Anales del Jardín Botánico de Madrid
Vol. 67(2): 103-112
julio-diciembre 2010
ISSN: 0211-1322
doi: 10.3989/ajbm.2254
Anatomy and fruit development in Schinopsis balansae
(Anacardiaceae)
by
Ana María González 1 & José Luis Vesprini 2
2
1
Instituto de Botánica del Nordeste, Corrientes, Argentina. anitama39@gmail.com
Facultad de Ciencias Agrarias, UNR. CONICET, Rosario, Argentina. jvesprin@unr.edu.ar
Abstract
Resumen
González, A.M. & Vesprini, J.L. 2010. Anatomy and fruit development in Schinopsis balansae (Anacardiaceae). Anales Jard. Bot.
Madrid 67(2): 103-112.
González, A.M. & Vesprini, J.L. 2010. Anatomía y desarrollo de los
frutos en Schinopsis balansae (Anacardiaceae). Anales Jard. Bot.
Madrid 67(2): 103-112 (en inglés).
Schinopsis balansae Engl. is a dioecious tree; reproduction is by
woody samaras containing a single seed. Fruit set is high, even
in isolated trees, empty fruits are frequent and account for a
high proportion of the total fruit production. To describe the
anatomy and the ontogeny of seeded and seedless fruits, flowers and fruits at different development stages from bagged and
pollen exposed flowers were studied. The development and the
anatomy of the pericarp in seeded and parthenocarpic fruits did
not differ. It consisted in an exocarp formed sensu lato from the
external epidermis of the ovary and some layers of the underlying parenchyma. The mature mesocarp was constituted by a
sclerified tissue and lysigenous channels. The fruit is of Anacardium type: the endocarp presented three sclerenchymatic and
a crystalliferous layer. The endocarp development was also sensu lato because it was formed from the epidermis and the hypodermis of the carpel. This organization agrees with the only
species described in the genus: S. haenkeana Engl. Schinopsis
balansae is able to produce parthenocarpic fruits in the absence
of pollination. Empty fruits from free exposed flowers presented
embryos arrested at different stages, although total absence of
an embryo was the most common condition. Parthenocarpy
seems to be a plesiomorphic trait within the Anacardiaceae, but
regardless of its origin, the maintenance of empty fruit production in a wind-dispersed samara is difficult to explain. Whether
parthenocarpy has an adaptive value, or is an evolutionary constraint remains unclear in Schinopsis.
Schinopsis balansae Engl. es un árbol dioico, cuya reproducción
se realiza a través de sámaras uniseminadas. La producción de
frutos es muy alta y una gran proporción de los mismos son vanos. Con la finalidad de describir la anatomía y ontogenia de los
frutos, con y sin semillas, se trabajó con flores y frutos en distintos estados de desarrollo. Se utilizaron inflorescencias femeninas
expuestas y embolsadas. El pericarpo de frutos con semilla y partenocárpicos presenta el mismo desarrollo y anatomía: está
constituido por un exocarpo formado sensu lato a partir de la
epidermis externa del ovario y algunas capas del parénquima
subyacente. El mesocarpo maduro está constituido por esclerénquima y canales lisígenos. Por las características del endocarpo, este fruto se clasifica como tipo Anacardium, presentando tres estratos esclerenquimáticos y uno cristalífero. El desarrollo del endocarpo también es sensu lato, ya que deriva de la epidermis e hipodermis del carpelo. Esta anatomía concuerda con
la descrita para S. haenkeana Engl., única especie estudiada de
este género. S. balansae desarrolla frutos partenocárpicos aunque no se produzca polinización. Los frutos vanos producidos
por polinización libre tienen embriones en distintas etapas de
desarrollo, no obstante, la condición más común es la ausencia
del mismo. La partenocarpia parece ser un carácter plesiomórfico en las Anacardiaceae, sin embargo la producción de frutos
vanos de dispersión anemófila es difícil de explicar. Si la partenocarpia tiene un valor de adaptación o es una restricción evolutiva sigue siendo poco claro en S. balansae.
Keywords: dioecy, dry fruits, fruit ontogeny, parthenocarpy,
agamospermy, samaras, quebracho, empty fruits, pericarp.
Palabras clave: dioecia, frutos secos, frutos vanos, partenocarpia, pericarpio, quebracho, sámaras.
Introduction
called “quebrachales”. Although most species of the
Anacardiaceae family has drupaceous fruits, many
of them adapted to different dispersers, S. balansae
fruits are woody samaras with a single oblong wing
Schinopsis balansae Engl. is a dioecious tree that
grows in Argentina, Bolivia, Paraguay and Brazil,
and is the dominant species of Argentine forests
104
A.M. González & J.L. Vesprini
and an ovoid seminiferous portion containing a single
seed.
Due to timber logging its populations have suffered
a marked reduction, and part of the area formerly covered by the species has been devoted to farming. Despite the economic, ecological and social importance
of the species, a basic knowledge on its biology is still
lacking. Schinopsis reproduces only by means of seeds
and so, to determine the factors constraining fertile
fruit production, how the samara develops, and
whether or not pollination is necessary, is of fundamental importance for developing strategies for the
conservation of this species.
Wannan & Quinn (1990) studied the fruit of 29
genera in the Anacardiaceae. They recognized two basic types of endocarp: the Spondias-type, which is
composed of a mass of irregularly oriented sclerenchyma and the Anacardium-type, characterized by
a lignified outer epidermis and discretely layered and
includes palisade like sclereids. Shinopsis haenkeana
Engl was the only species studied by Wannan &
Quinn (loc. cit.) and has been described and classified
as Anacardium-type. So far the anatomic structure of
the fruit in S. balansae remains unknown.
Empty fruits are frequent and can account for almost half the total fruit production (Alzugaray, 2005).
Fruit set is striking since before maturity the red samaras are easily observed on pistillate individuals. Notably, isolated trees can also yield huge amounts of
fruit as well as plants growing in dense populations,
and these observations led us to the hypotheses that
fruit development may be independent of pollination,
and/or that embryo abortion does not prevent fruit
formation.
Atypical fruit production is frequent in the Anacardiaceae: there are cases of parthenocarpy (development
of fruit without fertilization) and fruit development following embryo abortion. For instance, Grundwag &
Fahn (1969) observed post-fertilization embryo abortion in Pistacia vera L., and Shuraki & Sedgley (1994)
found that in P. vera funicle degeneration was the most
common cause to empty seeds. More recently, Polito
(1999) found that vascular transport to ovules is
blocked at the placenta or in the funicle in P. vera. Some
authors have explained the paradox of inviable fruits
from an evolutionary viewpoint, suggesting that parthenocarpy can reduce viable seed predation (Traveset, 1993; Verdú & García-Fayos, 1998, 2002). Other
studies on sterile seeds in the Anacardiaceae occur in
the genera Anacardium L., Mangifera L., Rhus L., and
Spondias L. (Peebles & Hope, 1937; Purseglove, 1968;
Young, 1972; Crane, 1975; Janzen, 1985; Chung &
Waller, 1986; Von Teichman & Robbertse, 1986).
The aim of the present research was:
1) To describe the anatomy and ontogeny of the
pericarp.
2) To determine if fruit development is unrelated to
fertilization, and if so, to compare pericarp development in both seeded and seedless fruits.
Material and methods
Pistillate and staminate individuals growing in natural populations located at “Las Gamas” station, near
Vera, Santa Fe province, Argentina, were studied.
Material from cultivated specimens in Corrientes City,
Argentina, was also studied. Vouchers of the specimens are deposited in the Herbarium of Instituto de
Botánica del Nordeste (CTES), Argentina.
In the natural population, inflorescences at flower
bud stage were bagged with two bags: an internal
plastic mesh -to keep the outer bag free from the flowers- and an external wax paper bag to impede pollen
flow. Bagging was carried out during two flowering
periods: 2004 and 2005. One group of flowers was left
in the bags until fruit dispersal, when we determined:
the number of abscised flowers, and undeveloped and
mature fruits.
Subsets of the flowers and fruits were collected at
different developmental stages from free exposed
flowers, bagged flowers, and also from unbagged
flowers of very isolated individuals from both populations. This material was fixed in FAA (5% formalin,
5% acetic acid, and 90% ethyl alcohol), dehydrated
in Johansen’s tert-butyl alcohol series and embedded
in paraffin (Johansen, 1940). Transverse (TS) and longitudinal (LS) sections, 10-12 µm, were stained with a
safranin-Astra blue combination (Luque & al., 1986).
Histochemical tests included the FeSO 4 (Ruzin,
1999) and IKI-H2SO4 methods for tannins (Jensen,
1962), and phloroglucinol for cellulose/lignin.
To check bagging efficiency and to detect pollen
grain presence or germination, some bagged and free
exposed flowers were fixed in FAA, stained with Aniline blue (Martin, 1949) and were observed using fluorescence microscopy. The observations, drawings and
photomicrographs were made with a Leica DM LB2
microscope. For scanning electron microscopy (SEM)
observations, fresh material was fixed in FAA, dehydrated in an acetone series, dried at critical point and
coated with gold-palladium. The observations and micrographs were made with a JEOL LV 5800 at 20 kV.
Results
The flowers of S. balansae were imperfect, pentamerous, heterochlamydeous, with an apotropous
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Fruit of Schinopsis balansae
ovule and an intrastaminal nectariferous disc. The gynoecium developed only in pistillate flowers, the
ovary was superior, ovoid, unilocular and laterally
compressed with three styles placed in the middle
portion of the ovary: a central one and two lateral
ones, with a capitate stigma each (Figs. 1 A, 2 A). Occasionally gynoecia possessed either two styles (11%)
or only one ventral style (4%) (Fig. 1 B). The ovary
contained a single anatropous ovule with sub apical
placentation (Figs. 1 C, 4 A).
Stamens in pistillate flowers have been reduced to
staminodes, with two thecae and longitudinal slits
(Fig. 1 B). These staminodes were not functional: the
105
sporogenous tissue and the endothecium were not
present (Fig. 1 D). In staminate flowers the anther
thecae were tetrasporangiate and contained pollen
grains. The endothecium developed as a subepidermal layer readily distinguished from the epidermis by
its bands of secondary wall (Fig. 1 E).
Ontogeny of the ovary wall
In a floral bud, the ovary wall consisted of an external uniseriate epidermis consisting of quadrangular
cells with conspicuous nuclei (Fig. 2 B, e). The middle
zone of the carpel was made up of several layers of
Fig. 1. A-C, scanning electron microscope photographs: A, pistil with three stigmata; B, pistil with only one stigma; C, longitudinal section of ovary showing the anatropous ovule. D, E, optical microscope photographs: D, transverse section of staminod from pistillate
flowers; E, transverse section of fertile stamen from staminate flowers, note the tetrasporangiate thecae with pollen grains. Scale bar:
A, B = 2.5 cm; C = 1cm; D, E = 10 µm.
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A.M. González & J.L. Vesprini
Fig. 2. Ontogeny of the ovary wall: A, pistillate flower; B, ovary in TS, previous to the ovule development; C, young ovary in transverse
section, corresponding to the shaded area in diagram D; E, external epidermis and stomata; F, internal epidermis; G, mature ovary in
transverse section. Abbreviations: c, secretory ducts; d, druses; e, external epidermis; h, vascular bundles; hip, hypodermis; i, internal
epidermis; me, external mesocarp; mi, internal mesocarp; pr, procambial tissue. Scale bar: A = 1 cm; B, C, E-G = 10 µm; D = 0.1 mm.
Anales del Jardín Botánico de Madrid 67(2): 103-112, julio-diciembre 2010. ISSN: 0211-1322. doi: 10.3989/ajbm: 2254
Fruit of Schinopsis balansae
parenchyma cells which were isodiametric and had
thin walls; several strands of procambial tissue were
present in the central portion of the carpel (Fig. 2 B,
pr). The protoderm formed an internal unistratified
epidermis; the sub epidermal layer was differentiated
in hypodermis also consisting of only one layer (Fig. 2
B, i, hip).
During the development of the ovary (Figs. 2 C; 4
A, B), the external epidermal cells continued dividing
anticlinally and then elongated radially forming a palisade of short cells of less than 12 µm high, with thin
walls and a smooth cuticle (Figs. 2 C, e; 4 B, e). These
cells, as seen from above, were polygonal and some
anomocytic stomata were differentiated (Fig. 2 E).
The number of cellular layers was higher in the middle zone of the carpel: cells remained with thin walls
and without intercellular spaces. In the external part
of the vascular bundles apparently lysigenous secretory ducts were formed (Fig. 2 C, h, c). Cells of the internal epidermis and the hypodermis divided periclinally, forming four layers (Figs. 2, C, hip, i; 4 B, hip, i).
On superficial view, the inner epidermis presented
polygonal cells without stomata (Fig. 2 F).
General cell size increased during anthesis, with an
extensive tannin deposition. Druses appeared in the
parenchyma exterior to vascular bundles (Fig. 2 G,
me, d).
At this point, the following tissues were already differentiated through the ovarian cavity: a) a uniseriate
external epidermis (Fig. 2 G, e), b) a mesocarp divided in two parts by the vascular bundles and the secretory ducts (Fig. 2 G, me, h, c, mi) a two-layered hypodermis and internal bistratified epidermis (Fig. 2 G,
hip, i).
Fruit development
Free exposed flowers always presented pollen
grains on the stigmata. In unbagged inflorescences, almost half the mature fruits did not produce a normal
seed. Some empty fruits presented seeds with undeveloped embryos arrested at different stages, although
the lack of an embryo was the most common condition. In the population of Santa Fe, fruits from freely
pollinated flowers containing normal seeds were
24.58 and 56.66% in 2004 and 2005, respectively. In
Corrientes city, 58.4% of the fruits were normal and
the remainders were empty.
No pollen was observed on the stigmata of sampled
flowers from the bagged inflorescences, confirming
the efficacy of the bagging treatment. All of the
bagged inflorescences produced some fruits. Within
the bags it was possible to distinguish the following: a
small proportion of flowers aborted at bud stage
107
(8.5%); flowers aborted at anthesis ranged from 27.35
to 41.01%; around 9% of very small fruits of 2-5 mm
were found, and around 40% of flowers produced
seemingly normal fruits. However, the percentage of
such fruits containing fully developed seeds was 2.5%
in 2004 and 8.5% in 2005.
In both analyzed flowers -free exposed and bagged
ones- the development of the pericarp was the same.
After the petal abscission there was a rapid increase in
the ovary size whilst styles and stigmata were still present. The apical portion of the ovary grew initiating
the wing (Fig. 3 A). The uniseriate external epidermis
formed the exocarp, the cells maintaining active anticlinal divisions and increased in size forming a palisade with cells of 20-25 µm high (Fig. 3 C, ex). As the
pericarp developed, the exocarp cells acquired a lobulated outline. New stomata were formed and the surrounding cells divided anticlinally (Fig. 3 D).
The cells of the middle zone of the carpel continued increasing in number and size forming the mesocarp, the external zone of which remained composed
of elliptic to spherical cells, filled with big starch granules with stellate hilum (Fig. 3 C, me). The cells of the
internal mesocarp acquired an irregular shape with a
lobulated outline, also filled with starch granules (Fig.
3 C, mi). Parenchyma cells close to the secretory ducts
collapsed and became flattened.
The endocarp at this time consisted of four layers,
which derived from the internal epidermis and the
carpel hypodermis (Figs. 3 C, en; 4 F). The cells of the
internal epidermis formed a palisade 20-25 µm high
with very vacuolated cells, although the nuclei remained in a central position (Fig. 4 F). Since the contour of these cells changed gradually from polygonal
to undulated, as seen from above (Figs. 3 E, 4 D), it is
very difficult to identify them in a transversal section
of the pericarp (Fig. 3 C, en). As the fruit continued
developing, the cells of the internal epidermis of the
endocarp became sclereids (Fig. 4 G). The remaining
three layers of the endocarp presented elongated cells
placed periclinally (Figs. 3 C, en, 4 G): two intermediate layers formed by sclereids, and an inner layer (derived from the hypodermis and in contact with the
mesocarp) formed by parenchymatic cells (Fig. 3 C).
The mature fruit wall (Fig. 3 B, F, G) presented the
following anatomical features:
a. Exocarp: was composed of the external epidermis, with thickened and lignified cells walls. A layer of
cuticle covered the whole surface (Fig. 3 G, ex). Seen
from above, these exocarp cells had a markedly undulate outline (Fig. 4 E). The 2-4 sub-epidermal layers
belonging to the external part of the mesocarp maintained their thin walls but they became suberized,
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A.M. González & J.L. Vesprini
Fig. 3. Fruit development: A, general view of the young fruit; B, mature fruit; C, young pericarp in transverse section; D, epicarp and
stomata; E, endocarp; F, diagram showing the arrangement of tissues in transverse section of fruit; G, mature pericarp in transverse
section; H, detail of sclereid of mesocarp. Abbreviations: c, secretory ducts, cr, prismatic crystal; en, endocarp; ex, exocarp; h, vascular
bundles; me, external mesocarp; mex, corky layer; mi, internal mesocarp; st, stomata; zt, transition zone. Scale bar: A, B = 0.5 cm;
C-E, G, H = 10 µm; F = 0.15 mm.
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Fruit of Schinopsis balansae
forming a corky layer (Fig. 3 F, G, mex). Following
fruit dispersal these suberized layers became dry and
collapsed to form, with the external epidermis, the exocarp (sensu lato); stomata were somewhat sunken in
relation to the epidermal cells.
b. Mesocarp: the external portion of the vascular
bundles remained as a zone of irregular cells with thin
walls forming intercellular spaces (Fig. 3 G, me). The
cells of the internal mesocarp differentiated into sclereids (Fig. 3 G, mi; H). The limit between both layers
of the mesocarp was due to the presence of vascular
bundles always externally associated with secretory
ducts and by a transition zone where the cells changed
gradually from lobulated to isodiametric, while cell
walls became thinner (Figs. 3 F, c, h; 3 G, zt, c, h).
When the fruit was mature and dry, the whole internal
mesocarp was formed by sclereids and constituted a
third part of the body of the pericarp (Fig. 3 F, mi).
The wing of the samara consisted of the external
mesocarp, consisting of parenchymatic cells and vascularized by a net of tiny vascular bundles.
c. Endocarp: was formed by four layers: the cells of
the internal epidermis formed a palisade of macrosclereids (Fig. 3. F, G, en). Both subepidermal layers also
differentiated into sclereids of quadrangular section
with branched pits (Figs. 3 G, en, 4 H). The cells of the
inner layer of the hypodermis, with thin cellulose cell
walls, each contained a conspicuous cuboid or prismatic crystal (Figs. 3 G, 4 C). This crystalliferous layer
appeared as a natural breaking zone since the three
layers of the endocarp remained attached to the seed
when the fruit was opened manually.
Empty fruits presented a seminal cavity delimited
by the three layers of the endocarp which contained a
vestigial ovule. A very long funiculus was present although it did not show any degree of nucellar development, suggesting the presence of an aborted embryo.
There were no differences in the initial pericarp development between bagged and unbagged flowers
nor did we detect anatomical or morphological differences between the pericarp of empty fruits and fruits
containing seeds. Before maturity, both kinds of fruits
were initially red, becoming brown when dry.
Discussion
According to the endocarp organization, Wannan
& Quinn (1990), proposed a classification of the different types of pericarp in the family Anacardiaceae.
The genus Schinopsis was classified as Anacardiumtype, based only in a brief description of S. haenkeana,
the endocarp was regularly guided and consists of
109
four layers of cells: an internal crystalliferous layer and
three external layers formed by sclereids in palisade.
The presence of a lignified outer epidermis is restricted to this type of endocarp. The exocarp of S. haenkeana was formed by 6 layers: the outer lignified epidermis plus 5 underlying parenchyma layers, the
mesocarp was almost completely lignified. The ontogeny of the fruit of S. haenkeana was not reported by
Wannan & Quinn (1990).
Following the Wannan & Quinn’s classification, the
endocarp of S. balansae presented an Anacardiumtype. The characteristic of their pericarp agrees with
the description of S. haenkeana.
According to Roth (1977) the endocarp or exocarp
could be formed by the epidermis of the ovary and its
immediate derivates, in which case it was referred to
as “sensu stricto”; if the endocarp/exocarp also includes layers of the mesocarp then it was referred to as
“sensu lato”. Our ontogenetic study confirm that the
exocarp and the endocarp in S. balansae were formed
sensu lato, i.e. the exocarp was made up of the external epidermis of the carpel, which contains cells with
thickened walls, and suberized layers originated from
the underlying parenchyma. The endocarp was
formed by four layers derived from the divisions of
the internal epidermis and carpel hypodermis, the latter not derived from the protodermis.
A feature of seeds contained within a lignified pericarp is that the function of the seed cover is transferred to the pericarp, and the episperm is relatively
undeveloped (Boesewinkel & Bouman, 1984). This
characteristic also appeared in the samaras of other
species i.e., Tipuana tipu (Benth.) Kuntze (Martins &
Oliveira, 2001) and Pterodon emarginatus Vogel,
Leguminosae (Oliveira & Paiva, 2005). In S. balansae
the protective functions of the seed coat are transferred to the endocarp, indeed the columnar cells of
the endocarp resemble the cells commonly encountered in seed testa. This transfer of functions has already been described as a generalized condition for
the family (Corner, 1976). The irregular arrangement
of the exocarp cells, with their lobulated shape and a
densely interlocked pattern, and their highly lignified
cell walls, explain the indehiscence of the S. balansae
fruit.
Schinopsis balansae is a parthenocarpic species: it
produces a great amount of fruits, many of them lacking seeds. Bagging treatment confirmed that the absence of pollen do not prevent fruiting. Self pollination is not possible because anthers of pistillated flowers -apparently normal- do not produce sporogenous
tissue and there is no pollen formation. The low percentage of seeded fruits produced under pollen exclu-
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A.M. González & J.L. Vesprini
sion should be checked for eventual maternal progeny
production.
Seed abortion at different development stages did
not lead to fruit abscission in this species. This is a
common condition in the family. Parthenocarpy in
Anacardiaceae is due to several causes: a) pre fertilization including funicle or embryo sac degeneration,
embryo sac absence, vascular transport to ovules
Fig. 4. Optical microscope photographs: A, longitudinal section of pistillate flower at anthesis; B, longitudinal section of ovary at floral anthesis; C, polarized light photograph of crystalliferous layer of endocarp; D, superficial view of endocarp; E, superficial view of
exocarp; F-H, transverse section of endocarp in three different stages of development, showing the formation of sclereids; F, endocarp in young fruit, phase of cell division; G, endocarp in immature fruit; H, endocarp in mature fruit. Abbreviations: e, external epidermis; hip, hypodermis; i, internal epidermis. Scale bar: A = 0.5 cm; B = 20 µm; C-H = 10 µm.
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Fruit of Schinopsis balansae
blocked at the placenta or in the funicle (Polito,
1999); b) lack of fertilization and of pollen tube penetration in the embryo sac (Shuraki & Sedgley, 1994)
and c) post fertilization failure of embryo development or cellularisation of the endosperm (Grundwag
& Fahn, 1969; Ram & al., 1976; Shuraki & Sedgley
1994).
In the current study we found normal fruits containing ovules, aborted embryos at different stages of
development and seeds. Further studies -ecological
and ontogenetic- will be necessary to figure out the
several steps and causes of seed abortion.
Parthenocarpy seems to be a plesiomorphic trait
within the Anacardiaceae. In other reported cases, in
species with fleshy, bird-dispersed fruits, it has been
suggested that empty fruit retention could be advantageous presumably in helping to attract the seed dispersers, and this may be the case in genera with endozoochorous drupe or drupaceous fruits (Traveset,
1993; Verdú & García-Fayos, 1998; 2002).
Stenospermocarpy is a term describing a small
seedless fruit caused by embryo abortion after fertilization. This characteristic was fully described in mango (Mangifera indica L.), and related to environment
conditions, specially the temperatures during pollination or early fruit set (Lakshminarayana & Aguilar,
1975; Davenport & Núñez-Elisea, 1983; Soule, 1985;
Whiley & al., 1988).
The occurrence of partenocarpic samara seems
rather difficult to explain. However, like Schinopsis,
species of the genera Astronium Jacq. and Loxopterygium Hook. fil. (Anacardiaceae) have wind dispersed
seeds, and the presence of parthenocarpy in these genera should be investigated. Whether parthenocarpy
has an adaptive value or otherwise an example of phylogenetic baggage remains unclear in Schinopsis.
Acknowledgements
We thank T. Ghiglione and G. Venturi for help with the English text. Research was carried out under grant PICT 11941 “Fondo para la investigación científica y tecnológica” to J. Vesprini.
Anonymous reviewers made helpful improvement to the manuscript.
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Associate Editor: S. Talavera
Received: 10-III-2010
Accepted: 7-X-2010
Anales del Jardín Botánico de Madrid 67(2): 103-112, julio-diciembre 2010. ISSN: 0211-1322. doi: 10.3989/ajbm: 2254