全球变暖背景下海洋浮游生物群落长期变化

张武昌; 赵苑; 刘红斌; 孙军

doi:10.3799/dqkx.2025.163

全球变暖背景下海洋浮游生物群落长期变化

doi: 10.3799/dqkx.2025.163

张武昌^{1, 2, 3, ,},
赵苑^{1, 2, 3, , ,},
刘红斌^{4, 6},
孙军⁵

1.
中国科学院海洋研究所, 海洋生态与环境科学实验室, 山东青岛 266000
2.
青岛海洋科技中心, 海洋生态与环境科学功能实验室, 山东青岛 266237
3.
中国科学院海洋大科学研究中心, 山东青岛 266000
4.
香港科技大学海洋科学系, 香港 999077
5.
中国地质大学(武汉)地质微生物与环境全国重点实验室, 湖北武汉 430078
6.
中国科学院‒香港科技大学三亚海洋科学综合(联合)实验室, 海南三亚 572000

基金项目:

国家重点研发项目 2022YFC3105301

国家自然科学基金项目 42476143

国家自然科学基金项目 42076139

详细信息

作者简介:
张武昌（1973-），男，研究员，博士，从事海洋浮游生物生态学研究. ORCID：0000-0001-7534-8368. E-mail：wuchangzhang@qdio.ac.cn

通讯作者:
赵苑，ORCID：0000-0002-8892-2100. E-mail：yuanzhao@qdio.ac.cn

中图分类号: P735
计量
- 文章访问数: 183
- HTML全文浏览量: 25
- PDF下载量: 18
- 被引次数: 0
出版历程
- 收稿日期: 2025-03-14
- 刊出日期: 2025-11-25

Long-Term Changes in Plankton Communities in Context of Global Warming

Zhang Wuchang^{1, 2, 3
,
,},
Zhao Yuan^{1, 2, 3
, ,
,},
Liu Hongbin^{4, 6},
Sun Jun⁵

1.
Laboratory of Marine Ecology and Environmental Sciences, Institute of Oceanology, Chinese Academy of Sciences, Qingdao 266000, China
2.
Laboratory for Marine Ecology and Environmental Science, Qingdao Marine Science and Technology Center, Qingdao 266237, China
3.
Center for Ocean Mega-Science, Chinese Academy of Sciences, Qingdao 266000, China
4.
Department of Ocean Science, The Hong Kong University of Science and Technology, Hongkong 999077, China
5.
State Key Laboratory of Geomicrobiology and Environmental Changes, China University of Geosciences (Wuhan), Wuhan 430078, China
6.
CAS-HKUST Sanya Joint Laboratory of Marine Science Research, Sanya 572000, China

摘要

摘要: 浮游生物是海洋生态系统的基础，其群落结构和物候变化影响生态功能和生物地理格局.全球变暖驱动海表升温，导致浮游生物分布区极向移动，其中分布区前缘移动较明显，核心和后缘变化较小.升温还促使部分物种出现春季现象提前、秋季现象延后的物候变化.不同浮游生物的分布区和物候变化幅度存在差异，影响群落边缘区的结构，但群落核心优势种类尚无显著变化.若未来海洋环流格局保持稳定，热带至亚极区的浮游生物群落核心发生剧变风险较低，极地群落优势物种在近百年内也难有变化.我国海区长期系统观测不足，关键海域数据缺乏.综合极地与近海证据，本文预测在升温不超过2 ℃的情形下，我国近海浮游生物群落核心区位置可能发生调整，但其结构实质性改变的风险较低.
- 浮游生物 /
- 海洋变暖 /
- 生物地理分布 /
- 群落 /
- 物候 /
- 气候变化 /
- 海洋学
Abstract: Plankton form the foundation of marine ecosystems, and changes in their community structure and phenology affect ecosystem function and biogeographical patterns. Global warming drives sea surface temperature increases, resulting in poleward shifts of plankton distribution, with the leading edge advancing more noticeably while the core and trailing edge remain relatively stable. Warming also causes phenological shifts in some species, such as earlier spring events and delayed autumn events. The extent of distribution and phenological changes varies among plankton groups, mainly affecting community structure at the margins, while no significant changes have been observed in the dominant species of community cores. If future ocean circulation patterns remain stable, the risk of major shifts in plankton community cores from the tropics to subarctic regions is low, and dominant species in polar communities are also unlikely to change substantially in the next century. Long-term systematic observation in Chinese seas is still insufficient, and key regions lack critical data. Synthesizing evidence from polar and coastal regions, this study predicts that if warming does not exceed 2 ℃, the positions of community cores in China's coastal plankton communities may shift, but the risk of substantial structural change remains low.
- plankton /
- ocean warming /
- biogeographic distribution /
- communities /
- phenology /
- climate change /
- oceanography

HTML全文

图 1 全球海洋浮游生物群落研究的代表性站区与海域

五角星为各研究选取的典型海区，阴影表示已开展CPR长期采样的监测区域（据本文引用文献绘制）

Fig. 1. Representative study sites and regions of marine planktonic community research worldwide

下载: 全尺寸图片幻灯片

图 2 生物的热性能曲线模式（改绘自Kefford et al., 2022）

Fig. 2. Schematic diagram of thermal performance curve of organisms (adapted from Kefford et al., 2022)

下载: 全尺寸图片幻灯片

图 3 目前和未来全球不同纬度陆地和海洋生物的温度安全裕度（改绘自Pinsky et al., 2019）

Fig. 3. Current and future global thermal safety margins for terrestrial and marine biota at different latitudes (adapted from Pinsky et al., 2019)

下载: 全尺寸图片幻灯片

图 4 2014—2020年间亚北极号角虫（Salpingella subarctica）分布区前缘的扩张（改绘自Wang et al., 2022a）

Fig. 4. Expansion of the leading edge of tintinnid Salpingella subarctica between 2014 and 2020 (adapted from Wang et al., 2022a)

下载: 全尺寸图片幻灯片

图 5 分布区向极移动和物候变化对浮游生物群集的影响

Fig. 5. Effects of poleward shifts in distribution areas and phenology changes on plankton assemblages

下载: 全尺寸图片幻灯片

参考文献(131)

Aarflot, J. M., Skjoldal, H. R., Dalpadado, P., et al., 2018. Contribution of Calanus Species to the Mesozooplankton Biomass in the Barents Sea. ICES Journal of Marine Science, 75(7): 2342-2354. https://doi.org/10.1093/icesjms/fsx221
Aberle, N., Bauer, B., Lewandowska, A., et al., 2012. Warming Induces Shifts in Microzooplankton Phenology and Reduces Time-Lags between Phytoplankton and Protozoan Production. Marine Biology, 159(11): 2441-2453. https://doi.org/10.1007/s00227-012-1947-0
Alcaraz, M., Felipe, J., Grote, U., et al., 2014. Life in a Warming Ocean: Thermal Thresholds and Metabolic Balance of Arctic Zooplankton. Journal of Plankton Research, 36(1): 3-10. https://doi.org/10.1093/plankt/fbt111
Atkinson, A., Hill, S. L., Pakhomov, E. A., et al., 2019. Krill (Euphausia Superba) Distribution Contracts Southward during Rapid Regional Warming. Nature Climate Change, 9(2): 142-147. https://doi.org/10.1038/s41558-018-0370-z
Atkinson, A., Hill, S. L., Reiss, C. S., et al., 2022. Stepping Stones towards Antarctica: Switch to Southern Spawning Grounds Explains an Abrupt Range Shift in Krill. Global Change Biology, 28(4): 1359-1375. https://doi.org/10.1111/gcb.16009
Atkinson, A., Siegel, V., Pakhomov, E., et al., 2004. Long-Term Decline in Krill Stock and Increase in Salps within the Southern Ocean. Nature, 432(7013): 100-103. https://doi.org/10.1038/nature02996
Balch, W. M., Gordon, H. R., Bowler, B. C., et al., 2005. Calcium Carbonate Measurements in the Surface Global Ocean Based on Moderate-Resolution Imaging Spectroradiometer Data. Journal of Geophysical Research: Oceans, 110(C7): 2004JC002560. https://doi.org/10.1029/2004JC002560
Basedow, S. L., Sundfjord, A., von Appen, W. J., et al., 2018. Seasonal Variation in Transport of Zooplankton into the Arctic Basin through the Atlantic Gateway, Fram Strait. Frontiers in Marine Science, 5: 194. https://doi.org/10.3389/fmars.2018.00194
Batchelder, H. P., Mackas, D. L., O'Brien, T. D., 2012. Spatial-Temporal Scales of Synchrony in Marine Zooplankton Biomass and Abundance Patterns: A World-Wide Comparison. Progress in Oceanography, 97/98/99/100: 15-30. https://doi.org/10.1016/j.pocean.2011.11.010
Batten, S. D., Abu-Alhaija, R., Chiba, S., et al., 2019. A Global Plankton Diversity Monitoring Program. Frontiers in Marine Science, 6: 321. https://doi.org/10.3389/fmars.2019.00321
Batten, S. D., Walne, A. W., 2011. Variability in Northwards Extension of Warm Water Copepods in the NE Pacific. Journal of Plankton Research, 33(11): 1643-1653. https://doi.org/10.1093/plankt/fbr065
Beaugrand, G., 2004. The North Sea Regime Shift: Evidence, Causes, Mechanisms and Consequences. Progress in Oceanography, 60(2/3/4): 245-262. https://doi.org/10.1016/j.pocean.2004.02.018
Beaugrand, G., Reid, P. C., Ibañez, F., et al., 2002. Reorganization of North Atlantic Marine Copepod Biodiversity and Climate. Science, 296(5573): 1692-1694. https://doi.org/10.1126/science.1071329
Bindoff, N. L., Cheung, W. W. L., Kairo, J. G., et al., 2019. Changing Ocean, Marine Ecosystems, and Dependent Communities. In: IPCC, ed., IPCC Special Report on the Ocean and Cryosphere in a Changing Climate. Cambridge University Press, Cambridge, 447-588. https://doi.org/10.1017/9781009157964.007
Borkman, D. G., Fofonoff, P., Smayda, T. J., et al., 2018. Changing Acartia spp. Phenology and Abundance during a Warming Period in Narragansett Bay, Rhode Island, USA: 1972-1990. Journal of Plankton Research, 40(5): 580-594. https://doi.org/10.1093/plankt/fby029
Both, C., Van Asch, M., Bijlsma, R. G., et al., 2009. Climate Change and Unequal Phenological Changes across Four Trophic Levels: Constraints or Adaptations? Journal of Animal Ecology, 78(1): 73-83. https://doi.org/10.1111/j.1365-2656.2008.01458.x
Brown, M., Kawaguchi, S., Candy, S., et al., 2010. Temperature Effects on the Growth and Maturation of Antarctic Krill (Euphausia superba). Deep Sea Research Part II: Topical Studies in Oceanography, 57(7/8): 672-682. https://doi.org/10.1016/j.dsr2.2009.10.016
Chaikin, S., Dubiner, S., Belmaker, J., 2022. Cold-Water Species Deepen to Escape Warm Water Temperatures. Global Ecology and Biogeography, 31(1): 75-88. https://doi.org/10.1111/geb.13414
Chavez, F. P., Ryan, J., Lluch-Cota, S. E., et al., 2003. From Anchovies to Sardines and Back: Multidecadal Change in the Pacific Ocean. Science, 299(5604): 217-221. https://doi.org/10.1126/science.1075880
Chivers, W. J., Edwards, M., Hays, G. C., 2020. Phenological Shuffling of Major Marine Phytoplankton Groups over the Last Six Decades. Diversity and Distributions, 26(5): 536-548. https://doi.org/10.1111/ddi.13028
Chivers, W. J., Walne, A. W., Hays, G. C., 2017. Mismatch between Marine Plankton Range Movements and the Velocity of Climate Change. Nature Communications, 8: 14434. https://doi.org/10.1038/ncomms14434
Chust, G., Castellani, C., Licandro, P., et al., 2014. Are Calanus spp. Shifting Poleward in the North Atlantic? A Habitat Modelling Approach. ICES Journal of Marine Science, 71(2): 241-253. https://doi.org/10.1093/icesjms/fst147
Conversi, A., Dakos, V., Gårdmark, A., et al., 2015. A Holistic View of Marine Regime Shifts. Philosophical Transactions of the Royal Society B: Biological Sciences, 370(1659): 20130279. https://doi.org/10.1098/rstb.2013.0279
Dalpadado, P., Ingvaldsen, R. B., Stige, L. C., et al., 2012. Climate Effects on Barents Sea Ecosystem Dynamics. ICES Journal of Marine Science, 69(7): 1303-1316. https://doi.org/10.1093/icesjms/fss063
Davis, A. J., Jenkinson, L. S., Lawton, J. H., et al., 1998. Making Mistakes When Predicting Shifts in Species Range in Response to Global Warming. Nature, 391(6669): 783-786. https://doi.org/10.1038/35842
Deutsch, C. A., Tewksbury, J. J., Huey, R. B., et al., 2008. Impacts of Climate Warming on Terrestrial Ectotherms across Latitude. Proceedings of the National Academy of Sciences of the United States of America, 105(18): 6668-6672. https://doi.org/10.1073/pnas.0709472105
Dulvy, N. K., Rogers, S. I., Jennings, S., et al., 2008. Climate Change and Deepening of the North Sea Fish Assemblage: A Biotic Indicator of Warming Seas. Journal of Applied Ecology, 45(4): 1029-1039. https://doi.org/10.1111/j.1365-2664.2008.01488.x
Duarte, C. M., Cebrián, J., Marbà, N., 1992. Uncertainty of Detecting Sea Change. Nature, 356(6366): 190. https://doi.org/10.1038/356190a0
Dupont, N., Bagøien, E., Melle, W., 2017. Inter-Annual Variability in Spring Abundance of Adult Calanus finmarchicus from the Overwintering Population in the Southeastern Norwegian Sea. Progress in Oceanography, 152: 75-85. https://doi.org/10.1016/j.pocean.2017.02.004
Edwards, M., Richardson, A. J., 2004. Impact of Climate Change on Marine Pelagic Phenology and Trophic Mismatch. Nature, 430(7002): 881-884. https://doi.org/10.1038/nature02808
Ekman, S., 1953. Zoogeography of the Sea, vol. 9. Sidgwick & Jackson, London, 417.
Ershova, E. A., Kosobokova, K. N., Banas, N. S., et al., 2021. Sea Ice Decline Drives Biogeographical Shifts of Key Calanus Species in the Central Arctic Ocean. Global Change Biology, 27(10): 2128-2143. https://doi.org/10.1111/gcb.15562
Falkenhaug, T., Broms, C., Bagøien, E., et al., 2022. Temporal Variability of Co-Occurring Calanus finmarchicus and C. helgolandicus in Skagerrak. Frontiers in Marine Science, 9: 779335. https://doi.org/10.3389/fmars.2022.779335
Field, C., Behrenfeld, M., Randerson, J., et al., 1998. Primary Production of the Biosphere: Integrating Terrestrial and Oceanic Components. Science, 281(5374): 237-240. https://doi.org/10.1126/science.281.5374.237
Fox-Kemper, B., Hewitt, H. T., Xiao, C., et al., 2021. Ocean, Cryosphere and Sea Level Change. In: Masson-Delmotte, V., Zhai, P., Pirani, A., et al., eds., Climate Change 2021: The Physical Science Basis. Contribution of Working Group Ⅰ to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change. Cambridge University Press, Cambridge, 1211-1362. https://doi.org/10.1017/9781009157896.011
García-Soto, C., Caesar, L., Cazenave, A., et al., 2021. Chapter 05 Trends in the Physical and Chemical State of the Ocean. In: United Nations, ed., World Ocean Assessment Ⅱ. United Nations, New York, 83-103. https://doi.org/10.18356/9789216040062
Grandremy, N., Bourriau, P., Daché, E., et al., 2024. Metazoan Zooplankton in the Bay of Biscay: A 16-Year Record of Individual Sizes and Abundances Obtained Using the ZooScan and ZooCAM Imaging Systems. Earth System Science Data, 16(3): 1265-1282. https://doi.org/10.5194/essd-16-1265-2024
Gushing, D. H., Dickson, R. R., 1977. The Biological Response in the Sea to Climatic Changes. In: Russell, F. S., Yonge, M., eds., Advances in Marine Biology, vol. 14. Academic Press, Cambridge, 1-122. https://doi.org/10.1016/S0065-2881(08)60446-0
Hampton, S. E., Gray, D. K., Izmest'eva, L. R., et al., 2014. The Rise and Fall of Plankton: Long-Term Changes in the Vertical Distribution of Algae and Grazers in Lake Baikal, Siberia. PLoS One, 9(2): e88920. https://doi.org/10.1371/journal.pone.0088920
Hare, S. R., Mantua, N. J., 2000. Empirical Evidence for North Pacific Regime Shifts in 1977 and 1989. Progress in Oceanography, 47(2-4): 103-145. https://doi.org/10.1016/S0079-6611(00)00033-1
Hastings, R. A., Rutterford, L. A., Freer, J. J., et al., 2020. Climate Change Drives Poleward Increases and Equatorward Declines in Marine Species. Current Biology, 30(8): 1572-1577. https://doi.org/10.1016/j.cub.2020.02.043
Hays, G. C., Richardson, A. J., Robinson, C., 2005. Climate Change and Marine Plankton. Trends in Ecology & Evolution, 20(6): 337-344. https://doi.org/10.1016/j.tree.2005.03.004
Helaouët, P., Beaugrand, G., 2007. Macroecology of Calanus finmarchicus and C. helgolandicus in the North Atlantic Ocean and Adjacent Seas. Marine Ecology Progress Series, 345: 147-165. https://doi.org/10.3354/meps06775
Henson, S. A., Cole, H. S., Hopkins, J., et al., 2018. Detection of Climate Change-Driven Trends in Phytoplankton Phenology. Global Change Biology, 24(1): e101-e111. https://doi.org/10.1111/gcb.13886
Henson, S. A., Sarmiento, J. L., Dunne, J. P., et al., 2010. Detection of Anthropogenic Climate Change in Satellite Records of Ocean Chlorophyll and Productivity. Biogeosciences, 7(2): 621-640. https://doi.org/10.5194/bg-7-621-2010
Hinder, S. L., Gravenor, M. B., Edwards, M., et al., 2014. Multi-Decadal Range Changes vs. Thermal Adaptation for North East Atlantic Oceanic Copepods in the Face of Climate Change. Global Change Biology, 20(1): 140-146. https://doi.org/10.1111/gcb.12387
Hinder, S. L., Manning, J. E., Gravenor, M. B., et al., 2012. Long-Term Changes in Abundance and Distribution of Microzooplankton in the NE Atlantic and North Sea. Journal of Plankton Research, 34(1): 83-91. https://doi.org/10.1093/plankt/fbr087
Hirche, H. J., Kosobokova, K., 2007. Distribution of Calanus finmarchicus in the Northern North Atlantic and Arctic Ocean-Expatriation and Potential Colonization. Deep Sea Research Part II: Topical Studies in Oceanography, 54(23-26): 2729-2747. https://doi.org/10.1016/j.dsr2.2007.08.006
Hoover, B. A., García-Reyes, M., Batten, S. D., et al., 2021. Spatio-Temporal Persistence of Zooplankton Communities in the Gulf of Alaska. PLoS One, 16(1): e0244960. https://doi.org/10.1371/journal.pone.0244960
Hu, S. N., Fedorov, A. V., 2020. Indian Ocean Warming as a Driver of the North Atlantic Warming Hole. Nature Communications, 11: 4785. https://doi.org/10.1038/s41467-020-18522-5
Jahn, A., Holland, M. M., 2013. Implications of Arctic Sea Ice Changes for North Atlantic Deep Convection and the Meridional Overturning Circulation in CCSM4-CMIP5 Simulations. Geophysical Research Letters, 40(6): 1206-1211. https://doi.org/10.1002/grl.50183
Jahn, A., Holland, M. M., Kay, J. E., 2024. Projections of an Ice-Free Arctic Ocean. Nature Reviews Earth & Environment, 5(3): 164-176. https://doi.org/10.1038/s43017-023-00515-9
Ji, R. B., Edwards, M., Mackas, D. L., et al., 2010. Marine Plankton Phenology and Life History in a Changing Climate: Current Research and Future Directions. Journal of Plankton Research, 32(10): 1355-1368. https://doi.org/10.1093/plankt/fbq062
Johannesen, E., Ingvaldsen, R. B., Bogstad, B., et al., 2012. Changes in Barents Sea Ecosystem State, 1970-2009: Climate Fluctuations, Human Impact, and Trophic Interactions. ICES Journal of Marine Science, 69(5): 880-889. https://doi.org/10.1093/icesjms/fss046
Jonkers, L., Hillebrand, H., Kucera, M., 2019. Global Change Drives Modern Plankton Communities away from the Pre-Industrial State. Nature, 570(7761): 372-375. https://doi.org/10.1038/s41586-019-1230-3
Jorda, G., Marbà, N., Bennett, S., et al., 2020. Ocean Warming Compresses the Three-Dimensional Habitat of Marine Life. Nature Ecology & Evolution, 4(1): 109-114. https://doi.org/10.1038/s41559-019-1058-0
Kaiser, P., Hagen, W., Bode-Dalby, M., et al., 2022. Tolerant but Facing Increased Competition: Arctic Zooplankton versus Atlantic Invaders in a Warming Ocean. Frontiers in Marine Science, 9: 908638. https://doi.org/10.3389/fmars.2022.908638
Kefford, B. J., Ghalambor, C. K., Dewenter, B., et al., 2022. Acute, Diel, and Annual Temperature Variability and the Thermal Biology of Ectotherms. Global Change Biology, 28(23): 6872-6888. https://doi.org/10.1111/gcb.16453
Kléparski, L., Beaugrand, G., Edwards, M., et al., 2022. Morphological Traits, Niche-Environment Interaction and Temporal Changes in Diatoms. Progress in Oceanography, 201: 102747. https://doi.org/10.1016/j.pocean.2022.102747
Kosobokova, K. N., 1999. The Reproductive Cycle and Life History of the Arctic Copepod Calanus Glacialis in the White Sea. Polar Biology, 22(4): 254-263. https://doi.org/10.1007/s003000050418
Kraft, A., Bauerfeind, E., Nöthig, E. M., 2011. Amphipod Abundance in Sediment Trap Samples at the Long-Term Observatory HAUSGARTEN (Fram Strait, ~79°N/4°E). Variability in Species Community Patterns. Marine Biodiversity, 41(3): 353-364. https://doi.org/10.1007/s12526-010-0052-1
Kraft, A., Bauerfeind, E., Nöthig, E. M., et al., 2012. Size Structure and Life Cycle Patterns of Dominant Pelagic Amphipods Collected as Swimmers in Sediment Traps in the Eastern Fram Strait. Journal of Marine Systems, 95: 1-15. https://doi.org/10.1016/j.jmarsys.2011.12.006
Kraft, A., Nöthig, E. M., Bauerfeind, E., et al., 2013. First Evidence of Reproductive Success in a Southern Invader Indicates Possible Community Shifts among Arctic Zooplankton. Marine Ecology Progress Series, 493: 291-296. https://doi.org/10.3354/meps10507
Kromkamp, J. C., van Engeland, T., 2010. Changes in Phytoplankton Biomass in the Western Scheldt Estuary during the Period 1978-2006. Estuaries and Coasts, 33(2): 270-285. https://doi.org/10.1007/s12237-009-9215-3
Kvile, K. Ø., Ashjian, C., Feng, Z. X., et al., 2018. Pushing the Limit: Resilience of an Arctic Copepod to Environmental Fluctuations. Global Change Biology, 24(11): 5426-5439. https://doi.org/10.1111/gcb.14419
Li, H. B., Xu, Z. Q., Zhang, W. C., et al., 2016. Boreal Tintinnid Assemblage in the Northwest Pacific and Its Connection with the Japan Sea in Summer 2014. PLoS One, 11(4): e0153379. https://doi.org/10.1371/journal.pone.0153379
Lindley, J. A., Daykin, S., 2005. Variations in the Distributions of Centropages chierchiae and Temora stylifera (Copepoda: Calanoida) in the North-Eastern Atlantic Ocean and Western European Shelf Waters. ICES Journal of Marine Science, 62(5): 869-877. https://doi.org/10.1016/j.icesjms.2005.02.009
Liu, H., Gong, X., 2024. Revisiting North Pacific Intermediate Water in the Modern Ocean. Earth Science, 49(8): 2914-2924 (in Chinese with English abstract). https://doi.org/10.3799/dqkx.2024.036
Longhurst, A. R., 2001. Pelagic Biogeography. In: Steele, J. H., Thorpe, S. A., Turekian, K. K., eds., Encyclopedia of Ocean Sciences, vol. 4. Academic Press, San Diego, 356-363. https://doi.org/10.1016/B978-012374473-9.00288-5
Mackas, D. L., Batten, S., Trudel, M., 2007. Effects on Zooplankton of a Warmer Ocean: Recent Evidence from the Northeast Pacific. Progress in Oceanography, 75(2): 223-252. https://doi.org/10.1016/j.pocean.2007.08.010
Mackas, D. L., Goldblatt, R., Lewis, A. G., 1998. Interdecadal Variation in Developmental Timing of Neocalanus plumchrus Populations at Ocean Station P in the Subarctic North Pacific. Canadian Journal of Fisheries and Aquatic Sciences, 55(8): 1878-1893. https://doi.org/10.1139/f98-080
Mackas, D. L., Greve, W., Edwards, M., et al., 2012. Changing Zooplankton Seasonality in a Changing Ocean: Comparing Time Series of Zooplankton Phenology. Progress in Oceanography, 97/98/99/100: 31-62. https://doi.org/10.1016/j.pocean.2011.11.005
Melle, W., Runge, J., Head, E., et al., 2014. The North Atlantic Ocean as Habitat for Calanus finmarchicus: Environmental Factors and Life History Traits. Progress in Oceanography, 129: 244-284. https://doi.org/10.1016/j.pocean.2014.04.026
Mészáros, L., van der Meulen, F., Jongbloed, G., et al., 2021. Climate Change Induced Trends and Uncertainties in Phytoplankton Spring Bloom Dynamics. Frontiers in Marine Science, 8: 669951. https://doi.org/10.3389/fmars.2021.669951
Michael, K., Suberg, L. A., Wessels, W., et al., 2021. Facing Southern Ocean Warming: Temperature Effects on Whole Animal Performance of Antarctic Krill (Euphausia superba). Zoology, 146: 125910. https://doi.org/10.1016/j.zool.2021.125910
Møller, E. F., Nielsen, T. G., 2020. Borealization of Arctic Zooplankton-Smaller and Less Fat Zooplankton Species in Disko Bay, Western Greenland. Limnology and Oceanography, 65(6): 1175-1188. https://doi.org/10.1002/lno.11380
Neukermans, G., Oziel, L., Babin, M., 2018. Increased Intrusion of Warming Atlantic Water Leads to Rapid Expansion of Temperate Phytoplankton in the Arctic. Global Change Biology, 24(6): 2545-2553. https://doi.org/10.1111/gcb.14075
Niehoff, B., Hirche, H. J., 2005. Reproduction of Calanus glacialis in the Lurefjord (Western Norway): Indication for Temperature-Induced Female Dormancy. Marine Ecology Progress Series, 285: 107-115. https://doi.org/10.3354/meps285107
Oliver, E. C. J., Burrows, M. T., Donat, M. G., et al., 2019. Projected Marine Heatwaves in the 21st Century and the Potential for Ecological Impact. Frontiers in Marine Science, 6: 734. https://doi.org/10.3389/fmars.2019.00734
Ono, A., Moteki, M., 2017. Spatial Distribution of Salpa thompsoni in the High Antarctic Area off Adélie Land, East Antarctica during the Austral Summer 2008. Polar Science, 12: 69-78. https://doi.org/10.1016/j.polar.2016.11.005
Oziel, L., Sirven, J., Gascard, J. C., 2016. The Barents Sea Frontal Zones and Water Masses Variability (1980-2011). Ocean Science, 12(1): 169-184. https://doi.org/10.5194/os-12-169-2016
Parmesan, C., Yohe, G., 2003. A Globally Coherent Fingerprint of Climate Change Impacts across Natural Systems. Nature, 421(6918): 37-42. https://doi.org/10.1038/nature01286
Pata, P. R., Galbraith, M., Young, K., et al., 2022. Persistent Zooplankton Bioregions Reflect Long-Term Consistency of Community Composition and Oceanographic Drivers in the NE Pacific. Progress in Oceanography, 206: 102849. https://doi.org/10.1016/j.pocean.2022.102849
Pata, P. R., Galbraith, M., Young, K., et al., 2024. Data-Driven Determination of Zooplankton Bioregions and Robustness Analysis. MethodsX, 12: 102676. https://doi.org/10.1016/j.mex.2024.102676
Perry, A. L., Low, P. J., Ellis, J. R., et al., 2005. Climate Change and Distribution Shifts in Marine Fishes. Science, 308(5730): 1912-1915. https://doi.org/10.1126/science.1111322
Piñones, A., Fedorov, A. V., 2016. Projected Changes of Antarctic Krill Habitat by the End of the 21st Century. Geophysical Research Letters, 43(16): 8580-8589. https://doi.org/10.1002/2016GL069656
Pinsky, M. L., Eikeset, A. M., McCauley, D. J., et al., 2019. Greater Vulnerability to Warming of Marine versus Terrestrial Ectotherms. Nature, 569(7754): 108-111. https://doi.org/10.1038/s41586-019-1132-4
Poloczanska, E. S., Brown, C. J., Sydeman, W. J., et al., 2013. Global Imprint of Climate Change on Marine Life. Nature Climate Change, 3(10): 919-925. https://doi.org/10.1038/nclimate1958
Poloczanska, E. S., Burrows, M. T., Brown, C. J., et al., 2016. Responses of Marine Organisms to Climate Change across Oceans. Frontiers in Marine Science, 3: 62. https://doi.org/10.3389/fmars.2016.00062
Polyakov, I. V., Alkire, M. B., Bluhm, B. A., et al., 2020. Borealization of the Arctic Ocean in Response to Anomalous Advection from Sub-Arctic Seas. Frontiers in Marine Science, 7: 491. https://doi.org/10.3389/fmars.2020.00491
Qi, Z. H., Shi, R. J., Dai, M., et al., 2021. A Review on Ecological Characteristics of Creseis acicula and Preliminary Analysis on Its Outbreak Triggers in Daya Bay. Journal of Tropical Oceanography, 40(5): 147-152 (in Chinese with English abstract). https://doi.org/10.11978/2020112
Richardson, A. J., 2008. In Hot Water: Zooplankton and Climate Change. ICES Journal of Marine Science, 65(3): 279-295. https://doi.org/10.1093/icesjms/fsn028
Schlüter, M. H., Kraberg, A., Wiltshire, K. H., 2012. Long-Term Changes in the Seasonality of Selected Diatoms Related to Grazers and Environmental Conditions. Journal of Sea Research, 67(1): 91-97. https://doi.org/10.1016/j.seares.2011.11.001
Schröter, F., Havermans, C., Kraft, A., et al., 2019. Pelagic Amphipods in the Eastern Fram Strait with Continuing Presence of Themisto Compressa Based on Sediment Trap Time Series. Frontiers in Marine Science, 6: 311. https://doi.org/10.3389/fmars.2019.00311
Schultz, M., Choquet, M., Tverberg, V., et al., 2023. Calanus helgolandicus-More Than a Guest in the North? Journal of Plankton Research, 45(1): 33-36. https://doi.org/10.1093/plankt/fbac070
Schultz, M., Nielsen, T. G., Møller, E. F., 2020. The Importance of Temperature and Lipid Accumulation for Initiation and Duration of Calanus hyperboreus Spawning. Journal of Plankton Research, 42(2): 159-171. https://doi.org/10.1093/plankt/fbaa003
Schwartzlose, R. A., Alheit, J., Bakun, A., et al., 1999. Worldwide Large-Scale Fluctuations of Sardine and Anchovy Populations. South African Journal of Marine Science, 21(1): 289-347. https://doi.org/10.2989/025776199784125962
Sheng, G. L., Tao, H. L., Song, S. W., et al., 2025. Applications of Ancient DNA Research in the Field of Geobiology. Earth Science, 50(3): 1105-1121 (in Chinese with English abstract). https://doi.org/10.3799/dqkx.2024.155
Shevchenko, O. G., Shulgina, M. A., Shulkin, V. M., et al., 2020. The Long-Term Dynamics and Morphology of the Diatom Thalassiosira Nordenskioeldii Cleve, 1873 (Bacillariophyta) from the Coastal Waters of Peter the Great Bay, Sea of Japan. Russian Journal of Marine Biology, 46(4): 284-291. https://doi.org/10.1134/S1063074020040069
Shi, Y. Q., Liu, Y. J., Shan, X. J., et al., 2025. Climate Change Induced First Record of Porpita porpita (Linnaeus, 1758) in the Yellow Sea, China. Marine Pollution Bulletin, 210: 117333. https://doi.org/10.1016/j.marpolbul.2024.117333
Słomska, A. W., Panasiuk, A., 2022. Environmental Conditions for the Successful Development of Salpa Thompsoni (Tunicata: Thaliaceae) Blastozooids and Embryos in the Atlantic Sector of the Southern Ocean. Marine Biology, 169(11): 138. https://doi.org/10.1007/s00227-022-04125-9
Słomska, A. W., Panasiuk, A., Weydmann-Zwolicka, A., et al., 2021. Historical Abundance and Distributions of Salpa Thompsoni Hot Spots in the Southern Ocean and Projections for Further Ocean Warming. Aquatic Conservation: Marine and Freshwater Ecosystems, 31(8): 2095-2102. https://doi.org/10.1002/aqc.3443
Southward, A. J., 1980. The Western English Channel-An Inconstant Ecosystem? Nature, 285(5764): 361-366. https://doi.org/10.1038/285361a0
Staten, P. W., Lu, J., Grise, K. M., et al., 2018. Re-Examining Tropical Expansion. Nature Climate Change, 8(9): 768-775. https://doi.org/10.1038/s41558-018-0246-2
Sunday, J. M., Bates, A. E., Dulvy, N. K., 2011. Global Analysis of Thermal Tolerance and Latitude in Ectotherms. Proceedings Biological Sciences, 278(1713): 1823-1830. https://doi.org/10.1098/rspb.2010.1295
Sunday, J. M., Pecl, G. T., Frusher, S., et al., 2015. Species Traits and Climate Velocity Explain Geographic Range Shifts in an Ocean-Warming Hotspot. Ecology Letters, 18(9): 944-953. https://doi.org/10.1111/ele.12474
Swadling, K. M., Constable, A. J., Fraser, A. D., et al., 2023. Biological Responses to Change in Antarctic Sea Ice Habitats. Frontiers in Ecology and Evolution, 10: 1073823. https://doi.org/10.3389/fevo.2022.1073823
Tachibana, A., Nomura, H., Ishimaru, T., 2019. Impacts of Long-Term Environmental Variability on Diapause Phenology of Coastal Copepods in Tokyo Bay, Japan. Limnology and Oceanography, 64(S1): S273-S283. https://doi.org/10.1002/lno.11030
Tarling, G. A., Freer, J. J., Banas, N. S., et al., 2022. Can a Key Boreal Calanus Copepod Species Now Complete Its Life-Cycle in the Arctic? Evidence and Implications for Arctic Food-Webs. Ambio, 51(2): 333-344. https://doi.org/10.1007/s13280-021-01667-y
Thackeray, S. J., Henrys, P. A., Hemming, D., et al., 2016. Phenological Sensitivity to Climate across Taxa and Trophic Levels. Nature, 535(7611): 241-245. https://doi.org/10.1038/nature18608
Thoman, R. L., Moon, T. A., Druckenmiller, M. L., 2023. Arctic Report Card 2023. NOAA, Washington D. C. . https://doi.org/10.25923/5vfa-k694
Usov, N. V., Khaitov, V. M., Kutcheva, I. P., et al., 2021. Phenological Responses of the Arctic, Ubiquitous, and Boreal Copepod Species to Long-Term Changes in the Annual Seasonality of the Water Temperature in the White Sea. Polar Biology, 44(5): 959-976. https://doi.org/10.1007/s00300-021-02851-2
Wang, C. F., Wang, X. Y., Xu, Z. Q., et al., 2022a. Planktonic Tintinnid Community Structure Variations in Different Water Masses of the Arctic Basin. Frontiers in Marine Science, 8: 775653. https://doi.org/10.3389/fmars.2021.775653
Wang, C. F., Xu, Z. Q., He, Y., et al., 2022b. Neritic Tintinnid Community Structure and Mixing with Oceanic Tintinnids in Shelf Waters of the Pacific Arctic Region during Summer. Continental Shelf Research, 239: 104720. https://doi.org/10.1016/j.csr.2022.104720
Wang, C. F., Xu, Z. Q., Liu, C. G., et al., 2019. Vertical Distribution of Oceanic Tintinnid (Ciliophora: Tintinnida) Assemblages from the Bering Sea to Arctic Ocean through Bering Strait. Polar Biology, 42(11): 2105-2117. https://doi.org/10.1007/s00300-019-02585-2
Wiltshire, K. H., Malzahn, A. M., Wirtz, K., et al., 2008. Resilience of North Sea Phytoplankton Spring Bloom Dynamics: An Analysis of Long-Term Data at Helgoland Roads. Limnology and Oceanography, 53(4): 1294-1302. https://doi.org/10.4319/lo.2008.53.4.1294
Winder, M., Berger, S. A., Lewandowska, A., et al., 2012. Spring Phenological Responses of Marine and Freshwater Plankton to Changing Temperature and Light Conditions. Marine Biology, 159(11): 2491-2501. https://doi.org/10.1007/s00227-012-1964-z
Woillez, M., Rivoirard, J., Petitgas, P., 2009. Notes on Survey-Based Spatial Indicators for Monitoring Fish Populations. Aquatic Living Resources, 22(2): 155-164. https://doi.org/10.1051/alr/2009017
Woodgate, R. A., Aagaard, K., Weingartner, T. J., 2005. Monthly Temperature, Salinity, and Transport Variability of the Bering Strait through Flow. Geophysical Research Letters, 32(4): 2004GL021880. https://doi.org/10.1029/2004GL021880
Xu, Z. L., Gao, Q., 2009. Labidocera Euchaeta: Its Distribution in Yangtze River Estuary and Responses to Global Warming. Chinese Journal of Applied Ecology, 20(5): 1196-1201 (in Chinese with English abstract).
Yamaguchi, R., Rodgers, K. B., Timmermann, A., et al., 2022. Trophic Level Decoupling Drives Future Changes in Phytoplankton Bloom Phenology. Nature Climate Change, 12(5): 469-476. https://doi.org/10.1038/s41558-022-01353-1
Yang, H., Lohmann, G., Krebs-Kanzow, U., et al., 2020. Poleward Shift of the Major Ocean Gyres Detected in a Warming Climate. Geophysical Research Letters, 47(5): e2019GL085868. https://doi.org/10.1029/2019GL085868
Yoshiki, T. M., Chiba, S., Sasaki, Y., et al., 2015. Northerly Shift of Warm-Water Copepods in the Western Subarctic North Pacific: Continuous Plankton Recorder Samples (2001-2013). Fisheries Oceanography, 24(5): 414-429. https://doi.org/10.1111/fog.12119
Zanna, L., Khatiwala, S., Gregory, J. M., et al., 2019. Global Reconstruction of Historical Ocean Heat Storage and Transport. Proceedings of the National Academy of Sciences of the United States of America, 116(4): 1126-1131. https://doi.org/10.1073/pnas.1808838115
Zhang, W., Li, J., Li, H., et al., 2025. A New Tintinnid Ciliate of Salpingella (Ciliophora: Spirotrichea) from the Subarctic North Pacific Ocean to Arctic Ocean, with Notes on Its Habitat. Zoological Systematics, 50(4): 293—301. https://doi.org/10.11865/zs.2025402
Zhang, W. C., Zhao, Y., Dong, Y., et al., 2021. Biogeography of Epipelagic Marine Plankton. Oceanologia et Limnologia Sinica, 52(2): 332-345 (in Chinese with English abstract). https://doi.org/10.11693/hyhz20200100211
刘辉, 宫勋, 2024. 现代北太平洋中层水研究进展综述. 地球科学, 49(8): 2914-2924. doi: 10.3799/dqkx.2024.036
齐占会, 史荣君, 戴明, 等, 2021. 尖笔帽螺(Creseis acicula)研究进展及其在大亚湾暴发机制初探. 热带海洋学报, 40(5): 147-152.
盛桂莲, 陶华林, 宋世文, 等, 2025. 古DNA研究在地球生物学领域的应用. 地球科学, 50(3): 1105-1121. doi: 10.3799/dqkx.2024.155
徐兆礼, 高倩, 2009. 长江口海域真刺唇角水蚤的分布及其对全球变暖的响应. 应用生态学报, 20(5): 1196-1201.
张武昌, 赵苑, 董逸, 等, 2021. 上层海洋浮游生物地理分布. 海洋与湖沼, 52(2): 332-345.