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A new species of Securidaca (Polygalaceae, Polygaleae) from Araracuara region, middle Caquetá river basin, Colombia Una nueva especie de Securidaca (Polygalaceae, Polygaleae) de la región de Araracuara, cuenca media del Río Caquetá, Colombia Gerardo A. Aymard C.1
ORCID: 0000-0001-9405-0508
Levi Andoque Andoque2
1UNELLEZ-Guanare, Programa de Ciencias del Agro y el Mar, Herbario Universitario (PORT), Mesa de Cavacas, estado Portuguesa 3350, Venezuela; Jardín Botánico de Bogotá José Celestino Mutis, Cl. 63 #68-95, Bogotá DC., Colombia. gerardo.aymard@gmail.com
2Comunidad Indígena San José, Clan Sol, Caño Aduche, departamento del Amazonas. Colombia.
Recibido: 28-12-2024
Aprobado: 12-02-2025
Publicado: 31-03-2025
Artículo de investigación
ABSTRACT
for identifying species of Securidaca to Colombia is also provided. Securidaca
Securidaca poeoesioehoeorum from poeoesioehoeorum is remarkable Araracuara region, at the middle for its four stamens, in an otherwise Caquetá River basin (northwestern predominantly eight stamens genus.
Amazon basin), Caquetá department, This new finding increases to 27 the Colombia, is described, illustrated, and number of species of the genus to its morphological relationships with an Colombia flora. This new taxon is the allied species are discussed. The new first plant species name that honors the species is similar to S. leiocarpa but Pøøsiøhø nation.
differs by the pubescence, shape, and size
of its leaves, outer and lateral enlarged Keywords: Araracuara,
Caquetá
sepals, the inflorescence, the keel petal, River basin, secondary wet forests, stamens number and fruit shape and size. Securidaca, Polygalaceae, Pøøsiøhø, A complete description, an illustration, Colombia information about habitat, phenology,
conservation status, the species name RESUMEN
etymology and an abstract in Pøøsiøhø Securidaca poeoesioehoeorum de la (Andoke) idiom are presented. A key región de Araracuara, cuenca media
Aymard C. y Andoque A.: Securidaca (Polygalaceae), especie nueva, Río Caquetá del Río Caquetá (noroeste de la cuenca nie ka dozkoi siọka ø įpøkø bø ni+ ke Amazónica), departamento del Caquetá, į´į. høma o´defø katpa oę høka. øñe bø Colombia, es descrita, ilustrada y sus keÁ pøsiøhø kaśofa´.
relaciones morfológicas con su especie
afín son discutidas. La nueva especie INTRODUCTION
es similar a S. leiocarpa pero difiere Securidaca L., nom. cons. is en el tamaño, forma y pubescencia a pantropical genus of Polygalaceae de las hojas, los sépalos externos y of about 67 species, of which ca. 61
laterales, las inflorescencias, la quilla (including the new species described (el pétalo inferior), el número de herein) are found in the Neotropics estambres y en la forma y tamaño de (Eriksen et al. 2000, Aymard et al. 2004, los frutos. Se presenta una descripción, Aymard 2022). A few species are found una ilustración, información acerca in Africa and Southeast Asia; the genus del hábitat, fenología, estado actual de is absent in Australia (World Checklist conservación, etimología del nombre of Vascular Plants; kew.org). In the de la especie, un resumen en el idioma Neotropics, Securidaca is most diverse en Pøøsiøhø (Andoke) y una clave de throughout the Amazon (presumed las especies del género en Colombia. centre of its diversification, sensu Securidaca poeoesioehoeorum es Anjos et al. 2024) and Orinoco River una especie notable por sus poseer basins, the Guayana Shield (Marques, cuatro estambres, en un género donde 1996, Aymard et al. 2004) and Andes predominan ocho estambres. Este bioregions.
nuevo hallazgo eleva a 27 el número
de especies del género para la flora de
Several species are found over
Colombia. Este nuevo taxon representa rocky slopes and oligotrophic soils la primera especie de planta que hace derived from the Precambrian crystalline honor a la nación Pøøsiøhø.
basement of the Guayana Shield (e.g.,
S. cacumina Wurdack, S. marginata Palabras clave: Araracuara, bosques Benth.), and drained by black waters húmedos secundarios, cuenca del río rivers (e.g., S. aquae-nigrae Aymard, S.
Caquetá, Polygalaceae, Pøøsiøhø, savannarum Wurdack) on white sandy Securidaca, Colombia
shrubby savannas and Caatinga forests
PØØSIØHØ Ka ñọifosí įsiakataį
(Aymard et al. 2004, Aymard 2022).
Some species have wide geographic
Ka´se
øo´defø,
өọpөa
føsí, distributions (e.g., S. divaricata Nees & meøtøseisiøhø fisi siøhø, ñ ọifesí ø Mart., S. diversifolia (L). S. F. Blake) in įpeikø økasiøhø bø (Cuenca del Río lowland “terra firme” (Anjos et al. 2024) Caquetá en Araracuara). nọí føsí øatíį and montane forests. The remaining siokapįį siøhø KAÁdt cono siøhø įsu species require more specific habitats, siøhø pañema į øke bø (Biodiversidad) like foothills in the low-medium to high aviøhø ye eį´ okasiø bø ke´Á pøsiøhø altitude (500–2500 m) mountains of the Andes (e.g., S. leiocarpa S. F. Blake,
81
S. planchoniana Killip & Dugand), infrageneric division has been along the Pacific Coast of Colombia and proposed. Marques (1996) identified Ecuador (e.g., S. calophylla (Poepp.) two taxonomically useful inflorescence S. F. Blake) and the dry forests (e.g., characters that separate species: 1) S. scandens Jacq.) as well. Others are flowers solitary or in short racemes endemic to particular geographical 0.5–2.5 cm long, with 5–15(-20) areas, such as some species found only flowers; and 2) inflorescences racemes in Caribbean islands (e.g., S. lophosoma or panicles, 2.5–45 cm long, with more Cheesman, sensu Ulloa Ulloa et al., than 15 flowers. After Wurdack (1972) 2018 Onwards), Mesoamerica (e.g., and Marques (1996) contributions, S. micheliana Chodat, sensu Morales only seven species had been described Quiros, 2014) or in Central to Southeast (Wallnöfer 1998, Eriksen et al. 2000, Brazil (e.g., S. ovalifolia A. St. Hil. & Aymard and Campbell 2007, Da Costa Moq., S. tomentosa A. St. Hil., sensu et al. 2013, Aymard 2022, Anjos et al.
Marques, 1996).
2024). In addition, the genus has been
treated for Flora of Peru (Macbride
Polygalaceae are monophyletic, divided 1950), Flora of Panama (Woodson et at present into four tribes: Carpolobieae al. 1969), Flora of Brazil (Marques B. Eriksen, Diclidanthereae Reveal, 1996), Flora of Ecuador (Eriksen et al.
Polygaleae Chodat and Xanthophylleae 2000), Flora of the Venezuelan Guayana Chodat (Pastore et al. 2017, Mota et al. (Aymard et al. 2004) and Manual de 2019, 2023). Securidaca is included Plantas de Costa Rica (Morales Quiros in the tribe Polygaleae characterized 2014).
by its papilionaceous flowers, with
the 5-merous corolla usually reduced While conducting
herbarium
to 3-merous (the keel formed by one research for updating the checklists of petal and the wings are inner sepals), Polygalaceae for the “Catálogo de plantas sometimes with a pair of rudimentary y Líquenes de Colombia” (Aymard and petals, and bicarpellate gynoceium with Freire-Fierro 2016) and “Catálogo de one ovule per fertile locule (Eriksen, la flora del medio Río Caquetá y áreas 1993a,b; Eriksen & Persson, 2007, adjacentes” (Sanchez 1997), a specimen Mota et al. 2023). Securidaca is further of a morphologically distinct species of distinguished by its habit-wise of woody Securidaca were detected, collected in lianas, herbaceous vines, rarely shrubs the Araracuara region, on the Middle or small trees (Eriksen & Persson 2007), Caquetá River basin. This biodiverse the keel with a crest, well developed area in the Colombian Amazon basin or obsolete, a one-seeded samara, and has had a considerable number of an appendaged seed (Aymard and studies on floristic, ecology, vegetation Campbell, 2007).
classification, soils, geomorphology,
and phytogeographic for more than
Securidaca
has
never three decades (e.g., Duivenvoorden et been monographed, and no formal al. 1988, Urrego 1991, Duivenvoorden
Aymard C. y Andoque A.: Securidaca (Polygalaceae), especie nueva, Río Caquetá
& Cleef 1994, Duivenvoorden & Lips the checklist: Catálogo de plantas y 1995, Duivenvoorden 1996, Sánchez Líquenes de Colombia (Aymard and 1997, Londoño-V. y Álvarez 1997, Freire-Fierro, 2016) was reviewed.
Arbeláez y Callejas 1999, Duque et al.
2001, Hoyos-Gómez et al. 2023, Urrego Type specimens of Securidaca species et al. 2024). Besides this amount of field involved in this study were examined work and interesting results, new taxa using on-line images from JSTOR
still appear in this region and adjacent Global Plants (https://plants.jstor.
areas (e.g., Córtes-Ballén 2014, Daly org/). In addition, International Plant
& Melo 2017, Oliveira et al. 2020, Names Index
(https://www.ipni.
Marcano-Berti et al. 2023, Da Silva et org /), the online botany collections al. 2024), including the new species of Smithsonian Natural Museum of described here.
Natural History (https://naturalhistory.
si.edu/research/botany) and Tropicos
This contribution increases to 27 the (http://legacy.tropicos.org/Home.aspx) number of Securidaca species known were also consulted to update the from Colombian flora, also four species current nomenclature and geographical of this genus are very probably to information.
be found in Colombia ( S. maguirei
Wurdack, S. prancei Wurdack, S. The specific terminology for vegetative pyramidalis Sprague & Sandwith and characters, vestiture
description,
S. savannarum; see Aymard (2022) inflorescences, flowers, and fruit and the species key below). The name morphology follow Font-Quer (2001), poeoesioehoeorum represents the first Harris and Harris (2006) and Eriksen & time that a plant, species honored the Persson (2007).
Pøøsiøhø (Andoke) nation.
RESULTS
MATERIALS AND METHODS
Securidaca poeoesioehoeorum Aymard,
This work is based on morphological sp. nov. (Figure 1) (using a dissecting stereomicroscope) TYPE: COLOMBIA. Caquetá: and herbarium studies in COAH, COL, Araracuara, Río Caquetá, bosques GH and HUA (herbarium codes after secundarios, aprox. 00º 36’ 50´Ś; 72º
Thiers 2024). Current bibliography was 23’ 03´´W, 120 m, 05 Septiembre 1991, scrutinized, essentially the Wurdackś Hans Vester & Bartolomé Castro 498
contributions (1957, 1972), Flora of (holotype: COAH-04300!).
Brazil (Marques 1996), the Flora of
Ecuador (Eriksen et al. 2000), the Flora Diagnosis of the Venezuelan Guayana (Aymard
et al. 2004) and the Colombian and Securidaca poeoesioehoeorum is similar Venezuelan Securidaca species key to S. leiocarpa, but morphologically included in Aymard (2022). Also, it differs from the latter in having oblong or obovate-oblong leaves (vs.
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ovate), chartaceous (vs. coriaceous or at the base of the inflorescences. Leaves subcoriaceous), black punctate, light- gradually diminishing in size from the brown on the lower surface (vs. not tip to below part of branch, 1.5–3.5 ×
punctate, golden-green), the base and 1.5–3.5 cm, oblong or obovate-oblong, apex rounded (vs. base truncate or chartaceous, shiny,
glabrescent-
sometimes short-attenuate, apex retuse punctate, black-brown on the upper or obtuse-retuse); the secondary veins surface, puberulent to glabrescent along 5–12, veinlets strongly reticulate (vs. midvein, black punctate, light-brown 4–6, veinlets inconspicuous) and petioles on the lower surface, erect pilose along 0.5–1 mm (vs. 1–2). Inflorescences the midvein and secondary veins; the axillary and along the branchlets, base and apex rounded; the margin 2–5 mm long, with 1–3 flowers (vs. entire, revolute, ciliate; the midvein terminal, l0–30 mm long with 8--20 flat above, prominent beneath, venation flowers), pedicels 0.5–1 mm long (vs. brochidodromous, the secondary veins 8–11); sepals densely pilose pubescent 5–12, very prominent on both surfaces, outside (vs. sparsely hirsute), lateral veinlets strongly reticulate forming a net enlarged sepal 5.5–6 mm long, claw ca. nervation. Petioles 0.5-1 × ca. 0.5 mm, 0.2 mm long, broadly ovate (vs. 9–10 not canaliculate, densely yellow pilose.
mm long, claw 0.5–1 mm long, almost Inflorescences axillary and along the circular), keel petal 4–5 mm long (vs. branchlets, unbranched, 2–5 mm long, 9–10 mm long), adaxial (lateral) petals with 1–3 flowers, rachis densely yellow ca. 1.8 × 1 mm, oblong-lanceolate (vs. pubescent, bracteoles at the base of ca. 5 × ca. 3, oblong), stamens 4 (vs. pedicels, ovate-lanceolate, ca. 1 × ca. 1
8) and samara with central spheroidal mm, pilose outside, glabrescent inside, body ca. 8 × ca. 5 mm, dorsal secondary pedicels 0.5–1 mm long, densely yellow wind 2–3 mm long, primary wing 3–3.5 pubescent. Sepals of the outer whorl
× 1–1.5 cm, membranaceous, brown, subequal, ca. 2 × ca. 2 mm, broadly adpressed pilose (vs. spheroidal body ovate or ovate, densely pilose pubescent ca. 5 × ca. 5 mm, dorsal secondary wind outside, glabrescent inside, margins ca. 1 mm, primary wing 2.5–3 × 1.2–1.5 ciliate, the outer sepals ca. 3.5 × ca. 2
cm, coriaceous, yellowish, glabrescent). mm, the inner sepal ca. 3.15 × ca. 1.7
mm, lateral enlarged sepal (petal-like)
Liana with flattened stems to sub- 5.5–6 × 3–3.5 mm, claw ca. 0.2 mm long herbaceous vine, lateral branches not white-purple, broadly ovate, glabrous forming sensitive hooks or looped on both sides, strongly reticulate-veined tendrils. Branches and branchlets terete, at the base, with 4–6 major veins, acute yellow patent-pubescent, sparsely at the apex. Corolla white-purple; keel pilose when mature, longitudinally petal 4–5 × 1.5–2 mm, claw ca. 2 mm striate, bark not flaking off when long, ovate, glabrous on both surfaces, mature; with short stalked glands, ca. strongly reticulate-veined, with 10–
0.2–0.5 mm long, glabrous, capitate in 12 major veins, without apical crest the top, located along the branchlets and (ecristate); adaxial (lateral) petals ca.
Aymard C. y Andoque A.: Securidaca (Polygalaceae), especie nueva, Río Caquetá 1.8 × 1 mm, oblong-lanceolate, glabrous department of Caquetá. The vegetation, on both sides; filament sheath ca. 2 soils, geology, and geomorphology of mm long, glabrous on both surfaces, this area have also been well studied filaments 4, ca. 2.2 mm long, free; (especially the sector from Araracuara anthers oblong, ca. 0.5 mm long. Ovary to the mouth of the Cahuinari river; ca. 2.5 mm long, glabrous. Samara 3.5– see the introduction). This research 4 × 1–1.5 cm, body spheroidal ca. 8 × was done through the establishment of ca. 5 mm, white adpressed pubescent, the Amazon herbarium in Araracuara strongly reticulate, dorsal secondary in 1982 and various research projects wind, obsolete, oblong, 2–3 mm long, conducted by the Tropenbos-Colombia sparsely pilose, primary wing 3–3.5 Foundation, the Colombian Corporation
× 1–1.5 cm, membranaceous, black- for the Amazon “Araracuara” and brown, entire, strongly veined (35–40 “Universidad Nacional de Colombia”
veins), sparsely adpressed pilose.
(Infante-Betancour and Rangel-Ch.
2018).
Phenology: This new species has been
collected with flowers and fruits in State of conservation: Currently, this September.
new species is only known from the type
collection, and it is reported here as rare
Taxonomic relationship: This new species. However, under IUCN (2022) species is remarkable by its four guidelines a single locality constitutes stamens, in an otherwise predominantly data deficient (DD) to determine eight stamens genus, and it has some its conservation status, the area of similarity with Securidaca leiocarpa, Occupancy (AOO) and the extent of S. F. Blake. Both taxa below to short Occurrence (EOO) as well (IUCN
racemes (0.5–2.5 cm long) informal 2022). Nevertheless, the conservation group and have bicolored leaves: drying of these wet forest of the middle black-brown on the upper surface, Caquetá basin in Colombia is not at risk light-brown ( S. poeoesioehoeorum) of deforestation and degradation.
and golden-green ( S. leiocarpa)
on the lower surface. Nonetheless, The area where S. poeoesioehoeorum S. poeoesioehoeorum differs from occurs is currently well protected by S. leiocarpa in the vegetative and regional initiatives such as the three reproductive characters discussed in the REDD+ projects named: “Aire de vida”
diagnosis and in the following key of (“Fivo Jaagava Komuya Jag+y+”) Securidaca species of Colombia.
in “Monochoa”; “Nuestro Aire de
Vida” (“Kai Komuya Jag+y+”) in
Distribution: The new species is known “Puerto Zábalo y Los Monos” and to occur in the Araracuara region, “CRIMA-Predio Putumayo y Andoke in secondary wet forest over clays de Aduche”. These three projects are soils, between 100-200 m elevation. efficiently managed by the Pøøsiøhø, The former is a region located in the Uitoto and Muinane nations, and middle Caquetá River basin, in the
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presently protect ca. 1,965.538 hectares by its complex cultural heritage, of non-fragmented forests. The latter multifaceted regional societies, locally activities are remarkable in this region dense populations, ancestral cultivation because Amazonian basin holds the techniques and the extensive use forest world’s largest structurally and diverse resources (Espinosa-Arango 1995, complex tropical rainforest, those are Estrada-Añokazi 2017). Also, Pøøsiøhø vital for earth’s ecosystem dynamic people were known by their neighbors processes (Laurance et al. 2018). as suppliers of quality stone axes, since Tropical forests are estimated to storage they knew and managed the site where ca. 1/4 of total carbon pools within the hardest stones (suitable for their land ecosystems. The latter condition manufacture), were found (Whiffen plays a fundamental role in combating 1915, Pineda-Camacho 1975). This climate change (Longo et al. 2016). In nation and other people located in the addition, Amazon rainforest has been middle Caquetá River are known as subject to climate-related disturbances “People of the Centre”, as the Pøøsiøhø such as forest fires, droughts, and think of themselves. They are composed danger anthropogenic activities like by seven ethnolinguistic groups of the deforestation, mineral extraction and Pøøsiøhø, Boran and Uitotoan linguistic logging in the past several decades with stocks, which share territory, a number the second highest gross forest loss of cultural traits and have a common globally (Csillik et al. 2024).
social and ceremonial organization in
spite of their spatial dispersion and
Etymology:
Securidaca linguistic heterogeneity (Echeverry & poeoesioehoeorum is named to Román-Jitdutjaaño 2013, Heckenberger honor the Pøøsiøhø (Andoke) nation. 2013). These people are experts in According to Art. 60.7 of the current ecological knowledge, including International Code of Nomenclature for advanced systems of river and forest algae, fungi, and plants (Turland et al. resource use including agroforestry 2018), a botanical name must not have (Denevan & Padoch 1988). However, diacritical sign (i.e., “ø”), therefore, “People of the Centre” were subjected here, the letter “ø” was changed to “oe”. to a brutal slavery system (mainly debt The Pøøsiøhø people are also known as peonage and forced labor) that forced
“Paasi-ahá” or “Andoke” in their own most native workers of middle Caquetá language, meaning “Gente de hacha” and Putumayo Rivers to gather the latex in Spanish, the “axe-people” in English of Hevea spp. (Euphorbiaceae) during (Landaburu 1976). The connotation the Amazon Rubber Boom perpetrated or word “Andoke” comes from the by Peruvian Amazon Company or Casa Uitoto language, which means people Arana, from 1885 to 1930 (Hardenburg who live in the hills (Nelson “Requi” 1921). Few if any native households in Andoque, pers. comm, Jan. 2025). This Northwestern Amazon basin escaped Amazonian nation was a pre-European from this “commerce” (Wasserstrom Amazonian
people
characterized 2017). Once, the Rubber Boom receded,
Aymard C. y Andoque A.: Securidaca (Polygalaceae), especie nueva, Río Caquetá the “People of the Centre” were again pubescent, sparsely pilose, puberulent exposed to colonization and invasion of or glabrescent … 6
their territories though the Araracuara
Agricultural Penal Colony (which 3. Leaves narrowly ovate to operated from 1938 to 1971), the coca ovate, or oblong, inflorescences plantations, fur trappers, armed conflict, 5–15(-20)-flowered … 4
and gold mining (Useche-Losada 1994, 3. Leaves ovate to broadly-ovate or Estrada-Añokazi 2017). Despite all orbicular-ovate; flowers solitary or these difficulties, the survivors grouped inflorescences with 2–6-flowered … 5
and reconstructed their ethnic identities
as we see them today (Estrada-Añokazi 4. Branchlets glabrous; leaves 15–22
2017). Currently, their ancestral cm long, coriaceous or subcoriaceous, territory spans the area between the veinlets inconspicuous on both sides; Caquetá and Putumayo Rivers in south- flowers yellowish to green; enlarged eastern Colombia, along the border lateral sepals subrotund, claw 2–3
with Peru ( sensu Cayón and Chacón mm long; pedicels 2–5 mm long; fruit 2022). The specific epithet name with a broadly wing … S. calophylla
( poeoesioehoeorum) represents the (Colombia: Antioquia,
Boyacá,
first time that a plant species honors the Caquetá, Nariño; Brazil, Ecuador, Perú) Amazonian Pøøsiøhø nation.
4 Branchlets minutely puberulous;
Key to the species of Securidaca of leaves 3–11 cm long; herbaceous to Colombia
chartaceous, veinlets forming a strongly
reticulate net nervation; flowers pink o
(Amended and modified from Aymard purple; enlarged lateral sepals broadly 2022)
oblong, claw ca. 1 mm long; pedicels
Species endemic to Colombia#
6–8 mm long; fruit with a narrowly,
lateral wing ... S. fragilis (Colombia: 1. Flowers solitary or in short racemes Antioquia, Caquetá; Ecuador) 0.5–2.5 cm long, with 1–15(-20) flowers
… 2
5. Leaves ovate, 5–8 × 2–4.5 cm,
puberulent on the abaxial surface,
1.Inflorescences racemes or panicles, margins revolute, petiole 2–2.5 mm 2.5–45 cm long, with more than 15 long; inflorescence with 1–3 flowers; flowers … 10
keel petal without a crest … S. uniflora
2. Branchlets glabrous or minutely (Colombia: Caquetá; Venezuela, Brazil, puberulous; samara wing always Guianas)
glabrous … 3
5. Leaves ovate to broadly-ovate or
2. Branchlets densely hirsute, pilose orbicular-ovate, 5–15 × 4–11 cm, pubescent or sparsely strigulose to glabrous on abaxial surface margins glabrescent when mature; samara wing not revolute, petiole 4–7 mm long; inflorescence with 3–6 flowers; keel
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petal crested … S. goudotiana# 8. Branchlets, petioles, and pedicels (Colombia: Antioquia, Cundinamarca, with golden pubescence; leaves elliptic Tolima, Valle)
to narrowly ovate, ovate to broadly
ovate or oblong-ovate; margins not
6. Leaves ovate, 2–4 × 1–2 cm, drying revolute; ovary glabrous … 9
black-brown on the upper surface,
golden-green on the lower surface; 9. Leaves elliptic to narrowly ovate, major wing 2–3 cm long … 7
chartaceous,
white-puberulent
on
the lower surface, base subcordate
6. Leaves ovate-elliptic, elliptic, ovate- or truncate; inflorescences 0.5–1 cm oblong or oblong-ovate, 5–11 × 3–6 long; outer sepals puberulent at the cm, drying brown-green on the upper middle externally; flowers white-surface, light green on the lower surface; yellow or green, keel crest 8–14 mm major wing longer than 5 cm … 8
long … S. prancei (Colombia: very 7. Leaves ovate, golden-green on probably in Amazonas, Guainía and the lower surface, base truncate or Vaupés departments; Venezuela, Brazil, sometimes short-attenuate, apex retuse Bolivia) or obtuse-retuse; inflorescence terminal, 9. Leaves ovate to broadly ovate l0–30 mm long (with 8--20 flowers), or oblong, subcoriaceous, yellow-pedicels 8–11 mm long; stamens 8; pubescent on the lower surface, base primary wing of fruit 2.5–3 × 1.2–1.5 rounded to slight obtuse; inflorescences cm, coriaceous, yellowish, glabrous … 2–2.5 cm long, outer sepals glabrous
S. leiocarpa (Colombia: Magdalena; or sparsely strigose externally; flowers Ecuador)
pink-purple, keel crest 4–6 mm long
7. Leaves oblong or obovate-oblong, … S. aff. S. purpurea (Colombia: light-brown on the lower surface, the Antioquia, Norte de Santander, Valle; base and apex rounded; inflorescence Venezuela) 2–5 mm long, axillary and along the 10. Leaves completely glabrous on both branchlets, with 1–3 flowers, pedicels surfaces … 11
0.5–1 mm long, stamens 4; primary
wing of fruit 3–3.5 × 1–1.5 cm, 10. Leaves sparsely hirtellous, strigulose, membranaceous, brown, adpressed pilose, pilosulous, puberulent, or pilose … S. poeoesioehoeorum# pubescent-hirsute to densely appressed-
(Colombia: Amazonas, Caquetá)
pubescent at least on one surface (more
evident along the midrib and secondary
8. Branchlets, petioles and pedicels with veins) … 13
white pubescence; leaves ovate-elliptic
or elliptic; margins revolute; ovary 11. Inflorescences compound by more sparsely strigulose to glabrescent … S. than 3 lateral racemes (paniculate);
savannarum (Colombia: very probably major fruit wing 1–1.5 cm long … S.
in Guainía and Vaupés departments; paniculata s. l. (Colombia: Amazonas, Venezuela)
Aymard C. y Andoque A.: Securidaca (Polygalaceae), especie nueva, Río Caquetá Antioquia, Chocó, Putumayo, Vaupés; ovary glabrous … S. paniculata s.
Venezuela, Brazil, Guianas, Perú)
l. (Colombia: Amazonas, Antioquia, Chocó, Putumayo, Vaupés; Venezuela;
11. Inflorescences compound by 1 or 2 Brazil, Guianas, Perú) lateral racemes; major fruit wing longer
than 2 cm long … 12
15. Leaves ovate, lanceolate, or
lanceolate-ovate;
inflorescences
12. Leaves ovate to broadly ovate, shiny unbranched racemes; flowers 8–16 cm on the upper surface; racemes 3–5 cm long; ovary pubescent …… 16
long; pedicels 8–10 mm long; samara
wings glabrous, secondary wing nearly 16. Leaves coriaceous; outer sepals fully confluent with the primary wing; symmetric, abaxially glabrescent; keel deciduous dry forests … S. scandens petal crest 1.2–2 mm long; samara (Colombia:
Antioquia,
Bolívar, 2–2.5 cm long, wings glabrescent … S.
Cesar, Magdalena, Santander, Valle; bialata (Colombia: Amazonas, Guianía, Venezuela)
Vichada; Venezuela, Brazil)
12. Leaves elliptic or ovate; racemes 16. Leaves chartaceous to subcoriaceous; 4–12 cm long, dull on the upper surface; outer sepals asymmetric, abaxially pedicels 5–6 mm long; samara wings pilose; keel petal crest 0.5–0.7 mm long; basally puberulent, sometimes sparsely samara 1.5–1.8 cm long, wings densely so; secondary wing noticeably cleft strigulose … S. longifolia (Colombia: from primary wing, acute; wet forest Amazonas; Venezuela, Brazil, Guyana,
… S. retusa (Colombia: Amazonas, Perú, Bolivia) Vaupés; Venezuela, Brazil, Guyana,
French Guiana)
17. Abaxial surface of the leaves
puberulent,
sparsely
hirtellous,
13. Secondary wing of the samara well strigulose, or pilosulous, more evident developed (samaras 2-winged) … 14
along the midrib and secondary veins
… 18
13. Secondary wing of the samara
weakly developed or obsolete (samaras 17. Abaxial surface of leaves hirsute, 1-winged) … 17
shortly-pilose, densely appressed-
pubescent or velutine, not pilosulous to
14. Racemes 10–30 cm long; samara glabrescent … 23
wings ca. 1 cm long … S. pendula
(Colombia: Meta, Vichada; Venezuela, 18. Keel petal without a crest or crest Brazil, Guyana)
reduced, 0.5–4 mm long, not reflexed
…. 19
14. Racemes 4–10 cm long; samara
wings 1.5–3.5 cm long … 15
18. Keel petal crested well developed;
crest 6–15 mm long, reflexed …… 21
15. Leaves oblong, sometimes elliptic
or obovate; inflorescences compound 19. Leaves coriaceous, margins revolute, (panicle); flowers 5–7 cm long;
89
glabrous on the adaxial surface; keel cm long …. 22
petal ciliate on the margins … S.
warmingiana (Colombia: Guianía, 22. Leaves elliptic-ovate, obtuse Vaupés; Venezuela, Brazil, Guyana, or subcordate at the base, sparsely Perú, Bolivia)
hirtellous to glabrescent on the adaxial
surface; inflorescences 8–45 cm long;
19. Leaves chartaceous or subcoriaceous, keel petal 10–11 mm long … S. speciosa
margins not revolute, pilosulous on the (Colombia: Guianía; Venezuela) adaxial surface: Keel petal not ciliate on
the margins … 20
22. Leaves ovate-oblong, sometimes
ovate, acute or short-attenuate at the
20. Branches and branchlets densely base, strigulose on the abaxial surface; hirsute; leaf blades oblong to narrowly inflorescences 4–12 cm long; keel elliptic or broadly oblong, 1–2(- petal ca. 8 mm long … S. diversifolia
3) cm wide, pilosule on both sides; (widespread in Colombia, SE-USA to inflorescences unbranched, keel petal Argentina) glabrous on both sides; lateral wings
glabrous … S. divaricata (Colombia: 23. Young branchlets, petioles, Amazonas, Caquetá, Cundinamarca, inflorescences, and pedicels densely Meta, Vaupés; Venezuela, Brazil, white-pubescent; petioles 10–15 mm Guianas, Ecuador, Perú, Bolivia)
long; ovary puberulent …. S. maguirei
(Colombia: very probably in Guainía
20. Branches and branchlets sparsely and Vaupés departments; Venezuela, strigulose; leaf blades ovate or oblong- Brazil, Perú) ovate, sparsely strigulose on both sides;
inflorescences with 2 lateral racemes; 23. Young branchlets, petioles, keel petal sparsely fine- setulose inflorescences, and pedicels not white-outside; lateral wings covered by golden pubescent; petioles 1–7 mm long; ovary trichomes outside … S. schlimii# glabrous or densely hirsute … 24
(Colombia: Bolívar, Norte de Santander; 24. Leaves two different sizes, the ones most likely found in Venezuela)
on the inflorescence rachis smaller
21. Leaves all one size; outer sepals than ones on main branches, with long glabrous abaxially; samara wing trichomes (0.7–1.5 mm long) on the 1–1.5 cm long … S. paniculata s. l. abaxial surface … 25
(Colombia: Amazonas, Antioquia, 24. Leaves all one size, trichomes ca.
Chocó, Putumayo, Vaupés; Venezuela, 0.5 mm long on the abaxial surface …
Brazil, Guianas, Perú)
26
21. Leaves two different sizes, the ones 25. Leaf blades on main branches ovate on inflorescence rachis smaller than to broadly ovate, coriaceous, sparsely ones on main branches; outer sepals pilulose to glabrous on the upper strigulose abaxially; samara wing 2–4 surface, lateral veins 4–6; enlarged
Aymard C. y Andoque A.: Securidaca (Polygalaceae), especie nueva, Río Caquetá lateral sepals ca. 8 × 6–8 mm, with 6–8 Antioquia, Boyacá, Cundinamarca, major veins; keel ca. 8 mm long, densely Guaviare, Magdalena, Norte de tomentose inside, with a well-developed Santander, Putumayo, Valle; Venezuela)
apical crest, 8–14 mm long; samara
primary wing 3.5–4 cm long, hirsute … 28. Leaves herbaceous to chartaceous;
S. coriacea (widespread in Colombia; flowers 9–12 mm long; outer sepals Panamá, Venezuela, Brazil, Guyana, glabrous or puberulent … 30
Surinam, Ecuador, Perú, Bolivia)
29. Leaves ovate to broadly ovate or
25. Leaf blades on main branches oblong, shiny on the upper surface, oblong to narrowly oblong or broadly yellow-pubescent on the lower surface; ovate, subcoriaceous, puberulent to outer sepals glabrous or sparsely strigose glabrous on the upper surface, lateral externally … S. purpurea (Colombia: veins 5–12; enlarged lateral sepals 5–6 Antioquia, Cauca; Venezuela)
× ca. 6 mm, with 14–16 major veins; 29. Leaves elliptic or ovate-elliptic, dull keel 4–5 mm long, glabrous outside, on the upper surface, velutinous on the without well-developed apical crest lower surface; outer sepals puberulent (ecristate), ca. 0.25 mm long; samara externally … S. amazonica# (Colombia: primary wing 2.5–3 cm long, sparsely Putumayo; most likely found in Brazil, pilose … S. aquae-nigrae (Colombia: Ecuador and Perú) Guianía, Venezuela)
30. Leaves densely puberulent on the
26. Leaves coriaceous; branches and abaxial surface; petioles 5–9 mm long; branchlets furfuraceous-pilose … S. ovary glabrous; samara wing ca. 4 cm
pyramidalis (Colombia: very probably long … S. pubescens (widespread in in Amazonas, Caquetá, Guainía and Colombia; Venezuela; Brazil, Guianas, Vaupés
departments,
Venezuela, Perú)
Guyana, Bolivia?)
30. Leaves densely pubescent or velvety-
26. Leaves chartaceous or subcoriaceous; pilose on the abaxial surface, especially branches and branchlets tomentose, along the vein; petioles 2–4 mm long; densely pilose or densely pubescent … ovary pubescent; samara wing 3–7 cm 27
long … 31
27. Leaves glabrescent, puberulent, 31. Leaves ovate to broadly ovate or shortly pilose or sparsely pilose on the oblong-ovate, margins not revolute, adaxial surface … 28
sparsely pilose to glabrous on the
27. Leaves dense pubescent or densely adaxial surface … 32
puberulent on the adaxial surface … 29 31. Leaves elliptic to ovate, margins 28. Leaves subcoriaceous; flowers 6–8 revolute, densely pubescent or strigose mm long; outer sepals densely pubescent on the adaxial surface … 33
… S. planchoniana (Colombia: 32. Scandent shrubs or herbaceous
91
vines; branchlets softly pilose, leaves of the communities of “CRIMA-oval, oblong-ovate or narrowly ovate, Predio Putumayo y Andoke de Aduche densely velvety-pilose on the abaxial REDD+ project, Amazonas and surface; outer sepals densely pilose Caquetá departments and the staff of outside … S. tenuifolia (Colombia: “Herbario Amazónico Colombiano”
Antioquia, Bolívar, Cundinamarca, – COAH (Dairon Cárdenas Lopez+, Magdalena;
Venezuela,
Panamá; Nicolas Castaño, Wilson Rodríguez
Trinidad and Tobago)
and Jorge M. Vélez). We also thank
the “Herbario Nacional de Colombia”
32. Woody lianas, branchlets densely (COL) and Harvard herbarium (GH) hirsute, leaves ovate to broadly ovate, for making their facilities available for pubescent on the abaxial surface; outer our research, to Alicia Micolta Cabrera sepals sparsely puberulent to glabrous and Pedro Posada for logistic support, except the margin outside … S. trianae to Walter Guerrero Palacios, Orlando (Colombia: Boyacá, Cundinamarca; Andoque, Nelson “Requi” Andoque Venezuela, Ecuador)
and Nelly Andoque for the first-hand
33. Leaves densely pubescent on the information about Pøøsiøhø nation, and adaxial surface; pedicel 4–6 mm long; the Pøøsiøhø forestry team for their outer sepals puberulous; keel petal field assistance. The first author is also spatulate-ovate, obtuse … S. volubilis grateful to Manuela A. Sánchez Quiroga (Colombia: Putumayo; Costa Rica, (JBB) for preparing the illustration, Panamá, Puerto Rico, Venezuela, Brazil, to Gustavo A. Romero-G. (AMES) Guianas, Ecuador, Perú, Bolivia)
for his help finding literature and his
invaluable comments on early versions
33. Leaves strigose on the adaxial of this text and Kanchi N. Gandhi (GH) surface; pedicel 6–8 mm long; outer by his advises relating to the species sepals densely pubescent; keel petal name. This work would not be possible obovate, acute … S. planchoniana without the International Plant Names (Colombia:
Antioquia,
Boyacá, Index (https://www. ipni.org/), JSTOR
Cundinamarca, Guaviare, Magdalena, Global Plants (https://plants.jstor.org/), Norte de Santander, Putumayo, Valle; Biodiversity Heritage Library website Venezuela)
(http://www.biodiversitylibrary.org/),
ACKNOWLEDGMENTS. The first and Tropicos (http://legacy.tropicos.org/
author is gratefully acknowledging the Home.aspx) databases and facilities.
support provided by Pøøsiøhø people
92
Aymard C. y Andoque A.: Securidaca (Polygalaceae), especie nueva, Río Caquetá Figure 1. Securidaca poeoesioehoeorum. A. Habit showing the adaxial leaf surface with the appressed pubescence, inflorescence and fruits. B. Fruit. C. Filament-sheath, lateral petals and the four stamens in lateral view. D. Keel crest in lateral view. E. Enlarged sepal (petal-like) in lateral view. F. Sepals. G. Branchlet and inflorescence showing the clavate glands, the bracteoles and the flower in lateral view. Illustration based on the type collection ( Vester & Castro 498).
93
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