Cano Et Al - 2013 - A Revision of Trithrinax PDF

Accepted by William J. Baker: 14 Oct. 2013; published: 8 Oct.
2013
1
PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Copyright 2013 Magnolia Press
Phytotaxa 136 (1): 153 (2013)
www.mapress.com/phytotaxa/
Article
http://dx.doi.org/10.11646/phytotaxa.136.1.1
A revision of the genus Trithrinax (Cryosophileae, Coryphoideae, Arecaceae)
NGELA CANO
1
, MATHIEU PERRET
2
& FRED W. STAUFFER
3
Conservatoire et Jardin botaniques de la Ville de Genve, Universit de Genve, Laboratoire de Systmatique vgtale et Biodiversit,
Ch. de lImpratrice 1, Case postale 60, 1292 Chambsy-Genve, Switzerland.
1
Angela.Cano@ville-ge.ch
2
Mathieu.Perret@ville-ge.ch
3
Fred.Stauffer@ville-ge.ch
Abstract
The palm genus Trithrinax is revised and three species and four varieties are accepted (T. brasiliensis var. brasiliensis, T.
brasiliensis var. acanthocoma, T. campestris, T. schizophylla var. schizophylla and T. schizophylla var. biflabellata comb.
nov.). This taxonomic treatment presents detailed and complete description of all taxa, based on the studies of natural
populations in Argentina, Brazil and Paraguay, the morphological analysis of herbarium specimens (including all
available types) and an exhaustive literature research. Illustrations are presented for all taxa and identification keys to the
species and varieties are proposed. Species descriptions include updated distribution maps, information about their
ecology, taxonomic notes and a compilation of common names and uses. The conservation status assessment reveals an
important level of threatening for all taxa, ranging from vulnerable (VU) to endangered (EN), mainly due to the decline
of the area of occupancy and the quality of habitat.
Introduction and historical survey
Trithrinax Martius (1837: 149) is a Neotropical palm genus distributed in the subtropical region of South
America. It is easily distinguished from other palms of the region by its relatively short size, palmate leaves,
persistent spiny leaf sheaths covering the trunk and solitary and hermaphrodite flowers. Trithrinax is placed in
the tribe Cryosophileae Dransfield et al. (2005: 561) of subfamily Coryphoideae Griffith (1844: 311;
Asmussen et al. 2006, Dransfield et al. 2008) and can be found in xeromorphic forests or open hilly areas,
where local people use their leaves to thatch and their fruits to prepare alcoholic beverages (Corra 1931,
Lorenzi et al. 2010).
The type species of the genus, Trithrinax brasiliensis Martius (1837: 150), was collected for the first time
by F. Sellow in southern Brazil and published by Martius in his magnificent palm treatment Historia Naturalis
Palmarum. In the third volume of that work Martius included a plate of T. brasiliensis (tab. 104). Burmeister
(1861: 48) described Copernicia campestris from Crdoba, Argentina, a species that was later transferred to
Trithrinax as T. campestris by Drude & Grisebach (in Grisebach 1879: 283). Trithrinax acanthocoma was
described by Drude (1878: 361) using a specimen collected by Glaziou in southern Brazil. The same author
described T. schizophylla in Martius Flora Brasiliensis (1882: 552) based on specimens collected by Weddell
and dOrbigny from Mato Grosso (Brazil) and Santa Cruz (Bolivia), respectively. The last species described
for the genus is T. biflabellata Barbosa Rodrigues (1899: 2), which was collected by J. D. Anisits from a
Paraguayan locality near Concepcin, and described in the Palmae Novae Paraguayense in 1899. Recently,
Quercellini et al. (2012) described two interesting cultivated individuals of Trithrinax (identified as T.
campestris), which are morphologically different from other taxa within the genus. They considered those
cultivated exemplars as a probable undescribed species.
CANO ET AL. 2 Phytotaxa 136 (1) 2013 Magnolia Press
Different treatments have been published for the genus, in most cases based only in herbarium material
(Drude 1882, Barbosa Rodrigues 1899, Beccari 1931, Pingitore 1978). Additionally, the genus has been
superficially treated in general floristic inventories including South American palms in general (Henderson et
al. 1995) or Brazilian palms (Lorenzi et al. 2010). Beside these efforts, and probably due to the difficulties
related to the collection of these palms (i.e. dry-xeric habitat, arborescent habit, strongly armed stems), there
have been relatively few taxonomic studies of this group, and it remains one of the few Neotropical palm
genera lacking a modern revision (Hahn 1990, Dransfield et al. 2008).
In order to contribute to the knowledge of this poorly studied genus we have undertaken an updated
monographic revision. This treatment includes the definition and description of the genus as well as the
species therein recognized. For this purpose we have visited natural populations of the species where
complete and informative samples were collected. This revision also includes the analysis of type and non-
type herbarium specimens, and exhaustive literature research. A compilation of common names and uses,
illustrations for all taxa, a key to the species based on vegetative and reproductive characters, a description of
the ecology of the species and updated distribution maps, imaging of the flowers employing scanning
electronic microscopy and an assessment of the conservation status for each taxa are also provided.
Material and methods
Fieldwork:Fieldwork in Florida, Brazil, Paraguay and Argentina was carried out in December 2010 and
FebruaryMarch 2011. In Florida, the botanical gardens of Fairchild Tropical Botanical Garden (FTBG), and
the Montgomery Botanical Center (MBC) were visited. Complete herbarium vouchers, photos and DNA
samples of Trithrinax and other Cryosophileae were collected in these institutions.
During the second trip natural populations of Trithrinax were visited and sampled in the provinces of
Chaco, Crdoba, Formosa, Jujuy, Salta and Santiago del Estero in Argentina, the states of Rio Grande do Sul
and Paran in Brazil and the departments of Alto Paran, Boquern and Presidente Hayes in Paraguay. Local
herbaria in those countries were visited in order to examine their collections of the genus.
Morphological analysis:Herbarium specimens were studied in the South American herbaria of
Argentina (CORD, CTES, SI; herbaria acronyms following Holmgren et al. 1990), Brazil (HAS, HPL, ICN,
MBM, SP, SPF) and Paraguay (FCQ, Itaip Binacional, PY), and in the European herbaria of G, P, FI, K and
BM. Further material was requested on loan from CTES, F, FTG, M, MO and NY. High quality images of
specimens from C and MVFA were also studied.
A detailed study of herbarium specimens and fresh material was carried out and the information gathered
included in a database. The measurements of the different organs were summarized as ranges or as means.
Normality of the data was tested with the Shapiro-Wilk test, a Fisher test was employed to compare the
variances of the data and the means were compared using a Students t test. Only well-developed leaves were
included in the morphologic analysis. Inflorescences at different developmental stages were studied (early
anthesis, full and late anthesis, presenting immature or mature fruits or after fructification) and the
measurement ranges of these organs include all possible stages. On the other hand, flowers were always
dissected at full anthesis for comparison.
The only fossil record described so far for Trithrinax consists of flowers found in the Dominican Republic
embedded in amber and identified as Trithrinax dominicana by Poinar (2002: 58). In order to evaluate the
identity of those flowers we contacted Dr. Poinar (Department of Entomology, Oregon State University), who
kindly provided additional pictures of the fossil. The images available from the paper and the additional
images showing in particular the gynoecium enabled us to complement the morphological description
presented by Poinar (2002).
Scanning electronic microscopy (SEM) of flowers:Flowers at full anthesis of all Trithrinax species
and varieties collected during the second fieldwork trip or sampled from herbarium specimens, were
dehydrated, critical-point-dried and sputter-coated with gold. Micrographs were obtained with a Zeiss DSM
Phytotaxa 136 (1) 2013 Magnolia Press 3 A REVISION OF THE GENUS TRITHRINAX (CRYOSOPHILEAE)
940A scanning electron microscope (Orion 6.60 Imaging System) at the Musum dHistoire Naturelle de la
Ville de Genve.
Distribution maps:Distribution maps are based on occurrences derived from 96 herbarium specimens
studied in the course of our work. Reports from recent publications (i.e. Moraes 2004; Gaiero et al. 2012)
were also included. When the coordinates were lacking in the specimen label, approximate coordinates were
found using Google Earth (6.0.1.2032 beta) based on the localities indicated in the label. Maps were
constructed using the layers available in the Digital Basemap of the Americas, prepared by Bletter et al.
(2003) from the New York Botanical Garden, using the software DIVA-GIS (Hijmans et al. 2005).
Conservation status assessment:The conservation status of the taxa were assessed by evaluating all
IUCN Categories and Criteria (IUCN 2001, IUCN 2010) but only the criteria A (population decline), B (size
of geographic ranges) and C (number of mature individuals, and population dynamics and structure) could be
practically applied. Criteria A and C were mainly analysed following our fieldwork observations. Criterion B
was examined using the software ArcView GIS 3.2 with the Conservation assessment tools extension Cats
(Moat 2007) and using the same coordinates that were employed to create the distribution maps. Following
the methodology suggested by Moat (2007) and using a grid of 10x10 km, we estimated the Extent of
Occurrence (EOO; defined as the distribution range of the taxon), the Area of Occupancy (AOO; defined as
the area within the EOO that is actually occupied by the taxon) and the AOO density. These results were
compared to the thresholds established by the IUCN and, considering also the evaluations of criteria A and B,
the different IUCN Categories (IUCN 2001) were assigned.
Morphology and anatomy
Stem:Species of Trithrinax present erect to slightly curved, solitary to clustered stems (Fig. 1) usually
covered by persistent tubular leaf-sheaths with strongly armed margins. Fire events are common in the
distribution area of Trithrinax, causing the loss of this protective device; the stem becomes naked and shows
only the leaf sheath scars. Reitz (1974) reported the presences of stolons for Trithrinax brasiliensis, but this
was not observed during our study. The anatomy of Trithrinax stems has not been studied in detail; however,
in a description of the stem anatomy of the genera belonging to tribe Cryosophileae, Tomlinson et al. (2011)
pointed out that for Trithrinax, the metaxylem of the central cylinder includes two narrow vessels, and that the
ground tissue in mature stems with parenchyma is highly compact.
Roots:Trithrinax species generally present a 2050 cm high, rarely to 3 m high, cone of adventitious
roots at the base of the stem (Fig. 2). These aerial roots are numerous, between 10 and 20 cm long and
between 0.2 and 1 cm of diameter. Old stems affected by fire or displaying old wound marks often show aerial
roots growing from the wounds, eventually serving as a protective mechanism for the stem.
Trithrinax roots were first described in a study published by Gillain (1900) in the course of a general
survey on the root anatomy of some species of Arecaceae and Pandanaceae. In that treatment he described the
epidermis and endodermis of Trithrinax brasiliensis, but no further analysis on the anatomical complexity was
provided. Drabble (1904) presented a wide study of the anatomy of palm roots, but contrary to the statement
of Tomlinson (1961), Trithrinax was not included in this study.
Seubert (1997) studied the morphology and the anatomy of the roots of Trithrinax brasiliensis including
informative figures of the surface view of the velamen with a few exodermal cells beneath, and a cross section
of the vascular cylinder of the species. For this palm she described the lateral roots as elongate, slender and
numerous; the rhizodermis cells were described as hexagonal in outline, wider than long and expanded
transversely to the root axis; the 1-layered velamen, presents cell walls unequally thickened, with outer cell
walls weakly thickened. Seubert described the 1-layered exodermis as not clearly distinguishable from the
outer cortex, and presenting cell walls equally and moderately thickened. The inner cortex of Trithrinax is
homogeneous, not divided into three zones, and the endodermis presents equally thickened cell walls. The
position of the vessels is restricted to the periphery.
FIGURE 1. Diversity of habits in Trithrinax. A. T. brasiliensis var. brasiliensis (Rio Grande do Sul, Brazil), scale bar: 40 cm; B. T.
campestris (Crdoba, Argentina), scale bar: 60 cm; C. T. schizophylla var. biflabellata (Presidente Hayes, Paraguay), scale bar: 20 cm;
D. T. schizophylla var. schizophylla (Salta, Argentina), scale bar: 70 cm.
FIGURE 2. Aerial root systems in Trithrinax. A. T. brasiliensis var. acanthocoma (Paran, Brazil), scale bar: 10 cm; B. T. brasiliensis
var. brasiliensis (Rio Grande do Sul, Brazil), scale bar: 20 cm; C. T. campestris (Santiago del Estero, Argentina), scale bar: 10 cm; D.
T. schizophylla var. biflabellata (Presidente Hayes, Paraguay), scale bar: 2 cm.
Tomlinson et al. (2011) included Trithrinax brasiliensis, T. campestris and T. schizophylla in their survey
of vegetative palm anatomy. This work confirms Seuberts (1997) observation that the genus presents a
homogeneous inner cortex, not differentiated in three zones and therefore lacking an aerenchyma in its middle
zone.
In a study of the mycorrhizal status of plant species in the Chaco Serrano Woodland from central
Argentina, Fracchia et al. (2009) reported root colonization of arbuscular mycorrhizal fungi in Trithrinax
campestris. A deeper study of this symbiotic relationship was carried out by Lugo et al. (2011), who described
and compared the roots colonization by arbuscular mycorrhizal fungi and dark septate endophytes in two
populations of Trithrinax campestris from Argentina, the first population being well conserved whereas the
other frequently affected by anthropogenic fires. This ecological analysis concluded that the fungal-root
relationship is destroyed by the fire events and that when the fungi are missing the palm produces a larger
amount of root hairs.
Leaves:Trithrinax species bear palmate leaves, nearly regularly divided up to mid-length of the blade
into numerous single-fold segments with induplicate plication and with shallowly to deeply bifid, apiculate to
sharp tips (Dransfield et al. 2008) (Fig. 3). The leaf of Trithrinax schizophylla is distinguished by its blade
divided almost to the base and forming two symmetrical halves. In well-conserved individuals observed in
wild populations (T. schizophylla and T. campestris), persistent marcescent leaves completely cover the stem.
The leaf-sheaths are tubular, persistent, smooth basally and reticulate towards the apex, the margin becoming
spiny (Fig. 3). The leaf sheaths spines originate when the softer tissues of the leaf sheaths decay, leaving the
interweaving skeletal network (Tomlinson 1961, 1962a, 1962b, 1990). The distal ends of the larger vascular
bundles project as stout and sharp spines, which ultimately become reflexed; these spines may protect the
immature inflorescences from predation by rodents or other medium size animals. The inflorescences are
short and surrounded by sharp spines at full anthesis and early stages of fruits development, and become
longer and clearly exposed when fruits are ripe. The persistent leaf-sheaths, as well as the marcescent leaf-
blades, may also be adaptations to fire, protecting the stem from direct flames; however this hypothesis
remains to be tested in the field.
Trithrinax petioles are adaxially shallowly channeled to rounded and abaxially rounded, the margins are
entire (Fig. 3), sharp and unarmed (Pingitore 1978, Dransfield et al. 2008). Tomlinson (1961) studied the leaf
anatomy of Trithrinax brasiliensis leaf axis (leaf sheath and petiole). It was described with an epidermis
presenting cubical cells, a 23-layered, lignified hypodermis, a peripheral chlorenchyma and scattered
vascular bundles. The adaxial hastula is triangular or deltoid, usually with a definite point (Fig. 3), and the
abaxial hastula is similar to the adaxial one although slightly smaller (Dransfield et al. 2008), or completely
absent. In a short note published by de Candolle (1913) a description of the adaxial and abaxial hastulas of
Trithrinax campestris was presented. The lamina was thoroughly studied by Tomlinson (1961), in particular
the histology and the anatomy of the epidermis, stomata, hypodermis, chlorenchyma, fibres, veins, transverse
commissures, ribs and expansions cells. More recently, Tomlinson et al. (2011) improved the study of the
vegetative anatomy in the genus by adding more species (i.e. T. campestris and T. schizophylla), as well as
information about the trichomes and the mesophyll of the lamina.
Tomlinson et al. (2011) distinguished Trithrinax from other genera of tribe Cryosophileae by its isolateral
lamina, adaxially and abaxially distributed stomata, knob-like or peltate, multicellular, and usually sunken
within the surface trichomes, veins presenting vascular bundles with abaxial fibre girder and expansion tissue
consisting in two bands adjacent to the abaxial rib.
Cell inclusions:Different types of cell inclusions have been reported for Trithrinax. Solereder & Meyer
(1928) reported the presence of raphide-sacs in the superficial indumentum of Trithrinax brasiliensis. In a
study including Trithrinax campestris and T. schizophylla, Guth (1970) found spherical silica bodies within
small cells (stegmata) associated to the leaf fibres, and excluded their presence in the epidermis. Bertoldi
(1971) classified the silica bodies observed in Trithrinax campestris as Globulita esferoequinulata.
Henderson (2006) reported the presence of spherical or ellipsoid silica bodies with spinulose margins,
distributed around the vascular bundles in the tissues of the seedlings of Trithrinax brasiliensis. The presence
of equidistant raphides and scarce tanniferous idioblasts was also reported in that work.
FIGURE 3. Leaf morphology in Trithrinax. A. Leaf sheaths and petioles of T. brasiliensis var. acanthocoma, scale bar: 10 cm; B. Leaf
sheaths tightly covering the stem of T. brasiliensis var. acanthocoma, scale bar: 20 cm; C. Entire leaf sheath of T. campestris, scale
bar: 10 cm; D. Entire leaf blade of T. brasiliensis var. acanthocoma, scale bar: 20 cm; E. adaxial view of hastula in T. brasiliensis var.
brasiliensis, scale bar: 1 cm; FI. Different types of segment apexes: F. T. schizophylla var. schizophylla, note deeply bifid and spiny
lignified apex, scale bar: 2 cm; G. T. campestris, note lignified-spiny apex, scale bar: 2 cm; H. T. brasiliensis var. brasiliensis, note
deeply bifid apex, scale bar: 5 cm; I. T. brasiliensis var. acanthocoma, scale bar: 1 cm.
Inflorescence and flowers:Trithrinax inflorescences (Fig. 4) are interfoliar, rather short to moderate
but never exceeding the length of the leaves, branched to two (rarely three) orders, robust and curved. They
are creamy-yellow at anthesis and green at fruit stage. The peduncle is short, the prophyll and two to five
peduncular bracts are morphologically similar, inflated, tubular at the base, expanded and split along one side
(Dransfield et al. 2008); the prophyll and the peduncular bracts release a sweetish fragrance during full
anthesis of the inflorescence. The rachis is longer than the peduncle and the rachis bracts are similar to the
peduncular bracts but becoming smaller towards the rachis apex, each subtending a first-order branch.
Branches are stout, curved, bearing small, triangular bracts subtending the rachillae, which are whorled to
spirally arranged, bearing small bracts each subtending a flower (Dransfield et al. 2008). Occasionally one or
two basal rachillae of the basal branches present further splitting. The inflorescences of Trithrinax campestris
were described in detail by Maldonado de Magnano (1973) with a schematic figure showing the inflorescence
architecture and a cross section diagram representing the arrangement of the prophyll, the peduncular bracts,
the rachis bracts and the branches (Fig. 4). Tomlinson (1990) considered the case of Trithrinax campestris as
an intermediate stage in the reduction in number of peduncular bracts in subfamily Coryphoideae, between the
cases of Sabal Adanson (1763: 495; with many bracts) and Chamaerops Linnaeus (1753: 1887; with one
bract) and used Trithrinax campestris as an example to describe a basic pattern regarding inflorescence and
flower structure in palms.
Trithrinax flowers appear sessile but Maldonado de Magnano (1973) highlighted the presence of a short
pedicel included in the perianth (Fig. 4). The three sepals are basally fused, fleshy, and slightly keeled
(Beccari 1907). The three petals are free, contorted or imbricate in bud (Beccari 1907, Tomlinson 1990). The
androecium is composed of six stamens, three are opposite to the sepals, and three are opposite and basally
adnate to the petals; the filaments are wide at the base, becoming much thinner towards the apex. The anthers
are versatile and dorsifixed, and their dehiscence is latrorse. The gynoecium is composed of three free,
uniovulate, carpels, with a short to long style (Beccari 1907). Ugolino Martelli revised and edited a
posthumous revision of Trithrinax by Odoardo Beccari (Beccari 1931) including detailed descriptions of the
flowers of each species, and precise line drawings of the different floral pieces. Morrow (1965) presented a
detailed morphological and anatomical description of Trithrinax brasiliensis flowers, in which he included
images of cross sections and longitudinal sections of the different floral organs. This study demonstrated that
the number of stamens and carpels may not be always six and three respectively, but rather varying from one
flower to another. A similar observation on the liability of reproductive pieces was pointed out by Maldonado
de Magnano (1973) concerning the gynoecium of Trithrinax campestris, which may have more than three
carpels. That study also described in detail the morphology and anatomy of the flowers of Trithrinax
campestris, including detailed information on the vascularization of the different floral whorls.
Fruits and seeds:Trithrinax fruits are one-seeded, globose, with an apical stigmatic remain (Fig. 4).
The epicarp is smooth and thin, the mesocarp is fleshy, and the endocarp is thin. The globose seed becomes
free from the endocarp at mature stage of the fruit and presents a circular hilum with ascending branches and
homogeneous endosperm (Fig. 4; Dransfield et al. 2008, Lorenzi et al. 2010).
Germination:A complete description of the germination and the seedlings of Trithrinax brasiliensis
was presented by Henderson (2006). The author included a cross section image of the eophyll midrib showing
the slightly protuberant midrib, the vascular bundle, the phloem strand, two metaxylem vessels, minor veins
attached to the adaxial epidermis and the multi-layered inner sheath. In her work, she described the position of
the seed remaining above the plumular-radicular node, the symmetric and straight plumular-radicular axis, the
stout and persistent primary root, and the induplicate marginal folds of the eophyll. A detailed anatomical
account of the simple and erect eophyll and its short petiole was also provided in this study. The latter
described the stomata as slightly sunken, the hypodermis as not distinct, the expansion cells as single-layered
and slightly larger than epidermal cells, the phloem strands as single and two metaxylem vessels as being
present. Henderson (2006) mentioned that some of the features shared by Trithrinax and other related genera
of the tribe Corypheae Martius (in Endlicher 1837: 252), as defined by Uhl & Dransfield (1987), are the single
cataphyll, and the entire, strongly plicate eophyll. According to Riffle & Craft (2003), fresh Trithrinax seeds
FIGURE 4. Reproductive organs in Trithrinax. A. Inflorescence structure (left hand) and longitudinal diagram of flower (right hand)
of T. campestris, after Maldonado de Magnano (1973); B. Inflorescences of T. brasiliensis var. brasiliensis at full anthesis, scale bar:
20 cm; C. Inflorescence of T. brasiliensis var. brasiliensis at full anthesis, scale bar: 10 cm; D. Fragment of rachilla showing insertion
of flowers at full anthesis in T. campestris, scale bar: 3 cm; E. Infructescences of T. campestris showing fruits at different
developmental stages, scale bar: 10 cm; F. Ripe fruit of T. brasiliensis var. acanthocoma, scale bar: 2 cm; G. LS of ripe fruit of T.
brasiliensis var. acanthocoma, note seed detached from the endocarp, scale bar: 2 cm. Credits C: K.P. Soares, D: J. Plumed.
germinate within 90 days, and completely dry seeds may take nine months or longer to germinate. The
germination of T. brasiliensis takes between 20 and 45 days (Hoyos & Braun 2001, Lorenzi et al. 2010), and
one year for T. schizophylla var. biflabellata (Lorenzi et al. 2010).
Karyology:Trithrinax karyology was first studied by Sat (1946), who reported a chromosome number
of 2n=36 for Trithrinax brasiliensis. For the same species Read (1963), found large chromosomes (13.5 m
long), and one year later (Read 1964) supported the report of Sat (1946). He also explained that
microsporogenesis of Trithrinax, and other related genera, occurs while the inflorescence is still deeply
enclosed within the tightly woven and spiny leaf sheath, which makes it difficult for karyologycal studies. Di
Fluvio (1966) studied the chromosome number of T. campestris and found 18 gametic chromosomes. Rser
(1994) published a systematic study of the chromosomes of the main palm groups in which T. brasiliensis and
T. campestris were included; the author confirmed previous reports of 2n=36 for Trithrinax, a chromosome
number shared by all the Cryosophileae members and with most of the Coryphoideae genera (Dransfield et al.
2008). Rser (1994) also explained that the metaphase chromosomes in Trithrinax continuously decrease in
size. Gaiero et al. (2012) presented a detailed karyotype analysis of T. campestris stating that the species
shows particularly large chromosomes (ranging from 3.11 to 8.02 m), the largest total chromosome length
for a diploid palm (185 m) and an exceptionally high genome size (2C = 17.15 0.07 pg) for the family.
They additionally highlighted the elevated heterochromatin amount present in the chromosomes, which
represents 39% of their total length. Gaiero et al. (2012) mentioned that small, clearly defined DAPI
+
bands in
the proximal and terminal regions of chromosomes 1, 4, 5, 14 and 18 are present in T. brasiliensis, whereas in
T. campestris these bands are only present in proximal regions. Another character useful to differentiate both
species is that DAPI
+
bands are larger and more numerous in T. campestris than in T. brasiliensis (Gaiero et al.
2012).
Palynology:The first description of Trithrinax pollen grains was published by Martius (1837). This
author described them as ellipsoideum, cum rimula longitudinali, meaning that the pollen grains were
ellipsoid with a longitudinal crease. The first modern description was based on Trithrinax campestris and
presented by Thanikaimoni (1966). This author presented a short morphologic description including
comments about the shape and the ornamentation of the exine. Thanikaimoni (1970) complemented our
knowledge of the pollen morphology in the genus by adding detailed measurements of the different axes, as
well as details on the thickness of the exine, the tectum, the collumelar layer and the foot layer. Sowunmi
(1972) included Trithrinax brasiliensis in a broad survey of palm pollen and its implications for taxonomy;
however no precise description of Trithrinax pollen was provided. Arbo (1974) carried out detailed SEM
studies of pollen grains in Trithrinax campestris and T. schizophylla. She presented a morphological
characterization of the pollen grains, including measurements of the polar axe, the diameters and the aperture
and the thickness of the exine.
Based on original data presented by Uhl & Dransfield (1987) and Harley (1996) on samples of T.
brasiliensis and T. campestris, Dransfield et al. (2008) characterized the pollen grains in the genus as
ellipsoidal, with slight to obvious asymmetry; the aperture is a distal sulcus; the ectexine is tectate, finely
perforate, perforate and micro-channelled, or perforate-rugulate; the aperture margin is slightly finer or
psilate; the infratectum is columellate; the longest axis measures 2545 m. SEM images of the distal face
and the coarsely perforate pollen surface were included in that publication.
Bauermann et al. (2010) compared pollen grains of the palm genera present in the state of Rio Grande do
Sul (RS, Brazil) and described different qualitative and quantitative attributes of pollen grains in Trithrinax
brasiliensis. In a study aiming to characterize the honey types produced in Entre Ros (Argentina), Fagndez
& Caccavari (2006) reported monofloral honey containing palm pollen grains of Trithrinax campestris. The
authors found that about 3% of the honey samples studied contained pollen of T. campestris.
Reproductive biology
Our knowledge about the reproductive biology of Trithrinax is still very sparse. Some of the populations of T.
brasiliensis and T. campestris visited between February and March 2011 had mixed reproductive stages.
Hence, in a single population and at the same period some individuals presented inflorescences at full anthesis
and others rather at an advanced fruiting stage. Flowering and fruiting periods seem highly variable. Pingitore
(1978) reported that the flowering period of T. brasiliensis var. brasiliensis is from January to March, whereas
for T. brasiliensis var. acanthocoma it is from December to March, for T. schizophylla var. schizophylla from
September to October and for T. schizophylla var. biflabellata from January to March. Lorenzi et al. (2010)
reported the fruiting period for T. brasiliensis var. brasiliensis from October to March, for T. brasiliensis var.
acanthocoma from August to February and for T. schizophylla var. biflabellata during November. Soares
(2009) pointed out that the flowering and fruiting periods of T. brasiliensis var. brasiliensis vary among the
populations of southern Rio Grande do Sul (Brazil), with flowering periods lasting from May to June or
November and fruiting periods occurring in March, October or November. Unfortunately most of the studies
cited above do not provide exact information on the localities to which this phenological data can be
attributed.
During our fieldwork, we could observe ants and wasps visiting inflorescences of T. brasiliensis at full
anthesis. Ants were also found in fruiting inflorescences of T. brasiliensis, constructing their nests inside the
peduncular and rachis bracts. Local people from Paran (Brazil) observed squirrels consuming fruits of T.
brasiliensis. Cows and goats were observed around wild populations of T. brasiliensis but there is no evidence
of seed dispersion by these animals.
Phylogeny and evolution
To date, few phylogenetic works have analysed the relationship of Trithrinax with the other genera composing
the tribe Cryosophileae. The analyses of plastid DNA sequences have so far provided little phylogenetic
resolution or low support within Cryosophileae (Asmussen & Chase, 2001, Asmussen et al. 2006). In these
analyses, Trithrinax campestris, the only species sampled for the genus, appears in a polytomy with the other
genera of the tribe. More recently, in a complete generic-level phylogeny of palms presented by Baker et al.
(2009), Trithrinax was placed as the sister genus of Chelyocarpus Dammer (1920: 395), but here again with
low support (bootstrap percentage: 58%). Improving resolution within Cryosophileae will probably require a
better taxonomic sampling and the use of additional nuclear markers. For example, Roncal et al. (2008)
managed to get moderate resolution of Caribbean Cryosophileae using PRK and RPB2 nuclear sequences.
Unfortunately, this work did not include sequences of Trithrinax. Preliminary analyses of PRK and RPB2
sequence data from samples collected in connection with this study support the monophyly of Trithrinax and
place the genus in a basal position in the phylogeny of the tribe, sister to Schippia Burret (1933: 867), with
69% bootstrap support. The low phylogenetic resolution obtained within Trithrinax did not help to distinguish
taxonomic entities at the specific and infraspecific levels.
Based on morphological and anatomical characters Trithrinax was considered as one of the least
specialized genera of palms (Maldonado de Magnano 1973, Moore & Uhl 1982, Uhl & Dransfield 1987,
Henderson et al. 1995) , as the character states found in the genus (i.e hermaphrodite, apocarpous gynoecium),
have been traditionally regarded as plesiomorphic for subfamily Coryphoideae. That view is, however, not
congruent with the most recent phylogenetic results (Baker et al. 2009). Indeed, the reconstruction of
evolutionary history of gynoecial characters showed that the apocarpy is a derived condition in palms that has
arisen independently up to four times, and that in Coryphoideae this character state has derived from the
plesiomorphic condition of postgenital syncarpy (Rudall et al. 2011).
Distribution and ecology
The genus Trithrinax is distributed in Argentina, Bolivia, Brazil, Paraguay and Uruguay (Fig. 5; Henderson et
al. 1995, Dransfield et al. 2008, Pintaud et al. 2008, Zuloaga et al. 2008, Gaiero et al. 2012). The genus is
present from 60 to about 1400 m of elevation. The northern limit of the distribution area is reached by T.
schizophylla, in the Bolivian department of Santa Cruz and the southern limit is reached by T. campestris in
Crdoba (Argentina). A collection of Herter (615, G) made in 1936 shows that that species was present in
Palmitas, Soriano (Uruguay), which may represent the southernmost locality for the genus, but no recent
reports from there were found. The Western limit of distribution is reached by T. campestris and T.
schizophylla in the Andean slopes. Towards the east the distribution area is reached by T. brasiliensis var.
acanthocoma in the Atlantic coast of southern Brazil. Trithrinax campestris and T. schizophylla have a broad
distribution area comprising northern Argentina, southern Bolivia and western Paraguay, whereas T.
brasiliensis is restricted to the southern states of Brazil and to a small population restricted to two individuals
in the Paraguayan Department of Alto Paran.
FIGURE 5. Geographic distribution of the species of the genus Trithrinax. T. brasiliensis (green circles); T. campestris (yellow
circles) and T. schizophylla (red circles).
The presence of T. brasiliensis in Uruguay was reported by Lombardo (1964) in the department of Treinta
y Tres (southeast) and by Lorenzi et al. (2010), although the latter author did not mention any precise
department for the country. The presence of this species in Uruguay is still to be confirmed as those authors
did not specify any detailed localities and no voucher specimens were provided.
An interesting distribution pattern was observed for the varieties recognized in T. brasiliensis and T.
schizophylla (Figs. 6 and 16). Both species present a predominant lowland variety: T. brasiliensis var.
brasiliensis (200400 m) and T. schizophylla var. biflabellata (100150 m), and a variety of much wider
altitudinal range, distributed from lowland to upland elevations: T. brasiliensis var. acanthocoma (450950
m) and T. schizophylla var. schizophylla (2001400 m).
FIGURE 6. Geographic distribution of Trithrinax brasiliensis and its two varieties T. brasiliensis var. brasiliensis (red circles) and T.
brasiliensis var. acanthocoma (yellow circles).
Regarding the distribution of the varieties of T. schizophylla an important gap was identified in the border
of Paraguay, Argentina and Bolivia (Fig. 16). Either the gap is due to an artifact of collection, possibly
explained by hard access conditions, implying that the palms are present there but have never been collected,
or the gap reveals a real vicariance in the distribution of the two varieties, probably due to environmental
barriers. An ecological explanation could be the dryer and warmer conditions of the region where the gap is
located, defined as the Dry Chaco with respect to the Humid Chaco present in the eastern region of central
Paraguay and northern Argentina (Olson et al. 2001).
The species of Trithrinax grow in open hilly or rocky areas, clear mid-height forests, un-flooded or
periodically flooded regions or xeromorphic thorny forests. The distribution area of Trithrinax occurs in the
subtropical region, in which the seasons of the year are more marked than in the tropical region. The species
of the genus are adapted to cold winters, sometimes with snow falls, and to warm summers. Boyer (1992)
reported a cold tolerance of T. brasiliensis to temperatures of -4C. For the same species Jacquemin (1999)
reported a tolerance of -6C and Meerow (2005) a tolerance of -5.6C without injuries. Trithrinax
schizophylla seems to be less tolerant and shows injuries when exposed to temperatures lower than -5C
(Meerow 2005). The more resistant species is T. campestris, with a tolerance of -10.6C according to Meerow
(2005) and from -8C to -10C according to Jacquemin (1999). Trithrinax brasiliensis and T. campestris are
highly tolerant to drought (Riffle & Craft 2003, Meerow 2005), whereas T. schizophylla is moderately
resistant (Meerow 2005). Adult Trithrinax individuals are also usually adapted to anthropogenic originated
fires. The populations of T. brasiliensis and T. campestris visited, frequently showed fire marks in the naked
stems, which lose their numerous leaf sheath layers, and their marcescent leaves. Baz (1933) explained that
the expansion of the agriculture forced local people in Entre Ros (Argentina) to destroy the original forests of
Trithrinax campestris using the fire to disarm the plants. After the burns, in short periods of time, new
leaves are generated by the palms (Bez 1933). In a study of the fungi-root relationship in T. campestris Lugo
et al. (2011) found that when fire destroys the symbiosis, the palm develops abundant root hairs as an
adaptation. However, seedlings are not resistant to fire episodes, showing that fire may represent an important
threat for the long-term survival for Trithrinax wild populations.
Conservation
The decline of the area of occupancy and of the quality of habitat represents the main threat for all taxa.
Trithrinax brasiliensis var. brasiliensis falls under the category of EN (endangered), and hence regarded
as the most threatened taxon within the genus. This species faces severe habitat destruction (criterion A3c),
very low population size (C2a(i)), small extent of occurrence (EEO: 7999 km
2
) small area of occupancy
(AOO: 600 km
2
), and fragmented area of occupancy (AOO density: 0.33). Trithrinax brasiliensis var.
acanthocoma also falls in the same category (criteria A3c and C2a(i)); however, this variety presents larger
EEO (151730 km
2
) and AOO (1100 km
2
). The AOO density for this taxon is 0.09, revealing an important
fragmentation of the Area of Occupancy.
The conservation status of Trithrinax campestris is vulnerable (VU) due to habitat destruction and high
levels of exploitation of the species (A3c and A3d), for which, according to local inhabitants, seeds are
massively collected for ornamental purposes. The estimated EOO for this species is 483515 km
2
, the AOO is
2800 km
2
and the AOO is highly fragmented (AOO density: 0.04).
The conservation status of T. schizophylla var. schizophylla and T. schizophylla var. biflabellata is
vulnerable (VU) due to a decline of the area of occupancy and the quality of habitat (A3c), with EOO, AOO
and AOO densities of 85515 km
2
, 1700 km
2
and 0.12. for T. schizophylla var. schizophylla and 141559 km
2
,
3300 km
2
and 0.24 for T. schizophylla var. biflabellata.
Previous conservation assessments that included species of Trithrinax were restricted to a specific
geographic region. In a study of the conservation status of Paraguayan palms, Gauto et al. (2011) proposed
IUCN red-list categories for T. brasiliensis var. acanthocoma (CR, based on criterion B1a) and T. schizophylla
var. biflabellata (LC, based on EOO and AOO estimates). The single population of T. brasiliensis var.
acanthocoma of Paraguay identified by Gauto et al. (2011) was visited by us during fieldwork in February
2011. During this visit we confirmed the high levels of threat of the taxon. In fact, only two extremely fragile
and old individuals were located. On the other hand, some of the figures obtained in our assessment for T.
schizophylla var. biflabellata are not congruent with those proposed by Gauto et al. (2011); the EOO of
232750 km
2
and the AOO of 177667 km
2
presented by these authors are very different from our results, which
include also the distribution area of the taxon in Argentina. Such difference is probably due to different
methods used for the estimations of EOO and AOO. Despite of the remarkable differences between the two
estimations, T. schizophylla var. biflabellata should be considered as a threatened species due to the heavy
destruction of the Gran Chaco region, the natural habitat of this palm, where the present rate of deforestation
is as high as 2142 ha/day (Palacios et al. 2012), proving that the decline of the area of occupancy of the taxon
is real.
Trithrinax campestris is probably the least threatened species in the genus, with a large Area of
Occupancy and moderately high population sizes. However, habitat destruction is also an important threat to
this species, known to be one of the most representative elements of the espinal forest type in Argentina
(Lewis et al. 2009). Espinal forests are dramatically declining due to agriculture expansion, over-grazing,
selective wood extraction for fuel or other uses (Lewis et al. 2006) and introduction of exotic invasive species
such as Broussonetia papyrifera (Linnaeus 1753: 986) L'Hr. ex Ventenat (1799: 547; Moraceae), Gleditsia
triacanthos Linnaeus (1753: 1056; Fabaceae) and Melia azedarach Linnaeus (1753: 348; Meliaceae; Lewis et
al. 2009). All types of forests, including the espinal forest where populations of Trithrinax campestris were
observed, are surrounded by extensive maize and soya fields.
In addition to extensive crop fields, other factors are closely related to the decline of the area of occupancy
and the quality of the habitat of Trithrinax species. Heards of cows or goats represent a threat due to two main
reasons: 1) In order to feed the cattle with young grass, local farmers intentionally burn large areas, killing
palm seedlings and severely injuring adults; 2) Cattle eat the fruits produced by adult palms as wells as the
leaves of the seedlings, the latter develop only in areas protected from cattle by fences or where the slopes are
very steep. We visited populations of Trithrinax brasiliensis var. brasiliensis where seedlings were completely
lacking and adult palms were represented only by extremely old individuals, suggesting low or no
regeneration of the population. In these localities cattle were always present.
Local inhabitants interviewed during our fieldwork in Brazil and Argentina pointed out that they use to
gather seeds from wild populations, and they are further sold for ornamental purposes, even commercialised
in international markets. Although the seed extraction from wild populations is difficult to quantify, this type
of activity should be closely monitored in order to evaluate its real impact on the populations. All these factors
should be taken into account if a protection strategy aiming to conserve these endemic palms is developed.
Since the IUCN categories presented here were assessed based on fieldwork observations and geographical
distribution based on extensive herbarium data, a deeper analysis, including studies of population dynamics, is
recommended to confirm our results.
Fossil record
The only fossil record described so far for Trithrinax consists of flowers found in the Dominican Republic
embedded in amber and identified as Trithrinax dominicana by Poinar (2002). In order to complement the
morphological description of the flowers and based on additional images of the fossils provided by Dr. Poinar,
we present the following description: the fossil flowers bear fused, glabrous sepals and free pubescent petals,
the latter three times longer than the sepals. Stamen filaments are free and about two times longer than the
sepals. Anthers are small, slightly ovate, dorsifixed and apparently versatile. The gynoecium is apparently
composed of three free pistillodes, each of these relatively small and topped by a short stigmatic branch.
The comparison of the fossil flowers with the flowers of extant Trithrinax species does not allow us to
support Poinars (2002) proposal of placing these fossil within Trithrinax. Although superficially similar,
some importat features are not shared between the fossil flowers and the flowers of extant species: 1) the
sepals of T. dominicana are considerably shorter than the ones observed in extant Trithrinax species (0.41
0.65 mm (Poinar 2002), vs. 13 mm, respectively), 2) the petals of T. dominicana are pubescent and not
imbricate, whereas extant Trithrinax species have glabrous, imbricate petals; 3) the anthers of T. dominicana
are conspicuously shorter than those of extant Trithrinax species (0.410.71 mm (Poinar 2002) vs. 14 mm
respectively); and 4) a detailed analysis of the gynoecium of T. dominicana suggests that the organs regarded
by Poinar (2002) as fertile carpels of a hermaphrodite flower may better correspond to pistillodes of a
staminate flower given their reduced degree of development and short size (although a possible reduction of
the size of the gynoecium due to dehydration caused by the fossilization process cannot be completely
excluded). Additionally, the stigmatic branches of the pistillodes of the fossil are clearly connivent whereas in
extant species of Trithrinax the styles and stigmas are clearly distinct and reflexed at anthesis.
In order to explore other possible taxonomic identities of the fossil we compared the floral morphology
displayed by other palm genera of Coryphoideae with similar geographic distribution to that shown by T.
dominicana. Our analysis led us to consider some affinities of the fossil with the flowers of Rhapidophyllum
H.Wendl. & Drude (in Drude 1876: 803) as defined by Dransfield et al. (2008). Indeed, this is the only genus
of Coryphoideae that shares with T. dominicana a New World distribution (in the Caribbean region and
southeastern North America), and staminate flowers with unfertile gynoecia composed of three small
apocarpic pistillodes. Among the characters displayed by Rhapidophyllum that do not match T. dominicana
could be mentioned its short filaments, anthers proportionally larger and its stigmatic branches reflexed at
anthesis, rather than connivent as observed in T. dominicana.
An accurate identification of this fossil is needed in order to understand evolutionary aspects related to
Neotropical Coryphoideae. So far neither the identity of the fossil flowers, nor the age of the Dominican
amber have been precisely established (Poinar 2002, Poinar pers. comm.). Thus, we believe that the use of this
fossil in order to calibrate phylogenies (see Crisp et al. 2010) or to explore hypotheses about the origins and
ancestral distribution patterns of palm groups (see Roncal et al. 2008 and Thomas & De Franceschi 2012)
could potentially offer misleading results.
Systematic treatment
Trithrinax Martius (1837: 149). Type:Trithrinax brasiliensis Mart.
Diodosperma Wendland (1878: 36). Type:Diodosperma burity H. Wendl. (Incertae Sedis)
Chamaethrinax H. Wendl. ex Pfister (1892: 46, treated as Chamathrinax). Type:Chamaethrinax hookeriana H. Wendl.
ex Pfister (Incertae Sedis).
Palms solitary to clustered, pleonanthic, hermaphroditic. Stem greyish, frequently covered with spiny leaf
sheaths, leaf scars prominent; aerial roots forming a basal cone, the roots rather smooth to rugose, light brown
or greyish. Leaves palmate; leaf sheath tubular, smooth at the base, reticulate with spiny margins towards the
apex, leaf sheath spines woody, square to rectangular in cross section; petiole flat to convex adaxially and
convex abaxially, smooth, light yellow-green or green, occasionally bearing a whitish indumentum; hastula
prominent, acute adaxially and inconspicuous, fragile and caducous abaxially; leaf blade coriaceous, lax to
rigid, loosely covered with a white-waxy indumentum, regularly segmented or biflabellate; leaf segments
pendant to erect, with a bifid, acute, sometimes lignified and spiny apex, plication induplicate. Inflorescences
interfoliar, branched into two (rarely three) orders, shorter than the leaves, erect at anthesis, pendant when
fruiting; peduncle dorsi-ventrally compressed in cross section, covered with a light brown-waxy indumentum,
frequently tightly enveloped by numerous layers of the persistent spiny leaf sheaths; prophyll covered with a
white-waxy indumentum towards the apex, coriaceous, reddish, opening laterally; peduncular bracts opening
laterally or bilaterally, slightly keeled, reddish, coriaceous, smooth towards the base and covered with a white-
waxy indumentum towards the acute apex; rachis similar to the peduncle at its basal portion, medial and
apical portions terete in cross section, smooth, the apical portion bearing several spirally arranged rachillae
and ending in a fertile rachilla; rachis-bracts subtending individual branches, basal rachis bracts similar to the
peduncular bracts, medial and apical rachis bracts smaller, weakly keeled, opening laterally, reddish, fragile,
smooth or with a white-waxy indumentum, their apices disintegrating late in the development of the
inflorescence; branches smooth, yellow (at flowering) or pale green (at fruiting), dorsi-ventrally compressed
and slightly channelled to terete in cross section, alternating or spirally inserted in the rachis; branches ending
in a fertile rachilla; rachillae whorled to spirally arranged, terete in cross section, smooth, pale yellow at
anthesis becoming green at fruiting stage, subtended by individual bracts, the latter small, inconspicuous,
deltoid, falling off at fruiting stage, occasionally one or two basal rachillae of the basal branches exhibiting
additional branching. Flowers apparently sessile, spirally arranged on the rachilla, pale yellow at full anthesis,
fragrant, each flower subtended by a small, inconspicuous, deltoid floral bract, which falls off at fruiting
stage; sepals 3, ovate, slightly keeled, acute, basally connate, margins entire; petals 3, ovate, slightly keeled,
imbricate, free, margins entire; stamens 6, free, basally adnate to the inner side of the petals; filaments longer
than the perianth and the gynoecium, clearly wider towards the base, much thinner towards the apex; anthers
dorsifixed, versatile; dehiscence latrorse; gynoecium 3-carpellate, apocarpous, ovary obovate to oblong, style
erect or curved, stigmatic branch papillose on its ventral side. Fruits one-seeded, globose, with an apical
stigmatic remain, epicarp smooth, thin, mesocarp fleshy, endocarp thin; seed globose, hilum circular, basal
with ascending branches, endosperm homogeneous, the seed detaching from the endocarp at mature stage of
the fruit. Eophyll simple, lanceolate.
Etymology:Tri: three; thrinax: generic name Thrinax Swartz (1788: 4). Martius (1837) identified in the
new genus a 3-carpellate gynoecium, which differed from the gynoecium of the genus Thrinax characterized
by its single carpel.
Distribution:Three species, distributed in subtropical South America, in southeastern Bolivia, northern
and central Argentina, western Paraguay, southern Brazil and western Uruguay (Fig. 5).
Taxonomic notes:Diodosperma was described by Hermann Wendland in 1878, based on seeds from
central-southern Brazil. The same year Drude compared Wendlands description with the seeds of Trithrinax
and found them very similar. However, it is only in 1882 that he formally placed Diodosperma as a synonym
of Trithrinax, a taxonomic decision that was also adopted by Beccari (1907) and by Dransfield et al. (2008).
Govaerts & Dransfield (2005) and Dransfield et al. (2008) considered Chamaethrinax as an invalid name and
cited page 19 of Pfisters publication. However, we consider that the name is valid because the genus is
correctly described on page 46, with further characters included in the key to the genera.
Key to the Species of Trithrinax
1. Peduncular bracts fibrous-reticulate (Fig. 17-E).......................................................................Trithrinax schizophylla.
- Peduncular bracts not fibrous-reticulate ...................................................................................................................... 2.
2. Leaf blade light to dark green; filaments as long as the anthers (Fig. 7). Brazil and Paraguay .....................................
.................................................................................................................................................. Trithrinax brasiliensis.
- Leaf blade pale green to greyish; filaments 2-3 times longer than the anthers (Fig. 12). Argentina and Uruguay........
................................................................................................................................................... Trithrinax campestris.
Multi-access key to the species of Trithrinax
Numbers refer to the numerical list of taxa (Appendix 1). Numbers in parenthesis indicate that the character is
only rarely found in that taxon.
Habitat
1. Geography: Country
a. Argentina (A)
2, 3a, 3b
b. Bolivia (Bo)
3a
c. Brazil (Br)
1a, 1b, (3b)
d. Paraguay (P)
1b, 3b
e. Uruguay (U)
2
2. Geography: Departments, Provinces or States
a. Chaco, Formosa (A), Mato Grosso do Sul (Br), Boquern, Concepcin, Presidente Hayes (P)
3b
b. Chaco, Crdoba, Corrientes, Entre Ros, San Luis, Santa Fe, Santiago del Estero, Tucumn (A),
Colonia, Paysand, Ro Negro, San Jos, Soriano (U).
2
c. Jujuy, Salta (A), Chuquisaca, Santa Cruz, Tarija (Bo)
3a
d. Rio Grande do Sul (Br)
1a, 1b
e. Santa Catarina, Paran (Br), Alto Paran (P)
1b
Vegetative characters
3. Habit
a. Solitary
1a, 1b
b. Clustering
(1a), (1b), 2, 3a, 3b
4. Marcescent leaves
a. Reaching the ground (covering the stem as a skirt)
2, (3a)
b. Not reaching the ground
1a, 1b, 3a, 3b
5. Stem height
a. 8m
1a, 1b, 2, 3a, 3b
b. > 8m
1b
6. Segments apex
a. Deeply bifid (as long or longer than of the total length of the segments)
1a, 3a, 3b
b. Entire to shortly bifid (shorter than of the total length of the segments)
1b, 2
c. Slightly to clearly spiny
1b, 2, 3a, (3b)
d. Not spiny
1a, (1b), (3a), 3b
Inflorescence
7. Peduncular bracts
a. Not fibrous-reticulate
1a, 1b, 2
b. Fibrous-reticulate
3a, 3b
FIGURE 7. Morphology of Trithrinax brasiliensis var. brasiliensis. A. Habit (Rio Grande do Sul, Brazil), scale bar: 30 cm; B. Leaf
bade, scale bar: 10 cm; C. Detail of hastula (adaxial view), scale bar: 2 cm; D. Segment apex, scale bar: 10 cm; E. Inflorescence at full
anthesis, scale bar: 10 cm; F. Scanning electronic micrograph of the flower, arrow pointing to the stigma, scale bar: 1 mm; G. Entire
infructescence showing immature fruits, scale bar: 10 cm.
Flowers
8. Sepals
a. Basal fusion < 2/3 of their length
1a, 1b, 2
b. Basal fusion 2/3 of their length
3a, 3b
9. Petals
a. Loosely imbricate
1a, 1b, 2
b. Strongly contorted
3a. 3b
10. Stamens
a. Filaments length = anthers length (ratio 1:1)
1a, 1b
b. Filaments length > anthers length (ratio 2:1 to 3:1)
2, 3a, 3b
11. Gynoecium
a. Ovary length = style length (ratio 1:1)
1a, 1b
b. Ovary length < style length (ratio 1:2 to 1:3)
2, 3a, 3b
Fruit
12. Size
a. Small (0.61.2 cm)
3a, 3b
b. Medium to large (1.54 cm)
1a, 1b, 2
1. Trithrinax brasiliensis Martius (1837: 150). Type:BRAZIL. Rio Grande do Sul, Sellow s.n. (holotype
M! (Fig. 8), isotypes B, destroyed, G!, P!, fragment ex B at FI!)
Palms solitary (rarely clustered). Stem 113 m tall 13.530 cm diameter (irregular in diameter when old),
greyish and often with associated lichens, leaf scars, separated 35 cm; aerial roots rather smooth, light brown
or greyish, 120 cm long 0.20.5 cm of diameter. Leaves 1535, lax to rigid; leaf sheath dark reddish or
brown, greyish and smooth towards the apex, leaf sheath spines light brown to dark reddish, 418.3 cm long
15 mm wide, rectangular in cross section; petiole 33108 cm long 0.93.5 cm wide; hastula 11.8 cm long
23.5 cm wide adaxially and ca. 1 cm long 23.5 cm wide abaxially; leaf blade, green, loosely covered
with a white-waxy indumentum more abundant abaxially than adaxially, leaf blade regularly segmented,
palman 0.49.5 cm (at the basal segments level) to 12.529.5 cm (median-apical segments level); leaf
segments 2746, with a lignified spiny or not lignified apex, basal segments 25.546.5 cm long 0.72 cm
wide, without or with an apical splitting of 0.944.5 cm, median segments 41101 cm long 1.53.4 cm
wide, with an apical splitting of 1.647 cm, apical segments 46113 cm long 1.53.7 cm wide, with an
apical splitting of 0.749 cm. Inflorescences 26, 4064 cm long, shorter than the leaves; peduncle 1020 cm
long 1.73.5 cm wide; prophyll up to 22 cm long; peduncular bracts 2 (13), 826 cm long, at fruiting stage
apart from each other 5.79.8 cm; rachis 3467 cm 0.43 cm wide at fruit stage, basal portion similar to the
peduncle but pale green, the apical portion bearing about 30 rachillae; basal rachis-bracts 1221.3 cm long,
medial and apical rachis bracts about 7.510.5 cm long; branches 56 per inflorescence; branches apart from
each other 3.212 cm, inserted 2070 with respect to the rachis axis, basal branches up to 39 cm long 20
cm wide, medial branches up to 31 cm long 28 cm wide, apical branches up to 27 cm long 21 cm wide;
rachillae 3552 per branch, inserted every 0.12 cm, subtended by bracts up to 0.5 cm long 0.10.2 cm
wide, basal rachillae 8.517 cm long 0.10.3 cm wide at fruiting stage, inserted at 7090 with respect to
the branches axis, median-apical rachillae 49 cm long 0.10.2 cm wide at fruiting stage, inserted at 4590
with respect to the branches axis. Flowers inserted every 0.10.5 cm, each subtended by a 0.10.2 cm long
0.1 cm wide floral bract; sepals 12 mm long 12 mm wide; petals 13 mm long 2 mm wide, not
imbricate; stamens filaments 35 mm long; anthers 34 mm long ca. 1 mm wide; gynoecium, carpels up to
2 mm long, ovary 1 mm long ca. 0.5 mm wide, obovate, style less than 1 mm long less than 1 mm wide.
Fruits 2.14 cm of diameter, pale yellow, epicarp smooth, thin, mesocarp fleshy, endocarp thin; seed 1.12.2
cm of diameter.
Etymology:brasiliensis: meaning Brazil, the home country of the material used by Martius for the
original description of this species.
Distribution and ecology:Distributed in southern Brazil in the states of Paran, Rio Grande do Sul and
Santa Catarina and in Paraguay, in the department of Alto Paran (Fig. 6). This species grows in lowland,
open areas or clear forests in hilly regions from 200-950 m of elevation.
Taxonomic notes:Trithrinax brasiliensis was the first species described for the genus on the base of a
collection made by Sellow in the southern Brazilian state of Rio Grande do Sul (RS). In 1878, Drude
described Trithrinax acanthocoma, based on a personal communication by Glaziou, who showed him this
ornamental palm cultivated in Rio de Janeiro and in some European parks and for which natural populations
were reported to be distributed in southern Brazil. Drude did not designate a type specimen in his publication;
however, in his treatment of the palms for the Flora Brasiliensis (Drude 1882) he complemented the
description of T. acanthocoma and designated as type specimen a collection of Glaziou (9014, P). The
definition of these two taxa has been controversial and they have been treated in different ways throughout
history. Mattos (1977) described the palms of the Brazilian state of Rio Grande do Sul, and presented a new
combination for Trithrinax acanthocoma, placing the taxon as a variety of T. brasiliensis. Unfortunately, this
important publication has not been cited in the most recent publications dealing with the genus Trithrinax. In
1995, Henderson et al. placed T. acanthocoma as a synonym of T. brasiliensis and the most updated treatment
of Brazilian palms by Lorenzi et al. (2010) reopened the debate by recognizing both names as independent
specific entities. To clarify this situation, we visited the natural populations of both taxa, we analysed
herbarium specimens, and we applied molecular phylogenetic methods (Cano et al. unpublished). The
resulting phylogeny lacked resolution at the specific and infraspecific levels (Cano et al. unpublished),
hindering any taxonomic decision. However, the morphological analysis of the specimens and fieldwork
revealed several differences between them, especially regarding the vegetative morphology (Tab. 1, key to the
varieties of the species), and contrasting distribution patterns between the two taxa (Fig. 6). Consequently, we
adopt Mattos (1977) decision and recognise two varieties within the species: Trithrinax brasiliensis var.
brasiliensis and T. brasiliensis var. acanthocoma.
Key to the varieties of Trithrinax brasiliensis
1. Leaf segments flexible and apically non-spiny, deeply bifid (1149 cm) ..................................... 1a. var. brasiliensis.
- Leaf segments rigid and apically spiny, entire to shortly bifid (0.79 cm) .................................1b. var. acanthocoma.
FIGURE 8. Holotype of Trithrinax brasiliensis, F. Sellow s.n (M), barcode: M0178215. Credits: herbarium G.
TABLE 1. Comparison of morphological and ecological traits between the two varieties of Trithrinax brasiliensis. For the mean
values of the quantitative characters, the number of measured organs is given in square brackets.
1
Information extracted from Lorenzi et al. 2010.
2
This range excludes two localities where the palms are suspected of being cultivated: Morro dos
Conventos (72 m) and Pikyry (250 m).
1a. Trithrinax brasiliensis var. brasiliensis (Fig. 7).
Stem 16 m tall; aerial roots forming a cone reaching up to 40 cm from the ground; leaves 1530, lax to
slightly rigid; leaf sheath spines 510 cm long; petiole 55108 cm long 0.92.5 cm wide; hastula ca. 1 cm
long ca. 2 cm wide adaxially and ca. 1 cm long ca. 2 cm wide abaxially; leaf blade thin, light green,
palman 0.42.0 cm (at the basal segments level) to 12.521.5 cm (median-apical segments level); leaf
segments 2743, flexible, with a bifid, acute, non-lignified apex, basal segments 25.546.5 cm long 0.71.6
cm wide, with an apical splitting of 11.744.5 cm, median segments 44.284 cm long 1.52.6 cm wide, with
an apical splitting of 1241 cm, apical segments 4694 cm long 2.13 cm wide, with an apical splitting of
1249 cm. Inflorescences 26. Flowers 5 mm long; sepals 1 mm long 1 mm wide, apex slightly acute;
petals 12 mm long 2 mm wide; filaments 3 mm long; anthers 3 mm long ca. 1 mm wide; gynoecium up
to 2 mm long, ovary 1 mm long ca. 0.5 mm wide, style ca. 1 mm long ca. 0.5 mm wide.
Distribution and ecology:Restricted to southern Brazil, in the southern region of the state of Rio
Grande do Sul (Fig. 6). The palm grows in open areas or poorly dense, up to 8 m high forests, in hilly or rocky
spaces. Also reported from riparian forests (Soares 2009). This variety is present in lowland areas from 200 to
400 m of elevation (Fig. 6).
Taxonomic notes:The fruits of T. brasiliensis var. brasiliensis observed during our fieldwork were
immature and measured 2.12.5 cm of diameter (Fig. 7). The seeds measured 1.11.2 cm of diameter
(obtained from Damazio-Silva s.n. (SP), a specimen with mature fruits). A different fruit size was given by
Lorenzi et al. (2010), who reported a diameter of 1.52.5 and included a photo in which some fruits are
mature, yellow, and measure about 3 cm of diameter.
Specimens examined:BRAZIL. Rio Grande do Sul: Sellow s.n. (FI ex B, M, P); about 70 km N of
Bag on the BR 153, 200 m, Gibbons & Spanner 19964 (M); Caapava do Sul, Guaritas, 249 m,
3050'06.4''S, 05330'37.4''W, 18 February 2011, Reis, Cano et al. 2611 (HBR); 74 km S de Caapava do Sul
(BR153, km 84), en Serrana, 7 December 1978, Krapovickas & Cristbal 34220 (CTES, G, ICN, K, MBM,
NY, SI); Caapava do Sul, 13 km al NW de Minas do Camaqua, 6 December 1978, Krapovickas & Cristbal
34207 (G, ICN); Quevedos, near Toropi's river, 286 m, 2923'41.3''S, 05401'19.9''W, 17 February 2011, Reis,
Cano et al. 2607 (HBR); Santo Anto, Santa Mara, 343 m, 2937'35.6''S, 05351'42.9''W, 19 February 2011,
Reis, Cano et al. 2613 (HBR); Saint-Hilaire 2764 (P). CULTIVATED. Brazil. So Paulo: Campinas, IAC, 5
Nov 2009, Lorenzi & Soares 6765 (HPL); Parque Estadual das Fontes do Ipiranga (Jardim Botnico), Perto do
var. brasiliensis var. acanthocoma Students t test **P<0.05
Stem length (m) 16 215
Aerial root cone height (m) up to 0.4 up to 3
Leaf blade colour
1
pale-green dark-green
Leaf blade thickness thin thick
Segments flexible rigid
Segments apex not lignified lignified-spiny
Mean number of segments 34.4 [18] 40.8 [11] -3.870 **
Mean medial segments length (cm) 54.6 [17] 66.7 [7] -2.145 **
Mean apical segments splitting (cm) 34.4 [17] 3.0 [10] 11.486 **
Elevation (m) 200400 450950
2
hidrofitrio, 25 November 1998, Kirizawa 3371 (SP); Jardim Botnico de So Paulo, Perto do porto histrico
e de bambuzal, 26 Mar 2001, Damsio-Silva s.n. (SP).
Local names:Carand-moroty, carand piranga, carand-uba, carandahy (Glassman 1972). Buriti,
buriti-palito, caranda, carand (Lorenzi et al. 2010).
Uses:The leaves are employed in woven handcrafts (Lorenzi et al. 2010) and as thatch. The fruits are
used to prepare an alcoholic beverage. It is rarely used for ornamental purposes but has potential in this area
(Lorenzi et al. 2010).
1b. Trithrinax brasiliensis var. acanthocoma (Drude) Mattos (1977: 15), (Fig. 9).
Trithrinax acanthocoma Drude (1878: 361). Type: BRAZIL. Rio Grande do Sul, Glaziou 9014 (holotype C! (Figs. 10
11); isotypes B, destroyed, F!, K!, P!, fragment ex B at FI!)
Stem 215 m tall; aerial roots forming a cone reaching up to 3 m from the ground; leaves 1535, rigid; leaf
sheath spines 7.218.3 cm long; petiole 3379 cm long 1.23.5 cm wide; hastula 1.51.8 cm long 1.53.5
cm wide, robust, adaxially and ca. 1 cm long ca. 1.53.5 cm wide abaxially; leaf blade, thick, dark green,
palman 0.89.5 cm (at the basal segments level) to 29.5 cm (median-apical segments level); leaf segments 35
46, rigid, with an acute, lignified-spiny apex, basal segments 3540.5 cm long 0.82 cm wide, entire or with
an apical splitting of 0.98.7 cm, median segments 50101 cm long 1.53.4 cm wide, with an apical
splitting of 1.66.2 cm, apical segments 63113 cm long 1.53.7 cm wide, with an apical splitting of 0.76
cm. Inflorescences 26. Flowers 78 mm longs; sepals 2 mm long 1 mm wide apex triangular; petals 23
mm long 2 mm wide; filaments 45 mm long; anthers 4 mm long ca. 1 mm wide; gynoecium up to 2 mm
long, ovary 1 mm long ca. 0.5 mm wide, style ca. 1 mm long ca. 0.5 mm wide. Fruits up to 4 cm of
diameter; seeds 1.92.2 cm diameter.
Etymology:acantho: spine; coma: foliage; Drude (1878) makes reference to the well-developed spines
in the leaf-sheaths of this taxon when comparing to T. brasiliensis.
Distribution and ecology:Distributed in Brazil, from northern Rio Grande do Sul to the state of
Paran. A small and extremely endangered population of this taxon is present near Ciudad del Este
(southeastern Paraguay) in the Refugio Biolgico Pikyry (Fig. 6). Commonly found in open areas and clear
forests in hilly regions, frequently associated with Araucaria Jussieu (1789: 413) forests (Reitz 1953). This
variety is also present in coastal regions of the states of Rio Grande do Sul (Lindeman et al. 1975) and Santa
Catarina, where it grows in saline conditions (Zocche et al. 2007). Trithrinax brasiliensis var. acanthocoma
grows at elevations from 450 to 950 m, although the coastal populations and one near Ciudad del Este, for
which an anthropogenic origin is suspected (Reitz 1974; Caballero-Marmori, pers. comm.), exist at lower
elevations.
Taxonomic notes: The fruits of T. brasiliensis var. acanthocoma collected by us in a mature stage were
pale yellow and measured 3.64 cm of diameter (Fig. 9), whereas Lorenzi et al. (2010) reported a diameter of
1.52.2 cm, and Beccari (1931) reported a diameter of 2.8 cm. The seeds collected by us had a diameter of
1.92.2 cm.
Specimens examined:BRAZIL. Paran: Imbituva, Pinho de Baixo, 15 July 1992, Pardini & Marcato
82 (SPF); Rio Grande do Sul: Glaziou 9014 (C, FI ex B, K, P); Bom Jesus, Margens do Rio Tainhas, 19 March
1999, Rossato 56 (MBM, NY); Farroupilha, 02920'4.6''S, 4943'23.2''W, 4 April 2001, Rossato 17 (MBM,
NY); Nova Petrpolis, Linha Brasil Fundos, Pedras do Silncio, 24 April 2006, Cossio et al. 67 (ICN); Pedras
Brancas, Municpio de Caixas, cerca de So Marcos, km 148, 28 May 1971, Mattos 15616 (SP); Itapeva,
Torres, 28 April 1985, Waechter & Baptista 2102 (HAS, ICN).PARAGUAY. Alto Paran: Ciudad del Este,
Pikyry, bosque costero del ro Paran, cercano al lago artificial Itaipu, 2512'33.20"S, 5430'56.10"W, 13
October 1995, Schinini & Caballero-Marmori 30221 (G); Itaipu Reservoir, Alto Parana River, Pikyry,
251248S, 543110W, 26 November 2002, Noblick et al. 5282 (FCQ); Campo Alegre, Pikyry, Refugio
Biolgico Karanday, 50 km NE de Hernandarias, 20 June 1995, Caballero-Marmori 3380 (ITAIPU).
FIGURE 9. Morphology of Trithrinax brasiliensis var. acanthocoma. A. Habit (Paran, Brazil), scale bar: 70 cm; B. Leaf blade, scale
bar: 20 cm; C. Detail of hastula (adaxial view), scale bar: 2 cm; D. Assymetrical segment apex, scale bar: 2 cm; E. Scanning electronic
micrograph of anthetic flower, arrow pointing to the stigma, scale bar: 1 mm; F. Ripe infructescence, scale bar: 10 cm; G. Ripe fruit,
scale bar: 1 cm; H. LS of a ripe fruit, scale bar: 1 cm; I. Seed, scale bar: 1 cm. Credits D: M. Faustino.
FIGURE 10. Holotype of Trithrinax acanthocoma, Glaziou 9014 (C), Credits: herbarium C. (1/2).
FIGURE 11. Holotype of Trithrinax acanthocoma, Glaziou 9014 (C), Credits: herbarium C. (2/2).
CULTIVATED. Brazil, Rio de Janeiro, Nova Friburgo, 17 July 1887, Glaziou 17343 (FI ex B, P); Nova
Odessa Coletado e culivado no Jd. Bot. Plantarum, 9 February 2009, Lorenzi & Soares 6763 (HPL);
Argentina, Corrientes, Santo Tom, Estancia Las Maras, ruta 14, 7 km S de Virasoro, 2 December 1970,
Maruak & Arbo 172 (CTES); Switzerland, Genve, Conservatoire et Jardin botaniques de la Ville de
Genve, rempotoir, 6 December 2010, Cano et al. 250 (G).
Local names:Buriti, carand, caranda (Reitz 1959, Mattos 1977, Lorenzi et al. 2010), carand falso,
surro (Glassman 1972) buriti-palito, caran, carand-moroti, carand-piranga, carandaba (Lorenzi et al.
2010).
Uses:Natural populations have become extremely reduced and some of the uses reported in the
literature probably no longer exist due to the lack of raw material. Corra (1931) reported that the stems are
employed to build fences or as torches; the leaves are used to make fans and brooms, and a strong fibre is used
to weave hats; a fibre is also extracted from the petioles in order to use in weaving; the fruits are used to
produce an alcoholic beverage, and edible oil is extracted from the seeds. This species is also cultivated for
ornamental purposes (Corra 1931, Lorenzi et al. 2010).
2. Trithrinax campestris (Burmeist.) Drude & Griseb. in Grisebach (1879: 283), (Fig. 12).
Copernicia campestris Burmeister (1861: 48). Type:Burmeister, 1881, pl. 9, fig. 20; Epitype (designated by
Krapovickas 2007): ARGENTINA. Crdoba: 3 km al E de Ballesteros, 12 March 1985, Krapovickas & Vanni 39776
(epitype CTES! (Figs. 1314), isoepitypes F!, MO).
Palms solitary or clustered with up to 6 stems. Stem 25 m tall 2030 cm diameter, greyish and often with
marks of bush fires, leaf scars separated 58 cm; aerial roots forming a basal cone reaching up to 30 cm from
the ground, the roots rather smooth, light brown, 10 cm long 0.3 cm of diameter. Leaves 2030, rigid; leaf
sheath up to 74 cm long, dark reddish and loosely covered with a white-waxy indumentum at the base, greyish
and glabrous towards the apex, leaf sheath spines up to 22, light brown, 3.321 cm long 36 mm wide;
petiole 1743 cm long 1.55.2 cm wide; hastula 1.22 cm long 2.23.5 cm wide, rigid adaxially and ca. 1
cm long 3 cm wide abaxially; leaf blade rigid, pale green, loosely covered with a white-waxy indumentum
on both sides, glossy, leaf blade with symmetric or asymmetric base, regularly segmented, palman 0.43.0 cm
(at the basal segments level) to 1233 cm (median-apical segments level); leaf segments 2034, with a bifid,
woody and spiny asymmetric apex, basal segments 1132.5 cm long 0.51.8 cm wide, with an apical
splitting of 0.57.8 cm, median segments 3165 cm long 1.84.2 cm wide, with an apical splitting of 2.613
cm, apical segments 4576 cm long 25.3 cm wide, with an apical splitting of 3.69.2 cm. Inflorescences 2
5, about 3050 cm long; peduncle up to 30 cm long 1.52 cm wide, light brown; peduncular bracts 4, 2735
cm long 5.58 cm wide, apically acute and rigid, at fruit stage apart from each other 9.510 cm; rachis 50
53 cm 0.42.5 cm wide at fruit stage, basal portion similar to the peduncle but pale green, medial and apical
portions, pale green, the apical portion bearing up to 20 rachillae; rachis-bracts basal rachis bracts 1621 cm
long 4.57 cm wide, medial and apical rachis bracts about 7 cm long 67 cm wide, pale yellow, smooth;
branches 56 per inflorescence; branches apart from each other 610.5 cm apart at fruiting stage, inserted at
1050 with respect to the rachis axis, basal branches up to 40 cm long 23 cm wide, medial branches up to
38 cm long 20 cm wide, apical branches up to 26 cm long 23 cm wide; rachillae 2839 per branch,
inserted every 0.12 cm, subtended by bracts 0.51cm long 0.10.2 cm wide, basal rachillae 915 cm long
0.2 cm wide at fruiting stage, inserted at 7090 with respect to the branches axis, median-apical rachillae
4.512 cm long 0.2 cm wide at fruiting stage, inserted at 3070 with respect to the branches axis. Flowers
inserted every 0.10.5 cm, each subtended by a 0.10.2 cm long 0.1 cm wide floral bract; sepals 3 (rarely 5,
Krapovickas & Vanni 33754, CTES), 2 mm long 12 mm wide; petals 3 (rarely 5, Krapovickas & Vanni
33754, CTES), 3 mm long 3 mm wide, not imbricate; stamens 6 (rarely 10, Krapovickas & Vanni 33754,
CTES); filaments 6 mm long; anthers 23 mm long ca. 1 mm wide; gynoecium 3-carpellate (rarely 5-
carpellate, Krapovickas & Vanni 33754, CTES), carpels up to 3 mm long, ovary 1 mm long ca. 0.5 mm
wide, oblong, style 2 mm long ca. 0.5 mm wide, stigma hook-shaped. Fruits 2.33 cm of diameter, pale
yellow, mesocarp 5 mm wide, bitter (Fig. 12). Beccari (1907) reported fruits 1.8 cm in diameter, with slightly
lignified, fragile endocarp. Beccari (1931) published a drawing of the fruits based on Lorentz 28 (B) showing
that in some cases all three carpels developed; seeds collected by us had a diameter of 1.7 cm and an intrusion
seed coat about 7 mm long, whereas Beccari (1907) reported a diameter of 1.21.4 cm. We judge these
differences to be taxonomically insignificant.
Etymology:campestris: that is found in the countryside, referring to its usual habitat.
Distribution and ecology:Central and northern Argentina, in the provinces of Chaco, Crdoba,
Corrientes, Entre Ros, San Luis, Santa Fe, Santiago del Estero and Tucumn (Fig. 15). In Uruguay it is
present in the departments of Paysand, Ro Negro (Gaiero et al. 2012), San Jos and Soriano, and based on
ancient collections the species could be also present in Colonia, although fieldwork efforts should confirm
whether these populations are still alive. These palms are present in open hilly areas and in xerophyte plane
Espinal forests of thorny trees and shrubs (Lewis et al. 2009), often affected by anthropogenic fires (Lewis et
al. 2009, Lugo et al. 2011).
Specimens examined:ARGENTINA. Crdoba: La Unin 3 km al este de Ballesteros, 5 April 1985,
Ferruchi 381 (G, CTES); La Unin, 3 km al E de Ballesteros, 153 m, 32 32' 60''S, 62 58' 60''W, 12 March
1985, Krapovickas & Vanni 39776, (CTES, F); Entre Chilibroste y Noetinger al lado de la carrilera del tren,
ruta 2, km 173, 117 m, 3221'08.2''S, 06224' 32.7'' W, 9 March 2011, Cano 277 (G, SI); Ro Segundo,
Manfredi, Estacin experimental INTA, 22 March 1978, Krapovickas & Vanni 33754 (CTES); Serrezuela, 20
March 1978, Kiesling 1210 (MO, SI); Cerca de Orcosu, Yerba buena etc., 1718 February 1876, Hieronymus
398 (P); Ro Seco, 17 January 1972, Hunziker et al. 21737 (CORD); Sobremonte, 30 January 1987, Hunziker
& Subilis 24958 and 24960 (CORD); Departamento de las Minas, February 1876, Hieronymus 412 (CORD);
Entre Ros: Federal, em beira de rodovia, lado esquerdo ca. 200 km de Uruguaiana, BR, 6 March 2008, Tsuji
2481 (HPL); La Paz, 14 km al sureste del acceso a Col. Avigdor, ruta12, 14 September 1986, Krapovickas &
Cristbal 40637 (CTES, G); Villaguay, Mojones Norte, 15 March 1985, Pedersen 14081 (MO); Concepcin
del Uruguay, 25 March 1875, Lorentz 879 (CORD); San Luis: Ayacucho, 28 December 1885, Kurtz s.n.
(CORD); Santiago del Estero: Pozo Hondo, a la entrada de la Estancia La Donosa, km 790, 7 km antes de
Pozo Hondo por la ruta 34, 270 m, 2713'24.8''S, 06427'51.0''W, 7 March 2011, Cano 276 (G, SI); Tucumn:
Leales, Los Puestos, ruta N9, 1 January 1971, Krapovickas & Cristbal 17399 (CTES); Leales, Los Puestos,
Ruta Nac N9, 3 May 1971, Krapovickas et al. 18551 (P).
URUGUAY. San Jos: Camino de entrada a Kiy por Libertad, antes del arroyo Mauricio, 12 September
1969, Del Puerto et al. 8671 (MVFA); Soriano: Palmitas, 1938, Herter 615 (G); Vera, March 1898, Berro 371
(MVFA); Vera, March 1913, Berro 6661 (MVFA); Unknown department: Banda oriental del Uruguay, 1816
1821, Saint-Hilaire 2464 (P).
Taxonomic notes:Trithrinax campestris was originally described as Copernicia campestris by Hemann
Burmeister in 1861. He provided a short description of the palm and its habitat, but did not designate a type
specimen. After the species was placed in the genus Trithrinax by Drude & Grisebach (Grisebach 1879),
Burmeister (1881) published a drawing of the palm habit in a complementary work, in which details of the
leaves and reproductive organs were missing. Burmeisters drawing was taken as the type of the species. In
Beccaris revision of Trithrinax (Beccari 1931), several herbarium specimens of T. campestris collected by
Lorentz and Niederlein were cited, some of them deposited in CORD and FI. Krapovickas (2007) designated
an epitype in order to complement Burmeisters drawing; in doing so, he chose a collection gathered in the
original locality specified by Burmeister (1861). The epitype collected by Krapovickas and Vanni presents all
the vegetative and reproductive organs necessary to accurately identify the taxon.
Local names:Caranday, carand (Bez 1933, del Vitto et al. 1994), carand de Entre Ros (Glassman
1972), palma (del Vitto et al. 1994), saro, caranda-i, palma carandilla (Cabral & Castro 2007).
FIGURE 12. Morphology of Trithrinax campestris. A. Habit (Santiago del Estero, Argentina), scale bar: 50 cm; B. Leaf blade, scale
bar: 10 cm; C. Detail of hastula (adaxial view), scale bar: 1 cm; D. Segment apex, scale bar: 2 cm; E. Scanning electronic micrograph
of anthetic flower, arrow pointing to the stigma, scale bar: 2 mm; F. Flower at full anthesis, arrow pointing to the stigma, scale bar: 2
mm; G. Infructescence showing fruits at different developmental stages, scale bar: 10 cm; H. Ripe fruit showing perianth remnants,
scale bar: 1 cm I. CS of the fruit and the seed, scale bar: 1 cm; J. Seed, scale bar: 1 cm; K. LS of seed showing the homogeneous
endosperm, scale bar: 1 cm.
FIGURE 13. Epitype of Copernicia campestris, Krapovickas & Vanni 39776 (CTES); barcode: CTES0000146. Credits: herbarium
CTES. (1/2).
FIGURE 14. Epitype of Copernicia campestris, Krapovickas & Vanni 39776 (CTES); barcode: CTES0000147. Credits: herbarium
CTES. (2/2).
FIGURE 15. Geographic distribution of Trithrinax campestris (yellow circles).
Uses:A mixture of sugarcane fibres and the cellulose extracted from this palm is used to fabricate
cardboard boxes (Luna 1977). The stems are used as posts and shingles due to the fact that plane boards can
be produced from them (Cabral & Castro 2007). The leaves are used to make brooms and brushes, and
extracted fibres are used to make ropes or baskets; these fibres were formerly used to fabricate typical
Argentinian sandals named alpargatas (Cabral & Castro 2007). The reticulate leaf sheaths are used to filter
the water and to make artisanal dolls and other handcrafts. The fruits are used to feed livestock (Bez 1933,
Cabral & Castro 2007) and also used to prepare an alcoholic beverage and oil. This species is also cultivated
for ornamental purposes.
3. Trithrinax schizophylla Drude (1882: 552). Lectotype (designated by Beccari 1931):BOLIVIA. Santa
Cruz: Chiquitos, September and October 1845, Weddell 3498. (lectotype P! (Figs. 1821), fragment ex P at
FI!); Syntype: Amrique Mridionale, dOrbigny s.n. (P!).
Palms solitary or clustered, up to 6 stems. Stem up to 5 m tall 5.515 cm diameter, greyish to pale brown,
leaf scars separated 23 cm; aerial roots forming a cone reaching up to 10 cm from the ground, the roots
smooth to rugose, 110 cm long 0.31 cm of diameter. Leaves 1225, lax to rigid; leaf sheath 1828 cm
long, dark reddish, greyish and smooth towards the apex, leaf sheath spines light brown, 631 cm long 14
mm wide; petiole 2291 cm long 0.53.2 cm wide; hastula 0.71 cm long 1.31.8 cm wide adaxially and
0.50.7 cm long 1.31.5 cm wide, bifid, abaxially; leaf blade, thin to thick, rigid to flexible, green, loosely
covered with a white-waxy indumentum abaxially and adaxially, leaf blade with symmetric or asymmetric
base, regularly segmented to biflabellate with a deep split at the middle of the blade forming two identical
portions, palman 0.12.0 cm (at the basal segments level) to 232 cm (median-apical segments level); leaf
segments 1536, pendant to erect, with a bifid, acute or spiny apex, basal segments 22.557 cm long 0.42
cm wide, entire or with an apical splitting of 0.849 cm, median segments 31.382 cm long 0.82.8 cm
wide, with an apical splitting of 1.249 cm, apical segments 3484 cm long 0.63.4 cm wide, with an apical
splitting of 3.559 cm. Inflorescences 13, 3574 cm long; peduncle 1036 cm long 1.03.3 cm wide, white
to light brown; prophyll up to 9.523 cm long, fibrous, pale-brown towards the apex; peduncular bracts 45,
13.520 cm long, pale-brown, smooth towards the base and fibrous towards the apex, the peduncular bracts at
fruiting stage apart from each other 4.57 cm; rachis 3045 cm 0.42 cm wide at fruit stage; basal rachis-
bracts 1315 cm long, medial and apical rachis bracts 5.511 cm long; branches 56 per inflorescence;
branches apart from each other 3.510 cm, inserted 1070 with respect to the rachis axis, basal branches 20
47 cm long 1122 cm wide, medial branches 1932 cm long 825 cm wide, apical branches 1128 cm
long 821 cm wide; rachillae 3775 per branch, inserted every 0.12.3 cm, bracts up to 0.5 cm long 0.1
0.3 cm wide, basal rachillae 8.518 cm long 0.10.3 cm wide, inserted at 4590 with respect to the
branches axis, median-apical rachillae 414 cm long 0.10.3 cm wide, inserted at 3090 with respect to the
branches axis. Flowers inserted every 0.10.8 cm; sepals ca. 2.5 mm long ca. 1.5 mm wide, basally connate
up to ca. 2 mm forming a cupule with three cuspidate tips; petals ca. 3.5 mm long 4 mm wide, strongly
imbricate; filaments 6 mm long; anthers 2 mm long ca. 0.5 mm wide; gynoecium up to ca. 3.5 mm long,
ovary 1 mm long ca. 0.5 mm wide, ovate, style ca. 2.5 mm long ca. 0.5 mm wide, erect or slightly curved.
Fruits 0.61.2 cm of diameter, fleshy, pale yellow; seed 0.51 cm of diameter.
Etymology:schizo: splitted, divided; phylla: leaf, in reference to the divided leaf blade.
Distribution and ecology:From southern Bolivia and northern Argentina, to central and western
Paraguay and southwestern Brazil (Fig. 16).
Taxonomic notes:Trithrinax schizophylla was described by Drude (1882) in Martius Flora
Brasiliensis, where he designated two syntypes: a collection of Weddell 3498 (prov. Mato Grosso finibus et
prov. Chiquitos), and dOrbigny s.n. (inter prov. Boliviae orientalis Chiquitos et montes pr. S. Cruz de la
Sierra). In his revision of Trithrinax, Beccari (1931) designated as lectotype of T. schizophylla the specimen
of Weddell 3498 from P. His decision was probably based on the fact that the specimen Weddell 3498
perfectly corresponds to the definition of the species, whereas the specimen of dOrbigny s.n. corresponds to a
mixed collection of a leaf-sheath of Trithrinax sp. and a leaf of Copernicia alba Morong (1893: 246).
Trithrinax biflabellata was described by Barbosa Rodrigues in his work Palmae Novae Paraguayenses
(1899), based on several unnumbered J. D. Anisits collections and one picture studied by him; in the same
publication a detailed line drawing was proposed. Anisits main palm specimens were part of the Barbosa
Rodrigues palm collections, which were destroyed by fire (Stafleu & Cowan 1976). Other material collected
by Anisits has been reported in BP and S (Kmmerle 1933, Lanjouw & Stafleu 1954); unfortunately no palm
material from this collector could be found in these herbaria (Z. Barina and M. Ehn pers. comm.).
Consequently, the only original material available is the drawing presented in the original publication (Fig.
23). The latter accurately represents most of the diagnostic characters of the taxon and hence designated here
as the lectotype of the species, following the ICBN articles 9.11 and 9.12.
Trithrinax biflabellata was placed under synonymy of T. schizophylla by Henderson et al. (1995). We
observed natural populations of both taxa (Trithrinax biflabellata in the Paraguayan and Argentinian Chaco
(type locality) and T. schizophylla in North-western Argentina), and we analysed a total of 41 herbarium
specimens from a broad range of localities. As a result, we noticed several morphological and biogeographical
differences that lead us to propose two varieties: T. schizophylla var. schizophylla and T. schizophylla var.
biflabellata. The main differences are summarized in Tab. 2 and in the key to the varieties of T. schizophylla.
A clear geographical differentiation for the two taxa is displayed in the distribution map including the two
varieties (Fig. 16).
FIGURE 16. Geographic distribution of Trithrinax schizophylla and its two varieties T. schizophylla var. schizophylla (yellow circles)
and T. schizophylla var. biflabellata (red circles).
TABLE 2. Comparison of morphological and ecological traits between the two varieties of Trithrinax schizophylla. For the mean
values of the quantitative characters, the number of measured organs is given in square brackets.
Character var. schizophylla var. biflabellata
Students t
test
*P<0,1
**P<0.05
Palm aspect robust slender
Leaf blade thickness thick thin
Segments rigid flexible
Segments apex normally lignified-spiny normally non-spiny
Mean petiole width (cm) 1.8 [14] 1.1 [27] 4.2599 **
Mean number of segments 28.5 [14] 20.2 [29] 7.5046 **
Mean segments width (cm) 1.7 [40] 1.2 [73] 4.3260 *
Elevation (m) 2501400 100150
Barbosa Rodrigues (1899), in his description of T. biflabellata, listed some differences between this taxon
and T. schizophylla. However, these do not allow an accurate differentiation of the taxa. Barbosa Rodrigues
cited as a distinguishing character the biflabellate leaves for T. biflabellata; however, this leaf-segmentation
type is present in both taxa, although more evident for T. biflabellata. Another character proposed by him was
the presence of a single carpel in T. biflabellata versus a 3-carpellate gynoecium in T. schizophylla. Our study
shows that all flowers dissected for both taxa consistently show 3 carpels. Barbosa Rodrigues was clearly
aware of the existence of two independent taxonomic entities but, probably due to a lack of material and
fieldwork, he could not sort out the variation in his description of T. biflabellata. The characters presented in
Tab. 2 allow a better differentiation of the two varieties.
Key to the varieties of Trithrinax schizophylla
1. Robust palms; leaf blade segments (22) 28 (36), mostly lignified-spiny at the apex; plants distributed at 2501400
m of elevation in SE Bolivia and NW Argentina (Salta and Jujuy) ................................................... var. schizophylla.
- Slender palms; leaf blade segments (15) 20 (26), mostly non-spiny at the apex; plants distributed at 100150 m of
elevation in Paraguay and NE Argentina (Chaco and Formosa) ..........................................................var. biflabellata.
3a. Trithrinax schizophylla var. schizophylla (Fig. 17).
Palms robust; Stem up to 5 m tall; leaves rigid; leaf sheath spines 1031 cm long; petiole 2251 cm long
0.73.2 cm wide; leaf blade thick, palman 0.52.0 cm (at the basal segments level) to 8.526 cm (median-
apical segments level); leaf segments 2236, rigid, with a bifid, acute, slightly spiny to lignified-spiny apex,
basal segments 22.556 cm long 0.42 cm wide, entire or with an apical splitting of 2.749 cm, median
segments 3682 cm long 1.02.8 cm wide, with an apical splitting of 15.549 cm, apical segments 39.584
cm long 0.83.4 cm wide, with an apical splitting of 15.559 cm. Inflorescences 13. Flowers 45 mm long;
sepals 12 mm long 1 mm wide; petals 2 mm long 12 mm wide; filaments 4 mm long; anthers 12 mm
long ca. 1 mm wide; gynoecium 2 mm long, ovary ca. 1 mm long ca. 0.5 mm wide, style ca. 12 mm long
ca. 0.5 mm wide. Fruits were not observed at mature stage during the present study but Moraes (2004)
described them as being yellow-green to black, 68 mm of diameter.
Distribution and ecology:Distributed from southeastern Bolivia in the departments of Chuquisaca,
Santa Cruz and Tarija, to northwestern Argentina in the provinces of Jujuy and Salta (Fig. 16); it grows in
xerophytic open areas and in mid-height thorny forests of the Chaco region, from 250 to 1400 m of elevation.
Specimens examined:BOLIVIA. Chuquisaca: Sud Cinti, ca. de la Comunidad de Las Abras, can
Piura, 1100 m, 2105'34''S, 6416'22''W, 23 October 2005, Lozano & Cardoso 1519 (MO); Luis Calvo,
municipio Villa Vaca Guzmn, Cantn Iguembe, Itaco, Trayecto Iguembe-Muyupampa, 1148 m, 2041'S,
06347'W, 19 October 2007, Jimenez et al. 552 (MO); Santa Cruz: Chiquitos, September and October 1845,
Weddell 3498 (FI ex P, P); Chiquitos, Pozo del Tigre, 135 km al este de Santa Cruz, propiedad de la empresa
ABAFA, alrededores del campamento principal, 250 m, 1734'S, 06157'W, 29 September 1994, Vargas &
Ortiz 3200 (F); Prov. Andres Ibanez, 5 km SE of Comunidad San Lorenzo, 320 m, 1749'30''S, 06250'W, 17
November 1990, Nee & Coimbra 39931 (MO); west Velasco, July 1892, Kuntze s.n. (NY); Provincia
Cercado, Cantn-Lugar: Cotoca, Don Lorenzo, Bola, 400 m, 13 April 1990, Moreno 30 (NY); 900 m,
2007.550'S, 06328.914'W, Gibbons & Spanner 19963 (M); Tarija: Burdet O'Connor, Berety, 7.9 km (by
road) from Entre Ros to Palos Blancos, 1040 m, 2126'01''S, 06400'03''W, 26 March 2007, Nee & Flores
54963 (MO); Burdet O'Connor, Rancho Saururo, 5 km S of highway to Villamontes, 30 km E of Entre Rios,
1200 m, 2130'S, 06400'W, 6 September 1987, Killeen 2721 (NY); lmite Prov. Chaco/ Burdet O'Connor.
Puerto Margarita, sobre el ro Pilcomayo, 450 m, 25 October 1983, Beck & Liberman 9758 (NY); Burdet
O'Connor, Tacuarenda, 35 km al E de Entre Ros, 1 May 1983, Krapovickas & Schinini 39081 (CTES, G).
FIGURE 17. Morphology of Trithrinax schizophylla var. schizophylla. A. Habit (Salta, Argentina), scale bar: 60 cm B. Leaf blade,
scale bar: 10 cm; C. Detail of hastula (adaxial view), scale bar: 1 cm; D. Segment apex, scale bar: 2 cm; E. Inflorescence at full
anthesis, note the fibrous-reticulate apex of the peduncular and rachis bracts (Krapovickas 18645, CTES), scale bar: 10 cm; F.
Scanning electronic micrograph of the flower, arrow pointing to the stigma, scale bar: 1 mm.
FIGURE 18. Lectotype of Trithrinax schizophylla, Weddell 3498 (P), barcode: P00725553. Credits: herbarium P. (1/4).
ARGENTINA. Jujuy: Ledesma, Chalicn, ruta 34, 10 May 1971, Krapovickas et al. 18645 (CTES, P, SI);
San Pedro de Jujuy abt. 20 km N of San Pedro, 500 m, 2402.250'S, 06447.905'W, Gibbons & Spanner
19962 (M); San Pedro de Jujuy, Depto. De Santa Brbara, about 40 km south of San Pedro del Jujuy on Santa
Brbara road, all along the roadside for about 10 km, 1400 m, 8 October 1938, Eyerdam & Beetle 22848 (G,
K); San Ramn de la Nueva Orn, ruta La Estrella-Siete aguas, 123 km al S de San Ramn de la Nueva Orn,
730 m, 2354'00''S, 6420'18''W, 13 December 2001, Marcato et al. 328 (SI, SPF); Depto. de Santa Brbara,
Palmeras, a 20 km de Vinalito, 22 October 1964, Cabrera & Fabris 16236 (K).
UNKNOWN COUNTRY: Amrique Mridionale [most probably Brazil], Orbigny s.n. (P)
Local names:In Bolivia these palms are known as palma de sao or sa, palmerita, palma chica, saro,
palma chuco, caryay-t, caryay-m (Moraes 2004), utsaho (Glassman 1972). In Argentina the common names
are saro or saho (Pingitore 1978).
Uses:The leaves are used to thatch; the segments of young leaves are used to weave hats (sombrero de
sa in Bolivia), small bags and fans. These palms are also cultivated for ornamental purposes (Moraes 2004).
3b. Trithrinax schizophylla var. biflabellata (Barb. Rodr.) An. Cano & F.W. Stauffer comb. nov. (Fig. 22).
Trithrinax biflabellata Barbosa Rodrigues (1899: 2). Type: Palm. Paraguay. Tab. 1; PARAGUAY. San
Salvador, ad Arroyo Porongo prope Togatiy et in Chaco inter flumina Pilcomayo et Negro, Anisits s.n.
(missing). Lectotype (designated here):Palm. Paraguay. Tab. 1 (Barbosa Rodrigues 1899; Fig. 23).
Palm slender. Stem up to 5 m tall; leaves flexible; leaf sheath spines 621 cm long; petiole 3191 cm long
0.51.8 cm wide; leaf blade coriaceous-thin, palman 0.12.0 cm (at the basal segments level) to 232 cm
(median-apical segments level); leaf segments 1526, flexible, with a bifid, acute, non-spiny to slightly spiny
apex, basal segments 26.557 cm long 0.51.3 cm wide, entire or with an apical splitting of 0.843.5 cm,
median segments 31.374 cm long 0.82 cm wide, with an apical splitting of 1.243 cm, apical segments
3480 cm long 0.62 cm wide, with an apical splitting of 3.546.5 cm. Inflorescences 13. Flowers 78 mm
longs; sepals 23 mm long 1 mm wide; petals 34 mm long 4 mm wide; filaments 6 mm long; anthers 2
mm long ca. 1 mm wide; gynoecium 34 mm long, ovary 1 mm long ca. 0.5 mm wide, style ca. 23 mm
long ca. 0.5 mm wide. Fruits were not observed in a mature stage during this work but Beccari (1931)
described them of 1 cm in diameter and Lorenzi et al. (2010) reported 0.81.2 cm., the latter also measured
seeds of 6 mm in diameter.
Etymology: bi: two; flabellata: fan-shaped. Making reference to the divided leaf blade.
Distribution and ecology:This palm is distributed in Paraguay and Argentina. In Paraguay it is present
in the departments of Alto Paraguay, Boquern, Presidente Hayes and known in the Department of
Concepcin only from the protologue. In Argentina it is present in the North-eastern provinces of Chaco and
Formosa (Fig. 16). A recently discovered population that may correspond to T. schizophylla var. biflabellata
was reported from Mato Grosso do Sul, Brazil, near the frontier of Brazil and Paraguay (Soares pers. comm.),
but an accurate identification of this palm is still needed. This variety grows in the lowland Chaco region, in
xeromorphic open areas or thorny forests, in terra firme or periodically flooded areas, from 100 to 150 m of
elevation.
Specimens examined:PARAGUAY: Alto Parana: Camino a Fuerte Olimpo, entre Fn. C. A. Lopez y
Fn. Torres, 15 May 1981, Krapovickas et al. 37435 (CTES); Estancia Laguna Ganso, 2234'3''S, 5936''51''W,
25 September 2002, Mereles et al. 8824 (CTES); Boquern: Campo Loro, 4 July 2003, Vogt 47 (FCQ);
Aproximadamente a 70 km de Filadelfia, terreno perteneciente a la Cooperativa Menonita, 116 m,
2224'25.5''S, 05920'38.2''W, 28 February 2011, Cano & Palacios 272 (FCQ, G); Filadelfia, Col. Fernheim
(Ro Verde?) parque de recreo para la salud mental, 15 September 1990, Vanni et al. 2161 and 2162 (G);
Filadelfia Col. Fernheim, 10 km E de Campo Grande, 2145'S, 059W, 15 September 1990, Vanni et al. 2156
(G); Filadelfia Coletado a cerca de 500 km de Ponta Por por estrada e de 120 km em linha reta da fronteira
FIGURE 22. Morphology of Trithrinax schizophylla var. biflabellata. A. Habit (Boquern, Paraguay), scale bar: 50 cm; B. Leaf
blade, scale bar: 10 cm; C. Detail of hastula (adaxial view), scale bar: 1 cm; D. Segment apex, scale bar: 20 cm; E. Inflorescence at
late anthesis, scale bar: 10 cm; F. Scanning electronic micrograph of the flower, arrow pointing to the stigma, scale bar: 1 mm.
FIGURE 23. Lectotype of Trithrinax biflabellata reproduced from Palm. Paraguay. (1899) Tab. 1.
com Brasil (regio de Porto Murtinho-MS), 20 November 2009, Lorenzi et al. 6780 (HPL); Newland km 412,
ruta Carlos A. Lpez, km 2 del cruce Loma Plata, camino a Newland 2234'41.58"S, 5949'52.23"W, 20
September 1989, Pea & Molas 221 (PY); Tyto Neuland-Campo Aceval, 23 July 1992, Mereles & Degen
4553 (FCQ); Entrada a la Estancia Pitiantuta, 2116'13''S, 5936'19''W, 8 February 2005, Vogt & Mereles 232
(FCQ); Colonia Mennonita, Parcialidad Sanapana, 2249S, 5949W, Mereles 4162 (CTES, FCQ, G).
Presidente hayes: Distrito Teniente Irala Fernndez, Estancia Salazar, potrero 5 (aproximadamente 1 km
despus del peaje interno de la Estancia), yendo hacia el norte, costado derecho de la carretera, 113 m,
2304'42.3''S, 05913'21.3''W, 27 February 2011, Cano & Palacios 271 (FCQ); Ruta Fortn General Bruguez,
entre km 5055, 25 July 1995, Pietrobom da Silva & Zardini 2155 (SPF); Monte Lindo, ruta trans-Chaco, km
210, 6 December 1995, Krapovickas & Cristbal 44186 (CTES, K, MO); Monte Lindo, 28 November 1988,
Caballero-Marmori 1643 (ITAIPU); Pozo Colorado, Estancia Victoria, ca. 40 km SE of Pozo Colorado,
collection site ca. 3.5 km E of the headquarters for Estancia Victoria, 131 m, 2343.763'S, 05833.782'W, 23
November 2002, Noblick & Rios-Otero 5273 (FTG); Ruta Transchaco, 21 km NW Tte. Irala Fernndez, 10
May 1994, Krapovickas et al. 45257 (CTES, G); De pozo Colorado, 93 km ante civitatem Concepcin, 4
March 1980, Bernardi 20166 (BM, G); Estancia La Perla, 2326'S, 05934'W, 14 October 1986, Pedersen
14637 (G); Estancia , 2416'21.0''S, 5833'27.0''W, 1 February 2005, de Egea & Pea-Chocarro 768 (BM);
Estancia Santa Elisa, 17 September 1994, Zardini & Guerrero 41281A (NY); Estancia Vanguardia,
2417'2.00''S, 5839'11.0''W, 13 September 2004, de Egea et al. 498 (BM); Pilcomayo river basin, trail to
Fortn General Bruguez, 100 m, 2455'25''S, 05811'07''W, 4 June 2000, Zardini & Guerrero 54705 (FTG).
Unknown department: Chaco, Hassler 2707 (FI ex B); Paraguay, S.E. en montes (orillas), Hassler 12701 (G).
ARGENTINA. Chaco: Castelli, General Gemes, 20 km al N de Asustado, Ruta 95 (camino de tierra), 24
April 1997, Castro 14 (CTES); Depto. de Ro Teuco, puente Saladillo, Ruta 95, a 115 km de Saenz Pea, 19
March 1952, Schulz 8302 (K); Formosa: Las Mercedes, ruta 81, 28 km NW de Piran, 8 June 1971,
Krapovickas et al. 19570 (F); 200 m, 2436.635'S, 06044.178'W, Gibbons & Spanner 19961 (M); Dep.
Patio, Colonia Muiz, a 8 km de Las Lomitas, 3 June 1985, DellArciprete 27 (CTES); Dep. Pirane, Loc.
Palo Santo, 6 December 1972, Maruak et al. 419 (CTES).
Local names:In Brazil locals call this palm carandilla, carandai (Lorenzi et al. 2010), or buriti (Cabral
& Castro 2007). In Paraguay it is known as karandilla (Gauto 2009) and in Argentina as carandillo (Pingitore
1978) or palma carandilla (Cabral & Castro 2007).
Uses:The stems are used as torches and in construction (Hahn 1990, Gauto 2009), the palm heart or
palmito is eaten and the leaves and the spines are used to make handcrafts such as baskets, fans and a variety
of useful objects (Arena 1981, Gauto 2009). The leaves are also used for thatch or strong fibre (Hahn 1990),
and the fruits are eaten by local people (Gauto 2009). This species is also cultivated for ornamental purposes
(Lorenzi et al. 2010).
Incertae Sedis
Chamaethrinax hookeriana H. Wendl. ex Pfister (1892: 46). Type:unknown, not present at GOET (M. E.
Reiner-Drehwald and G. W. F. Wagenitz, pers. comm.)
Govaerts & Dransfield (2005) considered Chamaethrinax hookeriana as an invalid name and cited the page
46 of Pfisters (1892) work. However, in this page a complete diagnosis of the genus and one single species is
proposed, which according to the ICBN rules is a valid publication. Pfister (1892) did not indicate a type
specimen for Chamaethrinax hookeriana and based his description on an individual of Brazilian origin
cultivated at Kew Gardens. He found the leaf anatomy of C. hookeriana and Trithrinax brasiliensis very
similar, except for the thickness of the stomata cuticle, which he described as thin in T. brasiliensis and rather
thick in C. hookeriana. Beccari (1907) placed Chamaethrinax hookeriana as a synonym of Trithrinax
campestris based on the suggestion he received from U. Dammer who studied a specimen of C. hookeriana in
B (he did not specify any details about that specimen). Despite of the lack of a type, we believe that C.
hookeriana may be probably a synonym of Trithrinax brasiliensis (instead of T. campestris; Beccari 1907,
Henderson et al. 1995, Govaerts & Dransfield, 2005), based on Pfisters description of the leaves of
Chamaethrinax hookeriana and on the Brazilian origin of the individual he studied (T. campestris has never
been reported in Brazil).
Diodosperma burity Wendland (1878: 118). Type:unknown, not present at GOET (M. E. Reiner-Drehwald
and G. W. F. Wagenitz, pers. comm.)
Diodosperma burity was described in 1878, based on seeds that Wendland received from central-southern
Brazil. Wendlands description of D. burity did not include any reference specimen. Drude (1878) compared
Wendlands description to the seeds of Trithrinax campestris and found them very similar. In 1882, Drude
explained that he could not find any differences between the seeds of Trithrinax and those described by
Wendland (1878) and expressed that the lack of other organs in the description of D. burity hindered the
possibility to assign it in one of the species of Trithrinax described so far. Nonetheless, he placed D. burity as
a probable synonym of T. schizophylla, adding a question mark beside the name. At that time, he had not seen
the fruits of the latter species and only had access to the description provided by Martius (1847). Given the
mentioned arguments, and based on our current knowledge of the distribution and the seed morphology of
Thritrinax, D. burity could be a synonym of Trithrinax brasiliensis or Trithrinax schizophylla. However,
without a type specimen, we are unable give an accurate determination for that name.
Excluded taxa
Trithrinax aculeata Liebm. ex Martius (1853: 320) = Cryosophila nana (Kunth in Humboldt et al. 1816: 299)
Blume ex Salomon (1887: 158).
Trithrinax chuco (Mart.) Walpers (1849: 1005) = Chelyocarpus chuco (Martius in Orbigny 1847: 45) Moore
(1972: 73).
Trithrinax compacta Griseb. & H. Wendl. in Grisebach (1866: 221) = Hemithrinax compacta (Griseb. & H.
Wendl.) Hook. f. in Bentham & Hooker (1883: 930).
Trithrinax mauritiiformis Karsten (1856: 244) = Sabal mauritiiformis (H. Karst.) Griseb. & H. Wendl. in
Grisebach (1864: 514).
Acknowledgements
This study was funded by the International Palm Society Endowment Found, the Augustin Lombard grant
from the SPHN Society of Geneva and by the Commission of the travel grant of the Acadmie des sciences
naturelles Suisse. For precious help during fieldwork we would like to thank H. Lorenzi (HPL), A. Reis
(UFSC), K. P. Soares, M. Virmond (and co-workers), R. Degen (FCQ), I. Gauto (Guyra Paraguay), C.
Palacios, P. Cacciali, R. Eaton (Estancia Salazar), G. Caballero-Marmori (ITAIPU), E. Rempel, Luis A.
Ventura, N. Deginani (SI) and M. Belgrano (SI). For help with the Scanning Electronic Microscopy we thank
A. Piuz from the Musum dHistoire Naturelle de la Ville de Genve. For allowing us to analyse their
collections we thank Z. Barina (BP), the C herbarium, L. Ariza and A. Anton (CORD), M. M. Arbo, A.
Krapovickas and R. O. Vanni (CTES), the F herbarium, R. Degen (FCQ), C. Nepi and R. M. Baldini (FI), C.
E. Lewis (FTBG), M. L. Abruzzi de Oliveira (HAS), H. Lorenzi (HPL), C. Carneiro (ICN), G. Caballero-
Marmori (ITAIPU), W. Baker and L. Gardiner (K), the M herbarium, J. Gregson (BM), P. Griffith and L.
Noblick (MBC), O. dos Santos (MBM), the MO herbarium, P. Gaiero (MVFA), the NY herbarium, T.
Haevermans (P), M. Ehn (S), N. B. Deginani and M. Belgrano (SI), M. C. Mamede (SP), R. de Mello-Silva
(SPF). For reviewing this article we thank S. Zona and F. Borchsenius. For various scientific and logistic
contributions to this work we thank K. P. Soares, D. Ouattara, A. Chautems (G), N. Fumeaux (G), C.
Chatelain (G), L. Nusbaumer (G), P. Perret (G), P. Boillat (G), O. Durbin (G) and M. Gonzalez. K. P. Soares,
M. Faustino and J. Plumed kindly provided some of the images used in this revision.
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Appendix 1. Current taxa accepted and list of exsiccatae
Numerical list of taxa
1.a. Trithrinax brasiliensis Mart. var. brasiliensis
1.b. Trithrinax brasiliensis Mart. var. acanthocoma (Drude) Mattos
2. Trithrinax campestris (Burmeist.) Drude & Griseb.
3.a. Trithrinax schizophylla Drude var. schizophylla
3.b. Trithrinax schizophylla Drude var. biflabellata (Barb. Rodr.) An. Cano & F.W. Stauffer
List of exsiccatae
Beck, St. G. 9758 (3a)
Bernardi, L. 20166 (3b)
Berro, M. B. 371, 6661 (2)
Caballero-Marmori, G. 1643 (3b); 3380 (1b)
Cabrera, A. L. 16236 (3a)
Cano, A. 250 (1b); 271, 272 (3b); 276, 277 (2)
Castro, M. 14 (3b)
Cossio, R. 67 (1b)
Damsio-Silva, E. s.n. (1a)
de Egea, J. 498, 768 (3b)
Del Puerto, O. 8671 (2)
DellArciprete, A. 27 (3b)
Eyerdam, W. 22848 (3a)
Ferruchi, M. S. 381 (2)
Gibbons & Spanner 19961 (3b); 19962, 19963 (3a); 19964 (1a)
Glaziou, A. F. M. 9014, 17343 (1b)
Hassler, E. 2707, 12701 (3b)
Herter, W. G. F. 615 (2)
Hieronymus, G. 398, 412 (2)
Hunziker, A. T. 21737, 24958, 24960 (2)
Jimenez, M. C. 552 (3a)
Kiesling, R. 1210 (2)
Killeen, T. 2721 (3a)
Kirizawa, M. 3371 (1a)
Krapovickas, A. 17399, 18551 (2); 18645 (3a); 19570 (3b); 33754 (2); 34207, 34220 (1a); 37435 (3b); 39081 (3a);
39776, 40637 (2); 44186, 45257 (3b)
Kuntze, C. E. O. s.n. (3a)
Kurtz, F. s.n. (2)
Lorenzi, H. 6763 (1b); 6765 (1a); 6780 (3b)
Lozano, R. 1519 (3a)
Marcato, A. C. 328 (3a)
Maruak, V. 172 (1b); 419 (3b)
Mattos, J. 15616 (1b)
Mereles, F. 4162, 4553, 8824 (3b)
Moreno, L. R. 30 (3a)
Nee, M. 39931, 54963 (3a)
Noblick, L. R. 5273 (3b); 5282 (1b)
Orbigny, A. C. V. M. D. d' s.n. (3a)
Pardini, R. 82 (1b)
Pedersen, T. M. 14081 (2) 14637 (3b)
Pea T. F. 221 (3b)
Pietrobom da Silva, M. R. 2155 (3b)
Reis, A., Cano, A., et al. 2607, 2611, 2613 (1a)
Rossato, M. 17, 56 (1b)
Saint-Hilaire, A. F. C. P. de 2464 (2) 2764 (1a)
Schinini, A. 30221 (1b)
Schulz, A. G. 8302 (3b)
Sellow, F. s.n. (1a)
Tsuji, R. 2481 (2)
Vanni, R. 2156, 2161, 2162 (3b)
Vargas, I. 3200 (3a)
Vogt, C. 47, 232 (3b)
Waechter, J. L. 2102 (1b)
Weddell, H. A. 3498 (3a)
Zardini, E. 41281A, 54705 (3b)

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