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Checklist of edaphic endemic plants in northeastern Iran: diversity, distribution patterns, and conservation

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Abstract

Soil variability is a key driver of plant diversity and endemism, yet the ecological roles of specific soil types remain poorly understood. In the Khorassan-Kopet Dagh floristic province of northeastern Iran and southern Turkmenistan, the coexistence of gypsum, calcareous, and serpentine soils provides a unique setting to explore these relationships. We carried out extensive field surveys and soil analyses across nine habitats to refine the classification of edaphic endemic plants and reassess their conservation status. Using elemental analyses and standard soil tests, we characterized chemical properties such as carbonates, organic matter, gypsum, pH, and conductivity. Our results documented 71 plant taxa from 30 families, including 35 regional endemics and six Iranian endemics. Based on soil affinity, species were grouped as gypsophiles (3%), gypsovags (20%), obligate calcicoles (25%), and facultative calcicoles (52%). Several species required reclassification from gypsum to calcareous specialists. Hemicryptophytes (45%) and chamaephytes (28%) were the dominant groups, especially among calcicoles, with the highest species richness found at elevations between 1,500 and 2,300 m. Updated distribution data revealed that facultative calcicoles are largely secure, while obligates remain vulnerable. This study offers the first integrated account of edaphic endemism in the region, providing essential insights for biodiversity monitoring and conservation under rising anthropogenic and climatic pressures.

Data availability

The datasets generated during this study are available from the corresponding author on reasonable request.

References

  1. Hulshof, C. M. & Spasojevic, M. J. The edaphic control of plant diversity. Glob Ecol. Biogeogr. 29, 1634–1650 (2020).

    Google Scholar 

  2. Cera, A. et al. Gypsum endemics accumulate excess nutrients in leaves as a potential constitutive strategy to grow in grazed extreme soils. Physiol. Plant. 174, e13738 (2022).

    Google Scholar 

  3. Rajakaruna, N. Lessons on Evolution from the Study of Edaphic Specialization. Bot. Rev. 84, 39–78 (2018).

    Google Scholar 

  4. Jenny, H. Factors of Soil Formation: A System of Quantitative Pedology (Courier Corporation, 1994).

  5. Mota, J. F., Garrido-Becerra, J. A., Merlo, M. E., Medina-Cazorla, J. M. & Sánchez-Gómez, P. The Edaphism: Gypsum, Dolomite and Serpentine Flora and Vegetation. in The Vegetation of the Iberian Peninsula (ed. Loidi, J.) vol. 13 277–354 (Springer International Publishing, 2017).

  6. Schimper, A. F. W. Plant-Geography upon a Physiological Basis (Clarendon, 1902).

  7. Noroozi, J. et al. Endemic diversity and distribution of the Iranian vascular flora across phytogeographical regions, biodiversity hotspots and areas of endemism. Sci. Rep. 9, 12991 (2019).

    Google Scholar 

  8. Kruckeberg, A. R. & Rabinowitz, D. Biological aspects of endemism in higher plants. Annu. Rev. Ecol. Syst. 447–479. https://doi.org/10.1146/annurev.es.16.110185.002311 (1985).

  9. Rivas-Goday, S. Flora serpentinícola española, nota primera (Edafismos endémicos del Reino de Granada). in 35 297–304 (1969).

  10. Shiba, M., Tate, T. & Fukuda, T. Serpentine Adaptation of Ligustrum japonicum Thunb.(Oleaceae) Based on Morphological and Anatomical Approaches. Int. J. Biol. 14, 10–18 (2022).

    Google Scholar 

  11. Wahba, M. M., Labib, F. & Zaghloul, A. Management of Calcareous Soils in Arid Region. Environ Model 2 (2019).

  12. Taalab, A., Ageeb, G., Siam, H. S. & Mahmoud, S. A. Some Characteristics of Calcareous soils. A review. Middle East. J. Agric. Res. 8, 96–105 (2019).

    Google Scholar 

  13. Cera, A., Montserrat-Martí, G., Ferrio, J. P., Drenovsky, R. E. & Palacio, S. Gypsum-exclusive plants accumulate more leaf S than non-exclusive species both in and off gypsum. Environ. Exp. Bot. 182, 104294 (2021).

    Google Scholar 

  14. Herrero, J. & Porta, J. The terminology and the concepts of gypsum-rich soils. Geoderma 96, 47–61 (2000).

    Google Scholar 

  15. Agriculture Organization of the United Nations. Soil Resources & Conservation Service. Management of Gypsiferous Soils. vol. 62 (Food & Agriculture Org., (1990).

  16. Gankin, R. & Major, J. Arctostaphylos myrtifolia, its biology and relationship to the problem of endemism. Ecology 45, 792–808 (1964).

    Google Scholar 

  17. Palacio, S. et al. Plants Living on Gypsum: Beyond the Specialist Model. Ann. Bot. 99, 333–343 (2007).

    Google Scholar 

  18. Zohlen, A. & Tyler, G. Soluble Inorganic Tissue Phosphorus and Calcicole–Calcifuge Behaviour of Plants. Ann. Bot. 94, 427–432 (2004).

    Google Scholar 

  19. Gigon, A. Springer,. A hierarchic approach in causal ecosystem analysis the calcifuge-calcicole problem in alpine grasslands. in Potentials and Limitations of ecosystem analysis 228–244 (1987).

  20. Clymo, R. S. An Experimental Approach to Part of the Calcicole Problem. J. Ecol. 50, 707 (1962).

    Google Scholar 

  21. Parmar, R. K., Arya, V., Gill, A. K. & Thakur, V. Reclaiming Calcicoles: New Insights into Lime Lovers. J. Pharmacol. Pharmacother. 16, 25–37 (2025).

    Google Scholar 

  22. Rorison, I. H. Some Experimental Aspects of the Calcicole-Calcifuge Problem: I. The Effects of Competition and Mineral Nutrition Upon Seedling Growth in the Field. J. Ecol. 48, 585 (1960).

    Google Scholar 

  23. Simpson, J. F. H. A Chalk Flora on the Lower Greensand: Its Use in Interpreting the Calcicole Habit. J. Ecol. 26, 218 (1938).

    Google Scholar 

  24. Cross, A. T. & Lambers, H. Calcicole–calcifuge plant strategies limit restoration potential in a regional semi-arid flora. Ecol. Evol. 11, 6941–6961 (2021).

    Google Scholar 

  25. Davis, S. D., Heywood, V. H. & Hamilton, A. C. Centres of Plant Diversity: Europe, Africa, South West Asia, and the Middle East Vol. 1 (World Conservation Union, 1994).

  26. Zohary, M. Geobotanical Foundations of the Middle East (Gustav Fischer, 1973).

  27. Noroozi, J. et al. Hotspots within a global biodiversity hotspot-areas of endemism are associated with high mountain ranges. Sci. Rep. 8, 10345 (2018).

    Google Scholar 

  28. Manafzadeh, S., Salvo, G. & Conti, E. A tale of migrations from east to west: The Irano-Turanian floristic region as a source of Mediterranean xerophytes. J. Biogeogr. 41, 366–379 (2014).

    Google Scholar 

  29. Memariani, F., Akhani, H. & Joharchi, M. R. Endemic plants of Khorassan-Kopet Dagh floristic province in Irano-Turanian region: diversity, distribution patterns and conservation status. Phytotaxa 249, 31 (2016).

    Google Scholar 

  30. Memariani, F. The Khorassan-Kopet Dagh Mountains. in Plant biogeography and vegetation of high mountains of central and south-west Asia 93–116 (Springer, 2020).

  31. Afshar-Harb, A. Geology of Kopet Dagh. Treatise Geol. Iran. 11, 1–275 (1994).

    Google Scholar 

  32. Djamali, M. et al. Application of the Global Bioclimatic Classification to Iran: implications for understanding the modern vegetation and biogeography. Ecol. Mediterr. 37, 91–114 (2011).

    Google Scholar 

  33. Djamali, M. et al. Ecological implications of Cousinia Cass. (Asteraceae) persistence through the last two glacial–interglacial cycles in the continental Middle East for the Irano-Turanian flora. Rev. Palaeobot Palynol. 172, 10–20 (2012).

    Google Scholar 

  34. IUCN Standards and Petitions Committee. Guidelines for using the IUCN red list categories and criteria. Version 16. Prep Stand. Petitions Comm (2024). https://www.iucnredlist.org/documents/RedListGuidelines.pdf

  35. Rechinger, K. H. Flora Iranica. vols 1–181 (Akademische Druck- u. Verlagsanstalt, Graz; vol. 175, Akademische Verlagsgesellschaft, Salzburg; vol. 176–181 (Verlag des Naturhistorischen Museums, 1963).

  36. Assadi, M., Maassoumi, A. A., Khatamsaz, M. & Mozaffarian, V. Flora of Iran. vols 1–184 (Research Institute of Forests and Rangelands Publications, Tehran (in Persian). (1988).

  37. Jones, J. Soil Analysis Handbook of Reference Methods (CRC, 2018).

  38. Artieda, O., Herrero, J. & Drohan, P. Refinement of the differential water loss method for gypsum determination in soils. Soil. Sci. Soc. Am. J. 70, 1932–1935 (2006).

    Google Scholar 

  39. Heiri, O., Lotter, A. F. & Lemcke, G. Loss on ignition as a method for estimating organic and carbonate content in sediments: reproducibility and comparability of results. J. Paleolimnol. 25, 101–110 (2001).

    Google Scholar 

  40. R Core Team. R: A Language and Environment for Statistical Computing (R Foundation for Statistical Computing, 2025).

  41. Wickham, H. Ggplot2: Elegant Graphics for Data Analysis (Springer, 2016).

  42. Atashgahi, Z., Ejtehadi, H., Mesdaghi, M. & Ghassemzadeh, F. Plant diversity of the Heydari Wildlife Refuge in northeastern Iran, with a checklist of vascular plants. Phytotaxa 340, 101 (2018).

    Google Scholar 

  43. Ghahremaninejad, F. Astragalus hekmat-safaviae (Fabaceae), a new species from Iran (2005).

  44. Hassanpour, S. et al. Molecular and morphological data reveal a new species of Sclerorhachis (Compositae, Anthemideae) and the reassessment of another species of the genus. Plant. Syst. Evol. 309, 10 (2023).

    Google Scholar 

  45. Memariani, F., Joharchi, M. R., Ejtehadi, H. & Emadzade, K. Contributions to the flora and vegetation of Binalood mountain range, NE Iran: Floristic and chorological studies in Fereizi region. J. Cell. Mol. Res. 1, 1–18 (2009).

    Google Scholar 

  46. Memariani, F., Joharchi, M. R. & Akhani, H. Plant diversity of Ghorkhod Protected Area, NE Iran. Phytotaxa 249, 118 (2016).

    Google Scholar 

  47. Moazzeni, H. et al. A taxonomic revision of the genus Aethionema s.l. (Brassicaceae) in Iran. Phytotaxa 356, 241 (2018).

    Google Scholar 

  48. Bachman, S., Moat, J., Hill, A. W., De La Torre, J. & Scott, B. Supporting Red List threat assessments with GeoCAT: geospatial conservation assessment tool. ZooKeys 117 https://doi.org/10.3897/zookeys.150.2109 (2011).

  49. Asri, Y., Sefidkon, F. & Rabie, M. Investigating the relationship between ecological parameters and vegetative and phytochemical indices of Seseli staurophyllum Rech.f. populations. J. Plant. Ecosyst. Conserv. 12, 174–194 (2024).

    Google Scholar 

  50. Zepner, L., Karrasch, P., Wiemann, F. & Bernard, L. ClimateCharts. net–an interactive climate analysis web platform. Int. J. Digit. Earth. 14, 338–356 (2021).

    Google Scholar 

  51. Pérez-García, F. J. et al. A first inventory of gypsum flora in the Palearctic and Australia. Mediterr. Bot. 39, 35–49 (2018).

    Google Scholar 

  52. Bahalkeh, K., Abedi, M., Palacio, S., Luzuriaga, A. L. & Escoderu, A. Study of Gypsum Ecosystems Species Adaptation in of North East of Iran (Case study: Khorasan Razavi Province). J. Plant. Ecosyst. Conserv. 13, 1–12 (2024).

    Google Scholar 

  53. Eftekhari, T., Assadi, M., Mahmoudi, S. & Dadkhahipour, K. Flora of the gypsiferous area in the Robat-Sefid (Khorasan province). Pajouhesh Sazandegi. 56 & 57, 81–94 (2003).

    Google Scholar 

  54. Memariani, F., Zarrinpour, V. & Akhani, H. A review of plant diversity, vegetation, and phytogeography of the Khorassan-Kopet Dagh floristic province in the Irano-Turanian region (northeastern Iran–southern Turkmenistan). Phytotaxa 249, 8–30 (2016).

    Google Scholar 

  55. Mota, J. F. et al. Shipwrecked on the Rock, or Not Quite: Gypsophytes and Edaphic Islands. Plants 13, 970 (2024).

    Google Scholar 

  56. Casby-Horton, S., Herrero, J. & Rolong, N. A. Gypsum soils—Their morphology, classification, function, and landscapes. Adv. Agron. 130, 231–290 (2015).

    Google Scholar 

  57. Escudero, A., Palacio, S., Maestre, F. T. & Luzuriaga, A. L. Plant life on gypsum: a review of its multiple facets. Biol. Rev. 90, 1–18 (2015).

    Google Scholar 

  58. Tazikeh, H., Khormali, F., Amini, A., Motlagh, M. B. & Ayoubi, S. Soil-parent material relationship in a mountainous arid area of Kopet Dagh basin, North East Iran. Catena 152, 252–267 (2017).

    Google Scholar 

  59. Memariani, F. Khorassan-Kopet Dagh Floristic Province (NE Iran, S Turkmenistan): an edaphic endemism hotspot in Irano-Turanian Region (2018).

  60. Palacio, S. et al. Recent and ancient evolutionary events shaped plant elemental composition of edaphic endemics: a phylogeny-wide analysis of Iberian gypsum plants. New. Phytol. 235, 2406–2423 (2022).

    Google Scholar 

  61. Alghamdi, S. A., Al-Ghamdi, F. A., El-Zohri, M. & Al-Ghamdi, A. A. M. Modifying of calcareous soil with some acidifying materials and its effect on Helianthus annuus (L.) growth. Saudi J. Biol. Sci. 30, 103568 (2023).

    Google Scholar 

  62. Bolan, N. et al. Distribution, characteristics and management of calcareous soils. Adv. Agron. 182, 81–130 (2023).

    Google Scholar 

  63. Chytrý, M. et al. Floristic diversity of an eastern Mediterranean dwarf shrubland: the importance of soil pH. J. Veg. Sci. 21, 1125–1137 (2010).

    Google Scholar 

  64. Mota, J. F. et al. Conceptual baseline for a global checklist of gypsophytes. Lazaroa 37, 7–30 (2016).

    Google Scholar 

  65. Sennikov, A. N. The plant world of the Khorassan-Kopet Dagh Floristic Province: A tribute to Eskandar Firouz. Phytotaxa 249, 5–7 (2016).

    Google Scholar 

  66. Knapp, H. D. On the distribution of the genus Cousinia (Compositae). Plant. Syst. Evol. 155, 15–25 (1987).

    Google Scholar 

  67. Mehregan, I. & Kadereit, J. W. taxonomic revision of Cousinia sect. Cynaroideae (Asteraceae, Cardueae). Willdenowia 38, 293–362 (2008).

    Google Scholar 

  68. Rajakaruna, N. & Boyd, R. S. Serpentine soils. Oxf. Bibliogr Ecol. https://doi.org/10.1093/OBO/9780199830060-0055 (2014).

    Google Scholar 

  69. Rozentsvet, O. A., Bogdanova, E. S., Tabalenkova, G. N. & Rozina, S. N. Morphological, Physiological, and Biochemical Characteristics of Adaptation of Calcephytes of the Genus Hedysarum. Contemp. Probl. Ecol. 14, 465–471 (2021).

  70. Persson, H. & Baitulin, I. O. Plant root systems and natural vegetation. in (Sv. växtgeografiska sällsk., (1996).

  71. Rudov, A. et al. Ecohydrological niche segregation among desert shrubs in a gypsum-calcareous formation, north-western Iran. Plant. Ecol. Divers. 16, 61–75 (2023).

    Google Scholar 

  72. Kurbanov, D. Flora of Kopetdagh. in Biogeography and ecology of Turkmenistan 105–128 (Springer, (1994).

  73. Atashgahi, Z., Erfanian, M. B., Moazzeni, H., Shemirani, G. & Pirani, A. Endemic cushions of the Khorassan-Kopet Dagh floristic province show differential responses to future climate change. Sci. Rep. 15, 16046 (2025).

    Google Scholar 

  74. Corlett, R. T. & Tomlinson, K. W. Climate Change and Edaphic Specialists: Irresistible Force Meets Immovable Object? Trends Ecol. Evol. 35, 367–376 (2020).

    Google Scholar 

  75. Berisha, N., Millaku, F., Krasniqi, E. & Gashi, B. Rare and Endangered Geophyte Plant Species in Serpentine of Kosovo. Ecol Balk 6 (2014).

  76. Chiarucci, A. & Baker, A. J. Advances in the ecology of serpentine soils. Plant. Soil. 293, 1–2 (2007).

    Google Scholar 

  77. Harrison, S. & Inouye, B. D. High β diversity in the flora of Californian serpentine’islands’. Biodivers. Conserv. 11, 1869–1876 (2002).

    Google Scholar 

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Acknowledgements

This work was conducted as part of the Ph.D. dissertation of the first author, supported by Grant number 3/60075, from the Office of the Vice-President for Research and Technology of Ferdowsi University of Mashhad and partly by European Union’s Horizon 2020 (H2020-MSCA RISE-777803). The authors wish to express deep gratitude to Mohammad Reza Joharchi in Herbarium of Ferdowsi University of Mashhad (FUMH) for helping with species determination. Thanks to Elaheh Tabasi for help with fieldwork and data collection.

Funding

This work was supported by Grant number 3/60075 from the Office of the Vice-President for Research and Technology of Ferdowsi University of Mashhad for H.R., and partly by the European Union’s Horizon 2020 (H2020-MSCA RISE-777803) for F.M.

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Authors and Affiliations

  1. Quantitative Plant Ecology and Biodiversity Research Lab, Department of Biology, Faculty of Sciences, Ferdowsi University of Mashhad, Mashhad, Iran

    Hossein Rahmaninia & Hamid Ejtehadi

  2. Department of Range and Watershed Management, Faculty of Natural Resources and Environment, Ferdowsi University of Mashhad, Mashhad, Iran

    Farshid Memariani

  3. Herbarium FUMH, Ferdowsi University of Mashhad, Mashhad, Iran

    Farshid Memariani

  4. Department of Range Management, Faculty of Natural Resources and Marine Sciences, Tarbiat Modares University, Noor, Iran

    Mehdi Abedi

  5. Instituto Pirenaico de Ecología (IPE-CSIC), Av. Nuestra Señora de la Victoria, 16, Jaca, Huesca, 22700, Spain

    Sara Palacio

Authors

  1. Hossein Rahmaninia
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  2. Hamid Ejtehadi
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  3. Farshid Memariani
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  4. Mehdi Abedi
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Contributions

Author contributions H.R., H.E., and F.M. conceived and designed the study; H.R. and F.M. collected and analyzed the data and H.R. wrote the first draft. H.E., F.M., M.A., and S.P. helped with revising drafts of the paper. All authors read and approved the final draft.

Corresponding authors

Correspondence to
Hamid Ejtehadi or Farshid Memariani.

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The authors declare no competing interests.

Ethical approval

All field studies and collection of wild plant material were performed in accordance with relevant institutional, national, and international guidelines under formal permission from the Iranian Natural Resources and Watershed Management Organization. Voucher specimens were deposited at the FUMH herbarium under accession numbers included in the manuscript.

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Rahmaninia, H., Ejtehadi, H., Memariani, F. et al. Checklist of edaphic endemic plants in northeastern Iran: diversity, distribution patterns, and conservation.
Sci Rep (2026). https://doi.org/10.1038/s41598-026-48907-3

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Keywords

  • Calciphytes
  • Edaphism
  • Gypsophytes
  • Irano-Turanian
  • Khorassan-Kopet Dagh

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