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Santos IAWA

et al. –Journal
Wood anatomy
34 (3), 2013:
of selected
313–323
Myrtaceae 313

WOOD ANATOMY OF MYRCIARIA, NEOMITRANTHES, PLINIA AND


SIPHONEUGENA SPECIES (MYRTEAE, MYRTACEAE)

Gabriel U.C. A. Santos1,*, Cátia H. Callado2, Marcelo da Costa Souza 3 and


Cecilia G. Costa 4
1 Colégio Pedro II, CSCII, Campo de São Cristóvão 177, 20921- 440 São Cristóvão,
Rio de Janeiro, RJ, Brazil
2 Universidade do Estado do Rio de Janeiro, Departamento de Biologia Vegetal, Instituto de Biologia
Roberto Alcantara Gomes, Rua São Francisco Xavier 524, PHLC - sala 224,
20550-900 Maracanã, Rio de Janeiro, RJ, Brazil
3 Museu Nacional / UFRJ, Departamento de Botânica, Quinta da Boa Vista,
20940-040 São Cristóvão, Rio de Janeiro, RJ, Brazil
4 Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Laboratório de Botânica Estrutural,
Rua Pacheco Leão 915, 22460-030 Jardim Botânico, Rio de Janeiro, RJ, Brazil
*Corresponding author; e-mail: guc_araujo@yahoo.com.br

abstract

Myrciaria, Neomitranthes, Plinia and Siphoneugena are closely related genera


whose circumscriptions are controversial. The distinctions between Myrciaria
vs. Plinia, and Neomitranthes vs. Siphoneugena, have been based on a few fruit
characters. The wood anatomy of 24 species of these genera was examined to
determine if wood anatomical features could help delimit the genera. It was
determined the four genera cannot reliably be separated by wood anatomy alone.
Characteristics seen in all four genera are: growth rings usually poorly-defined;
diffuse porous; exclusively solitary vessels, usually circular to oval in outline;
simple perforation plates; vessel-ray pits alternate and distinctly bordered; fibers
with distinctly bordered pits in radial and tangential walls, usually very thick-
walled; vasicentric tracheids typically absent; scanty paratracheal parenchyma,
sometimes unilateral, and diffuse to diffuse-in-aggregates; chambered crystal-
liferous axial parenchyma in many species, usually both prismatic and smaller
crystals; rays 1–4-seriate, uniseriate rays composed of upright/square cells,
multiseriate rays with procumbent body cells and 1 to many marginal rows of
upright /square cells; disjunctive ray parenchyma cells usually present.
Key words: Comparative wood anatomy, generic boundaries, South American
Myrteae, Plinia group.

INTRODUCTION

Myrtaceae has 132 genera and 5671 species, of which 49 genera and c. 2500 species
are in the tribe Myrteae (Lucas et al. 2007). The Myrteae comprises trees or occasional-
ly shrubs and has a Pantropical distribution (Lucas et al. 2007), with high species
diversity occurring along the eastern coast of Brazil, the Guayana Highlands and the
Caribbean (McVaugh 1968). Although the family is considered well-delimited and
© International Association of Wood Anatomists, 2013 DOI 10.1163/22941932-00000026
Published by Koninklijke Brill NV, Leiden
314 IAWA Journal 34 (3), 2013

easily recognizable in the field, the circumscription of its genera and species has been
debated since the 19th century (Landrum & Kawasaki 1997; Lucas et al. 2005). One of
the reasons for this confusion in generic boundaries is that many genera are distinguished
by a single or relatively few, often cryptic characters, such as the degree of fusion in the
calyx and in the embryo (Landrum & Kawasaki 1997; Salywon & Landrum 2007).
Generic boundaries within Myrteae have attracted attention in past decades. Lucas
et al. (2005, 2007) carried out molecular phylogenetic studies to clarify relationships
within the tribe. They concluded that the traditional three subtribe system based on
embryology and proposed by Berg (1855–56, 1857–59) is artificial, with subtribes Myr-
ciinae and Eugeniinae being polyphyletic. The Plinia group is one of the well-supported
monophyletic informal groups proposed by Lucas et al. (2007). It comprises four gen-
era: Myrciaria, Neomitranthes, Plinia and Siphoneugena. Myrciaria O. Berg has 25
species (Govaerts et al. 2012) that occur from northern Argentina to Mexico and the
Caribbean (Landrum & Kawasaki 1997). It is closely related to Plinia L., a genus dis-
tributed from Brazil and Peru to the West Indies and Cuba (Barrie 2004). The distinc-
tion between Myrciaria and Plinia is controversial, since only the persistence of the
calyx in the fruit distinguishes them, which has led some authors (e.g. Legrand & Klein
1978) to consider Plinia species as Myrciaria. Plinia is considered artificial by some
authors (e.g. Landrum & Kawasaki 1997); Barrie (2004) states that the range of the esti-
mates of the number of species of Plinia (6–40 species) indicates its poor generic limits.
Another indication is the controversial placement of species known as “jaboticabas”, a
small group of eight species treated either as Myrciaria (e.g. Landrum & Kawasaki
1997) or Plinia (e.g. Govaerts et al. 2012). Those species will be treated here as Plinia.
Neomitranthes Kausel ex D. Legrand comprises 14 species from the Brazilian Atlantic
Domain (Souza 2009). It is closely related to Siphoneugena O.Berg, a small genus of
9 species ranging from southern Brazil to the Antilles (Sobral & Proença 2006). The
distinction between them is also obscure, the principal difference being hypanthium
morphology (circumscissile below the staminal ring in Siphoneugena and above in
Neomitranthes) (Landrum & Kawasaki 1997).
Due to these controversies about the generic limits within the Plinia group, the
present study was undertaken to determine whether wood anatomical characters might
be useful for distinguishing the genera.

MATERIALS AND METHODS

Wood samples of 19 species were obtained either from field expeditions or from various
institutional wood collections, data on five species were obtained from the literature
(Table 1). In the field, all samples were taken at breast height, either cut with a saw or
sampled with an increment borer. Transverse, radial and tangential sections 20 –25 µm
thick were cut using a sliding microtome. Sections were bleached in 2–3% sodium
hypochlorite, stained in 1% Astra blue and then in 1% safranin in 50 % ethanol,
dehydrated, and mounted in synthetic resin. Macerations were prepared using Frank-
lin’s method, as modified by Kraus and Arduin (1997). Terminology, definitions and
measurements follow recommendations of an IAWA Committee (1989). Measurements
were taken using Image Pro-Plus 4.0.
Santos et al. – Wood anatomy of selected Myrtaceae 315

Table 1. Specimens details.

Genus / Species Registration number Specimen origin

Myrciaria O. Berg
M. disticha O. Berg RBw 8638 Linhares (ES), Brazil
RBw 9022 Santa Teresa (ES), Brazil
M. ferruginea O. Berg BOTw 854, BCTw 7939 Linhares (ES), Brazil
M. floribunda (H. West ex Willd.) RBw 8598, RBw 8599, Quissamã (RJ), Brazil
O. Berg RBw 8604
M. glazioviana (Kiaersk.) G.M. Barroso RBw 8755, RBw 8966, Rio de Janeiro (RJ), Brazil
ex Sobral RBw 8989
M. guaquiea (Kiaersk.) Mattos & RBw 8754 Niterói (RJ), Brazil
Legrand
M. strigipes O. Berg BOTw 853 Linhares (ES), Brazil
RBw 9024, RBw 9025 São Mateus (ES), Brazil
Neomitranthes D. Legrand
N. cordifolia (D. Legrand) D. Legrand RBw 8262 Sombrio (SC), Brazil
N. glomerata (D. Legrand) D. Legrand RBw 8263 Palhoça (SC), Brazil
N. obscura (DC.) N. Silveira RBw 8402 Rio de Janeiro (RJ), Brazil
RBw 9023 Santa Teresa (ES), Brazil
N. sp. (species unassigned) RBw 8639 Linhares (ES), Brazil
Plinia Plum. ex L.
P. cauliflora (Mart.) Kausel RBw 8751 Rio de Janeiro (RJ), Brazil
P. costata Amsh. Tw 37774, Tw 37797 Nickerie, Suriname
P. edulis (Vell.) Sobral RBw 8721, RBw 8749 Rio de Janeiro (RJ), Brazil
P. ilhensis G.M. Barroso RBw 8753 Niterói (RJ), Brazil
P. oblongata (Mattos) Mattos RBw 8981, RBw 8982 Santa Teresa (ES), Brazil
P. peruviana (Poir.) Govaerts RBw 8722, RBw 8723 Rio de Janeiro (RJ), Brazil
P. renatiana G.M. Barroso & RBw 9020, RBw 9021 Santa Teresa (ES), Brazil
A. L. Peixoto BCTw 18482 Linhares (ES), Brazil
Siphoneugena O. Berg
S. kiaerskoviana (Burret) Kausel RBw 7307, RBw 7538 Nova Friburgo (RJ), Brazil
S. reitzii D. Legrand RBw 8283 São Joaquim (SC), Brazil

RESULTS

We found that wood anatomy of the four genera is quite homogenous and so present a
general description for them. Summaries of the qualitative and quantitative features that
varied among species are in Tables 2 and 3, respectively. There is the possibility that
some individual species can be recognized by their wood anatomy, but more samples
need to be studied to verify this.
Growth rings – Usually poorly marked, rarely absent, described as distinct in P.
martinellii (Barros & Callado 1997) and P. rivularis (Santos 2012). Usually marked
by radially-flattened, thick-walled fibers and lower frequency of axial parenchyma;
some species also show lower frequency of vessels (Fig. 1A, D).
316 IAWA Journal 34 (3), 2013

Table 2. Summary of qualitative features that varied among Myrciaria, Neomitranthes,


Plinia and Siphoneugena woods.
Data are from direct observation, except when noted (1Santos 2012; 2Marchori & Muñiz 1987;
3Barros & Callado 1997; 4Paula et al. 2000). SVO = solitary vessel outline (A = angular, CO =
circular to oval); DHV = deposits in heartwood vessels (C = common, O = occasional, A =
absent/not observed); FWT = fiber wall thickness (T = thin to thick, VT = very thick); RMP =
rays with multiseriate portions as wide as uniseriate portions (Y = yes, N = no); crystals
(PC = in chambered axial parenchyma cells, E = crystals in enlarged axial parenchyma cells,
T = two sizes of crystals in the same chambered cell, A = absent/not observed); vitreous silica
(P = present in axial parenchyma cells; A = absent /not observed); * = unknown.

Species Samples SVO DHV FWT Ray width RMP Crystals Vitreous silica

M. cuspidata 1 1 CO O T 1; 2 O A A
M. disticha 2 CO C VT 1; 2 O T A
M. ferruginea 2 CO O VT 1; 2 N A A
M. floribunda 3 CO C T 1; 2–3 N A A
M. glazioviana 3 CO A T 1; 2 N A A
M. guaquiea 2 CO A VT 1; 2 N A A
M. strigipes 2 CO A VT 1; 2 Y A A
M. tenella 2 2 CO A T 1; 2–3 N A A

N. cordifolia 1 CO-A A VT 1; 2 O PC A
N. glomerata 1 CO A VT 1; 2 O PC A
N. obscura 2 CO A VT 1; 2 N PC A
N. sp 1 CO A VT 1; 2 O T A

P. cauliflora 1 CO-A A T 1; 2 N T A
P. costata 2 CO A VT 1; 2 N A A
P. edulis 2 CO A VT 1; 2–3 N A A
P. ilhensis 1 CO O VT 1 Y E A
P. martinellii 3 2 CO A T 1; 2–3 O PC P
P. oblongata 2 CO-A A T 1; 2 O PC A
P. peruviana 2 CO-A O T 1; 2 O T A
P. renatiana 3 CO O VT 1; 3–4 N A A
P. rivularis 1 1 CO A VT 1; 3 N T A

S. densiflora 4 1 * A T 1; 2–3 O A A
S. kiaerskoviana 2 CO A T/VT 1; 2–3 N T A
S. reitzii 1 A A VT 1; 3– 4 N T A

Vessels – All species diffuse-porous (Fig. 1A–E). Vessels predominantly solitary


(> 90%), only rarely tending to a diagonal to radial arrangement. Vessel outline mostly
circular to oval, in some species a tendency to an angular outline (Table 2). Perforation
plates simple. Intervessel pits rarely observed (since most species have all or almost
all vessels solitary), alternate; vessel-ray pits similar to intervessel pits (Fig. 1G–I);
vesturing, assumed to be present as is characteristic of the Myrtaceae, not obvious with
light microscopy, except in M. tenella and P. martinellii. A few species with occasional
Table 3. Quantitative features of Myrciaria, Neomitranthes, Plinia and Siphoneugena woods.
Data are from direct observation, except when noted (1Santos 2012; 2Marchori & Muñiz 1987; 3Barros & Callado 1997; 4Paula et al. 2000). All values represent
minimum-mean-maximum, except for axial parenchyma strand (minimum-maximum). * = unknown. Sp = number of samples; VFr = vessel frequency (vessels / mm²);
VEL = vessel element length (µm); VTD = vessel tangential diameter (µm); VWT = vessel wall thickness (µm); fiber length, fiber diameter = µm; Flu = fiber lumen
(µm); APS = axial parenchyma strand (in number of cells); RF = ray frequency (rays /mm); ray height, ray width = µm.

Species Sp VFr VEL VTD VWT Fiber length Fiber diameter FLu APS RF Ray height Ray width

M. cuspidata1 1 81-131-181 270-439-620 15-29-40 2-2,9-4 650-937-1190 10-13-17.5 * 3-8 26-29-32 50-239-700 *
M. disticha 2 20-61-123 316-679-953 23.7-47.8-66.8 2.1-3.5-5.8 741-1108-1415 11.8-16.0-20.7 2.2-3.5-7.9 3-9 9-17-27 117-319-786 9.3-15.5-23.3
M. ferruginea 2 62-85-123 403-645-951 17.3-35.9-48.0 2.1-3.1-3.8 729-1012-1406 11.2-15.9-20.9 2.2-3.1-5.2 4-7 9-16-19 117-245-678 9.5-15.7-22.2
M. floribunda 3 34-50-95 340-515-750 30.0-65.0-95.3 2.3-3.1-6.0 665-972-1330 12.0-16.2-21.1 2.4-4.8-6.5 4-6 15-21-26 129-258-559 17.6-25.7-35.5
M. glazioviana 3 110-152-238 288-512-724 19.9-30.5-56.5 2.5-3.3-4.8 645-974-1421 11.4-16.6-22.7 2.2-4.5-7.3 3-7 16-21-277 101-383-718 8.3-15.7-21.3
M. guaquiea 2 107-118-139 365-541-758 24.2-41.2-56.3 2.0-3.5-4.7 524-863-1249 11.7-16.2-20.4 2.1-4.2-8.2 4-6 7-11-16 138-251-457 11.3-15.5-23.2
M. strigipes 2 136-159-184 288-582-1055 16.7-39.6-58.7 1.8-2.8-3.8 578-1023-1438 12.0-18.3-25.0 1.8-3.6-9.5 6-8 11-14-19 117-235-439 9.5-18.3-32.4
M. tenella 2 2 112-177-240 270-468-670 17.5-29.5-36 1.3-1.6-2.1 810-1012-1360 8.7-11.4-15 * 3-7 23-28-33 42-203-767 6.3-13.0-22.5
N. cordifolia 1 76-101-139 306-438-821 63.4-77.9-112.5 2.0-3.0-4.3 520-799-1314 11.4-16.2-20.6 2.2-4.1-7.7 3-5 7-12-18 205-364-696 10.5-16.1-24.2
N. glomerata 1 24-38-49 481-602-751 27.4-49.4-81.3 3.1-4.2-6.6 759-1006-1314 14.4-19.2-27.6 1.6-4.2-8.5 3-4 10-14-17 168-403-637 9.5-18.4-29.5
N. obscura 2 25-27-36 370-640-902 46.5-72.9-105.8 2.4-3.6-5.9 691-1121-1382 11.8-18.6-27.6 1.3-2.4-7.6 5-8 9-14-18 161-282-538 13.9-19.7-28.9
N. sp 1 26-38-54 390-602-875 31.4-72.8-95.4 3.6-5.3-8.3 774-1050-1418 15.4-19.6-23.8 1.7-3.7-5.3 3-6 10-13-16 152-276-533 12.7-16.9-22.2
P. cauliflora 1 198-237-275 216-476-662 24.1-29.1-33.3 1.4-2.4-3.9 851-1104-1598 13.7-16.5-20.8 3.3-4.8-7.5 4-6 13-19-24 92-183-283 14.8-19.9-26.0
P. costata 2 63-114-169 211-533-833 20.7-31.9-42.2 1.9-3.0-4.3 534-919-1654 7.4-13.8-19.8 1.4-2.7-4.8 5-8 9-13-16 115-238-376 9.5-27.7-53.4
P. edulis 2 21-28-35 126-656-1056 47.4-89.9-141.6 2.3-3.6-4.7 849-1532-2047 12.6-19.7-27.2 3.2-6.1-8.6 4-6 14-19-24 167-368-646 20.5-26.4-37.4
P. ilhensis 1 88-107-135 350-514-730 19.0-36.9-51.3 2.1-3.1-4.8 630-1013-1439 14.4-17.8-22.4 1.3-3.1-5.7 5-14 16-19-23 57-267-609 6.5-10.4-14.8
P. martinellii 3 2 16-29-54 431-755-1201 30-59.8-140 2-2.9-4 856-1277-1704 14-22.1-95 2-6.1-24 4-14 5-10-13 174-373-839 8-16.3-24
Santos et al. – Wood anatomy of selected Myrtaceae

P. oblongata 2 198-233-264 193-473-781 23.2-30.0-44.5 1.9-2.6-4.7 801-965-1316 11.3-15.9-20.0 3.4-5.8-8.1 4-6 15-18-22 123-227-324 12.6-17.3-22.5
P. peruviana 2 160-171-208 271-431-657 22.7-32.3-41.9 1.3-2.1-2.7 600-953-1470 11.6-16.8-22.2 3.7-6.0-9.8 4-8 12-18-21 94.3-210-331 10.2-15.2-20.3
P. renatiana 3 0-5-11 310-535-888 61.4-126.1-177.1 3.4-5.0-6.7 765-1271-1806 12.7-19.5-26.5 2.4-5.5-9.8 3-8 9-13-16 138-282-668 20.0-27.7-40.4
P. rivularis 1 1 75-89-100 330-535-680 32.5-48-70 3.1-3.7-4.1 980-1316-1550 16.2-18.8-22.5 * 7-11 19-21-25 40-241-520 *
S. densiflora 4 1 13-15-18 250-346-400 81-93-103 * 1110-1320-* *-17.0-* * * 5-15-* 491-520-549 *-40-61
S. kiaerskoviana 2 7-13-17 338-567-862 49.5-130.0-235.4 1.6-2.5-3.4 1109-1272-1645 11.2-16.8-22.8 1.6-4.2-8.5 3-9 5-15-21 142-421-1047 13.9-32.4-53.3
S. reitzii 1 85-95-112 540-674-890 21.7-50.2-117.0 1.9-3.3-4.7 749-1130-1419 15.6-18.3-20.3 1.5-2.7-5.1 4-6 13-19-27 120-323-771 11.8-30.6-42.2
317
318 IAWA Journal 34 (3), 2013

Figure 1. For legends, see page 320.


Santos et al. – Wood anatomy of selected Myrtaceae 319

Figure 2. For legends, see page 320.


320 IAWA Journal 34 (3), 2013

yellow to orange gum-like deposits in their vessels, abundant only in M. disticha


and M. floribunda. Vessel frequency (vessels /mm 2) 5 (P. renatiana) to 237 (P. oblon-
gata), > 100 in many species. Mean tangential diameters from 29.1 µm (P. cauliflora)
to 130.0 µm (S. kiaerskoviana), most species < 50 µm. Mean vessel element lengths
346 µm (S. densiflora) to 755 µm (P. martinellii) (Table 3).
Fibers – All species with fiber-tracheids with distinctly bordered pits in both ra-
dial and tangential walls, in most species very thick-walled (Fig. 1A–C, E–F). Tracheids
reported only in M. cuspidata and P. rivularis (Santos 2012). Average fiber length
typically > 1000 µm, ranging from 799 (N. cordifolia) to 1532 (P. edulis) (Table 3).
Axial parenchyma – All species with scanty paratracheal parenchyma, sometimes
in a one-layered cap of unilateral paratracheal parenchyma (Fig. 1B–C). Apotracheal
parenchyma diffuse to diffuse-in-aggregates (Fig. 1A–F), occasionally in narrow bands
(1B). Strands of 3–14 cells (Table 3).
Rays – All species with two distinct populations of rays as defined by cellular com-
position (Fig. 2A-D): a) uniseriate rays, composed only of square/upright cells; and
b) 1–4 cell wide rays comprised of procumbent body cells and one to several marginal
rows of square/upright cells. Disjunctive ray parenchyma cells present. Rays typically
numerous, ranging from 10 (P. martinellii) to 28 rays/mm (M. tenella) (Table 3).
Crystals – Present in most species (Table 2). In some species, only one size of pris-
matic crystals in chambered axial parenchyma cells; in many species, two sizes of crys-
tals in the same cell (prismatic plus smaller crystals, Fig. 2 F, G). This latter feature
difficult to assess because smaller crystals might be cryptically present among pris-
matic (larger) crystals. In P. ilhensis crystals occur in slightly inflated parenchyma cells
(Fig. 2 H).
Silica – Vitreous silica in axial parenchyma cells reported only in P. martinellii
(Santos 2012).

Figure 1. – A–F: Transverse sections of Myrciaria and Plinia woods. Diffuse porosity, solitary
vessels, very thick-walled fibers, diffuse to diffuse-in-aggregates apotracheal parenchyma, scanty,
sometimes unilateral paratracheal parenchyma. – A: M. glazioviana. Growth ring boundaries
marked by radially-flattened fibers (arrows). – B: M. guaquiea. – C: M. strigipes. Unilateral
paratracheal parenchyma (arrow). – D: P. oblongata. – E: P. edulis. – F: P. renatiana. Indistinct
growth ring boundary marked by a single row of radially-flattened fibers (arrow). – G–I: Radial
sections. Alternate vessel-ray pits. – G: M. disticha. – H: M. ferruginea. – I: M. guaquiea. —
Scale bars: A, E = 210 µm; B, C, D = 90 µm; F = 80 µm; G, H, I = 30 µm.

Figure 2. Longitudinal sections of Myrciaria, Plinia and Siphoneugena woods. – A–E: Tan-
gential sections. Uniseriate rays composed of upright /square cells (thick arrows), 1–4-seriate
rays composed of procumbent cells in the body and marginal rows of upright /square cells
(thin arrows). – A: P. ilhensis. Exclusively uniseriate rays. – B: M. disticha. Uniseriate and
biseriate rays. – C: P. edulis. Uniseriate and 2–3-seriate rays. – D: P. renatiana. Uniseriate rays
and 3– 4-seriate rays. – E: Radial section of P. renatiana. Rays composed of procumbent cells
in the body and marginal rows of upright /square cells. – F–H: Crystals. – F: P. peruviana. –
G: S. kiaerskoviana. – H: P. ilhensis. — Scale bars: A = 55 µm; B, C = 160 µm; D, E = 120 µm;
F = 40 µm; G = 20 µm; H = 90 µm.
Santos et al. – Wood anatomy of selected Myrtaceae 321

DISCUSSION

All aforementioned features are consistent with previous descriptions of Myrtaceae


woods (e.g. Metcalfe & Chalk 1950; Ingle & Dadswell 1953), except for some slight
differences. First, Metcalfe and Chalk (1950) indicate that vasicentric tracheids are a
typical feature of the family, but these were only observed in two species, M. cuspidata
and P. rivularis (Santos 2012). Their absence, however, might not be uncommon in
tribe Myrteae – e.g. Marques (2010) reported vasicentric tracheids in only 3 out of 8
Eugenia species studied.
The occurrence of two populations of rays with distinct cell compositions, as seen
in the species we studied, is not a feature described as such in Metcalfe and Chalk’s
(1950) family description. This feature has been described before in Myrtaceae, e.g.,
Wagemann (1948), Ingle & Dadswell (1953), Ragonese (1977) and Barros et al. (2001).
This character is common in the tribe Myrteae (Santos et al., in preparation). Although
this feature was not recognized in Siphoneugena densiflora by Paula et al. (2000), it is
clearly seen in their figure. We also observed it in P. martinellii (in slides from RBw),
although Barros and Callado (1997) did not describe it; our observations of this fea-
ture also differ from Record & Hess (1943) for Myrciaria.
Among the species studied, species complexes have been recognized on the basis
of similarities in external morphology. Species in these complexes also have similar
wood anatomy (Table 2 & 3). Myrciaria guaquiea, M. strigipes and M. glazioviana
are in one complex and they have similar anatomy. The “jaboticabas” species analyzed
(P. cauliflora, P. oblongata and P. peruviana) share similar features (narrow and very
numerous vessels, with angular outline; 1–2-seriate rays). On the other hand, N. cor-
difolia, a species not considered part of this “jaboticabas” complex, is quite similar
wood anatomically.
Some wood characters may be diagnostic for some of the species: silica was only
observed in P. martinellii; exclusively uniseriate rays and crystals in somewhat en-
larged parenchyma cells was only observed in P. ilhensis (although only one sample
of P. ilhensis was analyzed). However, these diagnostic features are few. Most of the
qualitative features that varied among species are environmentally influenced to some
degree (e.g. growth ring boundary distinctiveness – Evert 2006), subjective (e.g. rays
with multiseriate portions as wide as uniseriate portions) or have some probability to
be overlooked (e.g. mineral inclusions).
Due to this homogeneity in wood anatomy, it is not possible to use wood characters
to distinguish Myrciaria from Plinia, or Neomitranthes from Siphoneugena. In fact, it is
not possible to separate any of the four genera. This homogeneity is another indication
of their close relationship, as is the similarity in plant morphology. A similar situation
for four genera of the subtribe Myrciinae was found by Dias-Leme et al. (1995), who
concluded that wood anatomical characters could not segregate Calyptranthes, Gomide-
sia, Marlierea and Myrcia, The wood anatomical data supported merging these genera
into a single genus, as proposed earlier by McVaugh (1968). A recent molecular-based
phylogeny (Lucas et al. 2011) supports those views, and inclusion of these genera in
Myrcia is being proposed (Lucas & Sobral 2011).
322 IAWA Journal 34 (3), 2013

On the other hand, homogeneity may indicate that wood anatomy provides limited use-
ful information for the systematics of the Plinia group, or even Myrteae. An analysis
of wood anatomical data of other Myrtean genera described in literature – e.g. some
Eugenia (Détienne & Jacquet 1993; Marques 2010) and some Myrcia (Barros et al.
2001) – shows that their wood structure is often very similar to that of Myrciaria,
Neomitranthes, Plinia and Siphoneugena. Santos (2012) studied the wood anatomy of
26 species of southern Brazilian Myrtaceae and did not find any characters that could
separate genera (except for Myrceugenia, based on pit morphology). Lucas et al. (2007),
based on unpublished wood anatomical data, state that the type of perforation plates is
possibly the only taxonomically useful character for distinguishing Myrteae genera.
They found that most of Myrtean genera have exclusively simple perforation plates,
except for five genera in which some scalariform perforation plates occur (Luma, Myr-
ceugenia, Myrteola, Ugni and Neomyrtus), and two taxa with mixed simple, reticulate
or scalariform perforation plates (Lophomyrtus and Blepharocalyx cruckshanksii)
(Lucas et al. 2007). Schmid and Baas (1984) recorded scalariform perforations in 40
species belonging to seven genera in the family as a whole, six of them Myrtoideae
and Tepualia belonging to the Leptospermoideae.
A comprehensive wood anatomical survey of Myrteae is needed to clarify the sys-
tematic importance of its wood anatomy, since the wood structure of many groups is
poorly known. It seems clear, however, that within the Plinia group wood anatomy
alone does not aid in delimiting genera.

ACKNOWLEDGEMENTS

The authors express their gratitude to FAPERJ, CNPq and CAPES for grants and fellowships; MN/
UFRJ and MBML for field expeditions; JBRJ and UERJ for laboratory structure; ICMBio for research
licence; to Maria J. Miranda (BCTw), Carmen Marcati (BOTw), Neusa Tamaio (RBw), and Hans
Beeckman (Tw) for wood samples; J.A.T. Glória (UERJ) and F. Santos (JBRJ) for technical support;
and the many colleagues that helped in field expeditions, especially A.A.M. de Barros (UERJ), A.G.
Oliveira and L.F.T. de Menezes (UFES), F.Z. Saiter (IFES), A.Z. Monico (ESFA), M.D.M. Vianna
Filho (MN/UFRJ) and A.F.P. Machado (UEFS).

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Accepted: 5 April 2013