Comparative Anatomy of the Wood of Ab/Es P/Nsapo and Its Two Moroccan Varieties

IAWA Journal, Vol. 28 (3), 2007: 285-299

COMPARATIVE ANATOMY OF THE WOOD OF AB/ES P/NSAPO AND ITS TWO MOROCCAN VARIETIES

Luis Garcia Esteban, Paloma de Palacios, Antonio Guindeo and Francisco Garcia Fernandez Universidad Politecnica de Madrid, Escuela Tecnica Superior de Ingenieros de Montes, Departamento de Ingenieria Forestal, Ciitedra de Tecnologia de la Madera, Ciudad Universitaria, s/n, 28040 Madrid, Spain - Corresponding author: [email protected]

SUMMARY

This study describes the structure of the wood of Abies pinsapo from samples taken from its three natural distribution areas in the Iberian Pe ninsula (Sierra de Grazalema, Sierra de las Nieves and Sierra Bermeja) and compares them with the varieties from the north of Africa, Abies pinsapo var. marocana from the Talassemtane mountains and A. pinsapo var. tazaotana from the Tazaout mountains. All the samples were col lected in their regions of provenance. To put the results into perspective, a comparison was also made with the wood of Abies alba and A. nu midica. The wood of the Iberian A. pinsapo and of its two varieties from the Rif mountains in Morocco is anatomically similar, and there are no qualitative differences that enable the wood to be differentiated except for the presence of resin deposits in the tracheids adjacent to the rays in the samples from Grazalema. Quantitatively, for tracheid diameter and tracheid length there are statistically significantly differences (p<0.05) between those of Spanish provenance and the Moroccan varieties, but for tracheid pit diameter, largest diameter of cross-field pits and tall ray fre quency the samples from Sierra Bermeja have more in common with the African samples. Key words: Abies pinsapo, A. pinsapo var. marocana, A. pinsapo var. tazaotana, A. alba, A. numidica.

INTRODUCTION

Abies is one of the most common coniferous taxa in the Mediterranean Basin, with eight species, one subspecies and two varieties: A. alba Miller, A. borisii-regis Mattfeld, A. cephalonica Loudon, A. cilicica Antoine & Kotschy, A. nebrodensis (Lojac.) Mattei, A. nordmanniana (Steven) Spach,A. nordmanniana subsp. equi-trojani (Asch. & Sint. ex Boiss.) Coode & Cullen, A. numidica de Lannoy ex Carriere, A. pinsapo Boissier, A. pin sapo var. marocana (Trabut) Ceballos & Bolanos, A. pinsapo var. tazaotana (C6zar ex Huguet del Villar) Pourtet (Farjon 1998). Although some authors cite the presence of the pinsapo fir in the province of Granada (Liu 1971; Farjon 1990), it is really only found in the provinces ofüidiz and Malaga, in the western part of the Betica range, specifically in the area known as Serrania de

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Ronda. In the province of Malaga its geographical distribution is in two differentiated areas: on the one hand, the small area of some 40 ha in Los Reales de Sierra Bermeja, a single enc1ave on peridotites from 1200 to 1400 m, very near the Strait of Gibraltar; and on the other hand, the forests of Sierra de las Nieves, with some 3000 ha distrib uted over the municipalities of Ronda, Tolox and Yunquera, at altitudes ranging from 1000 to 1800 m, in this case on limestone substrates. There are also minor spots in the Alcor, Caparain, Real, Istan, Rio Verde and Gialda ranges. In the province of Cadiz the species occurs in the Sierra deI Pinar, in the municipality of Grazalema, in an area of approximately 500 ha on limestone substrates, at altitudes ranging from 1000 to 1650 m. There are also specimens and isolated groves in the western part of Monte Prieto, on the slopes of EI Monton and the northern slope of Zafalgar and of Los Pinos (Fig. 1) (Ceballos & Vicioso 1933; Ceballos & Ruiz de la Torre 1979).

Sevilla

I Ronda - 3 Yunquera Grazalema II! Cacl iz .. Tolox Malaga 2. Sierra Bermeja Estepona North Atlanti c Ocean Mediterranean Strait cf Gibraltar Sea

Tazaout , 5 Chefchaouen ~ Talassentane 4 0 40 km

Figure l. Principal forests of Abies pinsapo and its African varieties: 1: Sierra de Grazalema; 2: Sierra Bermeja; 3: Sierra de las Nieves; 4: Talassemtane; 5: Tazaout.

The so-called Rifian firs (from the Rif mountains) are located in the north of Mo rocco. Abies pinsapo var. marocana is distributed from 1450 to 1900 m following the southeast-northwest mountain chain from the western part of the Beni Selman region along the massifs of Tunian, Bab-Tizimando, Isilan and Tisuka as far as mount Magot (Chefchaouen). It is found in northern and eastern exposures, over an area of 2600 ha. Abies pinsapo var. tazaotana is located on mount Tazaout in the Beni Sey-yel region, northwest ofTalambot, over an area of 1500 ha at altitudes ranging from 1280 to 1800 m (Fig. 1 & 2) (Ceballos & Martfn Bolafios 1928; Sanchez Cozar 1946; Charco 1999).

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The wood anatomy of Abies pinsapo and its two varieties has previously been de scribed by several researchers. However, the studies were based on a limited number of samples, mostly of unknown provenance or in some cases belonging to branches, as their biometry shows. Krauss (1864) in Viguie et Gaussen (1928), included A. pinsapo in an identification key, distinguishing between A. cephalonica, A. sibirica, A.fraseri, A. alba, A. pinsapo, A. nordmanniana, A. cilicica andA. balsamea. Castellarnau (1880) made a very thorough description of A. pinsapo, although it was based on a single sample from the Serrania de Ronda. Saint-Laurent (1932) made another description of A. pinsapo var. marocana, without stating its provenance. Greguss (1955) describ ed A. pinsapo, similarly failing to mention its provenance. Peraza (1964) described A. pinsapo var. tazaotana from samples collected by the Spanish Forest Service in the Rif and proposed the separation of Abies alba from the group Abies pinsapo and A. pinsapo var. tazaotana, on the basis of the higher frequency of rays over 30 cells high in Abies alba. The present authors, Garda Esteban et al. (2002), previously de scribed two samples of A. pinsapo from specimens provided by the Malaga Forest Ser vice. While there are very few descriptions of A. pinsapo var. marocana and var. tazaotana, Saint-Laurent (1932) andPeraza (1964), there are many more in the case of A. pinsapo, although a divergence in data can be observed, particularly in the average ray height in number of cells, one of the characteristic features of the genus Abies: 3-30 (Kraus 1864 in Viguie etGaussen 1928), 1-30 (Castellarnau 1880),1-8 (Greguss 1955),2-30 (Peraza 1964),4-15 (Richter & Dallwitz 2000), 1-10 (Garda Esteban et al. 2002). Descriptions with such a low number of cells most likely correspond to samples taken from juvenile wood. The objectives of this study are to describe, for the first time, the anatomy of the wood of A. pinsapo using samples from the three natural areas in the Iberian Peninsula; to describe the two Moroccan varieties using samples from their two places of origin; to compare the anatomy of the wood of A. pinsapo and its two varieties; and to analyse whether the variation found constitutes a distinct pattern.

MATERIALS AND METHODS

The material used for this study was collected in natural forests of Abies pinsapo in both the Iberian Peninsula and the north of Africa, in Morocco. In Spain sampies were taken from the three natural areas: Sierra deI Pinar de Grazalema (API-5) in the province of Cadiz, Los Reales de Sierra Bermeja (AP6-1O) and La Nava de San Luis in Parahuta de la Sierra de las Nieves (APll-l5), in the province of Malaga. In Morocco the samples of A. pinsapo var. marocana were taken from the Talassemtane National Park (TLI-5), and of A. pinsapo var. tazaotana from Mount Tazaout (TZI-5). Five trees were cut in each zone, all adult specimens over 70 years old and repre sentative of the forest. The study included a total of 25 trees, 15 from Spain and 10 from Morocco. In order to locate the natural forests, the publications of Ceballos and Martin Bolafios (1930) were used for the province of Cadiz and of Ceballos and Vicioso

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Figure 2. Pinsapo fir forest in its ecological optimum in Tazaout. - Figure 3. Sapwood and heart wood similar (Grazalema).

(1933) for the province of Malaga. For the forests in Morocco the studies of Ceballos and Mart{n Bolanos (1928) and Sanchez C6zar (1946) were used. Abies alba sampies were also taken in Mount Martinier-La Mascarina in the munici pality of Forcayo EI Comato (AAI-5), in the province of Huesca, and sampies of A. numidica from the wood collection ofthe Royal Botanic Garden, Kew (K-Jw 18393) and the Nationaal Herbarium Nederland (UNI92) were also used for comparison with the pinsapo fir wood, as the Mediterranean fir group is a c10sed group and is very differentiated from the groups in Asia and North America (Kormutak et al. 2004). In addition, this group supports the hypothesis of hybridisation in the fir species (Conte & Cristofolini 2003). Microtome seetions were made using standard methods, and the descriptions were made in accordance with the IAWA Committee (2004). The sampies were observed by using light microscopy and scanning electron mi croscopy SEM mod. JEOL JSM-6380, both without staining and after staining with safranine and Sudan 4 in order to stain the resin (Jane 1970). The test pieces for SEM were prepared following the method described by Heady and Evans (2000). The tracheid length was measured following Ladell's indirect method (Ladell1959). The measurements of tangential tracheid diameter, tracheid length, diameter of tra cheid pits, height and frequency of rays and size of cross-field pits were made on the microtome seetions using the WinCell image analysis programme. The biometry was done on 5 preparations from each tree, in all cases on mature wood from a basal disc

Downloaded from Brill.com09/30/2021 01:45:23PM via free access Garcfa Esteban et ai. - Wood anatomy oi Abies pinsapo 289 between rings 70 and 100. On each preparation 35 random measurements were made (5 trees x 5 preparations x 35 measurements = 875 measurements/provenance). The number of pits per cross-field was measured on the five preparations from each tree, in ten different rays. The measuring of the height and frequency of the rays was done on all the rays contained in one square millimetre on the tangential seetion. An analysis of variance was made in order to determine the significance level in the variables of tangential tracheid diameter, tracheid length, ray height and frequency, number of rays per square millimetre, diameter of tracheid pits, largest diameter of the pits in the cross-fields and the number of pits in the cross-fields. The Lilliefors normality test and Bartlett's homogeneity of variance test were done, both for a significance level of95 % (p < 0.05). As in all cases at least one ofthe condi tions was not met, the ANOVAKruskal-Wallis test was used to analyse the samples. To determine statistically significant differences between provenances LSD tests were done using the data from the ANOVA Kruskal-Wallis test. The statistical calculations were done with the MATLAB Y.6.5 Release 13 programme, for a significance level of 95 %.

Anatomical description and biometry Cross section (Fig. 3-8). Sapwood and heartwood colour similar, although the heartwood colour is quite marked in the fresh wood, particularly in the Sierra Bermeja sampie, which has shades of red (Fig. 3). Fresh wood has a fetid odour. Growth ring boundaries distinct (Fig. 3). Abrupt transition from earlywood to latewood (Fig. 4), with the transition being more pronounced in the wood from Sierra Bermeja (Fig. 5) where the rings are narrower. The Moroccan varieties show a transition closer to that of Sierra Bermeja, although the number of cells in the latewood is greater: 8-15 in var. marocana and 6-10 in var. tazaotana. Tracheids rectangular in trans verse section. Tangential wall pits generally present in the tracheids near the growth ring boundary (Fig. 6a). Axial parenchyma present but sparse, generally in marginal position, in iso lated cells with high cellular content (Fig. 6b). Resin canals absent. Occasional presence of traumatic resin canals, surrounded by subsidiary cells with high cellular content and crystals (Fig. 7). Resin deposits in the tracheids adjacent to the rays only observed in the samples from Grazalema (Fig. 8). Tangential seetion (Fig. 9-12). Rays of over 30 cells lall. Rays almost exclusively uniseriate. Rays 2-seriate in part, constituting less than 10 % of the total (Fig. 11). Trans verse end walls ofaxial parenchyma cells nodular (Fig. 12). Radial seetion (Fig. 13-18). Cross-field pits taxodioid (Fig. 13). Trabecula common (Fig. 14). Tracheid pits in radial walls of earlywood predominantly biseriate, in oppo site arrangement (Fig. 15). Horizontal walls of ray parenchyma cells distinctly pitted. Indentures present (Fig. 16). Calcium oxalate crystals in ray parenchyma cells pris matic, located in both the marginal and interior rows of the rays, more abundantly in the former (Fig. 17:a). Irregularly shaped cells present in the marginal rows ofthe rays (Fig. 17: b & 18). Tori well defined, not scalloped, but extensions present (Fig. 19). Warty layer present (Fig. 20). All biometrie data are summarised in Table 1. (Text continued on page 294)

Downloaded from Brill.com09/30/2021 01:45:23PM via free access Table I. Biometry of species of the genus Abies from the western Mediterranean, including A. alba from the Pyrenees and A. numidica.

1 8 A.pinsapo A.pinsapo A.pinsapo A. alba A. numidica Feature var. tazaotana var. marocana (mean value ± SD and range) Grazalema Sierra Sierra de las Tazaout Talassemtane Pyrenees Collections Bermeja Nieves

Tangential tracheid diameter (11m) 46.9 ± 9.4 44.4 ± 9.8 44.9 ± 10.8 51.4 ± 7.3 62.0 ± 6.3 50.4 ± 7.6 38.9 ± 4.1 (23.4-76.0) (22.7-71.3) (25.3-73.4) (30.1-67.2) (42.5-76.7) (27.3-74.6) (28.7-44,6)

Diameter tracheid pits (11m) 18.5 ± 1.9 20.2 ± 1.7 18.6 ± 2.7 20.7 ± 2.6 21.2 ± 2.1 22.1 ± 2.3 16.2 ± 1.7 (12.7-24.6) (16.0-25.3) (7.5-28.6) (9.8-29.6) (10.5-26.0) (17.7-28.6) (12.0-19.5)

Tracheid length (11m) 2591 ± 793 3154 ± 861 2946 ± 934 3518 ± 976 3405 ± 1255 2813±721 2593 ±452 ( 1335-4625) (1500-7250) (1420-6167) (1550-7500) (1500-5670) (1536-4750) (1833-3900)

Most frequent average height in 8 4 5 7 7 5 9 number of cells (1-40) (1-36) (1-50) (1-30) (1-25) (1-48) (1-17)

Most frequent average height of 169.7 ± 24.6 95.3 ± 8.9 114.0 ± 12.2 133.7 ± 12.4 134.0 ± 15.9 158.9 ± 14.1 165.6 ± 12.6 rays (11m)

Number of rays per mm2 24.4 ±4.0 23.6 ± 2.9 23.8 ±4.6 22.9 ± 3.8 23.5 ± 3.2 30.6 ± 5.2 30.6 ± 3.8 ......

Downloaded fromBrill.com09/30/2021 01:45:23PM (18-32) (17-30) (13-32) (16-31) (18-31) (18-42) (27-35) ~ Largest diameter cross-field pits (11m) 7.5 ± 1.2 7.1 ± 1.2 6.6 ± 1.1 7.2± 1.1 7.2 ± 1.1 7.1 ± 1.0 7.5 ± 1.7 ;J> ..... (4.7-10.9) (3.6-11.3) (3.5-9.8) (3.4-10.7) (1.0-11.1) (4.14-9.6) (4.66-10.7) 0 8= Smallest diameter cross-field pits (11m) 4.4 ± 1.2 4.2 ± 1.1 3.8 ±0.9 3.0 ± 0.9 2.6 ± 0.9 3.3 ± 1.5 5.2± 1.4 ? (1.9-7.3) (1.3-7.5) (1.7-6.8) (0.9-6.0) (1.1-5.2) (1.12-6.8) (2.1-8.5) ~ ~ Number of pits per cross-field 1.6 ± 0.6 1.9 ± 0.6 1.9 ± 0.7 1.9 ± 0.6 1.8 ± 0.6 1.8 ± 0.6 1.4 ± 0.5 00 (1-4) (1-4) (1-4) (1-4) (1-3) (1-4) (1-2) ,--, ~ via freeaccess ~ N umber of rays / mm 2 > 30 cells 0.025 ± 0.041 0.002 ± 0.009 0.021 ± 0.047 0.004 ± 0.017 O.O±O.O 0.040 ± 0.054 O.O±O.O 0 0 -.J Garcfa Esteban et al. - Wood anatomy of Abies pinsapo 291

Figures 4-8. Features of Abies pinsapo. Cross section. - 4: Growth rings (Grazalema). - 5: Growth rings (Sierra Bermeja). - 6a: Tracheid pits in the tangential walls of the tracheids; 6b: Axial parenchyma cell (Sierra Bermeja). - 7: Traumatic resin canals (Grazalema). - 8: Resin plugs in tracheids (Grazalema). - Scale bars for 4 & 5 = 300 ~m; for 6, 7 & 8 = 100 ~m.

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Figures 9-12. Features of Abies pinsapo. Tangential seetion. - 9: Rays (Sierra Bermeja). - 10: Rays (Grazalema). - 11: Biseriate rays (Grazalema). - 12: Axial parenehyma with nodular transverse end walls (Sierra Bermeja). - Seale bars for 9 = 300 11m; for 10 & 11 = 100 11m; for 12 = 50 11m.

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Figures 13-18. Features of Abies pinsapo. Radial seetion. - 13: Taxodioid cross-field pits (Sierra Bermeja). - 14: Trabecula (Grazalema). - 15: Biseriate tracheid pits (Sierra Bermeja). - 16: Indentures (Sierra de las Nieves). - 17a: Crystals in ray parenchyma cells (Grazalema); 7b: Irregularly shaped cells. - 18: Irregularly shaped marginal cells (Sierra Bermeja). - Scale bars for 13,15, 16 & 17 =50 !-lm; for 14 & 18 = 100 !-lm.

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Figure 19 & 20. SEM. - 19: Very weakly scalloped tori. - 20: Warty layer.

DISCUSSION

The genus Abies shows very pronounced wood anatomical homogeneity (Jacquiot 1955; Peraza 1964), and the wood of its species cannot be differentiated anatomically (Schweingruber 1990). However, many authors have proposed keys based on very dif ferent criteria to differentiate the species of Abies (Krauss 1864 in Viguie et Gaussen 1928; Castellarnau 1880; Penhallow 1896; Saint-Laurent 1932; Wiesehuegel 1932; Record 1934; Phillips 1948; Brown etal. 1949; Greguss 1955; Kukachka 1960; Peraza 1964; Core et al. 1979). The Abies pinsapo wood from the three Spanish provenances and its two varieties from the Rif mountains are anatomically similar, and there are no qualitative differ ences to allow them to be differentiated from each other except for the presence of resin deposits in the tracheids adjacent to the rays in the sampies from Grazalema, similar to those described in Araucariaceae (Iones 1912; Patton 1927; Heady et al. 2002). The presence of resin deposits in this provenance may be related to the high resin content in the ray parenchyma, which is much higher than in any of the other provenances, and the subsequent overflow of the resin into the tracheid lumina through the cross-field pits (Heady et al. 2002; Garcfa Esteban et al. 2005). While resin deposits are common in Araucariaceae and some genera of Podocarpaceae and Cupressaceae, they are very rare in Pinaceae, e.g. Picea sitchensis (Kukachka 1960). Although Sargent (1902) used wood colour to differentiate A. nobilis and A. mag nifica from the rest of the American firs, the more reddish colour of the Sierra Bermeja provenance is probably influenced by the edaphic substrate made up of peridotites. Wiesehuegel (1932) used the early-Iatewood transition ofthe rings in theAmerican species as a diagnostic feature, although it should not be considered as such, as these variations are a function of climatic conditions (Core et al. 1979). In fact, Sierra Ber meja has the least favourable ecological conditions of all the sites where A. pinsapo grows. On the other hand, the ecological optimum in the Moroccan forests (Sanchez C6zar 1946) explains the significantly wider diameter of the tracheids than those of the Spanish provenances (Fig. 21: a). This behaviour is also seen when the mean tracheid length values are analysed (Fig. 22).

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As for the tracheid pit diameter and the largest diameter of the cross-field pits, the results show an affinity between the Sierra Bermeja provenance and the two Moroccan ones. The number of pits per cross-field is a homogeneous feature for all the provenances except in the case of the GrazaIema provenance (Fig. 21: b, c, d).

a b -e- -- 2 ...... 2

1l () <:: "<:: '"<:: 3 ...... '"<:: 3 -<>- > > "E! "E! e. e. 4 4 -<>- 5 -- 5 -<>- 40 45 50 55 60 65 18 18.5 19 19.5 20 20.5 21 21.5 tangential diameter 01 tracheids diameter 01 tracheid pits

e d --e- --<>-

2 2 --<>--

() () "<:: "<:: '"<:: 3 --e- '"<:: 3 --<>- > > "E! "E! e. e. 4 4 --<>--

5 ---e--- 5 --e-

6.4 6.6 6.8 7 7.2 7.4 7.6 7.8 1.4 1.5 1.6 1.7 1.8 1.9 2 largest diameter 01 cross-field pits number 01 pits per cross-field

2 2

() () 3 "<:: "<:: <:: <:: '" 3 '" 4 > "E! " e. ee. 5 4 6

21.5 22 22.5 23 23.5 24 24.5 25 25.5 -0.01 0 0.01 0.02 0.03 0.04 0.05 number 01 rays per mm 2 number 01 rays per mm 2 > 30 cells Figure 21. LSD statistical analysis. Provenance of species: 1: Abies pinsapo (Sierra de Grazalema); 2: A. pinsapo (Sierra Berrneja); 3: A. pinsapo (Sierra de las Nieves); 4: A. pinsapo var. tazaotana (Tazaout); 5: A. pinsapo var. marocana (Talassemtane); 6: A. alba (Pyrenees); 7: A. numidica (collections). - a: Tangential diameter of tracheids. - b: Diameter of tracheid pits. - c: Largest diameter of cross-field pits. - d: Number of pits per cross-field. - e: Number of rays per mm2. - f: Number of rays per mm2 > 30 cells.

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Sierra de Grazalema o

Sierra Bermeja 2 o

Sierra de las Nieves 3 o ~ r:::: ro Tazaout r:::: 4 o e~ Cl. Talassentane 5 o

(A. alba) Pirineos 6 o

(A. numidica) collections 7 o

2.6 2.8 3 3.2 3.4 3.6 3.8 tracheid length

Figure 22. LSD statistical analysis. Tracheid length.

Ray size parameters are usually considered to be good indicators for Abies (Wiese huegel1932; Greguss 1955) and have been used by some authors for characterising its species (Castellamau 1880; Wiesehuegel 1932; Phillips 1948; Greguss 1955; Peraza 1964). In A. pinsapo the number of rays per square millimetre does not show significant differences (Fig. 21:e), but the frequency ofrays over 30 cells high does. This value is used by some authors as a threshold for differentiating A. alba from the other Abies species (Jacquiot 1955; Peraza 1964). Analysis ofthe maximum ray height allows two clusters to be distinguished: on the one hand the pinsapo firs from Grazalema and Sierra de las Nieves, and on the other hand the pinsapo firs from Sierra Bermeja and the two Moroccan varieties. The values place the woods from Grazalema and Sierra de las Nieves closer to A. alba than to the Rifian subspecies (Table 1). The rays in the wood from Sierra Bermeja are intermediate in height between those from Grazalema-Sierra de las Nieves and the Tazaout-Talassemtane. The Iberian pinsapo firs attain higher values of ray height than the Rifian firs. Analysis of the number of rays with over 30 cells shows that the Grazalema-Sierra de las Nieves provenance is closer to A. alba, but yet statistically different from each other, while the Sierra Bermeja provenance once again forms a cluster with the Mo roccan firs and even with A. numidica (Fig. 21: f). The Moroccan provenances do not have very high rays in spite of the fact that they are located in the ecological optimums

Downloaded from Brill.com09/30/2021 01:45:23PM via free access Garcfa Esteban et al. - Wood anatomy of Abies pinsapo 297 of the pinsapo fir forests, and therefore a relation between maximum ray height and ecological conditions seems to be ruled out. The wood structure of the Rifian firs is statistically different from the structure of A. numidica in their maximum ray height, although there are no significant differences in terms of the number of rays of over 30 cells between the Rifian and the Aigerian firs. Resin canals were only observed in the wood from Grazalema, but their presence should not be considered to have diagnostic value either, as the canals in Abies appear in response to injuries (Anderson 1897; Jeffrey 1905; Rhoades 1923; Chamberlain 1935; Jane 1970). Calcium oxalate crystals were present in the ray parenchyma in all the provenances, both in the marginal and submarginal rows. Their presence cannot, in this case, be used with diagnostic value to differentiate provenances, although it can be used to characterise the pinsapo fir woods, as in the case ofmost ofthe Abies species (Greguss 1955; Jane 1970; Core et al. 1979; IAWA Committee 2004). Ray tracheids were not observed in any of the provenances, and although some authors maintain that some species of Abies can contain ray tracheids (Penhallow 1907 in A. balsamea; Thompson 1912 in A. homolepis and A. veitchii; Jane 1970 in Abies spp.), their presence is related to injuries (Jeffrey 1917; Chamberlain 1935; Phillips 1948). The presence of irregularly shaped cells, generally in the marginal position, is usual in pinsapo fir wood, particularly in the wood from Grazalema and Las Nieves. Thompson (1912) referred to these cells as degenerate cells and considered them a characteristic element ofthe genus Abies. Later on, Chrysler (1915) also cited them in Cedrus. The warty layer on the tracheid walls is present in all the provenances. No quantita tive or qualitative differences are observed between the provenances. With the exception oftwo biometric variables (tracheid diameter and tracheid length), it can be stated that, in general, the wood structure of the two Moroccan varieties is similar to the firs of the Iberian Peninsula, particularly the Sierra Bermeja provenance. This provenance could perhaps present an intermediate phyletic stage between the provenances from Grazalema and Sierra de las Nieves on the one hand, and the Mo roccan firs on the other, in the same way as A. nebrodensis does between A. alba and A. numidica (Duc ci et al. 1999; Parducci et al. 2001; Conte & Cristofolini 2003).

ACKNOWLEDGEMENTS

We would like to express our gratitude to the following: The Forestry Engineers Miguel A. Martfn Casillas and Jose L6pez Quintanilla for assistance in col lecting the sampies of A. pinsapo in the Parque Natural de la Sierra de Las Nieves and in the Paraje Natural de Sierra Bermeja in the province ofMalaga, and the Parque Natural de la Sierra de Grazalema in the province of Cadiz. The public organisations of the Royaume du Maroc Haut Commissariat aux Eaux et Forets et ala lutte contre la desertification, Direction Regionale des Eaux et Forets du Rif aTetouan, Service Provincial des Eaux et Forets du Chefchaouen. In particular to Amrani Mohamed, Ingenieur et Chef des Eaux et Forets du Rif a Tetouan and Assim ElHoussain, Chef du Distriet du Chefchaouen, who provided assistance in the collection of samples of A. pinsapo in the forests of Talassemtane and Tazaout in Morocco.

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The Huesca Forest Service for providing assistance in the collection of samp1es of A. alba in mount Martinier-La Mascarina in the municipality ofForcayo EI Comato. The Director of the Royal Botanic Gardens, Kew, and of the Nationaal Herbarium Nederland, for providing sampIes of A. numidica. Marc Philippe and Pieter Baas for critical reading of the manuscript and valuable suggestions.

REFERENCES

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IAWA Committee. 2004. IAWA List of microscopic features for softwood identification. IAWA J. 25: 1-70. Jacquiot, e. 1955. Atlas d'anatomie des bois des coniteres. Centre Technique du Bois, Paris. Jane, EW. 1970. The structure of wood. Ed. 2. A & C Black, London. Jeffrey, E.C. 1905. The comparative anatomy and phylogeny of the Coniferales. Part. 2. The Abietineae. Memoirs Boston Soc. Nat. Hist. Vol. 6. Jeffrey, E.e. 1917. The anatomy of woody plants. The University of Chicago Press, Chicago. Jones, W.S. 1912. The structure of the timbers of some comrnon genera of coniferous trees. Quart. J. For. 1-23. Kormutak, A., B. Vookova, B. Ziegenhagen, H.Y. Kwon & Y.P. Hong. 2004. Chloroplast DNA variation in some representatives of the Asian, NorthAmerican and Mediterranean firs (Abies spp). Silvae Genetica 53: 99-104. Kukachka, B.E 1960. Identification of coniferous woods. Tappi 43: 887-896. Ladell, J. T. 1959. A new method of measuring tracheid length. Forestry 32: 124-125. Liu, T. 1971. A monograph of the genus Abies. Publ. Dept. Forestry, College of Agriculture, Taiwan University, Taipei. Parducci, L., A. E. Szmidt, M. M. Ribeiro & A. D. Drouzas. 2001. Taxonomic position and origin of the endemic Sicilian fir Abies nebrodensis (Lojac.) Mattei based on allozyme analysis. Forest Genetics 8: 119-127. Patton, R.T. 1927. Anatomy of Australian coniferous timbers. Proc. Roy. Soc. Vict. n.s. 40: 1-16. Penhallow, D.P. 1896. Taxaceae and Coniferae. Trans. Roy. Soc. Canada 2: 33-56. Penhallow, D.P. 1907. A manual of the North American gymnosperms. Ginn Boston, Boston. Peraza, C. 1964. Estudio de las maderas de coniferas espafiolas y de la zona norte de Marruecos. IFIE, Madrid. PhilIips, E. W. J. 1948. Identification of softwoods by their microscopic structure. Forest Products Research Bulletin No. 22. Department of Scientific and Industrial Research, London. Record, S.J. 1934. Identification ofthe timbers oftemperate NorthAmerica. J. Wiley & Sons, New York. Rhoades, A. S. 1923. The formation and pathological anatomy of frost rings in conifers injured by late frosts. U.S. Dept. Agric. Bull. 1131. Richter, H.G. & M.J. Dallwitz. 2000. Comrnercial timbers: descriptions, illustrations, identifica tion, and information retrieval. http://biodiversity.uno.edu/delta!. Saint-Laurent, J. deo 1932. Etudes sur les caracteres anatomiques du bois et du liber secondaire dans quelques essences forestieres du Maroc demierement recoltees par M. le Dr. R. Maire (avec les planches XIII-XV). Bull. Station de Recherches Forestieres du Nord de I' Afrique. 2: 49-60. Sanchez C6zar, S. 1946. EI Abies deI Tazaout. Revista de la Real Academia de Ciencias de Madrid. Vol. XL: 449-468. Sargent, C.S. 1902. The silva of North America: 105-193. Houghton, Mifflin & Co. New York. Schweingruber, EH. 1990. Anatomy of European woods. Swiss Federal Institute of Forestry Research, Stuttgart. Thompson, w.P. 1912. Ray tracheids in Abies. Bot. Gaz. 53: 331-338. Viguie, T. & H. Gaussen. 1928. Revision du genre Abies. Travaux du Laboratoire Forestier de Toulouse, Tome II, Art. I. Wiesehuegel, E.G. 1932. Diagnostic characteristics of the xylem of the North American Abies. Bot. Gaz. 43: 55-70.

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IAWA List of Microscopic Features for Hardwood Identification byan IAWA Committee; edited by E.A. Wheeler, P. Baas and P. E. Ga on

Publi hed for the International A ociation ofWood Anatomi ts First printing 1989, Third printing 1999. 112 pp., 190 half-tone illustrations Paperback - Price: USO or EUR 25.00 (excl. handling and postage)

Thi IAWA publication sets the standard for wood anatomical terminology and for the use of micro copic characters in wood identification. Altogether 163 wood anatomical de criptors are defined, annotated and illustrated with light micrographs. In addition, 57 non-anatomical descriptors are included and discussed. Thi IAWA Li t offers guidance to everyone who wants to interpret de criptive informa tion from the va t wood anatomicalliterature or who needs to include de criptive wood anatomical information in original publications. Thanks to the concise definitions and high-quality illustration , the IAWA Hardwood List i al 0 very suitable for teaching purpo es.

IAWA List of Microscopic Features for Softwood Identification

by an lAWA Committee; edited by H.G. Richter, O. Grosser, I. Heinz and P.E. Gasson

Publi hed for the International Association of Wood Anatomi t 2004.70 pp., 76 half-tone illustrations [Reprinted from IAWA Journal 25: 1-70] Paperback - Price: USO 30.00 or EUR 25.00 (excl. handling and postage)

Thi IAWA publication enumerate ,define, illustrates and annotates 124 characters that are relevant forthe microscopic de cription and identification ofthe secondary xylem of trees and hrub belonging to the Pinop ida (conifers) and Ginkgoop ida. It i a long awaited companion of the highly successful IAWA List of Micro copic Features for Hardwood ldentification, published in 1989, and now a much-used manual for teaching hardwood identification all over the world. The beautiful micrographs by Oietger Gro ser (Munich) illu trate each feature and make the IAWA Softwood Li st a user-friendly intro duction to the recognition of all ubtle micro copic softwood features. The numerical coding adopted makes the li st also uitable to model databases and inter active, computer-assisted identification keys on. Above all this Li twill be a u eful tool for training students in the arts of softwood identification and de cription.

To be ordered /rom: lAWA Office elo ationaal Herbarium ederland I Leiden braneh PO Box 9514 - 2300 RA Leiden - The etherlands Fax: +31 71 5273511 - E-mail: [email protected]

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