| تعداد نشریات | 285 |
| تعداد نشریات سازمان | 83 |
| تعداد شمارهها | 3,133 |
| تعداد مقالات | 34,850 |
| تعداد مشاهده مقاله | 50,288,172 |
| تعداد دریافت فایل اصل مقاله | 89,439,798 |
Phylogeny and species delimitation in the Medicago rigidula (L.) All. complex based on nrDNA sequence data | ||
| The Iranian Journal of Botany | ||
| دوره 31، شماره 2 - شماره پیاپی 62، اسفند 2025، صفحه 177-193 اصل مقاله (1.52 M) | ||
| نوع مقاله: Research Paper | ||
| شناسه دیجیتال (DOI): 10.22092/ijb.2025.369441.1523 | ||
| نویسندگان | ||
| Zahra Ghorbani1؛ Iraj Mehregan* 2؛ Mostafa Assadi3؛ Ernest Small4 | ||
| 1Department of Plant Sciences, Faculty of Biological Sciences, Kharazmi University, Tehran, Iran. | ||
| 2Department of Biology, SR.C., Islamic Azad University, Tehran, Iran | ||
| 3Research Institute of Forests and Rangelands, Agricultural Research Education and Extension Organization (AREEO), Tehran, Iran. | ||
| 4Biosystematics Research Institute, Agriculture Canada, K1A OC6, Ottawa, Ontario, Canada. | ||
| چکیده | ||
| Annual species of the genus Medicago L. (Fabaceae) are ecologically significant components of Mediterranean vegetation. The Medicago rigidula complex, comprising M. rigidula, M. rigiduloides, M. sinskiae, and M. constricta, has undergone substantial taxonomic revision recently, but species boundaries remain unclear due to overlapping morphological traits, recent divergence, and selfing reproductive strategies. To clarify species relationships within the complex, we conducted an integrative analysis of populations sampled from diverse regions of Iran. Floral morphology (34 characters), anatomical features (17 characters), palynology, and seed micromorphology were studied using light and SEM microscopy. Morphological and anatomical differences, particularly in floral characters, pollen aperture types, seed surface vesiculation, and pod vascular bundle shape, helped discriminate among taxa. Phylogenetic analyses based on combined ITS and ETS nuclear ribosomal DNA sequences supported the distinctiveness of M. constricta and M. sinskiae, while also validating the separation of M. rigidula and M. rigiduloides despite partial morphological overlap. A new ETS primer pair, designed specifically for the core Medicago group, improved amplification success and resolution. Cluster and PCA analyses of floral characters further supported species-level distinctions. Although clinal variation was observed in pod morphology, integration of multiple data sources enabled more robust species delimitation. Our findings underscore the importance of combining multilocus molecular data with detailed morphological and anatomical traits to resolve taxonomic complexity in recently diverged, selfing taxa. An updated identification key for the M. rigidula complex is provided, contributing to the systematics within the group. | ||
| کلیدواژهها | ||
| Annual medics؛ Iran؛ Mediterranean region؛ primer design؛ speciation؛ taxonomy | ||
| عنوان مقاله [English] | ||
| فیلوژنی و تعیین حدود گونه ها در کمپلکس گونه ای Medicago rigidula (L.) All. بر اساس داده های توالی nrDNA | ||
| نویسندگان [English] | ||
| زهرا قربانی1؛ ایرج مهرگان2؛ مصطفی اسدی3؛ ارنست اسمال4 | ||
| 1گروه علوم گیاهی، دانشکده علوم زیستی، دانشگاه خوارزمی، تهران | ||
| 2گروه زیست شناسی، دانشکده علوم پایه دانشگاه آزاد اسلامی واحد علوم و تحقیقات، تهران، ایران | ||
| 3موسسه تحقیقات جنگلها و مراتع کشور، سازمان تحقیقات، آموزش و ترویج کشاورزی، تهران | ||
| 4موسسه تحقیقاتی بیوسیستماتیک، وزارت کشاورزی کانادا، اوتاوا، اونتاریو، کانادا | ||
| چکیده [English] | ||
| گونههای یکساله جنسMedicago (تیره باقلائیان) از اجزای مهم پوشش گیاهی در ناحیه مدیترانهای به شمار میروند. مجموعهی Medicago rigidula که شامل گونههای M. rigidula، M. rigiduloides، M. sinskiae و M. constricta میباشد، در سالهای اخیر دستخوش بازنگریهای تاکسونومیکی قابل توجهی شده است. با این حال، به دلیل شباهتهای ریختشناسی، واگرایی افراد و سامانه تولیدمثل خودگشن، مرزهای گونهای در این مجموعه همچنان مبهم باقی ماندهاند. بهمنظور روشنسازی روابط گونهای، جمعیتهایی از مناطق مختلف ایران مورد بررسی قرار گرفتند. ریختشناسی گل (۳۴ صفت)، ویژگیهای تشریحی (۱۷ صفت)، ساختار دانه گرده و میکرومورفولوژی بذر با استفاده از میکروسکوپ نوری و الکترونی روبشی (SEM) مطالعه شد. تفاوتهای ریختی و تشریحی بهویژه در صفات گل، نوع منفذ دانههای گرده، ساختار سطح بذر و شکل دستههای آوندی در نیام، موجب تمایز بین گونهها گردید. تجزیههای فیلوژنتیکی مبتنی بر توالیهای ترکیبی ITS و ETS، موقعیت مجزای گونههای M. constricta وM. sinskiae را تأیید کردند و همچنین جداییM. rigidula و M. rigiduloides را با وجود همپوشانیهای جزئی در ریختشناسی تأیید نمودند. آغازگر جدیدی برای ناحیه ETS طراحی شد که موجب بهبود موفقیت در تکثیر و افزایش وضوح نتایج گردید. تحلیلهای خوشهای و PCA نیز از تفکیک گونهای حمایت کردند. با وجود مشاهده تغییرات تدریجی در ریختشناسی نیام، تلفیق دادههای مولکولی و ریختی امکان تعیین دقیقتر حدود گونهها را فراهم ساخت. کلید شناسایی بروز شده برای مجموعه M. rigidula نیز ارائه شده است. | ||
| کلیدواژهها [English] | ||
| ایران, تاکسونومی, طراحی پرایمر, گونه زایی, مناطق مدیترانه ای, یونجه های یکساله | ||
| مراجع | ||
|
Allioni, C. 1785: Flora Pedemontana sive Enumeratio Methodica Stirpium Indigenarum Pedemontii (Vol. 1). J. M. Briolus. -Turin, J.M. Briolus. Azizian, D., & Cutler, D.F. 1982: Anatomical, cytological and phytochemical studies on Phlomis L. and Eremostachys Bunge (Labiatae). -Bot. J. Linn. Soc. 85(4): 249–281. https://doi.org/10.1111/j.1095-8339.1982.tb00373.x Baldwin, B.G., & Markos, S. 1998: Phylogenetic utility of the external transcribed spacer (ETS) of 18S-26S rDNA: Congruence of ETS and ITS trees of Calycadenia (Compositae). -Mol. Phylogenet. Evol. 10(3): 449–463. https://doi.org/10.1006/mpev.1998.0545 Bayat, M., Assadi, M., Small, E., & Mehregan, I. 2021a: Investigating the taxonomic relationships and geographical distribution range of Medicago rigidula and M. rigiduloides in Iran. -Iran. J. Bot. 27(2): 115–129. https://doi.org/10.22092/ijb.2021.354838.1328 Bayat, M., Assadi, M., Small, E., & Mehregan, I. 2021b: Molecular studies of Iranian populations support the morphology-based taxonomic separation of Medicago rigidula and M. rigiduloides. -Phytotaxa 518(4): 281–299. https://doi.org/10.11646/phytotaxa.518.4.5 Bena, G. 2001: Molecular phylogeny supports the morphologically based taxonomic transfer of the “medicagoid” Trigonella species to the genus Medicago L. -Amer. J. Bot. 88(3): 466–475. https://doi.org/10.2307/2657114 Bena, G., Jubier, M.F., Olivieri, I., & Lejeune, B. 1998a: Ribosomal external and internal transcribed spacers: Combined use in the phylogenetic analysis of Medicago (Leguminosae). -J. Mol. Evol. 46(3): 299–306.https://doi.org/10.1007/pl00006306 Bena, G., Lejeune, B., Prosperi, J.M., & Olivieri, I. 1998b: Molecular phylogenetic approach for studying life-history evolution: The ambiguous example of the genus Medicago L. -Proc. Roy. Soc. B: Biol. Sci. 265(1401): 1141–1151. https://doi.org/10.1098/rspb.1998.0410 Bena, G., Prosperi, J.M., Lejeune, B., & Olivieri, I. 1998c: Evolution of annual species of the genus Medicago: A molecular phylogenetic approach. -Mol. Phylogenet. Evol. 9(3): 552–559. https://doi.org/10.1006/mpev.1998.0493 Branca, A., Paape, T., Zhou, P., Briskine, R., Farmer, A.D., Mudge, J., ... & Young, N.D. 2011: Whole-genome nucleotide diversity, recombination, and linkage disequilibrium in the model legume Medicago truncatula. -Proc. Natl. Acad. Sci. 108(42): E864–E870. https://doi.org/10.1073/pnas.1104032108 Douzery, E.J.P., Pridgeon, A.M., Kores, P., Linder, H. P., Kurzweil, H., & Chase, M.W. 1999: Molecular phylogenetics of Diseae (Orchidaceae): a contribution from nuclear ribosomal ITS sequences. -Amer. J. Bot. 86(6): 887–899. https://doi.org/10.2307/2656709 Downie, S.R., Katz-Downie, D.S., Rogers, E.J., Zujewski, H.L., & Small, E. 1998: Multiple independent losses of the plastid rpoC1 intron in Medicago (Fabaceae) as inferred from phylogenetic analyses of nuclear ribosomal DNA internal transcribed spacer sequences. –Can. J. Bot. 76(5): 791–803. https://doi.org/10.1139/b98-056 Doyle, J. J., & Doyle, J. L. 1987: A rapid DNA isolation procedure for small quantities of fresh leaf tissue. -Phytochem. Bull. 19: 11–15. Endress, P.K. 1994: Diversity and evolutionary biology of tropical flowers. Cambridge University Press. -Cambridge, Cambridge University Press. Erdtman, G. 1960: Pollen walls and angiosperm phylogeny. -Bot. Not. 113: 41–45. Felsenstein, J. 1988: Phylogenies from molecular sequences: Inference and reliability. -Ann. Rev. Genet. 22: 521–565. https://doi.org/10.1146/annurev.ge.22.120188.002513 Greene, S. L., Kisha, T. J., & Yu, L. X. 2020: Genetic diversity and population structure of Medicago species in relation to their center of diversity. -Crop Sci. 60(6): 3152–3164. https://doi.org/10.1002/csc2.20265 Heyn, C.C. 1963: The annual species of s. -Jerusalem, Magnes Press, The Hebrew University. Heyn, C.C. 1970: Medicago [annuals]. In P. H. Davis (Ed.), Flora of Turkey and the East Aegean Islands (Vol. 3, pp. 494–510). Edinburgh University Press. -Edinburgh, Edinburgh University Press. Heyn, C.C. 1984: Medicago. In K.H. Rechinger (Ed.), Flora Iranica (Vol. 157, Papilionaceae II, pp. 253–274). Akademische Druck- u. Verlagsanstalt. -Graz, Akademische Druck- u. Verlagsanstalt. Linnaeus, C. 1753: Species Plantarum (Vol. 2). Impensis Laurentii Salvii. -Stockholm, Impensis Laurentii Salvii. Maddison, D.R., & Maddison, W.P. 2005: MacClade 4: Analysis of phylogeny and character evolution. Version 4.08a. Sinauer Associates. -Sunderland, Sinauer Associates. Maureira-Butler, I.J., Pfeil, B.E., Muangprom, A., Osborn, T.C., & Doyle, J.J. 2008: The reticulate history of Medicago (Fabaceae). -Syst. Biol. 57(3): 466–482. https://doi.org/10.1080/10635150802172168 Mehregan, I., Rahiminejad, M.R., & Azizian, D. 2002: A taxonomic revision of the genus Medicago L. (Fabaceae) in Iran. -Iran. J. Bot. 9(2): 207–221. Mehregan, I. & Small, E. 2023: The genus Medicago L. In: M. Assadi & A.A. Maassoumi (eds.). Flora of Iran no. 177: 261-364. Miller, P. 1768: The Gardeners' Dictionary (8th ed.). Printed for the author. -London, Printed for the author. Posada, D., & Crandall, K. A. 1998: Modeltest: Testing the model of DNA substitution. -Bioinformatics 14(9): 817–818. https://doi.org/10.1093/bioinformatics/14.9.817 Punt, W., Hoen, P. P., Blackmore, S., Nilsson, S., & Le Thomas, A. 2007: Glossary of pollen and spore terminology. -Rev. Palaeobot. Palynol. 143(1–2): 1–81. https://doi.org/10.1016/j.revpalbo.2006.06.008 Ronfort, J., Bataillon, T., Santoni, S., Delalande, M., David, J. L., & Prosperi, J. M. 2006: Microsatellite diversity and broad scale geographic structure in a selfing species: Medicago truncatula. -Genetics 172(1): 373–386. https://doi.org/10.1534/genetics.105.044354 Ronquist, F., Huelsenbeck, J.P., & Van der Mark, P. 2005: MrBayes 3.1 Manual. Draft 5/17/2005. Shaffer, H.B., & Thomson, R.C. 2007: Delimiting species in recent radiations. -Syst. Biol. 56(6): 896–906. https://doi.org/10.1080/10635150701772563 Scotland, R.W., Olmstead, R.G., & Bennett, J. R. 2003: Phylogeny reconstruction: the role of morphology. -Syst. Biol. 52(4): 539–548. https://doi.org/10.1080/10635150390223620 Siol, M., Prosperi, J. M., Bonnin, I., & Ronfort, J. 2008: How multilocus genotypic pattern helps to understand the history of selfing populations: A case study in Medicago truncatula. -Heredity 100(6): 517–525. https://doi.org/10.1038/hdy.2008.7 Small, E. 1987a: A taxonomic study of the “medicagoid” Trigonella (Leguminosae). -Can. J. Bot. 65(6): 1199–1211. https://doi.org/10.1139/b87-166 Small, E. 1990: Medicago rigiduloides, a new species segregated from M. rigidula. -Can. J. Bot. 68(12): 2614–2617. https://doi.org/10.1139/b90-330 Small, E., Brookes, B. & Crawford, E.J. 1990: Intercontinental differentiation in Medicago rigidula. –Can. J. Bot. 68: 2607-2613. Small, E. 2011: Alfalfa and relatives: Evolution and classification of Medicago. NRC Research Press. -Ottawa, NRC Research Press. Small, E., & Jomphe, M. 1989: A synopsis of the genus Medicago (Leguminosae). -Can. J. Bot. 67(11): 3260–3294. https://doi.org/10.1139/b89-405 Small, E., Lassen, P., & Brookes, B.S. 1987: An expanded circumscription of Medicago (Leguminosae, Trifolieae) based on explosive flower tripping. -Willdenowia 16(2): 415–437. Soltis, D.E., & Soltis, P.S. 2000: Contributions of plant molecular systematics to studies of molecular evolution. -Plant Mol. Biol. 42(1): 45–75. https://doi.org/10.1023/A:1006380618351 Steele, K. P., & Wojciechowski, M. F. 2003: Phylogenetic analyses of tribes Trifolieae and Vicieae, based on sequences of the plastid gene, matK (Papilionoideae: Leguminosae). In B.B. Klitgaard & A. Bruneau (Eds.), Advances in legume systematics (Part 10, pp. 355–370). Royal Botanic Gardens, Kew. -Kew, Royal Botanic Gardens. Steele, K.P., Ickert-Bond, S.M., Zarre, S., & Wojciechowski, M.F. 2010: Phylogeny and character evolution in Medicago (Leguminosae): Evidence from analyses of plastid trnK/matK and nuclear GA3ox1 sequences. -Amer. J. Bot. 97(7): 1142–1155. https://doi.org/10.3732/ajb.1000009 Swofford, D.L. 2002: PAUP: Phylogenetic analysis using parsimony* (and other methods). Version 4. Sinauer Associates. -Sunderland, Sinauer Associates. Wojciechowski, M.F., Lavin, M., & Sanderson, M. J. 2003: A phylogeny of legumes based on sequences of the plastid matK gene. In Botany 2003: Aquatic and Wetland Plants: Wet and Wild, Abstracts (p. 99). Wojciechowski, M.F., Lavin, M., & Sanderson, M.J. 2004: A phylogeny of legumes (Leguminosae) based on analysis of the plastid matK gene resolves many well-supported subclades within the family. Am. J. Bot, 91(11): 1846–1862. https://doi.org/10.3732/ajb.91.11.1846 Yoder, J.B., Briskine, R., Mudge, J., Farmer, A., Paape, T., Steele, K., & Young, N. D. 2013: Phylogenomic analysis of Medicago species reveals clear species boundaries and evidence of reticulate evolution. -BMC Evol. Biol. 13, 131. https://doi.org/10.1186/1471-2148-13-131 Zareei, R., Small, E., Assadi, M., Mehregan, I. 2022: Genetic structure of Medicago sinskiae using microsatellite data reveals its fast expansion throughout western and southwestern Iran. -Collect. Bot. 41: e2002. https://doi.org//10.3989/collectbot.2022.v41.002 | ||
|
آمار تعداد مشاهده مقاله: 105 تعداد دریافت فایل اصل مقاله: 96 |
||