Organization for Flora Neotropica
Aiphanes (Palmae) Author(s): Finn Borchsenius and Rodrigo Bernal Source: Flora Neotropica, Vol. 70, Aiphanes (Palmae) (Dec. 16, 1996), pp. 1-94 Published by: New York Botanical Garden Press on behalf of Organization for Flora Neotropica Stable URL: http://www.jstor.org/stable/4393869 Accessed: 13/01/2009 13:13 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=nybg. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit organization founded in 1995 to build trusted digital archives for scholarship. We work with the scholarly community to preserve their work and the materials they rely upon, and to build a common research platform that promotes the discovery and use of these resources. For more information about JSTOR, please contact
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FLORA
NEOTROPICA MONOGRAPH 70
AIPHANES (PALMAE) FINN BORCHSENIUS AND RODRIGOBERNAL
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Publishedfor Organization for Flora Neotropica by The New York Botanical Garden New York Issued 16 December 1996
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Copyright? 1996 The New York Botanical Garden Publishedby The New YorkBotanicalGarden Bronx, New York 10458 Compositionby Eisner/MartinTypographics Manufacturedby Braun-Brumfield,Inc. InternationalStandardSerial Number0071-5794 TM
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Printedin the United States of America using soy-based ink on acid-freepaper MetropolitanLife Foundationis a leadershipfunderof The New YorkBotanicalGarden'sScientific PublicationsProgram. Library of Congress Cataloging in Publication Data Flora neotropica. - Monograph no. 1 - New York: Published for Organization for Flora Neotropica by the New York Botanical Garden, 1968v.:ill.; 26 cm. Irregular. Each issue has distinctivetitle. Separatelycataloguedand classified in LC before monographno. 40. ISSN 0071-5794 = Flora neotropica. 1. Botany-Latin America-Classification-Collected works. 2. BotanyTropics-Classification-Collected works. 3. Botany-Classification-Collected works. I. Organizationfor FloraNeotropica. II. New YorkBotanicalGarden.
QK205.F58
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Libraryof Congress ISBN 0-89327-407-0
[8508]
85-647083 MARC-S
AIPHANES (PALMAE) FINN BORCHSENIUS RODRIGO BERNAL
Tableof Contents Abstract/Resumen ............................................................ Introductionand TaxonomicHistory .............................................. Morphology .................................. ...................... ....... Stem and Habit ............................................................ Roots ................................................................... Indumentand Armature. ................... ................... ......... Leaves............ ........................................... Inflorescence............................................................ Fruits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Germinationand Seedling .............................. ...................... Pollen Morphology ................................................... . Anatomy . ....................................... ......................... Roots....... .................................... .................. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..... Stem . . . . . . . . .2.. . .... ......... Leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hairs and spines .................................. ...................... .. Flowers ................... ................... ........................... ChromosomeNumbers ........................................................ Distributionand Ecology ...................................... ...... .... . ReproductiveBiology ............................................. Inter-and IntragenericRelationships .................................. ..... ... Systematic Treatment ......................................................... Doubtful Names and ExcludedTaxa .............................................. Acknowledgements ........................................................... LiteratureCited . .................... ................. ....................... NumericalList of Taxa ............... ............................. ......... List of Exsiccatae ............................................................ Index of Local Names ........... ...................... ................... Index of Scientific Names of Plants........................................ ...... Index of Scientific Names of Animals .............................................
1 2 4 4 6 6 8 11 14 15 16 22 22 22 22 23 24 26 26 30 32 33 87 88 88 90 91 92 93 94
ABSTRACT Borchsenius, F. (Biological Institute,Departmentof SystematicBotany,AarhusUniversity,Nordlandsvej 68, DK-8240 Risskov, Denmark)and R. Bernal (Institutode Ciencias Naturales,Universidad Nacional de Colombia,Apartado7495, Bogota, Colombia.Aiphanes(Palmae).FloraNeotropica Monograph70: 1-95. 1996.-The neotropicalpalm genus Aiphanes(Arecoideae:Cocoeae:Bactridinae) is revised.Thegenus is characterizedby the combinationof emergentspines, praemorsepinnae, protandrousinflorescences, and pistillateflowers with petals for half their length. Its taxonomic history, morphology, pollen morphology, anatomy, chromosome numbers, distributionand ecology, reproductivebiology, and inter-andintragenericrelationshipsare discussed.We recognize22 species, distributedin the Lesser Antilles, Trinidad,Venezuela,and along the Andes south to Bolivia, with a concentrationof species in western Colombia. Most species of Aiphanes are understorypalms in humid forest types, from 0 to 3000 m above sea level. Aiphanesspicata, from Peru, is describedas new to science. Aiphanes kalbreyeriand A. fbsteriorumare reducedto subspecies underA. hirsuta, and a new subspecies, A. hirsutasubsp. intermedia,is described.
RESUMEN Borchsenius, F. (Biological Institute,Departmentof SystematicBotany,AarhusUniversity,Nordlandsvej 68, DK-8240 Risskov, Denmark)and R. Bernal (Institutode Ciencias Naturales,Universidad Nacional de Colombia,Apartado7495, Bogota, Colombia.Aiphanes(Palmae).Flora Neotropica Monograph 70: 1-95. 1996.-Se revisa el g6nero neotropico de palmas Aiphanes (Arecoideae: Cocoeae: Bactridinae).El g6nerose caracterizapor la combinaci6nde aguijones,pinnaspraemorsas,
Flora Neotropica
2
inflorecenciasprotandricas,y flores pistilladascon petalos unidoshastala mitad.Se discute la historia taxonomica,morfologia,palynologia, anatomfa,numerode cromosomas,distribuci6ny ecologia del genero. Se reconocen21 especies, distribuidasen las Antillasmenores,Trinidad,Venezuelay a lo largo de los Andes desde PanamahastaBolivia, con unaconcentraci6nmayorde especies en Colombiaoccidental. La mayoriade las especies son palmasde sotobosquecreciendoen bosques humedos,desde el nivel de mar hasta 3000 m. Se describeAiphanes spicata como nueva especie. Se reducenAiphanes kalbreyeriy A. fosterioruma subespecies bajoA. hirsuta,y se describeuna nueva subespecie,A. hirsuta subsp. intermedia.
INTRODUCTION AND TAXONOMIC HISTORY Aiphanes is a neotropical genus of understory palms numbering22 species. It is easily distinguished from other palms by its spiny habit and praemorse pinnae,a charactercombinationotherwisefoundonly in a single otherspecies, Bactris caryotifolia.But this latter species differs in having protogynous,not protandrous, inflorescences and pistillate flowers with connate sepals and petals. Aiphanes is restrictedto the Antilles and western South America, and is most diverse in the northwesternAndes, where 13 species occur. A few species are relatively widespread,but many are narrowendemics. Two species are widely plantedgardenornamentals.The genus belongs to the subtribe Bactridinae in tribe Cocoeae of subfamily Arecoideae (Uhl & Dransfield, 1987). Little has been publishedon Aiphanes.The only attempt to treat the genus as a whole is the rather sketchy overview by Burret(1932b), which provides no key and which gives descriptionsfor only those species he described as new. Galeano and Bernal (1987) treatedthe northwestColombianspecies, and BorchseniusandBalslev (1989) the Ecuadoreanones. General studies of palm morphology,anatomy,and palynology have tendedto include only the two cultivated species (underseveralnames), and the diversity within the genus has largely been overlooked. The first observationsof species belonging to what is now known as the genus Aiphanes were made by the French missionaryand botanistCharlesPlumier, who made threetripsto the West Indiesbetween 1689 and 1695. Plumiermadedrawingsand descriptionsof two species which he called "Palmadactylifera,aculeata, fructu corallino, major" and "Palmadactylifera, aculeata,fructu corallino, minor"(both included in A. minima (Gaertner)Burret in this study). The same West Indianspecies of Aiphanes was described by Jacquin (1763) under the name "Palma Grigri Martinicensibus".In 1779 the Spanish botanistJose Celestino Mutis, director of the Real Expedici6n Botanicaal Nuevo Reino de Granada,madea very detailed descriptionof what we now know as Aiphanes
lindeniana.He also directedthe paintingof plates of inflorescencesandinfructescencesof this species and anotherwhich probablycorrespondsto A. aculeata. In 1791 the German gardenerJoseph Gaertnerincluded seeds of A. minima in his book De Fructibus et Seminibus Plantarum under the name Bactris minima,the oldest name in Aiphanes that fulfills the requirementsfor a validly publishedname. The generic nameAiphanes was first used by Carl Ludwig von Willdenow in a lecture held at the KoniglischenAcademiederWissenschaftenzu Berlin in 1801, publishedin 1806 in German,and in 1807 in a French translation.The name was derived from Greekai (always), and phaneros (evident, visible, or conspicuous).Ironically,species of Aiphanesare generally very hardto spot and find in dense vegetation and, accordingly, are among the most poorly collected neotropicalpalms. Willdenowdescribeda single species, A. aculeata, based on materialcollected by the Austrian gardener Franz Bredemeyer in Caucagua near Caracas, Venezuela. The herbarium specimen (if there ever was one) was soon lost from Willdenow's herbarium,as noted by Martius(1847). Bredemeyerhad collected in Venezuelafrom 1786 to 1788, after which he returnedto Vienna where in 1793 he became chief gardener of Schonbrunn BotanicalGarden.In 1809, N. J. Jacquindescribeda palm originating from Caracas and cultivated in Schonbrunnunder the name Caryota horrida. From the descriptionit is clear that this was a species of Aiphanes, and it seems likely that this palm was the same individual or at least had the same origin as Willdenow's A. aculeata. Bredemeyer had most likely broughtseeds with him from Venezuela.Fruit productionof A. aculeata is usually abundant,and seeds are relativelyeasy to germinate. In 1816, Humboldt, Bonpland, and Kunth described A. aculeata once more, this time under the name Martinezia caryotifolia, based on cultivated material from Monte Quindiu, an old name for CordilleraCentralin Colombia.They also describeda new species of Aiphanes,A. praga [= Euterpepraga (H.B.K.) Sprengel].The genus Martineziahad been describedby Ruiz and Pav6n (1794) with a diagnosis
Introductionand TaxonomicHistory that mentionedamong othercharacters,six bractsand a monoecious inflorescence with each pistillate flower placed between two staminate ones. They described no species but mentioned two names, M. ciliata and M. abrupta. In 1798, Ruiz and Pav6n describedfive species of Martinezia,none of which fulfilled the generic diagnosis, as does indeed no known species of palms. Three of the five species were monoecious but had only two bracts, viz., M. ciliata [= Bactris ciliata (Ruiz & Pav.) Mart.],M. interrupta [= Geonomainterrupta(Ruiz & Pav.) Mart.],and M. ensiformis [= Euterpe ensiformis (Ruiz & Pav.) Mart.].The last two-M. lanceolata [= Chamaedorea lanceolata (Ruiz & Pav.) Kunth] and M. linearis [= Chamaedorealinearis (Ruiz & Pav.)Mart.]-had six bracts but were dioecious. Ruiz and Pav6n had also collected Aiphanes aculeata in Peru (Pavons.n., M), but they did not describethis species. The inconsistency of Martinezia Ruiz & Pav6n was soon recognized, and already in 1823 Martius abandonedthe genus in its originalsense, andtook up the name in the sense of Humboldt,Bonpland, and Kunth basing it on Martinezia caryotifolia. During the next years all species originallydescribedin Martinezia were transferredto other genera, the last by Kunth(1841). Kunthcited the genus as "Martinezia Humb. et Kunth (nec Ruiz & Pav6n),"and changed the generic diagnosis so as to fit Martineziacaryotifolia, the only remaining species. In 1847 Martius completed this approach to clear the generic concepts, and synonymized Aiphanes with Martinezia Humboldt.& Kunth.Aiphanes aculeata was placed in synonymy of a new name, Martineziaaiphanes, the diagnosis of which was taken directly from Willdenow.Bactris minima was placed in synonymy of anothernew name, Martineziacorallina, based on Plumier'sdrawings and notes to "Palmadactylifera, aculeata,fructu corallino, major" The result of Martius'sand Kunth'semendationof Martinezia was that the name Aiphanes was, with a few exceptions, out of use until 1932. Karsten(1857) erected the genus Marara to suit two species from Colombia, M. bicuspidata [= A. aculeata] and M. erinacea [= A. erinacea]. The generic name was the vernacular name of M. bicuspidata. Marara was sunk by Wendland(1878), who transferredall known species of Martinezia and Marara to Aiphanes. Wendland'seffort to reestablishthe name Aiphanes was, however, ignored, and Martineziaremainedthe name in use. Cook (1901) erected two genera, Curima and Tilmia (based on the close relatives Aiphanes minima and A. aculeata, respectively), as
3 his contributionto generalconfusion. Both were sunk by Burret(1932b). The last species of Martineziawere published by Burret (1932a). Shortly after, he changed his mind about this genus and, later the same year (Burret, 1932b), synonymized Martinezia with Aiphanes, therebyputtingan end to the confused history of the name. TodayMartineziaRuiz & Pav. is considereda synonym of Euterpe Gaertner(for a discussion of nomenclature,see Burret,1934; Moore, 1963). Martinezia in the emended sense of Martiusand Kunth, and as used by variousauthorssince, is synonymous with Aiphanes. Burret's (1932b) revision of Aiphanes contains a totalof 32 species, including5 dubiousones. Of these, 17 were describedfor the first time, based mostly on collectionsby the GermanbotanistW. Kalbreyer,who travelled in northernColombia between 1877 and 1881. Burretdivided the genus into two subgenera, Macroantheraand Brachyanthera.Burrethad a narrow species concept and described virtually every specimen as a separate species. Given the circumstancesunderwhich he worked,however,it is difficult to see whatelse he could havedone.All he hadto base his conclusionson were a few often incompletespecimens picked at randomfrom the large variationcharacteristicof many species of Aiphanes.Moreover,he had virtuallyno field experiencethat could give him an impressionof the variabilityof naturalpopulations. A largenumberof the species includedin Burret'srevision were representedonly by unicate collections kept in Berlin, and the fire of this herbariumduring WorldWarII destroyedall existing materialof 13 of the 32 species. Since 1932, 15 species havebeen described(Burret, 1937, 1940; Bailey, 1943a, 1949; Dugand, 1944; Moore, 1951;Gentry,1981; Galeano& Bernal, 1985; Borchsenius et al., 1989; Borchsenius & Balslev, 1989), raising the total numberof species to 47. No one has attemptedto revisethe genus since Burret,but regional treatmentsof varying detail exist. Bailey (1949) provideda synopsisof the WestIndianspecies. MacBride (1960) supplied English translations of Burret'sdescriptionsin his treatmentof the palms in Peru but added no new information.Aiphanes has been treatedfor the florasof Panama(Bailey, 1943b); Puerto Rico (Little & Wadsworth,1964); Barbados (Gooding et al., 1965); Martiniqueand Guadeloupe (Fournet,1978);the LesserAntilles (Read, 1979); and Venezuela(Wessels Boer, 1988). Contemporarytaxonomic treatmentsof Aiphaneshave been providedfor westernAntioquia[Galeano& Bernal, 1987 (5 spp.)]
Flora Neotropica
4 and for Ecuador [Borchsenius & Balslev, 1989 (12 spp.)]. Other importantcontributionshave been the resolutionof typificationproblemsconcerningspecies based on lost Kalbreyercollections (Bemal, 1986; Beral et al., 1989). The present revision recognizes 22 species, including 1 new.
MORPHOLOGY Stem and Habit Stems are branchedor unbranchedand vary from short and subterraneanto >20 m tall and up to 20 cm in diameter.Differences in stem branchingand stem developmentresult in a numberof differenthabits in the genus, including solitary subcanopypalms (Fig. 1A), caespitose or solitaryunderstorypalms (Fig. lB, 1C), and acaulescent palms (Fig. 1D). Individual stems are conspicuously ringed with leaf scars and armedwith flattened,grey or black spines insertedin bands or spirals below the nodes. Internodelength is variable and probably reflects differences in growth rate. In small, solitary understory palms such as Aiphanes chiribogensis, internodesare typically very short, only 1-2 cm long, whereas individualsof caespitose species such as A. eggersii in open areas may have internodesup to 15 cm long on the basal partof the stem (Skov,Borchseniuset al. 64735). Ranges of stem heights of floweringindividualsrecorded in the different species form a continuumin the genus as a whole among both caespitose and solitary species (Fig. 2). Two species, Aiphanes acaulis and A. spicata, can be characterizedas acaulescent, having a shortentirelysubterraneanstem;this habitis also found in some populationsof A. ulei and A. weberbaueri. In the lattertwo species, acaulescentpopulations appear to be more frequent as one moves away from the Andes and into lowland Amazonas. Seven species are solitary,understorypalms with unbranched, <10 m tall stems. This number is noteworthy since that habit is uncommon in neotropical palms where understoryspecies tend to be caespitose. Two species, A. grandis and A. minima, develop a solitary, usually >10 m tall stem. The remaining 11 species are all caespitose understorypalms. There is a geographicaldifference in the distributionof habit types: in Amazonian Peru the acaulescent habit is dominant,and only one of the four species occurring in this area (A. aculeata) develops a stem >2 m tall; in the Andes in western Colombia and Ecuador,the caespitose, caulescent, understoryhabit is dominant
and is representedin 9 of the 15 species occurringin thatregion. Stem branchingresultsfrom the formationof axillary shoots, normallyfrom basal nodes just above or below the ground,but sometimes also distally on the stem (Fig. 3A). Dependingon the frequencyof sideshoot formation and the speed with which these develop, a series of caespitose habits occur, ranging from essentially one-stemmed palms with one or a few basal suckers to caespitose palms with up to 20 densely clustered stems. In some cases the whole series can be found within a single species; in Ecuador and southernColombia, Aiphanes hirsuta typically develops only one or two, to 10 m tall stems, andone or morebasal suckers,whereasindividualsin lowland areas of the Choc6 departmentin Colombia frequentlyhave 10 or more, <5 m tall stems. The development of the caespitose habit is to some extent under environmentalcontrol; in western Ecuador, plants of A. erinacea in forest seldom develop more thanfour stems, whereasin open areasand on pasture they frequentlyhave 10 or 20. Aiphanes gelatinosa apparentlyoccurs in both a solitary and a caespitose form, and caespitose individualsof the usually solitary A. ulei and A. weberbaueri are occasionally found (Balslev 69053; Skov & Borchsenius 74770). Polymorphismwith respect to habit has also been recordedin other palms, e.g., Socratea salazarii and Dictyocaryumptariense (Henderson,1990). In other cases, solitary individuals of normally caespitose species represent an early developmental stage in which stem branchinghas not yet been developed; thus A. eggersii may flower while having a single stem, 2 m tall, whereas older plants typically have severalstems, 4-6 m tall. Branchedstems may have two functions:they increase probabilityof survivalof the plantand serve as a means of vegetative reproduction.In a forest in westernEcuadorwhere A. erinacea was abundantin certain areas, it was remarkablethat not a single seedling could be found, althoughmatureplantswere flowering continuously.All small plants encountered representedregrowthfrom brokenand fallen stems or otherwiseseverelydamagedolder plants.The maintenance of relatively high populationdensities of this species seemed to rely more on new shoot formation and recovery after damage than on reproductionby seeds. The oppositesituationwas observedat the same locality in the solitaryA. tricuspidata,which had an extremelylow densityof adultindividualsbut a greater abundanceof seedling plants.Vegetativereproduction through stolon formation has been observed in A.
5
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FIG. 1. Habits.A. Solitarysubcanopypalm,Aiphanesgrandis(Madsen86934). B. Caespitoseunderstorypalm,A. erinacea (Borchsenius& Luteyn91423) C. Solitaryunderstorypalm,A ulei (Balslev et al 62045). D. Acaulescentunderstorypalm, A. spicata (Kahn& Borchsenius26S1).
macroloba.In northwesternEcuadorwe observedthat lateralshoots originatingfrom basalstem nodes developed a thin,recliningstem, which at some distance(<1 m) from the mother plant formed adventitiousroots
anchoringthe daughterplant;above the anchoragethe stem thickenedand became erect, giving rise to a new individual(Fig. 3B). In southernChoc6 we observed thatside-shootson distalnodes of decumbentstems of
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Flora Neotropica
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FIG. 2. Stemheightrangesof floweringindividuals of Aiphanes. A. macroloba soon developed adventitiousroots and startedto segregatefrom the motherstem when leaves were about20 cm long. Roots Above-ground adventitious roots are formed by most species, and sometimes these form a distinctive cone basally on the stem. Formationof adventitious roots from distal stem nodes is common in A. macroloba and has also been observed in specimens of A. erinacea with decumbentstems (Skov& Borchsenius 64757). Adventitiousroots vary in color from grey to reddishbrown.They are 5-15 mm in diameter,often branched,and have numerouswhite, prickly lenticels of endogenous origin. The latter were termed pneumatorhiza by Frangi and Ponce (1985), who concluded thatthey functionin aerationof roots in waterlogged soils. Indument and Armature An indument of densely positioned hairs usually covers the surface of unexpandedleaves and most structuralparts of young inflorescences,particularly
the peduncularbract and peduncle. The indumentis normally caducous and is nearly lacking on older structures.The tomentose strip commonly found abaxially along the distal marginof the pinnaemarksthe part that was exposed before the leaf expanded. A thick, white indument of hairs, each with a muchbranchedfilamentof bladder-likecells, characterizes Aiphaneseggersii,A. lindeniana,andothers.A brown, thinnerindumentof hairs,each with a scaly filament of laterallyfused, ratherthick-walledcells, is found in A. erinacea and severalother species. A darkbrown, very thin,almostpaint-likeindumentoccursin A. ulei. The hairsconstitutingthis indumenthave filamentsof one or a few, spindle-shaped,weakly sclerifiedcells. Spines are a very characteristicfeatureof Aiphanes and otherBactridinae,and they are found on virtually all partsof the palms.Armatureis particularlywell developedon the stem (Fig. 4A), leaf base (Fig. 4B), and peduncle and less so on the petiole, leaf rachis, and peduncularbract. Tomlinson (1990) recognized two categoriesof spines as externalstructuresin the palm family: spines derived from other organs (modified roots,pinnae,leafbasefibers,etc.) andemergencesderived from superficialtissues. The spines in Aiphanes and other Bactridinae are of the latter type. Such
7
Morphology
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8
Flora Neotropica
emergent spines are also found in the paleotropical palm groups Oncospermatinaeand Calameae.Armature is typically more dense in young plants than in older ones, and some genera-e.g., Nephrospermaare spiny only in theirjuvenile stages. Reducedarmature is also found in some species of Bactris, particularly the entire-leaved ones which only have small spines distally on the marginsof the entireleaves. Spines in Aiphanes vary from <1 mm to >25 cm long. Larger spines consist of a pale, cushion-like foot and a rigid, flattened filament terminatingin a sharp point. A gradualtransitionis evident from the smallest spinules to the largest spines; but, for descriptive purposes, small, soft, hair-like structures have been termed "spinules,"and larger,rigid ones have been termed "spines."Spines develop on unexposed surfaces.At first they are pale and appressedto the surface on which they develop. Upon exposure they become rigid and erect. The color normally changes to black, but in A. erinacea, A. simplex, and A. tricuspidata they remain yellow. Spines on the stem of A. aculeata and A. eggersii are often grey. Erection results from the swelling of tissues in the foot region (Rudolph, 1911), and the spines normally end up being retrorse.Armature(including spinules) is highly variablewithin both populationsand species and is generally of little taxonomic value. Tomlinson (1990) suggested that emergent spines in palms are a protectionagainstherbivory.They may also function as a protection against fruit predation by rodents and other small animals, preventingthese from climbing the palm (also see under "Reproductive Biology"). Leaves Most species have spirally arrangedleaves. The phyllotactic patternis 2/5 in A. chiribogensis and A. ulei, and this numbermay be the most common in the genus. Three species-A. linearis, A. verrucosa, and A. lindeniana-have distichous leaves. Distichy of leaves characterizesall palm seedlings (Tomlinson, 1990) but is normally soon superseded by spiral arrangement.In adult plants, distichousphyllotaxyis found in species of several distantly related palm genera(Catoblastus,Oenocarpus,Orania, Wallichia), but its function (if any) is not understood.Rather,retention of distichy in adult palms seems maladaptive, as it leads to mutual shading of leaves (Tomlinson, 1990). Observations of A. verrucosa in southern Ecuadorshowed that the plane of the crown was not oriented in any particulardirection.Furthermore,the
orientationof the crown appearedto shift with time, for distichy was not perfect and the upper leaf was oriented slightly differently from the lower ones. Leaves arrangedin threeverticalrows are found in A. leiostachys, a patternknown also in the Madagascan Neodypsisdecaryi and relatedspecies. The numberof leaves in undamagedcrowns varies from 3 to >20 and appears to be consistent on the populationlevel. In a populationof 31 individualsof Aiphanes verrucosa, the number of leaves ranged from three to five, with four the most common number; 10 individualsof A. chiribogensissurveyednear Chiriboga in western Ecuador had from seven to eight leaves. In some cases the numberof leaves in the crown depends on the age of individuals. In a populationof A. minima in Barbados,young plants with 4-5 m tall stems had 10-14 leaves, whereasmature plants had up to 20 leaves (S. Carrington,pers. comm.). When the numberof leaves in the crown is low, they are usually erect and arching;when leaves are more numerous,they are generally spreadingand the lower ones recurved. The leaf sheaths are tubularin initial stages, enclosing the primordiaof the subsequentlydeveloping leaves. As a leaf ages, two irregular lateral splits form in a basipetal direction and the sheath opens almost to the base. The ventraltissues cut off by the splits soon fall off. The petiole is inserted some distance below the apex of the sheath, and in young leaves a prominent ligule is present. This disintegrates as the leaf matures, but a remnantis always evident, indicatingthe dividing point between sheath and petiole. Leaf abscission is clean in all species except for A. hirsuta subsp.fosteriorum, where the distal partof the stem is often covered with persistent sheaths. General morphologyof leaf bases in palms was treatedin some detail by Tomlinson(1962), who found that most bactrid palms have ligulate leaf bases splitting in the same way as described above for Aiphanes ("Phoenix type"). Exceptions were Desmoncus and some species of Bactris, in which the sheathremainstubularand is little modified with age ("Calamustype"). The petiole varies in length from a few centimetersto 1 m or more. Shortpetioles are channelled above like the sheath, and long ones are generally roundedto terete for most of their length. When the petiole is short, the leaf sheath bends away from the stem and resumes petiolar function, forming what has been termed a "pseudopetiole" (Tomlinson, 1962). The rachis is distally projectedinto a narrow filamentequal in length to the bifid partof the apical
Morphology segment. In older leaves this projection normally breaks off. The lamina is normally pinnately divided into a several-ribbedapical segment and a numberof oneribbed, reduplicatepinnae (Fig. 5A-C). In one species, Aiphanesmacroloba,the laminais entire.Entire leaves are frequently encountered in understory palms, e.g., species of Geonoma and Bactris. The apex of the pinnae (or the outer margin in entire leaves) is praemorse,and in this characterthe pinnae of Aiphanesdiffer from those of the otherBactridinae (with one exception). Praemorsepinnae are known from several distantly related palm groups in the neotropics:Chamaedoreatenerrima,all memebersof tribe Iriarteeae,Aiphanes, and Bactris caryotifolia. The mechanism causing the praemorsecondition is unstudied,but it seems to be the resultof a necrosisof apical tissues probably initiated at an early point in ontogeny. Evidence supporting this interpretation comes from observationsof wilted tissue adjacentto the apical margin of pinnae in unexpandedleaves. The pinnae vary in shape from lanceolateto broadly cuneate, and shape is often highly variablewithin a single species. The term "cuneate" although strictly referring to leaf base shape, has traditionallybeen used in descriptionsof Aiphanes pinnae and this use is continued here; the term equals Uhl and Dransfield's (1987) "wedge-shaped."In A. aculeata andA. eggersii the pinnae widen abruptly near the apex, resulting in an inrolled condition of the pinnae (A-shapedat base, u-shaped at apex) and a characteristic ragged appearanceof the leaves. The apex of the pinnae is variouslyshaped and often providesa good characterfor species recognition. It may be incised, truncate, oblique, lobulate, bicuspidate, or tricuspidate. The distal margin is normally projectedinto a finger-likeextension. The pinnae are regularlyinserted along the rachis or, more often, grouped.Withineach groupthe proximal pinnae are shorter, relatively wider, and more erectthanthe distal,andoften the groupsforma series of horizontallyoriented pinna "fans"along the erect or archingrachis. Regularlyinsertedpinnaeare more or less equal in size and shape and held in one plane. The shape of the pinnaevariesin accordancewith the groupingpattern:densely clusteredpinnaetend to be lanceolateor linear,whereaspinnaeinsertedin remote groupsof few normallyare broadlycuneate;regularly insertedpinnae are always more or less linear.These correlations,as well as the three-dimensionalarrangement of the pinnae,reflect a generaltendencyto minimize mutualpinnashadingwithin the leaf.
9 The overall diversity of leaf structurecan be expressedin the form of two morphologicalseries. The first leads from leaves with numerous,densely clustered, lanceolate, or linear pinnae (Fig. 5A), via leaves with fewer, cuneatepinnae, insertedin groups of 4-6, to leaves with few, often broadlycuneatepinnae inserted in widely spaced pairs or triplets (Fig. 5B). The second leads from stronglygroupedto subregularly or regularly inserted pinnae, usually in association with a change in shape from cuneate to linear (Fig. 5C). Such a transition occurs within several species (A. hirsuta,A. gelatinosa, A. parvifolia, A. weberbaueri),whereas only two species (A. minima, A. acaulis) always have regularly pinnate leaves. Leaf structureappearsin manycases to be related to habit. Densely clustered,linear pinnae characterizesome relativelylarge species as well as some medium-sized ones occurring at high altitudes (A. grandis,A. linearis, A. lindeniana),whereas broadly cuneate pinnae inserted in remote pairs or triplets characterizemost small understory species. Regularly pinnate leaves are found in large (A. minima), medium-sized (A. hirsuta), and small species (A. weberbaueri,A. acaulis). In A. linearis andA. parvifolia our observationssuggest that leaf morphology may depend on habitat, so that plants growing in clayey soil tend to have cuneate pinnae, whereas plantsfrom rockyplaces or growingdirectlyon rocks have linearpinnae. Evidence of environmentalinfluence on leaf morphologycomes from nearbycollections of A. lindenianagrowing <20 m apartbut under different light regimes (Bernal & Galeano 1334, 1335). The first,growingin an open place, had a crippled appearancewith very shortleaves and extremely clustered pinnae; the second, growing in shade, had largerleaves more like the averageof the species. Leaf armatureis highly variableon both an interand intraspecificlevel. The sheath is always densely spiny, whereas the petiole and rachis may be completely unarmed(rarely)or densely spiny. In general, spines become fewer and shortertowardthe apex of the leaves.The midribof the pinnaeusuallybearsone or a few large spines abaxiallyand sometimes also a row of thin spines adaxially.In A. minima spines are sometimes also found on minor veins, especially on the abaxialside. Juvenilefoliage is usuallymore spiny than adult leaves. The pinnae vary from glabrousto densely spinulose on both sides, often within single species. Usually they are more or less concolorous,or only slightly paler abaxially,but in A. ulei they are often stronglydiscolorous,green adaxiallyand silverish abaxially.
10
Flora Neotropica
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FIG. S. Leaves. A. A. Densely narrowpinnae, al. 62012). et Ml. B. Remotely clustered, nmnrow Densely clustered, (Balslev ?~ 62012). B. Aiphanes eggersii pinnae, Aiphan?~ eggersii (Balslev grouped, Remotely grouped, FIG. 5. Leaves. broadpinnae, A. ule Bs al. er et al. 62045). insertedpinnae in one one plume, A. spicatm 6204:. .C. & Brc!~?~ius Borchsenius pinnae,A. (Kahn& Subregularlyinsemted pinnaein plane,A. spicata (Katm broad M?ei (Bolslev C. Subregularly D. Simple A. macroloba macroloba(Skor & Borchscni,s Borchsenius64747). blade,A. 2646). D. (Sko>v Simple blade, 64747). 2646). &~
11
Morphology Inflorescence Aiphanes are pleonanthic and monoecious. Inflorescences are bisexual, interfoliar,erect to drooping duringanthesis,often becomingmoreor less recurved in fruit. Normallyonly one inflorescenceis produced per node, but A. gelatinosa often has three inflorescences per node, sharinga common three-chambered prophyll.Fisher and Moore (1974) suggestedthatthe ontogenetic patternbehind multiple inflorescencesin Aiphanes is the same as in Calyptrocalyx.In this a single inflorescence meristemdevelops two or three apices simultaneously,each giving rise to a single inflorescence, and the three-chamberedprophyllarises from fusion of the separateprophyllsin an early stage of ontogeny. This patterndiffers from that found in Arenga, Wettinia,Catoblastus,and many species of Chamaedoreawheremultipleinflorescencesariseas a centrifugalseries of new apices on an expandedhypopodiumon eitherside of the centralapex (Fisher& Moore, 1974). Inflorescences are usually branchedto one order, and consist of a main axis divided into peduncleand rachis and a number of rachillae (flower-bearing branches).Aiphanesacaulis, A. spicata, andA. macroloba apparentlyalways have-and A. simplexusually has-unbranched (spicate) inflorescences. The condition is rare in A. chiribogensis, and the same individual may produceboth unbranchedand branched inflorescences (see notes to the species). Inflorescences branchedto two ordersoccasionallyoccurin A. erinacea,A. aculeata, A. minima,andA. tricuspidata. A curious instance of second-orderbranchingwas recordedin an individualof A. erinacea growing in a pasture in western Ecuador.When first collected in October 1987, inflorescences had simple racillae (Skov,Borchsenius,et al. 64708); when a second collection was made in October1989, inflorescenceshad up to 20 second-orderrachillaeon each of the lower branches(Borchsenius& Luteyn91423). The peduncle is rounded to somewhat dorsiventrally flattened, and bears a prophyll and a single peduncularbract. Additional, rudimentarypeduncular bractsare sometimes present.The pedunclevaries in thickness from 3 to 50 mm. It has a conspicuous brown to white indument, and is normally densely armed with spines proportionalin size to that of the peduncle itself. The peduncle keeps elongating after the inflorescence has been liberatedand also during fruit development. Some species have a relatively short peduncle and bear their flowers among the leaves; in these species the infructescence usually
ends up being pendulous. Others have a long erect peduncle exserting the inflorescence above the crown, and in these cases the infructescenceremains erect. The prophyll is flat adaxially, rounded abaxially, and winged at the margins.It is generallyalmost hiddenby the sheathof the subtendingleaf. The prophyll encloses the developing inflorescence in its early stages, but is soon pierced abaxially near the apex by the elongatingpeduncularbract.The peduncularbractvariesin texturefrom thick and woody (A. linearis, A. aculeata) to thin and pergament-likedisplaying the contour of the enclosed rachillae (A. chiribogensis).The fully developed inflorescence is liberatedby the formationof a split in the peduncular bract.The split is adaxial in A. chiribogensis and A. erinacea, but in A. eggersii it seems to be abaxial as in otherBactridinae.After splittingis completed, the peduncularbract becomes marcescent. Thin bracts normally soon disintegrate and fall partly off, whereasthick ones are persistent. In the simplest type of inflorescence the rachillae are spirallyinsertedwith shorterand shorterintervals from base to apex, and theirlength decreasesin a linear manner(Fig. 6B). The length of the triad-bearing part of the rachilla likewise decreases linearly, and the distal rachillaeare entirely staminate.The apical rachilla (i.e., the flower bearing terminalpart of the rachis)is generallylongerthanthe subapicalones, and it sometimesbears a few pistillateflowers at its base even when subapicalrachillaeare entirely staminate. The basic inflorescencestructureis modified in various ways: 1) by the presenceof a basalflowerlesspart, 2) by shorteningof the rachillaerelativeto the length of the rachis, 3) by thickeningof the proximaltriadbearingpartof the rachillae,and 4) by adpressionof the rachillaeto the rachis.The developmentof a basal bare portion of the rachillae is observed in many species, most notably in A. chiribogensis and A. duquei, where the bare partof the proximalrachillae is often longerthanthe flower-bearingpart(Figs. 6D, 22A). Short,fastigiaterachillaecharacterizeA. parvifolia (Fig. 6E). Thickeningof the androgynouspartof the rachillaeis sometimes observed,either in combination with large pistillateflowers (A. linearis) or in combination with adpression of the rachillae (A. gelatinosa,A. ulei). In A. verrucosaapparentthickening results from the developmentof large pistillate flowerscoveredby the subtendingbractof the flowergroup. Adpressed rachillae are characteristicof A. gelatinosa andA. ulei (Fig. 6C). In these last two, the rachillae are partly adnate to the rachis and bear densely packedflower-groupsabaxially and laterally
12
Flora Neotropica
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FIG. 6. Inflorescence. A. Newly opened inflorescence of Aiphanes eggersii (Skov et al. 64735). B. Inflorescence of A. erinacea in staminateanthesis with spreadingslender rachillae(Borchsenius91421). C. Old inflorescence of A. ulei with appressedrachillaethat are partlyadnateto rachis(Balslev et al. 62045). D. Not fully expandedinflorescenceof A. chiribogensis in staminateanthesiswith widely separatedbasal rachillaethathave a long basal flowerless part(Borchsenius91440). E. Inflorescenceof A. parvifolia with short fastigiaterachillae(Bernal et al. 1752).
but no flowers abaxially, and the general impression of such inflorescences is often more that of a giant spike than that of a branchedinflorescence. The basal flowering unit of Aiphanes is a triad of one pistillate and two staminate flowers. On distal partsof the inflorescence the triadsare substitutedby dyads of staminateflowers or, near apex, often a few monads. Tetradsof two pistillate and two staminate flowers occur together with triads on the basal rachillae of A. deltoidea, and Read (1979) reporteda similar arrangementin A. minima.Each flower-group is subtendedby a vestigial, rim-likebract,which may be very low, <1 mm long, or may cover the entire flower-groupbefore anthesis,as in A. tricuspidata.In addition,each flower is subtendedby a bracteole.The bracteolesare usually small, but in species with large
pistillate flowers they may resemble the pistillate sepals, as in A. duquei andA. verrucosa.The flowergroups,and especially the pistillateflowers, are often more or less sunken into pits in the rachillae.Apparently sunken flowers may also result from the partial covering of the flower-groupby its subtendingbract or by the bending of the rachilla around the flower group(as in A. chiribogensis),and it is often very difficult to delimit the exact causes of sunken or otherwise hidden flowers. Staminate flowers (Fig. 7A, B) are subsessile or pedicellate.They have three free, carinate, membranous sepals; three free or basally connate, valvate, fleshy petals; six stamens borne in two whorls of three;and a rudimentarypistillode.They display little morphologicalvariationexcept in size and color. The
Morphology
13
FIG. 7. Flowers. A. Cream staminate flowers of Aiphanes aculeata with linear anthers(Bernal & Borchsenius 1959). B. Purplestaminateflowers of chiribogensis with oval anthers(Borchsenius91440). C. Pistillate flowers of A. aculeata with erect corolla lobes (Bernal & Borchsenius1959). D. Pistillateflowers of A. erinacea with spreadingto recurvedcorolla lobes (Skov et al. 64739).
sepals are the first organsto maturein the flower,and they areof nearlythe same size in all species. In larger staminate flowers they are much shorter than the petals and normallydistinct in the matureflower bud; in small staminateflowers they cover the whole bud before anthesis and are clearly imbricate.Petals are spreading to recurved at anthesis as the result of a swelling of basal tissues in the petals, filaments,and receptacle. The filaments are relatively short, never longer than the petals, and basally connate in a low ring. The anthersare introrseto latrorse,inflexed or vertical in bud, but often bend back at anthesis to a more or less horizontalposition with the slits upward. They vary in size from 0.3 to 4 mm, normallyin correlation with petal size. Ranges of anther lengths recordedin differentspecies form a continuumin the genus as a whole (Fig. 8). The pistillode is small, globose or trifid, and functions as a nectary (see "Anatomy"). Color variation of staminate flowers is poorly knowndue to few field observations.Two maingroups of colors may be distinguished:predominantlycream or yellow flowers and flowers with a purpleelement (at least in bud). Often, staminateflowers change in color just priorto anthesis.We observed a change in the color of staminate flowers of A. aculeata near Tamarain northeasternColombia; unopened flower buds were green at first, later creamish yellow; open flowers retainedthat color abaxially,while adaxially the color varied from dense yellow in the center, via creamishyellow, to pure white at the petal apices. We
also observedchanges in A. hirsutanear San Jos6 del Palmarin the departmentof Choc6, Colombia;staminate flower buds were pale violet changing to nearly white at anthesis; open flowers were yellow in the centre, with petals dirty white with pale violet apices and anthersdeep purple.In A. erinacea, flower buds are reddish,rose, or pale violet, whereasopen flowers normally are pure white with yellow anthers. In A. simplexthe color variesfrombrownishviolet in budto dirty white at anthesis.In A. chiribogensis,and probably also A. tricuspidata,flowers are deep purpleboth in bud and at anthesis.Orangestaminateflowers have been reportedin A. deltoidea (field notes to type collection), but how this color relates to the two main groupsdiscussed above is unknown. Differences in size and color, the main trends of diversification in staminate flowers, appear to be related to differences in pollination mode, further discussed in the section "ReproductiveBiology." Pistillateflowers (Fig. 7C, D) consist of three free, broadly imbricate, membranous to cartilaginous sepals;threevalvateor rarelyimbricate,fleshy petals, basally connate for half their length; six staminodes fused in an acutely lobed to nearly truncatestaminodial cup; and a syncarpous, three-carpellatepistil with three sessile stigmas that spreadat anthesis. Pistillate flowers are incompletely developed when the inflorescence is liberated,and they increase significantly in size during staminateanthesis. After pistillate anthesis, the connate part of the corolla and the staminodialcup keeps growing, whereas the corolla
14
Flora Neotropica
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lobes remain unchanged. The perianthsplits irregularly at fruit maturation.Color of pistillate flowers is even less well known than thatof the staminate,but it seems in general to follow the same pattern.Exceptions do occur, as in A. deltoidea, which, accordingto collectors' notes, has orange staminate flowers and green pistillate ones. The pistil may differ slightly in color from the petals, often being more greenish. Small amounts of nectar are secretedjust below the stigmas and at the mouth of the central stylar canal between the stigmas. Fruits Fruitsof Aiphanes are one-seeded drupeshaving a thick, hard endocarp with three distinct germination pores. Most species have globose, red fruits(Fig. 9A), but some variationoccurs. Ellipsoidfruitscharacterize A. macroloba. Stigmatic residues are always apical, and many species have more or less rostratefruits. Small, rostratefruits (Fig. 9B) characterizeA. erinacea and A. ulei. Long-rostratefruits are found in A. duquei. Fruitsof A. grandis are dull green at maturity, resembling not fully mature fruits of Acrocomia. Populations with white fruits are encounteredin A.
verrucosa,A. aculeata, and A. hirsuta;purple-fruited populations,in A. hirsuta,A. gelatinosa, and A. aculeata (A. praemorsa Martius, 1847). The size of the fruits is relativelysmall comparedto other Cocosoid palms, rangingfrom 5 to 25 mm in diameter. The epicarp is smooth, spinulose (A. grandis, A. hirsuta, rarely A. aculeata), or spiny (A. linearis). The mesocarp is normally orange and mealy-fleshy, with a sweet taste. The mesocarpof A. aculeata has one of the highest carotene contents recordedin the plantkingdomand is furtherrich in protein(Balick & Gershoff, 1990). In white-fruitedpopulations of A. aculeata in Bolivia, the mesocarp is hard, dry, and tasteless as in immature fruits. This suggests that white fruits may be the result of mutationsinhibiting the enzymatic apparatusnormally involved in mesocarp maturationand developmentof the red color in the epicarp. In A. verrucosa the mesocarp remains attachedto the rachillabut cracks open at maturityto drop the nut. The endocarp(Fig. 9C) is 0.5-2 mm thick, brown to black, and very hard at maturity.It has three distinct lateral germinationpores, each surroundedby radiatingappressedfibers arrangedin an asterisk-like pattern.The surface may be nearly smooth as in A.
15
Morphology
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lindeniana, shallowly pitted as in A. aculeata and A. minima,or deeply pitted as in A. hirsuta subsp.fosteriorum.The endocarpis often more or less turbinate, especially in species with sunkenpistillateflowers;in A. linearis it is often angulate from mutual pressure when fruits are densely packed. The seed is light brown with a thin testa. The endospermis white and homogeneous, often with an irregularcavity in the interior;in A. linearis and A. grandis the cavity is large, and the endosperm is representedby a thin layer that lines the endocarp,as in Cocos. The endosperm has a sweet taste resembling coconut, and the seeds of A. aculeata, A. eggersii, andA. minimaare eaten locally. Hegnauer(1986) reportedan oil content of 65% in endosperm of A. minima (as A. acanthophylla); Balick and Gershoff
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(1990) reportedan oil content of 37% in endosperm of A. aculeata (as A. caryotifolia),of which the major component(63%) was lauricacid. The embryo is obconical, 0.5-1 mm long and lateral, with the apex pointingtowardone of the three germinationpores. Germination and Seedling Germinationin Aiphanes is adjacent-ligular(Uhl & Dransfield, 1987). Braun (1968) reported the germinationtime of A. aculeata in the BotanicalGarden in Caracas,Venezuela, to be 65 days. The first eophyll is entire, bifid at apex, with praemorseouter margin, and often densely spiny (Fig. 9D). Subsequent leaves graduallydevelop more pinnae until the maturestage is reached(Tomlinson, 1960).
Flora Neotropica
16
POLLENMORPHOLOGY Pollen of Aiphaneshas been studiedas partof general surveys of pollen morphologyin palms based on light microscopy (Thanikaimoni, 1970; Sowunmi, 1972). A descriptionbased also on scanningelectron microscopy(SEM) is providedby Uhl andDransfield (1987). In the presentstudy,pollen of all species was examined by means of SEM; in variable species, pollen from a representativesampleof specimenswas studied.A full list of herbariummaterialexaminedis providedat the end of this section. Pollen grains of Aiphanes are monosulcateto meridionosulcate, rarely trichotomosulcate,globose to elliptic, rarely triangular,20-30 pm along the polar axis, and 20-30 pm in diameter.The exine is semitectate to tectate, and often providedwith supratectal processes including short or long spines, warts, or more or less fusing clavae. The diversity in exine structuremay be summarizedin the form of five main categories: 1. Exine tectate, weakly sculptured,finely reticulate, foveolate-fossulate, or perforate (Figs. 10A-F, 11A, B, 12A-G): A. aculeata, A. deltoidea, A. duquei, A. eggersii, A. gelatinosa (in part,Fig. 13F), A. leiostachys, A. lindeniana, A. minima,A. ulei, A. verrucosa, and A. weberbaueri. This type is also common in other Bactridinae (Uhl & Dransfield, 1987) and probablythe least advanced.Variationsappear to reflect relative degrees of fusion of the elements making up the tectum. Pollen of A. aculeata varies from reticulatewith broad muri (Fig. 12B) to foveolate-fossulate (Fig. 12C). Pollen of A. ulei is smooth, with a sparsely perforateexine (Fig. 12G). Grains are very difficult to examine as they tend to collapse duringthe preparationfor SEM, butlight microscopy of pollen of Balslev et al. 60733 indicatesa variationfrom smooth to somewhat rugulate.Pollen of A. deltoidea and some specimens of A. weberbaueri (Figs. 11B, 12E) is ratherstrongly sculptured and forms a transitionto the following type. 2. Exine tectate, strongly sculptured,with numerous warts or clavae, often laterallyfused resultingin a rugulate surface (Figs. 11C-E, 12J-L, 13A,B 14B,C): A. acaulis, A. chiribogensis,A. erinacea, A. hirsuta in part (Fig. 14B,C), A. linearis, and A. simplex. Least sculpturedis the pollen of A. simplex(Fig. 12J), which is transitionalto the formertype. A geographically isolated and morphologically deviating collection of A. erinacea (Moore 9470) has pollen with scatteredsupratectalwartsand spines (Figs. 11F, 12M) correspondingto the following type.
3. Tectum perforate with scattered, supratectal wartsor spines (Figs. 11F, 12M-0, 13E, 14A, D-F): A. gelatinosa (in part, Fig. 13E), A. hirsuta (in part, Fig. 14A, D-F), A. macroloba,A. parvifolia, and A. tricuspidata.This type appearsto be closely related to the previous,and spines appearto be elaborations of the wartspresentin both types. The close relationship is also indicatedby the occurrenceof pollen of both types in A. erinacea (discussed above) and A. hirsuta (Fig. 14). 4. Exine semitectateto tectate, reticulatewith circular lumina (Figs. 12H, 13C):A. spicata. This type is probablyderived from pollen of type 1, as indicated by the variation in pollen of A. weberbaueri (Kahn& Borchsenius2554) which sometimes is transitional to thatof A. spicata. 5. Exine semitectate,coarsely reticulatewith irregularly shaped lumina (Figs. 121, 13D): A. grandis. This type, too, is probablyderivedfrom pollen of type 1, throughdecreasedlateralfusion of exine elements. Pollen grainsof A. grandisshow some variationin the density of the reticulum and sometimes approach those of A. aculeata (Fig. 12B), althoughthe muriare much narrower. Some species present a considerablevariabilityin pollen morphology, particularlythe related, poorly understoodA. gelatinosa and A. hirsuta. In the first the presence or absence of supratectalspines (Fig. 13E, F) is correlatedwith other morphologicaldifferences, and perhapsthe species as presently circumscribedincludestwo distincttaxa (see notes A. gelatinosa). An equal variationoccurs in A. hirsuta: pollen of subsp. kalbreyeri from Cordillera Central (type locality) is rugulateand devoid of spines (Fig. 14C), whereas pollen of the same subspecies from Cordillera Centralhas numerouslong spines (Fig. 14D); pollen of one specimen of subsp. intermedia is coarsely verrucate (Fig. 14E), whereas that of a nearby collection bears a varying number of fine spines (Fig. 14F); pollen of subsp. fosteriorum is strongly rugulate(Fig. 14B) without spines. Without TEM studies of wall stratificationand knowledge of the ontogeny of the wall in these species, it is not possible to explainthis extremevariationnor to judge its taxonomic significance. Aiphanes certainly appearsto be a promisingobject for furtherpalynological studies. Exine structureand ornamentationis much more diverse in Aiphanesthan in other Bactridinae.Pollen of Acrocomia is subtriangularor rarely elliptic, trichotomosulcate or rarely tetrachotomosulcate or monosulcate, with foveolate-fossulate or reticulate,
Pollen Morphology
17
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FIG. 10. Pollen A Aiphanes eggersi (Balslev et al 62012) B A minima (Beard 484) C A aculeata (Nee 36870). D. A. duquei (Bernal & Devia 1540). E. A. Iindeniana(Henderson & Bernal 125) F. A. leiostachys (Henao et at. 299). A x2226;B, Cx2121; D x1734; E, Fx2121.
tectate exine; pollen of Gastrococos is subtriangular, trichotomosulcate or rarely tetrachotomosulcate, with coarsely reticulatetectate exine; pollen of Bactris is elliptic, monosulcate or rarely trichotomosulcate, with verrucate, foveolate-fossulate, or finely reticulatetectate exine; pollen of Desmoncus is ellip-
tical, monosulcate, with foveolate-fossulate, tectate exine; and pollen of Astrocaryumis elliptic, subtriangular or circular,monosulcate or rarely trichotomosulcate, with finely reticulate, tectate exine (Uhl & Dransfield, 1987). It has been suggested that diversity in exine ornamentation is highest in beetle-
18
Flora Neotropica ,=,,,.j.. ........
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pollinated genera, but in the Bactridinaethe situation have rather uniform, mostly weakly sculptured appears to be reversed: the beetle-pollinatedgenera pollen. The same is the case in other Cocoeae, e.g., Acrocomia, Bactris, Desmoncus, and Astrocaryum Elaeis (beetle-pollinated; Syed, 1979) and the
Pollen Morphology
19
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FIG. 12. Pollen surfaces. A. Aiphanes minima (Beard 484). B. A. aculeata (Bental & Borchsenius 1959). C. A. aculeata (Krukof 11130). D. A. duquei (Bernal & Devia 1540). E. A. weberbaueri (Borchsenius & Pedersen 91427). E A. weberbaueri (Kahn & Borchsenius 2554). G. A. ulei (Balslev 62200). H. A. spicata (Kahn & Borchsenius 2646). I. A. grandis (Steyermark 54185). J. A. simplex (Galeano et al. 262). K. A. linearis (Gentry et al. 40805). L. A. chiribogensis (Borchsenius 91442). M. A. erinacea (Moore 9470). N. A. hirsuta ssp. intermedia (Dransfield 4854). O. A. gelatinosa (Bemnal & Galeano 901). All x6068.
Attaleinae in which beetle pollination appearsto be tive Biology,"Aiphanes seems to comprise a variety common (Balslev & Henderson, 1987; Anderson et of pollinationmodes involving bees, differentgroups al., 1988). As discussed in the section on "Reproduc- of flies, and, to a minorextent, beetles. A comparison
20
Flora Neotropica
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FIG. 13. Pollen.A. Aiphaneschiribogensis (Borchsenius 91442).B. A. linearis(Gentryet al 40805).C. A.spicata(Kahn & Borchsenius2646). D. A. grandis(Steyermark 54185).E. A. gelatinosa(Bernal& Galeano901). F. A. gelatinosa
(Madison et al. 7187). A x2226; B x1734; C-E x2121' F x1980.
of pollen morphology and presumed pollination mechanism in the genus leads to the tentative suggestion that species pollinatedby bees andbeetles (A.
aculeata, A. eggersii) have less-sculptured pollen than do those species that are fly-pollinated (A. erinacea, A. chiribogensis).
Pollen Morphology
21
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Pollen material examined: A. acaulis: Bernal & Galeano 71 (holotype, COL);A. aculeata: Bernal & Borchsenius 1959 (AAU), Henderson & Solomon 520 (NY), Krukoff11130 (NY), Nee 36870 (NY); A. chiribogensis:
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Borchsenius91442 (topotype,AAU), Steyermark52832 (F); A. deltoidea: Tessmann4709 (lectotype, G); A. duquei: Bernal & Devia 1540 (COL); A. eggersii: Balslev et al. 62012 (AAU); Skov et al. 64735 (AAU); A. erinacea:
Flora Neotropica
22 Moore9470 (BH),Balslevet al. 62007 (AAU);A. gelatinosa:Bernal& Galeano901 (topotype,COL),Madisonet 54185 (holotype, al. 7187 (SEL);A. grandis:Steyermark F);A. hirsutasubsp.hirsuta:Allen1870 (holotypeof A. of fuscopubens,MO),Bernal& Galeano251 (holoneotype A. monostachys, COL),Foreroet al. 7224 (COL);A. hirsusta subsp. intermedia:Dransfield4854 (BH), Cuatrecasas 23885 (F);A. hirsutasubsp.kalbreyeri:Bernal& Tobdn1393(isoneotype, AAU);Bernalet al. 958 (FTG);A. Balslevet al. 62102;Skovet al. hirsutasubsp.fosteriorum: 64819 (AAU);A. leiostachys:Henaoet al. 299 (holoneotype,COL);A. lindeniana:Bernal& Galeano849 (AAU), Henderson& Bernal125 (NY);A. linearis:Gentryet al. 40805 (MO);A. macroloba:0llgaardet al. 57620(AAU); A. minima:Little16305(NY),Beard484 (holotypeof A. luciana,BH);A. parvifolia:Callejas4238 (NY);A. simplex: Galeanoet al. 262 (COL);A. spicata:Kahn&BorchSkovet al. senius2646 (paratype, AAU);A. tricuspidata: 64836(holotype,AAU);A. ulei:Balslevet al. 62200,60733 (AAU);A. verrucosa:Skovet al. 64734(AAU);A. weberbaueri:Borchsenius& Pedersen91427 (AAU),Kahn& Borchsenius 2554 (AAU).
ANATOMY Roots Drabble (1903) studied the anatomy of roots of Aiphanes aculeata and A. lindeniana (as Martinezia caryotifoliaand M. lindeniana,respectively).Tomlinson (1961) studied roots of A. minima (as A. erosa). Roots of Aiphanes have a well-developedexodermis. The cortex is parenchymatous,with irregular air spaces, numerousstrandsof non-vascularfibers, and frequentmucilage canals. Drabble(1903) also found individual lignified elements in the cortex of A. aculeata. The stele is medullated, with occasional medullaryvessels. The endodermisis lignifiedandthe pericycle is 1-3-layered. Examinations of roots of A. macroloba (Skov & Borchsenius 64747), A. ulei (Balslev et al. 60742, seedling), and A. weberbaueri(Skov & Borchsenius 64770) showed high levels of infectionwith vesiculararbuscular mycorrhiza. Formation of vesicles and arbuscles was observed in the inner parenchymatous cortex, close to the stele. Stem Stems have an outerepidermis,surroundinga layer of cortex, and a central cylinder of ground parenchyma with scattered vascularbundles (atactostele). The outer partof the centralcylinderconsists almost entirely of black, sclerified fibers, lending rigidity to the stem. Cross sections of stems show some variability in the relative width of the cortex and the
fibroussheath.In most species the cortex is very narrow or nearlyabsent,as in A. simplex (Fig. 15A), but in A. macroloba the cortex is ca. 5 mm thick (Fig. 15B), unusually well developed for palm stems in general (Tomlinson,1990). Leaves Koop (1907) examinedthe laminaof Aiphaneslindeniana (as Martinezia lindeniana); Solereder and Meyer(1928), thatof A. aculeata (as M. caryotifolia). Tomlinson(1961) describedleaf anatomyof A. minima (as A. erosa) andA. aculeata (as A. caryotifolia). In the presentstudy, transversesections of pinnae of all species have been studied.The resultsare summarizedin the followinggeneraldescription(see Fig. 16): Lamina dorsiventral.Epidermis 1-layered, cells rhombohedralto spindle-shaped;outer walls weakly to strongly cutinized;anticlinalwalls generally with periodic thickenings(resemblingpearls on a string), occasionally sinuous. Stomata found predominantly on the abaxialside, not in distinctfiles, sometimeselevatedor slightly sunken.Hypodermis1-layeredbelow each surface,cells generallytwice as wide as those of the epidermis,transverselyoblique-rhomboidin surface view, and orientedat right angles relativeto the epidermis.Substomatalchamberssurroundedby up to 9 cells presentin abaxialhypodermis.Chlorenchyma uniform or with a 1-3-layered adaxial palisade. Raphidecells common. Nonvascularfibers <5 pm in diam., non-sclerified, assembled in strands 10-100 pm in diam., arrangedin 1-4 layers;each strandlined with numerousstegmatawith hat-shapedsilica bodies. Veinsin the abaxialmesophyll;outersheathparenchymatous,incompletearoundmajorveins; inner sheath sclerified, complete, one- to several-layered;phloem of majorveins dividedinto 2-4 strands. Severalpoints in leaf anatomyappearto be related to ecology. Elevatedstomatathatperhapsincreasethe transpiration characterize A. macroloba, a small understorypalm thatoccursin the lowermoststrataof wet premontane rain forest from northwestern Ecuador to Choc6 in Colombia. Aiphanes eggersii from dry tropical forest in western Ecuador has slightly sunken stomata.A thick cuticle is found in several species from the understoryof very humid forest such as A. gelatinosa, A. chiribogensis,and, to some degree, A. macroloba.This characteris absent fromA. aculeata andA. eggersii, both of which occur in dry forest. This may perhaps suggest that cutinization functions as a protection against leaking of nutrients in areas of very high rainfall. Normally,
23
Anatomy
S
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FIG. 15. Stem anatomy.A. X.s. of stem of Aiphanes simplex (Bernal & Borchsenius 1954). Note thick layer of fibers (black) surroundedby a thin cortex (light). B. X.s. of stem of A. macroloba(Bernal & Borchsenius1961). Note relatively thin layer of fibers (black) surroundedby a thick cortex (light).
cutinization is interpretedas a xeromorphicfeature (Roth, 1984) that protects leaves from desiccation. Anatomicalstudiesof Phytelephantoidpalms (Barfod, 1991) gave a result similar to that observed in Aiphanes. Thick cuticles were found in the genera AmmandraandAphandra,both of which occur in wet forest, whereas the cuticle was much thinner in Phytelephas, although some species of this genus occur in seasonal or dry forest. Leaf anatomy in Aiphanes is not markedlydifferent from that of other Bactridinae.Tomlinson(1961) noted that some species of Bactris have branching sclereids in the mesophyll, but B. gasipaes and relatives lack such sclereids, and these species are anatomicallyvery similar to Aiphanes.
relativesclerificationand organizationof the filament cells. Transitionalforms are common, e.g., structures with typical spine structureat base, but terminatingin a branchedchain of weakly sclerified cells. Larger spines often have hairs at the margins,where a gradual transitioncan be found from the cell type typical of the spine filament to that typical of hair filament. Most emergences are, however, easily classified as either spines or hairs. For this reason the two categories are discussed separatelybelow. Hairsemergefrom a small pulvinusof more or less globose, relatively thick-walled cells. The hair itself is often divided into a uni- or multiseriate stalk of moreor less square,thick-walledcells, and a filament of bladder-liketo spindle-shaped,weakly sclerified cells organizedas a tuft of branchedcell rows, or as a flattened scale. Hairs with uniseriate stalk are comHairs and Spines mon on the lamina of all species and appear to be Hairsand spines are relatedstructuresderivedfrom characteristicof all Bactridpalms (Tomlinson, 1961). superficial tissues. Both emerge from a group of Hairs with a multiseriatestalk ending in a branched globose thick-walled cells, forming a small to large tuft of thin-walledcells are typical componentsof the pulvinus on the surface to which they are attached. indumentson sheath,leaf axis, and structuralpartsof The difference between the two types of emergences the inflorescence. Scattered, isolated hairs on is that hairs terminatein a loose, often-branchedfila- rachillaeof A. eggersii are unstalked,with a globose mentof unsclerifiedor weakly sclerified,bladder-like to peltateterminalstructurecomposed of small, bladto spindle-shapedcells, whereasspines terminatein a der-likecells often filled with tannins. The anatomy of spines of A. aculeata (as Mardense, hard, unbranchedfilament of spindle-shaped, stronglysclerifiedcells. The distinctionbetween hairs tineziacaryotifolia)was describedby Rudolph(1911). and spines is not absolute but is, rather,a questionof The foot consists of moreor less globose, thick-walled
24
Flora Neotropica
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FIG. 16. eafanatmy (all 2O2),adxial sid of pinne up. A. .s. of pnna of Aphanes vrrucosa howing sall vein (v) andseveral thin strands ofnon-vascularfibers (f) arrangedin 2-4 layers (Balslev et aL 64531). B. Majorvein of same.~~~~~~~ Noteoutersclerenchymatoussheath(light), and phloem divided into 2-3 strandsby sclerenchymatouspartitions.C. X.s. of~~~~~~~~
arrangedin one layer (Skovet aL 64735).
FIG. 16. Leaf anatomy(all x202), adaxial side of pinnae up. A. X.s. of pinnaof Aiphanesverrucosa showing small veins (v) and several thin strandsof non-vascularfibers (f) arrangedin 2-4 layers (Balslev et at. 64531). B. Majorveinof same. Note outer scierenchymatoussheath (light), and phloem divided into 2-3 strandsby sclerenchymatouspartitions.C. X.s. of in 1-2 layers (Skovet at. A. chiribogensis, showing small vein (v) and severalthin strandsof nonvascularfibers (f) arranged 64822). D. X.s. of A. eggersii small vein (v) with transversevein at bottom and two thick strandsof nonvascularfibers (f) arrangedin one layer (Skov et at. 64735).
cells. The smallest spinules have a filamentconsisting Ecuador(Borchsenius91421). All flowers examined of a single cell. Largespines develop some differenti- were at anthesisor in a statejust priorto this. ation between peripheralcells, which are strongly Staminate flowers (Fig. 17A, B): Sepals consist of sclerified,and internalcells, which are moreparenchy- 1-2 layers of narrowcells. Petals are supplied by a matous,and frequentlybecome vasculated. single vascularbundle, with two metaxylem vessels; they have an epidermis, 5-6 layers of ground parenchyma with many raphide and tannin cells, and a Flowers layer of non-vascularfiber strands located near the The following, predominantlyhistological,descrip- abaxial epidermis; the inner epidermis consists of tion is based on anatomicalsections of staminateand large cells with dense cytoplasm and large nuclei, pistillate flowers of A. erinacea collected in western perhapsindicatinga secretory function. Stamens are
25
Anatomy
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FIG. 17. Floralanatomy.A. X.s. of staminateflowerof Aiphaneseggersii (Skov& Borchsenius64735) showing antherwith fourpollen sacs andconnectivewith numeroussclereids(black)(x52). B. Staminateflowerof A. erinacea(Borchsenius91421), x.s. throughcentral part of pistillode showing nectary (x126). C-F. Pistillateflowers of A. erinacea (Borchsenius91421). C. X.s. throughbasalpartof pistillateflowershowingtree locules surroundedby an innerlayerof sclereids(black)andan outer layerwith many fiberstrands(light);note also fused petalsand staminodialcup (outerring)containingnumerousnonvascular fiberstrands(f) (x22). D. Longitudinalsection throughbasal partof pistil, showing pendulousorthotropousovules and central stylar canal (c) lined with secretorytissues (x52). E. X.s. throughmiddle partof pistil showing numerousnonvascularfiber strands(f), sclereids(black),centralstylarcanal,andthreeseptalnectaries(n) (x22). F. Closeupof septalnectarycanal (x126).
26 supplied by a single vascularbundle with 3-4 metaxylem vessels; filaments are 10-15 cells in diameter and consist mainly of ground parenchyma;sporangium walls are 1-2 cell layers thick; the pollen sacs contain numerousraphidecells in additionto pollen; the connective is rich in sclereids. The pistillode is partly submergedin the fusion productof the basal parts of the filaments, the receptacle, and the basal part of the petals; it contains septal nectaries structurallysimilarto those found in pistillateflowers. Pistillate flowers (Fig. 17C-F): Sepals are4-5 cell layers thick and contain a layer of non-vascularfiber strands.The basal half of the petals is fused with the basal partof the staminodialcup, and it is not possible to distinguish which cells belong to which organ; the joint structureconsists almost entirely of fiber strands held together by a little parenchyma.The petal valves are predominantlyparenchymatous,with some sclereids. The distal half of the staminodialcup is parenchymatouswith a single layer of fiber strands and some sclereids. The pistil consists of threefertile, connate carpels, each bearing a single laterally attached ovule in a basal locule. Running from the basal locules to the stigmas are a centralstylar canal and three channels lined with secretory tissue forming a septal nectary.The basal 1/3 of the pistil, surroundingthe ovules, consists of a dense outerlayer of very small parenchymatouscells, followed by a loose sheathof sclereids.The middle '/ consists of an outer sheath of non-vascular fiber strands, and an inner sheath of sclereids. The distal /3 consists almost entirely of fibers and raphides,held togetherby parenchymatous cells. Staminateflowers of A. chiribogensis(Borchsenius 91440) and A. eggersii (Skov, Borchsenius, et al. 64735) examined for comparisonwere little different in anatomy.Likewise, pistillateflowers of A. chiribogensis were anatomicallysimilar to those of A. erinacea, describedabove. Comparativeanatomyof the palm gynoecium was describedby Uhl and Moore (1971). The gynoecium of Aiphanes has the same fundamentalstructureas the gynoecium of Bactris except that septal nectaries are absent in the latter. Septal nectaries similar to those in Aiphanes are commonly encountered in palms, includingthe Cocosoid genus Butia. The most striking feature of pistillate flower anatomy in Aiphanes is the large amountof protectivetissues in the form of fibers, sclereids, and raphides.The phenomenon is probablyrelatedto the protandrouscondition of inflorescences and consequent prolonged exposure of the pistillate flowers priorto anthesis.
FloraNeotropica CHROMOSOME NUMBERS Chromosomecounts are given in the literaturefor two species.Aiphanesminima:n = 15 (Read, 1966; as A. erosa),n = 18 (Gassner,1941; as Martineziaerosa); andA. aculeata: n = 15 (Read, 1966; as A. caryotifolia); n = 16 (Eichhom, 1953; Sharma& Sarkar,1957; Sato, 1946; all as Martineziacaryotifolia).Accurate chromosomecounts in palms are difficult to obtain, and the apparentintraspecificvariation in chromosome numberof the two species may be an artifact. The most recent chromosome counts (Read, 1966) statedn = 15 for both species examined.This number has also been reportedfor the BactridgeneraAcrocomia, Gastrococos, and Astrocaryum(Uhl & Dransfield, 1987). The generalnumberin Bactris appearsto be n = 14, butn = 15 has been reportedin B. gasipaes. The most common state in otherCocoid palms is n = 16, but n = 15 has been reportedin Syagrus, n = 80-100 in Jubaeopsis, and n = 586+ in Voanioala (Johnson, 1989; Johnsonet al., 1989). Chromosome numbers in palms are believed to form a dysploid series decreasingfrom n = 18 (the ancestralnumber) to n = 13 in some groups(Uhl & Dransfield,1987).
DISTRIBUTION AND ECOLOGY Aiphanesis distributedin the LesserAntilles, Trinidad, and along the Andes from Venezuelato Bolivia, from sea level to about 2800 m elevation (Figs. 18, 19). One species reaches Panama,but otherwise the genus is not representedin CentralAmerica.The eastwarddistributionis restrictedto the westernmostparts of the Amazonbasin;it just scarcelyreachesBrazil in the borderareawith Peru.Reportsof A. aculeata from BritishGuyana(Schomburgk,1848) and from different parts of southern Venezuela (Braun & Chitty, 1987; Humboldtet al., 1816) are not documentedby herbariumvouchers.The maincentreof species diversity lies in western Colombia and Ecuador;a minor centreis found in northeasternPeru. Aiphanes aculeata is the most widely distributed species in the genus, occurring from Trinidad to Bolivia with a gap from middle Colombia to central Peru.It occursboth in tropicaldry forest and in morehumidforest types, but the gap in its distributionarea coincides with the partof the Amazon basin that receives the highest amountof precipitation.Aiphanes minimaandA. eggersii,both close relativesof A. aculeata, are endemic to the Lesser Antilles and the coastalplain of Ecuador,respectively.The firstoccurs
Distribution and Ecology
27
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F'IG.18. Distributionmaps. A. Aiphanesaculeata (circles);A. eggersii (stars);A. minima(triangles).B. A. hirsutasubsp. hirsuta (circles); subsp. hirsuta and subsp. kalbreyeri(squares);subsp. intermedia(triangles);subsp. fosteriorum (stars); (cG,irs)A. chisribtognsi (trangls); A.paeduqueit (squres); A. lgeiostahy (stars A im (rage). B .hruasbp subsp. hirsuta, subsp. intermedia, and subsp. fosteriorum (circle, star, and triangle superimposed). C. A. -weberbaueri A. A. A. (circles); chiribogensis(triangles); duquei (squares); leiostachys (star).
both in seasonal forest and in humid,premontaneforest in the interiorof St. Luciaand Dominica;the latter is found in areasreceiving as little as 500 mm annual precipitation and in which semi-deciduous Ceibadominated forest constitutes the natural vegetation. Aiphanes lindeniana is distributedin the Colombian CordillerasOrientaland Central.It is often a conspic-
uous element in montanecloud forest between 1900 and 2800 m elevation. The closely relatedA. verrucosa is endemic to the central Andes in southern Ecuador,where it occupies similarhabitats. The remaining species can be divided into two groups:those distributedeast of the Andes and those distributedwest of the Andes.
28
Flora Neotropica
'9,'~'v
A
B
C
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The first group consists of four species. Aiphanes ulei ranges from southernColombiato northernPeru, reaching 1850 m in the Andes; eastward it just reaches the westernmost parts of Brazil. The poorly
.0
knownA. deltoidea appearsto have a similardistribution. Aiphanes weberbaueri ranges from southern Ecuador to central Peru, reaching 1950 m in the Andes; eastwardit extends at least as far as Iquitos.
29
Distributionand Ecology Table I Distributionof taxa with main occurrencein the ColombianCordilleraOccidental and the Choc6 North Pan CCA A. hirsuta subsp. hirsuta A. leiostachys A. hirsuta subsp. kalbreyeri A. parvifolia A. linearis A. acaulis A. hirsuta subsp. intermedia A. simplex A. macroloba A. duquei A. hirsuta subsp.fosteriorum A. gelatinosa A. erinacea A. tricuspidata A. chiribogensis A. grandis
x
x x x x x
6
5
4
x
x
x
x
x x
x
3
x
x
x x x
x x
2
1
0
x
x
x x x x
x x x x x
-1
-2
x
x
-3
South -4
x x
x x
x x
x x x x x
x x x
x
x x
x x
in Colombia; Pan:Panama; Centralin dept.Antioquia CCA:Cordillera beltsalong remaining figuresall referto latitudinal CordilleraOccidentalin Colombiaand Ecuador;6: northof 6? northernlatitude;5: between 6? and 5? northernlatitude,etc.; -1: between 0? and 1? southernlatitude,etc.; x: occurrencedocumentedby a herbariumvoucher.
The last species, A. spicata, is known only from two nearbysites in the easternPeruvianAndes at ca. 1800 m elevation. The second group includes 13 species distributed along the westernAndeanCordillerafrom Panamato southern Ecuador.The central parts of this area receive up to 9000 mm precipitationequally distributed over the year (Cuatrecasas,1957), northernEcuador receives ca. 5000 mm (J0rgensen& Ulloa, 1989), and southern parts receive less. The latitudinaldistribution of the species of this group along the western Andean Cordillerais shown in Table I. Many appear to be narrowendemics:A. leiostachys is known from a single locality in the northernpart of Cordillera Central in Colombia, at 800-900 m; A. acaulis is known from two localities in the central Choc6, between 150 and 700 m; A. duquei is known from a small area in the departmentsof Valle and Cauca, at 2000-2600 m; A. chiribogensis and A. grandis are endemic to western Ecuador, at 1700-2100 m and 1000-2000 m, respectively. Others have somewhat wider distributions:A. hirsuta is found from Panama to northernEcuador(butrepresentedby differentsubspecies), at 100-2200 m; A. macroloba occurs from northernColombia to Ecuador, at 500-1400 m; A. linearis andA. parvifolia occur from northernto central Colombia, at 1900-2300 and 1500-2000 m, respectively; A. tricuspidata is found in the Pacific
lowlands and the Andean foothills from Valle del Caucato southernEcuador.Aiphanes simplexoccurs throughoutthe Rio Caucabasin;it crosses Cordillera Occidentalin the two places, nearFrontinoin Antioquia, and west of Cali where its distributionmeets that of the closely relatedA. erinacea, ranging from here south to central Ecuador.The latter also has a small isolated population on the eastern Andean slopes in Ecuador,between 1900 and 2100 m. The highest number of Aiphanes species is found in a zone between 3? and 4? northernlatitude,apparently the result of a superimpositionof species with main distributionsnorth and south of this zone. In that region, roughly correspondingto the Colombian departmentof Valle, 9 of the 24 taxa recognized in this study can be found (includingA. macroloba, which, presumablyoccurs in this region). The altitudinalranges of the 22 species are shown in Figure 20. A calculationof the numberof species occurringin each 100 m altitudinalintervalfrom sea level to 2800 m elevationgives the resultthatbetween 8 and 11 species occur in each 100 m interval from 100 m to 2100 m elevation,and thereafterthe number declines. If species are classified according to their mean altitudinaloccurrence,i.e., the average of the maximum and minimum elevation of their occurrences, the following conclusion is reached. Four species-A. minima,A. eggersii, A. tricuspidata,and
30
Flora Neotropica
3000
t2000
1000 ______
__
E . 0A
i
____
i
FIG. 20. Altitudinalrangesof species of Aiphanes.
A. acaulis-have mean altitudinaloccurrences<600 m, and have in no case been recorded at elevations >700 m. Five species-A. spicata, A. chiribogensis, A. linearis, A. duquei, andA. lindeniana-have mean altitudinaloccurrencesbetween 1800 and 2350 m and have not been recorded at elevations <1500 m. The remainingspecies have mean altitudinaloccurrences between 750 and 1500 m, and several have wide altitudinal ranges, e.g., A. aculeata (0-1700 m), A.
dantin patchesof tropicaldry forest in the department of Casanarein Colombia and to be dominantin secondaryvegetationon slopes in NorYungasin Bolivia. Madsen (pers. comm.) found thatA. grandis, though very local, occurredat high densities on the eastern Andean slopes in southernEcuador.Most species are found on well-drained, clayey or rocky soils. Aiphanes ulei occurs both in terrafirme and in periodically inundated varzea forest. Aiphanes weberweberbaueri (150-1950 m), A. ulei (200-1850 m), baueri occurs both on lateriticsoil and on white sand and A. hirsuta (100-2200 m). in the northernpart of Peruvian Amazonas. Many Little is known about populationstructureand eco- species appearto be unableto survivein disturbedforlogical preferenceof the species, but most appearto est or open areas, particularlyacaulescentor solitary occur in very low densities. Kahn and Mejia (1991) understorypalms such as A. ulei, A. weberbaueri,A. found seven individualsof A. ulei in a 0.5 ha plot of parvifolia, and A. tricuspidata. Some caespitose lowland rain forest in the PeruvianAmazon, a result species, such as A. erinacea and A. hirsuta, may surthat correspondswell with the general impressionof vive on pasturewhere plantstend to have many stems low densities of this species coming from field obser- and yellowish green leaves andto producemany more vations in Amazonian Ecuador.Borchsenius (1993) inflorescences than in closed vegetation. Seedling found 25 individualsof A. erinacea in an area of ca. plantshave neverbeen observedin open areas. 0.25 ha of disturbedpremontanerainforest in western Ecuador,but in the remainingparts of the forest the species was much less abundant.Only a few species REPRODUCTIVE BIOLOGY have been seen to occur at high densities:A. simplex Most species flower throughoutthe year, and buds, may sometimes dominate the shrub layer in forest around m in in 2000 the Rio Cauca basin andinfructescencesin all phenological inflorescences, patches Colombia;A. aculeata has been observedto be abun- stages can usually be found simultaneously within
Biology Reproductive populations.The continuity in flowering can be explained by the relativelyconstantclimatic conditions throughoutthe year in most parts of the distribution area. The only example of pronouncedseasonality comes from our observations,madein June 1990, of a white-fruitedpopulationof A. aculeata aroundCaranavi in Bolivia. Numerous individuals had old infructescences from which the fruits had fallen, and only a few infructescenceswith fruitswere found. No buds or inflorescenceswere encountered,andthe only collection with flowers of this populationwas madein December (Henderson & Solomon 520). The strong seasonality appearsto be a local phenomenon,since flowering collections from adjacent areas at higher altitudes have been made in July (Moraes & Balslev 836, 842). More knowledge of local differences in flowering season would be interesting,since such differences may function as genetic barriersbetween adjacentpopulationsof a species such as white- and red-fruitedpopulationsof A. aculeata in Bolivia. The time lapse between subsequentinflorescences on single shoots is largelyunknown.One observation of a populationof A. eggersii in a pasturein western Ecuador showed that the next inflorescence bud in several cases opened shortly after the last inflorescence had finished anthesis. With a period of ca. 25 days from inflorescence liberationto end of anthesis (Borchsenius, 1993), it can be estimatedthatinflorescences on one shoot may have been initiatedat intervals as short as one month.In contrast,individualsof the solitaryA. chiribogensisin montanerainforest in western Ecuador never had simultaneous inflorescences and infructescences on a single plant. Total anthesis time per inflorescence in this species is at least 80 days (Borchsenius, 1993), and fruitdevelopment may well be equally slow. Thus it is likely that this species produces only one or at most two inflorescences per plant per year. Although the estimates are crude, they neverthelessgive an idea of the variation in floweringintensitythatmay be expectedin the genus. Inflorescences of Aiphanes are protandrouswith non-overlappingstaminateand pistillate phases, and individualplants rarely have more than one inflorescence at a time (this occursonly in caespitosespecies, where different shoots simultaneouslymay bear inflorescences).The separationof male andfemalefunction of individualsin time (dichogamy,temporaldioecism) is very common in monoeciouspalms. Cruden (1988) concludedthattemporaldioecism could be interpretedas a mating system favouring xenogamy, comparableto self-incompatibilityor heterostyly.
31 Borchsenius(1993) describedsingle-inflorescence phenology and insect visitation of A. chiribogensis, A. eggersii, andA. erinacea in western Ecuador.The flowering patternsuggested acropetalmaturationof flower-groupsand inflorescences. First, one staminate flower of each flower-groupopened; only when the first flower had fallen from all flower-groupsdid the second ones startto open. Apartfrom this, no particular sequence of opening was observed. Pistillate flowers opened in an acropetalorder.Flowers of both sexes producedsmall amountsof nectar,but no scent was detected.Staminateanthesisof A. eggersii began immediatelyafterthe inflorescencewas liberated,and lasted 8-10 days; pistillate anthesis began one week laterand lastedabouta week; flowers were visited by bees andwasps,anda high percentageof the staminate flowers containedmicromothlarvae(Lepidoptera);it was concluded that bees were the pollinators, with possible participationof wind. Inflorescences of A. erinacea developed more slowly; staminateanthesis began immediatelyor up to 6 days after the inflorescence was liberatedand lasted 17-21 days; pistillate anthesis began 4-8 days later and lasted 7-9 days; flowers were visited by hundredsof insects, predominantlyflies; potentiallyimportantpollinatorsincluded fruit flies (Drosophilidae),hover flies (Syrphidae), biting midges (Ceratopogonidae),and leaf beetles (Chrysomelidae);no bees were observed.The slowest inflorescence developmentwas found in A. chiribogensis; the peduncularbract split gradually over 5 days or more; staminate anthesis began 7-14 days laterand lasted 40-50 days; pistillateanthesis started >15 days laterand lasted at least 10 days; insect visitors were few-fruit flies (Drosophilidae)were most common and other visitors included fungus gnats (Mycetophilidae,Sciaridae),midges (Cecidomyiidae, Ceratopogonidae),and micromoths(Lepidoptera);no bees or hoverflies were observed. Thereare a few otherobservationsrelatingto flowering biology of Aiphanes. Sweetly scented flowers have been noted in A. grandis (field notes to Steyermark54185) andA. minima(Hoyos & Braun, 1984). Listabarth(1992) observedin Peruthat staminateanthesis of A. aculeata lasted 15-20 days; it was followed by a pauseof 2-4 days beforepistillateanthesis began;all pistillateflowers became receptivemore or less simultaneouslyand remained so for 2-3 days; pollinating agents were wind, bees (Hymenoptera: Meliponidae), weevils (Curculionidae), and bugs (Hemiptera).We have observedbees visiting flowers of both sexes of A. aculeata nearYopal in Colombia; Curculionidae and Nitidulidae were common in
32 staminateflowers but not observedon pistillateones; furthermore, a large proportion of the staminate flowers contained larvae of micromoths (Lepidoptera).Beetles have been observedin staminateflowers of A. aculeata in Bolivia (Henderson & Solomon 520). We observedflies and weevils on inflorescences and flowers of A. simplex in Valle in Colombia.The weevils had the same colour and size as unopened flowerbudsand were difficult to spot without careful examination. The observations suggest that a series of related pollination mechanisms occur in the genus and that these are due to differencesin speed of inflorescence development and flower morphology.The combination of relatively large staminateflowers with linear anthers and rapid inflorescence development (less than one month total anthesis time) appearsto be associatedwith pollinationby bees, sometimeswith participation of beetles (Coleoptera),bugs (Hemiptera), and wind, whereasthe combinationof small staminate flowers with minute oval anthersand slower inflorescence developmentappearsto be associatedwith pollinationby flies. The colourof the flowersmay also be related to pollination mechanism. Large staminate flowers are generallycream or yellow, whereassmall staminate flowers often are more or less purple. Finally, a more specific plant-pollinatorrelationship, involving weevils mimicking staminate flowerbuds, may occur in A. simplex. Henderson(1986) suggested that fly pollinationin palms is most common in understorypalms and often involves a slower developmentof the inflorescence.If pollination mechanism, as discussed above, can be judged from staminateflower morphology,Aiphanes seems to confirmthis suggestion.Large,white to yellow staminateflowers with linear anthers,apparently associated with bee pollination, characterizeseveral subcanopyor canopy species (A. grandis,A. minima, A. aculeata, A. eggersii) as well as certain mediumsized species occurringat high altitudes(A. duquei,A. linearis, partlyA. lindeniana),whereassmall, mostly white to purple staminateflowers with minute, oval anthers, apparently associated with fly pollination, characterize most small understory species (A. acaulis, A. chiribogensis,A. deltoidea,A. erinacea,A. parvifolia,A. tricuspidata,A. ulei, A. weberbaueri). The time requiredfor fruit developmentis largely unknown. In western Ecuador we observed that an inflorescence of A. erinacea had full-sized though still immature fruits only 25 days after the end of pistillate anthesis.A consistentlyhigh fruitset is seen in A. aculeata, A. minima,andA. eggersii. In A. ulei,
FloraNeotropica A. macroloba, and A. simplex, most inflorescences appear to abort entirely, and mature fruits are very rarely seen. Observationsof A. chiribogensis at its type locality in westernEcuadorshowed that fruitset was highly variable. Several completely aborted inflorescences were observed:in others, fruit set varied from one or a few fruitson an entireinfructescenceto 58 fruits on an infructescencewith an initial number of ca. 230 pistillateflowers. Maturefruitsof A. aculeata are eaten by squirrelsthat are able to climb the stem in spite of the spines. In northeasternColombia we observednumerouscrackedendocarpslying scattered on the groundbelow fruitingindividuals,but it was not determinedwhich animal was responsible. The fruits and seeds of this species are rich in vitamins andenergy (Balick & Gershoff, 1990) and probably an attractivefood source for various animals. Thus it seems likely thatthe retrorsespines on stems, sheaths,and inflorescencesserve, at least in part,as a protection against seed predation. The bright-red fruits of A. aculeata are eaten and dispersed by the oilbird, Steatornis caripensis (Snow, 1962; Snow & Snow, 1978).
INTER- AND INTRAGENERIC RELATIONSHIPS Uhl and Dransfield(1987) placedAiphanesin subtribe Bactridinae (Arecoideae: Cocoeae), together with the genera Acrocomia, Gastrococos, Bactris, Desmoncus, and Astrocaryum.The Bactridinae(the Bactris alliance of Moore, 1973) are a strictly neotropicalgroup distinguished from the remaining Cocoeae by the presenceof emergentspines on most, or at least some, partsin all members.Anatomically the Bactridinaeare distinct in having stegmata with hat-shapedsilica bodies (vs. spherical in other Cocoeae) and in having very narrownonvascularfibers, less than5 pm in diameter,assembledin wide strands in the mesenchyma(Tomlinson, 1961). The characters thattraditionallyhave been weighted most in the alignment of the six genera are the arrangementof flowers on the rachillaeand especially the degree of connation of the perianth parts in flowers of both sexes. In this view Acrocomia is the least advanced genus, showing no connation in perianth parts of flowers of neithersex; Aiphanes and Gastrococosare somewhat more advanced, having partly connate pistillate petals; and Bactris, Desmoncus, and Astrocaryum form the most advanced group in the subtribe, having pistillate petals connate in a tubular,
33
SystematicTreatment three-lobedto truncatecorolla. The above view was advocatedby Moore (1967), who suggestedthatGastrococos and Aiphanes were sister-genera, sharing characterssuch as staminateflowers mostly in dyads distally on the rachillae(vs. mostly single in Acrocomia); pistillate petals half connate and valvate (vs. free and imbricate);staminodesconnatein a cup and adnateto the corolla tube (vs. free from corolla);and erect stamen filaments (vs. inflexed in Acrocomia). Inflexed filaments are, however,known in Aiphanes, as are staminodesfree of the petals, imbricatecorolla valves, and solitarystaminateflowers (apicallyon the rachillae). Aiphanes is morphologically a well-defined unit, and there seems no reason to doubt that it is monophyletic. The charactersdefining Aiphanes are praemorse pinnae (apomorphic), protandrousinflorescences (possibly apomorphic;see below), staminate flowers with free sepals and nearly free petals, and pistillate flowers with free sepals and half-connate petals. Among the Bactrid palms, praemorsepinnae occur only in Aiphanes and in Bactris caryotifolia, where they must have evolved independently.If the above considerationsof phyletic relationshipsin the Bactridinaeare correct, it is likely that a shift from protogynyto protandryoccurredin the differentiation of Aiphanes. The genera Acrocomia, Bactris, Desmoncus, and Astrocaryumhave all been shown to be protogynous and beetle pollinated (Essig, 1971; Bullock, 1981; Beach, 1984; Burquez et al., 1987; Scariotet al., 1991; Listabarth,1992). Unfortunately, no information about sexual system or pollination mechanism is available for Gastrococos. The shift from protogynyto protandryprobablyoccurredas an adaptationto bee or fly pollination,the predominant mode of pollinationin Aiphanes. Having studieddistribution and pollination syndromes in Ecuadorean palms, Borchsenius (1993) provideddata suggesting that beetle pollination is common in lowland palms whereas most montane genera are bee or fly pollinated. If this is correct,it seems likely that the shift in pollinationmode and in the timing of sexual functioning occurred as, or at least provided, an adaptation to the montane environmentwhere Aiphanes is most diversified. Burret(1932b) divided Aiphanes into two subgenera, Macroanthera and Brachyanthera.The differences between the two are summarizedbelow (translated from Burret'sdiagnoses): Subgen. Macroanthera. Flower clusters scarcely sunken. Male flowers pyramidal or conical, higher
thanwide, mostlyconspicuouslyacuminate,acute;
andsolidat base; sepalsnotmuchrounded-extended acuminate,acute; petalsmoreor less linear-oblong, antherslinear.Femaleflowers:staminodial cup with six narrow, teeth. conspicuous Flowerclusters,particuSubgen.Brachyanthera. larlymale flowers,not muchsunkeninto the axis. Maleflowersmostlysmall,oftenbroaderthanwide, roundedat apex;sepalsthin,neverextendedat base, butmostlyslightlycarinate; petalsmostlyovate,obtuse to rounded;anthersvery short,suborbicular or oval.Femaleflowers:staminodial cupwithtriangular, moreor less indistinct teeth. The characterspresentedby Burretare not of much value in separatingsubgeneric entities in Aiphanes. Floral differences are vague and refer to variable characters.The degree of sunkenness is a character that is difficult to define, and in reality there is no differencebetween the flower insertionpatternof A. eggersii (placed in Macroanthera) and A. weberbaueri (placed in Brachyanthera).The main character, antherslinear or very short, displays continuous variationin the genus as a whole (Fig. 8), and several species are quite variablein this respect. Burretreferred 11 species to subgenus Macroanthera, here reducedto three:A. aculeata, A. eggersii, and A. minima. These species are closely related, judged from overall similarity,particularlyin inflorescence and fruit morphology. Aiphanes aculeata andA. eggersii are both characterizedby pinnae that widen abruptlynearthe apex, a characterstate that is sometimesexpressedalso in A. minima(Ekman1484, US), andthis seems to be a synapomorphyunitingthe three.Anothersynapomorphyis the presenceof characteristicpeltate hairs on the rachillae.Thus there is little doubtthatthe threespecies form a monophyletic group,butrecognitionof this groupas a separatesubgenus is problematic,since it would leave a highly heterogeneoussecond subgenus, not characterizable in otherways thannegatively,i.e. being those species not belonging to subg. macroanthera.For these reasons, Burret's subgenericdivision has not been followed here.
SYSTEMATICTREATMENT 1. Aiphanes Willdenow, Samml. Deutsch. Abh. Konigl. Akad. Wiss. Berlin 1803: 250. 1806. Type. Aiphanesaculeata Willdenow. Auct.(nonRuiz& Pav6n). Martinezia Linnaea28: 389. 1857.Lectotype(Moore, MararaKarsten, 1963). MararaerinaceaKarsten[= Aiphaneserinacea H. Wendland]. (Karsten)
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CurimaCook,Bull.TorreyBot. Club28: 561. 1901.Type. CurimacolophyllaCook [= Aiphanesminima(Gaertner) Burret]. Tilmia Cook, Bull. TorreyBot. Club 28: 565. 1901. Lectotype(Moore,1963).Tilmiacaryotifolia(Humboldt, Bonpland & Kunth) Cook [= Aiphanes aculeata Willdenow]. Monoecious,pleonanthic,acaulescentor caulescent understorypalms, solitaryor caespitose,with up to 20 stems. Stems armed with rings of black, applanate spines inserted in bands or spirals below the nodes, sometimes becoming unarmedwith age. Leaves spirally arrangedor distichous; sheath open nearly to base in older leaves, densely armedwith black or yellow spines; petiole nearly absent or well-developed, armedlike sheath or unarmed;rachis roundedbelow, ridged adaxially, unarmedor spiny; lamina reduplicate, entire,or paripinnatelydividedwith up to 70 pinnae per side, pinnae regularly inserted or grouped, lanceolateto broadlycuneate,apex (or outermarginof entire lamina) praemorse;midribof pinnae normally with 1 to severalrigid spines abaxially.Inflorescences interfoliar,protandrous,erector curving,often becoming recurvedor pendulousin fruit,spicateor branched to 1 or occasionally 2 orders;prophyllshort, lanceolate, winged, more or less enclosed in the leaf sheath; peduncularbractlong and slender,unarmedor spiny, persistent or soon disintegrating;peduncle normally longer than rachis, more or less spiny; rachillaeup to 300, spreading, fastigiate, or appressed, sometimes with a long basal flowerless part;fertile partof proximal rachillaewith triadsof 1 pistillateand 2 staminate flowers for ca. /2of theirlength, sometimesalso with
a few tetradsof 2 pistillate and 2 staminateflowers, distally with dyads of staminateflowers, or near apex with a few single ones; distal rachillae staminateor with a few triads at base. Flowers unisexual, trimerous. Staminatesepals free or shortly connate, membranous, imbricate, carinate, often cap-shaped and enclosing the entirebud before anthesis;petals free or shortlyconnate,fleshy, ovate-acuminate,valvate;stamens 6, in 2 whorls,inflexed in bud, filamentsbasally connatein a ring,antherslatrorseto introrse;pistillode small,trifid.Pistillateflowersgenerallylargerthanthe staminate;sepals free, cartilaginous,ovateto reniform, imbricate;petals fleshy, connate for /2 their length, lobes acute-acuminate,valvate,or occasionallyimbricate; staminodes6, fused in an acuminatelylobed to nearly truncatestaminodialcup, rarely incompletely fused; pistil conical, glabrous or spinulose, with 3 sessile stigmas;ovules 3, sub-basal.Fruit globose or ellipsoid, red, or more rarelywhite, orange,or purple at maturity,glabrous,blackspinulose,or golden spiny; mesocarp fleshy to dry; endocarpblack at maturity, hard,smooth or variouslypitted-grooved,with 3 subequatorialgerminationpores each surroundedby an asteriskof applanatefibers;seed 1, globose-irregular, brown;endospermwhite, homogeneous,with a small to largecentralcavity.Embryolateral,conical.Eophyll simple, bifid, spiny,with praemorseoutermargin. Distribution. Twenty-two species distributed in the LesserAntilles, Trinidad,Venezuela,Panama,and along the Andes from Colombia to Bolivia, reaching 3000 m elev. in Colombia. Most species are found in Colombia (15) and Ecuador(11).
Key to the Species of Aiphanes 1.
Inflorescenceunbranched. 2. Leaves entire or divided into a large top segment and 1-4, 1-ribbedpinnae per side; Colombia and Ecuador. ............................................................... 14. A. macroloba 2. Leaves pinnatelydivided, with >6 pinnae per side. 3. Pinnae linear or nearly so, 6-18 times as long as wide; plants acaulescent. 4. Flowers purple;petals of staminateflowers 3-4 mm long; anthers0.3-0.4 mm long; Colombia (Choc). ................... .................................... 1. A. acaulis 4. Flowers greenish yellow; petals of staminateflowers 1.5-2.5 mm long; anthers1.1-1.4 mm long; Peru (San Martin)................................................ 18. A. spicata 3. Pinnae cuneate, 1.3-4.5 times as long as wide; plants caulescent,with a >1 m tall stem. 5. Stem caespitose, 1-2 cm diam.; leaf sheathusually with yellow spines; pinnae truncate to incised at apex; Colombia .............................................. 17. A. simplex 5. Stem solitary,3-6 cm diam.; leaf sheathwith black spines; pinnae oblique, rarely incised at apex (inflorescencenormallybranched);Ecuador.................... 3. A. chiribogensis I. Inflorescencebranched. 6. Rachillae appressedto rachis, or fastigiateand then very short. 7. Rachillae fastigiate, very short, slender;NW Colombia............................. 16. A. parvifolia 7. Rachillae appressedto and often partlyadnatewith rachis,thickenedin the basal part. 8. Middle pinnae45-60 cm long, linearto narrowlycuneate,5-12 times as long as wide; inflorescences 1 or 3 per node; W of the Andes......... .... ................... 8. A. gelatinosa
Systematic Treatment
Middle pinnae 15-35 cm long, cuneateto broadlycuneate, 1-4 times as long as wide; 20. A. ulei inflorescences 1 per node; E of the Andes ....................................... Rachillae spreadingor pendulous. 9. Proximalrachillaependulous,with flowers for <2/3 of their length. 10. Rachillae 10-25 mm diam. in pistillatepart,covered with black spines; pistillate flowers 5. A. duquei 13-20 mm long; Colombia ................................................. 10. Rachillae 2-6 mm diam. in pistillatepart,shortly spinulose or with scatteredspines; 3. A. chiribogensis pistillate flowers ca. 4 mm long; Ecuador ................................. 9. All rachillaespreading,with flowers for >23 of their length. 11. Pistil spinulose; fruit spinulose or spiny. 13. A. linearis 12. Fruitcovered with black to golden spines; leaves distichous;Colombia .......... 12. Fruitwith small black spinules;leaves polystichous. 13. Pinnaelanceolateto linear,widest in the middle or equally wide in the middle and at the apex. 14. Stem solitary, 10-21 m tall; fruitdull green, 20-23 mm diam.; anthers 9. A. grandis 3-4 mm long; SW Ecuador ..................................... 14. Stem caespitose, 4-10 m tall; fruitred, 9-11 mm diam.;anthers0.9-1.2 10. A. hirsuta mm long; NW Colombia........................................ 13. Pinnae cuneate,widest at apex. 15. Pinnaeabruptlywidening at apex; stem solitary;anthers2.5-3 mm long; 2. A. aculeata N Colombia (CordilleraOccidental).............................. 15. Pinnaenot abruptlywidening near apex; stem caespitose;anthers<1 10. A. hirsuta mm long; W Colombia(CordilleraCentraland Oriental) .............. 11. Pistil glabrous;fruit glabrous. 16. Pinnaeregularlyinserted;acaulescentor with a large solitarystem. 17. Stem 0-2 m tall, 2-6 cm diam. pinnae<30 cm long; E Ecuadorand Peru... 22. A. weberbaueri 17. Stem 2-21 m tall, 6-20 cm diam.;pinnae(30-)50-80 cm long; West Indies .... 15. A. minima 16. Pinnaeinsertedin groups;acaulescentor with a solitaryor caespitose stem. 18. Pinnae lanceolateto linear,widest in the middle or equally wide in the middle and at apex. 7. A. erinacea 19. With yellow and black spines; pinnae>50 cm long; W Colombia........ 19. With black spines; pinnae <50 cm long. 20. Stem solitary,0-2 m tall; leaves polystichous;E Ecuadorand Peru, 22. A. weberbaueri below 2000 m ....................................... 20. Stem caespitose, 2-7 m tall; leaves often distichous;Colombiato S Ecuador,usually above 2000 m altitude. 21. Fruitgreenishwhite, becoming brownverrucoseat apex; 21. A. verrucosa pistillateflowers 9-11 mm long; S Ecuador ............... 21. Fruitred, smooth;pistillateflowers 3-7 mm long; Colombia .... 12. A. lindeniana 18. Pinnae narrowlyto broadlycuneate,widest at apex. 22. Pinnae usually abruptlywidening nearapex; anthers>1 mm long. 23. Stem solitary;pinnae20-40 per side; middle pinnae 8-25 cm wide at apex, usually tricuspidate;Trinidadto Bolivia, not in Ecuador ..... 2. A. aculeata 23. Stem caespitose;pinnae(30-)50-65 per side; middle pinnae 4-9 6. A. eggersii cm wide at apex, irregularlypraemorse;W Ecuador ............... 22. Pinnaenot abruptlywidening near apex; anthers<1 mm long. 24. Pinnaetricuspidate;stem solitary;usually with some yellow spines; 19. A. tricuspidata Colombiaand Ecuador,W of the Andes..................... 24. Pinnaenot tricuspidate;stem solitaryor caespitose;spines black or yellow. 25. Pinnaetruncateto incised at apex (shorteralong the midrib than along the proximalmargin). 26. Stem solitary,usually <2 m tall; pinnae<5 cm wide at 19. A. weberbaueri apex; SE Ecuadorto Peru ........................ 26. Stem caespitose, usually >2 m tall; pinnae up to 15 cm wide at apex; Colombiato Ecuador. 27. Pinnaeinsertedin groupsof 4-5, usually with a row of slenderspines adaxiallyon the midrib; 12. A. lindeniana N Colombia ............................... 27. Pinnaesubregularlyinsertedor in groupsof 2-3, neverwith a row of spines adaxiallyon the midrib. 28. All spines black; rachillaeglabrous;pinnae insertedin lax groupsof 2-4, in one plane; 11.A. leiostachys Colombia(MagdalenaValley)............... 8.
6.
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Flora Neotropica
36
28. Some spines yellow; rachillaecovered with brownto purplespinules;pinnae insertedin widely separated groupsof 2-3, usually in severalplanes; Colombia (W of CordilleraCentral)and Ecuador. 29. Rachillae>15; SW Colombiaand Ecuador. .. 7. A. erinacea 29. Rachillae<15 (if middle pinnae >40 cm long then see notes to A. erinacea); Colombia .... 17. A. simplex 25. Pinnaeoblique to lobed at apex (at least as long along the midrib as along the proximalmargin). 30. Pinnaeconspicuouslylobed at apex; endocarpwith deep 10. A. hirsuta pits; SW Colombiato NW Ecuador . ................... 30. Pinnaeoblique to weakly lobed at apex; endocarpnot with deep pits. 31. Pinnaeusually in groups of >4, with a row of slender spines adaxiallyon the midrib;caespitose with up to 10 stems, to 7 m tall; Colombia,usually above 2000 m................... ................ 12. A. lindeniana 31. Pinnaein groupsof 2-3, or subregularlyinserted, never with spines adaxiallyon the midrib;solitary or caespitose, often acaulescent;SE Colombiato Peru, below 2000 m. 32. Petiole 90-105 cm long; leaf rachis 140-190 cm long; rachillae50-60, slender;Colombia, Brazil, and Peru ................................ 4. A. deltoidea 32. Petiole 13-46 cm long; leaf rachis 56-120 cm long; rachillae8-35, thickenedin the androgynouspart; Ecuadorand Peru ....................... 22. A. weberbaueri
Multi-access Key to the Species of Aiphanes Numbersreferto the numericallist of taxa. Numbersin parenthesesindicatethatthe characteris found only rarely in that taxon. Numbersin squarebracketsindicatethat informationaboutthe characteris lacking. Habitat 1. Altitude a. <700 m 1, 2, 4, 6, lOa, 14, 15, 19, 20, 22 b. 700 m-2000 m 2, 3, 4, 6, 7, 1Oa,14, 15, 19, 20, 22 c. >2000 m 3, 5, 7, 10c, 12, 13, 17, 21 2. Geography a. Lesser Antilles 15 b. Venezuelaand Trinidad 2 c. Panama lOa d. Colombia 1, 2, 4, 5, 7, 8, 1Oa,10b, 10c, 10d, 11, 12, 13, 14, 16, 17, 19, 20 e. Ecuador 3, 6, 7, 8, 9, 10d, 14, 19, 20, 21, 22 f. Peru 2,4, 18, 20, 22 g. Brazil 2,4,20 h. Bolivia 2 i. Amazonianlowlands (<1000 m) 2, 4, 20, 22 j. Lowlands west of the Andes (<1000 m) 1, 6, 7, 8, 10a, 10d, 14, 16, (17), 19
Systematic Treatment Vegetative characters 3. Habit a. Solitary 1, 2, 3, 4, 5, 8, 9, (10c), (10d), 15, 16, 18, 19, 20, 22 b. Caespitose 4, 6, 7, 8, 10a, 10b, 10c, 10d, 11, 12, 13, 14, 17, 21, 22 4. Stem height a. Subterranean(acaulescentpalms) or <0.5 m tall 1, 3, 4, (14), 18, 20, 22 b. 0.5-4 m tall (2), 3, 4, 5, 6, 7, O1a,12, 14, 15, 16, 17, 19, 20, 21, 22 c. 4-10 m 2, 5, 6, 7, 8, (9), 10a, 10b, 10c, 10d, 11, 12, 13, 15, 16, (17), (19), 20, 21 d. >10 m tall (2), 9, 15 5. Stem thickness a. <4 cm diam. [1], 3, 4, 5, 7, 1a, 11, 12, 14, 16, 17, 18, 19, 20, 21, 22 b. 4-9 cm diam. [1], 2, 3, 4, 5, 6, 7, 8, IOa, 10b, 10c, 10d, 12, 13, 15, 18, 19, 20, 21, 22 c. >9 cm diam. 2, 8, 9, 10c, 10d, 12, 13, 15 6. Stem armature a. Spines <10 cm long all b. Spines >10 cm long 5, 8, 9, 10a, 10b, 10c, 10d, 13, 15 7. Phyllotaxis a. Leaves polystichous all except 13, 21 b. Leaves distichous 12, 13, 21 8. Leaf division a. Leaves simple 14 b. Leaves pinnatelydivided all 9. Color of spines on sheath a. Black all b. Grey 2,4,6 c. Yellow 7, 17, 19 10. Length of leaf rachis a. <80 cm 1, 3, 10a, 12, 14, 16, 17, 18, 19, 20, 22 b. 80-150 cm 3, 4, 5, 6, 7, 8, 10a, 10b, 11, 12, 13, 15, 19, 20, 21, 22 c. >150 cm 2, 4, 6, 7, 8, 9, 10b, 10c, 10d, 12, 13, 15, 20 11. Numberof pinnae per side a. <10 3, 7, O1a,14, 16, 17, 20, 22 b. 11-24 1, 3, 4, 5, 7, 8, O1a,11, 12, 15, 16, 17, 18, 19, 20, 22 c. >24 1, 2, 5, 6, 8, 9, O1a,10b, 10c, 12, 13, 15, 21 12. Arrangementof pinnae, grouping a. Regularlyinserted 1, 8, 14, 15, 22 b. In groups of >6 (2), 6, 9, (10b), 12, 13, 21
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Flora Neotropica c. In groups of 4-6 2, (3), 5, 6, (7), 8, 9, 10a, 10b, 10c, 10d, 11, 12, 13, (17), (20), 21, (22) d. In groups of 2-3 (2), 3, 4, 5, 7, 10a, 10d, (11), 12, 16, 17, 19, 20, 22 13. Three-dimensionalarrangementof pinnae a. In one plane 1, (3), 8, 10b, 11, 14, 15, (16), 22 b. In differentplanes 2, 3, 4, 5, 6, 7, 8, 9, 10a, (10b), 10c, 10d, (11), 12, 13, 14, 16, 17, 19, 20, 21, 22 14. Shape of middle pinnae a. Lanceolateto linear,widest in the middle or equally wide in middle and at apex 1, 8, 9, 10b, 12, 13, (14), 15, 16, (17), 18, 21, 22 b. Narrowlycuneate, widest at apex, >4.5 times as long as wide 5, (6), 7, 8, 10a, 10c, 11, 12, 13, 14, (15), 16, 17, 18, 22 c. Cuneateto broadlycuneate, <4.5 times as long as wide (2), 3, 4, 7, 10a, 10c, 10d, 11, 12, 14, 16, 17, 19, 20, 22 d. Trumpetshaped, abruptlywidening at apex 2, 6, (15) 15. Shape of pinna apex a. Truncate,rect 1, (2), (6), 10a, 10c, 12, 16, 17 b. Oblique 3, 4, 5, 8, 9, 10b, 12, 13, 14, 15, 16, 18, 21, 22 c. Incised (3), 7, 8, (10a), 11, 12, 17, 20,22 d. Lobulate 4, 10a, 10b, 10c, 10d, 15, 16, (22) e. Tricuspidate 2, 19 f. Irregular 2, 6 16. Length of middle pinnae a. <20 cm 1, 3, 4, 10a, 12, 14, 16, 17, 18, 19, 20, 22 b. 20-40 cm 1,2,3,4,5,6,7, 10a, 10d, 11, 12, 13, 15, 16, 17, 18, 19,20,21,22 c. 40.1-60 cm 2, 6, 7, 8, 10a, 10b, 10c, 10d, 13, 15 d. >60 cm 7, 9, 10c, 13, 15 17. Pinna armature a. With a row of slender spines adaxiallyon the midrib 1, 9, 12, 14, 15, 21 b. Withouta row of slender spines adaxially on the midrib 1, 2, 3, 4, 5, 6, 7, 8, 10, 11, (12), 13, (14), 15, 16, 17, 18, 19, 20, 22
Inflorescence 18. Ramificationof inflorescence a. Unbranched 1, (3), 14, 17, 18 b. Branched 2, 3, 4, 5, 6, 7, 8, 9, 10a, 10b, 10c, 10d, 11, 12, 13, 15, 16, (17), 19, 20, 21, 22 19. Textureof peduncularbract a. Thin, soon dissolving in fibers 1, 3,4, 14, 17, 18,20,22 b. Coriaceous,dissolving in fibers 2, 5, 7, 10a, 10b, 10c, 10d, 11, 15, 16, 19 c. Thick, coriaceous to woody, persistent 2, 6, 8, 9, 12, 13, 15, 21 20. Length of inflorescencerachis (or spike) a. <20 cm 1, 3, 7, 10a, 12, 13, 14, 16, 17, 18, 19, 22
Systematic Treatment b. 20-50 cm 1, 2, 3, 4, 6, 7, 8, 10a, 10b, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20, 21, 22 c. >50 cm 2, 3, 5, 7, 9, 10c, 10d, 13, 15, 19, 22 21. Numberof rachillae a. <20 3, (7), 8, 10a, 11, 12, 15, 16, 17, 19,22 b. 20-49 2, 3, 4, 6, 7, 8, 10a, 10b, 12, 13, 15, 16, 19, 20, 22 c. 50-70 2, 4, 5, 6, 7, 10c, 10d, 12, 13, 15, 16, 19, 21 d. >70 2, 6, 7, 9, 10c, 10d, 13, 15, (16) 22. Position of rachillae a. Spreading 2, (3), 4, 5, 6, 7, 9, 10a, 10b, 10c, 10d, 11, 12, 13, 15, 17, 19, 20, 21,22 b. Fastigiateor appressed 8, 16, 20 c. 1-4 basal rachillaependulous,widely separated 3, 5 23. Length of longest rachilla a. <30 cm 2, 6, 8, 10a, 11, 12, 13, 15, 16, 19, 20, 21, 22 b. >30 cm 2, 3, 4, 5, 6, 7, 9, 10a, 10b, 10c, 10d, 11, 12, 13, 15, 19, 22 24. Length of basal flowerless partof proximalrachillae a. <6 cm 2, 4, 6, 7, 8, 9, 10a, 10b, 10c, 10d, 11, 12, 13, 15, 16, 19, 20, 21, 22 b. 6-15 cm 3, 7, 10b, 10d, 12, 16, 22 c. >15 cm 3, 5, (7), (12) 25. Rachilla armature a. Withoutspinules or spines 2, 6, 11, 15, (20) b. With spinules, without spines 1, (2), 3, 4, 7, 9, 10a, 10c, 10d, 12, 13, 14, 16, 17, 18, 19, 20, 21, 22 c. With spines 2, 5, 8, 10a, 10b, 10c Flowers 26. Color of staminateflowers a. White, cream, yellow, or greenishyellow, neverrose or violet in bud 2, 6, 9, (10b), 11, 12, 14, 15, 17, 18, 20, 21 b. Rose or purple,at least in bud, often white at anthesis 1, 3, 5, 7, 10a, 10b, 10c, 10d, 11, 12, 13, 16, 17, 19, 22 c. Orange 4, 14 27. Length of anthers a. >2 mm 2, 5, 9, 15, 21 b. 1-2 mm 2, 5, 6, 10b, 12, 13, 15, 18, 21 c. <1 mm 1, 3, 4, 7, 8, 10a, 10c, 10d, 11, 12, 13, 14, 16, 17, (18), 19, 20, 22 Fruit 28. Color a. Red to orange [1], 2, 3, [4], 5, 6, 7, 10a, 10b, 10c, 10d, [11], 12, 14, 15, 16, 17, [18], [20], 22 b. Brownishred and speckled [1], [4], [11], [18], 19
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Flora Neotropica
c. White or greenish white [1], 2, [4], (10a), [11], [18], [20], 21 d. Purpleto black [1], (2), [4], (10a), [11], [18], [20], 22 e. Dull green [1], [4], 9, [11], [18] f. Golden due to spine cover 13 29. Diameter a. <10 mm [1], [4], 5, 7, lOa, [11], 14, 16, 17, 18, 19, 20, 22 b. 10-15 mm [1], 2, 3, [4], 5, 8, 10a, 10b, 10c, [11], 12, 14, 15 c. 15.1-20 mm [1], 2, [4], 6, 9, 10d, [11], 12, 15 d. >20 mm [1], 2, [4], 9, O1d,[11], 13, 21 30. Indument(equally well observedon pistil) a. Glabrous 1,2,3,4,5,6,7,8, 10a, lOd, 11, 12, 14, 15, 16, 17, 18, 19,20,21,22 b. With spinules (2), 9, 10a, 10b, 1Oc c. With spines 13 31. Endocarp a. Smooth to moderatelypitted [1], 2, 3, [4], 5, 6, 7, 8, 9, 10a, lOd, [11], 12, 13, 14, 15, 16, 17, [18], 19, 20, 21, 22 b. Longitudinallygrooved and/orwith deep pits [1], [4], 10c, 10d, [11], [i8]
1. Aiphanes acaulis Galeano& Beral, Principes29: scattered short spinules; peduncle 12-88 cm long, 20, fig. 1 p. 21. 1985. Type. COLOMBIA.Choco: 3-5 mm diam. at junction with rachis, with scattered Mun. de El Carmen de Atrato, road Medellin- short spinules;spike 15-31 cm long, 5-6 mm diam., Quibd6, km 151, VeredaEl Doce, Rio Atrato, left with minute,brown spinules, with triadsfor ca. /2 of bank, 700 m, 5 Jan 1980 (fl), Bemal & Galeano 71 the length, the remainingpart with staminatedyads; (holotype, COL; isotypes, HUA, K). flower groups sunken,pistillateflowers sunkenfor /2 of their length or more. Staminateflowers purple, Solitary, acaulescent.Leaves 8-10, erect and arch- 1.5-2.5 mm long; sepals arched,carinate,1-1.5 x ca. ing, in vivo darkgreen; sheath 11-16 cm long, with a 2 mm; petals shortly connate at base, 1.5-2 x 1-2 brown, scaly indument,covered with purplish,to 0.5 mm; filamentsca. 0.3 mm long, anthersoval, 0.3-0.4 mm long spinules, usually also with a few black, x 0.3-0.5 mm; pistillode and receptacle forming a short spines; petiole 12.5-50 cm long, with indument disc-shaped structure,ca. 0.6 mm high and 1.2 mm and spinules as on the sheath;rachis 42-75 cm long, diam.Pistillateflowers ca. 3 mm long; sepals broadly densely covered with purplishblack, to 5 mm long ovate to reniform,1-1.5 x 3-4 mm; petals connatefor spinules, sometimes with a few, to 2 cm long spines; ca. /2 of their length, valvate distally, 2-2.5 x 2-2.5 pinnae 18-30 per side, regularlyinserted, all in one mm; staminodialcup 1.5-2 mm high, truncatewith 6 plane, linear or very narrowly cuneate, 6-10 times minuteteeth;pistil glabrous.Fruitnot seen. as long as wide, strongly plicate, rigid, truncateIllustrations. Fig. 11D (pollen), 19E (distribution praemorse at apex, with a 1-4 cm long finger-like 21M (middle pinnae); Galeano and Bernal, on the distal adaxial side often map), projection margin, with a row of black, thin, 5-10 mm long spines along 1985: 21, fig. 1. the midrib, abaxial side with scatteredsimilar spines Distribution and habitat. Endemic to western and 1-3 rigid spines on the basal part of the midrib; basal pinnae 7-24.5 x 0.71 cm; middle pinnae 10-27 Colombia,where only two small populationshave so x 1.7-2.5 cm; apical pinnae 1-3 ribbed, 6.5-15.5 x far been located, both in the departmentof Choc6, in 4-6 cm. Inflorescenceerect, spicate; prophyll 16-26 wet lowlandto premontanerain forest at 150-700 m. x 0.7-1.1 cm; peduncularbract37-94 cm long, with (text continuedon page 46)
SystematicTreatment
41
B
20cm
A
FIG. 21. A-E Middle pinnae. A. Aiphanesmacroloba(Skov & Borchsenius64747). B. A. linearis (Gentryet al. 40805). C. A. minima(Lee s.n.). D. A. lindeniana (Gentryet al. 52978). E. A. eggersii (Skov & Borchsenius64735).
Flora Neotropica
42
20 cm
/
ri'
FIG. 21 F-J. Middle pinnae.F. Aiphanesgrandis (Madsen86927). G. A. gelatinosa (Barfod60001). H. A. duquei(Bernal & Devia 1540). I. A. gelatinosa (Bernal & Galeano 901). J. A. leiostachys (Henao et al. 299).
43
SystematicTreatment
. ._..~-
~.
N
,
~
20 cm
K
ML
M
rAd
FIG. 21 K-Q. Middle pinnae. K. Aiphanes chiribogensis (Skov et al. 64822). L. A. simplex (Cuatrecasas 18377). M. A. acaulis (Bernal & Galeano 71). N. A. tricuspidata(Borchsenius & Luteyn 91425). 0. A. erinacea (Borchsenius 91421). P.A.. aculeata, Skov et al. 64839). Q. A. parvifolia (Callejas et al. 4238).
44
Flora Neotropica
CM
?20
a
~~~~~~~~~~~~~~~~~~~~~.
a~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ ~ ~
N~
R
~~~~~~~~~~r
',r
Tid
FIG. 21 R-W. Middle pinnae. R. Aiphanes spicata (Kahn & Borchsenius2651). S. A. ulei (Balslev et al. 62045). T. A. weberbaueri(Asplund14667). U. A. weberbaueri(Kahn& Borchsenbius2546). V. A. weberbaueri(Borchsenius& Pedersen 91427). W. A. deltoidea (Kahn & Borchsenius2556).
45
SystematicTreatment
A'~~~~~~~.
-
-
0/-00,~ ~
~ ~
--I
~~~~~~~~~~~..? .~,
"/
~~~~~AA
'"~~~~~~~~~~~~~~~~~~~~~~~
20 cm
Y FIG. 21 X-Z, AA. Middle pinnae of Aiphanes hirsuta. X. Subsp.fosteriorum(Skov et al. 64819). Y. Subsp. intermedia (Dransfield4854). Z. Subsp. kalbreyeri(Bernal & Tobdn1393). AA. Subsp. hirsuta(Bernal & Tobdn1396).
46
FloraNeotropica
Mun.de Pt). Lectotype(here designated).Martius,Palmetum Specimensexamined.COLOMBIA.CHOC6: tab.2, fig 1., 1847. km 151, Orbignianum, El Carmende Atrato,road Medellin-Quibd6, in ex Martius)H. Wendland veredaEl Doce, Rio Atrato,left bank,700 m, 5 Jan 1980 Aiphanestruncata(Brongniart Les Palmiers230. 1878. de Dentherghem, (fl), Bernal& Galeano72 (COL,HUA);Mun.de Quibd6, Kerchove SanFranciscode Ich6,Rio Ich6,left bank,150 m, 11 Jul MartineziaelegansLinden& H. Wendland,Linnaea28: 351. 1857.Type.Colombia("NewGrenada"): 1981 (fl), Galeano & Bernal 454 (COL, HUA, K); ca. 5 km (juv),Funck ? "Martinezia on the rd. to Medellin,150 m, 7 Jul 1986 & Schlims.n. (holotype,L, sheetannotated E of Tutunendo (infl), Bernal et al. 1079 (AAU, COL, FTG).
elegans Herm. Wendl., Neu Grenada,Linden coll."; iso-
Lindencoil."). type,? K, sheetlabeled"NeuGrenada, in H. Wendland Aiphanes acaulis is very characteristicin its acau- Aiphaneselegans(Linden& H. Wendland) LesPalmiers230. 1878. de Denterghem, lescent habit, linear or narrowly cuneate, regularly Kerchove Mararabicuspidata Karsten,Linnaea28: 390. 1856.Type. inserted, strongly plicate, rigid pinnae, and spicate Barinas:BetweenBarinasand Barinitas, VENEZUELA. inflorescence. In life form and general appearanceit WU). (fl), Karstens.n. (lectotype,heredesignated, resembles the PeruvianA. spicata; the differencesbe- Martineziaernestii Burret,Notizbl. Bot. Gart. BerlinuleiDammer, Dahlem11:327. 1932(basedon Martinezia tween the two are discussed underthe latter(see also Nom. Illeg.,Notizbl.Konigl.Bot. Gart.Berlin59: 266. Table II). 1915 non M. ulei Dammer,Verh.Bot. VereinsProv. 48: 127.1907).Type.PERU.Madrede Dios: Brandenburg Alto Acre, SeringalAuristella,Ule 116b (Bt). Neotype 2. Aiphanes aculeata Willdenow, Samml. Deutsch. (here designated).PERU.Madrede Dios: Tambopata, Abh. K6nigl. Akad. Wiss. Berlin 1803: 251. 1806; Barsola,Rio Piedras,350 m, 17 Jan 1967 (fl), Vargas Willdenow,Mem. Acad. Roy. Sci. Berlin 1804: 33. 18694(holoneotype, BH;isoneotype,CUZ-n.v.). 1807. Type. VENEZUELA. Miranda:Near Cauca- Aiphanesernestii (Burret)Burret,Notizbl. Bot. Gart. Berlin-Dahlem 11:560. 1932. gua, Bredemeyer s.n. (Bt). Neotype (here desig- Martinezia Burret,Notizbl. Bot. Gart. Berlinkillipii nated).VENEZUELA.Miranda:Dtto. Bri6n, along Rio Santander: Dahlem11:326. 1932.Type.COLOMBIA. andJaboncillo,800-1500 quebradaAgua Bendita,a tributaryto Rio Aricagua, Surata,betweenBucaramanga 2.3 km E of Pueblo Seco, 4.6 km E of Aricagua,75 m, Jan 1927 (fl, fr), Killip & Smith 16362 (holotype, Bt; NY;isotypes,A, BH,GH). lectotype,heredesignated, & Mar 1973 24-25 m, (fl), Steyermark Espinoza Burret,Notizbl.Bot.Gart.Berlin106916 (holoneotype, BH; isoneotypes, NY, VEN). Aiphaneskillipii(Burret) Dahlem11:561. 1932. Sprengel,Syst.Veg.2: 140. AiphanesorinocensisBurret,Notizbl.Bot. Gart.BerlinEuterpeaculeata(Willdenow) Orinoco(fem Dahlem11:560. 1932.Type.VENEZUELA: 1825. fl), Rusby&Squiress.n.(holotype,Bt;lectotype,heredesMartineziaaculeata (Willdenow)Klotzsch,Linnaea20: 455. 1847. ignated,NY). 77. MartineziaaiphanesMartius,Palmetum Orbignyanum 1847. Solitary;stem 3-10 m tall, 6-10 cm diam., in culMararaaculeata(Willdenow) Karsten,Fl. Columb.2: 143, tivated specimens up to 15 m tall and 15 cm diam., tab.175,figs. 10-15. 1866. armed with grey to black, to 10 cm long, flattened CaryotahorridaJacquin,Fragm.Bot.20. 1809. spines on the intemodes. Leaves 10-15, spreading, Bot. Gart. Notizbl. Aiphaneshorrida(Jacquin)Burret, the lower ones recurved;sheath60-90 cm long, with 11:575. 1932. Berlin-Dahlem Martinezia caryotifolia Humboldt, Bonpland, & Kunth, spines similar to those on stem; petiole almost lackNov. gen. et sp. 1: 305. 1816 ("caryotaefolia"). Type. or to 25 cm long, armedlike sheath,but spines COLOMBIA.Tolima: Ibague,Humboldt& Bonplands.n. ing up rachis 175-200 cm long, with a white or grey, shorter; (holotype,P). Marara caryotifolia (Humboldt, Bonpland, & Kunth) caducous indument, armed with numerous black, Karsten,Fl. Columb.2: 134,t. 170. 1866. applanate spines, to 5 cm long; pinnae 25-40 per Aiphanes caryotifolia (Humboldt,Bonpland, & Kunth) H. side, insertedin groups of (2-)4-6(-10), in different Les Palmiers230. Wendland in Kerchovede Denterghem, planes, groups occupying 8-18 cm along the rachis, 1878. Tilmiacaryotifolia(Humboldt, Bonpland,& Kunth)Cook, separated by 10-25 cm, pinnae abruptly widening Bull.TorreyBot.Club28: 565. 1901. near apex, 1-5 times as long as wide, tricuspidateor BactrispraemorsaPoeppigex Martius,PalmetumOrbigrarely truncate-bicuspidateat apex, glabrouson both nyanum66. 1847.Type.PERU.San Martin:Tocacheon or sides, minutely spinulose abaxially,rarelydensely RioHuallaga,Jul1830,Poeppigs.n.(holotype,M,sterile). Aiphanespraemorsa(Poeppigex Martius)Burret,Notizbl. covered with long spinules on both sides, often with 11:560. 1932. Bot.Gart.Berlin-Dahlem a row of black, to 5 mm long spines along the marMartineziatruncataBrongniartex Martius,Palmetum gins; basal pinnae quite variable, 9-28 x 2-8 cm; 75, tab.2, fig. 1, and tab.28 C, fig. 1-7. middle Orbignianum pinnae 21-52 x 8-25 cm; apical pinnae 4-7 1847.Type.BOLIVIA.Cochabamba: Yungasdel Palmar de la Palma"),Rio de Chajro,Rio Suri,Rio San ribbed,10-23 x 9-25 cm. Inflorescenceerect or curv("Yungas Mateo,Rio Reuni6n,1830-1833,d'Orbigny8 (holotype, ing, pendulousin fruit,once or rarelytwice branched;
47
SystematicTreatment Table II Aiphanesspicata andA. acaulis compared Character
A. spicata
A. acaulis
Insertionof pinnae Numberof pinnae per side Middle pinnae size (cm) Middle pinnae 1/wratio Spines adaxiallyon midrib Color of flowers
Groupeddistally 14-16 17-22 x 1-2.5 10-18 Absent Green-yellow
Regularthroughout 18-30 10-27 x 1.7-2.5 6-12 Absent or to 20 mm long Purple
Reticulate,smooth 4-5 Ca. 2000
Perforate,clavate Ca. 3 150-700
Stam.petalslength(mm) Antherasize (mm) Pollen exine Pist. flowers length (mm) Altitude (m)
3-4 1.1-1.4x 0.9-1.1
prophyll,peduncularbractandpedunclewith a dense, white or grey indument;prophyll30-35 cm long, 3-8 cm wide; peduncularbract insertedjust above prophyll or on the distal half of peduncle, 80-140 cm long, 4-8 cm wide, coriaceousto woody, unarmedor spiny; peduncle 30-150 cm long, 3-20 mm diam. at junction with rachis, densely armed with black, to 5 cm long spines; rachis 20-75 cm long, unarmedor proximallyarmedlike peduncle,rarelyspiny throughout; rachillae 25-170, unarmedor rarelyspiny,with a peltate indument,often with distinct swellings below the flower-groups; basal rachillae 8-40 cm long, sometimes with a basal flower less part up to 6 cm long, with triadsfor 2/3 of their length or less, distally staminate,sometimes only with a few triadsclustered near the base; apical rachillae 3-10 cm long, staminate, or sometimes with a few triads at base; each triad subtendedby a small bract,sometimescovering the pistillateflower for up to /4 of its length;eachdyad subtended by a minute or up to 1 mm long bract. Staminateflowers white, greenish yellow, or yellow, sometimes fragrant,superficial;sepals nearly free or partlyconnate,narrowlytriangular,carinate,neverexceeding >/4 of the petals, 1-2 mm long, with a minute spur at base; petals free or briefly connate at base, 2.5-5.5 mm long; filaments 1-2 mm long, flattened; anthers linear, (1.1-)1.5-3.3 mm long, 0.6-1 mm wide, connectivedark;pistillode0.5-1 mm high,trifid. Pistillate flowers superficial,or sunken into shallow pits in the rachillae;sepals 1.5-5 mm long, imbricate, dark, cartilaginous;petals 3-7 mm long, almost free or connate for 1/2 their length, corolla lobes often imbricate basally, valvate distally; staminodialcup 2-4 mm high, deeply lobed, lobes acute;pistil glabrousor occasionally covered with pale, soft spinules. Fruits bright red, rarely orange or white, (10-)16-23 mm
1.5-2.5 0.3-0.4 x 0.3-0.5
diam., smooth or occasionally with scattered,caducous spinules, 1-2 mm long; mesocarporange,mealyfleshy; endocarp(9-)13-20 mm diam.,nearlysmooth to shallowlypitted. Illustrations.Figs.4A (stemspines),7A, C (flowers), 9D (seedling plant), 10C, 14B, C (pollen), 18A (distributionmap), 21P (middlepinnae);Humboldtet al., 1825: tab. 699; Martius,1839: tab. 161, fig. 1, 1-12; Martius,1847: tab. 2, fig. 1, and tab. 28 C, figs. 1-7 (as Martinezia truncata); Drude, 1882: tab. 85; Hooker,1886: tab. 6864 (all as Martineziacaryotifolia); Karsten,1866: tab. 170 (as Mararacaryotifolia); Galeano & Bemal, 1987: fig. 5; Balslev & Moraes, 1989: figs. 8, 9, pp. 27, 28 (both as Aiphanescaryotifolia). Also photosin severalpopularbooks on palms: Blomberry& Rodd, 1982: fig. 47; Braun, 1968: fig. 17; Hoyos & Braun,1984: figs. 3, 5; Braun& Chitty, 1987: 47, 48, underthe synonymsA. caryotifolia and A. elegans. Local names and uses. Macagiiita, macahuite (northernVenezuela); corozo del Orinoco, corozo anchame (Bolivar); marara (western Venezuela); mararay,mararave(Llanos of Colombia); cubarro (easternColombia);chonta,pujamo,chascaraza,charascal, corozo chiquito, corozo colorado, pujamo (westernColombia);gualte(Narifio);corozo (Colombia, Ecuador); chonta ruro (Ecuador); cocos rura (Bolivia). Fruitsare edible and sold in many markets in the Magdalena and Cauca valleys in Colombia. The mesocarpis extremely rich in carotene,the precursorof vitaminA, and the seed is rich in oil (Balick & Gershoff, 1990). In some parts of the Llanos in Colombia the endocarpsof this species are used to play games.
48 Distribution and habitat. Widely distributedin western South America, from Trinidad to Bolivia. The total distributionarea is divided into two subunits. In the north it occurs from Trinidad,along the Coastal Cordillerain Venezuela (0-300 m), the eastern slopes of Cordillera de M6rida and Cordillera Orientalin Colombiasouthto the departmentof Meta (to 1500 m), and in the valleys of Rio Magdalenaand Rio Cauca, where it is commonly planted and naturalized;due to almost complete deforestationof these areas naturalpopulationscan no longer be found, but there is no doubt that the interandeanvalleys of Colombia are partof the naturalrange of the species. It has also been reported from British Guyana (Schomburgk, 1848), upper Orinoco, Rio Atabapo, and Rio Cassiquiarein Venezuela (Humboldtet al., 1816), as well as El Dorado and furthersouth on Rio Caroni (Braun & Chitty, 1987), but there are no herbariumvouchers to document these reports.The southern distributionalrange includes eastern Peru, from the departmentof San Martin and southward throughHuanucoand Madrede Dios, reaching 1300 m in the departmentof Cuzco, westernBrazil,andthe easternAndean slopes in Bolivia reaching 1700 m in the departmentof La Paz and 900 m in Santa Cruz. Aiphanesaculeata has a preferencefor seasonalto dry tropical forest but occurs also in humid forest types. The gap in the distribution,from southeasternColombia to northernPeru, neverthelesscoincides with the part of the upper Amazon richest in precipitation. Aiphanesaculeata is widely plantedas a gardenornamentaland in botanicalgardensthroughoutthe world. Specimens examined. COLOMBIA. ANTIOQUIA:
Ituango,HaciendaSanJuande Rodas,LaHelenaForest,30 Mar1980,Sanchez12 (HUA);rd.Venecia-Bolombolo, km 3, 1400 m, 9 Dec 1979 (fl, fern fl, fr), Bernal & Galeano 50
La Doradaon Rio Magdalena, Jul (COL,HUA).CALDAS:
1898 (st), Thereses.n. (M). CASANARE: Tabl6nde Tamara,
FincaLa Laguna,610 m, 10 Mar1991 (fl, fr), Bernal& Borchsenius1956 (AAU, BH, COL,NY); rd. Tabl6nde km9, 700 m, 10 Mar1991(fl, fr),Bernal Tamara-Tmnara,
& Borchsenius 1959 (AAU, BH, COL, NY). CUNDINAMARCA: Junca, 1400 m, Nov 1856 (st), Triana1767 (COL); Sasaima, vereda las Mercedes, 1300-1400 m, 27 May 1976 (juv), Garcia Barriga 20933 (COL). META:Villavicencio, 300 m, Jan 1856 (st), Triana1765 (COL). QUINDIO: Quimbaya, vereda Palermo, finca La Cascada, 1150 m, 3 Aug 1990 (im fr), Velez et al. 2474 (HUQ). TOLIMA:Mariquita, 1845 (st), Linden1202 (G); El Libano,trailto Murillo, 1470 m, 10 Dec 1939 (fl), Garcia Barriga 8443 (COL). VALLE: Las Juntasdel Dagua (fl, fer fl), Lehmann5290 (K); E of
Zarzal,fromRioAguaBonitato RioVieja(femfl),Pennelet al. 8576 (A, NY, PH);haciendaMediaCanoa,Rio Paila, between Caserio La Paila and Zarzal, 1000 m, 6 Apr 1986 (fem fl, fr), Gentry et al. 54157 (K, MO); Puerto Caldas, 800-900 m (fl), Killip & Hazen 11039 (US); Caicedonia,
FloraNeotropica 1300 m, 27 Jan 1946 (fl), Duque 4587 (COL). UNKNOWN DEPARTMENT:(fr), Dawe s.n. (K); Doyle 12 (BH); (fl), Pur-
die s.n. (K); Unknowncollector (K); Triana730 (BM); Isla Brava(Magdalena),Oct 1875 (st), Andre400 (F, K, NY). VENEZUELA. BARINAS:Rd. Barinas-Barinitas,near Barinitas (fl, fr), Wessels Boer 1991 (NY, U). MIRANDA: Cerros del Bachiller, near E end, margins of Quebrada Corozal, S of Santa Cruz, 50 m, 16 Mar 1978 (fl, im fr), Steyermark& Davidse 116220 (MO, VEN); Dtto. Paez, fila El Guapo,nearrepresaEl Guapo, 1 Jun 1977 (fl), Gonzales & Davidse 872 (BH, MO, VEN); ParqueNacional Guatobo, ravine between Alcabala station at Macanilla and Fila de Culebra(Montbrun),between Santa Teresa and Altagracia de Orituco,28 Nov 1961 (st), Steyermark90137 (NY,VEN). SUCRE:Peninsulade Paria, vicinity of CristobalCol6n (= Macuro), the Balc6n, Jan-Feb 1923 (fl), Broadway 623 (GH, NY); (st), 754 (GH, NY). TRINIDAD.MaracasValley,3 Feb 1927 (st), Wall122 (S). PERU. Cuzco: Prov. Calca, Pucara,Valle Lares, 1300 m, 18 Apr 1966 (fr), Vargas17375 (BH); Prov. La Convenci6n, RosarioMayo, 1050 m, 16 Oct 1968 (fl), Chdvez311 (BH); Sahuayaco,850 m, 18 Jan 1947 (fl), Vargas6333 (F). HUANUCO:Prov.Pachitea,Dtto. Honoria,Bosque Nacional de Iparia, along Rio Pachitea near campamiento Miel de Abeja, 1 km above the village Toumavistaand ca. 20 km above the confluence with Rio Ucayali, 300-400 m, 26 Jun 1967 (fl, fr), Schunke 2078 (F). MADRE DE DIOS: Prov. Mand,ParqueNacional del Mand, Rio Mani, Cocha Cashu station, 350 m, 5 Jul 1984 (st), Foster 9579 (USM); Prov. Tahuamanu,Iberia, beside Rio Tahuamanu,180 m, 8 Jun 1960 (st), Moore et al. 8564 (BH). UNKNOWN DEPARTMENT: (fem fl), Pav6n s.n. (M). BRAZIL. ACRE:Mun. de Mancio Lima, Rio Moa, 5 min. downstreamsfrom confluence of Rio Azul, 17 Oct 1989 (im fr), Hendersonet al. 1145 (NY); Xapuri,Seringal Cachoeira,35 km SSE of Xapuri, 13 Oct 1989 (st), Pinard 847 (NY). BOLIVIA. BENI:Rurrenabague,Oct-Dec 1930 (fl, im fr), Fleishmann367 (S). LA PAZ:Prov.Itturalde,San Buena Ventura,500 m, 18 Apr 1901 (fem fl), R. S. Williams394 (BM, K, NY); Copacabana,10 km S of Mapiri,850-950 m, 8 Oct 1939 (fl), Krukoff11130 (GH, NY); 850 m, 25 Nov 1906 (st), Buchtien344 (A, G); Tuiri over the left bank of Rio Mapiri,500 m, 12 Sep 1939 (fl, fem fl), Krukoff10907 (A, F, MO, NY); Guanay,610 m, May 1866 (fl), Rusby2862 (NY); Prov. Nor Yungas, rd. La Paz-Caranavi, 1 km past Yolosa, 1330 m, 9 Jul 1987 (fl), Moraes & Balslev 836 (AAU, LPB); 28 km from Challalla & 10 km before San Silverio, 870 m, 10 Jul 1987 (fr), Moraes & Balslev 837 (AAU, LPB); km 12 before Caranavi,900 m, 9 Jun 1990 (infl), Moraes, Borchseniuset al. 1314 (AAU, LPB); km 11 before Caranavi,750 m, 5 Dec 1985 (fl), Henderson & Solomon 520 (NY); rd. Caranavi-Guanay,km 2, 600 m, 8 Jun 1990 (fr), Moraes,Borchseniuset al. 1312 (AAU, LPB); km 20, 600 m, 7 Jun 1990 (fr), Moraes, Borchseniuset al. 1311 (AAU, LPB); Alto Israel, 28 km from Caranavi,turning left from the rd. towardsAlto Beni, 1700 m, 11 Jul 1987 (fl, fem fl), Moraes & Balslev 842 (AAU, LPB); Prov. Sur Yungas, rd. Chulumani-Ocobaya,km 5, 1700 m, 28 Jun 1985 (fl), Beck 12115 (LPB); San Bartolomenear Calisana in valley of Rio Boopi, 850 m, Jul 1939 (im fr), Krukoff 10021 (F, NY); Cafamino, MulfordBiological Exploration of the Amazonbasin, 1300 m, Jul 1921 (fl), White405 (NY). PANDO:Prov. Nicolas Suarez, Mukden,2000 m, Sep 1979
SystematicTreatment (st), Izawa 36 (MO). SANTACRUZ:Prov. Florida, 1 km W
900 m, of Bermejo,0.5 kmW of bridgeoverRioColorado, 8 Dec 1988(im fr),Nee 37066(NY);Prov.Ichilo,ca. 3-4 km S of SanRafaeland0.5 km N of SanSalvador,11 km (byair)SWof VillaGermanBusch,600 m, 19Nov 1988(fl, fr), Nee & Salidas 36870 (NY).
MATERIAL.Bahamas:Nassau,9 Jan CULTIVATED 1932,Loomiss.n. (US).Belgium:Bruxelles,HortoLinden-
iano,Patins.n. (K). Bermuda: Autorbridges.n. (BH). Brazil: Rio de Janeiro,Bailey & Bailey529 (BH);Jan 1924, Bailey & Bailey 547 (BH); 1883, Glaziou 13292 (BR, FI, G, K, LE). Ceylon: PeradeniyaGarden,Johnston1642, 1664, 1674 (B). Cuba: Cienfuegos, Botanic GardenLimones Soledad,Jack 8742 (NY). Ecuador: Pichincha, Sto. Domingo, 20 Nov 1987 (fl, fr), Skov,Borchsenius,et al. 64839 (AAU). Eng-
colland:RoyalBotanicGardens, Kew,May1885,unknown Beccaris.n. lectors.n. (K), Indonesia:GartenBuitenzorg, (FI); Johnston1586, 1586a, 1587a (B), Java;Bogor Botanical Garden,Herb.Beccari 46, 74, 75, 92 (FI). Panama:, Bal-
boaHeights,Cook&Martin17 (US);30 m,20 May1986,de Nevers7745(NY);CanalZone,SummitGardens, Bailey432 (BH); 16 Dec 1974,Mori & Kallunki3727 (MO).Peru: BotanicalGardenof TingoMaria,640 m, 27 Aug 1983(fl), Kahn1684(USM).Singapore:Botanical Garden, Baileys.n. (BH); Furtados.n. (K); Johnston1626 (B); unknowncollec-
tor(K).Trinidad:Kuntze1062(NY);Maracas valley,10Apr 1920, Britton & Hazen 1634 (GH, K, NY). United States:
Fairchild TropicalGarden(FTG),Moore6015(BH);22 Jul 1963,Read905(BH,FTG);31 Oct1978,Fantz3467(FTG); RoyalPalmGardenMiami,May 1919,Dahlgrens.n. (F); Hawaii,Honolulu,Iltis H-56 (BH);Judds.n. (BH); US Botanical Garden,Nov 1937, unknowncollector s.n. (BH).
Venezuela:Barinitas,26 Jun 1956, Bernardi3327 (NY); Bolivar,plantedin CiudadBolivaron theOrinoco,70 m, 3 cultivated Mar1921(fl),Bailey&Bailey1649(BH);Caracas, in theJardinBotanico,Brauns.n.(BH,NY,U, VEN). Aiphanes aculeata is the most widely distributed species in the genus, and correspondinglyvariablein morphology.It is distinguishedby the thick, solitary stem, the abruptlywidening, bi- or tricuspidatepinnae, typically inserted in groups of 4-6, and 2.5-5.5 mm long staminate flowers with linear anthers. Flowers are somewhat greenish in bud, yellow to white at anthesis. Fruits are normallybright red, but occasionally individualswith yellow, orange,or even white fruits are encountered.Aiphanesaculeata consists of two geographically separatedsubunits, one Venezuelan-Colombian,the otherPeruvian-Bolivian. The northernsubunitis ratheruniformmorphologically, with the exception of three specimens: 1) the type of A. killipii Burretfrom near Bucaramangain Colombia, which has a spinulose pistil and fruit but otherwise correspondsin all respects to A. aculeata; 2) the type of A. orinocensis, collected on the lower Orinoco (without further locality), which has relatively few, large pistillate flowers aggregatedon the basal part of the rachillae, an arrangementotherwise known only from Peruvian and Bolivian specimens
49 [A. truncata(Martius)H. Wendland];3) a collection made from a planted tree in Ciudad Bolivar on the Orinoco in Venezuela(Bailey & Bailey 1649) with a densely twice-branchedinflorescence. The southernsubunitis more variable,and appears to include two main morphologicalforms. The first, includingthe type of A. ernestiiBurret,is knownfrom the lowlands in southeasternPeru, adjacentareas in Brazil, and Nor Yungasin Bolivia. It is characterized by more or less hirsute,tricuspidatepinnae,peduncular bract inserted near the base, rachillae without swellings below the flower groups, and relatively small flowers. Although in Peru fruits are red and around Caranaviin Bolivia fruits are white, plants from these two areas are otherwise identical. The second form, correspondingto A. truncata(Martius) H. Wendland,is knownfrom the departmentof Cuzco in Peruandthe easternAndeanslopes in Bolivia from the departmentof La Paz to that of Santa Cruz. It is characterizedby narrower,truncate-bicuspidate,glabrouspinnae,peduncularbractinserteddistally on the peduncle, rachillae with strong swellings below the flower groups,and largerflowers. The holotype of A. aculeata was lost alreadyin the beginningof the 18thcentury(Martius,1847), and no isotypes exist. A collection from coastal Venezuela (Steyermark106916) has been chosen as neotype. This collection, including an inflorescence in staminate anthesis,has been preferredto a sterilecollection closer to the type locality (Steyermark90137). Aiphanes elegans (Linden & H. Wendland) H. Wendland(basionymMartineziaelegans) was based on a collection by Funck and Schlim from "Neu Grenada"without furtherlocality, including only a juvenile leaf. No collection by Funck and Schlim annotatedA. elegans has been found in the consulted herbaria.However, a specimen at L, labeled "Martinezia ? elegans, Herm.Wendl.,Nova Grenada,Linden coll.," matches the original description to such detail that it must be the holotype itself. Funck and Schlim sold all their materialto Linden's establishment in Luxembourg,from where specimens were latersold to variousprivateand official herbaria.It is likely that specimens bought in this way could have been annotated"Lindencoll." A similarly annotated specimen at K is most likely an isotype. Aiphanesernestii was publishedas a new name for Dammer's (1915) homonym Martinezia ulei; Dammerhad publishedthe name M. ulei (= Aiphanes ulei) in 1907. The type was a collection made by Ule at SeringalAuristella(not localized with certainty)on Rio Alto Acre in the border area between Brazil,
50 Bolivia, and Peru. The holotype at B was destroyed and no isotypes exist. A collection from Tahuamanu in Madre de Dios, Peru (Vargas18694), ca. 50 km S of Rio Alto Acre, has been chosen as neotype. Martineziatruncata [= Aiphanes truncata(Brongniart ex Martius)H. Wendland]was based on a collection by d'Orbigny (no. 8) from "Yungas de la Palma" (= Yungas del Palmar?, an area in dept. Cochabamba,Bolivia, 17? 08' S, 65? 30' W). Unfortunately,the herbariumof d'Orbigny (in P) is disorganized and much of the materialhas been lost. The cited collection could not be found on requestand is presumably lost. Instead the original illustrationby d'Orbigny (Martius, 1847: tab. 2, fig. 1) has been chosen as lectotype. 3. Aiphanes chiribogensis Borchseniusand Balslev, Nordic. J. Bot. 9: 386, fig. 2. 1989. Type. ECUADOR. Pichincha:Old rd. from Quito to Sto. Domingo de los Colorados,km 59, past Chiriboga, Las Palmeras, 10 min. walk along trail S from rd., 1900 m, 13 Nov 1987 (fl), Skov, Borchsenius & Bang Klitgaard 64822 (holotype, AAU; isotypes, COL, K, NY, QCA, QCNE). Solitary.Stem0-3 m tall, 3-6 cm diam., sometimes partlydecumbent,fiercely armedwith blackspines, to 6 cm long. Leaves 5-9, erect and arching,lower ones curving downwards;sheath,petiole, andrachiswith a brown, scaly, caducous indument;sheath 20-40 cm long, violet inside, covered with brown or black spines, to 9 cm long; petiole 12-45 cm long, green, with scatteredspines, 4-8 cm long; rachis48-100 cm long, green, unarmedor black spinulose, often with scattered, black spines, 4-5 cm long; pinnae 12-17 per side, insertedin groupsof 1-3, groupsoccupying 7-14 cm along the rachis, separatedby 12-16 cm, pinnae almost in one plane or in different planes, cuneate, 2.5-4.5 times as long as wide, obliquely or rarely incised-praemorseat apex, coarsely dentate, with an up to 5 cm long, 5-10 mm wide finger-like projectionon the distal margin,darkgreen adaxially, pale green abaxially, glabrous or spinulose on both sides, sometimes with short black spines along the margins;basal pinnae 8-14 x 0.5-3 cm; middle pinnae 10-24 x 4-7 cm; apical pinnae4-6 ribbed,6-27 x 5-16 cm. Inflorescence interfoliar,branchedto 1 orderwith pendulousrachillae,or rarelyspicate;prophyll 20-30 cm long, 1-1.5 cm wide, somewhatviolet; peduncularbract 60-100 cm long, 2-3 cm wide, thin, pale rose to violet, with a sparsewhite indument, unarmed or with black spinules; peduncle 2-90 cm
FloraNeotropica long, 3-6 mm diam. at junction with rachis, light green to pale violet, with a thin, white or light brown indumentandnumerousspines of very unevenlength, the majority 2-5 mm long, the longest to 6.5 cm, sometimesonly with shortspines;rachis 10-90(-150) cm long, the terminalpartforming a 10-40 cm long spike; rachillae up to 30, inserted at long intervals; basal rachillaoften insertedalmost at base of peduncle, 38-90(-150) cm long, without flowers on the basal (14-)/2-2/3 of the length, in this part flattened, 2-3 mm wide, glabrous,unarmed,flower-bearingpart 17-32(-50) cm long, more or less thickened,2-6 mm diam., minutely spinulose, often strongly flexuose distally, the proximal 3-2/3 with triads, distally with staminatedyads or near apex singles; apical rachillae 12-40(-100) cm long, withoutflowers for 2-20(-60) cm, the flower-bearingpart staminateor with triads for up to /3 of the length;flower groups more or less hidden due to strong flexing of rachillae. Staminate flowers deep purple with yellow anthers, ca. 2 mm long; sepals free, whitish, imbricate,covering 3/4 of the petals,ca. 2 x 3 mm; petals connatefor /4 of their length, valvatedistally,ca. 2 x 2 mm, recurvedat anthesis; filaments ca. 0.5 mm long, anthers slightly longer than broad, 0.6-0.8 x 0.6-0.7 mm; pistillode minute, sunken into the swollen, ca. 0.5 mm thick receptacle.Pistillateflowers pinkish violet with rose pistil, ca. 4 mm long; sepals free, imbricate,ca. 2 x 3-4 mm; petals connatefor /2-2/ of their length, valvate distally, ca. 4 x 3 mm, corolla lobes rounded, recurvedat anthesis;staminodialcup ca. 3 mm high, minutelytoothed;pistil 3-4 mm high, glabrous.Fruit bright red, 10-12 mm diam. (conserved in alcohol); exocarpglabrous,smooth; mesocarpyellow, fibrousfleshy, mealy; endocarp ca. 8 x 9 mm, globose to slightly acute at base, prominentlypitted-grooved. Illustrations. Figs. 6D (inflorescence),7B (staminate flowers), 12A, 14L (pollen), 16C (leaf anatomy), 18C (distributionmap), 21K (middle pinnae), 22A (inflorescence);Borchsenius& Balslev, 1989: fig. 2. Distribution and habitat. Endemic to western Ecuador, where it grows in premontane to lower montanemoist to wet forest at 1500-2100 m altitude. It is fairly common in primaryor slightly disturbed forest in an area in the northwesternpart of the province of Pichincha. It has never been found in secondary forest or open areas. The area around Molleturoin the provinceof Azuay,where the species was collected by Steyermarkin 1943, is heavily deforested, and an attempt to refind A. chiribogensis there was unsuccessful.
SystematicTreatment
51
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FIG.
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Aiphanes
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FIG.22. A. Aiphaneschiribogensi.s, inflorescence(Skov et al. 64822).B. A. duquei,inflorescence(Bernal& Devia1540). Specimens examined. ECUADOR. AZUAY:Between Rio Blanco and Rio Norcay on rd. between Chacanceoand Molleturo, 1520 m, 4 Jun 1943 (fl), Steyermark52832 (F). PICHINCHA: Rd. Quito-Pto. Quito, km 61, 1710 m, 15 Nov 1989 (imm fr), Borchsenius & Luteyn91422 (AAU, QCA,
4. Aiphanes deltoidea Burret, Notizbl. Bot. Gart. Berlin-Dahlem11: 568. 1932. Type. PERU. Loreto: Confluence of Rio Santiago and Rio Marafion,160 m, 8 Dec 1924 (fl), Tessmann4709 (holotype, Bt; QCNE);old rd.Quito-Sto.Domingo,km59, LasPalmeras, lectotype, here designated, G; isotype, single 2000 m, 3 Dec 1989(fr),Borchsenius 91441(AAU,QCA); pinna, F). San (fl) 91442(AAU);1.5kmN of LasPalmeras, quebrada 91440(AAU, Luis,2040 m, 3 Dec 1989(fl), Borchsenius Solitary,or caespitose with 2 large stems and sevQCA, QCNE);below Chiriboga,2000 m, 13 Aug 1980 eral small ones. Stem 0.1-2 m tall, ca. 6 cm in diam., al. Holm-Nielsen et 24787 (imm fr), (AAU). armedwith grey (black when dried), flat spines, to 6 cm long, standingat right angles to the trunk.Leaves Aiphanes chiribogensis is unique in its solitary habitand pendulousrachillae,the proximalones with 10-12, erect and arching; sheath ca. 30 cm long, a long basal flowerless part. The only other species armed with spines similar to those on trunk;petiole that has developed this feature to a similar or even 90-105 cm long, green, with a thin, brown,caducous strongerdegree is the ColombianA. duquei.The vari- indument,and scattered,black spines, to 6 cm long; ability in inflorescence structureobserved within the rachis 115-190 cm long, with indument and spines type population of A. chiribogensis is remarkable. like the peduncle,but spines fewer;pinnae 11-14 per Spicate inflorescences are apparentlymade only by side, inserted in groups of 2-3, in different planes, small individuals, and the degree of branchingis, to groupsoccupying 3-5 cm along the rachis, separated some extent, under environmentalcontrol. An indi- by 20-35 cm, pinnae cuneate, 1.5-3 times as long as vidual observed with spicate inflorescences in 1989 wide, lobulateor obliquelypraemorseat apex, with an had a bifurcateinflorescence in March 1991. Another up to 5 cm long finger-like projectionon the distal individual that had inflorescences with six rachillae margin, glabrous adaxially, shortly spinulose and in 1989 had an inflorescence with 13 rachillae in roughabaxially;basal pinnaeca. 28 x 2.5 cm; middle 1991. However, some of the largest individuals pinnae 19-32 x 8-14 cm; apical pinnae 3-4 ribbed, observed at the type locality producedvery large, up 20-25 x 13-28 cm. Inflorescenceinterfoliar,curving, to 2 m long inflorescences with only three or four branchedto 1 order;prophyll20-30 cm long, ca. 2 cm rachillae each more than 1 m long, whereas other wide; peduncularbract ca. 90 cm long, 2 cm wide, large individuals produced more densely branched thin, soon withering, unarmed, with a light brown, inflorescences with up to 30 rachillae,of which half caducous indument;peduncle 55-170 cm long, 4-8 or so were entirely staminate.Thus the branchingpat- mm diam. at junction with rachis, almost unarmedor ternof inflorescences seems partlyundergenetic and covered with black spines, to 1 cm long; rachis30-45 cm long; rachillae 49-60, minutely spinulose; basal partlyunder environmentalcontrol.
52 rachillae 40-45 cm long, ca. 3 mm wide at base, taperingto <1 mm diam. at apex, proximalhalf with triadsor tetradsof 2 pistillateand 2 staminateflowers, distal half with dyads of staminate flowers; apical rachillae 6-8 cm long, staminate;triads and tetrads sunkeninto shallow,elongate cavities in the rachillae, each subtendedby a minute bract,forming a <1 mm high rim aroundthe cavity; dyads superficial.Staminate flowers orange, ca. I x 1.5 mm; sepals half as long as petals, not imbricate,ca. 1 x 0.5 mm; petals briefly connate at base, 1.5-2 x 1-1.5 mm; filaments ca. 0.5 mm long, basally fused in a ring aroundthe pistillode; anthersalmost square,0.3-0.4 x 0.3 mm; pistillode globose, ca. 0.4 mm diam. Pistillateflowers light green, ca. 2 x 3 mm (before staminateanthesis); sepals broadlyovate, membranous,imbricate,almost enclosing petals, ca. 2 x 3-4 mm; petals connate for half theirlength,valvatedistally,2-2.5 x 2-2.5 mm in bud, ca. 4 x 4 mm in young fruit, corolla lobes rounded,recurvedat anthesis;staminodialcup nearly truncate,ca. 1.5 mm high in bud,3 mm in young fruit; pistil glabrous.Fruitsnot seen.
FloraNeotropica
rachis (vs. 13-46 cm and 56-120 cm, respectively,in A. weberbaueri).Inflorescences of A. deltoidea are correspondinglylarger,with 50-60, slender rachillae (vs. 5-35, often thickened rachillae in A. weberbaueri). Furthermore,A. deltoidea has orange staminate flowers, (vs. purplein A. weberbaueri).The two collections from the type locality demonstratethatA. deltoidea is variablein habit.The type specimen was caespitose, with two 1.5-2 m tall stems and several smaller ones, whereas a specimen collected in May 1990 (Kahn & Borchsenius 2556) was solitary and almost acaulescent. There are no collections of mature fruits, but field notes to Moore et al. 8358 state that immaturefruits were green, 22 mm long and 19 mm wide; dry immaturefruits in the material measure ca. 10 x 7 mm. The original description stated that fruits were small, sweet, and edible, but extant materialdoes not include fruits. Fruits of A. weberbaueriare ca. 12 x 10 mm, and purple. Size differences similar to those discussed above areknownin otherspecies, and should perhapsnot be overemphasized.As seen at its type locality, A. deltoidea nevertheless gave the impression of being Illustrations. Figs. 19B (distributionmap), 21W clearly distinct from A. weberbaueri,with which it (middle pinnae). was co-occurring.For this reason it has been maintained as a separatespecies. Local names and uses. Shicashica (Peru). Fruits areedible, "smallbut sweet"(notes to type collection). 5. Aiphanes duquei Burret, Notizbl. Bot. Gart. Distribution and habitat. The distributionof A. Berlin-Dahlem13: 492. 1937. Type. COLOMBIA. deltoidea is insufficientlyknown, but the few collecValle:Basin of Rio Cali, CordilleraOccidental,2000 tions indicate that it is widely distributedin the westm, 1 Nov 1936 (fem fl), Duque 393 (holotype,B). ern part of the Amazon basin along the Andean foothills, from southernColombia to Peru, reaching Solitary.Stem4-5 m tall, ca. 5 cm diam.,gray,with 1650 m in centralPeru. rings of retrorse,black, to 15 cm long spines that fall off in exposed areas.Leaves 8-9, erect to spreading; Rio sheath 72 cm Specimens examined. COLOMBIA. AMAZONAS: long, proximal part with a yellow betweenLakeCarijonaandLakepescado,300 Cahuinari, tomentumand densely covered with spinules, surface 9 1988 & Galeano Mirana 1676 m, Sep (st), (AAU,COL). PERU.AMAZONAS: Prov.Bagua(limitwithLoreto,Alto soft to the touch, the distalmost30 cm bending away Amazonas),mouthof Rio Santiago,160 m, 21 May 1990 from stem, almost unarmed:petiole 4-25 cm long, (imm fr), Kahn & Borchsenius2556 (AAU, COL, NY, with a yellow to brown, caducous indument,brownProv.TingoMaria,aboveTeaGardens USM).HUANUCO: spinulose adaxially,with few, black spines, to 10 cm headquarterson rd. from Tingo Maria to Divisoria, 1500-1650m, 27 Apr 1960 (immfr), Mooreet al. 8358 long; rachis 91-110 cm long, with an indumentlike ORLORETO:Rio Ucayali, (probably thaton the petiole,and scattered,blackspines, to 3 cm (BH, USM). UCAYALI also cultivatedin the BotanicalGardenof MuseuGoldi, long, unarmeddistally, adaxial side sometimes with Para,Brazil),3 Jun1908(fl), Huber3411(FI). shortpurplespinules;pinnae 23-35 per side, inserted BRAZIL.AMAZONAS: Rio Javary,SanAntoniode Boa in groupsof (1-)3-6 separatedby 9-15 cm, in differ6 Dec 1874 1070 Traill Vista, (st), (BH, GH, K). ent planes,narrowlycuneate,4.4-7.8 times as long as Aiphanes deltoidea is incompletely known, and wide, obliquelypraemorseat apex,glabrousor shortly more collections are needed to give a picture of its spinuloseon both sides, usually with no spines on the variation.It is closely relatedto A. weberbaueri,from midrib abaxially; basal pinnae linear, 17.5-22 x which it can be distinguishedby its largerleaves with 0.5-0.6 cm; middle pinnae28-40 x 5.5-7 cm; apical a 90-105 cm long petiole and a 140-190 cm long pinnae6-9 ribbed, 19-20 x 10-18 cm. Inflorescence
SystematicTreatment drooping,branchedto 1 order,the basal 1-3 rachillae pendulous, the remainingspreading;prophyll45-58 cm long; peduncularbract 120-196 x ca. 6.5 cm, coriaceous,with a brown,scaly indument,unarmedor with purplespinules;peduncle 22-94 cm long, 0.5-1 cm diam. at junction with rachis,densely armedwith brown to black, often flexuose spines, 1-3 cm long; rachis 123 cm long, armed like peduncle, but spines fewer; rachillae 54, the basal ones inserted at large intervals along the rachis, only the basal 8 rachillae with pistillate flowers; basal rachilla 115-150 cm long, without flowers for 3/5 of the length, in this part as thick as the rachis and armedsimilarly,proximally appressedto the rachisin the prophyll,giving the impression that the peduncle is forked, flower-bearing part 43-61 cm long, with densely packed triads for
53 Distribution and habitat. Endemicto a very small area in the WesternCordilleraof Colombia, in the departmentsof CaucaandValle,in cloudy,lower montane wet and rain forest, at 2000-2600 m. It has not been found duringrecentsearchesat the type locality and may now be restricted to the national parks Farallonesde Cali and Munchique,and the area between them,an areaof <200 km2. Specimensexamined:COLOMBIA.CAUCA: Parque NacionalNaturalMunchique, betweenLa RomeliaandEl 81, 2600 m, 5-6 Apr 1989 (fl, fr), Bernal& Devia 1540 (COL,TULV).
Aiphanes duquei is very characteristicin its extremelylong, densely spinose, basal rachillae,with a long basal flowerless part, 13-20 mm long pistillate flowers (the largest in the genus), and long-rostrate t/2-2/ of the length, in this part ca. 1-2.5 cm diam. and fruits.The unusual developmentof the basal rachillae covered with black, flexuose spines between the misled Burret(1937) to interpretthe inflorescence as flower groups, distally slender,with staminatedyads; having a branched pedunclegiving rise to two partial subbasal rachilla inserted 38-69 cm above the basal inflorescences, a spicate, androgynous one and a one, abouthalf as long as this, withoutflowersfor half branched,staminateone. ActuallyA. duqueijust repthe length;middlerachillae35-42 cm long, staminate, resents the extreme of a tendency toward developwithout flowers for half their length; apical rachillae ment of basal sterile partof the rachillaeobserved in 7-14 cm long, staminate, with a 0.5-1.5 cm long severalother species, most notablyA. chiribogensis. flowerless part; triads sunken into deep pits in the The type of A. duqueiat B is a hanging sheet, with rachillae, each subtendedby an acuminatebractthat a leaf and a basal rachilla with pistillate flowers, in covers the pistillateflower for half its length or more, relativelygood condition. No isotypes are located at the two associatedstaminateflowers borneon up to 6 COL or FAUC, where most of Duque's collections mm long pedicels lifting them out of the pit; dyads are kept. slightly sunken,composed of a proximalsessile and a distal shortly pedicellate staminateflower. Staminate flowers purple,4-5.2 mm long; sepals connatefor 1'/ 6. Aiphanes eggersii Burret, Notizbl. Bot. Gart. of theirlength, gibbous at base, 2-2.5 mm long, lobes Berlin-Dahlem 11: 563. 1932. Type. ECUADOR. narrowlytriangular,slightly imbricateat base; petals Manabi: El Recreo, 14 Feb 1897, Eggers 15480 free, valvate, 3-4.5 x 1.5 mm; filaments 1-1.5 mm (holotype, B; isotypes, GH, O, S). long, antherslinear, 1.5-2.1 mm long, thecae free and Caespitose, with up to 10 stems, each 1-6 m tall, sagittate basally; pistillode trifid. Pistillate flowers 13-20 mm long, each subtendedby one or two 10-12 7-8 cm diam., light grey, armed with black or grey, mm long, 6 mm wide bracteoles,muchresemblingthe flat, to 10 cm long spines at the internodes.Leaves sepals; sepals imbricate, acuminate,minutely spinu- 7-10, erect and arching, older leaves recurving; lose at base, 10-14 x 6-10 mm;petalsconnatefor half sheath40-75 cm long, with a thick, white tomentum, theirlengthor less, valvatedistally, 12-18 x 4-6 mm, armedwith black or grey, flat spines, to 10 cm long; middle part sometimes minutely spinulose abaxially, petiole 0-10 cm long, similar to the sheath; rachis staminodial cup free of 115-205 cm long, with a white, caducoustomentum, lobes triangular-acuminate; mm teeth ca. 1 mm high, linear- armedwith many black spines, to 5 cm long; pinnae ca. 5 petals, high, acuminate;pistil 6-12 mm high, glabrous.Fruitsred, (30-)50-65 per side, inserted in groups of (2-)4smooth, globose, long-rostrate,9-12 mm diam., ros- 10(-14), in differentplanes, groupsoccupying 10-20 trum4-6 mm long; endocarpturbinateto subglobose, cm along the rachis, separatedby 7-12 cm, pinnae ca. 9 mm diam. narrowly trumpet-shaped,abruptly widening near apex, 3-7 times as long as wide, inrolled in vivo, Illustrations. Figs. 10D, 14D (pollen), 18C (distri- irregularlypraemorseat apex, glabrous or spinulose butionmap),21H (middlepinnae),22B (inflorescence). on both sides, adaxially often with a row of slender
54 spines on the midrib,abaxiallywith 1 to severalblack spines basally on the midrib;basal pinnae 13-34 x 0.5-3 cm; middle pinnae20-45 x 4-9 cm; apical pinnae 6-8 ribbed, 14-22 x 10-17 cm. Inflorescence erect or curving, pendulous in fruit, branchedto 1 order;prophyll,peduncularbract,and peduncle with a thick white indument;prophyll 19-45 cm long, 3-6 cm wide; peduncularbract inserted 5-23 cm above prophyll, 80-145 cm long, 6-10 cm wide, woody, almost unarmedto densely armed;peduncle 42-137 cm long, 6-15 mm diam. at junction with rachis, densely armedwith black spines, to 2 cm long; rachis 35-48 cm long, unarmedor basally armed like the peduncle; rachillae 35-75, unarmed,with a sparse, peltate indument;basal rachillae20-35 cm long, with triads for l/2 of their length, staminatedistally; apical rachillae5-10 cm long, staminate;each flower group subtended by a minute, inconspicuous bract. Staminateflowers yellow, superficial;sepals 1-2 mm long, never exceeding /4 of the petals; petals 3-3.5 mm long; filaments 1-1.5 mm long, anthers 1.2-1.8 mm long, 0.8-1.1 mm wide; pistillode ca. 0.5 mm high, trifid. Pistillate flowers yellow with brown sepals, sunkeninto shallow pits in the rachillae;sepals ovate, 2.5-3 mm long; petals connate for half their length, valvate distally, acute, ca. 4 mm long, opening to ca. 45? at anthesis;staminodialcup ca. 3 mm high, acuminately lobed to nearly truncate; pistil glabrous. Fruits bright red, globose, 18-20 mm diam.; endocarp 13-14 mm diam., shallowly pitted. Illustrations. Figs. 5A (leaf), 6A (inflorescence), 9A (fruits), 10A (pollen), 16D (leaf antomy), 17A (flower anatomy), 18A (distributionmap), 21E (middle pinnae), 23A (habit). Local name and uses. Corozo (Ecuador).Mesocarp and endospermedible. Distribution and habitat. Endemic to the dry parts of the coastal plain of Ecuador below 600 m altitude. Common in the province of Manabi. The naturalhabitat is semi-deciduous, dry Ceiba forest, but it is often found in pasturesor sometimes planted near houses. Specimens examined. ECUADOR.EL ORO:Near Puyango,600 m, 15 Aug 1978 (fem fl), Daly 101 (NY).
MANABf: Upper Rio Jama valley, rd. between San Isidro
and San Jacinto,nearestero Capa Perra,300 m, 4 Nov 1982 (fr), Pearsall 1070 (AAU); 10 km N of Canoaon the Pacific coast, 3 km from the coast, 50 m, 26 Mar 1987 (fl, fer fl,
et al. 62012 (AAU,NY,QCA);2 fr),Balslev,Borchsenius
km N of San Vicente town center on Pacific coast, ca. 1 km inland, 50 m, 25 Mar 1986 (imm fr), Balslev,Borchseniuset
Flora Neotropica al. 62011(AAU,NY);rd.Jipijapa-Guayaquil, km 46, 300 et al. 64735 m, 4 Oct 1987(fl, fer fl), Skov,Borchsenius, Ana,km3, 250m, 5 Oct1987(fem (AAU);rd.Olmedo-Sta. 64736(AAU). fl), Skov& Borchsenius Aiphanes eggersii is closely relatedto A. aculeata from which it can be distinguishedby its caespitose habitandits more numerous,narrowerpinnae.Its leaf anatomyis verycharacteristicin thatthe non-vascualar fibersare arrangedin a single layer of very thick bundles, a patternotherwise found only in some specimens of A. lindeniana. The holotype of A. eggersii in B consists of a single staminateflower. The isotypes include only leaf fragments.
7. Aiphanes erinacea (Karsten) H. Wendland in Kerchovede Denterghem,Les Palmiers230. 1878. MararaerinaceaKarsten,Linnaea28: 391. 1857. Type. COLOMBIA. 1000m, Nariiio:nearBarbacoas, Pipulquer, Karstens.n. [lectotype,(Imschanitzkaya, 1987),LE;isotype,WU]. Caespitose, with 1-5, or in open areas up to 15, stems, each 1.5-5 m tall, 2.5-5 cm diam., sometimes partlydecumbent,with many black spines, to 10 cm long. Leaves 3-8, erect and arching;sheath, petiole, and rachis with a brown, caducous indument;sheath 16-50 cm long, covered with brownor yellow spines, to 5 cm long; petiole 15-40(-55) cm long; rachis 80-165 cm long, normally covered with yellow spines, to 1 cm long; pinnae (8-)10-19 per side, insertedin groupsof 2-3(-4) separatedby 10-20 cm, in different planes, strongly plicate, narrowly to broadly cuneate, 1.3-4.5 times as long as wide, incised-praemorseat apex, symmetrical around the midrib,with a 2-9 cm long finger-likeprojectionon the distal margin, adaxially glabrous or nearly so, abaxially and on marginsminutely spinulose, sometimes both sides covered with long, yellow spinules, midrib abaxially with 0-3 yellow spines, ca. 3 cm long: basal pinnae 6-30 x 0.5-8 cm; middle pinnae 12-42 x 5-15 cm; apical pinnae 3-6 ribbed, 14-32 x 5-25 cm. Inflorescenceerect or arching,once- or occasionally twice-branched;most parts with a brown, caducousindument;prophyll26-73 cm long, 1-4 cm wide; peduncularbract up to 185 cm long, 2-5 cm wide, coriaceous,brownspinulose;peduncle 80-210 cm long, 5-10 mm diam. at apex, densely armedwith slender, yellow spines, to 1(-2.5) cm long; rachis 13-90 cm long, covered with brown to purple spinules; rachillae (4-)15-180, densely covered with
Systematic Treatment
_
-
/:
55
.. .
-44.
::
:11~i
.:...:.::...:.
... "',:"'-..' !..
A. Aiphanes eggersii, young,single-stemmed plant(Skovet al. 64735) B. A. grandis (Madsen 86934) flower F?IG.23.
_ -_~.
_
6
.?
_?
S
1_~~~l';?
::id ?r?i i
_
_
_y
,
?
grandis (Madse 86934),lowr (Skv :~i- t a. 4735) 1l(lsai?lBY. A.1~ ~ `'?~J'i~ i~. sigestme 1962) ~ p lant~~~~~~~ eggersEi, FIG. 23. A. Aiphunes individual & Borchsenius (Bernalyoung,~ single-stemmed
ing ad fruting ndiviual.C. A.hirsua subp. klbreyri, sngle-temmd (rheis16) iniiul(enl& sigestme
indvidua. D. . hisuta ubsp.interedia
ing and fruitingindividual.C. A. hirsutasubsp kalbreyeri, individual.D. A. hirsutasubsp"intermedia. single-stemmed brown to purple spinules, (0.1-)0.5-1.5 mm long, basal rachillasometimes withoutflowers for up to 18 basal rachillae30-55 cm long, occasionally branched cm and then insertedwell below the next, the followwith up to 20 secondaryrachillae,to 25 cm long, the ing without flowers for 1-5 cm, the fertile part with
FloraNeotropica
56 triads for 1/-2- of the length, in this part 2-3 mm diam., distally staminate, ca. <1 mm diam.; apical rachillae 5-20 cm long, staminate,the apical rarely with a few triadsat base; triads sunkeninto pits in the rachillae, covering the pistillate flowers for ca. >2/3 their length, each triad subtendedby a 1-2 mm high bract;dyads superficialor slightly sunken,subtended by a minute bract. Staminateflowers rose to pale violet in bud,normallypurewhite at anthesis,1-2 mm long; sepals cap-shaped, carinate, imbricate, 1-1.5 mm long; petals 1-2 mm long; filaments0.3-0.8 mm long, basally connate in a low ring, anthers0.3-0.5 x 0.5-0.7 mm; pistillode minute, sunken into the swollen receptacle. Pistillate flowers 4-5 mm long, white with a slightly greenish pistil, petals turning pink at the end of anthesis; sepals 2.5-3 mm long; petals 4-5 mm long, with minute brown spinules abaxially on the basal half, connate for
/2-2/3,
valvate
distally; staminodial cup 2-3 mm high, truncate, adnateto petals;pistil ca. 4 mm high, glabrous.Fruits darkred, soon turningbrownor black, ca. 7 x 8 mm, shortly rostrate;endocarpca. 5 x 7 mm, prominently pitted-grooved. Illustrations. Figs. 1B (habit), 3A (stem branching), 6B (inflorescence), 7D (pistillate flowers), 9B (fruits), 11E-F, 14M (pollen), 17B-F (flower anatomy), 19B (distributionmap), 210 (middle pinnae); 24 (drawing);Karsten,1866: tab. 175, figs. 1-9. Distribution and habitat. Along the western Andean slopes from the departmentof Narifo in southern Colombia to the Ecuadoreanprovince of Cotopaxi, with a single (old) record from the province of Azuay. An isolated, deviating collection from the Pacific coast near Buenaventurain the Colombiandepartmentof Valle (see discussion in the notes) indicates that the distributionmay extend further north throughthe departmentof Cauca, an area poorly exploredfor palms. In westernEcuadorA. erinacea is common in primaryand somewhatdisturbed premontaneforest at 700-1500 m, occasionally up to 2000 m. Individualsare often left over on cattle pasture. The single Ecuadoreanpopulation east of the Andes is found in primaryor slightly disturbedforest at 1900-2100 m.
beyond Junin, 700 m, 14 Nov 1946 (fl), Foster & Foster DEPARTMENT: 2164a (BH). UNKNOWN Miraflores, 1 Jan 1906 (st), unknowncollector #13 (BH); unknown locality (fl), Purdie 34 (K). ECUADOR. AZUAY:Puentesde Guarumal,600-900 m, 1875 (st), unknowncollector s.n. (P). CARCHI:El Pailon,
ca. 45 km below Maldonadoalong foot path to Tobar
Donoso, 800 m, 28 Nov 1979 (fl, imm fr), Madison & Besse 7125 (BH, SEL); Chical, km 12 below Maldonadoon the Rio San Juan, 1200 m, 26 May 1978 (fr), Madison et al. 4567 (BH, SEL); Maldonado, 1450-1650 m, 1 Jun 1978 Tenefuerte, (fer fl), Madisonet al. 4838 (SEL). COTOPAXI: km 52-54 on rd. Quevedo-Latacunga,800-900 m, 9 Apr 1984 (st), Dodson & Thurston14204 (MO, NY). ESMERALDAS: New rd. from LitatowardsSan Lorenzo,km 19, 900 m, 15 Oct 1987 (fl), Skov& Borchsenius64750 (AAU). LOS RIos: Km 11 on rd. PatriciaPilar-La Centinela on Montafias de Ila, across Rio Palenque from the Biological Station, 560 m, 20 Jan 1977 (fem fl), Dodson & Young6627
km30-34, 1950-2100m, 4 Rd.Baeza-Tena, (SEL).NAPO:
May 1986 (fl), Balslev et al. 62084 (AAU, NY); 21 Oct 1987 (imm fr), Skov & Borchsenius64757 (AAU); (fer fl),
64758(AAU);ibid.,17 kmS of Cossanga,2100m, 6 May 1987 (fl), Balslev et al. 62495 (AAU). PICHINCHA: Toachi, 480 m, Oct 1883 (fl), Sodiro 187 (P); old rd. Quito-Sto. Domingo de los Colorados,6-11 km fromjoint to the new rd., 1300-1500 m, 22 Mar 1986 (fl), Balslev et al. 62007
(AAU, NY, QCA, QCNE);7 Oct 1987 (ab fr), Skov & Borchsenius64738 (AAU); (fem fl), 64739 (AAU, QCA); 25 Nov 1987 (fr), Skov & Borchsenius64840 (AAU); ibid.,
23 kmfromChiriboga, Estaci6nLa Palma,ca. 2000 m, 20 Jun1982(immfr),Balslev2738(NY,QCA);rd.Quito-Pto. Quito,km 113,10kmN of therd.,ENDESAforestreserve, 700 m, 27 Dec 1983 (fl), Balslev & Balsecca 4666 (AAU, NY); 29 Dec 1983, Argiiello & Betancourt353 (QCA); 20 Feb 1984 (fl), Argiiello 418 (K, QCA); 10 Nov 1989 (fem
91421(AAU,COL,NY,QCA,QCNE);rd. fl), Borchsenius
Quito-Pto. Quito,km 92, finca Limon Real, 1250 m, 21 Sep 1987 (fl, fr), Skov,Borchsenius,et al. 64708 (AAU); 15 Nov 89 (fern fl), Borchsenius & Luteyn 91423 (AAU, COL, QCA, QCNE).
Aiphanes erinacea is characterizedby its caespitose habit;usually yellow spines on leaf sheaths and leaf rachis;plicate,incised praemorsepinnae,inserted in widely separatedgroups of 2-3 or rarely 4; and branchedinflorescenceswith densely spinulose, slender rachillae. It is closely related to A. simplex; the differences are discussed under that species. A geographicallyisolated collection from the Pacific coast near Buenaventurain Colombia (Moore 9470) deviates in having 50-69
cm long, narrowly cuneate
pinnae, 9-14 times as long as wide vs. 14-32 cm long, 1.5-4.5 times as long as wide for Ecuadorean and inflorescences with only 8-14 VALLE: examined. COLOMBIA. populations, Specimens Agua Dulce, an island in BuenaventuraBay, 12 Feb 1967 (fl), rachillae,to 40 cm long, and insertedat largeintervals Moore et al. 9470 (BH). NARINO:La Planada,finca Salazar, the 19-37 cm long rachis; this specimen also 7 km above Ricaurte,1750 m, (fl), Gentryet al. 35064 (COL, along has spiny pollen, whereas Ecuadoreanpopulations MO); Piedrancha,Reserva NaturalLa Planada, 1500-1800 m, 29 Apr 1988 (fr), Benavides9642 (MO);ibid., 1800 m, 22 have pollen with densely positioned, sparsely fusing Nov 1986, Bernal & Hammel 1328 (COL, MO, PSO); supratectalclavae. Futureinvestigations,particularly
Systematic Treatment
57
50 cm
-
,
C,A.E
!
fig.,
FIG. 24. Aiphanes erinacea. A. Habit. B. Leaf. C. Staminate flowers. D. Pistillate flowers. E. Fruit. A, B, Skov & Borchsenius64757; C, E, Skov et al. 64708; D, Skov & Borchsenius64739.
58 in the poorly explored departmentof Cauca, will show whether this collection represents a separate species or whether the above circumscriptionof A. erinacea has to be widened.
8. Aiphanes gelatinosa H. E. Moore, Gentes Herb.8: 227, 228, t. 93. 1951. Type. COLOMBIA.Narifo: Beyond Junin, 700 m, 14 Nov 1946, Foster & Foster 2164 (holotype, BH; isotypes, A, COLleaves only). Solitaryor caespitose. Stems4-9 m tall, 5.5-10 cm diam., basally with up to 100 cm long prop roots; internodesarmedwith black, retrorse,to 20 cm long spines. Leaves 4-11, erect and arching, lower ones borne horizontally; sheath and petiole 60-160 cm long, armed with black spines, to 12 cm long, these fewer distally and only to 2 cm long, petiole sometimes only a few cm long; rachis 110-400 cm long, with a grey or brown, scaly, caducous indument, armedwith numerousspinules andfew to manyblack spines, to 2 cm long, fewer and shorter distally; pinnae 16-30 per side, regularlyinserted, or in lax groups of 2-4 separatedby up to 14 cm, in one or several planes, weakly to strongly plicate, often very rigid, linear to narrowlycuneate, 5-12 times as long as wide, obliquely praemorse at apex, or incisedpraemorseand then symmetricalaroundthe midrib, with a 0-3 cm long finger-likeprojectionon the distal margin,glabrousor with scatteredscales and minute spinules abaxially; basal pinnae 20-30 x 2-2.5 cm; middle pinnae48-60 x 3.5-14 cm; apical pinnae 3-4 ribbed,33-45 x 11-12 cm. Inflorescencesinterfoliar, 1 or 3 per node, then sharinga common, 3-chambered prophyll, curving, becoming pendulous,branchedto 1 order with appressedrachillae, sometimes encased in jelly in fruit; prophyll 50-72 x 7-8.5 cm; peduncular bract ca. 60 cm long or more, to 10 cm wide, thick, unarmedor spiny at apex, with a thick, brown, caducous indument; peduncle 60-110 cm long at anthesis, 5-10 mm in diam. at apex, in fruitup to 240 cm long, densely armed(at least distally) with brownish black spines, to 2(-5) cm long; rachis ca. 30 cm long, armedlike peduncle,but spines shorterdistally; rachillae 15-30, white at anthesis, appressed to rachis, the distal ones somewhat spreading; basal rachillae to 20 cm long, armed with black, crimped spines to 1 cm long, the proximal half thickened,to 1.5 cm wide, with densely packedtriads,basally only with flowers abaxially, distal half slender, 2-3 mm diam., with densely packed staminatedyads; apical
Flora Neotropica rachillae 8-10 cm long, slender, staminate; triads sunken into deep pits, staminateflowers borne marginally on 2-3 mm long pedicels; dyads borne in shallow cavities, surroundedby fused, ca. 1 mm tall bracts, the proximalflower of each dyad sessile, the distal with a ca. 1 mm long pedicel. Staminateflowers white or cream, 1.5-2.5 mm long; sepals narrowly triangular,not overlapping,1-2 x 0.5-1 mm; petals basallyconnate,valvate,2-2.5 x 1.5-2 mm; filaments ca. 0.5 mm long, anthersslightly longer than wide, 0.4-0.7 x 0.5-0.7 mm; pistillode minute,sunkeninto the swollen, 0.7-0.8 mm thick receptacle. Pistillate flowers white, with greenovaryandpale pink stigmas, ca. 7 mm long; sepals broadlyovate, imbricate,2-6 mm long, much shorterthanpetals and then enclosed in the flower pit, or enclosing the petals entirely; petalsverythick,connatefor /3 of theirlength,valvate distally,ca. 6-7 x 5 mm;staminodialcup 3-4 mm tall, nearly truncate;pistil glabrous.Fruits red with black apex, brown or shiny black at maturity,subglobose, somewhatacuteat base, 12-15 x 13-17 mm, green on basal unexposed part, rostrate 1-2 mm; endocarp 11-14 x 13-16 mm, moreor less acuteat base, prominently pittedand irregularlygrooved. Illustrations. Figs. 12E-F, 140 (pollen), 19D (distributionmap), 21G-I (middle pinnae), 25 (drawing). Distribution and habitat. Found on the western Andean slopes in southern Colombia and northern Ecuador,in premontanerain forest, at 800-1650 m. 4 km Specimensexamined:COLOMBIA.NARINO: pastJuninon rd.to Altaquer,1200m, 6 Oct 1985(fl, imm fr),Bernal& Galeano901 (AAU,COL);(fr),904 (COL). VALLE: Old rd. Cali-Buenaventura, km 50, 1 km W of Queremal,12 Mar 1975 (fem fl, fr), Anderson31 (BH); 1200-1300m, 22 Mar1988 ibid.,6-7 kmbelowQueremal, (fl, fr),Bernal& Prado1448(COL,FTG,TULV). ECUADOR. CARCHI:Environs of Maldonado, 1450-1650 m, 30 May 1978 (fr), Madisonet al. 4804 (SEL);El Pailon,ca. 45 km belowMaldonado alongfoot pathto TobarDonoso,800m, 30 Nov 1979(fl, fr),Madison & Besse 7187 (BH, K, QCA,SEL);trailGualpiAlto-La Guana,km3, on highestpoint,1000m, 15 May 1985(fem fl, fr),Barfod& Skov60003(AAU,COL,MO,NY). Aiphanes gelatinosa is easily recognized by its large leaves with linear or narrowlycuneate pinnae and its large, often multiple inflorescences with appressed,basally thickenedrachillae.The name refers to the gelatinous substancein which infructescences are sometimesencased.Aiphanesgelatinosa is known in two morphologicalforms that may representdistinct taxa (Table III). Plants with caespitose stems,
SystematicTreatment
59
.
FIG. 25. Aiphanes gelatinosa. a. Pinnae x0.30; b. Inflorescencex0.20; c. Basal portionof rachilla xl.20; d, e. Pistillate flower, lateralview and expandedx2.38; f. Fruitx1.79; g. Endocarpx1.79. Reproducedfrom Moore, Gentes Herb.8, 1951.
Flora Neotropica
60 Table III Intraspecificvariationof Aiphanesgelatinosa Character
A. gelatinosa s. str.
"triplex"
Habit Numberof leaves Leafrachis Pinnashape ratio Pinnalength/width Pinnastructure Pinnaapexshape Inflorescences Pollen Pistillatesepals Fruitssize Fruitscolor Infructescences
Caespitose 4-6 110-150 Linear 10-12 Weaklyplicate Obliquelypraemorse Onepernode Withlongspines 6 mmlong 12-13 x 13-14 mm Redwithblackapex Encasedinjelly
Solitary 8-11 170-400 cuneate Narrowly 5-8 Stronglyplicate Incisedpraemorse Threepernode Withoutspines 2 mmlong Ca. 15 x 17 mm Brownto shiningblack Notencasedinjelly
Specimensexamined:
A. gelatinosa s. str. (1 locality): Foster2164 (type), Bernal 901; 904 "triplex"(2 localities): Anderson31, Bernal 1448, Madison 7187; 4804; Barfod 60003.
single inflorescences,and spiny pollen (A. gelatinosa s.str.) are known only from the type locality of the species, in the Colombiandepartmentof Narinfo,right between the two areas from which the solitary form with multiple inflorescences and pollen without spines, here referredto as "triplex,"is known(Colombian departmentof Valle and northernEcuador,respectively). Knowledge of the species from more localities seems necessaryin orderto judge the taxonomic significance of these differences, particularly considering the great morphological variation that occurs in relatedspecies, e.g., A. hirsuta.The production of single or multiple inflorescence could be a questionof age and perhapsenvironmentalconditions of individualplants, as suggested by observationsof Paralinospadixhollrungii, an Arecoid palm native to New Guineawhich has multipleinflorescencesapparently of the same kind (Fisher& Moore, 1974). In this species, young plants produce a single inflorescence per node, whereas older plants producetwo or three inflorescences sharing a common, chambered prophyll. The difference in pollen morphologybetween the two forms of A. gelatinosa shouldbe viewed in the light of the variability in exine ornamentationthat occurs in the relatedA. hirsuta, where spiny, clavate, and rugulatepollen occur.
1860-1950 m, 27 Aug 1943 (fl), Steyermark54185 (holotype, F).
Solitary.Stem 10-21 m tall, to 17 cm diam., armed with black spines, often unarmed basally. Leaves 7-10, spirally arranged,spreading; sheath >70 cm long, with a thick, white indument, densely armed with black, applanatespines, to 12 cm long; petiole <3 cm long; rachis200-250 cm long, 3-4 cm wide at base, with an indument like that on the sheath, fiercely armed with black and yellow spinules and numerous applanate spines, to 4 cm long, fewer towardapex;pinnae insertedin groups of 5-8, these occupying 11-17 cm along the rachis, separatedby 5-8 cm, pinnaemultiranked,linear-lanceolate,widest in the middle, 10-12 times as long as wide, obliquely praemorseat apex, ecaudate,adaxial side with a row of short, thin spinules on the midrib, otherwise glabrous, abaxial side minutely spinulose to almost glabrous, margins lined with white hairs and short spinules; basal pinnae 18-20 x 1-3 cm; middle pinnae 60-80 x 5.5-8 cm; apical pinnae 5-6 ribbed, 20-45 x 2-13 cm. Inflorescence erect at anthesis, pendulous in fruit; prophyll ca. 9 cm wide, woody, armed with black, 1-2 cm long spines; peduncular bractca. 130 cm long, ca. 8 cm wide, woody, persistent, densely armedwith black, 1-2 cm long spines; peduncle ca. 50 cm long, ca. 2 cm diam. at junction 9. Aiphanes grandis Borchsenius& Balslev, Nordic with rachisat anthesis,thickerin fruit,densely armed J. Bot. 9: 388, fig. 3. 1989. Type. ECUADOR. El with black spines, ca. 1 cm long; rachis ca. 70 cm Oro: Cordillerade Dumari,WSW of Sambotambo, long, white at anthesis, with a thin white or later
SystematicTreatment brown, caducous indument, unarmed; rachillae ca. 200, white at anthesis, with an indumentas that on the rachis, unarmed;basal rachillae 30-50 cm long, without flowers on the basal 2-4 cm, with triadsfor 1/3 of the length, in this part4-5 mm diam., distally ca. 1 mm diam., with staminatedyads; apical rachillae 10-20 cm long, with triadsfor 1-4 cm, otherwise staminate;flower groups sunkeninto shallow cavities in the rachillae, subtended by a 1-2 mm tall bract. Staminateflowers 5-6 mm long, those of triadswith a 2-4 mm long pedicel, those of dyads shortly pedicelled; sepals imbricate, ca. 3 x 1 mm; petals free, valvate, 5-6 x 3 mm; filaments ca. 2 mm long, anthers linear, sagittate at base, 3-4 x 1 mm; pistillode distinct, trifid, ca. 0.5 mm high. Pistillate flowers 6-9 mm long; sepals broadlyimbricate,ca. 5 x 8 mm; petals connatefor /2 of the length,7-9 x 4-5 mm, lobes acute, slightly imbricate; staminodes acuminate,arrangedin 2 rings of 3 each, episepalous ones 6-7 mm long, epipetalous ones 4-5 mm long, connate with episepalous ones for 2-4 mm; pistil 6-7 mm high, densely covered with pale, at anthesis black, ca. 1 mm long spinules. Fruit dull green, 2023 mm diam. (dry), covered with ca. 1 mm long, black spinules, that easily fall off; mesocarpthin, <1 mm thick in dry fruits; endocarp globose, almost smooth. Illustrations. Figs. 1A (habit), 12D, 141 (pollen), 19E (distribution map), 21F (middle pinnae), 23B (habit);Borchsenius & Balslev, 1989: fig. 3. Distribution and habitat. Endemicto the western Andean slopes in Ecuador.It grows in premontane moist forest at 1000-2000 m, apparentlymost abundant around 1500 m. It is at least sometimes left in cleared areas, where it seems unable to regenerate. Though geographically very restricted, it is often abundanton both its localities, but very local. Specimens examined. ECUADOR. LOJA:NearOrianga, 1100 m, 6 Mar 1990 (fr), Madsen et al. 86927 (AAU). EL
ORO:NearSambotambo, 1450m, 6 Mar1990(fr),Madsen et al. 86934(AAU). Aiphanes grandis is easily distinguishedby its up to 21 m tall, solitary stem; linear, grouped, multirankedpinnae; 6-7 mm long staminateflowers with linear anthers, and spinulose pistil and fruits. It resembles A. linearis in its linear, grouped pinnae, fierce armature,and spinulose pistil, but the latter differs in having basally thickened rachillae and fruits covered with golden spines; furthermoreA. linearis is caespitose and has distichous leaves.
61 Aiphanes grandis appears to be the most primitive species in the genus with respect to floral morphology; pistillate flowers have imbricate petals and incompletelyconnate staminodes. 10. Aiphanes hirsuta Burret, Notizbl. Bot. Gart. Berlin-Dahlem11: 573. 1932. Type. COLOMBIA. Antioquia:Amalfi, La Vivora("LaViborra"),15001700 m, 11 May 1880, Kalbreyer1657 (holotype, Bt). Neotype (Bemal et al., 1989). COLOMBIA. Antioquia:Mun. de Amalfi, 10-11 km E of village on rd. to Medellin, 1650 m, 25 Sep 1987, Bernal & Tobon1396 (holoneotype,COL;isoneotypes,AAU, BH, FTG, HUA, K, MO, NY). Caespitose,with 1-8(-20) stems, each (2-)3-10 m tall, 2.5-10 cm diam., armed with black spines, to 12(-25) cm long, distal half of stem rarely covered with persistentold leaf sheaths.Leaves4-8, erect and arching;sheath 20-90 cm long, covered with brown spines, to 12 cm long; petiole 9-100 cm long, green, unarmedto densely armed with spinules and spines similar to those on sheath; rachis 86-194 cm long, with a grey to brown, scaly, caducous indument, unarmed to densely covered with black spinules, sometimes with manybrownto black spines, to 4 cm long; pinnae 9-40 per side, inserted in dense to lax groupsof (1-)2-9 separatedby 5-21 cm, in one plane or more often in differentplanes, more or less rigid, weakly to stronglyplicate, linear to broadlycuneate, 1-14 times as long as wide, truncateto lobulatepraemorse, or occasionally slightly incised praemorseat apex, with a 1-6 cm long finger-likeprojectionon the distal margin, adaxial side glabrous, abaxial side rough,or both sides sparselyto densely covered with up to 5 mm long, yellow spinules; basal pinnae 17.5-45 x 1-13 cm; middlepinnae 11-65 x 5-22 cm; apical pinnae 22-45 x 7-32 cm. Inflorescenceerect or more often curving,branchedto 1 order;prophyll 25-52 cm long, 1.5-4.5 cm wide; peduncularbract 70-133 cm long, 2.5-12 cm wide, with a brown, scaly, caducous indument,often covered with brown to black spinules, sometimes armed with brown to violet spines, to 2 cm long; peduncle(21-)43-170 cm long, 2-9 mm diam. atjunction with rachis, minutely spinulose or sparsely to very densely armed with numerousshort spinules and brown spines, to 3(-5) cm long; rachis 17-100 cm long, unarmedor armed like peduncle, spines fewer and shorter distally; rachillae 9-100, white to light brown at anthesis, densely covered with light brown, purple, or black spinules, minute or to 2.5 mm long; basal rachillae
Flora Neotropica
62 23-90 cm long, sometimes with an up to 18 cm long flowerless part, with triads for 1/2-2/ of the remaining
length, in this part 2-5 mm diam. at anthesis, often thicker in fruit, distally ca. 1-2 mm in diam., with staminate dyads; apical rachillae 11-40 cm long, staminate or in inflorescences with <30 rachillae often with a few triadsat base; fruitingrachillaeoccasionally very thick and fastigiate; triads inserted in shallow depressions in the rachillae, subtendedby a short rim-like bract covering the pistillate flower for ca. '/ of its length; dyads slightly sunken, subtended by an up to 1 mm long narrow bract. Staminate flowers 1.5-3.4 mm long, purpleto white, sometimes yellow in the center, with white to yellow anthers; sepals narrowlytriangular,imbricate,keeled, 1-3 mm long; petals almost free or basally connate, valvate, ovate to oblong, 2-3 mm long; filaments 0.7-1 mm long; anthersnearly square to linear, 0.5-1.2 x 0.40.7 mm; pistillode 0.5-1 mm high, trifid. Pistillate flowers brownto violet, 3-7 mm long; sepals rounded to broadly ovate, almost enclosing the petals before anthesis, 4-6 mm long, often with scatteredminute spinules adaxially; petals connate for /3-V/2of their length, valvate distally, 3-8 mm long, basally with minute, pale, or yellowish spinules abaxially, lobes triangular,rounded-acuteto acuminate,spreadingto recurved at anthesis; staminodialcup 2-5 mm high, acutely lobed to nearly truncate,basal half adnateto corolla tube, distal part appressedto the pistil; pistil 3-7 mm high, glabrous, or more often spinulose especially at base. Fruits darkred to purple,or occasionally white, globose, 7-20 mm in diam., sometimes shortly rostrate,glabrousor with many minute, black spinules; endocarp 6-18 mm diam., small endocarps finely pitted, especially distally, larger ones deeply pitted,sometimes longitudinallygrooved and 3-lobed in x.s., more rarely without pits; seed globose to very irregular.
tributional range, with a general tendency to increase in size of all structures toward the south. It is here divided into four subspecies. Key to the subspecies
of Aiphanes
hirsuta
1. Pinnaelinear,subregularlyinsertedor in lax groupsof 3-9, nearlyin one plane; inflorescencerachisand rachillaedensely armedwith black, crimpedspines... .. subsp. kalbreyeri 1. Pinnaenarrowlyto broadlycuneate, insertedin groupsof 2-5, in different planes;inflorescencerachis and rachillae unarmedor armed. 2. Petiole 9-35 cm long; leaf rachis66135 cm long; middle pinnae 11-45 cm long; rachillae<30; fruit red to purple, spinulose, 7-11 mm diam. ........ subsp. hirsuta 2. Petiole 43-100 cm long; leaf rachis 210260 cm long; middle pinnae 37-100 cm long; rachillae60-108; fruit red, glabrous or spinulose, 11-22 mm diam. 3. Pinnaenarrowlycuneate;fruit spinulose, 11-13 mm diam.;endocarp with 3 longitudinalfurrowsand with few pits.............. subsp. intermedia 3. Pinnaecuneate;fruit glabrous, 1622 mm diam.;endocarpglobose to somewhatfurrowed,with many subsp.fosteriorum deep pits ...............
lOa. Aiphanes hirsuta subsp. hirsuta Aiphanes monostachysBurretemend. Bernal, Brittonia38: 68. 1986;Burret,Notizbl.Bot. Gart.Berlin-Dahlem11: 574. 1932 (leaves only). Type. COLOMBIA.Antioquia:Murri, 1000 m, 21 Jul 1880, Kalbreyer1832 (Bt). Neotype (Bernal et al., 1989). COLOMBIA.Antioquia:Mun. de Frontino, Rio Cuevas,950 m, 18 Mar 1982 (fl, fr), Bernal & Murrm, Galeano251 (holoneotype,COL;isoneotype,HUA). Aiphanes pachyclada Burret, Notizbl. Bot. Gart. BerlinDahlem 11: 574. 1932. Type. COLOMBIA.Antioquia:Rio Rosario, 1600-1800 m, 27 May 1880, Kalbreyer 1701 (Bt). Neotype (Bernal et al., 1989). COLOMBIA. Antioquia: Mun. de San Roque, rd. San Roque-Santo Domingo, km 8, quebradaSanta Barbara,1600 m, 22 Sep 1987 (fl), Bernal & Tobdn1392 (holoneotype, COL; isoneotypes,AAU). AiphanesfuscopubensBailey, Gentes Herb.6: 209, 210, fig. 107. 1943. Type. PANAMA. Panama: above Campana, 600-800 m, 1 Jul 1939 (fl), Allen 1870 (holotype, MO; isotypes, BH, BM).
Distribution and habitat. From Panama and along the western slopes of CordilleraOccidentalin Colombiato north-westernEcuador,reachinginto the northernand eastern slopes of CordilleraCentralin the departmentof Antioquiain northwesternColombia. An understoryelement in premontaneor monStems 2-8(-20), each 4-10 m tall. Leaves 4-8; tane rain forest, between 600 and 2200 m, but it has sheath 20-45'cm long; petiole 9-35 cm long, densely also been recordedfrom lowland areas in northwest- armed with spinules and spines; rachis 86-135 cm ern Colombia at 100 m. long, unarmed or spiny; pinnae 9-26 per side, in groups of 2-5, spaced by 15-21 cm, in different
Aiphanes hirsuta is one of the most complex and planes, narrowly to broadly cuneate, 1-6 times as variable species in the genus. It includes a morpho- long as wide, nearly glabrous to densely covered with logically continuous series of forms throughits dis- yellow spinules, truncate to lobulate-praemorse at
SystematicTreatment apex, rarely slightly incised-praemorse;middle pinnae 11-45 x 5-16 cm. Peduncle43-120 cm long, 2-9 mm diam. at apex, minutely spinulose or sparselyto densely covered with brownspines, to 3(-5) cm long; rachis 17-46 cm long, unarmedor spiny; rachillae 9-28, densely covered with brown to black spinules, 0.5-1.5 mm long; basal rachillae 23-40 cm long, basally withoutflowers for 0.5-2.5 cm, often strongly thickened in the androgynous part, with closely inserted triads. Staminateflowers 1.5-3.4 mm long; anthers rectangular to shortly linear, 0.5-0.7 x 0.4-0.5 mm. Pistillateflowers 4.5-7 mm long, often acuminate; pistil ca. 3 mm high, glabrous or with some minute spinules at base. Fruit red to purple, globose, ca. 7-11 mm diam., glabrous to minutely spinulose; endocarpglobose, 6-9.5 mm diam., 0.5-1 mm thick, with numerousfine pits apically. Illustrations. Figs. 13A (pollen), 18B (distribution map), 21AA (middle pinnae); Motiska et al., 1984: fig. 8 (as A. pachyclada). Distribution and habitat. Known from premontane forest in Panama at 600-1400 m, the eastern slopes of northernCordilleraCentralin Colombia at 1600-1900 m, and westernslopes of CordilleraOccidental at 100-1500 m. Specimens examined. PANAMA. BOCA DEL TORO:
Fortuna Damarea,continental divide,ridgetrailto unnamed peakE of oleoductord., 1200 m, 1 Aug 1984 (immfr), Churchill 5874(MO).COCLE:N of ElValle,betweenCerro andCerroGaital,1000m, 18 Jul 1982(fer fl), Caracoral Knappet al. 6060(CAS);footof CerroPilon,aboveElValle deAnton,700 m, 27 Mar1969(st),Porteret al. 4388(MO); continentaldivideaboveEl Cope, 700 m, 27 Nov 1985
(infl), de Nevers et al. 6405 (NY); El Valle de Ant6n, Cerro Gaital, 26 Nov 1985 (infl), de Nevers et al. 6366 (NY). DARIEN:Serraniade Pirre, 900 m, 18 Jan 1985 (fer fl), Henderson & Contraires099 (NY); Cerro Pirre, ridge top nearRancho Plastico, 1200 m, 10 Jul 1977 (imm fr), Folsom 4325 (MO); ibid., just S of Pirre, 10 Jul 1977 (fr), Folsom 4513 (MO); ParqueNacional del Darien,CerroMali, ca. 22 km E of Pucuro, 1350 m, 23 Oct 1987 (imm fr), de Nevers et al. 8416 (CAS); Alto de Nique, southernmostpeak of CerroPirremassif, exactly on the Panama/Colombiaborder, 1400 m, 19 Apr 1980 (st), Gentry et al. 28658 (BH). PANAMA: Panama,(fem fl), Nee 11606 (MO); Comarcade San Blas, CerroBrewster,850 m, 21 Apr 1985 (st), de Nevers et al. 5395 (NY); CerroCampanaalong trail to summit, 22 Jun 1972 (st), Croat 17194 (MO); CerroJefe, 5 km S of the summit,4 Feb 1973 (imm fr), Busey & Croat268 (MO); ibid., near summit, 10 Mar 1973 (infl), Croat22684 (MO); ibid., 3 km beyond tower, 13 Dec 1987 (st), Henderson &
63 & Galeano349 (COL,HUA);Mun. de Mutata,rd. to 100-150m, 8 Dec 1982 (fl), Bernal& Pavarandogrande, Galeano440 (COL);Mun.de SantaRita, 7 km NE of Guatape,1900m, 16Aug 1980(fl), Galeano& Bernal238 (COL,HUA); 17 Sep 1987 (fl), Bernal & Tobdn1378 (AAU,BH, COL,HUA,NY); Mun.de Urrao,Rio Polo, 1500m, 16 Jun 1982 (fl), Bernal& Galeano348 (COL, HUA). CHOC6:Mun. de El Carmende Atrato, rd. km 150,veredaEl Doce,RioEl Aguil6n, Medellin-Quibd6 640 m,4 Jan1980(fl, immfr),Bernal& Galeano56 (COL, RioTorito, HUA),57 (COL);Mun.de SanJosedel Palmar, 900 m, 12Mar1980(fl), Foreroet al. 7224(COL). This subspeciesis the most variableof the four. In Panamaand the northernparts of CordilleraCentral there is a strong tendency toward thickening of the rachillae,densely insertedtriads,andpistillateflowers with conspicuously acuminatesepals and petals. In extreme cases inflorescencesmay approachthose of A. gelatinosa in structure.On the western slopes of CordilleraOccidental these tendencies are less pronounced,and plantsfrom the southernChoc6 (Forero 7224) form a transitionto subsp.fosteriorumdiffering only in size. Pollen of A. hirsutasubsp.hirsutaappear to be ratheruniform,with tectate,perforateto rugulate exine with supratectalwartsor spines. The confusion regardingtypificationof A. monostachys, A. leiospatha, and A. macroloba has been elucidatedby Bernal (1986). Aiphanes monostachys Burretwas based on a mixed collection includingthe leaves of A. monostachysand the misplacedinflorescence of what Burretdescribedas A. macroloba;thus the misleading name. The real inflorescence of A. monostachyswas probablyassociatedwith the leaves of a Geonomoidpalm (Kalbreyer1607) anddescribed by Burret as a separate species, A. leiospatha (see "Doubtfullnames and Excludedtaxa"). 10b. Aiphanes hirsuta subsp. kalbreyeri (Burret) Borchsenius& Bernal stat. nov. AiphaneskalbreyeriBuffet, Notizbl. Bot. Gart.BerlinDahlem11: 572. 1932. Type.COLOMBIA. Antioquia: 1650-1750m, 10 May Amalfi,LaVivora("LaViborra"), 1653 (holotype,Bt). Neotype(Bernalet 1880,Kalbreyer Mun.de Amalfi,queal., 1989).COLOMBIA. Antioquia: bradaLaVivora,6 kmE of village,1650m, 24 Sep 1987 COL;iso(fl, fer fl), Bernal& Tobon1393(holoneotype, neotypes,AAU,BH,FTG,HUA,K, NY).
Stems up to 12, each 4-10 m tall, 5-8 cm diam., densely armed with black, to 25 cm long spines. Herrera722 (NY). Leaves 4-6; sheath 70-90 cm long, densely armed; COLOMBIA. ANTIOQUIA:Mun. de Cocorm, hwy. petiole 14-32 cm long, densely armed with black Medellin-Bogota, km 63, quebradaEl Bihao, 1900 m, 26 to 7 cm long; rachis 125-194 cm long, yellow Apr 1980 (fl), Bernal et al. 171 (COL); Mun. de Frontino, spines, Murri,Rio Venados,750-850 m, 19 Jun 1982 (fl, fr), Bernal to brown spinulose, abaxially with few black spines;
64
FloraNeotropica
pinnae 30-40 per side, in lax groupsof 3-9, nearlyin Central(Bernal & Tobon1393) has pollen with ruguone plane, linear, 10-14 times as long as wide, late exine (Fig. 13C);pollen of Bernal et al. 958, coloblique to lobulatepraemorseat apex, glabrousadax- lected in CordilleraOccidental,have perforateexine ially, with short spinules abaxially; middle pinnae with numerouslong, supratectalspines (Fig. 13D). 48-65 x 3.5-5 cm. Peduncle 21-107 cm long, ca. 10 mm in diam. at junction with rachis, densely armed with brown, to 2 cm long spines; rachis 20-44 cm 10c. Aiphanes hirsuta subsp. intermedia Borchsenius & Bemal, subsp. nov. Type. COLOMBIA. long, densely covered with crimped, to 1.5 cm long San Jose del Palmar,500 m E of town, 1300 Choc6: toward shorter rachillae apex; spines, becoming 15 Mar 1991 (fl, fer fl, fr), Bernal & Borchm, cm with an basal rachillae 40-53 to 22-38; long, up senius 1962 (holotype, COL; isotypes, AAU, 10 cm long basal flowerless part,2-3 mm diam. in the CHOCO,NY). androgynouspart,covered between the flower groups with 0.5-1 cm long, crimped, black spines, distally A subsp. hirsuta petiolo 43-80 cm longo, pinnis very slender. Staminateflowers violet to white, with 44-81 cm longis, inflorescentia ramis 60-108, et yellow anthers, 2.5-3 mm long; anthers linear, fructu 11-15 mm diametro endocarpo longitudinal0.9-1.2 x 0.5-0.7 mm. Pistillate flowers violet, 5-6 iter exaratodiffert. mm long; sepals rounded; petals acuminate; pistil sparsely covered with minute, applanate spinules. Stems 1-4, each 2-8 m tall, 6-10 cm diam.,densely Fruitglobose, 11-13 mm diam.;epicarpwith minute, armed. Leaves 5-7; sheath 40 cm long, densely black spinules; endocarp 10-12 mm diam., with armed; petiole 43-80 cm long; rachis 220-242 cm numerousshallow to deep pits, especially at apex. long, unarmed;pinnae 31-32 per side, in ratherlax of 3-5 separated by 13-22 cm, narrowly groups Illustrations. Figs. 13C-D (pollen), 18B (distribucuneateto cuneate,3.5-8 times as long as wide, finely tion map), 21Z (middle pinnae), 23C (habit). plicate, glabrousadaxially,minutely spinulose abaxiLocal names. Cirqui (Antioquia). ally, truncateto lobulate-praemorseat apex; middle pinnae 44-81 x 14-15 cm. Peduncle 15-85 cm long Distribution and habitat. Known from the north- at flowering, to 120 cm in fruit, 8-25 mm diam. at eastern slopes of the Colombian CordilleraCentral apex, covered with black spines, to 3 cm long; rachis (type locality), and the western slopes of Cordillera 50-75 cm long, with numerousspines, to 1 cm long Occidental,between 1300 and 2000 m, often left over proximally, fewer and shorter distally; rachillae on pasture. 60-108, densely covered with brownspinules,ca. 0.3 mm long, and with some short spines proximally; Mun. Specimens examined. COLOMBIA. ANTIOQUIA: basal rachillae 40-84 cm long, without flowers for de Frontino,Murm,rd. Nutibara-La Blanquita,1900-1950 m, 3 Jan 1982(fl, fer fl, immfr), Galeano& Bernal461 2-8 cm, ca. 4 mm diam. in the androgynouspart,ca. (COL,HUA),462 (COL);Mun.de Urrao,quebradaLa 2 mm diam. distally. Staminateflowers ca. 1.5 mm to Rio Calles,1300-1400m, 14 Jun Agudelo,a tributary violet to white, anthers purple, pollen 1982, Bernal & Galeano 330 (COL, HUA). RISARALDA: long (dry), PuebloRico, Rio Taiba,1350m, 13 Mar1986(fl, fem fl, orange.Pistillateflowers 4-5 mm long (dry), purple; immfr),Bernalet al. 958 (COL,FTG). sepals rounded;petals acute; pistil with whitish yellow spinules proximally.Fruit dark red, 11-13 mm This subspecies includes plants with linear pinnae diam.; epicarp minutely spinulose; endocarp with 3 held more or less in one plane, densely armedinflodeep broadlongitudinalfurrows,almost withoutpits. rescences with relativelyfew, long, slender rachillae, Illustrations. Figs. 13E-F, 14N (pollen), 18B (disand small spinulose fruitswith a globose endocarp.In the northernpart of CordilleraCentral it co-occurs tributionmap), 21Y (middle pinnae), 23D (habit). with subsp.hirsuta,andthe two give the impressionof Distribution and habitat. Westernslopes of Corbeing distinct in this area. In CordilleraOccidentalit dillera Occidental in the departmentsof Choco and occurs at higher altitudesthan subsp. hirsuta.To the Valle in central Colombia, 1300-2200 m, often left and leaves inflorescences show a transouth, gradual sition to subsp. intermedia,found in Choc6 andValle. on pasture. The subspecies is well characterizedmorphologically, Specimens examined. COLOMBIA.CHOC6:Rd. but its pollen display a surprisingvariationin exine Cartago-San JosddelPalmar, ca. 10kmE of SanJose,1800 ornamentation.The neotype collected in Cordillera m, 2 Apr1976(fl, fr),Dransfield et al. 4854(BH,K);ibid.,
65
SystematicTreatment 1600m, 11Nov 1985(fl), Lozanoet al. 4903(AAU,COL); (fl, fr), Lozanoet al. 4905 (AAU, COL). VALLE:rd. Cartago-SanJose del Palmar,2 km from dividingline betweenValleandChoc6,2000m, 16 Mar1991(fi, femfl, imm fr), Bernal & Borchsenius1963 (AAU, COL,NY, TULV);basinof RioDagua,RioSanJuan,km52-53 below Dagua,1300-1500m, 19 Mar 1947 (fl, fr), Cuatrecasas 1800m, 13 Apr1989(fl, 23885 (F);rd.Trujillo-Cristales, fem fl, imm fr), Bernal& Devia 1555 (COL,TULV);rd. Danubio,1300-1420m, 24 Mar1988(fl, fr), Queremal-El Bernal& Prado1455(COL,TULV). This subspecies includes plants with long, narrowly cuneate pinnae, spiny inflorescences with 60-90 rachillae,and small to medium-sizedfruitwith longitudinallygrooved endocarphaving few pits. In the northit forms a transitionto subsp. kalbreyeri;in the south it forms a transitionto subsp.fosteriorum, which has more broadly cuneate pinnae, less spiny inflorescences, larger, glabrous fruits, and a more globose endocarp with numerous deep pits. In the centralChoc6, subsp. intermediaoccurs side by side with subsp. hirsuta, but at higher altitudes.Pollen of A. hirsuta subsp. intermedia has perforate, tectate exine and is spineless or provided with numerous supratectalwarts or spines (Figs. 13E-F, 14N). lOd. Aiphanes hirsuta subsp. fosteriorum (H. E. Moore) Borchsenius& Bernal, stat. nov.
Illustrations. Figs. 13B (pollen), 18B (distribution map),21X (middlepinnae);Moore, 1951:226, fig. 92. Distribution and habitat. WesternAndean slopes from the departmentof Valle in Colombiato northern Ecuador,at 900-1300 m. Specimens examined. COLOMBIA.Narifio: Rd. km 10, 1200m, 4 Oct 1985(fl, fr),Bernal Junin-Altaquer, & Galeano895 (AAU,COL,K). VALLE: Rio Chanchos, fromjunctionwithRioCalima,Campment 300 m upstream Calima III, 450-600 m, 16-19 Feb 1989 (fl), Bernal et al. 1521 (COL).
Rd.Lita-SanLorenzo,km ECUADOR.ESMERALDAS: 8-18, 800-900 m, 14 May 1986(fl, fr), Balslev,Borchsenius, et al. 62102 (AAU,NY, QCA);13 Sep 1987 (juv), Skov & Borchsenius64742 (AAU); 12 Nov 1987 (fl, fem fl, imm fr), Skov,Borchsenius,et al. 64819 (AAU, QCA); rd.
Lita-BuenosAires, km 12, 1400 m, Dodson & Gentry 17582 (QCNE).
This subspecies includes the largest plants in the species, with cuneate,lobulatepraemorsepinnae and weakly armedinflorescenceswith up to 100 rachillae. It is set apartby several characters:glabrous pistil; large fruits, ca. 16-22 mm diam, with deeply pitted endocarp;and pollen with fusing, supratectalclavae giving a rugulatesurface(Fig. 13B).
11. Aiphanes leiostachys Burret,Notizbl. Bot. Gart. Berlin-Dahlem11: 570. 1932. Type. COLOMBIA. AiphanesfosteriorumH. E. Moore,GentesHerb.8: 225, Narifo:Altaquer, Antioquia:Buenavista,Wald, 850 m, 18 Feb 1880, 226, fig. 92. 1951.Type.COLOMBIA. 1150 m, 10 Nov 1946,Foster& Foster2117 (holotype, Kalbreyer1429 (Bt). Neotype (Bernalet al., 1989). COLOMBIA.Antioquia:Mun. de San Carlos,3 km BH;isotypes,A, COL). N of village, quebradaLa Chorrera,1100 m, 22 Mar Stems 1-2 plus several suckers,to 10 m tall, 6-10 1981, Henao et al. 299 (holoneotype,COL; isoneocm diam., armedwith black, to 8 cm long spines, distypes, AAU sterile, HUA). tally with old leaf sheaths.Leaves 4-6; sheath60-80 cm long, armedwith black,to 10 cm long spines;petiCaespitose,with up to 10 stems, these 3.5-5 m tall, ole 60-100 cm long, green, unarmed;rachis210-260 ca. 3 cm diam., gray, sparsely armed with slender cm long, unarmed;pinnae 17-24 per side, somewhat spines, to 3 cm long. Leaves 11, horizontally held, golden-green,in groupsof (2-)3-4, in differentplanes, arrangedin 3 vertical rows; sheath ca. 18 cm long, cuneate, 2.5-3 times as long as wide, lobulate prae- with a dense light brown indument,densely armed morse at apex, glabrous adaxially, rough abaxially; with black, to 1 cm long spines; petiole 14-16 cm middle pinnae 37-58 x 15-22 cm. Peduncle 110-170 long, green, with a brown,caducousindument,armed cm long, 1-1.5 cm diam. at apex, with scattered with manyblack spines, to 5 cm long; rachis 102-108 brown, to 5 cm long spines; rachis 60-100 cm long, cm long, with indumentand spines like sheath, but unarmedor with scatteredspines like those on pedun- spines fewer; pinnae 17-20 per side, subregularly cle; rachillae 50-100, densely brown-spinulose;basal inserted in lax groups of 2-4, in one plane, plicate, rachillaeto 95 cm long, basally withoutflowers for up rigid, cuneate, 3-6 times as long as wide, incisedto 10 cm, 3-4 mm diam in the androgynouspart. praemorse,with an up to 7 cm long finger-likeproStaminateflowers pale violet, 1.5-2 mm long; anthers jection on the distal margin, adaxial side glabrous, 0.6-0.8 mm long. Pistillateflowers 6-7 mm long; pis- abaxial side very minutely spinulose, with a single til glabrous.Fruitrose-red,globose, 16-22 mm diam; black, 3-4 cm long spine basally on the midrib;basal pinnae 12-17 x 2-4 cm; middle pinnae 21-37 x 6-7 endocarp15-19 mm diam., globose, deeply pitted.
FloraNeotropica
66 cm; apical pinnae 2-3 ribbed, 24-32 x 6-8.5 cm. Inflorescenceinterfoliar;prophyllgreen, 38 x 1 cm; peduncular bract ca. 122 cm long, 1.7 cm wide, brown,minutelyspinulose, spineless;peduncle87-96 cm long, 4 mm diam. at junction with rachis,covered with brown, ca. 3 mm long spinules, armedwith few to many,black, to 5 cm long spines; rachis34-45 cm long, with a loose whitish indument, unarmed; rachillae ca. 19, light green, with an indumentlike that on rachis, unarmed;basal rachillae 28-33 cm long, ca. 2 mm diam., with triadsfor /2 the length,distally staminate;apical rachillaeca. 8 cm long, staminate. Triadsinsertedin pits in the rachillae,exposed, pistillateflowers sunkenfor ca. V2theirlength.Staminate flowers white, with a purplish brown corolla, 1.5-2 mm long; sepals narrowlytriangular,ca. 1 x 0.5 mm; petals ovate-acuminate,free, 1.5-2 x 1.5-2 mm; filaments flattened, 0.5-0.9 mm long, anthers oval, 0.3-0.4 x 0.3-0.5 mm; pistillode trifid, ca. 0.5 mm tall. Pistillate flowers (before anthesis) 2.5-3 mm long; sepals broadlyovate, ca. 1 x 2-3 mm, enclosed in the floral pit; petals 2.5-3 x 2-2.5 mm, connatefor 1/3 of their length, valvatedistally;staminodialcup ca. 1.5 mm tall, truncate,teeth indistinct;pistil ca. 2 x 1 mm, glabrous,stigmas white. Fruitsnot seen. Illustrations. Figs. O1F(pollen), 18C (distribution map), 21J (middle pinnae). Aiphanes leiostachys is at presentrepresentedonly by one collection, the neotype, collected in premontane wet forest at the northernend of the Cordillera Centralin Colombia,whereA. leiostachysis rarein a largely deforested area. The species was considered endangeredby Bernal (1989). In the caespitose habit and the incised praemorse,cuneatepinnaeit recallsA. erinacea, but pinnae of A. leiostachys are subregularly inserted instead of strongly grouped, borne in one plane instead of several, and very rigid, resembling those of A. lindeniana more than those of A. erinacea. Furthermore,the rachillaeof A. leiostachys are glabrous, whereas those of A. erinacea are densely spinulose. Inflorescences of A. leiostachys are remarkablein having flowers rightfrom the bases of the rachillae and in their lack of spinules. Pollen (Fig. 1OF)resembles that of A. lindeniana. 12. Aiphanes lindeniana (H. Wendland)H. Wendland in O. de Kerchove de Denterghem,
Les
Palmiers230. 1878. MartinezialindenianaH. Wendland,Linnaea28: 349. 1857. Type. COLOMBIA.Norte del Santander:Pamplona,2000
m, Funck& Schlim1655 (lectotype,annotatedby N. Imschanitzkaja, LE). AiphanesconcinnaH. E. Moore,GentesHerb.8: 223, 224, Cundinamarca: Near fig. 91. 1951.Type.COLOMBIA. 3000 m, 12 Oct 1946,Foster& Foster1870 Fusagasuga, (holotype,BH;isotype,A). Caespitose,with up to 10 stems, rarely 1-stemmed. Stems 1.5-7 m tall, 3-7(-10) cm diam., armed with black spines, to 10 cm long. Leaves 4-10, distichous or more rarely polystichous;sheath 15-42 cm long, densely armedwith black spines, to 12 cm long; petiole 6-56 cm, with a white or brown, caducous indument,andnumerousblack spines, to 8 cm long; rachis 38-172 cm long, with indumentandarmaturelike that on the petiole, but spines fewer and shorter;pinnae 18-45 per side, insertedin groupsof 2-7 separatedby up to 11 cm, lanceolateto narrowlycuneate,(3.5-)511 times as long as wide, often strongly plicate, truncate-or obliquelypraemorseat apex, with an up to 2.5 cm long finger-likeprojectionon the distal margin, nearlyglabrousor spinulose,especially abaxially, margins lined with 1-2 mm long spinules, midrib adaxially with a row of thin spines, to 1 cm long, abaxiallywith 1-6 rigid spines, to 2.5 cm long; basal pinnae6.5-20.5 x 0.6-1.5 cm; middlepinnae 16-34 x 1-7 cm; apicalpinnae2-7 ribbed,8-18 x 2-13.5 cm. Inflorescenceinterfoliar,erect, branchedto 1 order; prophyll 21-47 x 1.1-3.4 cm; peduncular bract 57-142 cm long, 2-6 cm wide, thick, with a thin, brown, caducous indument, unarmedor with black spines, to 3 cm long; peduncle 32-100 cm, with a brown indument, sparsely to densely armed with black spines; rachis 18-46 cm long, unarmed or armed for a variable part of the length with black, crimped spines, to 1 cm long; rachillae 16-68, densely covered with minute, brown or ferruginous spinules, ca. 0.1 mm long; basal rachillae 16-42 cm long, without flowers for 0.5 to 26 cm basally, with triads for /2-2/3 of the remaininglength, in this part 2-5 mm diam.,distallyca. 1 mm diam., with dyads of staminateflowers; apical rachillae 3.5-14 cm long, staminate;flower groups slightly sunken into the rachillae;triadssubtendedby a small bract,covering up to 1/4 of the pistillateflower;dyads subtendedby a minute triangularbract. Staminateflowers white to violet, 2-3 mm long; sepals imbricate,briefly connate at base, 2-3 mm long; petals 2-3 mm long, nearly free; filamentsca. 1 mm long, anthersalmost square to linear, 0.6-1.4 x 0.6-0.8 mm; pistillode small, trifid. Pistillate flowers 3-7 mm long; sepals imbricate, 3.5-7 mm long; petals connate for /2 of their length, valvate distally, ca. 4 mm long; staminodial
SystematicTreatment
67
(st), Rangelet al. 5948 (COL);ParqueNaturalRegional Ucumari,2200-2450m, 14 Jun1989(fr),Gonzalezet al. 1685 (COL);Mun.de SantaRosa,betweenTermalesand Paramode SantaRosa,2500 m, 18 Jul 1980(fl), Idroboet al. 9728(COL).Santander:Charala, rd.Duitama-Virolin, km 50-55, veredaEl Taladro,2250-2300m, 6 Dec 1978 Illustrations. Figs. 10E (pollen), 19D (distribution (fl),Diaz1634(COL);rd.to Piedecuesta, 7 kmS of rd.from to Pamplona,2200 m, 18 Mar 1987 (fl), Bucaramanga map), 21D (middle pinnae), 26 (drawing). Bernal & Galeano1352 (AAU, COL).TOLIMA: Santa Local names. Cuaro, cuvaro, mararai(Cundina- Isabel,cerroPurima,2540 m, 1 Aug 1980(femfl), Idrobo et al. 10400(COL). marca). Aiphaneslindenianais variablewith respectto size Distribution and habitat. Endemic to Colombia, and form of the pinnae, size of the inflorescence, and where it is widely distributedalong the Cordilleras the lengthof the basal flowerless partof the rachillae. Orientaland Central,from the junction of these and It is closely relatedto the south EcuadoreanA. verrunorthwardsto the departmentof Antioquiain Cordicosa, under which the differences are discussed. In lleraCentralandat leastto the departmentof Nortedel general termsA. lindeniana can be distinguishedby Santanderin CordilleraOriental.It grows in humid, its caespitose habit; fierce armatureof stem, leaves, frequentlycloudy zones between 1900 and 2700 m, and inflorescences;numerous,densely inserted,linear andit is often a conspicuouselementin the uppermon- to narrowlycuneate,often very rigid pinnae,normally tane forest. It is often conserved in deforestedareas, with a characteristicrow of thin spines adaxially on where it seems unableto regenerateby seeds. the midrib; glabrous pistil; and red fruits, with an almost smooth endocarp. Leaves are mostly distiSpecimens examined. COLOMBIA. ANTIOQUIA: but this characterpresentssome variation,and chous, km Mun.de Cocornm, 62, hwy.Medellin-Bogota quebrada LaRoca,2000m, 13Sep 1980(fl), Galeano&Bernal253, individualswith polystichousleaves are known from 254 (COL);Mun.de Medellin,veredaSantaHelena,12 km the northernpartsof CordillerasCentraland Oriental. E of Medellin,2400 m, 18 Dec 1980(fl, immfr),Galeano distichous and polystichous leaves et al. 331 (COL);Mun.de SantaRosade Osos,ca. 10 km Occasionally, in the same occur population(Gentry et al. 47724, NE of village,CerroSan Jose, 2630 m, 7 Jan 1985 (fl), Bernal & Galeano 849 (AAU, COL);Mun. de SantaRita, 5 47756). Leaf anatomy is also variable; specimens kmE of Guatape, 2000m, 17Sep 1987(fl),Bernal&Tob6n from the northernpartof CordilleraOriental(Bernal 1375 (AAU,COL);vicinityof La Uni6n,2600 m, 21 Aug & Galeano 1352; Grant10538) have the non-vascular 1948 (fl, fr), Barkley & Johnson 18C864 (COL, MEDEL). Mun.deArcabuco, 3 kmon rd.to LaPalma,2700 fiber strandsarrangedin a single layer of relatively BOYACA: m, 14 Mar 1987 (fl), Bernal & Galeano 1334 (AAU, COL), thick bundles,an arrangementotherwiseencountered 1335(COL);betweenMoniquira andArcabuco, 2150m,25 only in A. eggersii; specimens from localities further Feb 1940 (fl, fr), Perez-Arbelaez & Cuatrecasas 8173 south (Gentry 47756) have fibers arrangedin 2-3 (COL);quebradanearRio Cusiana,1900m, 23 Jul 1967 of thinnerbundles showing a transitionto the layers (imm fr), Jaramillo et al. 2750 (COL). CUNDINAMARCA: Las Mesitas,S side GuavioRiver,20 km NE of Gachala, patternfound in A. verrucosafrom southernEcuador. 2135 m, 30 Oct 1944(st), Grant10538(BH, COL,NY); 2700m, 18Apr1946 LosArenillos,nearrd.to Fusagasuga, 3367 (COL);SanAntoniode Tena, 13. Aiphanes linearis Burret, Notizbl. Bot. Gart. (fr), Duque-Jaramillo veredaChicaque,1900m, 15 Mar1980(fr),Idrobo9501 Berlin-Dahlem11: 577. 1932. Type. COLOMBIA. (COL);rd. fromZupata,6 km beforePacho,25 Feb 1985 Antioquia:Paramillo("Paranullo"),ca. 2300 m, 26 (fr), Galeano et al. 564 (COL). HUILA:La Plata, Finca Jul 1880, Kalbreyer1883 (Bt). Neotype (Bernal et E of VolcanPurace,100kmE of Popayan,near Merenberg CaucaborderE of Leticia,2200-2300m, 11Jul 1975(fr), al., 1989). COLOMBIA. Antioquia: Mun. de Diaz et al. 453 (COL); 5 Dec 1980 (fr), Croat51923 (MO); Frontino,rd. Nutibara-LaBlanquita,1800-1900 m, 7 Jul1984(fl, fr),Gentryet al. 47724(MO,NY);8 Jul1984 8 Jan 1982, Galeano & Bernal 488 (holoneotype, (fl, fr), Gentryet al. 47756 (MO,NY); 4 Apr 1986(fr), COL; isoneotype, HUA). et al. 53978 Gabinete, (K). HUILA-CAQUETA: Gentry 8473 2300-2450m, 22 Mar1940(fl, immfr),Cuatrecasas Dugand,Caldasia2: 455, fig. p. 456. Claras,2000 m, 7 Feb 1985 Aiphanesechinocarpa (COL);rd. Algeciras-Aguas BetweenSons6nand 1944. COLOMBIA. Antioquia: Type. 126 & Bernal Henderson 125, (COL, NY). QUINDIO: (fl), Nariio, near Rionegrito,2000-2200 m, 11 Dec 1943, rd. Salento-LaCeja,2520 m, 25 Nov 1986(immfr), G. Uribe-Uribe 683 (holotype,COL;isotypes,BH,F,NY). Mun. de Pereira, Arbeldez et al. 1842 (HUQ). RISARALDA: betweenEl CedralandLa Pastora,2250-2350m, 14 Jun Caespitose with up to 15 stems, these 5-10 m tall, 1989(fr),Galeanoet al. 2061 (COL);La Pastora,2320 m, 30 Jul 1989 (fr),M. P. Galeano199 (COL);14 Oct 1989 4-12 cm diam., densely armedwith black, applanate cup ca. 2 mm high, briefly dentate;pistil glabrous,ca. 3 mm long. Fruitsred or orange,subglobose,slightly rostrate,14-16 mm diam.;endocarp13-16 mm diam., slightly pittedtowardapex.
68
Flora Neotropica
10,1~~~~~~~~~~~~~~~~$
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,!-d
FIG. 26. Aiphanes lindeniana. a. Leaf x0.20; b. Inflorescence x0.30; c, d. Staminate flowers, face and back x3.57; e. Developing fruit x2.038; f. Fruitx1.079; g. Endocarpxl.79; h. Pinna x0.40. Reproducedfrom Moore, Gentes Herb. 8, 1951 (Aiphanes concinna).
SystematicTreatment spines, to 35 cm long. Leaves 5-9, distichous, erect and arching;sheath 46-220 cm long, densely armed with black, applanatespines, to 12 cm long; petiole 0-34 cm long, armed like the sheath;rachis 99-239 cm long, covered with purple spinules, with black spines at least abaxially; pinnae 35-48 per side, inserted in groups of (1-)2-9 separatedby 4-15 cm, in differentplanes or sometimes distally in one plane, linear to narrowly cuneate, 6-12 times as long as wide, obliquely praemorse at apex, with a 0-2 cm long finger-likeprojectionon the distal margin,adaxially glabrous, abaxially glabrousor minutely spinulose, marginslined with short spinules; basal pinnae 23-64.5 x 1-6 cm; middle pinnae 23-91 x 4-9 cm; apical pinnae 2-6 ribbed, 14-55 x 1.5-15 cm. Inflorescence erect at anthesis,curvedin fruit, 1-2 m long; prophyll 15-92 cm long, abaxially densely covered with black spinules and spines, to 2 cm long; peduncular bract 57-200 cm long, 5.5-19 cm wide, thick, woody, persistent, densely covered with black and yellow spinules and fewer spines, to 2 cm long; peduncle 34-85 cm long, at anthesis2-3 cm in diam. at junction with rachis, thicker in fruit, nearly unarmed to densely armed with black spines, to 6 cm long, with 1-4 rudimentarypeduncularbracts, to 18 cm long; rachis 16-87 cm long, unarmed,with a thin brown indument;rachillae 35-80, densely spinulose, each subtended by a minute or up to 3.5 cm long bract;basal rachillae 16-56 cm long, basally without flowers for up to 4 cm, with triads for 1/4-1/3 of the length, in this partthickened,up to 10 mm diam.,distally slender, 1-2 mm diam., with densely packed dyads of staminateflowers; apical rachillae9-26 cm long, staminate;flower groups sunkeninto shallow to deep pits in the rachillae, subtended by a 1-2 mm long bract;each flower in addition subtendedby an up to 10 mm long bracteoleresemblingthe pistillate sepals. Staminateflowers violet, 2-4 mm long, those of triads largest, with an up to 10 mm long pedicel, those of dyads smaller,sessile or shortlypedicellate; sepals nearly free, narrowlytriangular,1.5-2.6 mm long; petals briefly connate at base, valvate, 2-3.5 mm long; filaments ca. 1.5 mm long, antherslinear, 1.-1.5 mm long; pistillode minute, trifid. Pistillate flowers white with violet apex, 9-15 mm long; sepals rounded, imbricate, covered with yellow spinules, 6-10 x 6-10 mm, often nearly enclosing the petals; petals connate for ca. /2 of theirlength,imbricatedistally, sparsely covered with yellow spinules on the basal half of the outer side, 9-16 x 6-9 mm, corolla lobes reflexed at anthesis;staminodialcup 7-13 mm high, almost truncate, adnate to corolla tube; pistil
69 11-13 mm long, densely covered with yellow, applanate spines, ca. 3 mm long. Fruits densely packed, 22-45 x 18-35 mm, angular-turbinatefrom mutualpressures,covered with golden, ca. 1 cm long spines; endocarpacute at base, 17-35 x 16-28 mm, shallowly pitted and irregularlyfurrowed. Illustrations. Figs. 12K, 13B (pollen), 19E (distribution map), 21B (middle pinnae), 27 (habit, fruits); Galeano & Beral, 1987: 31, fig. 7. Local names and uses. Chascaray,chirca, cirqui, corocito de agua, corozo de agiiita, (Antioquia); chonta (Valle). Seeds are edible and are consumed locally. Distribution and habitat. Endemic to Colombia, where it is found in the northernpart of Cordillera Centraland on the westernslopes of CordilleraOccidental from the departmentof Antioquia to the departmentof Valle, in humid to wet, lower montane forest, often cloud forest, at 1800-2600 m. Often left in pasture. Specimens examined. COLOMBIA. ANTIOQUIA: Mun.de Cocorna,rd. Medellin-Bogota, km 62, veredaLa Roca, 2000-2100 m, 19 Mar 1980 (fl, fr), Galeano& Bernal131 (AAU,COL,HUA,NY);Mun.de Itagii,basin of quebrada DofiaMaria,above2500 m, Jul 1942(fl, fr), km Londoios.n.(COL);Mun.de Peque,rd.Uramita-Peque 45-48, veredaLa Tumba,2500 m, 2 Apr 1983 (fl, fr), Bernal & Galeano 534 (COL, HUA); 4 Apr 1983 (fr),
Bernal& Galeano570 (COL,HUA);Mun.de Urrao,trail to ParqueNacionalNaturalLasOrquideas, Paramode San Pedro,2100 m, 23 Jun 1982(fr),Bernal& Galeano379 (COL,HUA).VALLE:Mun.de Bolivar,FincaLa Zulia,3 km N of Betania,1900m, 12 Apr 1989(fl, fr),Bernal& Devia1533(COL,TULV);HaciendaTokio,ca. 10 kmS of behindmicrowave tower,2000 m, 26 Feb 1983 Queremal, (fl), Gentryet al. 40805 (COL, MO).
Aiphanes linearis is one of the most conspicuous species in the genus, readily distinguished by the densely clustered,golden-spinyfruits.Vegetativelyit is characteristic in its caespitose habit, strongly armedstem with spines up to 35 cm long, and distichous leaves, with linear to narrowlycuneate, clusteredpinnae,borne in differentplanes.There is some variationin size and shape of the pinnae, from linear to narrowlycuneate, a kind of variationthat is very common in the genus. 14. Aiphanes macroloba Burret,Notizbl. Bot. Gart. Berlin-Dahlem 11: 576. 1932. Type. COLOMBIA. Antioquia: Cieneguetas ("Cinegetas"),1350-1650 m, 23 Jul 1880, Kalbreyer 1887 (Bt). Neotype
Flora Neotropica
70 ?
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B
of ValleandChoc6 on dividinglinebetweenthedepartments individual FIG. 27. Aiphaneslinearis.A. Single-stemmed on roadfromCartagoto SanJosedel Palmarin Colombia.B. Fruits(Galeano&Bernal131). (Bernal et al., 1989). COLOMBIA. Antioquia: Mun. de Frontino, Cieneguetas, 1000 m, 21 Mar 1982 (fl), Bemal & Galeano 279 (holoneotype, COL; isoneotype, HUA). Burret,Notizbl.Bot. Gart.BerlinAiphanesmonostachys Dahlem11: 574. 1932 (in part,inflorescenceonly, non 38: Burretemend.Bernal,Brittonia Aiphanesmonostachys 68. 1986). AiphaneschocoensisA. H. Gentry,Ann. MissouriBot. Gard.68: 112. 1981. Type. COLOMBIA.Choc6:Rio of Rio El Vallebetweenthebaseof Alto Mutat5,tributary del Bueyandmouthof river,100-150m, 9 Aug 1976(fl, fr), Gentry& Fallen17450(holotype,COL;isotypes,BH sterile,MO). Caespitose, with 1-5 stems, these up to 2 m tall, 2-3 cm diam., erect or decumbent, sparsely armed with black, to 3 cm long spines, often with suckers and adventitious roots both at base and at the distal nodes, sometimes just below the crown. Leaves 5-8, borne more or less horizontally, lamina entire or divided in a large many-ribbedtop segment and 2-3 one-ribbed basal pinnae; sheath 7-17 cm long, sparsely armedwith black, to 1 cm long spines; petiole 10-36 cm long, with a brown,scaly indumentand few brown, to 1 cm long spines; rachis 36-71 cm long, densely brown-spinulose abaxially; blade (or
apical pinna in divided leaves) 30-82 x 18-37 cm, with 13-20 ribs, bifid at apex for 10-19 cm, outer marginpraemorse,adaxialside with a row of fine, ca. 1 cm long soft spines along each rib, otherwise glabrous or sparsely spinulose, abaxial side densely covered with brown,ca. 0.5 mm long spinules; basal pinnae of divided blades one-ribbed, narrowly to broadly cuneate, to 18 cm long and 8 cm wide, obliquely praemorse at apex, with a 2-4 cm long finger-like projectionon the distal margin. Inflorescence interfoliar,erect, spicate; prophyll 11-21 cm long, 1-2 cm wide; peduncularbract38-81 cm long, 1-2 cm wide, with a brown indument, unarmed or with some brown, to 1 cm long spines; peduncle 34-120 cm long, 3-5 mm diam. at apex, with a light brown indument, and armed with numerous brown, ca. 3 mm long spinules, and sometimes also a few black, to 1 cm long spines; spike 12-28 cm long, 410 mm diam., brown to black spinulose, with triads for ca. 1/2of the length,distally with dyads of two staminate flowers or one pistillate and one staminate flower;flower groupsdeeply sunken.Staminateflowers orange or greenish yellow in bud, white at anthesis, 4-7 mm long, larger than the pistillate ones; sepals narrowlytriangular,connate for up to V2 their
71
SystematicTreatment length, 1.5-3 mm long; petals connate basally for 1-1.5 mm, valvatedistally,4-6.5 mm long; filaments ca. 1 mm long, anthers nearly square, 0.3-0.9 mm long; pistillode minute.Pistillateflowers 3.5-4.5 mm long; sepals free, imbricate, 2.7-3 mm long; petals briefly connate at base, valvate, 3.5-4.5 mm long; staminodialcup 1-1.5 mm tall, nearlytruncate;pistil ca. 2 mm long, 0.6 mm diam., glabrous.Fruit ellipsoid, rostrate,ca. 10 mm long, 7 mm diam., red or orange when ripe; endocarpsmooth. Illustrations. Figs. 3B (stem), 5D (leaf), 15B (stem anatomy), 19A (distributionmap), 21A (leaf).
teredwith fruits.Insteadit reproducesvegetativelyby means of suckers formed from both basal and distal nodes on the often procumbentstem, and through stolon formation. The confusionregardingtypificationandsynonymy of A. macrolobawas resolved by Bernal (1986). See also notes to A. hirsuta subsp. hirsuta.
15. Aiphanes minima (Gaertner) Burret, Notizbl. Bot. Gart.Berlin-Dahlem11: 558. 1932.
Fruct.Sem.PI.2: 269, t. 139,fig. BactrisminimaGaertner, 5. 1791. Type.Hermanns.n. (holotype,TUB, a single Local names and uses. Peganore (Coaiquer); endocarp in theseedcollectionof J. Gaertner). 70. Palmetum Martius, Orbignyanum palmito (Ecuador).The CoaiquerIndiansin Ecuador Bactrisacanthophylla 400 m, 1827(juv), RICO.Yrurena, 1844.Type.PUERTO eat the palm heart. Webbianum, Wydler192 (holotype,FI, n.v.,in herbarium Distribution and habitat. WesternColombia and photosin AAU;isotype,G). Ecuador from the departmentof Antioquia in the Martinezia acanthophylla (Martius) Beccari in Urban, Symb.Antill.8: 79. 1920. north, along the western slopes of CordilleraOcci- Aiphanes acanthophylla (Martius) Burret, Notizbl. Bot. dental and occasionally also in the Pacific lowlands 11:558. 1932. Gart.Berlin-Dahlem 66. 1844. south to the Province of Esmeraldas in northern BactriserosaMartius,PalmetumOrbignyanum ased on Palmadactylifera,aculeata,fructucorallino, Ecuador, in lowland and premontane rain forest, Nov.PI.Gen.3, tab.msc.42. 1703.Type. minor;Plumier, reaching ca. 1400 m in northwesternColombia and Plumier,tab. msc. 42. (holotype,BibliothequeCentrale, ca. 1000 m in northwesternEcuador. Paris,n.v.;isotype,K, uncolored copy). 76. corallinaMartius,Palmetum Martinezia Orbignyanum COLOMBIA. examined. ANTIOQUIA: Specimens 1844.Basedon Palmadactylifera,aculeata,fructucoraMun.de Frontino,Murri,Cieneguetas,1350-1650m, 21 llino, major;Plumier,Nov. PI. Gen. 3, t. msc. 39-41. 1703. Mar 1982 (st), Bernal & Galeano 268 (COL); Mun. de Type.Plumier,tab. msc. 39-41 (holotype,Bibliotheque to Rio a La Calles, Urrao,quebrada Agudelo, tributary Centrale, Paris,n.v.). 1300-1400m, 15Jun1982,Bernal& Galeano342 (COL). in Kerchovede H. Wendland corallina(Martius) Aiphanes CHOC6:Mun. de El Carmende Atrato,rd. MedellinLesPalmiers230. 1878. Denterghem, Quibd6km 150,veredaEl Doce,RioElAguil6n,680 m,Jul Curima corallina (Martius)Cook, Bull. Torrey Bot. Club 1979,Galeano& Bernal109 (COL);Mun.de SanJosedel 28:563. 1901. of RioHdbita,finca Martinezia erosa Palmar,basinof RioTorito,a tributary Linden, Catalogue no. 87: 5. 1871. 700 m, 7 Mar1980(fl, fr), Foreroet al. "LosGuaduales," (heredesignated). Specimencollectedin Horto 6863 (MO);20 Mar 1980 (fr), Foreroet al. 7535 (COL, Lectotype Dec 1871 (fr), Patin s.n. (lectotype, K). Lindeniano, MO);sameregion,Altodel Oso, 1300m, 14 Mar1991(fl), Aiphaneserosa(Linden)Burret,Notizbl.Bot.Gart.BerlinBernal & Borchsenius1961 (AAU, CHOCO,COL). ECUADOR. ESMERALDAS: San Marcos de los Coai-
Dahlem 11: 558. 1932.
CurimacolophyllaCook,Bull.TorreyBot.Club28:561,pl. queres and surroundingforests, on trail Chical-Tobar 48. 1901. Type. PUERTORICO. Bayamon(fl, fr), Donoso,600-1000m,24 Nov 1983(fl),Barfodet al. 48916 Underwood& Griggs 878 (holotype, US). (AAU, QCA, QCNE); Feb 1985 (fl), 0llgaard et al. 57620
Aiphanesluciana Bailey, Gentes Herb.8: 166, fig. 65. 1949. (AAU, QCA, QCNE);new rd. from Lita towardSan Type.SAINTLUCIA:Barrede l'Isle,24 Mar1948(fl, fr), Lorenzo,km 15-18, 800-900 m, 14 Oct 1987(st),Skov& Bailey440 (holotype,BH;isotype,BM). Borchsenius64747 (AAU, QCA, QCNE); 12 Nov 1987 (fl), Bailey,GentesHerb.8: 170,fig. 68. Aiphanesvincentiana Skov,Borchsenius.et al. 64818 (AAU, QCA, QCNE). 1949. Type.SAINTVINCENT.Cumberland Valley,in woodlandsandthickets(fl, fr),J. Beard230 (holoforest, Aiphanes macroloba is the only species in the type,BH). genus thathas an entireor almostentirelamina.In the north of its distributionalrange the lamina is someSolitary. Stem (2-)5-18 m tall, 6-20 cm diam., times divided into a large terminalsegmentand a few armedon the interodes with rings of black, flattened basal one-ribbed pinnae; furthersouth in Choc6 and spines, often becoming almost unarmed with age. in Ecuadorthe laminais always undivided.Staminate Leaves 10-20, spreading;petiole 15-110 cm long, flowers are peculiarin being largerthan their female armedwith black, to 8 cm long spines; rachis 130-counterparts. Fruits are unique in being elliptical 400 cm long, unarmedor with many black spines, to insteadof globose. The species is only rarelyencoun- 6 cm long; pinnae 18-34 per side, regularlyinserted,
72 4-10 cm apart,all in one plane, linear,or more rarely (PuertoRico) wideningat apex, 5-12 times as long as wide, obliquely praemorseto lobulate-praemorseat apex, + caudateon the distal margin,glabrouson both sides, abaxially unarmed,or with many black spines, to 3 cm long, adaxially often with a row of ca. 1 cm long spines on the midrib;basal pinnae 24-26 x 1-2 cm; middle pinnae 31-80 x 4-11 cm; apical pinnae 2- to several-ribbed,25-34 x 9-22 cm. Inflorescence interfoliar,curving, once or rarely twice branched; peduncularbract 60-190 cm long, 1.5-8 cm wide, coriaceous to woody, unarmedor densely spiny, with a grey or white, caducous indument;peduncle 28130 cm long, 3-22 mm diam. at junction with rachis, densely covered with black spines; rachis 25-150 cm long, unarmed;rachillae 12-300, with a peltateindument, often becoming glabrous;basal rachillae 10-50 cm long, with triads for ca. /2 of their length, in this part2-4 mm diameter,distally staminate,taperingto 1-2 mm diam.; apical rachillae5-15 cm long, staminate; triadsborne superficiallyor in a shallow cavity in the rachilla; dyads superficial. Staminateflowers creamish white to yellow, 3-4 mm long; sepals triangular, carinate,much shorterthan the petals, 0.6-3.5 mm long; petals nearly free oblong-acuminate,elongate, 3.4-6.1 mm long; filaments flattened, 0.9-1.8 mm long, antherslinear, sagittate at base, with dark connective, 1.8-2.4 x 0.5-0.9 mm;pistillodedistinct, trifid, 0.4-1 mm high. Pistillate flowers creamish white to yellow, 3-4 mm long; sepals broadlyovate, free, imbricate, 1.2-1.8 mm long; petals ovate-acute, connate for /2of their length, valvatedistally,3.3-3.8 mm long; staminodialcup 1.2-2.5 mm high, deeply acutely lobed to nearly truncate; pistil 2.8-3 mm high, glabrous.Fruitsred, 12-16 x 14-17 mm; mesocarp mealy-fleshy; endocarp 8-12 x 10-16 mm, weakly to prominentlypitted-grooved. Illustrations. Figs. 10B, 12A (pollen), 18A (distributionmap), 21C (middle pinnae), 28, 29 (drawings); Bailey, 1949: 169 (as A. erosa), 171 (as A. vincentiana), 172; Little & Wadsworth1964: 37 (as A. acanthophylla);Cook, 1901:, pl. 48 (as Curimacolophylla);also photosin severalpopularbookson palms, underthe synonymsA. acanthophyllaandA. erosa. Local names and uses. Grigri (Martinique, St. Vincent, Dominica, St. Lucia);chou picant,glouglou, glouglou rouge (Martinique);palmade coyor,coyore, coyure, coyora (Puerto Rico); macaw palm (Barbados). The seed is edible; fruitsare boiled and cracked open with a hammer(S. Carrington,pers. comm.).
FloraNeotropica Distribution and habitat. Widely distributed in the Lesser Antilles: Dominican Republic, Puerto Rico, Dominica, St. Vincent, St. Lucia, Martinique, Barbados, and Grenada. Also a widely cultivated ornamental palm. Aiphanes minima grows in different habitats. In the northern part of its range (Dominican Republic, Puerto Rico) it is found on limestone hills; in the southern part (Dominica, St. Vincent, St. Lucia, Barbados) it grows in subcanopy or understory of seasonal to wet forest, sometimes in deep shade. Specimens examined: DOMINICAN REPUBLIC. LA VEGA:CordilleraCentral,Bonao, Rio Maim6n, 250 m, 17 Dec 1930 (fl), Ekman 16429 (S); Peninsula de Samana, LagunaSamana, 100 m, 3 May 1930 (fl), Ekman14845 (G, K, NY, S, US). PUERTO RICO. Candelaria,nearBayamon(fr), Britton et al. 2857 (F, MO, NY, US); Fajardo and vicinity, Rio Arribe,2 Mar 1913 (fr), Britton& Shafer 1701 (F, MO, NY, US); Haystackhills, near Vega Baja, 1937 (juv), Horn s.n. (BH); Maricao, 30 Nov 1884 (juv), Sintenis 484 (BM, G, GH, K, US); PuebloViejo, 19 Jul 1914 (fl), Stevenson2105 (NY, US); Punas Apuesto, Vega Baja, Dec 1899 (st), Goll 1044 (NY, US); VegaBajaof Espinosa,Apr 1927 (fr), Barker s.n. (BH); nearToaAlta, 29 Mar 1923 (fl, fr), Britton& Britton 7854 (G, NY); nearToa Baja, 100 m, 2 Jun 1954 (fl, fr), Little 16305 (BM, GH, NY); near VegaAlta, May 1932 (fl, fr), Bailey 61 (BH, MO); near Hato Grande,31 Aug 1885 (fl), Sintenis2628 (BM, F, FI, G, GH, K, LE, M, MO, NY, US); near Yuncos, mount Hormigas, 19 Sep 1885 (fl, fr), Sintenis2500 (BM, F, Fl, G, GH, K, LE, M, MO, NY, US); vicinity of Utuado,7 Mar 1915 (st), Britton5230 (NY); 15 Mar 1906 (fl, imm fr), Britton& Cowell396 (F, NY); 19 Mar 1906 (imm fr), Britton& Marble396 (US); between Utuado and Arecibo, 13 Mar 1906 (st), Britton & Marble 789 (NY, LOCALITY: 7 Aug 1915 (st), FLS 7767 US); UNKNOWN (NY); 7 Mar 1923 (fl, imm fr), Britton& Britton7829 (US). DOMINICA. Layou River, Clark Hall Estate, 15 m, 16 Jun 1964 (fr), Ernst 1701 (US); W of Salybia in Carib Reserve, 1 Mar 1942 (fr), Taylor 126 (BH, GH); S slope above Beux Branchestributaryof Pagua River, 300 m, 13 Jun 1964 (fl, fr), Ernst 1673 (US); ca. V2mile W of Pagua River,20 Jun 1964 (fl, fr), Ernst 1709 (US). MARTINIQUE: Piton du Champflore,Nov 1867 (fl), Hahn 347 (BM, G, GH). UNKNOWN 1871 (fl), LOCALITY: Hahn 1135 (GH, K, US). SAINT LUCIA: Fonds St. Jaques,700 m, 19 Mar 1889 (fl), Remages.n. (BM, K, NY); MourneLacombe,250 m, 9 Dec 1943 (fr), J. Beard173 (BH); Bar d'Isle, above Castries, 2 Feb 1932 (fl, imm fr), Loomis35 (BH, US); Castries,Piton Flore, 15 Sep 1945 (fl, fr), J. Beard 484 (BH); Piton Flore, (fl), P Beard 1058, 1059 (MICH);ibid., 625 m, 16 Mar 1971 (fl), Howard& Weaver17935 (A); Quilesse, on trailto head of Murrayrd., 1 May 1950 (fl), Howard 11696 (BH); near Quilesse, 10 Apr 1938 (fl, fr), Box 1760 (BM). UNKNOWN LOCALITY: 1879 (fr), Murrays.n. (K). SAINT VINCENT. Cumberlandvalley, 27 Jan 1963 (fr), Lee s.n. (BH); Kingshill, seasonal forest, 27 Nov 1945 (fr), P Beard 1374 (MICH); unknownlocality (juv), Unknown collector 239 (K). BARBADOS. TurnersHall Woods, Jan 1890 (fl, imm fr), Eggers 7135 (Fl, K, US); (fr), 7175 (B); 17 Nov 1891 (juv),
Systematic Treatment
73
i -,
FIG. 28. Aiphanes minima. From left to right: middle pinnae in adaxial and abaxial view (x0.39), endocarp (xl.74), infructescence(x0.39), and peduncularbract(x0.39); PuertoRico. Reproducedfrom Bailey, Gentes Herb.8, 1949 (Aiphanes acanthophylla).
74
Flora Neotropica
FIG. 29. Aiphanes minima. From left to right: inflorescence with peduncularbract attached(x ca. 0.40), middle pinnae in adaxial view (x0.40), and fruit (x nearly 2.38); St. Lucia. Reproducedfrom Bailey, Gentes Herb. 8, 1949 (Aiphanes luciana).
75
SystematicTreatment Warming83 (C); 6 Feb 1922 (fl, fr), Bailey & Bailey 291 (BH); Nov 1945 (fl), J. Beard 624 (BH); 4 Mar 1948 (imm fr), Bailey 428 (BH); 19 Jun 1974 (fr), Read 74-218 (US); gulch near Ape's hill, 1 Feb 1922 (st), Bailey & Bailey 352 (BH); near Battisheba, base of cliffs, 22 Feb 1924 (fr), Miller 93; St. John, Newcastle, 09 Feb 91 (st), Carrington 1590, 1592, 1610, 1611 (AAU); (fl), 1595, 1596, 1609 (AAU); (fr), 1598, 1612 (AAU); St. Thomas, Welchmann Hall (juv), Carrington1613 (AAU). CULTIVATED MATERIAL. BARBADOS: Cardington College, May 1933, Johnstone s.n. (B): BELGIUM:, Horto Lindeniana, Herb. Beccari s.n. (FI), BRAZIL: Rio de Janeiro, Bailey & Bailey 594 (BH); Dahlgren 611704 (F), BRITISH GUYANA: Georgetown,PromenadeGardens,May 1922, Dahlgren s.n. (F), Ceylon; Peradeniya garden, Jul 1935, Johnston 1672 (B), CUBA: Cienfuegos, Soledad, Atkins Gardens,Moore 6102, 6118, 6129 (BH); Santiagode las Vegas, Jun 1938, Acuna s.n. (BH); 1 Apr 1938, Bailey PuertoPlata, Sep 1981, 337 (BH), DOMINICAN REPUBLIC: Zanoni et al. 16879 (NY), ENGLAND: Royal Botanic Gardens Kew, Kew Acc 533-58.53301 (BH, K); Leon s.n. (K), GERMANY:Palm seeds purchasedfromHaageand Schmidt, Belle Terre, Erfurt, Cornell 416253 (BH) GUADELOUPE: Duss 3815 (FI); INDONESIA: Java, Bogor Botanic Garden, Herb. Beccari 90 (FI); 411 (FI); Sumatra, Botanic Garden, Lirzing 12134 (K), Jamaica: Bailey 15014 (BH); Hope gardens, Unknown collector 86 (K), MARTINIQUE: Duss CanalZone, SummitGardens,Jul 1931, s.n. (FI), PANAMA: Bailey & Bailey 440 (BH), PUERTORICO:FederalExperiment Station, May 1922, Bailey 60 (BH); Dorado Beach Country Club, Jun 1968, Read & Woodbury2055a (US). SINGAPORE: Botanic Gardens, Unknown collector s.n. (BM); TRINIDAD:Mar 1921, Bailey & Bailey s.n. (BH). UNITEDSTATES: FairchildTropicalGarden,Hull H-2 (BH); Dec 1969, Hull 10068 (FTG); Jan 1965, Read 1357 (BH, FTG); Mar 1975, Hill 2632 (FTG); Moore 5849 (BH); Oct Caracas, Parque 1978, Fantz 3469 (FTG); VENEZUELA: Nacional el Pifiar,Delgaso 287 (G).
Aiphanesminimais easily distinguishedby its large, solitarystem, and large leaves with regularlyinserted, linear or nearly linear pinnae, held in one plane. It is closely related to A. aculeata, which differs in its cuneate, abruptlywidening, grouped pinnae that are borne in different planes. One cultivated specimen (Kew Acc 533-58.53301)
has tricuspidate pinnae that
widen abruptlyat the relativelywide apex, and is perhaps a hybrid between A. minima and A. aculeata; the
two species are commonly plantedtogetherin botanical gardens. Taxonomy of the West Indian species has been con-
fused, and severalnames have been in use. As treated here, the groupconsists of a single variablespecies,A. minima. This treatmentis consistent with the pattern of variabilityobserved in other species. In the latest complete treatmentof Aiphanes in the West Indies, Bailey (1949) recognized five species here treatedas one: A. acanthophylla in Puerto Rico, A. vincentiana
in St. Vincent, A. luciana in St. Lucia, A. erosa in
Barbados,andA. minimaascribedto Martinique.The key charactersfor separatingthese were inflorescence size, armatureof pinnae and peduncularbract, shape of pinnae(whetherwith parallelmarginsor widening at apex), and pitting of the endocarp.Armatureand pinna shape are variablecharacters,and comparison of the specimens availablefrom the differentislands show no differencesthatcould justify the recognition of more than one species. Careful examination of endocarpsfromfruitingspecimenson all islandsshow a gradualvariationfromnearlysmoothto prominently pittedandgrooved,andseparationof species basedon this characterwould be impossible. Specimens with long, slenderpeduncleand few rachillae,corresponding to A. luciana Bailey, are known from the understory of dense rain forest in the interiorof Dominica, St. Lucia, and Martinique,at altitudes between 300 and700 m. They show a continuousvariationfrom 12 to 85 rachillae, and the largest inflorescences thus approachthose typical for the species in other areas (100-320 rachillae), including parts of Dominica (Ernst 1701), and Martinique(A. corallina Martius). Variationin size of inflorescence is commonly observed in the genus, often in correlationwith differences in habitatand altitude, and in the absence of other differencesbetween A. minima and A. luciana, the latterhas been includedin synonymy. The name Bactris minima was based on a single endocarp sent to Joeseph Gaertnerby the collector Hermann,without any indication of the origin. The type shows a perfect resemblancewith endocarpsof specimens collected on St. Vincent, including the type of A. vincentianaBailey. The originaldescriptionof Martineziaerosa Linden is a very brief descriptionof a juvenile plant,just sufficient to place the name in the genus Aiphanes with certainty. No type was designated, and only the Antilles was cited as place of origin, without further detail.In a latercatalogue,Linden(1881) ascribedM. erosa ("eroza")to Martinique.A specimen collected by Patinin Horto Lindenianoin 1871, and annotated Martineziaerosa, has here been designatedas lectotype. Certainlythis specimen,whichincludesthreeendocarpsandfoliage froma young plant,mustrepresent Linden'sMartineziaerosa. Burret(1932b) transferred M. erosa to Aiphanesand provideda full description based on a specimenfrom TurnersHall Woods, Barbados (Eggers 7125). Though Burretactuallydid not typify the name, as pointedout by Read (1979), this specimen has been cited as type of A. erosa (Glassmann, 1972), and the name Aiphaneserosa has been used since for Barbadanpopulationsof A. minima.
76
Flora Neotropica
cm wide, with a thin, caducous, brown, scaly indument, unarmed;peduncle 50-95 cm long, 2-4 mm diam. at junction with rachis, densely covered with minute,brownto black spinules, sometimes also with scattered,to 2.5 cm long, soft spines; rachis 16-42 cm long, minutely spinulose; rachillae (15-)40-70 (-120), slender, flexuose distally, densely covered with less than0.5 mm long, brownto black spinules; basal rachillae 7-13(-26) cm long, more or less appressedto rachis, with flowers from the base, or rarely with a basal, up to 12.5 cm long sterile part, with triads for 2/3 of the length, the remaining part with staminate dyads; middle rachillae 2.5-5 cm 16. Aiphanes parvifolia Burret, Notizbl. Bot. Gart. long, staminate,or with triadsbasally;distal rachillae Berlin-Dahlem 11: 569. 1932. Type. COLOMBIA. extremely short, 0.5-2 cm long, staminate,or with a few triads at base; triads slightly sunken into the Antioquia: Alto Caldera, forest, 2000 m, 14 Jan 1312 1932, Kalbreyer (Bt). Neotype (here desig- rachis, each subtendedby a 1-2 mm long bractcovthe pistillate and the proximalstaminateflower COLOMBIA. nated). Antioquia:Mun. de San Luis, ering km 16 SE of village on hwy. Medellfn-Bogota, before anthesis;dyads superficial,each subtendedby vereda La Josefina, 800 m, 10 Dec 1989 (fl, imm a similar,smallerbract.Staminateflowers purpleoutwhite inside, 1.1-1.5 mm long, those of dyads fr), Bernal, Borchsenius& Ramirez1752 (holoneo- side, distinctly pedicellate; sepals free, imbricate,keeled, type, COL; isoneotypes, AAU, BH, HUA, JAUM, /-3/4 of the petals, 0.9-1.4 x 0.5-1 mm; covering K, MO, NY). petals connate for '/3 of their length, valvate distally, Solitary. Stem (0.1-)1.5-2(-6) m tall, 2.5-3 cm acute, 1-1.5 x 1-1.5 mm;filaments0.3-0.6 mm long; diam., brown, unarmed or with few black spines; anthersslightly broaderthan long, 0.2-0.4 x 0.3-0.4 internodesgreen towardapex. Leaves 4-10, erect and mm; pistillode trifid, fused with receptacle and base arching; sheath 10-18 cm long, with a light brown, of filaments into a rounded, dark structure,ca. 0.7 scaly indument, often exposing the green surface mm diam. Pistillate flowers purple, ca. 2 mm long, below, armed with brown to black, to 2.5 cm long subtendedby a roundedbracteolethatcovers half the spines; petiole 8-25 cm long, unarmedor with many flower; sepals arched,roundedto very broad,enclosblack, to 3 cm long spines, adaxially with a light ing petals for 4/ or more, 1.5 x 2-4 mm;petals almost brown to ferruginous indument; rachis (24-)36-65 free, acute, 1.5-2.5 x 1-1.5 mm; staminodialcup ca. cm long, with a brown, scaly, caducous indument, 1 mm high, 1 mm in diam., almost truncate;pistil ca. unarmedor with scattered,black, to 1 cm long spines; 2 mm high, 1 mm diam., glabrous.Fruitorangeto red pinnae 5-10(-16) per side, inserted in groups of when mature,globose or elliptical, slightly rostrate, 2(-3) separated by 9-10 cm, in different planes, 7-10 x 7 mm; endocarpturbinate-subglobose,somerarely almost regularlyinserted,narrowlyto broadly what foveolate. cuneate, rarely linear, 1-4.5(-18) times as long as Illustrations. Figs. 6E (inflorescence), 19C (distriwide, truncate, oblique, or lobulate praemorse at butionmap), 21Q (middle pinnae). apex, with a 2-5.5 cm long finger-likeprojectionon the distal margin, adaxially glabrous, dark green, Distribution and habitat. Endemicto a small area abaxially paler, glabrous to rough due to numerous in northwesternColombia, on the eastern slopes of microscopic spinules, distal marginsometimes lined CordilleraCentralfromAmalfi to San Luis in the dewith a row of ca. 3 mm long, black spinules, midrib partmentof Antioquia,and on the western slopes of abaxially with 0-2 spines basally; basal pinnae 6- CordilleraOccidentalnearPueblo Rico in the depart17.5 x 0.6-4.5 cm; middle pinnae 9-24 x (1.8-)4-8 ment of Caldas. Grows in the understoryof primary anddisturbedpremontaneforest, at 800-1700 m. Apcm; apical pinnae 2-8 ribbed, 11-24.5 x (0.6-)4-30 cm, normally considerablybroaderthan the remain- parentlyvery restricted,and not found in secondary ing. Inflorescenceerect or arching,projectedbeyond forestor open areas.Most areaswithinits distribution leaves, branchedto I order;prophyll10-25 x 0.5-1.4 range are subject to intense deforestation,makingA. cm, unarmed;peduncularbract60-75 cm long, 1-1.5 parvifolia an endangeredspecies (Bernal, 1989). Aiphanesminimaundoubtedlyincludesseveralgeographicallyor ecologically separatedraces, but until more substantialinformationabout inter- and intrapopulationalvariationand ecology becomes available, it hardly serves any purpose to recognize more than one taxon. Possible futuredivision in subspecifictaxa will, by the way, inevitablylead to seriousproblemsof typification,since it will most likely be impossibleto assign the single endocarpthatconstitutesthe type of A. minimato any one of these. A similar,but less serious, problemwill arise with the type of A. erosa.
SystematicTreatment
77
Mun. petiole 7-26 cm, with a thin, brown, scaly indument, Specimens examined. COLOMBIA. ANTIOQUIA: de Amalfi,veredaEl Oso, 13-16 km fromAmalfitoward unarmedor with yellow spines, to 6 cm long; rachis Medellin,1590m, Sep 1988(fr),Betancuret al. 909 (MO); 26-67 cm long, with an indumentlike thaton petiole, La Viborita,1670-1700m, 28 Feb Peldarforest,quebrada 1961(COLNY);Mun. unarmedor with scattered,yellow spines and spin1990(femfl), Galeano&Henderson de SanLuis,km 16 SE of SanLuison rd.Medellin-Bogota, ules; pinnae 9-16 per side, inserted in groups of veredaLa Josefina,800 m, Jun1987(fl, fr), Callejaset al. (1-)2-4 separatedby up to 12 cm, in differentplanes, &Hoyos cuneate to 4238(HUA,NY);3 Dec 1982(femfl), Herndndez broadlycuneate, 1.2-4.5 times as long as 632 (HUA);ibid., 20 Sep 1983 (immfr), Hoyos& Heror briefly incised praemorseat apex, truncate wide, & 27 Dec 1983 333 ndndez (JAUM);ibid., (femfl), Hoyos 832 (JAUM);ibid., 18 Feb 1984(fl), Hoyos& with an up to 7 cm finger-likeprojectionon the distal Herndndez rd.toward margin, glabrous on both sides or with few, minute 891 (JAUM);(femfl) 912, (JAUM); Herndndez hwy.,850 m, 24 spinulesabaxially,marginslined with small spinules; Aquitania,12 km fromMedellin-Bogota Nov 1988 (fem fl), Cogolloet al. 3760 (JAUM);top of PiedraCastrill6n,1700 m, 17 Apr 1971 (fem fl, immfr), basal pinnae 5.5-13 x 0.7-4 cm; middle pinnae Loaiza& Cogollo319 (HUA,JAUM);Mun.de SanCarlos, 8.5-27 x 3-9 cm; apical pinnae 2-4 ribbed,8.5-19.5 la Miranda, x 3.5-11 cm. Inflorescencespicateor rarelybranched Alto Samana,rd. Jardin-Miraflores, quebrada 700-800 m, 24 Oct 1989 (fem fl), Callejaset al. 8535 to 1 order,with to 12 rachillae; up prophyll9-34 x 1 PuebloRico,La Selva,1500m, 27 (COL,HUA).CALDAS: bract to 95 cm cm; peduncular up long, ca. 1 cm Jan1946(fl, fr),Sneidern5385 (S). wide, thin, with a white or brownindument,unarmed Aiphanes parvifolia is readily distinguishedby its or with few yellow spines; peduncle 21-77 cm long; solitary,brown,hardstem; small leaves (rachis33-70 2-3 mm diam. at apex, with a brown-violetindument, cm long); andespecially its extremelyshort,fastigiate unarmedor sparsely armed with yellow, to 15 mm rachillae.There is some variabilityin the shape of the long spines; spike 10-32 cm long, 3-4 mm diam., pinnae, which sometimes are completely linear and covered with short, yellow to violet spinules; regularlyinserted (Callejas 8535, Loiaza & Gogollo branched inflorescences with a 1.5-15 cm long 319, Sneidern5385), but morecommonlycuneateand rachis, bearing4-12 rachillae, 7-28 cm long, 1-1.5 grouped.This variationmightbe relatedto habitat,lin- mm diam., with spinules like the spike; rachillae or ear pinnaeoccurringin palmsgrowingin rockyplaces spike with triads for 1/2-2 of the length, distally or directly on rocks, and cuneate pinnae in palms staminate;triads sunken into pits in the rachillae, growing in clayey soil. A similar variationhas been each subtended by a bract covering the pistillate observedin other species such as A. linearis.Thereis flower before anthesis. Staminateflowers brownish also some variationin the basal rachillae,which are purple outside, becoming nearly white at anthesis, longer, more loosely arranged,and with a longer ster- white inside, with yellow anthers, 2-2.7 mm long; ile basal partin specimenswith linearpinnae. sepals narrowlytriangularto ovate, carinate, 1-1.5 x The circumscriptionabove correspondsto the diag- 1.5-2 mm; petals ovate, acuminate,2.1-2.5 x 1.2-2 nosis by Burret(1932b) based on the lost type, except mm; filaments0.6-1 mm long, anthersoval, 0.4-0.7 in one respect:Burretwrote "ramihaud numerosi,"a x 0.3-0.8 mm; pistillode trifid, distinct, ca. 0.5 mm strangecharacterizationof inflorescencesof A. parvi- high. Pistillateflowers 4-5 mm long; sepals broadly folia, which rathergive the impressionof havingvery ovate, 2-2.5 x 3-4 mm, /2 as long as petals, enclosed many, though very short, rachillae. Nevertheless, in the pit; petals 4-5 x ca. 3 mm, connatefor '/2 their there is no doubt that the neotype designatedhere is length, valvate distally, lobes recurved at anthesis; conspecific with the lost holotype collected some 30 staminodial cup 3-4 mm high, truncate, with 6 km NW of the neotype locality. minute teeth; pistil ca. 4 x 2.5 mm, glabrous. Fruits bright red, globose, strongly rostrate, 8-10 mm ca. 8 mm diam. pitted-grooved. 17. Aiphanes simplex Burret, Notizbl. Bot. Gart. diam.;endocarp Berlin-Dahlem 11: 567. 1932. Type. COLOMBIA. Illustrations. Figs. 11C, 12J (pollen), 15A (stem Antioquia:Rio Verde, 1500-1700 m, 28 Jul 1880, anatomy), 19B (distributionmap), 21L (middle pinKalbreyer 1864 (holotype, B, a single staminate nae); Galeano & Bernal, 1987: 37, fig. 8. flower). Distribution and habitat. Grows throughoutthe Rio Cauca basin in Colombia extending into the m these 2-4 with to 20 stems, tall, Caespitose, up 1-2 cm diam., with scatteredblack spines, to 4 cm upperRio Patia basin. It passes the CordilleraOccilong. Leaves 4-9, erect and arching;sheath 11-29 cm dental only in two places: in the northnear Frontino long, covered with yellow spines, to 4 cm long; (Rio Verde, type locality) and near Cali where the
78 passes are relatively low. Aiphanes simplex is commonly encounteredin patches of primaryor secondary forest, sometimes growing near small streams,at 800-2200 m, most abundantbetween 1600 m and 2100 m, where it may be a dominantcomponentof the shrublayer. Specimens examined. COLOMBIA. ANTIOQUIA:
Mun. de Medellin,5 km E of the city, quebradaSanta Helena, 1800 m, 30 Oct 1980 (imm fr), Galeano et al. 262
(COL,K), 266 (COL);rd.to SantaHelena,km 8-12, 1600 m, 26 Mar 1987 (fer fl), Callejas & Escobar 3280 (HUA,
NY);Fredonia,20 Dec 1920(fl), Toro852 (MEDEL,NY); Palmitas,quebrada Miserengo,1900-2000m, 18 Mar1979 (fl, imm fr), Galeano et al. 14 (COL, HUA). CAUCA:El
Tambo,basinof RioPatia,AltoLosAngeles,1100-1700m, 20 Aug 1949 (fl), Idrobo & Fernandez198 (COL); Palace,
Oct 1945(st), Yepes1070 (COL);Popayan,1700-1800m,
Nov 1889 (fer fl, fr), Lehmann B.T 383 (GH, K, NY).
Bremen,15 km N of Armenia,1950m, 29 Jan QUINDIO: 1989 (fem fl), Gentryet al. 65315 (HUQ,MO);Circasia, El Bosque,1185m, 21 Dec 1989 fincaEl Bosque,quebrada (fr), Velezet al. 817 (HUQ);VeredaBarcelonaAlta,finca BuenosAires,1605m, 7 Apr1991(immfr),Agudeloet al. 1126 (COL,HUQ n.v.);C6rdoba,VeredaEl JardinAlto, 1860 m, 31 Mar 1987 (fl), Arbeldez et al. 2424 (HUQ);
veredaLa Carmelita,1430m, 30 Jul 1987(fl), Quimbaya, Arbeldez 2216 (HUQ). RISARALDA: Pereira,La Suiza, trail to Callejones, 1950 m, 3 Jun 1989 (fl), Franco et al. 2672 (COL). VALLE:Alto Mercedes,2000 m (fl), Dryander1288
Flora Neotropica rachillae have been found in marginal areas of the species distributionalrange, i.e., the upper Rio Patia valley (Idrobo 198) and the western slopes of Cordillera Occidentalin the departmentof Valle (Bernal 1538), indicating a possible transitionto the closely relatedA. erinacea. The argumentsfor keeping the two as separatespecies arethe following: 1) Aiphanes simplexis sufficientlywell knownfor it to be said that in the Rio Cauca basin itself inflorescences are always spicate, and likewise A. erinacea is sufficiently known in Ecuadorand southernColombia for it to be said that spicate inflorescences do not occur there;and 2) individualsof A. simplex with branched inflorescences can still easily be referred to this species by their thin stems and truncatepraemorse, ratherthan incised praemorse,pinnae. Apart from a single atypical collection from the Pacific coast near Buenaventura,there are no collections of A. erinacea furthernorththan the departmentof Nariiio, but this may be due to the poor explorationof the montane forests in the departmentof Cauca.Thus it is possible that the distributionsof A. erinacea and A. simplex form a continuum and that the branched inflorescences sometimes encounteredin A. simplex signify that some exchange of genetic material occurs between the two.
(B);Argelia,veredaLasBrisas,2140 m, 21 Jan1983(fer fl), Francoet al. 1681(COL);Bitaco,1 Apr1959(fl, fr),De Leon126 (BH);Cali,basinof RioPichinde,PeniasBlancas, 1700-1900m, 1 Jul1938(femfl, immfr),Duque-Jaramillo 18. Aiphanes spicata Borchsenius& Bernal,sp. nov. 1320 (COL); 15 Jan 1947 (imm fr), Duque-Jaramillo4531 PERU. San Martin: Rioja (limit with (COL);8 May 1977(fl, fr),Benalcdzar& Silva11 (COL); Type. Amazonas, Bongara), km 150 on rd. from Bagua 1 Oct 1977(fr),Benalcdzar&Silva101 (COL);ibid.,3 km Grandeto Moyobamba,ca. 1800 m, 26 May 1990 N of El 18 on rd.to Pavas,1870 m, 19 Mar1988(fl, fr), Bernal1442 (COL,TULV);El Cairo,rd. La Carbonera- (fl, fem fl, fr), Kahn & Borchsenius2651 (holotype, PerlaRoja,km 3.5, 1950m, 16 Mar1991(fl, fr),Bernal& AAU; isotypes, NY, USM). Borchsenius1964 (AAU, BH, COL, NY, TULV);El Silencio,Yanaconas,1900-2200m, 28 Feb 1939(fer fl), Aiphanesacauli Galeano& Bernal similis sed pinKillip & Garcia 33804 (COL); Mun. de Yumbo, finca La nis aggregatiset floribusflavo virentibusdiffert. Samaria,NE of Darien,nearlago Calima,1700m, 14 Feb 1984(abfr),Juncosa2172(MO,NY);SantaHelena,above Solitary,acaulescentor with a horizontal,subterde CaliNationalPark,1970 Topacio,edgeof LosFarallones m, 11 Dec 1985, Gentryet al. 53102 (NY);Tulua,km 6 ranean stem, to 30 cm long, 2-5 cm diam. Leaves fromLa MoreliatowardMonteloro,1600m, 16 Mar1988 9-12, erect and arching;sheath 14-20 cm long, with (fl), Bernalet al. 1435 (COL,CUVC,FTG,TULV);Baga numerousblack spines, to 8 cm long; petiole 16-25 de la Tulia,1850m, 30 Oct 1944(fr), Cuatrecasas 18377 km 22, 1800m, 13 cm long, armed like sheath, but spines fewer; rachis (BH,COL,F); rd.Cali-Buenaventura, km 42-80 cm long, green, with a brown, caducous induOct 1983,Escobar2673 (HUA);rd.Rio Frio-Salonika ElAlmendronal, 21 Nov 1974(st),Mooreet al. ment,unarmedor with scattered,blackspines, to 5 cm 8, quebrada 10197(BH,COL);upperRio Calima,betweenEl Pitaland pinnae 14-16 per side, nearlyregularlyinserted La Cristalina, 830 m, 22 Feb 1989(fl), Bernalet long; quebrada or in groupsof 2-4, these occupying3-7 cm along the al. 1538 (COL, TULV). rachis, separatedby 9-11 cm, all pinnae more or less Aiphanes simplex is characterizedby its caespitose in one plane, stronglyplicate, middle pinnae 17-22 x habit, with up to 20, to 4 m tall, 1-2 cm thick stems, 1-2.5 cm, linearor narrowlycuneate, 10-18 times as cuneate, truncatepraemorsepinnae,yellow spines on long as wide, obliquelypraemorseat apex, with an up leaf sheaths,and normallyspicate inflorescence.Indi- to 2 cm long finger-likeprojectionon the distal marviduals with branchedinflorescences with up to 12 gin, both sides with scattered,peltate hairs and few
SystematicTreatment minutespinules,marginslined with black spinules,ca. 1 mm long. Inflorescence interfoliar,erect, spicate; prophyll 14-16 cm long, ca. 1 cm wide; peduncular bract 30-50 cm long, 1-1.5 cm wide, thin, with minute, black spinules, soon disintegrating;peduncle 45-110 cm long, 3-4 mm diam. at apex, green,with a thin, brown, caducous indument,unarmedor covered with black, thin, to 3 cm long spines; spike 14-26 cm long, 5-8 mm diam., with a brown, caducous indument, covered with minute, black spinules, basal 2/3 with triads, distally staminate;flower groups sunken into deep, elongate cavities in the spike, each subtended by a 3-4 mm long bract,covering the flower group before anthesis. Staminateflowers 3-4 mm long, 3-4 mm wide, yellow in center,with light green petals; sepals imbricate,carinate,covering the petals for 2/3 of their length, 3-3.5 x 4-5 mm; petals free, valvate, 3.5-4 x 2.5-3 mm; filamentsca. 1 mm long, anthers 1.1-1.4 x 0.9-1.1 mm; pistillode minute, trifid;receptacleswollen, ca. 0.5 mm thick. Pistillate flowers (at anthesis) 4-5 mm long, ca. 4 mm wide, light greenishyellow; sepals imbricate,broadlyovate, shorter than the petals, enclosed in the floral pit, 2.5-3.5 x 4-5 mm; petals connatefor /2 their length, valvatedistally,4-5 x 3-4 mm; staminodialcup truncate, 3-4 mm high; pistil ca. 4 mm high, 3 mm diam., glabrous.Fruits 10 mm long, 9 mm wide.
79 furthermore,the two species occur at different altitudes. The acaulescenthabit and the spicate inflorescences areconsideredderivedcharactersin Aiphanes, and the strikingmorphologicalsimilaritymay be due to convergenceratherthan to a common, widely distributedancestor.
19. Aiphanes tricuspidata Borchsenius, Bernal & Ruiz, Brittonia41: 156, fig. 1. 1989. Type. ECUADOR. El Oro:rd. fromMachalato Naranjal,km 33, 8 km E along dirt rd. leaving from Rio Bonito, 380 m, 19 Nov 1987, Skov, Borchsenius, Blicher Mathiesen, & Bang Klitgaard 64836 (holotype, AAU; isotypes, COL, K, NY, QCA, QCNE).
Solitary.Stem erect or sometimes procumbent,to 4.5 m tall, 2.5-6 cm diam.,light brown,hard,unarmed or with black spines, to 5 cm long, base frequently supportedby a cone of prop roots, up to 15 cm high. Leaves 8-10, erect and arching,lower ones recurving; sheath, petiole and rachis with a brown, caducous indument;sheath 23-37 cm long, armed with brown to yellow spines, to 5 cm long; petiole 8-18 cm long; rachis 74-120 cm long, spinulose, unarmedor with scattered,brown to yellow spines abaxially,to 5 cm long;pinnae 11-14 per side, insertedin groupsof 2 or rarely 3 separatedby 10-20 cm, in differentplanes, Illustrations. Figs. 1D (habit), 5C (leaf), 12H, broadlycuneate,1-3 times as long as wide, darkgreen 13C (pollen), 19C (distributionmap), 21R (middle adaxially,paler abaxially,strongly tricuspidate,with pinnae). an up to 9 cm long finger-likeprojectionon the distal Distribution and habitat. Known only from two margin, adaxial side smooth or rough, abaxial side or almost glabrous,midrib nearby localities in northeasternPeru, in Dictyo- minutely spinulose rarely to with 0-5 black, to 5 cm long yellow caryum lamarckianum-dominatedmontane forest at abaxially x 6-18 1-13 basal cm; middle pinnae spines; pinnae ca. 2000 m. 11-29 x 12-20 cm; apical pinnae 1-3 ribbed, quite Specimens examined. PERU. SANMARTIN:Prov.Rioja variable in size and shape, 8-23 x 6-30 cm. Inflo(limitwithAmazonas,Prov.Bongara):152kmfromBagua rescence interfoliar,erect to drooping,branchedto 1 Grande,ca. 1800 m, 26 May 1990 (fl, fer fl), Kahn& order,sometimeswith a few rachillaebranchedto sec2646 (AAU,COL,K, USM). Borchsenius ond order;prophyll20-56 cm long, 1-3.5 cm wide; from which A. resembles acaulis, peduncularbract 100-170 cm long, 2-2.5 cm wide, Aiphanesspicata it differs in size andcolor of flowers (TableII). Pollen with a brown caducous indument,unarmedor with of the two species is also different:that of A. acaulis some yellow spines, to 1 cm long; peduncle 67-148 has finely reticulate tectum with fusing supratectal cm long, 4-6 mm diam. atjunctionwith rachis,with a clavae (Fig. 11D), whereas that of A. spicata has a brown, scaly, caducous indument,and many yellow reticulate, smooth tectum (Fig. 13C). No other spines, to 1 cm long; rachis 15-55 cm long, unarmed species combines acaulescent habit, linear or nar- or basally armedlike that on the peduncle;rachillae rowly cuneatepinnae,and spicateinflorescences.The 12-52, minutelyspinulose,each subtendedby a small known populations of A. acaulis and A. spicata are bract,ca. 5 mm long; basal rachillae28-50 cm long, separatedby the Andean Cordillera and more than sometimes without flowers for up to 6 cm, with 2000 km, including some of the best-collected areas triadsfor ca. /2of the length, in this part2 mm diam., with respect to palms, in Ecuador and Colombia; distally ca. 1 mm diam., with staminatedyads; apical
80 rachillae 4-8 cm long, staminate; flower groups sunken into the rachillae, each triad subtended by a bract that covers the pistillate and the proximal staminate flowers before anthesis, each dyad subtended by a similar but smaller bract, covering the proximal flower. Staminateflowers 1-2 mm long, purplewith yellow anthers,the proximalof each triad distinctly pedicellate, the remaining more or less sessile; sepals imbricate, 1-1.5 mm long; petals free, valvate, 1.5-2.5 mm long; filaments of very uneven length, 0.2-1.5 mm long, anthers nearly square or slightly longer than wide, 0.5-0.8 x 0.5 mm; pistillode ca. 0.5 mm high. Pistillateflowers 2-4 mm long; sepals broadly imbricate,2.5-3 mm long; petals connate ca. '/2 of the length, valvate distally, 2-4 mm long; staminodialcup 1-2 mm high, truncate, adnate to corolla tube; pistil ca. 2 mm high, glabrous.Fruitsreddishbrown,speckled,globose, ca. 7 x 9 mm (conserved in alcohol), shortly rostrate; endocarp ca. 6 x 7 mm, irregularly grooved and longitudinallyfurrowed. Illustrations. Figs. 19C (distributionmap), 21N (middle pinnae), 30 (drawing); Borchsenius et al., 1989: fig. 1. Distribution and habitat. Found in the Pacific lowlands and the Andean foothills from the department of Valle in Colombia to the province of El Oro in southernEcuador.An understorypalm in primary rain forest, sometimes growing along streams, from sea level up to 650 m. Like other solitary understory species, it appearsto be very sensitive to changes in the environmentand unable to survive in open areas or secondaryforest. Specimens examined. COLOMBIA. CAUCA:Mun. de Guapi, ParqueNacional NaturalIsla de Gorgona, 1841 (fl),
Hindss.n. (K);ibid.,300 m, 7 Sep 1987(fl), Lozano5691, 5916 (COL).NARINO:Barbacoas,Rio Telembi,between andca. 15kmuptheriver,160m, 20 Nov 1986 Barbacoas (fl), Bernal& Hammel1321(AAU,BA,COL,K, MO,NY, La Trojita,downstream fromBajoCalima, PSO).VALLE: 5-50 m, Feb 1944 (fl), Cuatrecasas16492 (F); 20 m, 9 Apr 1976 (fl), Dransfield et al. 4868 (BH, K). UNKNOWN DEPARTMENT:unknown locality,Purdie s.n. (K). km33, 8 ECUADOR.EL ORO:Rd.Machala-Naranjal, km E along dirtrd. leaving from Rio Bonito, 300 m, 14 May 1987 (st), Balslev et al. 62521 (AAU); hills NE of La Avanzada on rd. towardLa Pifia, 500 m, 18 Nov 1987 (imm fr), Skov, Borchsenius, et al. 64829 (AAU). PICHINCHA: ENDESA forest reserve, km 113 on rd. Quito-Pto. Quito, km 10 N of rd., 650 m, 17 Nov 1989 (fl, fr), Borchsenius& Luteyn 91424 (AAU, BH, COL, FTG, K, MO, NY, QCA, QCNE, SEL); ibid., km 127, 2 km N of rd., 500 m, Sep 1987 (juv), Skov,Borchsenius,et al. 64706 (AAU).
Flora Neotropica Aiphanes tricuspidatais distinguishedby its solitary habit, often yellow spines on leaf sheath and rachis, broadlycuneate, tricuspidatepinnae, inserted in widely spaced pairs or triplets,and priorto anthesis almost completely hidden flower groups. In its habit and in its grouped,dark green, gently undulate pinnae, it resembles A. parvifolia, which differs in having obliquely to lobulate praemorsepinnae, and very short,fastigiaterachillae.
20. Aiphanes ulei (Dammer) Burret, Notizbl. Bot. Gart.Berlin-Dahlem11: 568. 1932. Martineziaulei Dammer,Verh. Bot. Vereins. Prov. 48: 127. 1907[nonM. uleiDammer(1915), Brandenburg nom.illeg.].Type.PERU.SanMartin:Cerrode Escalero, 1200-1300m, Nov 1902 and Jan 1903, Ule 6880 (Bt). PERU.Loreto:Prov.Requena, Neotype(heredesignated). lowerRio Ucayalivalley,nearGenaroHerrera,Kahn& K). Mejfa1916(holoneotype, AiphanesschultzeanaBurret,Notizbl.Bot. Gart.BerlinDahlem15: 36. 1940.Type.ECUADOR.Pastaza:Mera, primaryswampygalleryforestalongthePastaza,1000m, 3 Sep 1939,Schultze-Rhonhof2769 (holotype,B, sterile). Solitary.Stem 0-6 m tall, 2.5-5 cm diam., armed with black spines, to 7 cm long. Leaves 6-15, erect and arching;sheath 20-35 cm long, densely armed with black spines; petiole 5-70 cm long, with a thin, darkbrown,caducousindument,armedas the sheath, less toward apex; rachis 60-180 cm long, with an indument like that on petiole, black spinules, and scatteredblack spines, to 5 cm long; pinnae 9-14 per side, inserted in pairs or rarely triplets separatedby 10-20 cm, broadlycuneate, 1-3(-4) times as long as wide, sometimes strongly plicate along secondary veins, incised to bilobulate praemorseat apex, symmetricalaroundmidrib,distal marginecaudate,adaxial side dark green and glabrous, abaxial side pale greyish to silverish green, glabrousor minutely spinulose; basal pinnae 11-20 x 2-9 cm; middle pinnae 14-34 x 8-19 cm; apical pinnae 2-5 ribbed,22-25 x 5-26 cm. Inflorescence interfoliar, erect, normally exserted above crown, all parts with a thin, dark brown, scaly indument;prophyll 21-32 x 1-3 cm; peduncularbract65-100 cm long, unarmed,soon disintegrating; peduncle 50-100 cm long, 5-10 mm diam. at junction with rachis, densely armed with black or brown spines, to 1(-3) cm long, rarely almost unarmed; rachis 20-40 cm long, unarmed except at base; rachillae 30-40, strongly appressed to rachis, with densely packed flower groups abaxially, without flowers adaxially,covered with minute
81
Systematic Treatment
~I ,
I.:j
2m~1-:t 1.1
5
FIG. 30. Aiphanes tricuspidata.A. Habit.B. Middle pinnae. C. Basal and distal portionsof a rachillabefore anthesis. D. Detail of proximal portion of of rachilla showing flower arrangement;in the lower triad the subtending bract has been removedto show the flowers. E. Longitudinalsection throughtriad.F. Staminateflower before anthesis,opened to show the stamens. G. Basal portion of rachilla with developing fruits. Drawn from holotype, Skov et al 64836. Reproducedfrom Borchseniuset al., Brittonia41(2).
82 spinules; basal rachillae 10-15 cm long, adnate to rachis for 3-5 cm or sometimes all their length, with flowers from base, the proximal2/3with triads,in this part thickened and flattened, 6-7 mm wide, the remainingpartmore slender, 1-3 mm wide, with staminatedyads or singles; apical rachillae 1-4 cm long, staminate,briefly adnateto rachis, somewhatspreading, distally with flowers on all sides; flower groups sunken into pits, subtendedby fused bracts,forming a rim around the pit. Staminateflowers white to yellow, ca. 1 mm long; sepals cap-shaped,carinate,in bud almost completely enclosing the petals, 1-1.5 mm long; petals almost free, valvate, 1-1.5 mm long; filaments0.5-1 mm long, anthersnearly square,0.50.6 x 0.5-0.7 mm; pistillode minute, sunkeninto the swollen receptacle. Pistillate flowers 2-3 mm long; sepals imbricate, ca. 2 mm long; petals connate /2, valvatedistally, 2-2.5 mm long; staminodialcup ca. 1 mm high, nearly truncate; pistil 1.5-2 mm high, glabrous. Fruits globose, strongly rostrate,ca. 7 mm in diam., rostrum1-2 mm long.
Flora Neotropica
Anananch RBAE223 (NY); trail Taisha-Cangaime, 1 hr. walkW of Taisha,400 m, 13 May 1986 (fl, fr), Balslev et al. 62200 (AAU, NY); same trail, 30 km SE of Rio Pastaza, 400-600 m, 22 Mar 1986, Baker 6805 (NY); rd. MendezMorona, km 3 past Santiago, 450 m, 27 Nov 1989 (st), Borchsenius & Pedersen 91435 (AAU, QCA, QCNE). NAPO: Estacion ExperimentalINIAP-Napo, 5 km N of Coca, 250 m, 3 Jul 1986 (st), Palacios et al. 1174 (AAU); same region, Reserva FaunisticaEl Chuncho, 6 Oct 1987 (st), Cerdn & Palacios 2381 (AAU); rd. Coca-Loreto, km 2-10, Rio Payamino, 250 m, 14 Dec 1987 (st), Cer6n & Palacios 3032 (MO);Aiiangu,4 hr.downstreamfrom Coca, S bank of Rio Napo, 300 m, 25 Jun 1985 (infl), Skov & Pedersen 60601 (AAU); Aug 1985 (fem fl), Balslev et al. 60733 (AAU); (juv), 60581, 60742 (AAU); 11 Apr 1986 (infl), Balslev et al. 62045 (AAU); 4 Nov 1987 (fl, fer fl), Skov,Borchsenius,et al. 64787 (AAU); laguna Pafiacocha, 200 m, 19 Apr 1987 (infl), Balslev 62473 (AAU); same region, 2-3 km NE of village, 300 m, 6 Nov 1987 (infl), Skov, Borchsenius, et al. 64795 (AAU); Yasuni National Park, oil well Daimi, 200 m, 1 Jun 1988 (fl), Cer6n & Hurtado4268 (MO); same region, Daimi II, 29 May 1988 (st), Bergmann& Legaard 67217 (AAU); 30 May 1988 (fl), Bergmann& Lcegaard67220 (AAU, QCA); between Tena and Archidona,400 m, 9 Oct 1939 (fl, imm fr), Asplund 9174 (S); Misahualli,2-3 km N of the village, 3 Apr 1987 (st), Balslev 62480 (AAU); Reserva Biol6gica JatunSacha, Illustrations. Figs. 1C (habit),5B (leaf), 6C (inflo- km 8 from Misahualli, 500 m, 18 Dec 1986 (imm fr), rescence), 11A, 12G (pollen), 19A (distributionmap), Hammel 15984 (MO); 8 Nov 1987, Cer6n 2621 (MO); 30 Apr 1991 (fl), Ponce 108 (QCA). PASTAZA:S bank of Rio 21S (middle pinnae). Pastaza,opposite Mera, ca. 2 km E of bridge crossing the Local names. Jimenatfue ("chontadurode nutria"; river,1300 m, 25 Oct 1987 (infl), Skov& Borchsenius64776 (AAU); rd. from Mera-MadreTierra,km 5, 1150 m, 27 Oct Colombia, Witoto); pa m si mo ha (Ecuador,Kofan); 1987 (infl), Skov & Borchsenius64780 (AAU); rd. Puyochontilla (Ecuador). Arajuno,about 10 km E of Diez de Agosto, 1200 m, 28 Sep 1983, Balslev & Balslev 4418 (NY); same rd., 21 km NE of Distribution and habitat. Found in the upper Puyo-Macasrd.,3/4hr. walk fromEl Triunfo,1250 m, 24 Oct Amazon and on the eastern Andean slopes from 1987 (st), Skov & Borchsenius64771 (AAU). SUCUMBIOS: southernColombia to Peru, extending into the west- Durenoon Rio Aguarico,300 m, 10 May 1966 (infl), Pinkernmost parts of Brazil, reaching 1850 m altitudein ley 486 (BH, ECON); 30 Dec 1987 (infl), Cerdn & Cer6n 3116 (MO); ReservaFaunistica LagunaGrande, central Ecuador, and ca. 1000 m in northernPeru. 300 m, 27 Apr 1986 (juv), BalslevCuyabeno, et al. 62067 (AAU); 9 Dec Often common, though never abundant,in primary 1987 (infl), Blicher-Mathiesen& Bang-Klitgaard 62606 rain forest, sometimes also in patches of disturbedor (AAU); 19 Apr 1988 (infl), Balslev 69053 (AAU). TUNGUrd. Bafios-Mera,km 35, 1450-1550 m, 4 Sep 1976 even secondary forest. In Ecuador the species is RAHUA: (infl), 0llgaard & Balslev 9306 (NY). found both on terrafirme and in temporarilyflooded PERU. LORETO:Prov. Maynas,Yanomamo,Explorama forest. The species is one of the most widely dis- TouristCampon Rio Amazonasbetween Indianaand mouth of Rio Napo, 120 m, 27 Jul 1980 (st), Gentryet al. 29148 tributedand most frequentlycollected in the genus. (MO); Prov.Requena,GenaroHerrera,180 m, 8 Nov 1982, Specimensexamined.COLOMBIA.AMAZONAS: Que- Mejfa 125 (USM); same region, km 15 along the stream brada Perico, an affluent of Rio Igara-Parani,25 km S of "Copal"(fl), Kahn & Mejia 2060 (USM). SAN MARTfN: km 20, Cerro Los Monos on Rio Caqueta, 300 m, 19 Aug 1988 (st), Prov. San Martin,rd. Tarapoto-Yurimaguas, Galeano et al. 1494 (COL);20 Aug 1988 (fem fl), Galeano de Escalero, 980 m, 26 May 1960 (topotype, st), Moore et & Angulo 1510 (COL). CAQUETA: Morelia, 350 m, 30 Oct al. 8529 (BH, USM). BRAZIL. ACRE:Mun. Mancio Lima, upper Rio Moa 1941 (infl), Sneidern 1260 (S); 15 km N of San Antonio Getucha, 350 m, 1 Feb 1988 (st), Bernal & Galeano 1406 near IgapareVitor, 14 Oct 1989 (fr), Hendersonet al. 1133 Mun.Atalaiado Norte, Rio Javary,fron(NY). AMAZONAS: (COL). ECUADOR. MORONA-SANTIAGO: Rd. Macas-Alshi (9 tier between Brazil and Peru, Estiraodo Ecuador,150 m, 8 de Octubre),km 8-9 from GeneralProano, 1400 m, 15 Jul Jan 1989 (st), Hendersonet al. 846 (NY); 11 Jan 1989 (fl), 1985 (fl, imm fr), Balslev & Henderson60658 (AAU); rd. Hendersonet al. 868 (NY). Plan de Milagro-Gualaceo, km 2, 1850 m, 26 Sep 1987 Aiphanes ulei is characteristic in its solitary, often (infl), Skov,Borchsenius,et al. 64715 (AAU); near Taisha, vicinity of CentroTuutinEntsa, 330 m, 18 Sep 1985 (infl), acaulescent habit; cuneate, paired pinnae, greyish to
SystematicTreatment
83
50-65, minutelyspinulose,each subtendedby a light green bract,small or up to 4 cm long; basal rachillae to 22 cm long, with flowersnearlyfromthe base, basal 4-6 cm thickenedwith up to 8 triads, the remaining partslender,ca. 1 mm diam., with dyads of staminate flowers; apical rachillae to 10 cm long, staminate; triadsslightly sunken,each subtendedby an up to 8 mm long bractthatcoversthe pistillateflower for /2 its lengthor more;dyads superficial,each subtendedby a small bract, ca. 1 mm long. Staminate flowers creamishyellow, 3-5 mm long, those of triadsborne on a 6-8 mm long pedicel partlyadnateto the rachilla, those of dyads shortlypedicellate;sepals narrowlytriangular,distinct or slightly imbricate,covering /2 of the petalsor less, 1.5-3 mm long; petals free, valvate, 2.2-4 mm long; filaments ca. 1.5 mm long, anthers linear, 1.4-2.5 x 0.5-0.8 mm; pistillode distinct, ca. 0.5 mm high, trifid.Pistillateflowers 9-11 mm long, each subtendedby an up to 5 mm long bracteole; sepals ovate, imbricate,5-9 mm long; petals connate 21. Aiphanes verrucosa Borchsenius & Balslev, for 2-2/3 of their length, valvate distally, 9-11 mm Nordic. J. Bot. 9: 389, fig. 4. 1989. Type. Ecuador. long; staminodial cup 4-5 mm high, distinctly Zamora-Chinchipe:Rd. Loja-Zumba, 10 km N of toothed, adnateto corolla tube; pistil ca. 6 mm high, Valladolid, 2450 m, 16 May 1987 (fl, fr), Balslev, glabrous.Fruitgreenishwhite at maturity,globose, ca. Bergmann, Bernal & Galeano 62531 (holotype, 30 mm diam., first smooth, soon corky-verrucoseat AAU; isotypes, COL, K, NY, QCA, QCNE). apex;mesocarpdry,2-3 mm thick,crackingopen and Caespitose, with up to 6 stems and several basal persistenton the rachillaewhile the endocarpdrops; suckers.Stems 2-5 (-8) m tall, 4-5 cm diam.,fiercely endocarpca. 25 mm diam., shallowlypittedapically. armed with black spines, to 8 cm long. Leaves 3-5, Illustrations. Figs. 4B (spines), 9C (endocarp), normally4, distichous,erect and arching;sheath,peti- 16A,B (leaf anatomy), 19D (distributionmap), 31 ole, and rachis with a thick, white, caducous (drawing);Borchseniusand Balslev, 1989: fig. 4. indument;sheath40-50 cm long, covered with black Distribution and habitat. Endemicto a small area spines, to 10 cm long; petiole ca. 6 cm long; rachis 85-115 cm long, with numerousblack spines, to 5 cm of the Andes in southernEcuador,in primaryor dislong; pinnae 58-70 per side, inserted in groups of turbed montane forest, between 2200 and 2800 m. 4-11 separatedby 5-8 cm, in several planes, lanceo- The montane forests in these areas are subject to late, praemorseat apex, distal margin 1-2 cm longer burningand felling for firewood,and the palm poputhan the proximal, both sides with black spinules, lation at the type locality, currentlythe only known midrib adaxially with a row of thin spines, 2-7 mm population of A. verrucosa, will probably become long, abaxially with 0-4 rigid spines, to 4 cm long, extinct in the nearfuture.So far it has not been found marginswith a row of spinules, 2-4 mm long; basal inside the nearbyParqueNacionalde Podocarpus,but pinnae 19-31 x 0.5-1 cm; middle pinnae 35-40 x it likely occurs also within the park and will thus 1.5-2.5 cm; apical pinnae 2-7 ribbed, 11-14 x 1-5 probablybe preserved. cm. Inflorescenceerect at anthesis,curvedin fruiting Specimens examined. ECUADOR. ZAMORA-CHINstage, branchedto I order;prophyll,peduncularbract, CHIPE:Alongquebrada Achupallas,2500-2800m, 9 Oct and peduncle with a thick, white indument;prophyll 1943 (fl), Steyermark 54535 (F); Rd. Yangana-Valladolid 2840 cm long; peduncularbract 70-110 cm long, km 28, 2520 m, 24 Jan 1987 (imm fr), Barjfd et al. 60179 et al. 64734 inserted 2-5 cm above prophyll,light green in vivo, (AAU);30 Sep 1987(fl, fr),Skov,Borchsenius, (AAU,QCA). thick, with some spines, to 3 cm long; peduncle60-90 cm long, 0.5-1 cm diam. at junction with rachis, Aiphanes verrucosa is distinguished by its disarmed with numerous brown spines, 2-6 cm long; tichous leaves; high number(58-70) of densely clusrachis 20-26 cm long, minutely spinulose; rachillae tered, lanceolate pinnae; creamish yellow staminate silverish green on the abaxial side; and especially inflorescences with strongly appressedrachillaethat bear densely packed flower groups abaxially and no flowers adaxially. The only other species with such inflorescences is A. gelatinosa, from southwestern Colombia and adjacentEcuador,which differsin having much larger, often multiple inflorescences; and linear to narrowlycuneate pinnae, regularlyinserted or in lax groups of 2-4. The original collection of A. ulei (Ule 116b) from Cerrode Escalero nearTarapotoin the departmentof San Martin,Peru, is lost, and no isotypes exist. And unfortunately,the only existing collection from the type locality (Moore 8529) is sterile. As neotype we have insteadchosen a specimen from GenaroHerrera on the Rio Ucayali in the departmentof Loreto(Kahn & Mejia 1916), some 250 km east northeastof the type locality.
84
Flora Neotropica
10 cm
1ig. E-F
10mm
10mm
VIN
/
' M
I meter
A
"
'
.
B
F
FIG. 31. Aiphanes verrucosa. A. Habit. B. Infructescencewith prophylland peduncularbractattached.C. Basal partof rachilla at staminate anthesis. D. Basal portion of rachilla at pistillate anthesis. E. Staminateflower. F. Pistillate flower. G. Fruitwith cracking, verrucosemesocarpdisplayingendocarp.Drawnfrom holotype, Balslev et al. 62531.
flowers with linear, 1.4-2.5 mm long anthers;9-11 mm long pistillate flowers with glabrous pistil; and white fruits, ca. 30 mm diam., which soon become corky-verrucoseat apex. It is closely related to A. lindeniana distributedalong CordilleraOrientaland Centralin Colombia, and probablyrepresenta geographicallyisolated form of that species. 22. Aiphanes weberbaueri Burret, Notizbl. Bot. Gart. Berlin-Dahlem 11: 565. 1932. Type. Peru.
Huanuco: Rio Pozuzo, 1700 m, 20 Jul 1913 (fl), Weberbauer 6775 (holotype, Bt; lectotype, here designated, F; isotypes, GH, MOL-n.v., S). Aiphanes tessmannii Burret, Notizbl. Bot. Gart. BerlinDahlem 11: 564. 1932. Type. Peru.Amazonas:Confluence of Rio Santiago and Rio Marafion,160 m, 13 Oct 1924, Tessmann4281 (Bt). Neotype (here designated). PERU. Amazonas:Confluenceof Rio Santiago and Rio Marafion, 10 min. upstreamRio Santiago, W bank, 160 m, 20 May 1990 (st), Kahn & Borchsenius2546 (holoneotype, AAU; isoneotypes, NY, USM)
85
SystematicTreatment Solitary, rarely with a few suckers at base. Stem 0-1.5 m tall, 3.5-6 cm diam., armedwith blackspines on the internodes,to 5 cm long. Leaves 5-13, spreading; sheath 1542 cm long, armedwith black spines, to 5 cm long; petiole 10-46 cm long, green, armed like sheath,but spines fewer; rachis 55-120 cm long, green, almost glabrous to densely yellow or black spinulose, armedwith scattered,black spines, to 4 cm long; pinnae 6-24 per side, insertedin groupsof 2-3 separatedby spaces of up to 13 cm, often subregularly insertedon the distal half of the leaf, sometimes regularly insertedthroughout,more or less in one plane, linear to cuneate, 2.5-15 times as long as wide, oblique or incised praemorseat apex, with an up to 4 cm long finger-like projectionon the distal margin, both sides nearly glabrous to densely covered with yellow or black spinules, up to 5 mm long; basal pinnae 7-21 x 0.5-4 cm; middle pinnae 12-30 x 2-7 cm; apical pinnae3-7 ribbed,8-28 x 5-20 cm. Inflorescence interfoliar,erect to curving, branchedto 1 order; prophyll 15-33 cm long, 0.5-2.5 cm wide; peduncularbract 60-105 cm long, 1.5-2.5 cm wide, unarmed or spinulose, thin, soon disintegrating; peduncle 30-115 cm long, 2-7 mm diam. at junction with rachis,covered with thin, <1 cm long spines and spinules, these shorterdistally; rachis 6-30(-60) cm long, densely covered with short,<1 mm long, yellow to brown spinules; rachillae 5-34, often inserted at relativelylargeintervals,with spinuleslike the rachis; basal rachillae8-35 cm long, sometimeswith an up to 8 cm long basal sterile portion, the fertile part with triads for ca. /2 of the length, in this part 2-5 mm diam., often distinctly thickened, especially in fruit, distal half 1-2 mm diam., staminate;apical rachillae 2-17 cm long, staminate;flower groups inserted in shallow depressionsin the rachillae,pistillateflowers sometimes sunkenfor up to /2of their length.Staminateflowers purple, 1-2 x 1-2.5 mm; sepals narrowly triangularto broadly ovate, arched, carinate,nearly enclosing the petals in bud, 1-1.5 x 0.5-1.5 mm; petals ovate-acute, briefly connate at base, 1.3-3 x 1.5-2.5 mm; filaments0.3-0.5 mm long, anthersoval, 0.3-0.9 x 0.4-0.9 mm; pistillode sunken into the 0.4-0.7 mm thick receptacle. Pistillate flowers 2-4 mm long, 2.5-6 mm wide; sepals broadly ovate, nearly as long as the petals, imbricate,2.5-3 x 4-5 mm; petals connate for /2-2A of their length, valvate distally, 3.5-4.5 x 3-4 mm, valves rounded,recurved at anthesis; staminodialcup 2.5-3.5 mm tall, nearly truncate;pistil ca. 3 mm high, glabrous.Fruitsred to purple, globose to slightly elongate 7-10 mm diam.; endocarp6-9 mm diam., shallowly pitted.
Distribution and habitat. EasternAndean slopes from southernEcuadorto southernPeru up to 1950 m, extending into the Amazonianlowlands in northern Peru as far as Iquitos. At the confluence of Rio Santiagoand Rio Marafionthe species grows on lateritic soil, aroundIquitos and Rio Nanay it is found on white sand, but morphologicaldifferences correlated with these differencesin habitatare not evident. Illustrations. Figs. 11B, 12E, F (pollen), 18C (distributionmap),21T-V (middlepinnae),32 (drawing). Local name. Chontilla(Ecuador). examined. ECUADORMORONA-SANTIAGO: Specimens Plan de Milagro-Gualaceord., km 2, 1850 m, 26 Sep 1987 (st), Skov,Borchsenius,et al. 64714 (AAU); 26 Nov 1989 (fl, imm fr), Borchsenius & Pedersen 91427 (AAU, COL, K,
NearElTriunfoon rd.to MO,NY,QCA,QCNE).PASTAZA: Arajuno,21 km NE of Puyo-Macasrd., 1250 m, 24 Oct 1987, Skov& Borchsenius64770 (AAU, COL, QCA, USM).
PERU.Amazonas:Prov.Bagua,ca. 12-18trailkmE of La Pecain Serrania de Bagua,1800-1950m, 14 Jun1978 (fl), Gentry et al. 22935 (BH, F, MO); confluence of Rio
RioSantiago, SantiagoandRioMarafon,10 min.upstream Wbank,160m, 20 May1990(p fl, immfr),Kahn&Borchsenius2535 (AAU,USM);ibid.,in frontof militarycamp "Pinglo," 21 May 1990 (fl), Kahn & Borchsenius 2554
Prov.Chanchamay, (AAU,NY,USM).JUNfN: Chilpez,ca. 26 kmS of SanRamon,1720-1850m, 19 Oct 1982(fl, fr), Smith & Palacios 2643 (MO-n.v.,
USM); Schunke
aboveSanRam6n,1400-1700m, 8 Jun1929(fl), hacienda, Killip & Smith 24608 (F, NY). LORETO:Prov. Maynas,
forest,22 Nov 1940(imm vicinityof Iquitos,Quistococha, fr), Asplund 14667 (S); 180 m, 9 Sep 1957 (st), Ellenberg
2877 (U); rd. Iquitos-Quisto Cocha,km 8-9, 100 m, 15 May 1960 (st), Mooreet al. 8456 (BH, USM); Prov. Mishana,Rio NanayhalfwaybetweenIquitosand Santa Mariade Nanay,nearCampamento 1, 20 Mar1982(fl, fr), near Gentryet al. 36514(MO);vicinityof LagoLlanchama Rio Nanay,2 Aug 1972 (st), Croat18711 (MO).PASCO: Prov.Oxapampa, Paleazu,RioAltoIscozacin,Ozuzto Rio Pescado,400-500m, 12 May1985(fl), Foster& d'Achille 10101(F n.v.,USM).SANMARTIN: Prov.SanMartin,rd. km20, Cerrode Escalero,980 m, 26 Tarapoto-Yurimaguas, May1960(infl),Mooreet al. 8527(BH,USM). Aiphanesweberbaueriis characterizedby its solitary, often more or less acaulescent habit, inflorescences with few (5-34), often thick rachillae,and pistillate flowers with completely reflexed, rounded petals and truncatestaminodialcup, lending a very characteristicappearanceto inflorescences with pistillate flowers in or past anthesis.It is closely related to A. deltoidea; the differences are discussed under that species. Aiphanes weberbaueri is a complex and incom-
pletely knownspecies. In the Andes, variationappears to be limited. Plants from this area have linear to
86
Flora Neotropica
A
50cm
Imm
fig.
FIG. 32. Aiphanes weberbaueri. A. Habit. B. Leaf. C. Inflorescence with peduncular bract. D. Triads. E. Staminate dyads. F. Staminate flower, one petal removed. G. Developing fruit. Drawn from Borchsenius & Pedersen 91427.
Doubtful Names and ExcludedTaxa
87
disticha Linden, nomen nudum.Burretdid not see narrowlycuneate, grouped to regularlyinsertedpinthe originaldescriptionM. disticha in Linden'scatnae, and inflorescences with relatively few (13-30), often very thick rachillae.In the Amazon the species alogue, but based his discussion on the crude illustrationaccompanyingMartineziadistichaWallis ex. is more variable, and two forms can be recognized: one with linear,regularlyinsertedpinnaeandinfloresRegel. cences with ca. 30 slender rachillae (A. tessmannii Aiphanes gracilis Burret,Notizbl. Bot. Gart. BerlinDahlem 11: 566. 1932. Type. Peru. Loreto: MounBurret);and one with cuneatepinnaeinsertedin pairs or triplets,and inflorescenceswith 5-10 rachillaethat tains N of Moyobamba, 1300-1400 m, 28 Aug are thickenedin the androgynouspart.The type of A. 1904, Weberbauer4665 (holotype,Bt). The type of A. gracilis Burretis destroyedand no isotypes exist. tessmanniiis destroyed,and no isotypes exist. When the type locality was visited in May 1990, only one The type locality at Moyobambain Peruwas visited in June 1990, but withoutresult.The areais heavily sterile individual was found and collected (Kahn & Borchsenius2546, neotype).This plantwas very condeforesteddue to small-scaleagricultureanduncontrolledburningof the forest. Primaryforest northof spicuous in its regularly pinnate, long-spinulose leaves, and appearedto be clearly distinct from the Moyobamba,at ca. 1500 m, two hours walk from of with cuneate individuals the form Yantalo many pinnae unfortunatelydid not reveal any Aiphanes. growing at the same locality.Plantswith subregularly In the notes to A. gracilis Burretstated that it was or regularlyinserted,long-spinulosepinnaeof similar very similarto A. weberbaueri,but differedin havsize and shape are, however,known from the Andes ing smallerleaves, plicate,morerigid,less spinulose (Borchsenius & Pedersen 94427, Moore 8527), and pinnae, and rachillae with fewer, smaller spinules. the latter specimen includes an inflorescence correFurther,A. gracilis had apical pinnaesimilarin size and shapeto the middlepinnae,whereasthose in A. sponding well to Burret'sdescriptionof the inflorescence of A. tessmannii.Thus, in reality,it is impossiweberbaueriwere broaderthan these. The differble to separateA. tessmanniifrom A. weberbaueri,at ences discussed by Burret seem insignificant, and least with our presentknowledge. The mattershould apartfrom the inflorescence rachis being up to 60 be reinvestigated when more informationbecomes cm long, A. gracilis as knownfromBurret'sdescripavailable. tion falls within the circumscriptionof A. weberThe existing materialfrom Peru is insufficient to baueri in the presenttreatment. resolve the taxonomy of A. weberbauerisensu lato in Aiphanes leiospatha Burret, Notizbl. Bot. Gart. a satisfactory manner.The variation patternresemBerlin-Dahlem11: 571. 1932. Type. Colombia.Anbles that found in A. hirsuta, which is still incomtioquia: Kalbreyer 1607 (holotype, Bt). Aiphanes fact that there exist understood the leiospathawas basedon an inflorescenceassociated despite pletely with the leaf of a Geonomoidpalm. Most likely this more than twice as many,and more complete, collecwas the misplacedinflorescenceof A. monostachys tions of that species. Both show the same transition from linear, regularly inserted pinnae to cuneate, Burret (here treated as a synonym of A. hirsuta subsp. hirsuta), but this cannot be provenwith cergrouped ones, variationin indumentand apex shape of the pinnae, and variationin number,length, and tainty. See discussion in the notes to A. hirsuta of of the rachillae. both Also, degree thickening subsp. hirsuta. species seem to occur in distinct sympatricforms in Aiphanes praga Humboldt, Bonpland, and Kunth, some areas, whereasdifferencesbreakdown in other. Nov. gen. et sp. 1: 303. 1816. [= Euterpe praga (H.B.K.) Sprengel]. Martinezia antiochensis Linden, l'Illustr. Hort. 32. 1881, nomen nudum. DOUBTFUL NAMES AND disticha Linden, Catalogue no. 93: 32. Martinezia EXCLUDED TAXA 1875, nomen nudum.Lindenprovideda drawingof a juvenile plant (with polystichous leaves) but no Aiphanes caryotoides Blatter, J. Bombay Nat. Hist. Soc. 24: 678. 1916. Spelling variantof Aiphanes descriptionor diagnosis. It is likely that the name Martineziadistichareferredto Aiphaneslindeniana, caryotifolia (Humboldt, Bonpland, & Kunth) H. which occurs at high elevations in the Colombian Wendland(= Aiphanes aculeata Willdenow). CordillerasCentraland Oriental,but this cannot be Aiphanesdisticha (Linden)Burret,Notizbl. Bot. Gart. concluded with certainty. Specimens collected in Berlin-Dahlem11: 575. 1932. Based on Martinezia
88
FloraNeotropica
Horto Lindeniano and annotated Martinezia disticha are Aiphanes aculeata. Martinezia disticha Wallis ex Regel, Gartenflora28: 163, tab. 977, fig. 1. 1879. Homonymof Martinezia disticha Linden (1875). The crude illustration shows a caespitose palm with seven entire, distichous leaves, most likely Aiphanes lindeniana. No descriptionwas provided. Martineziagranatensis H. Wendlandin Kerchovede Denterghem,Les Palmiers. 1878, nomen nudum. Martinezia leucophceusHort., GardenersChronicle (May 8): 589. 1875, nomen nudum. Martinezia minor Linden, l'Illustr. Hort. 32. 1881, nomen nudum. MartineziaRoezlii Hort., GardenersChronicle (June 3): 735. 1876, nomen nudum. Martineziaulei Dammer,Notizbl. Konigl. Bot. Gart. Berlin-Dahlem59:266. 1915.Homonym[=Aiphanes ernestii (Burret)Burret].The name Martineziaulei had been publishedby Dammeralreadyin 1907 [= Aiphanesulei (Dammer)Burret]. Tilmiadisticha (Linden)Cook, Bull. TorreyBot. Club. 28: 565. 1901. See Martineziadisticha Linden.
ACKNOWLEDGMENTS The presentstudyis partof a Ph.D. thesis submitted to the Facultyof NaturalSciences, AarhusUniversity, Denmark.The Faculty of NaturalScience supported the projectwith a stipendto Finn Borchsenius. We are gratefulfor economic supportfrom Fiedlers Legat (Denmark), and the Danish Natural Science Council (grant no. 116848). Fieldwork in Ecuador was made possible through the collaboration with Pontificia UniversidadCatolica del Ecuador,Quito, and we are grateful to the director of the Biology Departmenttherefor providingworkingfacilities.The National Park Service of Ministeriode Agriculturay Ganaderiakindly helped with researchpermitswhich made collection of specimens in Ecuador possible. Fieldwork in Peru was carried out in collaboration with Francis Kahn, ORSTOMmission in Lima, who organized and financed transportation;fieldwork in Bolivia was made in collaboration with Monica Moraes,HerbarioNacionalde Bolivia, La Paz. We are very gratefulfor this help. We thankthe curatorsof the following herbariafor puttingmaterialat our disposal:A, AAU, B, BH, BM, BR, C, CAS, COL, ECON, F, FI, FTG, G, GH, HUA,
HUQ, JAUM, K, L, LE, M, MEDEL, MICH, MO, NY, 0, OXF,P, PH, QCA, QCNE, S, SEL, TULV,U, US, USM, VEN, W, WU. Anni Sloth helped us in the laboratory.Line drawings were made by KirstenTind. Many other people helped us in variousways, and we wish to thankBirgitte Bergmann, Sean Carrington,Gloria Galeano, Andrew Henderson,Peter M0ller J0rgensen,Francis Kahn, Jim Luteyn, Monica Moraes, HenrikBorgtoft Pedersen, and Susanne Renner. Special thanks goes to HenrikBalslev for his advice and supportthroughout this study.
LITERATURECITED Anderson, A. B., W. L. Overall & A. Henderson. 1988. Pollinationecology of a forest-dominantpalm (Orbignya phalerata Mart.) in northern Brazil. Biotropica 20: 192-205. Bailey, L. H. 1943a. New palms in Panama and others. Gentes Herb.6: 198-264. . 1943b. Flora of Panama (Palmaceae). Ann. Missouri Bot. Gard.30: 327-403. - . 1949. Palms uncertainand new. Gentes Herb. 8: 146-205. Balick, M. J. & S. N. Gershoff. 1990. A nutritionalstudy of Aiphanes caryotifolia (Kunth)Wendl. (Palmae) fruit: an exceptionalsource of vitaminA and high quality protein from tropicalAmerica.Econ. Bot. 8: 35-40. Balslev, H. & A. Henderson. 1987. The identity of Ynesa colenda (Palmae).Brittonia39: 1-6. & M. Moraes. 1989. Sinopsis de las palmeras de Bolivia. AAU Rep. 20: 1-107. Barfod, A. S. 1991. A monographicstudy of the subfamily Phytelephantoideae(Arecaceae). Opera Bot. 105: 1-73. Beach, J. H. 1984. The reproductivebiology of the Peach or "Pejibay6"palm (Bactris gasipaes) and a wild congener (B. porschiana) in the Atlantic lowlands of Costa Rica. Principes28: 107-119. Bernal, R. 1986. AiphanesmacrolobaandAiphanesmonostachys (Palmae).Brittonia38: 65-70. . 1989. Endangerment of Colombian palms. Principes33: 113-128. --, G. Galeano & A. Henderson. 1989. Neotypification of Colombian palms collected by W. Kalbreyer. Taxon38: 98-107. Blomberry, A. & T. Rodd. 1982. Palms. An informative, practicalguide to palms of the world: their cultivation, care, and landscapeuse. Angus and Robertson,London. Borchsenius, F 1993. Floweringbiology and insect visitation of three EcuadoreanAiphanes spp. Principes, in press. & H. Balslev. 1989. Threenew species of Aiphanes (Palmae) with notes on the genus in Ecuador.Nordic J. Bot. 9: 383-393.
Literature Cited , R. Bernal & M. Ruiz. 1989. Aiphanestricuspidata (Palmae), a new species from Colombia and Ecuador. Brittonia41: 156-159. Braun, A. 1968. Cultivatedpalms of Venezuela. Principes 12: 39-103. & F. D. Chitty. 1987. Palmas aut6ctonas de Venezuela.Caracas,Venezuela. Bullock, S. H. 1981. Notes on the phenology of inflorescences and pollinationof some rainforest palms in Costa Rica. Principes25: 101-105. Burquez, A., K. Sarukhan, & A. L. Pedroza. 1987. Floral biology of a primaryrainforest palm,Astrocaryummexicanum Liebm. Bot. J. Linn. Soc. 94: 407-419. Burret, M. 1932a. Palmae Neogeae I. Notizbl. Bot. Gart. Berlin-Dahlem11: 313-327. 1932b. Die Palmengattungen Martinezia und Aiphanes.Notizbl. Bot. Gart.Berlin-Dahlem11: 557-577. . 1934. Bactris und verwandte Palmengattungen. Repert.Spec. Nov. Regni Veg. 34: 167-253. . 1937. Palmae. In M. Burret, Plantae duqueanae. Notizbl. Bot. Gart.Berlin-Dahlem13: 489-495. . 1940. Palmae. In Diels, Neue Arten aus Ecuador III. Notizbl. Bot. Gart.Berlin-Dahlem15: 23-38. Cook, O. F. 1901. A synopsis of the palms of PuertoRico. Bull. TorreyBot. Club 28: 525-569, pl. 43-48. Cruden, R. W. 1988. Temporal dioecism: systematical breadth, associated traits, and temporal patterns. Bot. Gaz. 149: 1-15. Cuatrecasas, J. 1957. A sketch of the vegetation of the north-Andeanprovince. Proc. 8th Pacific Sci. Congress 4: 167-173. Dammer, U. 1907. Beitrage zur Flora der Hylaea nach den Sammlungen von Ule's Amazonas-Expedition.Palmae. Verh.Bot. VereinsProv. Brandenburg48: 118-129. . 1915. Palmae.In: R. Pilger, PlantaeUleanaenovae vel minus cognitae. Notizbl. K6nigl. Bot. Gart. Berlin 59: 261-268. Drabble, E. 1903. On the anatomy of the roots of palms. Trans.Linn. Soc. London,Bot. 6: 427-487. Drude, 0. 1882. Palmae. In: C. F. P. von Martius, Flora Brasiliensis 3(2): 253-584. Dugand, A. 1944. Palmasnuevasy criticasColombianasII. Caldasia2: 443-458. Eichhorn, A. 1953. Nouvelle contributiona l'etude caryologique des Palmiers. Rev. Cyt. et Biol. Ve6g. 18: 139-151. Essig, F. B. 1971. Observationson pollination in Bactris. Principes 15: 20-24. Fisher, J. B. & H. E. Moore, Jr. 1977. Multiple inflorescences in palms (Arecaceae):their developmentand significance. Bot. Jahrb.Syst. 98: 573-611. Fournet, J. 1978. Flore illustre6 des phanerogames de Guadeloupe et de Martinique.Institutde la Recherche Agronomique,Paris. Frangi, J. L. & M. M. Ponce. 1985. The root system of Prestoea montana and its ecological significance. Principes29: 13-19. Gaertner, J. 1791. De fructibuset seminibusplantarum.G. H. Schramm,Tubingen. Galeano, G. & R. Bernal. 1985. Aiphanes acaulis, a new species fromColombia.Principes29: 20-22.
89 & . 1987. Palmas del Departamento de Antioquia,Regi6n Occidental.UniversidadNacional de Colombia,CentroEditorial,Bogota. Gassner, G. G. 1941. OberBau der mannlichenBliiten und Pollenentwicklungeiniger Palmen der Unterfamilieder Ceroxylinae.Beih. Bot. Centralbl.61A: 237-276. Gentry, A. H. 1981. New species and a new combinationin Palmae, Theaceae, Araliaceae, Apocynaceae, and Bignoniaceae from the Choc6 and Amazonian Peru. Ann. MissouriBot. Gard.68: 112-121. Glassman, S. F. 1972. A revision of B. E. Dahlgren'sIndex to American Palms. Phanerogamarummonographiae, TomusVI. Verlagvon J. Cramer,Leutershausen. Gooding, E. G. B., A. R. Loveless, & G. R. Proctor. 1965. Floraof Barbados.H. M. StationeryOffice, London. Hegnauer, R. 1986. Chemotaxonomie der Pflanzen. BirkhauserVerlag,Basel. Henderson, A. 1986. A review of pollinationstudies in the Palmae.Bot. Rev. 52: 221-259. . 1990. Arecaceae. Part I. Introduction and the Iriarteinae.FloraNeotropica53: 1-101. Hooker, J. D. 1886. Martineziacaryotifolia.Bot. Mag. 112: t. 6854. Hoyos, J. F. & A. Braun. 1984. Palmas tropicales cultivadas en Venezuela. Monografia no. 33. Sociedad y Fundationla Salle de Ciencias Naturales,Caracas. Humboldt, A., A. Bonpland & C. S. Kunth. 1816. Nova generaet species plantarum1. Paris. &. 1825. Nova genera et species plantarum7. Paris. Imschanitzkaja, N. 1987. Palmae (Arecaceae) in herbario institute botanici nomine V. L. Komarov (Leningrad) conservatae. 2. Specimina authentica taxorum a H. Karsteniodescriptorum.Novosti Sist. Vyssh. Rast. 24: 26-42. Jacquin, J. 1763. Selectarum Stirpium Americanarum Historia.Vienna. - . 1809. FragmentaBotanica.Vienna. Johnson, M. A. T. 1989. An unusually high chromosome number in Voanioala gerardii (Palmae: Arecoideae: Cocoeae: Butiinae) from Madagascar.Kew Bull. 44(2): 207-210. -, A. Y. Kenton, M. D. Bennett & P. E. Brandham. 1989. Voanioalahas the highest chromosomenumberin the monocotyledons.Genome 32: 328-333. J0rgensen, P. M. and C. Ulloa. 1989. Estudios botanicos en la "Reserva ENDESA", Pichincha-Ecuador.AAU Reports22: 1-135. Kahn, F. & K. Mejia. 1991. The palm communities of two "terrafirme"forests in PeruvianAmazon. Principes 35: 22-26. Karsten, H. 1857. Plantae Columbianae. Linnaea 28: 241-282, 387-412. .1866. FloraeColumbianae2. Berlin. Koop, H. 1907. Anatomie des Palmenblattes.Beih. Bot. Zbl. 22: 85-143. Kunth, C. S. 1841. Enumeratioplantarum.Stuttgart. Linden, J. 1875. Catalogueno. 93: 32-33. . 1881. Plantes introduiteset mises pour la premiere fois dans le commerce par 1'6tablissementJ. Linden. L'Illustr.Hort. 28: 31-32.
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Listabarth, C. 1992. A survey of pollination strategiesin the Bactridinae(Palmae). Bull. Inst. fr. etudes andines 21: 699-714. Little, E. L. & F. H. Wadsworth. 1964. Common trees of PuertoRico. Agriculturehandbookno. 249, Washington, DC. MacBride, J. F. 1960. Palmae. Flora of Peru. Part 1, no. 2. Field. Mus. Nat. Hist. Bot. Ser. 13: 321-418. Martius, C. F. P. von. 1823. HistoriaNaturalisPalmarum2. Leipzig. . 1839. HistoriaNaturalisPalmarum3:181-260, tab. 134-164. Leipzig. . 1847. PalmetumOrbignianum.P. Bertrand,Paris. Moore, H. E. 1951. Various new palms. Gentes Herb. 8: 223-238. . 1963. The types and lectotypes of some Palm Genera. Gentes Herb.3: 245-274. . 1967. The genus Gastrococos(Palmae-Cocoideae). ~--Principes 11: 114-121. . 1973. The majorgroupsof palms and theirdistribution. Gentes Herb. 11: 27-141. Motiska, D., C. Anderson, & J. G. Fullington 1984. The renewed quest for high elevation palms. Principes 28: 55-63. Read, R. W. 1966. New chromosomecounts in the Palmae. Principes 10: 55-61. ---. 1979. Palmae. In: R. A. Howard(ed.), Floraof the Lesser Antilles. Arnold Arboretum,HarvardUniversity, Cambridge. Roth, I. 1984. Stratificationof tropical forests as seen in leaf structure.W. Junk,The Hague. Rudolph, K. 1911. Zur Kenntnis der Entfaltungseinrichtungen an Palmenblattern.Ber. Dtsch. Bot. Ges. 29: 39-47. Ruiz, H. & Pavon, J. 1794. Prodromusflorae peruvianaeet chilensis. Rome. --- & . 1798. Systema Vegetabilium Florae Peruvianaeet Chilensis. Madrid. Sato, D. 1946. Karyotype alteration and phylogeny, VI. Karyotype analysis in Palmae. Cytologia 14: 174-186. Scariot, A. O., E. Llleras, E. & J. D. Hay 1990. Reproductivebiology of the palmAcrocomiaaculeata in central Brazil. Biotropica23: 12-22.
Schomburgk, R. 1848. Fauna und Flora von British Guyana.Leipzig. Sharma, A. K. & S. K. Sarkar. 1957. Cytology of different species of palms and its bearing on the solution of the problems of phylogeny and speciation. Genetica 28: 361-488. Snow, D. W. 1962. The naturalhistory of the Oilbird,Steatornis caripensis, in Trinidad,W. I. Part 2. Population, breeding,ecology, and food. Zoologica 47: 199-221. & B. K. Snow. 1978. Palm fruits in the diet of the oilbird, Steatorniscaripensis. Principes22: 107-109. Solereder, H. & F. J. Meyer. 1928. Systematische Anatomieder Monocotyledonen.Berlin. Sowunmi, M. A. 1972. Pollen morphology of the Palmae and its bearing on taxonomy. Rev. Palaeobot. Palynol. 13: 1-80. Syed, R. A. 1979. Studieson oil palm pollinationby insects. Bull. Entomol.Res. 69: 213-224. Thanikaimoni, G. 1970. Pollen morphology,classification and phylogeny of Palmae.Adansonia10: 347-365. Tomlinson, P. B. 1960. Seedling leaves in palms and their morphological significance. J. Arnold Arbor. 41: 414-428. - . 1961. Anatomyof the monocotyledons.II. Palmae. ClarendonPress, Oxford. -- . 1962. The leafbase in palms. Its morphology and mechanicalbiology. J. ArnoldArbor.43: 23-50. . 1990. The structuralbiology of palms. Clarendon Press, Oxford. Uhl, N. W. & J. Dransfield. 1987. GeneraPalmarum.L. H. Bailey Hortoriumand the InternationalPalm Society, Ithaca,NY. & H. E. Moore. 1971. The palm gynoecium.Amer. J. Bot. 58: 945-992. Wendland, H. 1878. In 0. de Kerchovede Denterghem,Les Palmiers.Paris. Wessels Boer, J. G. 1988. Flora of Venezuela, Palmae. Pittieria17: 201-206. Willdenow, C. L. 1806. Uber einige neue Siidamerikanischen Palmen.Samml.Deutch.Abh. Konigl. Akad.Wiss. (Berlin) 1803: 246-256. . 1807. De quelques nouveaux palmiers de l'Amerique m6ridionale.M6m. Acad. Roy. Sci. Hist. (Berlin) 13: 26-42.
NUMERICALLIST OF TAXA 1. Aiphanes acaulis Galeano & Bernal 2. A. aculeata Willdenow 3. A. chiribogensis Borchsenius& Balslev 4. A. deltoidea Burret 5. A. duquei Burret 6. A. eggersii Burret 7. A. erinacea (Karsten)H. Wendland 8. A. gelatinosa H. E. Moore 9. A. grandis Borchsenius& Balslev 10. a. A. hirsuta Burretssp. hirsuta 10. b. A. hirsuta Burretssp. kalbreyeri(Burret) Borchsenius& Bernal 10. c. A. hirsuta Burretssp. intermediaBorchsenius& Bernal
10. d. A. hirsutaBurretssp. fosteriorum (H. E. Moore) Borchsenius& Bernal 11. A. leiostachysBurret 12. A. lindeniana(H. Wendland)H. Wendland 13. A. linearis Burret 14. A. macrolobaBurret 15. A. minima(Gaertner)Burret 16. A. parvifolia Burret 17. A. simplexBurret 18. A. spicata Borchsenius& Bernal 19. A. tricuspidataBorchsenius,Bernal & Ruiz 20. A. ulei (Dammer)Burret 21. A. verrucosaBorchsenius& Balslev 22. A. weberbaueriBurret
List of Exsiccatae
91
LIST OF EXSICCATAE Acufa, J., s.n. (15). Agudelo, C., 1126, 1428(17). Allen, P., 1870 (10a). Anananch,RBAE223 (20). Anderson,A. B., 31 (8). Andre, E., 400 (2). Arbeldez,G., 1842 (12); 1977, (17); 2216, 2424 (17). Argiiello, A., 353, 418 (7). Asplund, E., 9174 (20); 14667 (22). Autorbridge,J., s.n. (2). Bailey, L. H., 60, 61, 291, 337, 352, 428 (15); 432 (2); 440 (15); 529, 547 (2); 584, 594 (15); 1649, s.n. (2); s.n. (15); 15014 (15). Baker,M., 6805 (20). Balslev, H., 2738 (7); 4418 (20); 4666 (7); 60581, 60658, 60733, 60742 (20); 62007 (7); 62011, 62012 (6); 62045, 62067 (20); 62084 (7); 62102 (10d); 62200, 62473, 62480 (20); 62495 (7); 62521 (19); 62531 (21); 69053 (20). Barfod,A., 48916 (14); 60003 (8); 60179 (21). Barker,E. E., s.n. (15). Barkley,F. A., 18C864 (12). Beard, J., 173, 230, 484, 624 (15). Beard, P., 1058, 1059, 1374 (15). Benalcazar,C., 11, 101 (17). Benavides, 0., 9642 (7). Bergmann,B., 67217, 67220 (20). Bernal, R., 50 (2); 56, 57 (lOa); 72 (1); 171, 251 (lOa); 330 (10b); 348, 349 (lOa); 379 (13); 440 (lOa); 534 (13); 849 (12); 895 (10d); 901, 904 (8); 958 (10b); 1079 (1); 1321 (19); 1328 (7); 1334, 1335, 1352, 1375 (12); 1378, 1392 (lOa); 1393 (10b); 1406 (20); 1435, 1442 (17); 1448 (8); 1455 (10c); 1521 (10d); 1538 (17); 1540 (5); 1553 (13); 1555 (10c); 1956, 1959 (2); 1961 (14); 1962, 1963 (10c); 1964 (17). Bernardi,A. L., 3327 (2). Betancur,N., 909 (16). Blicher-Mathiesen,U., 62606 (20). Borchsenius, F. 91421 (7); 91422 (3); 91423 (7); 91424 (19); 91427 (22); 91435 (20); 91440, 91441, 91442 (3). Box, H. 1760 (15). Braun,A., s.n. (2). British Museum (BM), s.n. (15). Britton, N. L., 396, 396, 789 (15); 1634 (2); 1701, 2857, 5230, 7829, 7854 (15). Broadway,W. E., 623, 754 (2). Buchtien, 0., 344 (2). Busey, P., 268 (10a). Callejas, R., 3280 (17); 4238, 8535 (16). Carrington,S., 1590, 1592, 1595, 1598, 1609, 1610, 1611, 1612, 1613 (15). Cer6n, C., 2381, 2621, 3032, 3116, 3186, 4268 (20). Chavez, R., 311 (2). Churchill,H. W., 5874 (10a). Cogollo, A., 3760 (16). Cook, O. F., 17 (2). Cornell (BH), #13 (17); s.n. (2); 416253 (15). Croat,T. B., 17194(10a); 18711 (22); 22684 (lOa); 51923 (12). Cuatrecasas,J., 8473 (12); 16492 (19); 18377 (17); 23885 (10c).
Curtis,s.n. (14). Dahlgren,B. E., s.n. (2, 15); 611704 (15). Daly, D. C., 101 (6). Dawe, M. T., s.n. (2). De Leon, J., 126 (17); s.n. (15). Delgaso, E., 287 (15). Diaz, S., 453, 1634 (12). Dodson, C., 6627, 14204 (7); 17582 (10d). Doyle, 12 (2). Dransfield,J., 4854 (10c); 4868 (19). Dryander,1288 (17). Duque, J. M., 573 (5); 4587 (2). Duque-Jaramillo,J. M., 1320 (17); 3367 (12); 4531 (17). Duss, P., 3815, 4136, s.n. (15). Eggers, 7135, 7175 (15). Ekman,E. L., 14845, 16429 (15). Ellenberg,H., 2877 (4). Ernst,W. R., 1673, 1701, 1709 (15). Escobar,0., 2673 (17). FLS, 7767 (15). Fantz,3467 (2); 3469 (15). Firenze (F-Beccari herbarium),46, 74, 75; 90; 92; 411, 3128/26; 3128/29, s.n. (2). Fleishmann,367 (2). Folsom, J. P., 4325, 4513 (10a). Forero,E., 6863 (14); 7224 (10a); 7535 (14). Foster,M. B., 1870 (12); 2117 (10d); 2164a (7). Foster,R., 9579 (2); 10101 (22). Franco,P., 1681, 2672 (17). Funck, 1655 (12); s.n. (2, 12). Furtado,C. X., s.n. (2). Galeano, G., 14 (17); 238 (10a); 253, 254 (12); 262, 266 (17); 331 (12); 454 (1); 461, 462 (10b); 564 (12); 1494, 1510 (20); 1676 (4); 1961 (16); 2061 (12). Galeano,M. P., 199 (12). GarciaBarriga,8443, 20933 (2). Gentry,A. H., 17450 (14); 22935 (22); 28658 (10a); 29148 (20); 35064 (7); 36514 (22); 40805 (13); 47724, 47756 (12); 53102 (17); 53978, 53978 (12); 54157 (2); 65315 (17). Glaziou,A., 13292 (2). Goll, G. P., 1044 (15). Gonzdles,A. C., 872 (2). Gonzalez, F., 1685 (12). Grant,M. L., 10538 (12). Hahn,M., 347, 1135 (15). Hammel,B., 15984 (20). Henao, L., 299 (11). Henderson,A., 099 (10a); 125, 126 (12); 520 (2); 722 (10a); 846, 868, 1133 (20); 1145 (2). Hermann,s.n. (15). Hernindez,J. J., 632 (16). Hill, S. R., 2632 (15). Hinds, s.n. (19). Holm-Nielsen, L. B., 24787 (3). Horn, s.n. (15). Howard,R. A., 11696, 17935 (15). Hoyos, S., 333, 832, 891, 912 (16). Huber,3411 (4). Hull, D. A., H-2, 10068 (15). Humboldt,A. von, s.n. (2).
Flora Neotropica
92 Idrobo,J. M., 198 (17); 9501, 9728, 10400 (12). Iltis, H. H., 56 (2). Izawa, K., 36 (2). Jack, J. G., 8742 (2). Jaramillo,2750 (12). Johnston, H. A., 1586, 1626, 1664 (2); 1672 (15); 1674, 1586a, 1587a, 1642? (2). Johnstone,s.n. (15). Judd,A. F, s.n. (2). Juncosa,A., 2172 (17). Kahn,F., 1684 (2); 1916, 2060 (20); 2535, 2546, 2554 (22); 2556 (4); 2646, 2651 (18). Kalbreyer,W., 1864 (17). Karsten,H., s.n. (2, 7). Kew Gardens (K): Acc 533-58.53301; 37, 51 (2); 86, 239 (15); s.n. (2). Killip, E. P., 11039 (2); 16362 (kill); 24608 (22); 33804 (17). Knapp,S., 6060 (10a). Krukoff,B., 10021, 10907, 11130 (2). Kuntze, 0., 1062 (2). Lorzing, J. A., 12134 (15). Lee, R. E., s.n. (15). Lehmann,F. C., 5290 (2); B.T. 383 (17). Leningrad(LE), 56.1, 61.12 (2). Linden, J., 1202, s.n. (2). Listabarth,C., 11-14389 (2). Little, E., 16305 (15). Loaiza, C. A., 319 (16). Londofio,J. B., s.n. (13). Loomis, H. F., 35 (15); s.n. (2). Lozano, C., 4903, 4905 (lOc); 5691, 5916 (19). Madison, 4567 (7); 4804 (8); 4838, 7125 (7); 7187 (8). Madsen, J., 86927, 86934 (9). Mejia, K., 125 (20). Miller, G. S., 93 (15). Moore, H. E. Jr., 5849 (15); 6015 (2); 6102, 6118, 6129 (15); 8358 (4); 8456, 8527 (22); 8529 (20); 8564 (2); 9470 (7); 10197 (17). Moraes, M., 836, 837, 842, 1311, 1312, 1314 (2). Mori, S., 3727 (2). Murray,W. B., s.n. (15). Nee, M., 11606 (0Oa);36870, 37066 (2). Neill, D., 8783 (20). de Nevers, G., 5395, 6366, 6405 (10a); 7745 (2); 8416 (10a). d'Orbigny,A., 8 (2). Paris (P), s.n. (7) Perez-Arbalaez,E., 8173 (12). Palacios, W., 1174, 2947, 4297 (20). Patin, C., s.n. (2, 15). Pav6n, J., s.n. (2).
Pearsall,1070 (6). Pennel, F. W., 8576 (2). Pinard,847 (2). Pinkley,H. W., 486 (20). Poeppig, s.n. (2). Ponce, M., 108 (20). Porter,D. M., 4388 (lOa). Purdie,s.n. (7, 2, 19). Rangel, 0., 5948 (12). Read, R., 905 (2); 1357, 2055a, 74-218 (15). Remage, G. A., s.n. (15). Rusby,H. H., 2862, s.n. (2). Sanchez, D., 12 (2). Schultze-Rhonhof,H., 2769 (20). Schunke,J., 2078 (2). Sharp,s.n. (15). Sintenis, P., 484, 2500, 2628 (15). Skov, F., 60601 (20); 64706 (19); 64708 (7); 64714 (22); 64715 (20); 64734 (21); 64735, 64736 (6); 64738, 64739 (7); 64742 (10d); 64747 (14); 64750, 64757, 64758 (7); 64770 (22); 64771, 64776, 64780, 64787, 64795 (20); 64818 (14); 64819 (10d); 64829 (19); 64839 (2); 64840 (7). Slane, V., 965 (15). Smith,A. C., 2643 (22). Sneidern,K. von, 1260 (20); 5385 (16). Sodiro, 187 (7). Stevenson,2105 (15). Steyermark,J. A., 52832 (3); 54535 (21); 90137, 116220 (2). Taylor,D., 126 (15). Tessmann,G., 4281 (22). Therese, Princessof Bayern, s.n. (2). Toro,R. A., 852 (17). Trail,J., 1070 (4). Triana,J., 730, 1765, 1767 (2). Ule, E., 116b (2); 6880 (20). Underwood,L. M., 878 (15). Uribe-Uribe,L., 683 (13). Vargas,C., 6333, 17375, 18694 (2). Velez, M. C., 817 (17); 2474 (2). Wall, E., 122 (2). Warming,E., s.n. (15). Weberbauer,A., 4665 (22). Wessels Boer, J., 1991 (2). White, G., 405 (2). Williams, R. S., 394 (2). Wright,C., s.n. (15). Wydler,H., 192 (15). Yepes, S., 1070 (17). Zanoni,T., 16879 (15). 0llgaard, B., 9306 (20); 57620 (14).
INDEX OF LOCAL NAMES Names in parenthsisrefer to numbersin the NumericalList of Taxa. charascal(2) chascaray(13) chascaraza(2) chirca (13) chonta (2)
chontaruro(2) chontilla(20), (22) chou picant (15) cirqui (lOb), (13) cocos rura(2)
Indexof ScientificNamesof Plants corocito de agua (13) corozo (2), (6) corozo anchame (2) corozo chiquito (2) corozo colorado (2) corozo de agiita (13) corozo del Orinoco (2) coyora (15) coyore (15) coyure (15) cuaro (12) cubarro(2) cuvaro (12) glouglou (15) glouglou rouge (15) grigri (15)
93 gualte (2) jimena tfue (20) macagiiita(2) macahuite(2) macawpalm (15) marara(2) mararai(12) mararave(2) mararay(2) pa m si mo ha (20) palma de coyor (15) palmito (14) peganore(14) pujamo(2) shicashica(4)
INDEX OF SCIENTIFIC NAMES OF PLANTS Synonyms are in italics. Primarypage referencesare in boldface type. An asterisk(*) indicatesa page with an illustrationor map. Acrocomia, 14, 16, 18, 26, 32, 33 Aiphanes, 1, 2, 3, 6, 8, 9, 11, 12, 14, 15, 16, 19, 22, 23, 26, 29,30,31,32,33,34,36,75,87 acanthophylla, 15, 71, 72, 73*, 75 acaulis, 4, 6, 9, 11, 16, 18*, 21, 29, 30, 32, 34, 40, 43*, 78, 79 aculeata,2, 3, 4, 6, 7*, 8, 9, 11, 13*, 14, 15*, 16, 17*, 19*, 20, 21, 22, 23, 26, 27, 30, 31, 32, 33, 34, 35, 43*,46,48, 49, 54, 75, 87, 88 caryotides, 87 caryotifolia, 15, 22, 26, 46, 47, 87 chiribogensis,4, 6, 8, 11, 12*, 13*, 16, 19*, 20*, 21, 22,24*,26,27*,29,30,31,32,34, 35, 43*, 50, 51*, 53 chocoensis, 70 concinna, 66, 68 corallina, 3, 71, 75 deltoidea, 6, 12, 13, 14, 16, 21, 28*, 30, 32, 36, 44*, 51, 52, 85 disticha, 87 duquei, 6, 11, 12, 14, 16, 17*, 19*, 21, 27*, 29, 30, 32, 35, 42*, 51, 52, 53 echinocarpa, 67 eggersii, 4, 6, 8, 9, 10*, 11, 12*, 15*, 16, 17*, 20, 21, 22, 23, 24*, 25*, 26,29, 30,31,32, 33, 35, 41*, 53, 54, 55* elegans, 46, 47, 49 erinacea, 3, 4, 5*, 6, 7*, 8, 11, 12*, 13*, 14, 15*, 16, 18*, 19*, 20, 21, 24, 25*, 26, 28*, 29, 30, 31, 32, 33, 35, 36, 43*, 54, 56, 57*, 58, 66, 78 ernestii, 46, 49 erosa, 22, 26, 71, 72, 75, 76 fosteriorum, 1, 2, 65 fuscopubens, 22, 62 gelatinosa, 4, 6, 9, 11, 14, 16, 19*, 20*, 22, 28*, 29, 30, 34, 42*, 58,59*, 60, 63, 83 gracilis, 87 grandis,4, 5*, 6, 9, 14, 15, 16, 19*, 20*, 22, 28*, 29, 30, 31, 32, 35, 42*, 55*, 60, 61
hirsuta,1, 2, 4, 6, 9, 13, 14, 16, 21, 29, 30, 35, 36, 44*, 60, 61, 62, 87 subsp. fosteriorum,8, 15, 16, 20*, 22, 27*, 29, 44*, 62, 63, 65 subsp. hirsuta,20*, 22, 27*, 29, 44*, 62, 63, 64, 65, 71, 87 subsp. intermedia,1, 2, 16, 19*, 20*, 22, 27*, 29, 44*, 55*, 62, 64, 65 subsp. kalbreyeri,16, 20*, 22, 27*, 29, 44*, 55*, 62, 63,65 horrida,46 kalbreyeri,1, 2, 63 killipii, 46, 49 leiospatha, 63, 87 leiostachys,6, 8, 16, 17*, 22, 27*, 29, 30, 35, 42*, 65, 66 lindeniana,2, 6, 8, 9, 15, 16, 17*, 22, 27, 28*, 30, 35, 36, 41*, 54, 66, 67, 68*, 84, 87, 88 linearis, 8, 9, 11, 14, 15, 16, 19*, 20*, 22, 28*, 29, 30, 32, 35, 41*, 61, 67, 69, 70*, 77 luciana, 22, 71, 74*, 75 macroloba,5, 6, 7*, 9, 10*, 11, 14, 16, 22, 23*, 28*, 29, 30, 32, 34, 41*, 63, 69,71 mimima,2, 3, 4, 6, 8, 9, 11, 12, 15, 16, 17*, 19*, 22, 6, 29, 30, 31, 32, 33, 34, 35, 41*, 71, 72, 73*, 74*, 75, 76 monostachys,22, 62, 63, 70, 87 orinocensis, 46, 49 pachyclada, 62, 63 parvifolia,6, 9, 11, 12*, 16, 22, 28*, 29, 30, 32, 34, 43*, 76, 77, 80 praemorsa,46 praga, 2, 87 schultzeana,80 simplex, 6, 8, 11, 16, 18*, 19*, 22, 23*, 28*, 29, 30, 32, 34, 36, 43*, 56, 77, 78 spicata, 1, 2, 4, 5*, 6, 10*, 11, 16, 19*, 20*, 22, 28*, 29, 30, 34, 44*, 46, 47, 78, 79 subg. Brachyanthera,3, 33 subg. Macroanthera,3, 33
94
FloraNeotropica
Aiphanes (continued) tessmannii, 84, 87 tricuspidata,4, 6, 8, 11, 12, 13, 16, 22, 28*, 29, 30, 32, 35, 43*, 79, 80, 81* truncata,46, 49, 50 ulei, 4, 5*, 6, 8,9, 10*, 11, 12*, 14, 16, 18*, 19*, 22, 28*, 30, 32, 35, 44*, 49, 80, 82, 83, 88 verrucosa,6, 7*, 8, 11, 12, 14, 15*, 16, 22, 24*, 27, 28*, 30, 35, 67, 83, 84* vincentiana, 71, 72, 75 weberbaueri,4, 6, 9, 16, 18*, 19*, 22, 27*, 28, 30, 32, 33,35, 36, 44*, 52, 84, 85, 86*, 87 Ammandra,23 Aphandra,23 Arecoideae, 1, 2, 32 Arenga, 11 Astrocaryum,17, 18, 26, 32, 33 Attaleinae, 19 Bactridinae,1, 2, 6, 9, 11, 32, 33 Bactris, 8, 9, 17, 18, 23, 26, 32, 33 acanthophylla,71 caryotifolia, 2, 9, 33 ciliata, 3 erosa, 71 gasipaes, 23, 26 minima, 2, 3, 71, 75 praemorsa, 46 Butia, 26 Calameae, 8 Calyptrocalyx,11 Caryota horrida, 2, 46 Catoblastus,8, 11 Ceiba, 27, 54 Chamaedorea,11 lanceolata, 3 linearis, 3 tenerrima,9 Cocoeae, 18, 32 Cocos, 15 Curima,3, 34 colophylla, 34, 71, 72 corallina, 71 Desmoncus, 8, 17, 18, 32, 33 Dictyocaryum,4, 79 lamarckianum,79 ptariense,4 Elaeis, 18 Euterpe,3 aculeata, 46 ensiformis, 3 praga,2, 87
Gastrococos, 17, 26, 32, 33 Geonoma,9 interrupta,3 Iriarteeae,9 Jubaeopsis,26 Marara, 3, 33 aculeata, 46 bicupidata,3, 46 caryotifolia,46, 47 erinacea, 3, 33, 54 Martinezia,2, 3, 33 abrupta,3 acanthophylla,71 aculeata, 46 aiphanes, 3, 46 antiochensis, 87 caryotifolia,2, 3, 22, 23, 26, 46, 47 ciliata, 3 corallina, 71 disticha, 87, 88 elegans, 46, 49 ensiformis,3 ernestii, 46, 88 erosa, 26, 71, 75 granatensis, 88 interrupta,3 killipii, 46 lanceolata, 3 leucoplweus,88 lindeniana,22, 66 linearis, 3 minor, 88 roezlii, 88 truncata,46, 47, 50 ulei, 49, 80, 88 ulei (homonym),46, 49, 88 Nephrosperma,8 Neodypsis decaryi, 8 Oenocarpus,8 Oncospermatinae,8 Orania,8 Paralinospadixhollrungii,60 Phytelephas,23 Socrateasalazarii,4 Syagrus,26 Tilmia,3, 34 caryotifolia, 34, 46 disticha, 88 Voanioala,26 Wallichia,8 Wettinia,11
INDEX OF SCIENTIFICNAMES OF ANIMALS Cecidomyiidae, 31 Ceratopogonidae,31 Chrysomelidae,31 Coleoptera,32 Drosophilidae,31 Hemiptera,31, 32 Hymenoptera,31
Lepidoptera,31, 32 Mycetophilidae,31 Nitidulidae,31 Meliponidae,31 Sciaridae,31 Steatorniscaripensis,32 Syrphidae,31
