Evidence of asexual polyp development from the adult body in Stomolopus sp2

preprint OA: closed
Full text JSON View at publisher
Full text 98,331 characters · extracted from preprint-html · click to expand
Evidence of asexual polyp development from the adult body in Stomolopus sp2 | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Evidence of asexual polyp development from the adult body in Stomolopus sp2 Cintya Alejandra Nevárez-López, Raul Llera-Herrera, Juana López-Martínez This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7579160/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 5 You are reading this latest preprint version Abstract Stomolophus spp. is one of the commercially important jellyfish in the world; however, its life cycle has recently been described with new forms of asexual reproduction in the polyp stage, with the reversion process to ephyrae considered an advantage for increasing population size, especially under stressful conditions. In this study, we observed the generation of new polyp colonies directly from the bell margins of the medusae. Here we present the first evidence for back-transformation of tissues medusae into polyps in Stomolophus sp.2. The new way of asexual reproduction from medusae in the complex life cycle of this species reveals a long-range asexual mechanism of dispersion and potential of invasion, as well as of capacities to rejuvenation, the possibilities of immortality with reprogramming of differentiated cells and the increase of populations and blooms not only in this species but also in others that may have the same process of reversion. cannonball jellyfish Stomolophus polyps asexual reproduction Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Introduction Cnidarians are a complex and diverse group of invertebrates that include corals, anemones, hydras, and jellyfish. In some regions, the abundance of jellyfish has increased over the past decade, and certain species have expanded their distribution ranges (Aouititen et al., 2024; Fonfría et al., 2024; Chuan et al., 2024). Besides seasonal fluctuations, several factors have contributed to the dramatic global increase in jellyfish populations over the last two decades, such as eutrophication, overfishing, coastal development, and species translocation (Pitt et al., 2018; Schnedler-Meyer et al., 2018; Fernández-Alías et al., 2020). This increase in jellyfish populations poses several challenges in coastal areas, such as blockages in pipeline intakes for thermoelectric power generation and skin injuries for swimmers. Additionally, these animals are voracious predators of fish eggs and planktonic larvae, leading to strong competition for food and predation pressure on fish populations when jellyfish proliferate excessively (Purcell et al., 2013; Fernández-Alías et al., 2024). Despite these challenges, jellyfish play a crucial ecological role. They contribute between 3.7 and 6.8 billion metric tons of carbon biomass each year (Luo et al., 2020) and serve as food for sea turtles and habitat for other species (González-Carmian et al., 2014). The cannonball jellyfish ( Stomolophus spp.) is found in the Mexican Pacific and is commercially exploited. In recent years, massive blooms of up to 60 org/m 2 have been recorded (López-Martínez and Álvarez-Tello, 2013). Its distribution extends from the northern part of Sonora in the Upper Gulf of California region (Brotz et al., 2021) to Ecuador (Preciado, 2016). More recently, it has also been reported in U.S. waters, suggesting a potential northward shifts in its distribution due to climatic conditions like El Niño, which enhance its reproductive capacity and ability to colonize new areas (Gómez-Salinas, 2018; Nevárez-López et al., 2020; Gómez-Salinas et al., 2021). Despite extensive evidence of increased jellyfish abundances, the strategies employed by these animals to boost their populations are not yet fully understood. However, their complex metagenic life cycle may hold some insight. This species reaches sexual maturity at a length of 10 cm (Carvalho-Saucedo et al., 2011). It is dioecious, with external fertilization, resulting in planula larvae that settle on a substrate after approximately 48 hours if optimal conditions are met. Following settlement, the larvae undergo a series of transformations, and a polyp becomes visible after 5 to 7 days (Carvalho-Saucedo et al., 2011). There are several variations of asexual reproduction mechanisms, with strobilation being the most common, producing ephyrae. Under laboratory conditions, polyps that undergo strobilation can produce up to 20-25 ephyrae that develop into free-swimming jellyfish (López-Martínez et al., 2022). In their sessile stage, the polyps can also produce additional polyps and podocytes, which are smaller forms that can develop into new polyps. This variety of asexual reproduction strategies allows these animals to significantly increase their populations while in the sessile stage; with asexual reproduction potentially continuing for years before strobilation. We have successfully been able to maintain asexually reproducing polyps for at least 8 years under controlled laboratory conditions. In various jellyfish species, the life cycle is similar. For example, both the moon jellyfish ( Aurelia sp.1 ) and the immortal jellyfish ( Turritopsis dohrnii ) can produce new polyps without sexual reproduction or the generation of planula larvae, and this occurs independently of any specific stress process (He et al., 2015; Matsumoto et al., 2019). While the life cycle of the cannonball jellyfish exhibits a reversal mechanism, only ephyrae have been observed reverting to polyps (López-Martínez et al., 2022). The existence of a broader range of asexual reproductive strategies may have significant ecological and fisheries management implications, as well as open up novel trends in biomedical and ecological research. In this context, we report a novel strategy for generating asexual polyps directly from juvenile jellyfish. Methods The polyps of Stomolophus sp.2 polyps derived from a laboratory colony reported in López-Martínez et al (2022) and transported in marine water to ICMyL-UNAM Mazatlán (Sinaloa, Mexico) in 2022. They settled on translucent acrylic plates (3 cm x 5 cm) in a horizontal position in polycarbonate containers half-filled with 4 liters of filtered marine water without aeration for 7 days at room temperature (25°C). Once the organisms were firmly attached, the acrylic plates were vertically immersed in 8-L tanks (see supplementary material) with gentle aeration and were fed every 3 days with brine shrimp nauplii. Colony growth and strobilation were induced by increasing the temperature to 28°C. The new polyps grew on the acrylic plate, and all ephyrae were collected and transferred to a volumetric flask containing aerated seawater, which was fed daily with commercial brine shrimp nauplii (Great Salt Lake Artemia; Ogden, UT). After one month, 25 juvenile jellyfish (0.5 cm) were collected and transferred to a 15-liter acrylic Kreisel tank with continuous water flow, maintaining 35 ppm salinity and 7.5 mg L -1 dissolved oxygen (DO) with water exchanges every 25 days (see supplementary material). The jellyfish were fed daily to satiety with brine shrimp nauplii. We observed that the umbrella of one jellyfish exhibited noticeable thickening, which was noticeable to the naked eye. The organism was gently collected in a Petri dish and observed using an stereomicroscope (mod. SZ60; Olympus) equipped with a 51 MP FHD camera (V6; Amszoom) mounted directly in the ocular tube. Morphological changes were recorded every 2 days; similar observations were made for other jellyfish 4 days after the initial observation. All individuals exhibiting morphological changes were documented and measured for umbrella diameter and total length. Results The jellyfish grew to a total length of 3.5 ± 0.5 cm within three months, with an umbrella diameter of 1.8 ± 0.5 cm. Five juvenile jellyfish exhibited dense tissue growth in the margins of their umbrellas. In two cases, the organisms displayed thicker tissue and slight outgrowths in the ectoderm (Fig. 1). In one of the jellyfish, the ectoderm protrusions disappeared once the organism reached a length of 2 cm, and it continued to grow normally. In two other organisms, a different type of abnormal tissue was observed in the umbrella, causing it to stop growing. The edges of the abnormal tissue formed a colony of developing polyps, as clearly shown in Fig. 2. Although an amorphous mass was observed, individual mouths and groups of tentacles able to capture and ingest brine shrimp nauplii could be identified. After a month of development, the polyp colony exhibited its first collective strobilation process, during which at least three polyps released ephyrae (Fig. 3a). However, these initial ephyrae were stuck together, and were not able to survive for more than a couple days. Once the strobilation process was completed, the recovery of the polyp colony allowed for the separation into two distinct polyps, which underwent a new strobilation process after two weeks (Fig. 3b). The jellyfish ceased to grow, and after the second strobilation process, it lost its oral arms, and the bell decreased in size. One of the polyps was released and successfully settled on a glass substrate (Fig. 4). In contrast, the other polyps remained with the unreleased ephyra until the umbrella was reduced to the point of disappearing. The total duration from the transformation to the settling of the new polyp was 10 days. Initially, the polyp had excess tissue (Figure 4a); however, after settling, it underwent transformation and reabsorption, ultimately developing into a polyp with a mouth and short tentacles (Figure 4b). A week later, the polyp continued to develop, and its tentacles became relaxed and elongated (Figure 4c). More than a week later, a monodisk strobilation process occurred, with the ephyra free and developing into a typical juvenile medusa (Figure 5). After two weeks following the first strobilation, a new process occurred with polydisk strobilation, resulting in at least eight free ephyrae (Fig. 5). The life cycle, including this new form of asexual reproduction, is illustrated schematically in Figure 6. We currently have a healthy colony, which has grown under lab conditions without induction, reaching over 30 polyps, all asexually derived from the single settled polyp described above. This asexual polyp generation from the medusa’s umbrella was observed at least one more time in a separate cohort of juvenile jellyfish derived from the same initial population of polyps. In this second observation, five of over a hundred juveniles showed at least one polyp at day 20. Another important observation is that, in the new polyp colony derived from medusa tissue, 15% of the medusae generated through the normal strobilation process also produced polyps on the umbrella in the following month, before reaching 5 cm of diameter. Discussion Reversion of the life cycle in Stomolopus sp.2 In this report, we describe for the first time the direct generation of a new colony of polyps in the umbrella of Stomolophus sp. 2 , a phenomenon that has not been documented previously. A significant finding is the development and growth of colonies of polyps in the umbrella of Stomolophus sp. 2 , ocurring without any apparent induction of stress. The life cycle of this species primarily involves multiple asexual reproductions from polyps and sexual reproduction in the medusa stage (López-Martínez et al., 2022). The discovery and redundant observation of a new process of asexual reproduction from the medusa stage indicate that this may be common, though not predominant, for this species. This could enhance their capacity to invade new areas, as jellyfish are carried by ocean and coastal currents. As a result, the emergence and eventual establishment of polyp populations could occur in different locations through this mechanism, rather than relying solely on external sexual reproduction. Additionally, this study observed a tissue repair process was observed, which has not been previously reported for this species. Tissue recovery has been noted in other jellyfish, associated with the restoration of "symmetrization" and body part reorganization (Lewis, 2018; Gamero-Mora et al., 2019). The ability to regenerate tissue is rare in nature, and understanding this process could have numerous applications for humans, from its use in biomedical research to comercial cosmetology. In this sense, the cannonball jellyfish could serve as a new model organism for such research. In many jellyfish species, the life cycle is very complex, with asexual reproduction primarly occurring during the polyp stage. This can lead to the formation of many new polyps within a short period, often linked to rising temperatures (Fernández-Alías et al., 2020). Although the rates of increase are not well-defined, the presence of polyps in the medusa stage has only been observed in Aurelia sp. 1 (He et al., 2015). In another jellyfish species, Turritopsis dohrnii , commonly known as the "immortal" jellyfish, the processes of reversion and rejuvenation has been extensively documented. The medusa stage can revert into a polyp form under unfavorable conditions, subsequently settling to form a new colony (Piraino et al., 1996). Exploring converging ontogenetic paths in jellyfish that lead to polyp generation could provide new perspectives on ecological and evolutionary processes in the animal kingdom. Recent studies on the genome and transcriptome of Turritopsis dohrnii have identified specific genes associated with development and rejuvenation. It has been shown that the cyst stage is crucial for these processes, and genes related to telomerase maintainence, repair mechanisms, mitotic cell division, and cellular differentiation are transcriptionally active at both the medusa and polyp stages (Matsumoto et al., 2019). Similar processes have been identified in Aurelia sp. 1 (Fuch et al., 2014), and this may also be the case in Stomolophus sp. 2 . Further studies on their genome and transcriptome are warranted; epigenetic detection through whole-genome bisulfite sequencing (WGBS) could provide insights into the mechanisms controlling shifts in the life cycle, and how environmental stressors may induce this life stage shortcuts. Furthermore, in nature, evolutionary processes are influenced by several factors, including mutations, gene flow, and inbreeding. In this context, species that reproduce only asexually have less adaptive potential than those that reproduce sexually (Orive et al., 2017). This is due to the irreversible accumulation of deleterious mutations, which can lead to genome degradation and reduced fitness, limiting the efficiency of natural selection (Otto, 2021). However, this is primarly relevant for species models with only one mode of reproduction; in species with both modes, the mechanisms are combined. DNA recombination through sexual reproduction generates new gene combinations that are tested in new local environments. Asexual reproduction, contrarly, allows fittest individuals to reproduce as clones, preserving and enabling the spread of those gene combinations across the population (de Meeûs et al., 2007). In jellyfish, the complex life cycle and occurrence of two types of reproduction may enhance the evolutionary advantage and ability to adapt to changing environments, along with their dispersal capabilities. Implications in dispersal and invasiveness potential One of the remarkable characteristics of jellyfish is their considerable plasticity in development and rapid adaptive responses to spatial and temporal environmental fluctuations. This allows them to colonize a wide variety of habitats (Thein et al., 2013). The ability of jellyfish polyps to prevail in multiple adverse environmental conditions, together with their rapid asexual reproductive processes, enables them to multiply in large numbers efficiently (Arai, 2009; Lee et al., 2018). Our findings on polyp generation in sexually immature medusae reveal novel mechanisms of polyp dispersion, which provide a significant advantage over other potential invaders. Polyps are resilient life stages of the species that can proliferate over short distances, even under adverse conditions faced by the free-swimming juvenile and adult stages. The ability to generate several free-swimming juvenile means that a relatively small population of polyps can act as a “population pool” with highly multiplicative potential when favorable conditions arise, all without requiring mature organisms or sexual reproduction. When medusae are carried by ocean currents or ballast water, free-floating polyps can be transported over long distances, potentially establishing new populations in regions where environmental conditions are favorable. In projected scenarios of climate change, the abundance of jellyfish populations is likely to increase. Moreover, factors such as eutrophication in new areas and overfishing, which reduce the number of natural predators, further increase the likelihood of blooms forming or persisting (Goldstein et al., 2020). The appearance of jellyfish in new regions and subsequent blooms highlight the extensive dispersal and invasive potential of these animals. Species such as Aurelia spp. and Nemopilema nomurai in Japan have caused significant ecological problems (Kawahara et al., 2006; Feng et al., 2024). Many studies on blooms focus on population dynamics and damage control, while others examinate the ecophysiology of polyps to understand environmental factors triggering significant proliferations (Zang et al., 2022). In another group of cnidarians, the hydrozoan Eudendrium carneum is recognized as an invasive species with high potential for global spread. It is commonly observed in harbors, yacht clubs, and in fouling communities (Gónzalez-Duarte et al., 2016). Its life cycle, though primarily sessile, allows for reproduction in both sexual and free-living stages, allowing its expansion into new areas (Puce et al., 2005). Other species like Phyllorhiza punctata , Cassiopea andromeda , Rhopilema nomadica , and Aurelia spp . have been reported to cover extensive areas in short time frames, leading to their classification as invasive species, usually related to maritime transport rather than biological dispersal mechanisms (González-Duarte et al., 2016). Currently, predictions about jellyfish blooms and dispersal remain uncertain due to unclear biological processes and environmental factors, coupled with ocean degradation that may be favorable to jellyfish growth (Fernández-Alías et al., 2024). The implications for jellyfish life cycle and bloom formation are significant, offering new insights into jellyfish population dispersion and increase, particularly in the absence of sexual reproduction or maritime shipping as part of the fouling community. Molecular mechanisms underlying the reversed cycle Another perspective from this reversal mechanism is related to tissue regeneration and cellular differentiation. In the hydrozoan Turritopsis nutricula , the differentiation of polyps from adult tissue involves a cellular transdifferentiation process occuring in both epidermal and gastrointestinal differentiated cells (Pirano et al., 1996). More recently, in the jellyfish Podocoryne carnea , it was found that muscle tissue cells can change from striated to smooth or even differentiate into functional neurons. This transdifferentiation is linked to differential expression of the Piwi-like gene, which is associated with stem cells in multicellular organisms (Seipel et al., 2004). This suggests that these organisms are cappable of reprogramming cells within the same tissue. In the moon jellyfish Aurelia sp. 1 , a reversion cycle from medusa to polyp has been reported, the development of polyps within the umbrella (He et al., 2015). However, the specific genes and cellular mechanisms involved in this reversal process have yet to be elucidated. While differential gene expressions at various life stages does not necessarily indicate a regeneration process or pluripotentiality, several Wnt genes are found to be overexpressed in the planula, strobila, and medusa stages (Brekhman et al., 2015). These genes are implicated in embryogenesis, proliferation, and cell migration (Miller, 2002). Most importantly, this gene family is involved in maintaining the undifferentiated state of stem cells and regulating transcriptional signaling for proliferation and differentiation (Nusee, 2008). Although the presence of these genes does not fully indicate the activity of undifferentiated cells capable of initiating cycle reversal processes, it suggests that these organisms may possess the capacity to perform such reversals at certain life stages. Most of the provided discussion related to the distribution of the polyps in the wild contain a significant speculative component, mainly because this life stage remains cryptic and elusive to ecological research. This is true not only for the cannonball jellyfish but for most of the species that alternate between asexual and sexual reproduction. Therefore, the development of laboratory models, along with advances in field research tools such as environmental DNA and miniaturized underwater drones, would help to understand the implications of the ability to propagate as medusae-derived polyps both seasonally and geographically. Finally, the significance of our findings shows the need to explore into different cellular and physiological mechanisms in these groups of organisms. Given that jellyfish have a long evolutionary history, it would be interesting to understand the adaptations that have helped them survive and invade new environments, their multiple ways of reproducing are certainly one of them. Declarations Acknowledgements CAN-L thanks to the SECIHTI (Ministry of Science, Humanities, Technology, and Innovation; Government of Mexico) for the postdoctorate fellowship (CVU 413245). We are grateful with Beatriz Yañez Rivera for her kind assistance on image documentation. Author contributions Conceived and designed the experiments: NLC LHR Performed the experiments: NLC. Analyzed the data: NLC. Contributed materials/analysis tools: LHR JLM. Wrote the manuscript: NLC LHR JLM. All the authors revised and approved the final version of the manuscript Funding This work was mainly funded by internal grants provided by IMCyL-UNAM. Conflict of interest: The authors have no conflicts of interest to declare. Ethics approval The rearing and lab culture of polyps and jellyfish do not require research ethics approval at the institution where these studies were conducted. Consent to participate: Not applicable. Consent for publication: Not applicable. References Aouititen M, Ravibhanu A, Ang SC, Magabanda-Mouanda DC, Luan X (2024) New records of two jellyfish species Rhizostoma luteum (Quoy and Gaimard 1827) and Cotylorhiza tuberculata (Macri 1778) in the Moroccan northwest Mediterranean coast. Discov Life 54(5). https://doi.org/10.1007/s11084-024-09649–2 Arai MN (2009) The potential importance of podocysts to the formation of scyphozoan blooms: a review. Hydrobiologia 616:241–246. http://doi.org/10.1007/s10750-008-9588–5 Brekhman V, Malik A, Haas B, Sher N, Lotan T (2015) Transcriptome profiling of the dynamic life cycle of the scyphozoan jellyfish Aurelia aurita. BMC Genomics 16:74. https://doi.org/10.1186/s12864-015-1320-z Brotz L, Cisneros-Montemayor AM, Cisneros-Mata MA (2021) The race for jellyfish: Winners and losers in Mexico’s Gulf of California. Mar Policy 134:104775. https://doi.org/10.1016/j.marpol.2021.104775 Chuan CH, Venmathi-Maran BA, Yap TK, Cheong KC, Syed-Hussein MA, Saleh E (2021) New Records of Cubozoan and Scyphozoan Jellyfish from Sabah Waters, Malaysia. Diversity 13(9):420. https://doi.org/10.3390/d13090420 Feng S, Xu X, Wan A, Sun S (2024) Differences in. Nemopilema nomurai colony Prolif biofouling invasion Var situ polyp densities Jiaozhou Bay China J Exp Mar Biol Ecol 574:152006. https://doi.org/10.1016/j.jembe.2024.152006 Fernández-Alías A, Concepción M, Quispe JI, Sabah S, Pérez-Ruzafa A (2020) Population dynamics and growth in three scyphozoan jellyfishes, and their relationship with environmental conditions in a coastal lagoon. Estuar Coast Shelf Sci 243:106901. https://doi.org/10.1016/J.ECSS.2020.106901 Fernández-Alías A, Marcos C, Pérez-Ruzafa A (2021) Larger scyphozoan species dwelling in temperate, shallow waters show higher blooming potential. Mar Pollut Bull 173(B):113100. https://doi.org/10.1016/j.marpolbul.2021.113100 Fernández-Alías A, Marcos C, Pérez-Ruzafa A (2024) The unpredictability of scyphozoan jellyfish blooms. Front Mar Sci 11:1349956. https://doi.org/10.3389/fmars.2024.1349956 Fonfría ES, Straehler-Pohl I, Stampar SN, Collins AG, Hoverd WA, Bordehore C (2024) Copula lucentia sp. nov., a new box jellyfish (Cnidaria: Cubozoa: Carybdeida) from Western Mediterranean Sea. Mar Biodivers 54:68. 10.1007/s12526-024-01460–6 Fuchs B, Wang W, Graspeuntner S, Li Y, Insua S, Herbst EM, Dirksen P, Böhm AM, Hemmrich G, Sommer F, Domazet-Lošo T, Klostermeier UC, Anton-Erxleben F, Rosenstiel P, Bosch TC, Khalturin K (2014) Regulation of polyp-to-jellyfish transition in Aurelia aurita. Curr Biol 24(3):263–273. https://doi.org/10.1016/j.cub.2013.12.003 Gamero-Mora E, Halbauer R, Bartsch V, Stampar SN, Morandini AC (2019) Regenerative Capacity of the Upside-down Jellyfish Cassiopea xamachana. Zool Stud 58:e37. https://doi.org/10.6620/ZS.2019.58–37 Goldstein J, Steiner UK (2020) Ecological drivers of jellyfish blooms - The complex life history of a 'well-known' medusa (Aurelia aurita). J Anim Ecol 89(3):910–920. https://doi.org/10.1111/1365–2656.13147 Gómez-Salinas LC (2018) Dinámica poblacional de la medusa Stomolophus meleagris (L. Agassiz, 1862) durante el evento El Niño 2015–2016 en la Laguna Las Guásimas, Sonora, México. Dissertation Thesis. Universidad Nacional Autónoma de México. https://hdl.handle.net/20.500.14330/TES01000769386 Gómez-Salinas LC, López-Martínez J, Morandini AC (2021) The Young Stages of the Cannonball Jellyfish (Stomolophus sp. 2) from the Central Gulf of California (Mexico). Diversity 13(6):229. https://doi.org/10.3390/d13060229 González-Carman V, Botto F, Gaitán E, Albareda D, Campagna C, Mianzan H (2014) A jellyfish diet for the herbivorous green turtle Chelonia mydas in the temperate SW Atlantic. Mar Biol 161:339–349. https://doi.org/10.1007/s00227-013-2339–9 González-Duarte MM, Megina C, López-González PJ, Galil B (2016) Cnidarian Alien Species in Expansion. In: Goff S, Zvy, Editors D (n.d.) (eds) The Cnidaria, Past, Present and Future. Springer International Publishing. http://doi.org/10.1007/978-3-319–31305–4_10 He J, Zheng L, Zhang W, Lin Y (2015) Life Cycle Reversal in Aurelia sp.1 (Cnidaria, Scyphozoa). PLoS ONE 10(12):e0145314. https://doi.org/10.1371/journal.pone.0145314 Houliston E, Leclère L, Munro C, Copley RR, Momose T (2022) Past, present and future of Clytia hemisphaerica as a laboratory jellyfish. Curr Top Dev Biol 147:121–151. https://doi.org/10.1016/bs.ctdb.2021.12.014 Kawahara M, Uye SI, Ohtsu K, Iizumi H (2006) Unusual population explosion of the giant jellyfish Nemopilema nomurai (Scyphozoa: Rhizostomeae) in East Asian waters. Mar Ecol Prog Ser 307:161–173. http://doi.org/10.3354/meps307161 Lewis-Ames C (2018) Medusa: A Review of an Ancient Cnidarian Body Form. In: Kloc M, Kubiak J (eds) Marine Organisms as Model Systems in Biology and Medicine. Results and Problems in Cell Differentiation, vol 65. Springer, Cham. https://doi.org/10.1007/978-3-319–92486–1_7 López-Martínez J, Álvarez-Tello F (2013) The jellyfish fishery in Mexico. Agri sci 4:57–61. https://doi.org/10.4236/as.2013.46A009 López-Martínez J, Álvarez-Tello FJ, Porchas-Cornejo MA, Nevárez-López CA, Muhlia-Almazán A, Urías-Padilla KV (2023) Multiple reproduction forms in the polyps of the cannonball jellyfish Stomolophus sp. 2: Probable life-cycle reversal. J Exp Zool Ecol Integr Physiol 339(3):239–252. https://doi.org/10.1002/jez.2673 Luo JY, Condon RH, Stock CA, Duarte CM, Lucas CH, Pitt KA, Cowen RK (2020) Gelatinous Zooplankton-Mediated Carbon Flows in the Global Oceans: A Data‐Driven Modeling Study. Global Biogeochem Cycles 34:e2020GB00670. https://doi.org/10.1029/2020GB006704 Matsumoto Y, Piraino S, Miglietta MP (2019) Transcriptome Characterization of Reverse Development in Turritopsis dohrnii (Hydrozoa, Cnidaria). G3. Genes Genomes Genet 9(12):4127–4138. https://doi.org/10.1534/g3.119.400487 Miller JR (2022) The Wnts. Genome Biol 3(1):REVIEWS3001. https://doi.org/10.1186/gb–2001-3-1-reviews3001 Nevárez-López C, Sánchez-Paz A, López-Martínez J, Llera-Herrera R, Muhlia-Almazán A (2020) Metabolic response of the cannonball jellyfish Stomolophus meleagris upon short-term exposure to thermal stress. J Sea Res 166:101959. https://doi.org/10.1016/j.seares.2020.101959 Nusse R (2008) Wnt signaling and stem cell control. Cell Res 18(5):523–527. https://doi.org/10.1038/cr.2008.47 Piraino S, Boero F, Aeschbach B, Schmid V (1996) Reversing the Life Cycle: Medusae Transforming into Polyps and Cell Transdifferentiation in Turritopsis nutricula (Cnidaria, Hydrozoa). Biol Bull 190(3):302–312. https://doi.org/10.2307/1543022 Pitt K, Lucas C, Condon RH, Duarte CM, Stewart-Koster B (2018) Claims That Anthropogenic Stressors Facilitate Jellyfish Blooms Have Been Amplified Beyond the Available Evidence: A Systematic Review. Front Mar Sci 5:451. https://doi.org/10.3389/fmars.2018.00451 Preciado M (2016) Pesquería artesanal de la medusa Stomolophusmeleagris en el estuario interno del Golfo de Guayaquil 2014. Master thesis. Universidad de Guayaquil. http://repositorio.ug.edu.ec/handle/redug/22212 Purcell JE, Baxter EJ, Fuentes VL (2013) Jellyfish as products and problems of aquaculture. In: Advances in Aquaculture Hatchery Technology. Eds. Allan G, Burnell G. Woodhead Publishing Series in Food Science, Technology and Nutrition, Cambridge, United Kingdom, 404–430 pp. https://doi.org/10.1533/9780857097460.2.404 Schnedler-Meyer NA, Kiørboe T, Mariani P (2018) Boom and Bust: Life History, Environmental Noise, and the (un)Predictability of Jellyfish Blooms. Front Mar Sci 5:257. https://doi.org/10.3389/fmars.2018.00257 Seipel K, Yanze N, Schmid V (2004) The germ line and somatic stem cell gene Cniwi in the jellyfish Podocoryne carnea. Int J Dev Biol 48(1):1–7. https://doi.org/10.1387/ijdb.15005568 Thein H, Ikeda H, Uye S-I (2013) Ecophysiological characteristics of podocysts in Chrysaora pacifica (Goette) and Cyanea nozakii Kishinouye (Cnidaria: Scyphozoa: Semaeostomeae): effects of environmental factors on their production, dormancy and excystment. J Exp Mar Biol Ecol 446:151–158. http://doi.org/10.1016/j.jembe.2013.05.013 Zang W, Zhang F, Chi X, Sun S (2022) Relationship Between Asexual Reproduction of Aurelia coerulea Polyps and Jellyfish Blooms Under the Influence of Temperature Dynamics in Winter and Spring. Front Mar Sci 9:888656. https://doi.org/10.3389/fmars.2022.888656 van Walraven L, Driessen F, van Bleijswijk J, Bol A, Luttikhuizen PC, Coolen JWP, Bos OG, Gittenberger A, Schrieken N, Langenberg VT, van der Veer HW (2016) Where are the polyps? Molecular identification, distribution and population differentiation of Aurelia aurita jellyfish polyps in the southern North Sea area. Mar Biol 163:172. https://doi.org/10.1007/s00227-016-2945–4 Supplementary Files supplemantaryreversion.tif Cite Share Download PDF Status: Under Review Version 1 posted Editorial decision: Revise and Resubmit 24 Oct, 2025 Reviewers agreed at journal 12 Sep, 2025 Reviewers invited by journal 12 Sep, 2025 Editor assigned by journal 11 Sep, 2025 First submitted to journal 09 Sep, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-7579160","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":514214718,"identity":"2ef4468a-d54c-4b58-ad2f-24b5a11bc02e","order_by":0,"name":"Cintya Alejandra Nevárez-López","email":"","orcid":"","institution":"Instituto de Ciencias del Mar y Limnología: Universidad Nacional Autonoma de Mexico Instituto de Ciencias del Mar y Limnologia","correspondingAuthor":false,"prefix":"","firstName":"Cintya","middleName":"Alejandra","lastName":"Nevárez-López","suffix":""},{"id":514214719,"identity":"f018372b-33f3-487c-b215-09247c0767d4","order_by":1,"name":"Raul Llera-Herrera","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABDElEQVRIiWNgGAWjYHACxgMMBRZAmvkAwwMGBh6ICAFwgMFAAkixJTAkQLQwEKuFxwCkhYGgFvP2MwYgLXJAxscHCTUMMvzTgCKMbXfkdRu40ySwaJE5kwPWYixzJnezQcIxBh6J2zkgLc8Mtx3g3WyARYsEA0RL4gyG3G0SCWxA50mDtRxmBGrZ+ACbFv43YC31M/jfPJNI+IfQYg/UsgGbpyQkILYkABlsEoltCC2JOG2ReFZwIMFAwnCGxDNjg8Q+CaBf0oAi5w4nbzuMwy/8yRsffKiwkQcyHj748M3Gnn82SKTssO22473bsIUYGCQgBwcDAwc0gphxqccE7Nh8MApGwSgYBSMYAAAfL2MIq9qpoQAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0000-0001-9873-9732","institution":"UNAM ICMYL: Universidad Nacional Autonoma de Mexico Instituto de Ciencias del Mar y Limnologia","correspondingAuthor":true,"prefix":"","firstName":"Raul","middleName":"","lastName":"Llera-Herrera","suffix":""},{"id":514214720,"identity":"1252ef0b-9222-4db0-926e-a6f887a61ec4","order_by":2,"name":"Juana López-Martínez","email":"","orcid":"","institution":"CIBNOR: Centro de Investigaciones Biologicas del Noroeste SC","correspondingAuthor":false,"prefix":"","firstName":"Juana","middleName":"","lastName":"López-Martínez","suffix":""}],"badges":[],"createdAt":"2025-09-10 06:00:38","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7579160/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7579160/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":91724038,"identity":"833011ab-346c-4e54-9934-c622bb59be2e","added_by":"auto","created_at":"2025-09-19 14:46:54","extension":"tif","order_by":0,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":2345837,"visible":true,"origin":"","legend":"","description":"","filename":"Fig1.tif","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/b08f031ca04760328b6ce2c6.tif"},{"id":91724039,"identity":"1ca7a926-1850-4484-9529-4c920846cca3","added_by":"auto","created_at":"2025-09-19 14:46:54","extension":"tif","order_by":2,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1928557,"visible":true,"origin":"","legend":"","description":"","filename":"Fig2.tif","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/06014dc1319abe406bcf5ed2.tif"},{"id":91723627,"identity":"1b33fa5d-4454-4331-9144-a2686300fb64","added_by":"auto","created_at":"2025-09-19 14:38:54","extension":"tif","order_by":3,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":402681,"visible":true,"origin":"","legend":"","description":"","filename":"Fig4.tif","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/1cb329bbd55942690178412a.tif"},{"id":91723623,"identity":"dc1335bf-5736-440c-a114-e8ab1be3f35c","added_by":"auto","created_at":"2025-09-19 14:38:54","extension":"tif","order_by":4,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":6825961,"visible":true,"origin":"","legend":"","description":"","filename":"Fig5.tif","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/383d91c4cc7226d6d98bb923.tif"},{"id":91724036,"identity":"a29fa555-ccd4-4d81-808b-231a399d2d78","added_by":"auto","created_at":"2025-09-19 14:46:53","extension":"tif","order_by":6,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1726251,"visible":true,"origin":"","legend":"","description":"","filename":"Fig3.tif","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/0347ada0c4e25e1ee27b60b3.tif"},{"id":91723548,"identity":"a962d499-7a97-42b7-9542-4aa172ed4090","added_by":"auto","created_at":"2025-09-19 14:38:51","extension":"xml","order_by":7,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":7577,"visible":true,"origin":"","legend":"","description":"","filename":"mabiMABID2500465.xml","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/df564374e1a1ae913fdd68c6.xml"},{"id":91723538,"identity":"832ba841-0eb9-44ab-9c90-771b6efd89e0","added_by":"auto","created_at":"2025-09-19 14:38:50","extension":"xml","order_by":8,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1176,"visible":true,"origin":"","legend":"","description":"","filename":"MABID250046514248.go.xml","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/c7f8435a8c7c386dbdc428a4.xml"},{"id":91723635,"identity":"33d51e13-a21d-4254-a1bc-9ddf20657455","added_by":"auto","created_at":"2025-09-19 14:38:55","extension":"xml","order_by":9,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":758,"visible":true,"origin":"","legend":"","description":"","filename":"MABID2500465Import.xml","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/cf24de7b19694bce0240bfe0.xml"},{"id":91723630,"identity":"5b4e8851-eb8f-4821-82d7-89f25828a997","added_by":"auto","created_at":"2025-09-19 14:38:54","extension":"xml","order_by":11,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":99262,"visible":true,"origin":"","legend":"","description":"","filename":"MABID25004650enriched.xml","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/eb3c55ca7193fd396fbd865a.xml"},{"id":91723613,"identity":"199820bc-b7fd-4a8f-86d0-b59405e835b8","added_by":"auto","created_at":"2025-09-19 14:38:53","extension":"tif","order_by":12,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":2345837,"visible":true,"origin":"","legend":"","description":"","filename":"Fig1.tif","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/ba55ff4bf0380b9d8bb07c48.tif"},{"id":91723546,"identity":"07615732-3ee4-4137-bceb-471089070f5d","added_by":"auto","created_at":"2025-09-19 14:38:51","extension":"tif","order_by":13,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1928557,"visible":true,"origin":"","legend":"","description":"","filename":"Fig2.tif","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/4e5f7b48ebeea47b8729d60e.tif"},{"id":91723624,"identity":"549ca715-283c-4487-bc02-4ee470fa3ddf","added_by":"auto","created_at":"2025-09-19 14:38:54","extension":"tif","order_by":14,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":402681,"visible":true,"origin":"","legend":"","description":"","filename":"Fig4.tif","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/e734a06ccc5d173a53a7850f.tif"},{"id":91723637,"identity":"7794612b-8ebb-4a18-a5b9-02f1f3fa8bfa","added_by":"auto","created_at":"2025-09-19 14:38:55","extension":"tif","order_by":15,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":6825961,"visible":true,"origin":"","legend":"","description":"","filename":"Fig5.tif","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/5864894cd9519a25234cdba9.tif"},{"id":91723617,"identity":"38882be2-670d-4b9e-a60e-26a415964ddc","added_by":"auto","created_at":"2025-09-19 14:38:53","extension":"eps","order_by":16,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":333625,"visible":true,"origin":"","legend":"","description":"","filename":"Fig6.eps","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/1ca1930d88cdd7603fefc8a0.eps"},{"id":91723619,"identity":"c698b1f8-bdb4-407a-87b3-289ede1724be","added_by":"auto","created_at":"2025-09-19 14:38:54","extension":"tif","order_by":17,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1726251,"visible":true,"origin":"","legend":"","description":"","filename":"Fig3.tif","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/3459d757533a6c6111420108.tif"},{"id":91723618,"identity":"e500d471-3341-489a-ac4c-34442c0bdd55","added_by":"auto","created_at":"2025-09-19 14:38:54","extension":"png","order_by":18,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":222457,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineFig1.png","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/6c9d66cbe5172c5a36e3087b.png"},{"id":91723620,"identity":"fa7233a4-c3cb-4358-a739-03fc0950f24c","added_by":"auto","created_at":"2025-09-19 14:38:54","extension":"png","order_by":19,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":170997,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineFig2.png","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/16c9a1e3d1ce74fdabc01808.png"},{"id":91723634,"identity":"d2af1144-6dd8-4f68-ae64-20cab416b925","added_by":"auto","created_at":"2025-09-19 14:38:55","extension":"png","order_by":20,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":44358,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineFig4.png","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/d5a445722bdb0baca4d07cde.png"},{"id":91723632,"identity":"7ddd8417-63c7-4b29-b4f1-4271c86c22c8","added_by":"auto","created_at":"2025-09-19 14:38:54","extension":"png","order_by":21,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":430421,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineFig5.png","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/f7bcd2a4e80e1f0d1a1fffc2.png"},{"id":91723631,"identity":"68ca111d-8014-4b92-9670-fcb51a2d3341","added_by":"auto","created_at":"2025-09-19 14:38:54","extension":"png","order_by":22,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":168426,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineFig3.png","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/0b2606aa51f155c456cfa375.png"},{"id":91723615,"identity":"891b6249-ab7d-428d-95c5-8e98e891099d","added_by":"auto","created_at":"2025-09-19 14:38:53","extension":"xml","order_by":23,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":97624,"visible":true,"origin":"","legend":"","description":"","filename":"MABID25004650structuring.xml","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/3dab369faa8f7f65993a74a4.xml"},{"id":91723628,"identity":"32f2c6f4-c8a3-45d4-9f66-4264a5c7617e","added_by":"auto","created_at":"2025-09-19 14:38:54","extension":"html","order_by":24,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":108783,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/b5b23050264206211be58280.html"},{"id":91723629,"identity":"38b59a83-1ee5-4ab6-b115-ef80d242d7b2","added_by":"auto","created_at":"2025-09-19 14:38:54","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":2063880,"visible":true,"origin":"","legend":"\u003cp\u003eAnomalous tissue in the umbrella of \u003cem\u003eStomolophus sp. 2\u003c/em\u003e. A. Complete organisms with different types of abnormal tissue, B. zoomed-in image of abnormal tissue, C. zoomed-in image of small bumps. Scale bar= 1mm.\u003c/p\u003e","description":"","filename":"Fig12.png","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/68ea7d8b1149fcaa60662dec.png"},{"id":91723625,"identity":"77671997-47a8-4ffa-aab0-4e4ad4c57889","added_by":"auto","created_at":"2025-09-19 14:38:54","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":1728643,"visible":true,"origin":"","legend":"\u003cp\u003eColony of polyps in the umbrella of \u003cem\u003eStomolophus sp. 2\u003c/em\u003e. A. Complete organisms with polyps in the umbrella, B. Colony of polyps. Scale bar= 1mm.\u003c/p\u003e","description":"","filename":"Fig21.png","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/82b24a0b43b0e386b68d4b47.png"},{"id":91724037,"identity":"362aa373-708b-41e1-a454-5be27a178f5b","added_by":"auto","created_at":"2025-09-19 14:46:53","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":1534191,"visible":true,"origin":"","legend":"\u003cp\u003eColony of polyps undergoing the strobilation process in the umbrella of \u003cem\u003eStomolophus sp. 2\u003c/em\u003e. A. First strobilation process, B. Second strobilation process. Ephyrae is denoted with arrowhead, and polydisk segmentation is denoted by dashed lines. Scale bar= 1mm.\u003c/p\u003e","description":"","filename":"Fig31.png","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/0dffb7b31dbd4ba8a7ac2158.png"},{"id":91723585,"identity":"38fed1c0-6979-4edf-8824-1709ba1c2d13","added_by":"auto","created_at":"2025-09-19 14:38:52","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":356192,"visible":true,"origin":"","legend":"\u003cp\u003eNew polyp of jellyfish \u003cem\u003eStomolophus\u003c/em\u003e sp. 2. A. Released polyp, B. Settled polyp, C. Developed polyp. Scale bar A=1 mm, B,C=0.5 mm.\u003c/p\u003e","description":"","filename":"Fig4.png","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/6bccefbcbaf683755862c7d2.png"},{"id":91723633,"identity":"2be9c56e-ac72-45fb-beac-f47d02ba6fcf","added_by":"auto","created_at":"2025-09-19 14:38:55","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":5725799,"visible":true,"origin":"","legend":"\u003cp\u003eThe asexual cycle from a new polyp developing into medusa. A. Polyp, B. Early ephyra, C. Metephyra, D. Metephyra with oral arms, E. Early juvenile medusae. Scale bar A-D=0.5 mm, E=0.25 mm.\u003c/p\u003e","description":"","filename":"Fig5.png","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/759e15991742e25a869fb834.png"},{"id":91723554,"identity":"8c6b501b-7c3f-476e-a159-b7bca4b85867","added_by":"auto","created_at":"2025-09-19 14:38:52","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":138596,"visible":true,"origin":"","legend":"\u003cp\u003eSchematic life cycle of \u003cem\u003eStomolophus\u003c/em\u003e sp.2. Normal development stages were presented with black illustrations and arrows. In contrast, modifications of the typical life cycle described in this study are depicted in red (modification of life cycle, as per López-Martínez et al., 2022).\u003c/p\u003e","description":"","filename":"Fig6.png","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/ce5151aa895dce968a1bb790.png"},{"id":91724040,"identity":"2743da5a-06e8-408f-9dbe-d35c74470213","added_by":"auto","created_at":"2025-09-19 14:47:05","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":16054200,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/fa961521-f821-409b-a2ed-db741826026f.pdf"},{"id":91723612,"identity":"cc26267f-17ac-43e6-90fb-2455ab6649cf","added_by":"auto","created_at":"2025-09-19 14:38:52","extension":"tif","order_by":10,"title":"","display":"","copyAsset":false,"role":"supplement","size":3430440,"visible":true,"origin":"","legend":"","description":"","filename":"supplemantaryreversion.tif","url":"https://assets-eu.researchsquare.com/files/rs-7579160/v1/4ad142dbddb8bc3da479b6bd.tif"}],"financialInterests":"","formattedTitle":"Evidence of asexual polyp development from the adult body in Stomolopus sp2","fulltext":[{"header":"Introduction","content":"\u003cp\u003eCnidarians are a complex and diverse group of invertebrates that include corals, anemones, hydras, and jellyfish. In some regions, the abundance of jellyfish has increased over the past decade, and certain species have expanded their distribution ranges (Aouititen et al., 2024; Fonfr\u0026iacute;a et al., 2024; Chuan et al., 2024). Besides seasonal fluctuations, several factors have contributed to the dramatic global increase in jellyfish populations over the last two decades, such as eutrophication, overfishing, coastal development, and species translocation (Pitt et al., 2018; Schnedler-Meyer et al., 2018; Fern\u0026aacute;ndez-Al\u0026iacute;as et al., 2020). This increase in jellyfish populations poses several challenges in coastal areas, such as blockages in pipeline intakes for thermoelectric power generation and skin injuries for swimmers. Additionally, these animals are voracious predators of fish eggs and planktonic larvae, leading to strong competition for food and predation pressure on fish populations when jellyfish proliferate excessively (Purcell et al., 2013; Fern\u0026aacute;ndez-Al\u0026iacute;as et al., 2024).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eDespite these challenges, jellyfish play a crucial ecological role. They contribute between 3.7 and 6.8 billion metric tons of carbon biomass each year (Luo et al., 2020) and serve as food for sea turtles and habitat for other species (Gonz\u0026aacute;lez-Carmian et al., 2014).\u003c/p\u003e\n\u003cp\u003eThe cannonball jellyfish (\u003cem\u003eStomolophus\u0026nbsp;\u003c/em\u003espp.) is found in the Mexican Pacific and is commercially exploited. In recent years, massive blooms of up to 60 org/m\u003csup\u003e2\u003c/sup\u003e have been recorded (L\u0026oacute;pez-Mart\u0026iacute;nez and \u0026Aacute;lvarez-Tello, 2013). Its distribution extends from the northern part of Sonora in the Upper Gulf of California region (Brotz et al., 2021) to Ecuador (Preciado, 2016). More recently, it has also been reported in U.S. waters, suggesting a potential northward shifts in its distribution due to climatic conditions like El Ni\u0026ntilde;o, which enhance its reproductive capacity and ability to colonize new areas (G\u0026oacute;mez-Salinas, 2018; Nev\u0026aacute;rez-L\u0026oacute;pez et al., 2020; G\u0026oacute;mez-Salinas et al., 2021).\u003c/p\u003e\n\u003cp\u003eDespite extensive evidence of increased jellyfish abundances, the strategies employed by these animals to boost their populations are not yet fully understood. However, their complex metagenic life cycle may hold some insight. This species reaches sexual maturity at a length of 10 cm (Carvalho-Saucedo et al., 2011). It is dioecious, with external fertilization, resulting in planula larvae that settle on a substrate after approximately 48 hours if optimal conditions are met. Following settlement, the larvae undergo a series of transformations, and a polyp becomes visible after 5 to 7 days (Carvalho-Saucedo et al., 2011). There are several variations of asexual reproduction mechanisms, with strobilation being the most common, producing ephyrae. Under laboratory conditions, polyps that undergo strobilation can produce up to 20-25 ephyrae that develop into free-swimming jellyfish (L\u0026oacute;pez-Mart\u0026iacute;nez et al., 2022). In their sessile stage, the polyps can also produce additional polyps and podocytes, which are smaller forms that can develop into new polyps. This variety of asexual reproduction strategies allows these animals to significantly increase their populations while in the sessile stage; with asexual reproduction potentially continuing for years before strobilation. We have successfully been able to maintain asexually reproducing polyps for at least 8 years under controlled laboratory conditions.\u003c/p\u003e\n\u003cp\u003eIn various jellyfish species, the life cycle is similar. For example, both the moon jellyfish (\u003cem\u003eAurelia sp.1\u003c/em\u003e) and the immortal jellyfish (\u003cem\u003eTurritopsis dohrnii\u003c/em\u003e) can produce new polyps without sexual reproduction or the generation of planula larvae, and this occurs independently of any specific stress process (He et al., 2015; Matsumoto et al., 2019). While the life cycle of the cannonball jellyfish exhibits a reversal mechanism, only ephyrae have been observed reverting to polyps (L\u0026oacute;pez-Mart\u0026iacute;nez et al., 2022). The existence of a broader range of asexual reproductive strategies may have significant ecological and fisheries management implications, as well as open up novel trends in biomedical and ecological research. In this context, we report a novel strategy for generating asexual polyps directly from juvenile jellyfish.\u003c/p\u003e"},{"header":"Methods","content":"\u003cp\u003eThe polyps of \u003cem\u003eStomolophus sp.2\u003c/em\u003e polyps derived from a laboratory colony reported in L\u0026oacute;pez-Mart\u0026iacute;nez et al (2022) and transported in marine water to ICMyL-UNAM Mazatl\u0026aacute;n (Sinaloa, Mexico) in 2022. They settled on translucent acrylic plates (3 cm x 5 cm) in a horizontal position in polycarbonate containers half-filled with 4 liters of filtered marine water without aeration for 7 days at room temperature (25\u0026deg;C). Once the organisms were firmly attached, the acrylic plates were vertically immersed in 8-L tanks (see supplementary material) with gentle aeration and were fed every 3 days with brine shrimp nauplii. Colony growth and strobilation were induced by increasing the temperature to 28\u0026deg;C.\u003c/p\u003e\n\u003cp\u003eThe new polyps grew on the acrylic plate, and all ephyrae were collected and transferred to a volumetric flask containing aerated seawater, which was fed daily with commercial brine shrimp nauplii (Great Salt Lake Artemia; Ogden, UT). After one month, 25 juvenile jellyfish (0.5 cm) were collected and transferred to a 15-liter acrylic Kreisel tank with continuous water flow, maintaining 35 ppm salinity and 7.5 mg L\u003csup\u003e-1\u003c/sup\u003e dissolved oxygen (DO) with water exchanges every 25 days (see supplementary material). The jellyfish were fed daily to satiety with brine shrimp nauplii.\u003c/p\u003e\n\u003cp\u003eWe observed that the umbrella of one jellyfish exhibited noticeable thickening, which was noticeable to the naked eye. The organism was gently collected in a Petri dish and observed using an stereomicroscope (mod. SZ60; Olympus) equipped with a 51 MP FHD camera (V6; Amszoom) mounted directly in the ocular tube. Morphological changes were recorded every 2 days; similar observations were made for other jellyfish 4 days after the initial observation. All individuals exhibiting morphological changes were documented and measured for umbrella diameter and total length.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003eThe jellyfish grew to a total length of 3.5 \u0026plusmn; 0.5 cm within three months, with an umbrella diameter of 1.8 \u0026plusmn; 0.5 cm. Five juvenile jellyfish exhibited dense tissue growth in the margins of their umbrellas. In two cases, the organisms displayed thicker tissue and slight outgrowths in the ectoderm (Fig. 1).\u003c/p\u003e\n\u003cp\u003eIn one of the jellyfish, the ectoderm protrusions disappeared once the organism reached a length of 2 cm, and it continued to grow normally. In two other organisms, a different type of abnormal tissue was observed in the umbrella, causing it to stop growing. The edges of the abnormal tissue formed a colony of developing polyps, as clearly shown in Fig. 2.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eAlthough an amorphous mass was observed, individual mouths and groups of tentacles able to capture and ingest brine shrimp nauplii could be identified. After a month of development, the polyp colony exhibited its first collective strobilation process, during which at least three polyps released ephyrae (Fig. 3a). However, these initial ephyrae were stuck together, and were not able to survive for more than a couple days. Once the strobilation process was completed, the recovery of the polyp colony allowed for the separation into two distinct polyps, which underwent a new strobilation process after two weeks (Fig. 3b). The jellyfish ceased to grow, and after the second strobilation process, it lost its oral arms, and the bell decreased in size. One of the polyps was released and successfully settled on a glass substrate (Fig. 4). In contrast, the other polyps remained with the unreleased ephyra until the umbrella was reduced to the point of disappearing.\u003c/p\u003e\n\u003cp\u003eThe total duration from the transformation to the settling of the new polyp was 10 days. Initially, the polyp had excess tissue (Figure 4a); however, after settling, it underwent transformation and reabsorption, ultimately developing into a polyp with a mouth and short tentacles (Figure 4b). A week later, the polyp continued to develop, and its tentacles became relaxed and elongated (Figure 4c). More than a week later, a monodisk strobilation process occurred, with the ephyra free and developing into a typical juvenile medusa (Figure 5). After two weeks following the first strobilation, a new process occurred with polydisk strobilation, resulting in at least eight free ephyrae (Fig. 5). The life cycle, including this new form of asexual reproduction, is illustrated schematically in Figure 6. We currently have a healthy colony, which has grown under lab conditions without induction, reaching over 30 polyps, all asexually derived from the single settled polyp described above. This asexual polyp generation from the medusa\u0026rsquo;s umbrella was observed at least one more time in a separate cohort of juvenile jellyfish derived from the same initial population of polyps. In this second observation, five of over a hundred juveniles showed at least one polyp at day 20. Another important observation is that, in the new polyp colony derived from medusa tissue, 15% of the medusae generated through the normal strobilation process also produced polyps on the umbrella in the following month, before reaching 5 cm of diameter.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003e\u003cem\u003eReversion of the life cycle in\u003c/em\u003e Stomolopus sp.2\u003c/p\u003e\n\u003cp\u003eIn this report, we describe for the first time the direct generation of a new colony of polyps in the umbrella of \u003cem\u003eStomolophus sp. 2\u003c/em\u003e, a phenomenon that has not been documented previously. A significant finding is the development and growth of colonies of polyps in the umbrella of \u003cem\u003eStomolophus sp. 2\u003c/em\u003e, ocurring without any apparent induction of stress. The life cycle of this species primarily involves multiple asexual reproductions from polyps and sexual reproduction in the medusa stage (L\u0026oacute;pez-Mart\u0026iacute;nez et al., 2022). The discovery and redundant observation of a new process of asexual reproduction from the medusa stage indicate that this may be common, though not predominant, for this species. This could enhance their capacity to invade new areas, as jellyfish are carried by ocean and coastal currents. As a result, the emergence and eventual establishment of polyp populations could occur in different locations through this mechanism, rather than relying solely on external sexual reproduction.\u003c/p\u003e\n\u003cp\u003eAdditionally, this study observed a tissue repair process was observed, which has not been previously reported for this species. Tissue recovery has been noted in other jellyfish, associated with the restoration of \u0026quot;symmetrization\u0026quot; and body part reorganization (Lewis, 2018; Gamero-Mora et al., 2019). The ability to regenerate tissue is rare in nature, and understanding this process could have numerous applications for humans, from its use in biomedical research to comercial cosmetology. In this sense, the cannonball jellyfish could serve as a new model organism for such research.\u003c/p\u003e\n\u003cp\u003eIn many jellyfish species, the life cycle is very complex, with asexual reproduction primarly occurring during the polyp stage. This can lead to the formation of many new polyps within a short period, often linked to rising temperatures (Fern\u0026aacute;ndez-Al\u0026iacute;as et al., 2020). Although the rates of increase are not well-defined, the presence of polyps in the medusa stage has only been observed in Aurelia sp. 1 (He et al., 2015).\u003c/p\u003e\n\u003cp\u003eIn another jellyfish species, \u003cem\u003eTurritopsis dohrnii\u003c/em\u003e, commonly known as the \u0026quot;immortal\u0026quot; jellyfish, the processes of reversion and rejuvenation has been extensively documented. The medusa stage can revert into a polyp form under unfavorable conditions, subsequently settling to form a new colony (Piraino et al., 1996). Exploring converging ontogenetic paths in jellyfish that lead to polyp generation could provide new perspectives on ecological and evolutionary processes in the animal kingdom. Recent studies on the genome and transcriptome of \u003cem\u003eTurritopsis dohrnii\u0026nbsp;\u003c/em\u003ehave identified specific genes associated with development and rejuvenation. It has been shown that the cyst stage is crucial for these processes, and genes related to telomerase maintainence, repair mechanisms, mitotic cell division, and cellular differentiation are transcriptionally active at both the medusa and polyp stages (Matsumoto et al., 2019). Similar processes have been identified in \u003cem\u003eAurelia sp. 1\u003c/em\u003e (Fuch et al., 2014), and this may also be the case in \u003cem\u003eStomolophus sp. 2\u003c/em\u003e. Further studies on their genome and transcriptome are warranted; epigenetic detection through whole-genome bisulfite sequencing (WGBS) could provide insights into the mechanisms controlling shifts in the life cycle, and how environmental stressors may induce this life stage shortcuts. Furthermore, in nature, evolutionary processes are influenced by several factors, including mutations, gene flow, and inbreeding. In this context, species that reproduce only asexually have less adaptive potential than those that reproduce sexually (Orive et al., 2017). This is due to the irreversible accumulation of deleterious mutations, which can lead to genome degradation and reduced fitness, limiting the efficiency of natural selection (Otto, 2021). However, this is primarly relevant for species models with only one mode of reproduction; in species with both modes, the mechanisms are combined. DNA recombination through sexual reproduction generates new gene combinations that are tested in new local environments. Asexual reproduction, contrarly, allows fittest individuals to reproduce as clones, preserving and enabling the spread of those gene combinations across the population (de Mee\u0026ucirc;s et al., 2007). In jellyfish, the complex life cycle and occurrence of two types of reproduction may enhance the evolutionary advantage and ability to adapt to changing environments, along with their dispersal capabilities.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eImplications in dispersal and invasiveness potential\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eOne of the remarkable characteristics of jellyfish is their considerable plasticity in development and rapid adaptive responses to spatial and temporal environmental fluctuations. This allows them to colonize a wide variety of habitats (Thein et al., 2013). The ability of jellyfish polyps to prevail in multiple adverse environmental conditions, together with their rapid asexual reproductive processes, enables them to multiply in large numbers efficiently (Arai, 2009; Lee et al., 2018).\u003c/p\u003e\n\u003cp\u003eOur findings on polyp generation in sexually immature medusae reveal novel mechanisms of polyp dispersion, which provide a significant advantage over other potential invaders. Polyps are resilient life stages of the species that can proliferate over short distances, even under adverse conditions faced by the free-swimming juvenile and adult stages. The ability to generate several free-swimming juvenile means that a relatively small population of polyps can act as a \u0026ldquo;population pool\u0026rdquo; with highly multiplicative potential when favorable conditions arise, all without requiring mature organisms or sexual reproduction.\u003c/p\u003e\n\u003cp\u003eWhen medusae are carried by ocean currents or ballast water, free-floating polyps can be transported over long distances, potentially establishing new populations in regions where environmental conditions are favorable. In projected scenarios of climate change, the abundance of jellyfish populations is likely to increase. Moreover, factors such as eutrophication in new areas and overfishing, which reduce the number of natural predators, further increase the likelihood of blooms forming or persisting (Goldstein et al., 2020).\u003c/p\u003e\n\u003cp\u003eThe appearance of jellyfish in new regions and subsequent blooms highlight the extensive dispersal and invasive potential of these animals. Species such as \u003cem\u003eAurelia spp.\u003c/em\u003e and \u003cem\u003eNemopilema nomurai\u003c/em\u003e in Japan have caused significant ecological problems (Kawahara et al., 2006; Feng et al., 2024). Many studies on blooms focus on population dynamics and damage control, while others examinate the ecophysiology of polyps to understand environmental factors triggering significant proliferations (Zang et al., 2022). In another group of cnidarians, the hydrozoan \u003cem\u003eEudendrium carneum\u003c/em\u003e is recognized as an invasive species with high potential for global spread. It is commonly observed in harbors, yacht clubs, and in fouling communities (G\u0026oacute;nzalez-Duarte et al., 2016). Its life cycle, though primarily sessile, allows for reproduction in both sexual and free-living stages, allowing its expansion into new areas (Puce et al., 2005). Other species like \u003cem\u003ePhyllorhiza punctata\u003c/em\u003e, \u003cem\u003eCassiopea andromeda\u003c/em\u003e, \u003cem\u003eRhopilema nomadica\u003c/em\u003e, and \u003cem\u003eAurelia spp\u003c/em\u003e. have been reported to cover extensive areas in short time frames, leading to their classification as invasive species, usually related to maritime transport rather than biological dispersal mechanisms (Gonz\u0026aacute;lez-Duarte et al., 2016). Currently, predictions about jellyfish blooms and dispersal remain uncertain due to unclear biological processes and environmental factors, coupled with ocean degradation that may be favorable to jellyfish growth (Fern\u0026aacute;ndez-Al\u0026iacute;as et al., 2024). The implications for jellyfish life cycle and bloom formation are significant, offering new insights into jellyfish population dispersion and increase, particularly in the absence of sexual reproduction or maritime shipping as part of the fouling community.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eMolecular mechanisms underlying the reversed cycle\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eAnother perspective from this reversal mechanism is related to tissue regeneration and cellular differentiation. In the hydrozoan \u003cem\u003eTurritopsis nutricula\u003c/em\u003e, the differentiation of polyps from adult tissue involves a cellular transdifferentiation process occuring in both epidermal and gastrointestinal differentiated cells (Pirano et al., 1996). More recently, in the jellyfish \u003cem\u003ePodocoryne carnea\u003c/em\u003e, it was found that muscle tissue cells can change from striated to smooth or even differentiate into functional neurons. This transdifferentiation is linked to differential expression of the Piwi-like gene, which is associated with stem cells in multicellular organisms (Seipel et al., 2004). This suggests that these organisms are cappable of reprogramming cells within the same tissue. In the moon jellyfish \u003cem\u003eAurelia sp. 1\u003c/em\u003e, a reversion cycle from medusa to polyp has been reported, the development of polyps within the umbrella (He et al., 2015). However, the specific genes and cellular mechanisms involved in this reversal process have yet to be elucidated. While differential gene expressions at various life stages does not necessarily indicate a regeneration process or pluripotentiality, several Wnt genes are found to be overexpressed in the planula, strobila, and medusa stages (Brekhman et al., 2015). These genes are implicated in embryogenesis, proliferation, and cell migration (Miller, 2002). Most importantly, this gene family is involved in maintaining the undifferentiated state of stem cells and regulating transcriptional signaling for proliferation and differentiation (Nusee, 2008). Although the presence of these genes does not fully indicate the activity of undifferentiated cells capable of initiating cycle reversal processes, it suggests that these organisms may possess the capacity to perform such reversals at certain life stages.\u003c/p\u003e\n\u003cp\u003eMost of the provided discussion related to the distribution of the polyps in the wild contain a significant speculative component, mainly because this life stage remains cryptic and elusive to ecological research. This is true not only for the cannonball jellyfish but for most of the species that alternate between asexual and sexual reproduction. Therefore, the development of laboratory models, along with advances in field research tools such as environmental DNA and miniaturized underwater drones, would help to understand the implications of the ability to propagate as medusae-derived polyps both seasonally and geographically. Finally, the significance of our findings shows the need to explore into different cellular and physiological mechanisms in these groups of organisms. Given that jellyfish have a long evolutionary history, it would be interesting to understand the adaptations that have helped them survive and invade new environments, their multiple ways of reproducing are certainly one of them.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eCAN-L thanks to the SECIHTI (Ministry of Science, Humanities, Technology, and Innovation; Government of Mexico) for the postdoctorate fellowship (CVU 413245).\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eWe are grateful with Beatriz Ya\u0026ntilde;ez Rivera for her kind assistance on image documentation.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eConceived and designed the experiments: NLC LHR Performed the experiments: NLC. Analyzed the data: NLC. Contributed materials/analysis tools: LHR JLM. Wrote the manuscript: NLC LHR JLM. All the authors revised and approved the final version of the manuscript\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u0026nbsp;\u003c/strong\u003eThis work was mainly funded by internal grants provided by IMCyL-UNAM.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflict of interest:\u0026nbsp;\u003c/strong\u003eThe authors have no conflicts of interest to declare.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthics approval\u003c/strong\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe rearing and lab culture of polyps and jellyfish do not require research ethics approval at the institution where these studies were conducted.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent to participate:\u0026nbsp;\u003c/strong\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication:\u0026nbsp;\u003c/strong\u003eNot applicable.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAouititen M, Ravibhanu A, Ang SC, Magabanda-Mouanda DC, Luan X (2024) New records of two jellyfish species Rhizostoma luteum (Quoy and Gaimard 1827) and Cotylorhiza tuberculata (Macri 1778) in the Moroccan northwest Mediterranean coast. Discov Life 54(5). \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s11084-024-09649\u0026ndash;2\u003c/span\u003e\u003cspan address=\"10.1007/s11084-024-09649\u0026ndash;2\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eArai MN (2009) The potential importance of podocysts to the formation of scyphozoan blooms: a review. Hydrobiologia 616:241\u0026ndash;246. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://doi.org/10.1007/s10750-008-9588\u0026ndash;5\u003c/span\u003e\u003cspan address=\"10.1007/s10750-008-9588\u0026ndash;5\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eBrekhman V, Malik A, Haas B, Sher N, Lotan T (2015) Transcriptome profiling of the dynamic life cycle of the scyphozoan jellyfish Aurelia aurita. BMC Genomics 16:74. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1186/s12864-015-1320-z\u003c/span\u003e\u003cspan address=\"10.1186/s12864-015-1320-z\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eBrotz L, Cisneros-Montemayor AM, Cisneros-Mata MA (2021) The race for jellyfish: Winners and losers in Mexico\u0026rsquo;s Gulf of California. Mar Policy 134:104775. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.marpol.2021.104775\u003c/span\u003e\u003cspan address=\"10.1016/j.marpol.2021.104775\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eChuan CH, Venmathi-Maran BA, Yap TK, Cheong KC, Syed-Hussein MA, Saleh E (2021) New Records of Cubozoan and Scyphozoan Jellyfish from Sabah Waters, Malaysia. Diversity 13(9):420. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3390/d13090420\u003c/span\u003e\u003cspan address=\"10.3390/d13090420\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eFeng S, Xu X, Wan A, Sun S (2024) Differences in. Nemopilema nomurai colony Prolif biofouling invasion Var situ polyp densities Jiaozhou Bay China J Exp Mar Biol Ecol 574:152006. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.jembe.2024.152006\u003c/span\u003e\u003cspan address=\"10.1016/j.jembe.2024.152006\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eFern\u0026aacute;ndez-Al\u0026iacute;as A, Concepci\u0026oacute;n M, Quispe JI, Sabah S, P\u0026eacute;rez-Ruzafa A (2020) Population dynamics and growth in three scyphozoan jellyfishes, and their relationship with environmental conditions in a coastal lagoon. Estuar Coast Shelf Sci 243:106901. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/J.ECSS.2020.106901\u003c/span\u003e\u003cspan address=\"10.1016/J.ECSS.2020.106901\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eFern\u0026aacute;ndez-Al\u0026iacute;as A, Marcos C, P\u0026eacute;rez-Ruzafa A (2021) Larger scyphozoan species dwelling in temperate, shallow waters show higher blooming potential. Mar Pollut Bull 173(B):113100. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.marpolbul.2021.113100\u003c/span\u003e\u003cspan address=\"10.1016/j.marpolbul.2021.113100\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eFern\u0026aacute;ndez-Al\u0026iacute;as A, Marcos C, P\u0026eacute;rez-Ruzafa A (2024) The unpredictability of scyphozoan jellyfish blooms. Front Mar Sci 11:1349956. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3389/fmars.2024.1349956\u003c/span\u003e\u003cspan address=\"10.3389/fmars.2024.1349956\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eFonfr\u0026iacute;a ES, Straehler-Pohl I, Stampar SN, Collins AG, Hoverd WA, Bordehore C (2024) Copula lucentia sp. nov., a new box jellyfish (Cnidaria: Cubozoa: Carybdeida) from Western Mediterranean Sea. Mar Biodivers 54:68. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1007/s12526-024-01460\u0026ndash;6\u003c/span\u003e\u003cspan address=\"10.1007/s12526-024-01460\u0026ndash;6\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eFuchs B, Wang W, Graspeuntner S, Li Y, Insua S, Herbst EM, Dirksen P, B\u0026ouml;hm AM, Hemmrich G, Sommer F, Domazet-Lošo T, Klostermeier UC, Anton-Erxleben F, Rosenstiel P, Bosch TC, Khalturin K (2014) Regulation of polyp-to-jellyfish transition in Aurelia aurita. Curr Biol 24(3):263\u0026ndash;273. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.cub.2013.12.003\u003c/span\u003e\u003cspan address=\"10.1016/j.cub.2013.12.003\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eGamero-Mora E, Halbauer R, Bartsch V, Stampar SN, Morandini AC (2019) Regenerative Capacity of the Upside-down Jellyfish Cassiopea xamachana. Zool Stud 58:e37. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.6620/ZS.2019.58\u0026ndash;37\u003c/span\u003e\u003cspan address=\"10.6620/ZS.2019.58\u0026ndash;37\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eGoldstein J, Steiner UK (2020) Ecological drivers of jellyfish blooms - The complex life history of a 'well-known' medusa (Aurelia aurita). J Anim Ecol 89(3):910\u0026ndash;920. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/1365\u0026ndash;2656.13147\u003c/span\u003e\u003cspan address=\"10.1111/1365\u0026ndash;2656.13147\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eG\u0026oacute;mez-Salinas LC (2018) Din\u0026aacute;mica poblacional de la medusa Stomolophus meleagris (L. Agassiz, 1862) durante el evento El Ni\u0026ntilde;o 2015\u0026ndash;2016 en la Laguna Las Gu\u0026aacute;simas, Sonora, M\u0026eacute;xico. Dissertation Thesis. Universidad Nacional Aut\u0026oacute;noma de M\u0026eacute;xico. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://hdl.handle.net/20.500.14330/TES01000769386\u003c/span\u003e\u003cspan address=\"https://hdl.handle.net/20.500.14330/TES01000769386\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eG\u0026oacute;mez-Salinas LC, L\u0026oacute;pez-Mart\u0026iacute;nez J, Morandini AC (2021) The Young Stages of the Cannonball Jellyfish (Stomolophus sp. 2) from the Central Gulf of California (Mexico). Diversity 13(6):229. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3390/d13060229\u003c/span\u003e\u003cspan address=\"10.3390/d13060229\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eGonz\u0026aacute;lez-Carman V, Botto F, Gait\u0026aacute;n E, Albareda D, Campagna C, Mianzan H (2014) A jellyfish diet for the herbivorous green turtle Chelonia mydas in the temperate SW Atlantic. Mar Biol 161:339\u0026ndash;349. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s00227-013-2339\u0026ndash;9\u003c/span\u003e\u003cspan address=\"10.1007/s00227-013-2339\u0026ndash;9\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eGonz\u0026aacute;lez-Duarte MM, Megina C, L\u0026oacute;pez-Gonz\u0026aacute;lez PJ, Galil B (2016) Cnidarian Alien Species in Expansion. In: Goff S, Zvy, Editors D (n.d.) (eds) The Cnidaria, Past, Present and Future. Springer International Publishing. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://doi.org/10.1007/978-3-319\u0026ndash;31305\u0026ndash;4_10\u003c/span\u003e\u003cspan address=\"10.1007/978-3-319\u0026ndash;31305\u0026ndash;4_10\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eHe J, Zheng L, Zhang W, Lin Y (2015) Life Cycle Reversal in Aurelia sp.1 (Cnidaria, Scyphozoa). PLoS ONE 10(12):e0145314. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1371/journal.pone.0145314\u003c/span\u003e\u003cspan address=\"10.1371/journal.pone.0145314\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eHouliston E, Lecl\u0026egrave;re L, Munro C, Copley RR, Momose T (2022) Past, present and future of Clytia hemisphaerica as a laboratory jellyfish. Curr Top Dev Biol 147:121\u0026ndash;151. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/bs.ctdb.2021.12.014\u003c/span\u003e\u003cspan address=\"10.1016/bs.ctdb.2021.12.014\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eKawahara M, Uye SI, Ohtsu K, Iizumi H (2006) Unusual population explosion of the giant jellyfish Nemopilema nomurai (Scyphozoa: Rhizostomeae) in East Asian waters. Mar Ecol Prog Ser 307:161\u0026ndash;173. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://doi.org/10.3354/meps307161\u003c/span\u003e\u003cspan address=\"10.3354/meps307161\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eLewis-Ames C (2018) Medusa: A Review of an Ancient Cnidarian Body Form. In: Kloc M, Kubiak J (eds) Marine Organisms as Model Systems in Biology and Medicine. Results and Problems in Cell Differentiation, vol 65. Springer, Cham. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/978-3-319\u0026ndash;92486\u0026ndash;1_7\u003c/span\u003e\u003cspan address=\"10.1007/978-3-319\u0026ndash;92486\u0026ndash;1_7\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eL\u0026oacute;pez-Mart\u0026iacute;nez J, \u0026Aacute;lvarez-Tello F (2013) The jellyfish fishery in Mexico. Agri sci 4:57\u0026ndash;61. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.4236/as.2013.46A009\u003c/span\u003e\u003cspan address=\"10.4236/as.2013.46A009\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eL\u0026oacute;pez-Mart\u0026iacute;nez J, \u0026Aacute;lvarez-Tello FJ, Porchas-Cornejo MA, Nev\u0026aacute;rez-L\u0026oacute;pez CA, Muhlia-Almaz\u0026aacute;n A, Ur\u0026iacute;as-Padilla KV (2023) Multiple reproduction forms in the polyps of the cannonball jellyfish Stomolophus sp. 2: Probable life-cycle reversal. J Exp Zool Ecol Integr Physiol 339(3):239\u0026ndash;252. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/jez.2673\u003c/span\u003e\u003cspan address=\"10.1002/jez.2673\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eLuo JY, Condon RH, Stock CA, Duarte CM, Lucas CH, Pitt KA, Cowen RK (2020) Gelatinous Zooplankton-Mediated Carbon Flows in the Global Oceans: A Data‐Driven Modeling Study. Global Biogeochem Cycles 34:e2020GB00670. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1029/2020GB006704\u003c/span\u003e\u003cspan address=\"10.1029/2020GB006704\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eMatsumoto Y, Piraino S, Miglietta MP (2019) Transcriptome Characterization of Reverse Development in Turritopsis dohrnii (Hydrozoa, Cnidaria). G3. Genes Genomes Genet 9(12):4127\u0026ndash;4138. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1534/g3.119.400487\u003c/span\u003e\u003cspan address=\"10.1534/g3.119.400487\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eMiller JR (2022) The Wnts. Genome Biol 3(1):REVIEWS3001. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1186/gb\u0026ndash;2001-3-1-reviews3001\u003c/span\u003e\u003cspan address=\"10.1186/gb\u0026ndash;2001-3-1-reviews3001\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eNev\u0026aacute;rez-L\u0026oacute;pez C, S\u0026aacute;nchez-Paz A, L\u0026oacute;pez-Mart\u0026iacute;nez J, Llera-Herrera R, Muhlia-Almaz\u0026aacute;n A (2020) Metabolic response of the cannonball jellyfish Stomolophus meleagris upon short-term exposure to thermal stress. J Sea Res 166:101959. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.seares.2020.101959\u003c/span\u003e\u003cspan address=\"10.1016/j.seares.2020.101959\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eNusse R (2008) Wnt signaling and stem cell control. Cell Res 18(5):523\u0026ndash;527. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1038/cr.2008.47\u003c/span\u003e\u003cspan address=\"10.1038/cr.2008.47\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003ePiraino S, Boero F, Aeschbach B, Schmid V (1996) Reversing the Life Cycle: Medusae Transforming into Polyps and Cell Transdifferentiation in Turritopsis nutricula (Cnidaria, Hydrozoa). Biol Bull 190(3):302\u0026ndash;312. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.2307/1543022\u003c/span\u003e\u003cspan address=\"10.2307/1543022\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003ePitt K, Lucas C, Condon RH, Duarte CM, Stewart-Koster B (2018) Claims That Anthropogenic Stressors Facilitate Jellyfish Blooms Have Been Amplified Beyond the Available Evidence: A Systematic Review. Front Mar Sci 5:451. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3389/fmars.2018.00451\u003c/span\u003e\u003cspan address=\"10.3389/fmars.2018.00451\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003ePreciado M (2016) Pesquer\u0026iacute;a artesanal de la medusa Stomolophusmeleagris en el estuario interno del Golfo de Guayaquil 2014. Master thesis. Universidad de Guayaquil. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://repositorio.ug.edu.ec/handle/redug/22212\u003c/span\u003e\u003cspan address=\"http://repositorio.ug.edu.ec/handle/redug/22212\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003ePurcell JE, Baxter EJ, Fuentes VL (2013) Jellyfish as products and problems of aquaculture. In: Advances in Aquaculture Hatchery Technology. Eds. Allan G, Burnell G. Woodhead Publishing Series in Food Science, Technology and Nutrition, Cambridge, United Kingdom, 404\u0026ndash;430 pp. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1533/9780857097460.2.404\u003c/span\u003e\u003cspan address=\"10.1533/9780857097460.2.404\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eSchnedler-Meyer NA, Ki\u0026oslash;rboe T, Mariani P (2018) Boom and Bust: Life History, Environmental Noise, and the (un)Predictability of Jellyfish Blooms. Front Mar Sci 5:257. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3389/fmars.2018.00257\u003c/span\u003e\u003cspan address=\"10.3389/fmars.2018.00257\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eSeipel K, Yanze N, Schmid V (2004) The germ line and somatic stem cell gene Cniwi in the jellyfish Podocoryne carnea. Int J Dev Biol 48(1):1\u0026ndash;7. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1387/ijdb.15005568\u003c/span\u003e\u003cspan address=\"10.1387/ijdb.15005568\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eThein H, Ikeda H, Uye S-I (2013) Ecophysiological characteristics of podocysts in Chrysaora pacifica (Goette) and Cyanea nozakii Kishinouye (Cnidaria: Scyphozoa: Semaeostomeae): effects of environmental factors on their production, dormancy and excystment. J Exp Mar Biol Ecol 446:151\u0026ndash;158. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://doi.org/10.1016/j.jembe.2013.05.013\u003c/span\u003e\u003cspan address=\"10.1016/j.jembe.2013.05.013\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eZang W, Zhang F, Chi X, Sun S (2022) Relationship Between Asexual Reproduction of Aurelia coerulea Polyps and Jellyfish Blooms Under the Influence of Temperature Dynamics in Winter and Spring. Front Mar Sci 9:888656. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3389/fmars.2022.888656\u003c/span\u003e\u003cspan address=\"10.3389/fmars.2022.888656\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003evan Walraven L, Driessen F, van Bleijswijk J, Bol A, Luttikhuizen PC, Coolen JWP, Bos OG, Gittenberger A, Schrieken N, Langenberg VT, van der Veer HW (2016) Where are the polyps? Molecular identification, distribution and population differentiation of Aurelia aurita jellyfish polyps in the southern North Sea area. Mar Biol 163:172. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s00227-016-2945\u0026ndash;4\u003c/span\u003e\u003cspan address=\"10.1007/s00227-016-2945\u0026ndash;4\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"marine-biology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"mabi","sideBox":"Learn more about [Marine Biology](https://www.springer.com/journal/227)","snPcode":"227","submissionUrl":"https://submission.nature.com/new-submission/227/3","title":"Marine Biology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"cannonball jellyfish, Stomolophus, polyps, asexual reproduction","lastPublishedDoi":"10.21203/rs.3.rs-7579160/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7579160/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e\u003cspan type=\"ItalicSmallCaps\" class=\"ItalicSmallCaps\" name=\"Emphasis\"\u003eStomolophus\u003c/span\u003e \u003cspan type=\"SmallCaps\" class=\"SmallCaps\" name=\"Emphasis\"\u003espp. is one of the commercially important jellyfish in the world; however, its life cycle has recently been described with new forms of asexual reproduction in the polyp stage, with the reversion process to ephyrae considered an advantage for increasing population size, especially under stressful conditions. In this study, we observed the generation of new polyp colonies directly from the bell margins of the medusae. Here we present the first evidence for back-transformation of tissues medusae into polyps in\u003c/span\u003e \u003cspan type=\"ItalicSmallCaps\" class=\"ItalicSmallCaps\" name=\"Emphasis\"\u003eStomolophus\u003c/span\u003e \u003cspan type=\"SmallCaps\" class=\"SmallCaps\" name=\"Emphasis\"\u003esp.2. The new way of asexual reproduction from medusae in the complex life cycle of this species reveals a long-range asexual mechanism of dispersion and potential of invasion, as well as of capacities to rejuvenation, the possibilities of immortality with reprogramming of differentiated cells and the increase of populations and blooms not only in this species but also in others that may have the same process of reversion.\u003c/span\u003e\u003c/p\u003e","manuscriptTitle":"Evidence of asexual polyp development from the adult body in Stomolopus sp2","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-09-19 14:38:25","doi":"10.21203/rs.3.rs-7579160/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revise and Resubmit","date":"2025-10-25T03:08:05+00:00","index":"","fulltext":""},{"type":"reviewerAgreed","content":"","date":"2025-09-12T13:23:55+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-09-12T12:57:46+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-09-11T13:30:13+00:00","index":"","fulltext":""},{"type":"submitted","content":"Marine Biology","date":"2025-09-10T02:00:22+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"marine-biology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"mabi","sideBox":"Learn more about [Marine Biology](https://www.springer.com/journal/227)","snPcode":"227","submissionUrl":"https://submission.nature.com/new-submission/227/3","title":"Marine Biology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"36787016-8a97-4202-9a35-a828e8266107","owner":[],"postedDate":"September 19th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-04-25T16:42:15+00:00","versionOfRecord":[],"versionCreatedAt":"2025-09-19 14:38:25","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-7579160","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7579160","identity":"rs-7579160","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}

Text is read by the "Ask this paper" AI Q&A widget below. Extraction quality varies by source — PMC NXML preserves structure cleanly, OA-HTML may include some navigation residue, and OA-PDF can have broken hyphenation. The publisher copy (via DOI) is the canonical version.

My notes (saved in your browser only)

Ask this paper AI returns verbatim quotes from the full text · source: preprint-html

Answers must be backed by verbatim quotes from this paper's full text. Hallucinated quotes are dropped automatically; if no verbatim passage answers the question, we say so. How this works

Citation neighborhood (no data yet)

We don't have any in-corpus citations linked to this paper yet. This is a recent paper (2025) — citers typically take a year or two to land, and the OpenAlex reference graph may still be filling in.

Source provenance

europepmc
last seen: 2026-05-20T01:45:00.602351+00:00