FIRST REPORT OF RETICULATE PERFORATION PLATES IN THE MELASTOMATACEAE
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12 IAWA Journal
IAWA 35 (1),
Journal 2014:
35 (1), 12–18
2014
First report of reticulate perforation plates in the
Melastomataceae
Alessandra Ribeiro Guimarães 1,*, José Fernando A. Baumgratz 2 and
Ricardo Cardoso Vieira 3
1 Universidade Federal do Rio de Janeiro, Pós-graduação/Museu Nacional, Departamento de
Botânica, Quinta da Boa Vista, 20940-040 Rio de Janeiro - RJ, Brazil
2 Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rua Pacheco Leão 915,
222460-030 Rio de Janeiro - RJ, Brazil. Bolsista de Produtividade CNPq
3 Laboratório de Morfologia Vegetal da Universidade Federal do Rio de Janeiro, Depto de Botânica,
IB, CCS, BL A, Sala A1-108, Av. Brigadeiro Trompowsky s.n., 21941-590 Ilha do Fundão,
Rio de Janeiro - RJ, Brazil. Bolsista de Produtividade CNPq
* Corresponding author; E-mail: alessandrarguimaraes@yahoo.com.br
Abstract
For the first time, the occurrence of reticulate perforation plates in Melastomata-
ceae is reported. They were discovered in the course of a study of the wood
anatomy of the tribe Microlicieae. The occurrence and frequency of the types of
plates are discussed with respect to their structure, function, and environment.
The reticulate perforation plates seem to be the result of parallel evolution. Some
speculations are offered about their possible adaptive value.
Keywords: Functional adaptation; environment; Microlicieae; parallel evolu-
tion.
Introduction
In Myrtales, vessel elements with multiple perforation plates have been reported
in several Myrtaceae and two species of Lythraceae, Lagerstroemia and Cuphea
carthagenens (Metcalfe & Chalk 1950; Baas & Zweypfenning 1979; Schmid & Baas
1984; Kshirsagar & Vikos 2012). In other families of the order, only simple perforation
plates are known (Van Vliet & Baas 1984). Several hypotheses have been suggested
regarding the scalariform perforation plates. Van Vliet and Baas (1984), in a phylo-
genetic context, hypothesized that these plates could be an ancestral condition for the
family Myrtaceae as well as for the order Myrtales. However, their interpretation of the
sporadic occurrence of multiple perforation plates in Myrtales in the Baileyan sense
was questionable, in the absence of robust phylogenetic reconstructions as currently
available (APG III 2009). On the other hand, Schmid and Baas (1984) discussed the
occurrence of different types of perforation plates in Myrtaceae in relation to the dif-
ferent environments in which they occur and in a functional context.
Phylogenetic analyses of angiosperms have suggested the likelihood of reversals in
Baileyan transformations. Moreover, the parallel emergence of different vessel char-
© International Association of Wood Anatomists, 2014 DOI 10.1163/22941932-00000043
Published by Koninklijke Brill NV, Leiden
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via free accessGuimarães et al. – Reticulate perforation plates 13
acters as functional adaptations is well accepted (Baas & Wheeler 1996; Jansen et al.
2004; Olson 2012). The tribe Microlicieae (Melastomataceae) is currently considered
a monophyletic group and is represented by six genera with about 185 species, of
which over 90% are endemic to the Brazilian Cerrado biome (Almeda & Martins 2001;
Clausing & Renner 2001; Fritsch et al. 2004; Michelangeli et al. 2013). The present
study reports, for the first time in the family, the occurrence of vessel elements with
reticulate perforation plates, in species belonging to four genera: Chaetostoma DC.,
Lavoisiera DC., Microlicia D. Don and Rhynchanthera DC., all belonging to this
tribe.
Figure 1–6. Vessel elements with reticulate perforation plates (arrow) in species of Microlicieae
(Melastomataceae). – 1: Chaetostoma armatum. – 2: Microlicia isophylla. – 3: Lavoisiera
imbricata. – 4: L. crassifolia. – 5 & 6: Rhynchanthera serrulata. — Scale bars in 1, 3, 6 = 5 µm;
in 2 = 10 µm; in 5 = 20 µm; in 4 = 50 µm.
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Table 1. Microlicieae with wood having reticulate perforation plates and simples in vessel
elements, and data of the samples.
Climate-Temperature
Plates reticulate (%)
Phytophysiognomy
Climate-Humidity
All plates simple
Altitude
Biome
Habit
Species / Country of origin / wood collection
Chaetostoma albiflorum (Naudin) C.Kosch. &
A.B. Martins
Brazil, State of Minas Gerais
RBW (9346, 9591, 9592) A Sub Cr wa mh 1 1031–1102
C. armatum (Spreng.) Cogn.
Brazil, State of Minas Gerais
RBW (9593, 9594, 9595) A Sub Cr wa mh 1 1300
C. cupressimum (D. Don) C.Kosch. & A.B. Martins
Brazil, State of Minas Gerais
RBW (9340, 9598, 9602) × Sub Cr wa mh 1 1153
Lavoisiera chamaepitys A.St.-Hil. ex Naudin
Brazil, State of Minas Gerais
RBW (10135, 10136, 10137) A Sub Cr wa mh 1 1200
L. crassifolia Mart. & Schrank ex DC.
Brazil, State of Minas Gerais
RBW (10132, 10133, 10134) A Sub Cr wa mh 1 1250
L. glandulifera Naudin
Brazil, State of Minas Gerais
RBW (10141, 10142, 10143) B Sub Cr ht hu 1 1700
L. imbricata (Thunb.) DC.
Brazil, State of Minas Gerais
RBW (9716, 9717, 9718) C Sub Cr me hu 2 1600
L. imbricata (Thunb.) DC.
Brazil, State of Minas Gerais
RBW (9334, 9345, 9731) A Sub Cr wa mh 1 1300–1500
Microlicia avicularis Mart. ex Naudin
Brazil, State of Minas Gerais
RBW (9337, 9343, 9600) A Sub Cr wa mh 1 1100–1300
M. cordata (Spreng.) Cham.
Brazil, State of Minas Gerais
RBW (10162, 10163, 10164) A Sub Cr wa mh 1 1400
M. euphorbioides Mart.
Brazil, State of Minas Gerais
RBW (10156, 10157, 10158) × Sub Cr wa mh 1 1250
(continued on the next page)
Material and Methods
Species belonging to six genera were analyzed (Table 1). The perforation plates were
observed in material dissociated in Jeffrey’s solution (Johansen 1940), and in transverse
and longitudinal (radial and tangential) sections. A total of 4,600 counts were made
using longitudinal radial sections (in which the plates were best observed), to calculate
the percentage of occurrence of the reticulate plates. The characterization of vegeta-
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(Table 1 continued)
Climate-Temperature
Plates reticulate (%)
Phytophysiognomy
Climate-Humidity
All plates simple
Altitude
Biome
Habit
Species / Country of origin / wood collection
M. fasciculata Mart.
Brazil, State of Minas Gerais
RBW (9348, 9596, 9597) × Sub Cr wa mh 1 1100–1300
M. isophylla DC.
Brazil, State of Minas Gerais
RBW (10138, 10139, 10140) A Sub Cr wa mh 1 1200
M. subsetosa DC.
Brazil, State of Minas Gerais
RBW (9339, 9599, 9601) × Sub Cr wa mh 1 1100–1350
Rhynchanthera grandiflora (Aubl.) DC.
Brazil, State of Minas Gerais
RBW (9336, 9344, 9347) B Shr Ve wa mh 1 900–1200
R. serrulata (L.C. Rich.) DC.
Brazil, State of Pará
RBW (9603, 9604, 9605) B Sub Re wa hu 3 40
Stenodon suberosus Naudin
Brazil, State of Goiás
RBW (10165, 10166, 10167) × Shr Cr wa mh 1 700
Trembleya laniflora (D.Don) Cogn.
Brazil, State of Minas Gerais
RBW (9710, 9711, 9712) × Sub Cr wa mh 1 1400
T. parviflora (D.Don) Cogn. in Mart. & Eichler
Brazil, State of Minas Gerais
RBW (9333, 9713, 9714) × Shr Cs wa mh 1 1300–1600
T. tridentata Naudin
Brazil, State of Minas Gerais
RBW (9335, 9707, 9708) × Sub Cr wa mh 1 1300
Perforation plates reticulate: A (1–3 %); B (10–13 %); C (40– 45 %)
Habit: Sub (subshrubs); Shr (shrubs)
Phytophysiognomy: Cr (Campo rupestre); Ve (Vereda); Cs (Cerrado sensu stricto); Re (Restinga)
Climate-Temperature: wa (warm), me (mesothermic), ht (humid temperate)
Climate-Humidity: mh (semi-humid), hu (humid)
Biome: 1 (Cerrado); 2 (Mata Atlântica); 3 (Amazônia)
tion types follows Ribeiro & Walter (2008), and the data for the environments where
collections were made are provided below:
Cerrado Biome — The collections were made in the states of Minas Gerais and
Goiás in the following vegetation types: campo rupestre (montane savanna), vereda
(Buriti palm swamp) and cerrado sensu stricto (Table 1). The climate is hot, semi-humid
and strongly seasonal, with dry winters and rainy summers. Across the region there
is a wide variation in the annual mean temperature, ranging from 18 to 28 °C, and in
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precipitation with annual means of 800–2,000 mm, with an intense dry season during
the austral winter (April to September) (Dias 1992). However, the region of collection
in the campo rupestre, at 1,700 m altitude, has a humid temperate climate with dry
winters and mild summers, and a mean annual precipitation of 1,500 mm (Table 1)
(Antunes 1986; Brandão et al. 1994).
Atlantic Forest Biome — The collection was made in the state of Minas Gerais, in
campo rupestre vegetation, above 1,500 m altitude (Table 1). The climate is humid
mesothermal with dry cold winters (frosts and precipitation may occur sporadically)
and mild humid summers; the dry season is short. The mean annual rainfall is 1,900
mm and the mean annual temperature is between 17 and 20 °C (Menini-Neto et al.
2009; Valente et al. 2011).
Amazon Biome — The collection was made in the state of Pará, in restinga (coastal
dune scrub) vegetation (Table 1). The climate is hot year-round, with a mean annual
temperature of 26 °C and minimum temperatures above 18 °C, with a mean annual
rainfall of 2,500 mm (Silva et al. 2010).
Results
In the secondary xylem of the stem, vessel elements with only simple perforation plates
were observed in eight species (Table 1). In the remaining 11 species studied, reticulate
perforation plates were observed (Table 1, Fig. 1–6) in addition to the predominant
simple perforations. In the same vessel element, both simple and reticulate perforation
plates occur, either at each end or in lateral walls (Fig. 1–6). While perforation plates
are very inclined at oblique end walls (see Fig. 4 & 5), the degree of inclination may
vary. In all the vessel elements, the pits are vestured.
Discussion
According to literature records, Melastomataceae possess exclusively simple perfora-
tion plates (Record & Hess 1943; Metcalfe & Chalk 1950; Ter Welle & Koek-Noorman
1981; Van Vliet 1981; Van Vliet & Baas 1984; Medeiros & Morretes 1994; Marcon &
Costa 2000; Bosio et al. 2010). Our results clearly indicate that reticulate perforations
also occur in the family.
It is tempting to ascribe a functional and /or adaptive role to the occurrence of reticu-
late plates in the 11 species of Microliceae. However, with their generally low incidence
a functional role in hydraulic conductivity or safety can at most be marginal. Some
possible advantages and disadvantages have been attributed to the multiple perforation
plates. Wheeler and Baas (1991) and others suggested that a possible disadvantage may
be related to the low efficiency in relation to the intense water-flow demand imposed
by the high transpiration rates in hot dry regions. In this respect, the simple perforation
plates appear to be well adapted. Originally, this suggestion was supported by the stud-
ies of Schulte and Castle (1993) and Ellerby and Ennos (1998), wherein the additional
resistance offered to the flow of water by the scalariform perforation plates, from 2%
to 20% of the total resistance, is greater than that observed for the simple plates, of
2% to 5%. In this context, the predominance of simple perforation plates in 11 species
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of Melastomataceae in this study, and the presence of only simple plates in the other
eight species (Table 1) makes perfect sense.
Although the number of specimens studied by us was very low, it is striking that the
highest incidence of reticulate perforations (40–45%) is in one specimen of Lavoiseria
imbricata, the only sample collected from the Mata Atlântica, at relatively high altitude
where frosts do occur from time to time. The other specimen studied of this species
came from the hot, frost-free sub-humid Cerrado and had only 1–3% reticulate plates.
Whether in the Mata Atlântica specimen the reticulate perforations function in trapping
freeze-thaw embolisms requires the study of more samples and physiological experi-
mentation. The limited variation in the incidence of reticulate perforations in species
from the other specimens does not show a clear ecological pattern. Schmid and Baas
(1984) found highest incidences of scalariform perforation plates in Myrtaceae in spe-
cies from cool mesic environments, in our opinion possibly as a comparable response
to sporadic frost to the reticulate perforations in frost-prone L. imbricata.
Because the tribe Microlicieae is a monophyletic group (Almeda & Martins 2001;
Fritsch et al. 2004; Michelangeli et al. 2013), the results obtained in the present study
suggest that the origin of the reticulate perforation plate is a result of parallelism in the
species examined. Future comparative analyses, involving additional representatives
of Microlicieae and species of other tribes, and physiological studies on the role of
sporadic reticulate plates in hydraulic functioning may better clarify the evolution of
the multiple perforation plates in Melastomataceae.
Acknowledgements
We thank Pieter Baas and two anonymous referees for insightful comments and criticism. Their
recommendations were essential to the improvement of this study.
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