A revision of chestnut-leaved oak (Quercus castaneifolia C. A. Mey.; Fagaceae) in Hyrcanian Forests of Iran

Panahi, P.; Jamzad, Z.; Pourmajidian, M.R.; Fallah, A.; Pourhashemi, M.

A revision of chestnut-leaved oak (Quercus castaneifolia C. A. Mey.; Fagaceae) in Hyrcanian Forests of Iran

Authors

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: Panahi@rifr-ac.ir

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.

REFERENCES
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Bačić, T. (1981) Investigations of stomata of three oak species with light and scanning electron microscope. Acta Bot. Croat. 40: 85-90.
Baranski, M. (1975) An analysis of variation within white oak (Quercus alba L.). North Carol. Agri. Expe. St. Tech. Bull. 236, 176 p.
Blue, M.P. Jensen, R.J. (1988) Positional and seasonal variation in oak (Quercus; Fagaceae) leaf morphology. Amer. J. Bot. 75: 939-947.
Boissier, E. (1879) Flora Orientalis. Geneva et Basileae, 4: 1162-1174.
Burger, W.C. (1974) The species concept in Quercus. Taxon, 24: 45-50.
Bussotti, F. Grossoni, P. (1997) European and Mediterranean oaks (Quercus L.; Fagaceae): SEM characterization of the micromorphology of the abaxial leaf surface. Bot. J. Linn. Soc. 124: 183-199.
Camus, A. (1936-1938) Les chênes. Monographie du genre Quercus. Encyc. Econ. Sylvic. Lechevalier, Paris. 1: 557.
Colombo, P.M. Lorenzoni, F.C. and Grigoletto, F. (1983) Pollen grain morphology supports the taxonomical discrimination of Mediterranean oaks (Quercus, Fagaceae). Pl. Syst. Evol. 141: 273-284.
Djavanchir Khoie, K. (1967) Les chênes de L`Iran. PhD Dissertation, Universite de Montpellier, Faculte des Sciences, 221 p.
Erdtman, G. (1986) Pollen morphology and plant taxonomy, Angiosperms (an introduction to palynology). Leiden E. J. Brill, 553 p.
Faegri, K. and Iversen, J. (1964) Textbook of pollen analysis. Munksgaard, Copenhagen, 237 p.
Forest, Range and Watershed Organization of Iran (2003) Kimiaye sabz. F. R. W. O Publication, 368 p.
Halbritter, H. Weber, M. Zetter, R. FroschRadivo, A. Buchner, R. and Hesse, M. (2006) PalDat- Illustrated handbook on pollen terminology. Univ. of Vienna. 61 p.
Hardin, J.W. (1976) Terminology and classification of Quercus trichomes. J. Elisha Mitchell Sci. Soc. 92: 151-161.
Hardin, J.W. (1979a) Atlas of foliar surface features in woody plants, I. vestiture and trichome types of eastern north American Quercus. Bull. Torrey Bot. Club.106: 313-325.
Hardin, J.W. (1979b) Patterns of variation in foliar trichomes of eastern North American Quercus. Amer. J. Bot. 66: 576-585.
Harley, M.M. (1992) The potential value of pollen morphology as an additional taxonomic character in subtribe Ociminae (Ocimeae: Nepetoideae: Labiatae). In: Harley, R.M., Reynolds, T. (Eds.), Advances in Labiatae Science, Royal Botanic Gardens. Kew, Richmond, Surrey, UK, pp. 125-138.
Hedge, I.C. Yaltirik, F. (1982) Fagaceae. In: Davis, P.H. Edmondson, J.R. Mill, R.R. Tan, K. (eds.), Flora of Turkey and the east Aegean islands, Edinburg Univ. Press, 7: 657-683.
Jarvis, D.I. Leopold, E.B. and Liu, Y. (1992) Distinguishing the pollen of deciduous oaks, evergreen oaks, and certain Rosaceous species of southwestern Sichuan province, China. Rev. Palaeobot. Palynol. 75: 259-271.
Jones, J.H. (1986) Evolution of the Fagaceae: the implications of foliar features. Ann. Missouri Bot. Gard. 73: 228-275.
Kedves, M. Pàrdutz, À. and Varga, B. (2002) Experimental investigation on the pollen grains of Quercus robur L. Taiwania, 47: 43-53.
Koch, K. Barthlott, W. (2009) Superhydrophobic and superhydrophilic plant surfaces: an inspiration for biomimetic materials. Philos. T. Roy. Soc. A. 367: 1487-1509.
Lawrence, G.H.M. (1951) Taxonomy of vascular plants. Mac Millan Co., New York.
Llamas, F. Perez-Morales, C. Acedo, C. and Penas Merino, A. (1995) Foliar trichomes of the evergreen and semideciduous species of the genus Quercus (Fagaceae) in the Iberian Peninsula. Bot. J. Linn. Soc. 117: 47-57.
Liu, Y.S. Zetter, R. Ferguson, D.K. and Mohr, B.A.R. (2007) Discriminating fossil evergreen and deciduous Quercus pollen: A case study from the Miocene of eastern China. Rev. Palaeobot. Palynol. 145: 289-303.
Menitsky, G.L. (1971) Fagaceae. In: Rechinger, K.H. (ed.), Flora Iranica. Akademische Druck, Univ. of Verlagsanstalt Graze, Austria, 77: 1-20.
Meyer, C.A. (1831) Verzeichn. der pflanzen. Casp. Meer, St. Perersbourg. 44, p. 241.
Monoszon, M.K.H. (1962) Variations of the morphological characters of pollen from certain species of oak. Dokl. Akad. Nauk, S.S.S.R. 140, 165-168.
Muller, C.H. (1952) Ecological control of hybridization in Quercus: A factor in the mechanism of evolution. Evolution, 6: 147-161.
Nikolić, N.P. Merkulov, L.S. Krstić, B.D. and Orlović, S.S. (2003) A comparative analysis of stomata and leaf trichome characteristics in Quercus robur L. genotypes. Proc. Nat. Sci, Matica Srpska Novi. Sad. 105: 51-59.
Parsa, A. (1949) Flore de L`IRAN: Fagaceae. Imprinerie Mazaheri Tehran, 4: 1319- 1332.
Penas Merino, A. Llamas, F. Perez Morales, C. and Acedo, C. (1994) Aportaciones al Conocimiento del Genero Quercus en la Cordillera Cantabrica. I, Tricomas Foliares de las Especies Caducifolias. Lagascalia. 17: 311-324.
Punt, W. Blackmore, S. Nilsson, S. and Le Thomas, A., 1994. Glossary of pollen and spore terminology. LPP Contributions Series, vol. 1. LPP Foundation, Utrecht, The Netherlands.
Sabeti, H. (1994) Forests, trees and shrubs of Iran, Univ. of Yazd Publication, Iran, 876 p.
Safou, O. and Saint-Martin, M. (1989) Leaf trichomes of some perimediterranean Quercus species. Bull. Soc. Bot. France. 136: 291-304. S
careli-Santos, C. Herrera-Arroyo, M.L. Sanchez-Mondragon, M.L. GonzalezRodriguez, A. Bacon, J. and Oyama, K. (2007) Comparative analysis of micromorphological characters in two distantly related Mexican oaks, Q. conzattii and Q. eduardii (Fagaceae) and their hybrids. Brittonia, 59: 37-48.
Schwarz, O. (1935) Einige neue Eichen des Mediterrangebiets und Vorderasiens, Notizblatt des Königl. botanischen Gartens und Museums zu Berlin, Bd. 12, Nr. 114, pp. 461-469.
Smit, A. (1973) A scanning electron microscopical study of the pollen morphology in the genus Quercus. Acta Bot. Neerl. 22: 655-665.
Soepadmo, E. (1972) Fagaceae. Fl. Males, I, 7: 265-403.
Solomon, A.M. (1983a) Pollen morphology and plant taxonomy of white oaks in eastern North America. Amer. J. Bot. 70: 481-494.
Solomon, A.M. (1983b) Pollen morphology and plant taxonomy of red oaks in eastern North America. Amer. J. Bot. 70: 495-507.
Thamson, P.M. and Mohlenbrock, R.H. (1979) Foliar trichomes of Quercus subgenus Quercus in the eastern united states. J. Arnold Arbor. 60: 350-366.
Uzunova, K. and Palamarev, E. (1985) The foliar epidermis studies of Fagaceae Dumort. from the Balkan Peninsula. II. Quercus [Subgenera Sclerophyllodrys Schwarz and Cerris (Spach) Oersted]. Fitologia. 29: 3-20. U
zunova, K. and Palamarev, E. (1992a) Study of the leaf epidermis of the Balkan representatives of the Fagaceae Dumort. III. Quercus L. [Subgenus Quercus, sect. Roburoides (Schwarz) Schwarz and Dascia Kotschy]. Fitologia. 42: 22-48.
Uzunova, K. and Palamarev, E. (1992b) The Foliar epidermis studies of Fagaceae Dumort. from the Balkan Peninsula. IV. Quercus L. (Subgenus Quercus, sect. Robur Reichend.). Fitologia. 43: 3-30.
Uzunova, K. and Palamarev, E. (1993) An investigation of the leaf epidermis of the European (non-Balkan) species of the genus Quercus. Fitologia. 45: 3-15.
Uzunova, K. Palamarev, E. and Ehrendorfer, F. (1997) Anatomical changes and evolutionary trends in the foliar epidermis of extant and fossil Euro-Mediterranean oaks (Fagaceae). Pl. Syst. Evol. 204: 141-159.
Wang, P.L. and Chang, K.T. (1988) Pollen morphology of Cyclobalanopsis and its relation to Quercus. Acta Phytotax. Sin. 26: 282-289.
Zheng, Z.H. Wang, P.L. and Pu, F.D. (1999) A comparative study on pollen exine ultrastructure of Nothofagus and the other genera of Fagaceae. Acta Phytotax. Sin. 37: 253-258.

Keywords

References

Aas, G. (1998) Morphologische und ökologische Variation mitteleuropäischer Quercus Arten: Ein Beitrag zum Verständnis der Biodiversität. Libri Botanici, Bd. 19. IHW-Verlag Eching bei München, 221 pp.

Bačić, T. (1981) Investigations of stomata of three oak species with light and scanning electron microscope. Acta Bot. Croat. 40: 85-90.

Baranski, M. (1975) An analysis of variation within white oak (Quercus alba L.). North Carol. Agri. Expe. St. Tech. Bull. 236, 176 pp. Blue, M.P. Jensen, R.J. (1988) Positional and seasonal variation in oak (Quercus; Fagaceae) leaf morphology. Amer. J. Bot. 75: 939-947.

Boissier, E. (1879) Flora Orientalis. Geneva et Basileae, 4: 1162-1174. Burger, W.C. (1974) The species concept in Quercus. Taxon, 24: 45-50.

Bussotti, F. Grossoni, P. (1997) European and Mediterranean oaks (Quercus L.; Fagaceae): SEM characterization of the micromorphology of the abaxial leaf surface. Bot. J. Linn. Soc. 124: 183-199.

Camus, A. (1936-1938) Les chênes. Monographie du genre Quercus. Encyc. Econ. Sylvic. Lechevalier, Paris. 1: 557.

Colombo, P.M. Lorenzoni, F.C. and Grigoletto, F. (1983) Pollen grain morphology supports the taxonomical discrimination of Mediterranean oaks (Quercus, Fagaceae). Pl. Syst. Evol. 141: 273-284.

Djavanchir Khoie, K. (1967) Les chênes de L`Iran. Ph. D Thesis, Universite de Montpellier, Faculte des Sciences. 221 pp.

Erdtman, G. (1986) Pollen morphology and plant taxonomy, Angiosperms (an introduction to palynology). Leiden E. J. Brill, 553 pp.

Faegri, K. and Iversen, J. (1964) Textbook of pollen analysis. Munksgaard, Copenhagen, 237 pp. Forest, Range and Watershed Organization of Iran (2003) Kimiaye sabz.

F. R. W. O Publication, 368 pp. Halbritter, H. Weber, M. Zetter, R. Frosch-Radivo, A. Buchner, R. and Hesse, M. (2006) PalDat- Illustrated handbook on pollen terminology. Univ. of Vienna. 61 pp.

Hardin, J.W. (1976) Terminology and classification of Quercus trichomes. J. Elisha Mitchell Sci. Soc. 92: 151-161.

Hardin, J.W. (1979a) Atlas of foliar surface features in woody plants, I. vestiture and trichome types of eastern north American Quercus. Bull. Torrey Bot. Club.106: 313-325.

Hardin, J.W. (1979b) Patterns of variation in foliar trichomes of eastern North American Quercus. Amer. J. Bot. 66: 576-585.

Harley, M.M. (1992) The potential value of pollen morphology as an additional taxonomic character in subtribe Panahi et al., 157Ociminae (Ocimeae: Nepetoideae: Labiatae). In: Harley, R.M., Reynolds, T. (Eds.), Advances in Labiatae Science, Royal Botanic Gardens. Kew, Richmond, Surrey, UK, pp. 125-138.

Hedge, I.C. Yaltirik, F. (1982) Fagaceae. In: Davis, P.H. Edmondson, J.R. Mill, R.R. Tan, K. (eds.), Flora of Turkey and the east Aegean islands, Edinburg Univ. Press, 7: 657-683.

Jarvis, D.I. Leopold, E.B. and Liu, Y. (1992) Distinguishing the pollen of deciduous oaks, evergreen oaks, and certain Rosaceous species of southwestern Sichuan province, China. Rev. Palaeobot. Palynol. 75: 259-271.

Jones, J.H. (1986) Evolution of the Fagaceae: the implications of foliar features. Ann. Missouri Bot. Gard. 73: 228-275.

Kedves, M. Pàrdutz, À. and Varga, B. (2002) Experimental investigation on the pollen grains of Quercus robur L. Taiwania, 47(1): 43-53.

Koch, K. Barthlott, W. (2009) Superhydrophobic and superhydrophilic plant surfaces: an inspiration for biomimetic materials. Philos. T. Roy. Soc. A. 367: 1487-1509.

Lawrence, G.H.M. (1951) Taxonomy of vascular plants. Mac Millan Co., New York. Llamas, F. Perez-Morales, C. Acedo, C. and Penas Merino, A. (1995) Foliar trichomes of the evergreen and semi-deciduous species of the genus Quercus(Fagaceae) in the Iberian Peninsula. Bot. J. Linn. Soc. 117: 47-57.

Liu, Y.S. Zetter, R. Ferguson, D.K. and Mohr, B.A.R. (2007) Discriminating fossil evergreen and deciduous Quercus pollen: A case study from the Miocene of eastern China. Rev. Palaeobot. Palynol. 145: 289-303.

Menitsky, G.L. (1971) Fagaceae. In: Rechinger, K.H. (ed.), Flora Iranica. Akademische Druck, Univ. of Verlagsanstalt Graze, Austria, 77: 1-20. Meyer, C.A. (1831) Verzeichn. der pflanzen. Casp. Meer, St. Perersbourg. 44, p 241.

Monoszon, M.K.H. (1962) Variations of the morphological characters of pollen from certain species of oak. Dokl. Akad. Nauk, S.S.S.R. 140, 165-168.

Muller, C.H. (1952) Ecological control of hybridization in Quercus: A factor in the mechanism of evolution. Evolution, 6: 147-161.

Nikolić, N.P. Merkulov, L.S. Krstić, B.D. and Orlović, S.S. (2003) A comparative analysis of stomata and leaf trichome characteristics in Quercus robur L. genotypes. Proc. Nat. Sci, Matica Srpska Novi. Sad. 105: 51-59.

Parsa, A. (1949) Flore de L`IRAN: Fagaceae. Imprinerie Mazaheri Tehran, 4: 1319-1332.

Penas Merino, A. Llamas, F. Perez Morales, C. and Acedo, C. (1994) Aportaciones al Conocimiento del Genero Quercus en la Cordillera Cantabrica. I, Tricomas Foliares de las Especies Caducifolias. Lagascalia. 17: 311-324.

Punt, W. Blackmore, S. Nilsson, S. and Le Thomas, A., 1994. Glossary of pollen and spore terminology. LPP Contributions Series, vol. 1. LPP Foundation, Utrecht, The Netherlands. Sabeti, H. (1994) Forests, trees and shrubs of Iran, Univ. of Yazd Publication, Iran. 876 pp.

Safou, O. and Saint-Martin, M. (1989) Leaf trichomes of some perimediterranean Quercus species. Bull. Soc. Bot. France. 136: 291-304.

Scareli-Santos, C. Herrera-Arroyo, M.L. Sanchez-Mondragon, M.L. Gonzalez-Rodriguez, A. Bacon, J. and Oyama, K. (2007) Comparative analysis of micromorphological characters in two distantly related Mexican oaks, Q. conzattii and Q. eduardii (Fagaceae) and their hybrids. Brittonia, 59: 37-48.

Schwarz, O. (1935) Einige neue Eichen des Mediterrangebiets und Vorderasiens, Notizblatt des Königl. botanischen Gartens und Museums zu Berlin, Bd. 12, Nr. 114, pp. 461-469.

Smit, A. (1973) A scanning electron microscopical study of the pollen morphology in the genus Quercus. Acta Bot. Neerl. 22: 655-665.

Soepadmo, E. (1972) Fagaceae. Fl. Males, I, 7: 265-403. Solomon, A.M. (1983a) Pollen morphology and plant taxonomy of white oaks in eastern North America. Amer. J. Bot. 70: 481-494.

Solomon, A.M. (1983b) Pollen morphology and plant taxonomy of red oaks in eastern North America. Amer. J. Bot. 70: 495-507. 158 A revision of chestnut-leaved oak

Thamson, P.M. and Mohlenbrock, R.H. (1979) Foliar trichomes of Quercussubgenus Quercus in the eastern united states. J. Arnold Arbor. 60: 350-366.

Uzunova, K. and Palamarev, E. (1985) The foliar epidermis studies of FagaceaeDumort. from the Balkan Peninsula. II. Quercus [Subgenera SclerophyllodrysSchwarz and Cerris (Spach) Oersted]. Fitologia. 29: 3-20.

Uzunova, K. and Palamarev, E. (1992a) Study of the leaf epidermis of the Balkan representatives of the FagaceaeDumort. III. Quercus L. [Subgenus Quercus, sect. Roburoides (Schwarz) Schwarz and Dascia Kotschy]. Fitologia. 42: 22-48.

Uzunova, K. and Palamarev, E. (1992b) The Foliar epidermis studies of Fagaceae Dumort. from the Balkan Peninsula. IV. Quercus L. (Subgenus Quercus, sect. Robur Reichend.). Fitologia. 43: 3-30.

Uzunova, K. and Palamarev, E. (1993) An investigation of the leaf epidermis of the European (non-Balkan) species of the genus Quercus. Fitologia. 45: 3-15.

Uzunova, K. Palamarev, E. and Ehrendorfer, F. (1997) Anatomical changes and evolutionary trends in the foliar epidermis of extant and fossil Euro-Mediterranean oaks (Fagaceae). Pl. Syst. Evol. 204: 141-159.

Wang, P.L. and Chang, K.T. (1988) Pollen morphology of Cyclobalanopsis and its relation to Quercus. Acta Phytotax. Sin. 26: 282-289.

Zheng, Z.H. Wang, P.L. and Pu, F.D. (1999) A comparative study on pollen exine ultrastructure of Nothofagus and the other genera of Fagaceae. Acta Phytotax. Sin. 37: 253-258.