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Summary
Introduction
The genus Picconia belongs to the Oleaceae andt is endemic to the Macaronesian Region
with Picconia excelsa (Ait.) DC., occurring in Madeira and the Canary archipelago,
and Picconia azorica (Tutin) Knobl. in the Azores.
Picconia azorica is an evergreen tree with smooth, pale bark. The Portuguese ver-
nacular name of the species is pau-branco, meaning that its wood colour is white to
pinkish, though it becomes darker over time. It can grow up to 15 m in height and the
maximum diameter registered is 53 cm for a tree of undetermined age (Dias 2001),
1) Department of Agriculture, Forests, Nature and Energy (DAFNE), Universit degli Studi della
Tuscia, Via S. Camillo de Lellis, 01100 Viterbo, Italy.
2) Azores Regional Coordinator for Nature Conservation, Azorina, S.A. - Sociedade de Gesto
Ambiental e Conservao da Natureza, Rua do Galo 118, 9700-091 Angra do Herosmo,
Azores, Portugal.
*) Corresponding author [E-mail: schirone@unitus.it].
Associate Editor: Steven Jansen
376 IAWA Journal, Vol. 33 (4), 2012
but for a specimen with a diameter of 22 cm, an age exceeding 64 years was inferred
(sample held at the Museu Carlos Machado, Ponta Delgada, So Miguel Island). The
species is xerophytic, apt to colonize dry environments and is resistant to sea spray
(Dias 2001).
The first references to P. azorica in the Azores were written over a century after the
Portuguese settlement in 1440 by Frutuoso (1583), who reported the occurrence of
dense stands of this species on all nine islands. Only in 1844 Seubert and Hochstetter
provided the first botanical description of the plant under the denomination of P. excelsa,
considering it identical to the Picconia species found in Madeira and Canary Islands.
Later, the species was revised by Tutin (1953) and placed within another genus, with
the denomination of Notelaea azorica Tutin, presently recognized as Picconia azorica
(Tutin) Knobl. (1934). This taxonomic delimitation has recently been confirmed by
Ferreira et al. (2011) after genetic study. The species is currently present in all the
Azorean islands, with the exception of Graciosa (Schaefer 2002), widespread in small
populations of coastal and marginal sites. More than one thousand reproductive indi-
viduals occur in the archipelago, even if populations have been declining over the last
three decades because of harvesting without restocking, new plantations with exotic
species, utilization as wood fuel, clearing land for agricultural purposes, and because of
its valuable wood (Martn et al. 2008; International Union for Conservation of Nature,
IUCN 2011).
Indeed, P. azorica was of important social value in the past, being used to build
wooden wagons, agricultural machinery and tools, house beams, as well as luxury fur-
niture, especially in the period 15801640 (Frutuoso 1583; Martins 1981; Dias et al.
2007).
Nowadays, P. azorica is classified as Endangered (IUCN 2011) and protected under
the Habitat Directive (Annexes II and IV) (EC 1992) and Bern Convention (Annex I)
(Council of Europe 1993). The fragile status of P. azorica led it to be included in the
100 taxa (endemic and non-endemic) list of Macaronesian Flora priority species, for
which management and conservation actions are crucial for their survival and population
increment (Martn et al. 2008). These authors classified the species with an ecological
value of structuring species. Additionally, it plays an important ecological role as
part of the diet of two protected Azorean endemic bird species: Columba palumbus
azorica and Pyrrhula murina (Dias et al. 2007; SPEA 2008).
In 1988, Baas et al. investigated the wood anatomical features of the Oleaceae,
including P. excelsa, enabling wood identification of specimens belonging to this
family.
This study represents the first wood anatomical and technological investigation of
P. azorica based on both micro- and macroscopic analyses. Here we investigated the
number of vessels, tangential and radial diameters, wall thickness, vessel, tracheid,
libriform fibre and parenchyma strand lengths. Ray features and density were measured
and compared with P. excelsa. At the same time, colour, pH, wood density, compres-
sion and bending strengths, shrinkage, static quality factor, ash content and HHV were
investigated and compared with other Oleaceae and other hardwoods.
Caetano Ferreira et al. Wood of Picconia azorica 377
Microscopic investigations
Wood samples were investigated using both light (LM) and scanning electron mi-
croscopy (SEM).
Transverse, radial and tangential sections of 2025 m thick of each tree were
prepared from cubic samples of 515 mm wide, with the aid of a sliding microtome
(Reichert Jung 1150 Autocut). Sections were then cut, and stained with safranine (1%
in 50% ethanol). After staining, sections were washed with distilled water, dehydrated
with ethanol and mounted on slides with Canadian Balsam.
Observations were first carried out with a Wild M420 stereomicroscope, then with
LM using a Reichert-Jung Polivar 100. Pictures were taken with a Moticam 2500
5.0 M Pixel digital camera and analyzed using Motic Image PLUS 2.0 ML software
package.
Other samples were fixed with 4% paraformaldehyde + 5% glutaraldehyde, pH 7.2
in 0.1 M cacodylate buffer for 1 h at 4 C (Karnovsky 1965) to be scanned by SEM.
After rinsing overnight in the same buffer, the sections were post-fixed in cacodylate-
buffered 1% osmium tetroxide for 1 h, and sections were dehydrated in a graded
acetone series, dried by the critical point method using CO2 in a Balzers Union CPD
020, sputter-coated with gold in a Balzers MED 0.10 unit, and observed with a JEOL
JSM 5200 scanning electron microscope.
Cellular morphology was investigated also by microscopic observations on macer-
ated wood from 8 samples reduced to slivers about 1 mm thick and 2 cm long, and
placed in vials with 1:1 mixture of acetic acid and hydrogen peroxide (30 vol.) for
48 h. The vials were then placed in a boiling water bath for 48 h, the fluid was gently
removed and the slivers were rinsed several times in distilled water, and a saturated
sodium bicarbonate solution was added to buffer the acidity excess. After some rinsing,
distilled water was added again, the tubes were shaken vigorously to obtain xylem cell
separation, and some suspension drops were placed in microscope slide covered with
slips. The identification of the cellular elements was conducted using a Zeiss Axioskop
378 IAWA Journal, Vol. 33 (4), 2012
microscope, the measurements were performed with Axio Vision AC system, and the
anatomical descriptions were conducted following the IAWA list of microscopic features
for hardwood identification (Wheeler et al. 1989).
Figure 1. Disk (a) and tangential surface (b) of the Picconia azorica sample used in this study.
White solid circles indicate pH determination measurements with the respective values. White
solid squares marked with d and b and subscript numbers show the measurement performed for
defining the colour of the disk and the board respectively.
Caetano Ferreira et al. Wood of Picconia azorica 379
Results
Macroscopic features
Seasoned (air-dried) wood colour is yellowish verging on pale brown, and no dif-
ferentiation, checked by eye and under UV light, has been observed between heartwood
and sapwood (Fig. 1a). The odour was not appreciable on freshly exposed surfaces.
Annual growth rings are evident but sometimes false or incomplete, and a moderately
fine texture was observed.
Microscopic features
Picconia azorica growth rings are delimited by marginal parenchyma bands and
by medium thick-walled latewood fibres (Fig. 3b). The species has diffuse-porous
wood with grouped vessels, arranged in oblique to dendritic pattern (Y or X shapes)
(Fig. 3a).
Vessels are rounded to quadrangular in cross section (Fig. 3b, c), 47106 (21116)
per mm2; confidence limits at level < 0.05, calculated on N = 30 random vessels with
minimum and maximum values given in parentheses, are 4274 (2885) m for tan-
gential diameter and 1855 (1587) m for radial diameter, while the walls are 25 m
thick. A low percentage of solitary vessels was recorded, with vascular tracheids and
paratracheal parenchyma (occasionally apotracheally diffuse). Vessel member length
380 IAWA Journal, Vol. 33 (4), 2012
Figure 2. SEM illustration of pit membrane in radial section (a) bearing a torus (T) surrounded by
a margo (M) with the annulus (A) at the outline of the membrane; overview of pit membrane in
radial section (b) with tori (T) and hypothetical pseudo-tori (P). Scale bars: a = 1 m; b = 5 m.
Figure 3. Microscopic features of Picconia azorica as observed at light and scanning electron mi-
croscope. a: Grouped vessels in cross section arranged in oblique to dendritic pattern. b: Mar-
ginal parenchyma and medium thick-walled latewood fibres in cross section (arrow).
c: Rounded to quadrangular vessels in cross section. d: Bi-seriate and tri-seriate rays in tangential
section. e: Procumbent and erect body ray cells with square marginal cells in radial section.
Caetano Ferreira et al. Wood of Picconia azorica 381
f: Helical thickenings in a vessel element in tangential section. g: Diamond and cubic shaped
crystals in ray cells. h: Libriform fibre in macerated wood. i: Crystals of various shapes and
spherical bodies. Scale bars for c: 500 m; a: 200 m; b, d, e: 100 m; f: 50 m; i: 5 m.
382 IAWA Journal, Vol. 33 (4), 2012
Vascular tracheids (N = 270), 108383 (42565) m long, are associated with vessel
groups and normal narrow vessel elements.
Libriform fibres (N = 175) are 620921 (4221060) m long and very thick-walled
(Fig. 3h). Fibre to vessel length ratio (F/V) ranges between 2.1 and 3.3.
Axial parenchyma forms complete to incomplete sheaths (14 cells of adjoining
parenchyma cells sheathing part of the vessels) all around the vessel/tracheid groups,
in 16-seriate marginal bands. The axial parenchyma cells are square with a length
ranging from 41 m to 245 m (N = 91). Parenchyma strand length varies from 25
up to 6 cells long.
Rays 11 up to 13 per mm (816), 711 (412) cells high, uniseriate or bi-(tri-)seriate
(Fig. 3d), including heterogeneous II, occasionally III, and less frequently homogeneous
types (cf. Kribs 1935; Carlquist 2001). Body ray cells are procumbent or rarely erect,
with one row upright of square marginal cells (Fig. 3e).
Spherical starch grains were observed by SEM analysis together with crystals of
various shapes (cubic, diamond) and sizes in ray and axial parenchyma cells (Fig. 3g, i).
In comparison with P. excelsa described by Baas et al. (1988), the anatomical features
of P. azorica wood show differences in terms of dimension ranges of some elements
(e.g. vessel number and libriform fibre length), but not significantly so as to provide a
relevant distinction between these species at the anatomical level. Conversely, some
characteristics, such as the number of vessels per mm2, and eventually the type of rays
heterogeneity might be worthy of consideration in order to differentiate the two Pic-
conia species (see Table 1 for descriptive statistics, including mean values and standard
deviation of the measured microscopic features).
P. azorica P. excelsa
Mean SD UCL-LCL (min-max) UCL-LCL (min-max)
Number of vessels/mm2 76 30 47106 (21116) 3580
Tangential diameter (m) 58 16 4274 (2885) 4555 (2585)
Radial diameter (m) 36.3 18.5 1855 (1587) > 90
Wall thickness (m) 25 25
Vessel element length (m) 308 123 184431 (103592) 360420 (280690)
Tracheid length (m) 246 137 108383 (42565)
Libriform fibre length (m) 771 151 620921 (4221060) 9901190 (6501460)
Parenchyma cell length (m) 90 35 55125 (41245)
F/V ratio 2.13.3 2.62.8
Ray width (no. of cells) 12 12 (rarely 3)
Ray height (no. of cells) 9 2 711 (412) 610 (115)
Number of rays/mm 12 2 1113 (816) 1012 (914)
Ray type Heterogeneous II Heterogeneous III
(rarely III) (rarely II)
Caetano Ferreira et al. Wood of Picconia azorica 383
Table 4. Mean values for each parameter examined for the physical and mechanical charac-
terization and p-values from the analysis of variance at < 0.05 probability level (ANOVA
Kruskal Wallis non-parametric test).
LCL and UCL refer to upper and lower confidence limits stated at 95% confidence level.
which data are available in the literature (Della Patria & Omarini 2009; Genco et al.
2011). Results indicate that a* and b* values of P. azorica fall within the range of both
broadleaves, while L* is highest (Table 3).
Results from pH analysis showed homogeneous conditions between the sampled
disk and board with values from 4.63 to 5.02 on the first, 4.625.30 on the second
(Fig. 1a, b), and 4.805.20 on the specimens sections (not shown).
The density of the study samples allowed classifying P. azorica wood as hard, with
an average of 0.77 0.05 g cm-3 and 0.82 0.05 g cm-3 at 0% and 12% of moisture
content, respectively (Table 4). Radial, axial, tangential, and volumetric total shrink-
age tests r were 6.29 1.22%, 1.36 0.75%, 8.12 0.66%, and 13.9 1.45% respec-
tively.
The determined HHV mean value was 18.66 1.20 MJ kgdw-1, and the ash content,
in percentage on dry weight, resulted of 1.34 0.25% (Table 4).
Concerning the compression ( y) and the bending strengths ( b), average values of
61.85 6.68 MPa and 92.18 21.97 MPa were calculated, and the mean static quality
factor was 7.46 0.46 km (Table 4).
The comparison of physical and mechanical features among P. azorica samples
showed no statistical differences at < 0.05 probability level as assessed by p-values
attained by the ANOVA non-parametric Kruskal Wallis tests reported for each variable
in Table 4.
Conversely, the comparison with other species occurring in Europe and retrieved
from the literature (Table 5) showed that P. azorica wood density is similar to Olea
europaea L. among the Oleaceae, while it is higher than almost any other hardwoods
except for Quercus ilex L. and Quercus pubescens Willd. (Table 5).
Caetano Ferreira et al. Wood of Picconia azorica 385
Note: a Berti et al. (1979); b Giordano (1981); c Kitani & Hall (1989); d Berti et al. (1991); e Tsoumis
(1991); f Hellrigl (2006); g Telmo & Lousada (2011); h Zamorano et al. (2011); i Lo Monaco et al.
(2011b). MC = Moisture content.
386 IAWA Journal, Vol. 33 (4), 2012
Discussion
Generally, the observed anatomical features of Picconia azorica are similar to P. ex-
celsa, with some exceptions regarding the number of vessels per mm, and the ray type.
To explain these overall similarities we might hypothesize the recent geological events
of the Macaronesia Archipelagos causing a differentiation process between P. excelsa
and P. azorica not sufficient to have an effect on the wood anatomy (Schirone et al.
2010; Fernandez-Palacios et al. 2011; Ferreira et al. 2011; Schaefer et al. 2011; Triantis
et al. 2011). Overall, the characteristics that describe P. azorica wood anatomy are in
agreement with the group V defined by Baas et al. (1988). To this group belong the
genera Phillyrea, Picconia, Nestegis, Notelaea, Osmanthus, and Olea C, which are
characterized by having nonseptate libriform fibres, marginal parenchyma, vessels in
oblique to dendritic pattern, usually associated with vascular tracheids in a vasicentric
position, vessels with well-developed spiral thickenings, intervessel pits more than
6 m in diameter (Baas et al. 1988). Within the same group, the genera Picconia and
Osmanthus are distinguished by the occurrence of inter-vessel and inter-tracheary pits
with tori (Dute et al. 2008). At the same time, the opportunity to discriminate between
P. azorica and P. excelsa by investigating the taxonomic value of the torus dimensions
is still not confirmed (cf. Rabaey et al. 2008).
Concerning the macroscopic features of P. azorica, colour, grain and texture are
crucial characteristics for the aesthetic evaluation of its wood. The wood is aesthetically
pleasing and odourless, and thus suitable for furniture making and decorative applica-
tions. The slightly acid wood pH values, above the lower limit of metal corrosion of
4.0 4.3 found by Farmer (1967), do not differ from those in most of the studied woods
(Xing et al. 2004; He & Yan 2005). Thus the bonding capacity and the corrosiveness,
in particular under humid conditions, are similar to other species used for furniture
making (Packman 1960; Giordano 1981; Landi & Staccioli 1992).
Caetano Ferreira et al. Wood of Picconia azorica 387
Despite the investigations performed in the last 30 years regarding the physical
and mechanical properties of several Oleaceae (cf. Berti et al. 1979; Giordano 1981,
1988; Tsoumis 1991), no information about the genus Picconia was available before
this study.
Historical references attest to the woods hardness and its difficulty to work (Dias
et al. 2007). Indeed, Picconia azorica wood was observed to be heavy, falling within
those species with high density, and classified as hard according to Berti (1985).
Volumetric shrinkage, tangential/radial shrinkage ratio, resistance to compression
and bending, as well as static quality factor confirm the effectiveness of the wood in fur-
niture and pavement. For such purposes, P. azorica shows a performance more compa-
rable to Juglans regia than to other Oleaceae. On the other hand, the high density, the
considerable ash content and the calorific potential retrieved from P. azorica samples
classify the species as not recommendable for biomass, as also confirmed by the com-
parison with other Portuguese species (Telmo & Lousada 2011).
The history of P. azorica utilization demonstrated that this species was appreciated
by the inhabitants of the Azorean archipelago as an income resource since the first
settlements in the XV century (Dias et al. 2007). Indeed, Picconia azorica, together
with other endemic species as Juniperus brevifolia (Seub.) Antoine and Frangula
azorica Tutin, was recognized to play an essential role in making highly valuable fur-
niture. Unfortunately, five centuries of exploitation and the recent invasion by exotic
plant species in the Archipelago have led to a continuous reduction and fragmentation
of P. azoricas habitat on each island.
However, the assumptions presented in this study could be improved by further
investigation and experiments about the species biology (e.g. growth performance) and
technological features (e.g. response of the wood to polishing and varnishing). Other
applications taking into account its properties could then be explored, namely utiliza-
tion for cabinets, interior panelling, veneer, handles, toys, etc. Such novel uses could
boost a niche market and move the public awareness and the forest owners towards
restoring the ecosystems where P. azorica occurs in the wild.
Acknowledgements
The authors are indebted to Dr. Anna Rita Taddei of CIME (Centro Interdipartimentale di Microscopia
Elettronica), University of Tuscia, for the valuable support in providing SEM pictures of Picconia
azorica.
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