1
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
Mayo de 2013 2
TABLE OF CONTENTS
Abstract Introduction Materials and methods The genus Swartzia sect. Terminales
Species of sect. Terminales found 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
3
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 is quite 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. 4
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 5 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 been divided 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
Terminales is most diverse in Amazonia, a secondary radiation of the group has taken place in 6 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 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. 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.
7
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 8 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.
9
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).
10
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
14. Swartzia sp nov. 4 L.K. Ruiz, Torke & Mansano
11
KEY TO ESPECIES OF SWARTZIA SECT. TERMINALES IN COLOMBIA
1. 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 12
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 13
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
per fruit (up to 15)………………………….S. macrophylla
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TAXONOMIC TREATMENT
1. Swartzia amplifolia Harms, Notizbl. Bot. Gart. Berlin 9: 970. 1926.
Type collection. Peru. Loreto, Río Marañón from Iquitos to the mouth of 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. 15
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 16 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 17 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
(COL, FMB); Bajo San Juan, bh – T, Feb 1976 (st), Mahecha 9346 (UDBC); Hoya del Rio San 18
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., 19
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 20
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, 21
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
Terminales that occur in the Pacific lowlands and inter−Andean Valleys of Colombia
22
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, 23 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. 24
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. 25
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, 26 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. 27
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).
28
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).
29
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 30 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. 31
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 32 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),
Valencia helipuerto #50 (FMB).
33
Habitat.This species grows in tropical wet forest (bh−T), on low terrace of mature forest, also on dissected terraces near rivers, and in secondary 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).
34
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, 35
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.
36
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 37
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
S. argentea is found in Guiania and Vichada. 38
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 39 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 40 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 41
1944 (bud), (fl), (im fr), Gutierrez&Schultes 623 (COL, MEDEL, US); Mun. Solano,
Chiribiquete, a 20 minutos de la desembocadura del Río Amu y subiendo por el Río Cuñaré, 0°
13’39’’ N,72° 26’38’’ W, 350 m elev., 15 Feb 2001 (fr) Mendoza et al. 13014 (COAH, FMB).
Casanare: Mun. Aguazul, Vereda Cupiagua, 1000 m elev., 2 Oct 1997 (st) López & Arcila 3572
(COAH); Caño San Miguel, 12−13 Jan 1990 (st), Mahecha 6373 (UDBC); Mun. Paz de Ariporo, corr. La Hermosa, Finca Nicaragua, Terraza, 5° 36’ 23.8’’ N, 70° 15’ 35.7’’ W, 113 m elev., 31
Oct 2004 (st), Ramírez et al. 9099 (FMB); Mun. Paz de Ariporo, Corr. La Hermosa, Finca
Nicaragua, Laguna (Dique), 5° 35’ 45,4’’ N, 70°16 8,9 W, 113 m elev., 5 Nov 2004 (bud), (fl),
(im fr), Ramírez Arango & Morales 9147 (COAH, COL). Guainía: Mun. Inírida, Río Guaviare,
Laguna Negra, 4°1' 28,91"N, 68° 0,1'1,94"W [check coordinates], 6 Jul 2008 (fr), Cárdenas et al. 21557 (COAH). Guaviare: Mun. San José del Guaviare, Inspección El Capricho, carretera de
El Capricho a La Carpa; 200−250 m elev., 25 Oct 1995 (fr), Cárdenas et al. 6757 (COAH, NY);
Mun. San José del Guaviare, Vereda Barrancón, finca de Hermes Peña (Bh−T), margen derecha del Río Guaviare, 2° 38 15,3 N 72° 34 19,1 W, 200−400 m elev., Apr 1998 (fr), Cárdenas et al.
9542 (COAH); Mun. San José del Guaviare Sitio Caño Dorado 2° 32 N 72° 56, 300 m elev., 24
Jan 1990 (bud), (fl), (im fr) Marulanda & Márquez 1651 (HUA). Putumayo: Río Igará−Paraná,
100−600 m elev., 27 Jul 1977 (fr) Roa 1207 (UDBC). Vaupés: Estación Biológica Caparú, sur oriente del departamento del Vaupés, límites con el departamento de Amazonas y la República de Brasil, 1°5, S 69° 32 W, 200 m elev., Jun 1999 (fr) Palacios & Rodriguez 375 (COAH); Alto
Vaupés, alrededores de Miraflores, 300 m elev.,17 Feb 1944 (bud), (fl), (im fr) Gutierrez &
Schultes 823 (COAH, COL, MEDEL, NY); Alto Vaupés, Caño de Guaracú, 21 Feb 1944 (bud),
(fl), (im fr), Gutierrez & Schultes 877 (COL, MEDEL); Alto Vaupés, Caño Arara 23 Feb 1944
(im fr),Gutierrez&Schultes 888 (COL, MEDEL, US); Mun. Mitú, alonglower Rio Kubiyú, 11 Jul 42
1975 (fr), Zarucchi 1440 (COL, US); Mun. Mitú, along Río Vaupés, nearmouth of Río Kuduyarí,
25 Jun 1976 (fr), Zarucchi & Balick 1744 (COL, MO, US); Bank of Río Kubiyu, off Río Vaupés,
200 m elev., 25 Mar 1970 (im fr), Soejarto & Lockwood 2422 (COL, HUA, K); Lowerreaches of bank of Río Kudiyari, above the confluence with Río Vaupés, 200 m elev., 28 Mar 1970 (bud),
(fl), (im fr), Soejarto & Lockwood2452 (COL, HUA, K); Mun. Taraira, comunidad de Jotabeyá,
Río Apaporis, Caño Jotabeyá, entre la comunidad y la bocana (Río Apaporis), 0°3748S, 70° 12’
46’’ W, 100−300 m elev., 29 Mar 2009 (fr), Cárdenas et al. 22305 (COAH, COL). Vichada:
Terrirotio Faunístico del Tuparro, Río Tomo, 1 km north of El Centro Administrativo, 100 m elev., 23 Dec 1979 (bud), (fl), (im fr), Vincelli1017 (COL, FMB); Territorio Faunístico del
Tuparro. El Rio Tomo, 1 km. North of El Centro Administrativo, 100 m elev., 28 Feb 1979 (st),
Vincelli 1036 (FMB); Parque Nacional Natural "El Tuparro", alonglower Caño Peinilla, a tributary of the Río Tomo, 5°20’ N, 68°2’ W, 90−110 m elev., 7 Mar 1985 (im fr), Zarucchi&
Barbosa 3632 (COAH, FMB); Mun. Cumaribo, corregimiento Santa Rita. A 500 m al N del centro Administrativo del parque, 5° 21’ 48’’ N, 67° 51’ 36’’ W, 200 m elev., 2 Feb 2004 (fr),
Mendoza & Robles 15384 (FMB).
Habitat.This species grows in tropical wet forest (bh−T), on low terraces of mature forest, also on dissected terraces near rivers, and in secondary forest.
Conservation status. LC (Least concern, since this species has been collected in several
Amazonian countries). 43
Phenology. Swartzia leptopetala flowers from December to May (9 samples studied).
Fruiting can extend from February to October (11 samples studied).although young fruits have been found as early as December (12 samples studied).
Notes. Swartzia leptopetala is closely related to S. cardiosperma, from which it differs in the strigulose vs. minute tomentose pubescence, triangular and smaller stipules, stipels alate to trinagular vs. triangular to filiform, the abaxial surface strigulose to malpighious vs. glabrescent to sparcely pilose, flower buds umbonate to globose vs. globose, calyx segments 4 vs. 5, petal blade smaller (8−11 x 6.8−8 mm vs. 10−20 x 10−24 mm), stamen filaments shorter (6.5−9.8 mm long vs. 14−15.5 mm long), and smaller anthers (1.8−2 mm vs. 3−6.2 mm). In general, then, it can be said that reproductive structures in S. leptopetala are smaller than those of S. cardiosperma.
7. Swartzia macrophylla Willd. ex Vogel, Linnaea 11: 172. 1837.
Type collection: Colombia. 1803, A. von Humboldt 5652 (B, destroyed).
Tree, to 12 m tall; trunk to 28 cm in diam.; pubescence glabrous to strigulose, leaf−bearing branchlets terete, 4.5–5 mm thick at middle of internodes, glabrous to minute strigulose. Stipules caducous. Leaves imparipinnate whit 2–4 pais of opposite lateral leaflets, petioles pulvinate, usually unwinged, 19–33 mm long, 2–3 mm thick at middle, glabrous to minute strigulose, the pulvinus 7–11 x 3–4.5 mm long; rachis minutely caniculate, essentially terete, often minutely 44 stipellate at leaflets, 10–20 cm long, 1–2.2 mm thick at middle, glabrous to moderately minute−strigulose, the stipels ausentes a reducidas to tringulares , 0.8–1 mm long, minute strigulose; petiolules terete, 3–9 x 1.5–4.2 mm glabrous or minute−strigulose; leaflet blades chartaceous to subcoriaceous, elliptic to oblong−elliptic, 1.8–3.2 X as long as wide, those of the basal−most leaflets 2.5–32 x 1.5–10.5, those of the largest distal leaflets 8–20 x 3.5–12 cm, glabrous, the base acete to obtuse, the apex acute, the acumen acute, 3–12 mm long, the abaxial surface glabrescent, often sparingly minute−strigulose on the midrib, the adaxial surface glabrescent, the midrib and secondary veins immersed and often impressed adaxially, the secondary veins ca. 6–10 paired, initially ascending at ca. 40°–45° with respect to the midrib, brochidodromous. Inflorescences cauliflorus, erect simple racemose, borne on branches, 17–50 flowered. the flowes spirally arranged, the immature ones clustered distally on the inflorescence, the axes terete, often longitudinally ridged to curved, 11.5–42 cm long, densely tomentose; bracts triangular, 0.8–1.7 mm long, densely tomentose to pilose, persistent; pedicels terete, 2–6 mm long, 0.8–2 mm thick at middle, tomentose to pilose; bracteoles absent; flower buds ferrugineous, globose, 4–9 x 4–8, smooth, tomentulose to pilose. Calyx light green, actinomorphic, entire in bud, dehiscense, glabrous adaxially, densely tomentose to pilose abaxially, the segments 4, sub−equal, ovate to elliptic, basally truncate, apically acute to obtuse, recurved, 7–10 x 3.8–5 mm. Corolla monopetalous, the petal yellow, mostly glabrous, sparsely pilose, villose at base and on center vein adaxially, petal peristence, the claw similary a stem. 3–
4 mm long, 1 mm wide, the blade rounded or oblate, the base cordate, 8–11 x 14–17 mm the venation palmate. Androecium strongly zygomorphic, glabrous, the stamens dimorphic, of two distinct sizes, stamens 30–40 for the pooled 2–3 other, dimorphic, the filaments pale yellow, linear to curved, 6–9 mm long, glabrous the anthers orange, oblate, 1.5–2 x 1, glabrous the larger 45 stamens 2–3, at the end opposite to petal, glabrous to minute pilose in the base, the filaments pale yellow, linear to curved, 9–15 mm long, glabrous to minute pilose at the base, the anthers orange, oblong to elliptic, 3–3.2 x 1 mm, glabrous, the smaller stamens 30–40, forming several rows centrally on the floral axis, come from the same point, similar to a range, glabrous the filaments pale yellow, erect to undulating between the base and at apex, 6–9 mm long, glabrous, the anthers orange, oblate, 1.5–2 x 1 mm, glabrous. Gynoecium monopistillate, pale, sericeous to pilose; the gynophore terete, 4–6 mm long, 1 mm thick at middle, sericeous to pilose, the ovary arcuate−oblong, 10–13 mm x 1.8–2 mm wide at center, sericeous to pilose, the style terminal, strongly differentiated from the ovary, terete, 1.2–3 mm long, 0.5 mm thick, glabrous, the stigma truncate. Fruits coriaceous, strigulose to densely strigulose, dehiscente, the stipe aplanado, 2–4.2 mm long, strigulose, the body moniliform, 15–22 x 1.2–1.4 cm, smooth, strigulose. Seeds 5–9 per fruit ellipsoid to obovoid, 20 x 11 mm, the aril present, fimbriate, 20 x
7 mm long, covering middle of seed.
Geographic distribution (Fig 1). Swartzia macrophylla is known only from Colombia, where it grows in the departments of Antioquia, Caldas, Cundinamarca, Santander and Tolima, at altitudes ranging from 50 to 1750 m above sea level.
Specimens examined. Antioquia: Mun. Carepa, Finca Tulenapa, Kilometro 1, vía
Carepa−Apartado, 7°46'46''N, 76°45'52''W, 57 m elev., Dec 2009 (st) Díaz et al. 476 (HUA);
Mun. Carepa, Finca Tulenapa, Kilometro 1 vía Carepa−Apartady, 7°46'46''N, 76°45'52''W, 57 m elev., Dec 2009 (st), Díaz et al. 530 (HUA); Mun. El Bagre, Vereda Sabalito, Finca Las 46
Malvinas, llanura aluvial de inundación, bh−T, 7°51'59''N, 74°46'17''W, 60 m elev., 2 Aug 2009
(bud), Botero et al. 1216 (MEDEL); Mun. El Bagre, Finca La Sierrita, 6°0'22''N, 74°36'38''W, 50 m elev., Oct 2009 (st), Díaz et al. 141 (HUA); Mun. Remedios, Vereda El Costeñal, carretera
Remedios a Puerto Berrío, km 10, hacia Río Duguito, Oleoducto Central S. A., 6°55'N, 74°48'W,
800 m elev., Dec 1997 13−14 (st), Roldán et al. 2948 (HUA); Mun. San Luis, Corregimiento El
Prodigio, Vereda Las Confusas, Bmh−T, 6°3'N, 74°49'W, 650−1000 m elev., 6 Apr 2004 (st),
Sánchez et al. 492 (MEDEL). Caldas: Mun. La Dorada, vereda la Habana, bosque muy húmedo premontano tropical, fragmento de bosque intervenido, 250 m elev., 3 Feb 2000 (bud), Bustos
292 (COL); Mun. La Dorada, 200 m elev., Sep 1960 (fr), Espinal & Pérez 278 (COL); Mun. La
Victoria, sector del boque el Tigre, 5°33'13,7''N, 74°52'51,3''W, 600 m elev., Jul 2005 (bud), (fl)
(im fr), (fr), David et al. 1110 (HUA); Mun. La Victoria, sector del bosque el Tigre, plano sedimentario terciario, 5°33'18,1''N, 74°52'51,9''W, 10 Nov 2007 (fr), David et al. 2492 (JAUM);
Mun. Norcasia, Vereda La Guayana, Finca La Moscovita, margen derecha del Río La Miel, bh−T, bosque primario baja intervención, 5°31'16,9''N, 74°54'10,7''W, 425 m elev., 15 Jun 2001
(bud), Duque et al.2403 (HUA); Mun. Norcasia, proyecto hidroeléctrico la Miel, zona de embalse, 400−600 m elev., 13−31 Jul 1999 (bud), Vargas 6193 (HUA); Mun. Samaná, Reserva
Natural Río Manso. Finca Río Manso, P03−10−001#173, 5°41'45''N, 74°52'41''W, 160 m elev., 6
Apr 2001 (st) Jiménez et al. 404 (JAUM). Cundinamarca: Mun. La Esperanza, cafetales, 1280 m elev., Feb 1934 (fr), Garcia Barriga 290 (COL); Mun. La Esperanza, linea del ferrocarril a
Girardot, cafetales del Hotel, 1280 m elev., 2 Oct 1943 (bud), (fl), (im fr), (fr) Garcia Barriga
10887 (COL); Mun. San Francisco, Cordillera Oriental, Finca "El Carmero," El Tablazo entre
Subachoque y San Francisco, 1900−2100 m elev., 26 Jan 1944, (im fr), (fr) Garcia Barriga
11034 (COL); cerca a Santandercito (en el camino viejo), 1600 m elev., 15 Sep 1944 (bud), (fl), 47
(fr), Uribe 825 (COL); Santander: Mun. Cimitarra, Corregimento de Puerto Olaya, Hacienda El
Bosque Potrero Quito, bosque en colina, bosques tropicales del magdalena medio, 6°28,349'N,
74°20,893'W, 27 Aug 1999 (st) Idárraga et al. 1545 (COL); Mun. Girón, 1000 m elev., 18 Jul
1974 (fr), García Barriga & Jaramillo 20541−A (COL); Mun. Cimitarra, Puerto Arturo, bh−T
Humedal, 6°27'523"N, 74°20'965"W, 180 m elev., 31 Jul 1999 (st), (fl?), Idagarra et al. 1416
(HUA). Tolima: Mun. Mariquita, 6 Dec 1994 (st), Cortés−B et al. 1384 (UDBC); Mun. Ibagué,
Vda. Chucuny, 4°28'5''N, 75°4'25,8''W, 750 m elev., 17 Feb 2007 (fr) Pava & Esquivel 24
(COL). Without date, Transect # 1, Jan 1968 (st), Stanley et al. 22 (COL).
Habitat. Swartzia macrophylla grows in tropical forests; contrary to most species of the genus, it is not necessarily found near water courses.
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 July, August, September and October (4 samples studied). Nearly mature fruits were collected in January, July, August and October (4 samples studied) and mature fruits in February, June, July, September and November (7samples studied).
48
Notes. Swartzia macrophylla is closely related to S. amplifolia. As already stated under S. amplifolia, it is here suggested that a species complex including these two species probably exists and needs to be studied in more detail (see under S. amplifolia). S. macrophylla is also close to Swartzia sp.1 y Swartzia sp. 4 with which it is sympatric. The main differences between
S. macrophylla and the new species are the type of venation and the pubescence present in reproductive organs, as well as the number and length of the leaflets.
8. Swartzia magdalenae Britton & Killip, Ann. N.Y. Acad. 35: 191. 1936.
Type collection.J.J. Triana 457 (holotype US; isotype NY), valley of the Rio
Magdalena, Colombia.
Tree or shrub, pubescence glabrous, leaf–bearing branchlets terete, 3–5 mm thick at middle of internodes, glabrous. Stipules lanceolate, 3.1–11.5 mm long, pilulose. Leaves imparipinnate, with 4 pairs of opposite lateral leaflets; petioles basally pulvinate, terete, 1.5–1.8 cm long, glabrous, the pulvinus, 5.8–7mm long; rachis adaxially minute canaliculate between leaflet, stipellate, 8–13 cm long, glabrescent, the stipels filiform; petiolules terete, 1.8–2.8 mm long, glabrous or essentially so; leaflet blades chartaceous, elliptic or oblong elliptic, 2.5–3.5 X as long as wide, the general leaflets 6–10.2 x 2.1–3.7 cm, glabrous, the base acute to obtuse, the
ápex acute to obtuse, the acumen acute, 5.6–6.5 mm long, glabrous, the abaxial surface glabrescent to strigulose, the adaxial surface glabrescent to minute lustrous, the midrib depressed and higher order veins inconspicuous adaxially, salient abaxially. Inflorescences racemose to branched, axillary, ca. 30–flowered, the flowers spirally arranged, the immature ones densely 49 clustered distally on the inflorescence, the axes terete to curved, 5 cm long, densely pilose to tomentulose; bracts persistent, triangular sometimes falcata, 1.3–10.6 mm long, pilose to strigulose; pedicels terete, 1.8–6 mm long, tomentose to pilose; bracteoles absent; flower buds, globose, 6.8–7.2 x 3.3–4 mm, smooth, tomentose to strigulose. Calyx, actinomorphic, entire in bud, dehiscense, glabrous adaxially, tomentose to strigulose abaxially, the segments 3, subequal, more or less oblate–elliptic, 9.5 x 8 mm. Corolla monopetalous, the petal, glabrous, caducius, the claw similary a stem, 5.3 mm long, 1.11 mm wide, the blade rounded or oblate, 29 x 26.5 mm, the venation palmate, with ca. 8 primary veins, the central vein more robust than the others.
Androecium strongly zygomorphic, glabrous, the stamens dimorphic, of two distinct sizes, stamens > 65 for the pooled, 8 other, dimorphic the larger stamens 8, at the end opposite to petal, glabrous, the filaments linear, 12–13 mm long, the anthers elliptic, 3.8–5.1 x 0.6–0.7 mm, glabrous, the smaller stamens ca. 65, forming several rows centrally on the floral axis, glabrous the filaments, wavy, 8–8.5 mm long, glabrous, the anthers, elliptic, 2 x 0.6 mm, glabrous.
Gynoecium style elongate, glabrous; ovary arcuate–linear, ca 14 mm long, 2 mm wide, glabrous; gynophore sparingly villosulose, 6mm long. Fruits not seen. Seeds not seen.
Geographic distribution. Known only from Middle Magdalena river valley, but without a precise locality.
Specimens examined. Colombia. Valley of the Rio Magdalena, without date, 1851−1857
(bud), (fl), Triana 457 (US), Type of S. magdalenae (Holotype: US).
50
Habitat. Unknown.
Phenology. Unknown.
Conservation status. CRD2 (Critical danger; D2, very restricted distribution area. This species is known only from the type specimen, collected by Triana in the middle of the XIX century).
Notes. This poorly known species is similar to Swartzia santanderensis from which it differs in being basically glabrous or glabrescent (vs. tomentose or tomentulose), and in the shorter pulvinus (5.8−7mm long vs. 7.5−10 mm long) and petiole (1.5−1.8 cm long vs. 1.8−5.5 cm long). The two species share the same general area of distribution in the middle Magdalena river valley. This is an area that has been subjected to intensive human activity over many years, and a large part of it has been converted to cattle ranching and monocultural agriculture. While there still remain some forest relics scattered here and there, the area has been seriously degraded due to anthropic influence.
51
9. Swartzia santanderensis R.S. Cowan, Fl. Neotrop. Monogr. 1: 59. 1968.
Type collection. O. Haught 1638 (US 1740168), vicinity of Puerto Barrio, between
Carare and Magdalena Rivers, Santander, Colombia, Apr. 1935.
Tree, to 20 m tall; to 20 cm in diam.; pubescence tomentulose to minute tomentulose to strigulose often glabrous, leaf–bearing branchlets terete, 3–5.5 mm thick at middle of internodes, tomentulose. Stipules lanceolate, 5.5–12 mm long, pilose to tomentulose, often caducous.
Leaves imparipinnate, with 5–7 pairs of opposite lateral leaflets, petioles basally pulvinate, terete, 1.8–5.5 cm long, tomentose to striguloso or minute glabrous, the pulvinus, 7.5–10 mm long; rachis adaxially minute canaliculate between the leaflets, minutely stipellate, 11–27 cm long, tomentulose to strigulose sometimes glabrescent, the stipels filiform to triangular, 1.2–2.8 mm long, tomentulose, often caducous; petiolules terete, 1.5–2.8 mm long, tomentulose; eaflet blades subcoriaceous, elliptic to obovate–elliptic, 3.5–5.5 X as long as wide, those of the basal– most leaflets 5–19 x 2.2–3.8, those of the largest distal leaflets 9–15 x 2.5–3.5 cm, tomentose to glabrous or nearly so, lustrous, sometimes sparsely strigulose to tomentose on the midrib abaxial and adaxially, the base acute or obtuse, the ápex obtuse to redonded–obtuse, sometimes retuse, the acumen acute, sometimes briefly mucronate, 3–12.5 mm long, glabrous to sparcely pilose abaxially, the abaxial surface glabrescent to sparcely tomentulose to pilose, the adaxial surface glabrous or nearly so, minute lustrous, the midrib immersed and often impressed adaxially, raised abaxially and order venation minute impressed. Inflorescences cauliflorus, erect simple racemose, borne on branches, ca. 50– flowered, the flowers spirally arranged, the immature ones clustered distally on the inflorescence, the axes green, terete, often longitudinally ridged to curved, 13–38 cm long, strigulose and/or pilulose; bracts lanceolate, 2–6 mm long, strigulose, 52 often caducous; pedicels green terete, 12–20 mm long, strigulose to tomentose often ferruginous; bracteoles absent; flower buds green, globose, 5.5–8 x 5.5–6.5 mm, smooth, strigulose. Calyx green, actinomorphic, entire in bud, dehiscense, glabrous adaxially, densely strigulose abaxially, the segments 4, subequal, more or less elliptic, deflexed, 6–9.5 x 3.5–5 mm. Corolla monopetalous, the petal adaxial, clawed, yellow, mostly glabrous, sparsely pilose, villose at base and on center vein adaxially, peristent often caducius, the claw similary a stem, 2.5–2.8 mm long, 0.8–1.2 mm wide, the blade rounded or oblate, 9–14 mm long, 8–13 mm wide, the venation minute palmate, transparent. Androecium strongly zygomorphic, glabrous to pilose, the stamens dimorphic, of two distinct sizes, stamens ca. 40 for the pooled, 6 other, dimorphic, the filaments yellow the larger stamens 6, at the end opposite to petal, sparcely pilose, the filaments yellow, linear, 7–9 mm long, pilose to sparcely pilose, the anthers elliptic to apiculate, 3–3.4 x 0.6–0.8 mm, glabrous to pilose, the smaller stamens ca. 40, forming several rows centrally on the floral axis, come from the same point, similar to a range, glabrous the filaments yellow, wavy, 8–8.5 mm long, glabrous, the anthers, oblate to elliptic to apiculate, 1.8–3.4 x 0.4–0.8 mm, glabrous.
Gynoecium monopistillate, densely tomentose to pilose–strigulose; the gynophore terete 5–6.8 mm long, pilose to strigulose, the ovary arcuate–oblong, 6–12 mm long, pilose to strigulose, the style terminal, erete, 1.5–3.8 mm long, glabrous to minute pilose to strigulose, the stigma truncate. Fruits green, coriaceous, strigulose, dehiscente, the stipe terete, 6.5–18 mm long, strigolose, the body moniliform, 4.5–15 cm x 12–20 mm, smooth, strigulose, the persistent style terminal, terete, strigulose. Seeds 1–7 per fruit, ellipsoid to oblong, 18–22 x 12–19 mm, the aril present, fimbriate, 7 mm long, 4 mm wide, covering ca. at apex of seed.
53
Geographic distribution (Fig 1). This species is known only from Colombia, in the departments of Antioquia and Santander, middle Magdalena River valley region, growing at altitudes between 100 and 700 m.
Specimens examined. Antioquia: Mun. Puerto Berrío, La Unión, Hacienda La Unión
(bh−T), bosque secundario, colinas, 6° 23’ 41,5’’ N, 74°32’ 13,2’’W, 230 m elev.,21 Nov 2001
(st), Vélez et al.4636(HUA); Mun. Remedios, Chorro de Lágrimas. Carretera a Puerto Berrio, bh
– T, 9−12 Feb 1990 (st), Mahecha et al. 7630 (UDBC). Santander: Mun. Vélez, 8 km adentro de Campo Capote, 24 Jun1969 (bud), (fl), (fr), Cabrera672 (COL, CUVC); Mun. Puerto Berrio, between Carare and Magdalena Rivers, Carare Valley, growing along ridges, 100−700 m elev.,14 Apr1936 (bud), (im fr) Haught1638 (F, US), Type of S. santanderensis (US); Finca el
Nuevo Amanecer, carretera el Pedral, Puerto de Sogamoso, corregimiento de Puerto Wilches (S),
100 m elev.,30Jan1980 (bud), (fl), (im fr), Rentería et al. 2189 (COL, HUA, JAUM); Finca el
Champan, via El Pedral, Puerto de Sogamoso, corregimiento de Puerto Wilches (S), 100 m elev.,
31 Jan 1980 (bud), (fl), (fr), Rentería et al. 2211 (COL, HUA, JUAM); Campo Capote, 4 km SW of Campo Capote, rain forest with some recent clearing and logging, 6°38’ N, 73°55’, 100−200 m elev., 26 Mar 1971 (im fr), Nee& Morí 3730 (COL, HUA); Campo Capote, 4 km SW of
Campo Capote, rain forest with some recent clearing and logging, 6°38’N, 73°55’, 100−200 m elev.,26 Mar 1971 (st), Nee & Mori 3731 (COL, HUA); 15 leguas al SE de Barranca Bermeja, a
8 km de la margen izquierda del Río Opón, 200 m elev.,26Aug1954 (fr), Romero Castañeda
4726 (COL); 10 leguas al SE de Barranca Bermeja, a 8 km de la margen izquierda del Rio Opón,
200 m elev., 27 Aug1954 (fr), Romero Castañeda 4734(COL, US); Magdalena Valley, La 54
Gomez, 40 km E of Puerto Wilches, sparse forest, hotflats of gray and White sand, 7°24’ N,
73°33’ W, 150 m elev., 15 Apr 1944 (fr), Fosberg & Fassett 21786 (NY, US).
Habitat. This species grows in tropical wet forest (bh−T).
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. The species has been found in flower in January and June (3 samples studied) and in fruit from January to August (5 samples studied).
Notes. Swartzia santanderensis is closely related to Swartzia sp. 1 but is can be distinguished by its larger vegetative structures. Pubescence in S. santanderensis is tomentulose to minutely tomentulose to strigulose vs. ferrugineous, white, silvery or golden on some structures and mostly appressed in Swartzia sp. 1.
55
10. Swartzia schultesii R.S. Cowan, Fl. Neotrop. Monogr. 1: 58. 1968.
Type collection. R. E. Schultes 3948 (US 1953297, 1953298; isotype NY), vicinity of
La Chorrera, Rio Igaraparaná, Amazonas, Colombia, June 1942.
Tree or arborescent shrub to 4 m tall; pubescence tomentose to ferruginous, leaf– bearing branchlets terete, 7–8.6 mm thick at middle of internodes, tomentose. Stipules lanceolate, 6.2–15.2 mm long, tementulose to pilose, often caducous. Leaves imparipinnate, with
4 pairs of opposite lateral leaflets, petioles basally pulvinate, terete, 8 cm long, tomentose to ferruginous, the pulvinus, 10 mm long; rachis adaxially minute caniculate between leaflet, stipellate, 25–30 cm long, tomentose to ferruginous, the stipels filiform to triangular, 2–3 mm long, tomentulose; petiolules terete, 5.5–7.8 mm long, tomentose to pilulose; leaflet blades subcoriaceous, elliptic or ovate–elliptic, 2–3 X as long as wide, those of the basal–most leaflets
11–24 x 5–8.5, those of the largest distal leaflets 23–28 x 8–9 cm, denselly tomentose to glabrous or nearly so, lustrous, sometimes sparsely strigulose to tomentose on the midrib abaxial and adaxially, the base rounded–obtuse, the apex obtuse, sometimes acuminate, the acumen obtuse to rounded, 5–8.5 mm long, piloluse to tomentulose often strigulose, the abaxial surface densely tomentose to pillulose often strigulose, pubescence ferruginous, the adaxial surface glabrescent, often sparingly tomentose at midrib, sometimes lustrous, the midrib impressed adaxially, higher order veins immersed to slightly raised adaxially, all venation salient abaxially, the secondary veins 10–12–paired. Inflorescences cauliflorus, erect simple racemose, borne on branches, ca. 40– flowered, the flowers spirally arranged, the immature ones densely clustered distally on the inflorescence, the axes green, terete, often longitudinally ridged, 13 cm long, tomentose to strigulose; bracts triangular, 1.2–2.5 mm long, tomentose to strigulose, often 56 caducous; pedicels green terete, 5.5–25 mm long, tomentose to pilulose; bracteoles absent; flower buds green, globose, 7.2 –8 x 6–8.3 mm, smooth, tomentose to strigulose. Calyx actinomorphic, entire in bud, dehiscense, glabrous adaxially, stigulose abaxially, the segments 3–
4, subequal, more or less elliptic, recurved, 7–9 x 3.5–7 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, 4.41mim long, 1 mm wide, the blade oblate–orbicular, the base obtuse, 13.5 mm long, 9.5 mm wide, the venation minute palmate, transparent often the more robust central vein. Androecium strongly zygomorphic, glabrous to pilose, the stamens dimorphic, of two distinct sizes, stamens ca. 80 for the pooled, 6 other, dimorphic the larger stamens 6, at the end opposite to petal, pilose, the filaments pilose, the anthers elliptic, 5.2 x 0.6 mm, pilose, the smaller stamens ca. 80, forming several rows centrally on the floral axis often united at base glabrous to pilose the filaments wavy, 5–6.8 mm long, glabrous, the anthers, oblate to elliptic to apiculate 1.8–3 x 0.4–0.8 mm, glabrous sometimes pilulose. Gynoecium monopistillate, densely pilose to strigulose in bud. Fruits green, coriaceous, tomentulose to strigulose, dehiscente, the stipe terete, 12 mm long, strigolose, the body moniliform, 26 x 1.7 cm, smooth, tomentulose, the persistent style terminal, terete, tomentulose. Seeds 7 per fruit, ellipsoid to oblong, 16–20 x 8–14 mm, the aril present, frimbriate, 17.5 mm long, 4.5 mm wide, enclosing 3/4 of seed.
Geographic distribution (Fig 2). Swartzia schultesii is restricted to the departments of
Amazonas and Caqueta, in the Amazon region of Colombia. It grows at altitudes ranging from
200 to 350 above sea level. 57
Specimens examined.Amazonas; Río Caquetá, Araracuara, 29 Nov 1990 (st), Matapi
163 (COAH); Río Caquetá, Araracuara, 29 Nov 1990 (st) Vester & Matapi 163 (K); Araracuara,
Villazul, Río Caquetá, margen izquierda frente Isla Sumaeta, plano sedimentario terciario, 0° 34’
S, 72° 8’ W, 200−300 m elev., 11 Mar 1989 (st), Londoño et al. 1131 (COL); Quebrada El
Mochilero, afluente del Yarí, márgenes y zona de rebalse aluvial, 150 m elev., 24 Apr 1986
(bud) Galeano et al. 1133 (COAH, COL, MO, NY); Rio Igaraparaná, los alredadores de La
Chorrera, 180 m elev., 4−10 Jun 1942 (bud), (fl), Schultes 3948 (NY, US), Type of S. schultesii
(US 1953297, 1953298); Rio Caqueta, alrededores de Araracuara, Aeropuerto Araracuara, sabana y borde de rastrojo Norte, 10−22 Nov 1982 (bud), (fr), Idrobo et al. 11446 (COAH, COL,
NY). Caquetá: Solano, PNN Serranía de Chiribiquete, cuenca alta del Río Mesay, borde del Río
Mesay, borde del río al inicio de bosque de varilzal, 0° 14’ 24’’ N, 72° 56’ 2’’ W, 350 m elev., 1
Jan 2000 (fr) Prado et al. 565 (COAH); Araracuara, margen izquierda, Rio Caquetá, Trocha al
Yarí, 12 Sep 1989 (st), Torres Duque et al. 1048 (MO); Araracuara, bosque secundario, terrazas bajas, 0° 37’ S, 72° 23’ W, 1 Dec 1993 (st) Vester & Roman 835 (COAH); Araracuara, bosque secundario terrazas bajas, 0° 37’ S, 72° 23’ W, 28 Dec 1993 (fr), Vester & Roman 867 (COAH,
COL, K); Solano, Chiribiquete, Río Sararamano, Complejo cerro, 15 Apr 2000 (st), Mendoza et al. 8452 (FMB); Solano, Chiribiquete. Río Sararamano. Complejo cerro, bosque transicional entre Bosque inundable y Bosque de tierra firme, suelo arenoso, 11° 11’ N, 72° 36’ 20’’ W, 350 m elev., 15 Apr 2000 (st), Mendoza et al. 8460 (FMB); Solano, Chiribiquete, Río Sararamano, bosque de tierra firme, 10° 47’ N, 72° 37’ 24’’ W, 350 m elev., 15 Apr 2001 (st), Mendoza et al.
11011 (FMB); Solano, PNN Serranía de Chiribiquete, Cuenca media del Río Cuñare, bosque de tierra firme, 32° 4’ N, 72° 37’ 57.5’’ W, 350 m elev., 15 Nov 2000 (st), Mendoza et al. 11711
(FMB); Solano, Chiribiquete. a 20 minutos de la desembocadura del Río Amu y subiendo por el 58
Río Cuñaré, bosque de tierra firme, 12° 48’ N, 72° 25’ 25’’ W, 350 m elev., 15 Feb 2001 (st)
Mendoza et al. 12778 (FMB); Solano PNN Serranía de Chiribiquete, Cuenca alta del río Mesay,
bosque de terra−firme, 15° 38’ N, 72° 55’ 24’’ W, 350 m elev., 21 Jan 2000 (st), Mendoza et al.
13610 (FMB); Solano, PNN Serrania de Chiribiquete. Cuenca alta del río Mesay, bosque de
tierra firme, 14° 54’ N, 72° 56’ 5’’ W, 350 m elev., Jan 2000 (st), Mendoza et al. 13790 (FMB);
Solano, PNN Serranía de Chiribiquete, Cuenca alta del Río Mesay, filo cerca al escarpe cerca
hacia el río, 14° 32’ N, 72° 56’ 15’’ W, 350 m elev., 20 Jan 2000 (st) Mendoza et al. 13958
(FMB).
Habitat.This species grows in forest, inundable zones and margins of rivers.
Conservation status. LC (Least concern, since this species has been collected in several localites in Amazonian Colombia).
Phenology. Swartzia schultesii has been found in flower in the month of June (1 sample
studied) and in fruit from November to January (3 samples studied).
Notes. Swartzia schultesii is morphologically similar to S. santanderensis but it can be
distinguished by the larger leaflets which can be up to 28 cm long and 9 cm wide (vs. 19 cm
long, 3.8 cm wide), longer petiole (up to 8 cm long in S. schultesii vs. up to 5.5 cm long in S.
santanderensis. In addition, S. schultesii is restricted to the Amazon region while S. 59 santanderensis grows in the middle Magdalena river valley, in the departments of Antioquia and
Santander.
DESCRIPTION OF NEW SPECIES
Swartzia sp.1 L. K. Ruiz, Torke & Mansano, sp. nov. —TYPE: COLOMBIA.
Santander: near Carare, ca. 45 km SSW of Barrancabermeja, 6°40’N, 74°5’W, 100–200
m elev., 8 Nov. 1967 (bud, fr), J. de Bruijn 1602 (holotype: COL!; isotypes: K!, MO!,
NY!, US!). Fig. 4.
Tree to 15 m+ tall; trunk cylindrical, with woody protuberances in mature individuals exceeding 25 cm diameter; bark rough, exfoliating, grayish−brown; secondary phloem yielding red−oxidizing exudate; pubescence ferrugineous, white, silvery or golden, mostly of appressed, fairly straight to weakly twisting hairs, these generally 0.1–0.5 mm long; distal, leaf−bearing branchlets 1.1–3 mm thick at middle of internodes, densely ferrugineous−strigose to tomentulose, glabrescent. Leaves imparipinnate, with 3–4 (–5) pairs of lateral leaflets; stipules
2.5–8.3 x 0.9–1.6 mm, triangular to oblong−lanceolate, glabrous adaxially, densely strigose abaxially, glabrescent; petioles 0.9–2.7 cm long, 0.8–1.4 mm thick at middle, terete or subterete, longitudinally bicarinate adaxially toward apex, somewhat densely strigose, the pulvinus 1.8–8.8 x 1.1–3.8 mm; rachis 1.8–10 cm long, 0.6–1.4 mm thick at middle or segments, somewhat densely strigose, longitudinally bicarinate or marginate adaxially along segments, the ridges 60 terminating in stipels, the stipels 0.1–1.4 mm long, triangular to ovate−lanceolate, strigose; petiolules 0.7–3 x 0.5–1.5 mm, strigose; laminas 1.9–6 x longer than wide, 1.9–13.7 x 0.9–3.5 cm, the basal ones elliptic or obovate, the distal ones more elongate, usually oblong or narrowly obovate, the base usually acute, sometimes rounded−obtuse in the lowemost leaflets, the apex usually acute or acuminate, occasionally obtuse, the acumen sometimes shortly mucronate, the adaxial surface lustrous, mostly glabrous, with the primary vein strigose, the abaxial surface thinly strigose, more densely so on the major veins, the primary vein impressed and other venation immersed adaxially, all venation prominent abaxially, the secondary veins ca. 11–16 on each side of the primary vein, initially ascending at 30°–48°, fairly straight to somewhat upward curving, more rapidly so near margin, weakly brochidodromous. Inflorescences simple racemes, borne from the trunk and probably also from low−order branches, to ca. 30 flowered; axes 5.5–
18 cm long, 1.4–2.6 mm thick near base, thinly to somewhat densely strigulose; bracts triangular to ovate−lanceolate, 1.3–2.7 x 0.6–1.1 mm, glabrous adaxially, densely strigose abaxially, caducous; pedicels 10.5–13 x 1.2–1.4 mm, somewhat dorso−ventrally compressed, densely strigulose; bracteoles absent; flower buds 5.6–6.6 x 5.2–6.3 mm, globose to ellipsoid, shortly umbonate, somewhat densely ferrugineous−strigose. Calyx segments 4–6 in number, ca. 2.8–6.5 mm wide, recurved, glabrous adaxially, somewhat densely ferruginous−strigose abaxially.
Corolla monopetalous, the petal yellow, essentially glabrous adaxially, somewhat densely strigose on the claw and base of the primary veins abaxially; claw observed only in bud, ca. 1 mm long, perhaps 2 mm long at maturity; limb elliptic, basally obtuse, ca. 11 x 7.4 mm, the venation subpalmate. Androecium strongly zygomorphic, the stamens dimorphic of two size classes; larger stamens ca. 5–6 in number, abaxial, the filaments ca. 5.3–6.1 x 0.5 mm, somewhat dorso−ventrally compressed, only weakly dilated basally or medially, thinly sericeous, 61 particularly on the abaxial side, the anthers 3.6–3.9 x 0.7–0.9 mm, oblong−lanceolate in outline, the connective bluntly acute at apex, prolonged 0.6–0.7 mm belong the thecae, mostly glabrous, sparingly villous at base of connective abaxially; smaller stamens ca. 197, abaxial, glabrous; the filaments ca. 0.6 x 0.2 mm, terete to somewhat dorso−ventrally compressed, the anthers 1.6–2.6 x 0.6–0.8 mm, ovate to oblong, the connective bluntly acute, prolonged 0.4–0.6 mm beyond the thecae. Gynoecium unicarpellate; ovary stipe ca. 4.2 mm long and 0.6–0.7 mm thick at middle, terete, somewhat dilated at base, densely silvery to golden strigose−sericeous; ovary proper ca.
5.1 x 1.5 mm, arcuate−oblong in outline, laterally compressed, densely silvery to golden strigose−sericeous, the locule densely strigose; ovules ca. 15; style 2.2 x 0.5 mm, obliquely terminal, terete, glabrous toward apex; stigma weakly capitellate. Fruits 1–4 seeded, somewhat densely ferruginous−strigulose, more thinly so on seed chambers; stipe 6.2–9.6 x 1.2–1.9 mm, more or less terete; body 3.6–12.1 x 1.4–2 cm, falcate−moniliform, variously constricted between seeds, the constrictions 2.3–8.7 mm wide, the seed chambers ellipsoid, the base and apex acute to acuminate. Seeds 1.9–2.3 x 1.3–1.5 cm, ellipsoid to obovoid; aril ca. 7 mm wide, oblong−linear, semi−encircling seed along hilar side of longitudinal axis.
Distribution (Fig 3). Swartzia sp. nov. 1 is known from two collections from the Carare and Chucurí drainages, tributaries of the Magdalena River in the department of Santander.
Specimens examined. Santander: Campo Capote, 30 km E of Carare in the valley of the
Magdalena River, 300 m elev., 28 Sep 1977 (fl, fr), Gentry et al. 19969 (COL, HUA, MO, US); 62 near Carare, ca. 45 km SSW of Barrancabermeja, 6°40’N, 74°5’W, 100–200 m elev., 8 Nov.
1967 (bud, fr), Bruijn 1602 (COL!; K!, M!, NY!, US!). Type of Swartzia sp. 1.
Habitat. The species occurs in tropical moist forest. One collection was made in secondary forest.
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 (1 sample studied) and fruits (2 samples studied) have been observed in September and November.
Notes. Swartzia sp. nov. 1 is incompletely known; the two collections comprise vegetative organs, fruits, flower buds, and fragmentary material of anthesis flowers. However, data are sufficient to support its specific distinction from its closest relative, S. santanderensis R.
S. Cowan. The two species occur in the same part of the Magdalena Valley and can be easily distinguished from related Andean and Chocoan species of section Terminales in having proportionately narrower leaflets, longer floral bracts, a greater number of larger stamens, and a pubescent ovary locule. The new species differs from S. santanderensis in its less dense, more appressed pubescence on the branchlets, leaves and inflorescence, its more narrow stipules 63
(0.9−1.6 mm wide versus 3–6.1 mm), thinner petiole (0.8–1.4 mm thick at middle versus 1.5–2.8 mm), typically shorter leaf rachis (1.8–10 cm long versus 8–21.5 cm) and flower bracts (1.3–2.7 mm long versus 2.5–6.6 mm), and its more diminutive floral structures (e.g., the larger stamen filaments 5.3–6.1 mm long versus 7.5–12.5 mm; the ovary proper ca. 5.1 mm long versus 6.3–9 mm), hence the epithet. The fact that these characters vary discontinuously between the two species when both occur in sympatry, for example at Campo Capote (S. santanderensis collections: Cabrera 672, Nee & Mori 3731), gives further credence to our taxonomic decision.
Swartia sp. 2 L. K. Ruiz, Torke & Mansano, sp. nov.—TYPE: COLOMBIA. Santander:
near Carare, ca. 45 km SSW of Barrancabermeja, 6°40'N, 74°05'W, 100–200 m elev., 8
Mar 1967 (fl), Bruijn 1605 (holotype: COL!; isotypes: K!, MO!, NY!, US!)...... Fig. 5.
Tree to 6 m+; trunk to ca. 20 cm in diameter; bark rough, grayish−brown; secondary phloem with red exudate; pubescence mostly of appressed golden or ferrugineous trichomes, these mostly less than 0.2 mm long, often borne from a bulbous base, which persists after dehiscence of the apex; distal, leaf−bearing portion of branches ca. 2.5–4.5 mm thick at middle of internodes, papilose−strigulose, glabrescent. Leaves imparipinnate, with (2–) 3–4 pairs of opposite to subopposite lateral leaflets; stipules not seen, apparently caducous, but leaving oval scars, about 1.5 mm wide; petiole 2.1–6.8 cm long including the pulvinus, ca 1.7–3.2 mm thick at middle, terete, longitudinally bicarinate adaxially toward apex, papilose−strigulose, glabrescent, the pulvinus 5–9.2 x 2.2–4.7 mm; rachis 7–18.2 cm long, 1.3–3.1 mm thick at 64 middle of segments, more or less terete, but longitudinally bicarinate adaxially, the ridges terminating in a minute pair of stipels at the insertion of leaflet pairs, papilose−strigulose, glabrescent, the stipels less than 1 mm long, more or less triangular; petiolules 3.3–6.3 x 1.4–3.1 mm, papilose−strigulose, glabrescent; laminas 2.1–3.6 x as long as wide, 6.8–25.6 x 3.2–10.2 cm, thinly chartaceous, elliptic to somewhat obovate, broadly acute at base, shortly acuminate or acute at apex, the adaxial surface essentially glabrous, sometimes thinly strigulose on the midrib, the abaxial surface papilose−strigulose on the midrib and secondary veins, glabrescent, the primary vein depressed and other venation immersed adaxially, all venation raised abaxially, the major secondary veins ca. 10–15 on each side of the primary vein, most fairly straight and ascending at 30°–45° before curving upward and forming strong submarginal loops, these demarcating a prominent submarginal vein ca. 2–7 mm interior from the margin. Inflorescences simple racemes, borne from nodes along the trunk and likely from low−order defoliate branches below leaves, sometimes 2−several fasciculate, ca. 8–30 flowered; axes 9.5–34.5 cm long, 1.3–
4.1 mm thick, densely papilose−strigulose; bracts 0.7–1.5 x 0.7–1 mm, triangular, glabrous adaxially, densely papilose−strigulose abaxially; pedicels 20–39 mm long, ca. 1.8 mm thick at middle, dorso−ventrally compressed, densely papilose−strigulose; bracteoles absent; flower buds
6.6–10.7 x 5.5–10.1 mm, ellipsoid to globose, weakly umbonate, densely papilose−strigulose.
Calyx segments 4–5 in number, 3.3–7.2 mm wide, strongly recurved, glabrous adaxially, densely papilose−strigulose abaxially. Corolla monopetalous, the petal mostly or entirely glabrous, sometimes thinly strigulose on the claw abaxially; claw 2.7–4.4 mm long, ca. 1 mm wide at base and 2.5 mm wide at apex; limb 15.2–19.6 x 15.1–21.6 mm, broadly ovate to oblate, basally obtuse, sometimes obliquely so, shallowly emarginate at apex, the venation palmate, with ca. 7–9 primary veins, the central vein more robust than the others. Androecium strongly zygomorphic, 65 the stamens dimorphic, of two size classes; larger stamens 2, adaxial, the filaments 15–19 mm long, 1.1–1.3 mm thick near base, yellow, dorso−ventrally compressed, apically tapering, thinly sericeous abaxially, the anthers 5.3–6.3 x 1.1–1.3 mm, oblong, the connective acute at apex and prolonged ca. 0.3 mm beyond the thecae, glabrous; smaller stamens ca. 162, abaxial, glabrous, the filaments 9.4–13.6 x 0.1–0.2 mm, yellow, free for all or most of their length, the anthers 1.5–
2.1 x 0.8–1.1 mm, ovate to oblong−elliptic in outline, sometimes oblique at base, the connective acute, rounded or obtuse at apex and prolonged 0.1–0.4 mm beyond the thecae. Gynoecium unicapellate; ovary stipe 4.5–6.6 mm long, ca. 1 mm tick at middle, terete, dilated at base, densely strigose; ovary proper 19–24 x 1.3–1.5 mm, acuate−linear, laterally compressed, densely strigose, the locule glabrous; ovules ca. 20; style 5–9.5 mm, ca. 0.5 mm thick at middle, terminal, terete, partially strigose at base, progressively glabrous toward apex; stigma truncate. Fruits green, ca. 1–9 seeded; stipe ca. 1 cm long and 1.5 mm thick at middle, terete, graduating into the prolonged base of body; body 6–25 cm long, moniliform, linear−lanceolate to lanceolate−elliptic in outline, often falcate, strongly attenuate at base and apex, variously constricted between seeds, the most severe constrictions ca. 2 mm wide, the chambers or subchambers ellipsoid, 1.4–1.8 cm wide, the surface smooth or finely rugose, strigose, somewhat densely so on constrictions and at base and apex, glabrescent on seed chambers, the style persistent.
Geographic distribution (Fig 3). Swartzia sp. nov. 2 is known from the departments of
Antioquia and Santander, where it has been collected in scattered localities between 50 and 500 m elevation in the central Magdalena Valley, on both sides of the river, and also in the lower drainage of the Cauca River, near its confluence with the Nechi River. The apparent disjunction 66 may be an artifact of sampling since the low mountain passes that separate the uppermost Nechi drainage from the Magdalena Valley have been little collected.
Specimens examined. Antioquia: Mun. Nechi, Vereda El 14, entre Mina El 14 y La
Roca del Camino, 8°94’60”N, 74°46’14”W, 60 m elev., 6 Mar 2010 (bud), Rodríguez et al. 6594
(MEDEL); Vereda Santa Elena, cerca de Mina Pablo Camino a Mina Vieja, 8°83’N,
74°46’18”W, 60 m elev., 8 Mar 2010 (fl, fr), Rodríguez et al. 6616 (MEDEL), 8 Mar 2010 (bud),
Rodríguez et al. 6617 (MEDEL); Mun. Puerto Nare, límites con la Vereda El Prodigio (Mun.
San Luis), Vereda Serranías, 6°6’N, 74°48’W, 300–500 m elev., 27 Sep 1990 (fl), Cardenas et al. 3014 (JUAM, MO); Mun. San Luis, Corregimiento de El Prodigio, 6°6’N, 74°48’W, 350–580 m elev., 24 Jun 1990 (fl), Cardenas et al. 2854 (JUAM, MO). Santander: near Carare, ca. 45 km SSW of Barrancabermeja, 6°40'N, 74°05'W, 100–200 m elev., 8 Mar 1967 (fl), Bruijn 1605
(COL!; K!, MO!, NY!, US!). Type of Swartzia sp. 2; Mun. Cimitarra, Vereda Toroba Alta, Finca
La Ponderosa y Vereda Pozo Tortugas, Finca El Cerrito, 6°12'34.7"N–6°12'32.7"N,
73°59'04.7"W, 258 m elev., 21 Sep 2005 (fl), Rodríguez et al. 215A (COL).
Habitat. The species occurs in wet lowland rainforest.
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). 67
Phenology. Flowers have been seen in March, June and September (5 samples studied)..
Nearly mature fruit were collected in March (1 sample studied).
Notes. The elongate pedicels, to which the epithet refers, serve to distinguish Swartzia sp.
2 from a closely related and sympatric species complex containing S. amplifolia, S. macrophylla, and several undescribed segregate species. It also differs from S. macrophylla in having a more prominent submarginal vein in the leaflets, and from S. amplifolia in having fewer pairs of secondary veins in the leaflets. Swartzia sp. 2 is also closely related to S. pinnata of Trinidad and northern Venenzuela, which has somewhat smaller flower parts, typically two pairs of lateral leaflets, with the secondary veins more strongly arcuate−ascending and less prominently brochidodromous.
Swartzia sp. 3 L. K. Ruiz, Torke & Mansano, sp. nov. —TYPE: COLOMBIA. Chocó:
Mun. Quibdó, barrio Margarita, 31 Sep 1984 (fl), W. A. Córdoba & F. García 354 (holotype:
COL!; isotype: CHOCO!,MO!). Fig. 6.
Tree to 10 m+; trunk to 15 cm in diameter; pubescence of appressed white, silvery or ferrugineous, fairly straight, elongate to scale−like trichomes with resinous bases, mostly less then 0.1 mm long, to 0.2 mm long on the androecium; distal, leaf−bearing portion of branches ca. 1–5 mm thick at middle of internodes, thinly papilose or strigulose, glabrescent. Leaves 68 unifoliolate; stipules ca. 0.5–0.7 x 0.5–0.6 mm, triangular, glabrescent, caducous; leaf stalk 2.5–
42 mm long, 1.2–3 mm thick at middle, usually with a prolonged "rachis" above the pulvinate base, occasionally wholly pulvinar, papilose or strigulose, glabrescent, the pulvinus 3.5–6.5 x
2.3–4mm, the "rachis" bicarinnate−marginate adaxially, the ridges terminating in vestigial stipels; petiolule 1.5–5x 1.5– 4.2 mm, pulvinular, papilose or strigulose, glabrescent; lamina 1.8–
2.8 x longer than wide, 11.3–32.8 x 5.3–13.2 cm, chartaceous, elliptic to somewhat ovate or obovate, obtuse or rounded to broadly acute at base, shortly acuminate to broadly acute or obtuse at apex, the adaxial surface essentially glabrous, sometimes briefly pilosulose on the primary vein, the abaxial surface essentially glabrous, but minutely scurfy, glabrescent, the venation, immersed adaxially, with the primary vein and often the secondary veins depressed, all venation raised abaxially, the secondary veins 9−22 on each side of the primary vein, initially ascending at
25°–45°, fairly straight or weakly upward−curving, near margin more rapidly upward−curving and forming prominent submarginal loops ca. 2–4.5 mm from margin. Inflorescences simple racemes borne from defoliate, annotinous or somewhat older portion of branches below leaves, ca. 5–30 flowered; axes 8.5–20 cm long, 1.2–2.9 mm thick near base thinly to densely strigulose; bracts 0.4–1 x 0.6–0.8 mm, triangular to broadly ovate, densely−papilose strigulose, glabrescent; pedicels 3.7–7.8 x 1.1–1.9 mm, dorso−ventrally compressed, densely strigulose; bracteoles absent; flower buds 6.9–12 x 7.3–11 mm, more or less globose, shortly umbonate, somewhat thinly papilose or scurfy−strigulose. Calyx segments 3–4 in number, 3.4–10.4 mm wide, green, strongly recurved, glabrous adaxially, somewhat thinly papilose or scurfy−strigulose abaxially.
Corolla monopetalous; petal yellow, mostly glabrous, sometimes with a few appressed hairs on the claw; claw 2.3–4.2 mm long, 0.7–1.3 mm wide at base, 1.4–1.9 mm wide at apex; limb 17.3–
23.7 x 19–21.6 mm, broadly ovate, orbicular or oblate, basally obtuse, truncate, or rounded, 69 shallowly retuse at apex, the venation sub−palmate with 7–9 primary veins. Androecium strongly zygomorphic, the stamens dimorphic of two size classes; larger stamens 2, abaxial, the filaments ca. 24 mm long, 1–1.5 mm thick near base, dorso−ventrally compressed, basally dilated, sparsely strigose toward base abaxially, the anthers 5.6–6.6 x 1.1–1.3 mm, oblong in outline, glabrous; smaller stamens ca. 90–135, adaxial, glabrous, the filaments 7–15.5 x 0.1–0.3 mm, laterally compressed, apically tapering, free or basally fused, occasionally for nearly half of total of length, the anthers (0.7–) 1.2–2.5 x 0.7–1.5 mm, ovate, elliptic or oblong in outline, often asymmetric at base, often apiculate, with the connective prolonged 0.1–0.2 mm beyond the thecae. Gynoecium unicarpellate; ovary stipe 5.7–7.6 mm long, ca. 0.8–0.9 mm thick at middle, terete, somewhat bulbous at base, densely silvery strigulose except at extreme base; ovary proper
14–17.9 x 1.6–1.9 mm, arcuate−linear, somewhat dilated basally, laterally compressed, densely silvery strigulose, the locule glabrous; ovules ca. 16; style 3.8–5.9 x ca. 0.5 mm terminal, terete, densely silvery strigulose at base, progressively glabrous toward apex; stigma weakly bulbous.
Fruits usually several−seeded; stipe ca. 1 cm long, terete densely strigulose; body to ca. 23 cm long, moniliform, linear−lanceolate to lanceolate−elliptic in outline, densely strigulose at base, otherwise somewhat thinly and minutely papilose, strongly attenuate at base and apex, variously constricted between seeds, the most severe constrictions ca. 3 mm wide, the chambers or subchambers ellipsoid, ca. 1.8 cm wide. Seeds ca. 2.7–3 x 1.6 cm, obovoid; aril more or less conical, the base nearly circular, ca. 11–13 mm in diameter at base, 12.5–14 mm from base to apex, forming an apical "cap" on the seed, reportedly light green, transparent.
70
Geographic distribution (Fig 3). Swartzia sp. nov. 3 is known from the departments of
Antioquia and Chocó, where it has been collected at a handful of sites in the drainage of the San
Juan River and the adjacent upper drainage of the Atrato River in the Chocó ecoregion.
Specimens examined. Antioquia: Mun. Vigia del Fuerte, corregimiento de San Miguel, cuenca de la Quebrada Dengado, afluentes Quebrada Nipundu y Nipunducito, bosques comunitarios sector Nipunducito−Dejengado, 6°33’45.2”N, 76°47’14.8”W, 53 m elev., 29 Apr
2005 (fl), López et al. 10358 (COL, UDBC). Chocó: Río Munduidó, afluente del Río Atrato, alrededores de Altagracia, 40 m elev., 3 May 1975 (fl), Forero et al. 1500 (COL);
Quibdo−Tutunendo road, ca. 3 km W of Tutunendo, Transect 6, 80 m elev., 6 Jan 1981 (st),
Gentry et al. 30218 (JAUM, MO); Quibdo−Tutunendo road, ca. 3 km W of Tutunendo, Transect
9, 5°46’N, 76°35’W, 80 m elev., 7 Jan 1981 (st), Gentry et al. 30216 (JAUM, MO); alrededores de Condoto, 31 Aug 1955 (fr), Idrobo 1900 (COL).
Phenology. Collections with flowers have been made from April to May and in
September (3 samples studied), suggesting that flowering may be sporadic or bimodal. Fruits are known from August (1 sample studied).
Conservation status. ENB2b (iii): Endangered; distribution area small, fragmented or changing; B2b (iii), area occupied estimated at less than 500 km2, severely fragmented, or 71 known to exist only in 5 localities; (iii) area, extension or habitat quality. For example the type locality in Quibdo has been totally replaced by a residential area.
Notes. Swartzia sp. nov. 3 is the only species of section Terminales with strictly unifoliolate leaves. In the context of the genus, the condition occurs fairly often in the seedling stage, but appears as a stabilized state at maturity only in the new species and in distantly related species groups belonging to sections Recurvae, Unifoliolatae and Possira. The new species is perhaps most closely related to S. macrophylla, a species distributed on the other side of the
Cordillera Occidental of the Andes Mountains in the valley of the Magdalena River. In addition to the unifoliolate (versus 3–4 jugate in S. macrophylla) leaves, the thinly papilose or scurfy−stigulose (versus densely strigulose) flower buds set the new species apart. Several collections made in the Darien region (Cárdenas 817, 1071, 1410; see Cárdenas−López, 2003) to the north of the distribution of Swartzia sp. nov. 3 seem to represent a closely related as yet undescribed species with two pairs of lateral leaflets.
Swartzia sp. 4 L. K. Ruiz, Torke & Mansano, sp. nov. —TYPE: COLOMBIA.
Antioquia: Mun. Turbo, carretera Tapón del Darién, sector Rio Leon−Lomas Aisladas, km 11, 20 m elev., 24 Oct 1983 (fl, fr), Brand & González 491 (holotype: COL−285832, 286162!; isotypes:
HUA!, JUAM!, MO!). Fig. 7.
72
Tree to ca. 30 m tall; secondary phloem with red exudate; pubescence mostly ferrugineous and composed of appressed to semi−erect hairs less than 0.2 mm long, these rapidly dehiscing, but the bulbous bases persisting, giving the pubescence a papilose or scurfy appearance, the trichomes on the petal, androecium and gynoecium longer, usually appressed, straight to somewhat twisting, occasionally to 0.8 mm long on the androecium; distal, leaf−bearing branchlets 2–6.5 mm thick at middle of internodes, thinly to densely papilose−strigulose, glabrescent. Leaves with (3–) 4 pairs of opposite lateral leaflets; stipules
0.5–2.1 x 0.6–1.1 mm, triangular to ovate, glabrous adaxially, densely strigulose−papilose abaxially, glabrescent, caducous; petioles 2.3–9.4 (–12) cm long, 1–3.4 (–4.5) mm tick at middle, terete, often bicarinate adaxially toward apex, papilose−strigulose, glabrescent, the pulvinus 3.8–
11.4 (–17.7) x 2.2–4.9 (–6.8) mm; rachis (3.8–) 7–18 (–24.5) cm long, 1–3.2 (–4.2) mm tick at middle of segments, usually longitudinally bicarinate or marginate adaxially, papilose−strigulose, glabrescent, the stipels vestigial or not apparent; petiolules 2.1–8.5 x 1–3.5 mm, pulvinular, papilose−strigulose, glabrescent; laminas 1.5–3.4 x longer than wide, (3.6–)
5.2–24.5 (–33.6) x (2–) 2.3–8.7 (–12.6) cm, the basal ones smaller and less elongate than the others, elliptic to somewhat obovate or oblong, the base acute to obtuse, the apex weakly acuminate or acute to obtuse, the acumen usually rounded at the tip, the adaxial surface essentially glabrous, but usually somewhat papilose or scurfy−strigulose on the primary vein, glabrescent, the abaxial surface minutely papilose or scurfy−strigulose, particularly on high order veins, glabrescent, the venation immersed adaxially, with the primary vein depressed, all venation raised abaxially, major secondary veins ca. 10–15, initially ascending at 25°–40° (–
48°), fairly straight to gradually upward curving, more strongly so near margin, forming submarginal loops, particularly in the distal half of leaflet, but lacking a clearly demarcated 73 submarginal vein. Inflorescences simple racemes, often 2–several fasciculate, borne from annotinous or older defoliate branches or from the trunk, ca. 15–50−flowered; axes 9–28 cm long, 2–3.7 mm thick near base, densely ferrugineous−papilose to scufy−strigulose; bracts 0.6–
1.6 x 0.9–1.5 mm, triangular to broadly ovate, abaxially convex, densely palilose to strigulose; pedicels 4.7–9.7 mm long, 1.3–2.3 mm thick at middle, dorso−ventrally compressed, apically dilated, densely ferrugineous−papilose to scurfy−strigulose; bracteoles absent; flower buds 5.1–
7.4 x 5.1–7.7 mm, globose, to somewhat ellipsoid or oblate−ellipsoid, densely ferrugineous papilose to scurfy−strigulose. Calyx segments 4–6 in number, 3–7.1 mm wide, recurved, glabrous adaxially, densely papilose to scurfy−strigulose abaxially. Corolla monopetalous, the petal yellow, densely to thinly sericeous on the claw and base of veins abaxially, sparingly so or glabrous adaxially; claw (1.9–) 3–4.6 mm long, 0.7–0.9 mm wide at base, 1–2.1 mm wide at apex; limb ovate or elliptic to oblate−ovate, the base cordate to truncate, 10.4–18.7 x 10–18.7 mm, the venation subpalmate. Androecium strongly zygomorphic, the stamens dimorphic of two size classes; larger stamens 2, abaxial, the filaments 7.3–10.3 mm long, 0.8–1.6 mm thick at base, brownish−green, dorso−ventrally compressed, basally dilated, whitish−sericeous to lanate, typically densely so at base and glabrescent toward apex, the anthers 3.5–4.5 x 0.7–1.2 mm, oblong in outline, the connective and often the thecae sparingly sericeous; smaller stamens 126–
176, adaxial, the filaments 5.4–12.4 mm long, 0.1–0.3 mm thick at base, orange, terete to somewhat dorso−ventrally compressed, the anthers 1.2–2.4 x 0.6–1.2 mm, ovate, elliptic or oblong in outline, often apiculate, with the connective prolonged to ca. 0.1 (–0.2) mm beyond the thecae, essentially glabrous, but often with a few long hairs on the connective. Gynoecium unicarpellate; ovary stipe 2.6–4.4 mm long, 0.6–0.9 mm thick at middle, terete, dilated at base, densely golden pilosulose or strigulose; ovary proper 4.1–6.2 x 1.5–1.9 mm, inequilaterally 74 elliptic or oblong−elliptic in outline, densely golden−strigulose, the locule glabrous; ovules 5–9; style 1.3–3.9 x 0.5–0.7 mm, terminal, arcuate, basally dilated, terete, densely strigulose at base, progressively glabrous toward apex; stigma truncate to weakly bulbous. Fruits 1–ca. 4 seeded; stipe 4.2–8.2 x 1.9–3.1 mm, terete, dilated at base; body 4.7–14 x 1.9–2.6 mm, inequilaterally elliptic to oblong in outline, often falcate when multiseeded, variously constricted between seeds, the most severe constrictions to ca. 1 cm wide, the base and apex usually acute, somewhat densely ferrugineous−papilose. Seeds ca. 2.8–3 x 1.8 cm, ellipsoid to obovoid; aril ca. 1.3 cm in diameter, forming an apical "cap" on the seed.
Geographic distribution (Fig 3). Swartzia sp. nov. 4 is known from the departments of
Antioquia and Chocó and is apparently endemic to the Darien area of the northern Chocó ecoregion, where it occurs in the drainage of the Leon River and the lower drainage of the Atrato
River.
Specimens examined. Antioquia: Turbo, 2 km después de Barranquillita, 80 m elev., 9
Jul 1981 (st), Brand & Cogollo 80 (HUA, JAUM, MO); 16 km de Apartadó hacia Chigorodó, margen izquierda, orilla de carretera, 80 m elev., 10 Jul 1981 (fr), Brand & Cogollo 117 (HUA,
JAUM, MO); Vereda Bohios, finca la cabaña, camino los bajos, 30 m elev., 5 Aug 1985 (fr),
Renteria & Cárdenas 4362 (JAUM, MO); Turbo, Tapón del Darién, 50 m elev., 16 Apr 1985
(fr), E. Renteria et al. 3818 (COL, JUAM, MO); Urabá−Chigorodó−Malagón, Caño Malagón, 10 m elev., 22 Mar 1986 (fr), Renteria et al. 4707 (MO); Urabá−Chigorodó−Malagón, sector de la
Quebrada La Puerca y Malagón, 10 m elev., 24 Mar 1986 (fl), Renteria et al. 4747 (JAUM, 75
MO); Mun. Chigorodó, 23−27 km NW de Chigorodó, en la vía a el Rio León, a orilla del Rio
León, 7°37’N, 76°42’W, 50 m elev., 17 Dec 1990 (fl), Callejas et al. 9710 (HUA, NY); Mun.
Chigorodó, Vereda Malagón, Caño Malagón abajo (El Cocuelo), 220 m elev., 11 Jan 1986 (im fr), Renteria et al. 4570 (JAUM, MO); Mun. Mutatá, Las Lomas Aisladas, carretera
Panamericana, 50 m elev., 23 Jan 1985 (fl), Cuadros 2040 (MO, US); Mun. Turbo, carretera
Tapón del Darién, sector Río Leon−Lomas Aisladas, km 29, 20 m elev., Brand & Ascanio 476
(MO, JUAM); Mun. Turbo, carretera Tapón del Darién, sector, Rio Leon−Lomas Aisladas, km
11, 20 m elev., 24 Dec 1983 (fl), Brand & Escobar 729 (HUA, JAUM, MO). Chocó: between
Rio Sucio and La Nueva, May 1967 (fl), Duke 9763 (NY); Rio Truando, between La Nueva and
La Esperanza, 6 Feb 1967 (fr), Duke 9884 (NY, US); upper Rio Truandó, between La Teresita and mouth of Rio Ramón, 20 Jan 1974 (fl), Gentry 9439 (MO, NY); Truando, Dec 1857 (fl),
Schott 6 (NY); Mun. Riosucio, margen derecha del Río Truandó, entre Riosucio y Calle Larga,
22 Aug 1986 (bud), Espina et al. 2166 (COL, HUA, MO); Mun. Riosucio, corregimiento de
Truandó, 6 km arriba de la confluencia del Chintadó, en el Rio Truandó, en las orillas de éste, 21
Oct 1956 (fl), Romero Castañeda 6086 (COL, MO, NY).
Habitat. The existing collections have been made in pluvial rainforest, often in poorly drained or periodically inundated habitats near rivers or streams.
Phenology. Flowering has been observed in June and July (8 samples studied) and fruiting from January to August (7 samples studied).
76
Conservation status. ENB2b (iii) = 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 qualityMost collections of this species were made before the year 1990.
Notes. Swartzia sp. nov. 4 is a morphologically distinctive species that is probably most closely related to the species complex that includes S. amplifolia, S. macrophylla and several undescribed segregate species. It differs from all of these in having dense, sericeous−lanate pubescence on the relatively short filaments of the larger stamens, a smaller, more compact ovary with relatively few ovules, and typically fewer seeds per fruit. The species occurs sympatrically with an apparently undescribed species with a much more elongate gynoecium, longer filaments of the larger stamens, and two pairs of lateral leaflets, with fewer, less prominently brochidodromous secondary veins (Cárdenas 817, 1071, 1410; see discussion under
Swartzia sp. nov 1).
Acknowledgements
The author thanks the curators of the herbaria cited for giving her access to their collections. I am particularly thankful to Dairon Cárdenas−López, Luis Fernando Jaramillo, Alvaro Cogollo,
Alvaro Idarraga, Hurtado, Noberto Lopez, Dino Tuberquia, Felipe Cardona, Carlos Parra, Julio
Betancur, Eva Ledezma, Fabio García and Lauren Raz for facilitating my travels in Colombia 77 and for help in different aspects of my work. Dr. Santiago Madriñán made available for my study the laboratory and library facilities at the Universidad de los Andes and provided help and assistance in many ways during the course of the investigation. A special word of thanks to Dr.
Enrique Forero for his encouragement, his teachings and his support through the years, and to
Dr. Ben Torke por all his teachings and for allowing me to visit The New York Botanical Garden and other botanical institutions in the United States of America through a six-month scholarship.
To Adolfo Jara, doctoral student and Alvinxon Castro, former student, Universidad de los Andes, for the friendship and assistance. I am grateful to my family for their help and encouragement.
Financial support for work on Swartzia has been provided by a grant from the National Science
Foundation (DEB−0918498).
78
Table 1. IUCN categories and criteria for endangered species. (From Cardenas & Salinas, 2007).
79
Table 2. Selected diagnostic characters for the species of Swartzia section Terminales that occur in the Pacific lowlands and inter−Andean Valleys of Colombia
Species No. Leaflet Pedicel Bracteo No. Larger Ovary Ovary
lateral lamina length les larger stamen proper locule
leaflet length/ (mm) length stamen filamen length pubesc
pairs width (mm) s t (mm) ence
ratio length
(mm)
S. amplifolia Harms 3−5 2.5−4.2 3−12 1.3−1.8 2 8−22 18−28 –
S. sp. 1 L. K. Ruiz, 3–5 1.9–6 10.5– 1.3–2.7 5–6 5.3–6.1 ca. 5.1 +
Torke & Mansano 13.0
S. sp. 2 L. K. Ruiz, 2–4 2.1–3.6 20–39 0.7–1.5 2 15–19 19–24 _
Torke & Mansano
S. sp. 3 L. K. Ruiz, 0 1.8–2.8 3.7–7.8 0.4–1.0 2 ca. 24 14– _
Torke & Mansano 17.9
S. sp. 4 L. K. Ruiz, 3–4 1.5–3.4 4.7–9.7 0.6–1.6 2 7.3– 4.1– _
Torke & Mansano 10.3 6.2
S. magdalenae Britton 4 2.5−3.5 1.8−6 − 8 12−13 14 −
& Killip
S. macrophylla Harms 2−4 1.8−3.2 2−6 0.8−1.7 2−3 9−15 10−13 _
S. santanderensis R. 4–7 2.1–6.2 10.8– 2.5–6.6 3–7 7.5– 6.3– +
S. Cowan 18.6 12.5 9.0
80
LITERATURE CITED
Britton, N. L. & E. P. Killip.1936. Mimosaceae and Caesalpiniaceae of Colombia. Ann. New
York Acad. Sci. 35: 191−192.
Calderón, E., G. Galeano & N. García (eds.) 2002. Libro rojo de plantas fanerógamas de
Colombia. I. Chrysobalanaceae, Dichapetalaceae y Lecythidaceae. Libros rojos de
especies amenazadas de Colombia. Intituto Alexander von Humboldt, Instituto de
Ciencias Naturales – Universidad Nacional de Colombia, Ministerio del Medio
Ambiente. Bogotá, Colombia. 218 p.
Calderón, E., G. Galeano & N. García (eds.) 2005. Libro rojo de plantas de Colombia. II.
Palmas, frailejones y zamias. Serie Libros rojos de especies amenazadas de Colombia.
Instituto Alexander von Humboldt, Instituto de Ciencias Naturales – Universidad
Nacional de Colombia, Ministerio de Ambiente, Vivienda y Desarrollo Territorial.
Bogotá, Colombia. 454 p.
Calderón-Sáenz, E. (ed.) 2007. Libro rojo de plantas de Colombia. VI. Orquídeas, primera
parte. Libros rojos de especies amenazadas de Colombia. Instituto Alexander von
Humboldt, Ministerio Ambiente, Vivienda y Desarrollo Territorial. Bogotá, Colombia.
828 p.
Cárdenas, D. & Salinas, N. 2007. Libro rojo de las plantas de Colombia. Especies 81
maderables amenazadas: Primera Parte. Volumen IV. Serie Libros rojos de especies
amenazadas de Colombia. Instituto Alexander von Humboldt, Instituto de Ciencias
Naturales – Universidad Nacional de Colombia, Ministerio de Ambiente, Vivienda y
Desarrollo Territorial. Bogotá, Colombia.
Cardoso, D., L. P. de Queiroz, R. T. Pennington, H. C. de Lima, E. Fonty, M. F.
Wojciechowski & M. Lavin. 2012. Revisiting the phylogeny of papilionoid legumes:
Newinsights from comprehensively sampled early−branching lineages. American Journal
of Botany 99: 1991−2013.
Cowan, R.S.1968. Flora Neotropica Monograph, vol. 1, Swartzia (Leguminosae,
Caesalpinioideae, Swartzieae). Hafner,New York.
Forero, E. 2005. Introducción. En: Forero, E. & C. Romero (eds.), Estudios en Leguminosas
colombianas.Col. Jorge Alvarez Lleras 25: 399−406. Publ. Acad. Colomb.Ci. Ex. Fis.
Nat., Inst. A. v. Humboldt, Inst. Ciencias Naturales y Red Lat. Botánica.Bogotá, D. C.
García, N. (ed.). 2005. Libro rojo de plantas de Colombia. V. Las Magnoliáceas, Las
Miristicáceas y Las Podocarpáceas. Serie Libros rojos de especies amenazadas de
Colombia. Instituto Alexander von Humboldt, Instituto de Ciencias Naturales –
Universidad Nacional de Colombia, Ministerio de Ambiente, Vivienda y Desarrollo
Territorial. Bogotá, Colombia.
García, N. & G. Galeano (eds.) 2006. Libro rojo de plantas de Colombia. III. Las bromelias, las 82
labiadas y las pasifloras. Serie Libros rojos de especies amenazadas de Colombia.
Instituto Alexander von Humboldt, Instituto de Ciencias Naturales – Universidad
Nacional de Colombia, Ministerio de Ambiente, Vivienda y Desarrollo Territorial.
Bogotá, Colombia. 679 p.
Linares, E. L. & J. Uribe-Meléndez 2002. Libro rojo de briófitas de Colombia. Libros rojos de
especies amenazadas de Colombia. Instituto de Ciencias Naturales – Universidad
Nacional de Colombia, Ministerio del Medio Ambiente. Bogotá, Colombia. 232 p.
Torke, B. M. & B. A. Schaal.2008. Molecular phylogenetics of the species−rich neotropical
GenusSwartzia (Leguminosae−Papilionoideae) and related genera of the swartzioid clade.
American Journal of Botany 95: 215–228.
Torke, B. M.& V. F. Mansano.2009. A phylogenetically based sectional classification of
Swartzia (Leguminosae−Papilionoideae). Taxon 58: 913–924.
Torke, B. M.& N. A. Zamora.2010. Notes on Swartzia (Leguminosae) in Central America
preliminary tothe Flora Mesoamericana, with descriptions of two new species from Costa
Rica. Brittonia62:
83
FIGURE LEGENDS
Figure 1. Distribution map, species of Central and Western Colombia.
Figure 2. Distribution map, species of Eastern Colombia.
Figure 3. Distribution map, new Colombian species of Swartzia sect. Terminales.
Figure 4. Swartzia sp. nov. 1
Figure 5. Swartzia sp. nov. 2
Figure 6. Swartzia sp. nov. 3
Figure 7. Swartzia sp. nov. 4
84
Figure 1. Distribution map, species of Central and Western Colombia.
85
Figure 2. Distribution map species of Eastern Colombia.
86
Figure 3. Distribution map, new Colombian species of Swartzia sect. Terminales. 87
Figure 4. Swartzia sp. nov. 1
88
Figure 5. Swartzia sp. nov. 2
89
Figure 6. Swartzia sp. nov. 3
90
Figure 7. Swartzia sp. nov. 4
91
LIST OF EXSICCATAE
Aldana et al.P-1 420 (1).
Barbosa s.n. (1).
Barclay et al. 3687 (1).
Bergeron 234-61 (4); 540-715 (4).
Bergeron & Román 568-790 (4); 624-817 (4).
Betancur et al. 13952 (4).
Botero et al. 1216 (7).
Brand & Ascanio 476 (14).
Brand & Cogollo 80 (14); 117 (14).
Brand & Escobar 729 (14).
Bruijn 1602 (11); 1605 (12).
Bustos 292 (7).
Cabrera 672 (9). 92
Callejas 328 (1); 487 (1); 4651 (1).
Callejas et al. 3203 (1); 9265 (1); 9710 (14).
Callejas & Atehortúa 359 (1).
Cárdenas et al. 2854 (12); 3014 (12); 4506 (4); 6757 (6); 8109 (5); 8134 (4); 8487 (5); 9542 (6);
9857 (5); 20353 (2); 21557 (6); 22141 (5); 22305 (6).
Cárdenas & Arenas 16727 (2).
Castañeda 4923 (1).
Castaño & Betancur 1559 (5).
Castro et al. 1658 (4); 1738 (5).
Cogollo et al. 8895 (1).
Córdoba et al. 581 (4).
Cortés et al. 323 (4); 1384 (7).
Cuadros 2040 (14).
Cuatrecasas 17737 (1); 19896 (1).
David et al. 1110 (7); 2492 (7).
Defler 696 (4).
Díaz et al. 141 (7); 476 (7); 530 (7).
Dueñas & Cruz 1100 (1). 93
Duke 9763 (14); 9884 (14); 11598 (1).
Dulmen AvD391 (6).
Duque et al. 139 (4); 1048 (5); 2403 (7).
Espina et al. 2166 (14).
Espinal & Pérez 278 (7).
Faber-Langendoen et al. 388 (1).
Faber-Langendoen & Rentería 93 (1); 977 (1); 1054 (1); 1400 (1); 1426 (1); 1857 (1) 293 (1);
314 (1).
Fonnegra & Roldán 2693 (1).
Fonnegra et al. 6958 (1); 7030 (1); 7818 (1); 8692 (1).
Forero et al. 1500 (13); 4800 (1).
Fosberg & Fassett 21786 (9).
Galeano et al. 1133 (10); 1336 (4); 4477 (1).
Garcia-Barriga 290 (7); 10887 (7); 11034 (7).
García-Barriga & Jaramillo 20541-A (7).
Gentry 9439 (14); 19969 (11); 20075 (1); 30216 (13); 30218 (13); 35427 (1);
57027 (1); 57040 (1).
Gentry & Sanchez 64913 (4). 94
Giraldo & Zea 66 (1).
Gutierrez & Schultes 623 (6); 823 (6); 888 (6);
Haught 1446 (1); 1638 (9).
Hodge 6507 (1).
Idágarra et al. 1416 (7); Idárraga et al. 1545 (7).
Idrobo 1900 (13).
Idrobo et al. 11446 (10).
Jaramillo & Palacios 7897 (6).
Jiménez et al. 404 (7).
Lawrence 495 (1).
Londoño et al. 1131 (10).
López et al. 5516 (4); 6282 (4); 6794 (3); 10358 (13).
López & Arcila 3572 (6).
Mahecha & Jiménez 7921 (1).
Mahecha 6373 (6); 9346 (1); 9736 (1).
Mahecha et al. 7630 (9); 7708 (1).
Marulanda & Márquez 1651 (6). 95
Matapi 163 (10).
Mendoza et al. 8452 (10); 8460 (10); 11011 (10); 11711 (10); 12778 (10); 13014 (6); 13610
(10); 13790 (10); 13958 (10).
Mendoza & Robles 15384 (6).
Monsalve 567 (1).
Nee & Morí 3730 (9); Nee & Mori 3731 (9).
Palacios et al. 375 (6).
Palacios & Rodriguez 375 (6).
Pava & Esquivel 24 (7).
Prado et al. 565 (10); 5295 (2); 5305 (2); 5315 (2); 5731 (2).
Ramírez Arango & Morales 9147 (6).
Ramírez et al. 9099 (6).
Rentería et al. 32 (1); 1850 (1); 2018 (1); 2189 (9); 2211 (9); 2234 (1); 3818 (14); 4570 (14);
4707 (14); 4747 (14).
Renteria & Cárdenas 4362 (14).
Rodríguez et al. 215A (12); 6594 (12); 6616 (12); 6617 (12).
Roldán et al. 2948 (7).
Romero-Castañeda 4726 (9); 4734 (9); 4796 (1); 4994 (1); 5428 (1); 6086 (14). 96
Rosselli & Ronderos s.n. (4).
Rudas et al. 4354 (5); 4767 (5); 5117 (5); 5363 (5); 5425 (5); 5442 (5).
Sánchez et al. 218 (6); 278 (6); 492 (7) 1184 (5); 1298 (5); 5063 (4).
Schott 6 (14)
Schultes 3948 (10); 19400 (5).
Schultes & Cabrera 14998 (4); 15168 (6); 15172 (4); 15457 (3).
Soejarto & Lockwood 2422 (6); 2452 (6).
Stanley et al. 22 (7).
Torres et al. 1048 (10).
Triana 457 (8).
Tuberquia et al. 2561 (1).
Uribe 825 (7); Uribe 2926 (4).
Urrego et al. 1703 (6).
Valencia 50 (4).
Vargas 6193 (7).
Vélez et al. 4636 (9).
Vester & Matapi 163 (10). 97
Vester & Roman 835 (10); 842 (4); 851 (4); 867 (10).
Vincelli 1017 (6); 1036 (6).
Viña 395 (4).
Zarucchi 1440 (6).
Zarucchi & Balick 1744 (6).
Zarucchi & Barbosa 3632 (6).