notes on oenocarpus (palmae) in the colombian amazon

Brittonia, 43(3), 1991, pp. 154-164. �9 1991, by the New York Botanical Garden, Bronx, NY 10458-5126

N O T E S O N O E N O C A R P U S ( P A L M A E ) I N T H E C O L O M B I A N A M A Z O N

RODRIGO G. BERNAL, GLORIA GALEANO, AND ANDREW HENDERSON

Bernal, Rodrigo G., Gloria Galeano (Instituto de Ciencias Naturales, Univer- sidad Nacional de Colombia, Apartado 7495, Bogota, Colombia) and Andrew Henderson (New York Botanical Garden, Bronx, NY 10458-5126). Notes on Oenocarpus (Palmae) in the Colombian Amazon. Brittonia 43: 154-164. 1991- Five species of Oenocarpus from the Rio Caquet~, in the Colombian Amazon, are discussed. Two of them, Oenocarpus simplex and Oenocarpus makeru are described as new. Oenocarpus bacaba var. parvus is shown to be a synonym of O. balickii; this species and O. minor are recorded for the first time in Colombia. Some comments on the poorly known O. circumtextus are also given. The new findings support the inclusion of Jessenia in Oenocarpus.

Se discuten cinco especies de Oenocarpus de la regi6n del rio Caquetfi, en la Amazonia Colombiana. Dos de elias, Oenocarpns simplex y Oenocarpus makeru se describen como nuevas. Oenocarpus bacaba var. parvus es considerada un sin6nimo de O. balicldL y esta especie, junto con O. minor, es registrada por primera vez en Colombia. Se da informaci6n adicional sobre O. circumtextus, una especie muy poco conocida. Los nuevos hallazgos respaldan la uni6n de Jessenia y Oenocarpus. Key words: Palmae, Oenocarpus, Jessenia, Colombian Amazon.

Nine species were recognized by Baliek (1986) in the most recent revision o f the Jessenia-Oenocarpus complex. Two new varieties o f Oenocarpus bacaba C. Martius f rom Venezuela were later added by Wessels Boer (1988), and a new species o f Oenocarpus C. Martius from Peru was described by Kahn (1990). Four species were reported by Balick to occur in Colombia: Jessenia bataua (C. Martius) Burret, Oenocarpus circumtextus C. Martius, Oenocarpus bacaba, and O. mapora Karsten, the last with two subspecies. As a result o f Balick's revision there has been a revival o f interest in the group, particularly in its economic potential. Consequently there are abundant new collections. Recent exploration o f the area o f La Pedrera, on the Rio Caquefft in the Colombian Amazon, in particular has revealed the existence o f at least seven species in this group in that region: Jessenia bataua, Oenocarpus bacaba, O. bah'ckii Kahn, O. minor C. Martins, O. circumtextus, O. makeru, and O. simplex, the latter two described here as new. Oeno- carpus mapora Karsten, a c o m m o n species throughout the northwestern Amazon , was not seen during the short visit to this area; nevertheless, it has been collected upstream along the Rio Caquetfi (Galeano, 1991), and its occurrence in the La Pedrera area seems likely. In any case, the presence o f seven species within such a restricted area represents the highest diversity for this group known so far.

La Pedrera is located on the lower Rio Caquelfi in Colombia, some 30 km from the Brazilian border. The region is most ly covered with tropical wet forest ( IGAC, 1977) usually developed on alluvial, acidic soils. Interspersed are rather large areas o f rocky outcrops o f granite or quartzite, marked by a peculiar flora that is physiognomical ly very different f rom that o f the surrounding forest. The physi- ognomy o f the flora is actually very similar to that o f the mounta ins and savannas o f the Guayana Highland (Huber, 1989). The area is poorly known botanically.

C o m m e n t s on some o f the species are made below, including description o f the new taxa.

1. Oenocarpus s implex Bernal, Galeano, & Henderson, sp. nov. (Fig. 1)

TYPE: COLOMBIA. Amazonas: ca 2 km along the trail f rom La Pedrera to Tarapac~, 230 m, 10 Mar 1990, G. Galeano, R. Bernal, A. Henderson, N. Espejo, & S. Churchill 2027 (HOLOTYPE: COL!; ISOTYPES: AAU!, BH!, K!, NY!).

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156 BRITTONIA [VOL. 43

Foliis s'implicibus spadicibusque simplicibus velfurcatis valde distincta.

Stems cespitose, 3-4 m tall, 1.5-1.8 cm diam, with 2 or 3 main stems and several basal shoots, smooth, brown, conspicuously ringed, the internodes 1.5-3 cm long. Leaves 5-8, reddish when young, erect; sheath 41-45 cm long, purplish to green, fibrous on the margins, with purplish scales; petiole 24-28 cm long, 5 mm diam, subterete, with deciduous scales like those of the leaf sheath; rachis 63-67 cm long; blade entire, narrowly cuneate, apically bifid for ca 1/~ its length, 76-91 cm long, 1 4-18.6 cm at the widest point shghtly below the bifurcation, the lobes ovate, cucuUate, 15.5-16.5 cm long, including a narrow acumen, 3-4 cm long; primary veins 8-11 on each side, forming an angle of 6-8* with the rachis, abaxial surface whitish, waxy, with scattered, very short, white trichomes. Inflo- rescence interfoliar, erect at anthesis, pendulous in fruit, spicate or bifid, up to 73 cm long; prophyll bicarinate, acute at apex, chartaceous, 15-20 cm long, ca 1 cm wide, with scattered, purplish scales; peduncular bract cylindrical in the bud, expanded and fiat at anthesis, 48-66 cm long, including a 3-4 cm long umbo, 1.8 cm wide, subcoriaceous, striate, with purplish scales; peduncle 16-37 cm long, 2-3 mm thick, with ferrugineous scales, occasionally bifid at apex for 10-19 cm, and then the inflorescence with 2 pedunculate rachillae; rachilla 21-25 cm long, 3-5 mm in diam, with deep pits, yellowish at anthesis, reddish in fruit, with ferrugineous scales; staminate flowers ovate-lanceolate, 3-4.5 x 1.5 mm; sepals connate at base, sometimes connate into a flattened, triangular calyx with concave sides, or into a strongly 3-carinate calyx, the lobes triangular, 0.8 x 0.5 mm; petals ovate-lanceolate, thick, 3 x 1.2-1.5 mm; stamens 6, the filaments undulate, inflexed at the apex, 1-1.5 mm long, anthers 1-1.5 mm long, bifid and inequilateral at base, the connective projected into a short point; pistillode 0.5 mm long; pistillate flowers ovoid when immature, ca 2.5 x 2 mm; sepals broadly imbricate, suborbicular, 2.5 mm high; petals broadly imbricate, suborbicular, cucullate, 1.5 mm high; gynoecium 1 mm long; fruit oblong-ellipsoid, 2.2-2.7 x 1.3-1.4 cm, with a prominent, cylindrical, slightly eccentric stigmatic residue, the epicarp purplish-black at maturity, with a thin layer of wax, the mesocarp with fiat fibers; seed ovoid, 1.5 x 0.9 cm, the raphe branches loosely reticulate; endosperm homogeneous; eophyll in outline cuneate to obovate, bifid for 1/4 its length, the lobes long-acuminate and sometimes cucullate, whitish abaxially.

Oenocarpus simplex is quite unlike any other species in the genus because of its small, undivided leaves and simple or bifid inflorescences, as well as its slender, canelike habit. However, its membership in Oenocarpus is unquestionable because of its leaves glaucous below, rachillae with deep depressions and becoming ferrugineous after anthesis, sinuous filaments, and ellipsoid fruits with apical stigmatic residue (Table I). The long peduncle and persistent peduncular bract, as well as the slightly bifid eophylls, and the shape of the staminate sepals place this species in subgenus Oenocarpopsis (Burret) Balick, although its stem does not have the fibrous vestiture of leaf sheaths that is typical of O. circumtextus. The two known species in this subgenus (O. circumtextus and O. simplex) are re- markably different from one another, and from all species placed in subgenus Oenocarpus. In their small size, their leaves simple or with few broad pinnae, and their long pedunculate inflorescences, O. circumtextus and O. simplex evoke species of Prestoea, although, as pointed out above, their obvious relationships are with Oenocarpus. Also, the bifid eophylls are much more like those of Prestoea; species of subgenus Oenocarpus have eophylls with usually four pinnae radiating from a short rachis (Fig. 2a).

Oenocarpus simplex is a common palm in the area behind La Pedrera, where

1 9 9 1 ] BERNAL ET AL.: OENOCARPUS 157

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it grows in the unders tory o f ma tu re forest. I t was also observed in ma tu re forest near the m o u t h o f the Rio Miriti-paranfi. Al though s~veral individuals were seen, none o f t h e m had pinnate leaves, which suggests that s imple leaves are a constant character in this species.

2. O e n o c a r p u s m a k e r u Bemal , Galeano & Henderson , sp. nov. (Fig. 2)

TYPE: C O L O M B I A . Amazonas : Rio Caquetfi, near the Chorro C6rdoba , trail to the savanna, ca 250 m, 13 Mar 1990, G. Galeano, R. Bernal, A. Henderson, N. Espejo, & S. Churchill 2070 (HOLOT~E: COL!; ISOTYPES: AAU[, BH[, HUA! , K~, NYD.

1991] BERNAL ET AL.: OENOCARPUS 159

Palma solitaria. Pinnae eodem piano insertae, subtus indumento tenui laxo cereo laniformi obsitae. Fructus ovoideus. Seminis endospermum ruminatum.

Stem solitary, 5-8 m tall, 7-8 cm diam, grayish, longitudinally fissured. Leaves 12; sheath 65-68 cm long, purplish, coarsely fibrous; petiole 15-25 cm long, covered with a gray, scaly indumentum, the latter more or less persistent adaxially, more or less deciduous abaxially; rachis 2.43 m long, with scaly indumentum like that of the petiole; pinnae 65 on each side, regularly arranged and in 1 plane, linear-lanceolate, acuminate, strongly plicate along secondary veins, concolorous, glabrous adaxially, covered abaxially with a thin, loose, wool-like, waxy indumentum, and sparsely beset with minute, white trichomes; basal pinnae 46.5- 50.4 x 0.8-1.5 cm; middle pinnae 73-75.5 x 5-5.3 cm; apical pinnae 29 • 1 cm. Inflorescence interfoliar; prophyll not seen; peduncular bract inserted 2.5-3 cm above prophyll, covered (at least in young bud) with purplish scales; peduncle 6.2-7.5 cm long, the distance between scars ofprophyll and peduncular bract 2.8- 3 cm, the axis 9.5-16 cm long from the scar of peduncular bract (i.e., distal part of peduncle + rachis), circular in transverse section; rachiUae 107-125, the longest ones 56-60 cm; flowers seen only in early bud; staminate flowers with sepals connate at base, the lobes triangular, carinate; petals free; stamens 6; pistillate flowers not seen; fruits ovoid, with apical stigmatic residue, 2.2-2.6 • 1.4-1.7 cm, the epicarp smooth, purplish-black at maturity, the fibers ca 0.2 mm wide; seed slightly ovoid to almost ellipsoid, 1.8-1.9 • 1.4-1.5 cm; endosperm white, with abundant brown ruminations; embryo well-developed, to ca 1.4 cm long; eophyll with 4 narrow pinnae palmately radiating from a very short rachis.

Common name: makeru (Yukuna). Oenocarpus makeru can be distinguished from all other species in the genus

through its ruminate endosperm. It can be recognized also by the combination of a solitary stem, regularly arranged pinnae with an abaxial thin, wool-like, waxy indumentum, and ovoid fruits.

Oenocarpus makeru is known only from a population at the type locality where it is an abundant palm growing in the forest bordering an area ofcaatinga. Although the dried specimen available for study lacks developed flowers, we have described it as a new species since it is quite different from any other taxon in the group notably through its ruminate endosperm. Within the Jessenia-Oenocarpus complex ruminate endosperm is known only in Jessenia bataua; all hitherto known species of Oenocarpus (sens. strict.) have homogeneous endosperm. In fact, the nature of the endosperm has been used by Balick (1986) as one of the main features distinguishing the genera. Since in all other respects the described plant is a typical Oenocarpus (sens. strict.), it might be thought that this population represents a hybrid between Jessenia and Oenocarpus. Although hybrids have been reported to occur in this complex (Balick, 1988), this does not seem to be the case for this species as there is no morphological character in this population that appears to be of hybrid nature. Indumentum on the undersurface of pinnae lacks the typical sickle-shaped scales characteristic of Jessenia bataua; eophylls are not bifid as in Jessenia, but they have four slender pinnae radiating from a very short rachis, as in all species of Oenocarpus subgenus Oenocarpus; and the number of stamens is six, typical ofOenocarpus (sens. strict.). Furthermore, known hybrids involving Jessenia bataua have a reduced endosperm and a small embryo (M. Balick, pers. comm.), whereas O. makeru has a large, well-developed embryo within the ruminate endosperm. Seedlings were abundant under the palm from which the specimens were collected.

The finding of Oenocarpus makeru is of great significance in understanding the relationships between Jessenia and Oenocarpus. Species in this group were initially


placed in one genus by Martins (1837), who first described Oenocarpus, and named five of the species currently recognized in the complex. Later, Karsten (1857) established the genus Jessenia, and described J. polycarpa, which he compared with Oenocarpus, particularly O. bataua C. Martins. He distinguished Jessenia polycarpa through the larger number of stamens, the apical (vs eccentric) stigmatic residue, and the trilobate (vs tripartite) calyx of the male flowers. Two species of Jessenia were added by later workers in the nineteenth century (Grisebach, 1864; Engel, 1865). Burrer (1928) reappraised the group and accepted two genera, trans- ferring Martius' O. bataua to Jessenia. Thus, Buffet was the first author to clearly contrast both genera, which he distinguished in a few characters, viz the endosperm, the vestiture of the undersurface of pinnae, the number of stamens, and the projection of the connective beyond the anther.

Recognition of two genera in this complex was questioned by Wessels Boer (1965, 1972, 1988), but he was not followed by other students of palms. Balick (198 6), in a monographic study of these genera, continued to keep them separate, mostly following the same criteria used by Burrer, and also including some in- formation on the chromatographic patterns offlavonoids. An additional difference used were eophylls, which are bifid in Jessenia bataua, and have four pinnae radiating from a short rachis in all species of Oenocarpus for which seedlings were known prior to this study. In Table I, morphological characters of Jessenia, subgenus Oenocarpus, and the species O. simplex, O. circumtextus, and O. makeru are contrasted.

Data now available from these two new species as well as observations on Oenocarpus circumtextus, show that some of the character states used to define Jessenia also occur in species of Oenocarpus, thus obscuring the distinction. Ru- minate endosperm occurs in Oenocarpus makeru; bifid eophyUs are now known to characterize subgenus Oenocarpopsis; and undulate filaments and a produced connective occur in Oenocarpus simplex. However, it should be noted that the nature of the connective is not consistent. In both O. simplex and O. circumtextus individual flowers can contain anthers with and without a projected connective. The remaining differences, while fully meaningful at the specific level, could scarcely be used to separate two genera. The number of stamens is variable in many genera throughout the family, and even in genera with mostly six-staminate flowers, there occur species with different number of stamens, e.g., Geonoma triandra (Burret) Wessels Boer (Wessels Boer, 1968), Astrocaryum triandrum Gale- ano, Bernal and Kahn (Galeano-Garcrs et al., 1988). Finally, the difference between "wiry" and "knitting needle-like" fibers or between apically inflexed and apically curved filaments, and the shape of trichomes do not seem to justify distinction at the generic level. In conclusion, we consider that Jessenia bataua, the only species currently recognized in the genus, must be treated under Oeno- carpus, as originally postulated by Martius (1837).

3. OENOCARPUS MINOR C. Martius

Oenocarpus minor C. Martius, Hist. Nat. Palm. 2: 25-26, t. 27. 1823. TYPE: BRAZIL. Amazonas: Manaus, [1819-1820], C. Martius 3121b (rtoLoa'~E: M!).

Specimens examined: COLOMBIA. AmaTonas: Rio Caquetfi, near Chorro C6rdoba, ca 250 m, 12 Mar 1990, G. Galeano et al. 2051 (COL, NY); Rio Miriti-paran~, near the mouth, ca 200 m, 14 Mar 1990, G. Galeano et al. 2081 (COL, NY).

This species was described from the Manaus area in the Brazilian Amazon and it has been subsequently reported (Balick, 1986) from several localities in the northwestern Amazon of Brazil. Literature records from the Amazon basin in

199 1 ] BERNAL ET AL: OENOCARPUS 161

neighboring countries were considered by Balick (1986) as likely to have been based on misinterpretation or confusion with related species.

In the area of La Pedrera, O. minor was found at two localities, one on the Rio Caquefft, and the other one near the mouth of Rio Miriti-paran~i. At both places it grows in well-drained soils in dense forest.

Balick (1986) recognized two subspecies of O. minor, subsp, minor, and subsp. intermedius. He distinguished them mainly in the solitary (minor) versus cespitose (intermedius) habit and in the number of rachillae (25-35 in minor vs 54-72 in intermedius). Our field observations in the region of La Pedrera revealed that both solitary and cespitose palms can be found in the same area and a correlation between habit and number of rachillae could not be found. Palms with cespitose stems had about 32-35 rachillae in the individuals we collected, suggesting that these characters cannot be used to separate subspecies.

Oenocarpus minor itself seems to be very closely related to O. mapora Karsten and there is no qualitative character that can be used to distinguish them. Balick separated them mainly on the size of the inflorescence axis and on the arrangement of the leaflets on the rachis but he himself described variation in the latter character thus leaving inflorescence axis as the only reliable character for recognizing both taxa. Abundant collections from the Amazon basin made after Balick's monograph was published, including the Colombian collections here reported, show that the length of the inflorescence axis is very variable in both species and should not be considered a reliable distinguishing character. However, it is usually possible to distinguish both entities in the field, mainly by their overall size, O. minor nor- mally being a smaller palm with a slender stem, and its inflorescence with a smaller number ofrachillae. It is probably significant that the areas of distribution of both species scarcely overlap. We do not discard the possibility that O. minor is just a reduced form of O. mapora and that they could be better treated as a single species.

4. OENOCARPUS BALICKII Kahn

Oenocarpus balickii Kahn, Candollea 45(1): 351. 1990. TYPE: PERU. Loreto: Prov. Requena, Bajo Ucayali, Jenaro Hen'era, 25 Jul 1989, F. Kahn et al. 2380 (HOLOTYPE: USM--n.v.; ISOTYPES: AAU, G, K, NY, P--n.y.).

Oenocarpus bacaba var. parvus W. Boer, Pittieria 17: 130. 1988. TYPE: VENEZUELA. Amazoaas: near San Carlos de Rio Negro, J. ;Vessels Boer 2276 (HOLOTYPE: U).

Specimens examined: COLOMmA. Amazonas: La Pedrera, ca 2 km on the road to Tarapacfi, 230 m, 10 Mar 1990, G. Galeano et al. 2026 (COL, HUA, NY); 11 Mar 1990, G. Galeano et aL 2033 (COL, HUA, NY); Leticia, 7 km N on the road to Tarapac~, 17 Mar 1990, G. Galeano et al. 2117 (COL, HUA, NY). V~V.Z~LA. Amazonas: brazo Casiquiare, near Solano, 8 Feb 1969, J. Wessels Boer 2408 (NY, paratype of Oenocarpus bacaba var. parvus). PERU. Loreto: Prov. Requena, Bajo Ucayali, Jenaro Herrera, 130 m, Sep 1984, F. Kahn & K. Mejfa 1723 (NY, paratype ofOenocarpus balickiO; 25 Jun 1989, F. Kahn et al. 2381 (COL, paratype of Oenocarpus balickiO; 14 Sep 1987, R. V~squez & N. Jaramillo 9579 (NY). BRAZlL. Acre: Mun. Mancio Lima, Serra do Moa, near the Rio Moa, ca 350 m, 12 Oct 1989, .4. Henderson et al. 1118 (NY); Mun. Mancio Lima, road from Cruzeiro do Sul to Mancio Lima, AC 317, 19 Oct 1989, .4. Henderson et al. 1147 (NY). Amazonas: Mun. Atalaia do Norte, Rio Javari, Paumari (or Natavidad), 4~ 70~ 5 Jan 1989, .4. Henderson et al. 834 (NY). Rondonia: Mun. Porto Velho, along BR 364, 6.5 km ENE of C6rrego Raiz and 27 km ENE of junction with BR 325, 27 km from Abtma, 9~ 'S, 65~ 140 m, 17 Apr 1987, M. Nee 34886 (NY).

This taxon was first recognized as an entity different from typical Oenocarpus bacaba by Wessels Boer (1972), although the name O. bacaba var. parvus was validated later (Wessels Boer, 1988). Wessels Boer distinguished it from O. bacaba var. bacaba through the smaller size of all its parts, especially the inflorescence. More recently, Kahn (1990) described Peruvian specimens as a new species,


Fios. 3 & 4. Oenocarpus circumtextus. 3. Habit at La Pedrera. 4. Inflorescence and fibrous leaf sheaths on stem.

Oenocarpus balickii. Study o f para types o f O. bacaba var. parvus and o f O. balickiL as well as the protologues and collections f rom Co lombia and Brazil reveal that they all belong to the same taxon, closely related to O. bacaba. Besides the obv ious difference in size, bo th taxa can be readily dist inguished in the field through the a r rangement o f p innae which are most ly pendulous in O. bacaba and radiate in m a n y directions in O. balickii. This character gives O. balickii the overal l appearance o f some species o f Syagrus C. Martius. Because o f these differences we are following K a l m (1990) in recognizing this t axon as a distinct species for which the n a m e O. balickii is the correct one.

Oenocarpus balickii is a c o m m o n species in the forests behind La Pedrera where it grows together with O. simplex. I t was also found to be c o m m o n near Leticia and has been collected often in other areas in Venezuela, Peru, and Brazil, which suggests that it is widespread throughout the Amazon .

5. OENOCARFUS CmCUMTEXTUS C. Mart ins (Figs. 3 & 4)

Oenocarpus circumtextus C. Martius, Hist. Nat. Palm. 2: 26, t. 26, figs. 3, 4. 1823. TYPE: COLOM- BIA. Amazonas: Cerro de la Pedrera, [1820], C. Martius s.n. (HOLOa~'E: M!).

Specimens examined: COLOMaIA. Amazonas: La Pedrera, Rio Caquet~, Cerro Yupati, 300 m, 8 Mar 1990, G. Galeano et al. 1974 (AAU, BH, COL, HUA, K, NY); La Pedrera, Rio CaquetA, opposite Cerro Yupati, 9 Mar 1990, G. Galeano et al. 1997 (AAU, BH, COL, HUA, K, NY).

This species was descr ibed f rom the Cerro de La Pedrera, also known as Cupat i or Yupat i , where it was first collected in 1820. Only two further collections seem to have been made , bo th at the same locality (Balick, 1986). The new collections f r o m the area o f L a Pedrera, as well as observa t ions o f the large populat ions, allow for a be t ter unders tanding o f this species and some new data can be added to Balick 's (1986) description. Oenocarpus c ircumtextus is a solitary pa lm, 3-6 m

1991] BERNAL ET AL.: OENOCARPUS 163

tall, with the t runk complete ly covered by a dense net o f in te rwoven fibers f rom the leaf sheath. The crown has 7 or 8 leaves with a petiole 72-75 cm long; the rachis is 175-180 c m long, and has 16-19 pinnae on each side. In florescences are interfoliar and erect, except for the pendulous rachillae. Fruit ing rachillae are reddish. EophyUs are cuneate, bifid at apex for slightly m o r e than ~ their length.

Because the few known specimens all came f rom the same small cerro, some authors considered this species to be endemic to Cerro Yupat i (Huber, 1910). However , we found it also in an area o f caatinga across the Rio Caquetfi, and according to local informants , the species is found on several rocky outcrops in the region. These consist o f granite or quartzi te r emnan t s o f the G u a y a n a Shield ( IGAC, 1977). Soils are very shallow and der ived f rom the parent material , but there is a thick h u m u s layer present. This physical rel ief gives the vegetat ion o f these cerros a comple te ly different appearance f rom the surrounding forest. A sclerophyllous, low shrubby vegeta t ion occurs, se ldom reaching more than 8 m tall. The m o s t abundan t families are Lauraceae, Melas tomataceae , Clusiaceae, Myrsinaceae, Apocynaceae , and Rubiaceae. Epiphytes are very c o m m o n , particularly orchids, bromel iads , and aroids, and the ground layer is m a d e up o f abundant lichens, mosses , sedges, and ferns. In some parts o f these Cerros O. circumtextus is the d o m i n a n t plant, fo rming dense stands.

Acknowledgments

We are very grateful to Biologist Ale jandro Santacruz ( I N D E R E N A ) for his valuable help and hospital i ty during our visit to La Pedrera; we also thank our guide Jos6 Y6pez Matap i for his assistance and invaluable knowledge. Henderson gratefully acknowledges the suppor t o f the Nat iona l Science Founda t ion (Grant #BSR-8822201). Fie ldwork in C o l o m b i a was suppor ted in par t by a grant to Michael Balick f rom the Uni ted States Agency for In ternat ional Deve lopment . We are very grateful for this support , and thank Michael Balick, Henr ik Balslev, John Dransfield and Natal ie Uhl for thei r review o f the manusc r ip t and Silvio Ferngndez and T o n y Salazar for their excellent illustrations.

Li tera ture Cited

Balick, M.J. 1986. Systematics and economic botany of the Oenocarpus-Jessenia (Palmae) Complex. Advances Econ. Bot. 3: 1-140.

1988. Jessenia and Oenocarpus: neotropical oil palms worthy of domestication. FAO Plant Production and Protection Paper 88. Rome.

Burrer, M. 1928. Die Palmengattungen Oenocarpus Mart. und Jessenia Karst., nebst Bemerkungen zu Archontophoenix H. Wendl. et Drude (einscliesslich Loroma O. F. Cook). Notizbl. Bot. Gart. Berlin-Dahlem 10: 291-312.

Engel, F. 1865. Palmae novae eolumbianae. Linnaea 33: 665-692. Galeano, G. 1991. Las palmas de la regi6n de Araraeuara Estudios en la Amazonia Colombiana.

Vol. 1. Tropenbos, Bogotfi, Colombia. Galeano-Garc6s, G., R. G. Bemal & F. Kahn_ 1988. Una nueva especie de Astrocaryum (Palmae)

de Colombia. Candollea 43(1): 271-283. Grisebach, A. 1864. Flora of the British West Indian Islands. London. Huber, J. 1910. Sobre urea collec~o de plantas da regi~o de Cupaty (rio Japurfi-Caquet~). Bol. Mus.

Paraense Hist. Nat. 7: 283-307. Huber, O. 1989. Shrublands of the Venezuelan Guayana. Pages 271-285. In: L. B. Holm-Nielsen

et al., editors. Tropical forests. Academic Press, London. IGAC (Instituto Geografico Agustin Codazzi). 1977. Zonas de vida o formaciones vegetales de

Colombia. Bogota. Kahn, F. 1990. I.as palmeras del Arbor6tum Jenaro Herrera (Provincia de Requena, Departamento

de Loreto, Peril). Contribuci6n al estudio de la flora y de la vegetaei6n de la Amazonia peruana. XVII. Candollea 45: 341-362.

Karsten, H. 1857. Plantae columbianae. Linnaea 28: 387-463.

164 8RITTONIA [VOL. 43

Martius, C. F. P. von. 1837. Historia naturalis palmarum. Vol. 2. Genera et species. T. O. Weigel, Leipzig.

Wessels Boer, J .G . 1965. The indigenous palms of Suriname. E. J. Brill, Leiden. 1968. The Geonomoid palms. Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., Tweede

Reeks 58: 1-202. 1972. Clave descriptiva de las palmas de Venezuela. Acta Bot. Venez. 6: 299-362. 1988. Palmas indigenas de Venezuela. Pittieria 17: 1-332.

B O O K R E V I E W S

Diseases and Disorders of Ornamen ta l Palms. Edi ted by A. R. Charles and T. K. Brosehat. APS Press, 3340 Pilot K n o b Rd., St. Paul, M N 55121-2097. ISBN 0-89054-119-1. 1991 .64 pp. US orders $27, n o n - U S orders $34 (paper).

This book contains details of 30 diseases and 27 physiological disorders of ornamental palms. It is a very high quality production, both the text and the 208 color photographs. Under each of the diseases and disorders there are details of symptomology, causal organ- isms, occurrence, species affected, diagnostic techniques, and prevention and treatment. I only noticed one little mistake, on p. 37 the palm captioned as Syagrus romanzoffiana is actually Cocos nucifera. This book will be very useful for any grower of palms in this country. -- ANDm~W HVAqDEmSON, New York Botanical Garden.

Kedondon, Ambarel la , Amra . The Spondiadeae (Anacardiaceae) in Asia and the Pacific Area (with some notes on introduced American species). By A. J. G. H. Kos te rmans . Yayasan T u m b u h - t u m b u h a n yang berguna Jilid I [Foundat ion Useful Plants o f Asia VoL I]. H e r b a r i u m Bogoriense, Jalan Juanda 22, Bogor, Indonesia . ISSN not given. 1991. 100 pp. $6 (paper).

Kosterman's revision ofSpondias L. and allied genera in Asia advocates a much narrower definition of Spondias than the earlier treatments by Airy Shaw and Forman [Kew Bull. 21(1): 1-19. 1967] and Ding Hou [Flora Malesiana, ser. 1, 8(3): 395-577. 1978]. His restrictive definition of Spondias makes the tribe Spondiadeae more amenable to cladistic analysis. The morphology of the endocarp is one of the more important characters used by Kostermans in segregating species from Spondias into the genera Allospondias (Pierre) Stapf, Evia Comm. ex Blume, Haplospondias Kosterm., and Solenocarpus Wight & Am. Haplospondias is a new genus described by Kostermans in order to accommodate the simple-leaved Spondias haplophylla Airy Shaw & Forman. Two of Kosterman's taxonomic decisions that may be particularly controversial are his placement of Spondias dulcis Par- kinson in the genus Evia, and his splitting of Spondias pinnata (L.f.) Kurz into several species. The Asian genera Dracontomelon Blume, Koordersiodendron Engi., Pegia Colebr., and Pleiogynium Engl., traditionally included in the Spondiadeae, are not covered by this revision. Students of the Anacardiaceae will need to consult this important paper.--Jomq MrrcrmLL, New York Botanical Garden.

notes on oenocarpus (palmae) in the colombian amazon

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