histÓria taxonÔmica e filogenia da pupunha
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CHAPTER 1
INTRODUCTION
The pupunha (Bactris gasipaes Kunth) is the only
domesticated neotropical palm species and is found only as
a cultivated plant (Fig. 01, a, b). It was selected and
artificially distributed in the past by the Amerindians,
well before the first Europeans arrived in the Americas
(Clement, 1988). Today, the artificial selection and
distribution of pupunha has been conducted mainly by
research centers and commercial planters. As a result, its
current distribution ranges from Northern Honduras, in
Central America, to southeastern Bolivia and the Atlantic
coast of Brazil, in South America (Fig. 02). It grows more
easily in the lowlands and humid areas with short dry
seasons, although it is possible to find it in the Andean
foothills of Bolivia, Ecuador, Colombia and Venezuela
(Mora-Urp, 1984).
In the past, several specimens of cultivated pupunha
were described as distinct species or varieties. A few
wild palms, also called pupunha by local people, were
described as species and considered to be closely related
to the domesticated plants. With a few exceptions, all
described species and varieties of the cultivated and wild
pupunha share two main characters. They all have large
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stems, the biggest in the genus Bactris, and flattened
fibers adnate to the endocarp. The cultivated plants are
easily recognized by theirs large fruits, usually more than
3 cm long, and ovoid shape. The wild species are
vegetatively very similar to the cultivated plants, but
usually have much smaller globose fruits.
The main products obtained from the cultivated pupunha
are edible fruits and the palm heart, which rivals in
quality that extracted from Euterpe palms (Fig. 03, a, b).
The fruits are widely sold and are eaten after cooking.
They also can be used for oil extraction, flour
preparation, and animal feed (Clement, 1995). More
recently palm heart extraction is becoming the most
important economic use of this palm. In Brazil pupunha is
now reaching the status of an industrial crop due to the
establishment of large plantations for palm heart
extraction (Fig. 04).
Although important economically, there are
several disagreements regarding the taxonomy of pupunha.
It is not totally clear what are the correct names to be
applied to the cultivated and wild species, how many of the
species can be considered as valid, synonyms, or simple
varieties. It is also unknown what taxon or taxa are
related to the domestication process that gave rise to the
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cultivated pupunha and one important question is still
unanswered: in what place was pupunha domesticated?
Due to the confusion related to its taxonomy, origin,
and ancestors involved in its domestication, there is an
unavoidable necessity to present a detailed and careful
account of the taxonomic history and origin of pupunha.
For this reason, this study is presented in two distinct
sections. The first section is dedicated to a review and
discussion of the several aspects related to the taxonomic
history and origin of pupunha. The second section
presents: (a) a review and discussion of the recent
taxonomic and cladistic studies of Bactris and the
placement of pupunha and related species, (b) a discussion
of the classification system for cultivated and wild
pupunha developed by agronomists, and (c) a cladisitic
analysis of pupunha and related species that was performed
during the development of the present study.
The results of the cladistic analysis are used to
propose a new phylogenetic arrangement for the cultivated
and wild species, and to indicate the most likely place
where pupunha was domesticated.
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SECTION I
HISTORY AND ORIGIN OF PUPUNHA (Bactris gasipaes Kunth).
CHAPTER 2
TAXONOMIC HISTORY OF PUPUNHA (Bactris gasipaes Kunth) AND
ALLIED SPECIES.
Introduction
The binomial Bactris gasipaes was first used by Kunth
(1816) to describe cultivated plants found in Colombia by
Humboldt and Bonpland. However, the taxonomic history of
pupunha began eighteen years before Kunths publication,
when Ruiz and Pavn (1798) described Martinezia ciliata
from Peru. It is not clear, however, if Ruiz and Pavn
described their species from cultivated or wild plants.
Martinezia Ruiz & Pavn was rejected as a genus name for
pupunha species and M. ciliata was transferred to Bactris
by Martius (1826).
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Martius (1824) created the genus Guilielma and
described G. speciosa from cultivated plants he had found
in the eastern Brazilian Amazon. The genus Guilielma was
widely accepted at that time and many palm species and
varieties were described or transferred under this name in
the following years. This situation persisted until the
beginning of the 1960's, when Guilielma was replaced by
Bactris as a valid genus in several publications (MacBride,
1960; Wessels Boer, 1965, 1988; Uhl and Dransfield, 1987).
However, some non-taxonomist researchers working with the
cultivated pupunha started challenging this new proposition
suggesting that Guilielma could not be discarded as a valid
genus at all (Mora-Urp, 1984; Clement, 1988). The
discussion about the validity of Guilielma was definitely
over when Sanders (1991) published the first cladistic
study of Bactris in which he included Guilielma. Despite
being criticized, Sanders results regarding the monophyly
of Bactris are accepted by current palm taxonomists
(Henderson, 1995; 1996; Henderson et al. 1995; d'Granville
1997). The reason for this acceptance is that Bactris is
not monophyletic if Guilielma is segregated. In fact,
Bactris would have to be divided into eight or more poorly
defined genera in order to keep Guilielma apart. Sanders
gave the name Guilielma to the clade that included the
pupunha species, both cultivated and wild.
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In the following pages I will review and discuss all
species and varieties of pupunha described in the past
(Guilielma clade Sanders). In this review I will also
include Bactris setulosa, Bactris riparia, and all taxa
currently accepted as synonyms of these species by Wessels
Boer (1988), Henderson (1995) and Henderson et. al.,
(1995). The reason for this inclusion is explained in the
topics I discuss each of these species.
The taxa of the cultivated and wild pupunha
The species and varieties here included have been
considered by several authors (Clement, 1988; Henderson,
1995; Henderson et. al., 1995; Mora-Urp, 1944, 1994, Mora-
Urp and Clement, 1981; Wessels Boer, 1965, 1988) as the
ones that better represent the several forms of the
cultivated and wild pupunha. The historical aspects and
the taxonomic disagreements over the correct use of the
names are discussed in most cases.
Martinezia ciliata Ruiz & Pavn, Systema veg. 275.
1798. Bactris ciliata (Ruiz and Pavn) Martius, Hist.
Nat. Palm. 2: 95, t. 71 fig. 3, 1826. Guilielma ciliata
(Ruiz and Pavn) Wendland in Kerchove, Palm. 246.
1878. Type. Peru. Hunuco: Pozuzo, s.d., Pavn s.n.
(Type at MA, not seen).
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This was the first pupunha to be described. Ruiz and
Pavn, however, used the genus name Martinezia and included
under it several species that are now known to belong to
other palm genera, such as M. ensiformes (=Prestoea
ensiformis), M. linearis (=Chamaedorea linearis)and, M.
interrupta (=Geonoma interrupta). As a result, Martinezia
Ruiz and Pavn was rejected, and is now recognized only as
a synonym of Prestoea (Uhl and Dransfield, 1987). Martius
(1826) studied the type of M. ciliata and transferred it to
Bactris. It is unknown, however, the reasons why Martius
did not transferred M. ciliata to the genus Guilielma,
which he himself had created. He probably did not noticed
the endocarps of Pavn's specimen were similar to his G.
speciosa, although of a smaller size. The transfer of M.
ciliata to Guiliema was made later by Wendland (in
Kerchove, 1878).
B. ciliata is still poorly known and, besides the
type, apparently no additional collections from the type
locality have been made. Bernal (1989) believe it is the
same species as B. gasipaes Kunth and proposed the
conservation of the later name, arguing it has been used
for a long time in the botanical and agricultural
literature. However, it is difficult to conclude if the
endocarps of the type specimen (illustrated in Martius,
1826) are from cultivated or wild plants. Ruiz and Pavn
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did not note in the description if the fruits were edible,
a common characteristic of all domesticated pupunha. Ruiz
(in Dahlgren, 1940) commented only about the uses of the
palm heart, reported to be tender and tasty, raw or
cooked, but reported to be coarse. Palm hearts of
cultivated pupunha are known by their high quality, while
many wild pupunha are scarcely reported as having edible
palm heart. Ruiz noted the uses of the palm trunk, but
this does not help decide if the palm is domesticated or
wild. The final answer for this puzzle will be achieved
only when new collections from the type locality is made.
Bactris gasipaes Kunth, in Humboldt, Bonpland &
Kunth, Nov. gen. sp. 1: 302. 1816. Guilielma
gasipaes (Kunth) L. H. Bailey, Gentes Herb. 2: 187,
1930. Type specimen. Colombia, Tolima: Ibagu, n.d.,
Bonpland s.n. (Type at P, not seen; F neg. 38701).
Kunth described this species from material collected
in Colombia by Bonpland and Humbold during their voyage to
tropical America. The epithet is derived from the common
name "gachipaes" given to the species in that region of
Colombia. For many years it was accepted to attribute to
Humbold, Bonpland and Kunth (H.B.K.) the description of
this species, however, it is now known that Kunth was the
one who really wrote the description, therefore the correct
authority for the name must be given to him alone.
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Guilielma Martius, Palm Fam. 21, 1824.
Martius never gave or discussed the reasons why he
decided to create Guilielma as a new genus apart from
Bactris, already known for almost 50 years. He knew that
Kunth had published B. gasipaes from Colombia and it was a
cultivated species, similar to the first Guilielma species
described by himself as G. speciosa (see discussion about
this species).
The adoption of Guilielma as a valid genus in the past
century was, perhaps, due to Martius influence in palm
taxonomy at that time. Karsten (1857) was the only one to
challenge Martius proposition. Wendland (in Kerchove,
1878) transferred several species from Bactris to
Guilielma, but never discussed the reasons why he decided
to do so. Furthermore, his publication is essentially a
list of previously published palm species. Burret (1934),
on the other hand, in his taxonomic treatment of Bactris
decided to keep Guilielma as a segregated genus and gave
data to support it. Sanders (1991) showed, however, that
Burret had found only two reliable characters to maintain
his proposition: a massive stem and an endocarp with adnate
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bands. I consider that massive stems can not be used as a
character to segregate these genera since non-Guilielma
species of Bactris, such as B. setulosa, B. riparia, B.
guineensis and all the Antillean species, can have a
massive stems and were never described under the genus
Guilielma. In following this logic, Astrocaryum, a closely
related genus to Bactris, would have to be segregated
because it also has species with massive or slender stems.
The endocarp with an adnate band is characteristic almost
unique to all species originally described as Guilielma,
but it can be also found in some specimens of B. setulosa
and B. riparia. It would be very difficult to justify
segregating Guilielma on the basis of only one good
character.
It is very common to find in papers, especially those
by agronomists (Mora-Urp, 1984; Clement, 1988; Silva,
1994) suggestions that the controversy about the validity
of Guilielma is not resolved, and that the "modern
tendency" is to accept this genus as a synonym of Bactris.
In my opinion Sanders (1991), has exhausted this
discussion, and I cannot see reason for any further
suggestion of Guilielma being a segregated genera. Mora-
Urp (1984) presented a incorrect discussion of the pupunha
taxonomy. I think Mora-Urp's publication deserves some
comments to prevent unaware researchers, especially those
unfamiliar with taxonomy, for being confused by wrong
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concepts regarding Bactris. First: not all species of
Bactris have a staminoidal ring. This character is found
in only a small group of black-fruited species (the
Pyrenoglyphys group). There are Bactris species not
closely related to the Guilielma clade without a
staminoidal ring and variously colored fruits, such as red,
yellowish and even black fruits. Second: crispate leaves
are throughout Bactris, and are not related to a specific
group of species.
Guilielma speciosa Martius, Hist. Nat. Palm. 2: 82.
1824. Bactris speciosa (Martius) Karsten, Linnaea
28: 402. 1857. Type. Brazil. South of Par, n.d.,
Martius s.n.(Type at M, not seen).
This was the first species described under Guilielma.
Martius considered B. gasipaes Kunth as synonym of this
species, in a clear transgression of the nomeclatural
rules. Spruce (1871) commented that speciosa was not the
epithet for this species, and Guilielma was
indistinguishable from Bactris. Martius' G. speciosa is a
nomen illegitimun according to article 63.1 of the ICBN
(Bernal, 1989). If B. gasipaes Kunth was placed as a
synonym of G. speciosa, in order to follow the Code,
Martius should have made the combination G. gasipaes, a
correction proposed by Bailey (1930) more than 100 years
later.
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Karsten (1857) transferred G. speciosa to Bactris when
he described his variety chichagui. Karsten understood
that all species without androecium structures in the
corolla of the pistillate flowers should have been placed
under Bactris. Therefore, if B. speciosa (Mart.) Karsten
is taken alone, it is a superfluous name (as implied by
Wessels Boer, 1965; 1988), because G. speciosa was
described with B. gasipaes as a synonym, and two names can
not be used to represent a single taxon.
G. speciosa represents the eastern Amazonian
population of the cultivated pupunha and is closely related
to the small-fruited B. insignis (G. insignis) from
Bolivia, according to Mora-Urp (cited in Clement, 1988).
Bactris speciosa (Martius) Karsten var. chichagui
Karsten, Linnaea 28: 402. 1857. Guilielma gasipaes
(Kunth) L. H. Bailey var. chichagui (Karsten)
Dahlgren, Field Mus. Nat. Hist. Bot. 14: 185, 1936.
Type. Colombia. Magdalena and Cauca. (Type
destroyed?).
As commented above, Martiu's G. speciosa is a nomen
illegitimun. Should Karsten's var. chichagui be considered
also a nomen illegitimun? No according to the article 68.2
of the ICBN.
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Guilielma speciosa Martius var. flava Barb. Rodr.
Enum. Palm. Nov. 23, 1875. Guilielma gasipaes
(Kunth) L. H. Bailey var. flava (Barb. Rodr.) L. H.
Bailey, Gentes Herb. 2: 187, 1930. Lectotype
designated by Henderson (1995): Barb. Rodr., Sert.
Palm. Brasil. 2: t. 52b. 1902.
Barbosa Rodrigues was the first to propose a
classification of the cultivated varieties of pupunha.
However his system was never adopted because it is based on
fruit color, which is not a good character to segregate
cultivated traits of pupunha (Mora-Urp, 1984). As implied
by the name, this variety is characterized by its yellowish
(gold color) fruits.
Guilielma speciosa Martius var. coccinea Barb.
Rodr., Enum. Palm. Nov. 23, 1875. Guilielma gasipaes
(Kunth) L. H. Bailey var. coccinea (Barb. Rodr.) L. H.
Bailey, Gentes Herb. 2: 187, 1930. Lectotype
designated by Glassman (1972): Barb. Rodr., Sert. Palm.
Brasil. 2: t. 52c. 1902.
This variety has red fruits.
Guilielma speciosa Martius var. mitis Drude. in
Martius, Fl. Brasil.: Cyclanthaceae et Palmae I, Fasc.
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85, vol. 3(2): 363. 1881. Type. Brazil. Rio de
Janeiro, cultivated, 26 Dec 1887, A. Glaziou 17342.
(Type: P, not seen)
This variety was proposed by Drude using as the main
character to segregate it the lack of spines in the stems.
It seems Drude was trying to complete the range of
varieties initially proposed by Barbosa Rodrigues, who
never described any spineless variety of pupunha. It is
worth noting that Glaziou collected the type specimen in
Rio de Janeiro from cultivated plants, probably from
material originally imported from the Amazon.
Guilielma speciosa Martius var. ochracea Barb. Rodr.
Vellosia 1, ed. 1: 40, 1888. Guilielma gasipaes
(Kunth) L. H. Bailey var. ochracea (Barb. Rodr.) L. H.
Bailey, Gentes Herb. 2: 187, 1930. Lectotype
designated by Glassman (1972): Barb. Rodr., Sert. Palm.
Brasil. 2: t. 52d. 1902.
This variety was described as having a dark-yellowish
color.
Guilielma insignis Martius, Palm. Orbign., in
d'Orbigny, Voy. dans l'Amer. Mr. 7(3): 71-73. 1847.
Bactris insignis (Martius) Bailon, Hist. Pl. 13: 305.
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1895. Type. Bolivia. Beni. Moxos, s.d., d'Orbigny 18
(Holotype at P, not seen)
Guilielma insignis was found by Alcides d'Orbigny in
several localities of Beni and Santa Cruz, in Bolivia.
Martius (1847) gave a detailed description for this
species, but the accompanying illustrations he presented
are not precise. The leaves in plates X and XXIX are drawn
as being in the same plane, a character not found in
Guilielma. The fruit, although showing the yellowish color
common to many of the cultivated pupunha, has an obovoid
shape, conflicting with his description "magnitude et
figura ovi gallinacei minoris."
Saldias (1991) collected and studied several specimens
of pupunha, either cultivated, semi-cultivated and growing
wild from Santa Cruz, Beni, Cochabamba, and La Paz, in
Bolivia, places that were partially visited by d'Orbigny.
He was able to identify B. gasipaes and B. dahlgreniana,
but not B. insignis (=G. insignis). From his data it is
possible to understand why he could not find G. insignis:
it has a fruit similar in size and shape to the smaller
cultivated pupunha that belongs to the microcarpa or
mesocarpa landraces proposed by Mora-Urp and Clement
(1989). Therefore, B. insignis is similar to B. gasipaes,
but its fruits are at the low range of size.
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Guilielma utilis Oersted, Vidensk. Meddel. Kjoeb.
1858: 46, 1859. Bactris utilis (Oersted) Benth. &
Hook. ex Halm. in Goodman & Salvin Bio. Cent. Amer.
Bot. 3: 413. 1885. Type. Costa Rica, n.d., Oersted
s.n. (Type at C, not seen).
This was the only Central American taxon described as
a closely related species to B. gasipaes. However, since
it is known that pupunha is believe to have been introduced
to Costa Rica as early as 2,300 BC (Mora-Urp, 1994),
Oersteds species is in fact a cultivated pupunha
(mesocarpa landrace sensu Mora-Urp & Clement, 1989)
introduced to Central America. Therefore, this species is
a synonym of B. gasipaes Kunth.
The epithet utilis has been applied in different ways
by agronomists. Mora-Urp (1994) uses it to describe one
of the four western varieties of the pupunha mesocarpa
landrace. Clement (1995), on the other hand, lumped all
known pupunha ladrances into two subspecies of B. gasipaes
(?) naming one of them utilis. The other subspecies is
called speciosa and includes an undescribed and unpublished
variety of Karstens G. chontaduro Triana var. chichagui
known locally in Colombia as "chinamato."
Bactris caribaeaa Karsten, Linneae 28: 403. 1857.
Guilielma caribaeaa (Karsten) Wendland in Kerchove,
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Palm. 246, 1878. Type. "Crescit ad pedem montis
glacialis St. Marthae in planitie sicca et calida
Maracaibensis et St. Marthensi." No date, Karsten
(?)(Type at LE?, not seen).
Dugand (1940) suggested Karstens description of the
type locality does not make geographic sense. He also
suggested, perhaps based on his field experience in that
region, that B. caribaea should be placed as a synonym of
G. macana (=B. macana). Clement (1988) decided to follow
Burret (1933-1934) and Glassman (1972) by recognizing both
species, detracting Dugands proposition of synonymy. It
is necessary, however, to make clear that Clement (1988)
apparently did not understood Karstens (1857) reasons for
describing a "true" Guilielma species under the name Bactris
as was the case for this taxon (see discussion of Guilielma
of Karsten).
Guilielma chontaduro Triana, Nuev. jen. i spec. Neo-
Gran. 15, 1854. Type. Colombia. "Entre Pedra de Moler
i Cartago" (Colombia). Type. Not designated. Guilielma
gasipaes (Kunth) L. H. Bailey var. chontaduro (Triana)
Dugand, Caldasia 1: 63, 1940.
Dugand (1976) gave additional data regarding the type
locality for this species, found in the Cauca River valley
and Caldas. Triana's fruit description says: "fructus
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aurantiaci, flavi, velcorallini, magnitudine Guilielma
speciosa, pericarpo carnoso eduli crasso." Therefore, it
is possible to conclude that this is a small-fruited and
cultivated pupunha. Clement (1988) placed this species as
a synonymy of B. gasipaes. Mora-Urp (1994), on the other
hand and without further discussion, considered B.
chontaduro (=G. chontaduro?) as one of the species that
gave rise to the cultivated pupunha. Perhaps the common
name of Trianas species "chontaduro de los naturales" or
the high altitude at which it was found (1,200m) led
researchers to think about this possibility. It is worth
to note that B. chontaduro proposed by Mora-Urp is a name
found nowhere in the literature, and can not be used,
unless validly published.
Guilielma macana Martius, in d'Orbigny, Voy. dans
l'Amer. Mr. 7(3): 74. 1847. Bactris macana (Martius)
Pittier, Man. Plant. Usual. Venz., 276. 1926. Type.
Venezuela. Zulia, Maracaibo, n.d., Ple s.n. (Type at
P?, M?, not seen).
This species was described from the Maracaibo region
in Venezuela, which is close to the Colombia border.
Karsten (1857) described his B. caribaeaa from plants
collected in an unknown place near the same type locality
of G. macana (either at the Colombian or Venezuelan side of
the border). Although Karsten's species description is far
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more complete than that of Martius, the data presented by
the later is within the range of Karsten's description,
making it clear that they had described similar species.
Pittier (1926) transferred G. macana to Bactris and
commented that it was a poorly known species. Perhaps the
lack of an illustration for Martiuss (and also Karsten's)
species contributed to this situation.
Guilielma matogrossensis Barb. Rodr., Palmae
Matogross. 33. 1898. Lectotype designated by
Glassman (1972): Barb. Rodr., Sert. Palm. Brasil.
2: t. 46b. 1903.
This species was found by Barbosa Rodrigues (1898)
without flowers and fruits, in the "virgin forests of the
morro do Capito-mr, at the margin of Rio da Casca,
tributary of Rio Manso, itself a tributary of Rio Cuiab,
in Serra da Chapada, Mato Grosso," located 70-90 kilometers
north of the city of Cuiab. "It is found deep in the
forest, usually solitary and far from each other, reaching
the forest canopy." From his description it is clear that
Barbosa Rodrigues definitely saw a species matching the
main vegetative characters of the Guilielma sensu Martius:
massive stem (10-12 m long and 12 cm in diameter), long
internodes (30 cm long at the base) fully covered by black
spines, plumose leaves, with sheath, petiole, and rachis
covered by a whitish tomentose layer. The fruits,
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according to the information Barbosa Rodrigues obtained
from his field assistant, were red like "pitanga," globose,
and no more than 2 cm, a size much smaller when compared to
the cultivated B. gasipaes and even the small-fruited B.
insignis from Bolivia. Based on this information, Barbosa
Rodrigues suggested G. matogrossensis could be the ancestor
of the cultivated pupunha, arguing that the seeds may have
migrated through the Madeira and Tapajs Rivers to the
Amazonas River, and there evolved into the cultivated
pupunha.
When Barbosa Rodrigues renamed his G. matogrossensis
as B. coccinea (see discussion under B. riparia), it was
generally accepted that G. matogrossensis was really a
Bactris sensu strictu (Clement, 1988; Henderson, 1995).
However, my opinion is that G. matogrossensis is de facto a
wild pupunha from the western Amazon, similar to the later
published G. microcarpa Huber (=B. dahlgreniana), also
found in Rondonia, Brazil, by Clement et al. (1989) and
Santa Cruz, Bolivia, by Saldias (1991). A collection from
the type locality of G. matogrossensis is needed to clarify
its status as a true member of the wild pupunha species.
Guilielma microcarpa Huber, Bol. Mus. Goeldi 4: 476,
1904. Type. Brazil. Amazonas: Boca do Acre. Type
specimen not designated (Type at MG?, not seen).
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Bactris dahlgreniana Glassman, A Revision of B. E.
Dahlgren's Index of American Palms, 150. 1972.
This species was found by Huber in several localities
near the town of Boca do Acre, Amazonas, Brazil, during his
trip to the upper Purus River in March-May of 1904. Huber
had already found, some years before, similar plants
(although sterile) in the vrzeas of the Chipurana River in
pampa Sacramento, between the Ucayali and Huallaga Rivers,
in Peru. He suggested that the geographic distribution of
this species included a large area, from the Huallaga and
upper Juru River to the Madeira River basin. More
recently Clement et al. (1989) extended this distribution
some 250 km eastward to the Madeira River, to the Ouro
Preto do Oeste region, in Rondnia, Brazil.
Huber (1904), discussed the differences between G.
matogrossensis and G. microcarpa, suggesting that they
could be synonyms. He was the first to propose a hybrid
origin for the cultivated pupunha, with G. microcarpa and
G. insignis as the parents.
Glassman (1972) transferred all Guilielma species to
Bactris. Not being able to use the binomial B. microcarpa
(already used by Spruce, 1871), he renamed G. microcarpa as
B. dhalgreniana.
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The Guilielma species published by Karsten
Karsten (1857) proposed a new system of classification
for some genera in Bactridinae. He emended the description
of Guilielma Martius charact. emend. calyx et corolla
gampophyllus and interpreted it in a completely different
point of view, proposing that species belonging to this
genus should have sterile androecium structures in their
pistillate flowers, and the ones lacking such structures
should be members of Bactris. Karstens proposal would
result in the placement of all previously described
Guilielma species as members of Bactris and he did
published two "true" Guilielma species under the generic
name of Bactris: B. speciosa var. chichagui (a variety of
the G. speciosa Martius) and B. caribaea. Therefore, all
Guilielma species described by Karsten were never intended
to be part of Guilielma Martius, as it has been suggested
by several authors (Mora-Urp and Clement, 1981; Mora-Urp,
1984; Clement, 1988).
Karsten described under the name Guilielma three small
statured species he found in northern Colombia and
Venezuela: G. piritu, G. tenera and G. granatensis. These
species shared basically two common features: pistillate
flowers with sterile androecium structures, either as a jug
or staminoidal ring or as distinct staminodes, and black or
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violaceous fruits, characters never found in the Guilielma
Martius.
Guilielma piritu Karsten, Linnaea 28: 397, 1857.
Bactris piritu (Karsten) Wendland in Kerchove, Palm.
234, 1878. Type area. "in regionibus calidis, planis,
argillosis, Venezuelae et Novo-Granada (Colombia)." No
type specimen designated.
The type locality given by Karsten is vague and a
precise geographic placement of the area referred it is
very difficult. Bactris piritu was considered by Wessels
Boer (1988) to be a synonym of B. guineensis, a proposal
accepted by Henderson et. al., (1995). B. guineensis it is
known today to be distributed in the llanos of northern
Venezuela and Colombia, and along the Pacific coast of
Central America. According to Karsten (cited by Wessels
Boer, 1988), this taxon is distinct from Bactris minor
Jacq. (=B. guineensis) by its pistillate flowers and stem
and petiole indument, being almost identical for the
remaining characters. Braun (1983, 1995) and Braun and
Chitty (1987) consider B. piritu and B. guineensis to be
distinct species, and their plates show B. piritu to be a
much smaller palm.
Guilielma tenera Karsten, Linnaea 28: 399, 1857.
Bactris tenera (Karsten) Wendland in Kerchove, Palm.
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234, 1878. Type area. Colombia. Rio Magdalena. No type
specimen designated.
When Karsten published G. tenera (1857) he discussed
its relationship with B. brongniartii and B. maraja.
Wendland (in Kerchove, 1878) transferred G. tenera to
Bactris sensustrictu without discussing the reasons for
it, as well as to what group of species it was related.
Burret (1933-1934) included all previously described
species of Bactris with androecium structures in the calyx
and corolla of pistillate flowers in the genus Pyrenoglyphis
Karsten, making the combination P. tenera. Wessels Boer
(1965; 1988) and Galeano and Bernal (1987), however,
maintained B. tenera (=G. tenera) as synonym of B. maraja,
a conflicting proposal since B. maraja was originally
described by Martius without flowers and the fruit
description does not mention the presence of a staminoidal
ring. Recently, Henderson (1995) maintained that B.
brongniartii is a good species, distinct from B. maraja on
the basis of the pinnae shape (linear x sigmoidal) and the
presence of a staminoidal ring, more easily found in the
fruit perianth. As a result, B. tenera was placed as a
synonym of B. brongniartii.
Guilielma granatensis Karsten, Linnaea 28: 400, 1857.
Bactris granatensis (Karsten) Wendland in Kerchove,
Palm. 234, 1878. Type specimen. "Locis umbrosis,
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humidis calidisque Venezuelae et Novo Granatae
[Colombia] habitans." Type specimen destroyed at
Vienna.
The description of the type locality for this species
is also vague and cannot be located precisely in Colombia
or Venezuela. It is considered by Henderson et. al. (1995)
a synonym of B. pilosa Karsten. B. pilosa is a member of a
large group of Bactris that have purple-black fruit lacking
androecium structures in the pistillate corolla. Since B.
granatensis Karsten was originally described as having
these structures, the present placement of this species
under the name B. pilosa does not seems to be an acceptable
decision.
The species of Sanders Antillean clade
In 1991 Sanders proposed a group of Bactris species
found in the Greater Antilles as members of a clade he
named Antillean. Salzman and Judd (1995) revised the
members of this clade and suggested it was formed by three
species, B. cubensis, B. jamaicana and and B. plumeriana,
and one ambiguous name. In both studies the Antillean
clade is considered a sister clade to the species of
pupunha (Guilielma clade) and together they form the Non-
Ocreate clade. The phylogenetic relationship of these two
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clades is supported by several synapomorphies according to
Sanders (1991), but geographically they appear to have
evolved independently from each other.
Bactris cubensis Burret, Kungl. Svenska Akademiens
Handlingar 6(7): 25. 1929. Type. Cuba. Oriente:
Sierra de Nipe, in charrascalles ad Ro Piloto,
29 Jul 1914, Ekman 2286 (Holotype at S, not seen).
This species is restricted to the eastern part of
Cuba, where it usually grows in forested areas. According
to Salzman and Judd (1995), its fruits approach the form of
the small-fruited B. macana.
Bactris jamaicana L. H. Bailey, Gentes Herb. 4: 177.
1938. Lectotype designated by Salzman and Judd, 1995:
Jamaica. St. Catherine Parish: Gilbralter, on
Gibralter Road in pasture of Frank Roper State, 10 Nov
1935, Bailey 216 (Lectotype at BH, not seen).
Bactris jamaicana is restricted to Jamaica where it
grows in sub-montane forest and wet savannas. Salzman and
Judd (1995) considered it the most distinctive of the
Antillean Bactris species. The fruits of B. jamaicana that
I have examined at NY are very similar in shape to those of
B. riparia, and their endocarps have an unusually large
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fertile pore, a character useful to identify the specimens
of this species.
Bactris plumeriana Martius, in d'Orbigny, Voy. dans
l'Amer. Mr. 7(3): 64. 1847. Type. Martinique or
Guadaloupe. Plumier, Tab. 43-45. Nova Plantarum
Americanarum Genera, Paris, Museum, Bibliotheque
Centrale.
Although Martius had expressed doubts if this species
was originally collected in two of the Lesser Antilles
islands, B. plumeriana seems now endemic only to the
Hispaniola. This species grows in forest or disturbed
areas.
Bactris chaetophylla Martius, in d'Orbigny, Voy. dans
l'Amer. Mr. 7(3): 71. 1847. Type. Dominican
Republic. Herb. Vetenatti nunc Webbii.
Martius described this species but placed question
marks on the name. It is probably a synonym of B.
plumeriana, according Salzman and Judd (1995).
Bactris setulosa and related taxa
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Galeano and Bernal (1987) discussed the presence of
adnate and flattened fibers in an unidentified species
(Bactris sp. 2) they collected in Antioquia, Colombia
(Bernal, Galeano & Bolivar, 693). The presence of
flattened fibers adherent to the endocarp is one of the
apomorphies that defines the Guilielma clade proposed by
Sanders (1991). I had the opportunity to examine Bernals
collection at NY and it is, without any doubt, B. setulosa.
This species and closely related taxa are characterized by
their large size, similar to the cultivated pupunha, and
have an Andean distribution, where they can be found up to
1.700 m of elevation, in Ecuador (H. Balslev 4288).
Galeano and Bernal (1987) suggested their unknown species
(=B. setulosa) resembled slightly B. gasipaes and B.
riparia. Henderson et. al., (1995) suggested this species
as related to B. gasipaes.
Bactris setulosa Karsten, Linnaea 28: 408. 1857. Type.
Venezuela. Carabobo: Cumbre de Valencia, Puerto
Cabello, s.d., Karsten s.n. (Type at LE, not seen).
This species is very common at high elevations in the
Andes, but can also be found at low elevations in Colombia,
Ecuador, Venezuela, and Trinidad and Tobago. It has a
series of characters that make easy its identification:
pinnae very wide (sometimes up to 9cm), leaf spines
clustered, spines on the stem arranged in regular rings,
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and fruits obovoid with the perianth usually entire.
According to A. Henderson (pers. com.) the broad pinnae of
B. setulosa is an uncommon feature in Bactris and, when
combined with the clustered spines on the leaves, are the
best characters to identify herbarium specimens of this
species. Two collections from Ecuador (Skov et al. 64.824;
Balslev & Steere 3121) and one from Surinam (Wessels Boer
506) have very distinct staminate flowers, with 9-12
stamens. In Wessels Boers samples seems to have occurred
a labeling mistake because the specimen collected in
Suriname by Wessels Boer (# 506) was supposedly taken from
forest camp, Coppenname Riverside, land floods with high
water, tidal influence, loamy soil, an environment
completely different from that cited for the two remaining
species collected in Ecuador, and unlikely to be typical
for a species belonging to the B. setulosa group.
Bactris cuvaro Karsten, Linnaea 28: 406. 1857. Type.
Colombia. Meta: Villavicencio. (Type at LE, not seen).
This species is poorly known, and was considered
synonym of B. setulosa by Henderson et. al. (1995).
Bactris cuesa Grisebach & H. Wendland in Grisebach,
Fl. British West Ind. Type. Trinidad. Maracas, s.d.,
Crueger s.n. (Holotype at K, not seen).
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B. cuesa is considered a synonym of B. setulosa by
Wessels Boer (1988) and Henderson et. al. (1995). Bailey
(1947) presented illustrations of B. cuesa and B.
sworderiana, which were originally described without
illustrations. From Baileys description it is possible to
conclude they are closely related species. As a matter of
fact, the fruit illustrations are basically the same, with
a slight and insignificant difference in the shape.
Bactris falcata Johnston, Proc. Amer. Acad. Arts 40:
683. 1905. Type. Venezuela. Margarita Island: Ro
Assuncin, 16 Jul 1903, J. Johnston 220 (Holotype at
GH).
The Margarita Islands are adjacent to continental
Venezuela and it would not be unusual to find B. setulosa
growing there. The type specimen of B. falcata consists of
the apex of a young leaf and a small section of the stem,
similar to several specimens of B. setulosa at NY. In
Johnson's description it is clear that the spines on the
stem are arranged in regular rings and the fruits are
broader than long, typical features of B. setulosa.
Bactris sworderiana Beccari, Repe. Spec. Regni Veg.
16: 437. Type. Tobago. Caledonia, s.d., W. E.
Broadway 4014 (Holotype at FI, not seen).
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Wessels Boer (1988) followed Burret (1933-1934) and
considered B. sworderiana and B. setulosa distinct species
on the basis of fruit shape (obovoid x depressed globose)
and the perianth size. All of the samples of plants
belonging to this group that I have examined at NY,
however, have obovoid fruits with or without a flattened
(depressed) apex. This small variation may have led
researchers to give the depressed-globose shape for the
fruits that present the flattened apex. Perianth size in
palm fruits are known to show some degree of variation when
they are not fully developed.
Beccari's description also has many of the typical
characters of B. setulosa. He suggested that B. sworderiana
has some differences form B. falcata, including broader
pinnae. This is irrelevant since in Johnson's description
of B. falcata a measure for the "segment intermediis" was
not given. Therefore Beccari's comparison was made with
the lower pinnae in Johnson's description, which are only 3
cm wide x 4 cm of the intermediate pinnae described by
Beccari. Henderson et. al., (1995), also considered this
species a synonym of B. setulosa.
Bactris kalbreyeri Burret, Repe. Spec. Regni Veg.
34: 231. 1934. Type. Colombia. Antioquia: Guadalito,
670-1000 m, 7 Feb 1880, W. Kalbreyer 1394 (Holotype
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destroyed at B, Holoneotype at COL, designated by
Bernal et al., 1989, not seen).
This species is considered synonym of B. setulosa by
Henderson et. al., (1995). Burret (1933-1934) had
originally placed his own species as incertae sedis,
because he did not have fruits or flowers to make a
complete description. To make the situation worse, the
type collection of this species was destroyed in Berlin
during the Second World War and Burrets description does
not show any illustration for the species. In the original
description the number of pinnae on each side of the leaf
is only 23, a value far below the 40-65 accepted by
Henderson et. al., (1995) for B. setulosa. Bernal et al.
(1989) designated a Holoneotype for B. kalbreyeri and
admitted they failed to trace the original place of
collection in Guadalito or Guadualito. With so much
missing information, it would be better to consider this as
incertaesedidspecies, as Burret did.
Bactris circularis L. H. Bailey, Gentes Herb. 7: 388.
1947. Type. Trinidad. St. Andrews: Melajo, 26 Feb
1946, L. H. Bailey 158 (Holotype at BH, not seen).
This species was decribed and illustrated by Bailey in
his Palmae Indigenae Trinitensis et Tobagensis (1947).
Wessels Boer (1988) comments that Bailey's B. circularis is
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a synonym of B. setulosa because the spine rings on the
stem can not be used to identify specific differences.
Bactris bergantina Steyermak, Fieldiana Bot. 28 (1):
71. 1951. Type. Venezuela. Anzategui: along Ro Leon,
northeast of Bergantin, 500m, 20 Feb 1945, J.
Steyermark 61039. (Holotype at F, not seen).
Steyermark (1951) comments that this species is
separated from the closely related B. cueso and B.
sworderiana by the flattened or nearly flattened spines on
the rachis of the petiole and the scales on the lower
surface of the pinnae. These are non-reliable characters
and should not be used to segregate species in Bactris. In
almost all other characters Steyermark's species is similar
to B. setulosa, including the clustered spines on the leaf
and the broad pinnae (4 cm wide). B. bergantina is
considered a synonym of B. setulosa by Wessels Boer (1988)
and Henderson et. al. (1995).
Bactris riparia
Like the species of B. setulosa, some herbarium
specimens of B. riparia have an endocarp with fiber
flattened and adnate to the endocarp, similar to those
found in pupunha. B. riparia and the species considered to
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be synonym of it are of a large size, although usually
smaller than pupunha. The fruits are depressed-globose and
reddish. The crown of leaves is a feature that makes them
very similar to the wild and cultivated pupunha, as noticed
by Martius in the original description. Its geographic
distribution overlaps that of the cultivated and wild
pupunha in the lowlands of the Amazon basin. Henderson
(in prep.) propose it as close to B. gasipaes.
Bactris riparia Martius, Hist. Nat. Palm. 2: 97. 1826
Type. Brazil. Amazonas: Rio Japur, s.d., C. Martius
s.n. (Holotype, M, not seen).
Martius described this species without fruits and
noticed similarities with pupunha in the vegetative habit
"habitu fere frondium Guilielma speciosa." As implied by
its name, this is a species frequently found close to the
margin of lakes, Rivers, and small streams, having a
widespread distribution in the lowlands of the Amazon. In
Colombia (Dugand, 1940) it is known as pupua-brava.
Bactris inundata Martius, in d'Orbigny, Voy. dans
l'Amer. Mr. 7(3): 58. Type. Bolivia. Beni: Moxos,
s.d., A d'Orbigny 24 (Holotype at P, not seen).
Alcides d'Orbigny found this species in Beni, Bolivia,
and in the adjacent Mato Grosso, Brasil. The fruits were
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described and an illustration provided (Pl. 27b, 6, 7 and
8). Herbarium specimens of plants collected in central
Amazon (the type locality of B. riparia) are similar to
those found in Bolivia and Mato Grosso, especially the
fruits, the habit, and the staminate flowers. B. inundata
is considered a synonym of B. riparia by Henderson (1995).
Bactris littoralis Barb. Rodr., Enum. Palm. Nov. 36.
1874. Type area. Brazil. Amazonas: cercanias de
Parintins. Lectotype (designated by Glassman, 1972):
Barb. Rodr., Sert. Palm. Brasil. 2: t. 32a and 33.
Barbosa Rodrigues species is similar to Martius B.
riparia not only in the vegetative and reproductive traits,
but also in the habit.
Bactris coccinea Barb. Rodr., Contr. Jard. Bot. Rio
de Janeiro 4: 110. 1907. Lectotype designated by
Glassman (1972): Barb. Rodr., Contr. Jard. Bot. Rio
de Janeiro 4: t. 24c.
When Barbosa Rodrigues described Guilielma
matogrossensis (1898) he did not see its fruits, but only
young and closed buds. He arranged to have the mature
fruits sent to him in Rio de Janeiro as soon as they became
available. However, seems it took almost eight years for
him to get the fruits. After he examined them, he notice
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they were not of Guilielma type and decided to transfer his
G. matogrossensis from Guilielma to Bactris. Since he
could not use the binomial Bactris matogrossensis (he has
used already it for another species) he applied the name B.
coccinea. From Barbosa Rodrigues drawings it is possible
to conclude the fruits he examined in 1907 were similar of
B. riparia. This is corroborated by the fact that this
taxon can be found in Mato Grosso and in the adjacent
Bolivian territory, as reported by dOrbigny in the notes
that follow the description of B. inundata (=B. riparia).
Henderson (1995) consider B. coccinea and B. inundata to be
synonyms of B. riparia.
From Barbosa Rodrigues description this species is a
mix between the vegetative features of a Guilielma taxon
(G. matogrossensis) and the fruits of a Bactris (strictu
sensu) species.
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CHAPTER 3
THE ANCESTORS AND PLACE OF DOMESTICATION OF PUPUNHA (Bactris
gasipaes Kunth)
There are disagreements about the origin of the
cultivated pupunha, the parent species, number of times it
was domesticated, and the place were it occurred. Some
authors suggest pupunha is of monophyletic origin (Barbosa
Rodrigues, 1898; Burret, 1933-1934; Clement, 1988;
Henderson, 1995), while a few others states it has a
polyphyletic origin (Huber, 1904; Mora-Urp, 1993). Mora-
Urp (1983) support the hypothesis of multiple
domestication events while Clement (1988) believe in a
single event followed by artificial dispersion. There are
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also a dispute about the area were the domestication
process took place. For single or multiple events the most
frequently cited region includes a large area that follows
the Andes cordilleras, from northern to the Southeastern
South America.
Wallace (1853) had already observed "this palm appears
to be indigenous to the countries near the Andes, on the
Amazon and Rio Negro it is never found wild." Spruce
(1871), who traveled from the Amazon River mouth to the
Andes, comments when he asked local people in central
Amazon about the origin of pupunha, they pointed westward
and said "From the Cordilleras."
Barbosa Rodrigues (1898), who also worked in the
Amazon valley and had found only cultivated pupunha,
believed the wild pupunha could not be found in Brazil but
in some area near the Andes of Bolivia and Peru. He was
surprised when he found his G. matogrossensis near the city
of Cuiab, in an area relatively close to the type locality
of the small fruited and cultivated B. insignis, from the
adjacent Bolivia. Barbosa Rodrigues species was apparently
a primitive type of the cultivated pupunha, and he
hypothesized the seeds of G. matogrossensis could have
migrated throughout the Madeira and Tapajs River, reaching
the Amazonas River, where they acclimated, acquiring along
the time the features of the cultivated plants he had found
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there. Barbosa Rodrigues G. matogrossensis was,
therefore, the first westward wild pupunha described.
I do not discard Barbosa Rodrigues hypothesis about
seed migration in the Tapajs River. The fact only
mesocarpa and macrocarpa landraces are found along the
middle course of the Solimes River (Mora-Urp and Clement,
1989), has put a question mark over the origin of the
microcarpa landrace found in Par, originally described by
Martius as G. speciosa. Mora-Urp (1984) suggested this
microcarpa landrace as being closely related to the
cultivated B. insignis Martius, from Bolvia, and believe
their seeds had been carried through the Mamor-Madeira
River (Bolvia) to the Amazon in Brazil.
Huber (1904) was the first to suggest the pupunha had
a hybrid origin, with G. microcarpa and G. insignis being
the parent species. This would explain the variations
already identified on the cultivated plants, such as fruit
size, color, mesocarp composition sometimes oily or
starchy, and the reduction or the spineless states found in
some plants. The most striking feature he discussed,
however, was the seed abortion (seedless fruits), perhaps
suggesting hybrids could not produce fertile descendent.
Huber did not accept the natives as being capable of
selecting plants because, as he states, they practiced a
primitive agriculture. He thought the appearance of
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"accidental" hybrids in the dispersal zones of the parents
species, located someplace in the upper Purus, or Beni or
Mamor Rivers basin, gave the opportunity for the natives
to choose the improved ones for cultivation, and later
dispersal to the north and eastern South America.
Clement et al. (1989) studied the phenotypic variation
of two populations of B. dahlgrenian from Acre and
Rondonia, in the southwestern Amazon of Brazil, and
compared them with one population of the mesocarpa landrace
Pastaza from Ecuador and another possible B. dahlgreniana
population, from Peru. Their results suggest B.
dahlgreniana could be the ancestor of the cultivated
pupunha because this species is vegetatively similar to the
cultivated pupunha and differences are present only in the
vegetative parts. Although they have studied only the
fruits, a tendency towards the increment of infructescence
weight and a decrease of fruit number was identified. This
trend would be expected in the case pupunha was
domesticated from B. dahlgreniana by human selection.
Although Clement et al. (1989) do not state conclusively
that B. dahlgreniana is the ancestor of pupunha, their
results match Spruce's (1871) expectations about how the
wild pupunha would look like: "with fruits so much smaller
and drier than what it has become by long cultivation as to
be not easily recognizable."
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Mora-Urp (1984) suggested different kinds of
cultivated pupunha had distinct origin. Some are simple
wild plants poorly improved, while others had hybrid
origin. He conclude saying pupunha does not have a common
center of origin since it is native to an extensive region
and the domestication process took place several times
along this area, hence, pupunha is a synthetic species.
Mora-Urp (1993, 1994) gives a list of the nine species he
believe are related to the multiple origin of pupunha.
Four of these species were validly described in the past as
B. ciliata, B. insignis, B. macana, and B. caribaea. Four
are undescribed species and one, B. chontaduro, is a name
not found in the literature. According to this author
pupunha evolved independently from each of these nine
species around their type locality. He comments the
ancestor for the macrocarpa landrace Vaups, which has the
biggest fruit among the cultivated pupunha, is still to be
described since all the wild pupunha found to date have
only small fruits.
To follow Mora-Urps logic one has to admit, e. g.,
that B. macana and B. caribaea must have been used by
natives in the Northern South America as a parent species
of some cultivated trait of pupunha. Considering they have
such small fruits (the smallest among the wild pupunha),
the expected cultivated variety derived from them must had
to had small fruits, and would be classified as a
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microcarpa landrace type. However, Mora-Urp (1993) map of
distribution of the cultivated varieties of pupunha in
South America does not shows any cultivated landrace in
northern South America that overlaps the type locality of
the wild B. caribaea and B. macana, and this is a clear
indication that domestication of pupunha was not developed
in that area.
Several authors (Galeano and Bernal, 1987; Henderson
et. al., 1995; Henderson, 1995; Henderson, com. per.) have
suggested B. setulosa and B. riparia as new members of
the pupunha group on the basis of characters I already
discussed. These two species cannot be considered to have
a direct relationship to the cultivated pupunha, as are the
cases of the wild species, such as B. macana and B.
dahlgreniana, but may have been hybridizing with them,
wherever they grow allopatrically, as suggested by Clements
(1995). This author discussed phylogenetics relationship
in pupunha using Harlan's & Wet's (1971) gene pool concept.
He considers all cultivated pupunha as the primary gene
pool (GP-1), while the wild species and spontaneous
populations of pupunha are classified as the secondary gene
pool (GP-2). The GP-2 species are known to hybridize
freely with the cultivated B. gasipaes when found growing
close to it (Mora-Urp, 1984, 1994) All the remaining
species of Bactris are suggested as to represent the
tertiary gene pool (GP-3) and may hybridize naturally with
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pupunha. I think only a partial number of Bactris species
could be considered as member of the GP-3 proposed by
Clement. This group would include only the orange,
yellowish or reddish fruited species that share common
features of their staminate and pistillate flowers with
pupunha species, such as non-spiny calyx and corolla, and
B. setulosa and B. riparia are members of this group.
SECTION II
THE PHYLOGENY OF PUPUNHA (Bactris gasipaes Kunth) AND
ALLIED SPECIES
CHAPTER 4
TAXONOMIC AND CLADISTICS STUDIES OF Bactris Jacq.
AND THE PLACEMENT OF PUPUNHA (Bactris gasipaes Kunth) AND
RELATED SPECIES.
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The more recent taxonomic study of Bactris and related
genera was published by Burret (1933-1934). In this study
Guilielma Martius was kept segregated and the species of
Guilielma of Karsten, as well as all other Bactris with
androecium structures in the pistillate corolla, were
transferred to the genus Pyrenoglyphys Karsten. The
remaining species were dumped in the genus Bactris sensu
strictu, that was divided into two sub-genus and several
sections. He recognized seven species in Guilielma,
including G. utilis and G. insignis, species obviously
described from samples of cultivated populations of B.
gasipaes. Burret's treatment was a mix of ideas proposed
previously by several taxonomists and the result are
considered a compendium (Sanders, 1991), a catalog of names
(de Nevers et. al., 1996) or a synopsis (Salzman and Judd,
1995), in which 257 species are listed (38 of them
described as new by Burret), and no key or illustrations
were provided.
Using Burret's treatment as a starting point, Sanders
(1991) produced the first cladistic study of Bactris in
which the segregated genera Guilielma and Pyrenoglyphis
were lumped together to form a unique and large genus under
the name Bactris. He studied 139 herbarium specimens that
represented 61 species of Bactris sensustrictu, Guilielma
and Pyrenoglyphis. He scored 106 morphological characters,
although a large number of multistate characters were
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broken down into separate binary characters. Sanders'
result shows that Bactris is composed of four "roughly
equivalent major clades" that do not correspond to any of
Burret's segregated genera. Regarding the Guilielma
species, Sanders used samples that represented only a
fraction of the species accepted by Burret (B. gasipaes, B.
dahlgreniana and B. macana), and they formed a small clade
he named Guilielma. This clade alone is equivalent to the
genus Guilielma Martius and in Sanders analysis it was
placed together with the Antillean clade to form one of the
major clades of Bactris, the Non-Ocreate clade.
From Sanders results (Fig. 5), the clade Guilielma
diverges from Bactris only as a member of the Non-Ocreate
clade and can not be segregated as the genus Guilielma
Martius alone without all the other equivalent clades being
equally considered. In this case, according to Sanders,
Bactris would be fragmented into eight or more poorly
defined genera.
Sanders was criticized because there is a difficulty
in interpreting the characters he used in his analysis (de
Nevers et. al., 1996). Hanh (1993) suggest the
shortcomings of Sanders study were due to the fact he has
"relied on original species descriptions for character
state assessment, redundancy of true characters,
multistates characters broken down into separate binary
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characters, and considerable homoplasy in most characters."
From my point of view all these problems are related to the
fact he did not have the access to a good monograph of
Bactris and used a very small number of specimens while
searching for the most reliable characters, giving a
generalized impression he was unable to cover successfully
all the morphological variation present in Bactris. In
some cases he did not have the opportunity to examine
samples of common and widespread species, as was the case
of B. riparia and this may explain the fact the placement
of some species and even clades (Balonophora) in Sanders
analysis are presently disputed (de Nevers and Henderson,
1996). Besides these problems, Sanders results are of
good resolution regarding the monophyly of Bactris and this
has finished the dispute about the validity of Guilielma
as a good genus.
Salzman and Judd (1995) reviewed the species of the
Antillean clade proposed by Sanders and studied its
phylogenetics relationship with the sister clade Guilielma.
In their cladistic analysis they used morphological and
anatomical characters, but the data used to assess the
characters states of the Guilielma clade were partially
obtained from several bibliographical sources, while most
of the material used for the study of the Antillean species
were collected by them in the field. They also did not
specify the species members of the clade Guilielma they
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used. The results (Fig. 06), confirmed Sanders hypothesis
about the relationship of the Guilielma and the Antillean
clade as being sister clades.
Henderson (1995) treated only the Bactris found in the
Amazon and did not recogniz Guilielma as a valid genus.
According to him there are only two pupunha species growing
in the Amazon basin. The domesticated B. gasipaes in which
he include as synonym several varieties, G. speciosa and G.
insignis, and the wild B. macana, with G. microcarpa under
it. The key he provided places B. setulosa in the same
group of pupunha on the basis of stem size.
CHAPTER 5
THE AGRONOMISTS SYSTEM OF CLASSIFICATION FOR THE
CULTIVATED AND WILD SPECIES OF PUPUNHA.
Mora-Urp and Clement (1989) have traveled extensively
in the Amazon basin, Andes and Central America, analyzing
different pupunha populations and collecting fruits for
germplasm banks. They have proposed (Clement, 1988; Mora-
Urp, 1994; Mora-Urp and Clement, 1989) a non-taxonomic,
but practical and efficient system to classify the
cultivated species of pupunha, including the wild plants
they believe are related to its domestication process.
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Their system is mainly based on the fruit size. They have
divided all populations of cultivated and wild pupunha in
the occidental group, when found West of the Andes, and
oriental, when found at the East. So far they have
classified the cultivated and wild pupunha in three major
landraces (they do not use the term variety in agronomic
sensu), named microcarpa, mesocarpa and macrocarpa. They
have been studying several populations in the field and
each of them is characterized and classified into one of
the major landraces. There are 17 studied populations,
named as cultivated variety by Mora-Urp (1994) (Table I)
or landraces by Clement (1995).
Cultivated and wild populations of pupunha are known
to hybridize freely when in contact (Mora-Urp, 1984) and
it may be the explanation for the high diversity of the
species in character such as fruit color, size and content,
spines presence and plant size. Being now widely
cultivated in farmlands, and with an intense process of
seed exchange taking place, the occurrence of germplasm
contamination, landraces hybridization and genetic erosion
are the most likely results. A possible impact of this
process at long term would be the ruin of Mora-Urp and
Clement's system of classification.
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CHAPTER 6
A CLADISTIC ANALYSIS OF PUPUNHA (Bactris gasipaes Kunth)
AND RELATED SPECIES: A MORPHOLOGICAL AND ANATOMICAL APPROACH
Introduction
This analysis was performed using morphological and
anatomical data and the new information is expected to
contribute to a better understanding of two important
aspects related to the taxonomy and history of pupunha: (a)
the phylogenetic relationship within the domesticated and
wild group of pupunha, and their relationship with the
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species suggested to be closely related to them, and (b)
the possible place were the domestication event of the
cultivated species was developed. The hypothesis is that
cladistic analysis can be used to elucidate taxonomic
problems at species level and at the same time the results
of the phylogenetic reconstruction would allow a more
precise definition of the place were domestication events
occurred.
Material and Methods
A total of seven Bactris species were used in the
present analysis (Table II). The two Desmoncus and two
Astrocaryum species were chosen as the source of
information for building the outgroup data set. I decided
to follow Henderson (1995) and Henderson et. al. (1995)
taxonomic arrangement for the species of pupunha. The
cultivated species will be represented by B. gasipaes and
the wild species by B. macana. This means that all
herbarium specimens belonging to these species found at NY
will be sorted into two distinct piles and the further
anatomical and morphological surveys of each group will
show if there is any necessity to subdivide them before
completing the data matrix and running the cladistic
analysis. The same procedure was applied to B. setulosa
and B. riparia specimens. For the Antillean species I
followed the recent treatment of Salzman and Judd (1995).
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The cladistic analysis was performed with an IBM-
compatible computer, using Wagner parsimony method to
analyze the characters. The programs Nona 1.1 (Goloboff,
1993) and Hennig86 1.5 (Farris, 1988) were used to generate
the trees. In Nona the commands Hold100, Hold/20 and
Mult*20 were used to search for the trees. The trees were
saved in Nona(sv* command), retrieved and generated in
Hennig86. Dada 1.0 (Nixon, 1995) and Clados 1.1 (Nixon,
1992) were used to examine and manipulate the data matrix
and the trees.
The outgroup comparison was used to asses the polarity
of the characters (Stevens, 1980; Watrous & Wheeler, 1981;
Madison et al., 1984). The outgroup used to assess the
polarity of the Bactris species analyzed here was
Desmoncus. In the cases when Desmoncus was too variable or
the character is considered to be an autapomorphy (stem
habit, modified leaves, e.g.), characters states were
obtained from Astrocaryum. Sanders (1991) and Salzman and
Judd (1995) discussed in more details the aspects of choice
of outgroup when cladistic analysis of Bactris were
performed.
The characters used in the analysis are mostly
qualitative and were searched among the outgroup and the
in-group taxa. Quantitative characters were used only when
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a clear gap or discontinuity were detected. Multistate
character were considered to be unordered. Thirty four
morphological and anatomical characters were selected in
the analysis of the nine terminal taxa resulting from the
anatomical and morphological surveys (Table III). The
plesiomorphic or ancestral state is coded as 0 and the
apomorphic condition as 1, 2, or 3, in the cases of the
multi-states characters. The data matrix is presented in
the Table IV.
Only herbarium specimens were used during the present
study. One hundred and fifty six collections from the NY
herbarium were examined in a morphological and anatomical
survey of characters (Table V). Each survey involved the
use of a specific number of herbarium specimens, as well as
specific methods. The morphological survey used only the
usual and conventional methods and a short description of
it is given. On the other hand, the anatomical survey
required a longer time and the use of several techniques to
prepare multiple samples that were used in determining the
more reliable anatomical features that were incorporated as
characters state in the computerized analysis. A detailed
account of the anatomical survey is given for two reasons.
The first is to allow future replication of the methods
here employed. The second is derived from the difficulties
I had in collecting all the necessary methods and
protocols, which are scattered over several different
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publications and not always were developed to be used
specifically with palm samples, requiring several
modifications.
Morphological Survey
Morphological observations were performed throughout a
careful examination of 156 herbarium specimens (table V).
A stereomicroscope Bausch & Lomb, with magnification of up
to 30x was used to study the comparative data for
vegetative, floral (staminate and pistillate) and fruiting
structures. All flowers and fruits were rehydrated prior
to their examination.
Anatomical survey.
Leaf clearing
The procedure recommended by Martens and Uhl (1994)
was mostly followed and minor changes were made (see
Appendix 01). The samples used are listed in Table V. All
procedures were done in small glass vials. Some samples
required a longer time in 5% NaOH (in a oven at 58C), and
in some cases up to one week. When the solution became
very dark it was changed for a new one, in most cases after
the first 24 hours. The samples were washed in deionized
water (three times, 15-30 min. each), and placed in one-
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third strength commercial bleach, for a period of up to 30
min. After being washed again in deionized water, they
were stained in aqueous Safranin (0.5%) for a minimum
period of 30 min. or up to overnight. The samples were
washed and dehydrated for final mount using kleermount.
Flower Clearing
A total of 3-4 flowers (staminate and/or pistillate)
were cleared for each specimen sampled. The specimens
and flowers types cleared are listed in the Table V, and
were cleared according to the protocol described in the
Appendix 02.
The flowers were dissected only after the staining
procedure, immediately before the slide preparation. This
was necessary in order to avoid the loss of floral parts
during the process, that require many changes of solutions.
Some flowers with thick petals, such as B. gasipaes, B.
macana and B. setulosa, required a longer time in 5% NaOH
and resulted very difficult to handle due to the excessive
maceration. The tannin rich staminate flowers of B.
setulosa 2 (W. Boer 506, H. Balslev and Steere 3121 and F.
Skov et. al. 64.824) required to two changes of the NaOH
solution.
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The long washing time required after the bleach
treatment(at least 30 min., step 7 of the protocol) was
necessary to ensure good water infiltration and complete
bleach removal. If the bleach is not completely removed
from the flower tissues, the staining procedure with
Safranin will not work properly.
Leaf Anatomy
Epidermal peels maceration, leaf clearings and cross
section were the techniques used in this part of my study.
The species and specimens sampled are listed in Table V.
Sampling
For the accuracy and uniformity of data collection,
only the middle portion of leaflets located along the
middle part of the leaves were sampled. Cross-section
samples included the midrib and one leaflet margin, and
measured 0.5-1 cm wide. Sample for epidermal peels were 1
cm and required one untrimmed leaflet margin. For
clearings, a 3 cm long section of leaflet with midrib and
one of the margins was used. In all cases one or two
additional spare sample was taken, rehydrated and kept in
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Glycerin-Alcohol ready to be processed in case of material
loss.
The sampling procedure was done using a scissors and
most of the samples for cross section and epidermal peels
were trimmed at this point. However, if the material was
too brittle, cutting was difficult and usually resulted in
longitudinal fissures on the lamina and/or midrib
partition. To avoid this problem a larger piece of the
leaflet (4-5 cm long) was taken, and the samples trimmed
only after the rehydration procedure. When necessary, the
cross section samples taken from B. setulosa (which has the
widest leaflet) were subdivided in order to included the
midvein.
Rehydration and fixing
The rehydration solution used was that recommended by
Martens and Uhl (1984). Samples were rehydrated overnight
or up to 24 hours in a vacuum chamber (12 p.s.i.) and
finally washed in three changes of deionized water (15
minutes each). They were fixed in FAA (formalin, acetic
acid, 50% ethanol) for 48 hours.
Epidermal peels
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The methodology of O'brien and MaCully (1981) was
chosen, following the recommendations of Pinheiro (1997).
The samples used are listed in Table V. Small glass vials
were used during most of the procedure (see Appendix 03).
The samples were placed in Jeffrey's solution, in half-
filled glass vials. The epidermis were easily separated
after 4-5 days and no additional cleaning with paintbrush
was necessary. Only one change of Jeffrey's solution was
made, usually after 24-48 hours. The vials were carefully
shaken once a day during the treatment period, since we
found it helpful for the epidermis and mesophyll
separation. To avoid the loss of samples during the
washing procedure a small glass pipette was used to
exchange the solutions. If the second Jeffrey's solution
was too dark, being impossible to see the samples, water
was added to dilute and clear it, making easy the use of
the pipette and diminishing the possibility of sucking up
the epidermis. The samples were moved to petri-dishes
after the staining step with 0.5% aqueous Safranin, and
processed there until the final slide mounting. Although
recommended, the use of paintbrush to move the sample from
the petri-dish to the slide is a very difficult task,
usually resulting in partial damage to the sample or even
total loss. To avoid it the better solution we found was
to fill up (to the top) a petri-dish with Hemo-De, dip
partially the slide and push the sample over it with the
use of a paintbrush. Some drops of Hemo-De on the slide
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surface is helpful to position the epidermis in the desired
position on the slide surface.
Leaf cross section
The methodology described in Martens and Uhl (1984)
was modified and used to prepare the cross sections. The
samples used are listed in Table V. Samples were left 6-15
days in a 2:1 mixture of glycerin-alcohol 70% (1:6) and
Hidrofluoric Acid (HF). Due to its dangerous nature, HF
must be handled with extreme care under the hood and small
plastic vials must be used during this part of the
treatment. The length of time the samples can be treated in
HF without being damaged is quite variable. All species of
Bactris used in this study were left in the HF solution 6-9
days, and some, such as B. gasipaes and B. setulosa, 15
days without problems. Desmoncus (used as outgroup in the
cladistics analysis) was damaged after 6 days in the
solution. At the end of HF treatment the samples were
transferred to plastic vials (with small holes on the
sides) and washed 24 hours in running water to eliminate
all traces of HF. As it is known, even small amounts of HF
are enough to destroy the samples after they are mounted in
the slides.
The samples were transferred to small glass vials and
embedded in Paraplast Plus according to a protocol using a
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Hemo-De series (Appendix 04). For each of the changes in
the Hemo-De series (up to the 100% Hemo-De step) the vials
were kept in a vacuum chamber (12 p.s.i.) for 1 hour to aid
the penetration of the solutions. Since Hemo-De is less
volatile than Toluene or Butyl Alcohol, two additional
changes of 100% melted paraplast, at 24 hour intervals,
were made in the oven. The resulting blocks were trimmed
(exposing the leaf edge) and soaked for additional
softening in glycerin-alcohol 70% (1:6) for 2-5 weeks.
This last step is of great importance for a successful leaf
sectioning. All samples sectioned without this further
softening were of very bad quality and not suitable for
mounting in slides. In the last series of cross section
preparation a 2:5 mixture of glycerin:alcohol 70% was used
as the further softening agent, also with good results.
The blocks were sectioned with an A. O. Spencer 820
rotary microtome, and the knives sharpened after use.
Sections of 15-20 were obtained and mounted in glass
slides. Problems of sections falling off the slides
reported by Pinheiro (1997) were not observed during this
study. Only Haupt adhesive was used, and a thin layer was
spread out with the finger on the slide surface. Abundant
Formalin 4% was placed over the adhesive and the section
transferred to the slide with the help of a scalpo. The
slides were transferred to a warming plate (43C) and after
the sections were stretched, the excess of Formalin was
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eliminated using bulbilous paper. The slides were left up
to three hours in the warming plate and then transferred to
trays for an overnight period. To avoid the formation of
air bubbles in the sections it is important that the
warming plate does not overheat.
The sections were stained using a modified Safranin-
Astra Blue protocol (Appendix 05), developed by P. Endress
(Switzerland?). Although Martens and Uhl (1984) and
Pinheiro (1997) recommended the use of 0.5% Safranin-Fast
Green, we found it not satisfactory and expensive (it uses
absolute Alcohol to stop the fast green stain action).
Unlike Fast Green, Astra Blue does not stain the adhesive,
and therefore there it is no necessary to scrape the slides
with a razor blade, risking to damage the sections. Astra
Blue also gives a better time control of the stain
intensity, which is not the case when using Fast Green,
which requires a maximum of 20 seconds, sometimes less.
Several slides were overstained by Fast Green, and this
stain usually takes the place of Safranin, resulting in a
totally green section. The sections were mounted with
Kleermount and the slides were left on a warming plate for
2-3 days. A Nikon light microscope was used to observe and
take all the pictures presented in this study.
Results
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The cladisitic analysis of the resulting nine terminal
taxa of Bactris generated a single most parsimonious tree
with 69 steps, a consistency index (C.I.) of 0.65 and a
retention index (R.I.) of 0.69 (Fig. 07).
Although I have started the characters survey giving
the name B. macana for all herbarium specimens of the wild
pupunha, the morphological and anatomical studies show two
distinct and isolated populations for this species, treated
separately in the analysis. The northern South America
population bears the name Macana1 in the data matrix and
tree, and the southwestern Amazonian population is named
Macana2. The reason for the splitting was the presence of
a high number of polymorphic characters which occured when
all specimens were grouped together as an isolated
terminal taxon. The splitting of B. setulosa was also
necessary and it was done because two very distinct
staminate flower types were found among the herbarium
specimens examined. These specimens also had a strong
variation in the arrangement of the fruit endocarp fibers.
The resulting two terminal taxa originating from the
splitting of B. setulosa were not associated with any
specific populations or geographic areas.
The monophyly of the Guilielma and the Antillean
clades of Sanders was confirmed, but they are not placed
together to form the Non-Ocreate clade as observed in
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Sanders (1991), and Salzman and Judd (1995). In my study
the Antillean clade is the sister clade of all other taxa
analyzed, while the Guilielma Clade forms a more inclusive
clade grouped with B. riparia.
A new clade comprising B. setulosa plus B. riparia-
Guilielma is the sister clade of the Antillean clade. This
new clade is supported by two synapomorphies; the anther
with folded filaments and the lack of adaxial non vascular
fiber bundles, or if they are present, they do not show the
columnar or pentagonal shape typical of the taxa belonging
to the Antillean clade (Fig. 08, a, b).
The Guilielma clade is the best supported clade and
presents four synapomorphies, one of them, unique to this
clade, is the staminate petals with fibers heavily branched
apically (Fig. 09, a, b). The in-group arrangement in this
clade shows that B. gasipaes is the sister species to the
southwestern Amazonian population of B. macana (Macana 2).
These two species share two synapomorphies; the endocarp
irregularly oblong in lateral view and the fertile pore
displaced and located near the top of the endocarp (Fig.
10, a). B. gasipaes can be distinguished from the
remaining taxa of the Guilielma clade only by their fruit
with a characteristic ovoid shape (character 23) (Fig. 10,
b). The northern South America populations of B. macana
(Macana1) are defined by one reversal character state
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(character 120) and the homoplasy represented by the
distinct palisade mesophyll, also found in the clade
comprising the Jamaican and Hispaniola taxa (Fig. 11, a,
b).
The clade B. riparia-Guilielma is supported by two
unique synapomorphies; pinnae with a pendulous apex and the
presence of buttresses touching the hypodermis in the small
veins (Fig. 12, a, b). The B. riparia clade is
differentiated by two apomorphies; the square-shaped
adaxial hypodermal cells and the ellipsoidal shape of its
primary veins.
The Setulosa clade is supported by two unique
synapomorphies; the spines on the stem forming distinct
rings close to the nodes and the trullate or obtrullate
shape of their primary veins (Fig. 13, a, b). Setulosa1 is
segregated by the presence of one apomorphy; the fruits
with an obovoid shape, a condition that can be partially
explained by the fact Setulosa2 was scored as missing (?)
for this character. All four homoplasia and the reversal
character 12 (120) that segregate Setulosa 2 from its
sister taxon are related to its specific staminate flowers.
The in-gro
top related