A new series and three new species of Passiflora subgenus Astrophea from the Guianas CHRISTIAN FEUILLET
Feuillet, C. (Department of Systematic Biology-Botany, MRC-166, Smithsonian Institution, Washington, DC 20560-0166, U.S.A.; email: bihorel@ concentric.net). A new series and three new species of Passiflora subgenus Astrophea from the Guianas. Brittonia 54: 18-29. 2002.--A new series, Passiflora ser. Carneae, and three new species are described in Passiflora subg. Astrophea: P. ascidia from Guyana and Venezuelan Guayana; P. balbis from Guyana, Venezuelan Guayana, and Amazonian Brazil; and P. saulensis from central French Guiana. A key to the 19 Guianan species of Passiflora subg. Astrophea is provided. Key words: Passifloraceae, Passiflora subg. Astrophea, Brazil, French Guiana, Guyana, Suriname, Venezuela. Feuillet, C. (Department of Systematic Biology-Botany, MRC-166, Smithsonian Institution, Washington, DC 20560-0166, U.S.A.; email: bihorel@ concentric.net). A new series and three new species of Passiflora subgenus Astrophea from the Guianas. Brittonia 54: 18-29. 2002.--Une nouvelle srrie, Passiflora ser. Carneae, et trois esprces nouvelles de Passiflora sous-genre Astrophea sont drcrites: P. ascidia du Guyana et de la Guyane vrnrzurlienne, P. balbis du Guyana, de la Guyane vrnrzurlienne et d'Amazonie brrsilienne, et P. saulensis du centre de la Guyane fran~aise. Une c16 permet d'identifier les 19 esp~ces guyanaises de Passiflora sous-genre Astrophea.
I n t e n s i v e c o l l e c t i n g in G u y a n a and French Guiana by several teams o f botanists (funded mainly by CAY, NY, R U, US) has increased the number o f species k n o w n to occur in the Guianas. In the Passifloraceae the number of collections and species increased dramatically since the Flora o f the Guianas (French Guiana, Guyana, and Suriname) program was created in the early 1980s, both by the extension to the Guianas of already k n o w n species and by the description o f new species. Killip (1938) cited Guianan collections for nine species of Passiflora subg. Astrophea, and Escobar (1994) cited in the key a Guianan distribution for 13 of them, including species described recently (Feuillet & Cremers, 1984; Escobar, 1994; Feuillet, 1994). In the present paper 19 species are recognized for the three Guianas, including three new species described herein.
Subgenus
Astrophea (DC.)
Mast.
Passiflora subg. Astrophea c o m p r i s e s about 60 species. The taxa are lianas, trees, or shrubs with stipules that are small and early deciduous. The petioles have two glands, adaxial at the apex of the petiole, or abaxial and at least partly on the base of the midrib. The leaf blades are simple and unlobed. The inflorescences are often 1-2flowered and sessile, derived from a simple cyme, and the terminal flower is transformed into a tendril. The inflorescences are in the leaf axils or sometimes borne from old growth on specialized short shoots with m u c h r e d u c e d , d e c i d u o u s leaves; these flowering shoots borne off w o o d y stems we will here call "inflorescences." The bracts are recaulescent on the pedicels and similar to the minute bracteoles; the pedicels are usually articulated. The flowers (hypanthi-
Brittonia, 54(1), 2002, pp. 18-29. 9 2002, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
ISSUED: 29 May 2002
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um, sepals, and petals) are white to red. The hypanthium (floral tube) is campanulate, funnel-shaped, or cylindric. The coronal filaments are yellow to orange. The limen is lacking. The fruits are 3-costate or hexagonal or round in cross section, usually with a chartaceous or brittle pericarp. The four species with long inflorescences in this paper (P. ascidia sp. nov., P. balbis sp. nov., P. longiracemosa Ducke, and P. saulensis sp. nov.) f o r m a well-defined group with two other species that have short inflorescences, P. amoena L. K. Escobar and P. fuchsiiflora Hemsl. This group of six Guianan species deserves recognition as a new series in subgenus Astrophea. The series can be separated f r o m other species of Astrophea by the following combination of characters: foliar nectary glands swollen, situated adaxially at the apex of the petiole; floral tube, sepals, and petals pink to red; leaf blades ovate or suborbicular to lanceolate, apex obtuse or rounded, often emarginate, base round to cordate. The leaves look remarkably alike and differ f r o m those o f the other species in the subgenus.
Passiflora ser. Carneae Feuillet, set. nov. TYPE: Passiflora ascidia Feuillet, sp. nov. Passiflora ser. Carneae in subg. Astrophea pertinens; foliis ovatis vel oblongatis, interdum ellipticis, infra glauca, basi obtusa, rotundata vel cordata, apice obtuso vel emarginato, inflorescentiis caulinis, tubo floris carneo, filamentis coronae brevibus, operculo propre basin vel usque ad trientem hypanthii, erecto differt. Lianas glabrous, with anomalous secondary growth and strong tendrils, reaching the canopy. Leaf: petiole with two adaxial glands at the apex; lamina ovate, oblong or rarely elliptic, base obtuse, rounded or cordate, apex obtuse or emarginate, glabrous, abaxially glaucous. Inflorescences borne on w o o d y stems, usually < 2 m from the ground. Flower: h y p a n t h i u m and perianth pink to red; corona shorter than in m o s t species o f subgenus Astrophea, the larger corona filaments 1 c m long or less; operculum erect, borne in the basal 1/3 of the hypanthinm. The species with short inflorescences ( < 5 c m long) have longer corona filaments (first row ca. 7 - 1 0 m m long). The species
19
with long inflorescences (10 cm long and m u c h more) have shorter corona filaments (first row 2 - 6 m m long). The tendency o f the i n f l o r e s c e n c e to r e s u m e v e g e t a t i v e growth at the apex has been observed in P. ascidia, P. balbis, and P. saulensis. A revision of Passiflora subg. Astrophea was prepared by the late Linda Escobar and the manuscript is now being completed and expanded by Katie Hansen (TEX). The revision will include a new assessment of the sections and will m o s t likely result in the description of new sections; therefore, it is better to refrain from placing too precisely the new series in the possibly obsolete sect i o n a l structure i n h e r i t e d f r o m Killip (1938). Nevertheless, one should k n o w that Killip included all the red-flowered species o f subg. Astrophea in sect. Botryastrophea (Harms) Killip. The type of this section, P. spicata Mast., has foliar nectary glands situated abaxially at the juncture of the petiole and the blade, leaf apex acuminate, base attenuate, and the smaller rows of corona filaments 4 - 5 m m long (0.1-1.5 m m in ser. Carneae). The new series is probably close to the red-flowered P. nuriensis Steyerm. f r o m the state of Bolivar, Venezuela, that has the same kind of foliar nectaries, but small flowers and acute to acuminate, coriaceous leaves, Etymology.--The a d j e c t i v e " carneus" means flesh-colored, referring to the pink to red flowers of the species in set. Carneae.
Species with Long Inflorescences When Ducke (1922) described Passiflora longiracemosa he cited two collections as syntypes, Ducke 16968 f r o m Lago do Salgado (Par~, Brazil) and Kuhlmann 2803 f r o m Rio Branco (Roraima, Brazil). In his Revision of the American Species of Passifloraceae, Killip (1938) selected Ducke's specimen deposited at M G as a lectotype. Besides K u h l m a n n ' s specimen, Killip cited three additional collections, Krukoff 1141 f r o m the Upper Cupary River (Parfi, Brazil), Im Thurn 84 f r o m the Rio Kukenfin (Bolivar, Venezuela) and Tate 207 from Mt. R o r a i m a (Bolivar, Venezuela). Additional collections have been identified as P. longiracemosa since then. The study of the
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syntypes, the specimens cited by Killip, and their comparison with new collections f r o m G u y a n a shows that specimens belonging to three species have been referred to P. longiracemosa. The two other species are described here, P. balbis f r o m Guyana, Venezuelan G u a y a n a (Amazonas, Bolivar) and Roraima (Brazil), and P. ascidia f r o m G u y ana and Venezuelan G u a y a n a (Bolivar). A third new species o f the same group, P. saulensis, is k n o w n from central French Guiana.
Passiflora ascidia C. Feuillet, sp. nov. (Fig. 1A-D) TYPE: G U Y A N A . Potaro-Siparuni: 2 - 3 k m N of Kato Village store-airstrip, o f f K a t o - K u r u k a b a r u rd., 4 ~ 59~ 6 9 0 - 7 5 0 m, 17 M a r 1989, L. J. Gillespie 8 3 1 (HOLOTYPE: B R G ; ISOTYPES: M O , NY,
US). Passiflora ascidia in subg. Astrophea pertinens, P. longiraeernosae affinis; tubo florali inflato, lobi brevibus differt.
L i a n a r e a c h i n g the c a n o p y , g l a b r o u s throughout. Young stems terete. Tendrils long and thin w h e n young. Stipules not seen, soon deciduous. Leaves: petiole 1-2 c m long, with 2 adaxial glands at the edge of the blade; blade ovate to lanceolate, 7 13 • 3.5-8 cm, obtuse or rounded to emarginate at the apex, rounded at the base, the margin entire, drying dark brown with pale veins adaxially and paler with dark veins abaxially, 6 - 8 main lateral veins on each side of the midrib, tertiary venation somewhat scalariform. Inflorescence 12-25 c m long, c a u l i f l o r o u s . P e d i c e l s 2 0 - 3 0 m m long, joined 2 - 4 m m from the base. Flowers pink outside ("bright p i n k " or "fluorescent pink," according to collectors), pink and white inside; hypanthium 2-2.7 c m long, round and dark at the base, then narrowly barrel-shaped, narrow at the throat; sepals lanceolate, 8 - 9 • 3.5 m m ; petals similar to the sepals, about 1/4 smaller; corona in 3 rows, the outermost of scattered 2-2.5 m m long filaments, the middle row of 0.5-1 m m long filaments or like the innermost, the innermost row of 0.1-0.2 m m tubercle; o p e r c u l u m attached near the base
[VOL. 54
of the hypanthium, mostly filamentous, 7 m m long; disk whitish; androgynophore ca. 4.5 c m long; stamen filaments 4 - 5 m m long, anthers dorsifixed, rectangular, 4 • 1.5-2 m m ; o v a r y 2.5-3 m m long, elliptic, glabrous; styles 2.5 m m long; stigmas ca. 1 m m wide. Fruit not seen. Distribution.--Passiflora ascidia h a s been collected in G u y a n a (Potaro-Siparuni) and Venezuela (Bolivar). It inhabits forest at 3 0 0 - 1 3 0 0 m and seems to be m o r e c o m m o n in low forests and slope forests. Etymology.--The specific epithet is a word in apposition, plural of " a s c i d i u m , " meaning " p i t c h e r s " in Latin. The flowers look like little pitchers with their inflated hypanthium and short lobes. Additional specimens examined. VENEZUELA. Bolivar: Rio Kukenfin, lm Thurn 84 (BG, BM, K, US); Arapuru, river side, 1250-1300 m, 1 Jan 1928, Tare 207 (NY). GUYANA. Potaro--Siparuni: 2 krn S of Paramakatoi, trail to Paramaktipu, 04~ 59~ 8001100 m, 21 Feb 1996, Clarke 1162 (BRG, US); 2-2.5 km W of Kato, along Chiung R., 4~ 59~ 650-675 m, 25 Mar 1989, Gillespie 880 (BRG, US); Pakaraima Mtns., summit of Cipo Mtn., 4~ 60~ 1250 m, 29 Jan 1993, Henkel 1106 (BRG, MO, NY, US); Pakaraima Mtns., Upper Ireng R. watershed, Sukabi R., 1-2 km upstream from Ando Falls, 5~ 60~ 800 m, 21 Oct 1994, Henkel 6010 (BRG, MO, US); Pakaraima Mtns., Upper Ireng R., 12 km upstream from Catch-a-Cow, 5~ 59~ 305 m, 16 Oct 1994, Mutchnick 50 (BRG, US).
Passiflora balbis C. Feuillet, (Fig. 2)
sp. nov.
TYPE: G U Y A N A . Upper T a k u t u - U p p e r Essequibo: N Rupununi, 4 - 5 k m f r o m Sura m a village, near Burro-Burro R., 1 0 0 - 1 5 0 m, 20 Feb 1990, C. Feuillet 10620 (HOLOTYPE: BRG; ISOTYPES: MO, NY, US). Passiflora balbis in subg. Astrophea pertinens, P. longiracemosae affinis; petiolo breviore, tubo florali angusto, corona nigra, secunda serie porcata, sine tuberculo vel filamento differt.
L i a n a r e a c h i n g the c a n o p y , g l a b r o u s throughout. Young stems terete, b e c o m i n g w o o d y with irregular secondary growth. Tendrils pinkish red when young. Stipules not seen, soon deciduous. Leaves: petiole ca. 3 c m long, with 2 adaxial glands at the edge of the blade; blade ovate to lanceolate, 10-11 • 5 - 6 . 5 cm, rounded to emarginate
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21
lcm
D
lcm
lem~
FIG. 1. A-D. Passiflora ascidia. A. Leafy stem. B. Cauline inflorescence. C. Flower. D. Flower, longitudinal section. E. P. fuchsiiflora flower, longitudinal section. F. P. longiracemosa flower, longitudinal section. (A-D from Gillespie 831, US and pickled material; E: from Cremers 15169, US; F: from Hoffman 347, US.)
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[VOL. 54
[ lcrn
[ ,0m[
lmm
D
E
,ore[ F
~~
~
D
FIG. 2. Passiflora balbis. A. Leafy stem. Bo Petiolar nectaries. C. Flowering stem. D. Flower, longitudinal section. E. Fruits. F. Fruit, longitudinal section. (A, B from Jansen-Jacobs et al. 2249, US; C-F from Feuillet 10620, US, pickled material, and photographs by the author.)
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at the a p e x , r o u n d e d at t h e base, s o m e t i m e s slightly cuneate around the petiolar glands, the m a r g i n entire, d r y i n g d a r k b r o w n w i t h pale veins adaxially and paler with dark veins abaxially, 6-8 main lateral veins on e a c h side o f the m i d r i b , t e r t i a r y v e n a t i o n scalariform. Inflorescences numerous, cauliflorous, 1 - 6 m l o n g , p r o d u c e d at the n o d e s of old woody stems, mostly from ground l e v e l to 2.5 m a b o v e it, s o m e t i m e s l o o s e l y b r a n c h e d , b r a n c h e s 5 c m to 2 m long. P e d i c e l s ca. 15 m m l o n g , j o i n t e d 4 - 5 m m f r o m the base. F l o w e r s p i n k ; h y p a n t h i u m c y l i n d r i c a l , 4.5 c m l o n g , ca. 5 m m d i a m . , r o s e red; s e p a l s p i n k o r l i g h t g r a y outside, p i n k o r p u r p l e to d a r k g r a y i n s i d e , 1 2 - 2 5 m m l o n g (to 60 m m l o n g in fruit), 5 - 6 m m wide; petals usually somewhat more pink t h a n the sepals; c o r o n a in 2 r o w s , the o u t e r r o w o f filaments ca. 3 m m long, d a r k p u r p l e to b l a c k i s h , the i n n e r r o w a r i d g e o r a series o f t u b e r c l e s 0.3 m m thick; o p e r c u l u m b o r n e 6 - 7 . 5 m m f r o m the b a s e o f t h e h y p a n t h i u m , a deeply 5-10-cleft membrane 5 mm long, b e a r i n g at a p e x 1 - 2 . 5 m m l o n g filaments; disk white; androgynophore 4.5-5 cm long, p a l e p i n k ; s t a m e n f i l a m e n t s ca. 5.5 m m long, pale pink, anthers dorsifixed, rectangular, 3 . 5 - 4 • 1.5 m m , y e l l o w ; o v a r y slightly obovoid, whitish, glabrous; styles p a l e p i n k to b l a c k i s h red, 3 - 4 m m l o n g ; s t i g m a s y e l l o w to g r e e n i s h , ca. 1 m m w i d e . Fruit pendent, oblong ellipsoid, somewhat h e x a g o n a l in section, t a p e r i n g at b a s e , r o u n d to t a p e r i n g , d a r k p u r p l i s h g r e e n turni n g to d a r k red, 6 - 1 0 X 2 . 5 - 5 c m ; s e e d s o b l o n g , s l i g h t l y c u r v e d , 6 • 3 - 4 m m , flatt e n e d , ca. 1 m m t h i c k at m a r g i n a n d < 2 m m t h i c k in the m i d d l e , m a r g i n o f e a c h h a l f crenate, the s w o l l e n , p a l e s u t u r e b e i n g in a d e p r e s s i o n b e t w e e n t h e c r e n a t e m a r g i n s , sides reticulate-foveate. M o s t l i k e l y the i n f l o r e s c e n c e s m a y r e a c h e v e n g r e a t e r l e n g t h t h a n r e c o r d e d here. T h e h i g h m e a s u r e m e n t g i v e n reflects the l i m i t o f m y p a t i e n c e at f o l l o w i n g a n d m e a s u r i n g this l o n g i n f l o r e s c e n c e ( F e u i l l e t 1 0 6 2 0 ) , ent a n g l e d w i t h m a n y others, g r o w i n g m o r e o r less h o r i z o n t a l l y t h r o u g h t h e d e n s e s h r u b b y u n d e r s t o r y v e g e t a t i o n . T h e tips o f the inflor e s c e n c e s t e n d to r e s u m e v e g e t a t i v e g r o w t h . Distribution and ecology.--Passiflora b a l b i s is k n o w n f r o m B r a z i l ( R o r a i m a ) ,
23
Guyana (Potaro-Siparuni, Upper TakutuUpper Essequibo), and Venezuela (Amaz o n a s , Bolfvar). It i n h a b i t s f o r e s t at 7 5 1650 m a n d s e e m s to b e m o r e c o m m o n in s e a s o n a l l y f l o o d e d f o r e s t s and s u m m i t forests. Etymology.--The s p e c i f i c e p i t h e t is a w o r d in a p p o s i t i o n , " b a l b i s " (Gr.) b e i n g the s t r i n g or r i b b o n s t r e t c h e d a c r o s s the t r a c k that is c u t b y the w i n n e r o f a race. It d e s c r i b e s the v e r y l o n g i n f l o r e s c e n c e s stretchi n g t h r o u g h the u n d e r s t o r y shrubs a c r o s s the collector's way. Common name.--Kurashidi people, amarrar (Cruxent 42). Additional specimens examined. VENEZUELA. A m a z o n a s : Near Base R. (Carlo Negro), SE base of
Cerro Duida, 215 m, 23 Aug 1944, Steyermark 57915 (US). Bolivar: Gran Sabana, near Rio Kukenfin, base of Mount Roraima, 1175-1300 m, 24 Sep 1944, Steyermark 58566 (US); Rio Karuai, at the base of the Sororopfin-tepui, W of La Laja, 1220 m, 29 Nov 1944, Steyermark 60763 (US); Medio Caura, Salto de Para, 250 m, 4 Mar 1939, Williams 11369 (US). GUYANA. Potaro--Siparuni: Pakaraima Mtns., Mt. Wokomung, Wusupubaru Creek, 2 km from Suruwabaru Creek, 5~ 59~ 975-1125 m, 13 Feb 1993, Henkel 1362 (BRG, MO, NY, US); Pakaraima Mtns., Mt. Wokomung, summit ridge of Ka-mie-wah pinnacle, 5~ 59~ 1550-1650 m, 17 Nov 1993, Henkel 4499 (BRG, US); 4-5 km N of Surama village, trail to Burro-Burro & Surama R., 4~ 59~ 75 m, 1 May 1992, Hoffman 1530 (BRG, MO, NY, U, US); Pakaraima Mtns., Upper Ireng R., 1-2 km upstream from Catch-a-Cow, 5~ 59~ 305 m, 16 Oct 1994, Mutchnick 51 (BRG, US). Upper Takutu-Upper Essequibo: Rio Ugueto, 340 m, 1 Jan 1952, Cruxent 42 (NY); Kuma, 15 km SE of Lethem, foot of the Kunuku Mtns., 3~ 59~ 120-150 m, 7 Jul 1989, GiUespie 2033 (BRG, MO, NY, US); S Rupununi Savanna, Aishalton Mtns., 2 km S from Aishalton village, 2~ 59~ 200-400 m, 12 Oct 1993, Henkel 3445 (AAU, BRG, MO, NY, US); S Rupununi Savanna, 15 km SSE from Aishalton, 2~ 59~ 200 m, 19 Nov 1993, Henkel 3488 (BRG, MO, NY, U, US); S Rupununi Savanna, Ororkar bush, SE of Aishalton, 2~ 59~ 250 m, 8 Feb 1994, Henkel 3725 (BRG, MO, US); S Rupununi, Toot R., 25 krn SE of Aishalton village, 2~ 59~ 250 m, 27 Apr 1994, Henkel 3822 (BRG, MO, US); Kanuku Mtns., near Rupununi R., Puwib R., 3~ 59~ 80-100 m, 19 Feb 1985, JansenJacobs et aL 191 (U, US), 303 (U, US); Kanuku Mtns., Nappi R. Head, SE slope of Mount Tsjiri Brue, 3~ 'N, 59~ 150-250 m, 2 Nov 1987, Jansen-Jacobs et al. 652 (NY, U, US); Kanuku Mtns., Nappi R. Head, Camp 1, 3~ 59~ 130 m, 4 Nov 1987, Jansen-Jacobs et al. 696 (U); E Kanuku Mtns., NE of Warimure, 3~ 59~ 200-500 m, 25 Jan 1991, Jansen-Jacobs et aL 2249 (U, US); Kusad Mts.,
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2~ 59~ 250-450 m, 29 Sep 1992, JansenJacobs et al. 2683 (U, US); Bushmouth Shea to Kwitaro R., Bowl Cr., 2~ 59~ 200 m, 29 Aug 1995, Jansen-Jacobs et al. 4927 (U); Rewa R., "near camp 2 at foot of Spider Mtn.," 3~ 58~ 220 m, 15 Sep 1999, Jansen-Jacobs et al. 5917 (U, US); Rewa R., Spider Mtn., 3~ 58~ 400-500 m, 20 Sep 1999, Jansen-Jacobs et al. 6019 (U, US); foothills of NE Kanuku Mtns., near Moco-Moco village, 3~ 59~ 100 m, 26 Oct 1979, Mass & Westra 3831 (MO, NY, U); Kanuku Mtns., Nappi R., 14 Oct 1935, Myers" 5374 (K, US); W extremity of the Kanuku Mtns., drainage of the Takutu R., 250 m, 4 - 2 2 Mar 1938, Smith 3116 (A, E MO, NY, R US). BRAZIL. Roraima: "Prope Cachoeira do Rio Branco," Dec 1912, Kuhlmann 2803 (G-BOIS, K, MG, RB, S, U, US; photograph MO); foothills of Serra da Lua, 2~ 60~ 12 Jan 1969, Prance et al. 9228 (INPA); SEMA, Ecological Station, llha de Maracfi, 3~ 61~ 21 Mar 1987, Ratter et al. 5799
(us).
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green spots; seeds (when immature) similar in shape and ornamentation to P. balbis. Distribution.--Passiflora longiracemosa has been found in Brazil (Parfi) and G u y a n a (Upper Takutu-Upper Essequibo). It inhabits forest at ca. 2 0 0 - 2 5 0 m. This distribution is disjunct and it is likely that collections will be made in southwestern Suriname or in the state of Roraima, Brazil. E t y m o l o g y . - - T h e specific e p i t h e t describes the long racemose inflorescence, reaching 60 cm long. Additional specimens examined. Guyana. U p p e r T a k u t u - U p p e r Essequiho: Acarai Mtns., 10 km S of Sipu R., l~ 58~ 1100 m, 2 Sep 1998, D. Clarke 7304 (BRG, US); SE Kanuku Mtns., trail beside Crabwood Creek, ca. 6 km NE of Makawatta Mtn., 3~ 59~ 200-220 m, 21 Oct 1991, Hoffman 347 (BRG, US).
PASSIFLORA LONGIRACEMOSA Ducke, Archiv.
Jard. Bot. Rio de Janeiro 3: 221; pl. 23. P a s s i f l o r a s a u l e n s i s C. Feuillet, sp. nov. 1922. TYPE: BRAZIL. Parfi: For&, ~t (Fig. 3) l'est du lac Salgado, bas Trombetas, TYPE: F R E N C H GUIANA. Near Saiil, 1918, A. Ducke 16968 (LECTOTYPE: des53~ ignated by Killip, 1938, MG; ISOLECTO- Mont Galbao, SE Peak, 3~ ca. 700 m, 13 Sep 1994, B. Boom 10861 TYPES: G , GH, S, US (Fig. 1F) (HOLOTYPE: NY). Liana reaching the canopy, glabrous. Passiflora saulensis in subg. Astrophea pertinens, P. Young stems terete. Tendrils thick. Stipules not seen, soon deciduous. Leaves: petiole longiracemosae affinis; tubo florali infundibuliformi, nec ventricoso differt. 6 - 9 cm long, with 2 adaxial glands at the edge of the blade; blade broadly ovate, 1 0 Liana, glabrous throughout. Young stem 20 • 12-18 cm, rounded to emarginate at terete. Tendrils thick. Stipules narrowly trithe apex, broadly and shallowly cordate at angular, minute, 0.2-0.3 m m long, early dethe base, the margin entire, drying dark ciduous. Leaves: petiole 2-3 cm long, with brown adaxially, glaucous abaxially, 6 - 8 2 adaxial glands at the limit with the blade; main lateral veins on each side of the mid- blade ovate to suborbicular, 10-11 • 6 - 8 rib, tertiary venation scalariform. Inflores- cm, obtuse at the apex and slightly emarcence to 60 cm long. Pedicels 12-20 m m ginate, rounded at the base, margin entire, long, jointed 5-7 m m from the base. Flow- drying dark green with lighter brown veins ers red or bright pink; hypanthium cylin- adaxially, and light g r e e n with d a r k e r drical, ventricose at the base, 3-4.5 cm b r o w n veins a b a x i a l l y ( g l a u c o u s g r e e n long; sepals triangular, 11-13 • 3 - 4 ram; when fresh), 4 or 5 pairs of main lateral petals somewhat smaller; corona in 2 rows, veins, tertiary veins not obvious, intermethe outer row o f filaments 3 m m long, the diate between scalariform and forming a inner of tubercles; operculum borne near loose net. Inflorescence cauliflorous, 1 0 - 8 0 the base of the hypanthium, 9 - 1 0 m m long, cm long, the longer one resuming vegeta5-lobed, lobes fimbrillate at apex; andro- tive growth on the apical 10 cm, nodes 1gynophore 4 - 5 cm long; stamens filaments flowered; pedicels 18-24 m m long, jointed 4 - 5 mm long, anthers dorsifixed, rectan- 6-8 mm from the base; bracts minute. gular, 4 - 5 • 2 ram, apiculate; ovary 4 - 6 Flowers red: hypanthium 3.5-4 cm long, m m long, glabrous; styles 3 - 4 m m long; not ventricose at the base, gently funnelstigmas 2 m m wide. Fruit (when young) shaped, ca. 3 times as wide at the throat pyriform, 4 • 2 cm, red with many small than at the base; sepals 15-30 m m long,
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B
lern
FIG, 3. Passiflora saulensis. A. Leafy stem. B. Cauline inflorescence. C. Flower, longitudinal section. (From Boom 10861, NY,)
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rounded at the apex; petals 12-25 m m long; corona in 3 rows, the outmost row of flattened filaments, rounded at apex, slightly curved, 6 m m long, the inner 2 rows o f filaments, 1.5 and 0.5 m m long respectively, somewhat flattened and irregular in shape; operculum inserted 1.3 cm from the base o f the hypanthium, filamentous, 5 - 8 m m long; androgynophore 5-5.5 c m long; stamen filaments flat, 6 m m long, anther dorsifixed, 7.5 • 4 m m , rectangular; ovary 7 - 8 • 3 m m , styles 3 m m long, stigmas 4.5 m m diam. Fruit not seen. Distribution.---Passiflora saulensis is k n o w n only by the type collection f r o m the top of Mont Galbao, east of SaUl (central French Guiana), near 700 m. Etymology.--The type specimen was collected near SaUl in central French Guiana. This is the center o f intensive collecting by The N e w York Botanical Garden under the direction of Scott A. Mori.
Species with Short Inflorescences Two species of the new series b l o o m on short stems; these inflorescences grow f r o m w o o d y stems and are often shorter than the floral tubes (Fig. 4B). PASSIFLORA AMOENA L. K. Escobar, Syst. Bot. 19: 203. 1994. TYPE: F R E N C H G U I A N A . K a w Mtns., Rt. D6, 26 k m E of Roura, C a m p Caiman, 11 N o v 1986, D. E. Stone 3962B (HOLOTYPE: D U K E ) . (Fig. 4)
Passiflora amoena is unique in a few characters. The pink flowers are thick and rubbery in texture, m o r e so than P. glandulosa Cav. of subg. Passiflora. The yellow outer corona filaments are narrowly triangular and inserted on the floral tube by an angle of their narrow side, thus pointing outward at anthesis (Fig. 4D). The o b o v o i d fruits are green, maturing brownish, but entirely covered with a pink bloom. Passiflora amoena has been c o m m o n l y collected in French Guiana, with a handful of specimens f r o m G u y a n a and Suriname.
[VOL. 54
PASSIFLORA FUCHSIIFLORAHemsl., H o o k e r ' s Icon. P1. 26: pl. 2553. 1898. TYPE: G U Y ANA. D e m e r a r a R., G. S. Jenman 6540 (HOLOTYPE: K; ISOTVPES: B, B R G , NY). (Fig. 1E) The thin f o w e r s o f P. fuchsiiflora are described on herbarium labels as pink, red, or pinkish violet. The orange outer corona filaments are ligulate and erect. The narrowly obovoid fruits are green, maturing reddish brown. This species has been m o s t frequently collected in Guyana, with a few specimens coming f r o m Brazil, Suriname, and French Guiana.
Species Affinities I n s i d e ser. Carneae, m o s t c h a r a c t e r s demonstrate the similarity between the six species, but we can rely on the type of inflorescence and the structure of the corona and the operculum. These are v e r y stable characters and therefore their variations are significant in assessing affinities between species. The two species with short inflorescences have the operculum borne in the floral tube near the base. T h e y are arguably the two closest species and can be separated as follows: the operculum of P. amoena is a timbrillate m e m b r a n e and the outer filaments of the corona are narrowly triangular, wider near the base; the operculum of P. fuchsiiflora is nearly entirely filamentous and the outer filaments of the corona are ligulate, wider at or a b o v e the middle. A m o n g the four species with long inftorescences there is m o r e variation. The operculum is a long-fimbrillate m e m b r a n e and borne near the base o f the floral tube in Passiflora ascidia; it is membranous, 5 - 1 0 cleft, and fimbriate, in P. balbis and P. longiracemosa; it is nearly entirely filamentous in P. saulensis. In P. balbis and P. saulensis the operculum is borne in the floral tube about 1/3 f r o m the base, which allows for a larger nectary c h a m b e r at the base of the tube.
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F
] 2cm
'~176 C
1o" E 2cm
j A
1oo[
E3
[3 FIG. 4. Passiflora amoena. A. Leafy stem. B. Flowers at the base of the plant. C. Flower. D~ Flower, longitudinal section. E. Fruits. (From Granville et al. 10806, US, and from photographs by the author.)
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[VOL.
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Key to the Guianan species of Passiflora subg. Astrophea 1.
1.
Foliar nectary glands at apex o f petiole, adaxial, swollen, not part o f the blade. 2. Leaf blades not glaucous abaxially; perianth white or whitish. 3. L e a f blades rufo-tomentose abaxially; h y p a n t h i u m funnel-shaped. (Suriname, French Guiana, N Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. candida (Poepp. & Endl.) Mast. 3. L e a v e blades glabrous; h y p a n t h i u m cylindrical. 4. L e a f blade membranaceous. (French Guiana) ................. P. plumosa Feuillet & C r e m e r s 4. L e a f blade coriaceous. 5. Sepal/hypanthium length ratio > 2:1. 6. L e a f blades 12-35 c m long; floral tube 1 5 - 2 0 m m long. ( G u y a n a ) ........... ................................................................................................... P. maguirei Killip 6. Leaf blades 6 - 9 c m long; floral tube 8 - 9 m m long. (Venezuela, Guyana) ........ ...................................................................................................... P . c a r d o n a e Killip 5. Sepal/hypanthium length ratio <1.5:1. 7. Leaf blades 3-12 c m wide, not undulate at margin, margin with a dark band when dry, with 8 or 9 m a i n lateral veins. (French G u i a n a to Peru) .............. ...................................................................................... P . c i t r i f o l i a (Juss.) Mast. 7. Leaf blades 2 - 4 c m wide, strongly undulate at margin, margin without a dark band w h e n dry, with 1 2 - 1 5 m a i n lateral veins. ( G u y a n a ) ......................... .................................................................................. P. quelchii N. E. B r o w n 2. Leaf blades glaucous abaxially, at least w h e n fresh; perianth pink to red . . . . . . . . . . . . . (P. ser. C a r n e a e ) 8. O p e r c u l u m inserted near the base of the hypanthium. 9. Inflorescence short, 1-4 c m long; h y p a n t h i u m ventricose at the base; outer corona filam e n t s 7 - 1 0 m m long. 10. Petiole 1.5-5 cm long; perianth fleshy; outer corona filaments pointing outward, ca. 7 m m long, narrowly triangular, wider near the base, pink at base, yellow at apex; fruit reddish green to brownish under a pink bloom. (Guyana, Suriname, French Guiana) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. amoena L. K. Escobar 10. Petiole 5 - 1 0 cm long; perianth membranaceous; outer corona filaments erect, ca. 10 m m long, subdolabriform, wider in the upper half, acuminate, orange; fruits green to red-brown. (Guyana, Suriname, French Guiana) . . . . . . . . . P . f u c h s i i f l o r a Hemsl. 9. Inflorescence long, 1 0 - 6 0 0 c m long, occasionally blooming before reaching 10 c m long; h y p a n t h i u m ventricose at the base or wider near the middle; outer corona filaments 2 - 3 m m long. 11. Petiole 6 - 9 c m long; h y p a n t h i u m cylindrical, ventricose at base; corona in 2 rows. (Guyana, N Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. longiracemosa Ducke 11. Petiole 1-2 c m long; h y p a n t h i u m wider near the middle, not ventricose at base; corona in 3 rows. (Venezuelan Guayana, Guyana) ..................... P. ascidia Feuillet 8. O p e r c u l u m inserted about 1/3 up from the base of the h y p a n t h i u m 12. H y p a n t h i u m cylindrical, ventricose at the base, long and narrow, sometimes s o m e w h a t curved; corona in 2 rows, outer row filaments ca. 3 m m long. (Venezuelan Guayana, Guyana, N Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P . b a l b i s Feuillet 12. H y p a n t h i u m funnel-shaped; corona in 3 rows, outer row filaments ca. 6 m m long. (French Guiana) ........................................................ P. saulensis Feuillet Foliar nectary glands at base of blade, abaxial, on midrib and/or lamina, scar-like. 13. Perianth orange-red. (Venezuelan Guayana, Guyana, N Brazil) ................. P. securiclata Mast. 13. Perianth white. 14. Peduncles 2 0 - 6 0 c m long, pedicels 3.5-7 c m long. (Guyana, Suriname, French Guiana) ...................................................................................... P. leptopoda Harms 14. Peduncles lacking or 1-2 c m long; pedicels 0 . 5 - 4 c m long. 15. Leaves densely pubescent on both surfaces. (Guyana, French Guiana, N Brazil) -.................................................................................. P. ceratocarpa E Silveira 15. L e a f blade essentially glabrous on adaxial surface. 16. Leaves 4 - 7 cm long, rounded to broadly elliptic or cuneate, blunt or emarginate at apex, reticulate venation forming abaxially a thick raised net. (Venezuela, Guyana) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. sclerophylla Harms 16. Leaves usually 8-25 c m long, elliptic, ovate or obovate, acuminate to emarginate at the apex, venation not reticulate. 17. Leaves obovate, with 12-15 pairs of major lateral veins. (Guyana, Suriname, French Guiana, to Peru) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. costata Mast. 17. Leaves ovate or elliptic to lanceolate, with < 1 3 pairs of major lateral veins.
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FEUILLET: ASTROPHEA 18. 18.
29
Leaves elliptic to lanceolate, apex obtuse to acute. (Venezuela, Guyaria, French Guiana) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. o v a t a Martin ex DC. Leaves widely ovate, apex acuminate. (Guyana, French Guiana) .... .............................................................. P. k a w e n s i s Feuillet
Acknowledgments
Literature Cited
I thank the curators of the herbaria CAY, NY, R and U for lending the material in their care and allowing m e to work in their institution. The author wants to thank Cathy Pasquale-Johnson for the excellent illustrations. This paper has been improved by the careful reviewing and insightful suggestions by Katie Hansen, John MacDougal, and an anonymous reviewer. This article is published as Studies on the Flora of the Guianas no. 95; it is n u m b e r 52 in the Smithsonian's Biological Diversity of the Guianas Program publication series.
Ducke, A. 1922. Plantes nouvelles ou peu connues de la r6gion amazonienne. IIe partie. Archiv. Jard. Bot. Rio de Janeiro 3: 3-269. E s c o b a r , L. K. 1994. Two new species and a key to P a s s i f l o r a subg. A s t r o p h e a . Syst. Bot. 19: 203-210. Feuillet, C. 1994. Two new species of P a s s i f l o r a (Passifloraceae) from French Guiana. Novon 4(3): 236241. - & G. C r e m e r s . 1984. P a s s i f l o r a c e a e nouvelles ou m6connues de Guyane fran~aise. Proc. Kon. Ned. Akad. Wetensch. C. 87(4): 377-386. Killip, E. P. 1938. The American species of Passifloraceae. Publ. Field Mus. Nat. Hist., Bot. Ser. 19: 1613.