The Unsung Heroes of the Deep: Exploring the Crucial Contributions of Sea Spiders (Pantopoda) to the Balance and Diversity of Aquatic Ecosystems.
1. Michel Murwanashyaka, Kibogora polytechnic, Other, Rwanda
2. Lihua Jiang, Zhejiang Ocean University, Doctor, China
3. Xiaojun Yan, Zhejiang Ocean University, Associate Professor, China
The Pantopoda, commonly known as sea spiders, has a significant impact on the equilibrium and diversity of aquatic ecosystems. This study elucidates their critical, yet frequently undervalued, roles within these environments. As pivotal predators of sessile invertebrates, sea spiders play a crucial role in regulating prey populations and shaping community dynamics. Their bioturbation activities facilitate enhanced nutrient cycling, and certain species uniquely engage in symbiotic relationships with methanotrophic bacteria, thereby assimilating methane-derived carbon into their tissues. This finding necessitates a reevaluation of marine food webs and has consequential implications for climate science. With over 1300 described species, sea spiders exhibit remarkable diversity across a range of habitats, encompassing both shallow coastal zones and deep-sea ecosystems. Nonetheless, populations of Pantopoda are increasingly threatened by climate change manifested through ocean acidification, elevated water temperatures, and habitat degradation, as well as pollution (including chemical contaminants, plastic debris, eutrophication, and acoustic disturbances) and overfishing practices such as bottom trawling, bycatch, and ensuing trophic cascades. In response, there is an urgent need for the implementation of effective conservation strategies. These should include the establishment of marine protected areas, continued research initiatives aimed at species identification and ecological functions, and assessment of the impacts of climate change. Furthermore, enhanced public awareness and education are paramount. Only through a concerted effort involving scientific inquiry, policy development, and public engagement can we safeguard the long-term viability of these crucial organisms and the health of the marine ecosystems they inhabit.
Sea spiders ecosystem resilience protected areas genetic diversity aquatic ecosystem
Sea
spiders, despite their frequently overlooked status, constitute integral
components of diverse marine ecosystems. Their predatory behaviors serve to
regulate invertebrate populations, thereby contributing to overall biodiversity
and ecosystem stability. Furthermore, their involvement in nutrient cycling, especially
the distinctive methane-consuming symbiosis observed in certain species, underscores
their significant influence on biogeochemical processes. However, these
essential organisms are confronted with substantial threats. Climate change,
pollution, and overfishing, particularly through destructive practices such as bottom
trawling, are exerting severe pressures on sea spider populations. Additionally,
phenomena such as ocean acidification, elevated water temperatures, and habitat
degradation directly impact their physiological processes and survival
prospects. Pollution from various anthropogenic sources introduces toxins and
degrades the habitats of sea spiders. Overfishing practices employed by
fisheries disrupt food webs, resulting in significant bycatch. Consequently, it
is imperative for conservationists, researchers, and policymakers to act with
urgency to formulate and enforce effective conservation strategies. The
establishment and expansion of marine protected areas are critical for the
preservation of sea spider habitats. Ongoing research initiatives that focus on
species identification, ecological roles, and the impacts of climate change are
essential for informing conservation efforts. Moreover, enhancing public
awareness and promoting educational initiatives are vital for fostering support
for sustainable practices and ensuring the long-term viability of these often-overlooked
organisms. Only through a concerted effort that integrates scientific
understanding, policy implementation, and public engagement can we secure the
future of sea spiders and the health of the marine ecosystems they inhabit.
1. Aagaard, A., Liu, S., Tregenza, T., Braad Lund, M., Schramm, A., Verhoeven, K. J. F., . . . Bilde, T. (2022). Adapting to climate with limited genetic diversity: Nucleotide, DNA methylation and microbiome variation among populations of the social spider Stegodyphus dumicola. Molecular ecology, 31(22), 5765–5783. doi:https://doi.org/10.1111/mec.16696 Andriuzzi, W. (2025). Methane fuel for sea spiders. Nature Ecology & Evolution, 9(8), 1308–1308. doi:https://doi.org/10.1038/s41559-025-02820-6 Anupama Sapkota. (2023). Competition Interaction- Definition and Types with Examples. Arango, C. (2001). Sea spiders (Pycnogonida) from the Great Barrier Reef, Australia, feed on fire corals and zoanthids. Memoirs of the Queensland Museum, 46. Arango, C. (2018). Sea Spiders: Class or suphylum Pycnogonida. australasian-arachnology. Arango, C., & Brodie, G. (2003). Observations of predation on the tropical nudibranch Okenia sp. by the sea spider Anoplodactylus longiceps Williams (Arthropoda: Pycnogonida). Veliger -Berkeley-, 46, 99–101. Arango, C. P. (2001). Sea spiders (Pycnogonida) from the Great Barrier Reef, Australia, feed on fire corals and zoanthids. Memoirs of the Queensland Museum, Nature, 46(2), 656–656. Retrieved from https://search.informit.org/doi/10.3316/informit.T2025071100006400768837818 https://search.informit.org/doi/full/10.3316/informit.T2025071100006400768837818 https://search.informit.org/doi/pdf/10.3316/informit.T2025071100006400768837818 Arango, C. P., & Wheeler, W. C. (2007). Phylogeny of the sea spiders (Arthropoda, Pycnogonida) based on direct optimization of six loci and morphology. Cladistics, 23(3), 255–293. doi:https://doi.org/10.1111/j.1096-0031.2007.00143.x Arnaud, F., & Bamber, R. N. (1988). The Biology of Pycnogonida. In J. H. S. Blaxter & A. J. Southward (Eds.), Advances in Marine Biology (Vol. 24, pp. 1–96): Academic Press. Arreguín-Sánchez, F. (2022). Ecosystem-Based Fisheries Management: Selected Examples. In F. Arreguín-Sánchez (Ed.), Holistic Approach to Ecosystem-Based Fisheries Management: Linking Biological Hierarchies for Sustainable Fishing (pp. 125–146). Cham: Springer International Publishing. Bachelot, B., Uriarte, M., & McGuire, K. (2015). Interactions among mutualism, competition, and predation foster species coexistence in diverse communities. Theoretical Ecology, 8. doi:https://doi.org/10.1007/s12080-015-0251-2 Bain, B. (2003). Larval types and a summary of postembryonic development within the pycnogonids. Invertebrate Reproduction & Development, 43, 193–222. doi:https://doi.org/10.1080/07924259.2003.9652540 Ballesteros, J. A., Setton, E. V. W., Santibáñez-López, C. E., Arango, C. P., Brenneis, G., Brix, S., . . . Sharma, P. P. (2021). Phylogenomic Resolution of Sea Spider Diversification through Integration of Multiple Data Classes. Mol Biol Evol, 38(2), 686–701. doi:https://doi.org/10.1093/molbev/msaa228 Bamber, R. (2009). Two new species of Sericosura Fry & Hedgpeth, 1969 (Arthropoda: Pycnogonida: Ammotheidae), and a reassessment of the genus. Zootaxa, 2140, 56–68. doi:https://doi.org/10.11646/zootaxa.2140.1.7 Begon, M., & Harper, J. L., & Townsend, C.R. (1996). Ecology: Individuals, populations and communities. Blackwell Science. Behrens, W. (1984). Larvenentwicklung und Metamorphose von Pycnogonum litorale (Chelicerata, Pantopoda). Zoomorphology, 104(5), 266–279. doi:https://doi.org/10.1007/BF00312008 Birader, K. (2023a). Genetic Diversity and the Adaptation of Species to Changing Environments. Journal of Biodiversity & Endangered Species. Birader, K. (2023b). Genetic Diversity and the Adaptation of Species to Changing Environments. Branco, V. V., & Cardoso, P. (2020). An expert-based assessment of global threats and conservation measures for spiders. Global Ecology and Conservation, 24, e01290. doi:https://doi.org/10.1016/j.gecco.2020.e01290 Bravo, M., Mueller, H.-G., Arango, C., Tigreros, P., & Melzer, R. (2009). Morphology of shallow-water sea spiders from the Colombian Caribbean (Arthropoda: Pycnogonida). Spixiana, 32, 9–34. Brenneis, G., Frankowski, K., Maaß, L., & Scholtz, G. (2023). The sea spider <i>Pycnogonum litorale</i> overturns the paradigm of the absence of axial regeneration in molting animals. Proceedings of the National Academy of Sciences, 120(5), e2217272120. doi:https://doi.org/10.1073/pnas.2217272120 Cano-Sánchez, E., & López-González, P. J. (2016). Basal articulation of the palps and ovigers in Antarctic Colossendeis (Pycnogonida; Colossendeidae). Helgoland Marine Research, 70(1), 22. doi:https://doi.org/10.1186/s10152-016-0474-7 Cardinale, B. J., Duffy, J. E., Gonzalez, A., Hooper, D. U., Perrings, C., Venail, P., . . . Naeem, S. (2012). Biodiversity loss and its impact on humanity. Nature, 486(7401), 59–67. doi:https://doi.org/10.1038/nature11148 Chen, R., Tang, H., Zhao, F., Wu, Y., & Huang, Y. (2016). Food availability and initial relative abundance determine the outcome of interspecific competition between two different‐sized cladocerans. International Review of Hydrobiology, 101, 105–112. doi:https://doi.org/10.1002/iroh.201501832 Crooker, A. (2008). Sea Spiders (Pycnogonida). In J. L. Capinera (Ed.), Encyclopedia of Entomology (pp. 3321–3335). Dordrecht: Springer Netherlands. Dal Bó, B., Guo, Y., Mayr, M. J., Pereira, O. S., Levin, L. A., Orphan, V. J., & Goffredi, S. K. (2025). Methane-powered sea spiders: Diverse, epibiotic methanotrophs serve as a source of nutrition for deep-sea methane seep <i>Sericosura</i>. Proceedings of the National Academy of Sciences, 122(26), e2501422122. doi:https://doi.org/10.1073/pnas.2501422122 Davenport, J., Blacstock, N., & Davies, D. A. Y., M. (1987). Observations on the physiology and integumentary structure of the Antarctic pycnogonid Decolopoda australis. Journal of Zoology, London, 211, 451–465. Davila, C. M. (2025). Why Biodiverse Ecosystems Are More Resilient to Climate Change. Wildlife nomads. de HARO, A. (2008). Ecological distribution of pycnogonids on the Catalan coast. Zoological Journal of the Linnean Society, 63(1-2), 181–196. doi:https://doi.org/10.1111/j.1096-3642.1978.tb02097.x De, K., Siliwal, M., Uniyal, V. P., & Hussain, S. A. (2022). Spiders as bio-indicators of habitat disturbance in the riparian zone of the Ganga river: a preliminary study. Tropical Ecology, 63(2), 209–215. doi:https://doi.org/10.1007/s42965-021-00192-z Deines, P., Bodelier, P. L., & Eller, G. (2007). Methane-derived carbon flows through methane-oxidizing bacteria to higher trophic levels in aquatic systems. Environ Microbiol, 9(5), 1126–1134. doi:https://doi.org/10.1111/j.1462-2920.2006.01235.x Dietz, L., Dömel, J. S., Leese, F., Lehmann, T., & Melzer, R. R. (2018). Feeding ecology in sea spiders (Arthropoda: Pycnogonida): what do we know? Front Zool, 15, 7. doi:https://doi.org/10.1186/s12983-018-0250-4 Du, Y., Sun, J., & Zhang, G. (2021). The Impact of Overfishing on Environmental Resources and the Evaluation of Current Policies and Future Guideline. Dudnik, Z., & Kremenetskaia, A. (2025). Pycnogonida collection of the Shirshov Institute of Oceanology, Russian Academy of Sciences. Biodiversity Data Journal, 13. doi:https://doi.org/10.3897/BDJ.13.e142496 Dunlop, J., & Arango, C. (2005). Pycnogonid affinities: A review. Journal of Zoological Systematics & Evolutionary Research, 43, 8–21. doi:https://doi.org/10.1111/j.1439-0469.2004.00284.x Easman, E. S., Abernethy, K. E., & Godley, B. J. (2018). Assessing public awareness of marine environmental threats and conservation efforts. Marine Policy, 87, 234–240. doi:https://doi.org/10.1016/j.marpol.2017.10.030 Edgar, G. J., Stuart-Smith, R. D., Willis, T. J., Kininmonth, S., Baker, S. C., Banks, S., . . . Thomson, R. J. (2014). Global conservation outcomes depend on marine protected areas with five key features. Nature, 506(7487), 216–220. doi:https://doi.org/10.1038/nature13022 Edgecombe, G. D., Wilson, G. D. F., Colgan, D. J., Gray, M. R., & Cassis, G. (2000). Arthropod Cladistics: Combined Analysis of Histone H3 and U2 snRNA Sequences and Morphology. Cladistics, 16(2), 155–203. doi:https://doi.org/10.1111/j.1096-0031.2000.tb00352.x Fage, L. (1949). Classe de Pycnogonides. In Grasse, P. P. (Ed). Traite de Zoologie, 906–941. Fontefrancesco, M. F., & Sidsaph, H. (2019). Food Web and Food Network: Role of Food System Ecological Interconnectedness in Achieving the Zero Hunger Goal. In (pp. 1–11). Fry, W. G. (1965). The feeding mechanisms and preferred foods of three species of Pycnogonida. Bulletin of the British Museum of Natural History, Zoology, 12, 195–233. Fry, W. G. H., J. W. (1969). Pycnogonida: Colossendeidae, Pycnogonidae, Endeidae, Ammotheidae. New Zealand Department of Scientific and Industrial Research Bulletin. 7, 1–139. Geldmann, J., Manica, A., Burgess, N. D., Coad, L., & Balmford, A. (2019). A global-level assessment of the effectiveness of protected areas at resisting anthropogenic pressures. Proc Natl Acad Sci U S A, 116(46), 23209–23215. doi:https://doi.org/10.1073/pnas.1908221116 Giribet, G. (2000). A Review of Arthropod Phylogeny: New Data Based on Ribosomal DNA Sequences and Direct Character Optimization. Cladistics, 16, 204–231. doi:https://doi.org/10.1006/clad.1999.0128 Giribet, G., Edgecombe, G. D., & Wheeler, W. C. (2001). Arthropod phylogeny based on eight molecular loci and morphology. Nature, 413(6852), 157–161. doi:https://doi.org/10.1038/35093097 Gordon, I. (1944). Pycnogonida. British and New Zealand Antarctic Research Expeditions, 1–72. Hedgpeth, J. W. (1948). The Pycnogonida of the Western North Atlantic and the Caribbean. Proceedings of the US National Museum, 98, 157–342. Hedgpeth, J. W. (1949). Report on the Pycnogonida collected by the Albatross in Japanese waters. Proceedings of the US National Museum, 97, 233–321. Hedgpeth, J. W. (1954). Bottom communities of the Gulf of Mexico. Hedgpeth JW. (1947). On the evolutionary significance of the Pycnogonida. . In: Smithson Misc Collect. Helfer, H. S., E (1935). Pantopoda. In Bronn H. G. (Ed) Dr. H. G. Bronns Klassen und Ordnungen des Tierreichs. Leipzig: Akademische Verlagsgesellschaft m.b.H, 1–314. Hoban, S., Archer, F. I., Bertola, L. D., Bragg, J. G., Breed, M. F., Bruford, M. W., . . . Hunter, M. E. (2022). Global genetic diversity status and trends: towards a suite of Essential Biodiversity Variables (EBVs) for genetic composition. Biol Rev Camb Philos Soc, 97(4), 1511–1538. doi:https://doi.org/10.1111/brv.12852 Hoek, P. P. C. (1881). Report on the Pycnogonida, dredged by H.M.S. Challenger during the years 1873-76. By Dr. P.P.C. Hoek. Hojo, M. K. (2022). Evolution of chemical interactions between ants and their mutualist partners. Current Opinion in Insect Science, 52, 100943. doi:https://doi.org/10.1016/j.cois.2022.100943 Hooper, J. (1981). Some aspects of the reproductive biology of Paraparallene avida Stock (Pycnogonida: Callipallenidae) from northern New South Wales. Australian zoologist, 20, 473–484. Hu, X., Wu, X., Zhou, Q., Niklas, K. J., Jiang, L., Eisenhauer, N., . . . Sun, S. (2024). Warming causes contrasting spider behavioural responses by changing their prey size spectra. Nature Climate Change, 14(2), 190–197. doi:https://doi.org/10.1038/s41558-023-01918-8 Hulme-Beaman, A., Dobney, K., Cucchi, T., & Searle, J. B. (2016). An Ecological and Evolutionary Framework for Commensalism in Anthropogenic Environments. Trends Ecol Evol, 31(8), 633–645. doi:https://doi.org/10.1016/j.tree.2016.05.001 Jager, M., Murienne, J., Clabaut, C., Deutsch, J., Guyader, H. L., & Manuel, M. (2006). Homology of arthropod anterior appendages revealed by Hox gene expression in a sea spider. Nature, 441(7092), 506–508. doi:https://doi.org/10.1038/nature04591 Janshi, T., Sakariya, K., & Kamleshbhai, B. (2023). Major Threats to Marine Ecosystem-A Global Issue. 4, 110–113. Jarvis, J. H., & King, P. E. (1972). Reproduction and development in the pycnogonid Pycnogonum littorale. Marine Biology, 13(2), 146–154. doi:https://doi.org/10.1007/BF00366565 Jarvis, J. H., & King, P. E. (1975). Egg development and the reproductive cycle in the pycnogonid Endeis laevis. Marine Biology, 33(4), 331–339. doi:https://doi.org/10.1007/BF00390571 Jennings, S., Dinmore, T., Duplisea, D., Randall, K., & Lancaster, J. (2002). Trawling disturbance can modify benthic production processes. Journal of Animal Ecology, 70, 459–475. doi:https://doi.org/10.1046/j.1365-2656.2001.00504.x Jump, A. S., Marchant, R., & Peñuelas, J. (2009). Environmental change and the option value of genetic diversity. Trends in Plant Science, 14(1), 51–58. doi:https://doi.org/10.1016/j.tplants.2008.10.002 King, P. E. (1973). Pycnogonids. London: Hutchinson and Co. King, P. E., & Jarvis, J. H. (1970). Egg development in a littoral pycnogonid Nymphon gracile. Marine Biology, 7(4), 294–304. doi:https://doi.org/10.1007/BF00750822 Li, G., Fang, C., Watson, J. E. M., Sun, S., Qi, W., Wang, Z., & Liu, J. (2024). Mixed effectiveness of global protected areas in resisting habitat loss. Nature Communications, 15(1), 8389. doi:https://doi.org/10.1038/s41467-024-52693-9 Liu, S., Chen, J., Gan, W., Schaefer, D., Gan, J., & Yang, X. (2015). Spider foraging strategy affects trophic cascades under natural and drought conditions. Scientific Reports, 5(1), 12396. doi:https://doi.org/10.1038/srep12396 Liu, Y., Melzer, R. R., Haug, J. T., Haug, C., Briggs, D. E. G., Hörnig, M. K., . . . Hou, X.-g. (2016). Three-dimensionally preserved minute larva of a great-appendage arthropod from the early Cambrian Chengjiang biota. Proceedings of the National Academy of Sciences, 113(20), 5542–5546. doi:https://doi.org/10.1073/pnas.1522899113 Liu, Y., Wang, J., Wu, Z., Li, S., & Dai, W. (2024). Identification of biodiversity priority conservation areas in China by integrating genetic, species and ecosystem diversity. Biological Conservation, 300, 110854. doi:https://doi.org/10.1016/j.biocon.2024.110854 Loman, J. C. C. (1908). Die Pantopoden der Siboga-Expedition. Siboga Expeditie Monographie, 40, 1–88. Manuel, M., Jager, M., Murienne, J., Clabaut, C., & Guyader, H. L. (2006). Hox genes in sea spiders (Pycnogonida) and the homology of arthropod head segments. Development Genes and Evolution, 216(7), 481–491. doi:https://doi.org/10.1007/s00427-006-0095-2 Mapstone, B. D., Little, L. R., Punt, A. E., Davies, C. R., Smith, A. D. M., Pantus, F., . . . Jones, A. (2008). Management strategy evaluation for line fishing in the Great Barrier Reef: Balancing conservation and multi-sector fishery objectives. Fisheries Research, 94(3), 315–329. doi:https://doi.org/10.1016/j.fishres.2008.07.013 Mathewson, S. (2016). Sea Spiders: Abnormally Large-Growing Arctic Sea Spiders Surprise Scientists. Maxmen, A., Browne, W. E., Martindale, M. Q., & Giribet, G. (2005). Neuroanatomy of sea spiders implies an appendicular origin of the protocerebral segment. Nature, 437(7062), 1144–1148. doi:https://doi.org/10.1038/nature03984 Maxwell, J., Gan, Y. M., Arango, C., Doemel, J. S., Allcock, A. L., van de Putte, A. P., & Griffiths, H. (2022). Sea spiders (Arthropoda, Pycnogonida) from ten recent research expeditions to the Antarctic Peninsula, Scotia Arc and Weddell Sea - data. Biodivers Data J, 10, e79353. doi:https://doi.org/10.3897/BDJ.10.e79353 Mercier, A., & Hamel, J.-F. (1994). Deleterious effects of a pycnogonid on the sea anemone Bartholomea annulata. Canadian Journal of Zoology-revue Canadienne De Zoologie - CAN J ZOOL, 72, 1362–1364. doi:https://doi.org/10.1139/z94-179 Miyazaki, K. M., T. (1993). A case of intersexuality in the sea spider, Cilunculus armatus. Zoological Science Tokyo, 10, 127–132. Myers, R. A., & Worm, B. (2003). Rapid worldwide depletion of predatory fish communities. Nature, 423(6937), 280–283. doi:https://doi.org/10.1038/nature01610 Nakamura, K. (1981). Post embryonic development of a pycnogonid, Propallene longiceps. Journal of Natural History, 15(1), 49–62. doi:https://doi.org/10.1080/00222938100770041 Nakamura, K., & Sekiguchi, K. (1980). Mating Behavior and Oviposition in the Pycnogonid Propallene longiceps. Marine Ecology Progress Series, 2, 163–168. Retrieved from https://www.int-res.com/abstracts/meps/v2/meps002163 Nonić, M., & Šijačić-Nikolić, M. (2019). Genetic Diversity: Sources, Threats, and Conservation. In W. Leal Filho, A. M. Azul, L. Brandli, P. G. Özuyar, & T. Wall (Eds.), Life on Land (pp. 1–15). Cham: Springer International Publishing. Olano, J. P., Weller, P. F., & Guerrant, R. L., & Walker, D. H, . (2011). Principles of Parasitism: Host-Parasite Interactions. In Tropical Infectious Diseases. Elsevier Inc, 1–7. doi: https://doi.org/10.1016/B978-0-7020-3935-5.00001-X Papadopoulos, N., Kulkarni, S. S., Baranyi, C., Fromm, B., Setton, E. V. W., Sharma, P. P., . . . Brenneis, G. (2025). The genome of a sea spider corroborates a shared Hox cluster motif in arthropods with a reduced posterior tagma. BMC Biol, 23(1), 196. doi:https://doi.org/10.1186/s12915-025-02276-x Papadopoulos, N., Kulkarni, S. S., Baranyi, C., Fromm, B., Setton, E. V. W., Sharma, P. P., . . . Brenneis, G. (2025). The genome of a sea spider corroborates a shared Hox cluster motif in arthropods with a reduced posterior tagma. BMC Biology, 23(1), 196. doi:https://doi.org/10.1186/s12915-025-02276-x Parniske, M. (2000). Intracellular accommodation of microbes by plants: a common developmental program for symbiosis and disease? Current Opinion in Plant Biology, 3(4), 320–328. doi:https://doi.org/10.1016/S1369-5266(00)00088-1 Pauly, D., Watson, R., & Alder, J. (2005). Global trends in world fisheries: impacts on marine ecosystems and food security. Philos Trans R Soc Lond B Biol Sci, 360(1453), 5–12. doi:https://doi.org/10.1098/rstb.2004.1574 Polidoro, B., Livingstone, S., Carpenter, K. E., Hutchinson, B., Mast, R., Pilcher, N., . . . Valenti, S. V. (2008). Status of the world's marine species. Wildlife in a Changing World-an Analysis of the 2008 IUCN Red List of Threatened Species, 1–7. Powers, R. P., & Jetz, W. (2019). Global habitat loss and extinction risk of terrestrial vertebrates under future land-use-change scenarios. Nature Climate Change, 9(4), 323–329. doi:https://doi.org/10.1038/s41558-019-0406-z Pringle, R. M. (2017). Upgrading protected areas to conserve wild biodiversity. Nature, 546(7656), 91–99. doi:https://doi.org/10.1038/nature22902 Rilov, G., Mazaris, A. D., Stelzenmüller, V., Helmuth, B., Wahl, M., Guy-Haim, T., . . . Katsanevakis, S. (2019). Adaptive marine conservation planning in the face of climate change: What can we learn from physiological, ecological and genetic studies? Global Ecology and Conservation, 17, e00566. doi:https://doi.org/10.1016/j.gecco.2019.e00566 Rodríguez-Rodríguez, D., & Martínez-Vega, J. (2022). Chapter Three - Ecological effectiveness of marine protected areas across the globe in the scientific literature. In C. Sheppard (Ed.), Advances in Marine Biology (Vol. 92, pp. 129–153): Academic Press. Romero, N. A. (2024). Sea Spiders - Types, Characteristics and Habitat. Sabroux, R., Corbari, L., & Hassanin, A. (2023). Phylogeny of sea spiders (Arthropoda: Pycnogonida) inferred from mitochondrial genome and 18S ribosomal RNA gene sequences. Mol Phylogenet Evol, 182, 107726. doi:https://doi.org/10.1016/j.ympev.2023.107726 Sala, O. E., Chapin, F. S., 3rd, Armesto, J. J., Berlow, E., Bloomfield, J., Dirzo, R., . . . Wall, D. H. (2000). Global biodiversity scenarios for the year 2100. Science, 287(5459), 1770–1774. doi:https://doi.org/10.1126/science.287.5459.1770 Salgotra, R. K., & Chauhan, B. S. (2023). Genetic Diversity, Conservation, and Utilization of Plant Genetic Resources. Genes (Basel), 14(1). doi:https://doi.org/10.3390/genes14010174 Seema Virbhan Keswani. (2025). The Effect of Environmental Pollution on Spiders: A Review of Impacts and Implications. . doi:https://doi.org/10.5281/zenodo.15254773. Sharma, P. P., & Gavish-Regev, E. (2025). The Evolutionary Biology of Chelicerata. Annu Rev Entomol, 70(1), 143–163. doi:https://doi.org/10.1146/annurev-ento-022024-011250 Sharma, S., Hampton, S. E., & Kimirei, I. (2024). Chapter 31 - Inland Waters: The Future of Limnology is Interdisciplinary, Collaborative, Inclusive, and Global. In I. D. Jones & J. P. Smol (Eds.), Wetzel's Limnology (Fourth Edition) (pp. 1045–1061). San Diego: Academic Press. Sherwood, J., Walls, J., & Ritz, D. A. (1998). Amathamide alkaloids in the pycnogonid, Stylopallene longicauda, epizoic on the chemically defended bryozoan, Amathia wilsoni. Pap. Proc. R. Soc. Tasman., 132. doi:https://doi.org/10.26749/rstpp.132.65 Sophia M. (2024). unveiling-the-mysteries-of-sea-spiders-their-role-and-importance-in-marine-ecosystems. The national. Staples DA. (2005). Pycnogonida (pycnogonids) In: Rohde K, editor. . Marine parasitology. Collingwood: CSIRO Publishing, 222–226. Staples, D. A. W., J. E. (1987). Associations between pycnogonids and hydroids. In Bouillion, J. (Ed) Modern trends in the systematics, ecology and evolution of hydroids and hydromedusae. Oxford: Oxford University Press, 215–226. Stock, J. H. (1954). Papers from Dr. Mortensen's Pacific Exped. 1914-1916 LXXVII. Pycnogonida from the Indo-West Pacific, Australian, New Zealand waters. Videnskabelige Meddelelser fra Dansk naturhistorisk Foreningen, 116, 1–168. Stock, J. H. (1975). Copepoda associated with West Indian Actiniaria and Corallimorpharia. Studies on the Fauna of Curaçao and other Caribbean Islands, 48(1), 88–118. Stock, J. H. (1994). Indo-West Pacific Pycnogonida collected by some major oceanographic expeditions. Beaufortia, 44(3), 17–77. Sumaila, R., & Tai, T. (2020). End Overfishing and Increase the Resilience of the Ocean to Climate Change. Frontiers in Marine Science, 7. doi:https://doi.org/10.3389/fmars.2020.00523 T, S. (2024). Aqua Complex Conservation and Strategies for Sustainable Management. J Aquac Res Dev, 15, 853. Tarhan, L., Pippenger, K., Cribb, A., Zill, M., Phelps, W., Droser, M., . . . Clapham, M. (2025). Tracking bioturbation through time: The evolution of the marine sedimentary mixed and transition layers. Science advances, 11, eadu7719. doi:https://doi.org/10.1126/sciadv.adu7719 Thorogood, R., Mustonen, V., Aleixo, A., Aphalo, P. J., Asiegbu, F. O., Cabeza, M., . . . Vanhatalo, J. (2023). Understanding and applying biological resilience, from genes to ecosystems. npj Biodiversity, 2(1), 16. doi:https://doi.org/10.1038/s44185-023-00022-6 Tomaschko, K. H., Wilhelm, E., & Bückmann, D. (1997). Growth and reproduction of Pycnogonum litorale (Pycnogonida) under laboratory conditions. Marine Biology, 129(4), 595–600. doi:https://doi.org/10.1007/s002270050201 Wang, Z., Leung, K. M. Y., Sung, Y.-H., Dudgeon, D., & Qiu, J.-W. (2021). Recovery of tropical marine benthos after a trawl ban demonstrates linkage between abiotic and biotic changes. Communications Biology, 4(1), 212. doi:https://doi.org/10.1038/s42003-021-01732-y Wilhelm, E., Bückmann, D., & Tomaschko, K. H. (1997). Life cycle and population dynamics of Pycnogonum litorale (Pycnogonida) in a natural habitat. Marine Biology, 129(4), 601–606. doi:https://doi.org/10.1007/s002270050202 Wolfe, J. M. (2017). Metamorphosis Is Ancestral for Crown Euarthropods, and Evolved in the Cambrian or Earlier. Integr Comp Biol, 57(3), 499–509. doi:https://doi.org/10.1093/icb/icx039 Wudu, K., Abegaz, A., Ayele, L., & Ybabe, M. (2023). The impacts of climate change on biodiversity loss and its remedial measures using nature based conservation approach: a global perspective. Biodiversity and Conservation, 32(12), 3681–3701. doi:10.1007/s10531-023-02656-1
Michel Murwanashyaka: Writing an original draft, writing a review & editing. Lihua Jiang: Supervision, writing a review & editing. Xiaojun Yan : Supervision, funding.
This work is supported by the National Key Research and Development Program of China (2023YFD2401905).
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The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.
This is to thank Kibogora Polytechnic University, Rwanda, for its partnership with Zhejiang Ocean University, China.
Data
supporting the findings of this study are available within the article.