Brittonia, 41(1), 1989, pp. 44-52. �9 1989, by the New York Botanical Garden, Bronx, NY 10458-5126

THREE NEW SPECIES OF DALECHAMPIA (EUPHORBIACEAE) FROM VENEZUELAN GUAYANA

W . SCOTT ARMBRUSTER

Armbruster, W. Scott (Department of Biology & Wildlife and Institute of Arctic Biology, University of Alaska, Fairbanks, AK 99775-0180). Three new species of Dalechampia (Euphorbiaceae) from Venezuelan Guayana. Brittonia 41: 44-52. 1989.--Three new species of Dalechampia are described from the Guayanan region of Venezuela. Daleehampia attenuistylus belongs to section Rhopalostylis and may be most closely related to D. parvibracteata Lanj. Dalechampia mega- carpa belongs to section Dioscoreifoliae, although it is not especially close to any species in the section. Dalechampia papillistigma is also an isolated member of section Dioscoreifolieae.

Eastern Venezuela is a floristically rich and topographically varied region of South America. Although poorly collected in the past, improvements in the road system have made possible more thorough botanical exploration in recent years. On field trips to this region in 1985 and 1987-1988 I collected fertile material of three undescribed species ofDalechampia L. One of these species (D. papillistig- ma) had previously been collected in fruit from this region. The second species (D. megacarpa) had previously been collected in fruit only in one location in Territorio Federal Amazonas, Venezuela, and was not expected in eastern Bolivar. The third species (D. attenuistylus), as far as I know, had not been collected before.

The work in Venezuela forms part of a larger revisionary study of sections Rhopalostylis (Klotzsch) Pax & K. Hoffm. and Dioscoreifoliae Pax and K. Hoffm., to which the three new species belong. It is necessary, however, that the new Venezuelan species be published in advance of the rest of the project so that the genus can be treated in its entirety in the soon-to-be published Flora of the Venezuelan Guayana (Steyermark et al., in prep.).

Dalechampia attenuistylus Armbruster, sp. nov. (Fig. 1) Differt ab aliis speciebus sectionis proprii Rhopalostylis floribus staminatis 10, stylis attenuatis non

clavatis, stigmatibus chlorinis, et bracteis involucralibus viridibus stipuliformibus 3-10 mm longis.

Clambering vine; stems twining, terete, 0.5-1.2 mm thick at growing tips, strig- ulose with persistent whitish incurved hairs ca 0.1 mm long. Leaves alternate, simple; stipules spreading-reflexed, commonly deciduous, lanceolate-falcate (ovate on fertile short-shoots), 3-8 mm long, 1-2 mm wide, sparsely strigulose, striate- veined; petioles terete, 1-4.5 cm long, strigulose as stem; stipels at base of blade 2, lanceolate, 1-2 mm long, 0.4-0.6 mm wide, the point of attachment surrounded by cluster of ca 12 glands; blades chartaceous, ovate, unlobed, 3-10 cm long, 2- 7 cm wide, sparsely strigulose above and densely strigulose below with minute whitish hairs 0.05-0.1 mm long; base shallowly cordate, the sinus 5-10 mm deep, 3-5 mm wide at petiole, 10-25 mm wide at outer margin of blade; apex cuspidate; margins plane, entire to sinuate, with glandular processes 0.1-0.2 mm long; prin- cipal veins 5 (7) from base, the secondaries ascending, the tertiaries scalariform, sparsely strigulose with whitish incurred hairs ca 0.1 mm long. Pseudanthial inflorescences solitary on reduced axillary short-shoots 2.5-8 cm long at anthesis, pubescent as stem, with 1 or 2 nodes each with reduced stipules, the leaf usually deciduous; or pseudanthial inflorescences borne on determinate shoots, 10-20 cm long with 2 or 3 nodes, each with a persistent or deciduous leaf, 1 pseudanthial inflorescence terminal, with additional pseudanthial inflorescences on axillary

1989] ARMBRUSTER" DALECHAMPIA 45

2 m m

L. 1 c r n J

5 mm

Fto. 1. Dalechampia attenuistylus. A. Habit. B. Pseudanthial inflorescence. C. Staminate pleio- chasium. D. Pistillate cymule. E. Fruit.

short-shoots produced from nodes; distance between ultimate node and proximal involucral bract 12-40 mm at anthesis in both arrangements. Involucral bracts green at anthesis, unlobed, lanceolate, sparsely strigulose with small incurring hairs, the base cuneate, the apex acuminate; proximal bract (subtending pistillate involucel) 4-10 mm long, 1-3 mm wide, the margin plane, glandular-dentate, the prominent veins 5 from base; distal bract (subtending staminate involucel) 3-6 mm long, 0.5-2 mm broad, the margin plane, entire, the prominent veins 3 from base; stipules of proximal bract lanceolate-falcate, 2-3 mm long, 1-1.5 mm wide,

46 BRITTONtA [VOL. 41

strigulose, the apex acute, striate with 5 or 6 parallel veins, with several glands at point of attachment, those of distal bract similar except 2.8-3.5 mm long. Staminate cymule with densely puberulent peduncle 3.5-7.5 mm long, 1.2-1.5 mm thick at anthesis; involucel of 2 pairs of more or less decussate, free bracts, these broadly ovate, 1.5-2.5 mm wide, puberulent at base, indistinctly striate- veined, the apex more or less acute, the margins entire, ciliate; staminate flowers 10 (11), arranged in a 4-branched pleiochasium with 1 terminal flower, 3 branches with usually 3 flowers each, and a fourth (= dorsal, opposed pistillate cymule) arm usually sterile (rarely 1-flowered), each arm subtended by 2-5 lanceolate or ob- lanceolate bractlets, the tip blunt to acute, sometimes broader than limb, often deeply toothed and glandular-laciniate, elsewhere margins entire to laciniate. Sta- minate flowers on sparsely strigose pedicels 3-5 mm long, ca 0.7 mm thick, articulate near top; calyx sparsely strigulose, splitting into 3 or 4 obovate, acute, spreading segments, ca 2 mm long, ca 1 mm wide; staminal column glabrescent, 2-2.5 mm long, 0.2-0.3 mm thick; stamens 8-10 (11), the filaments hirsute with spreading hairs on outer surface, ca 0.2 mm long, the anther sacs ca 0.4 mm long. Pistillate cymules sessile; involucel of 1 abaxial bract and 2 adaxial bractlets, these puberulent near base, glabrescent elsewhere, the margins entire, ciliate, striate- veined; abaxial (proximal) bract simple, triangular, 3-4 mm long, 2-3 mm wide; adaxial (lateral, distal) bractlets simple, lanceolate, 1.5-2.3 mm long, 0.5-1 mm wide. Pistillate flowers 3, on pedicels 2.2-3.7 mm long, ca 1 mm thick at anthesis, strigulose-puberulent; sepals 6, narrowly lanceolate, 2.4-4 mm long, 0.5-0.8 mm wide, sparsely strigulose, the margins entire, ciliate; ovaries 1.1-1.5 mm high, 1.5-2 mm in diam, tomentose; stylar column more or less straight, subcylindrical, 7-9 mm long, 1.5-2.6 mm thick, thickest at ca 1/3 of length, gradually tapering to a blunt point distally, the tip with pit below center, the base of style puberulous, the stigmatic surface glabrous, covering distal 2/3 of stylar column. Pedicels in fruit 12-23 mm long; pistillate sepals persistent, similar to at anthesis. Capsules 7-8 mm in diam, strigulose; seeds subspherical, slightly flattened, ca 4 mm in diam, 3 mm thick, the surface slightly and irregularly pusticulate, mottled white and dark brown, the hilum round, ca 0.2 mm in diam.

TYPE: VENEZUELA. ESTADO BOLIVAR: Dtto. Roscio, in riparian forest ca 2 km S Santa Elena de Uairrn, 4~ 61"7'W, ca 860 m elev., 20 Dec 1987, W. S. Armbruster, S. Keller & 311. Matsuki 87-153 (I-IOLOTYPE: MO; ISOTYPES" ALA, MY, NY, VEN).

Dalechampia attenuistylus appears to have been collected only once, but may occur elsewhere in the riparian forests of extreme southeastern Venezuela and northern Brazil. Its epithet refers to the unusual shape of the stylar column, which is widest at about one-third its length and then tapers gradually to a blunt tip. All other members of the section to which it belongs (and, in fact, nearly all other members of the genus) have stylar columns that are widest near the tip or are cylindrical throughout.

Dalechampia attenuistylus appears to be related to other members of the section Rhopalostylis Pax & K. Hoffm. (Pax & Hoffmann, 1919). It is like other members of this apparently primitive section in that the distal involucral bract is small and stipule-like, the pistillate flowers have six unlobed sepals, and the staminate pleio- chasium has a more-or-less decussate organization and lacks the large secretory gland. From D. micrantha Poeppig of Amazonia and Guayana it differs in usually a tapered rather than clavate style and in only 10 staminate flowers in three fertile pleiochasial arms rather than 13 flowers in four arms. From D. olympiana Kuhlm. & W. Rodrigues of northern Amazonia it differs in a tapered rather than clavate style, sparsely strigulose rather than orange-lanate stems, petioles, and veins, and in pseudanthial inflorescences long-pedunculate rather than subsessile. From D.

1989] ARMBRUSTER" DALECHAMPIA 47

parvibracteata Lanj. of Guayana, perhaps its closest relative, it differs in yellowish- green instead of whitish stigmatic surfaces, in the tapered rather than clavate style, and in involucral bracts, both proximal and distal, being small and stipule-like.

Dalechampia megacarpa Armbruster, sp. nov. (Fig. 2)

Differt ab aliis speciebus generis caapsulis maturis 35-40 mm latis, seminibus 9-11 mm diametro, et bracteis involucralibus lanceolatis, 10-15 (20) m m longis, 4-6 mm latis, viridibus.

Clambering vine; stems twining, terete, 1-1.5 mm thick at growing tips, the young shoots densely strigose-tomentose with persistent, slightly curved, whitish hairs 0.1-0.4 mm long. Leaves alternate, simple; stipules spreading-reflexed, com- monly deciduous, lanceolate to ovate, 2.5-5 mm long, 1.2-1.5 (1.7) mm wide, striate-veined, the margins entire, ciliate, the abaxial surface sericeous, the adaxial sparsely strigose; petioles terete, (1.5) 3.5-10 cm long, 1-2.5 mm thick, pubescent as stem; stipels at base of blade 2-4, minute, ca 0.2 mm long, glandular; blades chartaceous, unlobed, ovate, (4) 7-27 cm long, 3-20 cm wide, the abaxial surface strigose with whitish hairs ca 0.3 mm long and tomentose with shorter brown hairs, the adaxial surface glabrous; base of leaf truncate to deeply cordate; apex acuminate; margins plane, sinuate with glandular processes ca 0.1 mm long; major veins 5 from base, strigose with whitish hairs, the secondaries ascending, the tertiaries scalariforrn. Pseudanthial inflorescences 1 to several on reduced axillary short-shoots, these at anthesis 1-3 cm long, pubescent as stem, with 1 or 2 nodes, each with leaf and axillary bud, often producing second pseudanthial inflorescence, or commonly leaf deciduous and only stipules and axillary bud persistent; distance between ultimate node and proximal involucral bract 5-10 mm at anthesis; su- pernumerary buds in leaf axils of long-shoot commonly developing into 1 or 2 additional inflorescence-bearing short-shoots, these smaller than first. Involucral bracts pale green at anthesis, subequal, unlobed, lanceolate or oblanceolate, 10- 15 (20) mm long, 4-6 mm wide, sparsely strigose below, glabrescent above, the base attenuate, the margins revolute, glandular-undulate, ciliate, the prominent veins 1 or 3 from base; stipules of proximal bract reflexed, ovate-falcate, 3-5 mm long, 1-2 mm wide, the abaxial face sparsely strigose, the adaxial face glabrescent, the apex subacute, the margin plane, entire, ciliate; stipules of distal bract similar but up to 2.5 mm wide. Staminate cymule with stout, densely strigose peduncle, 2-4.2 mm long, 1-1.8 mm thick; involucel of 2 pairs of more or less decussate, free bracts; bracts depressed-obovate, cupped adaxially, 2.6-3.2 mm long, 4.5- 5.5 mm wide, striate-veined, the margins weakly erose, minutely ciliate; staminate flowers 7-9, arranged in branched pleiochasium with 1 terminal flower and 3 arms, each arm 2- or 3-flowered, subtended by 2-4 strap-shaped or square, blunt bractlets, these 2-2.5 mm long, O. 1-2.5 mm wide, the apical margins laciniate, the laciniae 0.5-1 mm long, O. 1 mm thick, resiniferous, secreting a greenish-blue resin; bractlets of fourth (sterile) pleiochasial-arm 2 or 3, similar but up to 3.5 mm wide and rolled together. Staminate flowers on stout pedicels 1-4 mm long, ca 1 mm thick, articulate near top, sparsely puberulent; calyx sparsely hispidulous, splitting into 4--6 lanceolate-ovate, acute, spreading segments 1-2 mm long, 0.8- 0.9 mm wide; staminal column with only a few hairs near top, 1.5-2.1 mm long, 0.2-0.4 mm thick; stamens 8-22, the filaments hirsute with spreading hairs on outer surface, ca O. 1 mm long, the anther sacs 0.2-0.3 mm long. Pistillate cymules sessile; involucel of 1 abaxial bract and 2 adaxial bractlets, these sparsely puber- ulent, the margins erose, ciliate, striate-veined; abaxial (proximal) bract simple, depressed obovate, 1-2 mm long, 1.5-3 mm wide; adaxial (distal) bractlets simple, squarish, 1-2 mm long, 1-2 mm wide. Pistillate flowers 3, on short pedicels 0.3- 1 mm long at anthesis; sepals 6, puberulent, ovate, 1-1.5 mm long, 0.4-1.0 mm

48 BRITTONIA [VOL. 41

A cm

B 5 mm

J ~ D

t.---.J I I 5 mm 1 cm

FIG. 2. Dalechampia megacarpa. A. Habit. B. Pseudanthial inflorescence. C. Pistillate cymule. D. Fruit.

wide, the margins entire, ciliate; ovaries hirtellous, 0.7-1.1 mm high, 0.7-1.3 mm in diam; stylar column straight, cylindrical, of nearly equal diam throughout length, 4-6 mm long, 0.8-1 mm thick, hirtellous only at extreme base, the glabrous stigmatic surface covering nearly entire length, the tip blunt, undilated, foveolate. Pistillate pedicels in fruit 9-13 mm long; pistillate sepals triangular, 2-3 mm long, 1.6-1.8 mm wide, sparsely pubescent, ciliate. Capsules 35-40 mm in diam, with massive woody walls 3-6 mm thick, sparsely strigose; seeds flattened-oval, 9-11 mm in diam, 8-8.5 mm thick, the surface smooth, with sparse small hairs, dark brown with gray mottling, the hilum round, ca 2 mm in diam.

TYPE: VENEZUELA. ESTADO BOLIVAR: Dtto. Roscio, 3-7 km NW San Isidro (Krn 88), ca 100 m elev., 11 Dec 1985, W. S. Armbruster, A. L. Herzig & L. D. Otero 85-115 (HOLOTVPE: MO; ISOrVPES: ALA, MY, NY, VEIN).

Additional specimens examined: VENEZUELA. TEeOUTOlUO FEDERAL AMAZONAS: Dtto. Rio Negro, carretera San Carlos--Solano, 6-7 km S de Solano, 120 m elev., 7 Feb 1977, Morillo & Villa 5442

1989] ARMBRUSTER: DALECHAMPIA 49

(MY, VEN); 15 km NE San Carlos de Rio Negro, 1"56'N, 67~ 120 m elev., 16 May 1979, Liesner 7471 (VEN). ESTADO BOLIVAR: Dtto. Roscio, 6.5 km W of San Isidro (Krn 88) on mining road to Rio Cuyuni, 19 Dec 1987, Armbruster, Keller & Matsuld 87-151 (ALA).

Dalechampia megacarpa has been collected in only two places: near San Isidro, Bolivar, and near San Carlos, Amazonas; but it could be expected elsewhere in wet lowland forests o f northeastern Amazo n basin and the Guianas. This seeming disjunction o f over 700 km is probably due to inadequate collecting. It occurs as a climbing vine at the edge o f pr imary forest or its secondary margins.

The epithet megacarpa refers to the capsules which are nearly twice the diameter o f those of the species with the next largest fruit (D. pentaphylla Lam.) and some four t imes or more the diameter o f the capsules o f nearly all other species of Dalechampia. The seeds are also unusually large. The evolut ionary significance o f the large fruits and seeds remains obscure.

The four free involucellar bracts o f the s taminate cymule and the distinct resin gland composed of lacinia te resiniferous bractlets indicate placement ofD. mega- carpa in section Dioscoreifoliae both as circumscribed by Pax and Hoffmann (1919) and Armbrus te r (1988). It does not appear, however, to be closely related to any other species in the section. The greenish-blue resin, small involucral bracts, cylindrical styles, and simple, unlobed leaves suggest affinity with D. schottii Greenman o f the Yucatan Peninsula, Mexico. The entire, deltoid pistillate sepals (in fruit), large fruits with thick, woody cocci, and straight, cylindrical styles, however, suggest affinity with D. pentaphylla Lam. o f southern Brazil. The rela- tionship, however, is close with neither species, and the similarities may be only superficial.

Dalechampia papillistigma Armbruster , sp. nov. (Fig. 3)

Differt ab aliis speciebus generis foliis trifoliolatis, bractcis involucralibus spathulatis, parvis, 5-8 mm longis, stylis capitatis, et stigmatibus papillatis.

Clambering vine; stems twining, terete, ca 1 m m thick at growing tips, strigose with persistent, yellowish to whitish, incur red hairs 0.1-0.5 m m long. Leaves alternate, palmately trifoliolate (rarely 4- or 5-foliolate), with 5 major veins from base (1 in central leaflet, 2 each in lateral leaflets), these sparsely strigose as stem abaxially, densely tomentose-strigose with yellow hairs adaxially, the secondary veins ascending, the tertiaries scalariform; stipules spreading reflexed, persistent or deciduous, lanceolate, 3-7 m m long, 1-2 m m wide, the abaxial surface sparsely strigose, the adaxial surface finely and sparsely puberulent, striate-veined, with several glands at a t tachment o f stipule to stem; petioles terete, 2 -10 cm long, 0.9- 2.3 m m thick, strigose as stem, except near a t tachment o f leaflets where tomentose on adaxial surface; petiolules terete, 2.5-6.5 m m long, 0.8-2.1 m m thick, strigose as stem abaxially, densely tomentose with yellow hairs adaxially; stipels at base o f petiolules 2, lance-linear, 2-3 m m long, ca 0.3 m m wide, with a few glands at point o f a t tachment to petiolule; blade o f central leaflet chartaceous, oblanceolate to obovate, 4-15 cm long, 2-7 cm wide, glabrescent on both surfaces, the base cuneate, the apex acuminate, the margins sinuate to shallowly serrate, ciliate, with glandular processes ca 0.1 m m long; lateral leaflets similar but lanceolate to ovate, the base asymmetric. Pseudanthiat inflorescences solitary on reduced axillary short- shoots 2-3 cm long at anthesis, these puberulent to strigose, with 1 or 2 nodes, each represented by a pair o f stipules; distance between ult imate node and prox- imal involucral bract 15-20 m m at anthesis. Involucral bracts green at anthesis, subequal, slightly concave adaxially, unlobed, spathulate with distinct l imb and claw, 5-8 m m long, 3-4 m m wide, the l imb 3-4 m m long, 3-4 m m wide, cuneate at base, the claw 2-4 m m long, 1-2 m m wide; surface o f involucral bracts pu-

50 ]}gITTONtA [VOL. 41

A 11 c m ~

B J

~ mm

t m J 5 m m

E F

5 m m

F~G. 3. Dalechampia papillistigma. A. Habit. B. Pseudanthial inflorescence, front view. C. Pseud- anthial inflorescence, lateral view. D. Staminate pleiochasium. E. Pistillate cymule, abaxial view. F. Pistillate cymule, adaxial view. G. Fruit.

berulent; margins o f claw plane, ciliate, the margin o f l imb plane, with gland- t ipped laciniae along basal half, erose along distal half; bracts more or less striate- ve ined with 1-3 indistinct m a i n veins f rom base; stipules o f proximal (pistillate) involucral bract triangular-falcate, 1.5-2.8 (3.6) m m long, 1-2 (2.4) m m wide,

1989] ARMBRUSTER: DALECHAMPIA 51

with both faces obscurely puberulent, the margins entire, sparsely ciliate, the apex subacute, striate-veined; stipules of distal (staminate) involucral bracts similar to those of proximal but larger, 2-3 (3.8) mm long, 1.5-2.2 mm wide, and tip rounded. Staminate cymule with short, stout, puberulent peduncle 0.6-2.0 (2.8) mm long, ca 2 mm thick; involucel of 2 pairs of more or less decussate, free bracts, these depressed obovate, 3.2-5.4 mm long, 4.9-8.0 mm wide, puberulent, striate-veined, the margins erose, sparsely ciliate; staminate flowers 10, arranged in branched pleiochasium with 1 terminal flower and 3 arms, the fourth arm sterile; each fertile arm 3-flowered, subtended by 3-6 strap-shaped, rectangular to obtriangular bractlets, 3-5 mm long, 0.5-3.5 mm wide, the margins at tip with terete, glandular, resiniferous laciniae 1-2 mm long, 0.1-0.2 mm thick; bractlets of sterile (dorsal) pleiochasial arm similar, but 4-6 arranged concentrically, the largest wrapping from adaxial side, the smallest to the center on the abaxial side. Staminate flowers on short stout pedicels 1-3 mm long, 1-1.5 mm thick, these articulate near top, puberulent; calyx triangular, subacute, the spreading segments 1.6-2.8 mm long, 0.5-3.5 mm wide; staminal column short and stout, 0.5-1 mm long, 1-2.5 mm thick, glabrous; stamens (60) 70-80 (95), the filaments ca 0.5 mm long, with a few short hairs, the anther sacs 0.2-0.35 mm long. Pistillate cymules sessile; involucel of 1 bract, this depressed obovate, 2-4 mm long, 3.5-4.5 mm wide, puberulent, striate-veined, the margin erose to dentate, sparsely ciliate. Pistillate flowers 3, on pedicels 0-0.6 mm long at anthesis; sepals 6 (5-7), 2-4 mm long, ca l mm wide, deeply divided longitudinally into 2-4 lobes, or rarely unlobed, the margins entire, sparsely ciliate; ovaries 1-1.5 mm high, 1-2 mm in diam, finely strigulose; stylar column puberulent, more or less straight, cylindrical with distinct capitate tip, 2.7-3.3 mm long, 0.5-1 mm thick, the tip ca 2 mm in diam, shaped like a swollen annulus, with deep central pit on ventral side of end, the stigmatic surface densely papillate, confined to swollen tip of style. Pistillate pedicels in fruit 2-12 mm long; pistillate sepals usually 2-4-lobed, 3-4 mm long, 1-2 mm wide, hispidulous. Capsules 8-9 mm in diam, puberulent, trigonous, each lobe projecting up 2-3 mm beyond the attachment of the style; seeds sphe- roidal, 3-4 mm in diam, the surface slightly roughened, with 3 or 4 faint ridges on the side opposite the hilum, brown with white mottling, the hilum triangular, 0.8-1 mm long.

TYPE: VENEZUELA. ESTADO BOLIVAR: Dtto. Roscio, 20.1 km SW of Santa Elena de Uair6n, 14.1 km W of junction with road to Icabarfi, 4~ 61 ~ 15'W, ca 930 m elev., 21 Dec 1987, W. S. Armbruster, S. Keller & M. Matsuki 87-155 (HOLOTYPE: MO; ISOTYPES: ALA, MY, NY, VEN).

Additional specimens examined: VENEZUELA. ESTADO BOLIVAR: Dtto. Roscio, alredadores de Santa Elena de Uair6n, 25 Apr 1944, Lasser 1454 (VEN); Dtto. Roscio, 5.2 km S Santa Elena de Uair6n, 4"37'N, 61~ ca 860 m elev., 21 Dec 1987, Armbruster, Keller & Matsuki 87-154 (ALA).

Dalechampia papillistigma has been collected only in the vicinity of Santa Elena de Uair6n, in extreme southeastern Bolivar, Venezuela, where it occurs in and at the edge of riparian forests in savanna.

Dalechampia papillistigma is distinct from all other congeneric species that I have examined in having a papillate rather than smooth stigmatic surface. The free staminate involucellar bractlets, and laciniate, resiniferous bractlets united into a distinct resin gland indicate that it belongs in the section Dioscoreifoliae as circumscribed by Armbruster (1988), but would be placed in section Triphyllae in the artificial treatment of Pax and Hoffmann (1919). Within section Diosco- reifoliae, D. papillistigma appears to be quite isolated. It differs from other mem- bers of the section in having relatively small spathulate involucral bracts, pal- mately 2-4-lobed pistillate sepals, and an annulus-like, papillate stigmatic tip.

52 BRITTONIA [VOL. 41

Acknowledgments

I thank A. L. Herzig, S. Keller, M. Matsuki, and L. D. Otero for help in the field, S. Keller for measuring specimens, K. Holmes for Figure 2, M. Matsuki for Figures 1 and 3, and four reviewers for their helpful comments . This work was suppor ted by the National Science Foundat ion (Grant No. BSR-850931).

Literature Cited

Armbruster, W.S. 1988. A new species and synopsis ofDalechampia (Euphorbiaceae) from Costa Rica. Syst. Bot. 13: 303-312.

Pax, F. & K. Hoffmann. 1919. Euphorbiaceae--Dalechampieae. Pages 1-59. In: A. Engler, editor. Das Pflanzenreich IV. 147. XII (Heft 68). Leipzig, Verlag von Wilhelm Engelmann.

B O O K REVIEW

Passage of Darkness. The Ethnobiology of the Hai t i an Zombie. By Wade Davis. Univers i ty o f Nor th Carol ina Press, P.O. Box 2288, Chapel Hill, NC 27514. ISBN 0-8078-1776-7 (cloth), 0-8078-4210-9 (paper). 1988. 344 pp. $29.95 (cloth), $9.95 (paper).

This book is an outstanding example o f an ethnobiological investigation focused on a mos t interesting subject, the Hait ian zombie. Davis, an anthropologist and biologist by training, immerses h imsel f in the social and cultural setting necessary to carry out his research. The reader is partner to Davis ' scientific research as he embarks on what turns out to be a remarkable journey into the natural and supernatural world of the zombie. One br ief anecdote deserves ment ion here, and will speak to those who carry out fieldwork on the international scene. When Davis asks one o f his informants to bring him a zombie, the man returns with a clay jar filled with a zombie "spir i t ." When asked why he did not bring a real zombie " . . . with complete sincerity he [the informant] asked how on earth I would get a zombie o f the flesh through U.S. Customs." Unfortunately, this project has become somewhat controversial , and the resulting debate has tended to ob- scure the fact that this work is one o f the more interesting, comprehensive, and thought-provoking in recent times. It serves as an inspirat ion to all ethnobiologists, and shows why practitioners in this field must address their research topics with the broadest possible view and with a complete understanding o f the perceptions o f the people under study. The biologist looking for long lists o f plants and animals will be disappointed. This is not the point o f Davis ' book. Rather, he offers a most remarkable and well documented vista into the ethnobiology of the Hai t ian zombie phenomenon.--MICHAEL J. BALICK, New York Botanical Garden.