Passiflora boenderi (Passifloraceae), a New Egg-Mimic Passionflower from Costa Rica John M. MacDougal Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. [email protected] Abstract. Passiflora boenderi, a new species of infra purpurata vel atrovinosa, hiloha; lobis lateralibus aeutis vel acuminatis, interdum rotundatis, lobo centrali Passifloraceae endemic to Costa Rica, is described obsolete, angulo inter lobos laterales 11—62°, marginibus and illustrated. It is assigned to section Decaloba integris, neetariis aureis. Flos parvus, petalis 3—6 mm lon- DC. in the species group that includes P. gilberti- gis; coronae filamentis biseriatis, exterioribus luteolis 3.5- ana J. M. MacDougal and P. ornithoura Masters. 5.5 mm longis; opereulo plicato; androgynophoro 2.9-4 This rare small-flowered species is notable for its mm longo; semina (7 vel) 8 ad 10-sulcata. intensely colorful and variegated leaves, with rows Vine 1.5—5 m, minutely puberulent throughout of conspicuous golden egg mimics. at the eernuous shoot lip. glabrescent below; stems Key words: Costa Rica, egg mimic. Passiflora, ± terete, striate. Stipules 1.7—3.5 X 0.2—0.3 mm. Passifloraceae. linear-triangular, subfalcate; petioles 1.2—3.8 cm, eglandular; leaf blades 4— 12(— I 7) cm long in out¬ For more than a century, a collection of a Costa line, 4.0—8.0(—9.5) cm wide, 2.4-7(—9.0) cm along Rican passionflower with bilobed purple-tinged central vein, lateral veins 4.0-8(-9.2) cm, entire, leaves with yellow stripes and dots has hidden in glabrous or glabrescent except margins minutely herbaria under different names. Only after plants lightly strigillose and sometimes with a few tri- of this taxon were collected alive and then culti¬ chomes on the larger veins abaxially, variegated vated have we been able to study the fresh flowers adaxiallv with light yellow along the three major and sec that it is an undescribed species, perfectly veins, especially on lateral lobes, (often deep) pur¬ morphologically distinct. While close relatives in ple or red-purple abaxially. truncate elliptic to ob- section Decaloba have similar variegated leaves, ovate to widely obovate (to very widely obovate) in this new species is most dramatic, with leaves dark general outline, bilobed %—V2(-Zf) their length, the green above and purple or red-purple beneath, with lateral lobes triangular to lanceolate, acute to acu¬ light yellow stripes along the two major lateral veins minate, the very apex sometimes rounded, the cen¬ and conspicuous bright golden spots in two lines. tral lobe absent or nearly so, sometimes represented It is a choice plant for ornamental cultivation for by a mucronate cusp less than 5 mm long, the angle its foliage, but is a local endemic in primary rain between the lateral lobes (14—)23—50(—62)° (juve¬ forest in central Costa Rica and is of conservation nile plants with leaves less deeply lobed). the ratio concern. of lateral to central lobe length 1.7—2.3(—2.5); lam¬ inar nectaries (5 to)8 to 13(to 16), borne between Passiflora boenderi J. M. MacDougal, sp. nov. the main veins, appearing bright yellow or yellow- TYPE: U.S.A. Florida: Broward Co., Coconut orange adaxiallv, and yellow-orange with purple Creek, cultivated at Butterfly World, Ltd., border abaxially, the larger leaves often with a sin¬ 1992—2002, type specimens collected Nov. gle nectary proximal (exmedial) to each lateral vein 2001, R. Roender 361, grown from plants col¬ at base of lamina; prophyll of vegetative bud 1, lected 19 May 1992 in Costa Rica. Alajuela: lanceolate to narrowly triangular. Peduncles (1)2 W of main road San Ramon to Bajo Rodriguez, per node, 1—3 cm, uniflorous; bracts 3, 1—2.8 X between Rio Cataratas and Rfo San Lorenzo 0.1—0.15 mm. linear-triangular, usually early ne¬ on side rd. to Colonia Palmarena, “near dead crotic and stramineous. Flowers light yellow-green, volcano,” ca. 1 km N of Finca Santa Cecilia, the corona yellowish with purple or purple-red ba- 10°13'43"N, 84°33'34"W, BOO m (holotype, sally and/or distally; flowers with little or no de¬ CR; isotypes, BM. CAS, INB. GOET, K. tectable odor, borne sub-horizontally at anthesis; MKXU, MO. TEX). Figures I. 2. floral stipe (3.5—)4.0—7.0 mm (6.5—9 mm in fruit); hypanthium 5.5—7 mm diam.; sepals 7.0— 11 (— 14) Species scandens ad Passifloram sectionem Decalobam pertinens. Folia petiolo eglanduloso; lamina supra vittata. X 3.0—5.0 mm, triangular-oblong to narrowly ovate- Novon 13: 454-458. 2003. Volume 13, Number 4 MacDougal 455 2003 Passiflora boenderi from Costa Rica Figure 1. Leaves of Possiflora boenderi (clone of type material, Boender 36]). Egg-mimic laminar nectaries can be seen as round light spots on the leaves. 456 Novon dres 70); styles 4—5 mm long including stigmas, green and unmarked, or with a faint overall flush of purple, the stigmas 1.5—2 mm diam. Fruit 11 — 21 X 10—20 mm, (ellipsoid) widely ellipsoid to subglobose, estipitate, purple-black with glaucous bloom, the mesocarp light green to white; arils 7— 8 mm long, transparent white to very pale trans¬ parent orange, gelatinous, sweet, not or only slightly fruity, nearly odorless; seeds 3.5—4.2 X (2.4—)2.7— 3.0 X 1.4—1.7 mm, transversely sulcate with (t)8 to 10 sulci, the intervening ridges strongly sculp¬ tured and verrucose, the funiculus conspicuous and white on fresh seed; seeds per fruit 24 to 67 (N = 10); germination epigeal. Ecology. Passiflora boenderi is known from only two sites in the Caribbean drainage of central Figure 2. Flower of Passiflora boenderi with inner coro¬ Costa Rica at 725—800 m elevation. These are nal series well expressed (clone of type material, Boender nearly identical habitats at the elevational transi¬ B6I). tion of very wet to pluvial premontane lorest. At both sites nearly all trees and branches are fes¬ triangular, obtuse to rounded, w ith no apical or sub- tooned with bryophytes, and the ground is often apical horn, light yellow-green outside, slightly pal¬ saturated and muddy. The vines are found growing er light yellow-green or green-yellow inside, at up and into small trees on steep slopes, and on anthesis reflexed (to wrapped under hypanthium); large shrubs at forest edges. Narrowly endemic spe¬ petals (2.7—)3.0—6.0 X 2.0—3.0 mm. triangular-ob¬ cies such as this passionflower, restricted to very long to broadly triangular-oblong, broadly obtuse, small ranges, are typically in danger of extirpation strongly reflexed at anthesis, sometimes erase api- by habitat conversion, and are therefore of special eally, very pale green (green-white) to [tale yellow- conservation concern. At the field site of the col¬ green; coronal filaments 2-seriate, the outer coronal lection of the type clones, P. boenderi was associ¬ filaments (31 to)35 to 43 (samples N = 10), 3.5— ated with or near P. ambigua Hemsley, P. lobata 5.5 mm long, at base narrow and greenish suffused (Fillip) J. M. MacDougal, P guatemalensis S. Wat¬ with light purple or red-purple, or dull purple son, P costaricensis Fillip, P vitifolia Funth, and whole proximal half, conspicuously geniculate near P. oerstedii Masters (R. Boender, pers. comm.). At middle, conspicuously thicker and dull to light yel¬ the Cariblanco site 1 found P capsularis L and P. low in middle or distal half, sometimes also purple lobata. or purple-brown distally, but always with yellow or Phenology. Flowering plants have been col¬ pale yellow at least in middle, at anthesis the outer lected in January, April, and August through No¬ corona forming a shallow bowl or saucer with a vember. strongly spreading and reflexed edge, the filament This newT species was first collected more than tips pointing downward, the anther-corona clear¬ 125 years ago by A. R. Endres and was cited by ance 2.5—3.5 mm; inner coronal series inconspic¬ Hemsley (1880: 481) as Passiflora, sp. (I1, capsu¬ uous, few to many, ea. 7 to 39 per flower, 1.0—1.8 laris [L.] ajf.)." The specimens are without exact mm, capillary, elavate to capitellate, dull light pur¬ locality. One of Endres’s known collecting sites is plish with yellow head; operculum 2.0 mm, plicate, Quebrada Verde near San Ramon, very close to the pale greenish, flushed pale violet or red-purple be¬ tvpe locality. We now know it from two localities, low the white apex; androgynophore 2.9-4.0 mm but it is rare at both. Earrv Gilbert’s 1978 collection long, pale green basally, suffused purplish or red- was the second discovery of the species, and was purple proximally, the free portions ol the staminal brought into cultivation for a short while but per¬ filaments 2.6—3.5 mm long, red-purple or green suf¬ ished before studies could be made of it. Despite fused with pale purple; anthers 2.0—2.3 mm long, repeated searches by botanists, the species was not purple-edged, the pollen yellow; ovary 1.5—2.0 X found again until 1984. In 1991 .Andres Vega found 1.3—1.5 mm, widely (to very' widely) ellipsoid, it near San Ramon, took cuttings, and reared the bright green, minutely puberulent in all specimens butterfly Heliconius cydno from eggs and larvae on seen except one where the ovary is glabrous (En- the plant. He guided Ron Boender there the next Volume 13, Number 4 MacDougal 457 2003 Passiflora boenderi from Costa Rica year and assisted getting more living material, some Fruit and aril observations were made by the au¬ of which ultimately furnished the type specimens. thor from ten fresh fruits produced in cultivation Clones of the type collection of P. boenderi were by Ron Boender through manual cross-pollination introduced to horticulture in late 1992 and had of several individuals of the type material. spread to Europe by 1995. The collection number Etymology. This species is named for Ronald of the type clones has been cited variously as Boen- Boender. founder of the Passiflora Society Interna¬ der 361, BW361, and BW92-361. Several recent tional, and also president for many years. As prime popular books on passionflowers have included this mover of that society, since 1989 Ron has brought species under a nomen nudum, and color photo¬ together several hundreds of persons interested in graphs can be seen in Vanderplank (1996: 61; the genus Passiflora, both amateurs and profession¬ 2000: 60—61). Kloek (1996: 105—106), and Ulmer als. He established a seed bank, living collections and Ulmer (1997: 110). database, newsletter, and annual national and in¬ Passiflora boenderi is assigned to subgenus De- ternational meetings. In this way, he has set the caloba (DC.) Reichenbaeh sect. Decaloba DC. on foundation for a forum that has truly benefited sci¬ the basis of its plicate operculum, cernuous shoot ence and the diffusion of knowledge of this plant tips, transverse testal sculpturing, and position of group. In Coconut Creek, Florida, he maintains the laminar nectaries. It is part of the species group world’s largest collection of germplasm of Passiflor- that includes 1\ ornithoura, P. gilbertiana, P. ape- aceae. Additionally, Ron brought this new species tala Killip. P. jorullensis HBK, P. mexicana Jussieu, into horticulture for closer study, unselfishly spread and P. affinis Engelmann. Most similar is an un¬ it widely, and hounded me for more than a year described relative of P. ornithoura from the moun¬ with carefully collected evidence that it was new. tains of Chiapas and Guatemala represented by Through his studies and educational presentation Matilda 3971. That can have remarkably similar of both butterflies and their host plants, he contin¬ leaves, but P. boenderi differs by its longer outer ues to champion and support numerous conserva¬ corona (3.5—5.5 mm vs. 1.8—3 mm), a slightly short¬ tion activities in North, Central, and South Amer¬ er androgynophore (2.9-4.0 mm vs. 3.4—-4.9 mm), ica. and a usually pubescent ovary versus an always glabrous ovary. The bilobed variegated leaves of P Paratypes. COSTA RICA. Locality unknown: 2500 ft.. boenderi can be similar also to l1. gilbertiana and “Costa Rica,” [anno 1867-1875], Endres 70 (BM, K). Al- P apetala, which both grow in Costa Rica. Passi¬ ajuela: along Rio Sarapiquf near Ua Virgen tie Socorro flora boenderi can be distinguished from P. gilber¬ near Cariblanco, Apr. 1978, L. E. Gilbert s.n. (TEX); cam- tiana by its shorter androgynophore (3—4 mm vs. ino Colonia Virgen del Socorro, Cariblanco de Sarapiquf, 6.2-9 mm), its shorter outer corona (3.5—5.5 mm 4 Jan. 2000, Kay & Vega 196 (MO); cuttings of Boender vs. 5—7 mm) that is strongly dilated proximally (vs. 361 from Costa Rica cultivated 1992—2002 at MO Cli- matron® greenhouse, 8 Sep. 1993, MacDougal 4697 (CR, filiform), and its habitat (elevations of 725—800 m MO); rd. to l.a Virgen de Socorro Irorn Rte. 9, near small vs. 1600—2300 m). From P. apetala it may be rec¬ cascade along the rd. on E side of the Rfo Sarapiquf, ca. ognized by its often longer outer corona (3.5—5.5 10°15.4'N, 84° 10.3'W, 29 Aug. 1994, I). Smith 1106 mm vs. 2-5 mm) that is strongly dilated proximally (DUKE); Sarapiquf, Cariblanco, carnino a Virgen del So¬ (vs. filiform), its longer petals (2.7-6 mm vs. absent corro, 8 Oct. 1992, A. Vega s.n. (CR, USJ). or to 2.5 mm), a usually pubescent ovary vs. always Acknowledgments. Great appreciation is due glabrous ovary, and its habitat (elevations of 725— Andres Vega, the indefatigable field worker and 800 m vs. 1280—3270 m). flic angles between the Costa Rican passionflower expert who helped re¬ leaves’ lateral lobes are usually wider in P. apetala discover this species and arranged for the collec¬ and narrower in P. gilbertiana. but there is overlap. The inner corona was variably expressed in the tion of the type material. I am grateful to the Pas¬ flowers of the six individuals I examined closely, siflora Society International for support for field varying from 7 to almost 40. This is similar to the studies and to Mario Posla for field assistance. 1 variable expression of the inner corona in its very thank Armando Estrada for checking the herbaria close relative, P. gilbertiana, where their number in CR, INB, and USJ, and Alexander Rodriguez of can vary greatly even on one plant (MacDougal, INB for his assistance. John Vanderplank of the 1989). The conspicuous bright golden laminar nec¬ National Collection of Passiflora in Britain gener¬ taries closely resemble Heliconius butterfly eggs ously offered resources and shared photographs (Gilbert, 1982), and I consider this species to have that assisted in preparing the description of the one of the most conspicuous eases of egg-mimicry species. The curators of BM, DUKE, K, and TEX known (pers. obs.). generously offered specimens on long-term loan. 458 Novon Literature Cited section Decaloba (Passifloraceae) from Costa Rica. Ann. Gilbert, L. E. 1982. The coevolution of a butterfly and a Missouri Bot. Card. 76: 608—614. vine. Sci. Amer. 247: 110—121. Ulmer, T. & B. Ulmer. 1997. Passionsblumen: Eine lasz- Hemsley, W. B. 1880. Biologia Centrali-Americana, Bot¬ inierende Gattung. Laupenmiihlen Druck, Witten. any, Vol. 1(6). 11. II. Porter and Dulau, London, Vanderplank, J. 1996. Passion Flowers, 2nd ed. MIT kloek, P. 1996. Das grosse Buch der Passionsblumen. Press, Cambridge. Lagerstroemia-Verlag, Hamburg. -. 2000. Passion Flowers, 3rd ed. MIT Press, Cam¬ MacDougal, J. M. 1989. Two new species of Pass if! ora bridge.