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Issue: Vol. 714 No. 3: 13 Aug. 2025
Type: Article
Published: 2025-08-13
DOI: 10.11646/phytotaxa.714.3.1
Page range: 197-213
Abstract views: 508
PDF downloaded: 10

Significance of basal-most leaf anatomy in taxonomy of Gagea (Liliaceae): a study of species from Kurdistan, Iraq

Department of Biology, Faculty of Sciences, University of Guilan, Rasht, Iran
Department of Biology, Faculty of Sciences, University of Guilan, Rasht, Iran
Department of Biology, Faculty of Sciences, University of Guilan, Rasht, Iran
flora of Iraq leaf morphology Liliales south-western Asian flora Tulipeae Monocots

Abstract

In this study, anatomical characteristics of the basal-most leaf of 11 species of the Gagea from the Kurdistan region of Iraq were assessed. To assess the value of quantitative and qualitative variables in describing the interspecific and intersectional relationships, factor analysis of mixed data (FAMD) analysis was performed followed by a hierarchical clustering on principal components (HCPC). The results showed that the basal-most leaf anatomical features were taxonomically informative and can be used as important criteria for the delimitation of sections and species of the genus. Some characters of the basal leaf, in particular the shape, apex, presence and number of angular collenchyma bundles, width of the leaf blade and thickness of the upper and lower epidermis, were of value for the differentiation of the studied taxa. Finally, an identification key was established on the basis of these anatomical characteristics.

References

  1. Aitchison, J.E.T. (1880) On the flora of the Kuram Valley, & c., Afghanistan. Botanical Journal of the Linnean Society 18: 1–113. https://doi.org/10.1111/j.1095-8339.1880.tb00458.x
  2. Akyol, Y., Kocabaş, O., Yetişen, K., Bozdağ, B. & Özdemir, C. (2015) Morphological and anatomical investigations of some Gagea Salisb. species (Liliaceae) in Turkey. Journal of Biological Research and Development 5: 1–10.
  3. Bayer, E. & González, G.L. (1989) Nomenclatural notes on some names in Gagea Salisb. [Liliaceae]. TAXON 38: 643–645. https://doi.org/10.2307/1222655
  4. Boissier, E. (1881) Flora Orientalis, vol. 5. Georg, Basel, 868 pp.
  5. Borsuk, A.M., Roddy, A.B., Théroux‐Rancourt, G. & Brodersen, C.R. (2022) Structural organization of the spongy mesophyll. New Phytologist 234: 946–960. https://doi.org/10.1111/nph.17971
  6. Caparelli, K.F., Peruzzi, L. & Cesca, G. (2006) A comparative analysis of embryo-sac development in three closely related Gagea species (Liliaceae), with some consideration on their reproductive strategies. Plant Biosystems 140: 115–122. https://doi.org/10.1080/11263500600756090
  7. Chen, X. & Turland, N.J. (2000) Lloydia Reichenbach. In: Wu, Z.Y. & Raven, P.H. (Eds.) Flora of China 24. Science Press, Beijing and Missouri Botanical Garden Press, p St. Louis, pp. 121–123.
  8. Dasgupta, S. & Deb, D.B. (1983) Taxonomic revision of the genus Gagea Salisb. (Liliaceae) in India and adjoining regions. Journal of the Bombay Natural History Society 83: 78–97. [https://biostor.org/reference/153513]
  9. Davlianidze, M.T. (1972) Conspectus of the Caucasian representatives of the genus Gagea Salisb. I. Notulae Systematicae ac Geographicae Instituti Botanici Thbilissiensis 29: 1–71.
  10. Davlianidze, M.T. (1973) Conspectus of the Caucasian representatives of the genus Gagea Salisb. II. Notulae Systematicae ac Geographicae Instituti Botanici Thbilissiensis 30: 1–62.
  11. Davlianidze, M.T. (1976) Kavkazaskie Predstaviteli Roda Gagea Salisb. Metsniereba, Tbilisi, pp.1–160.
  12. Ezamakhshi, B.A.M., Faghir, M.B. & Zamani, A. (2024) Three records of Gagea for the flora of Iraq. Rostaniha 25: 173–180. https://doi.org/10.22092/bot.j.iran.2024.367188.1400
  13. Fedtschenko, B.A. (1932) Flora Turkmenii, vol.1. Izd. Turkmenskogu fililala, Akademiia nauk SSSR, Ashkhabad, 392 pp.
  14. Feinbrun-Dotham, N. (1986) Flora Palaestina: Alismataceae to Orchidaceae, vol. 4. Israel Academy of Sciences and Humanities, Jerusalem,‏ 469 pp.
  15. Greilhuber, J., Ebert, I., Lorenz, A. & Vyskot, B. (2000) Origin of facultative heterochromatin in the endosperm of Gagea lutea (Liliaceae). Protoplasma 212: 217–226. https://doi.org/10.1007/BF01282922
  16. Grossheim, A.A. (1935) Gagea caroli-kochii. In: Komarov, V.L. (Ed.) Flora of the U.S.S.R., vol. 4. Izdatel’stvo Akademii Nauk SSSR, Leningrad, pp. 736.
  17. Heyn, C.H. & Dafni, A. (1971) Studies in the genus Gagea (Liliaceae) I. The platyspermous species in Israel and neighbouring areas. Israel Journal of Botany 20: 214–233.
  18. Heyn, C.H. & Dafni, A. (1977) Studies in the genus Gagea (Liliaceae) II. The non-platyspermous species from the Galilee, the Golan heights and Mt. Hermon. Israel Journal of Botany 26: 11–22.
  19. Hu, Z.M., Zhao, C.H., Zhao, Y.Y. & Liu, J.X. (2021) Pollen morphology of Liliaceae and its systematic significance. Palynology 45: 531–568. https://doi.org/10.1080/01916122.2021.1882601
  20. IPNI (2024) International plant names index. Published on the Internet http://www.ipni.org, The Royal Botanic Gardens, Kew, Harvard University Herbaria & Libraries and Australian National Herbarium. Available from: https://ipni.org/ (Retrieved 11 September 2024)
  21. Ivanova, L.A. & P’yankov, V.I. (2002) Structural adaptation of the leaf mesophyll to shading. Russian Journal of Plant Physiology 49: 419–431. https://doi.org/10.1023/A:1015513607202
  22. Koch, K. (1849) Beiträge zu einer Flora des Orientes (Fortsetzung): Gagea. Linnaea 22: 226–231.
  23. Kosenko, V.N. (1999) Contributions to the pollen morphology and taxonomy of the Liliaceae. Grana 38: 20–30. https://doi.org/10.1080/001731300750044672
  24. Kurbaniyazova, G.T., Levichev, I.G. & Kadirov, U.K. (2022) The species of the genus Gagea Salisb. is distribution in the flora of the Urgut region (Uzbekistan). American Journal of Plant Sciences 13: 1183–1195. https://doi.org/10.4236/ajps.2022.139080
  25. Lê, S., Josse, J. & Husson, F. (2008) FactoMineR: An R package for multivariate analysis. Journal of Statistical Software 25: 1–18. https://doi.org/10.18637/jss.v025.i01
  26. Levichev, I.G. (1990) The synopsis of the genus Gagea (Liliaceae) from the western Tien Shan. Botanicheskii Zhurnal 75: 225–234. (In Russian)
  27. Levichev, I.G. (1999a) Cooperation in genus Gagea. Optima Newsletter 34 (e): 12.
  28. Levichev, I.G. (1999b) The morphology of Gagea Salisb. (Liliaceae) I. Subterranean organs. Flora 194: 379–392. https://doi.org/10.1016/S0367-2530(17)30929-5
  29. Levichev, I.G. (2001) New species of the genus Gagea Salisb. (Liliaceae) from western regions of Asia. Turczaninowia 4: 5–35. (In Russian)
  30. Levichev, I.G. (2006) A review of the Gagea (Liliaceae) species in the flora of Caucasus. Botanicheskii Zhurnal 91: 917–951.
  31. Levichev, I.G. (2013) Structural features of shoots in Lloydia, Gagea, Kharkevichia (Liliaceae) as evolutionary variability of the modules of mesome nature in monocotyledons. Botanical Journal 98: 409–452. https://doi.org/10.1134/S1234567813040010
  32. Lin, M., Qiu, J., Xie, K. & Tan, D. (2023) Palynological features and taxonomic significance for 16 species of Gagea (Liliaceae) from Xinjiang, China. PhytoKeys 225: 53–68. https://doi.org/10.3897/phytokeys.225.101518
  33. Linnaeus, C. (1753) Species plantarum, vol. 1. Salvius, Stockholm, 530 pp.
  34. Liu, X., Wang, X., Chen, H., Chen, K., Mo, W., Yuan, Y., Zhu, J., Wang, R. & Zhang, S. (2023) Is the multi-species variation in leaf anatomical traits along the environmental gradient modulated by herbaceous functional groups? Ecological Indicators 154: 110880. https://doi.org/10.1016/j.ecolind.2023.110880
  35. Mabberley, D.J. (1997) The plant book. Cambridge University Press, Cambridge, 858 pp.
  36. Melchior, H. (1964) Reihe Liliiflorae, In: Melchior, H. (Ed.) Engler’s Syllabus der Pflanzenfamilien 2, 12th edition. Borntraeger, Berlin, pp. 513–542.
  37. Metcalfe, C.R. & Chalk, L. (1979) Anatomy of the dicotyledons, vol. 1. Oxford University Press, Oxford, 318 pp.
  38. Mistchenko, P.I. (1908) Gagea alexeenkoana. Trudy Botanicheskago Sada Imperatorskago 9: 76.
  39. Novikov, A. (2021) Comparative floral morphology and anatomy of Gagea s. str. and Lloydia. Acta Agrobotanica 74: 1–10. https://doi.org/10.5586/aa.7412
  40. Pallas, P.S. (1776) Reise durch Verschiedene Provinzen des Russischen Reichs 3: App. 727.
  41. Pascher, A.A. (1904) Übersicht über die Arten der Gattung Gagea. Sitzungberichte des Deutschen naturwissenschaflich-medicischen Vereins für Böhmen ‘Lotos’ 24: 116–118.
  42. Pascher, A.A. (1905) Conspectus gagearum Asiae. Bulletin of the de la Societe Imperiale des Naturalistes de Moscou 19: 353–375.
  43. Peruzzi, L. (2003) Contribution to the cytotaxonomical knowledge of Gagea Salisb. (Liliaceae) sect. Foliatae A. Terracc. and synthesis of karyological data. Caryologia 56: 115–128. https://doi.org/10.1080/00087114.2003.10589314
  44. Peruzzi, L. (2008) Hybridity as a main evolutionary force in the genus Gagea Salisb. (Liliaceae). Plant Biosystems 142: 179–183. https://doi.org/10.1080/11263500701873042
  45. Peruzzi, L. (2012a) Nomenclatural novelties at sectional level in Gagea (Liliaceae). Atti della Società Toscana di Scienze Naturali, Memorie, serie B, 118: 23–24.
  46. Peruzzi, L. (2012b) Chromosome diversity and evolution in Gagea (Liliaceae). Bocconea 24: 147–158.
  47. Peruzzi, L. (2016) A new infrafamilial taxonomic setting for Liliaceae, with a key to genera and tribes. Plant Biosystems 150: 1341–1347. https://doi.org/10.1080/11263504.2015.1115435
  48. Peruzzi, L. & Aquaro, G. (2005) Contribution to the cytotaxonomical knowledge of Gagea Salisb. (Liliaceae). II. Further karyological studies on Italian populations. Candollea 60: 237–253.
  49. Peruzzi, L., Peterson, A., Tison, J.M. & Peterson, J. (2008a) Phylogenetic relationships of Gagea Salisb. (Liliaceae) in Italy, inferred from molecular and morphological data matrices. Plant Systematics and Evolution 276: 219–234. https://doi.org/10.1007/s00606-008-0081-4
  50. Peruzzi, L., Peterson, A., Tison, J.M. & Peterson, J. (2008b) On the phylogenetic position and taxonomic value of Gagea trinervia (Viv.) Greuter and Gagea sect. Anthericoides A.Terracc. (Liliaceae). Taxon 57: 1201–1214. https://doi.org/10.1002/tax.574013
  51. Peterson, A., Levichev, I.G. & Peterson, J. (2008) Systematics of Gagea and Lloydia (Liliaceae) and infrageneric classification of Gagea based on molecular and morphological data. Molecular Phylogenetics and Evolution 46: 446–465. https://doi.org/10.1016/j.ympev.2007.11.016
  52. Peterson, A., John, H., Koch, E. & Peterson, J. (2004) A molecular phylogeny of the genus Gagea (Liliaceae) in Germany inferred from non-coding chloroplast and nuclear DNA sequences. Plant Systematics and Evolution 245: 145–162. https://doi.org/10.1007/s00606-003-0114-y
  53. Peterson, A., Levichev, I.G., Peterson, J., Harpke, D. & Schnittler, M. (2011) New insights into the phylogeny and taxonomy of Chinese species of Gagea (Liliaceae)—speciation through hybridization. Organisms Diversity and Evolution 11: 387–407. https://doi.org/10.1007/s13127-011-0059-x
  54. Peterson, A., Harpke, D., Peterson, J., Harpke, A. & Peruzzi, L. (2019) A pre‐Miocene Irano‐Turanian cradle: origin and diversification of the species‐rich monocot genus Gagea (Liliaceae). Ecology and Evolution 9: 5870–5890. https://doi.org/10.5061/dryad.97np7bt
  55. Peterson, A., Harpke, D., Levichev, I.G., Beisenova, S., Schnittler, M. & Peterson, J. (2016) Morphological and molecular investigations of Gagea (Liliaceae) in southeastern Kazakhstan with special reference to putative altitudinal hybrid zones. Plant Systematics and Evolution 302: 985–1007. https://doi.org/10.1007/s00606-016-1313-7
  56. P’yankov, V.I. & Kondrachuk, A.V. (2003) Basic types of structural changes in the leaf mesophyll during adaptation of eastern Pamir plants to mountain conditions. Russian Journal of Plant Physiology 50: 28–35. https://doi.org/10.1023/A:1021976132165
  57. POWO (2024) Plants of the World online. Facilitated by the Royal Botanic Gardens, Kew. Available from: https://powo.science.kew.org/ Retrieved 28 November 2024.
  58. Qiu, J., Lin, M. & Tan, D. (2023) Taxonomic implications of leaf morphology and epidermal anatomy for 14 species of Gagea (Liliaceae) from Xinjiang, China. Botanical Studies 64: 33. https://doi.org/10.1186/s40529-023-00405-9
  59. R Core Team (2024) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available from: https://www.R-project.org/
  60. Rastegari Khah, L., Heidarain, M. & Masoumi, S.M. (2023) Pollen micromorphology of some species of Gagea in Iran and its taxonomic implication. Rostaniha 24: 109–120. [https://dorl.net/dor/20.1001.1.16084306.2023.24.2.3.4]
  61. Rawat, G.S. & Pangtey, Y.P.S. (1987) Floristic structure of snowline vegetation in central Himalaya, India. Arctic Alpine Research 19: 195–201. https://doi.org/10.2307/1551252
  62. Rechinger, K.H. (1986) Six new species of Gagea (Liliaceae) from the flora Iranica area. Plant Systematic and Evolution 153: 287–292. https://doi.org/10.1007/BF00983695
  63. Rix, E.M. (1984) Gagea Salisb. In: Davis, P.H. (Ed.) Flora of Turkey and the east Aegean islands 8. Edinburgh University Press, Edinburgh, pp.312–327.
  64. Salisbury, R.A. (1806) On the characters of a distinct genus hitherto confounded with Ornithogalum, and called Gagea; with some remarks on the importance of the inflorescence in distinguishing genera. Annals of Botany 2: 553–557.
  65. Schultes, J.A. (1829) Gagea. In: Roemer, J.J. & Schultes, J.A. (Eds.) Systema vegetabilium 7. Cottae, Stuttgart, 753 pp.
  66. Sezer, O. & Yildiz, A.C. (2021) Pollen morphology studies on some Gagea Salisb. (Liliaceae) species from Turkey. Pakistan Journal of Botany 53: 1073–1077. https://doi.org/10.30848/PJB2021-3(21)
  67. Sezer, O., Yildiz, A.C., Koyuncu, O., Yaylaci, K. & Erkara, I.P. (2018) Investigation on pollen morphology of two Gagea Salisb. taxa from Turkey. Communications Faculty of Sciences University of Ankara Series C Biology 27: 232–237. https://doi.org/10.1501/commuc_0000000219
  68. Stapf, O. (1885) Beiträge zur Flora von Lycien, Carien und Mesopotamien. Gerold, Vienna, 100 pp.
  69. Stroh, G. (1937) Die Gattung Gagea Salisb. Beihefte zum Botanischen Centralblatt 57: 485–520.
  70. Tekşen, M. & Erkul, S.K. (2015) The synopsis of the genus Gagea (Liliaceae) in Turkey. Phytotaxa 230: 101–129. https://doi.org/10.11646/phytotaxa.230.2.1
  71. Terashima, I. (1992) Anatomy of non-uniform leaf photosynthesis. Photosynthesis Research 31: 195–212. https://doi.org/10.1007/BF00035537
  72. Terracciano, A. (1904) Gagearum novarum diagnoses. Bollettino del Real Giardino Botanico di Palermo 2: 1–7.
  73. Terracciano, A. (1905) Revisione monografica delle Gagea della flora spagnola. Boletín de la Sociedad Aragonesa de Ciencias Naturales 4: 188–253.
  74. Terracciano, A. (1906) Gagearum species florae orientalis. Bulletin de l’Herbier Boissier 2: 105–120.
  75. Tison, J.M. (2009) An update of the genus Gagea Salisb. (Liliaceae) in the Iberian Peninsula. Lagascalia 29: 7–22. Available from: http://hdl.handle.net/11441/55708
  76. Tison, J.M., Peterson, A., Harpke, D. & Peruzzi, L. (2013) Reticulate evolution of the critical Mediterranean Gagea sect. Didymobulbos (Liliaceae) and its taxonomic implications. Plant Systematics and Evolution 299: 413–438. https://doi.org/10.1007/s00606-012-0731-4
  77. Turczaninov, N.S. (1844) Plecostigma paucifolium Turcz. In: Trautwetter, E.R. (Ed.) Plantarum imagines descriptiones floram Russicam. Schweizerbart Stuttgart, pp. 8–9.
  78. Uphof, J.C. (1958) A review of the genus Gagea Salisb. I. Plant Life 14: 124–132.
  79. Uphof, J.C. (1959) A review of the genus Gagea Salisb. II. Plant Life 15: 152–161.
  80. Uphof, J.C. (1960) A review of the genus Gagea Salisb. III. Plant Life 16: 163–176.
  81. Villars, D. (1787) Histoire des Plantes de Dauphiné, tome 2. The author, Grenoble, 690 pp.
  82. Vvedensky, A.I. (1946) Botanicheskie Materialy Gerbariya Botanicheskogo Instituta Imeni V. L. Komarova Akademii Nauk S.S.S.R. 9: 238.
  83. Wendelbo, P. (1985) Gagea. In: Guest, E. & Townsend, C.C. (Eds.) Flora of Iraq 3. Ministry of Agriculture of the Republic of Iraq, pp. 65–75.
  84. Wendelbo, P. & Rechinger, K.H. (1990) Gagea. In: Rechinger, K.H. (Ed.) Flora Iranica 165. Akademische Druck-und Verlagsanstalt, Graaz, pp.13–57.
  85. Willis, J.C. (1980) A dictionary of the flowering plants and ferns. Cambridge University Press, Cambridge, 1346 pp.
  86. Zarrei, M. & Zarre, S. (2003) A new species of Gagea (Liliaceae) from Iran. Nordic Journal of Botany 23: 269–274. https://doi.org/10.1111/j.1756-1051.2003.tb00391.x
  87. Zarrei, M., Zarre, S., Wilkin, P. & Rix, M. (2007) Systematic revision of the genus Gagea Salisb. (Liliaceae) in Iran. Botanical Journal of the Linnean Society 154: 559–588. https://doi.org/10.1111/j.1095-8339.2007.00678.x
  88. Zarrei, M., Wilkin, P., Ingrouille, M.J., Zarre, S. & Chase, M.W. (2010) The systematic importance of anatomical data in Gagea (Liliaceae) from the Flora Iranica area. Botanical Journal of the Linnean Society 164: 155–177. https://doi.org/10.1111/j.1095-8339.2010.01081.x
  89. Zarrei, M., Wilkin, P., Ingrouille, M.J. & Chase, M.W. (2011) A revised infrageneric classification for Gagea (Tulipeae; Liliaceae): insights from DNA sequence and morphological data. Phytotaxa 15: 44–56. https://doi.org/10.11646/phytotaxa.15.1.6
  90. Zarrei, M., Wilkin, P., Fay, M.F., Ingrouille, M.J., Zarre, S. & Chase, M.W. (2009) Molecular systematics of Gagea and Lloydia (Liliaceae; Liliales): implications of analyses of nuclear ribosomal and plastid sequences for infrageneric classification. Annals of Botany 104: 125–142. https://doi.org/10.1093/aob/mcp103
  91. Zuccarini, J.G. (1843) Bulbillaria gageoides. Abhandlungen der Mathematisch-Physikalischen Classe der Königlich Bayerischen Akademie der Wissenschaften 3: 230–231.

How to Cite

Ezamakhshi, B.A.M., Faghir, M.B. & Zamani, A. (2025) Significance of basal-most leaf anatomy in taxonomy of Gagea (Liliaceae): a study of species from Kurdistan, Iraq. Phytotaxa 714 (3): 197–213. https://doi.org/10.11646/phytotaxa.714.3.1