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Accepted by B.J. Sinclair: 10 Aug. 2012; published: 19 Sept. 2012
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2012 · Magnolia Press
Zootaxa 3489: 81–88 (2012)
www.mapress.com/zootaxa/
Article
81
urn:lsid:zoobank.org:pub:9D26AF16-B9E1-4E37-99B3-75703DFA986C
Systematic position of the monotypic Azorean genus Falbouria Dyte with notes
on the definition of Chrysotus Meigen (Diptera: Dolichopodidae)
RENATO SOARES CAPELLARI1,2 & DALTON DE SOUZA AMORIM1
1Universidade de São Paulo, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Departamento de Biologia, Av.
Bandeirantes 3900, 14040–901, Ribeirão Preto, SP, Brasil
2Corresponding author: rscapellari@gmail.com
Abstract
The systematic position and the generic status of the monotypic genus Falbouria Dyte (Dolichopodidae: Diaphorinae) are
evaluated. The type-species of the genus, F. a co ren si s (Parent), endemic to the Azores, is re-described, and hypopygial
and oviscapt morphology illustrated in detail for the first time. Falbouria is kept as a separate genus, but close to Chrysotus
Meigen, which is certainly paraphyletic (or even polyphyletic) in its present composition. A brief discussion on the
delimitation of Chrysotus is provided and its laesus-group of species is suggested as the core clade of the genus.
Key words: Diptera, Dolichopodidae, Diaphorinae, Balfouria, Falbouria, Chrysotus, Palaearctic Region, Azores
Introduction
Parent (1933) proposed the genus Balfouria for an endemic Azorean species, B. acorensis Parent, based on a single
male specimen. He included the species in the Diaphorinae on the basis of its general habitus, strong bristles on the
end of abdomen and wing venation, distinguishing it from the remaining genera of the subfamily by the presence of
two “postvertical” setae. The genus-group name was later verified to be preoccupied in Mollusca (Crosse 1884)
and the replacement name Falbouria was proposed by Dyte (1980). Maslova and Negrobov (1996) placed
Falbouria in the Argyrini (sensu Negrobov 1986), as sister to the remainder of the tribe (i.e., the genera Argyra
Macquart, Keirosoma Van Duzee, Pseudargyra Van Duzee and Somillus Brèthes), a position later endorsed by
Grichanov (1998).
Quite recently an unpublished manuscript by C.E. Dyte came to our attention, in which he pointed out that
Frey’s (1945) treatment of the Azorean Diptera overlooked earlier papers by Parent (1933, 1934). Frey (1945)
recorded Chrysotus xanthoprasius from the Azores, a species originally described by Bezzi (1906) from Eritrea
(Afrotropical Region). Dyte, however, suspected that the species studied by Frey (1945) could actually be F.
acorensis. We have examined Frey’s specimens of “C. xanthoprasius”, currently in the collection of the Finish
Museum of Natural History (Finland), and the holotype of F. acorensis, confirming Dyte’s suspicion. In this paper,
Frey’s specimens and additional material of the same species recently collected in São Miguel Island (Azores) were
used to re-describe F. acorensis, which allowed considering the problem of the systematic position Falbouria and
the delimitation of Chrysotus Meigen.
Material and methods
The morphological nomenclature used here follows mainly Cumming and Wood (2009). Body length was
measured from the insertion of the antenna to the apex of the abdomen. Wing length was measured from the base to
the apex of the wing, while the width was measured at the widest point of the wing, both given as ranges.
Measurements of the podomeres are representative ratios and are given according the formula: trochanter + femur,
CAPELLARI & AMORIM
82 · Zootaxa 3489 © 2012
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tibia, tarsomeres 1, 2, 3, 4, 5. The CuAx ratio represents the length of the dm-cu crossvein/distal section of the anal
vein.
The following abbreviations are used in the description and figures: I, II, III: pro- meso-, metathoracic legs; A1,
anal vein; C, costal vein; cerc, cercus; CuA, basal part of anterior branch of cubital vein; Cx, coxa; dm-cu, discal
medial-cubital crossvein; F, femur; lel, lateral epandrial lobe; M, medial vein; MSSC, male secondary sexual
character(s); pgon, postgonite; ph, phallus; R, radial vein; St, sternite; Tg, tergite; VSur, ventral lobe of surstylus; t1-
5, tarsomeres 1 to 5. While describing the hypopygium, ‘dorsal’ and ‘ventral’ refer to the morphological position
prior to genitalic rotation and flexion; as such, the top of the drawings in lateral view is actually ventral on the
specimens. Habitus and wing photographs were produced using a Leica DC camera attached to a Leica MZ16
stereomicroscope, AutoMontage and Photoshop CS4.
The male holotype of Falbouria acorensis is housed in the Natural History Museum, London, UK (BMNH),
while additional material used in this study belongs to the Silesian Museum, Opava, Czech Republic (SMOC)
(collected by J. Rohá
č
ek), and the Finish Museum of Natural History, Helsinki, Finland (UZMH) (collected by R.
Frey and R. Storå).
Genus Falbouria Dyte
Balfouria Parent, 1933: 74. Type-species: Balfouria acorensis Parent, 1933: 75 (by monotypy). Preoccupied by Balfouria
Crosse, 1884 [Mollusca].
Falbouria Dyte, 1980: 223. Type-species: Balfouria acorensis Parent (automatic). Replacement name for Balfouria Parent,
1933.
Diagnosis (Male). Face obliterated by contiguous eyes (MSSC); scape of antenna bare, postpedicel subtriangular,
apical stylus bi-articulate near base. Acrostichals biseriate, 6 pairs of dorsocentrals. Upper part of proepisternum, in
front of anterior spiracle, bare. FII and FIII with row of anterior setae near apex; claws present in all legs, pulvilli of
usual normal size (not developed as MSSC). Tg6 pilose, hind margin with strong setae, longer than those on St8;
Tg7 and St7 strongly reduced. Hypopygium encapsulated; surstylus as a single lobe; lateral epandrial lobe reduced;
cercus short.
Comments. In spite of the rows of anterior setae at the apex of FII and III, these are not true pre-apical setae,
as those seen in Sympycninae and other subfamilies of Dolichopodidae. Indeed, Falbouria shows a typical
Chrysotus-like habitus and, in most respects, F. acore nsis could be regarded as a Chrysotus with stronger
chaetotaxy. Parent’s (1933) claim that two postverticals—“postverticaux” in Parent (1938) are paravertical setae in
Cumming and Wood (2009)—were of generic value to separate his genus proved to be misleading, as also
displayed by other diaphorines. Strong setae on femora and tibiae seem more useful to recognize the genus while
comparing it with species of Chrysotus (but see discussion below).
Falbouria acorensis (Parent)
(Figs. 1–7)
Balfouria acorensis Parent, 1933: 75 (fig. 1).
Falbouria acorensis (Parent): Dyte, 1980: 223. Grichanov, 2010: figs. 8–9 (habitus of holotype).
Diagnosis. As for the genus.
Material examined. HOLOTYPE
♂
, AZORES, Pico Island, Serra Gorda, 24.vii.1929, J. Balfour-Browne
leg. (BMNH). Additional material: 1
♂
, 4
♀
, São Miguel Island: 2 km S Pico da Pedra, 37º47'N, 25º35'W,
1.xi.2006 (sweeping undergrowth of deciduous forest) (SMOC); 1
♀
, Caldeira Velha (near Ribeira Grande),
37º47'N, 25º30'W, 7.ix.2006 (sweeping undergrowth of broadleaved forest) (SMOC); 1
♀
, same data except
Lombadas, Ribeira Grande river, 37º47'N, 25º27'W, 5.ix.2006 (sweeping riverside vegetation) (SMOC); 1
♀
, Sete
Cidades env., 37º51'N, 25º48'W, 31.viii.2006 (sweeping over meadow) (SMOC); 1
♂
, 1.5 km SE Sete Cidades,
37º51'N, 25º47'W (on refuse heap in mixed forest) (SMOC); 1
♀
, Sete Cidades, Lagoa Verde, 8.ix.2006 (sweeping
undergrowth of deciduous forest) (SMOC); 3
♂
, Sete Cidades, 17.v.1938 (UZMH); 2
♀
, same data except Santo
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FALBOURIA AND DEFINITION OF CHRYSOTUS
António, Atalhada res., 37º51'N, 25º13'W, 6.ix.2006 (sweeping undergrowth of laurel forest) (SMOC); 1
♀
, Furnas,
19–21.v.1938 (UZMH); 1
♀
, Furnas, 23.vii–1.viii.1938 (UZMH); 1
♀
, Lagoa do Congro, 21.v.1938 (UZMH). Faial
Island: 1
♂
, Caldeira, 4.vii.1938 (UZMH). Flores Island: 1
♀
, Caldeira Seca, 21.vi.1938 (UZMH); 1
♀
, Fazenda
Ribeira, 8.vi.1938 (UZMH); 1
♀
, Mato, 15.vi.1938 (UZMH). São Jorge Island: 2
♂
, Ribeira do Salto, 23.vi.1938
(UZMH); 1
♂
, Ribeira Funda, 23.vi.1938 (UZMH).
FIGURES 1–3. Falbouria acorensis (Parent). 1, male habitus. 2–3, wings: 2, male. 3, female. Scale bars: 1.0 mm.
Re-description. Male (Fig. 1). Body length, 2.8–3.5 mm. Wing length, 2.8–3.2 mm, width, 1.1–1.2 mm.
Head. Frons light green, with some bluish reflections, about 0.3 of head width, converging ventrally; face mostly
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obliterated by contiguous eyes, reduced to small triangle below antennae, concolorous with frons; eyes with ventral
facets enlarged (MSSC); palpus yellow, with black apical seta; proboscis brown, with brownish setulae; pair of
divergent, proclinate strong ocellar setae; pair of minute post-ocellar setae; pair of convergent, proclinate strong
vertical setae; pair of paravertical setae, almost as long as dorsalmost post-oculars; single row of white post-
oculars, 7 dorsalmost post-oculars black. Antenna light brown; dorsal surface of scape bare; pedicel with crown of
apical setae, dorsalmost longer; postpedicel sub-triangular, pubescent; arista-like stylus bi-articulated near base,
also pubescent. Thorax. Mesonotum and most of anepisternum and anepimeron metallic green with coppery
reflections, with some pruinosity laterally; katepimeron brownish; metepimeron yellow. Acrostichal setae
distinctly biseriate, extending to fifth pair of dorsocentrals; 6 pairs of dorsocentrals; scattered setulae on anterior
surface of scutum, around second pair of dorsocentrals; 1 strong and 2 smaller post-pronotals; upper part of
proepisternum, in front of anterior spiracle, bare, lower part with 2 setae, ventral one stronger; 1 pre- and 1 sutural
intra-alar setae; 1 pre- and 2 post-sutural supra-alar setae; 1 post-alar and 1 tiny hair anteriad; 2 notopleurals; pair
of strong medial scutellars and 1 pair smaller laterad, about one fourth length of medial scutellars. Wing (Fig. 2).
Membrane hyaline, slightly brownish anteriorly, veins brown. C ending at wing apex. R1 ending at basal fifth. R2+3
ending at apical 4/5. R4+5 and M slightly diverging at apex. CuAx ratio: 0.3. A1 present. Anal angle undeveloped.
Lower calypter and halter yellowish. Legs. I: 42, 36, 29, 10, 9, 6, 4. II: 49, 48, 28, 14, 10, 6, 3. III: 59, 60, 18, 17,
11, 6, 4. Mostly yellow, CxII, apical 1/4 of FIII, apex of tibia III, and tarsus III brown. Claws present in all legs;
pulvilli undeveloped. I. Anterior surface of CxI covered by black setae, lateral margin and outer edge with stronger
setae; FI with 1–2 small anterior setae at apex; tibia I with 2–4 dorsal setae, 1 antero-dorsal seta at basal fourth, 2
small ventral setae on apical half and 3 small apical setae; It1 with short ventral setae. II: FII with row of 4
conspicuous anterior setae on apical half and 1 postero-ventral at apex; tibia II with 4–5 antero-dorsal setae, 3–4
postero-dorsal setae, 2 ventral setae on apical half and 4 apicals; IIt1 with short ventral setae. III: CxIII with 1 large
lateral setae at base and 1 smaller at apex; FIII with 3–4 conspicuous anterior setae on apical 1/4, with row of
antero-ventral setae, 3 apicalmost setae stronger; tibia III with 3–4 antero-dorsal setae, 4–5 postero-dorsal setae and
4 apicals; ventral surface with row of 5–6 more conspicuous setae. Abdomen. Cylindrical, Tg1–3 yellow
(sometimes only Tg1 or Tg1–2), otherwise shinning dark green, with coppery reflections; St1–5 brownish, St1
lighter, St6 concealed. Tg1–6 covered by black setae, those on hind margins longer (setae on Tg6 extending beyond
apex of abdomen and longer than setae on St8; Fig. 1). St8 only with short setae, without strong bristles (as in
species of Diaphorus). Hypopygium (Figs. 4–5). Globular, brown, partially hidden by Tg6, foramen left lateral.
Surstylus as a single lobe (=VSur), laterally flattened, with a small, drop-like spine at apex. Lel reduced, with 3
setae. Dorsal surface of distal part of phallus rugose; proximal end of phallus with a short projection, arising dorsad
sperm pump. Pgon pilose, extending between cerci. Cercus short, brownish.
Female. Body length, 2.8–3.5 mm. Wing length, 2.7–3.7 mm, width, 1.1–1.4 mm. Similar to male except for
MSSC and as noted. Head. Face pruinose, as broad as narrowest width of frons; palpus brown, covered by black
setulae. Antennal postpedicel trapezoid. Wing (Fig. 3). Legs. I: 40, 37, 22, 10, 6, 4, 3. II: 50, 49, 28, 12, 9, 5, 3. III:
61, 60, 18, 13, 9, 5, 4. Abdomen. Narrowing towards apex, yellow band restricted to lateral margins of Tg1–2.
Setae on hind margin of Tg5 brownish and delicate. Oviscapt (Figs. 6–7). Tg8 and St8 divided into two rod-like
sclerites; acanthophorites with 3 spines and 1 lateral seta each.
Comments. As stated above, part of the material examined in this paper was misidentified by Frey (1945) as
the Afrotropical dolichopodid species Chrysotus xanthoprasius. Hence, records of that species (as “C.
xanthoprasinus”) from Azores by Negrobov (1991) and Yang et al. (2006) should be corrected. Apparently, this
also applies to the Italian records for this species (C.E. Dyte, unpublished manuscript). Therefore, C. xanthoprasius
must no longer be regarded as having a Palaearctic extension.
Discussion
Maslova and Negrobov (1996) placed Falbouria in the Argyrini (sensu Negrobov 1986), but this is not supported
by the re-description of the type-species of the genus made here. General external morphology as well as
hypopygial characters of F. acorensis show a number of features shared with Achradocera Becker, Chrysotus
Meigen and Lyroneurus Loew. Some of these features have been used by Robinson and Vockeroth (1981) to
distinguish Chrysotus and Achradocera from Diaphorus Meigen (see couplet 16 in their key), such as the upper
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FALBOURIA AND DEFINITION OF CHRYSOTUS
part of proepisternum bare, male face narrowed or parallel-sided, male Tg6 setose, sometimes with strong marginal
setae, and setae on St8 not longer nor stronger than those on Tg6. Additionally, hypopygial characters seen in
Falbouria and regarded as apomorphic within the diaphorines (sensu Ulrich 1981) are also found in Achradocera,
Chrysotus and Lyroneurus. These include epandrial lobe reduced, dorsal lobe of the surstylus absent, ventral lobe
of the surstylus usually with an apical spine, postgonites pilose, and proximal part of the phallus with a short
projection. So there is no doubt about Falbouria being a Diaphorinae.
The fact that Falbouria probably belongs to a clade with these other diaphorine genera, but does not fit into the
diagnoses neither of Achradocera nor of Lyroneurus raises the question of keeping Falbouria as a separate genus
or merging it into Chrysotus. This decision actually involves the problem of circumscription of Chrysotus, a still
ill-defined genus, with about 300 species worldwide. Part of this problem has to do with the recognition of a clade
within the genus Chrysotus including its type-species, the Palaearctic Musca nigripes Fabricius. The type-
specimen of this species, however, is presently unrecognizable: there is a single wing of the type left on the pin
(currently housed in the Natural History Museum of Denmark, Copenhagen; T. Pape, pers. comm.). The
designation of a neotype would help to fix the identity of this species, but in the presence of a type, even if
unrecognizable, an application to the I.C.Z.N. would be mandatory. This is, of course, beyond the scope of this
paper. Some remarks on the identity of Musca nigripes, however, can be made here in order to shed at least some
light on the systematic position of Falbouria.
Kowarz (1875) (about dating of Kowarz’s works, see Pont 2011), while describing Chrysotus varians Kowarz,
noted similarities between his species and Meigen’s concept of C. nigripes. He considered that Meigen’s
interpretation of Fabricius’ species was likely based on a mix of specimens of C. microcerus Kowarz and C.
varians—currently synonyms of C. gramineus Fallén (Negrobov 1991). In his words (Kowarz 1875: 472):
“Chr. nigripes F. bei Meigen scheint mir eine Mischart von Chr. microcerus m[ihi]. und obigem
[Chr.] varians zu sein; Meigen's Beschreibung wenigstens passt viel eher auf die beiden
letztgenannten als auf irgend eine andere der bisher bekannten Arten, besonders aber auf
[Chr.] varians, wenn man die „metallisch grüne" Körperfärbung und die „ziegelrothe oder
rostgelbe" Färbung der Hinterbeine auf das Männchen desselben bezieht. Ich war desshalb im
Zweifel, ob für [Chr.] varipes [sic, =nigripes] nicht der ältere Fabricius'sche Name zu wählen
sei (…)”
[“Chr. nigripes F. as recognized by Meigen seems to me a mixed species of Chr. microcerus
m[ihi]. with the above described [Chr.] varians; Meigen’s description at least fits much more in
both above mentioned than any other of the known species, but particularly in [Chr.] varians,
when the “metallic green” body colour and the “brick-red or the rust-yellow” colour of the hind
legs refers to males of that species. I was, hence, in doubt if the older Fabricius’ name varipes
[sic, =nigripes] should not be chosen (…)”]
Kowarz (1875) proceeds with his inspection considering Fabricius’ (1794) description of Musca nigripes
itself, concluding that both Chrysotus laesus Wiedemann and C. amplicornis Zetterstedt fit better with Fabricius’
description of C. nigripes:
“(…) da jedoch sowohl die Benennung als auch die Beschreibung Fabricius' (…) höchstens
auf Chr. laesus Wied. oder auf Chr. amplicornis Ztt., nicht aber auf die von Meigen
beschriebene Art hindeutet, und weil mir die Bestimmung Wiedemann's nicht genügende
Sicherheit bot, so entschied ich mich für einen neuen Namen.”
[“(…) however, because Fabricius’ designation and description fit better in Chr. laesus Wied.
or in Chr. amplicornis Ztt., rather than Meigen’s species, and also because Wiedemann’s
determination [of Chr. laesus] does not seem safe enough to me, I decided to propose a new
name.”]
Based on Kowarz’s (1875) considerations, it seems reasonable to assume that Fabricius’ Musca nigripes may
be a senior-synonym of Chrysotus laesus (C. amplicornis as synonym; see Negrobov 1991). Indeed Negrobov
(1991) listed Fabricius’ species under C. laesus as “a questionable synonym”, while Yang et al. (2011) treated both
as synonyms.
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FIGURES 4–7. Falbouria acorensis (Parent). 4–5, male hypopygium: 4, left lateral. 5, ventral. 6–7, female oviscapt: 6, dorsal.
7, lateral. Scale bars: 0.1 mm.
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FALBOURIA AND DEFINITION OF CHRYSOTUS
Wei and Zhang (2010) revised and re-defined Negrobov’s (1980) laesus-group of Chrysotus, and considered
the posterior angular projection of the epandrium as a major diagnostic feature, apomorphic in all these species. We
studied specimens of C. gramineus from England and Germany, and confirmed that this species also possesses the
angular projection of the epandrium (although not so pronounced), contrasting with the illustration by Kahanpää
and Grichanov (2006: fig. 1b). If confirmed that C. gramineus shows some variation of this character, the species
can also be regarded as belonging to the laesus-group sensu Wei & Zhang (2010). As such, both suspicions of
Kowarz (1875) about the identity of Musca nigripes would lead to its inclusion in the laesus-group, most probably
C. laesus itself. Therefore it seems justifiable to regard the Palaearctic laesus-group as the core clade of Chrysotus
for taxonomical purposes, very likely including the unrecognizable type-species of the genus. Maybe
synonymization of Musca nigripes under C. laesus proves to be warranted in the future, in order to preserve the
well-established usage of C. laesus over Fabricius’ name priority.
Falbouria clearly does not fit in the diagnosis of the laesus-group (sensu Wei & Zhang 2010). The angular
epandrial projection and apical phallic modifications (another apomorphic condition in the group and also in C.
gramineus) are both absent in F. a co ren s is (Figs. 4–5). Besides, setae on the posterior margin of Tg6 longer than
those arising from St8 are seen in some Neotropical species of Chrysotus—as C. angustifrons (Robinson), C.
robustus (Robinson), and C. wirthi (Robinson) (see Capellari & Amorim 2010)—and in Falbouria (Fig. 1), but not
in any species of the laesus-group. Robinson (1964) noted that these setae are very likely valuable as evidence of
relationships. These conditions seem to be obviously apomorphic, but acceptance of a clade including Falbouria
and the above mentioned Neotropical species of Chrysotus still depends on a wider study of the genus. Particularly
F. a co re ns is displays long setae covering body as a whole and, as such, interpretation on the setation of Tg6 could
be biased, i.e., this apomorphy could be convergent. We have also examined specimens identified by Richard Frey
as Chrysotus barretoi Becker from Madeira Island (UZMH), which he supposed to be close to his “Chrysotus
xanthoprasius” (Frey 1945), but they lack long marginal setae on Tg6 and major conspicuous setae on legs as well.
This condition was also verified in one unidentified species of Chrysotus from São Miguel Island (SMOC).
The conclusion so far is that the type-species of Falbouria probably does not belong to the core clade of
Chrysotus (i.e., including the type-species of the genus), and F. a co re ns is better fits elsewhere, in one of the other
species groups of Chrysotus. This evidently makes Chrysotus paraphyletic, but the genus is already paraphyletic
regarding other diaphorine genera, as Achradocera (Bickel 2000) and Lyroneurus (Capellari & Amorim 2010).
Even though we do not conceptually endorse the presence of paraphyletic taxa in classifications, making Falbouria
synonym of Chrysotus without a comprehensive study of the latter would reduce the information now available
within the group, with no real benefit to the overall classification of the subfamily. So for the time being we suggest
keeping the generic status of Falbouria. The division of Chrysotus in the future into smaller, monophyletic
subgroups with generic status probably will demand the use of the name Falbouria. Further investigation on other
species from the Macaronesian region may show if any of them are close to F. a co re ns is , nested in a clade aside
from the core group of Chrysotus.
Acknowledgements
Dan Bickel (Australian Museum, Sydney) shared his copy of the unpublished manuscript by C.E. Dyte. He and
Scott Brooks (Canadian National Collection, Ottawa) provided useful suggestions on an early version of the
manuscript. Jind
ř
ich Rohá
č
ek (SMOC) and Pekka Vilkamaa (UZMH) facilitated loan of specimens. Peter Chandler
(England) and Hans Meyer (Christian-Albrechts-Universität zu Kiel, Germany) kindly donated specimens of
Palaearctic Chrysotus gramineus and C. laesus for examination. Erica McAlister (BMNH) facilitated access to the
holotype of Falbouria acorensis. Thomas Pape (Natural History Museum of Denmark) provided valuable
information on Fabricius’ type of Musca nigripes. Neal Evenhuis (Bishop Museum, Hawai’i) helped with dating of
Kowarz’s work. Maria Isabel P.A. Balbi carefully prepared the specimens for examination. During preparation of
this paper, RSC benefited from FAPESP grants 2003/10274-9 and 2008/58224-3, while DSA was supported by a
research fellowship from CNPq (314371/2009-5).
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