Revision of the genus Paragynoxys (Asteraceae, Senecioneae–Tussilagininae)

Revision of the genus Paragynoxys (Asteraceae, Senecioneae-Tussilagininae) ALEJANDRA CORREA

Correa, Alejandra (Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Apartado Atreo 7495, Bogot& Colombia; e-mail: alemoy@tutopia. corn). Revision of the genus Paragynoxys (Asteraceae, Senecioneae-Tussilagininae). Brittonia 55: 157-168. 2003.---A revision of the neotropical genus Paragynoxys of the Asteraceae (tribe Senecioneae) is presented. This group is a distinct entity characterized by a pachycaulous arborescent habit, hornogamous discoid capitula, the presence of white corollas with limbs divided to the base, sagittate anthers, and style branches with enlarged conical tips. In the present treatment, 12 species are accepted. Descriptions of the genus and species, a key to species, synonyms, a new combination (Paraealia lopezii, based on Paragynoxys lopezii), and distribution data are presented. Key words: Paracalia, Paragynoxys, Tussilagininae, Senecioneae, Asteraceae, Colombia, Venezuela. Correa, Alejandra (Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Apartado Atreo 7495, Bogotfi, Colombia; e-mail: alemoy@tutopia. com). Revision of the genus Paragynoxys (Asteraceae, Senecioneae). Brittonia 55: 157-168. 2003.--Se presenta una revisi6n del gtnero neotropical Paragynoxys (Asteraceae, Senecioneae-Tussilagininae). Este taxtn se caracteriza por el hfibito arborescente paquicaule, y por presentar capftulos hom6gamos discoideos, corolas blancas con el limbo dividido hasta la base, anteras sagitadas y ramas estigmfiticas con los extremos alargados y ctnicos. En el presente tratamiento, se aceptan 12 especies. Adem~is, se presentan las descripciones del gtnero y de las especies, una clave de las especies, sin6nimos, una combinacitn nueva (Paracalia Iopezil, basado en Paragynoxys lopezii), y datos de distribuci6n.

Paragynoxys is a rather distinct unit within the subtribe Tussilagininae (Senecioneae: Asteraceae). This genus is distributed in northern South America in the Eastern Cordillera, in depts. Norte de Santander, Santander, and Boyac~i, Colombia; in a restricted area of the Cordillera de Mtrida, western Venezuela; in the Sierra del Perij~i and the Sierra Nevada de Santa Marta, Colombia; and in the western Andes of the departments of Antioquia and C h o c t , Colombia (Figs. 1, 2). The plants can be found at 1 5 0 0 - 4 0 0 0 m, in the montane rain forest, subpfiramo, or p ~ a m o . The first species of this genus were described within Senecio (Cuatrecasas, 1944).

Later Cuatrecasas (1951) created sect. Paragynoxys, which was distinguished from other species of Senecio by its floral structures and striking habit as follows: 1) Pachycaulous trees, the unbranched or slightly branched form with its terminally bunched panicles is typical of this genus; 2) Homogamous discoid capitula (except P. meridana and P. venezuelae, which are often heterogamous) with few involucral bracts and flowers, and with a rudimentary calyculus; 3) Whitish corolla with the limb divided to its base; 4) Sagittate anthers, similar to those of Gynoxys; and 5) Style branches with enlarged conical tips, like those of Gynoxys, but with the conical or lanceolate form less pronounced.

Brittonia, 55(2), 2003, pp. 157-168. 9 2003, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.

ISSUED: 30 June 2003

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f _

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t+,=

/_ ? FIG. 1. Distribution of Paragynoxys angosturae, P. cuatrecasasii, P. magnifolia, P. martingrantii, P. neodendroides, and P. santurbanensis. In 1955, Cuatrecasas elevated the section to a genus. He circumscribed the genus and presented a key to the eight species known at the time. Since then, several additional species have been transferred to P a r a g y noxys or described as new (Cuatrecasas, 1961, 1978, 1984, 1990). In the present paper 12 species o f P a r a g y n o x y s are recognized. The key was adapted by modifying and adding some species to the key of Cuatrecasas (1955). One species is reduced to synonymy and two others are excluded from the genus. Taxonomic Treatment

PARAGYNOXYS (Cuatrec.) Cuatrec. P a r a g y n o x y s (Cuatrec.) Cuatrec., Brittonia

8: 153. 1955. Senecio sect. Paragynoxys Cuatrec., Fieldiana, Bot. 27(2): 72. 1951. TYPE: Senecio neodendroides

Cuatrec. Trees, ca. 3 - 1 5 m; stem simple or fewbranched. L e a v e s alternate, petiolate, ab-

FIG. 2. Distribution of Paragynoxys corei, P. meridana, P. steyermarkii, P. undatifolia, P. uribei, and P. venezuelae.

axial surface c o m m o n l y with a t o m e n t o s e or woolly indumentum. C a p i t u l e s c e n c e s paniculate. C a p i t u l a h o m o g a m o u s (except P a r a g y n o x y s m e r i d a n a and P. v e n e z u e l a e , which usually have heterogamous capitula), discoid; involucre cylindrical, phyllaries 5-8, w e a k l y biseriate or uniseriate, coriaceous or subcoriaceous; receptacle alveolate, denticulate. F l o r e t s 5 - 1 1 ( - 1 2 ) , hermaphroditic or, by abortion o f the androecium, pistillate; corolla actinomorphic, white, glabrous, limb deeply 5lobed, lobes linear, revolute, marginal and medial veins prominent; anthers with the base sagittate, caudate, terminal appendage often lanceolate, endothecial tissue with cells oblong-hexagonal, nodes polarized, pollen spheric, tricolpate, echinate, filament collar cylindrical, filament sligthly thickened; style base dilated, style branches linear, contorted, adaxially complanate, stigmatic areas marginal, continuous, thickened, confluent, abaxially papillate-pilose, distally sligthly attenuate, apex more or less thickened, acute or ob-

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CORREA: ASTERACEAE, SENECIONEAE-TUSSILAGININAE

tuse, conical, central papillae longer, stylopodium denticulate. A c h e n e s h o m o m o r phic, oblong, glabrous, 10-veined, 5 - 1 0 costate, attenuate towards the base, apex

159

slightly dilated; pappus bristles biseriate, persistent, scabrous, fused at base, stramineous. C h r o m o s o m e number: 2n = 80 (Nordenstam, 1977).

Key to species of Paragynoxys Capitula o f 6-11(-12) florets; involucral phyllaries (5-)6-8; corolla tube usually longer than the corolla lobes. 2. Leaves extremely woolly-tomentose abaxially; capitula of 6-8 florets. 3. Leaves ovate or elliptic-ovate; indumentum o f thin, curled, entangled hairs; involucral phyllaries (5-)6 ........................................................................... P. n e o d e n d r o i d e s 3. Leaves oblong; i n d u m e n t u m o f thick, appressed hairs; involucral phyllaries 8 ............... ......................................................

P. s a n t u r b a n e n s i s

2. Leaves sparsely villous abaxially; capitula o f 10-11(-12) florets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P . u r i b e i Capitula o f 5(-6) florets; involucral phyllaries always 5; corolla tube usually shorter than the corolla lobes. 4. Leaves glabrescent or barely curled-pubescent abaxially. 5. Involucre minutely pubescent or glabrous. 6. Leaves elliptic, 20-30 cm long, base rounded or oblique, apex acute, deeply emarginate or obtuse; female florets are often produced through abortion of the anthers, staminodia filiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. venezuelae 6. Leaves obovate, 9-15 cm long, base cuneate, apex rounded; florets always hermaphroditic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. steyermarkii 5. Involucre tomentose or villous. 7. Leaves elliptic-oblong, base cuneate, margin entire; capitula pedicellate, pedicels ca. 8 m m long; involucre tomentose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. angosturae 7. Leaves oblanceolate, base cordate or auriculate, margin subrepand-dentate; capitula subsessile, pedicels ca. 3 m m long; involucre villous . . . . . . P. c u a t r e c a s a s i i 4. Leaves tomentose to woolly abaxially (soft to touch). 8. Petioles tuberculate-verrucose. 9. Leaves coriaceous, densely woolly abaxially; petioles 8-16 m m thick, rigid. 10. Leaves subovate-elliptic, margin slightly undulate; involucre glabrous; florets always hermaphroditic ...................... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .P . u n d a t i f o l i a 10. Leaves ovate, margin flat; involucre woolly-tomentose; female florets are often produced through abortion of the anthers, staminodia filiform. P. meridana 9. Leaves papery, sparsely woolly-tomentose abaxially; petioles 4 - 5 mm thick, flexuous. ....................................................................................................................................... P. magnifolia

8.

Petioles smooth, not tuberculate-verrucose. 11. Leaves 15-25 cm long, base rounded-truncate or obtuse, petioles ca. 4 cm long; capitulescences ca. 40 cm long . . . . . . P. c o r e i 11. Leaves 45-65 cm long, base oblique, deeply cordate or sub-cordate, petioles ca. 10 cm long; capitulescences ca. 60 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P . m a r t i n g r a n t i i

PARAGYNOXYS ANGOSTURAE trec.

(Cuatrec.) Cua-

Paragynoxys angosturae (Cuatrec.) Cuatrec., Brittonia 8: 154. 1955. a n g o s t u r a e Cuatrec., Feddes Repert. Spec. Nov. Regni Veg. 55: 132. 1953. TYPE: COLOMBIA. Antioquia: Angostura (just outside town), in ravine filled with huge granite boulders, 2000 m, 11 Mar 1944, F . R . F o s b e r g 2 1 6 0 3 (HOLOTYPE: US; ISOTYPE: US).

Senecio

Trees to 7 m, terminal branches densely tomentose. Leaves with petioles of 2-15 cm long, tomentose, blade obovate-elliptic-ob-

long, 10-45 • 4-21.5 cm, coriaceous, base cuneate, apex subacute, rounded or obtuse, abaxial surface tomentose, adaxial surface glabrous, reticulate veined with 9 - 1 0 secondary veins on each side of midrib, margins entire, slightly revolute. Capitulescences thyrsoid-paniculate. Capitula homogamous, discoid; involucre cylindrical, phyllaries 5, weakly biseriate, subcoriaceous, outer phyllaries oblong, 6-8 • 1.5-2.5 mm, tomentose, apex acute, inner phyllaries elliptic-oblong, 6-7.5 X 2-3 mm, medially tomentose, apex obtuse, margins scarious. Florets 5, hermaphroditic, corolla 7.5-9.5 mm

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BRITTONIA

long, tube 3 . 5 - 4 m m long, limb 4 - 5 . 5 m m , lobes linear, ca. 5 m m long, apex acute; anthers ca. 3 m m long, base sagittate, caudate, terminal appendage lanceolate, slightly acute; style branches with the apex slightly obtuse, dorsally pilose. A c h e n e s oblong, ca. 3 m m long; pappus 8 - 1 0 m m long. Distribution a n d e c o l o g y . - - T h i s species occurs in the Depts. Antioquia and C h o c 6 , Colombia, at 1 7 5 0 - 2 5 0 0 m, in the m o n t a n e rain forest (Fig. 1). C o m m o n n a m e . - - P a t a gallina. E t y m o l o g y . - - T h i s species was n a m e d for the t o w n o f Angostura, Dept. Antioquia, Colombia. P a r a g y n o x y s a n g o s t u r a e is similar in capitular m o r p h o l o g y to P. cuatrecasasii; they can be easily separated by the leaves which are oblong-elliptic, with the base cuneate and the m a r g i n entire (vs. oblanceolate, with the base cordate or auriculate, and the margin subrepand-dentate). In addition, these two species are f r o m different localities: P. a n g o s t u r a e is f r o m northwestern Colombia, and P. cuatrecasasii is f r o m western Venezuela. Additional specimens examined: COLOMBIA. Antloquia: Amalfi, Vereda La Blanquita, 1750 m, 25 Jun 1997, Alzate et al. 270 (F); Concepci6n, Vereda Pel~iez-San Bartolom6, Finca E1 Cardal, 2030 m, 23 Jul 1997, Cardona et al. 266 (F). Choc6: San Jos6 del Palmar, Cordillera de San Miguel, vertiente oriental, trocha entre mesa de Cerro del Torr~i y filo de cumbre del Cerro Panam{t, 2450-2500 m, 25 Aug 1988, Silverstone-Sopkin et al. 4760 (F). PARAGYNOXYS COREI (Cuatrec.) Cuatrec. corei (Cuatrec.) Cuatrec., Brittonia 8: 154, fig. I I - L . 1955.

Paragynoxys

Senecio corei Cuatrec. Feddes Repert. Spec. Nov.

Regni Veg. 55: 136-137. 1953. TYPE: COLOMBIA. Antioquia: Alto E10so, N. of Yarumal, 2320 m, E. L. Core 624 (HOLOTYPE:F; ISOTYPE: US). Trees, to 5 m; terminal branches thick, densely w o o l l y - t o m e n t o s e . L e a v e s with petioles ca. 5 c m long, tomentose, thick, rigid, blade oblong-elliptic or elliptic, 1 5 - 2 5 x 8 12 cm, coriaceous, base rounded-truncate or obtuse, apex rounded, abaxial surface w o o l ly-tomentose, adaxial surface slightly scabrous, reticulate veined with 7 - 1 0 second-

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ary veins in each side o f the midrib, margins unequal but entire, n a r r o w l y revolute. C a p i t u l e s c e n c e s thyrsoid-paniculate. Capitula h o m o g a m o u s , discoid; involucre subcylindrical; phyllaries 5, w e a k l y biseriate, subcoriaceous, outer phyllaries oblong, 6 - 8 • 2 - 2 . 2 m m , densely tomentose, apex acute, inner phyllaries o b l o n g - elliptic, 6 - 8 • 2 . 5 - 4 m m , medially tomentose, apex obtuse, m a r g i n s scarious. Florets 5, h e r m a p h roditic, corolla 8 - 1 0 m m long, tube 3 - 4 . 5 m m long, limb 5 - 5 . 5 m m long, lobes linear, ca. 5 m m long, apex acute; anthers ca. 3.5 m m long, base sagittate, caudate, terminal a p p e n d a g e lanceolate, acute; style b r a n c h e s with the apex slightly acute, dorsally papillose-pilose. A c h e n e s oblong, ca. 3.5 m m ; pappus 8 - 1 0 m m long. Distribution a n d e c o l o g y . - - T h i s species occurs in Dept. Antioquia, C o l o m b i a , at 1 4 9 0 - 2 6 5 0 m, in the m o n t a n e rain forest (Fig. 2). Etymology.--Named by Cuatrecasas (1953) for its collector, Earl L. C o r e ( 1 9 0 2 1984), w h o m a d e significant collections f r o m C o l o m b i a in 1 9 4 3 - 1 9 4 4 . P a r a g y n o x y s corei resembles P. martingrantii; P. corei has leaves 1 5 - 2 5 c m long, with r o u n d e d - t r u n c a t e or obtuse base (vs. leaves 4 5 - 6 5 c m long, with oblique or cordate base); in P. corei, the capitulescences are ca. 40 c m long, with robust b r a n c h e s and pedicels (vs. capitulescences ca. 60 c m long, with thin and slender branches and pedicels). Additional specimens examined: COLOMBIA. Antioquia: San Vicente, Aug 1952, Brother Daniel 4376 (COL, US); Amalfi, Vereda Peldar, Mina La Viborita, 1490-1600 m, 3 Oct 1992, Fonnegra et al. 4523 (COL, US); Santa Rosa de Osos, Vereda Minavieja, 10 road-kin to Carolina, 2500 m, 26 Feb 2001, Marrugo 79 (COL); 22 road-km beyond Santa Rosa de Osos on road to Yarumal, 2650 m, 26 Nov 1988, McPherson & Rolddn 13290 (MO, NY). PARAGYNOXYS CUATRECASASIIRufz-Terfin & L6pez-Figueiras. cuatrecasasii Rufz-Ter~in & L6pez-Figueiras, Revista Fac. Farm. Univ. A n d e s 14: 14. 1974. TYPE: V E N E Z U E L A . M6rida: Dist. Rangel, c u e n c a del Rfo A r a c a y , afluente del Santo D o -

Paragynoxys

2003]

CORREA: ASTERACEAE, SENECIONEAE-TUSSILAGININAE

mingo, al norte de la poblaci6n de Las Piedras, a orillas del camino hacia Giftgay, 2 5 5 0 - 2 7 0 0 m, 16 Dec 1972, L. R u f z - T e r d n et al. 8 2 5 8 (HOLOTYPE: M E R E n.v.; ISOTYPE: US). Trees, to 18 m. L e a v e s with petioles 1.57 cm long, densely tomentose; blade oblanceolate, 11.5-53 • 3.5-23 cm, coriaceous, base subcordate or auriculate, apex acute, abaxial surface glabrous except the villose reticule, adaxial surface glabrescent, reticulate veined with 18-22 secondary veins on each side o f the midrib, margins subrepanddentate, teeth callous-tftangular. Capitulescences paniculate. Capitula homogamous, discoid; involucre turbinate; phyllaries 5, uniseriate, coriaceous, elliptic or oblanceolate, 6 - 7 x 2.5-3 mm, villose, apex acute. Florets 5, hermaphroditic, corolla 9 - 1 2 mm long, tube ca. 4.5 m m long, limb ca. 5.5 m m long, lobes oblong or subelliptic-oblong, ca. 4 m m long, apex acute; anthers ca. 2 m m long, base sagittate, caudate, terminal appendage lanceolate, acute; style branches with the apex ovate-deltoid, dorsally puberulent-papillose. A c h e n e s turbinate, ca. 2 mm; pappus 5-7.5 m m long. Distribution a n d e c o l o g y . - - T h i s species occurs in the states of M f r i d a and Trujillo, Venezuela, at 2 4 0 0 - 2 7 0 0 m, in the montane rain forest (Fig. 1). Etymology. This species was named by Rufz-Terfin and L6pez-Figueiras (1974) in honor of Jos6 Cuatrecasas (1903-1996), one of the most important botanists involved with the flora o f northern South America. P a r a g y n o x y s c u a t r e c a s a s i i differs from the other species by its oblanceolate leaves with cordate or auriculate bases and subrepand-dentate margins.

Additional specimens examined: VENEZUELA. M~rida: Las Piedras, hoya del Rfo de Pueblo Llano, 2600 m, 24 Oct 1969, Cuatrecasas et aL 28161 (F); Sierra Nevada de Santo Domingo, Loma de E1 Baho, abajo de Valencia, 2900 m, l Mar 1973, Cuatrecasas et aL 28621 (US). Trujillo: Departamento de Bocon6, Parque Nacional Guaramacal, 2400 m, 13 Dec 1995, N. Cuello et al. 1217 (NY); Depart. Bocon6, Parque Nacional Guaramacal, road from Bocon6 to Guaramal, SE of Bocon6, ca. 15 km from the post of the park guards, 3 Nov 1998, Dorr et al. 8476 (NY).

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PARAGYNOXYS MAGNIFOLIACuatrec. P a r a g y n o x y s m a g n i f o l i a Cuatrec., Brittonia

8(2): 154, 156. 1955. TYPE: V E N E Z U E LA. Mfrida: Culata, 2135 m, May 1847, N. F u n c k & L. J. S c h l i m 1522 (HOLOTYPE: R n.v.; ISOTYPE" G). Trees, to 10 m. L e a v e s with petioles 1 0 20 cm long, densely tomentose; blade widely elliptic-ovate, obovate-oblong or ovateoblong, 2 5 - 7 0 X 1 1 - 4 6 cm, papery, base deeply cordate, apex rounded or obtuse, abaxial surface tomentose, adaxial surface grabrous except for the tomentose midrib, reticulate veined with 7 - 8 secondary veins in each side of the midrib, margins almost entire with minute callous teeth. Capitulescences thyrsoid-paniculate. Capitula homogamous, discoid, involucre cylindrical, phyllaries 5, uniseriate, subcoriaceous, oblong, 6 - 7 • 2.2-3.2 mm, glabrescent, apex obtuse or subacute, ciliate. Florets 5, hermaphroditic, corolla ca. 9 mm long, tube 3 . 5 - 4 m m long, limb ca. 5 mm long, lobes linear, ca. 4.5 m m long, apex acute; anthers ca. 3 m m long, base sagittate, caudate, terminal appendage lanceolate, acute; style branches w i t h the apex acute, dorsally pilose. A c h e n e s oblong, ca. 3 m m long; pappus, 8 m m long. Distribution a n d e c o l o g y . ~ h i s species is found in the states of Mfrida and T4chira, Venezuela, at 2 1 3 5 - 2 9 9 0 m, in the montane rain forest and subp4ramo (Fig. 1). Common name.--Cafiaflote.

I am in agreement with Cuatrecasas (1955) when he stated that P a r a g y n o x y s m a g n i f o l i a is most similar to P. meridana; however, it differs by having papery leaves that are only sparsely tomentose abaxially (vs. coriaceous and densely woolly-tomentose) and by the thin, and flexuous petioles (vs. thick and rigid). The type of capitulum is also important: h o m o g a m o u s in P. magnifolia (vs. heterogamous). Additional specimens examined: VENEZUELA. La Mucuy, 2400 m, Sep 1956, Aristeguieta, 2636 (NY). T~ichira-M~rida: entre Portachuelo de la Grita y Pregnnero, 2940-2990 m, 28 May 1980, Cuatrecasas et al. 29004 (US). M~rida:

162 PARAGYNOXYS MARTINGRANTII

BRITTONIA

(Cuatrec.)

Cuatrec. P a r a g y n o x y s m a r t i n g r a n t i i (Cuatrec.) C u a trec., B r i t t o n i a 8: 156. 1955. Senecio martingrantii Cuatrec., Feddes Repert. Spec. Nov. Regni Veg. 55: 139. 1953. TYPE: COLOMBIA. Magdalena: Sierra de Perij~, Casacar~ Valley, 23 km east of Codazzi, 2 km from the Venezuelan border, 2450 m, M. L. Grant 10949 (HOLOTYPE: F; ISOTYPES: NY, US). Trees, to 14 m. Leaves with petioles ca. 10 c m long, d e n s e l y woolly-tomentose; blade oblong-elliptic, 4 5 - 6 5 • 1 5 - 2 6 cm, coriaceous, base oblique, deeply cordate or r o u n d ed-cordate, apex rounded, abaxial surface woolly-tomentose, adaxial surface glabescent except for the midrib tomentose, reticulate veined, secondary veins 1 0 - 1 6 in each side of the midrib, m a r g i n s entire. Capitulescences thyrsoid-paniculate. Capitula h o m o g a r n o u s , discoid; involucre cylindrical; phyllaries 5, weakly biseriate, subcoriaceous, outer pbyllades oblong, ca. 6 X 2 m m , curled-tomentulose or glabrate, the apex subobtuse, i n n e r phyllaries oblong-elliptic, ca. 6 • 3 m m , curled-tomentulose or glabrate medially, apex obtuse, m a r g i n s scarious. Florets 5, h e r m a p h roditic, corolla 7 - 9 m m long, tube 2 . 5 - 4 m m long, limb ca. 4.5 m m long, lobes linear, ca. 4 m m long, apex acute; anthers ca. 3 m m long, base sagittate, candate, terminal appendage lanceolate, acute; style branches with the apex acute, dorsally papillose-hirsutulous. A c h e n e s oblong, ca. 3 m m long; pappus ca. 8 m m long. D i s t r i b u t i o n a n d e c o l o g y . - - T h i s species occurs in Depts. Cesar, M a g d a l e n a a n d Norte de Santander, C o l o m b i a , a n d the states o f M6rida, T~ichira, a n d Zulia, V e n e z u e l a , at 2 4 5 0 - 3 9 6 0 m, i n the m o n t a n e rain forest, subpS_ramo, a n d pS_ramo (Fig. 1). Common name.--Colmenero. E t y m o l o g y . - - T h i s p l a n t was n a m e d b y C u a t r e c a s a s (1953) after its collector, Martin L. G r a n t ( 1 9 0 7 - 1 9 6 8 ) w h o was a m e m ber o f the c o l l e c t i n g t e a m s e a r c h i n g for alternate sources o f q u i n i n e d u r i n g the W o r l d W a r II. See P a r a g y n o x y s corei for a d i s c u s s i o n o n h o w P. m a r t i n g r a n t i i differs.

Additional specimens examined: COLOMBIA. Ce-

[VOL. 55

sar: Pgramo de Sabana Rubia, 3250 m, 22 Jun 1987, Cuadros 3721 (US). Magdalena: Sierra Nevada de Santa Marta, Cuchilla de San Lorenzo, en cercanfas a la Lagunita, por la via que conduce a Cerro Kennedy, 2475 m, Barbosa & Rangel 178 (COL); Sierra de Perija, E of Manaure, Quebrada de Floridablanca, 27002800 m, 12 Nov 1959, Cuatrecasas & Romero-Castageda 25249 (COL, F, US); Cerro Kennedy, near top of highest peak in N massif of Sierra Nevada de Santa Marta, 2600 m, 26 Aug 1986, Gentry & Cuadros 55536 (US); Sierra Nevada de Santa Marta, 2620 m, 14 Jan 1989, Gentry" & Cuadros 64669 (US); San Lorenzo-Sierra Nevada de Santa Marta, camino a la antena de la Televisora, 2100 m, 14 Jul 1969, N. de L6pez 290 (COL); Santa Marta, San Lorenzo, 2240 m, 9 Sep 1969, Lozano 1031 (COL). Norte de Santander: Regi6n de Ocafia, entre La Marfa y Jurisdicciones, 2600 m, 25 Sep 1969, Cuatrecasas & Rodrlguez 27923 (COL, US). Norte de Santander-Cesar: Lfnea divisoria entre Deptos., 20 kmal sur de Abrego, Las Jurisdicciones (Cerro de Oroque), 3700-3960 m, 19 May 1969, Garcfa Barriga & Jaramillo Mej& 19713 (COL, US); lfnea divisoria entre Deptos. de Santander del Norte y Cesar, 20 km al sur de Abrego, Las Jurisdicciones (Cerro Oroque), 3700 m, 24 Jul 1974, Garcia Barriga & Jaramillo Mejs 20738 (US). VENEZUELA. M6rida: Pgramo de las Coloradas, entre La Capilla y El Aserrucho (Santa Cruz de MoraE1 Molino), 2700-2750 m, 27 May 1980, Cuatrecasas 28996 (NY). T~iehira: Depto. de Jaugueri, 5 km E Zumbador on road to Queniquea, 2820 m, 31 Mar 1979, Berry 3416 (NY, US). Zulia: Distrito de Perij~, Sierra de Perija, Serrania de Valledupar, on international boundary, headwaters of Rio Guasare, 27003300 m, 10 Jul 1974, Tillett 747-1050 (NY, US).

PARAGYNOXYS MERIDANA (Cuatrec.) Cuatrec. P a r a g y n o x y s m e r i d a n a (Cuatrec.) Cuatrec., B r i t t o n i a 8: 156. 1955. Gynoxys verrucosa Badillo, Bol. Soc. Venez. Ci. Nat. 10: 312. 1947. Non G. verrucosa Wedd. Senecio steyermarkii Cuatrec., Fieldiana, Bot. 27(1): 32. 1950. Non S. steyermarkii Greenm. Senecio meridanus Cuatrec., Fieldiana, Bot. 27(2): 38. 1951. TYPE: VENEZUELA. M6rida: P~amo de Pozo Negro between San Jos6 and Beguilla, 2590-3220 m, 3 May 1944, J. A. Steyermark 56268 (HOLOTYPE:NY; ISOTYPES:E US). Trees, up to 8 m. L e a v e s with petioles 1 0 12 c m l o n g , m o d e r a t e l y v e r r u c o s e a n d d e n s e l y w o o l l y - t o m e n t o s e , blade ovate, 36-50 x 2 7 - 3 6 cm, coriaceous, base d e e p l y cordate, apex obtuse, l o w e r surface d e n s e l y w o o l l y - t o m e n t o s e , verrucose at the midrib, the l o w e r surface glabrescent except for the strongly t o m e n t o s e midrib, reticulate veined, 8 - 1 0 s e c o n d a r y veins in each side o f the midrib, m a r g i n s entire. Capitulescences pa-

2003]

CORREA: ASTERACEAE, S E N E C I O N E A E - T U S S I L A G I N I N A E

niculate. Capitula often heterogamous or rarely homogamous, discoid; involucre cylindrical; phyllaries 5, uniseriate, membranous, elliptic-linear, 7.5 • 2.5-3 mm, lightly tomentose-woolly, apex attenuate, slightly acute, ciliolate. Florets 5, hermaphroditic or pistillate, corolla 7-10 mm long, tube 3 - 4 mm long, limb ca. 6 m m long, lobes linear, ca. 5.5 mm long, apex acute; anthers ca. 3 mm long, base slightly auriculate, terminal a p p e n d a g e lanceolate, acute, s o m e t i m e s abortive but with persistent filaments; style branches with the apex obtuse, dorsally papillose-pilosulous. Achenes fusiform, ca. 4.5 m m long; pappus 9 m m long. Distribution and ecology.~This species is found in the states of Mrrida and Tfichira, Venezuela, at 2 4 0 0 - 3 2 2 0 m, in the montane rain forest, subpfiramo, and p~iramo (Fig. 2).

Common name.--Hojiancho. Etymology. This species was named for the state of Mrrida, Venezuela. Paragynoxys meridana is similar to P. undatifolia, but it can be distinguished by its ovate leaves, with flat margin (vs. subovate-elliptic and undulate), woolly-tomentose involucre (vs. glabrous), and presence of female florets, developed through abortion of the anthers (vs. hemaphroditic). Additional specimens examined: VENEZUELA. M~rida: Pgramo de San Josr, Zanjdn del Cupfs, 3100 m, 1 Feb 1973, Cuatrecasas et al. 28459 (US); Pfiramo del Cupfs (San Josd-Beguilla), 3250 m, 22 Mar 1967, L6pez Figueiras 8609 (US); P~amo de Cupis a partir de La Piedra de Pirela, San Jos6 y Mucutny, 3200 m, 22 Mar 1967, Ruiz Terdn & Lrpez Figueiras 3847 (NY). Tfichira: El Portachuelo, hacia Queniquea, 2400 m, Sep 1956, Aristeguieta 2553 (E NY); Pgramo del Zumbador, 3100 m, 1 Oct 1969, Cuatrecasas et al. 27995 (US); Dstr. Jauregui, 5 km E of Zumbador on road to Queniquea, 2820 m, 31 Mar 1979, Berry 3416 (US).

PARAGYNOXYS NEODENDROIDES

(Cuatrec.)

Cuatrec.

Paragynoxys neodendroides (Cuatrec.) Cuatrec., Brittonia 8: 156, figs. 1A-D, 4. 1955. Senecio neodendroides Cuatrec., Notas FI. Colomb.

6: 19. 1944. TYPE: COLOMBIA. Santander: Cordillera Oriental, P~iramo de la Rusia, vertiente noroeste, 3300-3500 m, 4 Aug 1940, J. Cuatrecasas 10435 (HOLOTYPE: COL; ISOTYPE: F).

Trees, to 15 m. Leaves with petioles 11-

163

13 cm long, d e n s e l y t o m e n t o s e ; blade ovate-elliptic, 3 0 - 4 0 • 20-21 cm, coriaceous, base cordate, apex attenuate, obtuse, lower surface densely woolly-tomentose, upper surface glabrous except for the woolly-tomentose midrib, reticulate veined, 1517 secondary veins in each side of the midrib, margins entire. Capitulescences corymbose-paniculate. Capitula homogamous, discoid; involucre cylindric-campanulate; p h y l l a r i e s 5 - 6 , uniseriate, c o r i a c e o u s , ovate-oblong, ca. 8 • 3 mm, densely tomentose, apex acute. Florets 7-9, hermaphroditic, corolla ca. 12 mm long, tube ca. 6 mm long, limb ca. 5.5 m m long, lobes linear, ca. 5 mm long, apex acute; anthers ca. 4 ram, base appendiculate, terminal appendage lanceolate, acute; style branches with the apex obtuse, dorsally penicillate-appendiculate. Achenes oblong, ca. 3 m m long; pappus, 9 mm long. Distribution and ecology.--This species occurs in depts. Boyacfi and Santander, Colombia, at 3 2 0 0 - 3 5 0 0 m, in the pfiramo (Fig. 1).

Common name.--Gamuzo. Etymology.--The name given this species by Cuatrecasas (1944) is from the Greek neo- (new-) and dendroideus (tree-like), meaning a young tree. Paragynoxys neodendroides is similar to P. santurbanensis, but they can easily be separated by the leaf characters: ovate (vs. oblong); thin curled, intricate hairs (vs. thick and appresed); and the number o f phyllaries: (5-)6 (vs. 8). Additional specimens examined: COLOMBIA. Boyaefi: Cordillera Oriental, Arcabuco, pfiramos vecinos a la poblaci6n, 3200 m, 6 Jun 1969, Camargo & Huertas 6802 (COL); Cordillera Oriental, Pfiramo de la Rusia, Duitama, vereda E1 Carmen, Km 28, carretera a Charal~i, 3300 m, 30 Sep 1973, D{az Piedrahita 355 (COL); Duitama, E1 Huerto, 3000, 5 Mar 1964, Espin a l & Montenegro 1576 (COL); alrededores de El Rosal, 3100 m, 1 Jun 1958, Romero-Castafieda 6804 (COL). Santander: Onzaga, alto entre Onzaga y Soafit, 3350 in, 4 Aug 1958, Jaramillo Mejfa et al. 865 (COL, US); Onzaga, vereda Chaguacfi, flanco oriental del pfiramo de Guantiva, 3370 m, 6 Aug 1958, Jaramillo Mejla et al. 916 (COL); Onzaga, vereda Chaguac~, monte bordeando el Rio Chaguac~i, 3040 m, 31 Oct 1987, Orozco et al. 1848 (COL); Onzaga, vereda Chaguacfi, 3200 m, 3 Apr 1976, Torres 660 (COL). PARAGYNOXYS

Cuatrec.

SANTURBANENSIS

(Cuatrec.)

164

BRITTONIA

Paragynoxys santurbanensis ( C u a t r e c . ) Cuatrec., Brittonia 8: 156. 1955. Senecio santurbanensis Cuatrec., Feddes Repert. Spec.

Nov. Regni Veg. 55: 145. 1953. TYPE: COLOMBIA. Santander: P~amo de Santmbfin, vert. W., bajando hacia Bucaramanga, 3100 m, J. Cuatrecasas & H. Garcs Barriga 10326 (ItOLOTYPE: F).

Trees, to 10 m. Leaves with petioles 8-9 cm long, densely tomentose; blade oblongelliptic, 2 5 - 3 0 • 12-14 cm, coriaceous, base cordate, apex obtuse, lower surface densely tomentose, upper surface glabrous except for the tomentose midrib, reticulate veined, ca. 12 secondary veins on each side o f the midrib, the margins entire. Capitulescences thyrsoid-paniculate. Capitula homogamous, discoid; involucre cylindrical; phyllaries 8, weakly biseriate, subcoriaceous, outer phyllaries lanceolate, ca. 7.5 • 2 mm, densely tomentose, apex subacute, inner phyllaries ovate, ca. 7 • 3 mm, tomentose medially, apex subacute, margins scarious. Florets 8, hermaphroditic, corolla ca. 10 m m long, tube ca. 7 m m long, limb ca. 3.3 m m long, lobes ca. 3 m m long, linear, apex acute; anthers ca. 4 m m long, base slightly caudate, terminal appendage apiculate, obtuse; style branches with the apex obtuse, dorsally pilose. Achenes ca. 3.5 m m long; pappus 10 m m long. Distribution and ecology.--This species is only known from the type locality in the Pgtramo de Santurb~n, Dept. Santander, Colombia, at ca. 3000 m (Fig. 1). Etymology.--This species is named for the Pfiramo de Santurbfin, Colombia. See Paragynoxys neodendroides for a discussion on how it differs from P. santurbanensis. Additional specimen examined: COLOMBIA. Santander: Cordillera Oriental, P~ramo de Santurb~n, entre Bucaramanga y Berlin, 3175 m, 3 Jan 1960, Barclay & Juajibioy 10418 (US).

PARAGYNOXYS STEYERMARKII Cuatrec.

Paragynoxys steyermarkii Cuatrec., Phytologia 40: 34. 1978. TYPE: V E N E Z U E L A . Tfichira: Between Las Copas and Alto de Fila de Tierra Negra at the ridge dividing headwaters of Rivers Quinimarf, Riofrfo, Uribante, and Talco (Oir~), 2 8 7 0 - 2 8 8 0

[VOL. 55

m, 16 Jan 1968, J. A. Steyermark et al. 101014 (HOLOTYPE: U S ; ISOTYPE" VEN, n.v.).

Trees, to 3 m. Leaves with petioles 1230 m m long, tomentulose; blade obovateelliptic, 9 - 1 5 • 4 - 7 cm, coriaceous, base cuneate, apex obtuse, lower surface tomentose, u p p e r s u r f a c e glabrous, r e t i c u l a t e veined, 9 - 1 1 secondary veins on each side of the midrib, the margins sinuate or subentire, but with minute callous teeth. Capitulescences paniculate-corymbiform. Capitula homogamous, discoid; involucre cylindrical; phylaries 5, weakly biseriate, subcoriaceous, outer phyllaries oblong-elliptic, 9.5 x 2 mm, glabrous, apex slightly attenuate, subobtuse, slightly barbellate-ciliolate, inner phyllaries elliptic, 8.5 • 3 mm, glabrous, apex obtuse, margins scarious. Florets 5-6, hermaphroditic, corolla 11-12.5 m m long, tube ca. 6 m m long, limb ca. 5.5 m m long, deeply 5-lobed, lobes linear, ca. 5 m m long, apex acute; anthers ca, 3.5 m m long, base sagittate, deeply caudate, terminal a p p e n d a g e o b l o n g - a t t e n u a t e , acute; style branches with the apex somewhat attenuate, subacute, dorsally papillose-pilosulose. Achenes oblong, ca. 3 m m long; pappus 8 - 9 m m long. Distribution and ecology.~This species occurs in the state of Tfichira, Venezuela, at 2 8 7 0 - 2 8 8 0 m, in the montane rain forest (Fig. 2). Etymology.~This species was named after Julian A. Steyermark (1909-1988), an important contributor to Venezuelean botany. Paragynoxys steyermarkii resembles P. venezuelae from which it differs by its leaves 9 - 1 5 cm long, obovate, cuneate at base and rounded at apex (vs. 2 0 - 3 0 cm long, elliptic, rounded or oblique at base, and acute, deeply emarginate or obtuse at apex); florets always hermaphroditic in P. steyermarkii (vs. often female). PARAGYNOXYS UNDATIFOLIA Cuatrec.

Paragynoxys undatifolia Cuatrec., Proc. Biol. Soc. Wash. 74.15. 1961. TYPE: COLOMBIA. Magdalena: Sierra Nevada de Santa Marta, SE slope, H o y a del Rio Do-

2003]

CORREA: ASTERACEAE, SENECIONEAE-TUSSILAGININAE

nachu/, b e l o w S a b a n i t a D i r i c u n e , 3 2 0 0 m, 29 S e p 1959, J. C u a t r e c a s a s & R. Rom e r o - C a s t a t i e d a 2 4 4 8 5 (HOLOTYPE: U S ; ISOTYPE: C O L ) . Trees, 6 - 1 2 m. L e a v e s w i t h p e t i o l e s ca. 20 c m l o n g , s l i g h t l y v e r r u c u l o s e , d e n s e l y tomentose; blade subovate-elliptic, 50-70 • 2 8 - 4 0 cm, b a s e d e e p l y c o r d a t e , a p e x attenuate, obtuse, lower surface densely woolly-tomentose, upper surface glabrate e x c e p t for the d e n s e l y w o o l l y - t o m e n t o s e midrib, reticulate veined, 14-15 secondary v e i n s o n e a c h s i d e o f the m i d r i b , the m a r gins entire b u t s l i g h t l y a n d i r r e g u l a r l y undulate. C a p i t u l e s c e n c e s t h y r s o i d - p a n i c u l a t e . Capitula h o m o g a m o u s , d i s c o i d ; i n v o l u c r e c y l i n d r i c a l , p h y l l a r i e s 5, w e a k l y b i s e r i a t e , p a p e r y , o u t e r p h y l l a r i e s o b l o n g , 8.5 • ca. 2 ram, g l a b r o u s , a p e x s u b a c u t e , ciliate, inner phyllaries elliptic, 8 • 3-3.5 mm, glab r o u s , a p e x o b t u s e , m a r g i n s scarious, e r o s e ciliate. Florets 5, h e r m a p h r o d i t i c , c o r o l l a ca. 11 m m long, t u b e 5 . 5 - 6 m m l o n g , l i m b ca. 5.5 m m l o n g , l o b e s linear, ca. 5 m m l o n g , a p e x acute; a n t h e r s ca. 3 m m l o n g , b a s e s l i g h t l y sagittate, a u r i c u l a t e , t e r m i n a l a p p e n d a g e l a n c e o l a t e , acute; style b r a n c h e s w i t h the a p e x acute, d o r s a l l y p e n i c i l l a t e . A c h e n e s subtriangular, ca. 3 m m l o n g ; p a p p u s ca. 8 m m long. Distribution a n d e c o l o g y . - - T h i s s p e c i e s is o n l y k n o w n f r o m the S i e r r a N e v a d a d e Santa Marta, Dept. Magdalena, Colombia, at 3 2 0 0 - 3 4 5 0 m, in the pfiramo. H o w e v e r , C u a t r e c a s a s c l a i m e d that he s a w this s p e cies as l o w as 2 5 0 0 m, in the m o n t a n e r a i n f o r e s t (Fig. 2). S e e P a r a g y n o x y s m e r i d a n a for a d i s c u s s i o n on h o w it d i f f e r s f r o m P. undatifolia.

Additional specimens examined: COLOMBIA. Magdalena: Sierra Nevada de Santa Marta, alrededores de cabeceras del Rio Sevilla, 3330 m, 27 Jan 1959, Barclay & Juaijibioy 6719 (COL); entre San Pedro y cabeceras del Rio Sevilla, 3200 m, 31 Jan 1959, Barclay & Juaijibioy 6777 (COL, US); alrededores de la cabecera del Rio Ancho, P~iramo de Macotama, 3490 m, 30 Jun 1959, Barclay 6990 (COL); divorcio de aguas entre Riofrfo y la quebrada del pfiramo, en el filo Alto de la Cumbre, 3350 m, 23 Sep 1977, Jaramillo et al. 5522 (COL); 1 km al NW de la quebrada de la Laguna Rio Fifo, en direcci6n al Pico Jos6 Hilario, 3400 m, 31 Jul 1972, Forero & Kirkbride 627 (COL, NY); 1.5 km W of Laguna Rio Frfo on camino from San Pedro de la Sierra via Pico Jos6 Hilario, 3450

165

m, 31 Jul 1972, Kirkbride & Forero 1805 (COL, NY); Regi6n de Santa Marta, flanco oeste de la Sierra Nevada, Cidnaga, 16 Jan 1954, Romero-Castageda 4562 (COL); Ci6naga, regi6n de Santa Marta, flanco oeste de la Sierra Nevada, 17 Jan 1954, Romero-Castatieda 4578 (COL); flanco occidental de la Sierra Nevada de Santa Marta, 3140 m, 29 Jan 1959, Romero-Casta~eda 7148 (COL, US). PARAGYNOXYS URIBEI C u a t r e c . P a r a g y n o x y s uribei C u a t r e c . , P h y t o l o g i a 40(1): 33. 1978. TYPE: C O L O M B I A . B o yac~i: A r c a b u c o , 4 k m N E o f t o w n , 2 7 0 0 m, 8 Jun 1966, L. Uribe Uribe 5633 (HOLOTYPE: U S ; 1SOTYPE: C O L ) . Paragynoxys pileolanata S. Dfaz, Caldasia 12: 379, fig. 1. 1979. TYPE: COLOMBIA. Santander: Mun. Onzaga, vereda Chaguac& finca de Oliverio Mesa, 2820 m, 29 Mar 1976, J. M. Torres, G. Lozano, & S. Dfaz 500 (HOLOTYPE:COL).

Trees, 3 - 4 m. L e a v e s w i t h p e t i o l e s 6 - 1 3 c m long, d e n s e l y t o m e n t o s e ; b l a d e o v a t e o r e l l i p t i c - o v a t e , 34 • 1 6 - 1 8 cm, c o r i a c e o u s , base broadly rounded, oblique or cuneate, a p e x s u b a c u t e , l o w e r s u r f a c e d e n s e l y tomentose, upper surface glabrous, except for the t o m e n t u l o s e m i d r i b , r e t i c u l a t e v e i n e d , 1 2 - 1 3 s e c o n d a r y v e i n s o n e a c h side o f the m i d r i b , m a r g i n s r e v o l u t e , entire o r r a r e l y w i t h m i n u t e c a l l o u s teeth. C a p i t u l e s c e n c e s s u b t h y r s o i d - p a n i c u l a t e . Capitula h o m o g a mous, discoid; involucre cylindrical, phyllaries 8, w e a k l y b i s e r i a t e , s u b c o r i a c e o u s , o u t e r p h y l l a r i e s o b l o n g , 7.5 • 2.5 m m , densely woolly-tomentose, apex subacute, inner phyllaries elliptic, 7-7.3 • 3 mm, med i a l l y t o m e n t o s e , m a r g i n s scarious. Florets 10-11(-12), hermaphroditic, corolla 8.5-9 m m long, t u b e 3 . 5 - 4 m m long, l i m b 4 , 5 - 5 m m long, l o b e s linear, 3 . 5 - 4 m m l o n g , f u s e d ca. 1 m m at the b a s e , a p e x s u b a c u t e ; a n t h e r s ca. 2.5 m m l o n g , b a s e sagittate, c a u date, t e r m i n a l a p p e n d a g e o b l o n g , s l i g h t l y o b t u s e ; style b r a n c h e s w i t h the a p e x subacute, d o r s a l l y p a p i l l o s e - p i l o s e . A c h e n e s o b l o n g , 2 . 5 - 2 . 8 m m ; p a p p u s 7 - 8 m m long. Distribution a n d e c o l o g y . - - T h i s s p e c i e s o c c u r s in D e p t s . B o y a c f i and Santander, C o l o m b i a , at 2 7 0 0 - 2 9 0 0 m, in the m o n t a n e rain f o r e s t (Fig. 2). C o m m o n n a m e s . - - G o r r e l a n a , O r e j o , Totumo. E t y m o l o g y . ~ T h i s s p e c i e s w a s n a m e d af-

166

BRITTONIA

ter L o r e n z o U r i b e U r i b e ( 1 9 0 0 - 1 9 8 0 ) , an i m p o r t a n t C o l o m b i a n collector. P a r a g y n o x y s u r i b e i differs f r o m the o t h e r spe ci es o f the g e n u s by h a v i n g the h i g h e s t n u m b e r o f florets p e r c a p i t u l u m ( 1 0 - 1 2 ) and a c o n s t a n t n u m b e r (8) o f p h y l l a r i e s . T h e c o r o l l a has its l o b e s f u s e d to ca. 1 m m a b o v e the base. P a r a g y n o x y s p i l e o l a n a t a is h e r e c o n s i d e r e d a s y n o n y m o f P. u r i b e i . W h e n P. p i l e o l a n a t a w a s o r i g i n a l l y d e s c r i b e d (Dfaz, 1979), t h ere w a s little m a t e r i a l o f P. u r i b e i a v a i l a b l e . T h e t y p e had d i s t i n c t i v e l e a f sha p e an d r e t i c u l a t i o n , c a p i t u l e s c e n c e s , ind u m e n t u m , an d capitula, N o w , w i t h m o r e m a t e r i a l f r o m t h e type l o c a l i t y o f P. p i l e o l a n a t a , it has b e c o m e e v i d e n t that the characters u s e d to separate P. p i l e o l a n a t a are not distinct. T h e r e f o r e , w e (Dr. D f a z a n d I) c o n c l u d e t h e y are the s a m e species. Additional specimens examined: COLOMBIA. Boyacsi: Duitama, corregimiento El Carmen, vereda Santa Helena, cerca al Rfo La Rusia, finca La Sierra, 2700 m, 6 Nov 1993, Betancur et aI. 4173 (COL); Arcabuco, La Cumbre, 2300 m, 8 Oct 1980, Lozano & Ds 3722 (COL); Km 49, alto de El Venado, via Duitama Charal~i, 29 Jul 1992, Neira & Ni~o 10 (COL); Arcabuco, Vereda Piedras Blancas, finca Las Delicias, 2600-2750 m, 11 May 1996, Rangel et al. 13143 (COL); Hacienda de la Rusia, carretera de Duitama a Charal& 2900 m, 12 Oct 1959, Uribe Uribe 3404 (COL, US); Carretera de Duitama a Charal~i, frente a la hacienda de La Rusia, 3000 m, 25 Apr 1965, Uribe Uribe 5227 (COL); Virolin, finca La Sierra, 2600 m, 29 Jan 1969, Villa 3 (COL). Boyaea-Santander: Cerro Berlin, entre Arcabuco y La Palma, 2780-2800 m, 28 Mar 1973, Cuatrecasas et al. 28675 (COL, US); lfmites entre Deptos. Santander y Boyaczi, corregimiento de Virolfn, finca La Sierra, 2500-2600 m, 18 May 1976, Lozano et al. 2631 (COL). Santander: Charal~i, vereda Santa Helena, predio La Sierra, margen izquierdo aguas abajo del Rfo La Rusia, Santuario de Fauna y Flora de Guanetfi, alto del Rio Fonce, 2480-2550 m, 8 Oct 1998, Cadena et al. 117 (COL); Charal~, Vereda Santa Helena, predio La Sierra, margen izquierdo aguas abajo del Rfo La Rusia, Santuario de Fauna y Flora de Guanet~i, alto del Rio Fonce, 2440 m, 16 Dec 1998, Cadena et al. 186 (COL); Charahl, vereda E1 Talado, Km 50-55 carretera Duitama Virolfn, 22502300 m, 6 Dec 1978, Dfaz Piedrahita 1608 (COL); Carretera Arcabuco La Palma, 3 Jul 1971, Guarfn & Villarreal 1054 (COL); Charal& Vereda Santa Helena, 2800 m, 12 Feb 1999, Stancik & Medina 2402 (COL).

PARAGYNOXYS VENEZUELAE trec.

(Badillo)

Cua-

[ V O L . 55

Paragynoxys venezuelae (Badillo) Cuatrec.,

B r i t t o n i a 8: 156, Fig. 2 M - Q .

1955.

Cacalia venezuelae Badillo, Bol. Soc. Venez. Ci.

Nat. 10:319. 1947. Senecio venezuelae (Badillo) Cuatrec., Fieldiana

Bot. 27(1): 31. 1950. TYPE: VENEZUELA. M6rida: Arriba de San Isidro Alto, 1880 m, 14 May 1944, J. A. Steyermark 56560 (HOLOTYPE:VEN, n.v.; ~SOTYPES:E NY, US). T r e e s , to 3 m. L e a v e s w i t h p e t i o l e s 5 - 7 c m long, d e n s e l y p u b e s c e n t ; b l a d e e l l i p t i c 20-30 • 10-15 cm, coriaceous, base r o u n d e d or c o r d a t e , a p e x o b t u s e to r o u n d e d , lower surface somewhat pubescent, upper s u r f ace g l a b r o u s e x c e p t f o r the w o o l l y m i d rib, r e t i c u l a t e v e i n e d , 5 - 8 s e c o n d a r y v e i n s on e a c h side o f the m i d r i b , m a r g i n s entire, revolute. Capitulescences paniculate-subc o r y m b o s e . C a p i t u l a o f t en h e t e r o g a m o u s or rarely h o m o g a m o u s , d i s c o i d ; i n v o l u c r e cy lindrical; p h y l l a r i e s 5, u n i ser i at e, c o r i a c e o u s , o v a t e - o b l o n g , ca. 4.5 • 3 m m , pub e r u l en t , a p e x obtuse, m a r g i n s scarious. F l o r e t s 5, h e r m a p h r o d i t i c or pistillate, c o rolla ca. 8.5 m m long, tube ca. 3 m m l o n g , l i m b ca. 5.5 m m long, l o b es linear, ca. 5.5 m m l o n g , a p e x acute; anthers 2 . 5 - 3 m m long, b a s e s l i g h t l y auriculate, t e r m i n a l app e n d a g e l a n c e o l a t e , acute, o f t en a b o r t i v e but with the filaments persistent; style b r a n c h e s w i t h the a p e x obtuse, d o r s a l l y papillose-pilose. Achenes oblong, 2-3 mm l on g ; p a p p u s 7 - 7 . 5 m m long. Distribution and ecology.--This species o c c u r s in the states o f M 6 r i d a and Trujillo, V e n e z u e l a , an d Dept. N o r t e d e Santander, C o l o m b i a , at 1 8 8 0 - 3 0 0 0 m, in the m o n t a n e rain forest, subpfiramo, and pfiramo (Fig. 2). S e e P a r a g y n o x y s s t e r y e r m a r k i i : f o r a disc u s s i o n on it d i f f er s f r o m P. v e n e z u e l a e .

Additional specimens examined: COLOMBIA. Norte de Santander: La Playa de Beldn, Parque Nacional Natural de los Estoraques, 1900 m, 17 May 2001, Galindo et al. 456 (COL)j La Playa de Bel6n, vereda Peritama, Area Natural Unica los Estoraques, 1650 m, 20 Feb 2002, R. Galindo et al. 684 (COL); Toledo a Saravena, Km 35, vereda Miralindo, abajo del parador Santa In6s, 1 Nov 1994, Orozco et al 2967 (COL); Cucutilla, vereda E1 Carrizal, sector Sisavita, 3020 m, 2 Mar 2002, A. Pietro et al. 1129 (COL). VENEZUELA. M6rida: Santa Cruz de Mora-Canagufi, Jul 1971, Aristeguieta 7850 (E NY). Trujillo: Dist. de Carache, Pfiramo Turmal, below Pico Jab6n, ca. 3000 m, 27 May 1988, Dorr et al. 5316 (NY); Carache, camino desde la ilia del P~iramo del Avi6n

2003]

CORREA: ASTERACEAE, SENECIONEAE-TUSSILAGININAE

(SE del P~iramo de Cend6) hacia Mesa Arriba, 21002900 m, 14 Feb 1999, Duno de Stefano & Riina 709 (US); entre La Pena y Agua de Obispo, 22-28 km de Carache, 2400-2500 m, 1 Mar 1971, Steyermark 104976 (E NY, US). EXCLUDED SPECIES

Paragynoxys

lopezii ( D i l l o n & Sagast.) Cuatrec., P h y t o l o g i a 69(5): 314. 1990.

Paracalia lopezii ( D i l l o n & Sag~ist.) A. C o rrea, c o m b . nov.

Gynoxys Iopezii Dillon & Sag~ist., Brittonia 40(2): 223, fig. 2. 1988. TYPE:PERU. A. Ldpez & A. Sagdstegui 8160 (HOLOTYPE:HUT, n.v.; ISOTYPES:E US). This species was o r i g i n a l l y d e s c r i b e d b y D i l l o n & Sag~istegui (1988) i n Gynoxys. Later, C u a t r e c a s a s (1990) transferred it to Paragynoxys b e c a u s e o f its floral structure a n d its alternate leaves. However, I c o n s i d e r that this species b e l o n g s to Paracalia, w h i c h shares with Paragynoxys the s u b c o r i a c e o u s i n v o l u c r a l bracts, the shape o f the corolla, the sagittate b a s e o f the anthers, a n d the shape o f the stylar tips, b u t differs b y its s h r u b y habit. T h e p l a n t d e s c r i b e d b y D i l l o n a n d Sagfistegui is a shrub a n d has the floral structure o f Paracalia.

167

Therefore, I p r o p o s e that this species b e transferred b a c k to Gynoxys.

Acknowledgments I w a n t to t h a n k the curators a n d institutions w h o p r o v i d e d l o a n s a n d / o r facilities for this study: C O L , E G, M O , NY, and US. I gratefully a c k n o w l e d g e D a v i d L e n t z (Director o f the G r a d u a t e Studies P r o g r a m at N Y ) a n d the A n d r e w W. M e l l o n F o u n d a t i o n for the e x c e l l e n t g r a n t that m a d e p o s s i b l e this research. F i n a l l y , I t h a n k T h o m a s A. Z a n o n i (NY), J i m L. L u t e y n (NY), M i c h a e l D i l l o n (F), Bertil R. N o r d e n s t a m (S), E n r i q u e Forero ( C O L ) , a n d S a n t i a g o Dfaz Piedrahita ( C O L ) for the v a l u a b l e s u g g e s t i o n s a n d corrections that greatly i m p r o v e d the paper.

Literature Cited Cuatrecasas, J. 1944. Compositae. Not. F1. Colomb.

6: 16-36. --.. 1951. Studies on Andean Compositae II. Fieldiana, Bot, 27(2): 1-74. 9 1953. Neue und bemerkenserte andine Compositen. Feddes Repert. Spec. Nov. Regni Veg. 55: 120-153. --.. 1955. A new genus and other novelties in Compositae. Brittonia 8: 151-163. --.. 1961. Studies on Andean Compositae V. Proc. Biol. Soc. Wash. 74: 15-18. Paragynoxys regis (H. R o b i n s o n & Cuatrec.) --.. 1978. Miscellaneous notes on neotropical floH. R o b i n s o n & Cuatrec., N o v o n 2(4): 415. ra. Phytologia 40: 33-35. 1992. Gynoxys regis H. R o b i n s o n & Cua--. 1984. Observations of the genus Gynoxys in Ecuador Senecioneae, Asteraceae). Phytologia 56: 370. trec., Phytologia 56(5): 370. 1984. --.. 1990. Micellaneous notes on neotropical flora, XIX. Combinations in Senecioneae, Compositae. This species was transferred b y its auPhytologia 69: 313-315. thors ( R o b i n s o n & C u a t r e c a s a s , 1984, Diaz Piedrahita, S. 1979. Aportes a la flora de Co1992) to Paragynoxys b e c a u s e o f " t h e lack lombia, estudios en Compuestas III. Caldasia 12: 379-382. o f rays i n the capitula, the b l u n t tips o f the style b r a n c h e s , a n d the l o n g corolla lobes 9 Dillon, M. & A. Sagfistegui. 1988. Additions to South American Senecioneae (Asteraceae). Brittonia 40: separated to the b a s e o f the throat." H o w 221-228. ever, w h e n I e x a m i n e d the type s p e c i m e n I Nordenstam, B. 1977. Senecioneae and Liabeae-systematic review. Pages 799-830. In: V. H. Heyf o u n d the corolla u n d i v i d e d to its base, wood, J. B. Harborne & B. L. Turner, editors. The w h i c h is t y p i c a l o f Gynoxys, and the d i s c o i d biology and chemistry of the Compositae. Academhead, a l t h o u g h u n c o m m o n , is also p r e s e n t ic Press, London. in other species o f that g e n u s (e.g., G. camRobinson, H. & J. Cuatrecasas. 1984. Observations pii Cuatrec., G. dielsiana D o m k e , G. induta of the genus Gynoxys in Ecuador (Senecioneae, Asteraceae). Phytologia 56: 370. Cuatrec., G. littlei Cuatrec., G. longifolia - - . 1992. Additions to Aequatorium and Gynoxys Wedd., G. soukupii Cuatrec., a n d G. sub(Asteraceae: Senecioneae) in Bolivia, Ecuador, and hirsuta Cuatrec.). I n fact, C u a t r e c a s a s later Peril. Novon 2: 411-416. r e c o g n i z e d his m i s t a k e w h e n he m a r k e d the Rulz-Terfin, L. & M. L6pez-Figueiras. 1974. Revista Fac. Farm. Univ. Andes 14: 14-20. s p e c i m e n as p r o b a b l y n o t Paragynoxys.

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BRITTONIA

[ V O L . 55

Appendix SPECIES NAMES In this list the a c c e p t e d n a m e s in the p r e s e n t r e v i s i o n are in b o l d a n d the s y n o nyms are in italic.

Cacalia venezuelae s e e : Paragynoxys venezuelae Gynoxys lopezii, s e e : Paracalia lopezii, Gynoxys regis Gynoxys regis, s e e : Paragynoxys regis Gynoxys verrucosa, s e e : Paragynoxys meridana Paracalia lopezii, s e e : Paragynoxys lopezii, Gnoxys lopezii Paragynoxys angosturae Paragynoxys corei Paragynoxys cuatrecasasii Paragynoxys lopezii, s e e : Paracalia lopezii Paragynoxys magnifolia Paragynoxys martingrantii Paragynoxys meridana Paragynoxys neodendroides

Paragynoxys pileolanata, s e e : Paragynoxys uribei Paragynoxys regis, s e e : Gynoxys regis Paragynoxys santurbanensis Paragynoxys steyermarkii Paragynoxys undatifolia Paragynoxys uribei Paragynoxys venezuelae Senecio angosturae, s e e : earagynoxys angosturae Senecio corei, s e e : Paragynoxys corei Senecio martingrantii, s e e : Paragynoxys martingrantii Senecio meridanus, s e e : Paragynoxys meridana Senecio neodendroides, s e e : Paragynoxys neodendroides Senecio santurbanensis, s e e : earagynoxys santurbanensis Senecio steyermarkii, s e e : Paragynoxys meridana Senecio venezuelae, s e e : earagynoxys venezuelae