Propagation of corals from the Arabian Gulf in a land-based nursery: effects of fragment size and light color

Abstract

Despite the Arabian Gulf hosting the most heat-tolerant corals in the world, coral restoration in the region remains constrained by the lack of species−specific husbandry guidelines. We fragmented donor colonies of seven common Gulf scleractinian species into five size classes and reared 1,540 ramets for 130 days in a land−based nursery under three controlled light-color treatments (blue, green, and white). Daily skeletal growth was quantified using buoyant−weight increments, and survivorship was monitored across treatments. Six species achieved 100% survival, whereas Porites harrisoni showed reduced survivorship across all treatments. Coral fragments consistently grew faster than whole colonies—up to eightfold in Dipsastraea pallida. Growth responses were strongly species−specific: Coscinaraea monile, Cyphastrea serailia, and Psammocora stellata grew fastest at medium fragment sizes, while Platygyra daedalea and D. pallida exhibited peak growth in smaller fragments. Light color significantly affected growth in only four species: blue light consistently enhanced growth in C. monile, C. serailia, and P. stellata, whereas P. daedalea generally grew fastest under green light. Responses to the same light spectrum varied markedly among species, underscoring the need for tailored nursery protocols. These findings provide the first species−level benchmarks for optimizing fragment size and lighting for ex situ propagation of Gulf corals, offering practical pathways to improve production efficiency and scalability of coral restoration in thermally extreme environments.

1 Introduction

Coral reefs are among the most important ecosystems on earth, hosting nearly a quarter of all known marine biodiversity while occupying less than 1% of the seafloor (Souter et al., 2021). They also provide critical ecosystem services such as fisheries, tourism, and coastal protection. Unfortunately, their high sensitivity to ocean warming makes coral reefs extremely vulnerable to climate change. Ocean warming and acidification, resource extraction, and frequent marine heatwaves are projected to cause increased coral bleaching and mortality (Hoegh-Guldberg et al., 2018). The Arabian Gulf has already lost 40.1% of the living coral cover between 1996 and 2019 (Burt et al., 2021), and recent assessments show continued declines, particularly in Acropora−dominated communities, with shifts toward heat−tolerant genera such as Porites and Dipsastraea (Hadj-Hammou et al., 2025). As bleaching events become more frequent and recovery windows shorten, active restoration is becoming increasingly important for maintaining reef−building coral populations.

Active reef restoration includes coral transplantation, substrate enhancement, larval seeding, and coral gardening, which involves rearing corals in in situ or ex situ nurseries before outplanting (Bradshaw, 1996). These methods are widely applied, but their performance varies across taxa and environments. Most empirical evidence comes from fast−growing branching corals, whereas massive and submassive taxa remain underrepresented and often respond unpredictably to propagation (Boström-Einarsson et al., 2020). As restoration expands into new regions, including the Arabian Gulf, the lack of species−specific nursery protocols limits its scalability and long−term success. Ex situ nurseries provide controlled conditions for asexual propagation, ranging from basic flow-through systems using natural seawater and sunlight to highly regulated enclosed systems with artificially prepared seawater and artificial illumination. Recent evidence indicates that nursery rearing can reduce thermal tolerance (Alderdice et al., 2024), which poses additional challenges in thermally extreme environments like the Arabian Gulf.

Fragmentation is a natural mode of asexual reproduction where broken pieces of coral reattach and grow into genetically identical colonies (Lirman et al., 2010; Foster et al., 2007; Vaughan, 2021). Coral gardening applies this process in situ and ex situ. In microfragmentation, corals are cut to sizes of only a few polyps to a few centimeters. This accelerates growth, relative to larger fragments or whole colonies, though optimal sizes differ among species (Vaughan, 2021). Microfragments can be outplanted adjacent to each other and fuse into new colonies much faster than if they grow from single polyps (Forsman et al., 2021; Koch, 2021). Studies across branching (Rinkevich, 2005) and massive corals (Forsman et al., 2006; Page et al., 2018) show clear size−dependent effects on survival and growth. Other field and nursery experiments consistently highlight trade−offs between fragment size, survival, and growth (Lirman et al., 2010; Sen and M. Yousif, 2016; Knapp et al., 2022), reinforcing the need for species−specific fragmentation protocols, especially for massive corals.

Light strongly influences coral through its effects on the photosynthetic symbionts (Osinga et al., 2011). Within certain limits, corals can photoacclimate through morphological, physiological, and biochemical adjustments (Osinga et al., 2011; Rocha, 2013). However, changes beyond this range impair performance: reduced irradiance limits photosystem activity and electron transport (Stirbet and Govindjee, 2011; Bessell-Browne et al., 2017), decreasing photosynthesis and promoting bleaching, while excessive irradiance reduces growth and photosynthetic rates (Leal et al., 2016; Titlyanov and Titlyanova, 2002; Osinga et al., 2011). Extreme light intensity can also cause photoinhibition and formation of reactive oxygen species (ROS), leading to expulsion of symbionts from the coral host (i.e., coral bleaching; Smith et al., 2005). Species−specific light requirements further complicate the issue and underscore the need to carefully define irradiance levels for ex situ nurseries.

Light spectra also affect the physiological performance of corals. Blue and violet wavelengths often enhance growth, tissue condition, and protein activity, while longer wavelengths (green, yellow, or red) tend to reduce performance (Cheng et al., 2022). For example, Stylophora pistillata fragments exposed to blue light showed higher survival, symbiont density, and health status than those under red light (Wijgerde et al., 2014). Similarly, Montipora damicornis and Montipora verrucosa grow faster under blue and white light than under green or red light, and isolated symbionts exhibit parallel spectral responses (Kinzie et al., 1984). Recent experiments further show that spectral composition modulates symbiont physiology and host gene expression (Izumi et al., 2023), while reviews highlight the role of host pigments in shaping internal light fields (Ferreira et al., 2023).

Modern artificial lighting technologies enhance coral growth: light-emitting diode (LED) and light-emitting photon (LEP) systems outperform conventional fluorescent and metal−halide bulbs, with LEP lighting supporting high photosynthetic performance under elevated irradiance (Wijgerde et al., 2012; Rocha et al., 2013). Nonetheless, species differ widely in their spectral preferences due to differences in morphology, depth distribution, and symbiont assemblages. These factors remain largely untested for Arabian Gulf corals, which have evolved under extreme temperature and irradiance regimes and may therefore respond differently to specific light spectra.

Literature on coral restoration is dominated by fast−growing branching species (e.g., Acroporids) and a small number of massive corals from the Pacific and Atlantic Oceans (Page et al., 2018; Koval et al., 2020; Rivas et al., 2021). In contrast, the issue remains understudied in the Arabian Gulf (Vaughan and Burt, 2016; Boström-Einarsson et al., 2020), and most regional restoration projects still focus on artificial reefs (Bartholomew et al., 2022; Giraldes et al., 2026), rather than propagation−based approaches (Giraldes et al., 2025). Summer seawater temperatures in the Arabian Gulf exceed 35 °C during an average of 23 days per year (NOAA Coral Reef Watch, 2026), which would cause massive coral bleaching and mortality in reef systems elsewhere. Accordingly, the region is considered a “natural laboratory” for understanding coral responses to future ocean−warming conditions (Bouwmeester et al., 2020). Despite their resilience, Gulf corals lack systematic evaluation under controlled nursery conditions, particularly regarding optimal fragment sizes and species−specific light requirements. Addressing this gap is essential for developing scalable, evidence−based restoration strategies tailored to the region.

The objective of this study was, therefore, to advance coral fragmentation and husbandry techniques for land-based coral nurseries in the Arabian Gulf and to increase the technology readiness level (TRL) of coral gardening in the region (Roch et al., 2023; Schmidt-Roach et al., 2024). We experimentally tested the effects of fragmentation size on the survival and growth of seven common coral species in this region when exposed to three different light color spectra. The following hypotheses were tested: H1: coral fragments will have similar survivorship (H1a) but faster growth rates (H1b), relative to conspecific whole colonies. H2: within the same coral species, growth rate will be inversely related to size class. H3: exposure to different light color spectra will affect growth rates within the same coral species (H3a), and the same light color spectra will cause distinct growth responses across coral species (H3b).

2 Materials and methods

2.1 Extraction of corals

The corals used in this study were extracted from subsea construction sites for new pipeline corridors connecting the QatarEnergy LNG offshore facilities in the North Field to the onshore facilities in Ras Laffan Industrial City. The use of the recovered corals for research purposes was approved by the Ministry of Municipality and Environment (MME) of Qatar as part of the mitigation and compensation strategy for the environmental impacts of the subsea construction. On 27–28 April 2021, a team of scientific scuba divers collected 214 coral colonies, representing seven Scleractinian species (Supplementary Figure S1), from donor sites adjacent to the northern limit of Ras Laffan Port, at depths between 6.7 and 9.2 m (Giraldes et al., 2026). The extracted corals reflected the natural species assemblages at the donor sites and included the following: Anomastraea irregularis von Marenzeller, 1901; Coscinaraea monile (Forskål, 1775); Cyphastrea serailia (Forskål, 1775); Dipsastraea pallida (Dana, 1846); Platygyra daedalea (Ellis & Solander, 1786); Porites harrisoni (Veron, 2000); and Psammocora stellata (Verrill, 1866). These species are among the most widespread in the EEZ of Qatar, as in many shallow reefs in the Gulf, and tend to be the dominant among their congenerics (Fanning et al., 2021; Bouwmeester et al., 2022). Accordingly, they are priority candidates for refining species−specific propagation protocols.

2.2 Coral husbandry

The experimental system comprised six glass aquaria (300 cm × 65 cm × 40 cm) with individual sumps, providing a total capacity of 910 L (780 L main tank + 130 L sump). Support trays were built using PVC pipe and plastic egg crate to hold the coral fragments, preventing direct contact with the tank bottom and optimizing water flow. Corals were periodically rotated within the tank, and trays were replaced during routine maintenance to prevent attachment and reduce fouling on the exposed skeletons. In terms of life support systems (LSS), each aquarium was equipped with a protein skimmer (MANTIS Tornado), two wave makers (MANTIS Tourban), three overhead LED light panels (ORPTEK Atlantik V4), a UV filter (EHEIM), and a return pump (EHEIM Compact ON 5000). Sumps contained filter socks, mechanical mesh media, and biological filtration (ceramics, coral skeletons, and bioballs). The system operated in a semiopen mode, with ~ 10% daily water replacement using natural seawater. Water temperature was continuously monitored using electrodes (Neptune Apex) connected to individual tank sumps and regulated by a heat exchange unit and two air-conditioning units in the room. Status checks for tanks, LSS, and the health status of the corals in each one of the tanks were done daily. Salinity was measured using a refractometer. Water quality parameters (pH, alkalinity, nitrite, nitrate, ammonia, phosphate, calcium, and magnesium) were regularly measured using commercial water analysis test kits (Tropic Marine, JBL, and AquaForest). Corals were fed twice per week using a combination of commercial coral foods (VITALIS Mixed Reef Food and LPS Coral Pellets: 2.06 g per tank) and microalgae (Nannochloropsis oceanica: 16.98 × 10⁶ cells mL⁻¹, 1 L per tank) in rotation.

2.3 Experimental setup

Each coral colony (unique genotype [genet]) received a unique three-digit tag and was photographed. An angle grinder was used to cut the donor colonies into pieces of 20–25 cm², which were subsequently fragmented to smaller sizes using a band saw (Gryphon C-40 CR Tall) and assigned to four size classes: (extra small [XS; 1.6–2.5 cm²], small [S; 3.8–6.1 cm²], medium [M; 8.3–11.2 cm²], and large [L; 15.1–22.0 cm²]; Supplementary Table S1). Each genet produced three to 46 genetically identical fragments (ramets), contributing to one or more size classes within that species (Supplementary Table S2). Polyp diameter varies substantially among the seven species: P. harrisoni (0.8–1 mm), P. stellata (1–1.5 mm), C. serailia (2–3 mm), A. irregularis (3–6 mm), P. daedalea (4–5 mm), C. monile (5–15 mm), and D. pallida (8–9 mm) (Riegl et al., 2012; Salimi et al., 2018). To avoid cutting fragments smaller than a single polyp, the four species with larger polyps (A. irregularis, C. monile, D. pallida, and P. daedalea) were not included in the XS size class. Whole colonies were also included in the experimental design to compare growth with fragments of the same species. For Psammocora stellata, all eight donor colonies required fragmentation; therefore, the two largest fragments were retained as whole−colony proxies. Due to their limited number and because they were not intact colonies, these proxies were excluded from statistical analyses but are reported in the Supplementary Materials.

A total of 148 genets and 1,540 ramets were included in the experiment. Each tank received 98–101 genets and 251–262 ramets (Supplementary Table S2). All ramets were labeled with unique five−digit codes (genet + ramet) using electrical cable tags fixed with cyanoacrylate glue, and those that were unstable were mounted onto ceramic tiles using marine epoxy or cyanoacrylate glue (Supplementary Table S3). Scaled photographs of each ramet were taken at the start and end of the experiment. Initial perimeter and surface area were measured using TagLab image analysis software to define the size ranges per species and size class (Supplementary Table S1). Ramets of the same species and size class were evenly distributed across tanks in a haphazard manner.

Corals were fragmented over 20 days in May 2021, followed by a 5-week healing period before the start of the experiment in July 2021. During the healing/acclimation period, lights were programmed to achieve a balanced spectral output, with 70% intensity in all channels. Apart from this, coral husbandry conditions (i.e., temperature, water flow, photoperiod, and feeding regime) matched the subsequent experimental phase. Whenever commensal or parasitic organisms or signs of disease were detected during these 5 weeks, affected ramets were dipped in antiseptic Lugol solution to prevent spread (Klinges et al., 2023). Overall, only 6.6% of the ramets were treated. The experiment lasted for a total of 130 days, from July to October 2021.

2.4 Light color treatments

During the experimental period, the ORPTEK lights were programmed to achieve three distinct light color treatments (green, blue, and white):

• Green: 80% channel 1 (451 and 517 nm) + 20% other channels.

• Blue: 80% channel 2 (452 nm) + 20% other channels.

• White: 80% channel 4 (414 nm) + 20% other channels.

Each light treatment was assigned to two replicate tanks. Photosynthetically active radiation (PAR) was measured in each tank at coral height using a Neptune Systems PAR Monitoring Kit. Lights turned on at 06:00, reached target intensity by 07:00 (sunrise simulation), maintained constant intensity until 17:00, and dimmed until 18:00 (sunset simulation). The constant intensity from 07:00 to 17:00 was intentionally maintained to ensure that spectral composition, not irradiance, was the sole experimental variable. Natural midday peak irradiances typical of Gulf reefs were not reproduced to avoid photoinhibition stress. The chosen intensity range provided standardized and physiologically safe conditions under which to test the effects of light color.

2.5 Determination of buoyant weight as a proxy for skeletal growth

Changes in buoyant weight were measured as a proxy for coral skeletal growth (Supplementary Figure 2). According to Jokiel et al. (1978), the buoyant weight technique minimizes damage to the corals because they do not have to be removed from the water. Furthermore, buoyant weight is unaffected by the amount of water or air retained in the skeleton, the amount of tissue or mucus, and the biomass of commensal or parasitic organisms on and within the skeleton. All pumps and wave-makers were turned off during measurements to eliminate turbulence, but for periods no longer than 2 h. Colonies and fragments were weighed at the start and end of the experiment. For fragments mounted on tiles, the initial buoyant weight of the tiles was measured separately and deducted from all subsequent measurements. Equations 1, 2 denote the calculations for relative buoyant weight increment (RBWI) and daily skeletal growth rate (DSGR), respectively.

2.6 Data analyses

Survival of P. harrisoni was assessed using Kaplan–Meier survival curves and log-rank tests. For growth analyses, ramet−level DSGR values were averaged per species × size class × tank and analyzed independently for each species using two-way analysis of variance (ANOVA), with light color and size class as orthogonal fixed factors and tanks as replicates (n = 2). Assumptions of normality and homogeneity of variances were tested using Kolmogorov–Smirnov and Cochran’s tests, respectively. The datasets showed moderate departures from the assumptions, which could not be resolved by statistical transformations. However, ANOVA was retained due to the balanced design and the importance of interpreting the interaction terms (Quinn and Keough, 2002). Student–Newman–Keuls (SNK) tests were used for multiple comparisons on significant main effects or interactions. Differences between means were considered statistically significant where p < 0.05. All statistics and plots were generated using SigmaPlot v11.0 (Systat Software Inc.).

3 Results

All physical and chemical seawater parameters were maintained within natural ranges for Arabian Gulf coral reefs, with an average temperature of 26.6 °C ± 0.06 °C and salinity of 44.4 °C ± 0.08 °C. Other chemical properties of seawater are reported in Supplementary Table S5.

3.1 Effects on survival

Six of the seven species (A. irregularis, D. pallida, C. monile, C. serailia, P. stellata, and P. daedalea) recorded 100% survival during the 130-day experiment. In contrast, P. harrisoni showed substantially reduced survival (55%). No significant difference was observed in P. harrisoni survival rate among size class or light color treatments (Figure 1; Kaplan–Meier analysis). Nevertheless, whole colonies survived at lower rates (31%) than fragments (62%).

Figure 1

3.2 Effect of fragment size on growth

Size class significantly affected skeletal growth rates in A. irregularis, D. pallida, C. monile, and C. serailia (Table 1). In all species where analyses were possible, whole colonies consistently registered the smallest growth rates among all size classes (Figure 2; Supplementary Table S6), although these differences were not always significant (Supplementary Table S7). Peak growth rates for D. pallida (up to 0.14% day⁻¹) and P. daedalea (up to 0.46% day⁻¹) were recorded in small fragments (6.08 and 5.30 cm² surface area, respectively; Supplementary Tables 1, S6). In contrast, peak growth in C. monile (up to 0.11% day⁻¹), C. serailia (up to 0.21% day⁻¹), and P. stellata (up to 0.13% day⁻¹) was consistently recorded in medium fragments (10.81, 9.22, and 9.17 cm² surface area, respectively; Supplementary Table S1, S6). P. harrisoni and P. stellata were excluded from certain size−class analyses due to low survivorship and insufficient whole−colony representation, respectively.

Figure 2

3.3 Effect of light color on growth

PAR measured at coral height did not differ significantly among light color treatments (Supplementary Table S4), with mean values of 124 µmol ± 1.2 µmol m⁻² s⁻¹ (green), 141 µmol ± 17.7 µmol m⁻² s⁻¹ (blue), and 114 µmol ± 1.1 µmol m⁻² s⁻¹ (white). These similar irradiance levels corroborate that differences in skeletal growth reflect spectral effects rather than light intensity. Light color significantly affected C. monile, C. serailia, and P. stellata, and produced a significant size class × light color interaction in P. daedalea (Table 1; Supplementary Table S8). In C. monile, C. serailia, and P. stellata, blue light consistently produced the highest growth rates across all size classes (Figure 3; Supplementary Table S6). P. daedalea grew fastest under green light in all size classes, except whole colonies (Figure 4). However, light color only had significant effects on the growth of P. daedalea in the small and medium size classes (Supplementary Table S9), which explains the significant interaction term (Table 1). White light produced the lowest growth in P. stellata, C. serailia, and P. daedalea (Figure 3), while C. monile registered the smallest growth under green light. Growth of P. harrisoni was not significantly affected by light color or size class (Table 1), which is consistent with its reduced survivorship across all treatments.

Figure 3

Figure 4

4 Discussion

4.1 Effects on survival

Hypothesis 1a, stating that coral fragments would have similar survival rates relative to conspecific whole colonies, was supported for all species except Porites harrisoni. Whole colonies and fragments of Anomastraea irregularis, Coscinaraea monile, Cyphastrea serailia, Psammocora stellata, Platygyra daedalea, and Dipsastraea pallida had 100% survival during the 130-day experiment. This high survivorship demonstrates that the nursery conditions and husbandry protocols were effective for these six species across all treatments. P. harrisoni was the only species that sustained mortality, and this occurred uniformly across light color and size class treatments (Figure 1), indicating that its poor performance reflects species−specific sensitivities rather than treatment effects. A previous study on P. lutea reported 100% survivorship after 12 weeks under ex situ conditions (Ng et al., 2012), but PAR levels in that study were more than double those used in our experiment (375 and 114–141 µmol m⁻² s⁻¹, respectively). This difference in irradiance likely explains the contrasting outcomes and reinforces that P. harrisoni, being a shallow, high−irradiance species (Bouwmeester et al., 2022), may experience light limitation in ex situ systems operated under conservative PAR levels. The columnar morphology of P. harrisoni compared to the massive form of P. lutea (Terraneo et al., 2021) may further contribute to lower survivorship of whole colonies due to self−shading. This could also have contributed to increased mortality relative to fragments (Figure 1). In fact, given their small size and simple shape, fragments had their entire surface area directly exposed to light, while whole colonies were partially shaded due to their more complex growth forms. Additionally, lower genetic diversity among whole colonies (two to three genets per tank) relative to fragments (six to 13 genets per tank) may also have amplified their sensitivity to light limitation or other stressors.

4.2 Effects of fragment size on growth

Hypothesis 1b, predicting that fragments would grow faster than conspecific whole colonies, was corroborated for all species where tests were possible. Whole colonies consistently exhibited the slowest skeletal growth, consistent with patterns previously documented for both branching and massive corals (Forsman et al., 2015; Koch, 2021; Vaughan, 2021). Small fragments of P. daedalea and D. pallida grew six and eight times faster, respectively, than whole colonies, while medium fragments of C. monile and C. serailia grew six and five times faster than whole colonies (Supplementary Table S1). Growth rate of medium-sized fragments of A. irregularis and P. stellata also consistently exceeded whole colonies (1.6 × and 1.4 × faster, respectively), although most of these differences were not statistically significant (Table 1; Supplementary Table S7). Accordingly, our study has conclusively shown that, under the same conditions, whole colonies generally have slower growth than conspecific fragments of optimal size. This difference in growth rates was observed both in terms of skeletal growth (i.e., buoyant weight increments; Figures 2, 4; Supplementary Table S6), as well as in the surface area of live tissue (Supplementary Figure 1). Nevertheless, measuring the surface area of the fragments from the photos taken at the end of the experiment was not possible due to the new tissue spreading to the underside of the fragments. The enhanced growth of fragments is consistent with the “healing response”, whereby fragmentation stimulates accelerated tissue and skeletal deposition to recover the cut surface (Vaughan, 2021). Several potential mechanisms have been proposed to explain this response (i.e., physiological, hormonal, genetic, or evolutionary), but the issue remains unresolved and requires further investigation (Koch, 2021).

Hypothesis 2, predicting an inverse relationship between fragment size and growth rate within species, was only partially supported. Small fragments grew fastest in P. daedalea and D. pallida, whereas C. monile, C. serailia, and P. stellata grew fastest when cut into medium-size fragments (Figures 2, 4). A. irregularis also showed faster growth rates as medium or large fragments, although differences were not significant. Variation among species likely reflects differences in polyp size, morphology, and energy allocation strategies, which are known to modulate the balance between healing and skeletal accretion following fragmentation (Forsman et al., 2006; Page et al., 2018; Koch, 2021). For P. harrisoni, reduced survivorship limited interpretation, but medium−sized fragments exhibited the fastest growth under blue and green light (Supplementary Table S6). Comparison with earlier studies on Porites spp (Forsman et al., 2006; Ng et al., 2012). suggests that optimal size thresholds vary not only across coral taxa but also across environmental contexts, reinforcing the need for species− and region−specific guidelines.

The predicted inverse size–growth relationship (H2) was grounded in previous work showing that small fragments, down to single polyps, often grow fastest (Wijgerde et al., 2012; Forsman et al., 2015; Sen and M. Yousif, 2016; Koch, 2021). This effect has been attributed to the “healing−driven” growth theory (Koch, 2021), according to which the greater the proportion of polyps damaged during fragmentation, the faster the growth of the fragment in order to accelerate recovery. Smaller fragments have a proportionally larger cut surface area relative to their total tissue volume and are therefore often predicted to exhibit faster growth rates during recovery, a pattern corroborated in previous studies (Forsman et al., 2015; Page et al., 2018). The mixed outcomes observed in this study, however, imply that the healing−driven growth advantage might only apply within species−specific size ranges. In some of the species with larger polyps (P. daedalea and D. pallida), the optimal threshold for accelerated growth was in the size range of 5–6 cm², which was assigned as “small” in our experiment. The reasons why the response predicted in H2 did not extend to all species remain unclear; however, some of the observed variability may reflect methodological artifacts. Notably, P. daedalea and D. pallida had the lowest proportion of small fragments mounted on tiles (7% and 3%, respectively; Supplementary Table S3), and these species also exhibited the highest growth rates within that size class (0.30 and 0.12% day⁻¹, respectively; Supplementary Table S6). In contrast, other species had a much higher proportion of small fragments mounted on tiles (27%–98%; Supplementary Table S3) and correspondingly lower growth rates (0.04%–0.10% day⁻¹; Supplementary Table S6). Fragment mounting on tiles may, therefore, have increased competition with encrusting organisms, potentially suppressing growth (Craggs et al., 2019; Neil et al., 2024; Ramsby et al., 2024). Resolving this potential artifact will require further investigation of fragment healing and growth across species with different morphologies under varying substrate availability, types, and orientations.

4.3 Effects of light color on growth

Hypothesis H3a, predicting that light color would affect growth within species, was supported for C. monile, C. serailia, P. daedalea, and P. stellata. Skeletal growth in these four species was significantly affected by light color, although unambiguous alternatives to the null hypothesis were only identified for P. stellata (Figure 3C, blue = green > white) and for the smaller size class of P. daedalea (Supplementary Table S9; green > blue > white). Blue light maximized growth in C. monile, C. serailia, and P. stellata (Figure 3), consistent with the more efficient water penetration of shorter wavelengths, particularly for deeper−reef taxa, such as P. stellata (Bouwmeester et al., 2022). In contrast, C. serailia is mainly associated with shallow reefs, where the full visible spectrum is available. Previous studies on other Cyphastrea species have reported higher calcification and skeletal extension under increased PAR levels (Howells et al., 2018). While PAR did not differ significantly among treatments in the present study, blue light was associated with slightly higher mean PAR (141 µmol m⁻² s⁻¹) than white light (114 µmol m⁻² s⁻¹). This suggests that the growth pattern observed for C. serailia may have been marginally influenced by light intensity; however, given the intentionally narrow PAR range across treatments, spectral composition was likely the dominant driver of the observed response. In contrast, A. irregularis was found to maximize growth under white light, while fragments of P. daedalea had maximum growth under green light. Although reduced survivorship obscured growth patterns for P. harrisoni, the maximum growth rates for fragments were also consistently recorded under green light (Supplementary Table S6). These trends, although not statistically significant for all species and Size-Classes, are consistent and cannot be related to the PAR levels, so can only be driven by light color.

P. daedalea is the most widespread coral species in the EEZ of Qatar and the dominant species among its congenerics in the majority of reefs (Bouwmeester et al., 2022). Accordingly, the adaptation to different light spectra could be attributed to the variable niches in which this species occurs (Burt et al., 2019). The pattern observed for blue and green light in P. daeadalea appears to follow the predicted inverse relationship between size and growth (Figure 4; Supplementary Table S6), although this only translated into an unambiguous alternative to the null hypothesis under green light (Supplementary Table S9, S > M > L > WC). This pattern is broken under white light, which explains the significant Size Class × Light Color interaction. Overall, these trends emphasize that growth responses to light spectra cannot be generalized across taxa and that optimal nursery lighting must be species−specific.

Hypothesis 3b, predicting that the same light spectra would produce different growth responses across species, was supported. In fact, the three light color treatments generated variable and even opposing growth patterns for different coral species. Light color significantly affected skeletal growth for C. monile, C. serailia, P. daedalea, and P. stellata, but not for A. irregularis, D. pallida, and P. harrisoni (Table 1). Although PAR levels did not differ significantly among light color treatments (Supplementary Table S4), the fact that three out of seven species (C. monile, C. serailia, and P. stellata) had their fastest growth rates under blue light (Figure 3) cannot be completely disassociated from the fact that blue light had the highest PAR level (141 µmol m⁻² s⁻¹). The apparent lack of response of D. pallida to light color was unexpected, given that this species is mainly associated with deeper reefs (10–25 m, Bouwmeester et al., 2022), where blue light prevails. This may reflect physiological flexibility or acclimation to the experimental setting.

As previously mentioned, green light generated the maximum growth rates for P. daedalea and P. harrisoni, but also the minimum growth rates for A. irregularis and C. monile. White light tended to generate the smallest growth increments in most species, but maximum growth for A. irregularis was also recorded under white light. Discriminating the physiological mechanisms underpinning these variations in growth responses of different coral species to the same light color is beyond the scope of the present study. Nevertheless, it seems safe to assume that the light requirements of each coral will be driven by the photosynthetic efficiency of its endosymbionts, which will in turn be determined by their abundance and community structure (Titlyanov and Titlyanova, 2002; Izumi et al., 2023). Symbiont composition will be influenced by the depth at which the corals occur, since light attenuation will play a critical role in determining the available light spectrum. Such physiological and ecological variations underscore the challenge of defining universal light regimes for ex situ nurseries.

Finally, we acknowledge that genotype is a significant driver of coral survival and growth, which was not explicitly resolved in our experimental design. Accordingly, we cannot exclude the possibility that intraspecific variability could also have influenced our results, and this is more likely to have occurred for the species and size classes represented by the smallest number of genotypes (Supplementary Table S2).

5 Conclusion

Most coastal and shallow coral reefs in the Arabian Gulf are collapsed or heavily degraded, sustaining only a residual coral community paired with increasing algal cover (Burt et al., 2021). The primary cause of these losses is recurrent coral bleaching due to the increasing frequency and duration of marine heat waves (Burt et al., 2019; Paparella et al., 2019). The few coral species remaining in these reefs are the best adapted and most resilient to heat stress and disease outbreaks (Aeby et al., 2024) and, therefore, should be among the primary candidates for coral restoration efforts. Research on these heat-resilient corals is of strategic importance to understand and maximize their adaptation potential to future climate change.

Our results demonstrate that fragmentation significantly accelerates growth in several common Gulf coral species, as fragments consistently exhibit higher growth rates than whole colonies under the same nursery conditions. Optimal fragment size was strongly species−specific, highlighting that a single “best” fragment size cannot be universally applied across Gulf taxa. Light color also affected coral growth but only for some species and size classes. Different coral species showed variable and even opposing responses under the same light color treatment. Collectively, these results provide the first species−level benchmarks for optimizing fragment size and light spectrum for land−based propagation of Arabian Gulf corals. Such evidence−based parameters can reduce production time and cost, improve nursery efficiency, and support the scaling of restoration programs.

Advancing restoration practices in the extreme environment of the Arabian Gulf is of critical relevance for the future of coral reefs globally. This work contributes to raising the TRL for coral gardening approaches tailored to thermally extreme reef systems. Future research should investigate additional drivers of nursery performance, such as heterotrophic feeding, substrate interactions, and genotype effects, as well as explore the physiological and molecular mechanisms underlying species−specific responses. Such work will further enhance the effectiveness and scalability of restoration initiatives across the region.

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