Flora 240 (2018) 89–97

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Flora
journal homepage: www.elsevier.com/locate/flora

How to distinguish cavities from ducts in Casearia Jacq. (Salicaceae): T

Anatomical characterization and distribution
Valéria Ferreira Fernandesa,b, Marcela Thadeoc, Valdnéa Casagrande Dalvid,
Ronaldo Marquetee,f, Julianna Xavier de Brito Silvaa, Luana de Jesus Pereiraa,
⁎
Renata Maria Strozi Alves Meiraa,
a
Universidade Federal de Viçosa, Departamento de Biologia Vegetal, Programa de Pós-Graduação em Botânica, Av. P.H. Rolfs, s/n, Campus Universitário, CEP: 36.570-
900, Viçosa, Minas Gerais, Brazil
b
Programa de Pós-graduação em Botânica, Departamento de Ciências Biológicas, Universidade Estadual de Santa Cruz, Ilhéus, 45.662-900, Bahia, Brazil
c
Universidade Estadual de Maringá, Departamento de Biologia, Av. Colombo, 5790, Jardim Universitário, CEP 87.020-900, Maringá, Paraná, Brazil
d
Instituto Federal de Educação, Ciência e Tecnologia Goiano – Campus Rio Verde, Rod. Sul Goiana Km 01, Zona Rural, CEP: 75. 901-970, Caixa Postal 66, Rio Verde,
Goiás, Brazil
e
Instituto Brasileiro de Geografia e Estatística, Av. República do Chile 500, 7° andar, CEP: 20.031-170, Rio de Janeiro, Rio de Janeiro, Brazil
f
Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rua Pacheco Leão, 915, CEP: 22.460-030, Rio de Janeiro, Rio de Janeiro, Brazil

A R T I C L E I N F O A B S T R A C T

Edited by: Alessio Papini Pellucid punctuations and lines in the leaf blade are common in Casearia, but for some of its species these
Keywords: structures are mentioned as inconspicuous, not visible or obscure. Cavities and ducts have been anatomically
Leaf blade described and correlated with pellucid punctuations and lines, respectively, in only six species from the genus;
Pellucid punctuations however, the distinction between these types of secretory structures remains unclear. Therefore, we aimed to
Lines anatomically describe the pellucid punctuations and lines in leaves of 47 species of Casearia. Leaf blade samples
Secretory structures were obtained from herbarium and field collected material and processed according to standard methods for
light microscopy analysis. Cavities and ducts were absent in only four of the 47 species (C. commersoniana, C.
javitensis and C. spruceana of section Piparea, and C. eicheleriana of section Casearia). In the remaining species,
cavities corresponded to pellucid punctuations, while ducts corresponded with lines. The majority of species
have both structures randomly distributed throughout the leaf blade, however, in C. bahiensis, these structures
are concentrated along the leaf margin, while in C. aquifolia and C. crassinervis they were predominant along the
midrib. Our data show that anatomical analyses are essential for distinguishing between cavities and ducts. We
also demonstrate that the distribution of cavities and ducts provides useful information for the taxonomy of
Casearia at both the species and section levels.

1. Introduction
Cavities (sacs) and ducts (canals) are internal secretory structures of
plants that consist of a space (lumen) bounded by a layer of secretory
cells (epithelium), which produce the secretion (Cutter, 1978; Fahn,
1979; Lersten and Curtis, 1989; Evert, 2006). Among previous anato-
mical descriptions of cavities and ducts, Fahn (1979) considered
structures with a more or less spherical lumen as secretory cavities
while those with an elongated lumen as ducts; Cutter (1978) referred to
cavities that are more or less isodiametric as glands while those that are
considerably elongated in one plane as ducts; and Evert (2006) re-
cognized that secretory cavities are short secretory spaces and ducts are
long secretory spaces. However, the lack of a clear method for
evaluating the dimensions of the lumen has made distinguishing ducts
and cavities difficult. As a result, alternative denominations have been
used, such as: elongate cavity in Porophyllum lanceolatum (Monteiro
et al., 1995); oil reservoir in Solidago canadensis (Lersten and Curtis,
1989); and tubular cavities in Eupatorium rugosum (Lersten and Curtis,
1986), species belong to Asteraceae, and Casearia decandra (Thadeo
et al., 2009), species belong to Salicaceae and secretory spaces as sy-
nonyms of cavities and ducts in species of the clade Dipterygeae, Le-
guminosae (Palermo et al., 2017).
Cavities and ducts are found in several organs, such as leaves, stems,
roots and flowers (Lersten and Curtis, 1989; Ciccarelli et al., 2001;
Sant'Anna-Santos et al., 2006; Ciccarelli et al., 2008). They are im-
portant taxonomic characters for many plant families (Solereder, 1908;

⁎
Corresponding author.
E-mail address: rmeira@ufv.br (R.M. Strozi Alves Meira).

https://doi.org/10.1016/j.flora.2018.01.007
Received 9 August 2017; Received in revised form 11 January 2018; Accepted 16 January 2018
Available online 31 January 2018
0367-2530/ © 2018 Elsevier GmbH. All rights reserved.
V. Ferreira Fernandes et al. Flora 240 (2018) 89–97

Table 1
Distribution of cavities and ducts in the leaf blade of species of Casearia Jacq. (Salicaceae).

Cavities/Ducts Species Absent Presence

SECTIONS Numerous and Numerous and randomly Concentrated in the midrib In the In the medulla
concentrated on the leaf distributed in the leaf and rarely present in the phloem in the midrid
margin blade mesophyll region

Casearia (35/61) C. aculeata x

C. altiplanensis x
C. aquifolia x
C. arborea x x
C. bahiensis x
C. combaymensis x
C. corymbosa x
C. cotticensis x
C. crassinervis x
C. decandra x
C. duckeana x
C. eichleriana x
C. espirotosantensis x
C. fasciculata x
C. grandiflora x x
C. guianensis x
C. hirsuta x
C. lasiophylla x
C. manausensis x x
C. mariquitensis x
C. melliodora x
C. mestrensis x x
C. negrensis x
C. nigricans x
Casearia (35/61) C. nitida x
C. oblongifolia x
C. obovalis x
C. paranaensis x
C. pauciflora x
C. pitumba x
C. resinifera x x
C. rufescens x
C. rupestris x
C. sessiliflora x
Crateria (3/4) C. selloana x x
C. obliqua x
C. sylvestris x x
Endoglossum (1/1) C. tremula x
Gossypiorpermum (3/3) C. gossypiosperma x
C. luetzelburgii x
C. praecox x
Guidonia (2/4) C. bartlettii x
C. spinescens x
Piparea (3/3) C. commersoniana x
C. javitensis x
C. spruceana x

Metcalfe and Chalk, 1950), but their usefulness is limited by the lack of
a clear distinction between them. For instance, cavities are common in
Asteraceae (Anderson and Creech, 1975; Castro et al., 1997), Legumi-
nosae (Teixeira et al., 2000; Sartori and Tozzi, 2002; Palermo et al.,
2017), Rutaceae (Turner et al., 1998; Bennici and Tani, 2004;
Muntoreanu et al., 2011), Myrtaceae (Fontenelle et al., 1994; Cardoso
et al., 2009) and Hypericacacae (Curtis and Lersten, 1990; Ciccarelli
et al., 2001), whereas ducts have been reported in Anacardiaceae (Joel
and Fahn, 1980; Machado and Carmello-Guerreiro, 2001; Lacchia and
Guerreiro, 2009), Asteraceae (Sacchetti et al., 1997; Castro and
Demarco, 2008) and Salicaceae (Thadeo et al., 2009, 2014).
Reports by taxonomists for presence of pellucid punctuations and
lines on leaf blades is normally based on stereomicroscopic analysis
(Torres and Yamamoto, 1986; Tozzi, 1989; Marquete and Vaz, 2007;
Marquete and Mansano, 2013, 2016), and such structures are generally
recognized as being cavities and ducts, respectively (Thadeo et al.,
2009, 2014). However, to confirm the presence of cavities and ducts,
and to distinguish them from each other, anatomical studies are ne-
cessary (Thadeo et al., 2009, 2014). Based on anatomical analysis, the
glandular pellucid dots of Lonchocarpus were recognized as secretory
cavities in 19 of 23 analyzed species, while in the remaining four spe-
cies they corresponded to large intercellular spaces that lacked secre-
tory epithelium, and which were termed false secretory cavities
(Teixeira et al., 2000).
Clearly the distinction between lumen length and width will facil-
itate the establishment of a parameter for recognizing cavities and
ducts, but in order to achieve this a three-dimensional view is needed. A
three-dimensional view can be obtained by using three different visual
planes (paradermical, longitudinal and cross sections) or serial sections,
as well as cleared samples.Taxonomic descriptions have cited Casearia
as having pellucid punctuations and lines on the leaf blade (Sleumer,
1980; Torres and Yamamoto, 1986; Marquete and Vaz, 2007; Marquete
and Mansano, 2013, 2016), which have been described as being cavities
and ducts, respectively, in just six of the 180 species of the genus
(Thadeo et al., 2009, 2014). In this context, this study addresses the
following two questions: Are cavities, ducts, or both, common among
species of Casearia? How can cleared samples and anatomical sec-
tioning aid in distinguishing between cavities and ducts? The goal of

90
V. Ferreira Fernandes et al.
this study was to anatomically characterize the pellucid punctuations
and lines in leaves of 47 species of Casearia, including representatives
from all of its sections (sensu Sleumer), and to assess their distributional
patterns, which may provide taxonomically useful characters.
2. Material and methods
Leaf samples of 47 species of Casearia, including representatives of
all of its sections (sensu Sleumer), were collected from herbarium ma-
terial or in the field (C. sylvestris; Appendix A). The herbarium samples
were rehydrated (Smith and Smith, 1942) and stored in 70% ethanol.
For field collected material, mature leaves from the fifth node were
91
Flora 240 (2018) 89–97
Fig. 1. Presence and distribution of cavities and ducts
on the leaf blade of species of Casearia. (cleared
samples). a–d. Cavities and ducts absent in C. com-
mersoniana (a), C. javitensis (b), C. spruceana (c) and
C. eichleriana (d). e–h. Cavities (white arrows) and
ducts (black arrows) sparsely distributed on the leaf
blade of C. combaymensis (e), C. rupestris (f), C. ru-
pestris (g) and C. aculeata (h). Bars = 300 μm.
collected from three specimens of C. sylvestris in Viçosa (Minas Gerais,
Brazil), and fixed in FAA (formalin, acetic acid and 50% ethanol, 1:1:18
by volume) for 48 h under a vacuum, and stored in 70% ethanol
(Johansen, 1940). Voucher specimens were deposited in the herbarium
of the Universidade Federal de Viçosa under the numbers VIC 44866
(Fernandes and Faria s.n) and VIC 44867 (Fernandes and Faria s.n).
Entire leaves and leaf fragments were cleared with 5% sodium hy-
droxide and 10% sodium hypochlorite and interspersed with distilled
water, rinsing at least three times (10 min, each). The samples were
then dehydrated, stained with 50% alcoholic fuchsin (Vasco et al.,
2014) and mounted in glycerin jelly (Johansen, 1940) in order to ob-
serve the distribution of cavities and ducts. Leaf blade fragments of the
V. Ferreira Fernandes et al.
middle region, the region between margin and midrib and the margin
were dehydrated in an ethanol series and embedded in methacrylate
(Historesin; Leica Instruments, Heidelberg, Germany). Paradermal and
cross sections of 5 μm were obtained using an automatic rotary mi-
crotome (Leica RM2265, Deerfield, IL, USA). The sections were stained
with toluidine blue at pH 4.7 (O’Brien and McCully, 1981), and
mounted on slides with synthetic resin (Permount, Fisher Scientific, NJ,
USA). Analysis and photographic documentation were performed using
a light microscope (Olympus AX70TRF; Olympus Optical, Tokyo,
Japan) equipped with a U-Photo system and a digital camera (Zeiss
AxioCamHRc; Zeiss, Göttingen, Germany).
The presence of cavities and ducts on leaf blades was compared with
taxonomic descriptions in order to confirm their correspondence to
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Flora 240 (2018) 89–97
Fig. 2. Presence and distribution of cavities (white
arrows) and ducts (black arrows) on the leaf blade of
species of Casearia (cleared samples). a, b. Cavities
and ducts densely distributed on the leaf blade de C.
manausensis (a) and C. grandiflora (b); c, d. Cavities
and ducts distributed densely (c) and sparsely (d) on
the leaf blade of C. arborea. e. Cavities and ducts
concentrated along the leaf margin of C. bahiensis;
Cavities and numerous ducts in C. negrensis (e) and C.
nigricans (f); and d. Cavities and ducts distributed on
the midrib of C. lasiophylla. Bars = 300 μm.
pellucid punctuations and lines, respectively (Sleumer, 1980; Marquete
and Mansano, 2016).
3. Results
Both cavities and ducts were found in 43 out of the 47 species
studied (Table 1), and were entirely lacking only in C. commersoniana,
C. javitensis and C. spruceana (sect. Piparea) and C. eichleriana (sect.
Casearia) (Fig. 1a–d, Table 1). In all 43 species, the cavity lumen had a
rounded shape or were elliptical, but less than three times longer than
wide, whereas ducts possessed an elongated lumen that was at least
three times longer than wide. (Fig. 1e–h)
In most species, cavities and ducts were found randomly distributed
V. Ferreira Fernandes et al.
Fig. 3. Cavities (white arrows) and ducts (black arrows) on the leaf blade of species of Casearia viewed in paradermal sections. a–f. Note the single-layered epithelium (ep) on the cavities
and ducts of C. altiplanensis (a), C. grandiflora (b), C. sessiflora (c) and C. sylvestris (d). Bars = 150 μm.
on the leaf blade, in a sparse (Fig. 1e–h) or dense manner (Fig. 2a–b).
However, variation in the distribution pattern was observed among
specimens from the same species (Fig. 2c–d). In C. bahiensis (section
Casearia) cavities and ducts were concentrated along the leaf margin
(Fig. 2e). Casearia negrensis and C. nigricans (section Casearia) had the
greatest number of ducts compared to all other 41 species (Fig. 2f–g).
Cavities and ducts on the midrib were observed in all species, as illu-
strated in Casearia lasioplyla (Fig. 2h).
A single layer of epithelium delimiting a lumen was detected in
paradermal (Fig. 3a–f) and cross sections (Fig. 4a–i), confirming that
the structures observed in cleared samples were indeed cavities and
ducts. In C. aquifolia and C. crassinervis, cavities and ducts are common
in the cortex of the midrib (Fig. 4a–b) but less so in the mesophyll
(Fig. 4c). It was not possible to observe these structures in cleared
samples of these species due to the coriaceous consistency of the leaves
and the large caliber of the midrib, and so histological sections proved
to be essential for identifying the structures, which possessed a narrow
lumen and 5–8 epithelium cells (Fig. 4a–c). Cavities and ducts were
93
Flora 240 (2018) 89–97
observed in the region of phloem (Fig. 4i) in six species, and on the
medulla of the midrib (Fig. 4j) in C. resinifera (Table 1). Cavities and
ducts were observed in parallel, perpendicular and oblique positions
relative to the midrib (Fig. 5).
In most species, the secretion within the lumen was preserved and
easily observed in cleared samples (Figs. 1e–h and 2 e–h). On rare oc-
casions the contents of the lumen had a slightly intense color
(Fig. 2b–d), but even so, the cavities and ducts were still easily ob-
served.
4. Discussion
In the present work, we proposed that only structures with a lumen
that is at least three times longer than it is wide be considered ducts.
This made it easier to distinguish cavities and ducts and to confirm the
correspondence between the pellucid punctuations and cavities, and
between lines and ducts. This clear distinction is important since the
presence and distribution of cavities and ducts have been used in
V. Ferreira Fernandes et al.
taxonomical studies (Castro et al., 1997; Thadeo et al., 2014; Bombo
et al., 2016).
The importance of cleared samples for performing a preliminary
survey for the presence and distribution of cavities and ducts on the leaf
blade was demonstrated in the present work and even in other plant
organs. It is a simple technique that allows the presence of cavities and
ducts and their patterns of distribution to be easily determined for a
large number of species. However, C. crassinervis and C. aquifolia pos-
sess coriaceous leaves with cavities and ducts on both the prominent
ribs and mesophyll, which were not detected in the cleared samples;
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Flora 240 (2018) 89–97
Fig. 4. Distribution of cavities and ducts (arrows)
along the midrib (a, b) and mesophyll (c–h) of species
of Casearia shown in cross sections of the leaves. a, b.
reduced lumen in C. aquifolia and C. crassinervis, re-
spectively. Note the epithelium with only five cells in
the detail; c. Cavity/duct located subepidermally in
C. crassinervis; d, e. Cavity/duct in the palisade par-
enchyma and interface between palisade and lacunar
parenchyma in C. rufescens and C. mariquitensis, re-
spectively; f. Duct in the interface between palisade
and lacunar parenchyma in C. sessiflora; g. Cavity/
duct in the interface between palisade and lacunar
parenchyma in C. spinescens, note the cavities side by
side; h. Subepidermal location of the cavity/duct in
C. grandiflora. i. Cavity/duct in the phloem region in
C. sylvestris; and j. Cavity/duct in the medulla of
midrib in C. resinifera. Bars = a–b, 200 μm; c–h,
100 μm; i–j, 100 μm.
thus, histological sections proved critical in enabling careful inter-
pretations.
4.1. Taxonomical importance of ducts and cavities in Casearia
Casearia is a Pantropical genus with almost 200 species, being ap-
proximately 75 distributed in a Subtropical America. This genus is di-
vided in six sections (Sleumer, 1980) and presents a complexity tax-
onomy, have been segregated from Flacourtiaceae to Salicaceae (Chase
et al., 2002), subfamily Samydoideae (Stevens, 2001 onwards).
V. Ferreira Fernandes et al.
Fig. 5. Drawings showing the distribution and orientation of cavities and ducts on the leaf blade of C. negrensis. a. Whole leaf. b. Detail of the middle region.
However, according Alford (2005, 2007) Casearia is the largest genus of
Samydaceae.
The presence, type and position of secretory structures are relevant
characters for taxonomy at different levels (Fahn, 1979; Castro et al.,
1997; Teixeira et al., 2000; Thadeo et al., 2014; Bombo et al., 2016) and
evolutionary investigations (Vitarelli et al., 2015). For instance, in As-
teraceae, the presence of ducts was common in genera belonging to the
tribes Astereae, Eupatorieae and Heliantheae, while they were absent in
Venonieae (Castro et al., 1997). In a similar way, species of the genus
Aldama (Asteraceae) bear a distinct secretory duct arrangement of the
midrib, which might assist in the diagnosis of this genus (Filartiga et al.,
2016). Based on the presence and absence of cavities on species of
Lonchocarpus (Leguminosae – Papilionoideae), it was suggested that the
subgeneric position of some species should be reevaluated (Teixeira
et al., 2000). Pellucid punctuations and lines are considered to be im-
portant diagnostic characters for species of Casearia (Torres and
Yamamoto, 1986; Marquete and Mansano, 2016). The presence of
cavities and ducts in Casearia is a common character since 43 of the 47
species studied, including representatives of all sections, have these
structures. These data are relevant since only six species have pre-
viously been anatomically described (Thadeo et al., 2014). Thus, our
results extend the anatomical database for Casearia and reinforce that
the presence, type and position of secretory cavities and ducts on the
leaf blade are taxonomically important characters for the genus, as
suggested by Thadeo et al. (2014).
Cavities and ducts were not observed in all species of the section
Piparea (C. commersoniana, C. javitensis, C. spruceana) and only in C.
eichleriana from section Casearia, according to the classification of
Sleumer (1980). These findings demonstrate the usefulness of anato-
mical descriptions in demonstrating the absence of pellucid punctations
and lines previously mentioned as inconspicuous, not visible or obscure
in section Piparea and in C. eichleriana (Sleumer, 1980; Marquete and
Mansano, 2016). The biological significance of this evidence should be
95
Flora 240 (2018) 89–97
investigated in future studies that address the phylogenetic distribution
and evolution of cavities and ducts in Caseria. Based on the presence
and absence of cavities for species of Lonchocarpus (Leguminosae –
Papilionoideae), it was suggested that the subgeneric position of some
species should be reevaluated (Teixeira et al., 2000). The taxonomic
descriptions of C. aquifolia, C. crassinervis and C. grandiflora cited pel-
lucid punctations and lines as being obscure, inconspicuous or even
absent (Sleumer, 1980; Marquete and Mansano, 2016). However, cav-
ities and ducts were easily found in cleared samples of C. grandiflora, as
well as in the anatomical sections of C. aquifolia and C. crassinervis.
These anatomical data also help to clarify discrepancies reported
with regard to the descriptions of some species of Casearia. In C. ba-
hiensis, Sleumer (1980) reported the occurrence of “… dense pellucido-
punctata et-lineata … ”, while Marquete and Mansano (2016) men-
tioned that such structures were small and sparsely distributed on the
blade. Our data show cavities and ducts distributed throughout the leaf
blade, but concentrated on the margin, confirming Sleumer’s observa-
tions and detailed description of their distribution. Furthermore, these
anatomical data corroborate the description of the occurrence of pel-
lucid punctations at the center of each areola of C. combaymensis, as
reported by Sleumer (1980).
Descriptions based on cleared samples and both paradermal and cross
sections allowed the careful measurement of lumen length and width for
distinguishing between cavities and ducts. The criterion used in the pre-
sent work is, in part, according to Cutter (1978), who considered cavities
as structures that are more or less isodiametric, whereas ducts are con-
siderably elongated in one plane. However, Cutter (1978) did not explain
how to measure lumen length or width. The three-dimensional recon-
stitution obtained in the present work by examining different planes of
view facilitated the careful distinction between cavities and ducts in a
more objective manner than has been done previously. In additional, this
method makes it possible to easily analyze a large number of species and
specimens as required in taxonomic studies.
V. Ferreira Fernandes et al.
5. Conclusions
Among the species of Casearia, both cavities and ducts exhibited
heterogeneous distributions and orientations on the leaf blade, and so
only examining histological sections could lead to misinterpretations.
On the other hand, observations based on cleared samples ensure the
distinction between cavities and ducts, since they show the length (and
width) of the lumen. From a taxonomic point of view, our results
provide valuable taxonomic characters for Casearia and allow the in-
clusion of the presence/absence and position of cavities and ducts in
combined analyses of morphological and molecular data. In future
studies, such characters can be evaluated in related genera to verify
their taxonomic usefulness in the family Salicaceae.
Acknowledgements
The authors thank CNPq (Conselho Nacional de Desenvolvimento
Cientifico e Tecnológico), CAPES (Brazilian Federal Agency for Support
and Evaluation of Graduate Education within the ministry of Education
of Brazil), FAPEMIG (Fundação de Amparo à Pesquisa do Estado de
Minas Gerais) and Floresta-Escola (SECTES/UNESCO/HidroEX/
FAPEMIG) for financial support. We also thank CNPq for providing a
research scholarship to R.M.S.A. Meira, and CAPES for providing a PhD
scholarship to V.F. Fernandes. We are grateful to the Laboratório de
Anatomia Vegetal and Patricia Fonseca and Aurora Dias for technical
assistance; and the anonymous reviewers for their constructive critiques
and thoughtful reports that improved the manuscript.
Appendix A
List of used materials. Species of Casearia and voucher information.
Section: Taxon: Voucher (Herbarium code). The herbaria acronyms:
CEPEC = Herbário do Centro de Pesquisa do Cacau; INPA = Instituto
Nacional de Pesquisas da Amazônia; NY = New York Botanical Garden;
RB = Jardim Botânico do Rio de Janeiro; IAC = Instituto Agronômico
de Campinas; UEC = Herbário da Universidade Estadual de Campinas;
VIC = Herbário da Universidade Federal de Viçosa. Collections fixed in
FAA (formaldehyde, acetic acid and 50% ethanol) are indicated by
asterisk (*).
Casearia – Casearia aculeata Jacq.: França et al. 3560 (CEPEC), Pott
et al. 8880 (RB), Heringer et al. 2305 (RB), Bernacci et al 790 (IAC),
Bertoni s.n (IAC); Casearia altiplanensis Sleumer: Silva et al. 5247 (RB),
Marquete and Mendonça 2848 (RB), Duarte et al. 185 (VIC); Casearia
aquifolia C. Wright: Iniaged s/n (NY), Shafer 4171 (NY), Roigy and
Mesa 72 (NY) Casearia arborea (Rich.) Urb.: Thomas et al. s.n (CEPEC),
Braga 1226 (RB), Lobão et al. 1309 (RB), Filardi et al. 814 (RB), Torres
et al. 1937 (IAC); Casearia bahiensis Sleumer : Fiaschi et al. 2325
(CEPEC), Mori et al. 12076 (CEPEC), Folli 3512 (RB), Folli 4360 (RB);
Casearia crassinervis Urban: Underwood & Ecule 1343 (NY), Carabia s.n
(NY), Casearia combaymensis Tuslane: Granville and Hoeselc s.n (NY),
Palacios 1405 (NY), Mello 3288 (INPA); Casearia corymbosa Kunth:
Smith 122 (NY), Ortiz 436 (NY), Stimson 5224 (NY); Casearia cotticensis
Uittien: Santos 553 (INPA), Davidson (10068) and Martinelli (INPA);
Casearia decandra Jacq.: Pereira 1156 (CEPEC), Forzza et al. 4356
(CEPEC), Jardim et al. 2849 (CEPEC), Lobão 1658 et al. (RB), Marquete
3377 et al. (RB), Torres s.n (IAC); Casearia duckeana Sleumer:
Albuquerque 67-46 (INPA); Casearia eichleriana Sleumer: Harley et al.
19200 (CEPEC), Jesus et al. 1311 (CEPEC), Zappi et al. 2091 (RB),
Martinelli 5827 (RB) Carvalho et al. 1116 (IAC); C. espiritosantensis R.
Marquete & Mansano: Siqueira 783 (RB); Casearia fasciculata (Ruiz &
Pav.) Sleumer: Krukroff´s 5148 (NY), Prance 30240 et al. (NY),
Krukoff´s 5174 (NY); Casearia grandiflora Cambess.: Thomas et al s.n,
(CEPEC), Belém and Mendes 27 (CEPEC), Forzza 3782 and Mello-Silva
(RB), Soavedra 417 et al. (RB), Bernacci 2248 (IAC), Árbocz 7203
(IAC), Marquete et al. 3166 (IAC); Casearia guianensis (Aubl.) Urb.:
Croat 14978 (NY), Riedel s.n (NY), Kerbbride et al. 1684 (NY), Harris
96
Flora 240 (2018) 89–97
10367 (NY), Howard 9705 (NY); Casearia hirsurta Sw. : Breteler 3675
(NY), Proctor 10183 (NY), Molina 3384 (NY); Casearia lasiophylla
Eichler: Wesenberg 132 et al. (RB), Lindeman 5334 and Haas (RB),
Echternacht 671 and Dornas (RB), Fernandes et al. 33439 (UEC),
Alencar and Oliveira 1557 (IAC), Torres et al. 749 (IAC), Torres et al.
743 (IAC), Torres and Figueiredo 409 (IAC); Casearia manausensis
Sleumer: Cid et al. 1879 (INPA), Mota 67 and Coêlho (INPA), Coelho
3201 (INPA), Monteiro 1330 et al. (INPA), Ferreira 1435 et al. (INPA);
Casearia mariquitensis Kunth: Evans 2575 and Kalmar (RB), Pereira
7888 et al. (RB), Tozzi et al. s.n (UEC), Bertoni 281 (IAC); Casearia
melliodora Eichler: Jardim et al. 1594 (CEPEC) Paixão et al. 197
(CEPEC), Macedo 12 (IAC), Macedo and Lima 470 (IAC); Casearia
mestrensis Sleumer: Silva 158 (CEPEC), Sarmento 644 (RB), Valente
2328 et al. (VIC), Melo 3245 and Franco (IAC); Casearia negrensis
Eichler: Duarte 6557 (RB), Amaral 100 et al. (INPA), Coelho 4121
(INPA), Amaral 750 et al. (INPA); Casearia nigricans Sleumer: Vieira 561
et al. (INPA); Casearia nitida L. Jacq.: Correll 43589 (NY), Brittom 5677
et al. (NY); Casearia oblongifolia Cambess.: Cavalcante 04 (RB), Lima
et al. 4568 (CEPEC), Siqueira 74 (RB), Marquete 1421 (IAC); Casearia
obovalis Poepp. ex Griseb.: Cid and Nelson 2810 (RB); Casearia para-
naensis Sleumer: Marchiori 501 et al. (UEC), Souza 111 (IAC), Zickel
et al s.n (IAC), Rosi et al. 602 (IAC), Anunciação et al. 243 (IAC);
Casearia pauciflora Cambess.: Braga 3923 et al. (RB), Nadruz 2341 et al.
(RB), Braga et al. 1622 (IAC); Casearia pitumba Sleumer: Amaral 760
et al. (INPA); Casearia resinifera Spruce ex Eichler: Ducke s.n (RB),
Ducke 1447 (NY); Casearia rufescens Cambess.: Pirani 5248 et al. (RB),
Carvalho and Silva s.n (IAC), Valente 2327 et al. (VIC); Casearia ru-
pestris Eichler: Pott 12235 et al. (RB), Marquete 3170 et al. (RB), Mota
2096 (VIC), Duarte and Silva 90 (VIC), Árbocz s.n (IAC); Casearia ses-
siliflora Cambess. : Cavalcanti 166 and Sá (RB), Cavalcanti 153 et al.
(RB); Casearia ulmifolia Vahl ex Vent. : Alencar et al., 1163 (UEC)
Hatschbach 61927 (CEPEC)
Crateria – Casearia obliqua Spreng. : Carvalho s.n (CEPEC), Hoehne
s.n (CEPEC), Carvalho s.n (RB), Marquete 4147 et al. (RB), Farney 4313
and Gomes (IAC),; Casearia selloana Eichler: Paula 693 (RB), Macedo
865 and Souza (IAC); Casearia sylvestris Sw.: *Fernandes and Pereira
s.n (VIC), * Fernandes and Pereira s.n (VIC), Lobão 272 et al. (RB),
Bovini et al. 802 (RB).
Endoglossum – Casearia tremula (Grisebach) Grisebach ex Wright:
Howard 10581 (NY), Arnoldo-Broeders 3793 (NY), Buntriy 5103 (NY),
Dugand 6324 (NY), Gentry and Cuadros 47607 (NY).Gossypiospermum
– Casearia luetzelburgii Sleumer: Farney 4160 and Gomes (IAC);
Casearia gossypiosperma Briq.: Scardino 1015 et al. (RB), Silva and
Barbosa 4865 (RB), Thomas et al. s.n (CEPEC), Duarte and Chiregatto
104 (IAC); Casearia praecox Griseb.: Lack, 5971 (NY), Stevens 22930
(NY), Burger and Gentry Jr. 9168 (NY).
Guidonia – Casearia bartlettii Lundell: Ortiz 877 (NY), Ortiz 978
(NY), Ortiz s.n (NY); Casearia spinescens (Sw.) Griseb.: Kuhlmann 6446
(RB), Stannard 826 and Arraís (NY), Britton 6875 (NY), Buntriy 6610
(NY), Curtiss 750 (NY), León 5056 (NY).
Piparea – Casearia commersoniana Cambess. : Peixoto et al. 955
(RB), Bovini et al. 1844 (RB), Peixoto et al. 354 (RB), Lima et al. 2196
(CEPEC), Santos and Alves 220 (CEPEC), Marquete et al. 3404 (IAC);
Casearia javitensis Kunth: Lopes 872 et al. (CEPEC), Carvalho et al. 1479
(CEPEC), Kollmann et al. 4079 (RB), Pereira 49 and Cardoso (RB), Silva
et al. 8532 (IAC), Torres et al. 1927 (IAC), Bernacci et al. 4611 (IAC);
Casearia spruceana Benth. ex Eichler: Pires 420 (IAC), Amaral 718 et al.
(INPA).
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