Abstract
Pollen and seed micro-morphological features of six species of Polygala distributed in Iran were examined using light (LM) and scanning electron microscope (SEM). The results show a great pollen morphological variation, which provides helpful data for species delimitation in Polygala. The pollen grains are monad, radially symmetrical, isopolar, polyzonocolporate, medium to large, oblate-spheroidal to prolate-spheroidal in shape. We observed four types of surface sculpture (psilate, psilate to granulate, rugulate-granulate and rugulate) and four types of apocolpial area (psilate with small lumina, reticulate with large lumina, micro-perforate and coarsely regulate and gemmate or granulate with lumina). Moreover, the importance of seed micro-morphological traits in species delimitation have demonstrated based on elaiosome appendages size, number of lobes and their sculpture, seed surface sculpture and trichome type.
References
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Arroyo-Cosultchi, G., Terrazas, T., Arias, S. & Arreola-Nava, H.J. (2006) The systematic significance of seed morphology in Stenocereus (Cactaceae). Taxon 55 (4): 983–992. https://doi.org/10.2307/25065693
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Bojnanský, V. & Fargašová, A. (2007) Atlas of seeds and fruits of Central and East-European flora: The Carpathian Mountains region. Springer Science & Business Media.
Bonilla-Barbosa, J., Novelo, A., Orozco, Y.H. & Márquez-Guzmán, J. (2000) Comparative seed morphology of Mexican Nymphaea species. Aquatic Botany 68 (3): 189–204. https://doi.org/10.1016/S0304-3770(00)00125-X
Bornmüller, J.F.N. & Gauba, E. (1942) Repertorium Specierum Novarum Regni Vegetabilis. Centralblatt für Sammlung und Veroffentlichung von Einzeldiagnosen neuer Pflanzen, 51. [Edited by Friedrich Fedde]. Berlin, 33 pp.
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Cosson, E.S. (1874) Bulletin de la Société Botanique de France, 20. Paris, 240 pp. https://doi.org/10.1080/00378941.1873.10825518
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Cullen, J. (1965) Polygala L. In: Davis, P.H. (Ed.) Flora of Turkey and the East Aegean Islands 1. Edinburgh University Press, Edinburgh, pp. 533–539.
de Candolle, A.P. (1824) Prodromus systematics, 1. Naturalis Regni vegetabilis, 326 pp.
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Erdtman, G. (1986) Pollen morphology and plant taxonomy: angiosperms (Vol. 1). Hafner Publishing Company, New York.
Eriksen, B. (1993) Phylogeny of the Polygalaceae and its taxonomic implications. Plant Systematics and Evolution 186: 33–55. https://doi.org/10.1007/BF00937712
Eriksen, B. & Persson, C. (2007) Polygalaceae. In Flowering Plants–Eudicots. Springer, Berlin, Heidelberg, pp. 345–363. https://doi.org/10.1007/978-3-540-32219-1_41
Eriksson, O., Friis, E.M. & Löfgren, P. (2000) Seed size, fruit size, and dispersal systems in angiosperms from the Early Cretaceous to the Late Tertiary. The American Naturalist 156 (1): 47–58. https://doi.org/10.1086/303367
Fagundez, J. & Izco, J. (2011) Seed morphology and anatomy of the Mediterranean pentamerous species of Erica (Ericaceae). Turkish Journal of Botany 35 (6): 643–651. https://doi.org/10.3906/bot-1006-22
Fischer, F.E.L., Meyer, C.A. & Trautvettler, E.R. (1838) Index quartum seminum, quae Hortus Botanicus Imperialis Petropolitanus promutua commutatione offert. Accedunt animadversiones botanicae nonnullae. St. Petersburg, 42 pp.
Forest, F., Chase, M.W., Persson, C., Crane, P.R. & Hawkins, J.A. (2007) The role of biotic and abiotic factors in evolution of ant dispersal in the milkwort family (Polygalaceae). Evolution: International Journal of Organic Evolution 61 (7): 1675–1694. https://doi.org/10.1111/j.1558-5646.2007.00138.x
Fritzsche, J. (1832) Beiträge zur Kenntniss des Pollen (Vol. 1). Stellin and Elbing, Nicolai, Berlin.
Furness, S.H. & Stafford, P.J. (1995) Polygalaceae. The Northwest European Pollen Flora. Review of Palaeobotany and Palynology 88 (1–4): 61–82.https://doi.org/10.1016/0034-6667(95)00019-T
Ghimire, B., Jeong, M.J., Suh, G.U., Heo, K. & Lee, C.H. (2018) Seed morphology and seed coat anatomy of Fraxinus, Ligustrum and Syringa (Oleeae: Oleaceae) and its systematic implications. Nordic Journal of Botany 36 (10): e01866.https://doi.org/10.1111/njb.01866
Heubl, G.R. (1984) Systematische Untersuchungen an mitteleuropaischen Polygala Arten. Mitteilungen der Botanischen Staatssammlung, Munchen.
Isaacs, M.J.H., Weitz, F.M. & Johnson, C.T. (1993) Seed-coat characteristics of selected southern African species of Polygala L.(Polygalaceae). South African Journal of Botany 59 (6): 592–596. https://doi.org/10.1016/S0254-6299(16)30675-5
Jalilian, N. (2005) Flora of Iran, No. 49: Polygalaceae. Agriculture Institute of forests and rangelands press. [In Persian]
Kerrigan, R.A. (2012) A treatment for Polygala of northern Australia. Australian Systematic Botany 25 (2): 83–137. https://doi.org/10.1071/SB08032
Klotzsch, F. & Perers, W.C.H. (1861) Polygalinae. Naturwissenschaften Reise Mossambique 1: 113–117.
Krachai, P., Chantaranothai, P. & Piwpuan, N. (2009) Pollen characteristics of Polygala, Salomonia and Xanthophyllum (Polygalaceae) in Thailand. Tropical Natural History 9 (1): 27–34.
Linnaeus, C. (1753) Species plantarum. L. Salvii, Holmiae [Stockholm], pp. 701–704.
Marques, M.C.M. (2003) Estudo taxonômico do gênero Polygala L. subgênero Ligustrina (Chodat) Paiva (Polygalaceae). PhD Thesis, Museu Nacional do Rio de Janeiro.
Mathew, L. & Teresa, M.V.M. (2010) Seed morphology and its systematic implications in Polygala L.(Polygalaceae). Phytomorphology 60 (3/4): 71–76.
Milby, T.H. (1976) Studies in the floral anatomy of Polygala (Polygalaceae). American Journal of Botany 63 (10): 1319–1326. https://doi.org/10.1002/j.1537-2197.1976.tb13217.x
Mohl, H. (1834) Über den Bau und die Formen der Pollenkôrner. Beiträge zur Anatomie und Physiologie der Gewächse. Bern: Ersters Heft.
Mohl, H. (1835) Sur la structure et lês formes dês graines de pollen. Annales des Sciences Naturelles 3: 220–236.
Nauman, C.E. (1981) Polygala grandiflora (Polygalaceae) Walter re-examined. SIDA, Contributions to Botany 1–18.
Paiva, J.A. & Dias, J.D.S. (1989) The pollen grain of Polygala fruticosa Berg. (Polygalaceae). Anales del Jardín Botánico de Madrid 47 (2): 377–385.
Paiva, J.A.R. & De Cuba, A.D.C. (1998) Polygalarum africanarum et madagascariensium prodromus atque gerontogaei generis’ Heterosamara ’Kuntze, a genere ’Polygala’ L. segregate et a nobis denuo recepti, synopsis monographica. Cyanus 1: 346.
Pastore, J.F.B., Abbott, J.R., Neubig, K.M., Van Den Berg, C. & Mota, M.C.D. (2019) Phylogeny and biogeography of Polygala (Polygalaceae). Taxon 68 (4): 673–691. https://doi.org/10.1002/tax.12119
Persson, C. (2001) Phylogenetic relationship in Polygalaceae based on plastid DNA sequences from the trnL-F region. Taxon 50: 763–779.
Punt, W., Hoen, P.P., Blackmore, S., Nilsson, S. & Le Thomas, A. (2007) Glossary of pollen and spore terminology. Review of palaeobotany and palynology 143 (1–2): 1–81. https://doi.org/10.1016/j.revpalbo.2006.06.008
Sarvi, A., Beygom Faghir, M. & Shahi Shavoon, R. (2020) Polygala mazandaranica (Polygalaceae), a new species from Mazandaran province, North Iran. Phytologia Balcanica 26 (3): 461–469.
Sarvi, A., Beygom Faghir, M. & Shahi Shavoon, R. (2021) Polygala guilanica (Polygalaceae), a new species from Guilan province, North Iran. Phytotaxa 513 (2): 141–151. https://doi.org/10.11646/phytotaxa.513.2.5
Shishkin, B.K. (1949) Flora of the U.S.S.R. series initiated by Komarov 1. pp. 188–204.
U?urlu Ayd?n, Z. (2020) Comparative capsule and seed morphology of Polygala L. (Polygalaceae) species in Turkey with implications for taxonomy. Plant Biosystems 154 (2): 189–196. https://doi.org/10.1080/11263504.2019.1588177
Villanueva, E. & Ramos, Á. (1985) Contribución al estudio polínico de” Polygala” L. (“ Polygalaceae”) en la Península Ibérica. In Anales del Jardín Botánico de Madrid 42 (2): 377–388.
Welwitsch, F. (1869) Sertum Angolense, sive stirpium quarundam novarum vel minus cognitarum. Transactions of the Linnean Society of London 27 (1): 1–94. https://doi.org/10.1111/j.1096-3642.1869.tb00202.x
Wodehouse, R.P. (1935) Pollen Grains. Their Structure, Identification and Significance in Science and Medicine. McGraw-Hill, New York.
Zori?, L., Merkulov, L., Lukovi?, J. & Boža, P. (2010) Comparative seed morphology of Trifolium L. species (Fabaceae). Periodicum biologorum 112 (3): 263–272.
Aldridge, J. (1842) Memoir to determine the use of pollen in natural classification. Journal of Botany 1: 575–601.
APG, I.V. (2016) The linnean society of London. Botanical journal of the linnean society 181: 1–20.
Arroyo-Cosultchi, G., Terrazas, T., Arias, S. & Arreola-Nava, H.J. (2006) The systematic significance of seed morphology in Stenocereus (Cactaceae). Taxon 55 (4): 983–992. https://doi.org/10.2307/25065693
Banks, H., Klitgaard, B.B., Claxton, F., Forest, F. & Crane, P.R. (2008) Pollen morphology of the family Polygalaceae (Fabales). Botanical Journal of the Linnean Society 156 (2): 253–289. https://doi.org/10.1111/j.1095-8339.2007.00723.x
Barthlott, W. (1981) Epidermal and seed surface characters of plants: systematic applicability and some evolutionary aspects. Nordic Journal of Botany 1 (3): 345–355. https://doi.org/10.1111/j.1756-1051.1981.tb00704.x
Blake, S.F. (1916) A revision of the genus Polygala in Mexico, Central America, and the West Indies, 47. Contributions from the Gray Herbarium of Harvard University, pp. 11–17. https://doi.org/10.5962/bhl.title.4348
Boissier, E. & Heldreich, T. (1853) Diagnoses Plantarum Orientalium novarum ser. 2, 1. Apud B. Herrmann, Lipsiae, 57 pp.
Bojnanský, V. & Fargašová, A. (2007) Atlas of seeds and fruits of Central and East-European flora: The Carpathian Mountains region. Springer Science & Business Media.
Bonilla-Barbosa, J., Novelo, A., Orozco, Y.H. & Márquez-Guzmán, J. (2000) Comparative seed morphology of Mexican Nymphaea species. Aquatic Botany 68 (3): 189–204. https://doi.org/10.1016/S0304-3770(00)00125-X
Bornmüller, J.F.N. & Gauba, E. (1942) Repertorium Specierum Novarum Regni Vegetabilis. Centralblatt für Sammlung und Veroffentlichung von Einzeldiagnosen neuer Pflanzen, 51. [Edited by Friedrich Fedde]. Berlin, 33 pp.
Castro, S., Silveira, P., Navarro, L., Paiva, J. & Coutinho, A.P. (2009) Pollen morphology of Chamaebuxus (DC.) Schb., Chodatia Paiva and Rhinotropis (Blake) Paiva (Polygala L., Polygalaceae). Grana 48 (3): 179–192. https://doi.org/10.1080/00173130902938428
Chodat, R. (1891) Monographia Polygalacearum. Premiere partie. Mémoires de la Société de Physique et d’Histoire Naturelle de Genève 7: 1–143.
Chodat, R. (1893) Monographia polygalacearum Vol. 31. Impr. Aubert-Schuchardt.
Chrtek, J. & K?ísa, B. (1977) Flora Iranica, Family Polygalaceae, No. 124. Akademische Druckund, Graz, pp. 1–11.
Cosson, E.S. (1874) Bulletin de la Société Botanique de France, 20. Paris, 240 pp. https://doi.org/10.1080/00378941.1873.10825518
Cronquist, A. & Takhtadzhi?a?n, A.L. (1981) An Integrated System of Classification of Flowering Plants. Columbia university press, pp. 248–250.
Cullen, J. (1965) Polygala L. In: Davis, P.H. (Ed.) Flora of Turkey and the East Aegean Islands 1. Edinburgh University Press, Edinburgh, pp. 533–539.
de Candolle, A.P. (1824) Prodromus systematics, 1. Naturalis Regni vegetabilis, 326 pp.
Dönmez, A.A., U?urlu Ayd?n, Z. & I?ik, S. (2015) Polygala turcica (Polygalaceae), a new species from E Turkey, and a new identification key to Turkish Polygala. Willdenowia 45 (3): 429–434. https://doi.org/10.3372/wi.45.45309
Erdtman, G. (1952) Pollen morphology and plant taxonomy. Angiosperms. Chronica Botanica. Waltham, Massachusetts.
Erdtman, G. (1986) Pollen morphology and plant taxonomy: angiosperms (Vol. 1). Hafner Publishing Company, New York.
Eriksen, B. (1993) Phylogeny of the Polygalaceae and its taxonomic implications. Plant Systematics and Evolution 186: 33–55. https://doi.org/10.1007/BF00937712
Eriksen, B. & Persson, C. (2007) Polygalaceae. In Flowering Plants–Eudicots. Springer, Berlin, Heidelberg, pp. 345–363. https://doi.org/10.1007/978-3-540-32219-1_41
Eriksson, O., Friis, E.M. & Löfgren, P. (2000) Seed size, fruit size, and dispersal systems in angiosperms from the Early Cretaceous to the Late Tertiary. The American Naturalist 156 (1): 47–58. https://doi.org/10.1086/303367
Fagundez, J. & Izco, J. (2011) Seed morphology and anatomy of the Mediterranean pentamerous species of Erica (Ericaceae). Turkish Journal of Botany 35 (6): 643–651. https://doi.org/10.3906/bot-1006-22
Fischer, F.E.L., Meyer, C.A. & Trautvettler, E.R. (1838) Index quartum seminum, quae Hortus Botanicus Imperialis Petropolitanus promutua commutatione offert. Accedunt animadversiones botanicae nonnullae. St. Petersburg, 42 pp.
Forest, F., Chase, M.W., Persson, C., Crane, P.R. & Hawkins, J.A. (2007) The role of biotic and abiotic factors in evolution of ant dispersal in the milkwort family (Polygalaceae). Evolution: International Journal of Organic Evolution 61 (7): 1675–1694. https://doi.org/10.1111/j.1558-5646.2007.00138.x
Fritzsche, J. (1832) Beiträge zur Kenntniss des Pollen (Vol. 1). Stellin and Elbing, Nicolai, Berlin.
Furness, S.H. & Stafford, P.J. (1995) Polygalaceae. The Northwest European Pollen Flora. Review of Palaeobotany and Palynology 88 (1–4): 61–82.https://doi.org/10.1016/0034-6667(95)00019-T
Ghimire, B., Jeong, M.J., Suh, G.U., Heo, K. & Lee, C.H. (2018) Seed morphology and seed coat anatomy of Fraxinus, Ligustrum and Syringa (Oleeae: Oleaceae) and its systematic implications. Nordic Journal of Botany 36 (10): e01866.https://doi.org/10.1111/njb.01866
Heubl, G.R. (1984) Systematische Untersuchungen an mitteleuropaischen Polygala Arten. Mitteilungen der Botanischen Staatssammlung, Munchen.
Isaacs, M.J.H., Weitz, F.M. & Johnson, C.T. (1993) Seed-coat characteristics of selected southern African species of Polygala L.(Polygalaceae). South African Journal of Botany 59 (6): 592–596. https://doi.org/10.1016/S0254-6299(16)30675-5
Jalilian, N. (2005) Flora of Iran, No. 49: Polygalaceae. Agriculture Institute of forests and rangelands press. [In Persian]
Kerrigan, R.A. (2012) A treatment for Polygala of northern Australia. Australian Systematic Botany 25 (2): 83–137. https://doi.org/10.1071/SB08032
Klotzsch, F. & Perers, W.C.H. (1861) Polygalinae. Naturwissenschaften Reise Mossambique 1: 113–117.
Krachai, P., Chantaranothai, P. & Piwpuan, N. (2009) Pollen characteristics of Polygala, Salomonia and Xanthophyllum (Polygalaceae) in Thailand. Tropical Natural History 9 (1): 27–34.
Linnaeus, C. (1753) Species plantarum. L. Salvii, Holmiae [Stockholm], pp. 701–704.
Marques, M.C.M. (2003) Estudo taxonômico do gênero Polygala L. subgênero Ligustrina (Chodat) Paiva (Polygalaceae). PhD Thesis, Museu Nacional do Rio de Janeiro.
Mathew, L. & Teresa, M.V.M. (2010) Seed morphology and its systematic implications in Polygala L.(Polygalaceae). Phytomorphology 60 (3/4): 71–76.
Milby, T.H. (1976) Studies in the floral anatomy of Polygala (Polygalaceae). American Journal of Botany 63 (10): 1319–1326. https://doi.org/10.1002/j.1537-2197.1976.tb13217.x
Mohl, H. (1834) Über den Bau und die Formen der Pollenkôrner. Beiträge zur Anatomie und Physiologie der Gewächse. Bern: Ersters Heft.
Mohl, H. (1835) Sur la structure et lês formes dês graines de pollen. Annales des Sciences Naturelles 3: 220–236.
Nauman, C.E. (1981) Polygala grandiflora (Polygalaceae) Walter re-examined. SIDA, Contributions to Botany 1–18.
Paiva, J.A. & Dias, J.D.S. (1989) The pollen grain of Polygala fruticosa Berg. (Polygalaceae). Anales del Jardín Botánico de Madrid 47 (2): 377–385.
Paiva, J.A.R. & De Cuba, A.D.C. (1998) Polygalarum africanarum et madagascariensium prodromus atque gerontogaei generis’ Heterosamara ’Kuntze, a genere ’Polygala’ L. segregate et a nobis denuo recepti, synopsis monographica. Cyanus 1: 346.
Pastore, J.F.B., Abbott, J.R., Neubig, K.M., Van Den Berg, C. & Mota, M.C.D. (2019) Phylogeny and biogeography of Polygala (Polygalaceae). Taxon 68 (4): 673–691. https://doi.org/10.1002/tax.12119
Persson, C. (2001) Phylogenetic relationship in Polygalaceae based on plastid DNA sequences from the trnL-F region. Taxon 50: 763–779.
Punt, W., Hoen, P.P., Blackmore, S., Nilsson, S. & Le Thomas, A. (2007) Glossary of pollen and spore terminology. Review of palaeobotany and palynology 143 (1–2): 1–81. https://doi.org/10.1016/j.revpalbo.2006.06.008
Sarvi, A., Beygom Faghir, M. & Shahi Shavoon, R. (2020) Polygala mazandaranica (Polygalaceae), a new species from Mazandaran province, North Iran. Phytologia Balcanica 26 (3): 461–469.
Sarvi, A., Beygom Faghir, M. & Shahi Shavoon, R. (2021) Polygala guilanica (Polygalaceae), a new species from Guilan province, North Iran. Phytotaxa 513 (2): 141–151. https://doi.org/10.11646/phytotaxa.513.2.5
Shishkin, B.K. (1949) Flora of the U.S.S.R. series initiated by Komarov 1. pp. 188–204.
U?urlu Ayd?n, Z. (2020) Comparative capsule and seed morphology of Polygala L. (Polygalaceae) species in Turkey with implications for taxonomy. Plant Biosystems 154 (2): 189–196. https://doi.org/10.1080/11263504.2019.1588177
Villanueva, E. & Ramos, Á. (1985) Contribución al estudio polínico de” Polygala” L. (“ Polygalaceae”) en la Península Ibérica. In Anales del Jardín Botánico de Madrid 42 (2): 377–388.
Welwitsch, F. (1869) Sertum Angolense, sive stirpium quarundam novarum vel minus cognitarum. Transactions of the Linnean Society of London 27 (1): 1–94. https://doi.org/10.1111/j.1096-3642.1869.tb00202.x
Wodehouse, R.P. (1935) Pollen Grains. Their Structure, Identification and Significance in Science and Medicine. McGraw-Hill, New York.
Zori?, L., Merkulov, L., Lukovi?, J. & Boža, P. (2010) Comparative seed morphology of Trifolium L. species (Fabaceae). Periodicum biologorum 112 (3): 263–272.