Ecological and behavioural responses of butterflyfishes to coral reef restoration

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S. Sudrajat, Andi Tanri Abeng, and 3 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8883345/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Coral restoration is increasingly applied to combat reef habitat degradation, yet its impacts on fish populations and behavioural dynamics remain underexplored. This study examines how coral restoration influences the population density and behaviour of butterflyfish (Chaetodontidae) – obligate corallivores and recognized bioindicators of reef health. We measured the abundance, diversity and aggressive territorial behaviour of butterflyfish at one of the world’s largest coral restoration projects (Bontosua Island, South Sulawesi, Indonesia), comparing naturally healthy reefs with those that had been restored after historic damage from dynamite fishing. Additionally, we compared these data with equivalent metrics from previous studies at other natural reefs in the Central Indo-Pacific region, to contextualise our data within a wider ecological framework. At Bontosua, coral restoration led to significant recovery of butterflyfish abundance within six years, with both restored and healthy reefs exhibiting high butterflyfish abundance but low species diversity and consistently low rates of aggressive interactions. On both natural and restored reefs, butterflyfish communities at Bontosua were dominated by a single species, Chaetodon octofasciatus – with low levels of both inter- and intra-specific aggressive behaviour. This contrasts with more diverse and competitive butterflyfish communities in other locations around the Central Indo-Pacific. We conclude that butterflyfish behavioural patterns were more sensitive to broad-scale regional variation than to restoration outcomes at this site. Butterflyfish behaviour may serve as a more effective indicator of regional ecosystem conditions than of localized restoration success. butterflyfish fish behaviour coral restoration coral rehabilitation Indonesia Central Indo-Pacific Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Introduction Coral reefs are among the world’s most biodiverse and productive ecosystems, yet they are increasingly threatened by both anthropogenic and natural stressors (Good & Bahr, 2021 ; Hughes et al., 2017 ; Hoegh-Guldberg & Bruno, 2010 ). These stressors often interact synergistically, exacerbating the decline of coral reef ecosystems (Castro-Sanguino et al., 2021 ; Pandolfi et al., 2003 ). The cumulative impact of these stressors has led to extensive damage across many of the world’s coral reefs, reducing biodiversity, compromising ecosystem services, and threatening the livelihoods of communities that depend on them. Reef-associated fishes are particularly affected by reef degradation, as many species rely on live coral for food, shelter, and breeding grounds (Cole et al., 2008 ; Jones et al., 2004 ), resulting in a strong positive relationship between coral cover and both fish abundance and diversity(Pratchett et al., 2011 ;Findley & Findley, 2001 ; Bell & Galzin, 1984 ). As coral cover declines, associated fish populations often decrease (Coker et al., 2014 ). Coral loss can also influence fish behaviour, for example by increasing aggression as individuals compete for limited resources(Kok et al., 2016 ; Pratchett et al., 2006 ;Graham, 2007 ) or decreasing aggression as the energy required to hold a territory outweighs the benefits. This is particularly evident in specialist species, such as obligate corallivores, which are especially vulnerable to changes in coral composition (Keith et al., 2018 ; Pratchett et al., 2014 ; Cole et al., 2008 ). Butterflyfish (family Chaetodontidae) are particularly dependent on healthy coral communities because many species in this family are obligate corallivores (Pratchett et al., 2008 ). Habitat degradation and the loss of live coral therefore pose a significant threat to butterflyfish, by reducing resource availability and disrupting biological interactions such as competition and predation (Keith et al., 2023 ; Coker et al., 2009 ). Among butterflyfish, aggression is known to vary in response to resource availability (Kok et al., 2016 ). When coral is scarce, aggression may be low because defending a territory is not worth the energetic effort if it does not contain valuable resources; for example, Keith et al ( 2018 ) found reduced aggression across the Central Indo-Pacific following a mass bleaching event. Conversely, aggression can also be low when coral is very abundant, because resources are readily available and easy to obtain, so defending a territory is not necessary (Peiman & Robinson, 2010 ). Aggression tends to peak when resources are at an intermediate level—limited but defensible—because individuals gain a clear benefit from protecting valuable foraging areas (Peiman & Robinson, 2010 ; Maher & Lott, 2000 ). As such, butterflyfish behaviour is likely to change in response to changes in coral cover. In response to the widespread degradation of coral reefs in recent decades, coral restoration efforts have increased around the world, as a strategy to support reef recovery and maintain ecosystem services (Bayraktarov et al., 2020 ; Hein et al., 2021 ). Coral reef restoration includes a wide range of different techniques such as rubble stabilisation (Leung et al., 2025 ), coral transplantation (Ferse et al., 2021 ), the use of artificial structures to promote reef accretion (Tanaya et al., 2025 ), and assisted evolution techniques aimed at enhancing coral resilience to both anthropogenic and natural stressors (van Oppen et al., 2015 ). These active reef restoration efforts seek to restore coral cover, physical reef structure, ecological function, and biodiversity (Boström-Einarsson et al., 2020 ). Restoration can produce conditions similar to those of natural coral reef habitats in some cases, but in other cases restored reefs can differ from naturally healthy reefs in aspects such as community composition, structural complexity, and ecological interactions (Vida et al., 2024 ; Lange et al., 2024 ; Lamont et al., 2022 ). Differences between natural and restored reefs may influence habitat suitability for corallivores and thus patterns of corallivory (Ladd & Shantz, 2020 ). Restored reefs are often dominated by fast-growing coral genera such as Acropora , due to their rapid growth and colonization rates (Hein, et al., 2020 a; Lirman et al., 2014 ). Acropora corals are a preferred food source for some butterflyfish species (Semmler et al., 2022 ). If restored reefs contain different amounts of preferred food sources, butterflyfish behaviour may also differ in restored environments (Hein et al., 2019 ). Understanding how butterflyfish respond behaviourally to restored versus natural reef habitats can provide valuable insight into the effectiveness of restoration efforts in replicating natural ecological conditions. This study aims to quantify butterflyfish behaviour in both natural and restored coral reef habitats. It focuses on two primary objectives: first, we tested whether there are differences in butterflyfish abundance and aggressive behaviour between natural healthy and restored reefs at one of the world’s largest coral restoration (the ‘Mars Assisted Reef Restoration System (MARRS)’ restoration site in Bontosua Island, South Sulawesi, Indonesia). Second, we compared these behavioural patterns with those observed in other reef systems across the Central Indo-Pacific (Keith et al. 2018 ), in order to contextualise these data within regional ecological variation. By examining the behaviour of obligate corallivorous species, this research contributes to a broader understanding of the ecological impacts of coral reef restoration and provides a more comprehensive framework for evaluating restoration success. Materials and methods Study sites This study compared three coral reef habitat types at Bontosua Island in the Spermonde Archipelago, South Sulawesi, Indonesia: degraded reefs, restored reefs, and naturally healthy reefs. Degraded reefs in this study had been severely impacted by historical human activities, including boat channel construction (30–40 years ago), blast fishing (30 years ago), and coral mining for infrastructure (20 years ago) (Williams et al., 2019 ; Sawall et al., 2013 ; Plass-Johnson et al., 2015 ). Such disturbances have damaged the coral framework, resulting in unstable rubble fields and a significant reduction in reef structural complexity. Unconsolidated rubble inhibits coral recruitment and impedes the formation of mature coral colonies (Cameron et al., 2016 ; Fox et al., 2003 ). To address this degradation, the MARRS Coral Reef Restoration Project ( www.buildingcoral.com ) initiated a restoration effort using hexagonal steel structures (Reef Stars) coated with resin and sand (Smith et al., 2021 ). These interconnected structures create a stable substrate for reattaching coral fragments, and restoration on Bontosua Island led to rapid increases in coral cover, fish abundance, and biomass within two to three years (Smith et al., 2021 ). Restored reefs in this study had been restored using this method for between one and six years. Healthy reefs in this study had historically remained in good condition, supporting thriving coral communities and showing no evidence of the historical disturbance that affected degraded reefs. The total study area included 29 reef sites, comprising 3 degraded sites, 7 healthy sites, and 19 restored sites. The restored sites were further classified into two age categories: 11 old restored sites (5–7 years post-restoration) and 8 young restored sites (1–3 years post-restoration). In addition to comparing habitat conditions at Bontosua Island, we also compared these data to other sites around the Central Indo-Pacific, using data from Keith et al. ( 2018 ). Data from Bali (Indonesia), Christmas Island (Indian Ocean, Australia), Iriomote (Japan), and Luzon (Philippines) (Fig. 1 ) were sourced from Keith et al. ( 2018 ) based on their representation of a range of natural healthy reef conditions across the Central Indo-Pacific region. In order to represent naturally healthy conditions, only data from before the mass bleaching event studied in Keith et al. ( 2018 ) was used. This regional comparison enabled us to contextualize patterns observed in our location (Bontosua) within a wider biogeographical framework. Butterflyfish abundance and diversity surveys Butterflyfish abundance and species diversity were quantified using underwater visual census (UVC) surveys at each study site. Surveys were conducted along transects that were 5m wide, with the transect length adjusted based on the size of the surveyed site. Transects were 50 m in length where habitat allowed, or 20 m, 33 m, and 40 m in length in smaller restoration areas. Transects were positioned parallel to the reef crest, targeting reef zones most frequently used by corallivorous butterflyfish. After transects were laid, an acclimation period (~ 1 min) was observed, during which the observer ensured that butterflyfish were behaving naturally and not disturbed by observer presence. Following this acclimation period, butterflyfish abundance was quantified by counting all individuals within 2.5 m of either side of the transect. Butterflyfish behaviour We quantified the frequency of interactions between butterflyfish individuals, and the aggression levels present within each interaction, following the same methods used in Keith et al. ( 2018 ). We observed interactions between focal butterflyfish individuals and other Chaetodon species, including both conspecifics and heterospecifics. Data were collected using SCUBA, and observers followed a modified U-shaped swimming pattern at each site to minimize the likelihood of repeatedly observing the same individuals. Behavioural observations were only carried out in healthy and restored habitat but not in degraded habitat, because there were not enough butterflyfish present on degraded reefs to ensure adequate sample size. We carried out behavioural observations in the same area as the abundance surveys, immediately after abundance surveys were carried out. The first butterflyfish encountered from any species was designated as the focal individual. Many Chaetodon species occur in pairs, and since no aggression was observed between partners, only one fish per pair was observed. Focal individuals were followed at a distance of 2–4 m for five minutes each. Encounters were recorded when another butterflyfish individual came within 1 m of the focal individual. At this distance, it was assumed that both individuals were aware of each other and had the potential to interact. Interactions were categorized as passive if no behavioural response could be observed, or aggressive if one or both individuals displayed signalling (dorsal fin raised, head down) or chasing behaviour. For each observation the time of day, water temperature, focal fish species, interacting fish species, distance at the point of encounter, and the size (cm) of both the focal and interacting fish were also recorded. Statistical analysis Butterflyfish abundance was modelled as a function of restoration age using a linear mixed model (LMM). The response variable was butterflyfish abundance (individuals m⁻²), with restoration age in years included as a fixed effect, and site as a random intercept to account for spatial non-independence. Degraded reefs were included with a restoration age of zero to provide a baseline for comparison, while healthy reefs were excluded from the model and compared visually on a graph as reference sites. Model residuals were inspected to goodness-of-fit. Live coral cover was modelled as a function of restoration age using the same fixed and random effect structure with butterflyfish abundance model, but using a beta error distribution with a logit link in a generalised linear mixed model (GLMM). Butterflyfish abundance at each location was analyzed using non-parametric methods because the data violated assumptions of normality and homogeneity of variance, as assessed by the Shapiro–Wilk test. Differences in butterflyfish abundance per square meter among location were first evaluated using a Kruskal–Wallis rank-sum test. When significant differences were detected, pairwise Wilcoxon rank-sum tests were performed to determine which location pairs differed, with p-values adjusted using the Bonferroni correction. The degraded site in Bontosua exhibited very low butterflyfish abundance and species diversity, with a small sample size (n = 6). As a result, statistical comparisons for this location could not be conducted due to insufficient statistical power. Butterflyfish species diversity was quantified using the Shannon–Wiener index (H′), which incorporates both species richness and evenness (Kunakh et al., 2023; Hurlbert, 1971). Differences in diversity among locations were tested using the non-parametric Kruskal–Wallis test. Significant results were followed by pairwise Wilcoxon rank-sum tests with Bonferroni corrections to identify differences between specific site pairs. To test for species dominance in the butterflyfish community, the abundance of the most common butterflyfish species was compared to the total abundance of all other butterflyfish species at each site. At each site, paired Wilcoxon signed-rank tests were used to compare the abundance of the dominant species against all other species. The frequency of butterflyfish interactions was analysed using generalized linear mixed-effects models (GLMMs) with a negative binomial distribution to account for overdispersed count data. Aggression was quantified as the proportion of interactions that were aggressive (bounded between 0 and 1) and analysed using GLMMs with a beta distribution and logit link. The response variable for the interaction frequency model was the number of interactions per focal fish, whereas for the aggression model it was the proportion of interactions classified as aggressive for each focal fish. In both models, interaction type (conspecific vs. heterospecific) and location (Bontosua Restored Old, Bontosua Restored Young, Bontosua Healthy, Bali, Christmas Island, Iriomote, and Luzon) were included as fixed effects. The two models differed in their random-effects structure. For interaction frequency, site was included as a random intercept to account for non-independence of observations within sites. For aggression proportion, both site and focal species were included as random intercepts to account for variation among sites and species-level differences in aggressive behaviour. All analyses were conducted in R v.4.4.0 (R Core Team, 2024), using the packages glmmTMB package (Brooks et al., 2017 ), emmeans (Lenth, 2023 ), DHARMa(Hartig, 2016 ) and ggplot2 (Wickham, 2016 ). Results Butterflyfish abundance in response to time since restoration The abundance of butterflyfishes in restored sites at Bontosua Island increased with time since restoration, from 0.02 individuals m⁻² at one year old site to 0.12 ± 0.03 individuals m⁻² at seven years old sites (Fig. 2 a). This overall pattern of increase was a significant positive trend (LMM: estimate = 0.015 ± 0.004 SE, z = 3.77, p < 0.001; Table S1 ), corresponding to an annual increase of ~ 0.015 individuals m⁻². Post-hoc contrasts between individual years showed that abundance at six-year-old reefs was significantly higher than in degraded reefs, but reefs restored younger than that age (1–5 years old) did not differ significantly from degraded reefs (Table S1 ). Seven-year reefs also exhibited higher abundances than degraded reefs, and were comparable with the healthy sites, though this effect was not statistically significant (Table S1 ). Together, these results indicate a clear positive trajectory in butterflyfish abundance with reef age, with significant recovery evident from approximately six years post-restoration. This positive trend in butterflyfish abundance aligned with a corresponding increase in coral cover. Live coral cover also increased strongly with restoration age (GLMM, β = 0.377 ± 0.057 SE; z = 6.65, p < 0.001). Coral cover increased from 2.8% at one-year restored sites to 57.4% ± 6.1% at seven-year restored sites, approaching levels found on healthy reefs (54.5% ± 4.4%) and exceeding those of degraded sites (10.3% ± 3.9%) (Fig. 2 b). Pairwise contrasts indicated that there was no significant difference between degraded and 1-year old restored reefs, but restored reefs of 2 years or older all had significantly higher coral cover than degraded reefs, with effect sizes strengthening with restoration age (Table S2). Comparing butterflyfish populations at Bontosua Island with other locations in the Central Indo-Pacific Butterflyfish abundance (individuals per m²) differed significantly among habitats across the Central Indo-Pacific (Kruskal–Wallis test: χ² = 47.13, df = 6, p < 0.001; Table S3). In particular, abundance at Bontosua Healthy and Bontosua Restored (Old) were significantly higher than at other location around the Central Indo-Pacific (Wilcoxon rank-sum test, W = 2716, p < 0.001; Fig. 3 A). But not different significant with Bontosua Restored (Young). Post-hoc pairwise Wilcoxon rank-sum test (Bonferroni-corrected) showed that Bontosua Healthy sites had significantly higher butterflyfish abundance than Christmas Island (W = 200, p < 0.01) and Iriomote (W = 253, p < 0.001), but did not differ significantly from Bali, Luzon, or either restored habitat at Bontosua. Among restored habitats, Bontosua Restored (Old) sites exhibited significantly greater butterflyfish abundance than Christmas Island (W = 436, p < 0.001), Iriomote (W = 558, p < 0.001), and Luzon (W = 530, p < 0.01). Bontosua Restored (Young) sites did not differ significantly from Bali, Christmas Island, Iriomote, or Luzon, nor from Bontosua Healthy or Bontosua Restored (Old) sites. No significant differences were detected between Bali and any Bontosua habitat (Healthy, Restored (Old), or Restored (Young)) (Fig. 3 A). Butterflyfish species diversity varied significantly across locations (Kruskal–Wallis test: χ² = 34.7, df = 6, p < 0.001; Table S4). Post-hoc pairwise comparisons using Bonferroni-adjusted Wilcoxon rank-sum tests revealed strong spatial differences in species diversity among locations. Diversity at Bontosua Restored (Old) sites was significantly lower than at both Bali (W = 292, p < 0.001) and Luzon (W = 115, p < 0.001), but did not differ from Christmas Island and Iriomote. A similar pattern was observed at the Bontosua Healthy sites, where diversity was significantly lower than at Bali (W = 126, p < 0.05) and Luzon (W = 41, p < 0.05), while showing no significant differences from Christmas Island, Iriomote, and the Bontosua Restored sites. Overall, both restored and healthy reefs at Bontosua exhibited reduced species diversity relative to Bali and Luzon, but were broadly similar to Christmas Island and Iriomote. Dominant species of butterflyfish Significant differences in butterflyfish species composition were observed among locations (Kruskal-wallis test, χ² = 27.1, df = 6, p < 0.001, Fig. 4 , Table S5). At Bontosua Restored (Old) sites, the most common species was significantly more abundant than all other species combined (Wilcoxon test, W = 470, p < 0.001, r = 0.467), and a similar dominance pattern was observed at the Bontosua Healthy sites (W = 64, p < 0.001, r = 0.155). In contrast, Bontosua Restored (Young) showed no significant dominance by any species. In contrast, the other locations exhibited a more even species composition, with the abundance of the dominant species either less than, or not significantly different from, the combined abundance of all other species (Fig. 4 ). There were significantly higher abundances of other species relative to the most common species at Bali (W = 41, p < 0.01, r = 0.124), Christmas Island (W = 175, p < 0.05, r = 0.155), and Luzon (W = 413, p < 0.05, r = 0.111). At Iriomote, there was no significant difference between the abundance of the most common species and the abundance of all other species (Wilcoxon signed-rank test, W = 623, p = 0.312, r = 0). Butterflyfish behaviour Across all locations except for Bontosua, heterospecific interactions occurred more frequently than conspecific interactions (GLMM, estimate = 1.06 ± 0.09, z = 12.15, p < 0.001, Fig. 5 a, Table S6). In Bontosua, conspecific interactions occurred more frequently than heterospecific interactions. This effect was strongest at Bontosua Healthy sites (GLMM, estimate = 2.37 ± 0.40, z = 5.95, p < 0.001), while at Bontosua Restored (Old) (GLMM, estimate = 0.68 ± 0.28, z = 2.45, p = 0.014) and Bontosua Restored (Young) showed no significant difference between interaction types (GLMM, estimate = 0.16 ± 0.25, z = 0.62, p = 0.538). In contrast, the other locations—Bali, Christmas Island, Iriomote, and Luzon, exhibited significantly higher frequencies of heterospecific interactions. Pairwise contrasts revealed significant effects for Bali (GLMM, estimate = − 2.71 ± 0.13, z = − 21.26, p < 0.001), Christmas Island (GLMM, estimate = − 2.66 ± 0.14, z = − 19.50, p < 0.001), Iriomote (GLMM, estimate = − 2.76 ± 0.14, z = − 20.34, p < 0.001), and Luzon (GLMM, estimate = − 2.50 ± 0.14, z = − 18.22, p < 0.001), indicating that butterflyfish tended to interact more often with individuals of other species than with conspecifics Conspecific interactions tended to be slightly more aggressive than heterospecific interactions across locations, although this overall effect was not statistically significant (GLMM: β = − 0.11 ± 0.52, z = − 0.21, p = 0.83). However, the model revealed significant interactive effects of interaction type and location on aggression frequency (Fig. 5 b). Post-hoc comparisons showed that conspecific aggression was significantly higher than heterospecific aggression in Bali (β = 2.64 ± 0.35, z = 7.57, p < 0.001), Christmas Island (β = 1.25 ± 0.37, z = 3.35, p < 0.005), Iriomote (β = 3.13 ± 0.39, z = 8.08, p < 0.001), and Luzon (β = 1.79 ± 0.35, z = 5.08, p < 0.001). In contrast, no significant differences between conspecific and heterospecific aggression were detected at the Bontosua sites, including Bontosua Healthy (β = 0.12 ± 0.53, z = 0.23, p = 0.821), Bontosua Restored (Old) (β = 1.11 ± 0.52, z = 0.21, p = 0.831), and Bontosua Restored (Young) (β = − 0.10 ± 0.37, z = − 0.27, p = 0.786). Discussion General findings This study focused on the behavioural ecology of butterflyfish (family Chaetodontidae) to evaluate the ecological outcomes of coral reef restoration on Bontosua Island, with comparative insight drawn from a broader regional context encompassing Bali (Indonesia), Christmas Island (Indian Ocean, Australia), Iriomote (Japan), and Luzon (Philippines). At the restored reefs at Bontosua, butterflyfish abundance increased progressively with restoration age and increasing coral cover. This pattern aligns with previous findings that link the presence and population growth of butterflyfish to live coral cover (Pratchett et al., 2008 ). The observed population growth likely reflects the increased availability of coral in the restored sites. Across both healthy and restored sites, particularly the older restored sites, butterflyfish communities on Bontosua Island exhibited patterns distinct from those in other locations. While both healthy and older restored reefs on Bontosua supported substantially higher butterflyfish abundances, their species diversity was markedly lower. This reduced diversity was driven by the strong dominance of a single species, Chaetodon octofasciatus , which comprised the majority of the butterflyfish community at Bontosua. Behavioural patterns were also distinctive at Bontosua Island compared to other locations around the Central Indo-Pacific. At Bontosua, aggressive interactions were infrequent, with levels of aggression consistently low across both restored and healthy reefs. This pattern may be attributable to the dominance of a single species ( C. octofasciatus ), resulting in reduced interspecific competition and relaxed competition among conspecifics. The consistently low aggression observed in C. octofasciatus may reflect weak territoriality or reliance on concealment rather than overt defence. In contrast, behavioural data from other locations revealed higher levels of aggression, possibly due to greater species diversity and increased interspecific competition at those sites. Abundance and behavioural patterns in restored reefs and healthy reefs in Bontosua Island Restoration efforts in Bontosua have significantly increased live coral cover and small-scale structural complexity of reef habitats compared to previously degraded conditions (Vida et al., 2024 ). These improvements are likely to be the driver of the observed increase in corallivorous fish (Santoso et al., 2022 ; Darling et al., 2017 ). The three-dimensional framework of live corals provides microhabitat for shelter, territorial behaviour, and spawning activities (Coker et al., 2014 ), and live coral serves as the primary food source for corallivorous butterflyfish (Cole et al., 2008 ; Pratchett, 2007 ; Rotjan & Dimond, 2010 ). While coral cover recovered substantially within the first three years of restoration, butterflyfish abundance required approximately five to seven years to reach levels comparable to healthy reefs, indicating a lag in the biological response of fish communities to habitat improvement. Similar lagged responses have been documented by (Hein et al., 2020 ), who reported gradual increases in fish abundance over several years following coral restoration sites, even after coral cover had already recovered. Comprehensive reviews also suggest that while coral cover may rebound from degradation within three years, full recovery of fish communities, particularly among specialist species, can take five to fifteen years (McClanahan et al., 2007 ). Despite the increase in abundance, butterflyfish species diversity remained low at all sites at Bontosua. Communities in Bontosua were dominated by a single species, C. octofasciatus . This pattern was also observed in healthy reefs, suggesting that such single-species dominance is not a product of restoration, but a characteristic of the reefs around this island. Comparable findings were reported from other islands within the Spermonde Islands, where C. octofasciatus emerged as the dominant species among 21 Chaetodontidae species surveyed across 13 locations (Putra et al., 2024 ). C. octofasciatus exhibits high resilience to fishing pressure, with a population doubling time of less than 15 months pressure making it a suitable indicator for monitoring rapid habitat changes even under conditions of intense exploitation (Froese & Pauly, 2010; Madduppa et al., 2014 ). These ecological traits are particularly relevant to the Spermonde region, including Bontosua Island, where destructive fishing practices continue to exert substantial pressure on reef ecosystems. Additional studies have demonstrated C.octofasciatus tolerance to elevated water turbidity (Bramasta et al., 2024 ) and its frequent occurrence in habitats dominated by rubble or dead coral substrates (Songploy et al., 2017 ). As the dominant species, C. octofasciatus likely underpins the consistently low levels of aggressive behaviour observed at Bontosua Island. This numerical dominance may reduce aggressive encounters between different species, resulting in consistently low aggression levels across both restored and healthy reefs. Although this species is an obligate corallivore (Cole et al., 2008 ; Allen et al., 1998 ; Michael, 2004 ), previous studies have demonstrated that it exhibits relatively weak territorial behaviour compared to other obligate corallivores such as C. baronessa and C. trifascialis (Blowes et al., 2017 ;Pratchett & Berumen, 2008 ). Field observations from Redang Island, Malaysia, indicate that C. octofasciatus typically occupies relatively small territories and frequently seeks refuge among coral branches (Ghaffar et al., 2006 ). Such patterns suggest that C. octofasciatus exhibits weak territoriality and relies more on concealment than aggressive defence. This aligns with our findings, in which the proportion of aggressive encounters involving C. octofasciatus was substantially lower than that observed for other species (Table S8). Moreover, the very low number of aggressive encounters recorded overall at Bontosua Island further supports the conclusion that C. octofasciatus exhibits low levels of aggression. Hence, these findings indicate that C. octofasciatus may be a non- or weakly territorial species. Reefs and butterflyfish behaviour patterns in other locations across the Central Indo-Pacific Biogeographic factors play an important role in shaping butterflyfish community structure and behaviour, in some regions, species diversity remains low despite high coral cover, reflecting ecological or historical constraints (Findley & Findley, 2001 ). The ecological similarity observed between restored and healthy habitats in Bontosua suggests that regional (biogeographic) conditions may exert a stronger influence on community structure than the restoration status of individual reefs. This is supported by the strong correlation between coral cover and fish abundance at Bontosua compared to other locations in the Central Indo-Pacific. These findings indicate that the response of butterflyfish to coral cover is location-dependent and may be influenced by local ecological factors such as habitat complexity, interspecific interactions, or environmental stressors, as well as by broader biogeographic constraints. Both restored and healthy reefs in Bontosua displayed butterflyfish abundance and social interaction patterns distinct from those reported in other parts of the Central Indo-Pacific. Lower species richness and reduced aggression observed in Bontosua may be influenced by multiple factors, including latitudinal gradients (Siqueira et al., 2016 ; González-Barrios et al., 2025 ), limited regional larval supply (van der Meer et al., 2013 ), historical habitat degradation (Jackson et al., 2001 ; Wilson et al., 2008 ), or reduced connectivity with more diverse source populations (Hobbs & Srinivasan, 2024 ). Although Bontosua is located within a globally recognized marine biodiversity hotspot, its low-diversity butterflyfish community structure may reflect regional-level biogeographic isolation or ongoing recovery from past disturbances. In particular, historical degradation events may have led to local extinctions or demographic bottlenecks that reduced species richness, particularly among less resilient or weakly dispersing taxa. Conclusion This study demonstrates that both restored, particularly older restoration sites and healthy reef habitats on Bontosua Island exhibit similar characteristics in their butterflyfish populations, including high butterflyfish abundance, low species richness, and consistent behavioural patterns marked by low aggression. The observed increase in butterflyfish abundance over time in restored areas suggests a positive response to restoration efforts, likely driven by improvements in coral cover and structural complexity. Compared to other sites in the Central Indo-Pacific, Bontosua stands out for its unusually high butterflyfish abundance and low species diversity, a pattern largely attributed to the dominance of C. octofasciatus . This low-diversity community structure, present in both healthy and restored reefs, may reflect limited regional larval supply, historical habitat degradation, or reduced connectivity to more diverse source populations. The dominance of a single species may also contribute to reduced interspecific competition, as C. octofasciatus is thought to be non- or weakly territorial. Overall, these findings suggest that reef restoration at Bontosua has successfully supported the recovery of butterflyfish populations to levels comparable with nearby healthy reefs. However, the patterns of aggressive behaviour appear to be shaped more strongly by broader regional ecological conditions than by the local effects of restoration alone. Declarations Author Contribution **Sera Maserati** : conceptualization, data curation, formal analysis, investigation, methodology, visualization, writing – original draft preparation and editing. **Timothy A.C. Lamont:** conceptualization, methodology, investigation, funding acquisition, supervision, writing – review & editing, project administration. **Jane C.S. Sudrajat** : investigation. **Andi Tanri Abeng** : investigation. **Permas B. Maulana:** investigation. **Sally A. Keith:** methodology, investigation. writing – review. **Tries B. Razak** : conceptualization, methodology, supervision, funding acquisition, writing – review & editing, project administration. Acknowledgement Survey data for this study were collected as part of the monitoring program of the Mars Coral Reef Restoration Project, in collaboration with Universitas Hasanuddin. We thank L. Damayanti, P. Mansell, and members of the Mars Sustainable Solutions monitoring team (M. E. Prasetya, P. B. Maulana, A. Hamka, A. Dwiyanto, A. M. A. Pratama, A. T. Abeng, Irwan, R. Madjid, E. Agiel, and Suandar) for their invaluable support with field logistics and monitoring activities. We are also grateful to the Department of Marine Affairs and Fisheries of South Sulawesi Province, the Government Offices of Kabupaten Pangkep, Pulau Bontosua, and Pulau Badi, as well as the community of Pulau Bontosua, for their support and cooperation. We thank S. Keith for helpful discussions and for contributions to data collection outside Indonesia. Fieldwork in Indonesia was conducted under a national research permit issued by BRIN (No. 108/SIP/IV/FR/2/2023), jointly held by T.A.C.L. as lead foreign researcher and T.B.R. as lead Indonesian host researcher, with ethical approval granted by BRIN and Lancaster University. We thank Prof. J. Jompa and Prof. R. A. Rappe at Universitas Hasanuddin for logistical assistance with permit and visa applications. Funding was provided by a research fellowship from The Royal Commission for the Exhibition of 1851 (awarded to T.A.C.L.), an international traveling fellowship from the Fisheries Society of the British Isles (awarded to T.A.C.L. and T.B.R.), and a Pew Fellowship in Marine Conservation (awarded to T.B.R.). Data Availability The datasets generated and/or analysed during the current study will be deposited in a public repository upon acceptance of the manuscript. References Allen, G. R., Steene, R., & Allen, M. (1998). A Guide to Angelfishes & Butterflyfishes. Odyssey Publishing, Perth , 250. Bayraktarov, E., Banaszak, A. T., Maya, P. M., Kleypas, J., Arias-Gonzalez, J. E., Blanco, M., Calle-Triviño, J., Charuvi, N., Cortes-Useche, C., Galvan, V., Salgado, M. A. G., Gnecco, M., Guendulain-Garcia, S. D., Delgado, E. A. H., Moraga, J. A. M., Maya, M. F., Quiroz, S. M., Cervantes, S. M., Morikawa, M., … Frias-Torres, S. (2020). 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S. Sudrajat","email":"","orcid":"","institution":"IPB University","correspondingAuthor":false,"prefix":"","firstName":"Jane","middleName":"C. S.","lastName":"Sudrajat","suffix":""},{"id":629477100,"identity":"a87d3d18-6220-421a-9bf1-17e492fd9e50","order_by":3,"name":"Andi Tanri Abeng","email":"","orcid":"","institution":"Mars Sustainable Solutions","correspondingAuthor":false,"prefix":"","firstName":"Andi","middleName":"Tanri","lastName":"Abeng","suffix":""},{"id":629477101,"identity":"8944ff9b-646f-480d-b5c0-8b263e3fc3f7","order_by":4,"name":"Permas B. Maulana","email":"","orcid":"","institution":"Mars Sustainable Solutions","correspondingAuthor":false,"prefix":"","firstName":"Permas","middleName":"B.","lastName":"Maulana","suffix":""},{"id":629477102,"identity":"859d9ef9-7a98-4776-adbc-6162666c1a51","order_by":5,"name":"Sally A. Keith","email":"","orcid":"","institution":"Lancaster University","correspondingAuthor":false,"prefix":"","firstName":"Sally","middleName":"A.","lastName":"Keith","suffix":""},{"id":629477103,"identity":"70c60708-cfbd-4b41-af51-841a525381fc","order_by":6,"name":"Tries B. Razak","email":"","orcid":"","institution":"School of Coral Reef Restoration (SCORES), Faculty of Fisheries and Marine Science, IPB University","correspondingAuthor":false,"prefix":"","firstName":"Tries","middleName":"B.","lastName":"Razak","suffix":""}],"badges":[],"createdAt":"2026-02-15 03:08:28","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-8883345/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-8883345/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":108181996,"identity":"60c15577-fe0d-4cd9-abb0-5817c227c95c","added_by":"auto","created_at":"2026-04-30 08:59:03","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":709673,"visible":true,"origin":"","legend":"\u003cp\u003eMap of the study area. (a) Location of Bontosua Island in South Sulawesi, alongside comparison locations from Keith et al. (2018); (b),(c), and (d) detailed maps of the study sites on Bontosua Island, with colored points indicating different coral reef habitats (healthy, degraded, restored) and varying restoration ages of 1, 2, 3, 5, 6, and 7 years. Photographs showing examples of the three habitat conditions in Bontosua: (e) healthy reef, (f) restored reef, and (g) degraded reef.\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-8883345/v1/f505c50eba5d42344a5ac45d.png"},{"id":108182185,"identity":"d83b0553-8115-4d78-bf1f-8cc2b6a81dcf","added_by":"auto","created_at":"2026-04-30 08:59:11","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":269524,"visible":true,"origin":"","legend":"\u003cp\u003eRecovery of butterflyfish populations and live coral cover over time since coral restoration in Bontosua. (a) Butterflyfish abundance (individuals per m²) and (b) live coral cover (%) both increased with restoration age. Blue lines show generalized linear mixed model (GLMM) fits testing the effect of time since restoration, with shaded areas indicating 95% confidence intervals. For healthy habitats, boxplots show the median (centre line), interquartile range (boxes), and full data range (whiskers).\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-8883345/v1/7d1dbd196641274eb211dbeb.png"},{"id":108045500,"identity":"dd6331ed-0287-4c4d-882e-eb7269ce73d5","added_by":"auto","created_at":"2026-04-28 19:25:10","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":1881713,"visible":true,"origin":"","legend":"\u003cp\u003e(a) Butterflyfish abundance and (b) species diversity across Bontosua and comparison locations from Keith et al. (2018), shown as boxplots. Bontosua healthy and restored sites exhibited the highest abundance and the lowest diversity of butterflyfish. Boxplots display the median (centre line), interquartile range (boxes), full range (whiskers), and individual transects (points). Different letters above boxplots indicate significant differences among habitats based on pairwise Wilcoxon rank-sum tests with Bonferroni correction (\u003cem\u003ep\u003c/em\u003e \u0026lt; 0.01). Bontosua degraded was excluded from statistical tests and diversity estimates due to very small sample size.\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-8883345/v1/35b2744470a7896bb903e5eb.png"},{"id":108045499,"identity":"7e5d2dd5-c222-47b2-b9a0-dd6a84a905fd","added_by":"auto","created_at":"2026-04-28 19:25:10","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":348595,"visible":true,"origin":"","legend":"\u003cp\u003eButterflyfish abundance across locations, highlighting the most abundant species at each site. Yellow bars represent the dominant (most abundant) species (labelled above each bar), while green bars show the combined abundance of all other species. Bar height indicates mean abundance (individuals/m²), with error bars representing the standard error. Significant differences between groups within each location were assessed using Wilcoxon signed-rank tests and are denoted by different letters above the bars.\u003c/p\u003e","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-8883345/v1/19c7f205896290b8283ad737.png"},{"id":108181218,"identity":"c52c2178-26d0-4793-99d1-5f09408a9eb7","added_by":"auto","created_at":"2026-04-30 08:58:25","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":604227,"visible":true,"origin":"","legend":"\u003cp\u003eButterflyfish behaviour in conspecific (same species) and heterospecific (different species) interactions. (a) Interaction frequency with each point representing an individual butterflyfish. (b) Aggression rate (% interactions involving chasing), with each point representing a site. Violin plots show data distribution; embedded boxplots indicate the median (centre line), interquartile range (boxes), and data range (whiskers). Different letters denote significant differences between interactions types within each location, based on generalized linear mixed models (GLMMs) with a beta distribution.\u003c/p\u003e","description":"","filename":"floatimage5.png","url":"https://assets-eu.researchsquare.com/files/rs-8883345/v1/0bca8109255313e5e1dac19a.png"},{"id":108491506,"identity":"d06b14c1-6d3a-48d0-a818-484276696bb2","added_by":"auto","created_at":"2026-05-05 09:54:13","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":3217908,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8883345/v1/531da6a6-788a-455a-9968-ae76d894fcd2.pdf"},{"id":108045497,"identity":"24056313-8f11-4791-9ad0-570e29ca2aeb","added_by":"auto","created_at":"2026-04-28 19:25:10","extension":"docx","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":27361,"visible":true,"origin":"","legend":"","description":"","filename":"Supplementarymaterial.docx","url":"https://assets-eu.researchsquare.com/files/rs-8883345/v1/f9ab4836d26745464ddad211.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Ecological and behavioural responses of butterflyfishes to coral reef restoration","fulltext":[{"header":"Introduction","content":"\u003cp\u003eCoral reefs are among the world\u0026rsquo;s most biodiverse and productive ecosystems, yet they are increasingly threatened by both anthropogenic and natural stressors (Good \u0026amp; Bahr, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Hughes et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Hoegh-Guldberg \u0026amp; Bruno, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). These stressors often interact synergistically, exacerbating the decline of coral reef ecosystems (Castro-Sanguino et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Pandolfi et al., \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). The cumulative impact of these stressors has led to extensive damage across many of the world\u0026rsquo;s coral reefs, reducing biodiversity, compromising ecosystem services, and threatening the livelihoods of communities that depend on them.\u003c/p\u003e \u003cp\u003eReef-associated fishes are particularly affected by reef degradation, as many species rely on live coral for food, shelter, and breeding grounds (Cole et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Jones et al., \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2004\u003c/span\u003e), resulting in a strong positive relationship between coral cover and both fish abundance and diversity(Pratchett et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2011\u003c/span\u003e;Findley \u0026amp; Findley, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Bell \u0026amp; Galzin, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1984\u003c/span\u003e). As coral cover declines, associated fish populations often decrease (Coker et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Coral loss can also influence fish behaviour, for example by increasing aggression as individuals compete for limited resources(Kok et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Pratchett et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2006\u003c/span\u003e;Graham, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2007\u003c/span\u003e) or decreasing aggression as the energy required to hold a territory outweighs the benefits. This is particularly evident in specialist species, such as obligate corallivores, which are especially vulnerable to changes in coral composition (Keith et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Pratchett et al., \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Cole et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2008\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eButterflyfish (family Chaetodontidae) are particularly dependent on healthy coral communities because many species in this family are obligate corallivores (Pratchett et al., \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Habitat degradation and the loss of live coral therefore pose a significant threat to butterflyfish, by reducing resource availability and disrupting biological interactions such as competition and predation (Keith et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Coker et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). Among butterflyfish, aggression is known to vary in response to resource availability (Kok et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). When coral is scarce, aggression may be low because defending a territory is not worth the energetic effort if it does not contain valuable resources; for example, Keith et al (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) found reduced aggression across the Central Indo-Pacific following a mass bleaching event. Conversely, aggression can also be low when coral is very abundant, because resources are readily available and easy to obtain, so defending a territory is not necessary (Peiman \u0026amp; Robinson, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Aggression tends to peak when resources are at an intermediate level\u0026mdash;limited but defensible\u0026mdash;because individuals gain a clear benefit from protecting valuable foraging areas (Peiman \u0026amp; Robinson, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Maher \u0026amp; Lott, \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). As such, butterflyfish behaviour is likely to change in response to changes in coral cover.\u003c/p\u003e \u003cp\u003eIn response to the widespread degradation of coral reefs in recent decades, coral restoration efforts have increased around the world, as a strategy to support reef recovery and maintain ecosystem services (Bayraktarov et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Hein et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Coral reef restoration includes a wide range of different techniques such as rubble stabilisation (Leung et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), coral transplantation (Ferse et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), the use of artificial structures to promote reef accretion (Tanaya et al., \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), and assisted evolution techniques aimed at enhancing coral resilience to both anthropogenic and natural stressors (van Oppen et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). These active reef restoration efforts seek to restore coral cover, physical reef structure, ecological function, and biodiversity (Bostr\u0026ouml;m-Einarsson et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Restoration can produce conditions similar to those of natural coral reef habitats in some cases, but in other cases restored reefs can differ from naturally healthy reefs in aspects such as community composition, structural complexity, and ecological interactions (Vida et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Lange et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Lamont et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eDifferences between natural and restored reefs may influence habitat suitability for corallivores and thus patterns of corallivory (Ladd \u0026amp; Shantz, \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Restored reefs are often dominated by fast-growing coral genera such as \u003cem\u003eAcropora\u003c/em\u003e, due to their rapid growth and colonization rates (Hein, et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2020\u003c/span\u003ea; Lirman et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). \u003cem\u003eAcropora\u003c/em\u003e corals are a preferred food source for some butterflyfish species (Semmler et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). If restored reefs contain different amounts of preferred food sources, butterflyfish behaviour may also differ in restored environments (Hein et al., \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Understanding how butterflyfish respond behaviourally to restored versus natural reef habitats can provide valuable insight into the effectiveness of restoration efforts in replicating natural ecological conditions.\u003c/p\u003e \u003cp\u003eThis study aims to quantify butterflyfish behaviour in both natural and restored coral reef habitats. It focuses on two primary objectives: first, we tested whether there are differences in butterflyfish abundance and aggressive behaviour between natural healthy and restored reefs at one of the world\u0026rsquo;s largest coral restoration (the \u0026lsquo;Mars Assisted Reef Restoration System (MARRS)\u0026rsquo; restoration site in Bontosua Island, South Sulawesi, Indonesia). Second, we compared these behavioural patterns with those observed in other reef systems across the Central Indo-Pacific (Keith et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), in order to contextualise these data within regional ecological variation. By examining the behaviour of obligate corallivorous species, this research contributes to a broader understanding of the ecological impacts of coral reef restoration and provides a more comprehensive framework for evaluating restoration success.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy sites\u003c/h2\u003e \u003cp\u003eThis study compared three coral reef habitat types at Bontosua Island in the Spermonde Archipelago, South Sulawesi, Indonesia: degraded reefs, restored reefs, and naturally healthy reefs.\u003c/p\u003e \u003cp\u003eDegraded reefs in this study had been severely impacted by historical human activities, including boat channel construction (30\u0026ndash;40 years ago), blast fishing (30 years ago), and coral mining for infrastructure (20 years ago) (Williams et al., \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Sawall et al., \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Plass-Johnson et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Such disturbances have damaged the coral framework, resulting in unstable rubble fields and a significant reduction in reef structural complexity. Unconsolidated rubble inhibits coral recruitment and impedes the formation of mature coral colonies (Cameron et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Fox et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2003\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTo address this degradation, the MARRS Coral Reef Restoration Project (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e\u003ca href=\"http://www.buildingcoral.com\" target=\"_blank\"\u003ewww.buildingcoral.com\u003c/a\u003e\u003c/span\u003e\u003cspan address=\"http://www.buildingcoral.com\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) initiated a restoration effort using hexagonal steel structures (Reef Stars) coated with resin and sand (Smith et al., \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). These interconnected structures create a stable substrate for reattaching coral fragments, and restoration on Bontosua Island led to rapid increases in coral cover, fish abundance, and biomass within two to three years (Smith et al., \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Restored reefs in this study had been restored using this method for between one and six years.\u003c/p\u003e \u003cp\u003eHealthy reefs in this study had historically remained in good condition, supporting thriving coral communities and showing no evidence of the historical disturbance that affected degraded reefs. The total study area included 29 reef sites, comprising 3 degraded sites, 7 healthy sites, and 19 restored sites. The restored sites were further classified into two age categories: 11 old restored sites (5\u0026ndash;7 years post-restoration) and 8 young restored sites (1\u0026ndash;3 years post-restoration).\u003c/p\u003e \u003cp\u003eIn addition to comparing habitat conditions at Bontosua Island, we also compared these data to other sites around the Central Indo-Pacific, using data from Keith et al. (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Data from Bali (Indonesia), Christmas Island (Indian Ocean, Australia), Iriomote (Japan), and Luzon (Philippines) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) were sourced from Keith et al. (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) based on their representation of a range of natural healthy reef conditions across the Central Indo-Pacific region. In order to represent naturally healthy conditions, only data from before the mass bleaching event studied in Keith et al. (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) was used. This regional comparison enabled us to contextualize patterns observed in our location (Bontosua) within a wider biogeographical framework.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eButterflyfish abundance and diversity surveys\u003c/h3\u003e\n\u003cp\u003eButterflyfish abundance and species diversity were quantified using underwater visual census (UVC) surveys at each study site. Surveys were conducted along transects that were 5m wide, with the transect length adjusted based on the size of the surveyed site. Transects were 50 m in length where habitat allowed, or 20 m, 33 m, and 40 m in length in smaller restoration areas. Transects were positioned parallel to the reef crest, targeting reef zones most frequently used by corallivorous butterflyfish. After transects were laid, an acclimation period (~\u0026thinsp;1 min) was observed, during which the observer ensured that butterflyfish were behaving naturally and not disturbed by observer presence. Following this acclimation period, butterflyfish abundance was quantified by counting all individuals within 2.5 m of either side of the transect.\u003c/p\u003e\n\u003ch3\u003eButterflyfish behaviour\u003c/h3\u003e\n\u003cp\u003eWe quantified the frequency of interactions between butterflyfish individuals, and the aggression levels present within each interaction, following the same methods used in Keith et al. (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). We observed interactions between focal butterflyfish individuals and other \u003cem\u003eChaetodon\u003c/em\u003e species, including both conspecifics and heterospecifics. Data were collected using SCUBA, and observers followed a modified U-shaped swimming pattern at each site to minimize the likelihood of repeatedly observing the same individuals. Behavioural observations were only carried out in healthy and restored habitat but not in degraded habitat, because there were not enough butterflyfish present on degraded reefs to ensure adequate sample size. We carried out behavioural observations in the same area as the abundance surveys, immediately after abundance surveys were carried out.\u003c/p\u003e \u003cp\u003eThe first butterflyfish encountered from any species was designated as the focal individual. Many \u003cem\u003eChaetodon\u003c/em\u003e species occur in pairs, and since no aggression was observed between partners, only one fish per pair was observed. Focal individuals were followed at a distance of 2\u0026ndash;4 m for five minutes each. Encounters were recorded when another butterflyfish individual came within 1 m of the focal individual. At this distance, it was assumed that both individuals were aware of each other and had the potential to interact. Interactions were categorized as passive if no behavioural response could be observed, or aggressive if one or both individuals displayed signalling (dorsal fin raised, head down) or chasing behaviour. For each observation the time of day, water temperature, focal fish species, interacting fish species, distance at the point of encounter, and the size (cm) of both the focal and interacting fish were also recorded.\u003c/p\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eButterflyfish abundance was modelled as a function of restoration age using a linear mixed model (LMM). The response variable was butterflyfish abundance (individuals m⁻\u0026sup2;), with restoration age in years included as a fixed effect, and site as a random intercept to account for spatial non-independence. Degraded reefs were included with a restoration age of zero to provide a baseline for comparison, while healthy reefs were excluded from the model and compared visually on a graph as reference sites. Model residuals were inspected to goodness-of-fit.\u003c/p\u003e \u003cp\u003eLive coral cover was modelled as a function of restoration age using the same fixed and random effect structure with butterflyfish abundance model, but using a beta error distribution with a logit link in a generalised linear mixed model (GLMM).\u003c/p\u003e \u003cp\u003eButterflyfish abundance at each location was analyzed using non-parametric methods because the data violated assumptions of normality and homogeneity of variance, as assessed by the Shapiro\u0026ndash;Wilk test. Differences in butterflyfish abundance per square meter among location were first evaluated using a Kruskal\u0026ndash;Wallis rank-sum test. When significant differences were detected, pairwise Wilcoxon rank-sum tests were performed to determine which location pairs differed, with p-values adjusted using the Bonferroni correction.\u003c/p\u003e \u003cp\u003eThe degraded site in Bontosua exhibited very low butterflyfish abundance and species diversity, with a small sample size (n\u0026thinsp;=\u0026thinsp;6). As a result, statistical comparisons for this location could not be conducted due to insufficient statistical power.\u003c/p\u003e \u003cp\u003eButterflyfish species diversity was quantified using the Shannon\u0026ndash;Wiener index (H\u0026prime;), which incorporates both species richness and evenness (Kunakh et al., 2023; Hurlbert, 1971). Differences in diversity among locations were tested using the non-parametric Kruskal\u0026ndash;Wallis test. Significant results were followed by pairwise Wilcoxon rank-sum tests with Bonferroni corrections to identify differences between specific site pairs.\u003c/p\u003e \u003cp\u003eTo test for species dominance in the butterflyfish community, the abundance of the most common butterflyfish species was compared to the total abundance of all other butterflyfish species at each site. At each site, paired Wilcoxon signed-rank tests were used to compare the abundance of the dominant species against all other species.\u003c/p\u003e \u003cp\u003eThe frequency of butterflyfish interactions was analysed using generalized linear mixed-effects models (GLMMs) with a negative binomial distribution to account for overdispersed count data. Aggression was quantified as the proportion of interactions that were aggressive (bounded between 0 and 1) and analysed using GLMMs with a beta distribution and logit link. The response variable for the interaction frequency model was the number of interactions per focal fish, whereas for the aggression model it was the proportion of interactions classified as aggressive for each focal fish.\u003c/p\u003e \u003cp\u003eIn both models, interaction type (conspecific vs. heterospecific) and location (Bontosua Restored Old, Bontosua Restored Young, Bontosua Healthy, Bali, Christmas Island, Iriomote, and Luzon) were included as fixed effects. The two models differed in their random-effects structure. For interaction frequency, site was included as a random intercept to account for non-independence of observations within sites. For aggression proportion, both site and focal species were included as random intercepts to account for variation among sites and species-level differences in aggressive behaviour.\u003c/p\u003e \u003cp\u003eAll analyses were conducted in R v.4.4.0 (R Core Team, 2024), using the packages \u003cem\u003eglmmTMB\u003c/em\u003e package (Brooks et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), \u003cem\u003eemmeans\u003c/em\u003e (Lenth, \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), DHARMa(Hartig, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) and ggplot2 (Wickham, \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eButterflyfish abundance in response to time since restoration\u003c/h2\u003e \u003cp\u003eThe abundance of butterflyfishes in restored sites at Bontosua Island increased with time since restoration, from 0.02 individuals m⁻\u0026sup2; at one year old site to 0.12\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 individuals m⁻\u0026sup2; at seven years old sites (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea). This overall pattern of increase was a significant positive trend (LMM: estimate\u0026thinsp;=\u0026thinsp;0.015\u0026thinsp;\u0026plusmn;\u0026thinsp;0.004 SE, \u003cem\u003ez\u003c/em\u003e\u0026thinsp;=\u0026thinsp;3.77, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001; Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e), corresponding to an annual increase of ~\u0026thinsp;0.015 individuals m⁻\u0026sup2;. Post-hoc contrasts between individual years showed that abundance at six-year-old reefs was significantly higher than in degraded reefs, but reefs restored younger than that age (1\u0026ndash;5 years old) did not differ significantly from degraded reefs (Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). Seven-year reefs also exhibited higher abundances than degraded reefs, and were comparable with the healthy sites, though this effect was not statistically significant (Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). Together, these results indicate a clear positive trajectory in butterflyfish abundance with reef age, with significant recovery evident from approximately six years post-restoration.\u003c/p\u003e \u003cp\u003eThis positive trend in butterflyfish abundance aligned with a corresponding increase in coral cover. Live coral cover also increased strongly with restoration age (GLMM, β\u0026thinsp;=\u0026thinsp;0.377\u0026thinsp;\u0026plusmn;\u0026thinsp;0.057 SE; \u003cem\u003ez\u003c/em\u003e\u0026thinsp;=\u0026thinsp;6.65, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Coral cover increased from 2.8% at one-year restored sites to 57.4% \u0026plusmn; 6.1% at seven-year restored sites, approaching levels found on healthy reefs (54.5% \u0026plusmn; 4.4%) and exceeding those of degraded sites (10.3% \u0026plusmn; 3.9%) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eb). Pairwise contrasts indicated that there was no significant difference between degraded and 1-year old restored reefs, but restored reefs of 2 years or older all had significantly higher coral cover than degraded reefs, with effect sizes strengthening with restoration age (Table S2).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eComparing butterflyfish populations at Bontosua Island with other locations in the Central Indo-Pacific\u003c/h3\u003e\n\u003cp\u003eButterflyfish abundance (individuals per m\u0026sup2;) differed significantly among habitats across the Central Indo-Pacific (Kruskal\u0026ndash;Wallis test: χ\u0026sup2; = 47.13, df\u0026thinsp;=\u0026thinsp;6, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001; Table S3). In particular, abundance at Bontosua Healthy and Bontosua Restored (Old) were significantly higher than at other location around the Central Indo-Pacific (Wilcoxon rank-sum test, W\u0026thinsp;=\u0026thinsp;2716, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001; Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eA). But not different significant with Bontosua Restored (Young).\u003c/p\u003e \u003cp\u003ePost-hoc pairwise Wilcoxon rank-sum test (Bonferroni-corrected) showed that Bontosua Healthy sites had significantly higher butterflyfish abundance than Christmas Island (W\u0026thinsp;=\u0026thinsp;200, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01) and Iriomote (W\u0026thinsp;=\u0026thinsp;253, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), but did not differ significantly from Bali, Luzon, or either restored habitat at Bontosua.\u003c/p\u003e \u003cp\u003eAmong restored habitats, Bontosua Restored (Old) sites exhibited significantly greater butterflyfish abundance than Christmas Island (W\u0026thinsp;=\u0026thinsp;436, p\u0026thinsp;\u0026lt;\u0026thinsp;0.001), Iriomote (W\u0026thinsp;=\u0026thinsp;558, p\u0026thinsp;\u0026lt;\u0026thinsp;0.001), and Luzon (W\u0026thinsp;=\u0026thinsp;530, p\u0026thinsp;\u0026lt;\u0026thinsp;0.01). Bontosua Restored (Young) sites did not differ significantly from Bali, Christmas Island, Iriomote, or Luzon, nor from Bontosua Healthy or Bontosua Restored (Old) sites. No significant differences were detected between Bali and any Bontosua habitat (Healthy, Restored (Old), or Restored (Young)) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eA).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eButterflyfish species diversity varied significantly across locations (Kruskal\u0026ndash;Wallis test: χ\u0026sup2; = 34.7, df\u0026thinsp;=\u0026thinsp;6, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001; Table S4). Post-hoc pairwise comparisons using Bonferroni-adjusted Wilcoxon rank-sum tests revealed strong spatial differences in species diversity among locations. Diversity at Bontosua Restored (Old) sites was significantly lower than at both Bali (W\u0026thinsp;=\u0026thinsp;292, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) and Luzon (W\u0026thinsp;=\u0026thinsp;115, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), but did not differ from Christmas Island and Iriomote. A similar pattern was observed at the Bontosua Healthy sites, where diversity was significantly lower than at Bali (W\u0026thinsp;=\u0026thinsp;126, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) and Luzon (W\u0026thinsp;=\u0026thinsp;41, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05), while showing no significant differences from Christmas Island, Iriomote, and the Bontosua Restored sites. Overall, both restored and healthy reefs at Bontosua exhibited reduced species diversity relative to Bali and Luzon, but were broadly similar to Christmas Island and Iriomote.\u003c/p\u003e\n\u003ch3\u003eDominant species of butterflyfish\u003c/h3\u003e\n\u003cp\u003eSignificant differences in butterflyfish species composition were observed among locations (Kruskal-wallis test, χ\u0026sup2; = 27.1, df\u0026thinsp;=\u0026thinsp;6, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001, Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e, Table S5). At Bontosua Restored (Old) sites, the most common species was significantly more abundant than all other species combined (Wilcoxon test, W\u0026thinsp;=\u0026thinsp;470, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001, r\u0026thinsp;=\u0026thinsp;0.467), and a similar dominance pattern was observed at the Bontosua Healthy sites (W\u0026thinsp;=\u0026thinsp;64, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001, r\u0026thinsp;=\u0026thinsp;0.155). In contrast, Bontosua Restored (Young) showed no significant dominance by any species.\u003c/p\u003e \u003cp\u003eIn contrast, the other locations exhibited a more even species composition, with the abundance of the dominant species either less than, or not significantly different from, the combined abundance of all other species (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). There were significantly higher abundances of other species relative to the most common species at Bali (W\u0026thinsp;=\u0026thinsp;41, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01, \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.124), Christmas Island (W\u0026thinsp;=\u0026thinsp;175, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05, \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.155), and Luzon (W\u0026thinsp;=\u0026thinsp;413, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05, \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.111). At Iriomote, there was no significant difference between the abundance of the most common species and the abundance of all other species (Wilcoxon signed-rank test, W\u0026thinsp;=\u0026thinsp;623, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.312, \u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eButterflyfish behaviour\u003c/h2\u003e \u003cp\u003eAcross all locations except for Bontosua, heterospecific interactions occurred more frequently than conspecific interactions (GLMM, estimate\u0026thinsp;=\u0026thinsp;1.06\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09, z\u0026thinsp;=\u0026thinsp;12.15, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001, Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ea, Table S6). In Bontosua, conspecific interactions occurred more frequently than heterospecific interactions. This effect was strongest at Bontosua Healthy sites (GLMM, estimate\u0026thinsp;=\u0026thinsp;2.37\u0026thinsp;\u0026plusmn;\u0026thinsp;0.40, z\u0026thinsp;=\u0026thinsp;5.95, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), while at Bontosua Restored (Old) (GLMM, estimate\u0026thinsp;=\u0026thinsp;0.68\u0026thinsp;\u0026plusmn;\u0026thinsp;0.28, z\u0026thinsp;=\u0026thinsp;2.45, p\u0026thinsp;=\u0026thinsp;0.014) and Bontosua Restored (Young) showed no significant difference between interaction types (GLMM, estimate\u0026thinsp;=\u0026thinsp;0.16\u0026thinsp;\u0026plusmn;\u0026thinsp;0.25, z\u0026thinsp;=\u0026thinsp;0.62, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.538).\u003c/p\u003e \u003cp\u003eIn contrast, the other locations\u0026mdash;Bali, Christmas Island, Iriomote, and Luzon, exhibited significantly higher frequencies of heterospecific interactions. Pairwise contrasts revealed significant effects for Bali (GLMM, estimate\u0026thinsp;=\u0026thinsp;\u0026minus;\u0026thinsp;2.71\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13, \u003cem\u003ez\u003c/em\u003e\u0026thinsp;=\u0026thinsp;\u0026minus;\u0026thinsp;21.26, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), Christmas Island (GLMM, estimate\u0026thinsp;=\u0026thinsp;\u0026minus;\u0026thinsp;2.66\u0026thinsp;\u0026plusmn;\u0026thinsp;0.14, \u003cem\u003ez\u003c/em\u003e\u0026thinsp;=\u0026thinsp;\u0026minus;\u0026thinsp;19.50, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), Iriomote (GLMM, estimate\u0026thinsp;=\u0026thinsp;\u0026minus;\u0026thinsp;2.76\u0026thinsp;\u0026plusmn;\u0026thinsp;0.14, \u003cem\u003ez\u003c/em\u003e\u0026thinsp;=\u0026thinsp;\u0026minus;\u0026thinsp;20.34, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), and Luzon (GLMM, estimate\u0026thinsp;=\u0026thinsp;\u0026minus;\u0026thinsp;2.50\u0026thinsp;\u0026plusmn;\u0026thinsp;0.14, \u003cem\u003ez\u003c/em\u003e\u0026thinsp;=\u0026thinsp;\u0026minus;\u0026thinsp;18.22, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), indicating that butterflyfish tended to interact more often with individuals of other species than with conspecifics\u003c/p\u003e \u003cp\u003eConspecific interactions tended to be slightly more aggressive than heterospecific interactions across locations, although this overall effect was not statistically significant (GLMM: β = \u0026minus;\u0026thinsp;0.11\u0026thinsp;\u0026plusmn;\u0026thinsp;0.52, z = \u0026minus;\u0026thinsp;0.21, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.83). However, the model revealed significant interactive effects of interaction type and location on aggression frequency (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eb). Post-hoc comparisons showed that conspecific aggression was significantly higher than heterospecific aggression in Bali (β\u0026thinsp;=\u0026thinsp;2.64\u0026thinsp;\u0026plusmn;\u0026thinsp;0.35, z\u0026thinsp;=\u0026thinsp;7.57, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), Christmas Island (β\u0026thinsp;=\u0026thinsp;1.25\u0026thinsp;\u0026plusmn;\u0026thinsp;0.37, z\u0026thinsp;=\u0026thinsp;3.35, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.005), Iriomote (β\u0026thinsp;=\u0026thinsp;3.13\u0026thinsp;\u0026plusmn;\u0026thinsp;0.39, z\u0026thinsp;=\u0026thinsp;8.08, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), and Luzon (β\u0026thinsp;=\u0026thinsp;1.79\u0026thinsp;\u0026plusmn;\u0026thinsp;0.35, z\u0026thinsp;=\u0026thinsp;5.08, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). In contrast, no significant differences between conspecific and heterospecific aggression were detected at the Bontosua sites, including Bontosua Healthy (β\u0026thinsp;=\u0026thinsp;0.12\u0026thinsp;\u0026plusmn;\u0026thinsp;0.53, z\u0026thinsp;=\u0026thinsp;0.23, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.821), Bontosua Restored (Old) (β\u0026thinsp;=\u0026thinsp;1.11\u0026thinsp;\u0026plusmn;\u0026thinsp;0.52, z\u0026thinsp;=\u0026thinsp;0.21, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.831), and Bontosua Restored (Young) (β = \u0026minus;\u0026thinsp;0.10\u0026thinsp;\u0026plusmn;\u0026thinsp;0.37, z = \u0026minus;\u0026thinsp;0.27, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.786).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eGeneral findings\u003c/h2\u003e \u003cp\u003eThis study focused on the behavioural ecology of butterflyfish (family Chaetodontidae) to evaluate the ecological outcomes of coral reef restoration on Bontosua Island, with comparative insight drawn from a broader regional context encompassing Bali (Indonesia), Christmas Island (Indian Ocean, Australia), Iriomote (Japan), and Luzon (Philippines). At the restored reefs at Bontosua, butterflyfish abundance increased progressively with restoration age and increasing coral cover. This pattern aligns with previous findings that link the presence and population growth of butterflyfish to live coral cover (Pratchett et al., \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). The observed population growth likely reflects the increased availability of coral in the restored sites. Across both healthy and restored sites, particularly the older restored sites, butterflyfish communities on Bontosua Island exhibited patterns distinct from those in other locations. While both healthy and older restored reefs on Bontosua supported substantially higher butterflyfish abundances, their species diversity was markedly lower. This reduced diversity was driven by the strong dominance of a single species, \u003cem\u003eChaetodon octofasciatus\u003c/em\u003e, which comprised the majority of the butterflyfish community at Bontosua.\u003c/p\u003e \u003cp\u003eBehavioural patterns were also distinctive at Bontosua Island compared to other locations around the Central Indo-Pacific. At Bontosua, aggressive interactions were infrequent, with levels of aggression consistently low across both restored and healthy reefs. This pattern may be attributable to the dominance of a single species (\u003cem\u003eC. octofasciatus\u003c/em\u003e), resulting in reduced interspecific competition and relaxed competition among conspecifics. The consistently low aggression observed in \u003cem\u003eC. octofasciatus\u003c/em\u003e may reflect weak territoriality or reliance on concealment rather than overt defence. In contrast, behavioural data from other locations revealed higher levels of aggression, possibly due to greater species diversity and increased interspecific competition at those sites.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eAbundance and behavioural patterns in restored reefs and healthy reefs in Bontosua Island\u003c/h2\u003e \u003cp\u003eRestoration efforts in Bontosua have significantly increased live coral cover and small-scale structural complexity of reef habitats compared to previously degraded conditions (Vida et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). These improvements are likely to be the driver of the observed increase in corallivorous fish (Santoso et al., \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Darling et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). The three-dimensional framework of live corals provides microhabitat for shelter, territorial behaviour, and spawning activities (Coker et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), and live coral serves as the primary food source for corallivorous butterflyfish (Cole et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Pratchett, \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Rotjan \u0026amp; Dimond, \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). While coral cover recovered substantially within the first three years of restoration, butterflyfish abundance required approximately five to seven years to reach levels comparable to healthy reefs, indicating a lag in the biological response of fish communities to habitat improvement. Similar lagged responses have been documented by (Hein et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), who reported gradual increases in fish abundance over several years following coral restoration sites, even after coral cover had already recovered. Comprehensive reviews also suggest that while coral cover may rebound from degradation within three years, full recovery of fish communities, particularly among specialist species, can take five to fifteen years (McClanahan et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2007\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eDespite the increase in abundance, butterflyfish species diversity remained low at all sites at Bontosua. Communities in Bontosua were dominated by a single species, \u003cem\u003eC. octofasciatus\u003c/em\u003e. This pattern was also observed in healthy reefs, suggesting that such single-species dominance is not a product of restoration, but a characteristic of the reefs around this island. Comparable findings were reported from other islands within the Spermonde Islands, where \u003cem\u003eC. octofasciatus\u003c/em\u003e emerged as the dominant species among 21 Chaetodontidae species surveyed across 13 locations (Putra et al., \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). \u003cem\u003eC. octofasciatus\u003c/em\u003e exhibits high resilience to fishing pressure, with a population doubling time of less than 15 months pressure making it a suitable indicator for monitoring rapid habitat changes even under conditions of intense exploitation (Froese \u0026amp; Pauly, 2010; Madduppa et al., \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). These ecological traits are particularly relevant to the Spermonde region, including Bontosua Island, where destructive fishing practices continue to exert substantial pressure on reef ecosystems. Additional studies have demonstrated \u003cem\u003eC.octofasciatus\u003c/em\u003e tolerance to elevated water turbidity (Bramasta et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) and its frequent occurrence in habitats dominated by rubble or dead coral substrates (Songploy et al., \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAs the dominant species, \u003cem\u003eC. octofasciatus\u003c/em\u003e likely underpins the consistently low levels of aggressive behaviour observed at Bontosua Island. This numerical dominance may reduce aggressive encounters between different species, resulting in consistently low aggression levels across both restored and healthy reefs. Although this species is an obligate corallivore (Cole et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Allen et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Michael, \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2004\u003c/span\u003e), previous studies have demonstrated that it exhibits relatively weak territorial behaviour compared to other obligate corallivores such as \u003cem\u003eC. baronessa\u003c/em\u003e and \u003cem\u003eC. trifascialis\u003c/em\u003e (Blowes et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2017\u003c/span\u003e;Pratchett \u0026amp; Berumen, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Field observations from Redang Island, Malaysia, indicate that \u003cem\u003eC. octofasciatus\u003c/em\u003e typically occupies relatively small territories and frequently seeks refuge among coral branches (Ghaffar et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). Such patterns suggest that \u003cem\u003eC. octofasciatus\u003c/em\u003e exhibits weak territoriality and relies more on concealment than aggressive defence. This aligns with our findings, in which the proportion of aggressive encounters involving \u003cem\u003eC. octofasciatus\u003c/em\u003e was substantially lower than that observed for other species (Table S8). Moreover, the very low number of aggressive encounters recorded overall at Bontosua Island further supports the conclusion that \u003cem\u003eC. octofasciatus\u003c/em\u003e exhibits low levels of aggression. Hence, these findings indicate that \u003cem\u003eC. octofasciatus\u003c/em\u003e may be a non- or weakly territorial species.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eReefs and butterflyfish behaviour patterns in other locations across the Central Indo-Pacific\u003c/h2\u003e \u003cp\u003eBiogeographic factors play an important role in shaping butterflyfish community structure and behaviour, in some regions, species diversity remains low despite high coral cover, reflecting ecological or historical constraints (Findley \u0026amp; Findley, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). The ecological similarity observed between restored and healthy habitats in Bontosua suggests that regional (biogeographic) conditions may exert a stronger influence on community structure than the restoration status of individual reefs. This is supported by the strong correlation between coral cover and fish abundance at Bontosua compared to other locations in the Central Indo-Pacific. These findings indicate that the response of butterflyfish to coral cover is location-dependent and may be influenced by local ecological factors such as habitat complexity, interspecific interactions, or environmental stressors, as well as by broader biogeographic constraints.\u003c/p\u003e \u003cp\u003eBoth restored and healthy reefs in Bontosua displayed butterflyfish abundance and social interaction patterns distinct from those reported in other parts of the Central Indo-Pacific. Lower species richness and reduced aggression observed in Bontosua may be influenced by multiple factors, including latitudinal gradients (Siqueira et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Gonz\u0026aacute;lez-Barrios et al., \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), limited regional larval supply (van der Meer et al., \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), historical habitat degradation (Jackson et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Wilson et al., \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), or reduced connectivity with more diverse source populations (Hobbs \u0026amp; Srinivasan, \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Although Bontosua is located within a globally recognized marine biodiversity hotspot, its low-diversity butterflyfish community structure may reflect regional-level biogeographic isolation or ongoing recovery from past disturbances. In particular, historical degradation events may have led to local extinctions or demographic bottlenecks that reduced species richness, particularly among less resilient or weakly dispersing taxa.\u003c/p\u003e \u003c/div\u003e"},{"header":"Conclusion","content":"\u003cp\u003eThis study demonstrates that both restored, particularly older restoration sites and healthy reef habitats on Bontosua Island exhibit similar characteristics in their butterflyfish populations, including high butterflyfish abundance, low species richness, and consistent behavioural patterns marked by low aggression. The observed increase in butterflyfish abundance over time in restored areas suggests a positive response to restoration efforts, likely driven by improvements in coral cover and structural complexity.\u003c/p\u003e \u003cp\u003eCompared to other sites in the Central Indo-Pacific, Bontosua stands out for its unusually high butterflyfish abundance and low species diversity, a pattern largely attributed to the dominance of \u003cem\u003eC. octofasciatus\u003c/em\u003e. This low-diversity community structure, present in both healthy and restored reefs, may reflect limited regional larval supply, historical habitat degradation, or reduced connectivity to more diverse source populations. The dominance of a single species may also contribute to reduced interspecific competition, as \u003cem\u003eC. octofasciatus\u003c/em\u003e is thought to be non- or weakly territorial.\u003c/p\u003e \u003cp\u003eOverall, these findings suggest that reef restoration at Bontosua has successfully supported the recovery of butterflyfish populations to levels comparable with nearby healthy reefs. However, the patterns of aggressive behaviour appear to be shaped more strongly by broader regional ecological conditions than by the local effects of restoration alone.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003e**Sera Maserati** : conceptualization, data curation, formal analysis, investigation, methodology, visualization, writing \u0026ndash; original draft preparation and editing. **Timothy A.C. Lamont:** conceptualization, methodology, investigation, funding acquisition, supervision, writing \u0026ndash; review \u0026amp;amp; editing, project administration. **Jane C.S. Sudrajat** : investigation. **Andi Tanri Abeng** : investigation. **Permas B. Maulana:** investigation. **Sally A. Keith:** methodology, investigation. writing \u0026ndash; review. **Tries B. Razak** : conceptualization, methodology, supervision, funding acquisition, writing \u0026ndash; review \u0026amp;amp; editing, project administration.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eSurvey data for this study were collected as part of the monitoring program of the Mars Coral Reef Restoration Project, in collaboration with Universitas Hasanuddin. We thank L. Damayanti, P. Mansell, and members of the Mars Sustainable Solutions monitoring team (M. E. Prasetya, P. B. Maulana, A. Hamka, A. Dwiyanto, A. M. A. Pratama, A. T. Abeng, Irwan, R. Madjid, E. Agiel, and Suandar) for their invaluable support with field logistics and monitoring activities. We are also grateful to the Department of Marine Affairs and Fisheries of South Sulawesi Province, the Government Offices of Kabupaten Pangkep, Pulau Bontosua, and Pulau Badi, as well as the community of Pulau Bontosua, for their support and cooperation. We thank S. Keith for helpful discussions and for contributions to data collection outside Indonesia. Fieldwork in Indonesia was conducted under a national research permit issued by BRIN (No. 108/SIP/IV/FR/2/2023), jointly held by T.A.C.L. as lead foreign researcher and T.B.R. as lead Indonesian host researcher, with ethical approval granted by BRIN and Lancaster University. We thank Prof. J. Jompa and Prof. R. A. Rappe at Universitas Hasanuddin for logistical assistance with permit and visa applications. Funding was provided by a research fellowship from The Royal Commission for the Exhibition of 1851 (awarded to T.A.C.L.), an international traveling fellowship from the Fisheries Society of the British Isles (awarded to T.A.C.L. and T.B.R.), and a Pew Fellowship in Marine Conservation (awarded to T.B.R.).\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eThe datasets generated and/or analysed during the current study will be deposited in a public repository upon acceptance of the manuscript.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAllen, G. R., Steene, R., \u0026amp; Allen, M. (1998). A Guide to Angelfishes \u0026amp; Butterflyfishes. \u003cem\u003eOdyssey Publishing, Perth\u003c/em\u003e, 250.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBayraktarov, E., Banaszak, A. T., Maya, P. M., Kleypas, J., Arias-Gonzalez, J. 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Exploitation and habitat degradation as agents of change within coral reef fish communities. \u003cem\u003eGlobal Change Biology\u003c/em\u003e, \u003cem\u003e14\u003c/em\u003e(12), 2796\u0026ndash;2809. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1365-2486.2008.01696.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1365-2486.2008.01696.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"butterflyfish, fish behaviour, coral restoration, coral rehabilitation, Indonesia, Central Indo-Pacific","lastPublishedDoi":"10.21203/rs.3.rs-8883345/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8883345/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eCoral restoration is increasingly applied to combat reef habitat degradation, yet its impacts on fish populations and behavioural dynamics remain underexplored. This study examines how coral restoration influences the population density and behaviour of butterflyfish (Chaetodontidae) \u0026ndash; obligate corallivores and recognized bioindicators of reef health. We measured the abundance, diversity and aggressive territorial behaviour of butterflyfish at one of the world\u0026rsquo;s largest coral restoration projects (Bontosua Island, South Sulawesi, Indonesia), comparing naturally healthy reefs with those that had been restored after historic damage from dynamite fishing. Additionally, we compared these data with equivalent metrics from previous studies at other natural reefs in the Central Indo-Pacific region, to contextualise our data within a wider ecological framework. At Bontosua, coral restoration led to significant recovery of butterflyfish abundance within six years, with both restored and healthy reefs exhibiting high butterflyfish abundance but low species diversity and consistently low rates of aggressive interactions. On both natural and restored reefs, butterflyfish communities at Bontosua were dominated by a single species, \u003cem\u003eChaetodon octofasciatus\u003c/em\u003e \u0026ndash; with low levels of both inter- and intra-specific aggressive behaviour. This contrasts with more diverse and competitive butterflyfish communities in other locations around the Central Indo-Pacific. We conclude that butterflyfish behavioural patterns were more sensitive to broad-scale regional variation than to restoration outcomes at this site. Butterflyfish behaviour may serve as a more effective indicator of regional ecosystem conditions than of localized restoration success.\u003c/p\u003e","manuscriptTitle":"Ecological and behavioural responses of butterflyfishes to coral reef restoration","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-04-28 19:25:01","doi":"10.21203/rs.3.rs-8883345/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"c367943f-aa70-4cae-a610-60ff1523365e","owner":[],"postedDate":"April 28th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2026-04-28T19:25:09+00:00","versionOfRecord":[],"versionCreatedAt":"2026-04-28 19:25:01","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8883345","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8883345","identity":"rs-8883345","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}