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
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FLORA
NEOTROPICA
MONOGRAPH
70
AIPHANES (PALMAE)
FINN BORCHSENIUS
AND RODRIGOBERNAL
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FLORA
NEOTROPICA
TO I1
TROPIC
OF
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CAPRICORN
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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
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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 .............................................
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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 discutela 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
westernEcuadorwhereA. 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
Morphology
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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
6
Flora Neotropica
20
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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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FiG. 3. Stembranching.
A. StemofAiphaneserinacea showing
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B. Stolon
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inA. macroloba(Skovet al. 64818).
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-X i_41~:~iliiijit':it
ru~~~~~~~~~~~~~~~~~~~~~~~~i
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...'.,;--J
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y
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FIG. 4. Armature.A. Stem spines of Aiphanesaculeata (Bernal & Galeano 50). B. Spines on distal partof stem and leafsheathof A. verrucosa (Skov et al. 62734).
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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S. Leaves.
FIG. 5.
Leaves. A.
A. Densely
narrowpinnae,
al. 62012).
et Ml.
B. Remotely
clustered, nmnrow
Densely clustered,
62012). B.
(Balslev ?~
Aiphanes eggersii
pinnae, Aiphan?~
eggersii (Balslev
grouped,
Remotely grouped,
FIG.
broadpinnae,
A. M?ei
ule (Bolslev
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
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
~~~~~~~~~~~Dig~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
~.....
:~
..
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 (Skovet 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
~~4~~
E3
~
a
~~"
.
--1.111
cl
jji.i
II~IG.}
1
A
lengtE
5
.
1
fc
S
,
i
FIG. 8. Antherlength ranges for species of Aiphanes.
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
s;.,~~~~~~~~~~~~~~~I
....
.
?..:..
FI.9
C.Salfut
..
.
........":ii'ii:i~iilliiiii:~::
:?
.....
rit.A
fA
rit
fApansegrii(ase
L680.D
rnca(kve
e
l 212.B
eeln
ln
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
nocrso
fA
. ercs
(ase
t l
23)
claa(enl&Brheis15)
(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
_
___~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~':
..
..........
.. ...
i~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ni
IVE1
;Kj
?
i
TI
* : ::
*:.
*~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
_
i
C~~~~~~~~~~~~~~~~~~~
Wi~~~~~~~~~~~~~~~~~~~~~~~~~: e}...:....~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
_
.:
.
;-
i...
X
X
.;
..
?J'.ls':?;
...
...
...
.....
.
.:.:.
ii??~~~~~~~~~~~~~~~~~~~~~~~~~
''
`it-,er;:4
....
?iI~i.i':Ciii:ia
------------
............
. ...
.
........
..... .
...
..:?
??
. .........
:i,i~~iikb;~~-
'" ''
I~--"~e~ussss~E~l~?Wia~
lr
~~~~~~~~~~~~~~~~~~
~~~
~~~
~~~~~
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
tal 21)
. .mnm
FI.1.Plln
(ase
.Apansegri
.A idnaa(edesn&Bra
D.A uui(enl&Dva14)
A226
22
,C
;Dx13;
,Fx22I
Ber
2)F
. .auet
8)
.lisahs(ea
(e
ta
67)
9)
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..
........
__
....
..
_
.. .
..
( C aleI a n o et al
...
9470).
. 26 2)...?
D. A
ac
ul s ( B r n l &
......
F..
..0
al
a
o
1)
.
. e r n a .. e.... (
.c .....
al l
v
t
l
6
00 )
a
... ...
-Li~~~~~~~~~i
.ii
..iiis.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~...
i:
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
9i~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
HiI~
.i
i:
?:::?i' ?;;i??::..
q
.i?
..
1;~~~~~~~~~~~~~~~~~~~~~~~~
ii.ji
LM
:???:,~~~~~~~~~~~~~~?
?:ic.:;r
'iib::::I~~~~~~~~~~~~~~~~~~~~~~~~:.Cal:1
A
........
...
..~~
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~;....
'
'i
i.~~~~~~~~A
:s
:?? :i
...
Y............
i~~~~~~~~~~~'
~
~
~~~~~~i??
*??'j ?;
:?
ii
i:.
??~~~~~~~~~I
FI.1.Plen
ue Bllv
.A wbraei(orheis&Peesn947.
.A
20)
.Apae
. A. rinaca (Baslev t al.62007. F. . eriacea
(Galeno etal. 22). D A.aaulis(Bernl & Gleano71).
947). x 980 B 21 1;C x433 D- x226
ipe
Moor
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
??~i
i
i~~i
a!
Xiu 5,!3k,ii:riii!:!,:
i
'.
:?:?
iiS~~~~~~~~~~~~~~i?:.~
~*- ~ ~~ ~ ~~~~~~~~!.
.:
'
s;. ..
.$,:::.....
i!
..
,i:.....
r_~~~~~~~~~~
~~~~~~~~~~.:..:'.
.... .....
Mi
:
'
:
c
v
'
.
..A
.
i
???':i
1.':::"':"
sjiH~~
:::,a~~~~i*ii!
..;i!iiiiic~~~~
. .........
:.....;
..
54185) J. A simplex(Galeanoet al 262) K. A lineari
(Gentryet al 40805) L.
A.
91442).
.-.
_ .. ;
e
_
-
.
chiribogensis(Borchsenius
M
e}I
:erinacea
(Moore9470) N A . hirsuta
ssp
intermedia(Dransfield4854). O.A. gelatinosa(Bernal& Gaeano 901). All x6068
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
_
.
.. :..
:~:'L
'7~~~~~~~~~~~~~
~'7
P~~~~~~~~~~~~~~~~~~~?~~~~.
.... .
=
;
,,#
.
.
....
.
.....b
.. ... .....'
.
L~:!~~:~i::~:!t:~i~f~i~il:"-.:,
/6::~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
2:'~
~' : .............
?W.-
~:?~~:
L
d.::'
?....
.. .. .....i.....
:'
p
:'
~'*:
.:
..
"
";~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
..':.~..
~....
.......
'
...'. : :.*? ,' ~:.~'.
:'
(:_?".
L. ?
4*
?
? ~.
..
..
...
.
~
:
... 1 .
..
13
~
?.i
.
:!iu4ttf
. ..
..
?:.
fti
~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~~~~~~~~~??q.~
...........~
...
FIG
'i .:R'
.... ....
~~~~~~~~~~~~~~?~
..
j
V
.
,,
.. ....
:
i.
..
Polen'A.Aipane
.c
..
'
N. .:...
:
:
....
..: . ::..:.
~~~~~~~~~~~~~~~~~~~~~~~~~~~.....
...:
.......... ..
4 ...::. ::..spa
...
.
D. A. grandis (Steyermark
54185).
E. A. g..e
.&......
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.
& Borchsenius
2646).
.
..........
.. al.7.1........
F
(Madison
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
.....ii'
;::~~~~~~~~~~~~~~~~~~~~~~~~~~~
:.~~:
i~~~~~~~~~~~li
~:. ~ ~ ~ ~ ~ ~ ~ ~
~~~~~~~~~~~~~~~~~?
~
i
?'
'41
ML
??:&.'~??9-?,
:: t,:
:'..i:.::
?~~~~~~
~ ,. ~ ~~~~~~~~~~~~~~~~~~~
:'.,"-'.~
?:;~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.
?~?.r:',~i.. ~. ~F. ii~?... ~
~ ~ ~ ~ ~ ~ ~ ::j~
~'~':.
:..
.;i...>.,
"'
?:
'*"3
:.I:.'.?
"'''~~~~~~~~~~~~~~~~~~
i
i:"~" .:
~
ai'iii,
::~~:'~:
':.':
?
?
?*
....
?; " ,
'DU"':::?
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~:~,~,'l.:::'
?;
. . .
i.
~ ~ ~ ~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~ ~~~~~~~~~~~~~~~~~~~~~-:a-s
--- -------- --
H
rii
~~~~~~~~~~P!
:i~~~~~~~~~~~~~~~~~~~~~~~~i
i
s?
..::.. .in?
:
~:'
,...::::.
Dans 484)
AB
N
FIG".
1.
.
': ....
.:
,.!::<..
.'::.,...<~
. . .
:.;:'.,.
r:;"
i...
x i
2
1
2
lCEx1'Fx74
'
.. B??x11;
:..":"
...........'*
.
25'. ....p.
3Bllv
'"&:Ga':n
45 '.'len.of..:._::.,:rsut.-'A :
.' ..yer (Bena 39;B~a
& '.areaa
o
?ne
:AF
D. Subs...,
:. C:5,~:F,
62102)~~~~~~~~~~:
<"::.'
1
':'ut
:
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:
na
et al.98.
B.Su
', .Sus.
. ?otrou
ine:ei
ta.
285
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
g
| |
|
l
|
,
.
,
.
.
m.
.
, .., . i
.
. ..:
...
! .
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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:i'iii:iiliiiiiiiii:i:
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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 (Skov et 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
~~4
i: i
FI.17
lra
+R
ntoy
.
Xi.o
tmnt
lwro
ipae
ges
So
ocsnis675
hwn
nhrwt
for olenscsan cnnciv
wthnueou
o A riaca Bocseiu
sleeis(bac)
x5) B Samnaeflwe
942)
x ?.truhcnra
ato
itlld
hwn
I6.CF
isilt
lwr
o .eiae
Bocsnu
12)
etay
oue
nine ae fslris(lc)ada
ue
C.Xs hog
aa ato itlaefowrsoigte
uruddb
lae ihmn
ie
n trioia
oeas
u otrrn)cnaiignmru
ovsua
tans(ih)
usdptl
fiber trands(f) ( ). D.ongituinal ectionthroug basa part f pisil, shwing pndulos orthtropou ovuls and entra
ise
x2.E.Xs
idl
itl hwn
ueos
ovsua
ihscear
ato
ie
stlrcnl()lie
hog
etr
aa x2)
stad
) .Coep
fslris(lc)
enrlsya
aaadtresptlncais()(
fspa
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 pistillateflowershowingtreelocules 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
-
4'
e5t'.j
--
4/
I
a
0~~~~~~~~~~~~~~~~
,'r
.
a~~~~~~~~~~~~
lo~~~~~~~~~~o
(
J
0~~~~~~~~~
a?
,r
:i~~~~~~~~~~~~~~I
/r'
A
~
~
~
~
~
'
~~5N~1EI
FI.1.Dsrbto
Ailune aclet
(crls;A
aps A.??~
sbp
irsutaandsusp kabeei(qae)
hist cice)
gesi(tr)Amnm
us.itrei(rage)
tinls
ist
us.fseirm(tr)
us
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
.j-0^
0_
,._
-'
-?
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
beltsalong
CCA:Cordillera
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.
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
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.
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)
34
FloraNeotropica
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,shortlyspinulose 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.
35
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
37
38
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
39
40
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
Pt). Lectotype(here designated).Martius,Palmetum
Mun.de
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
(fl), Steyermark
m,
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/w ratio
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
throughHuanuco and 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
*
.
???.
7
;Bii.ii:iilii:il,,l: ii'il,:il:::i I
;I:iiild:i:
.
..:
.
..:..
..
:.
:
~?
. . :. ..
...:... . . ....................
:
~~~~....~..........................
~~~~~~~~~~~~~~~~~~~~i'iilii::ii.lliiii;ii:iiiil:illii?i
~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~
~
'-?il??,:i.!i~~~~~~~~~~~~~~~~~~~~iiii;iLliCI
P!ll'~~~~~~~~~~~~~~~~~~~~~~~~~ii.:l~~~~~~~~~i:liii
.I:i'iiiiii'iiiiiiiiii
~~~~~
~~~........
~~~~~~~~~~~~~~..:
..............
..
.. .... .. .... .
... ...
.................
........~
....
....
..
. .... ..... ... ... .... .....
..
ilil!6;
.... ...
.. .... ....
... ....
n
.
iil:
.... ..il
...
Ei4
.
.. .. ....
...
.......
.
.
.
..~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
lllll:'
i,~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
??--??-~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
.. . . .. ... . ..
tCI
... .... .... .... ...
FIG.
22.
A.
(
...........
....FIS
Aiphanes
chirib~ensis,
inflorescence
(Skav
et
al,
64822),
B.
A,
duquei,
inflorescence
(Bernal
.... ....
&
D...... ....
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
91441(AAU,QCA);
2000 m, 3 Dec 1989(fr),Borchsenius
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 pinnae 6-9 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
high,
petals,
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.
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
Illustrations. Figs. 5A (leaf), 6A (inflorescence),
to 1 cm long; pinnae (8-)10-19 per side,
9A (fruits), 10A (pollen), 16D (leaf antomy), 17A spines,
insertedin groupsof 2-3(-4) separatedby 10-20 cm,
(flower anatomy), 18A (distributionmap), 21E (mid- in different
planes, strongly plicate, narrowly to
dle pinnae), 23A (habit).
broadly cuneate, 1.3-4.5 times as long as wide,
Local name and uses. Corozo (Ecuador).Meso- incised-praemorseat apex, symmetrical around the
midrib,with a 2-9 cm long finger-likeprojectionon
carp and endospermedible.
the distal margin, adaxially glabrous or nearly so,
Distribution and habitat. Endemic to the dry abaxially and on marginsminutely spinulose, someparts of the coastal plain of Ecuador below 600 m times both sides covered with long, yellow spinules,
altitude. Common in the province of Manabi. The midrib abaxially with 0-3 yellow
spines, ca. 3 cm
naturalhabitat is semi-deciduous, dry Ceiba forest, long: basal pinnae 6-30 x 0.5-8 cm; middle
pinnae
but it is often found in pasturesor sometimes planted 12-42 x 5-15 cm; apical pinnae 3-6 ribbed, 14-32 x
near houses.
5-25 cm. Inflorescenceerect or arching,once- or octwice-branched;most parts with a brown,
casionally
Specimens examined. ECUADOR.EL ORO:Near
caducous
indument;
prophyll26-73 cm long, 1-4 cm
600
15
1978
101
(fem fl), Daly
Puyango, m, Aug
(NY).
MANABf: Upper Rio Jama valley, rd. between San Isidro wide; peduncularbract up to 185 cm long, 2-5 cm
and San Jacinto,nearestero Capa Perra,300 m, 4 Nov 1982 wide, coriaceous,brownspinulose;peduncle 80-210
(fr), Pearsall 1070 (AAU); 10 km N of Canoaon the Pacific cm
long, 5-10 mm diam. at apex, densely armedwith
coast, 3 km from the coast, 50 m, 26 Mar 1987 (fl, fer fl,
slender,
yellow spines, to 1(-2.5) cm long; rachis
Borchsenius
et
al.
62012
fr),Balslev,
(AAU,NY,QCA);2
km N of San Vicente town center on Pacific coast, ca. 1 km 13-90 cm long, covered with brown to purple spininland, 50 m, 25 Mar 1986 (imm fr), Balslev,Borchseniuset ules; 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 pinnae 48-60 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,
subsp. hirsuta
spinulose, 7-11 mm diam. ........
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
rachillae
shorter
apex;
spines, becoming
15
Mar
1991 (fl, fer fl, fr), Bernal & Borchm,
cm
with
an
to
basal
rachillae
40-53
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~~~~~~~~~~~~~~~~$
~c-
,!-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
?
....
.,..
..,.:
i~~~
~;~~~~~~~~~~~~~~~~~~
....UIP
.i~~~~r
l.~3.
C"'~~~~~~~~:"
::
:i
i~~~~~~~~~~~~~i
-
'"
!C
s
,*?~~~~~~~~~~~~~~,
:i::
x~~Bs~~8~9!s ~ s~~~~:..~
n~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
?
e
-
.~?''.. i
:~.. ~ ~
':'.
~ ~
l
A
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
fructucora1844.Basedon Palmadactylifera,
aculeata,
Mun.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,"
Specimencollectedin Horto
Lectotype(heredesignated).
6863 (MO);20 Mar 1980 (fr), Foreroet al. 7535 (COL, Lindeniano,
Dec 1871 (fr), Patin s.n. (lectotype, K).
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) widening at 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
each subtendedby a 1-2 mm long bractcovrachis,
1312
1932, Kalbreyer
(Bt). Neotype (here desigthe
ering
pistillate and the proximalstaminateflower
de
San
Mun.
COLOMBIA.
Luis,
nated).
Antioquia:
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
of 2-4, these occupying3-7 cm along the
groups
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 pinnae 3-7 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 at Moyobambain Peruwas visited
the type locality was visited in May 1990, only one
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
pinnae
many
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 pinnae similarin 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'sdescription falls within the circumscriptionof A. weberavailable.
The existing materialfrom Peru is insufficient to
baueri in the presenttreatment.
resolve the taxonomy of A. weberbauerisensu lato in Aiphanes leiospatha Burret, Notizbl. Bot. Gart.
Berlin-Dahlem11: 571. 1932. Type. Colombia.Ana satisfactory manner.The variation patternresembles that found in A. hirsuta, which is still incomtioquia: Kalbreyer 1607 (holotype, Bt). Aiphanes
leiospathawas basedon an inflorescenceassociated
pletely understood despite the fact that there exist
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
Burret (here treated as a synonym of A. hirsuta
from linear, regularly inserted pinnae to cuneate,
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,
subsp. hirsuta.
degree
thickening
species seem to occur in distinct sympatricforms in Aiphanes praga Humboldt, Bonpland, and Kunth,
Nov. gen. et sp. 1: 303. 1816. [= Euterpe praga
some areas, whereasdifferencesbreakdown in other.
(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.
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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