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Botanical Journal of the Linnean Society, 2012, ••, ••–••. With 7 figures

A comparison of the wood anatomy of 11 species

from two cerrado habitats (cerrado s.s. and adjacent
gallery forest)
JULIA O. SONSIN1*, PETER E. GASSON FLS2, CLAUDIA F. BARROS3 and
CARMEN R. MARCATI4
1
Departamento de Recursos Naturais, FCA, PG Ciência Florestal, UNESP – Universidade Estadual
Paulista, Fazenda Experimental Lageado – Rua José Barbosa de Barros, n°1780, CP 237,
18603-970, Botucatu, SP, Brazil
2
Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK
3
Instituto de Pesquisas do Jardim Botânico do Rio de Janeiro, Rua Jardim Botânico, n°1008, Jd.
Botânico, 22460-030, Rio de Janeiro, Brazil
4
Departamento de Recursos Naturais, FCA, UNESP – Universidade Estadual Paulista, Fazenda
Experimental Lageado, Rua José Barbosa de Barros, n°1780, CP 237, 18603-970, Botucatu, SP, Brazil

Received 25 January 2012; revised 4 May 2012; accepted for publication 1 June 2012

A comparative study of the secondary xylem (wood) anatomy of 11 species (38 specimens) occurring in cerrado s.s.
and the adjacent gallery forest (both cerrado s.l. habitat) was made with the aim of identifying the anatomical
characteristics of ecological value and correlating them with the environmental conditions. The anatomical features
that vary, in general, between the two habitats are: growth ring distinctness (well or poorly defined); tyloses and
deposits (more abundant in cerrado specimens); gelatinous fibres (more evident in cerrado specimens and in
different patterns between habitats); variation in paratracheal and banded parenchyma (more abundant in
cerrado); and more cells per parenchyma strand in cerrado. In general, gallery forest specimens have wider vessels,
fewer vessels per square millimetre and larger intervessel pits, indicating more efficient water conduction, whereas
cerrado s.s. specimens are the opposite, with low vulnerability and mesomorphy indices, demonstrating greater
safety under conditions of water stress. © 2012 The Linnean Society of London, Botanical Journal of the Linnean
Society, 2012, ••, ••–••.

ADDITIONAL KEYWORDS: comparative wood anatomy – ecological wood anatomy – safety versus
efficiency.

INTRODUCTION da Flora do Brasil, 2011), 44% of which are endemic

Cerrado s.l. is considered to be the most biodiverse
savanna in the world (Oliveira & Marquis, 2002),
and is one of the 25 global hotspots – priority areas
for conservation (Mittermeier et al., 1999). It is
mostly restricted to Brazil and merges into a great
diversity of biomes, such as Amazon rainforest, Pan-
tanal, Atlantic forest and Caatinga. This enriches its
flora, about 11 500 plant species (Lista de Espécies
*Corresponding author. E-mail: jsonsin@yahoo.com.br
(Coutinho, 1990; Castro et al., 1999). The cerrado
biome has different habitats varying from open
savanna to savanna with scattered trees to cerrado
s.s. and cerradão (Coutinho, 2002). Along the rivers
there are gallery forests among other formations,
which are all considered part of cerrado s.l. (Eiten,
1972). According to Hammerle (2006), few species
occur regularly in both cerrado s.s. and gallery
forests. In São Paulo state, which includes our study
site, the remnants of this vegetation are in scattered
patches covering < 1% of the state (Durigan, Franco
& Siqueira, 2004b).

© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–•• 1
2 J. O. SONSIN ET AL.
Studies on ecological wood anatomy (e.g. Klaassen,
1999; Alves & Angyalossy-Alfonso, 2000, 2002; Lens
et al., 2004; Barros et al., 2006; Loepfe et al., 2007;
Pratt et al., 2007; see also Carlquist, 2010) are of
great importance in understanding the strategies
used by species to survive in different environments
and the relationship between safety and efficiency in
water conduction. Water availability and hydraulic
architecture in the wood may strongly influence
species distribution (Pockman & Sperry, 2000; Barros
et al., 2006), as well as altering photosynthetic capac-
ity, water usage efficiency and storage quantity in dry
periods (Hammerle, 2006). However, much is still
unknown about the ways in which plants adjust the
anatomy of their xylem tissues to meet transpiration
needs or to increase mechanical stability, in part
because studies have been based on relatively small
numbers of species and/or limited geographical
regions (Zanne et al., 2010).
Comparative studies of secondary xylem of cerrado
and other forest formation species (see Marcati,
Angyalossy-Alfonso & Benetati, 2001; Machado et al.,
2007) generally show that the main differences are
quantitative (vessel diameter and frequency, fibre
length and thickness, and ray width and frequency).
Qualitative differences are few and, for cerrado
species, largely apply to growth rings (Oliveira, 2007).
This article describes the differences in wood
anatomy of 11 species that occur in two different
habitats of cerrado s.l. (relatively dry cerrado s.s. and
wet gallery forest), relating the wood anatomical
features to these environments and verifying the
strategies used by plants to survive in the different
habitats.
MATERIAL AND METHODS
The study was carried out in a private cerrado s.l.
reserve of about 180 ha, located in the Palmeira da
Serra Ranch in the municipality of Pratânia
(22°82′01.8″S and 48°74′02.6″W, 715 m) in the mid-
western region of the state of São Paulo, Brazil
(Fig. 1; maps were compiled by DIVA-GIS software).
The species were collected in the cerrado s.s., which is
characterized by short xeromorphic trees and shrubs
with thick bark and leathery leaves, and by herba-
ceous ground cover, and in the adjacent wet gallery
forest alongside small streams (the canopy forms a
tunnel or gallery) in regions in which the interfluvial
zone is covered by cerradão, with standing water even
in the dry period.
The soils are similar in both habitats: acidic with
low organic matter, low phosphorus, potassium,
calcium and magnesium and high aluminium (higher
in cerrado). Both are considered sandy, but there is a
© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
greater quantity of clay in cerrado s.s. (see Milanez,
2007).
The climate diagram shows that both habitats have
a dry period of about 4 months with precipitation
below 60 mm (Fig. 2); average annual rainfall is
approximately 1306 mm and the average temperature
is 19.9 °C. Climate parameters were obtained by the
same software as described in Hijmans et al. (2005).
There are no differences in the climate between the
two habitats, which are adjacent.
Samples of 11 species (adult individuals) from 38
specimens in total, 23 from cerrado s.s. (Ce) and 15
from the gallery forest (GFo), were collected (Table 1),
with three to four specimens of each species. Large
branches were sampled above the base of the main
branch to avoid any irregular formation of that area,
and because destructive sampling of trunks is not
permitted in environmental reserve areas. Discs
of c. 3–6 cm in thickness were obtained from each
specimen.
Habit information follows the classification by
Durigan et al. (2004a) with adaptations: shrub (woody
plant branched from the base, without a defined
trunk); small tree (similar to a shrub in size, but with
a single trunk); tree (tree, the height of which is
always lower than the average canopy); emergent tree
(tree, the height of which is generally higher than the
average canopy). Vouchers were deposited at the
‘Irina Delanova de Gemtchujinicov’ Herbarium
(BOTU) of the Institute of Biosciences (IB). The wood
samples were deposited at the ‘Maria Aparecida
Mourão Brasil’ Wood Collection (BOTw) of the Univer-
sidade Estadual Paulista (UNESP) Faculty of Agro-
nomic Sciences (FCA) at Botucatu, São Paulo State.
Transverse (TS), tangential longitudinal (TLS) and
radial longitudinal (RLS) sections, 15–20 mm in thick-
ness, were double stained with aqueous 1% safranin
and aqueous 1% astra blue (Bukatsch, 1972) (1 : 9).
The innermost layer of gelatinous fibres is mainly
composed of cellulose, and astra blue staining makes
it easier to distinguish the different patterns of dis-
tribution. Histological slides were embedded perma-
nently in synthetic resin (Entellan®). The cells were
macerated using the method of Franklin (1945), modi-
fied by Kraus & Arduin (1997), and stained with
aqueous 1% safranin (Sass, 1958). Semi-permanent
slides were mounted in glycerin diluted in water
(1 : 1).
Qualitative and quantitative anatomical data were
obtained using an Olympus DP70, attached to a Leitz
DMRB microscope, and analySIS software. The Inter-
national Association of Wood Anatomists (IAWA)
hardwood list (IAWA Committee, 1989) was used as a
basis for the descriptions, and observations were
made on both sapwood and heartwood when present.
Growth rings were assessed as follows.
 COMPARATIVE WOOD ANATOMY OF CERRADO 3

Figure 1. Study area: 1, cerrado s.s.; 2, gallery forest. Maps were compiled by DIVA-GIS software. Satellite image from
Google Earth (2010) at an altitude of 3.58 km.

1. Present: the ring boundaries are visible.
2. Well defined: the ring boundaries are well defined
and easy to observe.
3. Poorly defined: the ring boundaries are not well
defined and difficult to observe.
4. Absent: no rings present.
For quantitative data, we made 30 measurements
or counts for fibre length, vessel element length,
vessel lumen diameter, fibre wall thickness, and
vessel frequency. For intervessel pit diameter, vessel-
ray pit diameter and rays per millimeter, we made
ten measurements or counts.
For vessel grouping, 100 counts were made for each
specimen, represented by one for solitaries, two for
pairs, three for multiples of three, and so on succes-
sively, and the average of these counts was calculated
following the method of Carlquist (1984). Vulnerabil-
ity (V = vessel element diameter/vessel frequency)
and mesomorphy (M = V ¥ vessel element length)
indices were calculated following Carlquist (1977).
Statistical analyses were performed with Statistica
software version 8.0. The Shapiro–Wilk test was used
to assess the normality of the samples (Zar, 1996).
Variance analyses with a nested design were used to
test variation between specimens of the same species

© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
4 J. O. SONSIN ET AL.
Figure 2. Climatic diagram for cerrado and gallery forest
(climate parameters were obtained using the same soft-
ware as described in Hijmans et al., 2005): , temperature
monthly maximum; , temperature monthly minimum;
, precipitation.
and individuals of the two habitats (Jonsson &
Malmqvist, 2000). Honestly significant difference
(HSD) Tukey tests were carried out for multiple com-
parisons. Principal components analysis (PCA) was
used to order species, qualitative and quantitative
wood anatomy features and climate features, showing
the factors with more variance (Ludwig & Reynolds,
1988).
RESULTS
WOOD ANATOMY DESCRIPTIONS
A summarized description with the main anatomical
features is given with the relative frequencies in
Table 2. A full description of the species is given in
Supporting Information, Appendix S1, because there
is no information in the literature about the wood
anatomy of these species in the cerrado, except for
Pera glabrata (Schott) Poepp. ex Baill. and Vochysia
tucanorum Mart. by Alves & Angyalossy-Alfonso
(2000, 2002). The descriptions cover TS, TLS and RLS,
but only TSs are shown in the plates, except for
Figure 5.
GENERAL WOOD ANATOMICAL DIFFERENCES
BETWEEN HABITATS
Individual species show anatomical differences
between the two habitats, but these are not universally
consistent. Growth ring distinctness was assessed as
© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
either well or poorly defined. Only Acosmium subel-
egans (Mohlenbr.) Yakovlev (Fig. 4E, F) had well-
defined growth rings with the same boundary marker
in all specimens from both cerrado and the gallery
forest. In Siparuna cujabana (Mart. ex Tul.) A.DC.
(Fig. 5C, D), growth rings were absent in the gallery
forest and present in cerrado. Overall, cerrado samples
had narrower vessel elements, higher vessel frequency
per square millimetre, smaller intervessel pits
(Fig. 6C, E, H, Table 3) and tyloses and deposits in
vessels were more frequent. Fibres were longer and
thicker walled in cerrado (Fig. 6A, F, Table 3). Gelati-
nous fibres were also more evident in cerrado speci-
mens (P. glabrata and Tibouchina stenocarpa (DC.)
Cogn., Figs 3G, H, 4G, H, Appendix S1), with different
patterns and abundance between specimens in the
two habitats. Paratracheal axial parenchyma pat-
terns were more diverse in cerrado s.s. and banded
parenchyma was more apparent in cerrado s.s.
species (Dimorphandra mollis Benth., Stryphnoden-
dron polyphyllum Mart., A. subelegans and V. tucano-
rum; Figs 4A–F, 5E, F, Table 2, Appendix S1). There
were more cells per axial parenchyma strand in
cerrado [Schefflera vinosa (Cham. & Schltdl.) Frodin &
Fiaschi, S. polyphyllum, A. subelegans, T. stenocarpa
and Sp. cujabana; Table 2]. Schefflera vinosa from
cerrado had wider radial canals than gallery forest
samples (46.7 mm and 21.3 mm, respectively; Fig. 5I, J).
Vessel arrangement showed a slight variation only
in P. glabrata (Fig. 3G, H). Intervessel pitting differed
in arrangement in A. subelegans and V. tucanorum.
Ray cell composition varied in Sp. cujabana.
STATISTICAL ANALYSES
There were statistically significant differences in
quantitative parameters between the two habitats for
all specimens, except for vessel element length and
vessel grouping index (Table 3, uppercase letters). In
general, gallery forest samples had wider vessels at
lower frequency per square millimetre, shorter fibres
with thinner walls, larger intervessel pits and fewer
rays per millimetre than cerrado samples of the same
species. Each species, however, showed a limited
range within its own taxon (Table 3, lowercase
letters) (Fig. 6A–H). This result is best shown in the
PCA (Fig. 7A, B), where all cerrado samples of a given
species were placed above those from the gallery
forest, except for A. subelegans, but they remained
close to each other.
In general, the means of the vulnerability and
mesomorphy indices from cerrado were lower (2.9 and
1295, respectively) than those of the gallery forest (5
and 2167, respectively). The vessel grouping index
was similar for both habitats (1.7 in cerrado s.s. and
1.6 in gallery forest).
 COMPARATIVE WOOD ANATOMY OF CERRADO 5

Table 1. Species from gallery forest (GFo) and cerrado s.s. (Ce)

Family Species Area BOTw Habit H (m) CAP (cm)

Anacardiaceae Tapirira guianensis GFo 1952 T 4.5 17

Aubl. 1956 10.0 80
Ce 1321 2.5 10
1965 1.7 10
Apocynaceae Tabernaemontana GFo 1958 T 9.0 35
catharinensis A.DC. Ce 1629 6.0 32
1634 4.0 30
1957 5.0 27
Araliaceae Schefflera vinosa GFo 1963 ST 5.5 27
(Cham. & Schltdl.) Ce 1324 3.5 47
Frodin & Fiaschi 1508 4.5 52
Fabaceae Dimorphandra mollis GFo 1990 T 4.5 25
Caesalpinioideae Benth. Ce 1351 4.0 35
2043 3.5 38
Fabaceae Mimosoideae Stryphnodendron GFo 1999 T 7.0 40
polyphyllum Mart. 2000 6.0 38
Ce 1352 4.0 40
1537 4.5 47
Fabaceae Acosmium subelegans GFo 1998 ST 4.2 30
Papilionoideae (Mohlenbr.) Yakovlev Ce 1342 3.5 35
2044 3.3 40
Melastomataceae Tibouchina stenocarpa GFo 2021 S 3.0 27
(DC.) Cogn. Ce 2018 2.1 10
1633 2.0 10
Peraceae (or Pera glabrata (Schott) GFo 1987 ET 4.5 23
Euphorbiaceae) Poepp. ex Baill. 1989 7.5 38
Ce 1339 3.5 22
1579 4.5 17
Rutaceae Zanthoxylum GFo 2036 T 4.0 17
rhoifolium Lam. Ce 1357 2.7 14
1560 3.5 16
Siparunaceae Siparuna cujabana GFo 2024 T 2.1 8
(Mart. ex Tul.) A.DC. 2159 2.2 10
Ce 2022 5.0 15
2023 3.0 10
Vochysiaceae Vochysia tucanorum GFo 2041 T 14.0 88
Mart. Ce 1368 6.4 42
2178 4.5 72

BOTw, ‘Maria Aparecida Mourão Brasil’ Wood Collection; H, tree height; CAP (CBH), tree circumference at breast height;
T, tree; ST, small tree; ET, emergent tree; S, shrub.

PCA (Table 4) showed that anatomical characteris-
tics vary within factors that together explain 77% of
the total variance (Fig. 7A, B). Only quantitative ana-
tomical characters were selected as the qualitative
and soil features showed low variance and therefore
had a low contribution in PCA. Axis 1 explains 41% of
the total variance and is influenced by fibre length,
vessel element length and fibre wall thickness. Axis 2
explains 23% of the total variance and is influenced
by intervessel and vessel-ray pit diameter, and axis 3
explains 13% of the total variance and is influenced
by rays per millimetre. Both Sp. cujabana and
Tabernaemontana catharinensis A.DC. remain more
distant, as the vessel frequency is higher than that in
other species.
DISCUSSION
Most species from cerrado s.s. and gallery forest have
growth rings, and this is to be expected, as both areas
experience precipitation of < 60 mm for 4 months,
which, according to Worbes (1995), is sufficiently long

© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
 6

Table 2. Summarized description of the cerrado s.s. and gallery forest species with the relative frequencies of the anatomical features

Percentage Vessels Defined by Growth rings

Thick-walled
J. O. SONSIN ET AL.

Radially and radially

Gelatinous flattened Bands/lines Thick-walled flattened
More than 2 or 3 Solitary Radial Diffuse fibres/ parenchyma of axial latewood latewood Poorly Well
4 vessels vessels vessels pattern porous bands cells parenchyma fibres fibres Absent defined defined CS Species

2 21 77 – * – – – – * – * – Ce Tp. guianensis (Fig. 3A)

2 22 76 – * – – – – * – – * GFo Tp. guianensis (Fig. 3B)
9 38 53 * * – – – * – – – * Ce Tb. catharinensis (Fig. 3C)
3 33 64 * * – – – * – – * – GFo Tb. catharinensis (Fig. 3D)
5 44 51 – * – – – – * – – * Ce Sf. vinosa (Fig. 3E, 5I)
2.5 33.5 64 – * – – – – * – – * GFo Sf. vinosa (Fig. 3F, 5J)
26 28 62 * * * – – – * – * * Ce P. glabrata (Fig. 3G)
23 42 35 * * * – – – * – * * GFo P. glabrata (Fig. 3H)
8 34 58 – * – * – – – * * – Ce D. mollis (Fig. 4A)
5 33 62 – * – * – – – – * – GFo D. mollis (Fig. 4B)
2 19 79 – * – – – – * – * * Ce S. polyphyllum (Fig. 4C)
3 20 77 – * – – – – * – * * GFo S. polyphyllum (Fig. 4D)
5 39 56 – * – – * – – – – * Ce A. subelegans (Fig. 4E)
3 57 40 – * – – * – – – – * GFo A. subelegans (Fig. 4F)
4 24 72 – * – – – – * – * – Ce T. stenocarpa (Fig. 4G)
4 26 70 – * – – – – * – * – GFo T. stenocarpa (Figs.4H, 5G)
1.5 36.5 62 – * – – – – * – – * Ce Z. rhoifolium (Fig. 5A)
3 38 59 – * – – – – * – * – GFo Z. rhoifolium (Fig. 5B)
10 39 51 * * – – * – * – * * Ce Sp. cujabana (Fig. 5C)
15 43 42 * * – – – – – * – – GFo Sp. cujabana (Fig. 5D, 5H)
1 23 76 – * – – * – – – * – Ce V. tucanorum (Fig. 5E)
0.5 38.5 61 – * – – * – – – * – GFo V. tucanorum (Fig. 5F)
27 100 9 9 27 9 64 9 64 64 Ce Percentage
27 100 9 9 18 9 55 9 64 45 GFo Percentage

© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
 Fibres Intervessel pits

With minutely VRP of two

bordered pits With simple distinct sizes
(visible, to minutely in the same Vestured Alternate, PP PP
Septate but <3 mm) bordered pits Tyloses ray cell VRPAS VRPSI pits Polygonal circular Scalariform multiple simple CS Species

1
* – * * * * – – * * – *1 * Ce Tp. guianensis (Fig. 3A)
* – * * * * – – * * – *1 * GFo Tp. guianensis (Fig. 3B)
* * – – – – * – – * – – * Ce Tb. catharinensis (Fig. 3C)
* * – – – – * – – * – – * GFo Tb. catharinensis (Fig. 3D)
1
* – * * * * – – – * * * * Ce Sf. vinosa (Fig. 3E, 5I)
* – * – * * – – – * * * * GFo Sf. vinosa (Fig. 3F, 5J)
– * – – * * – – – * – – * Ce P. glabrata (Fig. 3G)
1
– * – * * * – – – * – – * GFo P. glabrata (Fig. 3H)
– – * – – – * * * – – – * Ce D. mollis (Fig. 4A)
– – * – – – * * * – – – * GFo D. mollis (Fig. 4B)
– – * – – – * * * * – – * Ce S. polyphyllum (Fig. 4C)
– – * – – – * * * * – – * GFo S. polyphyllum (Fig. 4D)
– – * – – – * * – * – – * Ce A. subelegans (Fig. 4E)
– – * – – – * * * * – – * GFo A. subelegans (Fig. 4F)
* – * * * * – * – * – – * Ce T. stenocarpa (Fig. 4G)
* – * – * * – * – * – – * GFo T. stenocarpa (Figs. 4H, 5G)
– – * – – – * – – * – – * Ce Z. rhoifolium (Fig. 5A)
– – * – – – * – – * – – * GFo Z. rhoifolium (Fig. 5B)
1
– * – * * * – – – * – * * Ce Sp. cujabana (Fig. 5C)
1
– * – * * * – – – * – * * GFo Sp. cujabana (Fig. 5D, 5H)
– – * * – – * * – * – – * Ce V. tucanorum (Fig. 5E)
– – * * – – * * * – – – * GFo V. tucanorum (Fig. 5F)
36 27 73 45 45 45 55 45 27 91 9 27 100 Ce Percentage
36 27 73 36 45 45 55 45 45 82 9 27 100 GFo Percentage

© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
COMPARATIVE WOOD ANATOMY OF CERRADO
7
 8

Table 2. Continued

Parenchyma Axial
Rays strand parenchyma Fibres
J. O. SONSIN ET AL.

1–3 Exclusively Over Up to 2–4 In Extremely Gelatinous Very Thin- to Very

cells wide uniseriate 7 cells 6 cells cells bands Paratracheal Apotracheal rare fibres thick-walled thick-walled thin-walled CS Species

3
* – * * * – * – – * – * * Ce Tp. guianensis (Fig. 3A)
* – * * * – * – – * – – * GFo Tp. guianensis (Fig. 3B)
* – – – – – – – * * – * – Ce Tb. catharinensis (Fig. 3C)
* – – – – – – – * – – * – GFo Tb. catharinensis (Fig. 3D)
* – * * * – * – – – – * * Ce Sf. vinosa (Fig. 3E, 5I)
* – – – * – * – – – – * * GFo Sf. vinosa (Fig. 3F, 5J)
– * * * * * – * – * – * – Ce P. glabrata (Fig. 3G)
– * * * * * – * – * – * – GFo P. glabrata (Fig. 3H)
3
* – * * * * * – – * – * – Ce D. mollis (Fig. 4A)
* – * * * * * – – * * – – GFo D. mollis (Fig. 4B)
3
– * – * * – * – – * – * – Ce S. polyphyllum (Fig. 4C)
– * – – * – * – – * – * – GFo S. polyphyllum (Fig. 4D)
3
– * – * * * * * – * * – – Ce A. subelegans (Fig. 4E)
* – – – * * * * – * * – – GFo A. subelegans (Fig. 4F)
* – – – * – * * – * * * – Ce T. stenocarpa (Fig. 4G)
* – – * * – * * – * – * – GFo T. stenocarpa (Figs. 4H, 5G)
* – – – * – * – – – – * – Ce Z. rhoifolium (Fig. 5A)
* – – – * – * – – – – * – GFo Z. rhoifolium (Fig. 5B)
* – – * * – – * – – * * – Ce Sp. cujabana (Fig. 5C)
* – – – * – – * – – – * – GFo Sp. cujabana (Fig. 5D, 5H)
3
* – * * * * * – – * – * – Ce V. tucanorum (Fig. 5E)
* – * * * – * – – * – – * GFo V. tucanorum (Fig. 5F)
82 18 45 82 91 36 73 36 9 73 27 91 18 Ce Percentage
82 18 36 36 91 27 73 36 9 64 18 64 27 GFo Percentage

© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
 Inorganic Organic Secretory elements and
inclusions inclusions cambial variants PRC Rays

Deposits Canal of Radial 4–5 cells

in vessels traumatic origin canals Heterocellular Homocellular wide CS Species

1
* – – * * * – – Ce Tp. guianensis (Fig. 3A)
1
* – – * * * – – GFo Tp. guianensis (Fig. 3B)
* – – – * * – – Ce Tb. catharinensis (Fig. 3C)
* – – – * * – – GFo Tb. catharinensis (Fig. 3D)
2
– – – * * * – * Ce Sf. vinosa (Fig. 3E, 5I)
– – – * * * – – GFo Sf. vinosa (Fig. 3F, 5J)
* * – – – * – – Ce P. glabrata (Fig. 3G)
* * – – – * – – GFo P. glabrata (Fig. 3H)
* * – – – * * – Ce D. mollis (Fig. 4A)
1
* * – – – * * – GFo D. mollis (Fig. 4B)
1
* * – – – – * – Ce S. polyphyllum (Fig. 4C)
1
* * – – – – * – GFo S. polyphyllum (Fig. 4D)
* * – – * * – – Ce A. subelegans (Fig. 4E)
* * – – * * – – GFo A. subelegans (Fig. 4F)
* * – – – * – – Ce T. stenocarpa (Fig. 4G)
– – – – – * – – GFo T. stenocarpa (Figs. 4H, 5G)
– – * – – * – – Ce Z. rhoifolium (Fig. 5A)
– – * – – * – – GFo Z. rhoifolium (Fig. 5B)
1
– * – – * * – – Ce Sp. cujabana (Fig. 5C)
1
– * – – * * – – GFo Sp. cujabana (Fig. 5D, 5H)
1
* * * – – * – * Ce V. tucanorum (Fig. 5E)
1
* * * – – * – * GFo V. tucanorum (Fig. 5F)
73 64 18 18 45 91 18 18 Ce Percentage
64 55 18 18 45 91 18 9 GFo Percentage

© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
CS, collection site; Ce, cerrado s.s.; GFo, gallery forest; PP, perforation plate; VRPSI, vessel-ray pits with distinct borders; similar to intervessel pits in size and shape throughout the ray cell; VRPAS,
vessel-ray pits with much reduced borders to apparently simple; pits rounded or angular/horizontal to vertical; VRP, vessel-ray pits; PRC, perforated ray cells; *present; –, absent; 1few; 2abundant;
3
of varying types. The growth ring definition and markers, perforation plate type (simple and multiple), intervessel and ray-vessel pits, fibre thickness, axial parenchyma type and strand, and
ray width might occur within a species; therefore, the sum of the percentage exceeds 100%.
COMPARATIVE WOOD ANATOMY OF CERRADO
9
 10

Table 3. Statistical analyses (ANOVA design nested up to a level of 5% of probability) comparing quantitative data between species and habitats

Species CS FL (mm) VL (mm) IVPD (mm) VRPD (mm) VD (mm) FWT (mm) R/mm V/mm2 VGI V M
A ± SD A ± SD A ± SD A ± SD A ± SD A ± SD A ± SD A ± SD A

Tp. guianensis GFo 670 ± 181 B b,c,d 461 ± 77 A c,d,e,f 10 ± 1 B c,d 15.6 ± 5 B f,g 90 ± 18 B d,e,f 2.7 ± 1 B a,b 6 ± 1 B b,c,d 34 ± 10 B e,f 1.46 A a,b,c,d 2.7 1241
J. O. SONSIN ET AL.

Ce 792 ± 169 A d,e,f 494 ± 116 A e,f 8.4 ± 1 A b,c 14 ± 4 A e,f,g 70 ± 21 A b,c,d,e 3.5 ± 1 A b,c,d,e 8.5 ± 2 A g,h,i 31 ± 14 A d,e,f 1.58 A a,b,c,d 2.3 1130
Tb. catharinensis GFo 831 ± 174 B d,e,f 482 ± 101 A d,e,f 4±0Ba 3.4 ± 1 B a 51 ± 6 B a 4.6 ± 1 B d,e,f,g,h,i 8 ± 3 B e, f,g,h 122 ± 11 B i 1.44A a,b,c,d 0.4 202
Ce 811 ± 172 A d,e,f 514 ± 111 A f 4.4 ± 1 A a 4 ± 1 A a,b 43 ± 8 A a 5.3 ± 2 A i,j 9.4 ± 3 A h,i 157 ± 32 A j 1.75 A c,d,e 0.3 141
Sf. vinosa GFo 820 ± 106 B d,e,f 689 ± 134 A g,h 24 ± 11 B f 15 ± 4 B e,f,g 71 ± 15 B b,c,d,e 4 ± 2 B c,d,e,f 4±1Ba 31 ± 9 B d,e,f 1.44 A a,b,c,d 2.3 1596
Ce 833 ± 175 A e,f 726 ± 152 A h,i 13 ± 3 A e 17.5 ± 9 A f,g 64 ± 12 A a,b,c 5.2 ± 1 A h,i,j 4.6 ± 1 A a,b 32 ± 9 A e,f 2.08 A e,f 2 1468
D. mollis GFo 783 ± 146 B d,e 361 ± 76 A b,c 7±1Bb 7 ± 2 B b,c 137 ± 27 B i 3.5 ± 1 B a,b,c,d,e 7 ± 2 B c,d,e,f 13 ± 5 B b,c 1.62 A a,b,c,d,e 10.5 3799
Ce 896 ± 150 A e,f 365 ± 62 A b,c 6.3 ± 1 A a,b 7.5 ± 3 A c 103 ± 24 A f,g,h 5.5 ± 1 A i,j 9 ± 2 A g,h,i 20 ± 6 A a,b,c 1.9 A d,e 5.2 1910
S. polyphyllum GFo 555 ± 111 B a 309 ± 77 A a,b 8.6 ± 1 B b,c 6.9 ± 1 B b,c 115 ± 29 B h,i 2.4 ± 1 B a 9.5 ± 2 B h,i 8±5Ba 1.3 A a 13.8 4248
Ce 599 ± 120 A a,b,c 350 ± 68 A a,b,c 6.8 ± 1 A a,b 6.4 ± 1 A a,b,c 101 ± 22 A e,f,g,h 3 ± 1 A a,b,c 7.5 ± 1 A d, e,f,g 36 ± 8 A f 1.54 A a,b,c,d 2.8 992
A. subelegans GFo 852 ± 153 B e,f 221 ± 40 A a 8 ± 1 B b,c 8.3 ± 1 B c,d 88 ± 18 B d,e,f 5 ± 1 B g,h,i,j 6.2 ± 1 B b,c,d,e 27 ± 6 B c,d,e,f 1.73 A a,b,c,d,e 3.3 721
Ce 976 ± 207 A f 233 ± 28 A a 10.3 ± 2 A c,d,e 8 ± 2 A c,d 75 ± 16 A b,c,d,e 6±1Aj 8.4 ± 2 A f,g,h,i 25 ± 7 A c,d,e 1.74 A b,c,d,e 3.0 693
T. stenocarpa GFo 616 ± 92 B a,b,c,d 456 ± 97 A c,d,e,f 11.4 ± 3 B c,d,e 13 ± 6 B d,e,f,g 91 ± 17 B d,e,f,g 4.9 ± 1 B f,g,h,i 11 ± 2 B i,j 22 ± 5 B b,c,d 1.48 A a,b,c,d 4.1 1871
Ce 572 ± 133 A a,b 410 ± 119 A b,c,d 8.2 ± 2 A b,c 13.3 ± 6 A e,f,g 83 ± 18 A c,d,e 4.6 ± 1 A e,f,g,h,i 9.7 ± 2 A i 35 ± 9 A f 1.35 A a 2.3 961
P. glabrata GFo 979 ± 144 B f 721 ± 159 A h,i 11.7 ± 3 B d,e 12 ± 4 B d,e,f 80 ± 17 B c,d,e 4.2 ± 2 B d,e,f,g 13 ± 3 B j 23 ± 7 B b,c,d 2.42 A f 3.4 2484
Ce 964 ± 170 A e,f 600 ± 133 A f,g 12 ± 2 A d,e 12 ± 4 A d,e,f 77 ± 22 A b,c,d,e 4.6 ± 1 A d,e,f,g,h,i 10.5 ± 3 A i 23 ± 9 A b,c,d 1.87A c,d,e 3.4 2038
Z. rhoifolium GFo 777 ± 121 B d,e 416 ± 77 A c,d 4.5 ± 1 B a 4 ± 1 B a,b 80 ± 17 B c,d,e 3.5 ± 1 B a,b,c,d 7 ± 1 B d,e,f,g 34 ± 8 B e,f 1.43 A a,b,c 2.4 986
Ce 721 ± 166 A c,d,e 438 ± 94 A c,d,e,f 4±1Aa 3.5 ± 1 A a,b 66 ± 14 A a,b,c,d 4 ± 1 A c,d,e 7 ± 1 A c,d,e,f,g 35 ± 8 A f 1.38 A a,b 1.9 827
Sp. cujabana GFo 1495 ± 237 B g 794 ± 204 A h,i 7±1Bb 12 ± 3 B d,e 65 ± 13 B a,b,c,d 9.4 ± 2 B k 7.5 ± 2 B e,f,g 84 ± 15 B g 2.04 A e 0.8 617
Ce 1934 ± 392 A h 838 ± 179 A i 4.5 ± 1 A a 13 ± 6 A d,e,f 57 ± 11 A a,b 11 ± 2 A l 10 ± 1 A i 105 ± 16 A h 1.86 A c,d,e 0.5 456
V. tucanorum GFo 769 ± 101 B c,d,e 428 ± 72 A c,d,e 7±1Bb 7.7 ± 2 B c 115 ± 31 B g,h 3.6 ± 1 B b,c,d,e 5.6 ± 1 B a,b,c,d 27 ± 12 B c,d,e,f 1.39 A a,b 4.2 1796
Ce 645 ± 127 A a,b,c,d 398 ± 101 A b,c,d 6.2 ± 1 A a,b 6 ± 2 A a,b,c 102 ± 24 A e,f,g,h 4.3 ± 2 A d,e,f,g,h 5 ± 1 A a,b,c 16 ± 4 A a,b 1.26 A a 6.5 2580
F 0.000333* 0.641751 0.000155* 0.048263* 0.006012* 0.000001* 0.029633* 0.000000* 0.171712

*Statistically significantly different at a 5% significance level of probability.

CS, collection site; GFo, gallery forest; Ce, cerrado s.s.; FL, fibre length; VL, vessel element length; IVPD, intervessel pit diameter; VRPD, vessel-ray pit diameter;
VD, vessel lumen diameter; FWT, fibre wall thickness; R/mm, ray per millimetre; V/mm2, vessels per square millimetre; VGI, vessel grouping index;
V, vulnerability index; M, mesomorphy index; A, average; SD, standard deviation. Uppercase letters indicate the comparison of quantitative data between
habitats. Lowercase letters indicate the comparison of quantitative data between species and habitats.

© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
 COMPARATIVE WOOD ANATOMY OF CERRADO 11

Figure 3. Transverse sections: A, Tapirira guianensis from cerrado (Ce); B, Tp. guianensis from gallery forest (GFo); C,
Tabernaemontana catharinensis from Ce; D, Tb. catharinensis from GFo; E, Schefflera vinosa from Ce; F, Sf. vinosa from
GFo; G, Pera glabrata from Ce; H, P. glabrata from GFo. Arrows point to the limit of the growth rings; gf, gelatinous
fibres. Scale bars, 100 mm.

© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
12 J. O. SONSIN ET AL.

Figure 4. Transverse sections: A, Dimorphandra mollis from cerrado (Ce); B, D. mollis from gallery forest (GFo); C,
Stryphnodendron polyphyllum from Ce; D, S. polyphyllum from GFo; E, Acosmium subelegans from Ce; F, A. subelegans
from GFo; G, Tibouchina stenocarpa from Ce; H, T. stenocarpa from GFo. Arrows point to the limit from the growth rings;
gf, gelatinous fibres. Scale bars, 100 mm.

© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
 COMPARATIVE WOOD ANATOMY OF CERRADO 13

Figure 5. Transverse sections: A, Zanthoxylum rhoifolium from cerrado (Ce); B, Z. rhoifolium from gallery forest (GFo);
C, Siparuna cujabana from Ce; D, Sp. cujabana from GFo; E, Vochysia tucanorum from Ce; F, V. tucanorum from GFo.
(A–F) Arrows point to the limit from the growth rings; gf, gelatinous fibres. G, Septate parenchyma (arrows) in Tibouchina
stenocarpa. H, Dumb-bell-shaped perforation plates (arrows) in Sp. cujabana from GFo. I, Wider radial canals (arrows)
in Schefflera vinosa from Ce. J, Narrower radial canals (arrows) in Sf. vinosa from GFo. Scale bars: (A–F, I, J) 100 mm;
(G, H) 50 mm.

© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
14 J. O. SONSIN ET AL.

Figure 6. (A–H) Comparison of quantitative data between species in the different habitats: A, fibre length; B, vessel
diameter length; C, intervessel pit diameter; D, vessel-ray pit diameter; E, vessel diameter; F, fibre wall thickness; G,
ray/mm; H, vessels/mm2.

© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
 COMPARATIVE WOOD ANATOMY OF CERRADO 15

Figure 7. Principal component analysis: Ce (black circles), cerrado s.s.; GFo (white circles), gallery forest. A: FL, fibre
length; VL, vessel element length; IVPD, intervessel pit diameter; VRPD, vessel-ray pit diameter; VD, vessel lumen
diameter; FWT, fibre wall thickness; R/mm, ray per millimetre; V/mm2, vessels per square millimetre; VGI, vessel
grouping index. B: A, Tapirira guianensis; B, Tabernaemontana catharinensis; C, Schefflera vinosa; D, Pera glabrata; E,
Dimorphandra mollis; F, Stryphnodendron polyphyllum; G, Acosmium subelegans; H, Tibouchina stenocarpa; I, Zan-
thoxylum rhoifolium; J, Siparuna cujabana; K, Vochysia tucanorum.

for radial growth to cease and rings to be well defined.
Marcati, Oliveira & Machado (2006) also found a high
incidence of growth rings which correlated with the
presence of a distinct annual dry season in cerrado
species. Furthermore, A. subelegans has well-defined
growth rings with the same marker in all specimens
from both habitats, which suggests that, in some
species, growth ring formation could also be partly
genetically determined. A forthcoming book with
about 90 cerrado species (J. O. Sonsin, P. Gasson, S.
R. Machado, C. Caum & C. R. Marcati, unpubl. data)
will show many more examples of this phenomenon in
cerrado species.
Most of the species do not have a distinct vessel
arrangement, i.e. their vessels are diffusely arranged
and not in a distinctive tangential, radial or diagonal

© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
16 J. O. SONSIN ET AL.
Table 4. High value data of principal component analysis
Variable Factor 1
Fibre length (mm) -0.873252*
Vessel element length (mm) -0.821138*
Intervessel pit diameter (mm) -0.006320
Vessel-ray pit diameter (mm) -0.369329
Vessel lumen diameter (mm) 0.699444
Fibre wall thickness (mm) -0.851011*
Ray/mm -0.326246
Vessel/mm2 -0.615054
Vessel grouping index -0.689091
Explained variance 3.746296
Proportion of total 0.416255
*Features that contributed most for axes 1, 2 and 3.
pattern. This corresponds with c. 80% of dicot woods
worldwide (Wheeler, Baas & Rodgers, 2007). Tb.
catharinensis, P. glabrata and Sp. cujabana have
radial vessel patterns in both habitats, but only
P. glabrata shows a more distinct radial pattern in
riparian samples. A similar pattern occurs in the
same species from the Atlantic forest (Barros et al.,
2001), but was not shown by Detienne & Jacquet
(1983) in Amazonian samples.
All species have solitary vessels and vessels in
short multiples, as do about two-thirds of dicot woods
worldwide (Wheeler et al., 2007). The cerrado s.s. and
gallery forest species do not have tracheids or vascu-
lar tracheids. A few species (Tb. catharinensis, P. gla-
brata and Sp. cujabana) have fibres with distinctly
bordered pits (< 3 mm). Carlquist (1984) speculated
that such fibres would provide limited protection
against embolism, whereas a radial arrangement of
vessels is likely to be safer.
Tapirira guianensis, Sf. vinosa and Sp. cujabana
have both simple and multiple perforation plates in
both habitats, but, in Tp. guianensis, the multiple
perforation plates are less frequent. Scalariform per-
foration plates are thought to reduce water flow (Car-
lquist, 1975; Baas & Schweingruber, 1987), and
lumen flow resistance has been shown experimentally
to be double that of simple perforations by Christman
& Sperry (2010). The combination of simple and sca-
lariform plates is seen as an adaptation to environ-
ments with seasonal droughts (Dickison & Phend,
1985), although many taxa do not have both.
There are also variations between habitats in axial
parenchyma patterns and the number of cells per
strand. Axial parenchyma is more abundant in
cerrado samples of D. mollis, S. polyphyllum, A. sub-
elegans and V. tucanorum. The greater proportion of
axial parenchyma in cerrado is probably for storage
and mobilization of metabolites in adverse periods,
© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
Factor 2 Factor 3
0.085033 0.078278
-0.322656 -0.163623
-0.882585* -0.206513
-0.832519* -0.037471
-0.184896 0.410972
0.161213 0.038579
0.143112 0.756148*
0.585214 -0.347460
-0.245837 0.439393
2.066950 1.132893
0.229661 0.125877
which are less harsh in riparian samples (see Baas,
1982; Carlquist, 1988; Mauseth, 1988; Alves &
Angyalossy-Alfonso, 2002), but this generalization
does not apply to all species. Axial parenchyma is
scant or absent in both habitats in Tp. guianensis,
Sf. vinosa, T. stenocarpa and Tb. catharinensis. These
four species have septate fibres which are thought to
have a similar storage function to axial parenchyma
(Carlquist, 1988; Fahn, 1990; Dickison, 2000). Any
storage tissue, either parenchyma or septate fibres,
can support rapid flushes of growth, flowering and
fruiting when conditions are suitable (Braun, 1984;
Carlquist, 1988).
Parenchyma cells in abundance are present in some
species, such as S. polyphyllum and V. tucanorum,
which have > 70% of solitary vessels in both habitats.
Are these parenchyma cells involved in embolism
repair in vessels? Nardini, Lo Gullob & Salleo (2011)
hypothesized a process whereby parenchyma cells
directly adjacent to an embolized vessel can refill it,
even under negative pressure, following changes in
carbohydrate metabolism from soluble sugars to
insoluble starch. In addition, the authors stated that
the decrease in the starch content in the adjacent cell
would cause these cells to become strong sinks to
phloem, with consequent unloading of sugars and
water from the phloem and a bulk flow directed to the
refilling conduits. We do not have experimental evi-
dence for this.
Although each species has a suite of qualitative
anatomical characters that is consistent between the
two habitats, the quantitative features are signifi-
cantly different. Past studies have linked such differ-
ences with climate, soil type and precipitation
(Klaassen, 1999; Alves & Angyalossy-Alfonso, 2000;
Dickison, 2000). Cerrado samples have more abun-
dant narrower vessels, longer fibres with thicker
walls, smaller intervessel pits and more rays than
 COMPARATIVE WOOD ANATOMY OF CERRADO
gallery forest samples of the same species. The PCA
shows that the same species are close together
because, despite their quantitative differences, their
anatomy is similar, except with regard to water
transport.
The cerrado s.s. and gallery forest species show low
vessel indices of 1.7 and 1.6, respectively, suggesting
that vessel failure rarely occurs in these species (see
Carlquist, 1984). The presence of simple and multiple
perforation plates in vessels and perforated ray cells
in some species (Tp. guianensis, Sf. vinosa and Sp. cu-
jabana), and abundant axial parenchyma cells in
others (D. mollis, S. polyphyllum, A. subelegans and
V. tucanorum), may explain the low vessel index for
both cerrado s.s. and gallery forest species. They may
enhance hydraulic safety for these species, as would a
high frequency of narrow vessels.
In general, when present, tyloses are more frequent
in cerrado s.s. species (Tp. guianensis, Sf. vinosa and
T. stenocarpa) than in gallery forest. Tyloses can be
formed in reaction to a wide variety of causes, such as
frost, flooding, heartwood formation, wounding and
natural senescence (Chattaway, 1949; Davison & Tay,
1985; Parameswaran, Knigge & Liese, 1985; Cochard
& Tyree, 1990; Dute, Duncan & Duke, 1999; Sun
et al., 2007). All of these phenomena can stimulate
ethylene production according to Sun et al. (2007). In
grapevine, they found that wound ethylene produc-
tion was the cause of tylosis formation, and that
embolisms in vessels are not directly required for
wound-induced tyloses. In cerrado s.s. species, the
lack of water in the driest period could stimulate
the increase in ethylene production, whereas, in the
gallery forest, there is standing water, even in dry
periods.
Gelatinous fibre distribution is usually in distinct
bands or small groups spread throughout the second-
ary xylem, but exhibits considerable variation within
species between habitats. Gelatinous fibres are
usually considered to be a phenomenon of tension
wood on the upper side of branches (Esau, 1974 and
many other authors), and so their presence is to be
expected. The innermost layer of gelatinous fibres is
mainly cellulose, which is hydrophilic (Esau, 1965;
Mauseth, 1988; Fahn, 1990), suggesting that they
might also have a water storage function (Paviani,
1978). Although, to our knowledge, there is no direct
experimental evidence for water storage in gelatinous
fibres, their abundance in the wood of taxa growing in
dry environments suggests that this relationship is
worth exploring further. Milanez, Marcati & Machado
(2008, 2009) found that aluminium accumulates in
the gelatinous fibre walls and septate fibres, both
of pectic-cellulosic nature, of Melastomataceae. A
hypothesis considered by many authors is that the
means by which plants interact with the presence of
© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
17
aluminium seems to prevent its entry into the cell by
its deposition on the cell wall, by compartmentaliza-
tion in vacuoles or plastids, or by the formation of
aluminium chelates (Cuenca, Herrera & Merida,
1991; Britez et al., 2002; Milanez et al., 2009). There
is therefore some evidence that gelatinous fibres are
not only indicative of tension wood, but could also
sequester aluminium, which is potentially harmful to
plants.
The gallery forest samples are better optimized for
conductive efficiency (wider vessels, larger intervessel
pits, simple perforation plates) than are cerrado
samples, which have a strategy for safety reflected by
narrower vessels that are less efficient at water trans-
port, but also less prone to embolism (Baas, Wheeler
& Fahn, 1983; Schulte, 1999; Dickison, 2000; Tyree &
Zimmermann, 2002; Zanne et al., 2006; Loepfe et al.,
2007; Pratt et al., 2007).
The vulnerability and mesomorphy indices of
cerrado (2.9 and 1295, respectively) and gallery forest
samples (5 and 2167, respectively) correspond with
this hypothesis, but are based on observation and not
experiment.
CONCLUSIONS
We have demonstrated considerable differences in the
quantitative characteristics of the secondary xylem
among the 11 species which occur in both cerrado s.s.
and gallery forests, particularly in the vessel features.
The wood of the species from gallery forest is better
optimized for conductive efficiency (wider vessels,
larger intervessel pits, simple perforation plates and
higher vulnerability and mesomorphy indices) than
the wood from cerrado species, which have a strategy
for safety reflected in a higher frequency of narrower
vessels that are less efficient at transporting water,
but less prone to embolism. If, as seems likely, these
characteristics are at least partly genetically control-
led, when restoring cerrado habitats, it is important
to consider the ecological provenance of the trees to be
reintroduced, especially if they have a broad habitat
tolerance.
ACKNOWLEDGEMENTS
Thanks are due to the Coordination for the Improve-
ment of Higher Level Personnel (CAPES) and the
National Council of Technological and Scientific
Development (CNPq) for the sponsorship of this
research, to Liliane C. Pereira for technical assistance
in the laboratory, to Clemente José Campos who pro-
vided assistance in the field and to the Royal Botanic
Gardens, Kew, for general support and for hosting the
first author.
18 J. O. SONSIN ET AL.
REFERENCES
Alves ES, Angyalossy-Alfonso V. 2000. Ecological trends in
the wood anatomy of some Brazilian species. 1. Growth
rings and vessels. International Association of Wood Anato-
mists Journal 21: 3–30.
Alves ES, Angyalossy-Alfonso V. 2002. Ecological trends in
the wood anatomy of some Brazilian species. 2. Axial paren-
chyma, rays and fibres. International Association of Wood
Anatomists Journal 23: 391–418.
Baas P. 1982. Systematic, phylogenetic, and ecological wood
anatomy: history and perspectives. In: Baas P, ed. New
perspectives in wood anatomy. Leiden: Martinus Nijhoff
Publishers, 23–58.
Baas P, Schweingruber F. 1987. Ecological trends in the
wood anatomy of trees, shrubs and climbers from Europe.
Bulletin of the International Association of Wood Anatomists
8: 245–274.
Baas P, Wheeler E, Fahn A. 1983. Some ecological trends in
vessel characters. Bulletin of the International Association
of Wood Anatomists 4: 141–159.
Barros CF, Callado CH, Marcon ML, Costa CG, Cunha
M, Lima HRP, Marquete O. 2001. Madeiras da Mata
Atlântica: anatomia do lenho de espécies ocorrentes nos
remanescentes florestais do estado do Rio de Janeiro –
Brasil, v.2. Rio de Janeiro: Programa Mata Atlântica,
Instituto de Pesquisas Jardim Botânico do Rio de Janeiro.
Barros CF, Marcon-Ferreira ML, Callado CH, Lima
HRP, Da Cunha M, Marquete O, Costa CG. 2006.
Tendências ecológicas na anatomia da madeira de espécies
da comunidade arbórea da Reserva Biológica de Poço das
Antas. Rodriguésia 57: 443–460.
Braun HJ. 1984. The significance of the accessory tissues of
the hydrosystem for osmotic water shifting as the second
principle of water ascent, with some thoughts concerning
the evolution of trees. Bulletin of the International Associa-
tion of Wood Anatomists 5: 275–294.
Britez RM, Watanabe T, Jansen S, Reissmann CB, Osaki
M. 2002. The relationship between aluminium and silicon
accumulation in leaves of Faramea marginata (Rubiaceae).
New Phytologist 156: 437–444.
Bukatsch F. 1972. Bermerkungen zur Doppelfälbung
Astrablau-Safranin. Mikrokosmos 61: 33–36.
Carlquist S. 1975. Ecological strategies of xylem evolution.
Berkeley, CA: University of California Press.
Carlquist S. 1977. Ecological factors in wood evolution:
a floristic approach. American Journal of Botany 64: 887–
896.
Carlquist S. 1984. Vessel grouping in dicotyledon wood:
significance and relationship to imperforate tracheary ele-
ments. Aliso 10: 505–525.
Carlquist S. 1988. Comparative wood anatomy: systematic,
ecological and evolutionary aspects of dicotyledon wood.
Berlin: Springer Verlag.
Carlquist S. 2010. Caryophyllales: a key group for under-
standing wood anatomy character states and their
evolution. Botanical Journal of the Linnean Society 164:
342–393.
© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
Castro AAJF, Martins FR, Tamashiro JY, Shepherd GJ.
1999. How rich is the flora of Brazilian cerrados? Annals of
the Missouri Botanical Garden 86: 192–224.
Chattaway MM. 1949. The development of tyloses and secre-
tion of gum in heartwood formation. Australian Journal of
Scientific Research 2: 227–240.
Christman MA, Sperry JS. 2010. Single-vessel flow meas-
urements indicate scalariform perforation plates confer
higher flow resistance than previously estimated. Plant,
Cell and Environment 33: 431–443.
Cochard H, Tyree MT. 1990. Xylem dysfunction in Quercus:
vessel sizes, tyloses, cavitation and seasonal changes in
embolism. Tree Physiology 6: 393–408.
Coutinho LM. 1990. Fire in the ecology of the Brazilian
Cerrado. In: Goldammer JG, ed. Fire in the tropical biota,
ecosystem processes and global challenges. Berlin: Springer
Verlag, 81–105.
Coutinho LM. 2002. O bioma do cerrado. In: Klein AL, ed.
Eugen Warming e o cerrado brasileiro. São Paulo: Imprensa
Oficial do Estado, 77–92.
Cuenca G, Herrera R, Merida T. 1991. Distribution of
aluminium in accumulator plants by X-ray microanalysis
in Richeria grandis Vahl leaves from a cloud forest in
Venezuela. Plant, Cell & Environment 14: 437–441.
Davison EM, Tay FCS. 1985. The effect of waterlogging on
seedlings of Eucalyptus marginata. New Phytologist 101:
743–754.
Detienne P, Jacquet P. 1983. Atlas d’identification des bois
de l’amazonie et des regions voisines. Nogent sur Marne:
Centre Technique Forestier Tropical.
Dickison WC. 2000. Integrative plant anatomy. San Diego,
CA: Harcourt Academic Press.
Dickison WC, Phend KD. 1985. Wood anatomy of the
Styracaceae: evolutionary and ecological considerations.
Bulletin of the International Association of Wood Anatomists
6: 3–22.
Durigan G, Baitello JB, Franco GADC, Siqueira MF.
2004. Plantas do Cerrado Paulista: imagens de uma pai-
sagem ameaçada. São Paulo: Páginas & Letras Editora e
Gráfica.
Durigan G, Franco GADC, Siqueira MF. 2004. A ve-
getação dos remanescentes de cerrado no Estado de São
Paulo. In: Bitencourt MD, Mendonça RR, eds. Variedade de
conservação dos remanescentes de cerrado no Estado de Sao
Paulo. Sao Paulo: Annablume, 29–56.
Dute RR, Duncan KM, Duke B. 1999. Tyloses in abscission
scars of loblolly pine. International Association of Wood
Anatomists Journal 20: 67–74.
Eiten G. 1972. The cerrado vegetation of Brazil. Botanical
Review 38: 201–341.
Esau K. 1965. Plant anatomy, 2nd edn. New York: John Wiley
& Sons.
Esau K. 1974. Anatomia das plantas com sementes. São
Paulo: EDUSP.
Fahn A. 1990. Plant anatomy. Oxford: Pergamon Press.
Franklin GL. 1945. Preparation of thin sections of synthetic
resins and wood-resin composites, and a new macerating
method for wood. Nature 155: 51.
 COMPARATIVE WOOD ANATOMY OF CERRADO
Hammerle INC. 2006. Ecofisiologia comparativa de espécies
arbóreas de cerrado s.s. e mata de galeria. Tese de Douto-
rado, Universidade de Brasília.
Hijmans RJ, Cameron SE, Parra JL, Jones PG, Jarvis A.
2005. Very high resolution interpolated climate surfaces for
global land areas. International Journal of Climatology 25:
1965–1978.
IAWA Committee. 1989. List of microscopic features for
hardwood identification. Bulletin of the International
Association of Wood Anatomists 10: 219–332.
Jonsson M, Malmqvist B. 2000. Ecosystem process rate
increases with animal species richness: evidence from leaf-
eating, aquatic insects. Oikos 89: 519–523.
Klaassen R. 1999. Wood anatomy of the Sapindaceae.
International Association of Wood Anatomists Journal
Supplement 2: 1–214.
Kraus JE, Arduin M. 1997. Manual básico de métodos em
morfologia vegetal. Rio de Janeiro: Seropédica, EDUR.
Lens F, Luteyn JL, Smets E, Jansen S. 2004. Ecological
trends in the wood anatomy of Vaccinioideae (Ericaceae s.l.).
Flora 199: 309–319.
Lista de Espécies da Flora do Brasil. 2011. Available at:
http://floradobrasil.jbrj.gov.br/2011/index?mode=sv&group=
Root_.Angiospermas_&family=&genus=&species=&author=
&common=&occurs=1&region=&state=&phyto=3&endemic=
&origin=&last_level=subspecies&listopt=1&x=23&y=8.
(Acessed December 2011)
Loepfe L, Martinez-Vilalta J, Pinol J, Mencuccini M.
2007. The relevance of xylem network structure for plant
hydraulic efficiency and safety. Journal of Theoretical
Biology 247: 788–803.
Ludwig JA, Reynolds JF. 1988. Statistical ecology: a
primer on methods and computing. New York: John Wiley &
Sons.
Machado SR, Rodella RA, Angyalossy-Alfonso V, Marcati
CR. 2007. Structural variation in root and stem wood of
Styrax (Styracaceae) from Brazilian Forest and Cerrado.
International Association of Wood Anatomists Journal 28:
173–188.
Marcati CR, Angyalossy-Alfonso V, Benetati L. 2001.
Anatomia comparada do lenho de Copaifera langsdorffii
Desf. (Leguminosae-Caesalpinoideae) de floresta e cerradão.
Revista Brasileira de Botânica 24: 311–320.
Marcati CR, Oliveira JS, Machado SR. 2006. Growth rings
in cerrado woody species: occurrence and anatomical
markers. Biota Neotropica 6: 1–31. Available at: http://
www.Biotaneotropica.org.br/v6n3/pt/abstract?article+
bn00206032006> (Accessed 4 December 2006)
Mauseth JD. 1988. Plant anatomy. San Francisco, CA:
Benjamin Cummings Publishing.
Milanez CRD. 2007. Estudos anatômicos e ultra-estruturais
em Melastomataceae de cerrado. Tese de Doutorado, Uni-
versidade Estadual Paulista.
Milanez CRD, Marcati CR, Machado SR. 2008. Uma nova
interpretação do papel das fibras gelatinosas e septadas em
plantas acumuladoras de alumínio do cerrado. In: XXVIII
Encontro Regional de Botânica, 2008, Carangola – Minas
Gerais. Anais do XXVIII Erbo – Ambientes de Montanha:
© 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••
19
Pesquisa e Conservação. Universidade do Estado de Minas
Gerais.
Milanez CRD, Marcati CR, Machado SR. 2009. Sítios
de acúmulo de alumínio no lenho de três espécies de
Melastomataceae do cerrado. In: 60º Congresso Nacional de
Botânica, 2009, Feira de Santana – Salvador. Anais do 60º
Congresso Nacional de Botânica. Universidade do Estado da
Bahia.
Mittermeier RA, Myers N, Gil PR, Mittermeier CG. 1999.
Hotspots. Mexico City: CEMEX.
Nardini A, Lo Gullob MA, Salleo S. 2011. Refilling embol-
ized xylem conduits: is it a matter of phloem unloading?
Plant Science 180: 604–611.
Oliveira JS. 2007. Variações estruturais do lenho de espécies
de cerrado do estado de São Paulo. Dissertação de Mestrado,
Universidade Estadual Paulista.
Oliveira PS, Marquis RJ. 2002. The cerrados of Brazil:
ecology and natural history of a Neotropical savanna. New
York: Columbia University Press.
Parameswaran N, Knigge H, Liese W. 1985. Electron
microscopic demonstration of a suberized layer in the tylosis
wall of beech Fagus sylvatica and oak Quercus robur.
Bulletin of the International Association of Wood Anatomists
6: 269–271.
Paviani TI. 1978. Anatomia vegetal e cerrado. Ciência e
Cultura 30: 1076–1086.
Pockman WT, Sperry JS. 2000. Vulnerability to xylem
cavitation and the distribution of Sonoran desert vegetation.
American Journal of Botany 87: 1287–1299.
Pratt RB, Jacobsen AL, Ewers FW, Davis SD. 2007.
Relationships among xylem transport, biomechanics and
storage in stems and roots of nine Rhamnaceae species
of the California chaparral. New Phytologist 174: 787–
798.
Sass JE. 1958. Elements of botanical microtechnique. New
York, London: McGraw-Hill.
Schulte PJ. 1999. Water flow through a 20-pore perforation
plate in vessels of Liquidambar styraciflua. Journal of
Experimental Botany 50: 1179–1187.
Sun Q, Rost TL, Reid MS, Matthews MA. 2007. Ethylene
and not embolism is required for wound-induced tylose
development in stems of grapevines. Plant Physiology 145:
1629–1636.
Tyree MT, Zimmermann MH. 2002. Xylem structure and
the ascent of sap. Berlin: Springer.
Wheeler EA, Baas P, Rodgers S. 2007. Variations in dicot
wood anatomy: a global analysis based on the Inside Wood
database. International Association of Wood Anatomists
Journal 28: 229–258.
Worbes M. 1995. How to measure growth dynamics in
tropical trees: a review. International Association of Wood
Anatomists Journal 16: 337–351.
Zanne AE, Sweeney K, Sharma M, Orians CM. 2006.
Patterns and consequences of differential vascular sectori-
ality in 18 temperate tree and shrub species. Functional
Ecology 20: 200–206.
Zanne AE, Westoby M, Falster DS, Ackerly DD, Loarie
SR, Arnold SEJ, Coomes DA. 2010. Angiosperm wood
20 J. O. SONSIN ET AL.

structure: global patterns in vessel anatomy and their rela- Zar JH. 1996. Biostatistical analysis, 3rd edn. Englewood
tion to wood density and potential conductivity. American Cliffs, NJ: Prentice Hall.
Journal of Botany 97: 207–215.

SUPPORTING INFORMATION
Additional Supporting Information may be found in the online version of this article:
Appendix S1. Wood anatomical descriptions.
Please note: Wiley-Blackwell are not responsible for the content or functionality of any supporting materials
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