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A Revision of Chestnut-Leaved Oak (Quercus Castaneifolia C. A. Mey.; Fagaceae) in Hyrcanian Forests of Iran

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A Revision of Chestnut-Leaved Oak (Quercus Castaneifolia C. A. Mey.; Fagaceae) in Hyrcanian Forests of Iran Caspian J. Env. Sci. 2011, Vol. 9 No.2 pp. 145~158 ©Copyright by The University of Guilan, Printed in I.R. Iran CJES [Research] Caspian Journal of Environmental Sciences A revision of chestnut-leaved oak (Quercus castaneifolia C. A. Mey.; Fagaceae) in Hyrcanian Forests of Iran Z. Jamzad 1, M. R. Pourmajidian 2, A. Fallah 2, M. Pourhashemi 3 ,٭P. Panahi 1 1- Botany Research Division, Research Institute of Forests and Rangelands of Iran, P.O. Box 13185-116, Tehran, Iran 2- Sari Agricultural Sciences and Natural Resources University, Sari, Iran 3- Forest Research Division, Research Institute of Forests and Rangelands of Iran, Tehran, Iran * Corresponding author’s E-mail: [email protected] ABSTRACT Chestnut-leaved oak (Quercus castaneifolia C. A. Mey) is one of the most important native oaks of Iran distributed in the Hyrcanian Forests. The pure and mixed stands of it cover about 6.5% of these forests. The species represents morphological variations which have resulted in the description of several infraspecific taxa by different authors. Eight taxa were recognized as subspecies and varieties of Q. castaneifolia. In this survey, a set of quantitative and qualitative micro-morphological characteristics such as type of trichome, number and length of trichome rays, type of epicuticular waxes, type and shape of stomata, shape and sculptural features of pollen exine were studied using light microscopy and scanning electron microscopy. These characters were analyzed by the discriminant analysis method and combined with the macromorphological characters to designate the variation within the species and to evaluate the existing infraspecific taxa. Based on our results, the studied taxa are defined into four infraspecific taxa as follows: Q. castaneifolia subsp. castaneifolia var. castaneifolia, Q. castaneifolia subsp. castaneifolia var. minuta, Q. castaneifolia subsp. aitchisoniana, Q. castaneifolia subsp. undulate. The studied characteristics showed to be diagnostic for identification of the infraspecific taxa. Keywords: Hyrcanian Forests, Quercus castaneifolia, Pollen, Trichome, SEM. INTRODUCTION community gradually transforms into a Quercus is the largest genus in the family Zelkovo-Quercetum community. Quercus Fagaceae with about 300 (Lawrence 1951) - castaneifolia is found in composition with 600 species (Soepadmo 1972). This genus other woody species on the upper altitudes. includes evergreen and deciduous shrubs Pure and mixed stands of Q. castaneifolia and trees extending from cold latitudes to cover about 6.5% of the Hyrcanian Forests tropical Asia and Americas. It is the most (Forest, Range & Watershed Management common genus of Fagaceae in forests of Iran of Iran, 2003). (Sabeti 1994). Different species of oaks are This species shows morphological distributed in vast areas (about 6 million variations in leaf and acorn characteristics. ha) of Zagros, Arasbaran and Hyrcanian The delimitation of Q. castaneifolia, number Forests. The genus shows a remarkable of its subspecies and varieties varies morphological variation in these forests. according to different authors. At first, Quercus castaneifolia is one of the most Meyer (1831) described Q. castaneifolia C. A. important species of Iran`s native oaks, Mey. from the Caucasus and Hyrcanian distributed in the Hyrcanian Forests. As a Forests of Iran. Then Schwarz (1935) main species of the Hyrcanian forests, it studied the herbarium specimens collected forms two important forest communities in by Bornmüller from Bandar-Gaz forests in this zone: Querco-Buxetum in the Caspian Golestan Province, and Rasht forests in coast plains and Querco-Carpinetum in the Guilan province. He described a new lowlands (up to 700m a.s.l). With drier species namely Q. sintenisiana O. Schwarz, climate in the northeast of Iran, the later based on trichome characteristics, Three Online version is available on http://research.guilan.ac.ir/cjes 146 A revision of chestnut-leaved oak years later, Camus (1936-1938) reported for taxonomic purposes (Monoszon 1962; two subspecies including: Q. castaneifolia C. Smit 1973; Hardin 1976, 1979a, 1979b; A. Mey. subsp. eucastaneifolia A. Camus Thamson & Mohlenbrock 1979; Bačić 1981; and Q. castaneifolia C. A. Mey. sub sp. Colombo et al. 1983; Solomon 1983a, 1983b; aitchisoniana A. Camus in his monograph of Uzunova & Palamarev 1985, 1992a, 1992b, world's oaks. The most complete study of 1993; Jones 1986; Wang & Chang 1988; Quercus in Iran was done by Djavanchir Safou & Saint-Martin 1989; Jarvis et al. Khoie (1967) based on leaf and acorn 1992; Penas Merino et al. 1994; Llamas et al. morphology. He classified the Iranian oaks 1995; Uzunova et al. 1997; Zheng et al. 1999; into two subgenera; Quercus (lobed-leaved Kedves et al. 2002; Nikolić et al. 2003; species) and Complanate Djav.-Khoie Scareli-Santos et al. 2007). (dentate-leaved species), based on leaf In this research, we focused on the foliar venation. The subgenus Complanate was epidermis structure and pollen further divided into two sections; micromorphology of Q. castaneifoli as well Oligandrae Djav.-Khoie and Polyandrae as, its subspecies and varieties, introduced Djav.-Khoie based on number of stamens. by Djavanchir Khoie, to evaluate the According to this classification Q. taxonomic significance of these castaneifolia belongs to section Oligandrae. characteristics for the identification of the Djavanchir Khoie treated Q. sintenisiana O. infraspecific taxa. A complete survey on Schwarz and Q. castaneifolia C. A. Mey. habitats of these taxa, their distribution subsp. eucastaneifolia A. Camus as patterns, populations and ecological synonyms of the type species but he conditions were also carried out. accepted Q. castaneifolia C. A. Mey. subsp. aitchisoniana A. Camus as a separate taxon Materials and Methods and introduced 6 new taxa from the The majority of specimens used in this Hyrcanian Forests as follows: study were obtained during several field - Q. castaneifolia C. A. Mey. subsp. trips to areas according their geographic castaneifolia var. ellipsoidalis Djav.-Khoie ranges in Hyrcanian Forests of the country. - Q. castaneifolia C. A. Mey. subsp. At least 10 trees were studied for each castaneifolia var. minuta Djav.-Khoie taxon. Furthermore other previously - Q. castaneifolia C. A. Mey. subsp. incurvata collected specimens of the Herbarium of Djav.-Khoie Research Institute of Forests and - Q. castaneifolia C. A. Mey. subsp. Rangelands of Iran (TARI) and the subrotundata Djav.-Khoie Herbarium of Natural Resources Faculty, - Q. castaneifolia C. A. Mey. subsp. University of Tehran (NRF) were sampled. triangularis Djav.-Khoie Materials and collected data of some - Q. castaneifolia C. A. Mey. subsp. undulata individual plant samples of all studied taxa Djav.-Khoie are listed in table 1. The voucher specimens Menitsky in Flora Iranica (1971), are deposited in TARI. categorized Iran`s native oaks into Fully matured anthers and leaves were subgenus: Quercus, and two sections: collected in spring and summer. Leaf Quercus and Cerris. He placed dentate- samples were collected from open-grown leaved species in section Cerris and lobed- canopy branches that showed limited leaved species in section Quercus. morphological variations, resulting from According to his classification, Q. environmental factors such as exposure and castaneifolia was considered in section directions (Baranski 1975; Blue & Jensen 1988). The micromorphological features Cerris, subsec. Cerris and all infraspecific and quantitative characters of pollen grains taxa recognized by Djavanchir Khoie and were studied by Hitachi (S-4160) scanning Camus were treated as its synonyms. He electron microscope at an accelerating recognized only Q. castaneifolia C. A. Mey. voltage of 15 KV and photographed by subsp. castaneifolia from Iran. Olympus camera using a ×100 eyepiece. Leaf epidermal structures and pollen Pollen grains were prepared by the micromorphology have been often used for standard acetolysis method and then they classification and identification of Quercus were mounted on glycerin jelly and sealed species and based on the previous with paraffin wax for LM (Harley 1992). researches, these characteristics are helpful Measurements were taken from 50 grains of Panahi et al., 147 each examined taxon and the mean, range gold palladium for 5 minutes in a Humer ΙΙ and standard deviation were calculated for Sputtering Device. All leaf materials were polar axis (P) and equatorial diameter (E). prepared without pretreatment or critical The ratio of polar axis to equatorial point drying. Quantitative characters were diameter (P/E) was provided as an index of calculated from 50 representative trichomes pollen shape (Faegri & Iversen 1964). The and stomata per examined taxon. The exine thickness of pollen grains was trichome ray length was measured from the measured as well. For SEM observation, the point of divergence of the ray and at the pollen grains were mounted on aluminum base of the trichome. Terminology used for stubs, air dried at room temperature and trichome types is based on Hardin (1976, coated with gold. In the case of leaves, 1979a). Identification of wax layer types is epidermal structures such as trichome carried out according to Koch & Barthlott types, number and length of the trichome (2009). Terminology used for pollen shapes rays, dimension and location of stomata and features are based on Erdtman (1986), and type of epicuticular waxes were Punt et al. (1994) and Halbritter et al. studied by SEM. The leaf measurements (2006). Five quantitative
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