Abstract
This study investigated the spatial distribution of coastal foraminifera at four stations, identifying seven distinct species representing five genera and three families, with Ammonia beccarii caspica as the dominant species. The Bandar Gaz station exhibited the highest density, species richness, evenness, and Shannon diversity. Consequently, Bandar Gaz was selected for a controlled microcosm experiment examining benthic foraminiferal community responses to increased water temperature (24 °C, 27 °C, and 30 °C) over 60 days. While the highest total density and evenness were observed at 30 °C, the total number of species and Margalef and Shannon indices did not significantly differ among treatments. Temperature changes significantly altered community structure through shifts in species dominance. Ammonia species displayed resilience and increased dominance with higher temperatures, replacing other species. Elphidium advenum density decreased significantly at 30 °C, while Ammonia beccarri and Ammonia tepida increased in dominance with rising temperatures. These findings highlight temperature-driven alterations in foraminiferal assemblages, with implications for coastal ecosystem monitoring in the context of climate change.
Data availability
The datasets during and/or analyzed during the current study available from the corresponding author on reasonable request.
References
Maslin, M. Global Warming: A Very Short Introduction (Oxford University Press, 2008).
Venegas, R. M., Acevedo, J. & Treml, E. A. Three decades of ocean warming impacts on marine ecosystems: a review and perspective. Deep Sea Res. Part II. 212, 105318. https://doi.org/10.1016/j.dsr2.2023.105318 (2023).
Bagheri, H., Mahmudy Gharaei, M., Mousavi Harami, R. & Khanebad, M. Reconstruction of Caspian sea level changes using magnetic susceptibility during the last millennium. Iran. J. Mar. Sci. 48, 1452–1459 (2019).
Arpe, K., Molavi-Arabshahi, M. & Leroy, S. A. G. Wind variability over the Caspian Sea, its impact on Caspian seawater level and link with ENSO. Int. J. Climatol. 40, 6039–6054. https://doi.org/10.1002/joc.6564 (2020).
Lahijani, H. et al. Behravesh M. A note on the silent decline of the Caspian environment. Mar. Pollut. Bull. 205, 116551. https://doi.org/10.1016/j.marpolbul.2024.116551 (2024).
Roshan, G., Moghbel, M. & Grab, S. Modeling Caspian sea water level oscillations under different scenarios of increasing atmospheric carbon dioxide concentrations. J. Environ. Health Sci. Eng. 9, 24. https://doi.org/10.1186/1735-2746-9-24 (2012).
Azizpour, J. & Ghaffari, P. Low-frequency sea level changes in the Caspian sea: long-term and seasonal trends. Clim. Dyn. 61, 2753–2763. https://doi.org/10.1007/s00382-023-06715-9 (2023).
Taheri, M., Yazdani Foshtami, M., Manbohi, A. & Mira, M. Spatial distribution and effects of temperature rise on coastal free-living nematode community in the South Caspian sea. Mar. Pollut. Bull. 214, 117806. https://doi.org/10.1016/j.marpolbul.2025.117806 (2025).
Danovaro, R., Gambi, C. & Della Croce, N. Meiofauna response to organic matter enrichment in a mediterranean coastal lagoon. Hydrobiologia 445, 131–139 (2001).
Danovaro, R. et al. Deep-sea biodiversity in the mediterranean sea: which is the role of food availability? Deep sea research part I. Oceanogr. Res. Papers. 51, 1529–1542. https://doi.org/10.1016/j.dsr.2004.07.015 (2004).
Mojtahid, M., Nercessian, A., Debenay, J. P. & Buoncristiani, J. F. Modern benthic foraminifera distribution in the Arcachon Lagoon, French Atlantic coast: environmental significance. Estuar. Coast. Shelf Sci. 81, 81–93. https://doi.org/10.1016/j.ecss.2008.10.001 (2009).
Bouchet, V. M. P., Koho, K. A., McCarthy, F. M. G., Martin, R. A. & Martin, J. B. Benthic foraminiferal distribution in Bonne Bay, newfoundland: assessing environmental gradients and proxy potential. Mar. Micropaleontol. 96, 38–54. https://doi.org/10.1016/j.marmicro.2012.08.001 (2012).
Dolven, J. K., McCarthy, F. M. G., Collins, E. S., Chmura, G. L. & Patterson, R. T. Salt-marsh foraminifera as proxies for relative sea-level change: statistical (transfer function) vs. indicator zone approaches. J. Quat. Sci. 28, 369–380. https://doi.org/10.1002/jqs.2624 (2013).
Polovodova Asteman, I. & Nordberg, K. The influence of different ecological factors on benthic foraminiferal distribution in the Himmerfjärden Estuary, Northern Baltic sea. J. Micropalaeontol. 32, 173–187. https://doi.org/10.1144/jmpaleo2012-015 (2013).
Duffield, A., Patterson, R. T., Reinhardt, E. G., McKillop, I. K. & Roe, H. M. Distribution of intertidal foraminifera in the Pachena river salt marsh, British Columbia, canada: assessing their utility as sea-level indicators. Mar. Micropaleontol. 119, 22–36. https://doi.org/10.1016/j.marmicro.2015.06.001 (2015).
Nordberg, K., Eichler, P. P. B., Volkov, I. A. & Asteman, I. P. Modern benthic foraminifera in the sea of azov: distribution in relation to salinity, organic matter, and pollution. J. Foraminifer. Res. 47, 167–181. https://doi.org/10.2113/gsjfr.47.2.167 (2017).
Altenbach, A. V. Short term processes influencing the vertical distribution of foraminifera in a Sublittoral habitat. Mar. Micropaleontol. 20, 21–29. https://doi.org/10.1016/0377-8398(92)90024-9 (1992).
Alve, E. & Bernhard, J. M. Vertical migratory response of benthic foraminifera to controlled oxygen perturbations in an experimental marine ecosystem. Mar. Ecol. Prog. Ser. 116, 137–151 (1995).
Havach, S., Frontalini, F. & Coccioni, R. Benthic foraminifera as indicators of heavy-metal pollution in the lagoon of Venice, Italy. J. Foraminifer. Res. 31, 194–201. https://doi.org/10.2113/0310194 (2001).
Le Cadre, V. & Debenay, J. P. Morphological abnormalities in foraminifera: one tool for pollution assessment? Mar. Micropaleontol. 61, 178–196. https://doi.org/10.1016/j.marmicro.2006.08.003 (2006).
Elderfield, H., Vautravers, M. & Cooper, M. The relationship between benthic foraminiferal magnesium and seawater temperature. Earth Planet. Sci. Lett. 250, 208–223. https://doi.org/10.1016/j.epsl.2006.07.034 (2006).
Wit, J. C., Filipsson, H. L. & Platzman, D. J. Interactive effects of hypoxia and pH on the benthic foraminifera Ammonia tepida: growth rate, survival, and shell characteristics. Biogeosciences 10, 7479–7496. https://doi.org/10.5194/bg-10-7479-2013 (2013).
Schönfeld, J. Living Foraminifera in Shelf Seas (Cambridge University Press, 2018).
Dong, B., Kong, L., Guo, W., Zhan, B. & Li, X. Benthic foraminiferal response to environmental changes in the Pearl river Estuary, China. J. Mar. Syst. 195, 1–13. https://doi.org/10.1016/j.jmarsys.2019.03.001 (2019).
Goldstein, S. T. & Alve, E. Effect of increased bottom-water temperature on foraminiferal assemblages in a Norwegian Fjord. J. Foraminifer. Res. 41, 1–11. https://doi.org/10.2113/gsjfr.41.1.99 (2011).
Samant, R. & Prange, M. Climate-driven 21st century Caspian sea level decline estimated from CMIP6 projections. Commun. Earth Environ. 4, 357. https://doi.org/10.1038/s43247-023-01017-8 (2023).
Taheri, M., Darvish Bastami, K. & Yazdani Foshtomi, M. Meiofauna-sediment relationships in shallow water of the south Caspian Sea. In: INOC-International Congress “Estuaries & Coastal Protected Areas” ECPA 2014, 04–06 November 2014, Izmir–Turkey (2014).
Darvish Bastami, K. et al. Molamohyeddin N. Nematode community structure in relation to metals in the Southern of Caspian sea. Acta Oceanol. Sin. 36, 1–8. https://doi.org/10.1007/s13131-017-1114-z (2017).
Taheri, M., Darvish Bastami, K. & Yazdani Foshtomi, M. October. The role of the sediment conditions in shaping meiofauna spatial distribution in the shallow water of the south Caspian Sea. In First International Conference on Oceanography for West Asia, 30–31 (2017).
Bernhard, J. M. Distinguishing live from dead foraminifera: methods review and proper applications. Micropaleontology 46, 38–46 (2000).
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).
Folk, R. L. Petrology of Sedimentary Rocks (Hemphill Publishing Company, 1980).
APHA. Standard Methods for the Examination of Water and Wastewater (American Public Health Association, 2005).
Taheri, M., Braeckman, U., Vincx, M. & Vanaverbeke, J. Effect of short-term hypoxia on marine nematode community structure and vertical distribution pattern in three different sediment types of the North sea. Mar. Environ. Res. 99, 149–159. https://doi.org/10.1016/j.marenvres.2014.04.008 (2014b).
Taheri, M., Yazdani Foshtomi, M., Hamzeh, M. A., Manbohi, A. & Rahnama Haratbar, R. Effects of discarded garbage bags on intertidal free living nematode community. Aquat. Ecol. 58, 853–863. https://doi.org/10.1007/s10452-024-10110-9 (2024).
Gingold, R., Moens, T. & Rocha-Olivares, A. Assessing the response of nematode communities to climate change-driven warming: A microcosm experiment. PLoS ONE 8, e66653. https://doi.org/10.1371/journal.pone.0066653 (2013).
Lei, Y., Li, T., Jian, Z. & Nigam, R. Taxonomy and distribution of benthic foraminifera in an intertidal zone of the yellow Sea, PR china: correlations with sediment temperature and salinity. Mar. Micropaleontol. 133, 1–20. https://doi.org/10.1016/j.marmicro.2017.04.005 (2017a).
Lei, Y. L., Stumm, K., Wickham, S. A. & Berninger, U. G. Distributions and biomass of benthic ciliates, foraminifera and amoeboid protists in marine, brackish, and freshwater sediments. J. Eukaryot. Microbiol. 61, 493–508. https://doi.org/10.1111/jeu.12129 (2014).
Anderson, M. J., Gorley, R. N. & Clarke, K. R. Guide to Software and Statistical Methods (PRIMER-E Ltd., 2008).
Gooday, A. J. Benthic foraminifera (Protista) as tools in deep-water palaeoceanography: environmental influences on faunal characteristics. Adv. Mar. Biol. 46, 1–90. https://doi.org/10.1016/S0065-2881(03)46002-1 (2003).
Murray, J. W. Ecology and Applications of Benthic Foraminifera (Cambridge University Press, 2006).
Dubey, R., Saraswat, R. & Nigam, R. Mudbank off Alleppey, India: a Bane for foraminifera but not so for carbon burial. Sci. Total Environ. 634, 459–470. https://doi.org/10.1016/j.scitotenv.2018.03.369 (2018).
Lei, Y., Stumm, K., Wickham, S. A. & Berninger, U. G. Distributions and biomass of benthic ciliates, foraminifera and amoeboid protists in marine, brackish, and freshwater sediments. J. Eukaryot. Microbiol. 61, 493–508. https://doi.org/10.1111/jeu.12129 (2014).
Hale, R., Calosi, P., McNeill, L., Mieszkowska, N. & Widdicombe, S. Predicted levels of future ocean acidification and temperature rise could alter community structure and biodiversity in marine benthic communities. Oikos 120, 661–674. https://doi.org/10.1111/j.1600-0706.2010.19469.x (2011).
Meadows, A. S., Ingels, J., Widdicombe, S., Hale, R. & Rundle, S. D. Effects of elevated CO2 and temperature on an intertidal meiobenthic community. J. Exp. Mar. Biol. Ecol. 469, 44–56. https://doi.org/10.1016/j.jembe.2015.04.001 (2015).
Ingels, J. et al. Short-term CO2 exposure and temperature rise effects on metazoan meiofauna and free-living nematodes in sandy and muddy sediments: results from a flume experiment. J. Exp. Mar. Biol. Ecol. 502, 211–226. https://doi.org/10.1016/j.jembe.2018.03.009 (2018).
Vafeiadou, A. M. et al. Temperature-driven dynamics: unraveling the impact of climate change on cryptic species interactions within the Litoditis marina complex. PeerJ 12, e17324. https://doi.org/10.7717/peerj.17324 (2024).
Moens, T. & Vincx, M. Temperature, salinity and food thresholds in two brackish-water bacterivorous nematode species: assessing niches from food absorption and respiration experiments. J. Exp. Mar. Biol. Ecol. 243, 137–154. https://doi.org/10.1016/S0022-0981(99)00114-8 (2000).
Acknowledgements
We thank the anonymous reviewers for their constructive comments that improved the quality of the manuscript.
Funding
This work is based upon research founded by Iranian National Science Foundation (INSF) under project NO. 40266240.
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Hossein Bagheri: writing—original draft preparation, methodology, writing-review and editing. Mehrshad Taheri: methodology, software, writing-review and editing, visualization. All authors have read and agreed to the published version of the manuscript.
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Bagheri, H., Taheri, M. Responses of South Caspian coastal foraminifera to warming: spatial patterns and assemblage shifts.
Sci Rep (2026). https://doi.org/10.1038/s41598-026-38207-1
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DOI: https://doi.org/10.1038/s41598-026-38207-1
Keywords
- Caspian sea
- Temperature
- Foraminifera
- Sediments
Source: Ecology - nature.com
