Reef Foraminifera: Artificial vs. Natural Environments

Reef Foraminifera: Artificial vs. Natural Environments | 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 Reef Foraminifera: Artificial vs. Natural Environments FATIN IZZATI MINHAT, Hasrizal Shaari, Wan Nurzalia Wan Saelan, and 4 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-5280024/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 The success of artificial reefs is influenced by localised oceanographic factors including wave exposure, sedimentation rates, and proximity to natural reef environments. The Bidong Shipwreck, on the shallow shelf of the South China Sea, is a hotspot for meiobenthos. This study compared foraminifera distribution at the Bidong Shipwreck with that of the adjacent natural reef of Bidong Island. Foraminiferal assemblages at the shipwreck were less diverse. Amphistegina spp. were the most dominant genus at both study sites. Calcarinids—such as Calcarina and Neorotalia spp.—occurred only on the natural reef. Cluster analysis categorised the foraminiferal assemblages from both sites into three distinct groups based on substrate preference. Group A consisted of foraminiferal assemblages from the shallow reef area of Bidong Island with a sandy substrate, where Nummulites and Peneroplis spp. were indicator species. Group B represented a mix of stations from both the shipwreck and natural reef, characterised by water depths > 18 m and a muddy sand substrate, with Amphistegina spp. comprising ~ 80% of this assemblage. Group C displayed a more diverse distribution of foraminiferal species, including a mix of symbiont-bearing and heterotrophic taxa coexisting within the coarser sediments. Despite being over 500 years old, the Bidong Shipwreck does not exhibit a foraminiferal assemblage similar to that of the nearby natural reef. Differences in substrate type and water depth between these sites caused foraminiferal assemblage shifts. This suggests that although artificial reefs can have beneficial effects on fish biomass and meiobenthic communities, their impact on protists such as foraminifera can be markedly different. Figures Figure 1 Figure 2 Figure 3 1. Introduction Artificial reefs are human-engineered structures deliberately or inadvertently introduced into marine environments [ 1 , 2 ]. They are frequently used to enhance fish diversity, bolster fish biomass, and conserve coastal habitats [ 3 , 4 ]. The success of artificial reefs depends on localised oceanographic factors, including wave exposure, sedimentation rates, and proximity to natural reef environments [ 5 , 2 ]. Shipwrecks, the remnants of maritime vessels, are designated as inadvertent artificial reefs, and depending on their geographical location and temporal history, certain shipwrecks have been proven to be conducive to the establishment of microhabitats, serving as marine organism sanctuaries [ 2 , 6 , 7 ]. However, in certain instances, shipwrecks facilitate the colonisation of invasive species, thereby inducing perturbations in the natural ecology of the surrounding area [ 8 ]. The colonisation of species in artificial reefs or shipwrecks can influence adjacent benthic communities [ 2 , 9 ] and older shipwrecks appear to foster a more balanced ecosystem in conjunction with neighbouring natural ecosystems [ 2 ] Despite this, to date, there is limited research and understanding of the biotic components and colonisation of shipwrecks > 100 years old and their adjacent environments [ 9 ]. This is attributed to the relatively young age (mostly < 100 years old) and remote positioning of shipwrecks in natural reef environments in many places [ 2 , 5 ]. Additionally, numerous investigations into the benthic colonisation of shipwrecks or artificial reefs have concentrated on macrobenthic communities [ 6 – 8 ], lacking data on microbenthic communities, including protist foraminifera. In Malaysia, the Bidong Shipwreck provides an ideal setting for investigating the correlation between foraminiferal distribution in the shipwreck area and nearby reef ecosystems. Based on archaeological evidence, the Bidong Shipwreck may have occurred between 1500 and 1550 CE [ 10 ] making it older than 400 years. However, rapid sedimentation in the shallow shelf region where the shipwreck resides has resulted in the submergence of most vessels, leaving only archaeological artefacts, such as ceramics, visible on the seafloor surface. A recent investigation of meiobenthos colonisation around the Bidong Shipwreck indicated that it is a thriving hotspot for the meiobenthos community [ 11 ]. However, this study did not compare meiofaunal diversity between shipwrecks and nearby natural reefs. Consequently, to elucidate the response of benthic foraminiferal assemblages to artificial and natural reef environments, this study aimed to compare the benthic foraminiferal assemblage of the Bidong Shipwreck with that of the adjacent reef ecosystem on Bidong Island. Additionally, we explored the main environmental parameters that influence foraminiferal distribution between artificial and natural reef environments in the South China Sea. 2. Methodology The Bidong Shipwreck and Bidong Island are in the South China Sea on the east coast of Peninsular Malaysia. The area has a tropical climate with two monsoon events. The average water temperature and salinity ranges from 29 to 30 °C and 32 to 33ppt, respectively. The Bidong Shipwreck lies at a water depth of 18 m, approximately two nautical miles southwest of Bidong Island (05° 36.9021′ N and 103° 02.38995′ E) and is estimated to be 25 m long and 10 m wide. Based on archaeological evidence found at the shipwreck and absolute dating, it is estimated that the ship sunk between 1500 and 1550 CE [ 10 ] The shipwreck was discovered in 2013; however, the first scientific excavation was only conducted in 2017, and the second in 2022. For this study, nine samples were collected along two transects during the second excavation (Fig. 1 ). All sediment samples were collected by SCUBA divers at 50 m intervals along the transect. Bulk samples were collected from the top 10 cm of the surface sediment and scooped into a ziplock bag. For comparison, sediment samples were also collected from the natural reef of Bidong Island, less than 12 nautical miles from the shipwreck. Here, ten samples were collected by SCUBA divers at 20 m intervals along two transects perpendicular to the coastline. Samples from both sites were preserved in ethanol (> 80%) and transported to the laboratory for future analysis. 3. Laboratory analysis 3.1. Sediment analysis The sediment grain size composition was determined with the dry sieving method following Folk [ 12 ]. Meanwhile, the organic matter compositions were determined based on the loss ignition method based on Heiri et al. [ 13 ]. 3.2. Foraminifera analysis For the foraminifera analysis, ~ 10 cm 3 of sediment samples from both sites were washed on 125 and 63 µm sieves under tap water. The residue remaining on the sieves was transferred to a weighing boat and dried in an oven at 50°C for 24 h. Once completely dried, the samples were stored in pre-labelled bags for future analysis. Next, ~ 300 foraminifera specimens > 125 µm were picked from each sample. All specimens were sorted on microslides and identified to the genus level based on taxonomic references [ 14 , 15 ]. 3.3. Community composition and diversity Foraminiferal data from the natural reef were based on data published by Husain et al. [ 16 ] All data were screened before statistical analysis to avoid reworked specimens. To understand the community composition, distribution, and diversity of benthic foraminifera from both natural and artificial reefs, diversity indices, indicator values [ 17 ] and cluster analyses were performed using PAST v4.1 to understand the species-habitat associations [ 18 ] 4. Results Sediment analyses indicated that a sandy substrate dominated the natural reef, whereas coarser sands were observed around the shipwreck area (Table 1 ). A higher mud composition (8%) was also observed in the shipwreck. Nonetheless, organic matter accumulation was relatively higher around the natural reef. The water depth around the natural reef and shipwreck was between 7–26 and 18 m, respectively. Table 1 Water depth (m) and organic matter and sediment composition (%) observed at all 19 stations. The sites labelled as natural reef and artificial reef are situated at Bidong Island and the shipwreck, respectively. Station Site category Depth (m) OM (%) Mud Sand Gravel W3S1 Natural reef 7 2.9 0.0 92.1 7.8 W3S2 Natural reef 14 2.9 0.0 77.6 22.4 W3S3 Natural reef 21 3.0 0.0 85.6 14.3 W3S4 Natural reef 24 2.8 0.1 80.9 19.0 W3S5 Natural reef 26 2.8 0.1 81.4 18.5 W4S1 Natural reef 7 3.2 0.0 98.5 1.5 W4S2 Natural reef 12 2.9 0.1 94.9 5.0 W4S3 Natural reef 14 3.2 0.0 92.2 7.8 W4S4 Natural reef 18 2.3 0.0 90.3 9.6 W4S5 Natural reef 24 3.2 1.1 88.7 10.2 SH1 Artificial reef 18 0.9 8.5 75.8 37.5 SH2 Artificial reef 18 0.9 8.1 72.9 33.9 SH3 Artificial reef 18 0.5 8.0 78.0 34.5 SH4 Artificial reef 18 0.5 8.0 76.1 32.4 SH5 Artificial reef 18 0.0 8.7 63.0 33.0 SH6 Artificial reef 18 N/A 8.1 77.0 38.0 SH7 Artificial reef 18 0.6 8.1 74.4 35.1 SH8 Artificial reef 18 0.5 8.2 74.6 35.3 SH9 Artificial reef 18 0.7 8.0 86.0 32.2 The foraminiferal assemblages at the shipwreck were less diverse than those found on the reef (Table 1 ). The number of foraminiferal genera reported from the shipwreck and the reef ranged from 2 to 12 and 6 to 15, respectively. The highest Shannon–Wiener index was observed for W4S2 (H’= 2.34). Amphistegina spp. were the dominant genus at both sites. At the reef—where the water depth ranged between 7–26 m—the relative abundance of Amphistegina spp. was between 23–85%. At the shipwreck—with a water depth of 18 m—the relative abundance of Amphistegina spp. was notably higher (68–98%). Calcarinids such as Calcarina and Neorotalia spp. were completely absent from the shipwreck while on the natural reef, the relative abundance of Calcarina and Neorotalia spp. was between 1–14 and 1–5%, respectively. Porcelaneous foraminifera such as Quinqueloculina and Triloculina spp. were more abundant on the natural reef while the distribution of agglutinated groups, such as Sahulia and Textularia spp. was notably similar between the two sites. A Q-mode cluster analysis grouped all 19 stations into three groups (Fig. 2 ). Group A consisted of three stations from natural reefs characterised by indicator species, such as Nummulites (IndVal = 67%) and Peneroplis (IndVal = 51%) (Table 2 ). The water depth for Group A ranged between 7–18 m where a sandy substrate dominated. The foraminiferal assemblage was dominated by hyaline and porcelaneous groups, whereas agglutinated taxa were absent. Group B consisted of 11 stations, nine from the shipwreck and two from the deeper (> 20 m) parts of the natural reef—W3S5 and W4S5. The presence of two foraminiferal genera, Operculina and Spiroloculina , defined the foraminiferal assemblages in this group. This group represents foraminiferal assemblages, especially the hyaline group, which prefer deeper waters where the substrate is dominated by muddy sand. Group C represented five stations from shallow to deep (7–24 m) areas of the natural reef. This group is characterised with highly diverse foraminiferal assemblage (H’ > 1.6) that prefer a coarse sandy substrate. The indicator species that define this group belong to Heterolepa (IndVal = 80%), Reusella (IndVal = 69%), and Discorbinella spp. (IndVal = 60%) and the calcarinids. Table 2 The three cluster groups based on the Q-mode classical cluster analysis (Euclidean distance). For each group, the substrate type, organic matter composition (%), Shannon–Wiener diversity Index (H’), wall-type groups, and indicator species (IndVal) values are listed. Clusters Group A Group B Group C Stations Three stations from the natural reef Nine stations from the shipwreck and two from the natural reef Five stations from the natural reef Water depth 7–18 m 18–26 m 7–24 m Organic matter 2.3–3.2% 0–3.2% 2.9–3.2% Type of substrate Sandy Muddy sand Coarse sand Shannon–Wiener (H’) 1.07–1.51 0.10–1.31 1.65–2.34 Agglutinated (%) 0% 0–5% 0–10% Porcelaneous (%) 18–25% 0–22% 14–22% Hyaline (%) 75–82% 78–100% 68–82% Indicator genera Nummulites spp. (67%) Operculina spp. (66%) Heterolepa spp. (80%) (IndVal %, P < 0.05) Peneroplis spp. (51%) Spiroloculina spp. (46%) Reusella spp. (69%) Amphistegina spp. (36%) Discorbinella spp. (60%) Millieonella spp. (60%) Elphidium spp. (59%) Neorotalia spp. (59%) Calcarina spp. (53%) Planorbulinella spp. (51%) Using the canonical correspondent analysis (CCA) (Fig. 3 ) the cumulative variation of axis-1 was determined to be 59.4% (eigenvalue = 0.206) and that of axis-2 to be 29.3% (eigenvalue = 0.102). The substrate type exhibited the greatest influence on foraminiferal distribution at the natural reef and shipwreck sites. 5. Discussion 5.1 Foraminifera assemblages at the shipwreck The foraminifera assemblages shows that distribution at the shipwreck (H’= 0.10–1.31) is less diverse than that at the adjacent natural reef (H’ = 0.60–2.34). We believe that this is due to dynamic coastal processes in the shipwreck area, which form muddy sand with fewer hard substrates for attachment. Furthermore, significant sediment resuspension occurs in coastal areas between November and March each year due to the monsoon seasons, characterized by heavy rains and heightened wave energy [ 19 ] may influence foraminiferal diversity differences. This study revealed that the relatively high mud composition within the shipwreck area likely promoted the distribution of Amphistegina (68–98%), Operculina (2–6%), and Quinqueloculina (4–9%) spp. Amphistegina and Operculina spp. are both larger benthic foraminifera that host symbionts within their test. However, compared with other larger benthic foraminifera (i.e., Baculogypsina sphaerulata and several Calcarina spp.), [ 20 ] Amphistegina spp. and Operculina spp. may thrive in environments with lower light intensities, allowing them to inhabit deeper shelf areas or regions where high sediment suspensions influence light penetration [ 21 , 22 ]. Additionally, the presence of Operculina spp. in the coastal waters of Sulawesi suggests that this genus exhibits a higher tolerance to terrestrial sediments, a characteristic shared by the Bidong Shipwreck environment [ 22 ]. 5.2. Foraminiferal assemblages at Bidong Island Common genera of foraminifera reported on Bidong Island include Amphistegina , Calcarina , Elphidium , Heterolepa , Heterostegina , Quinqueloculina , Peneroplis , and Triloculina spp. [ 16 ]. The reef surrounding Bidong Island receives a moderate amount of nutrients because of human settlement and tourism activities on the island [ 25 ]. The close proximity between Bidong Island and mainland Peninsular Malaysia means that the reef surrounding this island is subject to terrestrial influences. In mesotrophic reefs, terrestrial input and substrate type strongly influence water transparency and nutrient availability [ 23 , 24 ]. The presence of heterotrophic foraminifera, such as Ammonia and Elphidium spp. in the reef area of Bidong Island may be linked to the enrichment of organic matter in the reef environment [ 20 , 22 ]. A study by Husain et al. [ 16 ] across 12 sites surrounding Bidong Island showed that heterotrophic species such as Elphidium and Textularia spp. were predominantly found in the shallowest sites, where the water depth is < 10 m. Notably, several of these sites are inhabited by humans, and their close proximity to the island suggests a potential influx of organic matter into the sediment through anthropogenic activities, thereby fostering an increase in heterotrophic taxa [ 20 ]. 5.3. Similarities and differences in foraminifera assemblages There has been a large increase in the number of artificial reefs worldwide since the 1960s [ 26 ]. Many of these have been intentionally introduced to boost fish biomass in the surrounding areas [ 27 ]. Consequently, many efforts to increase fish biomass have involved the deployment of artificial reefs in coastal areas [ 28 , 29 ]. Artificial reefs comprising accidental shipwrecks have been classified as unintentionally deployed [ 26 ] and proven to promote changes in the surrounding habitats [ 2 ]. However, unlike other shipwrecks which have been identified as artificial reefs, the Bidong Island shipwreck was completely submerged under the seafloor, making its discovery unique. Reports by local fishers on the high fish biomass in the area initiated preliminary efforts to study the area [ 30 ]. The shipwreck was predicted to have hit the bottom of the seafloor ~ 500 years ago [ 31 ]. Situated within the shallow Sunda Shelf region, the location of the shipwreck receives terrigenous sediment from the nearby Terengganu River at an approximate rate of 0.30 cm per year [ 32 ]. The sedimentation process caused the large body of the ship to be fully covered, with only ceramic artefacts from the shipwreck visible on the seafloor, marking the wreck area (Fig. 1 ). Ecological studies conducted by Ismailluddin et al. [ 33 ] and Abdullah et al. [ 11 ] reported a high density and diversity of benthic communities within the shipwreck area, including corals, nematodes, antrapod meiofauna, and foraminifera. Both studies indicate that benthic communities, particularly those favouring soft-bottom habitats, thrive around shipwrecks. However, these studies did not compare the diversity of these benthic communities with that of the adjacent natural reef area of Bidong Island. Therefore, our understanding of benthic communities in artificial and natural coral reefs remains incomplete. Here, cluster analysis was used to categorise the foraminiferal assemblages from natural reefs and shipwrecks into three distinct groups based on substrate preference. Group A consisted of foraminiferal assemblages from a shallow reef area on Bidong Island with a sandy substrate. Nummulites and Peneroplis spp. were identified as indicator species for this group, distinguishing them from the other two groups. Peneroplis spp., which host red algae as symbionts, have been documented in the shallowest reef areas of Okinawa [ 34 ] and the Spermonde Archipelago [ 23 ]. However, this species thrives in regions with high organic matter and low light availability, such as the Great Barrier Reef [ 20 ]. On Bidong Island, Peneroplis spp. are the most abundant in shallow reef areas (5–10 m), where they occur together with Amphistegina and Quinqueloculina spp. [ 16 ]. The second cluster, Group B, represents a mix of stations from the shipwreck and natural reef, characterised by water depths of > 18 m and a muddy sand substrate. This assemblage had the highest average relative abundance of Amphistegina spp. (80%). Noticeable occurrences of Operculina spp. were observed with increasing water depth and mud content. We suspect that sediment resuspension, especially in the shipwreck area, resulted in higher particulate matter in the water column, resulting in low light availability and thereby allowing Amphistegina and Operculina spp. to occur in highest abundance. Similarly, a study of foraminiferal assemblages at four different artificial reefs (wreck sites) offshore in Brunei revealed a significant increase in the abundance of Amphistegina and Operculina spp., particularly in areas where wrecks had been colonised by corals and sponges [ 35 ]. Additionally, Operculina spp. are known to be highly tolerant to terrestrial sediment influences, as documented in mesotrophic reefs in the Spermonde Archipelago [ 22 ]. The final assemblage identified (Group C) exhibited a more diverse distribution of foraminiferal species. This group includes a mix of symbiont-bearing and heterotrophic foraminifera taxa that coexist within coarser sediments. The assemblages are representative of reef flats close to the shoreline and reef slopes around Bidong Island. CCA analysis indicated that the variation in substrate types between the natural reef of Bidong Island and the artificial reef (shipwreck) was the primary factor influencing foraminiferal distribution. The foraminiferal assemblages in the deeper parts of the natural reef showed greater similarity to those recorded around the shipwreck. Studies suggest that with sufficient time, artificial reefs may become a good surrogate for the close natural reef environment, promoting balanced ecosystems between natural and artificial reefs [ 2 , 27 ]. However, despite being > 500 years old, the Bidong Shipwreck does not exhibit a foraminiferal assemblage similar to that of the nearby natural reef. The differences in substrate type and water depth between the shipwreck and natural reef caused a shift in foraminiferal assemblages. Additionally, the foraminiferal diversity around the shipwreck was notably lower than that in the natural reef environment. This suggests that while artificial reefs can have beneficial effects on fish biomass, their impact on protists, such as foraminifera, can be markedly different. 6. Conclusion Our findings demonstrate distinct differences in the composition and diversity of foraminiferal assemblages between the artificial reef at the Bidong Shipwreck and the natural reefs of Bidong Island. The artificial reef, despite being > 500 years old, harboured a less diverse foraminiferal assemblage, predominantly consisting of Amphistegina spp. with an absence of Calcarina spp. This is in stark contrast to natural reefs, which exhibit higher diversity, including species such as Nummulites and Peneroplis and a balance of symbiont-bearing and heterotrophic taxa. This study identified substrate type and water depth as the primary environmental parameters influencing foraminiferal distribution. The Bidong Shipwreck, located in deeper waters with higher sedimentation rates, supports the adaptation of foraminifera to lower light conditions and higher mud content. Conversely, natural reefs with their sandy substrates and varying depths support a more diverse and balanced assemblage of foraminifera. The artificial reef at the Bidong Shipwreck, although beneficial to certain benthic communities, does not replicate the foraminiferal diversity observed in natural reefs. The distinct environmental conditions of these habitats play a critical role in shaping the distribution and diversity of foraminiferal assemblages. Declarations Conflicts of interest No conflict of interest Funding This work was supported by the strategic research grant of UMT (UMT/SRG/2022/55384) Author Contribution F.I.M contributed to sampling design, methodology, manuscript drafting, and statistical analysis.H.S. oversaw field sampling, sample collection, and secured funding. W.N.W.S. assisted with sampling design and provided data from Bidong Island research. F.A.B.A. conducted laboratory and data analysis. M.N.A.M, N.S.S. and S.Y. contributed to manuscript editing. Acknowledgement We are grateful to Department of National Heritage (JWN), UZMA Archaeological Research Sdn. Bhd. (UAR) and Hidrokinetik Sdn. Bhd. We would like to thank the professional SCUBA divers of UMT, Kamaruzzaman Arifin, Mohd Sahimi Abdul Rashid, Mohd Forzi Zahari, and Mohd Fauzi Romeli that help with the sample collection. Thank you to boat crews of UMT Discovery II and the land-support team from JWN, UAR, and UMT. 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The preliminary dating of the Bidong shipwreck, Malaysia. Int J Nautical Archaeol. 2024:1–15. doi: 10.1080/10572414.2024.2323984 . Kamaruzzaman BY, Ong MC. The determination of 210Pb in the sediments: sedimentation rates from the Terengganu coastal waters, Malaysia. Curr World Environ. 2010;5(1):9–14. doi: 10.12944/CWE.5.1.02 . Ismailluddin SAM, Mohd Abdullah M, Ibrahim YS, Idris I. Benthic community diversity at marine archaeological site, Pulau Bidong, South China Sea. UMT JUR. 2021;3(3):93–106. doi: 10.46754/umtjur.2021.07.011 . Hohenegger J, Yordanova E, Nakano Y, Tatzreiter F. Habitats of larger foraminifera on the upper reef slope of Sesoko Island, Okinawa, Japan. Mar Micropaleontol. 1999;36(2–3):109–68. doi: 10.1016/S0377-8398(98)00030-9 . Goeting S, Briguglio A, Eder W, Hohenegger J, Roslim A, Kocsis L. Depth distribution of modern larger benthic foraminifera offshore Brunei Darussalam. Micropaleontology. 2018;64(4):299–316. doi: 10.47894/mpal.64.4.04 . Additional Declarations No competing interests reported. Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. 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-5280024","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":377781699,"identity":"e8b615be-2274-44e6-8a38-f64c6d19e5b6","order_by":0,"name":"FATIN IZZATI MINHAT","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA30lEQVRIiWNgGAWjYBAC9gYGhgMMDBIMBgwMjA8gYgn4tfAcYIZpYWYGajMgTgsYALWwSRCnhf38wcOFORZ55uznj1Xz5vxh4GfPMWC62YZHC08yw+GZ2ySKLXuS2W7zbjNgkOx5Y8Cci0eLPQNQC+82icQNB6BaDG7k4NfCw/8YquX8Y7ZikBZ7glokYLbcSGZjBtsiQVDLYwOolsfGknO3GfNInHlWcDjnHD6HJT7+zLutDuiwxIcf3m6Tk+NvT974OKcMtxZMM0DEAUY2ErRAwR/StYyCUTAKRsGwBQDT5E291AvTRAAAAABJRU5ErkJggg==","orcid":"","institution":"Universiti Malaysia Terengganu","correspondingAuthor":true,"prefix":"","firstName":"FATIN","middleName":"IZZATI","lastName":"MINHAT","suffix":""},{"id":377781700,"identity":"2b5c00bb-b1bf-4e3e-8696-afa3ddd97c90","order_by":1,"name":"Hasrizal Shaari","email":"","orcid":"","institution":"Universiti Malaysia Terengganu","correspondingAuthor":false,"prefix":"","firstName":"Hasrizal","middleName":"","lastName":"Shaari","suffix":""},{"id":377781701,"identity":"2d2d353f-2c19-44dc-9bef-6629f51b8936","order_by":2,"name":"Wan Nurzalia Wan Saelan","email":"","orcid":"","institution":"Universiti Malaysia Terengganu","correspondingAuthor":false,"prefix":"","firstName":"Wan","middleName":"Nurzalia Wan","lastName":"Saelan","suffix":""},{"id":377781702,"identity":"b0ea2301-63ec-49bf-99d1-92fef5c41e2a","order_by":3,"name":"Farah Amelia Bee Abdullah","email":"","orcid":"","institution":"Universiti Malaysia Terengganu","correspondingAuthor":false,"prefix":"","firstName":"Farah","middleName":"Amelia Bee","lastName":"Abdullah","suffix":""},{"id":377781703,"identity":"20df4f06-8140-4123-bd94-daf0f137630e","order_by":4,"name":"Muhammad Naim Abdul Malek","email":"","orcid":"","institution":"Universiti Sains Malaysia","correspondingAuthor":false,"prefix":"","firstName":"Muhammad","middleName":"Naim Abdul","lastName":"Malek","suffix":""},{"id":377781704,"identity":"0049c83d-9b24-4b76-9fc9-1a1ea7d6fd19","order_by":5,"name":"Shouye Yang","email":"","orcid":"","institution":"Tongji University","correspondingAuthor":false,"prefix":"","firstName":"Shouye","middleName":"","lastName":"Yang","suffix":""},{"id":377781705,"identity":"9df86aac-7f5e-47f2-840a-616468afa379","order_by":6,"name":"Nursyafiqah Syazana Sharudin","email":"","orcid":"","institution":"Universiti Malaysia Terengganu","correspondingAuthor":false,"prefix":"","firstName":"Nursyafiqah","middleName":"Syazana","lastName":"Sharudin","suffix":""}],"badges":[],"createdAt":"2024-10-17 06:08:09","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-5280024/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-5280024/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":69104139,"identity":"f47ed9de-fd1a-4a99-a428-929925d0a46b","added_by":"auto","created_at":"2024-11-15 16:48:03","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":715663,"visible":true,"origin":"","legend":"\u003cp\u003eShipwreck image obtained from a side-scan sonar. The two crossed transects were plotted on the Bidong Shipwreck in the South China Sea. The dotted line indicates the shipwreck shape.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-5280024/v1/2b6e085dc9105cb651f7fa94.png"},{"id":69104140,"identity":"66b59fd0-d160-47e0-8269-a55e3c20bb6e","added_by":"auto","created_at":"2024-11-15 16:48:03","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":30409,"visible":true,"origin":"","legend":"\u003cp\u003eDendrogram produced by Q-mode cluster analysis based on Euclidean distances for 19 sampling stations from both the natural reef and the shipwreck. The cluster analysis divided the samples into Groups A, B, and C\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-5280024/v1/b05b4f4b922776d999e6bb71.png"},{"id":69104137,"identity":"b5dcbcd0-1fb0-429e-802a-83eee16a2938","added_by":"auto","created_at":"2024-11-15 16:48:02","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":51741,"visible":true,"origin":"","legend":"\u003cp\u003eCanonical correspondence analysis (CCA) triplot based on species-environmental relationships between the artificial and natural reef. The total variances of both axes were 88.7 %\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-5280024/v1/dcabeae36da8a143a5306e28.png"},{"id":70345247,"identity":"a993e74d-d59f-4e11-be0d-9234fe148319","added_by":"auto","created_at":"2024-12-02 10:53:30","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1443985,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-5280024/v1/e11b706c-d849-43a5-8ccd-ecf7f610d3e2.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Reef Foraminifera: Artificial vs. Natural Environments","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eArtificial reefs are human-engineered structures deliberately or inadvertently introduced into marine environments [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e]. They are frequently used to enhance fish diversity, bolster fish biomass, and conserve coastal habitats [\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e]. The success of artificial reefs depends on localised oceanographic factors, including wave exposure, sedimentation rates, and proximity to natural reef environments [\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e]. Shipwrecks, the remnants of maritime vessels, are designated as inadvertent artificial reefs, and depending on their geographical location and temporal history, certain shipwrecks have been proven to be conducive to the establishment of microhabitats, serving as marine organism sanctuaries [\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e]. However, in certain instances, shipwrecks facilitate the colonisation of invasive species, thereby inducing perturbations in the natural ecology of the surrounding area [\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eThe colonisation of species in artificial reefs or shipwrecks can influence adjacent benthic communities [\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e] and older shipwrecks appear to foster a more balanced ecosystem in conjunction with neighbouring natural ecosystems [\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e] Despite this, to date, there is limited research and understanding of the biotic components and colonisation of shipwrecks\u0026thinsp;\u0026gt;\u0026thinsp;100 years old and their adjacent environments [\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e]. This is attributed to the relatively young age (mostly\u0026thinsp;\u0026lt;\u0026thinsp;100 years old) and remote positioning of shipwrecks in natural reef environments in many places [\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e]. Additionally, numerous investigations into the benthic colonisation of shipwrecks or artificial reefs have concentrated on macrobenthic communities [\u003cspan additionalcitationids=\"CR7\" citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e], lacking data on microbenthic communities, including protist foraminifera. In Malaysia, the Bidong Shipwreck provides an ideal setting for investigating the correlation between foraminiferal distribution in the shipwreck area and nearby reef ecosystems.\u003c/p\u003e \u003cp\u003eBased on archaeological evidence, the Bidong Shipwreck may have occurred between 1500 and 1550 CE [\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e] making it older than 400 years. However, rapid sedimentation in the shallow shelf region where the shipwreck resides has resulted in the submergence of most vessels, leaving only archaeological artefacts, such as ceramics, visible on the seafloor surface. A recent investigation of meiobenthos colonisation around the Bidong Shipwreck indicated that it is a thriving hotspot for the meiobenthos community [\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e]. However, this study did not compare meiofaunal diversity between shipwrecks and nearby natural reefs. Consequently, to elucidate the response of benthic foraminiferal assemblages to artificial and natural reef environments, this study aimed to compare the benthic foraminiferal assemblage of the Bidong Shipwreck with that of the adjacent reef ecosystem on Bidong Island. Additionally, we explored the main environmental parameters that influence foraminiferal distribution between artificial and natural reef environments in the South China Sea.\u003c/p\u003e"},{"header":"2. Methodology","content":"\u003cp\u003eThe Bidong Shipwreck and Bidong Island are in the South China Sea on the east coast of Peninsular Malaysia. The area has a tropical climate with two monsoon events. The average water temperature and salinity ranges from 29 to 30 \u0026deg;C and 32 to 33ppt, respectively. The Bidong Shipwreck lies at a water depth of 18 m, approximately two nautical miles southwest of Bidong Island (05\u0026deg; 36.9021\u0026prime; N and 103\u0026deg; 02.38995\u0026prime; E) and is estimated to be 25 m long and 10 m wide. Based on archaeological evidence found at the shipwreck and absolute dating, it is estimated that the ship sunk between 1500 and 1550 CE [\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e] The shipwreck was discovered in 2013; however, the first scientific excavation was only conducted in 2017, and the second in 2022. For this study, nine samples were collected along two transects during the second excavation (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). All sediment samples were collected by SCUBA divers at 50 m intervals along the transect. Bulk samples were collected from the top 10 cm of the surface sediment and scooped into a ziplock bag.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eFor comparison, sediment samples were also collected from the natural reef of Bidong Island, less than 12 nautical miles from the shipwreck. Here, ten samples were collected by SCUBA divers at 20 m intervals along two transects perpendicular to the coastline. Samples from both sites were preserved in ethanol (\u0026gt;\u0026thinsp;80%) and transported to the laboratory for future analysis.\u003c/p\u003e"},{"header":"3. Laboratory analysis","content":"\u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e3.1. Sediment analysis\u003c/h2\u003e \u003cp\u003eThe sediment grain size composition was determined with the dry sieving method following Folk [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e]. Meanwhile, the organic matter compositions were determined based on the loss ignition method based on Heiri et al. [\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e].\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e3.2. Foraminifera analysis\u003c/h2\u003e \u003cp\u003eFor the foraminifera analysis, ~\u0026thinsp;10 cm\u003csup\u003e3\u003c/sup\u003e of sediment samples from both sites were washed on 125 and 63 \u0026micro;m sieves under tap water. The residue remaining on the sieves was transferred to a weighing boat and dried in an oven at 50\u0026deg;C for 24 h. Once completely dried, the samples were stored in pre-labelled bags for future analysis. Next, ~\u0026thinsp;300 foraminifera specimens\u0026thinsp;\u0026gt;\u0026thinsp;125 \u0026micro;m were picked from each sample. All specimens were sorted on microslides and identified to the genus level based on taxonomic references [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e].\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e3.3. Community composition and diversity\u003c/h2\u003e \u003cp\u003eForaminiferal data from the natural reef were based on data published by Husain et al. [\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e] All data were screened before statistical analysis to avoid reworked specimens. To understand the community composition, distribution, and diversity of benthic foraminifera from both natural and artificial reefs, diversity indices, indicator values [\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e] and cluster analyses were performed using PAST v4.1 to understand the species-habitat associations [\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e]\u003c/p\u003e \u003c/div\u003e"},{"header":"4. Results","content":"\u003cp\u003eSediment analyses indicated that a sandy substrate dominated the natural reef, whereas coarser sands were observed around the shipwreck area (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). A higher mud composition (8%) was also observed in the shipwreck. Nonetheless, organic matter accumulation was relatively higher around the natural reef. The water depth around the natural reef and shipwreck was between 7\u0026ndash;26 and 18 m, respectively.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eWater depth (m) and organic matter and sediment composition (%) observed at all 19 stations. The sites labelled as natural reef and artificial reef are situated at Bidong Island and the shipwreck, respectively.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eStation\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSite category\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDepth (m)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eOM (%)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMud\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSand\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eGravel\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eW3S1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNatural reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e92.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e7.8\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eW3S2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNatural reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e77.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e22.4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eW3S3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNatural reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e85.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e14.3\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eW3S4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNatural reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e80.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e19.0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eW3S5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNatural reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e81.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e18.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eW4S1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNatural reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e98.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e1.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eW4S2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNatural reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e94.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e5.0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eW4S3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNatural reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e92.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e7.8\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eW4S4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNatural reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e90.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e9.6\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eW4S5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNatural reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e88.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e10.2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSH1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eArtificial reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e75.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e37.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSH2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eArtificial reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e72.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e33.9\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSH3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eArtificial reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e78.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e34.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSH4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eArtificial reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e76.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e32.4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSH5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eArtificial reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e63.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e33.0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSH6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eArtificial reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eN/A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e77.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e38.0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSH7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eArtificial reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e74.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e35.1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSH8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eArtificial reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e74.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e35.3\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSH9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eArtificial reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e86.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e32.2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eThe foraminiferal assemblages at the shipwreck were less diverse than those found on the reef (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The number of foraminiferal genera reported from the shipwreck and the reef ranged from 2 to 12 and 6 to 15, respectively. The highest Shannon\u0026ndash;Wiener index was observed for W4S2 (H\u0026rsquo;= 2.34). \u003cem\u003eAmphistegina\u003c/em\u003e spp. were the dominant genus at both sites. At the reef\u0026mdash;where the water depth ranged between 7\u0026ndash;26 m\u0026mdash;the relative abundance of \u003cem\u003eAmphistegina\u003c/em\u003e spp. was between 23\u0026ndash;85%. At the shipwreck\u0026mdash;with a water depth of 18 m\u0026mdash;the relative abundance of \u003cem\u003eAmphistegina\u003c/em\u003e spp. was notably higher (68\u0026ndash;98%). Calcarinids such as \u003cem\u003eCalcarina\u003c/em\u003e and \u003cem\u003eNeorotalia\u003c/em\u003e spp. were completely absent from the shipwreck while on the natural reef, the relative abundance of \u003cem\u003eCalcarina\u003c/em\u003e and \u003cem\u003eNeorotalia\u003c/em\u003e spp. was between 1\u0026ndash;14 and 1\u0026ndash;5%, respectively. Porcelaneous foraminifera such as \u003cem\u003eQuinqueloculina\u003c/em\u003e and \u003cem\u003eTriloculina\u003c/em\u003e spp. were more abundant on the natural reef while the distribution of agglutinated groups, such as \u003cem\u003eSahulia\u003c/em\u003e and \u003cem\u003eTextularia\u003c/em\u003e spp. was notably similar between the two sites.\u003c/p\u003e \u003cp\u003eA Q-mode cluster analysis grouped all 19 stations into three groups (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Group A consisted of three stations from natural reefs characterised by indicator species, such as \u003cem\u003eNummulites\u003c/em\u003e (IndVal\u0026thinsp;=\u0026thinsp;67%) and \u003cem\u003ePeneroplis\u003c/em\u003e (IndVal\u0026thinsp;=\u0026thinsp;51%) (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The water depth for Group A ranged between 7\u0026ndash;18 m where a sandy substrate dominated. The foraminiferal assemblage was dominated by hyaline and porcelaneous groups, whereas agglutinated taxa were absent. Group B consisted of 11 stations, nine from the shipwreck and two from the deeper (\u0026gt;\u0026thinsp;20 m) parts of the natural reef\u0026mdash;W3S5 and W4S5. The presence of two foraminiferal genera, \u003cem\u003eOperculina\u003c/em\u003e and \u003cem\u003eSpiroloculina\u003c/em\u003e, defined the foraminiferal assemblages in this group. This group represents foraminiferal assemblages, especially the hyaline group, which prefer deeper waters where the substrate is dominated by muddy sand. Group C represented five stations from shallow to deep (7\u0026ndash;24 m) areas of the natural reef. This group is characterised with highly diverse foraminiferal assemblage (H\u0026rsquo; \u0026gt; 1.6) that prefer a coarse sandy substrate. The indicator species that define this group belong to \u003cem\u003eHeterolepa\u003c/em\u003e (IndVal\u0026thinsp;=\u0026thinsp;80%), \u003cem\u003eReusella\u003c/em\u003e (IndVal\u0026thinsp;=\u0026thinsp;69%), and \u003cem\u003eDiscorbinella\u003c/em\u003e spp. (IndVal\u0026thinsp;=\u0026thinsp;60%) and the calcarinids.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eThe three cluster groups based on the Q-mode classical cluster analysis (Euclidean distance). For each group, the substrate type, organic matter composition (%), Shannon\u0026ndash;Wiener diversity Index (H\u0026rsquo;), wall-type groups, and indicator species (IndVal) values are listed.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colspan=\"3\" nameend=\"c4\" namest=\"c2\"\u003e \u003cp\u003eClusters\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eGroup A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eGroup B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eGroup C\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eStations\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eThree stations from the natural reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNine stations from the shipwreck and two from the natural reef\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFive stations from the natural reef\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWater depth\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7\u0026ndash;18 m\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e18\u0026ndash;26 m\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e7\u0026ndash;24 m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eOrganic matter\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2.3\u0026ndash;3.2%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u0026ndash;3.2%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.9\u0026ndash;3.2%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eType of substrate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSandy\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMuddy sand\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCoarse sand\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eShannon\u0026ndash;Wiener (H\u0026rsquo;)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.07\u0026ndash;1.51\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.10\u0026ndash;1.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.65\u0026ndash;2.34\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAgglutinated (%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u0026ndash;5%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0\u0026ndash;10%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePorcelaneous (%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e18\u0026ndash;25%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u0026ndash;22%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e14\u0026ndash;22%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eHyaline (%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e75\u0026ndash;82%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e78\u0026ndash;100%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e68\u0026ndash;82%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eIndicator genera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eNummulites\u003c/em\u003e spp. (67%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eOperculina\u003c/em\u003e spp. (66%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eHeterolepa\u003c/em\u003e spp. (80%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e(IndVal %, P\u0026thinsp;\u0026lt;\u0026thinsp;0.05)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003ePeneroplis\u003c/em\u003e spp. (51%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eSpiroloculina\u003c/em\u003e spp. (46%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eReusella\u003c/em\u003e spp. (69%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eAmphistegina\u003c/em\u003e spp. (36%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eDiscorbinella\u003c/em\u003e spp. (60%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eMillieonella\u003c/em\u003e spp. (60%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eElphidium\u003c/em\u003e spp. (59%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eNeorotalia\u003c/em\u003e spp. (59%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eCalcarina\u003c/em\u003e spp. (53%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003ePlanorbulinella\u003c/em\u003e spp. (51%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eUsing the canonical correspondent analysis (CCA) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e) the cumulative variation of axis-1 was determined to be 59.4% (eigenvalue\u0026thinsp;=\u0026thinsp;0.206) and that of axis-2 to be 29.3% (eigenvalue\u0026thinsp;=\u0026thinsp;0.102). The substrate type exhibited the greatest influence on foraminiferal distribution at the natural reef and shipwreck sites.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"5. Discussion","content":"\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003e5.1 Foraminifera assemblages at the shipwreck\u003c/h2\u003e \u003cp\u003eThe foraminifera assemblages shows that distribution at the shipwreck (H\u0026rsquo;= 0.10\u0026ndash;1.31) is less diverse than that at the adjacent natural reef (H\u0026rsquo; = 0.60\u0026ndash;2.34). We believe that this is due to dynamic coastal processes in the shipwreck area, which form muddy sand with fewer hard substrates for attachment. Furthermore, significant sediment resuspension occurs in coastal areas between November and March each year due to the monsoon seasons, characterized by heavy rains and heightened wave energy [\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e] may influence foraminiferal diversity differences. This study revealed that the relatively high mud composition within the shipwreck area likely promoted the distribution of \u003cem\u003eAmphistegina\u003c/em\u003e (68\u0026ndash;98%), \u003cem\u003eOperculina\u003c/em\u003e (2\u0026ndash;6%), and \u003cem\u003eQuinqueloculina\u003c/em\u003e (4\u0026ndash;9%) spp. \u003cem\u003eAmphistegina\u003c/em\u003e and \u003cem\u003eOperculina\u003c/em\u003e spp. are both larger benthic foraminifera that host symbionts within their test. However, compared with other larger benthic foraminifera (i.e., \u003cem\u003eBaculogypsina sphaerulata\u003c/em\u003e and several \u003cem\u003eCalcarina\u003c/em\u003e spp.), [\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e] \u003cem\u003eAmphistegina\u003c/em\u003e spp. and \u003cem\u003eOperculina\u003c/em\u003e spp. may thrive in environments with lower light intensities, allowing them to inhabit deeper shelf areas or regions where high sediment suspensions influence light penetration [\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e]. Additionally, the presence of \u003cem\u003eOperculina\u003c/em\u003e spp. in the coastal waters of Sulawesi suggests that this genus exhibits a higher tolerance to terrestrial sediments, a characteristic shared by the Bidong Shipwreck environment [\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e].\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003e5.2. Foraminiferal assemblages at Bidong Island\u003c/h2\u003e \u003cp\u003eCommon genera of foraminifera reported on Bidong Island include \u003cem\u003eAmphistegina\u003c/em\u003e, \u003cem\u003eCalcarina\u003c/em\u003e, \u003cem\u003eElphidium\u003c/em\u003e, \u003cem\u003eHeterolepa\u003c/em\u003e, \u003cem\u003eHeterostegina\u003c/em\u003e, \u003cem\u003eQuinqueloculina\u003c/em\u003e, \u003cem\u003ePeneroplis\u003c/em\u003e, and \u003cem\u003eTriloculina\u003c/em\u003e spp. [\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e]. The reef surrounding Bidong Island receives a moderate amount of nutrients because of human settlement and tourism activities on the island [\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e]. The close proximity between Bidong Island and mainland Peninsular Malaysia means that the reef surrounding this island is subject to terrestrial influences. In mesotrophic reefs, terrestrial input and substrate type strongly influence water transparency and nutrient availability [\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e, \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e]. The presence of heterotrophic foraminifera, such as \u003cem\u003eAmmonia\u003c/em\u003e and \u003cem\u003eElphidium\u003c/em\u003e spp. in the reef area of Bidong Island may be linked to the enrichment of organic matter in the reef environment [\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e]. A study by Husain et al. [\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e] across 12 sites surrounding Bidong Island showed that heterotrophic species such as \u003cem\u003eElphidium\u003c/em\u003e and \u003cem\u003eTextularia\u003c/em\u003e spp. were predominantly found in the shallowest sites, where the water depth is \u0026lt;\u0026thinsp;10 m. Notably, several of these sites are inhabited by humans, and their close proximity to the island suggests a potential influx of organic matter into the sediment through anthropogenic activities, thereby fostering an increase in heterotrophic taxa [\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e].\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003e5.3. Similarities and differences in foraminifera assemblages\u003c/h2\u003e \u003cp\u003eThere has been a large increase in the number of artificial reefs worldwide since the 1960s [\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e]. Many of these have been intentionally introduced to boost fish biomass in the surrounding areas [\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e]. Consequently, many efforts to increase fish biomass have involved the deployment of artificial reefs in coastal areas [\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e]. Artificial reefs comprising accidental shipwrecks have been classified as unintentionally deployed [\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e] and proven to promote changes in the surrounding habitats [\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e]. However, unlike other shipwrecks which have been identified as artificial reefs, the Bidong Island shipwreck was completely submerged under the seafloor, making its discovery unique. Reports by local fishers on the high fish biomass in the area initiated preliminary efforts to study the area [\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e]. The shipwreck was predicted to have hit the bottom of the seafloor\u0026thinsp;~\u0026thinsp;500 years ago [\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e]. Situated within the shallow Sunda Shelf region, the location of the shipwreck receives terrigenous sediment from the nearby Terengganu River at an approximate rate of 0.30 cm per year [\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e]. The sedimentation process caused the large body of the ship to be fully covered, with only ceramic artefacts from the shipwreck visible on the seafloor, marking the wreck area (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Ecological studies conducted by Ismailluddin et al. [\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e] and Abdullah et al. [\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e] reported a high density and diversity of benthic communities within the shipwreck area, including corals, nematodes, antrapod meiofauna, and foraminifera. Both studies indicate that benthic communities, particularly those favouring soft-bottom habitats, thrive around shipwrecks. However, these studies did not compare the diversity of these benthic communities with that of the adjacent natural reef area of Bidong Island. Therefore, our understanding of benthic communities in artificial and natural coral reefs remains incomplete.\u003c/p\u003e \u003cp\u003eHere, cluster analysis was used to categorise the foraminiferal assemblages from natural reefs and shipwrecks into three distinct groups based on substrate preference. Group A consisted of foraminiferal assemblages from a shallow reef area on Bidong Island with a sandy substrate. \u003cem\u003eNummulites\u003c/em\u003e and \u003cem\u003ePeneroplis\u003c/em\u003e spp. were identified as indicator species for this group, distinguishing them from the other two groups. \u003cem\u003ePeneroplis\u003c/em\u003e spp., which host red algae as symbionts, have been documented in the shallowest reef areas of Okinawa [\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e] and the Spermonde Archipelago [\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e]. However, this species thrives in regions with high organic matter and low light availability, such as the Great Barrier Reef [\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e]. On Bidong Island, \u003cem\u003ePeneroplis\u003c/em\u003e spp. are the most abundant in shallow reef areas (5\u0026ndash;10 m), where they occur together with \u003cem\u003eAmphistegina\u003c/em\u003e and \u003cem\u003eQuinqueloculina\u003c/em\u003e spp. [\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eThe second cluster, Group B, represents a mix of stations from the shipwreck and natural reef, characterised by water depths of \u0026gt;\u0026thinsp;18 m and a muddy sand substrate. This assemblage had the highest average relative abundance of \u003cem\u003eAmphistegina\u003c/em\u003e spp. (80%). Noticeable occurrences of \u003cem\u003eOperculina\u003c/em\u003e spp. were observed with increasing water depth and mud content. We suspect that sediment resuspension, especially in the shipwreck area, resulted in higher particulate matter in the water column, resulting in low light availability and thereby allowing \u003cem\u003eAmphistegina\u003c/em\u003e and \u003cem\u003eOperculina\u003c/em\u003e spp. to occur in highest abundance. Similarly, a study of foraminiferal assemblages at four different artificial reefs (wreck sites) offshore in Brunei revealed a significant increase in the abundance of \u003cem\u003eAmphistegina\u003c/em\u003e and \u003cem\u003eOperculina\u003c/em\u003e spp., particularly in areas where wrecks had been colonised by corals and sponges [\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e]. Additionally, \u003cem\u003eOperculina\u003c/em\u003e spp. are known to be highly tolerant to terrestrial sediment influences, as documented in mesotrophic reefs in the Spermonde Archipelago [\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eThe final assemblage identified (Group C) exhibited a more diverse distribution of foraminiferal species. This group includes a mix of symbiont-bearing and heterotrophic foraminifera taxa that coexist within coarser sediments. The assemblages are representative of reef flats close to the shoreline and reef slopes around Bidong Island. CCA analysis indicated that the variation in substrate types between the natural reef of Bidong Island and the artificial reef (shipwreck) was the primary factor influencing foraminiferal distribution. The foraminiferal assemblages in the deeper parts of the natural reef showed greater similarity to those recorded around the shipwreck.\u003c/p\u003e \u003cp\u003eStudies suggest that with sufficient time, artificial reefs may become a good surrogate for the close natural reef environment, promoting balanced ecosystems between natural and artificial reefs [\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e]. However, despite being \u0026gt;\u0026thinsp;500 years old, the Bidong Shipwreck does not exhibit a foraminiferal assemblage similar to that of the nearby natural reef. The differences in substrate type and water depth between the shipwreck and natural reef caused a shift in foraminiferal assemblages. Additionally, the foraminiferal diversity around the shipwreck was notably lower than that in the natural reef environment. This suggests that while artificial reefs can have beneficial effects on fish biomass, their impact on protists, such as foraminifera, can be markedly different.\u003c/p\u003e \u003c/div\u003e"},{"header":"6. Conclusion","content":"\u003cp\u003eOur findings demonstrate distinct differences in the composition and diversity of foraminiferal assemblages between the artificial reef at the Bidong Shipwreck and the natural reefs of Bidong Island. The artificial reef, despite being \u0026gt;\u0026thinsp;500 years old, harboured a less diverse foraminiferal assemblage, predominantly consisting of \u003cem\u003eAmphistegina\u003c/em\u003e spp. with an absence of \u003cem\u003eCalcarina\u003c/em\u003e spp. This is in stark contrast to natural reefs, which exhibit higher diversity, including species such as \u003cem\u003eNummulites\u003c/em\u003e and \u003cem\u003ePeneroplis\u003c/em\u003e and a balance of symbiont-bearing and heterotrophic taxa. This study identified substrate type and water depth as the primary environmental parameters influencing foraminiferal distribution. The Bidong Shipwreck, located in deeper waters with higher sedimentation rates, supports the adaptation of foraminifera to lower light conditions and higher mud content. Conversely, natural reefs with their sandy substrates and varying depths support a more diverse and balanced assemblage of foraminifera. The artificial reef at the Bidong Shipwreck, although beneficial to certain benthic communities, does not replicate the foraminiferal diversity observed in natural reefs. The distinct environmental conditions of these habitats play a critical role in shaping the distribution and diversity of foraminiferal assemblages.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eConflicts of interest\u003c/h2\u003e \u003cp\u003eNo conflict of interest\u003c/p\u003e\u003ch2\u003eFunding\u003c/h2\u003e \u003cp\u003eThis work was supported by the strategic research grant of UMT (UMT/SRG/2022/55384)\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eF.I.M contributed to sampling design, methodology, manuscript drafting, and statistical analysis.H.S. oversaw field sampling, sample collection, and secured funding. W.N.W.S. assisted with sampling design and provided data from Bidong Island research. F.A.B.A. conducted laboratory and data analysis. M.N.A.M, N.S.S. and S.Y. contributed to manuscript editing.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eWe are grateful to Department of National Heritage (JWN), UZMA Archaeological Research Sdn. Bhd. (UAR) and Hidrokinetik Sdn. Bhd. We would like to thank the professional SCUBA divers of UMT, Kamaruzzaman Arifin, Mohd Sahimi Abdul Rashid, Mohd Forzi Zahari, and Mohd Fauzi Romeli that help with the sample collection. Thank you to boat crews of UMT Discovery II and the land-support team from JWN, UAR, and UMT.\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eData will be made available upon request\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eEvans AJ, Lawrence PJ, Natanzi AS, Moore PJ, Davies AJ, Crowe TP et al. Replicating natural topography on marine artificial structures \u0026ndash; a novel approach to eco-engineering. 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Micropaleontology. 2018;64(4):299\u0026ndash;316. doi: \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.47894/mpal.64.4.04\u003c/span\u003e\u003cspan address=\"10.47894/mpal.64.4.04\" 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":false,"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":"","lastPublishedDoi":"10.21203/rs.3.rs-5280024/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-5280024/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe success of artificial reefs is influenced by localised oceanographic factors including wave exposure, sedimentation rates, and proximity to natural reef environments. The Bidong Shipwreck, on the shallow shelf of the South China Sea, is a hotspot for meiobenthos. This study compared foraminifera distribution at the Bidong Shipwreck with that of the adjacent natural reef of Bidong Island. Foraminiferal assemblages at the shipwreck were less diverse. \u003cem\u003eAmphistegina\u003c/em\u003e spp. were the most dominant genus at both study sites. Calcarinids\u0026mdash;such as \u003cem\u003eCalcarina\u003c/em\u003e and \u003cem\u003eNeorotalia\u003c/em\u003e spp.\u0026mdash;occurred only on the natural reef. Cluster analysis categorised the foraminiferal assemblages from both sites into three distinct groups based on substrate preference. Group A consisted of foraminiferal assemblages from the shallow reef area of Bidong Island with a sandy substrate, where \u003cem\u003eNummulites\u003c/em\u003e and Peneroplis spp. were indicator species. Group B represented a mix of stations from both the shipwreck and natural reef, characterised by water depths\u0026thinsp;\u0026gt;\u0026thinsp;18 m and a muddy sand substrate, with \u003cem\u003eAmphistegina\u003c/em\u003e spp. comprising\u0026thinsp;~\u0026thinsp;80% of this assemblage. Group C displayed a more diverse distribution of foraminiferal species, including a mix of symbiont-bearing and heterotrophic taxa coexisting within the coarser sediments. Despite being over 500 years old, the Bidong Shipwreck does not exhibit a foraminiferal assemblage similar to that of the nearby natural reef. Differences in substrate type and water depth between these sites caused foraminiferal assemblage shifts. This suggests that although artificial reefs can have beneficial effects on fish biomass and meiobenthic communities, their impact on protists such as foraminifera can be markedly different.\u003c/p\u003e","manuscriptTitle":"Reef Foraminifera: Artificial vs. Natural Environments","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-11-15 16:47:58","doi":"10.21203/rs.3.rs-5280024/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":"d15ce913-f1ac-4ccc-99a2-1a06b5a39b74","owner":[],"postedDate":"November 15th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2024-12-02T10:53:13+00:00","versionOfRecord":[],"versionCreatedAt":"2024-11-15 16:47:58","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-5280024","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-5280024","identity":"rs-5280024","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}