Studies in Swartzia (Leguminosae−Papilionoideae) of
Colombia with emphasis in sect. Terminales
LIZ KAREN RUIZ BOHÓRQUEZ
Director: Ph. D. Santiago Madriñan
Maestria en Ciencias Biológicas con énfasis en Botánica y Sistemática
Escuela de Post−Grados−Departamento de Biología
Universidad de los Andes
TABLE OF CONTENTS
Abstract Introduction
Materials and methods
The genus Swartzia sect. Terminales
Species of sect.
T
erminalesfound in Colombia
Key to species of Swartzia sect. Terminales in Colombia Taxonomic Treatment
1. Swartzia amplifolia Harms
2. Swartzia argentea Spruce ex Bentham 3. Swartzia cabrerae R.S. Cowan
4. Swartzia cardiosperma Spruce ex Bentham 5. Swartzia flavescens (Cowan) Suessenguth 6. Swartzia leptopetala Bentham
7. Swartzia macrophylla Willd. ex Vogel 8. Swartzia magdalenae Britton & Killip 9. Swartzia santanderensis R.S. Cowan 10.Swartzia schultesii R.S. Cowan
11.Swartzia sp nov. 1 L.K. Ruiz, Torke & Mansano 12.Swartzia sp nov. 2 L.K. Ruiz, Torke & Mansano 13.Swartzia sp nov. 3 L.K. Ruiz, Torke & Mansano 14.Swartzia sp nov. 4 L.K. Ruiz, Torke & Mansano
Acknowledgements Literature cited List of exsiccatae
Studies in Swartzia (Leguminosae−Papilionoideae) of Colombia with emphasis
in sect. Terminales
ABSTRACT
The genus Swartzia (Leguminosae−Papilionoideae−Swartzieae) includes nearly 200
species and is particularly diverse and abundant in the tropical forests of the Amazon basin, the Guianas and northern South America (Torke & Zamora, 2010); in Colombia it isquite diverse on both sides of the Andes, in the inter−andean valleys and in the low and humid lands. It grows from sea level to 2200 m altitude. Swartzia has been divided into 15 sections. Among them,
Terminalesis the largest and most diverse in Colombia; it can be recognized by the combination of pedicels without bracts, flowers with a yellow petal, the stipe and style reduced in relation to the ovary, with numerous seeds and moniliform ovary (Torke & Schaal 2008; Torke & Mansano, 2009). The research has centered on taxonomic studies and field observations of the Colombian species of Swartzia sect. Terminales. In the course of the investigation, material from 13 herbaria both in Colombia and the United States totaling ca. 2000 specimens, was studied. Variability of morphological characters was determined for each species by measuring both vegetative and reproductive organs. Field collections were carried out in Antioquia, the middle Magdalena valley and Vaupes. It is concluded that ten species–out of 23 species so far described for South America−grow in Colombia; in addition, four species are described as new to science and one species is reinstated.
Key words: Antioquia, Colombia, Magdalena River valley, Neotropics, Swartzia, Taxonomy, Vaupes.
INTRODUCTION
The plant family Leguminosae is one of the largest groups of Angiosperms (flowering plants) in the world, with about 730 genera and 19.300 species (Cardoso et al., 2012), it is cosmopolitan in distribution and is present in almost all of the earth´s biomes, from deserts to wet tropical forests. The Leguminosae can be herbs, shrubs or very large trees and are well represented in the Colombian flora (Forero, 2005).
A good example of this diversity is the genus Swartzia Schreber of the tribe Swartzieae , that includes nearly 200 species and is particularly abundant and diverse in the tropical forests of the Amazon basin, the Guianas and northern South America (Torke & Zamora, 2010); species of
Swartzia grow from sea level to 2200 m altitude, and several species can grow and coexist in the same forest zone, showing interesting and complex relations with other plants, pollinators seed dispersal agents, herbivores and even human beings. Given its species richness, its ecological complexities and ubiquity in the neotropical forests, the genus constitutes a model system to understand the origin and sustainability of plant diversity in the neotropics. Species of Swartzia
are also important ecological and biogeographic entities due to their high diversity and notable endemism in neotropical forests.
The genus Swartzia (Leguminosae) is highly diversified in the wet lowlands of Colombia on both sides of the Andes Mountains, as well as in the large valleys that separate the Andean Cordilleras. Documentation of this diversity, much of it threatened, has proceeded at a slow pace. The existing herbarium collections of Colombian Swartzia number fewer than 2000 and are too few to give more than a cursory view of nation−wide taxonomic diversity, morphological
variation and species distributions. To complicate matters, many collections are not deposited outside of Colombia, or even beyond the herbarium of origin. Nevertheless, data are substantially more numerous than they were when Richard Cowan undertook the last comprehensive taxonomic study of Swartzia. Collections made subsequent to the publication of his monograph (Cowan, 1968) account for approximately 80 percent of the existing material.
In addition to the revision by Cowan (l.c.) for Flora Neotropica, the genus has been studied in recent years and from different perspectives by Torke & Schaal (2008), Torke & Mansano (2009), and Torke & Zamora (2010). Britton & Killip (1936) mentioned 7 species for Colombia. The lack of information on Colombian taxa is evident in works like Cowan’s. As part of a long−term systematic study of the genus throughout its range, Torke and collaborators have examined much new material, along with older collections, in Colombian herbaria and abroad. They have concluded that approximately 65 species of Swartzia occur in Colombia, doubling the number recorded from the country by Cowan (1968). While some of the new records involve nomenclatural changes (e.g., Torke & Mansano, 2009) or represent range extensions of species that were known to Cowan or species that have been subsequently described in the literature; the majority are of species new to science.
The genus is characterized by being trees with unifoliolate or compound leaves, moniliform fruits and just one petal that may be absent in some cases. Swartzia has beendivided in 15 sections (Torke & Mansano, 2009). Among them, Section Terminales is the largest and most diverse in Colombia. It can be recognized by the combination of pedicels without bracts, flowers with a yellow petal, the stipe and style reduced in relation to the ovary, with numerous seeds and moniliform ovary (Torke & Schaal 2008; Torke & Mansano, 2009). Although section
the Pacific lowlands and inter−Andean valleys of Colombia, where a number of species are highly localized and threatened by habitat destruction, including the four species described herein.
In the course of the investigation, field collections were carried out in Antioquia, the middle Magdalena river valley and Vaupes. Part of the material collected has been instrumental in the process of identifying and describing four new species. As an additional result of the field work, a species of Swartzia was found growing at 2200 m altitude, the highest record for the genus in Colombia, making it a component of the pre−montane wet forest (“bosque humedo premontano”; bh−PM).
It is concluded that tenspecies – out of 23 species so far described for South America − grow in Colombia; in addition, four species are described as new to science and one species is reinstated. Each species was categorized regarding its conservation status following the criteria established by the International Union for the Conservation of Nature (IUCN).
On the basis of a detailed analysis of herbarium material and taking into consideration de geographic distribution of the species, the existence of the S. amplifolia / S. macrophylla
complex is proposed. This artificial complex is recognized on the basis of several morphological characters that are shared by both taxa; there are, however, marked differences in the geographic distribution of the populations involved. Additional field, herbarium and laboratory studies will be needed in order to sort out this complex, as well as to learn about plant−animal relationships, pollinators, and the influence of ants in the development of certain species of Swartzia.
MATERIALS AND METHODS
The research here presented was based mainly on the study of plant collections deposited in the following Colombian and U.S. herbaria: Herbario Nacional Colombiano (COL), Herbario Amazónico Colombiano (COAH), Herbario, Universidad Tecnológica del Chocó, Quibdó (CHOCO), Herbario, Universidad Distrital, Bogotá (UDBC), Instituto Alexander von Humboldt (FMB), Universidad de Antioquia (HUA), Jardín Botánico de Medellín (JAUM), Universidad Nacional sede Medellín (MEDEL), Universidad de los Andes (ANDES), Universidad del Cauca (CAUP), New York Botanical Garden (NY), United States National Herbarium (US) and Missouri Botanical Garden (MO).
Field work was carried out in Colombia during 2010; at that time, the author traveled to Antioquia, Vaupés and the middle Magdalena river valley to collect and observe in the field as many species of Swartzia as possible. The author then (2013) made a short field trip to the municipality of Caldas, department of Antioquia.
The present paper includes detailed descriptions of all 14 species found in Colombia, together with information on geographic distribution in the country, altitude above sea level, habitat, phenology, specimens examined and taxonomic notes. A key to the species of Swartzia
sect. Terminales found growing in Colombia is also included.
In addition, each species was categorized regarding its conservation status following the criteria established by the International Union for the Conservation of Nature (IUCN); these criteria (Table 1) have been applied to several groups of plants in the series of “red books” of threatened and endangered species of Colombia (Linares & Uribe-Meléndez, 2002; Calderón et
al., 2002, 2005; García, 2005; García & Galeano, 2006; Cárdenas & Salinas, 2007; Calderón-Sáenz, 2007). The process of categorization was carried out using information found in herbarium specimens as to locality, habitat, date of collection and so on.
The IUCN categories recognized here are: a) for species growing in Amazonia and Orinoquia, LC = least concern; b) for species found in the Magdalena river valley, Antioquia, Santander, Andean zone and Chocó, VU= Vulnerable; A2c, where A= rapid reduction of population size, 2 = Obvious reduction (observed, estimated, inferred or suspected) in the last 10 years or 3
generations, for reasons that may not be currently present, that are not well known or that can be reversed, and c= Reduction in extent of presence, area occupied and/or habitat quality and EN: Endangered; distribution area small, fragmented or changing; B2b (iii), area occupied estimated at less than 500 km2, severely fragmented, or known to exist only in 5 localities; iii) area, extension or habitat quality, and c) applied only to Swartzia magdalenae , CR = in critical danger; D2, very restricted distribution area.
Swartzia sect.Terminales (R.S. Cowan) Torke & Mansano Taxon 58 (3) 2009: 913–924 Swartzia subsect. Terminales R.S. Cowan, Fl. Neotrop. Monogr. 1: 13. 1968 ≡
Swartzia ser. Tounateoideae Benth.in Martius, Fl. Bras. 15(2): 15. 1870. Type: S. leptopetala Benth.
Lateral leaflets (1–)2–7(–9) jugate; inflorescences ramiflorous or cauliflorous, the bracts estipulate, the pedicels ebracteolate; petal yellow or absent; larger stamens 2–10, occasionally the stamens essentially isomorphic; gynoecium unipistillate, densely pubescent to glabrous, the ovary (2–)2.2–ca.10× as long as wide; the stipe 0.2–1 (–2.5)× as long as the ovary, the style terminal, 0.05–0.5× as long as the ovary, rarely essentially absent, the stigma usually punctiform, occasionally capitate or capitellate; fruits 1 to ca. 12 seeded, the body irregularly elliptic, obovate, oblong, or linear in outline, (1.4)2to ca. 25× as long as wide, usually not strongly compressed, often sub−compartmentalized, with the fruit wall variously constricted between seeds, the arils white, orange, red, or yellow.
Distribution: widespread and diverse in the Amazon basin, Guiana shield, Central America and Pacific lowlands of Colombia, also present in western Cuba and on southern Caribbean islands (Torke & Mansano, 2009).
SPECIES OF SECT. TERMINALES FOUND IN COLOMBIA
1. Swartzia amplifolia Harms
2. Swartzia argentea Spruce ex Bentham
3. Swartzia cabrerae R.S. Cowan
4. Swartzia cardiosperma Spruce ex Bentham
5. Swartzia flavescens (Cowan) Suessenguth
6. Swartzia leptopetala Bentham
7. Swartzia macrophylla Willd. ex Vogel
8. Swartzia magdalenae Britton & Killip
9. Swartzia santanderensis R.S. Cowan
10.Swartzia schultesii R.S. Cowan
11.Swartzia sp nov. 1 L.K. Ruiz, Torke & Mansano
12.Swartzia sp nov. 2 L.K. Ruiz, Torke & Mansano
13.Swartzia sp nov. 3 L.K. Ruiz, Torke & Mansano
KEY TO ESPECIES OF S
WARTZIAS
ECT.T
ERMINALES IN COLOMBIA1. Leaflet surface lustrous to semi−lustrous when dry.
2. Leaflet lower surface tomentulose, tomentose, bright pilose, ferruginous, white, silvery or golden, coriaceous or sub−coriaceous.
3. Leaflets abaxial and adaxial with different colorwhen dry. Fruit not constricted between the seeds, the body ellipsoidto ovoid.
4. Larger stamens 4, smaller stamens 60, seeds per fruit1−4..……..…..S. argentea
4. Larger stamens 2, smaller stamens number 30,
seeds per fruit 15………...…………S. flavescens
3. Leaflets abaxial and adaxial with equal color when dry.Fruit moniliform. 5. Petiole 8 cm long, petiolule 5.5−7.8 mm long, species restricted to the
Amazon region……….S. schultessii
5. Petiole 0.9−5.5 cm long, petiolule 0.7−3 mm long, species restricted to the Magdalena River valley.
6. Leaflets chartaceous, glabrescent……….……….S. magdalenae
6. Leaflets coriaceous to sub−coriaceous, pubescence tomentose to strigulose.
7. Pubescencetomentulose to minute tomentulose to strigulose, often glabrous, secondary venation prominulous on the upper surface of the leaflets; leaflets large, 5−19 cm long x 2.2−3.8 cm wide; rachis 11−27 cm long;smaller stamens <40……… ……….………..S. santanderensis
7. Pubescence ferrugineous, white, silvery or golden, mostly appressed; secondary venation plane on the upper surface of the leaflets; leaflets small, 1.9−13.7 cm long x 0.9−3.5 cm wide; rachis 1.8−10 cm long; smaller stamens>120 ..………...……S. sp. nov. 1 2. Leaflet lower surface minutely strigulose to minutely tomentose, glabrous or nearly so, chartaceous.
8. Stipules 1.4−1.6 mm long, strigulose; inflorescence 40−80 flowered; flower buds 3.5−5.8 x 3.5 mm; calyx segments 4; petal blade 8.8−11 cm long x 6.8−8 mm wide;larger stamen filaments 6.5−9.8 mm long, glabrous; ovary 4.5−5 mm long, densely tomentoseto strigulose………...…..….S. leptopetala
8. Stipules 3.8−6 mm long, tomentulose; inflorescence < 50 flowered; flower buds5−9.2 x 4.5−7.8 mm; calyx segments 5; petal blade 11−20 mm long x 10−24 mm wide,pilose; langer stamen filaments 14−15.5mm long, glabrous; ovary 6.5−10.5 mm long, pilose to tomentose………...….…S. cardiosperma
1. Leaflet surface opaque when dry.
9. Pubescence lanate ferruginous to dark brown in the rachis; species restricted to Vaupes……….……...……….……S. cabrerae
9. Pubescence glabrous to strigulose to tomentose in the rachis.
10. Leaves unifoliolate………..……….S. sp. nov. 3 10. Leaves compound.
11. Venation strongly brochidodromous and prominent on both the upper and lower surfaces, secondary veins penniveined with narrow spaces between the nerves………..………..……S. amplifolia
11. Venation strongly brochidodromous mainly on the lower surface, secondary veins with other types of venation, with wide spaces between the nerves
12. Pedicel 20−39 mm long………..….S. sp. nov. 2 12. Pedicel < 10 mm long.
13. Pubescence dense, sericeous−lanate in filaments of the longer stamens; smaller stamens 126–176; typically few seeds per fruit (1−4)………..………...S. sp. nov. 4 13. Pubescence glabrescent to minute pilose
at the base in filaments of longer stamens; smaller stamens 30−40; typically multiple
TAXONOMIC TREATMENT
1. Swartzia amplifolia Harms, Notizbl. Bot. Gart. Berlin 9: 970. 1926.
Type collection. Peru. Loreto, Río Marañón from Iquitos to the mouthof the Santiago at Pongo de Manseriche. 1924, G. Tessmann 4597(Lectotype F−fragment)
Tree, 15 m tall; pubescence glabrous to strigulose, leaf−bearing branchlets terete, 4 mm thick at middle of internodes, sparsely minute−strigulose to glabrescent. Stipules caducous. Leaves imparipinnate with 3−5 pairs of opposite lateral leaflets, petioles pulvinate, usually unwinged, 6.5−10 cm long, 3.7−4 mm thick at middle, glabrous or nearly so, sometimes sparsely minute−strigulose
,
the pulvinus, 6−11 x 3.3−5.5 mm; rachis minutely caniculate, essentially terete, oftenminutely stipellate at leaflets, 25−33 cm long, 2.6−3.3 mm thick at middle, glabrous to moderately minute−strigulose, the stipels triangular, 0.5 mm long, strigulose; petiolules terete, 5−93 x 2−3.6 mm glabrous or minute−strigulose; leaflet blades chartaceous to subcoriaceous, elliptic to oblong−obovate−elliptic, 2.5−4.2 X as long as wide, those of the basal−most leaflets 10−24.5 x 3−9.6, those of the largest distal leaflets 25−58 x 7−20 cm, glabrous, sometimes lustrous adaxially, the base acute to obtuse−rounded, the apex acute to obtuse often rounted, the acumen acute, 4−9.8 mm long, the abaxial surface glabrescent, often sparingly minute−strigulose on the midrib, the adaxial surface glabrescent sometimes minute lustrous, the midrib and secondary veins immersed and often impressed adaxially, the secondary veins ca. 8−28 paired, initially ascending at ca. 40°−45° with respect to the midrib, strongly brochidodromous.Inflorescences cauliflorus, erect simple racemose, borne on branches, ca. 40−flowered, the flowers spirally arranged, the immature ones densely clustered distally on the inflorescence, terete, often longitudinally ridged to curved, 10−30 cm long, densely strigulose; bracts triangular; 1.3−1.8 mm long, strigulose, persistent; pedicels terete, 3−12 mm long, 1.3−1.5 mm thick at middle, strigulose; bracteoles absent; flower buds, globose often umbonate, 2.7−10.6 x 2.8−9.5 mm, smooth, strigulose. Calyx green, actinomorphic, entire in bud, glabrous adaxially, densely strigulose abaxially, the segments 4, sub−equal, ovate to elliptic, basally truncate, apically acute to obtuse, recurved, 6.5−11.5 x 5.5−6.5 mm. Corolla monopetalous, the petal yellow glabrous, petal peristence, the claw similary a stem, 4 mm long, 1 mm wide, the blade rounded or oblate, the base cordate, 13−17 x 14−16 mm, the venation minute palmate. Androecium strongly zygomorphic, glabrous, the stamens dimorphic, of two distinct sizes, stamens ca. 80 for the pooled, 2 other, dimorphic, the filaments yellow, linear to curved the biggest, the smallest erect to minutely curved, ca. 8−22 mm long, glabrous the anthers brown, oblate, the larger stamens 2 (−4), at the end opposite to petal, glabrous, the filaments yellow, linear to curved, 17−22 mm long, glabrous to minute pilose, the anthers brown, oblong to elliptic, 5−5.5 x 1.3−1.4 m, glabrous, the smaller stamens ca. 80, forming several rows centrally on the floral axis, come from the same point, similar to a range, glabrous the filaments yellow, erect to recurved apex, 8−12 mm long, glabrous to minute strigulose−lustrous, the anthers brown, elliptic ovate apiculate, 1.3−1.5 x 0.7−1 mm, glabrous. Gynoecium monopistillate, minute pilose to strigulose; the gynophore terete, 4−6 mm long, 1 mm thick at middle, strigulose, the ovary arcuate−oblong, 18−28 mm x 1.8−2.6 mm wide at center, pilose to densely strigulose, the style terminal, strongly differentiated from the ovary, terete, 6−6.8 mm long, 0.6−0.8 mm thick, glabrous to sparcely strigulose, the stigma truncate. Fruits green, coriaceous, strigulose to
densely strigulose, dehiscente, the stipe sub−terete to flatted, 3.5 cm long, strigolose, the body moniliform, 22−40 x 1.4−2 cm, smooth, strigulose. Seeds 6−11 per fruit, ellipsoid to obovoid, 22.5−28 x 11−20 mm, the aril present, fimbriate, 10.7 mm long, semi−circular covering apex.
Geographic distribution (Fig 1). Swartzia amplifolia grows in Peru, Ecuador and central and western Colombia, in the departments of Antioquia, Boyacá, Chocó, Cundinamarca, Santander and Valle del Cauca, at altitudes ranging from 0 to 1725 m.
Specimens examined. Antioquia: Mun. Cáceres, cabecera municipal El Doce, Bajo Cauca, zona entre las Quebradas Puri y Corrales, 215 km de Medellín, Quebrada "Pité", creciendo a orilla de quebrada, bosque húmedo y muy húmedo tropical, transición, 200−400 m elev., 11 Jun 1977 (st), Callejas 328 (HUA); Cáceres, El Doce, bajo Cauca, entre las quebradas Puri y Corrales, 200−400 m elev. 28 Dec 1977 (fr), Callejas & Atehortúa 359 (HUA); Mun. Cáceres, cabecera municipal El Doce, Bajo Cauca, zona entre las Quebradas Puri y Corrales, 215 km de Medellín, bosque húmedo y muy húmedo tropical, transición, 200−400 m elev., 25 Mar 1978 (fr) Callejas 487 (COL, HUA); Mun. Cáceres, Vereda Cacerí, margen izquierda quebrada Cacerí, Reserva Natural Regional Refugio Bajo Cauca−Nechi, Selva Húmeda Tropical, 7°30’N, 75°8’W, 430−450 m elev., 11 Jun 1996 (st) Cogollo et al. 8895 (JAUM); Mun. Maceo, Cañón del río Alicante, vereda Alto Buenos Aires, bosque ripario, 6 Nov 2005 (fr), Tuberquia et al. 2561 (HUA); Mun. Maceo, Vereda Santa Bárbara, Vegas del Rio Alicante, hacía la desembocadura del Rio Guardasol, 6°32’38,9’’N, 74°38’24,9’’W, 580 m elev., 22 Nov 2002 (fl), (fr) Fonnegra et al. 7818 (HUA, MO); Mun. Puerto Berrío,vereda Alicante, Finca Penjamo, en la
via que conduce de San Juan de Bedout−La Cabaña, colecciones a lo largo de la quebrada Penjamo, bosque perturbado, 6°39’N, 74°32’W, 380−410 m elev., 1 Mar 1990 (fr) Callejas et al. 9265 (HUA); Mun. Puerto Berrio, corregimiento Calera, Planta Hidroelétrica Calera, margen izquierda de la quebrada “Malena”, 170−200 m elev., 20 Oct 1999 (st), Fonnegra et al. 6958
(HUA, MO); Mun. Puerto Berrío, camino a la cabecera del Municipio, (afluente de la quebrada "Malena"), 230 m elev., 21 Oct 1999 (st), Fonnegra et al. 7030 (HUA); Mun. Puerto Berrío,corregimiento La Cristalina,hacienda Los Angeles, quebrada El Cairo, 300 m elev., 24 Sep 2004, (fr), (fr) Fonnegra et al. 8692 (HUA); Mun. Puerto Berrío, mina Gringos, 1990 (st),
Mahecha & Jiménez 7921 (UDBC); Mun. Remedios, carretera Pto. Berrío: Chorro de Lágrimas, 9−12 Feb 1990 (st), Mahecha et al. 7708 (UDBC); Mun. Tarazá, corregimiento “El 12”, vía El 12−Barroblanco, finca Las Mercedes, a orilla de ríachuelo en bosque, ocasional, bht, 7°28’N, 75°24’W, 450 m elev., 14 Dec 1986, (im), (fr), Callejas et al. 3203 (HUA, K, NY); Mun. Zaragoza, La Planta, Nov−Dec 1982 (fr), Barbosa s.n. (FMB); Mun. Zaragoza, trayecto del Rio Tiquí a Zaragoza, bordeando la carretera, a orilla de río, bosques bordeando la carretera, 7°35’N, 74°50’W, 12 Jul 1987 (fl), Callejas 4651 (HUA, MO, NY); Mun. Zaragosa: Km−14 de la carretera Caceres−Zaragosa, Quebrada Bijagual 7°34’N, 75°10’W, 70 m elev., 26 Mar 1994 (fr),
Giraldo & Zea 66 (HUA); Mun. Zaragoza, Quebrada Cogüí, 2 km antes de la desembocadura del Rio Mata, en el Rio Porce, 150 m elev., 18 Mar 1989 (fr), Fonnegra& Roldán 2693 (HUA, MO, NY). Boyacá: Without date, “bosque reserva”, Aldana et al. P−1 420 (ANDES); El Umbo region of Mt. Chapón, extreme western part of Dept. Boyacá, NW of Bogota, high thick forest, 2500 ft elev., 1 Oct 1932, (fl), Lawrence 495 (A, BM, K, MO, US). Chocó: Mun. Nuquí, quebrada Chaquí, 5°40’N, 77°16’W, 200 m elev., 1994 Feb−Mar 1994 (st), Galeano et al. 4477
Juan, Rio Fujiadó, afluente del Rio San Juan, 4°36’N, 76°54W, 7 Apr 1979, (im), (fr), Forero et al. 4800 (COL, MO); trail between Rio Curiche and Camp Curiche 5−60 m elev., 25 May 1967, (im) (fr), Duke 11598 (NY). Cundinamarca: Mun. Villeta, vega del Río, 3 Oct 1972, (fr im),
Mahecha 9736 (UDBC); Cordillera Oriental: 19.5 km NW of Villeta along highway to Guaduas, growing on steep slope in finca, 1725 m elev., 14 Aug 1972, (fl), (fr im), Barclay et al.3687
(COL, FMB); Santander: Mun. Sabana de Torres: vereda la Puyana, cruce de las Quebradas la Gómez y la Puyana, 100 m elev., 13 Feb 2006 (st), Dueñas & Cruz 1100 (COL); Vicinity of Barranca Bermeja: Magdalena Valley, between Sogamoso and Colorado Rivers, Camp Zarzal, 100−300 m elev., 9 Dec 1934 (bud) (fr), Haught 1446 (NY); Puerto Araujo, 500 m elev., 21 Sep 1979, (fl) (fr), Rentería et al. 1850 (HUA, MEXU, MO); Puerto de Sogamoso, 80 m elev., 1 Nov 1979 (fr), Renteria et al. 2018 (COL, HUA, JUAM, MEXU, MO); Finca el Chapman: Via el Pedral, Puerto de Sogamoso, Corregimiento de Puerto Wilches, 100 m elev., 1 Feb 1980 (fr),
Rentería et al. 2234 (COL, HUA); 10 leguas al SE de Barranca Bermeja, a 8 km de la margen izquierda del Río Opón, 200 m elev., 1 Sep 1954 (st) Romero Castañeda 4796 (COL); 12 leguas al SE de Barranca Bermeja, a 5 km de la margen derecha del Río Opón, 200 m elev., 25 Sep 1954 (fr) Castañeda 4923 (COL); 15 leguas al SE de Barrancabermeja, a 3 km de la margen izquierda del Río Opón, 200 m elev., 8 Oct 1959 (st), Romero Castañeda 4994 (COL); Río Carare, 300−400 m elev., 5 Sep 1945 (st) Hodge 6507 (MEDEL); Magdalena Valley, Campo Capote, 30 km E of Carare, matureforest, being selectively logged, 300 m elev., 30 Sep 1977 (st), Gentry et al. 20075 (HUA, MO); Bucaramanga: Loma de Tigre, Sabana de Torres, 320 m elev., 10 Dec 1977 (fl) (fr) Renteria et al. 32 (MO). Valle del Cauca: Mun. Buenaventura: Costa del Pacífico: Rio Cajambre: Quebrada de Corosal, 0−5 m elev., 17 May 1944 (fr), Cuatrecasas 17737 (F); Costa del Pacífico: Bahía de Buenaventura, Quebrada de San Joaquín, 0−10 m elev.,
21 Feb 1946 (fr), Cuatrecasas 19896 (F, US); Mun. Buenaventura: Bajo Calima, concession, ca. 16 km NW of Buenaventura, at end of Gasolina Road, Juanchaco area 82, wet tropical forest on 20−45° slopes, 3° 50’ N 77° 0’, 50 m elev., 25 Apr 1987 (st), Faber−Langendoen et al. 293
(MO); Mun. Buenaventura: Bajo Calima, Concession, ca. 16 km NW of Buenaventura, at end of Gasolina Road, Juanchaco area, BV−82, wet tropical forest on 20−45° slopes, 3°50’N, 77°0’, 50 m elev., 25 Apr 1987 (st), Faber−Langendoen et al. 314 (MO); Mun. Buenaventura: Bajo Calima Concession, ca. 16 km NW of Buenaventura, at end of Gasolina Road, Juanchaco area, BV−82, wet tropical forest on 20−45° slopes, 3°50’N, 77°0’W, 50 m elev. 28 Apr 1987 (st),
Faber−Langendoen et al. 388 (MO); Mun. Buenaventura: Bajo Calima Concession. Ca 20 km NW of Buenaventura, ca 3km N of km 22.5 on "Hanz", 5°56’N, 77°8’W, 50 m elev., 21 Jun 1988 (st), Faber−Langendoen & Hurtado 1400 (JAUM); Mun. Buenaventura: Bajo Calima, Concession, ca. 20 km NW of Buenaventura, ca. 3 km N of km 22.5 on “Hanz”, remants of primary forest, slopes 5−25°, vegetation recently (less than one year) cut pluvial rainforest, 3°56’N, 77°8’W, 50 m elev., 22 Jun 1988 (st), Faber−Langendoen & Hurtado 1426 (JAUM, MO); Mun. Buenaventura: Bajo Calima, Concession. Ca 17 km NW of Buenaventura, 4 km N of km 22.5 ( mainroad) on "Hanz", plano sedimentario terciario, 3°58’N, 77°6’, 50 m elev., 27 Jul 1988 (st), Faber−Langendoen & Hurtado1857 (JAUM); Mun. Buenaventura: Bajo Calima, concession ca. 20 km N of Buenaventura, recently clearcut primary forest, 3° 40’ N 77°, 50 m elev., 10 Jul 1987 (st), Faber−Langendoen & Rentería 93 (JAUM); Mun. Buenaventura: Bajo Calima, Concession, ca. 15 km NW of Buenaventura, 1 km past Luchin/Lijal intersection, on Luchin, Juanchaco area, four year old secondary wet forest after clear cutting, 3°53N, 77°10W, 50 m elev., 18 Jun 1987 (st), Faber−Langendoen & Renteria 977 (MO); Mun. Buenaventura, Bajo Calima Concession, ca. 16 km NW of Buenaventura, and 2.2 km from Juanchaco gate, on
Tomar road, seconday wet forest after clear cutting, 4°0N, 77°10W, 50 m elev., 23 Jun 1987 (st),
Faber−Langendoen & Renteria 1054 (JUAM, MO); Mun. Buenaventura: Bajo Calima, ca. 10 km due N of Buenaventura, Carton de Colombia concession, transition between tropical wet and pluvial forest, 3°56’N, 77°8’W 50 m elev. 7 Dec 1981, (st), Gentry et al. 35427 (MO); Mun. Buenaventura: Bajo Calima, Juanchaco Palmeras area, ca. 10 km NW of Buenaventura, transition between wet and pluvial forest, 3°56’N, 77°8’W, 50 m elev., 17 Apr 1987 (st), Gentry et al. 57027 (CUVC, MO); Mun. Buenaventura: Bajo Calima, Juanchaco Palmeras area, ca. 10 km NW of Buenaventura, transition between wet and pluvial forest, 3°56’N, 77°8’W, 50 m elev., 18 Apr 1987 (st), Gentry et al. 57040 (CUVC, MO); Mun. Buenaventura, Bajo Calima; Concesión Pulpapel / Buenaventura, bosque pluvial tropical, 3°55’N, 77°0’W, 100 m elev., 27 Nov 1984, (fl), (fr), Monsalve 567 (CUVC, MO).
Habitat. The especies grows in primary forest, riparian forest, humid and very humid tropical forest in aluvial and clayey soils.
Conservation status. VUA2c (Vulnerable, where A= rapid reduction of population size, 2 = Obvious reduction in the last 10 years or 3 generations, and c= Reduction in extent of presence, area occupied and/or habitat quality).
Phenology. Flowers have been seen in March, July, August, September, October, November and December (10 samples studied). Nearly mature fruits were collected in March,
April, May, August, September, October, November and December (12 samples studied), and mature fruit in February, March, May, June, September, October, November and December (15 samples studied).
Notes. Cowan (1968) recognized three varieties within Swartzia amplifolia: S. amplifolia
var. amplifolia, S. amplifolia var. colombiana y S. amplifolia var. rigida. In this study the species has not been subdivided, but the existence of the S. amplifolia / S. macrophylla complex is proposed. This artificial complex is recognized on the basis of several morphologica characters, although there are morphological differences such as the strong brochidodromous venation of the leaflets, penniveined, glabrous and opaque secondary veins, and elongate and small leaves in S. amplifolia; in S. macrophylla the secondary veins are not penniveined, and they do not clearly reach the margin; the gynoecium in S. amplifolia is minutely pilose to strigulose, while in S. macrophylla is densely pilose. Table 2 shows a comparison of selected diagnostic characters for S. amplifolia, S macropylla and all other species of Swartzia section
2. Swartzia argentea Spruce ex Bentham in Martius, Fl. Bras. 15(2): 31. 1870.
Type collection. Brazil. Amazonas: Manáos, Apr. 1851, R. Spruce 1452 (Lectotype BM, isolectotype K)
Tree, to 20 m tall; to 20 cm in diam; pubescence pilose to villous; leaf–bearing branchlets terete, 5–10.8 mm thick at middle of internodes, pilose to villous. Stipules triangular, 4–5 x 2.8–3.1 mm, glabrous to minute villose, often caducous. Leaves imparipinnate, with 3–4 pairs of opposite lateral leaflets, petioles basally pulvinate, unwinged, 2.5–9.5 cm long, 2.5–5.5 mm thick at middle, villose to pilose, the pulvinus, 9–11 x 4–6.6 mm; rachis adaxially caniculate between leaflet pairs, sometimes marginate, rarely stipellate, 13–41 cm long, 2.6–3.8 mm thick at middle, villose to pilose; petiolules terete, 5–8 mm long, 2.5–5.6 mm thick at middle, villous to pilose; leaflet blades coriaceous, elliptic to oblong–elliptic, sometimes oblate to obovate, 2.4– 3 X as long as wide, those of the basal–most leaflets 7.5–31.5 x 4.2–11, those of the largest distal leaflets 10–30 x 5. 5–10.5 cm, villous to pilose, glabrous, sometimes lustrous adaxially, the base obtuse to rounded, sometimes acute, the ápex acute, obtuse to rounded, the acumen acute to retuse, sometimes minute mucronate, 2–13 mm long, the abaxial surface sericeous, hairs golden or silvery, the adaxial surface glabrescent, often sparingly minute–pilose at midrib, sometimes minute lustrous, the midrib and secondary veins immersed and often impressed adaxially, the secondary veins ca. 8–10 paired, initially ascending at ca. 40°–45° with respect to the midrib, strongly brochidodromous. Inflorescences cauliflorus, erect simple racemose, borne on branches, 15 to ca. 50 flowered, the flowers spirally arranged, the immature ones densely clustered distally on the inflorescence, the axes green, terete, often longitudinally ridged to curved, 9.5–43 cm long, villous to pilose; bracts triangular, 1.3–3.3 mm long, villous to pilose,
persistent; pedicels green, 9.4–33 mm long, 2.2–6.8 mm thick at middle, villous to pilose; bracteoles absent; flower buds green, globose, 4.3–9.8 x 5.5–9 mm, smooth, pilose to villose. Calyx green, actinomorphic, entire in bud, dehiscense, glabrous adaxially, densely pilose–villous abaxially, the segments 4, subequal, more or less elliptic, strongly recurved, 7.4–16 x 3.5–4.5 mm. Corolla monopetalous, the petal adaxial, clawed, yellow, densely pilose, adaxially at center venation, peristent often caducius, the claw similary a stem, 2.4–3 mm long, 1.5 mm wide, the blade rounded or oblate, the base cordate, 15–18.5 x 13–15.5 mm the venation minute palmate, transparent. Androecium strongly zygomorphic, glabrous to pilose, the stamens dimorphic, of two distinct sizes, stamens ca. 60 for the pooled, 4 other, dimorphic the langer stamens 4, at the end opposite to petal, glabrous often lanate to villous, the filaments yellow, linear short, 4.8–8 mm long, lanate to villous, the anther orange, oblate to elliptic, 3.3–3.9 x 1.1–1.4 mm, sparcely lanate to villous, the smaller stamens ca.60, forming several rows centrally on the floral axis, come from the same point, similar to a range, glabrous the filaments yellow, wavy, 4.8–9.8 mm long, glabrous, the anthers orange, oblate to elliptic, 1.5–2 x 0.6–0.75 mm, glabrous. Gynoecium monopistillate, pilose to strigulose; the gynophore terete, 2–4.2 mm long, 0.9–1.4 thick at middle, pilose to strigulose, the ovary arcuate–oblong, 4–7.2 mm x 2.5–3 wide at center, pilose to strigulose, the style terminal, recurved, terete, 0.6–1.5 mm long, 4.2–5 mm thick, pilose to strigulose often glabrous, the stigma truncate. Fruits brown often ferrugineus, coriaceous, densamente strigulose to pilose, dehiscente, the stipe sub–terete to flatted, 4–7.2 mm long, strigolose to pilose, the body ellipsoid to ovoid, 4–12 x 1.8–2.8 cm, smooth, pilose to strigulose, the persistent style terminal, terete to compressed, glabrous. Seeds 1–3 per fruit, brownish, ellipsoid to obovoid, 1.7–3.2 x 0.8–1.2 cm, the aril present, white often silver, fimbricate, 4.2– 5.2 mm long, covering ca. one–third of seed.
Geographic distribution (Fig 2). Swartzia argentea grows in the Amazon region of Brazil and Venezuela; in Colombia it has been collected in the departments of Guainia and Vichada, along the margins of the Orinoco and Inirida rivers, between 100 and 220 m above sea level.
Specimens examined. Guainía: Mun. Inírida, Río Inírida, sector entre Chorrobocón y Nabuquen, zona de rebalse; 3°9’6”N, 68°14’ W, 100 m elev., 3 Jan 2007 (fr) Cárdenas et al. 20353 (COAH); Mun. Inirida, Río Inirida, margen izquierda creciendo a la orilla de rio, áreas de rebalse 3° 53’ 48,5” N, 67°56’20”W, 120 m elev., 14 Jan 2005 (fr), Cárdenas & Arenas 16727
(COAH). Vichada: Mun. Cumaribo, resguardo unificado Selva de Mataven, sector Caño Zama, río Orinoco Bosque bajo del plano inundable del caño Mataven, Sitio BI−a, bosque bajo del plano inundable 4°32 11 N, 67° 54 32 W, 220 m elev., 15 Mar 2007 (st) Prieto et al. 5295
(FMB); Mun. Cumaribo, Resguardo unificado Selva de Mataven, sector Caño Zama, río Orinoco, bosque de terra−firme, 4° 32 11 N, 67° 54 32 W, 220 m elev., 15 Mar 2007 (fr) Prieto et al. 5305 (FMB); Mun. Cumaribo, resguardo unificado Selva de Mataven, sector Caño Zama, río Orinoco, 4° 32 11 N, 67° 54 32 W, 220 m elev., 15 Mar 2007 (st), Prieto et al. 5315 (FMB); Mun. Cumaribo, selva de Mataven, resguardo unificado, Selva de Mataven, cuenca baja del caño Mataven, sector Mataven, bosque alto del plano inundable del caño Mataven, sitio BI−b, 4° 30’ 28” N, 68° 3’ 32” W, 190 m elev., 28−29 Mar 2007 (fr) Prieto et al. 5731 (FMB).
Habitat. River margins, forest on periodically inundated grounds and “terra−firme” forest.
Conservation status. LC (Least concern, since this species has been collected in several Amazonian countries).
Phenology. Fruits have been observed between January and March (4 samples studied).
Notes. Cowan (1968) recognized Swartzia argentea as a species closely related to S. ulei; the leaves, however, are morphologicaly more similar to those of S. amplifolia. Cowan (l.c.) placed S. flavescens as a variety of S. argentea; these two taxa as treated here as separate entities, while recognizing that they are closely related. S. argentea can be distinguished by the golden or silvery pubescence of the undersurface of the leaflets and by the shorter stamens.
3. Swartzia cabrerae R.S. Cowan, Fl. Neotrop. Monogr. 1: 56. 1968.
Type collection: Colombia. Amazonas−Vaupés: La Playa, Raudal Yayacopi, Rio Apaporis, Feb 1952, R. E. Schultes and I. Cabrera 15457 (holotype US 2171604, 2171605; isotype NY).
Tree, pubescence lanate, ferruginous; leaf–bearing branchlets terete, 2.4–7.8 mm thick at middle of internodes, lanate, ferruginous. Stipules subulate to lanceolate, 4–8.8 mm long,
strigulose, persistent. Leaves imparipinnate, with 3–4 pairs of opposite lateral leaflets, petioles basally pulvinate, terete, 3–8.8 cm long, 1.3–3 mm thick at middle, tomentulose to strigulose, the pulvinus, 8.8–9.8 x 4.5–5; rachis adaxially minute caniculate between leaflet, stipellate, 16.5–35 cm long, 2–2.5 mm thick at middle, lanate to tomentulose the stipels filiform, 1.7–2.7 mm long, tomentulose to glabrous ; petiolules terete, 4.5–8.8 mm long, 2.6–3 thick at middle, tomentulose to strigulose; leaflet blades subcoriaceous, elliptic to oblong–elliptic, sometimes oblate to obovate, 2.8–3.2 X as long as wide, those of the basal–most leaflets 7.5–23 x 3–6.8, those of the largest distal leaflets 15–23 x 5–7 cm, glabrous or nearly so, sometimes sparsely strigulose on the midrib abaxial and adaxially, the base acute or obtuse, the ápex acute or obtuse, sometimes acuminate, the acumen acute to obtuse, 8–15 mm long, the abaxial surface glabrous or nearly so, sometimes sparsely strigulose at midrib, the adaxial surface glabrous or nearly so, strigulose at midrib, the midrib and secondary veins immersed and often impressed adaxially, the secondary veins ca. 8–10 paired, initially ascending at ca. 40°–45° with respect to the midrib, strongly brochidodromous. Inflorescences cauliflorus, erect to simple racemose, borne on branches, ca. 40– flowered, the flowers spirally arranged, the axes, terete, often longitudinally ridged to curved 13–18 cm long, strigulose; bracts triangular; 1.6–2.1 mm long, strigulose, persistent; pedicels terete, 4–6.5 mm long, strigulose; bracteoles absent; flower buds, umbonate to globose, 7–8 x 5.5–7 mm, smooth, strigulose. Calyx, actinomorphic, entire in bud, dehiscense, glabrous adaxially, strigulose abaxially, the segments 4, subequal, more or less elliptic, deflexed, 7–8.5 x 3.5–5 mm. Corolla monopetalous, the petal adaxial, clawed, yellow, mostly glabrous, sparsely pilose, villose at base and on center vein adaxially, caducius often peristent, the claw similary a stem, 1.6–2.2 mm long, 0.88 mm wide, the blade rounded or oblate, the base cordate, 8–10 x 7– 12 mm, the venation minute palmate, transparent from the more robust central vein.
Androecium strongly zygomorphic, glabrous to pilose, the stamens dimorphic, of two distinct sizes, stamens ca. 100 for the pooled, 6–8 other, dimorphic, the larger stamens 6–8, at the end opposite to petal, sparcely pilose, the filaments yellow, linear, 4.4–6.2 mm long, sparcely pilose, the anthers, elliptic to apiculate, 4.2–4.8 x 0.5–0.7 mm, glabrous, the smaller stamens ca. 100, forming several rows centrally on the floral axis, glabrous the filaments yellow, wavy, 3.8–6.5 mm long, glabrous, the anthers, oblate to elliptic to apiculate, 1.5–2.5 x 0.5 mm, glabrous. Gynoecium monopistillate, glabrous to pilose; the gynophore terete, 1–1.4 mm long, pilose at edge, the ovary oblate to elliptic, 2.2–2.5 mm long, pilose at edge, the style reduced. Fruits not seen. Seeds not seen.
Geographic distribution (Fig 2). Swartzia cabrerae is endemic to Colombia; so far known only from two collections, both from the department of Vaupes, at elevations of about 200 m above sea level.
Specimens examined. Vaupés: Mun. Mitú, por la carretera que conduce de Mitú a Monfort, en área extraida de la Reserva, km 7 al 35 de la carretera, suelos con altos contenidos de arenas blancas 1° 12’ 55,5’ N, 70° 11’ 36’’ W, 18 Sep 2000 (fl), López et al. 6794 (COAH); Rio Apaporis, radual Yayacopi (La Playa) and vicinity, 0° 5’ S, 70° 30’ W, 800 ft elev., 18 Feb 1952 (bud), (fl), (im fr), Schultes & Cabrera 15457 (GH, US). Type of S. cabrerae (holotype: US).
Habitat. The species is found growing on white sand and in riparian environments.
Conservation status. LC (Least concern. Originally known only from the type collection; several new population have been found in recent years).
Phenology. The species has been collected in flower in February and September (2 samples studied) and with immature fruits in February (1 samples studied).
Notes. Swartzia cabrerae may be closely related to S. santanderensis given the number of leaflets and leaflet shape; however, S. cabrerae can be recognized by the elliptic or oblong/elliptic leaflets, lanate pubescence, strongly ferrugineous or dark brown, and by the opaque (vs. lustrous) leaflet surface; in addition, S. cabrerae is restricted to the department of Vaupes. On the basis of the opaque appearance of the leaflet surface and the secundary venation it could be related to S. macrophylla.
4. Swartzia cardiosperma Spruce ex Bentham in Martius, Fl. Bras. 15(2): 33.1870.Type collection. R. Spruce 3361 (lectotype K, isolectotypes BM, BR, C, CGE, F, GOET, M, NY, RB, W).
Tree, to 4–15 m tall; to 10–20 cm in diam.; pubescence minute tomentose; leaf–bearing branchlets terete, 3–4.5 mm thick at middle of internodes, tomentose. Stipules triangular to lanceolate, 3.8–6 mm long, tomentulose, often caducous. Leaves imparipinnate, with 3–5 pairs of opposite lateral leaflets, petioles basally pulvinate, terete, 1.5–6 cm long, tomentulose, the pulvinus, 2.2–5.8 mm long; rachis adaxially caniculate between leaflet pairs, sometimes marginate, rarely winged, stipellate, 4.2–20 cm long, tomentulose, the stipels triangular to filiform, 2–2.5 mm long, minute tomentose, often caducous; petiolules terete sometimes canuculate, 1.8–2.6 mm long, tomentulose to strigulose; leaflet blades chartaceous, elliptic to oblong–elliptic, 3–4.7 X as long as wide, those of the basal–most leaflets 3.5–19 x 1.5–6.2 those of the largest distal leaflets 6–17 x 3.3–7 cm, glabrous or nearly so, strigulose on the midrib adaxially, sparcely pilose abaxially, the base acute to obtuse, the ápex acute or obtuse, the acumen sometimes retuse to mucronate, 5–12 mm long, glabrous sometimes strigulose to pilose, the abaxial surface glabrescent to sparcely pilose, the adaxial surface glabrous or nearly so, strigulose at midrib, the midrib depressed and higher order veins inconspicuous adaxially, salient abaxially. Inflorescences cauliflorus, erect to simple racemose, borne on branches, ca. 50– flowered, the flowers spirally arranged, the immature ones densely clustered distally on the inflorescence, the axes green, terete, often longitudinally ridged to curved, 13–22 cm long, strigulose; bracts triangular; 0.8–2.5 mm long, strigulose, persistent; pedicels terete, 3–27 mm long, strigulose to tomentose; bracteoles absent; flower buds green, globose, 5–9.2 x 4.5–7.8 mm, smooth, strigulose to tomentulose. Calyx green, actinomorphic, entire in bud, dehiscense, glabrous adaxially sometimes densely pilose at base, densely pilose–villous abaxially, the segments 5, subequal, more or less elliptic, deflexed, 7–9 x 4.5–5.5 mm. Corolla monopetalous, the petal adaxial, clawed, yellow, mostly glabrous, sparsely pilose, villose at base and on center
vein adaxially, caducius often peristent, the claw similary a stem, 2.2–7 mm long, the blade obovate to ovate, 11–20 x 10–24 mm, the venation minute palmate or subpalmate, with ca. 4–5 minute, lateral veins braching from the more robust central vein. Androecium strongly zygomorphic, glabrous, the stamens primarily of two sizes, stamens ca. 60 for the pooled, 4 other, dimorphic, the filaments yellow to purple, linear to curved, the larger stamens 4, at the end opposite to petal, pilose, the filaments yellow to purple, linear to curved, 14–15.5 mm long, pilose, the anthers orange, oblong to elliptic to apiculate, 3–6.2 mm long, glabrous, the smaller stamens ca. 60, forming several rows centrally on the floral axis, glabrous to tomentose the filaments yellow sometimes violet at base, wavy to erect, 7.5–11.8 mm long, glabrous to tomentose, the anthers orange, oblate to elliptic to apiculate, apex coriaceous, 2.5–3.2 x 0.5 mm, glabrous to tomentose abaxially. Gynoecium monopistillate, tomentose to pilose; the gynophore terete, 6–11 mm long, tomentose to pilose, the ovary arcuate–oblong, 6.5–10.5 mm long, pilose to tomentose, the style terminal, terete, 2–2.7 mm long, sparcely pilose to glabrous, the stigma truncate. Fruits green, coriaceous, densely strigulose, ferruginous, dehiscente, the stipe sub– terete to flatted, 1.5–2.8 mm long, strigolose, the body moniliform sometimes inequalleteral, 3–8 x 1.4–1.5 cm, smooth, densely strigulose, the persistent style terminal, terete, strigulose. Seeds 1–6 per fruit, ellipsoid to oblate, 11–15 x 8–9 mm, the aril present, fimbriate, 5–5.5 mm long, apex of seed.
Geographic distribution (Fig 2). Swartzia cardiosperma is a species known from eastern Colombia, in the departments of Amazonas, Caqueta, Casanare, Guainia and Vaupes, at elevations of under 500 m above sea level. It has also been collected in the adjacent Amazon regions of Venezuela, Brazil and Peru.
Specimens examined. Amazonas: Araracuara. Cuenca del medio Caquetá, Comunidad de Peña Roja, 27 May 1996 (fr) Rosselli & Ronderos s.n. (COAH); aproximadamente 300 m en dirección 245º de la margen derecha, bajando del Río Caquetá, 4.6 km aguas arriba de la boca del Quebradón de La Culebra, Plot 16, 0° 56 28 S, 71°47 5 W, 25 Aug 1997 (st) Sánchez et al. 5063 (COAH); Corregimiento de Tarapacá, Unidad SD1a, Parcelas 44 y 46, 2°34 29,5 S 70° 5 7,1 W, 16 Mar 1999 (fr) López et al. 5516 (COAH); Corregimiento de Tarapacá, cuenca del Caño Agua Blanquilla, tributario del Río Alegría, bosques con nivel freático superficial e inundaciones, 2° 34 29,5 S, 70° 5 7,1 W, 50 m elev., 11 Apr 2000 (fr) López et al. 6282
(COAH); Corregimiento departamental de la Pedrera, margen izquierda del Río Puré en límite con Brasil, 2.5 km. aguas arriba de la bocana Aguas Negras, bosque maduro sobre superficie alomada con pendientes del 15−30%, 2° 3 22 S, 69° 39 6 W, 190−220 m elev., 25 Jul 1997 (st)
Cárdenas et al. 8134 (COAH); Araracuara, Villa Azul, resguardo indígena Muinanae, terrazas altas disectadas en llanura aluvial, 0° 32 S, 72°6 W, 270−300 m elev., 1 Mar 1993 (st) Duque et al. 1391 (COAH); Amazonas−Vaupes. Rio Apaporis. Soratama (above mouth of Rio Kananarí) and vicinity, 0° 5 N, 70° 40 W, 900 ft elev., 28 Jan 1952 (st), Schultes& Cabrera 14998 (US); Rio Apaporis: Soratama (above mouth of Rio Kananarí) and vicinity, highlands, 0° 5 N, 70° 40 W, 900 ft elev., 6 Feb 1952 (fl), (im fr) Schultes & Cabrera 15172 (US). Caquetá: Araracuara, Trocha del Yari, Noreste de Araracuara, bosque primario, llanura aluvial del Amazonas, 0° 25 S, 72° 20 W, 160 m elev., 26 Mar 1994 (fr), Cárdenas et al. 4506 (COAH); Araracuara, Río Caquetá, camino de fenología, TA1 PII, terraza antigua, without date (st), Bergeron 234−61 (COAH); Araracuara, Río Caquetá, camino de cazería, TA2PVIII, without date (st), Bergeron 540−715 (COAH); Araracuara, Río Caquetá, camino de cazería, TA2PX, terraza antigua alta, without date (st), Bergeron & Román 568−790 (COAH); Araracuara, Río Caquetá, camino de
cazería, TA2PXI, terraza antigua alta, without date (st), Bergeron & Román 624−817 (COAH); Araracuara, trocha a Yari, parcelas experimentales de regeneración, 0° 25 S 72°20 W, 200 m elev.,22 Jan 1989 (st), Gentry & Sánchez 64913 (COAH, MO); Araracuara, bosque maduro, terrazas bajas 0° 37 S, 72°22 W, 2 Dec 1993 (st), Vester & Román 842 (COAH, K); Araracuara, bosque secundario, terrazas bajas, 0° 37 S, 72° 23 W, 15Dec 1993 (st), Vester & Román 851
(COAH); Araracuara, chagra antiguo de la colonia penal, entre la primera y la segunda isla, 22 May 1989 (fr), Wal. 267 (COAH). Casanare: Yopal, cercanias a la inspección el Palmar, Vereda El Aracal, bosque alto del plano inundable del caño Mataven, sitio BI−b, 5° 32 N, 72°22 W, 500 m elev., 9 Jul 1994 (st), Viña 395 (FMB). Guainia: Trocha Nubaquen−Palmira, 2° 50 N 68° 38 W, 25 Feb 1995 (st), Córdoba et al. 581 (COL); Maimachi, Serranía de Naquen. Alrededores del campamento La Planada. Parcela No. 2, zona aluvial, 2° 12 N, 68°12 W, 320 m elev., 10 Aug 1992 (st), Cortés et al. 323 NAQUEN 261 (UDBC).Vaupés: Mun. Taraira, comunidad Jotabeyá, resguardo Yaigoje−Apaporis, camino desde la comunidad a Alsacia, bosque primario y varillal, 0° 38’ S, 70° 11’ W, 150−250 m elev., 29 Mar 2009 (fr), Betancur et al. 13952 (COAH, COL, UDBC); Mun. Taraira, Estación Biológica Caparú within 3 km of the N bank of LagoTaraira, 1° 0 S, 69° 49 W, 200 m elev., 13 Mar 1990 (st), Defler 696 (COAH, MO); Carretera vía Monfort, Km 11.2, 22 Sep 1993 (st), Galeano et al. 1336 (COL); Mun. Mitú. Cucura, Finca La Maye, Propiedad de Jeny Quiñonez 1° 9’ 55,6’’N, 70 °8 ’ 37,8’’, 190 m elev., 21 May 2009 (bud), (im fr), Castro et al. 1658 (COAH); 15 km debajo de Mitú, selva de la orilla izquierda del Río Vaupés, 150 m elev., 28 Jan 1957 (bud), (fl) Uribe 2926 (COL); without date, 28 Jan 1988 (st),
Habitat.This species grows in tropical wet forest (bh−T), on low terrace of mature forest, also on dissected terraces near rivers, and insecondary forest.
Conservation status. LC (Least concern, since this species has been collected in several Amazonian countries).
Phenology. Flowers in January and February (2 samples studied)., fruits in March, April and May (9 samples studied).
Notes. Among the species of sect. Terminales growing in Colombia, Swartzia cardiosperma is closely related to S. leptopetala and S. santanderensis, by its canaliculate rachis, decurrent estipels and leaflets with lustrous appearance when dried; it differs from S. santanderensis in the pubescence glabrous or nearly so, and in the chartaceus vs. coriaceous leaflets. It can be distinguished from S. leptopetala by the longer stipules, 3.8−6 mm vs. 1.4−1.6 mm, the smaller number of flowers per inflorescence (ca. 50 flowered vs. 80 flowered), the number of calyx segments (5 vs. 4) and the ovary dimensions (6.5−10.5 mm long vs. 4.5−5 mm long).
5. Swartzia flavescens Suessenguth, Repert. Spec. Nov. Regni Veg.51: 200. 1942. Type collection. Brazil. Amazonas: Santa Maria, Rio Papory (tributary of Rio Vaupés), Dec 1928, P. Luetzelburg 23860 (M).
Swartzia argentea var. flavescens (Suessenguth) R.S. Cowan, Fl. Neotrop. Monogr.1: 70. 1968.
Tree, to 30 m tall; to 15 cm in diam; pubescence strigulose to densely strigulose, leaf−bearing branchlets terete, 4.2−6.5 mm thick at middle of internodes, strigulose. Stipules caducous. Leaves imparipinnate, with (2) 3−4 pairs of opposite lateral leaflets, petioles basally pulvinate, terete, 3.5−7.5 cm long, strigulose, the pulvinus, 1−14.2 mm long; raquis adaxially minute caniculate between leaflet, stipellate, 13−30 cm long, strigulose to densely strigulose sometimes surface lepidote, the stipels highly reduced or absent; petiolules terete, 4.5−8 mm long, strigulose; leaflet blades coriaceous to subcoriaceous, elliptic to oblong−elliptic, 2.5−3.2 X as long as wide, those of the basal−most leaflets 6.5−32 x 5−15, those of the largest distal leaflets 16−30 x 6−11 cm, glabrous to densely strigulose also pubescence bright, the base obtuse, the apex acute to obtuse, the acumen acute sometimes retuse, 3.5−8 mm long, glabrous to strigulose, the abaxial surface densely strigulose, pubescence bright, the adaxial surface minute lustrous, mostly glabrous, often sparingly minute strigulose on the midrib, the midrib and other venation prominent on both surfaces, the secondaries relatively straight and parallel, also brochidodromous. Inflorescences cauliflorus, erect to simple racemose, borne on branches, ca. 30−flowered, the flowers spirally arranged, the immature ones clustered distally on the inflorescence, the axes green, terete, often longitudinally ridged to curved, 9−21.5 cm long, strigulose; bracts triangular, 1.2−1.6 mm long, strigulose, persistent; pedicels green terete,
7.5−18 mm long, strigulose; bracteoles absent; flower buds green, globose, 6−7 x 5.5−7.5 mm, smooth, strigulose. Calyx green, actinomorphic, entire in bud, dehiscense, glabrous adaxially, densely strigulose abaxially, the segments 4 (−5), subequal, elliptic or obovate, 6.5−8 x 4.5−7.5 mm. Corolla not seen. Androecium strongly zygomorphic, glabrous to pilose, the stamens primarily of two sizes, stamens ca. 30 dimorphic, the filaments yellow the larger stamens 2, at the end opposite to petal, glabrous to pilose, the filaments yellow, 4.5 mm in bud, pilose, the anthers oblate, 3.2 mm long, glabrous, the smaller stamens ca. 30, forming several rows centrally on the floral axis, glabrous the filaments yellow, wavy to erect, ca. 6.5 mm long, glabrous, the anthers oblate, 2.2 x 0.5 mm, glabrous. Gynoecium monopistillate, densely pilose to strigulose; the gynophore terete, 4−4.5 mm long, pilose to strigulose, the ovary inequallateral, 4.5−6.5 mm long, 3.2−4 mm wide, densely pilose to strigulose, the style terminal, terete, 0.8−1.5 mm long, sparcely pilose−strigulose to glabrous, the stigma truncate. Fruits green, coriaceous, densely strigulose, hairs ferruginous with lepidote pubescence, strong and coriaceus, dehiscent, the stipe terete, 5−8 mm long, strigulose, the body ellipsoid to ovoid, 3.5−6.5 x 2.5−2.8 cm, lepidote−coriaceous, densely strigulose, the persistent style terminal, minutely terete, strigulose. Seeds numerous.
Geographic distribution (Fig 2). Swartzia flavescens grows in Amazonian Brazil and Venezuela. In Colombia is known only from the departments of Amazonas and Vaupes, growing below 400 m of altitude.
Specimens examined. Amazonas: Araracuara, margen izquierda Río Caquetá, trocha al Yarí, without date (bud), (fl), Duque et al. 1048 (COAH); Río Cahuinari, 7.5 km al oeste de sus bocas, tra98, 400 m elev., 14 Sep 1988 (st), Sanchez et al. 1184 (COAH); Río Cahuinari, 14 km al oeste de sus bocas, tra101, 400 m elev., 17 Sep 1988 (st), Sanchez et al.1298 (COAH); Corregimiento departamental de la Pedrera, margen izquierda del Río Puré en límite con Brasil, 2.5 km aguas arriba del Caño Castillo, 2° 5’ 0’’ S, 69° 37’ 23’’, 200 m elev., 24 Jul 1997 (st),
Cárdenas et al. 8109 (COAH); Corregimiento departamental de la Pedrera, margen derecha del Río Puré, Cananguchal, 2° 8’ 33’’ S, 69° 53’’ 16’ W, 198 m elev., 2 Aug 1997 (st) Cárdenas et al.8487 (COAH); Corregimiento Tarapacá, Río Putumayo, Caño Porvenir, en cercanías de Lago Centro, 2° 33’ 29,3’’ S, 70° 13’ 14,9’’ W, 200 m elev., 13 Dec 1998 (st) Cárdenas et al. 9857
(COAH); Corregimiento Tarapacá, PNN Amacayacu. Sector Lorena (R. Cotuhé), Parcela permanente #2. Cuadrado E3, 3° 2’ S, 70° W, 100 m elev., 27 Jun 1992 (st) Rudas et al. 4354
(FMB); Corregimiento Tarapacá. PNN Amacayacu. Sector Lorena (R. Cotuhé). Parcela permanente #2. Cuadrado E21.3, 2° S, 70° 3’ W, Jul 1992 (st), Rudas et al. 4767 (FMB); Corregimiento Tarapacá, PNN Amacayacu, Sector Lorena (R. Couthé), Parcela permanente # 2, Transecto F5−F4, 3° 2’ S, 70° 0’ W, 100 m elev., 9 Jul 1992 (st), Rudas et al. 5117 (COL); Corregimiento de Tarapacá, 3° 2’ S, 70° W, 100 m elev., 13 Jul 1992 (st), Rudas et al. 5363
(FMB); Corregimiento Tarapacá, PNN Amacayacu, Sector Lorena (R. Cotuhé), Parcela permanente # 2, Cuadrado U13, arbol 907, 3° 2’ S, 70° 0’ W, 100 m elev., 13 Jul 1992 (st),
Rudas et al. 5425 (COL, FMB); Corregimiento Tarapacá. PNN Amacayacu. Sector Lorena (R. Cotuhé). Parcela permanente # 2. Cuadrado U14, 100 m elev., 13 Jul 1992 (st) Rudas et al. 5442
(FMB). Amazonas – Vaupés: Soratama, without date, 2 Apr 1952 (fr), Schultes& Cabrera 16153 (NY, US). Vaupés. Inspección Departamental de Monfoth, Río Tapurí, frontera
Colombo−Brasilera, orillas del Río Papurí, río debajo de Monforth, 0° 39.741'−0 ° 37.226' N, 69° 42.573'−69 ° 37.226' W, 310−325 m elev., 5 Jul 2002 (fr), Castaño & Betancur 1559
(COAH, COL); Mun. Mitú, Pueblo Nuevo, Finca del Sr. Luis Alfredo Mancipe, 1° 8’ 38,7’’ N, 70° 8’ 12,7’’, 190 m elev., 22 Jun 2009 (st), Castro et al. 1738 (COAH); Mun. Pacoa, corregimiento Buenos Aires, margen izquierda Río Apaporis, raudal de Jirijirimo, antes del raudal, 0° 2’ 53’’ S, 70° 56 36 W, 100−300 m elev., 23 Mar 2009 (fr) Cárdenas et al. 22141
(COAH, COL); Río Vaupés, between Mitú and Javareté, AraráCachivera, 14−24 May 1958 (fr),
Schultes 19400 (COL).
Habitat. This species grows in “terra−firme” forest, also in sandy soils near rivers.
Conservation status. LC (Least concern, since this species has been collected in several Amazonian countries).
Phenology. This species has been found fruiting from March to July (4 samples studied).
Notes. Cowan (1968) placed Swartzia flavescens as a variety of S. argentea. These two taxa as treated here as separate entities (see note under S. argentea), In addition to the differences already stated, the fruit of S. flavescens has numerous (up to 15) seeds while S. argentea has between 1 and 4 seeds per fruit. S. flavescens is known to occur in Amazonas and Vaupes, while
6. Swartzia leptopetala Bentham, Jour. Bot. Hooker 2: 87. 1840.
Type collection: Brazil. Amazonas: near Obidos, Oct 1828, L. Riedel 34 (holotype K; isotype LE)
≡Tounatea leptopetala (Bentham) Taubert, Bot. Centralbl. 47: 391. 1891.
≡Tunatea leptopetala (Bentham) Kuntze, Rev. Gen. P1. 1: 211. 1891.
Swartzia discolor Poeppig & Endlicher, Nov. Gen. &Sp. 3: 62. 1845.
Swartzia fugax Spruce ex Bentham in Martius, Fl. Bras. 15(2): 30. 1870.
Tounatea fugax (S pruce ex Bentham) Britton, Bull. Torrey Club 16: 325. 1889.
Tounatea discolor (Poeppig & Endlicher) Taubert, Bot. Centralbl. 47: 391. 1891.
Tunatea discolor (Poeppig & Endlicher) Kuntze, Rev. Gen. PI. 1: 211. 1891.
Tunatea fugax (Spruce ex Bentham) Kuntze, Rev. Gen. PI. 1: 211. 1891.
Swartzia melanoxylon Ducke, Arch. Bot. Jard. Rio de Janeiro 3: 123. 1922.
Swartzia rotundata Cowan, Mem.N.Y. Bot. Gard.8: 116. 1953.
Tree, to 4–25 m tall; to 10–80 cm in diam.; pubescence strigulose; leaf–bearing branchlets terete, 3.2–9.8 mm thick at middle of internodes, strigulose. Stipules triangular, 1.4– 1.6 mm long, strigulose, often caducous. Leaves imparipinnate with 2–5 pairs of opposite lateral leaflets, petioles basally pulvinate, terete, 2.2–6.5 cm long, strigulose, the pulvinus, 0.6–18 mm long; rachis adaxially minute caniculate between leaflet, stipellate, 6.5–20 cm long, strigulose, the stipels alate to triangular, 0.8–3 mm long, densely strigulose; petiolules terete, 1.6–3.2 mm long, strigulose; leaflet blades chartaceous to subcoriaceous, elliptic to oblong elliptic, sometimes oblate to obovate, 2–2.8 X as long as wide, those of the basal most leaflets 1.8–13 x 1.2–6, those of the largest distal leaflets 6–13 x 2.5–5 cm, glabrous or nearly so, often
malpighious, sometimes sparsely strigulose on the midrib abaxial and adaxially, the base acute or obtuse, the ápex acute or obtuse, sometimes retuse, the acumen acute to obtuse, 3.5–6.8 mm long, glabrous or nearly so, sometimes sparsely malpighious to strigulose, the abaxial surface striguloso to malpighious, the adaxial surface glabrescent, often sparingly minute–strigulose on the midrib, the midrib depressed and higher order veins inconspicuous adaxially, salient abaxially. Inflorescences cauliflorus, erect to simple racemose, borne on branches, 40–80 flowered, the flowers spirally arranged, the immature ones densely clustered distally on the inflorescence, the axes green, terete, often longitudinally ridged to curved–pendulous, 4–27 cm long, strigulose; bracts triangular, 0.7–1.1 mm long, strigulose, persistent; pedicels terete, 2.5–10 mm long, 0.3–0.6 mm thick at middle, strigulose; bracteoles absent; flower buds green, umbonate to globose, 3.5–5.8 x 3–5 mm, smooth, strigulose. Calyx green to yellow to green olive, actinomorphic, entire in bud, dehiscense, glabrous adaxially, densely strigulose abaxially, the segments 4, subequal, more or less elliptic, deflexed, 6–8.6 x 2.5–4.8 mm. Corolla monopetalous, the petal adaxial, clawed, yellow, glabrous, caducius often peristent, the claw similary a stem, 1.8–2.8 mm long, the blade cuneate to obovate, 8.8–11 x 6.8–8 mm, the venation minute palmate, transparent. Androecium strongly zygomorphic, glabrous, the stamens primarily of two sizes, stamens ca. 50 for the pooled, 2–4(–8) other, dimorphic, the larger stamens 2–4(–8), at the end opposite to petal, glabrous, the filaments yellow, golden, linear, 6.5– 9.8 mm long, glabrous, the anthers orange elliptic, 1.8–2 mm long, glabrous, the smaller stamens ca. 50, forming several rows centrally on the floral axis, glabrous the filaments yellow, golden, wavy, 5–8.5 mm long, glabrous, the anthers orange, oblate to elliptic to apiculate, 0.8–1.5 x 0.6– 0.8 mm, glabrous. Gynoecium monopistillate, green tomentose to strigulose; the gynophore terete, 2.2–4.8 mm long, sparcely strigulose, the ovary oblate to elliptic, 4.5–5 mm long, densely
tomentose to strigulose, the style reduced, terminal, terete, 0.5–1.3 mm long, sparcely strigulose, the stigma trúncate. Fruits green, coriaceous, densely strigulose, hairs ferruginous with lepidote strong and coriaceus, dehiscente, the stipe sub–terete to flatted, 3–16 mm long, strigolose, the body inequallateal to ellipsoid–ovoid, 1.8–80.5 x 2.2–2.8 cm, smooth, densely strigulose, the persistent style terminal, compressed,strigulose. Seeds 1–3 per fruit.
Geographicdistribution (Fig 2). This species is widely distributed in eastern Colombia, in the departments of Amazonas, Caqueta, Casanare, Guainia, Guaviare, Putumayo, Vaupes and Vichada. It has been found at altitudes ranging from 20 to 1000 m above sea level.
Specimens examined. Amazonas: Río Caquetá, margen derecha, 4.6 km antes de boca de Quebrada Quinché, 20 m, TRA 59, 5 May 1988 (st), Sánchez et al. 218 (COAH); Río Caquetá, margen derecha, bajando 2 km arriba boca Quebrada Quinché; en orilla del río, trans 61, 7 May 1988 (st) Sánchez et al. 278 (COAH); Bocas del Río Bernardo, afluente del Río Caquetá, 160 m elev., 1 Jun 1984 (fr) Palacios et al. 375 (COAH, COL, VEN); Margen derecha del Río Caquetá−sur, 3 km arriba del Quebrada del Quinche, Apr 1990 (st), Urrego et al. 1703
(COAH); Río Yarí, arriba de su desembocadura en el Caquetá, ca. 200 m elev., 23 May 1984 (bud), (fl), (im fr), Jaramillo & Palacios 7897 (COAH, COL, VEN); al lado occidental del Rio Caquetá, entre las comunidades indígenas de Villa Azul y Peña Roja, 0° 37’ S, 72°7’ W, 200 m elev., 22 Feb 1996 (bud), Dulmen AvD391 (COAH, MO). Amazonas−Vaupes: Rio Apaporis: Soratama and vicinity, 0° 5’N,70°40’ W, 900 ft elev., 4 Feb 1952 (bud), (fl), (im fr) Schultes& Cabrera15168 (BM, NY, US). Caquetá: orillas del Rio Apoporis, cerca de Puerto Hevea, 16 Jan
