Comparative wood anatomy of Maytenus in Northwestern Argentina (South America)

Anatomía comparada del leño de Maytenus en el Noroeste de Argentina (Sudamérica)

Ana María Giménez¹, Juana Graciela Moglia¹, M.E. Figueroa¹, J.A. Díaz Zírpolo¹ and Federico Calatayu¹

¹ LAM (Laboratorio de Anatomía de Madera) Facultad de Ciencias Forestales, Universidad Nacional de Santiago del Estero (UNSE), Argentina. amig@unse.edu.ar

ABSTRACT

This paper is a comparative wood anatomy study of four species of the genus Maytenus living in Northwest Argentina: Maytenus vitis-idaea, M. viscifolia, M. spinosa and M. cuezzoi. The specimens were collected in Santiago del Estero and Salta, Argentina and wood samples are safeguarded in the collection of the LAM (Laboratory of Wood Anatomy), Faculty of Forestry of Santiago del Estero University (UNSE), Argentina. The terminology used followed the IAWA List of Microscopic Features for Hardwood Identification. The diagnostic features of wood anatomical characters were evaluated by employing statistical methods such as Cluster Analysis (CA) and Principal Component Analysis (PCA). PCA showed vessel diameter, fibre wall, and ray width to be significant variables. CA showed M. cuezzoi and M. viscifolia to have the highest affinity.

Key words: Celastraceae, Chaco, Hardwoods, Santiago del Estero.

RESUMEN

El presente trabajo es un estudio de anatomía comparada de madera de cuatro especies del género Maytenus del Noroeste Argentino: Maytenus vitis-idaea, M. viscifolia, M. spinosa y M. cuezzoi. Las muestras fueron recolectadas en Santiago del Estero y Salta, Argentina y se salvaguardan en la colección del LAM (Laboratorio de Anatomía de Madera), Facultad de Ciencias Forestales de la Universidad de Santiago del Estero (UNSE), Argentina. Se empleó la terminología de IAWA (Lista de caracteres anatómicos del xilema de angiospermas). Los caracteres anatómicos de madera fueron evaluados mediante métodos estadísticos tales como análisis de conglomerados (AC) y el Análisis de Componentes Principales (PCA). El PCA mostró como variables significativas el diámetro de vasos, el espesor de pared de las fibras y el ancho de radios. El CA mostró que M. cuezzoi y M. viscifolia tienen alta afinidad específica.

Palabras clave: Celastraceae, Chaco, angiospermas, Santiago del Estero.

INTRODUCTION

Celastraceae is a widely distributed family in the world, comprising 57 genera with 370 species in both hemispheres. There are several indigenous representatives in Argentina. They are usually small trees or shrubs (Dimitri, 1972). Celastraceae stem xylem anatomy has been described by Record and Hess (1943), Mennega (1997), Archer and Van Wyk (1993a). Mennega (1997) analyzed the anatomy of the subfamily Hippocrateoideae, which is distinguished by thin, long rays. Record and Hess (1943) studied the Genera: Celastrus, Zinowiewia, Elaeodendron, Goupia. Archer and Van Wyk (1993a; 1993b) made a comparative anatomical study of mature wood and bark of the subfamily Cassinoideae, comprising mainly southern African species of Cassine, Pleurostylia (17) and the three monotypic genera, Allocassine, Hartogiella and Maurocenia.

Metcalfe and Chalk (1983) recorded the presence of numerous small vessels, solitary, in radial multiples of 2-3, ring and semi-ring porous, occasionally with helical thickenings, simple perforation plate, alternate intervascular pits, very small, members of vessels of medium length to moderately long, are typical features. Very weakly defined vestures were seen in Celastraceae (Archer and Van Wyk, 1993a).

Rays are described in some genera to be exclusively uniseriate and homogeneous, in other heterogeneous multiseriate (3-4).

Maytenus is a genus of temperate and warm regions of South Asia (Yemen, Malaysia and Thailand), Africa (Canary Islands, northwest and northeast Ethiopia, and South Africa), and America from Mexico to Tierra del Fuego. It grows in a variety of climates, from tropical (77 sp. in Brazil) to subpolar (Hurrel and Bazzano, 2003).

In Argentina the Genus Maytenus comprises about eleven native species. Three are endemic shrubs and small trees (Boelke, 1992), distributed in the subtropical wet forest of Misiones, Yungas, Chaco semiarid forest and Patagonian Andean temperate forests (Zuloaga and Morrone, 1999).

Maytenus boaria (Maitén) is the most important tree of Patagonian Andean forests (Lourteig and Odonell, 1955). The species is used as a natural dye and its branches are an important source of foliage for sheep in the steppe (Tortorelli, 2009). M. spinosa, M. viscifolia and M. vitis-idaea are species of the semiarid forest of the Chaco region. They are indicative of saline soils and semi-arid areas. Giménez and Hernández (2008) found M. spinosa in 50% of the areas studied in Santiago del Estero, Province, M. vitis-idaea in 36% of them, and M. viscifolia at only one site (Sierras de Guasayan). Their uses include dye, fodder, food and to a lesser extent, as timber depending on the size of their trunks (Giménez et al., 2010). M. cuezzoi is a threatened endemic species (Cat. 5: restricted distribution plant and sparse populations with restricted distribution and sparse populations).

For the genus Maytenus, the outstanding features of the wood are: diffuse porous, small vessels, very numerous (20-40), solitary and in radial multiples, and sometimes helical thickenings. Axial parenchyma is very variable in type and number, commonly scattered or absent, diffuse and multiseriate or thin bands (Metcalfe and Chalk, 1983). Vessels with helical thickenings were cited M. boaria (Tortorelli, 2009).

Rays are heterogeneous, frequently with 2-4 vertical rows of cells, 1 to 3 seriate, less than 12/mm (Metcalfe and Chalk, 1983); M. boaria (-9 ray/mm) with 3-4 seriates (Tortorelli, 2009).

Fibres with distinctly bordered pits were cited in M. boaria (Tortorelli, 2009), M. acuminata (Metcalfe and Chalk, 1950), M. micrantha (Detienne and Jacquet, 1983).

Fibre-tracheids are elements of transition between tracheids and fibres. They are characterized by the presence of bordered pits, generally located in the radial walls, present in Apocynaceae, Celastraceae (some Genera) Myrtaceae, Sapotaceae, Zygophyllaceae among others. Carlquist (1988) described the imperforate tracheal elements of Catha, Elaeodendron, Celastrus and Maytenus as septate libriform fibres, septate fibre-tracheids and vasicentric tracheids.

Fibre-tracheid has been cited as an adaptive strategy in species of arid and semiarid areas, such as Aspidosperma quebracho-blanco (Apocynaceae) (Moglia et al., 2009); Monttea aphylla (Scrophulariaceae) (Giménez et al., 1998); Tabernaemontana catharinensis (Apocynaceae) (Giménez, 2004). Carlquist and Hoekman (1985) cite the presence of fibre-tracheids as typical element of the arid flora of California.

Several African species have been described anatomically: M. acuminata (Metcalfe and Chalk, 1950); M. micrantha (Detienne and Jacquet, 1983), M. senegalensis (Neumann et al., 2000). Perforated ray cells (PRC) are present in the stem xylem and in the roots of M. brasiliensis and M. obtusifolia (Joffily et al., 2007).

Sokal and Rohlf (1981) have shown the importance of multivariate statistical techniques in numerical taxonomy. Using multivariate techniques, Robertse et al. (1980) solved problems in variation of wood anatomical characters of South African Acacia. Somaratne and Heart (2001) established relationships among species of the genus Calophyllum, and Wickremasinghe and Heart (2006) did the same for Diospyros.

MATERIALS AND METHODS

The aim of the study is to analyze the comparative wood anatomy of four species which are little known of northwestern Argentina: Maytenus vitis-idaea. Griseb.; M. viscifolia Griseb.; M. spinosa (Griseb.) Lourteig & O'Donell and M. cuezzoi Legname.

The characteristics of the species studied are:

Maytenus viscifolia Griseb. (chasqui-yuyo) is an endemic unarmed shrub or small tree, 3 m - 7 m. tall, with a trunk up to 0,3 m in diameter, with persistent foliage. Its geographic distribution in Argentina is the semiarid Chaco region, in the following provinces: Salta, Tucumán, Santiago del Estero, Catamarca, La Rioja, Córdoba, San Juan (Gimenez and Moglia, 2003).

Maytenus spinosa (Griseb.) Lourteig & O'Donell (abriboca) is an endemic shrub with spiny branches and leathery leaves. It is native to Argentina and Uruguay (Digillio and Legname, 1966). The bark produces a pink brown dye (Stramiglioli, 2007).

Maytenus cuezzoi Legname (Legname, 1973; 1982) is an endemic unarmed tree, 3 m - 7 m tall and 0,25 m. in diameter at the base, with entirely glabrous pale green leaves, smooth bark, and red flowers. Its geographic distribution is the Yungas, in the temperate humid upper floor of the montane forest, between 1500 m and 2000 m altitude of the provinces of Salta and Jujuy. (Zuloaga and Morrone, 1999). The species has been found in the Los Toldos valley, Salta province, forming secondary forests dominated by Ilex argentina, M. cuezzoi and Roupala sp.; these associates indicate degraded forests (Brown and Grau, 2000). Samples were collected by the author of the species, Prof. P. Legname.

We sampled the four species of Maytenus from different habitats of northwestern Argentina. M. vitis-idaea and. M. spinosa were collected in Tala Atun, San Martín department, Santiago del Estero (latitude S 28° 44' 44'', longitude W 63° 11' 34''; altitude: 130 m, mean annual precipitation (MAP): 470 mm; mean annual temperature (MAT): 20 °C (Saavedra, 2004). M. viscifolia was collected in Guampacha, Sierras de Guasayán, (latitude S: 28° 15' 44'', longitude W 63° 34' 55'', altitude: 479 m, MAP: 563 mm; MAT: 19 °C) of Santiago del Estero province. M. cuezzoi was collected in the montane forest of Los Toldos, Salta province (latitude S: 22° 24', longitude W 64° 43', altitude: 1700 m, MAP: 1349 mm, MAT: 19 °C).

Fresh wood samples were collected along with herbarium vouchers. The material was collected from five wild-growing individuals of each species. 10 cm thick disk was cut from each tree at 0.3 m height. The samples are deposited in Laboratory of Wood Anatomy (LAM), Faculty of Forestry of Santiago del Estero University (UNSE).

Mature wood samples were fixed in 70% alcohol. Transverse, radial and tangential sections (12 μ-18 μ thick) were cut using a sliding microtome. The sections were doubly-stained with 1% fuchsin and astra blue (Roeser, 1972) and mounted permanently by Entellan. Wood material was macerated following Jeffrey's method (Johansen, 1940).

Terminology and determination of quantitative features follow the recommendations of the IAWA Committee (1989) and Tortorelli (2009). For vessel diameters, vessel element lengths, fibre lengths and ray height, 25 measurements were taken from each specimen and averaged.

The influence of environmental conditions on characteristics (vessel diameter and vessel number) associated with water conduction was evaluated through the vulnerability index. The vulnerability index (VI) was calculated to estimate susceptibility to damage during water conduction of wood, as proposed by (Carlquist, 1988): VI= Vd / Vmm; where: Vd =vessel diameter and Vmm = vessel per square mm. Numerous, narrow vessels give the plant protection against cavitation, especially in stress environments, while fewer, wider vessels are more susceptible to cavitation.

Multivariate statistical analyses were applied in the present study to trace the possible relationships between anatomical and ecological features. A preliminary approaches to treatment of the subject about which of the quantitative variables are associated with each of the species analyzed follows. Were used seven quantitative variables: vessel per square/ mm (Vmm); tangential diameter of vessel (Vd); vessel element length (Vl); fibre length (Fl); fibre wall thickness (Fw); ray height (Rh); ray width (Rw).

Non parametric analysis of variance (ANAVA) for repeated measures (Cody and Smith, 1991) and the Kruskal Wallis test (α = 0,05) was performed for 7 variables. Professional Program InfoStat was used for statistical analysis (INFOSTAT, 2008).The photomicrographs were taken with Zeiss Axiostar microscope and Sony video camera ExwaveHAD. Small blocks cut to produce transverse, radial and tangential surfaces were used for scanning electron microscopy (SEM).

RESULTS

Anatomical descriptions

Maytenus vitis-idaea (Fig. 1-8)

Growth rings distinct, marked by thick-walled fibres. Wood diffuse-porous, vessels solitary (75%) and in radial multiples of 2-4; occasionally with some clusters, 68 (50-85) per mm2; 28 (20-40) μm in diameter. Vessel element length 77 (45-125) μm. Perforations simple, intervessel pits alternate, small, 4 μm - 6 μm in diameter. Vessel-ray pits with distinct borders, similar to intervessel pits in size and shape. Fibre non-septate, thick- to very thick-walled, 165 (112-262).μm in length, with distinctly bordered pits; pit frequency on radial and tangential walls more or less equal, pits minute (2 μm - 5 μm). Axial parenchyma diffuse-in-aggregates and scanty paratracheal; 1-2 cells per parenchyma strand. Rays heterocellular with 2-4 rows of upright and/or square marginal cells, 13 (7-16) per mm, mostly 1 to 2 (occasionally 3) cells wide, 116 (150- 450) μm in height, not storied. Prismatic crystals occasionally present in upright and/or square ray cells and in short chains in axial parenchyma cells; one crystal per cell or chamber.

Maytenus viscifolia (Fig. 9-16) (Fig. 17-20)

Growth rings distinct, marked by thick-walled fibres. Wood diffuse porous. Vessels solitary (30%) and in radial multiples of 2-6; occasionally with some clusters, 84 (42-117). per mm². Vessels 36 (30-40) μm in diameter. Vessel element length 113 (65-172) μm. Perforations simple. Intervessel pits vestured, minute (3 μm - 5 μm), polygonal. Vessel-ray pits rounded with much reduced borders, to apparently simple.

Fibre 227 (135-282) μm in length, with thick to very thick cell walls. Fibre pits mostly conspicuously bordered, 2 μm - 5 μm in diameter, frequency on radial and tangential walls approximately equal. Apotracheal axial parenchyma diffuse-in-aggregates and scanty paratracheal vasicentric.

Maytenus cuezzoi (Fig. 21-25) (Fig. 26-30)

Growth rings distinct, marked by thick-walled fibres. Wood diffuse-porous, rarely appearing semi-ring-porous. Vessels with helical thickenings, in radial multiples of 2-5 (60%), solitary (25%), and occasionally clusters, 99 (62-172) vessels per mm². Vessels 32 (20-20) μm in diameter. Vessel element length 131 (77-182) μm. Perforations simple. Intervessel pits alternate, small (4 μm - 6 μm), vestured. Vessel-ray pits with distinct borders, similar to intervessel pits in size and shape.

Fibre 230 (162-325) μm in length with thick to very thick walls. Fibre pits mostly conspicuously bordered, 2-5 μm in diameter, frequency on radial and tangential walls approximatelyequal.

Apotracheal axial parenchyma diffuse-in-aggregates; scanty paratracheal, vasicentric, and unilateral.

Rays heterocellular with 2-4 rows of upright and/or square marginal cells, 13 (9-18) per mm, 1 to 3 cells wide, 460 (240-960) μm in height.

Prismatic crystals occasionally present in upright and/or square ray cells and in short chains in axial parenchyma cells; one crystal per cell or chamber.

Maytenus spinosa (Fig. 31-36)

Growth rings distinct, marked by thick-walled fibres. Wood diffuse-porous, rarely appearing semi-ring-porous. Vessels predominantly solitary (84%), in radial multiples of 2, and occasionally some clusters; 136 (87-185) per mm2. Vessels 20 (10-30) μm in diameter. Vessel element length 84 (40-117) μm; tails frequent at both ends. Perforations simple. Intervessel pits alternate, small, 4 μm - 6 μm in diameter. Vessel-ray pits with distinct borders, similar to intervessel pits in size and shape.

Imperforate tracheary elements consist of non-septate fibre with pits mostly conspicuously bordered, 153 (102-197) μm in length, 2-4 μm in diameter; density on radial and tangential walls approximately equal. Cell walls thick to very thick.

Rays heterocellular with 2-4 rows of upright and/or square marginal cells, 20 (14-24) per mm, 1 to 3 cells wide, 330 (130-690) μm in height.

Prismatic crystals occasionally present in upright and/or square ray cells and in short chains in axial parenchyma cells, one crystal per cell or chamber.

The most important anatomical features are summarized in Table 1

From PCA, factor 1 expresses 58% of the variability, 33% factor 2 and 0,06 factor 3 (Table 2). Factor 1 is considered practically significant, as explaining an important amount of the variability in the data. The variables that load significantly in Factor 1 are: vessel diameter (Vd), fibre length (Fl), and ray width (Rw); on Factor 2: vessel length (Vl), ray height (Rh), and fibre wall (Fw) load significantly. The species separate into different quadrants (Fig. 37).

Species appear as points and the variables as vectors. Species that appear in the same direction that a variable, may have high values for that variable and low value are plotted in the opposite direction. The correlations between variables can be interpreted through the angles between the vectors. Angles of 90 degrees indicates no correlation, angles (<90°) indicate a positive correlation and angles of > 90° negative correlation, angles close to 180° would show high negative correlation between the variables.

The histogram of vessel groupings shows that M. spinosa and M. vitis-idaea have more than 70% solitary vessels. M. cuezzoi and M. viscifolia have solitary, multiple and cluster vessels. M. vitis-idaea and M. viscifolia present a predominance of biseriate rays.

DISCUSSION

Our observations of Maytenus are in accordance with the studies of Metcalfe and Chalk (1983), Tortorelli (2009); Joffily et al., 2007.

M. spinosa and M. vitis-idaea have wide prevalence of solitary vessels (75%). This character is mentioned for M. acuminata (Metcalfe and Chalk, 1983), M. micrantha (Detienne and Jacquet, 1983), and M. senegalensis (Neumann et al., 2000). Vessel multiples of 2-3 is present in M. cuezzoi. M viscifolia exhibits vessels of all types including multiples of 2-5.

M. boaria had vessels solitary and in multiples of 2-6 (Tortorelli, 2009).

The tangential diameter of vessel varies significantly with the species. The vessels are extremely small, in M. spinosa (mean: 19,6 μm); and small in M. viscifolia. (35,6 μm).

The vessel frequency show highly significant differences between species; they are extremely numerous in M. spinosa; numerous in M. cuezzoi, M. viscifolia and M. vitis-idaea.

Vessels with helical thickenings were cited M. boaria (Tortorelli, 2009) and in M. cuezzoi, and vestured pits were observed in M. cuezzoi y M. viscifolia.

The vulnerability index predicts which species can live in arid (Carlquist, 1988). Xylem efficiency depends on the diameter and frequency of vessels in a given area.

VI index in the four sp is low (0,15/0,45), indicating a high specialization in water transport. Wood samples of M. spinosa and M. vitis-idaea were collected in Tala Atun (MAP: 470 mm and dry season of 5 months). M. spinosa (VI 0,15), presents a strategy to ensure water conduction with small, short and very numerous vessels.

M. vitis-idaea (IV 0,43) has very small short and moderately numerous vessels.

M. viscifolia living in Chaco serrano (MAP: 563 mm and 5-month dry season), adopts the strategy of increasing vessel diameter (VI: 0,45). The adaptive strategy of M. cuezzoi is the increase in vessel length, tangential diameter and vessels frequency, can be interpreted by the greater availability of water in the plant (1340 mm). The differences are statistically significant between the sp. for variables vessel diameter and vessel frequency.

The axial parenchyma in the four species show is diffuse in aggregates.

M. vitis-idaea and M. viscifolia have predominantly biseriate rays; M. spinosa and M. cuezzoi, uniseriate rays. The fibres have distinctly bordered pits, are very short, especially in M. spinosa, M.vitis-idaea and M. viscifolia have thick walls (over 60% of the outer diameter of the fibres).

In the Celastraceae, the presence of PRCs was only known to data for Cassine (Archer and Van Wyk 1993a). However, the authors did not consider the presence of this cell type diagnostic for the genus due to its rare occurrence and it was only found in M. viscifolia.

In arid regions, wood must support high negative pressures. In these regions, the morphology of vessels has adaptive value. There is a reduction of vessel diameter and length, increase in vessel frequency and degree of clustering, and presence of qualitative anatomical features, such as simple perforations. These adaptations increase safety against embolisms (Lindorf, 1994). Baas and Carlquist (1985) emphasize the presence of the following characters in xerophytes and halophytes: small vessels (29-53 μ), high vessel frequency (92-150), semi-ring porous, and presence of fibre-tracheids. All of these features (except porosity) are present in the species studied. The anatomical structure of the woods of this study is characteristic of species of arid zones, such as the dry Chaco region in which Maytenus is common. The smaller diameter and length of the vessels, and their high frequency, are some of the most notable features of the conductive elements. They are considered signs of adaptation to extreme aridity and tend to increase the safety, given the limited amount of water available (Lindorf, 1994; Carlquist, 1988; Moglia and Giménez, 1998; Giménez, 1993; Roth and Giménez, 1997; Roth and Giménez, 2006).

The sp M. viscifolia and M. cuezzoi not show statistical differences for 5 of the 7 considered variables (Kruskal Wallis). This is confirmed by the cluster analysis where both sp. form a group.

CONCLUSIONS

The species studied show the anatomical features typical of genus Maytenus.

The anatomical structure of the woods of this study is characteristic of species of arid zones.

The smaller diameter, shorter vessels and high frequency, are the most notable features of the conductive elements, considered signs of adaptation to extreme aridity

Quantitative variables that best explain the anatomical differences are: vessel diameter, fibre wall thickness, rays width. The distinctive anatomical features are the presence of PRC only in M. viscifolia, intervascular vestured pits in M. cuezzoi and M. viscifolia and helical thickenings in M. cuezzoi.

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Nota

This manuscript must be cited as: Giménez, A.M., J.G. Moglia, M.E. Figueroa, J.A. Díaz-Zírpolo and F. Calatayu. 2014. Comparative wood anatomy of Maytenus in Northwestern Argentina (South America). Madera y Bosques 20(2):95-110.