radosavljevic / PhytoKeys 127: 1–37 (2019)
Figure 8. Illustration of Cynometra steyermarkii. A Habit, flowering branch B habit, fruiting branch
C leaf base, showing basal acrodromous veins arising from leaflet pulvinus D inflorescence rachis, showing
bracts and bracteoles; pedicels removed to show structure E buds with imbricate bracts F flower G bract
H bracteoles I sepals J petal K longitudinal section of flower showing single ovule and much reduced
hypanthium; sepals, petals and stamens removed. A–K Steyermark et al. 106937, US.
The rise of Cynometra and the fall of Maniltoa: a generic re-circumscription and...
27
Figure 9. Photograph of the holotype of Cynometra steyermarkii, sheet one of two (Steyermark et al.
106937, US).
career, he made over 130,000 collections and greatly advanced our knowledge of the
Venezuelan flora.
Additional specimens examined. None.
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Aleksandar Radosavljevic / PhytoKeys 127: 1–37 (2019)
Figure 10. Photograph of the holotype of Cynometra steyermarkii, sheet two of two (Steyermark et al.
106937, US).
Notes. Cynometra steyermarkii is the only confirmed species of Cynometra known
from the forests of the Cordillera de la Costa in northern Venezuela and one of two
species of Venezuelan Cynometra growing outside of the Guayana region. Henri Pittier
The rise of Cynometra and the fall of Maniltoa: a generic re-circumscription and...
29
(1926) published an account of a Cynometra growing in the hills outside of Caracas,
Cynometra sphaerocarpa Pittier, for his Manual de las plantas usuales de Venezuela. While
the description is valid, according to the rules of nomenclature in effect at the time,
he failed to designate a type or list any collections that may correspond to this taxon.
In his 1958 revision, Dwyer was unable to locate any material that could be assigned
to this taxon and pointed out the limited utility of Pittier’s description – ‘Las hojas
inequilaterales, lanceadas, largamente atenuadas y glabras… Los frutos son subglobosos, de 3.5–4 cm. de diametro y contienen una sola semilla [Leaflets asymmetric,
lanceolate, largely attenuate and glabrous. The fruits are sub-globose, 3.5–4 cm in
diameter, and contain a single seed].’– which could very well describe any number of
Cynometra species, including C. steyermarkii. A full set of Pittier’s duplicates from this
publication should be deposited in the United States National Herbarium (US), but
after exhaustive searches in the US collections and a study of both Pittier’s archives
and the museum registrar’s records, I was unable to find any material that could be
attributable to C. sphaerocarpa. Curators at Herbario Nacional de Venezuela graciously
searched through material housed there, but were also unsuccessful. While it is possible
that C. steyermarkii and C. sphaerocarpa are the same taxon, without Pittier’s original
material, it is impossible to know for sure. Rather than leave this unresolved, it is more
useful to describe a new species with good type material.
This species bears a resemblance to Cynometra spruceana var. phaselocarpa Benth.
owing to the obtuse nature of the leaf apices. However, C. steyermarkii often has slightly acuminate apices. Furthermore, both the rachis and pedicels of the inflorescence are
much shorter in C. steyermarkii than in typical C. spruceana var. phaselocarpa.
4. Cynometra tumbesiana Rados., sp. nov.
urn:lsid:ipni.org:names:77199277-1
Figures 11, 12
Type. ECUADOR. El Oro: Bosque Petrificado Puyango, dirt track from information
centre towards the camping area near Río Puyango, 03°52'30"S, 80°05'01"W, 450 m
alt., 6 May 1997 [fl], B.B.Klitgaard et al. 507, (holotype K; isotype AAU n.v., LOJA
n.v., NY, QCNE n.v., US).
Description. Tree 10–25 m tall; bark grey-brown, lenticelate, inner bark red;
branchlets with short pubescence when young, becoming glabrous with age. Stipules
not seen. Leaves bifoliolate, axes ferrugino-puberulent when young, glabrous when
mature; petioles 4.5–6.5 mm long, transversely corrugated; petiolules 1.5–2.0 mm
long, inconspicuous; leaflets oblong-ovate to elliptic to oblong-obovate, occasionally slightly falcate or sub-trapeziform, strongly asymmetric, primary vein eccentric,
proximal side 1.8–2.5 times wider than distal, 4.1–7.9 cm long, 2.5–3.3 cm wide,
discolorous, abaxial surface sparsely pubescent on midvein, secondary veins and along
basal margin, adaxial surface glabrous, primary venation pinnate, secondary venation
brochidodromous-eucamptodromous, 2–3 (–4) basal acrodromous veins, decurrent to
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Aleksandar Radosavljevic / PhytoKeys 127: 1–37 (2019)
primary vein, prominent abaxially, flush to slightly raised adaxially, tertiary venation
visible on abaxial surface at 10× magnification, margins entire, apex acute, acuminate
(to 6.0 mm), retuse, mucronate, base oblique, distal side acute, convex to cuneate,
proximal side obtuse, concave to rounded, decurrent to petiolule, single laminar gland
present, abaxial, near basal margin of proximal lamina and insertion point of petiolule,
typically adjacent to tertiary veins, crateriform, 1.0 mm in diameter. Inflorescence a
cluster of (1–)2–3 axillary racemes, bracteate, axes ferrugino-puberulent at base, hairs
becoming scattered at distal end; peduncle together with rachis 4.5–8.0 mm long, flowers spirally arranged, 12–20 per raceme; pedicels 5.0–9.0 mm, pubescent initially but
soon glabrescent, accrescent in fruit; bracts subtending individual flowers, scale-like,
deciduous, leaving behind a lunate scar on the rachis, lustrous, brown, broadly elliptical, strongly convex 1.5–2.5 mm long, 1.5–2.5 mm wide, striate, abaxial surface with
scattered appressed pubescence at apex and along margins, glabrous adaxially; paired
bracteoles inserted 0.5–1.0 mm from base of pedicel, subopposite, oblong-lanceolate,
2.0–2.5 mm long, 1.0 mm wide, convex at apex, pubescent along margins and medial
abaxial surface. Flowers bisexual, radially symmetric, pentamerous, delicate; hypanthium cupular, 1.0–1.5 mm deep, fleshy, with a few scattered hairs on abaxial and adaxial
surface; sepals 4, imbricate, reflexed, unequal, adaxial sepal usually 2 times as wide as
the others, white, petaloid, broadly ovate to elliptical, 3.0–4.5 mm long, 1.5–4.0 mm
wide, striate, scattered pubescence at base; petals 5, erect, equal to subequal, white,
spathulate to oblanceolate, 3.5–5.5 mm long, 1.0–2.0 mm wide, glabrous but with
a tuft of hair at base of claw; stamens 10, filaments free, subequal, 5.5–7.5 mm long,
white, anthers dorsifixed, versatile, longitudinal dehiscence, suborbicular, to 1.5 mm
long, yellow-orange, glabrous; ovary centrally inserted in hypanthium, free, stipitate,
obliquely elliptical, 4.0–5.0 mm long, 2.0–2.5 mm wide, densely pilose, stipe 0.5–1.0
mm long, style apical, 3.0–4.0 mm long, glabrous, eccentric, geniculate, stigma capitate. Legume indehiscent, oblong, weakly apiculate, slightly compressed, up to 5.2 cm
long, 4.0 cm wide, 3.9 cm thick, surface of valves finely textured, granulose, wall of
pericarp up to 4.0 mm thick, deep brown colour at maturity. Seeds 1 per pod, filling
entire cavity, dark brown.
Distribution and ecology. Cynometra tumbesiana occurs in the seasonally dry
tropical forests of western and southern Ecuador and north-western Peru (a single
collection) at elevations between 100–800 m (Figure 3). These habitats are severely
threatened regionally and globally due to extensive human modification of the landscape and, as a result, this species now exists primarily as isolated fragments. While C.
tumbesiana is locally abundant at a few sites, it is currently known from less than 10
localities, several of which are within 5 km of each other.
Phenology. Flowering specimens have been collected in May and December to
January; fruiting specimens have been collected in August and January to February.
Cynometra tumbesiana is one of the few woody taxa in the dry forests that retain their
leaves during the dry season.
Etymology. The specific epithet refers to the Tumbes region, where the type specimen was collected and where many of the known localities occur.
The rise of Cynometra and the fall of Maniltoa: a generic re-circumscription and...
31
Figure 11. Illustration of Cynometra tumbesiana. A Habit B leaflet base, showing laminar gland C flower
D inflorescence rachis, showing bracts and bracteoles; pedicels removed to show structure E fruit, dissected to show single seed filling entire cavity. A–D Klitgaard et al. 507, K; B Neill & Núñez 10453, US.
Additional specimens examined. ECUADOR. El Oro: Bosque Petrificado Puyango, dirt track from information centre towards the camping area near Río Puyango,
03°52'30"S, 80°05'01"W, 450 m alt., 23 Aug 1996 [fr], B.B. Klitgaard et al. 325 (AAU
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Aleksandar Radosavljevic / PhytoKeys 127: 1–37 (2019)
n.v., K, LOJA n.v., NY, QCNE n.v.); 26 Feb 1997 [fr], B.B. Klitgaard et al. 424 (AAU
n.v., K, LOJA n.v., NY, QCNE n.v.). [Piñas]: Piedras, about 3 km. along new trail, 18
Jun 1943 [st], E.L. Little, Jr. 6622 (US). Guayas: [without specific locality] 2 Feb 1962
[fr], A.J. Gilmartin 551 (US). Guayaquil: Bosque Protector Cerro Blanco, 15 km west
of Guayaquil, summit area of Cerro Blanco, 2°10'S, 79°58'W, 370 m alt., 27 Feb 1996
[fr], D. Neill & T. Núñez 10453 (MO, US); Bosque Protector Cerro Blanco, along
road from visitor centre to “Cusumbo Top”, 80, 01 W, 2 10 S, 400 m alt., 7 Aug 1996
[im fr], D. Neill, T. Núñez & J. Machuca 10636 (MO); Bosque Protector Cerro Blanco, carretera a Salinas, km 15, 2°10'S, 79°58'W, 400 m alt., 21–25 Jan 1992 [fr], D.
Rubio & Galo Tipaz 2365 (MO). Isidro Ayora: Reserva Ecológica Manglares Churute,
carretera Guayaquil–Puerto Inca, sector norte del Cerro Masvale, 2°20'S, 79°50'W,
200–300 m alt., May 1993 [fl], T. Núñez & A. Hernández 147 (MO). Manabi: [Puerto
López]: Estero Perro Muerto, Machalilla National Park, below San Sebastian, 1°36'S,
80°42'W, 400–420 m alt., 23 Jan 1991 [fl], A. Gentry & C. Josse 72677 (MO); [San
Vicente]: [hacienda] El Recreo, [fl], H.F.A. von Eggers 15752 (US). PERU. Tumbes:
Zarumilla: Dtto. Matapalo, Campo Verde a 68 km de. Tumbes, 700–800 m alt., 24
Dec 1967 [fr], J. Schunke V. 2411 (F, NY, US).
Notes. This species, restricted to the few remaining fragments of dry tropical forest
in western Ecuador and the Tumbes region of Peru, has been mistakenly referred to as
Cynometra crassifolia Benth. for many years. However, closer examination shows it to
be quite distinct from this taxon. The type specimen of C. crassifolia was collected in
Brazil by Portuguese naturalist Alexandre Rodriques Ferreira during his exploration of
the Amazonian region of Brazil from 1783–1792. His collections, along with many
others housed at Lisbon, were expropriated by Étienne Geoffroy Saint-Hilaire and
transferred to Paris during Napoleon’s occupation of Portugal. There, it was seen by
George Bentham, who described Cynometra crassifolia in 1840. The primary differences
between C. tumbesiana and C. crassifolia are found in the inflorescences: the racemes of
C. crassifolia have larger flowers, longer pedicels and a more robust pedicel and rachis,
but fewer individual flowers than those of C. tumbesiana. The flowers of C. tumbesiana are indeed relatively small compared to the other neotropical Cynometra species,
though they are densely clustered on the short rachis of the inflorescence. The leaflets
of C. tumbesiana are also less distinctly acuminate than those of C. crassifolia and have a
less obtuse base; the base of the leaflet in C. crassifolia can appear to be almost truncate.
Some taxonomists have placed the Ecuadoran Cynometra within C. bauhiniifolia
Benth., with a few treating C. crassifolia as a synonym of C. bauhiniifolia (Neill et al.
1999). In the first case, while C. tumbesiana does bear a passing resemblance to some
forms of C. bauhiniifolia, the pod of C. bauhiniifolia is many times smaller and the
surface of the valves is corky and deeply rugose. In the latter case, it is difficult to find
justification for synonymising the two with the exception that C. bauhiniifolia has
been a dumping ground for hard to place taxa within the genus and does occasionally
occur in drier habitats. The leaflets, inflorescences and fruits all differ. Instead, the type
material of C. crassifolia seems to be very similar to C. longicuspis Ducke, a widespread
species from the moist lowland forests of Brazil.
The rise of Cynometra and the fall of Maniltoa: a generic re-circumscription and...
Figure 12. Photograph of the holotype of Cynometra tumbesiana (Klitgaard et al. 507, K).
33
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Aleksandar Radosavljevic / PhytoKeys 127: 1–37 (2019)
Cynometra tumbesiana is morphologically and ecologically similar to C. oaxacana
Brandegee from western and southern Mexico. The two can be distinguished by the
narrower and more acuminate leaflets and larger fruits in C. tumbesiana. The inflorescences of C. oaxacana are also slightly more robust and have a more obvious pubescence. Both species are found in dry habitats (uncommon amongst the neotropical
species of Cynometra), though C. tumbesiana is found in much drier sites.
Acknowledgements
I thank the directors, curators and support staff at the following herbaria for allowing me to view and borrow specimens: A, ATH, BM, BRI, CANB, F, IAN, INPA,
K, MG, MO, NY, P, RB, SING and US. I am very grateful to my thesis advisors
Vicki Funk (US) and Patrick Herendeen (CHC) for their suggestions to improve the
manuscript. Kenneth Wurdack (US) and Warren Wagner (US) provided invaluable
feedback during the preparation of this manuscript. I thank Alice Tangerini (US)
and her students, Mattias Lanas and Katherine Rudebusch for providing detailed
and illuminating illustrations of the new species. This project was conducted as part
of my doctoral research at the Chicago Botanic Garden, Northwestern University
and the National Museum of Natural History. The research was funded by grants
and fellowships from the American Society of Plant Taxonomists, Garden Clubs of
America, the Hunt Institute, the National Science Foundation (Doctoral Dissertation Improvement Grant DEB-1501199; co-PI Patrick Herendeen) and the Smithsonian Institution.
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