{ "componentChunkName": "component---src-templates-article-page-js", "path": "/journals/biology/micropub-biology-002242", "result": {"data":{"article":{"manuscript":{"id":"ce777782-86a8-458f-b93b-9fc7b39f9c81","submissionTypes":["new finding","materials and reagents","methodology"],"citations":[],"doi":"10.17912/micropub.biology.002242","dbReferenceId":null,"pmcId":null,"pmId":null,"proteopedia":null,"reviewPanel":null,"species":["other"],"integrations":[],"corrections":null,"history":{"received":"2026-06-10T19:18:47.388Z","revisionReceived":"2026-09-21T18:24:46.341Z","accepted":"2026-09-23T22:54:25.486Z","published":"2026-09-25T16:29:48.912Z","indexed":"2026-10-09T16:29:48.912Z"},"versions":[{"id":"0a8bfc06-47a4-468e-8b55-073800ffdea0","decision":"revise","abstract":"

The sea urchin Diadema antillarum is a restoration target due to its function as a keystone reef herbivore. However, restocking efforts are hindered by a lack of long-term tagging methods. We tested the effectiveness of calcein marking for urchin releases. Cultured individuals were tagged using a 150 mg/L calcein bath and monitored for 11 weeks. Dye markings remained fully visible in all individuals for two weeks but began to fade within three weeks. Only individuals with white spines maintained markings beyond five weeks. Unlike with other urchin species, calcein only seems suitable for short-term external tagging of D. antillarum.

","acknowledgements":"

Many thanks go to Joshua Patterson of the University of Florida and The Florida Aquarium for providing the urchins, and to Justin Voss, Ellery Lennon, and Holly White for helping maintain the urchins.

","authors":[{"affiliations":["Florida Fish and Wildlife Conservation Commission, Tallahassee, FL, United States"],"departments":["Fish and Wildlife Research Institute"],"credit":["conceptualization","dataCuration","formalAnalysis","investigation","methodology","project","software","validation","visualization","writing_originalDraft","writing_reviewEditing"],"email":"Benjamin.Belgrad@MyFWC.com","firstName":"Benjamin A.","lastName":"Belgrad","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":"0000-0003-2032-2210"},{"affiliations":["Florida Fish and Wildlife Conservation Commission"],"departments":["Fish and Wildlife Research Institute"],"credit":["conceptualization","dataCuration","fundingAcquisition","investigation","methodology","resources","writing_reviewEditing"],"email":"Lindsay.Spiers@MyFWC.com","firstName":"Lindsay J.","lastName":"Spiers","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":null},{"affiliations":["Florida Fish and Wildlife Conservation Commission"],"departments":["Fish and Wildlife Research Institute"],"credit":["conceptualization","fundingAcquisition","methodology","project","resources","validation","writing_reviewEditing"],"email":"William.Sharp@MyFWC.com","firstName":"William C.","lastName":"Sharp","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":null}],"awards":[],"conflictsOfInterest":"

The authors declare that there are no conflicts of interest present.

","dataTable":{"url":null},"extendedData":[],"funding":"

This work was supported by funds provided to the Florida Fish and Wildlife Conservation Commission’s Marine Resources Trust Fund.

","image":{"url":"https://portal.micropublication.org/uploads/7650f22ede66ffdeaf4062baee1b5bdf.png"},"imageCaption":"

Photographs of the same two urchins tagged with calcein under a) normal light, b) unfiltered UV light, and c) UV light viewed through a blue light filter. Fluorescence from tagging was best viewed under blue filtered UV light and was observed on either c) the spines or d) tube feet along the ambulacral plates. e) Number of urchins with only black spines or with some white spines still exhibiting fluorescent tagging each week (n = 13 urchins tagged total). f) Urchins with white spines that were either dyed with calcein (tagged) or undyed (control) 10 weeks after tagging. g) A black and white spine from a tagged and control urchin viewed under filtered UV light. h–l) Photo series spanning ~30 seconds of an urchin expelling fluorescent substance three days after tagging. Yellow arrow indicates initial material expulsion.

","imageTitle":"

Photographs of calcein tagged and control urchins and duration of tag visibility

","methods":"

Juvenile Diadema antillarum, spawned from wild-caught broodstock, were acquired from The Florida Aquarium's Center for Conservation, Apollo Beach, FL, US, in May 2024. Individuals were transported to the Florida Fish and Wildlife Research Institute’s South Florida Regional Lab in Marathon, FL, US, where they were nursed for approximately 21 months before commencing the experiment. During the nursery period, urchins were held in circular flow-through tanks (diameter = 155 cm, water height = 70 cm) under similar conditions to Sharp et al. (2023). Sand-filtered seawater was supplied from Florida Bay and matched natural abiotic conditions. Shelter was provided to urchins in the form of longitudinally halved terracotta pots (22 cm L × 22 cm W × 12 cm H) that mimicked natural coral reef crevice habitat, and urchins were fed satiating amounts of drift algae.

            Thirty-four urchins were haphazardly selected for tagging (n = 13; test diameter = 27–43 mm) and control treatments (n = 21; test diameter = 32–61 mm) to verify that untagged urchins do not fluoresce and determine if tagging influences survival. Urchins selected for tagging were distributed across three continuously aerated tanks containing a solution of sand-filtered seawater with 150 mg/L concentration of dissolved calcein. Two tanks held five urchins with 11 L calcein solution (Chemcruz source) while the third tank held three urchins with 7 L calcein solution (MP Biomedicals source). These volumes and urchin numbers were chosen based on perceived physiological tolerances, and the compounds from both sources produced the same results. Urchins were immersed in this dye bath for seven days (typical dye regimens use 100–150 mg/L for 24–48 h immersion). This extended period of immersion was performed because our preliminary experiments noted that fluorescence tagging disappeared quickly, and we wanted to identify the maximum amount of time tagging can be viewed externally on pure black-spined urchins, which comprise the majority of D. antillarum populations.

            After this tagging period, control and tagged urchins were placed into two separate 356-L rectangular aquaria (120 cm L × 75 cm W × 36 cm H) and maintained under similar conditions as the nursery period. Individuals were held for 11 weeks and checked at least weekly to determine if fluorescent markings were still observable. Markings were observed by shining a Sola NIGHTSEA UV dive light on individuals and viewing them through a NIGHTSEA blue light filter visor. The proportion of tagged individuals fluorescing over time was analyzed using a generalized linear model with a binomial distribution. Observations were treated as population-level repeated measurements, rather than independent individual trajectories because of the inability to track individuals across time. All statistical analyses were performed in R version 4.5.1.

","reagents":"

Reagent

Alternate names

Lot Number

Source

Calcein disodium salt

Fluorexon Sodium Salt

C2425

ChemCruz

Calcein

Fluorexon; Fluorescein-bis(methyliminodiacetic acid);

Bis[N,N−bis(carboxymethyl)aminomethyl]fluorescein;

3,3'-bis[N,N'-di(carboxymethyl)-aminomethyl]-fluorescein; Calcein W

S8883-2

MP Biomedicals

","patternDescription":"

Sea urchins comprise a global multimillion dollar fishery and are ecological engineers (Ogden et al., 1978; Stefánsson et al., 2017; FAO 2025). As their populations have declined, urchins are increasingly cultivated around the world for consumption and restoration purposes (FAO 2024; Rakaj et al., 2024; Smith et al., 2026). The long-spined sea urchin, Diadema antillarum, is a keystone herbivore native to the tropical Atlantic and Caribbean coral reefs (Randall et al., 1963; Carpenter 1981; Carpenter 1988; Lessios 2016). In the 1980s, an epidemic spread throughout the Caribbean and killed 93–98% of the urchin populations (Carpenter 1988; Lessios 1988). This was followed by an enormous increase in benthic macroalgae and a decrease in coral cover as the coral was outcompeted for space and light (Hughes et al., 1987). D. antillarum populations have only modestly recovered since this epidemic (~12% population pre-epidemic; Lessios 2016), and the species has recently become a target of restoration to aid coral recovery (Pilnick et al., 2021; Hylkema et al., 2022). In areas where D. antillarum populations have increased in abundance, there have been significant decreases in macroalgal cover and, in some areas, increases in coral recruitment (Edmunds and Carpenter 2001). The restoration effort has centered around supplementing wild populations with hatchery reared juveniles (Pilnick et al., 2021; Hudspeth et al., 2026), as natural recruitment is low and early life-stage predation is high (Lessios 2016; Hylkema et al., 2022). However, restocking efforts and research on the effectiveness of this approach are hampered by a lack of viable long-term tagging methods. Reliable long-term tagging is essential to obtain information that is largely lacking and fundamental to developing and optimizing restoration strategies for this species, such as post-release survival, individual growth rates, and movement patterns of outplanted urchins.

            Various methods of tagging urchins have been employed since the 1960s with mixed success (Ebert 1965; Ebert 2013; Santos et al., 2022; Tourón et al., 2023). Many internal physical tags such as decimal coded wire tags (CWTs) and passive integrated transponder (PIT) tags can cause increased mortality or decreased growth rates in urchins, especially in smaller individuals, while external physical markings such as painting spines or placing beads are quickly lost (Cipriano et al., 2014; Santos et al., 2022; Tourón et al., 2023). Chemically marking individuals with the fluorochrome dye calcein is a potential noninvasive and non-toxic means of tagging D. antillarum for long durations. The molecule typically binds irreversibly to calcium ions and becomes incorporated into the tissue and skeletal structure of animals as they grow, which causes the individual’s tissues to fluoresce under UV light (Johnson et al., 2013; Li et al., 2020). Although this method has been used to tag several species of urchins for months (Johnson et al., 2013; Rodríguez et al., 2016; Santos et al., 2022; Tourón et al., 2023), the technique has yet to be tested with D. antillarum. Here, we conducted a laboratory study to evaluate the efficacy of using calcein to tag D. antillarum for extended timeframes.

            Survival was 100% over the course of this study. Recently tagged D. antillarum fluoresced strongly under UV light and the markings were most visible when observed through a blue light filter (Figure 1a–c). Individuals differed in where tag fluorescence appeared on their body as fluorescence was either localized to their spines (Figure 1c) or tube feet along the ambulacral plates (Figure 1d). Tag fluorescence started to become difficult to detect visually after two weeks (Figure 1e; GLM; coefficient estimate = -0.66, z = -6.38, p < 0.0001). Most tagged urchins (n = 10 of 13) were indistinguishable from untagged control urchins (n = 21) after five weeks; however, all three urchins with white spines (n = 3 of 13) retained fluorescence for at least 11 weeks (Figure 1f). This continued fluorescence was limited to the white spines (Figure 1g), possibly either because the darker pigmentation within black spines prevented UV light from reaching the incorporated calcein, or it may have attenuated emission visibility, masking calcein fluorescence. However, further molecular studies are needed to identify the mechanisms underlying differences in spine fluorescence. Interestingly, we observed four individuals periodically releasing a fluorescent substance aborally three days after removal from the dye bath. This may indicate post-exposure clearance or redistribution of dye-associated material, though chemical confirmation was not performed (Figure 1h–l).

Importantly, unlike previous studies where calcein dye was visible externally on urchins for months (Rogers-Bennett 2003; Johnson et al., 2013; Santos et al., 2022; Correia et al., 2023; Tourón et al., 2023), dye markings only remained visible on D. antillarum for approximately two weeks — despite our use of a more intense than typical dye regimen. Such contrasting results are likely due to differences in pigmentation of those species as Paracentrotus lividus (Santos et al., 2022), Strongylocentrotus droebachiensis (Johnson et al. 2013), Strongylocentrotus franciscanus (Rogers-Bennett 2003), and Pseudechinus huttoni (Kirby et al., 2006) are substantially lighter in coloration than D. antillarum. In the dark-colored Diadema africanum (Rodríguez et al., 2016) and Arbacia lixula (Barrera 2018), extended tag visibility was only achieved by dissection and microscopic examination of the Aristotle’s lantern. Consequently, we conclude that calcein dye is unsuitable for tagging D. antillarum for long durations when individual survival is a priority or disturbance must be minimized, but calcein dye may be sufficient for short-term studies or when dissection is an option. These results highlight how even biologically conserved chemical reactions can have different outcomes across similar species and the necessity for species-specific studies.

","references":[{"reference":"

Barrera AM. 2018. Estimación del crecimiento del erizo de mar Arbacia lixula. Máster en Biología marina: Biodiversidad y conservación. San Cristóbal de La Laguna, Spain: Universidad de la Laguna. http://riull.ull.es/xmlui/handle/915/23646

","pubmedId":"","doi":""},{"reference":"

Carpenter RC. 1981. Grazing by Diadema antillarum (Philippi) and its effects on the benthic algal community. Journal of Marine Research. 39: 749-765. https://elischolar.library.yale.edu/journal_of_marine_research/1568

","pubmedId":"","doi":""},{"reference":"

Carpenter RC. 1988. Mass mortality of a Caribbean sea urchin: Immediate effects on community metabolism and other herbivores. Proc Natl Acad Sci U S A 85(2): 511-4.

","pubmedId":"16593907","doi":""},{"reference":"

Ciriano A, Burnell G, Cullotv S, Long S. 2014. Evaluation of 3 Tagging Methods in Marking Sea Urchin,Paracentrotuslividus, Populations under Both Laboratory and Field Conditions. Journal of Aquaculture Research & Development 05: 10.4172/2155-9546.1000276.

","pubmedId":"","doi":"10.4172/2155-9546.1000276"},{"reference":"

Correia MJo, Mega Lopes Pc, Santos PM, Jacinto D, Mateus D, Maresca F, et al., Costa. 2023. Pilot studies for stock enhancement of purple sea urchins (Paracentrotus lividus, Lamarck, 1816): usefulness of refuges and calcein marking for the monitoring of juveniles released into the natural environment. Aquatic Living Resources 36: 12.

","pubmedId":"","doi":"10.1051/alr/2023009"},{"reference":"

Ebert TA. 1965. A Technique for the Individual Marking of Sea Urchins. Ecology 46: 193-194.

","pubmedId":"","doi":"10.2307/1935273"},{"reference":"

Ebert TA. 2013. Growth and survival of postsettlement sea urchins. In: Lawrence J.M. (ed), Sea Urchins: Biology and Ecology. Academic Press, San Diego, California, pp. 83-117.

","pubmedId":"","doi":"10.1016/B978-0-12-396491-5.00007-1"},{"reference":"

Edmunds PJ, Carpenter RC. 2001. Recovery of Diadema antillarum reduces macroalgal cover and increases abundance of juvenile corals on a Caribbean reef. Proc Natl Acad Sci U S A 98(9): 5067-71.

","pubmedId":"11274358","doi":""},{"reference":"

FAO. 2025. Fishery and aquaculture statistics – yearbook 2023. Rome.

","pubmedId":"","doi":"10.4060/cd6788en"},{"reference":"

FAO. 2024. The State of World Fisheries and Aquaculture 2024 – Blue Transformation in action. Rome.

","pubmedId":"","doi":"10.4060/cd0683en"},{"reference":"

Hudspeth CG, Pilnick AR, Petrosino A, Smith JN, Patterson JT. 2026. Developing standard production diets for early juvenile Diadema antillarum sea urchins. Aquaculture Reports 47: 103430.

","pubmedId":"","doi":"10.1016/j.aqrep.2026.103430"},{"reference":"

Hughes TP, Reed DC, Boyle MJ. 1987. Herbivory on coral reefs: community structure following mass mortalities of sea urchins. Journal of Experimental Marine Biology and Ecology 113: 39-59.

","pubmedId":"","doi":"10.1016/0022-0981(87)90081-5"},{"reference":"

Hylkema A, Debrot AO, van de Pas EE, Osinga R, Murk AJ. 2022. Assisted Natural Recovery: A Novel Approach to Enhance Diadema antillarum Recruitment. Frontiers in Marine Science 9: 10.3389/fmars.2022.929355.

","pubmedId":"","doi":"10.3389/fmars.2022.929355"},{"reference":"

Johnson AS, Salyers JM, Alcorn NJ, Ellers O, Allen JD. 2013. Externally visible fluorochrome marks and allometries of growing sea urchins. Invertebrate Biology 132: 251-269.

","pubmedId":"","doi":"10.1111/ivb.12024"},{"reference":"

Kirby S, Lamare MD, Barker MF. 2006. Growth and morphometrics in the New Zealand sea urchin\n Pseudechinus huttoni\n (Echinoidea: Temnopleuridae). New Zealand Journal of Marine and Freshwater Research 40: 413-428.

","pubmedId":"","doi":"10.1080/00288330.2006.9517432"},{"reference":"

Lessios HA. 1988. MASS MORTALITY OF DIADEMA ANTILLARUM IN THE CARIBBEAN: What Have We Learned?. Annual Review of Ecology and Systematics 19: 371-393.

","pubmedId":"","doi":"10.1146/annurev.es.19.110188.002103"},{"reference":"

Lessios HA. 2016. The Great Diadema antillarum Die-Off: 30 Years Later. Ann Rev Mar Sci 8: 267-83.

","pubmedId":"26048480","doi":""},{"reference":"

Li H, Lei Y, Li T, Jian Z. 2020. Effects of calcein incorporation on benthic foraminiferal community under various concentrations and incubation durations. Marine Micropaleontology 157: 101874.

","pubmedId":"","doi":"10.1016/j.marmicro.2020.101874"},{"reference":"

Ogden JC, Lobel PS. 1978. The role of herbivorous fishes and urchins in coral reef communities. Environmental Biology of Fishes 3: 49-63.

","pubmedId":"","doi":"10.1007/BF00006308"},{"reference":"

Pilnick AR, O'Neil KL, Moe M, Patterson JT. 2021. A novel system for intensive Diadema antillarum propagation as a step towards population enhancement. Sci Rep 11(1): 11244.

","pubmedId":"34045538","doi":""},{"reference":"

Rakaj A, Grosso L, Fianchini A, Cataudella S. 2024. A sustainable no-kill sea urchin aquaculture method to obtain caviar. Nature Sustainability 7: 1038-1047.

","pubmedId":"","doi":"10.1038/s41893-024-01372-0"},{"reference":"

Randall JE, Schroeder RE, Stark WA. 1963. Notes on the biology of the echinoid Diadema antillarum. Caribbean Journal of Science. 4: 2-3. https://www.researchgate.net/publication/246980787_Notes_on_the_biology_of_the_echinoid_Diadema_antillarum

","pubmedId":"","doi":""},{"reference":"

Rodríguez A, Hernández JC, Clemente S. 2016. Efficiency of calcein tagging on juveniles of the sea urchins Diadema africanum and Paracentrotus lividus. Marine Ecology 37: 463-469.

","pubmedId":"","doi":"10.1111/maec.12287"},{"reference":"

Rogers-Bennett L. 2003. Modeling red sea urchin growth using six growth functions. Fishery Bulletin. 101: 614-626. https://escholarship.org/uc/item/8gn2v2xn

","pubmedId":"","doi":""},{"reference":"

Santos PM, Ruivo Quintella B, Jacinto D, Gomes A, Saldanha C, Lourenço Sl, et al., Lino Costa. 2022. Physical and chemical tagging methods for the sea urchin Paracentrotus lividus (Echinodermata: Echinoidea). Scientia Marina 86: e038.

","pubmedId":"","doi":"10.3989/scimar.05259.038"},{"reference":"

Sharp WC, Delgado GA, Pilnick AR, Patterson JT. 2023. Diurnal Sheltering Behavior of Hatchery-propagated Long-spined Urchins (Diadema Antillarum): a Re-examination Following Husbandry Refinements. Bulletin of Marine Science 99: 97-108.

","pubmedId":"","doi":"10.5343/bms.2022.0054"},{"reference":"

Stefánsson G, Kristinsson H, Ziemer N, Hannon C, James P. 2017. Markets for sea urchins: a review of global supply and markets. Skýrsla Matís. 45: 10-17.

","pubmedId":"","doi":"10.13140/RG.2.2.12657.99683"},{"reference":"

Smith XJ, Dempster T, Salini M, Francis D, Warren-Myers F. 2026. Commercial scale test of purple sea urchin roe enhancement aquaculture. Aquaculture 618: 743804.

","pubmedId":"","doi":"10.1016/j.aquaculture.2026.743804"},{"reference":"

Tourón N, Campos S, Costas Dn, Paredes Ea. 2023. Marking Methodologies for Sea Urchins: A Review. Journal of Shellfish Research 42: 10.2983/035.042.0116.

","pubmedId":"","doi":"10.2983/035.042.0116"}],"title":"

The efficacy of calcein as a tagging agent for the long-spined sea urchin, Diadema antillarum

","reviews":[{"reviewer":{"displayName":"Olaf Ellers"},"openAcknowledgement":false,"status":{"submitted":true}}],"curatorReviews":[]},{"id":"ab696100-956e-4c99-873a-2e8ffd89350c","decision":"accept","abstract":"

The sea urchin Diadema antillarum is a restoration target due to its function as a keystone reef herbivore. However, restocking efforts are hindered by a lack of long-term tagging methods. We tested the effectiveness of calcein marking for urchin releases. Cultured individuals were tagged using a 150 mg/L calcein bath and monitored for 11 weeks. Dye markings remained fully visible in all individuals for two weeks but began to fade within three weeks. Only individuals with white spines maintained markings beyond five weeks. Unlike with other urchin species, calcein only seems suitable for short-term external tagging of D. antillarum.

","acknowledgements":"

Many thanks go to Joshua Patterson of the University of Florida and The Florida Aquarium for providing the urchins, and to Justin Voss, Ellery Lennon, and Holly White for helping maintain the urchins.

","authors":[{"affiliations":["Florida Fish and Wildlife Conservation Commission, Tallahassee, FL, United States"],"departments":["Fish and Wildlife Research Institute"],"credit":["conceptualization","dataCuration","formalAnalysis","investigation","methodology","project","software","validation","visualization","writing_originalDraft","writing_reviewEditing"],"email":"Benjamin.Belgrad@MyFWC.com","firstName":"Benjamin A.","lastName":"Belgrad","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":"0000-0003-2032-2210"},{"affiliations":["Florida Fish and Wildlife Conservation Commission"],"departments":["Fish and Wildlife Research Institute"],"credit":["conceptualization","dataCuration","fundingAcquisition","investigation","methodology","resources","writing_reviewEditing"],"email":"Lindsay.Spiers@MyFWC.com","firstName":"Lindsay J.","lastName":"Spiers","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":null},{"affiliations":["Florida Fish and Wildlife Conservation Commission"],"departments":["Fish and Wildlife Research Institute"],"credit":["conceptualization","fundingAcquisition","methodology","project","resources","validation","writing_reviewEditing"],"email":"William.Sharp@MyFWC.com","firstName":"William C.","lastName":"Sharp","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":null}],"awards":[],"conflictsOfInterest":"

The authors declare that there are no conflicts of interest present.

","dataTable":{"url":null},"extendedData":[],"funding":"

This work was supported by funds provided to the Florida Fish and Wildlife Conservation Commission’s Marine Resources Trust Fund.

","image":{"url":"https://portal.micropublication.org/uploads/f607c75e6fda74cf3a296d4f7f0e8a4d.png"},"imageCaption":"

Photographs of the same two urchins tagged with calcein under a) normal light, b) unfiltered UV light, and c) UV light viewed through a blue light filter. Fluorescence from tagging was best viewed under blue filtered UV light and was observed on either c) the spines or d) tube feet along the ambulacral plates. e) Number of urchins with only black spines or with some white spines still exhibiting fluorescent tagging each week (n = 13 urchins tagged total). Urchins with black spines had fluorescence intensity notably decline and become harder to detect visually three weeks after tagging. f) Urchins with white spines that were either dyed with calcein (tagged) or undyed (control) 10 weeks after tagging. g) A black and white spine from a tagged and control urchin viewed under filtered UV light. h–l) Photo series spanning ~30 seconds of an urchin expelling fluorescent substance three days after tagging. Yellow arrow indicates initial material expulsion.

","imageTitle":"

Photographs of calcein tagged and control urchins and duration of tag visibility

","methods":"

Juvenile Diadema antillarum, spawned from wild-caught broodstock, were acquired from The Florida Aquarium's Center for Conservation, Apollo Beach, FL, US, in May 2024. Individuals were transported to the Florida Fish and Wildlife Research Institute’s South Florida Regional Lab in Marathon, FL, US, where they were nursed for approximately 21 months before commencing the experiment. During the nursery period, urchins were held in circular flow-through tanks (diameter = 155 cm, water height = 70 cm) under similar conditions to Sharp et al. (2023). Sand-filtered seawater was supplied from Florida Bay and matched natural abiotic conditions. Shelter was provided to urchins in the form of longitudinally halved terracotta pots (22 cm L × 22 cm W × 12 cm H) that mimicked natural coral reef crevice habitat, and urchins were fed satiating amounts of drift algae.

            Thirty-four urchins were haphazardly selected for tagging (n = 13; test diameter = 27–43 mm) and control treatments (n = 21; test diameter = 32–61 mm) to verify that untagged urchins do not fluoresce and determine if tagging influences survival. Urchins selected for tagging were distributed across three continuously aerated tanks containing a solution of sand-filtered seawater with 150 mg/L concentration of dissolved calcein. Two tanks held five urchins with 11 L calcein solution (Chemcruz source) while the third tank held three urchins with 7 L calcein solution (MP Biomedicals source). These volumes and urchin numbers were chosen based on perceived physiological tolerances, and the compounds from both sources produced the same results. Urchins were immersed in this dye bath for seven days (typical dye regimens use 100–150 mg/L for 24–48 h immersion). This extended period of immersion was performed because our preliminary experiments noted that fluorescence tagging disappeared quickly, and we wanted to identify the maximum amount of time tagging can be viewed externally on pure black-spined urchins, which comprise the majority of D. antillarum populations.

            After this tagging period, control and tagged urchins were placed into two separate 356-L rectangular aquaria (120 cm L × 75 cm W × 36 cm H) and maintained under similar conditions as the nursery period. Individuals were held for 11 weeks and checked at least weekly to determine if fluorescent markings were still observable. Markings were observed by shining a Sola NIGHTSEA UV dive light (emission spectra 360–380 nm) on individuals and viewing them through a NIGHTSEA blue blocking light filter visor (blocking wavelengths shorter than 415 nm).

","reagents":"

Reagent

Alternate names

Lot Number

Source

Calcein disodium salt

Fluorexon Sodium Salt

C2425

ChemCruz

Calcein

Fluorexon; Fluorescein-bis(methyliminodiacetic acid);

Bis[N,N−bis(carboxymethyl)aminomethyl]fluorescein;

3,3'-bis[N,N'-di(carboxymethyl)-aminomethyl]-fluorescein; Calcein W

S8883-2

MP Biomedicals

","patternDescription":"

Sea urchins comprise a global multimillion dollar fishery and are ecological engineers (Ogden et al., 1978; Stefánsson et al., 2017; FAO 2025). As their populations have declined, urchins are increasingly cultivated around the world for consumption and restoration purposes (FAO 2024; Rakaj et al., 2024; Smith et al., 2026). The long-spined sea urchin, Diadema antillarum, is a keystone herbivore native to the tropical Atlantic and Caribbean coral reefs (Randall et al., 1963; Carpenter 1981; Carpenter 1988; Lessios 2016). In the 1980s, an epidemic spread throughout the Caribbean and killed 93–98% of the urchin populations (Carpenter 1988; Lessios 1988). This was followed by an enormous increase in benthic macroalgae and a decrease in coral cover as the coral was outcompeted for space and light (Hughes et al., 1987). D. antillarum populations have only modestly recovered since this epidemic (~12% population pre-epidemic; Lessios 2016), and the species has recently become a target of restoration to aid coral recovery (Pilnick et al., 2021; Hylkema et al., 2022). In areas where D. antillarum populations have increased in abundance, there have been significant decreases in macroalgal cover and, in some areas, increases in coral recruitment (Edmunds and Carpenter 2001). The restoration effort has centered around supplementing wild populations with hatchery reared juveniles (Pilnick et al., 2021; Hudspeth et al., 2026), as natural recruitment is low and early life-stage predation is high (Lessios 2016; Hylkema et al., 2022). However, restocking efforts and research on the effectiveness of this approach are hampered by a lack of viable long-term tagging methods. Reliable long-term tagging is essential to obtain information that is largely lacking and fundamental to developing and optimizing restoration strategies for this species, such as post-release survival, individual growth rates, and movement patterns of outplanted urchins.

            Various methods of tagging urchins have been employed since the 1960s with mixed success (Ebert 1965; Ebert 2013; Santos et al., 2022; Tourón et al., 2023). Many internal physical tags such as decimal coded wire tags (CWTs) and passive integrated transponder (PIT) tags can cause increased mortality or decreased growth rates in urchins, especially in smaller individuals, while external physical markings such as painting spines or placing beads are quickly lost (Cipriano et al., 2014; Santos et al., 2022; Tourón et al., 2023). Chemically marking individuals with the fluorochrome dye calcein is a potential noninvasive and non-toxic means of tagging D. antillarum for long durations. The molecule typically binds irreversibly to calcium ions and becomes incorporated into the tissue and skeletal structure of animals as they grow, which causes the individual’s tissues to fluoresce under UV light (Johnson et al., 2013; Li et al., 2020). Although this method has been used to tag several species of urchins for months (Johnson et al., 2013; Rodríguez et al., 2016; Santos et al., 2022; Tourón et al., 2023), the technique has yet to be tested with D. antillarum. Here, we conducted a laboratory study to evaluate the efficacy of using calcein to tag D. antillarum for extended timeframes.

            Survival was 100% over the course of this study. Recently tagged D. antillarum fluoresced strongly under UV light and the markings were most visible when observed through a blue light filter (Figure 1a–c). Individuals differed in where tag fluorescence appeared on their body as fluorescence was either localized to their spines (Figure 1c) or tube feet along the ambulacral plates (Figure 1d). Fluorescence on completely black urchins followed similar intensity levels and duration regardless of whether it occurred on spines or tube feet. Tag fluorescence started to become difficult to detect visually in black spined urchins after two weeks and was unlikely to be observed under SCUBA diving conditions beyond this point (Figure 1e). Most tagged urchins (n = 10 of 13) were indistinguishable from untagged control urchins (n = 21) after five weeks; however, all three urchins with white spines (n = 3 of 13) retained fluorescence for at least 11 weeks (Figure 1f). This continued fluorescence was limited to the white spines (Figure 1g), possibly either because the darker pigmentation within black spines prevented UV light from reaching the incorporated calcein, or it may have attenuated emission visibility, masking calcein fluorescence. However, further molecular studies are needed to identify the mechanisms underlying differences in spine fluorescence. Interestingly, we observed four individuals periodically releasing a fluorescent substance aborally three days after removal from the dye bath. This may indicate post-exposure clearance or redistribution of dye-associated material, though chemical confirmation was not performed (Figure 1h–l).

Importantly, unlike previous studies where calcein dye was visible externally on urchins for months (Rogers-Bennett 2003; Johnson et al., 2013; Santos et al., 2022; Correia et al., 2023; Tourón et al., 2023), dye markings only remained visible on D. antillarum for approximately two weeks — despite our use of a more intense than typical dye regimen. Such contrasting results are likely due to differences in pigmentation of those species as Paracentrotus lividus (Santos et al., 2022), Strongylocentrotus droebachiensis (Johnson et al. 2013), Strongylocentrotus franciscanus (Rogers-Bennett 2003), and Pseudechinus huttoni (Kirby et al., 2006) are substantially lighter in coloration than D. antillarum. In the dark-colored Diadema africanum (Rodríguez et al., 2016) and Arbacia lixula (Barrera 2018), extended tag visibility was only achieved by dissection and microscopic examination of the Aristotle’s lantern. Consequently, we conclude that calcein dye is unsuitable for tagging D. antillarum for long durations when individual survival is a priority or disturbance must be minimized, but calcein dye may be sufficient for short-term studies or when dissection is an option. These results highlight how even biologically conserved chemical reactions can have different outcomes across similar species and the necessity for species-specific studies.

","references":[{"reference":"

Barrera AM. 2018. Estimación del crecimiento del erizo de mar Arbacia lixula. Máster en Biología marina: Biodiversidad y conservación. San Cristóbal de La Laguna, Spain: Universidad de la Laguna. http://riull.ull.es/xmlui/handle/915/23646

","pubmedId":"","doi":""},{"reference":"

Carpenter RC. 1981. Grazing by Diadema antillarum (Philippi) and its effects on the benthic algal community. Journal of Marine Research. 39: 749-765. https://elischolar.library.yale.edu/journal_of_marine_research/1568

","pubmedId":"","doi":""},{"reference":"

Carpenter RC. 1988. Mass mortality of a Caribbean sea urchin: Immediate effects on community metabolism and other herbivores. Proc Natl Acad Sci U S A 85(2): 511-4.

","pubmedId":"16593907","doi":""},{"reference":"

Ciriano A, Burnell G, Cullotv S, Long S. 2014. Evaluation of 3 Tagging Methods in Marking Sea Urchin,Paracentrotuslividus, Populations under Both Laboratory and Field Conditions. Journal of Aquaculture Research & Development 05: 10.4172/2155-9546.1000276.

","pubmedId":"","doi":"10.4172/2155-9546.1000276"},{"reference":"

Correia MJo, Mega Lopes Pc, Santos PM, Jacinto D, Mateus D, Maresca F, et al., Costa. 2023. Pilot studies for stock enhancement of purple sea urchins (Paracentrotus lividus, Lamarck, 1816): usefulness of refuges and calcein marking for the monitoring of juveniles released into the natural environment. Aquatic Living Resources 36: 12.

","pubmedId":"","doi":"10.1051/alr/2023009"},{"reference":"

Ebert TA. 1965. A Technique for the Individual Marking of Sea Urchins. Ecology 46: 193-194.

","pubmedId":"","doi":"10.2307/1935273"},{"reference":"

Ebert TA. 2013. Growth and survival of postsettlement sea urchins. In: Lawrence J.M. (ed), Sea Urchins: Biology and Ecology. Academic Press, San Diego, California, pp. 83-117.

","pubmedId":"","doi":"10.1016/B978-0-12-396491-5.00007-1"},{"reference":"

Edmunds PJ, Carpenter RC. 2001. Recovery of Diadema antillarum reduces macroalgal cover and increases abundance of juvenile corals on a Caribbean reef. Proc Natl Acad Sci U S A 98(9): 5067-71.

","pubmedId":"11274358","doi":""},{"reference":"

FAO. 2025. Fishery and aquaculture statistics – yearbook 2023. Rome.

","pubmedId":"","doi":"10.4060/cd6788en"},{"reference":"

FAO. 2024. The State of World Fisheries and Aquaculture 2024 – Blue Transformation in action. Rome.

","pubmedId":"","doi":"10.4060/cd0683en"},{"reference":"

Hudspeth CG, Pilnick AR, Petrosino A, Smith JN, Patterson JT. 2026. Developing standard production diets for early juvenile Diadema antillarum sea urchins. Aquaculture Reports 47: 103430.

","pubmedId":"","doi":"10.1016/j.aqrep.2026.103430"},{"reference":"

Hughes TP, Reed DC, Boyle MJ. 1987. Herbivory on coral reefs: community structure following mass mortalities of sea urchins. Journal of Experimental Marine Biology and Ecology 113: 39-59.

","pubmedId":"","doi":"10.1016/0022-0981(87)90081-5"},{"reference":"

Hylkema A, Debrot AO, van de Pas EE, Osinga R, Murk AJ. 2022. Assisted Natural Recovery: A Novel Approach to Enhance Diadema antillarum Recruitment. Frontiers in Marine Science 9: 10.3389/fmars.2022.929355.

","pubmedId":"","doi":"10.3389/fmars.2022.929355"},{"reference":"

Johnson AS, Salyers JM, Alcorn NJ, Ellers O, Allen JD. 2013. Externally visible fluorochrome marks and allometries of growing sea urchins. Invertebrate Biology 132: 251-269.

","pubmedId":"","doi":"10.1111/ivb.12024"},{"reference":"

Kirby S, Lamare MD, Barker MF. 2006. Growth and morphometrics in the New Zealand sea urchin\n Pseudechinus huttoni\n (Echinoidea: Temnopleuridae). New Zealand Journal of Marine and Freshwater Research 40: 413-428.

","pubmedId":"","doi":"10.1080/00288330.2006.9517432"},{"reference":"

Lessios HA. 1988. MASS MORTALITY OF DIADEMA ANTILLARUM IN THE CARIBBEAN: What Have We Learned?. Annual Review of Ecology and Systematics 19: 371-393.

","pubmedId":"","doi":"10.1146/annurev.es.19.110188.002103"},{"reference":"

Lessios HA. 2016. The Great Diadema antillarum Die-Off: 30 Years Later. Ann Rev Mar Sci 8: 267-83.

","pubmedId":"26048480","doi":""},{"reference":"

Li H, Lei Y, Li T, Jian Z. 2020. Effects of calcein incorporation on benthic foraminiferal community under various concentrations and incubation durations. Marine Micropaleontology 157: 101874.

","pubmedId":"","doi":"10.1016/j.marmicro.2020.101874"},{"reference":"

Ogden JC, Lobel PS. 1978. The role of herbivorous fishes and urchins in coral reef communities. Environmental Biology of Fishes 3: 49-63.

","pubmedId":"","doi":"10.1007/BF00006308"},{"reference":"

Pilnick AR, O'Neil KL, Moe M, Patterson JT. 2021. A novel system for intensive Diadema antillarum propagation as a step towards population enhancement. Sci Rep 11(1): 11244.

","pubmedId":"34045538","doi":""},{"reference":"

Rakaj A, Grosso L, Fianchini A, Cataudella S. 2024. A sustainable no-kill sea urchin aquaculture method to obtain caviar. Nature Sustainability 7: 1038-1047.

","pubmedId":"","doi":"10.1038/s41893-024-01372-0"},{"reference":"

Randall JE, Schroeder RE, Stark WA. 1963. Notes on the biology of the echinoid Diadema antillarum. Caribbean Journal of Science. 4: 2-3. https://www.researchgate.net/publication/246980787_Notes_on_the_biology_of_the_echinoid_Diadema_antillarum

","pubmedId":"","doi":""},{"reference":"

Rodríguez A, Hernández JC, Clemente S. 2016. Efficiency of calcein tagging on juveniles of the sea urchins Diadema africanum and Paracentrotus lividus. Marine Ecology 37: 463-469.

","pubmedId":"","doi":"10.1111/maec.12287"},{"reference":"

Rogers-Bennett L. 2003. Modeling red sea urchin growth using six growth functions. Fishery Bulletin. 101: 614-626. https://escholarship.org/uc/item/8gn2v2xn

","pubmedId":"","doi":""},{"reference":"

Santos PM, Ruivo Quintella B, Jacinto D, Gomes A, Saldanha C, Lourenço Sl, et al., Lino Costa. 2022. Physical and chemical tagging methods for the sea urchin Paracentrotus lividus (Echinodermata: Echinoidea). Scientia Marina 86: e038.

","pubmedId":"","doi":"10.3989/scimar.05259.038"},{"reference":"

Sharp WC, Delgado GA, Pilnick AR, Patterson JT. 2023. Diurnal Sheltering Behavior of Hatchery-propagated Long-spined Urchins (Diadema Antillarum): a Re-examination Following Husbandry Refinements. Bulletin of Marine Science 99: 97-108.

","pubmedId":"","doi":"10.5343/bms.2022.0054"},{"reference":"

Stefánsson G, Kristinsson H, Ziemer N, Hannon C, James P. 2017. Markets for sea urchins: a review of global supply and markets. Skýrsla Matís. 45: 10-17.

","pubmedId":"","doi":"10.13140/RG.2.2.12657.99683"},{"reference":"

Smith XJ, Dempster T, Salini M, Francis D, Warren-Myers F. 2026. Commercial scale test of purple sea urchin roe enhancement aquaculture. Aquaculture 618: 743804.

","pubmedId":"","doi":"10.1016/j.aquaculture.2026.743804"},{"reference":"

Tourón N, Campos S, Costas Dn, Paredes Ea. 2023. Marking Methodologies for Sea Urchins: A Review. Journal of Shellfish Research 42: 10.2983/035.042.0116.

","pubmedId":"","doi":"10.2983/035.042.0116"}],"title":"

The efficacy of calcein as a tagging agent for the long-spined sea urchin, Diadema antillarum

","reviews":[],"curatorReviews":[]},{"id":"4a0accde-1936-4aae-8e5f-7e3f06c74f31","decision":"publish","abstract":"

The sea urchin Diadema antillarum is a restoration target due to its function as a keystone reef herbivore. However, restocking efforts are hindered by a lack of long-term tagging methods. We tested the effectiveness of calcein marking for urchin releases. Cultured individuals were tagged using a 150 mg/L calcein bath and monitored for 11 weeks. Dye markings remained fully visible in all individuals for two weeks but began to fade within three weeks. Only individuals with white spines maintained markings beyond five weeks. Unlike with other urchin species, calcein only seems suitable for short-term external tagging of D. antillarum.

","acknowledgements":"

Many thanks go to Joshua Patterson of the University of Florida and The Florida Aquarium for providing the urchins, and to Justin Voss, Ellery Lennon, and Holly White for helping maintain the urchins.

","authors":[{"affiliations":["Florida Fish and Wildlife Conservation Commission, Marathon, FL, United States"],"departments":["Fish and Wildlife Research Institute"],"credit":["conceptualization","dataCuration","formalAnalysis","investigation","methodology","project","software","validation","visualization","writing_originalDraft","writing_reviewEditing"],"email":"Benjamin.Belgrad@MyFWC.com","firstName":"Benjamin A.","lastName":"Belgrad","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":"0000-0003-2032-2210"},{"affiliations":["Florida Fish and Wildlife Conservation Commission"],"departments":["Fish and Wildlife Research Institute"],"credit":["conceptualization","dataCuration","fundingAcquisition","investigation","methodology","resources","writing_reviewEditing"],"email":"Lindsay.Spiers@MyFWC.com","firstName":"Lindsay J.","lastName":"Spiers","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":null},{"affiliations":["Florida Fish and Wildlife Conservation Commission"],"departments":["Fish and Wildlife Research Institute"],"credit":["conceptualization","fundingAcquisition","methodology","project","resources","validation","writing_reviewEditing"],"email":"William.Sharp@MyFWC.com","firstName":"William C.","lastName":"Sharp","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":null}],"awards":[],"conflictsOfInterest":"

The authors declare that there are no conflicts of interest present.

","dataTable":{"url":null},"extendedData":[],"funding":"

This work was supported by funds provided to the Florida Fish and Wildlife Conservation Commission’s Marine Resources Trust Fund.

","image":{"url":"https://portal.micropublication.org/uploads/f607c75e6fda74cf3a296d4f7f0e8a4d.png"},"imageCaption":"

Photographs of the same two urchins tagged with calcein under a) normal light, b) unfiltered UV light, and c) UV light viewed through a blue light filter. Fluorescence from tagging was best viewed under blue filtered UV light and was observed on either c) the spines or d) tube feet along the ambulacral plates. e) Number of urchins with only black spines or with some white spines still exhibiting fluorescent tagging each week (n = 13 urchins tagged total). Urchins with black spines had fluorescence intensity notably decline and become harder to detect visually three weeks after tagging. f) Urchins with white spines that were either dyed with calcein (tagged) or undyed (control) 10 weeks after tagging. g) A black and white spine from a tagged and control urchin viewed under filtered UV light. h–l) Photo series spanning ~30 seconds of an urchin expelling fluorescent substance three days after tagging. Yellow arrow indicates initial material expulsion.

","imageTitle":"

Photographs of calcein tagged and control urchins and duration of tag visibility

","methods":"

Juvenile Diadema antillarum, spawned from wild-caught broodstock, were acquired from The Florida Aquarium's Center for Conservation, Apollo Beach, FL, US, in May 2024. Individuals were transported to the Florida Fish and Wildlife Research Institute’s South Florida Regional Lab in Marathon, FL, US, where they were nursed for approximately 21 months before commencing the experiment. During the nursery period, urchins were held in circular flow-through tanks (diameter = 155 cm, water height = 70 cm) under similar conditions to Sharp et al. (2023). Sand-filtered seawater was supplied from Florida Bay and matched natural abiotic conditions. Shelter was provided to urchins in the form of longitudinally halved terracotta pots (22 cm L × 22 cm W × 12 cm H) that mimicked natural coral reef crevice habitat, and urchins were fed satiating amounts of drift algae.

            Thirty-four urchins were haphazardly selected for tagging (n = 13; test diameter = 27–43 mm) and control treatments (n = 21; test diameter = 32–61 mm) to verify that untagged urchins do not fluoresce and determine if tagging influences survival. Urchins selected for tagging were distributed across three continuously aerated tanks containing a solution of sand-filtered seawater with 150 mg/L concentration of dissolved calcein. Two tanks held five urchins with 11 L calcein solution (Chemcruz source) while the third tank held three urchins with 7 L calcein solution (MP Biomedicals source). These volumes and urchin numbers were chosen based on perceived physiological tolerances, and the compounds from both sources produced the same results. Urchins were immersed in this dye bath for seven days (typical dye regimens use 100–150 mg/L for 24–48 h immersion). This extended period of immersion was performed because our preliminary experiments noted that fluorescence tagging disappeared quickly, and we wanted to identify the maximum amount of time tagging can be viewed externally on pure black-spined urchins, which comprise the majority of D. antillarum populations.

            After this tagging period, control and tagged urchins were placed into two separate 356-L rectangular aquaria (120 cm L × 75 cm W × 36 cm H) and maintained under similar conditions as the nursery period. Individuals were held for 11 weeks and checked at least weekly to determine if fluorescent markings were still observable. Markings were observed by shining a Sola NIGHTSEA dive light (emission spectra 420–470 nm, peaking at ~445 nm) on individuals and viewing them through a NIGHTSEA blue blocking light filter visor (blocking wavelengths shorter than 500 nm).

","reagents":"

Reagent

Alternate names

Lot Number

Source

Calcein disodium salt

Fluorexon Sodium Salt

C2425

ChemCruz

Calcein

Fluorexon; Fluorescein-bis(methyliminodiacetic acid);

Bis[N,N−bis(carboxymethyl)aminomethyl]fluorescein;

3,3'-bis[N,N'-di(carboxymethyl)-aminomethyl]-fluorescein; Calcein W

S8883-2

MP Biomedicals

","patternDescription":"

Sea urchins comprise a global multimillion dollar fishery and are ecological engineers (Ogden et al., 1978; Stefánsson et al., 2017; FAO 2025). As their populations have declined, urchins are increasingly cultivated around the world for consumption and restoration purposes (FAO 2024; Rakaj et al., 2024; Smith et al., 2026). The long-spined sea urchin, Diadema antillarum, is a keystone herbivore native to the tropical Atlantic and Caribbean coral reefs (Randall et al., 1963; Carpenter 1981; Carpenter 1988; Lessios 2016). In the 1980s, an epidemic spread throughout the Caribbean and killed 93–98% of the urchin populations (Carpenter 1988; Lessios 1988). This was followed by an enormous increase in benthic macroalgae and a decrease in coral cover as the coral was outcompeted for space and light (Hughes et al., 1987). D. antillarum populations have only modestly recovered since this epidemic (~12% population pre-epidemic; Lessios 2016), and the species has recently become a target of restoration to aid coral recovery (Pilnick et al., 2021; Hylkema et al., 2022). In areas where D. antillarum populations have increased in abundance, there have been significant decreases in macroalgal cover and, in some areas, increases in coral recruitment (Edmunds and Carpenter 2001). The restoration effort has centered around supplementing wild populations with hatchery reared juveniles (Pilnick et al., 2021; Hudspeth et al., 2026), as natural recruitment is low and early life-stage predation is high (Lessios 2016; Hylkema et al., 2022). However, restocking efforts and research on the effectiveness of this approach are hampered by a lack of viable long-term tagging methods. Reliable long-term tagging is essential to obtain information that is largely lacking and fundamental to developing and optimizing restoration strategies for this species, such as post-release survival, individual growth rates, and movement patterns of outplanted urchins.

            Various methods of tagging urchins have been employed since the 1960s with mixed success (Ebert 1965; Ebert 2013; Santos et al., 2022; Tourón et al., 2023). Many internal physical tags such as decimal coded wire tags (CWTs) and passive integrated transponder (PIT) tags can cause increased mortality or decreased growth rates in urchins, especially in smaller individuals, while external physical markings such as painting spines or placing beads are quickly lost (Cipriano et al., 2014; Santos et al., 2022; Tourón et al., 2023). Chemically marking individuals with the fluorochrome dye calcein is a potential noninvasive and non-toxic means of tagging D. antillarum for long durations. The molecule typically binds irreversibly to calcium ions and becomes incorporated into the tissue and skeletal structure of animals as they grow, which causes the individual’s tissues to fluoresce under UV light (Johnson et al., 2013; Li et al., 2020). Although this method has been used to tag several species of urchins for months (Johnson et al., 2013; Rodríguez et al., 2016; Santos et al., 2022; Tourón et al., 2023), the technique has yet to be tested with D. antillarum. Here, we conducted a laboratory study to evaluate the efficacy of using calcein to tag D. antillarum for extended timeframes.

            Survival was 100% over the course of this study. Recently tagged D. antillarum fluoresced strongly under UV light and the markings were most visible when observed through a blue light filter (Figure 1a–c). Individuals differed in where tag fluorescence appeared on their body as fluorescence was either localized to their spines (Figure 1c) or tube feet along the ambulacral plates (Figure 1d). Fluorescence on completely black urchins followed similar intensity levels and duration regardless of whether it occurred on spines or tube feet. Tag fluorescence started to become difficult to detect visually in black spined urchins after two weeks and was unlikely to be observed under SCUBA diving conditions beyond this point (Figure 1e). Most tagged urchins (n = 10 of 13) were indistinguishable from untagged control urchins (n = 21) after five weeks; however, all three urchins with white spines (n = 3 of 13) retained fluorescence for at least 11 weeks (Figure 1f). This continued fluorescence was limited to the white spines (Figure 1g), possibly either because the darker pigmentation within black spines prevented UV light from reaching the incorporated calcein, or it may have attenuated emission visibility, masking calcein fluorescence. However, further molecular studies are needed to identify the mechanisms underlying differences in spine fluorescence. Interestingly, we observed four individuals periodically releasing a fluorescent substance aborally three days after removal from the dye bath. This may indicate post-exposure clearance or redistribution of dye-associated material, though chemical confirmation was not performed (Figure 1h–l).

Importantly, unlike previous studies where calcein dye was visible externally on urchins for months (Rogers-Bennett 2003; Johnson et al., 2013; Santos et al., 2022; Correia et al., 2023; Tourón et al., 2023), dye markings only remained visible on D. antillarum for approximately two weeks — despite our use of a more intense than typical dye regimen. Such contrasting results are likely due to differences in pigmentation of those species as Paracentrotus lividus (Santos et al., 2022), Strongylocentrotus droebachiensis (Johnson et al. 2013), Strongylocentrotus franciscanus (Rogers-Bennett 2003), and Pseudechinus huttoni (Kirby et al., 2006) are substantially lighter in coloration than D. antillarum. In the dark-colored Diadema africanum (Rodríguez et al., 2016) and Arbacia lixula (Barrera 2018), extended tag visibility was only achieved by dissection and microscopic examination of the Aristotle’s lantern. Consequently, we conclude that calcein dye is unsuitable for tagging D. antillarum for long durations when individual survival is a priority or disturbance must be minimized, but calcein dye may be sufficient for short-term studies or when dissection is an option. These results highlight how even biologically conserved chemical reactions can have different outcomes across similar species and the necessity for species-specific studies.

","references":[{"reference":"

Barrera AM. 2018. Estimación del crecimiento del erizo de mar Arbacia lixula. Máster en Biología marina: Biodiversidad y conservación. San Cristóbal de La Laguna, Spain: Universidad de la Laguna. http://riull.ull.es/xmlui/handle/915/23646

","pubmedId":"","doi":""},{"reference":"

Carpenter RC. 1981. Grazing by Diadema antillarum (Philippi) and its effects on the benthic algal community. Journal of Marine Research. 39: 749-765. https://elischolar.library.yale.edu/journal_of_marine_research/1568

","pubmedId":"","doi":""},{"reference":"

Carpenter RC. 1988. Mass mortality of a Caribbean sea urchin: Immediate effects on community metabolism and other herbivores. Proc Natl Acad Sci U S A 85(2): 511-4.

","pubmedId":"16593907","doi":""},{"reference":"

Ciriano A, Burnell G, Cullotv S, Long S. 2014. Evaluation of 3 Tagging Methods in Marking Sea Urchin,Paracentrotuslividus, Populations under Both Laboratory and Field Conditions. Journal of Aquaculture Research & Development 05: 10.4172/2155-9546.1000276.

","pubmedId":"","doi":"10.4172/2155-9546.1000276"},{"reference":"

Correia MJo, Mega Lopes Pc, Santos PM, Jacinto D, Mateus D, Maresca F, et al., Costa. 2023. Pilot studies for stock enhancement of purple sea urchins (Paracentrotus lividus, Lamarck, 1816): usefulness of refuges and calcein marking for the monitoring of juveniles released into the natural environment. Aquatic Living Resources 36: 12.

","pubmedId":"","doi":"10.1051/alr/2023009"},{"reference":"

Ebert TA. 1965. A Technique for the Individual Marking of Sea Urchins. Ecology 46: 193-194.

","pubmedId":"","doi":"10.2307/1935273"},{"reference":"

Ebert TA. 2013. Growth and survival of postsettlement sea urchins. In: Lawrence J.M. (ed), Sea Urchins: Biology and Ecology. Academic Press, San Diego, California, pp. 83-117.

","pubmedId":"","doi":"10.1016/B978-0-12-396491-5.00007-1"},{"reference":"

Edmunds PJ, Carpenter RC. 2001. Recovery of Diadema antillarum reduces macroalgal cover and increases abundance of juvenile corals on a Caribbean reef. Proc Natl Acad Sci U S A 98(9): 5067-71.

","pubmedId":"11274358","doi":""},{"reference":"

FAO. 2025. Fishery and aquaculture statistics – yearbook 2023. Rome.

","pubmedId":"","doi":"10.4060/cd6788en"},{"reference":"

FAO. 2024. The State of World Fisheries and Aquaculture 2024 – Blue Transformation in action. Rome.

","pubmedId":"","doi":"10.4060/cd0683en"},{"reference":"

Hudspeth CG, Pilnick AR, Petrosino A, Smith JN, Patterson JT. 2026. Developing standard production diets for early juvenile Diadema antillarum sea urchins. Aquaculture Reports 47: 103430.

","pubmedId":"","doi":"10.1016/j.aqrep.2026.103430"},{"reference":"

Hughes TP, Reed DC, Boyle MJ. 1987. Herbivory on coral reefs: community structure following mass mortalities of sea urchins. Journal of Experimental Marine Biology and Ecology 113: 39-59.

","pubmedId":"","doi":"10.1016/0022-0981(87)90081-5"},{"reference":"

Hylkema A, Debrot AO, van de Pas EE, Osinga R, Murk AJ. 2022. Assisted Natural Recovery: A Novel Approach to Enhance Diadema antillarum Recruitment. Frontiers in Marine Science 9: 10.3389/fmars.2022.929355.

","pubmedId":"","doi":"10.3389/fmars.2022.929355"},{"reference":"

Johnson AS, Salyers JM, Alcorn NJ, Ellers O, Allen JD. 2013. Externally visible fluorochrome marks and allometries of growing sea urchins. Invertebrate Biology 132: 251-269.

","pubmedId":"","doi":"10.1111/ivb.12024"},{"reference":"

Kirby S, Lamare MD, Barker MF. 2006. Growth and morphometrics in the New Zealand sea urchin\n Pseudechinus huttoni\n (Echinoidea: Temnopleuridae). New Zealand Journal of Marine and Freshwater Research 40: 413-428.

","pubmedId":"","doi":"10.1080/00288330.2006.9517432"},{"reference":"

Lessios HA. 1988. MASS MORTALITY OF DIADEMA ANTILLARUM IN THE CARIBBEAN: What Have We Learned?. Annual Review of Ecology and Systematics 19: 371-393.

","pubmedId":"","doi":"10.1146/annurev.es.19.110188.002103"},{"reference":"

Lessios HA. 2016. The Great Diadema antillarum Die-Off: 30 Years Later. Ann Rev Mar Sci 8: 267-83.

","pubmedId":"26048480","doi":""},{"reference":"

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The efficacy of calcein as a tagging agent for the long-spined sea urchin, Diadema antillarum

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