Evidence of adaptive plasticity in the coral Echinopora spp. to different turbidity regimes

Evidence of adaptive plasticity in the coral Echinopora spp. to different turbidity regimes | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Evidence of adaptive plasticity in the coral Echinopora spp. to different turbidity regimes Muhammad Irsyad Abiyusfi Ghafari, Mohd Hanafi Idris, Ezmahamrul Afreen Awalludin, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7040520/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 06 Mar, 2026 Read the published version in Marine Biology → Version 1 posted 5 You are reading this latest preprint version Abstract Turbid reefs, once seen as marginal, are now considered potential refuges for laminar-shaped corals under climate stress. This study examines micro-morphological plasticity in two foliose coral species: Echinopora lamellosa and E. pacificus, across turbid and non-turbid sites in East Lombok, Indonesia. A total of 137 specimens from different depths (2 m and 4 m) and turbidity levels were analysed using ten diagnostic morphological traits. Principal component analysis explained >97% of the total morphological variance and revealed clear differentiation by site and depth. E. pacificus showed complete separation between populations, while E. lamellosa displayed partial overlap—suggesting environmentally driven plasticity. Deep colonies in non-turbid sites resembled shallow colonies in turbid areas, supporting the idea of adaptive morphometric responses rather than fixed traits. Outgroup validation using a geographically distinct population of the same species confirmed the observed patterns, strengthening the evidence for environment-related variation. The patterns observed may involve phenotypic plasticity or epigenetic changes, which warrants further investigation. These findings highlight the plastic ability of foliose corals to exploit either shallow or turbid habitats through their diverse morphological traits, underscoring their resilience and potential significance for future reef conservation, restoration and management efforts amid turbidity-driven climate change. Coral resilience foliose corals morphological plasticity sediment stress turbidity Figures Figure 1 Figure 2 Figure 3 Figure 4 INTRODUCTION Despite that the sediment has long been considered as among the most significant stressors to coral reef systems, increasing current evidence suggests that turbid waters might serve as an important reef conservation hotspot and corals exposed to enhanced turbidity could be more resilient to climate change impacts (Soares et al. 2019 ; Zweifler et al. 2021 ; Carlson et al. 2022 ). Theoretically, turbidity provides shelter for coral from two current major bleaching factors: the excessive UV exposure (Morgan et al. 2017 ) and the elevated sea water temperature (Löptien and Meier 2011 ; Oxenford and Vallès 2016 ), while at the same time it also holds detrimental potential to suffocate corals to bleach (Erftemeijer et al. 2012 ; Jones et al. 2020 ). It has led to disagreement among scientists on the actual effect of turbidity on coral reefs (see Sully and van Woesik ( 2020 ); Zweifler et al. ( 2021 ); and Lucas et al. ( 2023 )). Turbid coral reefs remain understudied due to the technical limitations associated with underwater visibility and imaging in sediment-rich environments. Increased turbidity and sedimentation pose varying levels of stress on different coral species, with some proving more sensitive than others. Most corals rely on morphological adaptations to cope with sediment settling on their surfaces. Among these, branching corals tend to accumulate the least sediment due to their complex three-dimensional structures, which effectively reduce the surface area where sediments can settle (Doszpot et al. 2019 ; Jones et al. 2019 ). In contrast, foliose corals, characterized by their broad, layered morphologies and horizontal surfaces, are more prone to sediment accumulation. Studies by Duckworth et al. ( 2017 ) revealed that foliose-shaped corals generally show the highest surface sediment cover under elevated sedimentation levels, making them particularly vulnerable to stress. This susceptibility has significant consequences. Foliose corals often suffer the greatest losses among coral morphologies due to sediment-induced anoxia, where trapped sediment blocks light and gas exchange, leading to tissue death (Fabricius et al. 2006 ; Duckworth et al. 2017 ; Jones et al. 2019 ). However, despite their apparent vulnerability, some foliose corals have shown unexpected tolerance to sedimentation. Some populations have been reported to persist under turbid conditions, including sediment-laden runoff and severe sedimentation stress near river mouth (Bachtiar and Hadi 2019 ; Guest et al. 2016 ; Syahrir et al. 2018 ). A study by Ghafari et al. ( 2024 ) found that fine-scale morphological variation—particularly in corallite structure—may play a key role in enabling foliose corals, such as those from the genus Echinopora , to survive in turbid environments. Here, our observations on foliose coral species suggest a degree of morphological plasticity that may help corals with this lifeform withstand varying turbidity levels. This study proposes that morphological plasticity particularly within multi-morphotypic coral species may be a critical adaptive mechanism enabling survival in turbid habitats previously deemed unsuitable due to sedimentation stress, possibly allowing them to exploit a wide range of turbid environment. MATERIALS AND METHODS Sampling Sites Four sampling sites were located on reef areas in East Lombok, Indonesia, and were selected to represent distinct turbidity conditions (Fig. 1 A). Turbidity level classification was based on quantitative assessment by Ghafari et al. ( 2024 ), resulting in the identification of non-turbid sites, i.e., ST1 and ST4 (TSS ± SD = 14.32 ± 1.61 ml L − 1 , visibility ± SD = 8.49 ± 0.45 m) and turbid sites, i.e., ST2 and ST3 (TSS ± SD = 29.05 ± 0.76 ml L − 1 , visibility ± SD = 4.35 ± 0.70 m). At each site, coral specimens were sampled at two depth regimes: shallow (2 m) and deep (4 m), to capture variation related to both water clarity and depth. Sampling, Data Collection and Processing The sampling process collected 114 specimens of Echinopora lamellosa and 23 specimens of E. pacificus (Fig. 1 B-C). Specimens taken from a depth of 2 m at each station are referred to as the ‘shallow population’, whereas those taken from 4 m depth are termed the ‘deep population’. Despite the small depth differences in East Lombok, Ghafari et al. ( 2024 ) reported the resulting turbidity levels vary significantly, potentially affecting the morphological adaptations of the coral colonies. Data processing was conducted on ten measurable and/or countable micro-morphological characters of each observed specimen, i.e., corallite area (CA), minimum (CD-min) and maximum (CD-max) diameter of corallite, average corallite diameter (CD-rate), corallite spacing (ICD), mouth-disk diameter (MD), exsert septal spacing (IES), polyp density (PD), polyp height (PH), and skeleton thickness (SkT), following the protocol by Ghafari et al. ( 2022 ). Measurement of PH and SkT required microscopic inspection, which was carried out by making cross-sectional cuts, as described by Ghafari et al. ( 2024 ). Statistical Analysis To reduce dimensional complexity and reveal underlying structure in the morphometric dataset, we employed principal component analysis (PCA), a method well-suited for capturing variance across correlated variables and emphasizing dominant trait combinations (Todd et al., 2001 ). The PCA was performed using BioVinci version 3.09, a user-friendly and statistically robust platform chosen for its integrated clustering features and visual interpretability. The clustering results were used to recognize potential phenotypic groupings across turbidity gradients, reflecting adaptive morphological responses of coral Echinopora spp. that inhabit varying turbidity levels of water. To validify the clustering result, additional data was also added to the PCA process as an outgroup. Additional data were collected from the West Lombok region, approximately 95 km from the study area (outgroup site: 8°43'0.06"S, 115°54'43.56"E). This dataset includes measurements of corallite diameter (CD) and corallite area (CA) of E. lamellosa (n = 30), two key morphometric parameters linked to turbidity-related stress (Ghafari et al. 2024 ). The outgroup site is characterized by relatively clear water (TSS ± SD = 13.05 ± 0.75 mL L⁻¹, visibility ± SD = 8.75 ± 0.43 m). RESULTS AND DISCUSSION The results show that there was a different grouping trend among samples following the fluctuation of water turbidity levels. The most salient outcome from our PCA was the separation between observed groups (among station groups or between depth regimes) in both Echinopora lamellosa and E. pacificus , as respectively shown in Fig. 2 and Fig. 3 . In E. pacificus , the first principal component (PC1) captures 97.52% of the total dispersion among observed samples. It is clear that the population of turbid (green symbol) is separated from the non-turbid, without showing any overlap between the two (Fig. 3 ). Likewise, the deep population (symbolized by '△') is isolated from the shallow population (symbolized by '○'), indicating strong morphometric differentiation across environments. However, grouping pattern is slightly different in E. lamellosa , where the ordination reveals overlap in the observed sample groups, with maximum variance up to 97.88% among samples. The overlap between groups indicates that most members (60.98%, n = 25) of the non-turbid population living in deeper regimes (4 m) exhibit similar characteristics to the turbid population inhabit in shallow water (2 m) (Fig. 2 ). This supports the assumption that the distinct micro-morphological characteristics observed in E. lamellosa from deeper waters are indeed suited to shallow-turbid environment. The overlaps zone might critical to signify the micro-morphological adaptation range of Echinopora spp. within different turbidity gradient and might reflect higher possibility for plasticity mechanisms rather than selection. Although there is faint overlap, the PCA result of E. lamellosa generally implies that the separation pattern remains clear among population, with the trend of grouping similar to what found in E. pacificus . Additional data were incorporate to validate the robustness of multivariate ordination by means of PCA. The results showed that the comparison data of Echinopora lamellosa (symbolized by '🔴') was clustered in similar zone with members of the non-turbid stations, especially for those that inhabit in the shallow waters (Fig. 4 ). This site is characterized by low turbidity levels, comparable to the non-turbid stations in the main study area. The grouping pattern is likely due to the similar turbidity characteristics shared between the outgroup habitat and the non-turbid stations, reflecting a distribution consistent with previously reported trends. This convergence between independently sourced outgroup data and non-turbid in-group samples strengthens the interpretation that turbidity is a key environmental driver of micro-morphometric variation in Echinopora spp. The fact that samples from a different region, but similar turbidity condition, fall within the same cluster as in situ non-turbid populations provides an external validation of the ordination pattern. However, it should be noted that this simulation only involves fewer variables (CA and CD). Despite this limitation, Fig. 4 illustrates a clear trend, indicating that the distribution of variation in micro-morphometric traits among Echinopora colonies corresponds to the turbidity gradient. This alignment enhances the credibility of the PCA-based conclusions and supports the broader claim that environmental conditions, such as turbidity, may play a role in shaping morphological expression. In general, it appears the grouping trend of Echinopora spp. based on modularity of their micro-morphological characteristics are discriminated by the turbidity level of water (among stations and between depths), as previously studied by Ghafari et al. ( 2024 ). Similar pattern of response to turbidity level was observed in other coral population, such as in Favia speciosa (Todd et al., 2001 ), Leptoria Phrygia (Stafford-Smith, 1990 ), Pocillopora spp (Soto et al., 2018 ; Jones et al., 2020 ). and Turbinaria spp. (Anthony et al., 2005 ; Sofonia et al., 2008). There are various factors that support the survivability of corals in turbid conditions, such as the efficiency of sediment rejection and the coral tolerance level to sediment exposure, which can be vary between species. Sediment rejection efficiency is not directly related to sediment tolerance. Some species are poor sediment rejectors but have high sediment tolerance (Stafford-Smith, 1990 ; Zweifler et al., 2021 ). Although the turbidity effect certainly differs between species, it is clear that the morphological aspect, from macro to very fine scales, is a crucial control on tissue damage from sediments (Duckworth et al., 2017 ). Building on this understanding, our findings contribute novel empirical evidence that foliose corals—previously considered highly susceptible to sedimentation due to their foliose lifeform—may, in fact, exhibit significant micro-morphological plasticity in response to turbidity gradients. This challenges the traditional view that laminar corals are uniformly vulnerable to sedimentation (see Fabricius et al. ( 2006 ), Erftemeijer et al. ( 2012 ), Duckworth et al. ( 2017 ), and Jones et al. ( 2019 )), and instead suggests that some multi-morphotypic species—like Echinopora spp.—may possess the capacity to structurally adapt to variable sedimentary regimes. The results imply that these corals are not only able to tolerate environmental stressors associated with turbidity, but may also exploit such habitats more effectively than previously assumed, highlighting their overlooked ecological potential in marginal reef zones. Given this evidence, a critical question arises: what mechanisms enable such morphological variability? While the observed patterns exhibited by Echinopora spp. in this study appear to be related to environmental factors—particularly turbidity—, the underlying mechanisms remain difficult to confirm. Indeed, the observed micro-morphological variations of the present studied Echinopora spp seem well-suited to distinct turbidity regimes, suggesting possible adaptation. However, this variability can be attributed to genetic differentiation, phenotypic plasticity or a combination of both, and often appears to be environmentally correlated (Kramer et al., 2020 ; Million et al., 2022 ). Coral genotypes with a phenotypic advantage in the certain local environment may be actively selected for specialization (Drury & Lirman, 2021 ), or the corals might have plastic phenotypes that respond to prevalent conditions (Kenkel & Matz, 2016 ; Fox et al., 2019 ). It is also plausible thatboth mechanisms may function together. Alternatively, morphological distribution may result from stochastic processes that can directly or indirectly effect genotypes in small or isolated populations, potentially leading to beneficial outcomes via genetic drift (Starger et al., 2010 ; van Oppen et al., 2015 ; Quigley et al., 2019 ). Studies have documented distinct plastic responses in other coral taxa (Bruno & Edmunds, 1997 ; Shaish et al., 2007 ), yet attributing such patterns to either plasticity or selection remains challenging due to the inherently heterogeneous nature of reef environments. Disentangling these influences is essential for understanding how corals like Echinopora spp. adapt morphologically across variable habitats. Several conjectures arise to explain the mechanism of the morphological adaptation pattern of Echinopora spp. Previous studies suggest that some population of Echinopora spp. in eastern Lombok waters (area of present study) may have experienced a bottleneck effect that influenced by various environmental stressor, including water turbidity (Bachtiar et al., 2021 ; Ghafari et al., 2022 ). In addition, phenotypic matching with molecular signature shows interrelated patterns between selected Echinopora gene and their displayed morphotypes (Ghafari, 2022 ), which might lead to assumption on possible genetic-driven plasticity. However, previous studies lack of identified genetic markers and the samples were taken directly from nature without treatment for propagation, thus it is still unable to confirm the adaptation mechanism experienced by Echinopora spp. Future reciprocal transplant experiments would be necessary to identify and to confirm whether specialization, phenotypic plasticity or genetic drift is responsible for the micro-morphological variation identified in this study. If turbidity is indeed a driver of micro-morphometric variation, an important next question is how specific morphological traits may confer adaptive advantages to corals under such conditions, both at the colony scale (e.g., sediment rejection or light capture) and at the reef scale (e.g., population persistence or growth). Equally important is determining whether these traits arise through phenotypic plasticity or reflect long-term specialization. Declarations ACKNOWLEDGEMENTS We gratefully acknowledge Prof. Imam Bachtiar for providing equipment and gear during field sampling, Vita Fitrianti for assistance with laboratory work, and Namira R. P. Muquita for improving the English language of this manuscript. We also sincerely thank BioVinci for generously providing free access to their platform for data processing and statistical analysis. We also would like to acknowledge the grant support of Badan Riset dan Inovasi Daerah (BRIDA) of West Nusa Tenggara Province, Indonesia (Grant no. 184.LPPNTB.X.2022) and Universiti Malaysia Terengganu, Malaysia (Grant no. UMT//PGRG/2024/55530). COMPLIANCE WITH ETHICAL STANDARDS Funding: This research was supported by Badan Riset dan Inovasi Daerah (BRIDA) of West Nusa Tenggara Province, Indonesia (Grant no. 184.LPPNTB.X.2022) and Universiti Malaysia Terengganu, Malaysia (Grant no. UMT//PGRG/2024/55530). Conflict of interest: The authors declare that they have no conflict of interest. Ethical approval: Coral tissue sampling was conducted in accordance with Indonesian national regulations, under permits issued by the Department of Marine Affairs and Fisheries (Dinas Kelautan dan Perikanan, DKP) of West Nusa Tenggara Province, and with coordination from local conservation authorities in Gili Sulat-Lawang MPA, East Lombok, Indonesia. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-7040520","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":492717516,"identity":"b306e7fc-a6ad-460a-af1f-bd1fef99f16f","order_by":0,"name":"Muhammad Irsyad Abiyusfi Ghafari","email":"","orcid":"","institution":"Life as Naturalist","correspondingAuthor":false,"prefix":"","firstName":"Muhammad","middleName":"Irsyad Abiyusfi","lastName":"Ghafari","suffix":""},{"id":492717517,"identity":"6458bb09-4336-46bc-9cb8-cc4cee76ff12","order_by":1,"name":"Mohd Hanafi Idris","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA6ElEQVRIiWNgGAWjYBACAxBRYSDB2MDMwPgAIpZAhJYzEC3MBhA+UVoYGBgbGBjYJIjSYs5+9uCHAwUWstvb2Z9V8+b8YeBnzzFgutmGW4tlT16yxAEDCeM5h3nMbvNuM2CQ7HljwJyLR4vBgRwD6Q8GEokzmHnYwFoMbuQQ0HL+jfGPA2At7M+KQVrsCWq5kWMmAdHCYMYMtkWCoJY3ZhYgvwAdZiw5d5sxj8SZZwWHc87hc1iO8Y0Df+pkZ/Aff/jh7TY5Of725I2Pc8pwa8EAPCDiACMbCVqg4A/pWkbBKBgFo2DYAgBYAE2jqmoSVAAAAABJRU5ErkJggg==","orcid":"","institution":"Universiti Malaysia Terengganu Fakulti Perikanan dan Sains Makanan","correspondingAuthor":true,"prefix":"","firstName":"Mohd","middleName":"Hanafi","lastName":"Idris","suffix":""},{"id":492717518,"identity":"213cc6b3-3125-40ea-b2bf-49791beb8bac","order_by":2,"name":"Ezmahamrul Afreen Awalludin","email":"","orcid":"","institution":"Universiti Malaysia Terengganu Fakulti Perikanan dan Sains Makanan","correspondingAuthor":false,"prefix":"","firstName":"Ezmahamrul","middleName":"Afreen","lastName":"Awalludin","suffix":""},{"id":492717519,"identity":"609518ff-212d-4c03-89de-334f009e2ed9","order_by":3,"name":"Abu Hena Mustafa Kamal","email":"","orcid":"","institution":"Universiti Malaysia Terengganu Fakulti Perikanan dan Sains Makanan","correspondingAuthor":false,"prefix":"","firstName":"Abu","middleName":"Hena Mustafa","lastName":"Kamal","suffix":""}],"badges":[],"createdAt":"2025-07-03 17:36:05","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7040520/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7040520/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s00227-026-04802-z","type":"published","date":"2026-03-06T15:59:13+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":88001666,"identity":"28061991-8ac9-4fef-9985-48d1dc4c0419","added_by":"auto","created_at":"2025-07-31 10:24:11","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":703335,"visible":true,"origin":"","legend":"\u003cp\u003eSampling location and number of samples per site. (A) Map showing sampled population on Gili Sulat-Lawang. Blue-coloured circle indicates the non-turbid stations (ST1=8°17'42\"S, 116°41'09\"E; ST4=8°19'26\"S, 116°42'31\"E), while green circle pointed out the turbid stations (ST2=8°20'20\"S, 116°43'42\"E; 8°18'01\"S, 116°41'46\"E). (B) The number of sampled colonies of \u003cem\u003eEchinopora lamellosa\u003c/em\u003e and (C) \u003cem\u003eE. pacificus\u003c/em\u003e in both turbid and non-turbid stations. Lighter region of pie chart indicates percentage of shallow population sampled (2 m), while darker region showing percentage of deep population sampled (4 m).\u003c/p\u003e","description":"","filename":"Fig.1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7040520/v1/cea924be74a4868634216a2a.jpg"},{"id":88001653,"identity":"0621f97e-e7dd-4c12-88fc-474b26081551","added_by":"auto","created_at":"2025-07-31 10:24:11","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":81039,"visible":true,"origin":"","legend":"\u003cp\u003eOrdination result of PCA on observed \u003cem\u003eEchinopora lamellosa\u003c/em\u003e samples. The graph indicates slightly separation between the non-turbid populations (ST1 and ST4) and turbid populations (ST2 and ST3) of both shallow (circle-shaped symbols) and deep (triangle-shaped symbols) regime. The similarities order from turbid – deep → turbid – shallow → non-turbid – deep → non-turbid – shallow. Take a note that the overlap occurs between turbid – shallow population and non-turbid – deep population.\u003c/p\u003e","description":"","filename":"Fig.2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7040520/v1/618bc061570f2658676df58f.jpg"},{"id":88001680,"identity":"58500a7f-3eb5-45ff-9894-1f3578d27578","added_by":"auto","created_at":"2025-07-31 10:24:13","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":62942,"visible":true,"origin":"","legend":"\u003cp\u003eOrdination result of PCA on observed \u003cem\u003eEchinopora pacificus\u003c/em\u003e samples. The graph indicates complete separation between the non-turbid populations (ST1 and ST4) and turbid populations (ST2 and ST3) of both shallow (circle-shaped symbols) and deep (triangle-shaped symbols) regime.\u003c/p\u003e","description":"","filename":"Fig.3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7040520/v1/966e6316a829b8599a068f74.jpg"},{"id":88001656,"identity":"de8b9aa4-246b-4b59-b1be-53e009c0625b","added_by":"auto","created_at":"2025-07-31 10:24:11","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":91913,"visible":true,"origin":"","legend":"\u003cp\u003eOrdination result of PCA on observed \u003cem\u003eEchinopora lamellosa\u003c/em\u003e with outgroup. The outgroup tends to clustered among non-turbid populations (ST1 and ST4), indicates their similarities. Thus, the graph successfully portrays separation between the non-turbid populations and turbid populations (ST2 and ST3) of both shallow (circle-shaped symbols) and deep (triangle-shaped symbols) regime\u003c/p\u003e","description":"","filename":"Fig.4.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7040520/v1/02a4abbd62020548358b51c3.jpg"},{"id":104251298,"identity":"10755a39-792f-4da8-9b94-501aa3d43cca","added_by":"auto","created_at":"2026-03-09 16:12:42","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1361160,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7040520/v1/d9780a2a-8b68-4533-8f34-1187ba41afdb.pdf"}],"financialInterests":"","formattedTitle":"Evidence of adaptive plasticity in the coral Echinopora spp. to different turbidity regimes","fulltext":[{"header":"INTRODUCTION","content":"\u003cp\u003eDespite that the sediment has long been considered as among the most significant stressors to coral reef systems, increasing current evidence suggests that turbid waters might serve as an important reef conservation hotspot and corals exposed to enhanced turbidity could be more resilient to climate change impacts (Soares et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Zweifler et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Carlson et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Theoretically, turbidity provides shelter for coral from two current major bleaching factors: the excessive UV exposure (Morgan et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) and the elevated sea water temperature (L\u0026ouml;ptien and Meier \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Oxenford and Vall\u0026egrave;s \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), while at the same time it also holds detrimental potential to suffocate corals to bleach (Erftemeijer et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Jones et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). It has led to disagreement among scientists on the actual effect of turbidity on coral reefs (see Sully and van Woesik (\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2020\u003c/span\u003e); Zweifler et al. (\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2021\u003c/span\u003e); and Lucas et al. (\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2023\u003c/span\u003e)). Turbid coral reefs remain understudied due to the technical limitations associated with underwater visibility and imaging in sediment-rich environments.\u003c/p\u003e\u003cp\u003eIncreased turbidity and sedimentation pose varying levels of stress on different coral species, with some proving more sensitive than others. Most corals rely on morphological adaptations to cope with sediment settling on their surfaces. Among these, branching corals tend to accumulate the least sediment due to their complex three-dimensional structures, which effectively reduce the surface area where sediments can settle (Doszpot et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Jones et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). In contrast, foliose corals, characterized by their broad, layered morphologies and horizontal surfaces, are more prone to sediment accumulation. Studies by Duckworth et al. (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) revealed that foliose-shaped corals generally show the highest surface sediment cover under elevated sedimentation levels, making them particularly vulnerable to stress.\u003c/p\u003e\u003cp\u003eThis susceptibility has significant consequences. Foliose corals often suffer the greatest losses among coral morphologies due to sediment-induced anoxia, where trapped sediment blocks light and gas exchange, leading to tissue death (Fabricius et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Duckworth et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Jones et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). However, despite their apparent vulnerability, some foliose corals have shown unexpected tolerance to sedimentation. Some populations have been reported to persist under turbid conditions, including sediment-laden runoff and severe sedimentation stress near river mouth (Bachtiar and Hadi \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Guest et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Syahrir et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). A study by Ghafari et al. (\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) found that fine-scale morphological variation\u0026mdash;particularly in corallite structure\u0026mdash;may play a key role in enabling foliose corals, such as those from the genus \u003cem\u003eEchinopora\u003c/em\u003e, to survive in turbid environments.\u003c/p\u003e\u003cp\u003eHere, our observations on foliose coral species suggest a degree of morphological plasticity that may help corals with this lifeform withstand varying turbidity levels. This study proposes that morphological plasticity particularly within multi-morphotypic coral species may be a critical adaptive mechanism enabling survival in turbid habitats previously deemed unsuitable due to sedimentation stress, possibly allowing them to exploit a wide range of turbid environment.\u003c/p\u003e"},{"header":"MATERIALS AND METHODS","content":"\u003cp\u003e\u003cb\u003eSampling Sites\u003c/b\u003e\u003c/p\u003e\u003cp\u003eFour sampling sites were located on reef areas in East Lombok, Indonesia, and were selected to represent distinct turbidity conditions (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eA). Turbidity level classification was based on quantitative assessment by Ghafari et al. (\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), resulting in the identification of non-turbid sites, i.e., ST1 and ST4 (TSS\u0026thinsp;\u0026plusmn;\u0026thinsp;SD\u0026thinsp;=\u0026thinsp;14.32\u0026thinsp;\u0026plusmn;\u0026thinsp;1.61 ml L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, visibility\u0026thinsp;\u0026plusmn;\u0026thinsp;SD\u0026thinsp;=\u0026thinsp;8.49\u0026thinsp;\u0026plusmn;\u0026thinsp;0.45 m) and turbid sites, i.e., ST2 and ST3 (TSS\u0026thinsp;\u0026plusmn;\u0026thinsp;SD\u0026thinsp;=\u0026thinsp;29.05\u0026thinsp;\u0026plusmn;\u0026thinsp;0.76 ml L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, visibility\u0026thinsp;\u0026plusmn;\u0026thinsp;SD\u0026thinsp;=\u0026thinsp;4.35\u0026thinsp;\u0026plusmn;\u0026thinsp;0.70 m). At each site, coral specimens were sampled at two depth regimes: shallow (2 m) and deep (4 m), to capture variation related to both water clarity and depth.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003cb\u003eSampling, Data Collection and Processing\u003c/b\u003e\u003c/p\u003e\u003cp\u003eThe sampling process collected 114 specimens of \u003cem\u003eEchinopora lamellosa\u003c/em\u003e and 23 specimens of \u003cem\u003eE. pacificus\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eB-C). Specimens taken from a depth of 2 m at each station are referred to as the \u0026lsquo;shallow population\u0026rsquo;, whereas those taken from 4 m depth are termed the \u0026lsquo;deep population\u0026rsquo;. Despite the small depth differences in East Lombok, Ghafari et al. (\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) reported the resulting turbidity levels vary significantly, potentially affecting the morphological adaptations of the coral colonies.\u003c/p\u003e\u003cp\u003eData processing was conducted on ten measurable and/or countable micro-morphological characters of each observed specimen, i.e., corallite area (CA), minimum (CD-min) and maximum (CD-max) diameter of corallite, average corallite diameter (CD-rate), corallite spacing (ICD), mouth-disk diameter (MD), exsert septal spacing (IES), polyp density (PD), polyp height (PH), and skeleton thickness (SkT), following the protocol by Ghafari et al. (\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Measurement of PH and SkT required microscopic inspection, which was carried out by making cross-sectional cuts, as described by Ghafari et al. (\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003eStatistical Analysis\u003c/h2\u003e\u003cp\u003eTo reduce dimensional complexity and reveal underlying structure in the morphometric dataset, we employed principal component analysis (PCA), a method well-suited for capturing variance across correlated variables and emphasizing dominant trait combinations (Todd et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). The PCA was performed using BioVinci version 3.09, a user-friendly and statistically robust platform chosen for its integrated clustering features and visual interpretability. The clustering results were used to recognize potential phenotypic groupings across turbidity gradients, reflecting adaptive morphological responses of coral \u003cem\u003eEchinopora\u003c/em\u003e spp. that inhabit varying turbidity levels of water.\u003c/p\u003e\u003cp\u003eTo validify the clustering result, additional data was also added to the PCA process as an outgroup. Additional data were collected from the West Lombok region, approximately 95 km from the study area (outgroup site: 8\u0026deg;43'0.06\"S, 115\u0026deg;54'43.56\"E). This dataset includes measurements of corallite diameter (CD) and corallite area (CA) of \u003cem\u003eE. lamellosa\u003c/em\u003e (n\u0026thinsp;=\u0026thinsp;30), two key morphometric parameters linked to turbidity-related stress (Ghafari et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The outgroup site is characterized by relatively clear water (TSS\u0026thinsp;\u0026plusmn;\u0026thinsp;SD\u0026thinsp;=\u0026thinsp;13.05\u0026thinsp;\u0026plusmn;\u0026thinsp;0.75 mL L⁻\u0026sup1;, visibility\u0026thinsp;\u0026plusmn;\u0026thinsp;SD\u0026thinsp;=\u0026thinsp;8.75\u0026thinsp;\u0026plusmn;\u0026thinsp;0.43 m).\u003c/p\u003e\u003c/div\u003e"},{"header":"RESULTS AND DISCUSSION","content":"\u003cp\u003eThe results show that there was a different grouping trend among samples following the fluctuation of water turbidity levels. The most salient outcome from our PCA was the separation between observed groups (among station groups or between depth regimes) in both \u003cem\u003eEchinopora lamellosa\u003c/em\u003e and \u003cem\u003eE. pacificus\u003c/em\u003e, as respectively shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e and Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eIn \u003cem\u003eE. pacificus\u003c/em\u003e, the first principal component (PC1) captures 97.52% of the total dispersion among observed samples. It is clear that the population of turbid (green symbol) is separated from the non-turbid, without showing any overlap between the two (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Likewise, the deep population (symbolized by '△') is isolated from the shallow population (symbolized by '○'), indicating strong morphometric differentiation across environments.\u003c/p\u003e\u003cp\u003eHowever, grouping pattern is slightly different in \u003cem\u003eE. lamellosa\u003c/em\u003e, where the ordination reveals overlap in the observed sample groups, with maximum variance up to 97.88% among samples. The overlap between groups indicates that most members (60.98%, n\u0026thinsp;=\u0026thinsp;25) of the non-turbid population living in deeper regimes (4 m) exhibit similar characteristics to the turbid population inhabit in shallow water (2 m) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). This supports the assumption that the distinct micro-morphological characteristics observed in \u003cem\u003eE. lamellosa\u003c/em\u003e from deeper waters are indeed suited to shallow-turbid environment. The overlaps zone might critical to signify the micro-morphological adaptation range of \u003cem\u003eEchinopora\u003c/em\u003e spp. within different turbidity gradient and might reflect higher possibility for plasticity mechanisms rather than selection. Although there is faint overlap, the PCA result of \u003cem\u003eE. lamellosa\u003c/em\u003e generally implies that the separation pattern remains clear among population, with the trend of grouping similar to what found in \u003cem\u003eE. pacificus\u003c/em\u003e.\u003c/p\u003e\u003cp\u003eAdditional data were incorporate to validate the robustness of multivariate ordination by means of PCA. The results showed that the comparison data of \u003cem\u003eEchinopora lamellosa\u003c/em\u003e (symbolized by '\u0026#128308;') was clustered in similar zone with members of the non-turbid stations, especially for those that inhabit in the shallow waters (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). This site is characterized by low turbidity levels, comparable to the non-turbid stations in the main study area. The grouping pattern is likely due to the similar turbidity characteristics shared between the outgroup habitat and the non-turbid stations, reflecting a distribution consistent with previously reported trends.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eThis convergence between independently sourced outgroup data and non-turbid in-group samples strengthens the interpretation that turbidity is a key environmental driver of micro-morphometric variation in \u003cem\u003eEchinopora\u003c/em\u003e spp. The fact that samples from a different region, but similar turbidity condition, fall within the same cluster as in situ non-turbid populations provides an external validation of the ordination pattern. However, it should be noted that this simulation only involves fewer variables (CA and CD). Despite this limitation, Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e illustrates a clear trend, indicating that the distribution of variation in micro-morphometric traits among \u003cem\u003eEchinopora\u003c/em\u003e colonies corresponds to the turbidity gradient. This alignment enhances the credibility of the PCA-based conclusions and supports the broader claim that environmental conditions, such as turbidity, may play a role in shaping morphological expression.\u003c/p\u003e\u003cp\u003eIn general, it appears the grouping trend of \u003cem\u003eEchinopora\u003c/em\u003e spp. based on modularity of their micro-morphological characteristics are discriminated by the turbidity level of water (among stations and between depths), as previously studied by Ghafari et al. (\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Similar pattern of response to turbidity level was observed in other coral population, such as in \u003cem\u003eFavia speciosa\u003c/em\u003e (Todd et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2001\u003c/span\u003e), \u003cem\u003eLeptoria Phrygia\u003c/em\u003e (Stafford-Smith, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1990\u003c/span\u003e), \u003cem\u003ePocillopora\u003c/em\u003e spp (Soto et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Jones et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). and \u003cem\u003eTurbinaria\u003c/em\u003e spp. (Anthony et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Sofonia et al., 2008). There are various factors that support the survivability of corals in turbid conditions, such as the efficiency of sediment rejection and the coral tolerance level to sediment exposure, which can be vary between species. Sediment rejection efficiency is not directly related to sediment tolerance. Some species are poor sediment rejectors but have high sediment tolerance (Stafford-Smith, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Zweifler et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Although the turbidity effect certainly differs between species, it is clear that the morphological aspect, from macro to very fine scales, is a crucial control on tissue damage from sediments (Duckworth et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eBuilding on this understanding, our findings contribute novel empirical evidence that foliose corals\u0026mdash;previously considered highly susceptible to sedimentation due to their foliose lifeform\u0026mdash;may, in fact, exhibit significant micro-morphological plasticity in response to turbidity gradients. This challenges the traditional view that laminar corals are uniformly vulnerable to sedimentation (see Fabricius et al. (\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), Erftemeijer et al. (\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), Duckworth et al. (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), and Jones et al. (\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2019\u003c/span\u003e)), and instead suggests that some multi-morphotypic species\u0026mdash;like \u003cem\u003eEchinopora\u003c/em\u003e spp.\u0026mdash;may possess the capacity to structurally adapt to variable sedimentary regimes. The results imply that these corals are not only able to tolerate environmental stressors associated with turbidity, but may also exploit such habitats more effectively than previously assumed, highlighting their overlooked ecological potential in marginal reef zones.\u003c/p\u003e\u003cp\u003eGiven this evidence, a critical question arises: what mechanisms enable such morphological variability? While the observed patterns exhibited by \u003cem\u003eEchinopora\u003c/em\u003e spp. in this study appear to be related to environmental factors\u0026mdash;particularly turbidity\u0026mdash;, the underlying mechanisms remain difficult to confirm. Indeed, the observed micro-morphological variations of the present studied \u003cem\u003eEchinopora\u003c/em\u003e spp seem well-suited to distinct turbidity regimes, suggesting possible adaptation. However, this variability can be attributed to genetic differentiation, phenotypic plasticity or a combination of both, and often appears to be environmentally correlated (Kramer et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Million et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Coral genotypes with a phenotypic advantage in the certain local environment may be actively selected for specialization (Drury \u0026amp; Lirman, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), or the corals might have plastic phenotypes that respond to prevalent conditions (Kenkel \u0026amp; Matz, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Fox et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). It is also plausible thatboth mechanisms may function together.\u003c/p\u003e\u003cp\u003eAlternatively, morphological distribution may result from stochastic processes that can directly or indirectly effect genotypes in small or isolated populations, potentially leading to beneficial outcomes via genetic drift (Starger et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; van Oppen et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Quigley et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Studies have documented distinct plastic responses in other coral taxa (Bruno \u0026amp; Edmunds, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Shaish et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2007\u003c/span\u003e), yet attributing such patterns to either plasticity or selection remains challenging due to the inherently heterogeneous nature of reef environments. Disentangling these influences is essential for understanding how corals like \u003cem\u003eEchinopora\u003c/em\u003e spp. adapt morphologically across variable habitats.\u003c/p\u003e\u003cp\u003eSeveral conjectures arise to explain the mechanism of the morphological adaptation pattern of \u003cem\u003eEchinopora\u003c/em\u003e spp. Previous studies suggest that some population of \u003cem\u003eEchinopora\u003c/em\u003e spp. in eastern Lombok waters (area of present study) may have experienced a bottleneck effect that influenced by various environmental stressor, including water turbidity (Bachtiar et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Ghafari et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In addition, phenotypic matching with molecular signature shows interrelated patterns between selected \u003cem\u003eEchinopora\u003c/em\u003e gene and their displayed morphotypes (Ghafari, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), which might lead to assumption on possible genetic-driven plasticity. However, previous studies lack of identified genetic markers and the samples were taken directly from nature without treatment for propagation, thus it is still unable to confirm the adaptation mechanism experienced by \u003cem\u003eEchinopora\u003c/em\u003e spp.\u003c/p\u003e\u003cp\u003eFuture reciprocal transplant experiments would be necessary to identify and to confirm whether specialization, phenotypic plasticity or genetic drift is responsible for the micro-morphological variation identified in this study. If turbidity is indeed a driver of micro-morphometric variation, an important next question is how specific morphological traits may confer adaptive advantages to corals under such conditions, both at the colony scale (e.g., sediment rejection or light capture) and at the reef scale (e.g., population persistence or growth). Equally important is determining whether these traits arise through phenotypic plasticity or reflect long-term specialization.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eACKNOWLEDGEMENTS\u003c/h2\u003e\u003cp\u003eWe gratefully acknowledge Prof. Imam Bachtiar for providing equipment and gear during field sampling, Vita Fitrianti for assistance with laboratory work, and Namira R. P. Muquita for improving the English language of this manuscript. We also sincerely thank BioVinci for generously providing free access to their platform for data processing and statistical analysis. We also would like to acknowledge the grant support of Badan Riset dan Inovasi Daerah (BRIDA) of West Nusa Tenggara Province, Indonesia (Grant no. 184.LPPNTB.X.2022) and Universiti Malaysia Terengganu, Malaysia (Grant no. UMT//PGRG/2024/55530).\u003c/p\u003e\u003cp\u003e\u003cstrong\u003eCOMPLIANCE WITH ETHICAL STANDARDS\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding:\u003c/strong\u003e This research was supported by Badan Riset dan Inovasi Daerah (BRIDA) of West Nusa Tenggara Province, Indonesia (Grant no. 184.LPPNTB.X.2022) and Universiti Malaysia Terengganu, Malaysia (Grant no. UMT//PGRG/2024/55530).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflict of interest:\u003c/strong\u003e The authors declare that they have no conflict of interest.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthical approval:\u0026nbsp;\u003c/strong\u003eCoral tissue sampling was conducted in accordance with Indonesian national regulations, under permits issued by the Department of Marine Affairs and Fisheries (Dinas Kelautan dan Perikanan, DKP) of West Nusa Tenggara Province, and with coordination from local conservation authorities in Gili Sulat-Lawang MPA, East Lombok, Indonesia. The standard, non-destructive biopsy techniques were used to minimize impact on coral colonies. These samples were previously partially analysed and published in Coral Reefs (DOI:10.1007/s00338-024-02579-5), and the current manuscript presents additional and distinct analyses from the same original sampling effort. All necessary approvals were obtained, and documentary evidence is available upon request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n \u003cli\u003eAnthony KR, Hoogenboom MO, Connolly SR (2005) Adaptive variation in coral geometry and the optimization of internal colony light climates. Funct Ecol 19:17\u0026ndash;26. https://doi.org/10.1111/j.0269-8463.2005.00925.x\u003c/li\u003e\n \u003cli\u003eBachtiar I, Ghafari MIA, Rahman I et al (2021) Neither coral- nor symbiont- genetic diversity may explain the resistance of the coral \u003cem\u003eEchinopora lamellosa\u003c/em\u003e to bleaching. J Trop Biodivers Biotechnol 6:66161. \u0026nbsp; https://doi.org/10.22146/jtbb.66161\u003c/li\u003e\n \u003cli\u003eBachtiar I, Hadi TA (2019) Differential impacts of 2016 coral bleaching on coral reef benthic communities at Sekotong Bay, Lombok Barat Indonesia. 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Coral Reefs 43:1819\u0026ndash;1830. https://doi.org/10.1007/s00338-024-02579-5\u003c/li\u003e\n \u003cli\u003eGhafari MIA, Litaay M, Agus R (2022) Variation on the morphological features of coral \u003cem\u003eEchinopora lamellosa\u003c/em\u003e population suggests corals survivorship mechanisms in Alas Strait, Indonesia. AACL Bioflux 15:1680\u0026ndash;1691\u003c/li\u003e\n \u003cli\u003eGuest JR, Tun K, Low J et al (2016) 27 years of benthic and coral community dynamics on turbid, highly urbanised reefs off Singapore. Sci Rep 6:36260. https://doi.org/10.1038/srep36260\u003c/li\u003e\n \u003cli\u003eJones R, Fisher R, Bessell-Bowne P (2019) Sediment deposition and coral smothering. PLoS One 14. https://doi.org/10.1371/journal.pone.0216248\u003c/li\u003e\n \u003cli\u003eJones R, Giofre N, Luter HM et al (2020) Responses of corals to chronic turbidity. 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AACL Bioflux 11:362\u0026ndash;378\u003c/li\u003e\n \u003cli\u003eTodd PA, Sanderson PG, Chou LM (2001) Morphological variation in the polyps of the scleractinian coral \u003cem\u003eFavia speciosa\u003c/em\u003e (Dana) around Singapore. Hydrobiologia 444:227\u0026ndash;235. https://doi.org/10.1023/A:1017570100029\u003c/li\u003e\n \u003cli\u003evan Oppen MJ, Oliver JK, Putnam HM et al (2015) Building coral reef resilience through assisted evolution. Proc Natl Acad Sci 112:2307\u0026ndash;2313. https://doi.org/10.1073/pnas.142230111\u003c/li\u003e\n \u003cli\u003eZweifler A, O\u0026apos;Leary M, Morgan K et al (2021) Turbid coral reefs: Past, present and future\u0026mdash;a review. Diversity, 13. https://doi.org/10.3390/d13060251\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"marine-biology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"mabi","sideBox":"Learn more about [Marine Biology](https://www.springer.com/journal/227)","snPcode":"227","submissionUrl":"https://submission.nature.com/new-submission/227/3","title":"Marine Biology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Coral resilience, foliose corals, morphological plasticity, sediment stress, turbidity","lastPublishedDoi":"10.21203/rs.3.rs-7040520/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7040520/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"Turbid reefs, once seen as marginal, are now considered potential refuges for laminar-shaped corals under climate stress. This study examines micro-morphological plasticity in two foliose coral species: Echinopora lamellosa and E. pacificus, across turbid and non-turbid sites in East Lombok, Indonesia. A total of 137 specimens from different depths (2 m and 4 m) and turbidity levels were analysed using ten diagnostic morphological traits. Principal component analysis explained \u0026gt;97% of the total morphological variance and revealed clear differentiation by site and depth. E. pacificus showed complete separation between populations, while E. lamellosa displayed partial overlap—suggesting environmentally driven plasticity. Deep colonies in non-turbid sites resembled shallow colonies in turbid areas, supporting the idea of adaptive morphometric responses rather than fixed traits. Outgroup validation using a geographically distinct population of the same species confirmed the observed patterns, strengthening the evidence for environment-related variation. The patterns observed may involve phenotypic plasticity or epigenetic changes, which warrants further investigation. These findings highlight the plastic ability of foliose corals to exploit either shallow or turbid habitats through their diverse morphological traits, underscoring their resilience and potential significance for future reef conservation, restoration and management efforts amid turbidity-driven climate change.","manuscriptTitle":"Evidence of adaptive plasticity in the coral Echinopora spp. to different turbidity regimes","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-07-31 10:24:02","doi":"10.21203/rs.3.rs-7040520/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revise and Resubmit","date":"2025-09-15T05:36:33+00:00","index":"","fulltext":""},{"type":"reviewerAgreed","content":"","date":"2025-07-30T13:57:53+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-07-29T16:01:22+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-07-15T08:56:48+00:00","index":"","fulltext":""},{"type":"submitted","content":"Marine Biology","date":"2025-07-14T21:45:36+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"marine-biology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"mabi","sideBox":"Learn more about [Marine Biology](https://www.springer.com/journal/227)","snPcode":"227","submissionUrl":"https://submission.nature.com/new-submission/227/3","title":"Marine Biology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"1fead646-e04c-4a52-85f3-b67542392cd2","owner":[],"postedDate":"July 31st, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2026-03-09T16:08:33+00:00","versionOfRecord":{"articleIdentity":"rs-7040520","link":"https://doi.org/10.1007/s00227-026-04802-z","journal":{"identity":"marine-biology","isVorOnly":false,"title":"Marine Biology"},"publishedOn":"2026-03-06 15:59:13","publishedOnDateReadable":"March 6th, 2026"},"versionCreatedAt":"2025-07-31 10:24:02","video":"","vorDoi":"10.1007/s00227-026-04802-z","vorDoiUrl":"https://doi.org/10.1007/s00227-026-04802-z","workflowStages":[]},"version":"v1","identity":"rs-7040520","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7040520","identity":"rs-7040520","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}