Desperate Larva or Death Before Dishonor: can old coral larvae replenish degraded reefs?

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Nepsis García, Margaret Miller, D. Abigail Renegar, Joana Figueiredo This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6874996/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 09 Feb, 2026 Read the published version in Coral Reefs → Version 1 posted 12 You are reading this latest preprint version Abstract The recovery of coral populations depends largely on larval recruitment. Coral larvae settle in response to environmental cues that indicate habitat quality. In the absence of suitable cues, lecithotrophic larvae may delay settlement until they die (Death Before Dishonor Hypothesis) or become increasingly less discriminatory (Desperate Larva Hypothesis). To examine which of these hypothesized strategies corals follow, we used larvae from three broadcast spawning reef-building corals, Diploria labyrinthiformis, Pseudodiploria clivosa , and Orbicella faveolata , and one brooding, weedy coral, Agaricia agaricites. Settlement behaviors of larvae of different ages, specifically 0, 7, 14, or 21 days after becoming competent, were examined in response to suitable (crustose coralline algae) and unsuitable ( Dictyota sp .) settlement cues. Potential latent effects of delayed metamorphosis on post-settlement survival and growth were also assessed. As larvae got older, broadcast spawning species decreased selectivity (‘Desperate Larva Hypothesis’), whereas the brooding species’ larvae never did (‘Death Before Dishonor Hypothesis’). Delayed settlement of the brooder A. agaricites produced latent effects on post-settlement survival and growth, while O. faveolata displayed latent effects of delayed settlement on post-settlement growth. The other broadcast spawners exhibited no latent effects of delayed settlement on post-settlement. This suggests “old” coral larvae from broadcast spawners have the potential to disperse widely to replenish degraded reefs. The brooding, weedy coral A. agaricites seems to have evolved to disfavor dispersal and maximize local retention, evidenced by the fast acquisition of competency and latent effects of delayed settlement combined with an early loss of competency. delayed settlement latent effects post-settlement recruitment Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction The capacity of coral populations to recover from disturbances depends greatly on their replenishment by coral larvae from nearby and distant populations (Lukoschek et al. 2013 ; Holbrook et al. 2018 ). Reef connectivity through coral larval dispersal allows the exchange of genes among locations and enhances reef resilience (Goodbody-Gringley et al. 2012 ). The onset and duration of the larval competency period determines dispersal time and, consequently, the extent of their distribution and connectivity (Figueiredo et al. 2022 ; King et al. 2023 ). The longer the larvae remain competent, the farther they can disperse and replenish distant populations. Brooding species typically release readily competent larvae, and thus tend to settle on or closer to their natal reef; while the larvae of broadcast spawning species take longer to develop and only acquire competency 1–6 days after spawning, reducing their local retention (Sammarco et al. 1989; Figueiredo et al. 2013 ). Corals have lecithotrophic larvae (i.e., sustained by energy reserves placed in the egg), and because maternal provisioning is limited, so is their competency period (Richmond et al. 2018 ). However, the larvae of some coral species, mostly brooders but not exclusively (see Baird et al. 2009 ), contain symbiotic zooxanthellae which can supplement the coral larval energy via photosynthesis and thus might extend the larva’s pelagic larval duration (Chamberland et al. 2017 ). Larval settlement is a crucial process in a coral’s life history since it can directly influence its successful recruitment and long-term chances to survive, grow, and eventually sexually reproduce (Ritson-Williams et al. 2020 ). Coral planulae have been shown to utilize specific cues to determine if a habitat is suitable to settle and metamorphose (Golbuu and Richmond 2007 ). Whereas bacterial films (Tebben et al. 2015 ) and crustose coralline algae (CCA) have been shown to induce settlement in coral larvae (Heyward and Negri 1999 ; Negri and Heyward 2000 ; Hadfield and Paul 2001 ; Negri et al. 2001 ; Ritson-Williams et al. 2016 ), some macroalgae and cyanobacteria species are known settlement inhibitors (Kuffner et al. 2006 ; Birrell et al. 2008 ; Arnold et al. 2010 ; Diaz-Pulido et al. 2010 ; Doropoulos et al. 2014 ; Webster et al. 2015 ; Ritson-Williams et al. 2020 ). Changes to the ecological composition of benthic communities on coral reefs, such as the dominance shift from stony corals to fleshy macroalgae (Arias-González et al. 2017 ), may jeopardize the settlement of coral larvae due to a lack of inducing settlement cues and/or the presence of inhibitory cues. As lecithotrophic larvae extend their larval period in the search for a suitable settlement site at the cost of depleting their energy reserves, they follow one of two pathways: decreased selectivity, known as the Desperate Larva Hypothesis (i.e. as larvae age, they become indiscriminate or “desperate” and settle on non-suitable substrates), or continued searching for a specific habitat or requirement to the point of depleting their energy reserves and perishing without settling and metamorphosing, known as the Death Before Dishonor Hypothesis (Bishop et al. 2006 ). The adoption of either of these pathways varies among species. The Desperate Larva Hypothesis was validated in some species in the phylum Bryozoa (Gribben et al. 2006 ), Arthropoda (Harvey and Colasurdo 1993 ; Miron et al. 2000 ; Thiyagarajan et al. 2002 ), Echinodermata (Rumrill 1989 ; Swanson et al. 2007), Annelida (Pechenik and Qian 1998 ), Mollusca (Pechenik 1984 ; Lima and Pechenik 1985 ; Carroll and Kempf 1990 ; Gibson 1995 ; Avila 1998 ) and Phoronida (Herrmann 1979 ; Herrmann 1995 ). Conversely, the Death Before Dishonor Hypothesis has only been validated in some phoronids, annelids and mollusks (Kempf 1981 ; Hadfield and Pennington 1990 ; Lambert and Todd 1994 ; Krug 2001 ; Santagata 2004 ; Botello and Krug 2006 ). Following one or the other strategy appears closely linked with the strictness of their post-settlement habitat requirements (i.e., generalists or specialists, Elkin and Marshall 2007 ), e.g., food and host specialists tend to follow the Death Before Dishonor Hypothesis. It remains unknown which pathway, desperate larva or death before dishonor, corals follow and if it varies depending on the species’ life history strategy. Delaying settlement can have latent effects on post-settlement survival (Miller 1993 ), growth (Pechenik et al. 2002 ), and reproduction success (e.g., lower fecundity, Miller 1993 ) of several invertebrates, including bryozoans (Wendt 1998 ; Sams et al. 2015 ), ascidians (Marshall et al. 2003 ), sponges (Maldonado and Young 1999 ), and mollusks (Roberts and Lapworth 2001 ). For benthic species, delaying settlement might reduce the energy reserves in the newly settled individuals, making them smaller and more vulnerable to fluctuations in food availability, predation and overgrowth (Qian and Pechenik 1998 ). Little is known about the effect of delaying larval settlement on corals. The only study to date found Acropora tenuis settled 2, 4 and 6 weeks after spawning did not exhibit latent effects on post-settlement survival and growth (Graham et al. 2013 ). Studies focusing on other species are needed to better understand how delayed settlement might affect scleractinians. This study aimed to first address whether the selectivity of larvae from four Caribbean Scleractinia coral species for suitable settlement habitats diminished with age. Secondly, we examined whether the age at which larvae settled has latent effects on their post-settlement survival and growth. The results of this study suggest that the paradigm that disturbed reefs cannot recover through larval replenishment may be, at least partly, flawed. Materials and Methods Study species and Coral collection The speciesused in this experiment are common in Florida’s Coral Reef and throughout the Caribbean, are susceptible to stony coral tissue loss disease (SCTLD), and have different reproductive modes. Diploria labyrinthiformis, Orbicella faveolata , and Pseudodiploria clivosa are reef-building, hermaphroditic, broadcast-spawning corals, while Agaricia agaricites is a resilient, weedy, hermaphroditic, brooding species (Richmond and Hunter 1990 ; Carlon 1999 ; Knowlton 2001 ). Twenty adult colonies of Agaricia agaricites (12–15 cm diameter) were collected from the Dania Erojacks, off the coast of Dania Beach (Florida, U.S.A.) via SCUBA a week before the new moon of May 2022, the expected larval release window (Van Moorsel 1983 ). The colonies were transported in coolers with seawater, wrapped in bubble wrap, and then housed in a 1500 L recirculating aquarium at Nova Southeastern University (NSU). The colonies were placed into individual 3L bowls within a recirculating tank, keeping the water level in the tank at ¾ of the bowl’s height to maintain water temperature. Filtered seawater was supplied to each bowl using a manifold system with rubber tubing at a rate of 1.3 L min − 1 . Water then exited the bowl through the bowl’s handle into PVC cups filled at least halfway with water, fitted with plankton mesh (100 µm) at the bottom, that allowed constant water change while larvae were retained. Colonies released larvae from 4 days before to 6 days after the new moon, which were collected from the cups daily at the end of the expected larval release window (between 2300 and 0700 h, Van Moorsel 1983 ). The larvae from each broadcast spawning species were donated by different institutions and transported to NSU (Appendix); their competency was inferred by initiation of downward swimming. Settlement Approximately 3000 larvae per species were equally and randomly split among six glass Mason jars filled with 500 mL of seawater and covered with parafilm with holes to reduce evaporation rates whilst allowing oxygenation. The jars were kept on an orbital shaker (Troemner™ Talboys™) at 130–180 rpm for the duration of the experiment to mimic ocean water movement and prevent settlement. Water changes were performed daily. We tested the effect of delayed settlement (larval age) on settlement success of coral larvae exposed to suitable (crustose coralline algae) and unsuitable ( Dictyota sp .) settlement cues, respectively mimicking “healthy reef” and “degraded reef” habitats. To do so, for each of these two treatments, ten replicates of 10 or 15 larvae were randomly collected from all six jars at four times points: 0, 7, 14, and 21 days after becoming competent, hereafter referred to as “larval age” (Appendix). Specifically, each replicate consisted of 10 or 15 larvae in a 150 mL glass jar filled with 100 mL of artificial seawater with one labelled 3.2 cm 2 ceramic settlement tile (Ocean Wonders) with the rough side facing up. For the healthy reef treatment, the tiles used were preconditioned in a CCA-covered aquarium with adult corals for 4 weeks to allow microbial biofilm to grow. A mixture of crustose coralline algae, (that previously induced settlement for multiple species) was scraped off the aquarium’s walls, collected in a 50 mL polypropylene (PP) tube (Falcon®), ground into a fine dust using a mortar and pestle, and sprinkled on top of the preconditioned tile. For the “degraded reef” treatment, a 0.5×1 cm piece of Dictyota spp. sp . from NSU tanks, a macroalga that inhibits coral settlement and reduces post-settlement survival (Olsen et al. 2014 ; Paul et al. 2011 ) was placed on top of the tile. All jars were placed into a 2.4 L Polystyrene hinged food container (PACTIV) to reduce evaporation rates but allow for oxygenation, and jars were topped off daily with RO-DI water to maintain a salinity of 35 ppt. Larvae were left with the tiles for 48 h, after which they were assessed under a dissecting microscope (Olympus SZ61). Corals were considered settled and metamorphosed when attached to the substrate and polyp-shaped. Post-settlement survival and growth assessments Once metamorphosis was complete, the location of each coral on the tiles was mapped to fate-track their survival and growth over time. Tiles were placed in a 42 L rectangular polypropylene (PP) tank with indirect sunlight (photosynthetically active radiation (PAR) of 4–5 µmol photons m − 2 s − 1 ), an air bubbler, and a Titanium heater (Finnex TH Deluxe). Corals were fed 5 mL of CoralAminos (Brightwell Aquatics) twice a week. The tank was topped off daily with DO water. (Sea)water was fully exchanged twice a week. Approximately 4 weeks after settlement, the newly settled corals that did not receive symbionts from a parent colony (i.e., vertical transmission), were inoculated with a cocktail of laboratory cultured Breviolum minutum , Durusdinium trenchii , Fugacium kawagutii and Symbiodinium microadriaticum . Artificial overhead lights (Hydra Twenty-Six HD) set to 20 µmol photons m − 2 s − 1 were added to the tank 2 weeks post-settlement. After this period, the juveniles were gradually acclimated into an indoor recirculating aquarium for grow-out, where adult colonies were also held, which allowed them to continue acquiring a diverse Symbiodiniaceae assemblage. The system consisted of two connected 454 L raceways with a common 341 L sump equipped with a Titanium heater (Finnex TH Deluxe), drop-in chiller (Aqua Logic Cy-3), Protein Skimmer (ASM G-3), two UV Sterilizers (Pentair AES EU25-U), phosphate reactor (PhosBan Reactor 550), calcium reactor (GEO with Sicce Syncra Pump), 25 kg of live rock, and a 45.7×30.5 cm perforated plastic basket full of bioballs. The system was set to mimic a truncated annual temperature cycle (23.3°C in February to 29.2°C in September) based on SECREMP data from Southeast Florida Reef Tract from 2007 to 2018 (Gilliam et al. 2019 ), excluding years when mass coral bleaching happened. In this system, the juveniles were positioned under LED lights (Radion® G5 XR30) set to a PAR 35 µmol photons m − 2 s − 1 , which were ramped up weekly by 10–15 µmol photons m − 2 s − 1 , until 200 µmol photons m − 2 s − 1 was reached. Once the juveniles were transferred into the recirculating aquarium, they were target fed 4 times a week with a 0.25 L slurry containing Reef-Roids (Polyp Lab ®), Oyster Feast (Reef Nutrition®), CoralAmino (Brightwell Aquatics), and live rotifers that were enriched with a blend of microalgae (Rotigrow Plus®) 1 h before being prepared for feeding. The water flow was stopped for 1 h to allow the corals to eat. The tiles were hand-cleaned weekly due to algal growth. Post-settlement survival and growth of juvenile corals were assessed monthly for three months by taking pictures of each tile with a digital camera (Olympus LC20) attached to a dissecting microscope (Olympus SZ61) and analyzed on cellSens imaging software (Olympus). When a juvenile coral was found with live tissue, it was marked as a living individual. Living individuals were measured by drawing a closed polygon around their live tissue (excluding tentacles; Fig. 1 ). Data analysis To test the effect of species, settlement cue treatment, and larval age on percent settlement, a Generalized Linear Model (GLM) with a binomial distribution (and then quasibinomial due to overdispersion) was performed in R using the package stats, with species ( Agaricia agaricites , Diploria labyrinthiformis , Orbicella faveolata , and Pseudodiploria clivosa ), settlement cue [healthy reef (conditioned tiles with ground CCA) and degraded reef (non-conditioned tiles with Dictyota sp.], and larval age (0, 7, 14, and 21 days after becoming competent) as fixed predictors and percent settlement as response variable. Since settlement significantly differed between species, GLMs were conducted separately for each species (same variables as described above excluding species), followed by multiple comparison post-hoc tests using the library emmeans. A higher settlement of newly-competent larvae (0 days after becoming competent) on tiles with CCA vs. Dictyota confirms that the CCA is the preferred settlement cue. An increased settlement on tiles with Dictyota as larvae age validates the Desperate Larva Hypothesis, while a sustained, low settlement validates the Death Before Dishonor hypothesis. An increase in settlement on tiles with CCA as larvae age suggests increased percent competency over time and/or that the CCA species used was sub-optimal cue for that species (and thus larvae are just settling more due to desperation). A decrease in settlement on tiles with CCA and/or Dictyota as larvae age is indicative of the onset of loss of competency. To analyze the effect of larval age on post-settlement survival over time for each species, a Survival Analysis Mantel–Haenszel log-rank test was performed using the R package survival, with larval age (levels: 0, 7, 14, and 21 days after becoming competent) as categorical predictor, and time as continuous predictor. For each coral, the time until the event of interest (mortality) occurred was recorded; if the coral remained alive at the end of the experiment, that time was recorded as right-censored. A lower post-settlement survival for corals whose settlement was delayed (i.e. older age) indicates latent effects on post-settlement survival. To test the effect of species and larval age on the post-settlement growth over time, a Generalized Linear Mixed-effects Model (GLMM) with a gamma log distribution was performed using R package glmmTMB, with species ( Agaricia agaricites , Diploria labyrinthiformis , Orbicella faveolata , and Pseudodiploria clivosa) and larval age (0, 7, 14, and 21 days after becoming competent) as fixed predictors, time as continuous predictor, coral individual as random predictor, and surface area as the response. Since species was found to be a significant predictor, the growth models were instead analyzed separately for each species, with all the other variables remaining the same. A lower post-settlement growth for corals which settlement was delayed (i.e. older age) indicates latent effects on post-settlement growth. All tests were run in R version 4.4.2 using RStudio version 2024.12.0. Results Settlement Percent settlement was significantly different among species (p < 2.2×10 − 16 ) and was significantly affected by both settlement cue (p < 2.2×10 − 16 ) and larval age (p = 4.3×10 − 3 ), with significant interactions between all variables (all p ≤ 0.002). Together, coral species, settlement cue and larval age explained almost 72% of the variance in larval settlement (R 2 = 0.717). For Agaricia agaricites , settlement cue (p < 2.2×10 − 16 ) and larval age (p = 3.6×10 − 4 ) had a significant effect on percent settlement, explaining over 78% of the variance in settlement (R 2 = 0.781), but they did not significantly interact (p = 0.42). When newly-competent larvae (larval age 0), were exposed to the conditioned tile with CCA, a very high percentage of larvae settled (mean ± SE: 91 ± 4%), but settlement progressively declined with each week settlement was delayed (79 ± 4%, 60 ± 4%, 50 ± 4% for larval ages 7, 14 and 21, respectively, Fig. 2 ), suggesting the onset of loss of competency. Regardless of the larva’s age, very few to no larvae settled on the tile with Dictyota sp. (3 ± 2%, 5 ± 5%, 0%, 0%, respectively for larval ages 0, 7, 14 and 21), validating the Death Before Dishonor hypothesis for this species. Diploria labyrinthiformis larvae settled significantly less on tile with Dictyota sp. than with CCA (p < 2.2×10 − 16 ). Larval age at the time of settlement did not significantly affect percent larval settlement (p = 0.071), but it significantly interacted with the settlement cue (p = 1.4 × 10 − 4 ), together explaining nearly 75% of the variance in settlement (R 2 = 0.748). When exposed to CCA, an average of 52 ± 3% of the newly competent (i.e. larval age 0) settled, versus only 5 ± 2% with Dictyota sp.. The average percentage settlement on the tile with CCA increased significantly when settlement was delayed by 7 days (73 ± 3%, p = 0.02) suggesting that not all larvae may have been competent at the beginning of the trial, i.e. by larval age 0 (as it had been assumed), with some larvae only acquiring competence afterwards and/or the settlement cue used was sub-optimal and the increased settlement with larval age was a result of desperation. Larval settlement remained high when delayed for 14 days (65 ± 8%), but it declined significantly if delayed for one further week (45 ± 7%, p = 0.04, Fig. 2 ), suggesting the onset of loss of competency is sometime within this period. As larvae aged, the average percent settlement on the tiles with Dictyota sp. remained low, but its average increased slightly with larval age, being significantly higher for larvae with 21 days of age (4 ± 2%, 11 ± 2% and 19 ± 2%, respectively, for larval ages 7, 14 and 21 days); suggesting this species follows the Desperate Larva hypothesis. The larvae of Orbicella faveolata displayed the lowest settlement among the studied species, yet they still settled more on the tiles with CCA vs. the tiles with Dictyota sp. . While percent settlement significantly differed among settlement cue treatments (p = 5.7×10 − 5 ) and larval ages (p = 1.4×10 − 2 ), the two variables only explained 28% of the variance in settlement (R 2 = 0.283) and they did not significantly interact (p = 0.08). Larvae exposed to CCA initially displayed with low settlement, but settlement progressively increased with larval age (12 ± 4%, 15 ± 7%, 26 ± 8%, and 30 ± 2%, respectively for larval ages 0, 7, 14 and 21 days, Fig. 2 ), being significantly higher when settlement was delayed by 21 days (compared to larvae with 0 and 7 days, p = 0.02 and p = 0.04, respectively), suggesting the CCA cue used was sub-optimal, and thus that the increased settlement was a result of desperation. In the presence of Dictyota sp. , settlement was even lower (7 ± 3% 5 ± 2%, 1 ± 1%, and 16 ± 4%, respectively for larval ages 0, 7, 14 and 21 days), with the settlement of larvae which were only exposed to the cue 21 days after acquiring competency also being significantly higher (compared to larvae of 0 and 7 days, p = 0.02 and p = 0.04, respectively), once more suggesting this species follows the Desperate Larva hypothesis. For Pseudodiploria clivosa , larval age (p = 3.7×10 − 6 ) and settlement cue (p = 4.7×10 − 13 ), significantly affected percent larval settlement, and significantly interacted (p = 5×10 − 5 ); together they explained 65% of the variance in percent settlement (R 2 = 0.65). The larval settlement in the presence of CCA was relatively higher (32 ± 8%, 43 ± 5% 29 ± 5% and 23 ± 5% for larvae exposed 0, 7, 15 and 21 days after acquisition of competency, respectively) (Fig. 2 ). Conversely, newly competent larvae did not settle in the tiles with Dictyota sp. (0 ± 0%); only larvae which settlement was delayed by 7 or more days settled on them (24 ± 5%, 3 ± 1%, 1 ± 1%, respectively for larval ages 0, 7, 14 and 21). The drastic increase in settlement on tiles with Dictyota sp. from larval age 0 to 7 validates the Desperate Larva hypothesis. The significant declines in percent settlement from larval age 7 to 21 days (p = 0.03) in the CCA treatment and from larval age 7 to 14 days (p = 2×10 − 3 ) in the Dictyota sp. treatment suggest the loss of competency starts within 7–14 days after it is acquired. Post-settlement There were no significant effects of larval age-at-settlement on the post-settlement survival of all broadcast spawning species tested (all p > 0.05). However, the post-settlement survival of Agaricia agaricites , the only brooding species, significantly differed among larval ages (ꭓ 2 = 34.5, df = 3, p = 2 × 10 − 7 ). For larvae settled 0 and 7 days after acquiring competency, post-settlement survival after 2 months remained high at 94 ± 2% and 91 ± 3%, respectively. For larvae whose settlement was delayed by 14 days, the estimated post-settlement survival decreased from 74 ± 6% to 64 ± 7% between months 2 and 3. Whereas for larvae whose settlement was delayed for 21 days, the post-settlement survival dropped to 50 ± 9% and 46 ± 9% on months 2 and 3, respectively (Fig. 3 ). The log rank test showed that the post-settlement survival of corals settled immediately after or 7 days after acquiring competency was not significantly different (p = 0.49) but was significantly higher than for corals whose settlement was delayed further (all p < 0.05); corals settled 21 days after acquiring competency also had significantly lower survival than corals whose settlement was delayed for 14 days (p = 0.04). Growth (change in surface area over time) significantly differed among species, with larval age and over time (all p < 2.2×10 − 16 , Fig. 4 ), with significant interactions between all variables (all p < 1×10 − 9 ). Time, larval age and species explained 76% of the variance in surface area of the corals, with coral individual only explaining less than 10% of the variance (Conditional R 2 = 0.858, Marginal R 2 = 0.76). The surface area of A. agaricites corals changed significantly over time and was significantly affected by the age of the larva at the time of settlement (both p < 2.2×10 − 16 , Fig. 4 ), with time and larval age significantly interacting (p < 2.2×10 − 16 ); larval age and time explained 66.5% of the variance in coral size, while differences between individual corals only explain 5.1% (Conditional R 2 = 0.716, Marginal R 2 = 0.665). This species exhibited significantly faster growth when settled within 7 days after acquiring competency than when their settlement was delayed for 14 or more days (all p < 0.001, Fig. 4 ), suggesting latent effects on post-settlement growth. The surface area of D. labyrinthiformis corals changed significantly over time (p < 2.2×10 − 16 ) and was significantly affected by delayed settlement (larval age) (p = 4.8×10 − 4 ), with the two significantly interacting (p = 2.9×10 − 9 ). However, larval age and time only explained 12.3% of the variance in size (Marginal R 2 = 0.123), and 41.7% of the variance being explained by differences among coral individuals (Conditional R 2 = 0.54), thus the differential growth patterns among larval ages are hard to interpret; regardless, there is no evidence of latent effect of delayed settlement on post-settlement growth (Fig. 4 ). The surface area of O. faveolata corals significantly changed over time (p < 2.2×10 − 16 ) and was significantly affected by larval age (p = 4.1×10 − 12 ), with the two predictors significantly interacting (p = 4.1×10 − 5 ) and explaining 36% the variance in coral surface area (Marginal R 2 = 0.362); coral individual explained 23% of the variance (Conditional R 2 = 0.591). Corals settled right after acquiring competency grew significantly faster than the ones which settlement was delayed for 7 or more days (all p < 0.006); the corals which settlement was delayed by 21 days grew significantly less than all the others (all p < 0.04). This suggests there are latent effects on delayed settlement on post-settlement growth of O. faveolata . Larval age at the time of settlement significantly influenced growth of P. clivosa (p = 0.003), with its surface area changing significantly over time (p < 2.2×10 − 16 ), and time and larval age significantly interacting (p = 0.002); however they only explained 17% of the variance in coral surface area (Marginal R 2 = 0.173); most of the variance, 47%, in surface area was explained by coral individual (Conditional R 2 = 0.64), thus, despite the differential growth patterns among larval ages, there is no clear evidence of latent effect of delayed settlement on post-settlement growth for this species (Fig. 4 ). Discussion Marked and unexpected differences in recruitment strategies were found among coral species. The brooding, weedy species Agaricia agaricites adhered to the Death Before Dishonor Hypothesis, i.e. their larvae refused to settle on a “degraded” substrate even when they became older. The larvae of this species are competent upon release, but lost competency quickly, within 7 days of release. Additionally, they suffered latent effects of delayed larval settlement on post-settlement survival and growth. This suggests A. agaricites evolved to disfavor dispersal and maximize local retention, and contrarily to what was expected, may not be good colonizers of degraded habitats, are likely just resistant once established. On the contrary, the reef-building, broadcast spawners Diploria labyrinthiformis , Pseudodiploria clivosa and Orbicella faveolata were found to follow the Desperate Larva Hypothesis, i.e. newly formed larvae settled significantly less in the absence of crustose coralline algae (CCA) and presence of Dictyota sp., however, as larvae became older, settlement in those same conditions increased, albeit sometimes later being offset by the onset of loss of competency. All spawners studied exhibited no latent effects of delayed settlement on post-settlement survival, and only O. faveolata displayed latent effects on post-settlement growth, suggesting their life history favors dispersal and that they may be able contribute to the replenishment of disturbed reefs, even if some will do so at the expense of hindered growth rates. Agaricia agaricites , commonly recognized as aweedy coral species, followed the Death Before Dishonor Hypothesis, displaying an early loss of competency, persistent reluctance to settle in the presence of an inhibitory cue, and suffered latent effects of delayed settlement on post-settlement survival and growth, suggesting it evolved to maximize local retention, and not long-distance dispersal. A similar pattern was observed in the brooder Agaricia humilis (Hartmann et al. 2013), hinting it may be widespread among brooders, which is surprising because this strategy is more common among food specialists (Santagata 2004 ), and Agaricia (and corals in general) are food generalists (Figueiredo et al. 2012 , Short et al. 2025 ). The larvae of brooders are competent to settle upon release, thus their local retention is commonly high (Figueiredo et al. 2013 ), limiting the proportion that disperses. Dispersal is further limited by early loss of competency as displayed by A. agaricites. The loss of competency is generally explained by the exhaustion of energy reserves in lecithotrophic larvae. However, Agaricia larvae have algal symbionts, and thus are not exclusively lecithotrophic. Still, the presence of algal symbionts does not necessarily translate into a prolonged competency period in stony corals (Nishikawa et al. 2003 ) and can in fact be a burden (Hartmann et al. 2019 ), particularly when the larvae are stressed (Chamberland et al. 2017 ). Mean water retention around reefs typically ranges between 0.5 and 6 days (Black et al. 1990 ; Black 1993 ; Cetina-Heredia and Connolly 2011 ), thus there is still scope for A. agaricites larvae to disperse (for at least up to 6 days) to farther reefs. However, A. agaricites exhibited latent effects of delayed settlement on post-settlement survival and growth. Together, these suggest this species does not prioritize energy allocation into dispersal, meaning that the benefits of local retention likely outweigh the advantages of dispersal. Weedy species thrive, not because of their high dispersal and connectivity, but rather because they can better withstand environmental stressors (Darling et al. 2012 ). Brooders produce larger, lipid-rich, ready to settle larvae, allowing them not to waste energy dispersing, and better sustain stressful conditions such as rising temperatures which accelerate the depletion of their energy reserves (Clarke and Fraser 2004 ; Harii et al. 2007 ; Figueiredo et al. 2014 ). Settling early guarantees they save more energy for post-settlement survival and growth, making them less likely to be preyed upon and overgrown by algae. It may be a winning strategy but is highly dependent on stock-recruitment relationships for population recovery, and thus making them vulnerable to catastrophic local disturbances. The broadcast spawning, reef-building corals D. labyrinthiformis and P. clivosa followed the Desperate Larva Hypothesis (Bishop et al. 2006 ), as they maintained competency longer, with settlement rates declining only after 7 or 14 days. Post-settlement survival and growth was not affected by larvae age at the time of settlement. This suggests they can disperse and potentially replenish more distant, disturbed reefs. As their larvae aged, they increasingly settled more in the “degraded reef” settlement cue treatment (i.e., on a tile with Dictyota sp .). This increase coincided with when they started losing competency (at 7–14 days old), apparent by the decrease in settlement in the CCA settlement cue treatment around that age. Adhering to the Desperate Larva Hypothesis agrees with findings by Elkin and Marshall ( 2007 ) that suggest this to be shared among generalist species (e.g., opportunistic heterotrophs) with lecithotrophic larvae. Other invertebrates’ non-feeding larvae, such as the sand dollar Dendraster excentricus , the free-living mollusk Alderia spp. and the bryozoan Watersipora subtorquata have also been documented to become less selective of settlement cues as they aged, which coincided with the depletion of energy reserves (Botello and Krug, 2006 ; Hodin et al. 2018 ; Marshall and Keough 2003 ).Extended competence over at least 7–14 days without compromising post-settlement survival nor growth suggests these species favor at least some dispersal and may potentially seed more distant disturbed reefs. Because D. labyrinthiformis and P. clivosa have a stress-tolerant life history strategy (Darling et al. 2012 ) with high fecundity but slow growth rates, there is likely a trade-off between their superior ability to colonize new habitats (generating conditions for other coral species to get established posteriorly) and their reduced competitiveness for space once others get established too. The broadcast spawning, reef-building O. faveolata also followed the Desperate Larva Hypothesis (Bishop et al. 2006 ), but delaying settlement hindered their post-settlement growth, suggesting dispersal may compromise its ability to compete for space. Despite displaying much lower settlement than the other species, the newly competent larvae of Orbicella faveolata still settled more on the “healthy reef” than on the “degraded reef” settlement cue treatment. It is uncertain whether this low settlement was caused by the lack of a suitable settlement cue and/or poor larval quality, however the first hypothesis seems more likely. The CCA that induce corals to settle are species-specific, and the one preferred by this species could have been absent from the mix provided in this study (Pechenik 1999 , Abdul Wahab et al. 2023 ). A significant increase in percent settlement when exposure to a tile was delayed by21 days after acquiring competency in both the “healthy reef” and the “ degraded reef ” treatments seems consistent with this hypothesis and is likely explained by desperation (in both cue treatments) rather than a developmental pattern of slowly advancing competency over 21 days. However, the low settlement together with the latent effects on post-settlement growth, can also indicate that larvae had low energy reserves, potentially caused by parental condition. Orbicella faveolata is a generalist species (strategy that can have some overlap with the competitive, weedy, and stress-tolerant life histories) or belongs to a sub-group of stress-tolerant species (Darling et al. 2012 ). Like generalists, O. faveolata does well in habitats where competition is limited by low levels of stress (evidenced by their increased growth rates after heat events in the Caribbean, in comparison to other species, Manzello et al. 2015 ; Muñiz-Castillo et al. 2019 ; Manzello et al. 2021 ), but is long-lived (Renegar et al. 2024 ), slow growing, thus akin of stress-tolerant species. Our results for O. faveolata suggest there may be a trade-off between its growth rates and dispersal potential to colonize new habitats. This study suggests that the paradigm that degraded reefs are unrecoverable (Roth et al. 2018 ) is at least partly flawed. For a disturbed reef to recover, three things need to happen: (1) larvae need to be produced and be transported to the degraded reef, (2) the larvae need to settle on degraded reefs, and (3) the settlers need to survive and grow to reproduce (realized recruitment). Even in highly disturbed systems, we often find that surviving corals are fecund (Renegar et al. 2024 ), and thus, in a well-connected reef system where coral density is high enough in at least a few key sites to guarantee fertilization success, disturbed reefs should continue to receive larvae dispersing from farther unaffected areas or produced by the surviving colonies (Gilmour et al. 2013 ). Increased nutrient run-off, which fuels algal growth, and overfishing of herbivores, which control it, have led reefs to become overgrown by algae. Because newly competent coral larvae are known to avoid settling in the presence of some algae, the current paradigm is that larvae would not settle on degraded reefs, leaving these with very low chances to recover naturally. However, this assertion is challenged by this study’s finding that older larvae of some reef-building coral species are less discriminatory and thus could potentially colonize degraded environments. The delayed settlement often has no latent effects on post-settlement survival and growth, which suggests some corals could potentially recruit to disturbed reefs. However, it is still possible that the competition with macroalgae will jeopardize the long-term survival and growth of the coral recruits in degraded habitats. A degraded reef will very likely offer worst conditions in a healthy reef for a coral recruit to survive and grow to sexual maturity, however if least some survive, recovery could happen, albeit at a slower pace. To fully understand if and to what extent degraded reefs can be replenished by the recruitment of older larvae, longer term post-settlement survival and growth in degraded vs. non-degraded environments need to be tested in situ . Existent literature suggests that reef recovery after disturbances such as mass bleaching is favored, among other things, by high coral recruitment (Gilmour et al. 2013 ; Graham et al. 2015 ). Recruitment to degraded reefs has indeed been observed (Roth et al. 2018 ). Even if only some reef-building, stress-tolerant corals could get established in degraded reefs, they might eventually produce settlement cues for the other coral species to settle there too (Da-Anoy et al. 2017 ). The coexistence of corals with different life histories has multiple benefits, such as enhanced tissue growth and survivorship, and macroalgal suppression (Clements and Hay 2019 ). From an active restoration perspective, if newly settled corals could survive and grow in degraded environments, seeding degraded reefs with older, desperate larvae may encourage settlement in these habitats and jumpstart their recovery. Still, with increasing CO 2 emissions, coral reefs have been experiencing more frequent and severe storms, giving little to no time for coral populations to replenish themselves before a new disturbance occurs (Buddemeier and Smith 1999 , Gouezo et al. 2019 ). Determining whether old larvae of a more comprehensive set of coral species might (re)populate distant reefs will change current predictions of realized recruitment and connectivity and contribute to the effective conservation and management of coral reefs. Declarations Competing Interests Statement: Authors have no financial interests directly or indirectly related to the work submitted for publication. Funding statement: We acknowledge the Florida Department of Environmental Protection and Nova Southeastern University’s President’s Faculty Research and Development Grant for funding support. Author Contribution N.G. collected the data. N.G. and J.F developed the research idea, analyzed the data and wrote the main manuscript. D.R. and M.M. assisted in the development of the methodology and edited the manuscript. J.F. secured the funding for the study. Acknowledgement We acknowledge the Florida Department of Environmental Protection and Nova Southeastern University’s President’s Faculty Research and Development Grant for funding support. We thank the UNCW REEF lab, the Florida Aquarium, and NOAA/CIMAS for the larval donations. We are also grateful to the Marine Larval Ecology and Recruitment Laboratory team for their support developing and executing this experiment, especially Rachel Ionata, Karagan Ross, Michael Hood, Morgan Short, Daisy Ponce, Ryan Chabotte, and Krista Laforest. Data Availability Data is provided within the manuscript or upon request to authors. References Abdul Wahab MA, Ferguson S, Snekkevik VK, McCutchan G, Jeong S, Severati A, Randall CJ, Negri AP, Diaz-Pulido G (2023) Hierarchical settlement behaviours of coral larvae to common coralline algae. 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Marine Ecology Progress Series 13: 273-283 Webster FJ, Babcock RC, Van Keulen M, Loneragan NR (2015) Macroalgae inhibits larval settlement and increases recruit mortality at Ningaloo Reef, Western Australia. PLoS ONE 10(4): e0124162 https://doi.org/10.1371/journal.pone.0124162 Wendt DE (1998) Effect of larval swimming duration on growth and reproduction of Bugula neritina (Bryozoa) under field conditions. The Biological Bulletin 195(2): 126-135 https://doi.org/10.2307/1542820 Appendix Appendix is not available with this version. Additional Declarations No competing interests reported. Cite Share Download PDF Status: Published Journal Publication published 09 Feb, 2026 Read the published version in Coral Reefs → Version 1 posted Editorial decision: Revision requested 07 Aug, 2025 Reviews received at journal 04 Aug, 2025 Reviews received at journal 26 Jul, 2025 Reviews received at journal 23 Jul, 2025 Reviewers agreed at journal 13 Jul, 2025 Reviewers agreed at journal 12 Jul, 2025 Reviewers agreed at journal 11 Jul, 2025 Reviewers agreed at journal 07 Jul, 2025 Reviewers invited by journal 07 Jul, 2025 Editor assigned by journal 04 Jul, 2025 Submission checks completed at journal 24 Jun, 2025 First submitted to journal 11 Jun, 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. 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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-6874996","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":481998468,"identity":"532b982e-91d2-474b-a0da-512c83bb96c8","order_by":0,"name":"Nepsis García","email":"","orcid":"","institution":"University of Michigan","correspondingAuthor":false,"prefix":"","firstName":"Nepsis","middleName":"","lastName":"García","suffix":""},{"id":481998469,"identity":"adcae669-4109-4d1b-b514-908332e4d1e5","order_by":1,"name":"Margaret Miller","email":"","orcid":"","institution":"SECORE International","correspondingAuthor":false,"prefix":"","firstName":"Margaret","middleName":"","lastName":"Miller","suffix":""},{"id":481998470,"identity":"7a4a7e15-ed70-481e-8a54-894208e3d544","order_by":2,"name":"D. Abigail Renegar","email":"","orcid":"","institution":"National Coral Reef Institute, Halmos College of Arts and Sciences, Nova Southeastern University","correspondingAuthor":false,"prefix":"","firstName":"D.","middleName":"Abigail","lastName":"Renegar","suffix":""},{"id":481998471,"identity":"dc33364a-b7f4-4cd5-969d-85f39417e22c","order_by":3,"name":"Joana Figueiredo","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA7ElEQVRIiWNgGAWjYBACxuaDDUBKAogTGB9ABQ3wa2lLhGthhinFr4WBLQHGSmCTIEoLcxtz82ueCot8fvbkY9UFFYflGdibt0ng08LYxthmzXNGwnJmz7O02zPOHDZs4DlWhl/L/MY2Y942CQODGzlmt3nb0hgbJHLMCNpizPtPwsAeqKUYqMW+Qf4NQS3Nj3kbgLYADWfmbbNJbJDgIWwL45xjEgYSZ54lS/OcsUlu40krtsCnxbCN/fGHNzV1BvztyQc/81RI2PazH954A6+WBgY2VGew4VMOAvLAqPlASNEoGAWjYBSMcAAANXFC1xZsRx4AAAAASUVORK5CYII=","orcid":"","institution":"National Coral Reef Institute, Halmos College of Arts and Sciences, Nova Southeastern University","correspondingAuthor":true,"prefix":"","firstName":"Joana","middleName":"","lastName":"Figueiredo","suffix":""}],"badges":[],"createdAt":"2025-06-11 21:53:14","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6874996/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6874996/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s00338-026-02825-y","type":"published","date":"2026-02-09T15:59:02+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":86432945,"identity":"63610d58-7a54-44b2-b3f4-bfb8613c2a31","added_by":"auto","created_at":"2025-07-10 14:58:48","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":196532,"visible":true,"origin":"","legend":"\u003cp\u003eMeasurement of a 2 month-old \u003cem\u003eDiploria labyrinthiformis\u003c/em\u003e on the cellSens (Olympus) software.\u003c/p\u003e","description":"","filename":"image1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6874996/v1/35dfadd4ca75461a951e7281.jpg"},{"id":86434268,"identity":"12564f9b-8587-487b-adec-9e271c2831b4","added_by":"auto","created_at":"2025-07-10 15:14:48","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":433278,"visible":true,"origin":"","legend":"\u003cp\u003eProportion of settlement of \u003cem\u003eAgaricia agaricites\u003c/em\u003e, \u003cem\u003eDiploria labyrinthiformis\u003c/em\u003e, \u003cem\u003eOrbicella faveolata\u003c/em\u003eand \u003cem\u003ePseudodiploria clivosa\u003c/em\u003e larvae with differing ages (0, 7, 14, and 21 days after acquiring competency) when exposed to the healthy – CCA (left, red) and degraded - \u003cem\u003eDictyota sp.\u003c/em\u003e (right, blue) treatments. The middle line of the boxplot represents the median, while the black circles represent the mean.\u003c/p\u003e","description":"","filename":"image2.png","url":"https://assets-eu.researchsquare.com/files/rs-6874996/v1/8f866540d39d255b5df6c30e.png"},{"id":86432080,"identity":"b42e0271-6d07-40ca-85cd-39200997c0fd","added_by":"auto","created_at":"2025-07-10 14:50:48","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":215110,"visible":true,"origin":"","legend":"\u003cp\u003eKaplan-Meier survival curves showing the effect of larval age-at-settlement on the proportion of post-settlement survival of \u003cem\u003eAgaricia agaricites; \u003c/em\u003en is the (initial) number of corals of each larval age-at-settlement monitored for post-settlement survival.\u003c/p\u003e","description":"","filename":"image3.png","url":"https://assets-eu.researchsquare.com/files/rs-6874996/v1/1e8ef06ff180deb0923f6b2f.png"},{"id":86432947,"identity":"0762f77a-ba45-4567-9a71-41e0b68f2f13","added_by":"auto","created_at":"2025-07-10 14:58:48","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":313540,"visible":true,"origin":"","legend":"\u003cp\u003eSurface area\u003cem\u003e \u003c/em\u003e(mm\u003csup\u003e2\u003c/sup\u003e) over time (months) of the corals of the four studied species settled at different larval ages, with the lines representing the best fit models and the different lines types representing the age larvae where settled (i.e. the number of days settlement was delayed): solid, dashed, dotted and dot-dashed lines respectively for larval ages 0, 7, 14 and 21 days.\u0026nbsp;\u0026nbsp;\u003c/p\u003e","description":"","filename":"image4.png","url":"https://assets-eu.researchsquare.com/files/rs-6874996/v1/930783ff0bf589a5f952c9e7.png"},{"id":102786673,"identity":"e59c9296-468c-42d8-9dca-07a53f811e37","added_by":"auto","created_at":"2026-02-16 16:14:32","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1699403,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6874996/v1/c60e20ea-b7db-4034-b6a1-e05e89808929.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Desperate Larva or Death Before Dishonor: can old coral larvae replenish degraded reefs?","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe capacity of coral populations to recover from disturbances depends greatly on their replenishment by coral larvae from nearby and distant populations (Lukoschek et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Holbrook et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Reef connectivity through coral larval dispersal allows the exchange of genes among locations and enhances reef resilience (Goodbody-Gringley et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). The onset and duration of the larval competency period determines dispersal time and, consequently, the extent of their distribution and connectivity (Figueiredo et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; King et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The longer the larvae remain competent, the farther they can disperse and replenish distant populations. Brooding species typically release readily competent larvae, and thus tend to settle on or closer to their natal reef; while the larvae of broadcast spawning species take longer to develop and only acquire competency 1\u0026ndash;6 days after spawning, reducing their local retention (Sammarco et al. 1989; Figueiredo et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Corals have lecithotrophic larvae (i.e., sustained by energy reserves placed in the egg), and because maternal provisioning is limited, so is their competency period (Richmond et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). However, the larvae of some coral species, mostly brooders but not exclusively (see Baird et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2009\u003c/span\u003e), contain symbiotic zooxanthellae which can supplement the coral larval energy via photosynthesis and thus might extend the larva\u0026rsquo;s pelagic larval duration (Chamberland et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eLarval settlement is a crucial process in a coral\u0026rsquo;s life history since it can directly influence its successful recruitment and long-term chances to survive, grow, and eventually sexually reproduce (Ritson-Williams et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Coral planulae have been shown to utilize specific cues to determine if a habitat is suitable to settle and metamorphose (Golbuu and Richmond \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Whereas bacterial films (Tebben et al. \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) and crustose coralline algae (CCA) have been shown to induce settlement in coral larvae (Heyward and Negri \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Negri and Heyward \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Hadfield and Paul \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Negri et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Ritson-Williams et al. \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), some macroalgae and cyanobacteria species are known settlement inhibitors (Kuffner et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Birrell et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Arnold et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Diaz-Pulido et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Doropoulos et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Webster et al. \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Ritson-Williams et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Changes to the ecological composition of benthic communities on coral reefs, such as the dominance shift from stony corals to fleshy macroalgae (Arias-Gonz\u0026aacute;lez et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), may jeopardize the settlement of coral larvae due to a lack of inducing settlement cues and/or the presence of inhibitory cues.\u003c/p\u003e\u003cp\u003eAs lecithotrophic larvae extend their larval period in the search for a suitable settlement site at the cost of depleting their energy reserves, they follow one of two pathways: decreased selectivity, known as the Desperate Larva Hypothesis (i.e. as larvae age, they become indiscriminate or \u0026ldquo;desperate\u0026rdquo; and settle on non-suitable substrates), or continued searching for a specific habitat or requirement to the point of depleting their energy reserves and perishing without settling and metamorphosing, known as the Death Before Dishonor Hypothesis (Bishop et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). The adoption of either of these pathways varies among species. The Desperate Larva Hypothesis was validated in some species in the phylum Bryozoa (Gribben et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), Arthropoda (Harvey and Colasurdo \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Miron et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Thiyagarajan et al. \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2002\u003c/span\u003e), Echinodermata (Rumrill \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e1989\u003c/span\u003e; Swanson et al. 2007), Annelida (Pechenik and Qian \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e1998\u003c/span\u003e), Mollusca (Pechenik \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e1984\u003c/span\u003e; Lima and Pechenik \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e1985\u003c/span\u003e; Carroll and Kempf \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Gibson \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Avila \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1998\u003c/span\u003e) and Phoronida (Herrmann \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1979\u003c/span\u003e; Herrmann \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e1995\u003c/span\u003e). Conversely, the Death Before Dishonor Hypothesis has only been validated in some phoronids, annelids and mollusks (Kempf \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e1981\u003c/span\u003e; Hadfield and Pennington \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Lambert and Todd \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Krug \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Santagata \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Botello and Krug \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). Following one or the other strategy appears closely linked with the strictness of their post-settlement habitat requirements (i.e., generalists or specialists, Elkin and Marshall \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2007\u003c/span\u003e), e.g., food and host specialists tend to follow the Death Before Dishonor Hypothesis. It remains unknown which pathway, desperate larva or death before dishonor, corals follow and if it varies depending on the species\u0026rsquo; life history strategy.\u003c/p\u003e\u003cp\u003eDelaying settlement can have latent effects on post-settlement survival (Miller \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e1993\u003c/span\u003e), growth (Pechenik et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2002\u003c/span\u003e), and reproduction success (e.g., lower fecundity, Miller \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e1993\u003c/span\u003e) of several invertebrates, including bryozoans (Wendt \u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Sams et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), ascidians (Marshall et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2003\u003c/span\u003e), sponges (Maldonado and Young \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e1999\u003c/span\u003e), and mollusks (Roberts and Lapworth \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). For benthic species, delaying settlement might reduce the energy reserves in the newly settled individuals, making them smaller and more vulnerable to fluctuations in food availability, predation and overgrowth (Qian and Pechenik \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). Little is known about the effect of delaying larval settlement on corals. The only study to date found \u003cem\u003eAcropora tenuis\u003c/em\u003e settled 2, 4 and 6 weeks after spawning did not exhibit latent effects on post-settlement survival and growth (Graham et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Studies focusing on other species are needed to better understand how delayed settlement might affect scleractinians.\u003c/p\u003e\u003cp\u003eThis study aimed to first address whether the selectivity of larvae from four Caribbean Scleractinia coral species for suitable settlement habitats diminished with age. Secondly, we examined whether the age at which larvae settled has latent effects on their post-settlement survival and growth. The results of this study suggest that the paradigm that disturbed reefs cannot recover through larval replenishment may be, at least partly, flawed.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003eStudy species and Coral collection\u003c/h2\u003e\u003cp\u003eThe speciesused in this experiment are common in Florida\u0026rsquo;s Coral Reef and throughout the Caribbean, are susceptible to stony coral tissue loss disease (SCTLD), and have different reproductive modes. \u003cem\u003eDiploria labyrinthiformis, Orbicella faveolata\u003c/em\u003e, and \u003cem\u003ePseudodiploria clivosa\u003c/em\u003e are reef-building, hermaphroditic, broadcast-spawning corals, while \u003cem\u003eAgaricia agaricites\u003c/em\u003e is a resilient, weedy, hermaphroditic, brooding species (Richmond and Hunter \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Carlon \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Knowlton \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). Twenty adult colonies of \u003cem\u003eAgaricia agaricites\u003c/em\u003e (12\u0026ndash;15 cm diameter) were collected from the Dania Erojacks, off the coast of Dania Beach (Florida, U.S.A.) via SCUBA a week before the new moon of May 2022, the expected larval release window (Van Moorsel \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e1983\u003c/span\u003e). The colonies were transported in coolers with seawater, wrapped in bubble wrap, and then housed in a 1500 L recirculating aquarium at Nova Southeastern University (NSU). The colonies were placed into individual 3L bowls within a recirculating tank, keeping the water level in the tank at \u0026frac34; of the bowl\u0026rsquo;s height to maintain water temperature. Filtered seawater was supplied to each bowl using a manifold system with rubber tubing at a rate of 1.3 L min\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e. Water then exited the bowl through the bowl\u0026rsquo;s handle into PVC cups filled at least halfway with water, fitted with plankton mesh (100 \u0026micro;m) at the bottom, that allowed constant water change while larvae were retained. Colonies released larvae from 4 days before to 6 days after the new moon, which were collected from the cups daily at the end of the expected larval release window (between 2300 and 0700 h, Van Moorsel \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e1983\u003c/span\u003e). The larvae from each broadcast spawning species were donated by different institutions and transported to NSU (Appendix); their competency was inferred by initiation of downward swimming.\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eSettlement\u003c/h3\u003e\n\u003cp\u003eApproximately 3000 larvae per species were equally and randomly split among six glass Mason jars filled with 500 mL of seawater and covered with parafilm with holes to reduce evaporation rates whilst allowing oxygenation. The jars were kept on an orbital shaker (Troemner\u0026trade; Talboys\u0026trade;) at 130\u0026ndash;180 rpm for the duration of the experiment to mimic ocean water movement and prevent settlement. Water changes were performed daily. We tested the effect of delayed settlement (larval age) on settlement success of coral larvae exposed to suitable (crustose coralline algae) and unsuitable (\u003cem\u003eDictyota sp\u003c/em\u003e.) settlement cues, respectively mimicking \u0026ldquo;healthy reef\u0026rdquo; and \u0026ldquo;degraded reef\u0026rdquo; habitats. To do so, for each of these two treatments, ten replicates of 10 or 15 larvae were randomly collected from all six jars at four times points: 0, 7, 14, and 21 days after becoming competent, hereafter referred to as \u0026ldquo;larval age\u0026rdquo; (Appendix). Specifically, each replicate consisted of 10 or 15 larvae in a 150 mL glass jar filled with 100 mL of artificial seawater with one labelled 3.2 cm\u003csup\u003e2\u003c/sup\u003e ceramic settlement tile (Ocean Wonders) with the rough side facing up. For the healthy reef treatment, the tiles used were preconditioned in a CCA-covered aquarium with adult corals for 4 weeks to allow microbial biofilm to grow. A mixture of crustose coralline algae, (that previously induced settlement for multiple species) was scraped off the aquarium\u0026rsquo;s walls, collected in a 50 mL polypropylene (PP) tube (Falcon\u0026reg;), ground into a fine dust using a mortar and pestle, and sprinkled on top of the preconditioned tile. For the \u0026ldquo;degraded reef\u0026rdquo; treatment, a 0.5\u0026times;1 cm piece of \u003cem\u003eDictyota spp. sp\u003c/em\u003e. from NSU tanks, a macroalga that inhibits coral settlement and reduces post-settlement survival (Olsen et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Paul et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) was placed on top of the tile. All jars were placed into a 2.4 L Polystyrene hinged food container (PACTIV) to reduce evaporation rates but allow for oxygenation, and jars were topped off daily with RO-DI water to maintain a salinity of 35 ppt. Larvae were left with the tiles for 48 h, after which they were assessed under a dissecting microscope (Olympus SZ61). Corals were considered settled and metamorphosed when attached to the substrate and polyp-shaped.\u003c/p\u003e\n\u003ch3\u003ePost-settlement survival and growth assessments\u003c/h3\u003e\n\u003cp\u003eOnce metamorphosis was complete, the location of each coral on the tiles was mapped to fate-track their survival and growth over time. Tiles were placed in a 42 L rectangular polypropylene (PP) tank with indirect sunlight (photosynthetically active radiation (PAR) of 4\u0026ndash;5 \u0026micro;mol photons m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003es\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), an air bubbler, and a Titanium heater (Finnex TH Deluxe). Corals were fed 5 mL of CoralAminos (Brightwell Aquatics) twice a week. The tank was topped off daily with DO water. (Sea)water was fully exchanged twice a week. Approximately 4 weeks after settlement, the newly settled corals that did not receive symbionts from a parent colony (i.e., vertical transmission), were inoculated with a cocktail of laboratory cultured \u003cem\u003eBreviolum minutum\u003c/em\u003e, \u003cem\u003eDurusdinium trenchii\u003c/em\u003e, \u003cem\u003eFugacium kawagutii\u003c/em\u003e and \u003cem\u003eSymbiodinium microadriaticum\u003c/em\u003e. Artificial overhead lights (Hydra Twenty-Six HD) set to 20 \u0026micro;mol photons m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003es\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e were added to the tank 2 weeks post-settlement.\u003c/p\u003e\u003cp\u003eAfter this period, the juveniles were gradually acclimated into an indoor recirculating aquarium for grow-out, where adult colonies were also held, which allowed them to continue acquiring a diverse Symbiodiniaceae assemblage. The system consisted of two connected 454 L raceways with a common 341 L sump equipped with a Titanium heater (Finnex TH Deluxe), drop-in chiller (Aqua Logic Cy-3), Protein Skimmer (ASM G-3), two UV Sterilizers (Pentair AES EU25-U), phosphate reactor (PhosBan Reactor 550), calcium reactor (GEO with Sicce Syncra Pump), 25 kg of live rock, and a 45.7\u0026times;30.5 cm perforated plastic basket full of bioballs. The system was set to mimic a truncated annual temperature cycle (23.3\u0026deg;C in February to 29.2\u0026deg;C in September) based on SECREMP data from Southeast Florida Reef Tract from 2007 to 2018 (Gilliam et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), excluding years when mass coral bleaching happened.\u003c/p\u003e\u003cp\u003eIn this system, the juveniles were positioned under LED lights (Radion\u0026reg; G5 XR30) set to a PAR 35 \u0026micro;mol photons m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003es\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, which were ramped up weekly by 10\u0026ndash;15 \u0026micro;mol photons m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003es\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, until 200 \u0026micro;mol photons m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003es\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e was reached. Once the juveniles were transferred into the recirculating aquarium, they were target fed 4 times a week with a 0.25 L slurry containing Reef-Roids (Polyp Lab \u0026reg;), Oyster Feast (Reef Nutrition\u0026reg;), CoralAmino (Brightwell Aquatics), and live rotifers that were enriched with a blend of microalgae (Rotigrow Plus\u0026reg;) 1 h before being prepared for feeding. The water flow was stopped for 1 h to allow the corals to eat. The tiles were hand-cleaned weekly due to algal growth.\u003c/p\u003e\u003cp\u003ePost-settlement survival and growth of juvenile corals were assessed monthly for three months by taking pictures of each tile with a digital camera (Olympus LC20) attached to a dissecting microscope (Olympus SZ61) and analyzed on cellSens imaging software (Olympus). When a juvenile coral was found with live tissue, it was marked as a living individual. Living individuals were measured by drawing a closed polygon around their live tissue (excluding tentacles; Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cdiv id=\"Sec6\" class=\"Section2\"\u003e\u003ch2\u003eData analysis\u003c/h2\u003e\u003cp\u003eTo test the effect of species, settlement cue treatment, and larval age on percent settlement, a Generalized Linear Model (GLM) with a binomial distribution (and then quasibinomial due to overdispersion) was performed in R using the package stats, with species (\u003cem\u003eAgaricia agaricites\u003c/em\u003e, \u003cem\u003eDiploria labyrinthiformis\u003c/em\u003e, \u003cem\u003eOrbicella faveolata\u003c/em\u003e, and \u003cem\u003ePseudodiploria clivosa\u003c/em\u003e), settlement cue [healthy reef (conditioned tiles with ground CCA) and degraded reef (non-conditioned tiles with \u003cem\u003eDictyota\u003c/em\u003e sp.], and larval age (0, 7, 14, and 21 days after becoming competent) as fixed predictors and percent settlement as response variable. Since settlement significantly differed between species, GLMs were conducted separately for each species (same variables as described above excluding species), followed by multiple comparison post-hoc tests using the library emmeans. A higher settlement of newly-competent larvae (0 days after becoming competent) on tiles with CCA vs. \u003cem\u003eDictyota\u003c/em\u003e confirms that the CCA is the preferred settlement cue. An increased settlement on tiles with \u003cem\u003eDictyota\u003c/em\u003e as larvae age validates the Desperate Larva Hypothesis, while a sustained, low settlement validates the Death Before Dishonor hypothesis. An increase in settlement on tiles with CCA as larvae age suggests increased percent competency over time and/or that the CCA species used was sub-optimal cue for that species (and thus larvae are just settling more due to desperation). A decrease in settlement on tiles with CCA and/or \u003cem\u003eDictyota\u003c/em\u003e as larvae age is indicative of the onset of loss of competency.\u003c/p\u003e\u003cp\u003eTo analyze the effect of larval age on post-settlement survival over time for each species, a Survival Analysis Mantel\u0026ndash;Haenszel log-rank test was performed using the R package survival, with larval age (levels: 0, 7, 14, and 21 days after becoming competent) as categorical predictor, and time as continuous predictor. For each coral, the time until the event of interest (mortality) occurred was recorded; if the coral remained alive at the end of the experiment, that time was recorded as right-censored. A lower post-settlement survival for corals whose settlement was delayed (i.e. older age) indicates latent effects on post-settlement survival.\u003c/p\u003e\u003cp\u003eTo test the effect of species and larval age on the post-settlement growth over time, a Generalized Linear Mixed-effects Model (GLMM) with a gamma log distribution was performed using R package glmmTMB, with species (\u003cem\u003eAgaricia agaricites\u003c/em\u003e, \u003cem\u003eDiploria labyrinthiformis\u003c/em\u003e, \u003cem\u003eOrbicella faveolata\u003c/em\u003e, and \u003cem\u003ePseudodiploria clivosa)\u003c/em\u003e and larval age (0, 7, 14, and 21 days after becoming competent) as fixed predictors, time as continuous predictor, coral individual as random predictor, and surface area as the response. Since species was found to be a significant predictor, the growth models were instead analyzed separately for each species, with all the other variables remaining the same. A lower post-settlement growth for corals which settlement was delayed (i.e. older age) indicates latent effects on post-settlement growth.\u003c/p\u003e\u003cp\u003eAll tests were run in R version 4.4.2 using RStudio version 2024.12.0.\u003c/p\u003e\u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e\u003ch2\u003eSettlement\u003c/h2\u003e\u003cp\u003ePercent settlement was significantly different among species (p\u0026thinsp;\u0026lt;\u0026thinsp;2.2\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;16\u003c/sup\u003e) and was significantly affected by both settlement cue (p\u0026thinsp;\u0026lt;\u0026thinsp;2.2\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;16\u003c/sup\u003e) and larval age (p\u0026thinsp;=\u0026thinsp;4.3\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e), with significant interactions between all variables (all p\u0026thinsp;\u0026le;\u0026thinsp;0.002). Together, coral species, settlement cue and larval age explained almost 72% of the variance in larval settlement (R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.717).\u003c/p\u003e\u003cp\u003eFor \u003cem\u003eAgaricia agaricites\u003c/em\u003e, settlement cue (p\u0026thinsp;\u0026lt;\u0026thinsp;2.2\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;16\u003c/sup\u003e) and larval age (p\u0026thinsp;=\u0026thinsp;3.6\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;4\u003c/sup\u003e) had a significant effect on percent settlement, explaining over 78% of the variance in settlement (R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.781), but they did not significantly interact (p\u0026thinsp;=\u0026thinsp;0.42). When newly-competent larvae (larval age 0), were exposed to the conditioned tile with CCA, a very high percentage of larvae settled (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SE: 91\u0026thinsp;\u0026plusmn;\u0026thinsp;4%), but settlement progressively declined with each week settlement was delayed (79\u0026thinsp;\u0026plusmn;\u0026thinsp;4%, 60\u0026thinsp;\u0026plusmn;\u0026thinsp;4%, 50\u0026thinsp;\u0026plusmn;\u0026thinsp;4% for larval ages 7, 14 and 21, respectively, Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), suggesting the onset of loss of competency. Regardless of the larva\u0026rsquo;s age, very few to no larvae settled on the tile with \u003cem\u003eDictyota sp.\u003c/em\u003e (3\u0026thinsp;\u0026plusmn;\u0026thinsp;2%, 5\u0026thinsp;\u0026plusmn;\u0026thinsp;5%, 0%, 0%, respectively for larval ages 0, 7, 14 and 21), validating the Death Before Dishonor hypothesis for this species.\u003c/p\u003e\u003cp\u003e\u003cem\u003eDiploria labyrinthiformis\u003c/em\u003e larvae settled significantly less on tile with \u003cem\u003eDictyota sp.\u003c/em\u003e than with CCA (p\u0026thinsp;\u0026lt;\u0026thinsp;2.2\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;16\u003c/sup\u003e). Larval age at the time of settlement did not significantly affect percent larval settlement (p\u0026thinsp;=\u0026thinsp;0.071), but it significantly interacted with the settlement cue (p\u0026thinsp;=\u0026thinsp;1.4 \u0026times; 10\u003csup\u003e\u0026minus;\u0026thinsp;4\u003c/sup\u003e), together explaining nearly 75% of the variance in settlement (R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.748). When exposed to CCA, an average of 52\u0026thinsp;\u0026plusmn;\u0026thinsp;3% of the newly competent (i.e. larval age 0) settled, versus only 5\u0026thinsp;\u0026plusmn;\u0026thinsp;2% with \u003cem\u003eDictyota sp..\u003c/em\u003e The average percentage settlement on the tile with CCA increased significantly when settlement was delayed by 7 days (73\u0026thinsp;\u0026plusmn;\u0026thinsp;3%, p\u0026thinsp;=\u0026thinsp;0.02) suggesting that not all larvae may have been competent at the beginning of the trial, i.e. by larval age 0 (as it had been assumed), with some larvae only acquiring competence afterwards and/or the settlement cue used was sub-optimal and the increased settlement with larval age was a result of desperation. Larval settlement remained high when delayed for 14 days (65\u0026thinsp;\u0026plusmn;\u0026thinsp;8%), but it declined significantly if delayed for one further week (45\u0026thinsp;\u0026plusmn;\u0026thinsp;7%, p\u0026thinsp;=\u0026thinsp;0.04, Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), suggesting the onset of loss of competency is sometime within this period. As larvae aged, the average percent settlement on the tiles with \u003cem\u003eDictyota sp.\u003c/em\u003e remained low, but its average increased slightly with larval age, being significantly higher for larvae with 21 days of age (4\u0026thinsp;\u0026plusmn;\u0026thinsp;2%, 11\u0026thinsp;\u0026plusmn;\u0026thinsp;2% and 19\u0026thinsp;\u0026plusmn;\u0026thinsp;2%, respectively, for larval ages 7, 14 and 21 days); suggesting this species follows the Desperate Larva hypothesis.\u003c/p\u003e\u003cp\u003eThe larvae of \u003cem\u003eOrbicella faveolata\u003c/em\u003e displayed the lowest settlement among the studied species, yet they still settled more on the tiles with CCA vs. the tiles with \u003cem\u003eDictyota sp.\u003c/em\u003e. While percent settlement significantly differed among settlement cue treatments (p\u0026thinsp;=\u0026thinsp;5.7\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;5\u003c/sup\u003e) and larval ages (p\u0026thinsp;=\u0026thinsp;1.4\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e), the two variables only explained 28% of the variance in settlement (R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.283) and they did not significantly interact (p\u0026thinsp;=\u0026thinsp;0.08). Larvae exposed to CCA initially displayed with low settlement, but settlement progressively increased with larval age (12\u0026thinsp;\u0026plusmn;\u0026thinsp;4%, 15\u0026thinsp;\u0026plusmn;\u0026thinsp;7%, 26\u0026thinsp;\u0026plusmn;\u0026thinsp;8%, and 30\u0026thinsp;\u0026plusmn;\u0026thinsp;2%, respectively for larval ages 0, 7, 14 and 21 days, Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), being significantly higher when settlement was delayed by 21 days (compared to larvae with 0 and 7 days, p\u0026thinsp;=\u0026thinsp;0.02 and p\u0026thinsp;=\u0026thinsp;0.04, respectively), suggesting the CCA cue used was sub-optimal, and thus that the increased settlement was a result of desperation. In the presence of \u003cem\u003eDictyota sp.\u003c/em\u003e, settlement was even lower (7\u0026thinsp;\u0026plusmn;\u0026thinsp;3% 5\u0026thinsp;\u0026plusmn;\u0026thinsp;2%, 1\u0026thinsp;\u0026plusmn;\u0026thinsp;1%, and 16\u0026thinsp;\u0026plusmn;\u0026thinsp;4%, respectively for larval ages 0, 7, 14 and 21 days), with the settlement of larvae which were only exposed to the cue 21 days after acquiring competency also being significantly higher (compared to larvae of 0 and 7 days, p\u0026thinsp;=\u0026thinsp;0.02 and p\u0026thinsp;=\u0026thinsp;0.04, respectively), once more suggesting this species follows the Desperate Larva hypothesis.\u003c/p\u003e\u003cp\u003eFor \u003cem\u003ePseudodiploria clivosa\u003c/em\u003e, larval age (p\u0026thinsp;=\u0026thinsp;3.7\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;6\u003c/sup\u003e) and settlement cue (p\u0026thinsp;=\u0026thinsp;4.7\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;13\u003c/sup\u003e), significantly affected percent larval settlement, and significantly interacted (p\u0026thinsp;=\u0026thinsp;5\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;5\u003c/sup\u003e); together they explained 65% of the variance in percent settlement (R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.65). The larval settlement in the presence of CCA was relatively higher (32\u0026thinsp;\u0026plusmn;\u0026thinsp;8%, 43\u0026thinsp;\u0026plusmn;\u0026thinsp;5% 29\u0026thinsp;\u0026plusmn;\u0026thinsp;5% and 23\u0026thinsp;\u0026plusmn;\u0026thinsp;5% for larvae exposed 0, 7, 15 and 21 days after acquisition of competency, respectively) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Conversely, newly competent larvae did not settle in the tiles with \u003cem\u003eDictyota sp.\u003c/em\u003e (0\u0026thinsp;\u0026plusmn;\u0026thinsp;0%); only larvae which settlement was delayed by 7 or more days settled on them (24\u0026thinsp;\u0026plusmn;\u0026thinsp;5%, 3\u0026thinsp;\u0026plusmn;\u0026thinsp;1%, 1\u0026thinsp;\u0026plusmn;\u0026thinsp;1%, respectively for larval ages 0, 7, 14 and 21). The drastic increase in settlement on tiles with \u003cem\u003eDictyota sp.\u003c/em\u003e from larval age 0 to 7 validates the Desperate Larva hypothesis. The significant declines in percent settlement from larval age 7 to 21 days (p\u0026thinsp;=\u0026thinsp;0.03) in the CCA treatment and from larval age 7 to 14 days (p\u0026thinsp;=\u0026thinsp;2\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e) in the \u003cem\u003eDictyota sp.\u003c/em\u003e treatment suggest the loss of competency starts within 7\u0026ndash;14 days after it is acquired.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003ePost-settlement\u003c/h3\u003e\n\u003cp\u003eThere were no significant effects of larval age-at-settlement on the post-settlement survival of all broadcast spawning species tested (all p\u0026thinsp;\u0026gt;\u0026thinsp;0.05). However, the post-settlement survival of \u003cem\u003eAgaricia agaricites\u003c/em\u003e, the only brooding species, significantly differed among larval ages (ꭓ\u003csup\u003e2\u003c/sup\u003e= 34.5, df\u0026thinsp;=\u0026thinsp;3, p\u0026thinsp;=\u0026thinsp;2 \u0026times; 10\u003csup\u003e\u0026minus;\u0026thinsp;7\u003c/sup\u003e). For larvae settled 0 and 7 days after acquiring competency, post-settlement survival after 2 months remained high at 94\u0026thinsp;\u0026plusmn;\u0026thinsp;2% and 91\u0026thinsp;\u0026plusmn;\u0026thinsp;3%, respectively. For larvae whose settlement was delayed by 14 days, the estimated post-settlement survival decreased from 74\u0026thinsp;\u0026plusmn;\u0026thinsp;6% to 64\u0026thinsp;\u0026plusmn;\u0026thinsp;7% between months 2 and 3. Whereas for larvae whose settlement was delayed for 21 days, the post-settlement survival dropped to 50\u0026thinsp;\u0026plusmn;\u0026thinsp;9% and 46\u0026thinsp;\u0026plusmn;\u0026thinsp;9% on months 2 and 3, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The log rank test showed that the post-settlement survival of corals settled immediately after or 7 days after acquiring competency was not significantly different (p\u0026thinsp;=\u0026thinsp;0.49) but was significantly higher than for corals whose settlement was delayed further (all p\u0026thinsp;\u0026lt;\u0026thinsp;0.05); corals settled 21 days after acquiring competency also had significantly lower survival than corals whose settlement was delayed for 14 days (p\u0026thinsp;=\u0026thinsp;0.04).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eGrowth (change in surface area over time) significantly differed among species, with larval age and over time (all p\u0026thinsp;\u0026lt;\u0026thinsp;2.2\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;16\u003c/sup\u003e, Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e), with significant interactions between all variables (all p\u0026thinsp;\u0026lt;\u0026thinsp;1\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;9\u003c/sup\u003e). Time, larval age and species explained 76% of the variance in surface area of the corals, with coral individual only explaining less than 10% of the variance (Conditional R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.858, Marginal R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.76).\u003c/p\u003e\u003cp\u003eThe surface area of \u003cem\u003eA. agaricites\u003c/em\u003e corals changed significantly over time and was significantly affected by the age of the larva at the time of settlement (both p\u0026thinsp;\u0026lt;\u0026thinsp;2.2\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;16\u003c/sup\u003e, Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e), with time and larval age significantly interacting (p\u0026thinsp;\u0026lt;\u0026thinsp;2.2\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;16\u003c/sup\u003e); larval age and time explained 66.5% of the variance in coral size, while differences between individual corals only explain 5.1% (Conditional R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.716, Marginal R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.665). This species exhibited significantly faster growth when settled within 7 days after acquiring competency than when their settlement was delayed for 14 or more days (all p\u0026thinsp;\u0026lt;\u0026thinsp;0.001, Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e), suggesting latent effects on post-settlement growth.\u003c/p\u003e\u003cp\u003eThe surface area of \u003cem\u003eD. labyrinthiformis\u003c/em\u003e corals changed significantly over time (p\u0026thinsp;\u0026lt;\u0026thinsp;2.2\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;16\u003c/sup\u003e) and was significantly affected by delayed settlement (larval age) (p\u0026thinsp;=\u0026thinsp;4.8\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;4\u003c/sup\u003e), with the two significantly interacting (p\u0026thinsp;=\u0026thinsp;2.9\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;9\u003c/sup\u003e). However, larval age and time only explained 12.3% of the variance in size (Marginal R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.123), and 41.7% of the variance being explained by differences among coral individuals (Conditional R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.54), thus the differential growth patterns among larval ages are hard to interpret; regardless, there is no evidence of latent effect of delayed settlement on post-settlement growth (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe surface area of \u003cem\u003eO. faveolata\u003c/em\u003e corals significantly changed over time (p\u0026thinsp;\u0026lt;\u0026thinsp;2.2\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;16\u003c/sup\u003e) and was significantly affected by larval age (p\u0026thinsp;=\u0026thinsp;4.1\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;12\u003c/sup\u003e), with the two predictors significantly interacting (p\u0026thinsp;=\u0026thinsp;4.1\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;5\u003c/sup\u003e) and explaining 36% the variance in coral surface area (Marginal R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.362); coral individual explained 23% of the variance (Conditional R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.591). Corals settled right after acquiring competency grew significantly faster than the ones which settlement was delayed for 7 or more days (all p\u0026thinsp;\u0026lt;\u0026thinsp;0.006); the corals which settlement was delayed by 21 days grew significantly less than all the others (all p\u0026thinsp;\u0026lt;\u0026thinsp;0.04). This suggests there are latent effects on delayed settlement on post-settlement growth of \u003cem\u003eO. faveolata\u003c/em\u003e.\u003c/p\u003e\u003cp\u003eLarval age at the time of settlement significantly influenced growth of \u003cem\u003eP. clivosa\u003c/em\u003e (p\u0026thinsp;=\u0026thinsp;0.003), with its surface area changing significantly over time (p\u0026thinsp;\u0026lt;\u0026thinsp;2.2\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;16\u003c/sup\u003e), and time and larval age significantly interacting (p\u0026thinsp;=\u0026thinsp;0.002); however they only explained 17% of the variance in coral surface area (Marginal R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.173); most of the variance, 47%, in surface area was explained by coral individual (Conditional R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.64), thus, despite the differential growth patterns among larval ages, there is no clear evidence of latent effect of delayed settlement on post-settlement growth for this species (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eMarked and unexpected differences in recruitment strategies were found among coral species. The brooding, weedy species \u003cem\u003eAgaricia agaricites\u003c/em\u003e adhered to the Death Before Dishonor Hypothesis, i.e. their larvae refused to settle on a \u0026ldquo;degraded\u0026rdquo; substrate even when they became older. The larvae of this species are competent upon release, but lost competency quickly, within 7 days of release. Additionally, they suffered latent effects of delayed larval settlement on post-settlement survival and growth. This suggests \u003cem\u003eA. agaricites\u003c/em\u003e evolved to disfavor dispersal and maximize local retention, and contrarily to what was expected, may not be good colonizers of degraded habitats, are likely just resistant once established. On the contrary, the reef-building, broadcast spawners \u003cem\u003eDiploria labyrinthiformis\u003c/em\u003e, \u003cem\u003ePseudodiploria clivosa\u003c/em\u003e and \u003cem\u003eOrbicella faveolata\u003c/em\u003e were found to follow the Desperate Larva Hypothesis, i.e. newly formed larvae settled significantly less in the absence of crustose coralline algae (CCA) and presence of \u003cem\u003eDictyota\u003c/em\u003e sp., however, as larvae became older, settlement in those same conditions increased, albeit sometimes later being offset by the onset of loss of competency. All spawners studied exhibited no latent effects of delayed settlement on post-settlement survival, and only \u003cem\u003eO. faveolata\u003c/em\u003e displayed latent effects on post-settlement growth, suggesting their life history favors dispersal and that they may be able contribute to the replenishment of disturbed reefs, even if some will do so at the expense of hindered growth rates.\u003c/p\u003e\u003cp\u003e\u003cem\u003eAgaricia agaricites\u003c/em\u003e, commonly recognized as aweedy coral species, followed the Death Before Dishonor Hypothesis, displaying an early loss of competency, persistent reluctance to settle in the presence of an inhibitory cue, and suffered latent effects of delayed settlement on post-settlement survival and growth, suggesting it evolved to maximize local retention, and not long-distance dispersal. A similar pattern was observed in the brooder \u003cem\u003eAgaricia humilis\u003c/em\u003e (Hartmann et al. 2013), hinting it may be widespread among brooders, which is surprising because this strategy is more common among food specialists (Santagata \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2004\u003c/span\u003e), and \u003cem\u003eAgaricia\u003c/em\u003e (and corals in general) are food generalists (Figueiredo et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2012\u003c/span\u003e, Short et al. \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The larvae of brooders are competent to settle upon release, thus their local retention is commonly high (Figueiredo et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), limiting the proportion that disperses. Dispersal is further limited by early loss of competency as displayed by \u003cem\u003eA. agaricites.\u003c/em\u003e\u003c/p\u003e\u003cp\u003eThe loss of competency is generally explained by the exhaustion of energy reserves in lecithotrophic larvae. However, \u003cem\u003eAgaricia\u003c/em\u003e larvae have algal symbionts, and thus are not exclusively lecithotrophic. Still, the presence of algal symbionts does not necessarily translate into a prolonged competency period in stony corals (Nishikawa et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2003\u003c/span\u003e) and can in fact be a burden (Hartmann et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), particularly when the larvae are stressed (Chamberland et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Mean water retention around reefs typically ranges between 0.5 and 6 days (Black et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Black \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Cetina-Heredia and Connolly \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), thus there is still scope for \u003cem\u003eA. agaricites\u003c/em\u003e larvae to disperse (for at least up to 6 days) to farther reefs. However, \u003cem\u003eA. agaricites\u003c/em\u003e exhibited latent effects of delayed settlement on post-settlement survival and growth. Together, these suggest this species does not prioritize energy allocation into dispersal, meaning that the benefits of local retention likely outweigh the advantages of dispersal. Weedy species thrive, not because of their high dispersal and connectivity, but rather because they can better withstand environmental stressors (Darling et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Brooders produce larger, lipid-rich, ready to settle larvae, allowing them not to waste energy dispersing, and better sustain stressful conditions such as rising temperatures which accelerate the depletion of their energy reserves (Clarke and Fraser \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Harii et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Figueiredo et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Settling early guarantees they save more energy for post-settlement survival and growth, making them less likely to be preyed upon and overgrown by algae. It may be a winning strategy but is highly dependent on stock-recruitment relationships for population recovery, and thus making them vulnerable to catastrophic local disturbances.\u003c/p\u003e\u003cp\u003eThe broadcast spawning, reef-building corals \u003cem\u003eD. labyrinthiformis\u003c/em\u003e and \u003cem\u003eP. clivosa\u003c/em\u003e followed the Desperate Larva Hypothesis (Bishop et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), as they maintained competency longer, with settlement rates declining only after 7 or 14 days. Post-settlement survival and growth was not affected by larvae age at the time of settlement. This suggests they can disperse and potentially replenish more distant, disturbed reefs. As their larvae aged, they increasingly settled more in the \u0026ldquo;degraded reef\u0026rdquo; settlement cue treatment (i.e., on a tile with \u003cem\u003eDictyota sp\u003c/em\u003e.). This increase coincided with when they started losing competency (at 7\u0026ndash;14 days old), apparent by the decrease in settlement in the CCA settlement cue treatment around that age. Adhering to the Desperate Larva Hypothesis agrees with findings by Elkin and Marshall (\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2007\u003c/span\u003e) that suggest this to be shared among generalist species (e.g., opportunistic heterotrophs) with lecithotrophic larvae. Other invertebrates\u0026rsquo; non-feeding larvae, such as the sand dollar \u003cem\u003eDendraster excentricus\u003c/em\u003e, the free-living mollusk \u003cem\u003eAlderia spp.\u003c/em\u003e and the bryozoan \u003cem\u003eWatersipora subtorquata\u003c/em\u003e have also been documented to become less selective of settlement cues as they aged, which coincided with the depletion of energy reserves (Botello and Krug, \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Hodin et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Marshall and Keough \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2003\u003c/span\u003e).Extended competence over at least 7\u0026ndash;14 days without compromising post-settlement survival nor growth suggests these species favor at least some dispersal and may potentially seed more distant disturbed reefs. Because \u003cem\u003eD. labyrinthiformis\u003c/em\u003e and \u003cem\u003eP. clivosa\u003c/em\u003e have a stress-tolerant life history strategy (Darling et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) with high fecundity but slow growth rates, there is likely a trade-off between their superior ability to colonize new habitats (generating conditions for other coral species to get established posteriorly) and their reduced competitiveness for space once others get established too.\u003c/p\u003e\u003cp\u003eThe broadcast spawning, reef-building \u003cem\u003eO. faveolata\u003c/em\u003e also followed the Desperate Larva Hypothesis (Bishop et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), but delaying settlement hindered their post-settlement growth, suggesting dispersal may compromise its ability to compete for space. Despite displaying much lower settlement than the other species, the newly competent larvae of \u003cem\u003eOrbicella faveolata\u003c/em\u003e still settled more on the \u0026ldquo;healthy reef\u0026rdquo; than on the \u0026ldquo;degraded reef\u0026rdquo; settlement cue treatment. It is uncertain whether this low settlement was caused by the lack of a suitable settlement cue and/or poor larval quality, however the first hypothesis seems more likely. The CCA that induce corals to settle are species-specific, and the one preferred by this species could have been absent from the mix provided in this study (Pechenik \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e1999\u003c/span\u003e, Abdul Wahab et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). A significant increase in percent settlement when exposure to a tile was delayed by21 days after acquiring competency in both the \u0026ldquo;healthy reef\u0026rdquo; and the \u003cem\u003e\u0026ldquo;\u003c/em\u003edegraded reef\u003cem\u003e\u0026rdquo;\u003c/em\u003e treatments seems consistent with this hypothesis and is likely explained by desperation (in both cue treatments) rather than a developmental pattern of slowly advancing competency over 21 days. However, the low settlement together with the latent effects on post-settlement growth, can also indicate that larvae had low energy reserves, potentially caused by parental condition. Orbicella faveolata is a generalist species (strategy that can have some overlap with the competitive, weedy, and stress-tolerant life histories) or belongs to a sub-group of stress-tolerant species (Darling et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Like generalists, \u003cem\u003eO. faveolata\u003c/em\u003e does well in habitats where competition is limited by low levels of stress (evidenced by their increased growth rates after heat events in the Caribbean, in comparison to other species, Manzello et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Mu\u0026ntilde;iz-Castillo et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Manzello et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), but is long-lived (Renegar et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), slow growing, thus akin of stress-tolerant species. Our results for \u003cem\u003eO. faveolata\u003c/em\u003e suggest there may be a trade-off between its growth rates and dispersal potential to colonize new habitats.\u003c/p\u003e\u003cp\u003eThis study suggests that the paradigm that degraded reefs are unrecoverable (Roth et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) is at least partly flawed. For a disturbed reef to recover, three things need to happen: (1) larvae need to be produced and be transported to the degraded reef, (2) the larvae need to settle on degraded reefs, and (3) the settlers need to survive and grow to reproduce (realized recruitment). Even in highly disturbed systems, we often find that surviving corals are fecund (Renegar et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), and thus, in a well-connected reef system where coral density is high enough in at least a few key sites to guarantee fertilization success, disturbed reefs should continue to receive larvae dispersing from farther unaffected areas or produced by the surviving colonies (Gilmour et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Increased nutrient run-off, which fuels algal growth, and overfishing of herbivores, which control it, have led reefs to become overgrown by algae. Because newly competent coral larvae are known to avoid settling in the presence of some algae, the current paradigm is that larvae would not settle on degraded reefs, leaving these with very low chances to recover naturally. However, this assertion is challenged by this study\u0026rsquo;s finding that older larvae of some reef-building coral species are less discriminatory and thus could potentially colonize degraded environments. The delayed settlement often has no latent effects on post-settlement survival and growth, which suggests some corals could potentially recruit to disturbed reefs. However, it is still possible that the competition with macroalgae will jeopardize the long-term survival and growth of the coral recruits in degraded habitats. A degraded reef will very likely offer worst conditions in a healthy reef for a coral recruit to survive and grow to sexual maturity, however if least some survive, recovery could happen, albeit at a slower pace.\u003c/p\u003e\u003cp\u003eTo fully understand if and to what extent degraded reefs can be replenished by the recruitment of older larvae, longer term post-settlement survival and growth in degraded vs. non-degraded environments need to be tested \u003cem\u003ein situ\u003c/em\u003e. Existent literature suggests that reef recovery after disturbances such as mass bleaching is favored, among other things, by high coral recruitment (Gilmour et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Graham et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Recruitment to degraded reefs has indeed been observed (Roth et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Even if only some reef-building, stress-tolerant corals could get established in degraded reefs, they might eventually produce settlement cues for the other coral species to settle there too (Da-Anoy et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). The coexistence of corals with different life histories has multiple benefits, such as enhanced tissue growth and survivorship, and macroalgal suppression (Clements and Hay \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). From an active restoration perspective, if newly settled corals could survive and grow in degraded environments, seeding degraded reefs with older, desperate larvae may encourage settlement in these habitats and jumpstart their recovery. Still, with increasing CO\u003csub\u003e2\u003c/sub\u003e emissions, coral reefs have been experiencing more frequent and severe storms, giving little to no time for coral populations to replenish themselves before a new disturbance occurs (Buddemeier and Smith \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1999\u003c/span\u003e, Gouezo et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Determining whether old larvae of a more comprehensive set of coral species might (re)populate distant reefs will change current predictions of realized recruitment and connectivity and contribute to the effective conservation and management of coral reefs.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eCompeting Interests Statement:\u003c/h2\u003e\u003cp\u003eAuthors have no financial interests directly or indirectly related to the work submitted for publication.\u003c/p\u003e\u003ch2\u003eFunding statement:\u003c/h2\u003e\u003cp\u003eWe acknowledge the Florida Department of Environmental Protection and Nova Southeastern University\u0026rsquo;s President\u0026rsquo;s Faculty Research and Development Grant for funding support.\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eN.G. collected the data. N.G. and J.F developed the research idea, analyzed the data and wrote the main manuscript. D.R. and M.M. assisted in the development of the methodology and edited the manuscript. J.F. secured the funding for the study.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eWe acknowledge the Florida Department of Environmental Protection and Nova Southeastern University\u0026rsquo;s President\u0026rsquo;s Faculty Research and Development Grant for funding support. We thank the UNCW REEF lab, the Florida Aquarium, and NOAA/CIMAS for the larval donations. We are also grateful to the Marine Larval Ecology and Recruitment Laboratory team for their support developing and executing this experiment, especially Rachel Ionata, Karagan Ross, Michael Hood, Morgan Short, Daisy Ponce, Ryan Chabotte, and Krista Laforest.\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eData is provided within the manuscript or upon request to authors.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAbdul Wahab MA, Ferguson S, Snekkevik VK, McCutchan G, Jeong S, Severati A, Randall CJ, Negri AP, Diaz-Pulido G (2023) Hierarchical settlement behaviours of coral larvae to common coralline algae. Scientific Reports 13(1): 5795. https://doi.org/10.1038/s41598-023-32676-4\u003c/li\u003e\n\u003cli\u003eArias-Gonz\u0026aacute;lez JE, Fung T, Seymour RM, Garza-P\u0026eacute;rez JR, Acosta-Gonz\u0026aacute;lez G, Bozec YM, Johnson CR (2017) A coral-algal phase shift in Mesoamerica not driven by changes in herbivorous fish abundance. PLoS ONE 12(4): e0174855 https://doi.org/10.1371/journal.pone.0174855\u003c/li\u003e\n\u003cli\u003eArnold SN, Steneck RS, Mumby PJ (2010) Running the gauntlet: Inhibitory effects of algal turfs on the processes of coral recruitment. Marine Ecology Progress Serie\u003cem\u003e,\u003c/em\u003e 414: 91\u0026ndash;105 https://doi.org/10.3354/meps08724\u003c/li\u003e\n\u003cli\u003eAvila C (1998) Competence and metamorphosis in the long-term planktotrophic larvae of the nudibranch mollusk \u003cem\u003eHermissenda crassicornis\u003c/em\u003e (Eschscholtz, 1831). Journal of Experimental Marine Biology and Ecology 231(1): 81-117 https://doi.org/10.1016/S0022-0981(98)00093-8\u003c/li\u003e\n\u003cli\u003eBaird AH, Guest JR, Willis BL (2009) Systematic and biogeographical patterns in the reproductive biology of scleractinian corals. Annual Review of Ecology, Evolution, and Systematics 40: 551\u0026ndash;571 https://doi.org/10.1146/annurev.ecolsys.110308.120220\u003c/li\u003e\n\u003cli\u003eBirrell CL, McCook LJ, Willis B L, Diaz-Pulido GA (2008) Effects of benthic algae on the replenishment of corals and the implications for the resilience of coral reefs. In: Gibson RN, Atkinson RJA, Gordon JDM (ed) Oceanography and Marine Biology: Annual Review, 46\u003csup\u003eth\u003c/sup\u003e edn. 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Scientific Reports 5(1): 1-11 https://doi.org/10.1038/srep10803 \u003c/li\u003e\n\u003cli\u003eThiyagarajan V, Harder T, Qian PY (2002) Relationship between cyprid energy reserves and metamorphosis in the barnacle \u003cem\u003eBalanus amphitrite\u003c/em\u003e Darwin (Cirripedia; Thoracica). Journal of Experimental Marine Biology and Ecology 280(1-2): 79-93 https://doi.org/10.1016/S0022-0981(02)00415-X\u003c/li\u003e\n\u003cli\u003eVan Moorsel GWNM (1983) Reproductive strategies in two closely related stony corals (\u003cem\u003eAgaricia\u003c/em\u003e, Scleractinia). Marine Ecology Progress Series 13: 273-283\u003c/li\u003e\n\u003cli\u003eWebster FJ, Babcock RC, Van Keulen M, Loneragan NR (2015) Macroalgae inhibits larval settlement and increases recruit mortality at Ningaloo Reef, Western Australia. PLoS ONE 10(4): e0124162 https://doi.org/10.1371/journal.pone.0124162\u003c/li\u003e\n\u003cli\u003eWendt DE (1998) Effect of larval swimming duration on growth and reproduction of \u003cem\u003eBugula neritina\u003c/em\u003e (Bryozoa) under field conditions. The Biological Bulletin 195(2): 126-135 https://doi.org/10.2307/1542820\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Appendix","content":"\u003cp\u003eAppendix is not available with this version.\u003c/p\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":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"coral-reefs","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"core","sideBox":"Learn more about [Coral Reefs](http://link.springer.com/journal/338)","snPcode":"338","submissionUrl":"https://submission.nature.com/new-submission/338/3","title":"Coral Reefs","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"delayed settlement, latent effects, post-settlement, recruitment","lastPublishedDoi":"10.21203/rs.3.rs-6874996/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6874996/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe recovery of coral populations depends largely on larval recruitment. Coral larvae settle in response to environmental cues that indicate habitat quality. In the absence of suitable cues, lecithotrophic larvae may delay settlement until they die (Death Before Dishonor Hypothesis) or become increasingly less discriminatory (Desperate Larva Hypothesis). To examine which of these hypothesized strategies corals follow, we used larvae from three broadcast spawning reef-building corals, \u003cem\u003eDiploria labyrinthiformis, Pseudodiploria clivosa\u003c/em\u003e, and \u003cem\u003eOrbicella faveolata\u003c/em\u003e, and one brooding, weedy coral, \u003cem\u003eAgaricia agaricites.\u003c/em\u003e Settlement behaviors of larvae of different ages, specifically 0, 7, 14, or 21 days after becoming competent, were examined in response to suitable (crustose coralline algae) and unsuitable (\u003cem\u003eDictyota sp\u003c/em\u003e.) settlement cues. Potential latent effects of delayed metamorphosis on post-settlement survival and growth were also assessed. As larvae got older, broadcast spawning species decreased selectivity (\u0026lsquo;Desperate Larva Hypothesis\u0026rsquo;), whereas the brooding species\u0026rsquo; larvae never did (\u0026lsquo;Death Before Dishonor Hypothesis\u0026rsquo;). Delayed settlement of the brooder \u003cem\u003eA. agaricites\u003c/em\u003e produced latent effects on post-settlement survival and growth, while \u003cem\u003eO. faveolata\u003c/em\u003e displayed latent effects of delayed settlement on post-settlement growth. The other broadcast spawners exhibited no latent effects of delayed settlement on post-settlement. This suggests \u0026ldquo;old\u0026rdquo; coral larvae from broadcast spawners have the potential to disperse widely to replenish degraded reefs. The brooding, weedy coral \u003cem\u003eA. agaricites\u003c/em\u003e seems to have evolved to disfavor dispersal and maximize local retention, evidenced by the fast acquisition of competency and latent effects of delayed settlement combined with an early loss of competency.\u003c/p\u003e","manuscriptTitle":"Desperate Larva or Death Before Dishonor: can old coral larvae replenish degraded reefs?","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-07-10 14:50:43","doi":"10.21203/rs.3.rs-6874996/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-08-07T18:02:32+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-08-04T13:56:22+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-07-26T15:51:33+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-07-23T23:51:57+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"52548956260649729801524621435743940822","date":"2025-07-14T03:04:01+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"331950466046106202447304266908013006930","date":"2025-07-13T03:29:32+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"334354307497985734042803594688813428578","date":"2025-07-11T14:56:49+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"294041194913892777029801217273391348299","date":"2025-07-07T20:52:02+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-07-07T12:47:13+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-07-05T01:09:24+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-06-24T14:26:55+00:00","index":"","fulltext":""},{"type":"submitted","content":"Coral Reefs","date":"2025-06-11T21:45:26+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"coral-reefs","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"core","sideBox":"Learn more about [Coral Reefs](http://link.springer.com/journal/338)","snPcode":"338","submissionUrl":"https://submission.nature.com/new-submission/338/3","title":"Coral Reefs","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"7548c88f-b959-4f98-8723-54754ccf27a7","owner":[],"postedDate":"July 10th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2026-02-16T16:13:50+00:00","versionOfRecord":{"articleIdentity":"rs-6874996","link":"https://doi.org/10.1007/s00338-026-02825-y","journal":{"identity":"coral-reefs","isVorOnly":false,"title":"Coral Reefs"},"publishedOn":"2026-02-09 15:59:02","publishedOnDateReadable":"February 9th, 2026"},"versionCreatedAt":"2025-07-10 14:50:43","video":"","vorDoi":"10.1007/s00338-026-02825-y","vorDoiUrl":"https://doi.org/10.1007/s00338-026-02825-y","workflowStages":[]},"version":"v1","identity":"rs-6874996","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6874996","identity":"rs-6874996","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}