Assessing the diversity of benthic foraminifera in coral reefs: a global perspective

Assessing the diversity of benthic foraminifera in coral reefs: a global perspective | 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 Assessing the diversity of benthic foraminifera in coral reefs: a global perspective Muhamad Naim Bin Abd Malek, Fabrizio Frontalini This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7863429/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 Utilizing a comprehensive dataset of species records from the Indo-Pacific and Atlantic regions, this work explores the diversity, distribution, and paleoenvironmental significance of benthic foraminifera across different coral reef ecosystems. Overall, 1,054 species of benthic foraminifera from coral ecosystems are recognized in this compilation, belonging to 371 genera, 123 families, and 13 orders. Although symbiont-bearing taxa such as Amphistegina , Heterostegina , and Calcarina are typical of reef environments, the dataset is dominated by small heterotrophic species. This compilation identifies several cosmopolitan taxa, including three opportunistic species ( Ammonia beccarii , Ammonia tepida , Elphidium advena ), with five symbiont-bearing ( Amphisorus hemprichii , Borelis pulchra , Borelis schlumbergeri , Heterostegina depressa , Peneroplis pertusus ) and 38 small heterotrophic species. European and American assemblages recorded the highest species diversity, with 391 and 356 species, respectively. The highest β-diversity was observed between the Atlantic and Pacific oceans (β = 0.83), followed by the Atlantic and Indian oceans (β = 0.81), whereas the Indian and Pacific oceans shared more species, with a lower β-diversity (β = 0.56). The findings highlight significant species turnover across continents and ocean basins, indicating high foraminiferal diversity and the influence of regional environmental gradients on assemblage composition in coral reef settings. Marine and Freshwater Ecology bleaching event symbiont-bearing carbonate biodiversity Indo-Pacific Atlantic Figures Figure 1 Figure 2 1. Introduction Coral reefs are among the most biologically diverse and ecologically important ecosystems on Earth, commonly referred to as the "rainforests of the sea" (Wooldridge, 2017 ). These complex bioconstructions, primarily composed of calcium carbonate laid down by corals and calcareous algae, support an immense array of marine life, housing an estimated 30% of all known marine species despite occupying less than 0.5% of the ocean floor (Bravo et al., 2021 ). They are primarily distributed in shallow tropical and subtropical waters, typically between latitudes 30°N and 30°S, with major reef systems located in the Indo-Pacific, Caribbean, and Red Sea regions (Ranjan et al., 2023 ). The Coral Triangle, located in the western Pacific Ocean, is commonly regarded as the global epicentre of marine biodiversity, which host a variety of species and endemisms (Pinheiro et al., 2019 ; Mills et al., 2023). Approximately 845 species of reef-building corals are distributed globally (DeVantier et al., 2020 ). These ecosystems, which offer ecosystem services like the availability of renewable resources like fisheries, shoreline protection from erosive processes, biogeochemical cycles like nitrogen fixation, and valuable cultural and recreational tourism opportunities, are essential to humankind (Elliff & Silva, 2017 ). Despite their importance, these vital ecosystems are facing unprecedented threats from both localized anthropogenic pressures and global climate change, threatening their structural integrity (Sous et al., 2017 ). The primary threats include ocean warming due to climate change, which leads to coral bleaching, ocean acidification, overfishing, coastal development, and pollution from land-based sources such as agricultural runoff and plastic debris. These stressors have contributed to widespread degradation of reef systems, with reports indicating that more than 50% of coral reefs globally have experienced significant decline over the past few decades (Aswani et al., 2015 ; Ranjan et al., 2023 ). The cumulative effect of these disturbances has pushed many coral reef systems to the brink of ecological collapse, which triggers urgent attention for comprehensive conservation and management strategies to enhance their resilience against future impacts (Chung et al., 2019 ). Coral cover and reef resilience have significantly decreased due to the increased frequency and severity of mass bleaching events, particularly those linked to strong El Niño phenomena (Heenan et al., 2017 ). Furthermore, reef habitats are still being harmed by destructive fishing methods like blast and cyanide fishing, especially in areas with weak enforcement capabilities (Carneiro & Martins, 2021 ). To mitigate this situation, several international and regional bodies are actively monitoring coral reef health and promote sustainable management. Some of the key organizations include the International Coral Reef Initiative (ICRI), the Global Coral Reef Monitoring Network (GCRMN), and the United Nations Environment Programme (UNEP), alongside regional programs such as the Coral Triangle Initiative (CTI). These governing bodies collaborate with national governments, NGOs, and research institutions in order to assess reef status, implement marine protected areas, and foster community-based conservation. However, effective management remains constrained due to limited funding, inconsistent enforcement, and the accelerating impacts of global climate change, urgent the need for coordinated international action to safeguard these vital ecosystems. Coral reefs are host to diverse assemblages of microorganisms, including coral-associated benthic foraminifera, a single-celled eukaryotes with calcareous shells that play a crucial role in reef sediment formation and biogeochemical cycling (Dawson et al., 2014 ; Langlet et al., 2020 ; Narayan et al., 2021 ). These foraminifera are particularly abundant in warm, shallow reef environments and have been commonly used as bioindicators of reef health due to their sensitivity to environmental changes such as water temperature, salinity, pH, and pollution levels (Hallock et al., 2003 ; A’ziz et al., 2021 ; Marín, 2023 ; Belart et al., 2025 ). Foraminiferal assemblages can reflect both natural variability and anthropogenic disturbances, making them essential in long-term reef monitoring programs (Prazeres et al., 2019 ). A decline in symbiont-bearing large benthic foraminifera, such as Amphistegina spp. and Marginopora spp., commonly parallels with coral bleaching events, signalling broader ecosystem stress (Girard et al., 2021 ). Inferring foraminiferal diversity in coral reef ecosystems enhances our understanding of micro- to macroscale ecological shifts and provides an additional layer of insight for conservation and restoration efforts (Hallock et al., 2003 ; Prazeres et al., 2019 ; Marín, 2023 ). Previously, biogeographical distribution of benthic foraminifera has been carried out from various settings (e.g., Culver & Buzas, 1999 ; Murray, 2001 , 2006 ; Gooday & Jorissen, 2011 ; Camacho et al., 2015 ; Frontalini et al., 2015 ; Kim et al., 2016; Förderer et al., 2018 ; Malek et al., 2021 ; Amao et al., 2019 , 2022 ; Mamo et al., 2023 ). However, despite these efforts, the compilation and synthesis of species-level data remain ambiguous, particularly due to regional biases, inconsistent taxonomic resolution, and limited integration of datasets across biogeographical provinces. The present study aims to bridge this gap by compiling and analyzing species-level occurrence records from a wide range of coral reef habitats across major ocean basins, thereby offering new insights into the global taxonomical diversity of benthic foraminifera in these sensitive ecosystems. This work is intended to provide a list of benthic foraminiferal species from coral reefs worldwide in order to assess the overall biodiversity and identify geographical pattern. 2. Materials and methods Compiling species-level data is a challenging and time-consuming task that is commonly hindered by inconsistencies in the reporting of species names in published studies. Scopus databases are used for collecting publication by using keywords such as “foraminifera”, “benthic foraminifera”, and “coral reefs”. The time frame was selected to include publication from the beginning of the year these keywords emerged. From the initial 428 documents retrieved, we excluded publications not directly related to benthic foraminifera in coral reef environments. To ensure consistency and relevance, we restricted the dataset to articles that specifically addressed species diversity and were published in the English language. The search strings used are shown below: (TITLE-ABS-KEY (benthic foraminifera) AND TITLE-ABS-KEY (coral reef)) AND PUBYEAR > 1980 AND PUBYEAR < 2025 AND (LIMIT-TO (DOCTYPE, "ar")) AND (LIMIT-TO (EXACTKEYWORD, "Foraminifera") OR LIMIT-TO (EXACTKEYWORD, "Coral Reef") OR LIMIT-TO (EXACTKEYWORD, "Benthic Foraminifera") OR LIMIT-TO (EXACTKEYWORD, "Coral Reefs") OR LIMIT-TO (EXACTKEYWORD, "Reefs") OR LIMIT-TO (EXACTKEYWORD, "Foraminifer") OR LIMIT-TO (EXACTKEYWORD, "Coral") OR LIMIT-TO (EXACTKEYWORD, "Reef") OR LIMIT-TO (EXACTKEYWORD, "Benthic Foraminifers") OR LIMIT-TO (EXACTKEYWORD, "Foraminifers") OR LIMIT-TO (EXACTKEYWORD, "Corals")) AND (LIMIT-TO (LANGUAGE, "English")). In some cases, authors chose not to include detailed species identifications in their work, either due to a focus on higher taxonomic levels or limitations in available taxonomic resources. This lack of uniformity complicates efforts to create comprehensive datasets and may lead to gaps in our understanding of the overall species diversity and distribution. To address these challenges, we selected publications for taxonomic review based on the following criteria: publications that include species names; publications that specify the sampling area as coral reefs; In addition, a compilation of species from cold-water reefs, as reported in the dissertation of Margreth (2010), was included. Systematic classification of the benthic foraminiferal species, genera, and the suprageneric taxonomic categories are adopted from the World Register of Marine Species (WoRMS, 2025). Each species and genus were verified against WoRMS, using only the accepted name. Open nomenclature species were not considered in the species lists compilation. The presence-absence list of taxa was compiled and used for the β diversity (differentiation) and γ diversity (regional) calculation (Al-Enezi et al., 2020 ). The β diversity was calculated using the PAlaeontological STatistics (PAST) data analysis software (version 4.02) (Hammer et al., 2001 ) and expressed as the global β diversity (βw) (Whittaker, 1960), while γ diversity is computed as the total species richness. The βw represents the variation in species composition, whereby the highest difference is indicated with the high value of βw (Al-Enezi et al., 2020 ). In addition, Jaccard dissimilarity index ( D J ) was calculated, which ranges from 0 to 1, and provided the degree of overlap between the datasets, with higher values suggesting greater similarity (Jaccard, 1912 ). 3. Results 3.1 Data compilation A total of 317 documents were retrieved using the search strings, and the publication and citation trends from 1981 to 2024, are presented in Fig. 1 . The trend fluctuates annually but shows an overall general upward pattern in both publications and citations. The trends are tentatively divided into three phases. The first phase, from 1981 to 2000 represents the formative years of research output, during which only a small number of papers were published annually, commonly fewer than five per year. This period reflects the early exploration of the field, with publications appearing sporadically and a relatively low cumulative citation count. The second phase between 2001 and 2012 marked as a period of steady growth. During this time, the annual publication rate gradually increased, consistently exceeding five papers per year, with several peaks reaching 10 to 15 publications. In the recent phase (i.e., 2013 to 2024), the period is characterized by higher productivity, with annual publications frequently ranging between 15 to 25 documents. The cumulative citation curve continued to climb steeply, surpassing 9,000 citations by 2024, reflecting both the maturity of the field and its wider impact. Despite some fluctuations in annual outputs, the overall trend shows sustained and intensive research interest during this phase. Following the search criteria, 24 publications of benthic foraminifera from coral reefs were selected comprising seven locations from Atlantic Ocean (i.e., Norway, Puerto Rico, Ireland, Alboran Sea, Brazil, Panama, and Columbia), seven from Pacific Ocean (i.e., Malaysia, New Caledonia, Australia, Philippines, Ecuador, Polynesia, and Bora bora) and five from Indian Ocean (i.e., Iran, Indonesia, Tanzania, Maldives and Mozambique) (Fig. 2 ; Table S1). Overall, the data collection recorded 1,054 species (γ diversity) of benthic foraminifera that are represented by 371 genera, 123 families, and 13 orders (Table 1 ; Table S2). The most diverse order belonged to Rotaliida, with a total of 65 families, followed by Lituolida (15 families) and Miliolida (13 families). Moderate diversification was observed in orders like Astrorhizida (nine families), Textulariida (six families), and Nodosariida (four families). The Order Loftusiida and Spirillinida was less common, with only two families, while the orders Robertinida, Allogromiida, Carterinida, and Vaginulinida were represented by a single family. Table 1 Benthic foraminiferal taxa recognized in coral reefs worldwide. Order Family Genus Species Allogromiida Allogromiidae 1 1 Astrorhizida Astrorhizoidea 1 1 Botellinidae 1 1 Hyperamminidae 1 1 Saccamminidae 3 5 Stannomidae 1 1 Rhabdamminidae 4 4 Hippocrepinellidae 1 1 Hyperamminidae 1 1 Psammosphaeridae 1 3 Carterinida Carterinidae 2 3 Loftusiida Haddoniidae 1 1 Globotextulariidae 2 2 Lituolida Adercotrymidae 1 2 Ammosphaeroidinidae 4 6 Trilocularenidae 1 1 Discamminidae 2 2 Haplophragmoididae 2 5 Hormosinellidae 1 1 Hormosinidae 3 7 Lituolidae 3 3 Nouriidae 1 1 Placopsilinidae 1 1 Prolixoplectidae 1 1 Remaneicidae 2 2 Spiroplectamminidae 3 3 Trochamminidae 10 19 Verneuilinidae 2 8 Miliolida Alveolinidae 2 5 Cornuspiridae 2 4 Cribrolinoididae 1 4 Fischerinidae 6 10 Hauerinidae 47 268 Miliamminidae 1 2 Miliolidae 1 1 Nubeculariidae 4 4 Ophthalmidiidae 3 3 Peneroplidae 7 23 Riveroinidae 1 5 Soritidae 6 10 Spiroloculinidae 2 38 Nodosariida Ammolagenidae 1 1 Lagenidae 5 16 Nodosariidae 6 9 Reophacidae 1 1 Polymorphinida Ellipsolagenidae 11 42 Glandulinidae 2 3 Polymorphinidae 5 12 Robertinida Robertinidae 4 6 Rotaliida Acervulinidae 5 8 Alabaminidae 3 3 Almaenidae 1 1 Ammoniidae 5 17 Anomalinidae 2 10 Amphisteginidae 1 6 Asterigerinidae 1 1 Asterigerinatidae 2 2 Astrononionidae 1 2 Baggininae 1 1 Bolivinitidae 9 32 Bolivinellidae 1 1 Buliminidae 3 8 Buliminoididae 1 1 Calcarinidae 6 18 Cancrisidae 3 10 Cassidulinidae 7 15 Cibicididae 7 28 Chilostomellidae 1 1 Cymbaloporidae 2 5 Discorbidae 2 4 Discorbinellidae 2 6 Elphidiellidae 1 2 Elphidiidae 3 41 Epistomariidae 3 5 Eponididae 7 14 Gavelinellidae 4 8 Glabratellidae 2 3 Globigerinitidae 2 2 Globobuliminidae 1 3 Heronalleniidae 1 1 Homotrematidae 2 3 Haynesinidae 1 4 Melonidae 1 3 Murrayinellidae 1 2 Mississippinidae 1 1 Nonionidae 7 16 Notorotaliidae 1 1 Nummulitidae 7 13 Pavoninidae 3 4 Pegidiidae 1 1 Placentulinidae 2 2 Planorbulinidae 3 4 Planulinidae 1 1 Pseudoparrellidae 3 3 Pulleniidae 1 6 Reussellidae 1 4 Rosalinidae 13 42 Rotaliidae 1 4 Siphoninidae 2 8 Siphogenerinoididae 3 4 Sphaeroidinidae 1 3 Stainforthiidae 2 3 Svratkinidae 1 2 Tortoplectellidae 1 2 Tosaiidae 1 1 Trimosinidae 1 1 Turrilinidae 1 2 Ungulatellidae 1 1 Uvigerinidae 2 9 Victoriellidae 1 2 Epistominidae 1 2 Ammodiscidae 2 2 Patellinidae 1 1 Planispirillinidae 1 3 Spirillinida Ammodiscidae 1 1 Spirillinidae 4 9 Textulariida Eggerellidae 5 33 Kaminskiidae 1 1 Olgiidae 1 1 Pseudogaudryinidae 5 8 Textulariidae 5 9 Valvulinidae 2 2 Vaginulinida Vaginulinidae 10 17 TOTAL 371 1,054 Bold families indicate symbiont-bearing type. Figure 2 : (A) Worldwide corals distribution modified from Allen Coral Atlas ( 2022 ). Red shaded colour showed cold-water corals while green shaded colour indicates the warm-water corals. (B) Map showing location of studied benthic foraminifera from coral reefs worldwide: Malaysia: 1 -Tioman Island, 2 - Bidong Island, 3 - Redang Island; Philippines: 4 - Palawan, 5 - Nogas Island; Indonesia: 6 - Tambelan Island, 7 - Kepulauan Seribu, 8 - Bali; Australia: 9 - Great Barrier Reef, 10 - Moreton Bay; 11 - New Caledonia; Polynesia: 12 - Niue; French Polynesia: 13 - Bora bora; 14 - Galapagos Island; Panama: 15 - Punta Galeta; Colombia: 16 - Isla Barú; Puerto Rico: 17 - Jobos Bay; Brazil: 18 - Pirangi and Maracajaú; Ireland: 19 - Rockall Bank; Norway: 20 - Norway Margin; 21- Alboran Sea; 22 - Qeshm Island; Maldives: 23 - Ari Atoll, 24 - Rasdhoo Atoll; Tanzania: 25 - Zanzibar; Ecuador: 26 - Galapagos Archipelagos. The family Hauerinidae represented the most taxonomically diverse group, comprising 47 genera, the highest recorded among benthic foraminifera family. Notably, it encompassed a wide range of genera, including Quinqueloculina (109 species), Triloculina (35 species), Pyrgo (17 species), and Miliolinella (14 species). Other diverse genera were Elphidium with 31 species and Textularia with 27 species as well as Rosalina (16 species), Bolivina (17 species), and Cibicides (12 species). Despite the occurrence of symbiont-bearing foraminifera in coral reefs, these families only represented a minor part of the species. Among them were reported Alveolinidae (i.e., Alveolinella , Borelis ), Peneroplidae (i.e., Coscinospira , Dendritina , Euthymonacha , Laevipeneroplis , Monalysidium , Peneroplis , Parasorites ), Amphisteginidae (i.e., Amphistegina ), Nummulitidae (i.e., Heterostegina , Nummulites , Planostegina , Operculina , Operculinella , Neoassillina , Planoperculina ), Calcarinidae (i.e., Baculogypsina , Baculogypsinoides , Calcarina , Neorotalia , Pararotalia , Schlumbergerella ), and Soritidae (i.e., Archaias , Amphisorus , Cycloputeolina , Cyclorbiculina , Marginopora , Sorites ). 3.2 Distribution of species 3.2.1 European assemblages The benthic foraminiferal assemblages on European coral reefs were studied at three locations in Norwegian margin reefs (Margreth, 2010; Spezzaferri et al., 2013 ), Rockall Bank, Ireland (Margreth, 2010; Morigi et al., 2011 ), and Alboran Sea (Margreth, 2010; Stalder et al., 2021 ). These studies site consisted of cold-water coral reefs assemblages. The total number of species were high with 391 species: 180 species in Rockall Bank, 278 species in Porcupine Seabight and 173 species in Alboran Sea. Dominant species in Rockall Bank were Epistominella exigua , Globocassidulina subglobosa , Cassidulina carinata , Cassidulina laevigata , Discanomalina coronata , while Porcupine Seabight with Alabaminella weddellensis , Adercotryma wrighti , Ehrenbergina carinata , Paratrochammina challenger , Planulina ariminensis , Spirillina vivipara , and Trochammina inflata . Meanwhile, in Alboran Sea, deep-water species such as Bulimina marginata, Bolivina dilatata, Bolivina striatula , Bolivina alata , Cassidulina crassa , C. laevigata , Rectuvigerina elongatastriata , Uvigerina peregrina , and Globobulimina doliolum were dominant. The cold-water coral assemblages mainly consisted of small heterotrophic and opportunistic species. 3.2.2 America assemblages Four locations of reefs have been studied, Jobos Bay, Puerto Rico (Marin et al., 2024), Pirangi and Maracajaú, Brazil (Eichler & de Moura, 2020 ), Punta Galeta, Panama (Elsa et al., 2024 ) and Galapagos Island, Ecuador (Humphreys et al., 2019 ). Overall, 356 species has been identified with Ecuador recorded highest species (n = 142 species), followed by Panama (n = 88 species), Puerto Rico (n = 73), and Brazil (n = 53 species). Only three species were found at all locations (i.e., Quinqueloculina laevigata , Quinqueloculina lamarckiana , and Triloculina trigonula ). The heterotrophic assemblages in American reefs were the most diverse dominated with genera like Quinqueloculina , Spiroloculina , and Triloculina , while symbiont-bearing taxa recorded such as Amphisorus hemprichii , Amphistegina gibbosa , Archaias angulatus , Borelis pulchra , Borelis schlumbergeri , Borelis clarionensis , Heterostegina antillarum , Heterostegina depressa , Heterostegina curva , Laevipeneroplis proteus , Peneroplis carinatus , Peneroplis pertusus , Sorites marginalis , Sorites marginata , Planogypsina acervalis , and Vertebrasigmoilina mexicana . 3.2.3 African assemblages Two sites were studied in the African region: the Zanzibar Archipelago (Narayan et al., 2022 ) and Bazaruto Archipelago, Mozambique (Langer et al., 2013 ). A total of 124 species of benthic foraminifera were found in Zanzibar, belonging to 56 genera while slightly lower species in Mozambique with 95 species belonging to 63 genera. Both Amphistegina lessonii and Amphistegina lobifera were dominant in the assemblage. Although a large number of small heterotrophic species was found, the number of symbiont-bearing and opportunistic taxa were almost similar. In Zanzibar, symbiont-bearing taxa were Alveolinella quoyi , A. hemprichii , A. lessonii , A. lobifera , Amphistegina papillosa , Amphistegina radiata , Coscinospira hemprichii , H. depressa , Marginopora vertebralis , Neorotalia calcar , Operculina ammonoides , P. pertusus , Peneroplis planatus , and Sorites orbiculus , while eight species were found in Mozambique (i.e., A. hemprichii , C. hemprichii , H. depressa , N. calcar , P. planatus , P. acervalis , S. orbiculus , and Sorites variabilis ). Opportunistic taxa were Ammonia beccarii , Ammonia tepida , Elphidium advena , Elphidium craticulatum , Elphidium crispum , Elphidium fichtelianum , Elphidium limbatum , Elphidium striatopunctatum , Elphidium macellum , Elphidium milletti , Euloxostomum pseudobeyrichi , Elphidium hispidulum , Reusella spinulosa , Sagrina zanzibarica , Sagrinella durrandii , and Sagrinella jugosa. 3.2.4 Asian assemblages The Asian region has the highest number of location studied for coral reefs foraminifera, including three coral reefs in Malaysia (Bidong Island (Husain et al., 2022 ), Tioman Island (A'ziz et al., 2021), Redang Island (Minhat et al., 2024 ), three in Indonesia (i.e., Riau (Junita et al., 2020 ), Kepulauan Seribu (Renema, 2008 ) and Bali (Renema, 2003 ), two in the Philippines (Palawan (Förderer & Langer, 2019 ) and Nogas Island (Gonzales et al., 2022 ), two at Maldives atolls (Parker & Gischler, 2011; Giraldo-Gomez et al, 2024), and one at each Iran (Maghsoudlou et al., 2021 ) and Egypt (Badr El Din & Hallock, 2024). It corresponded of total 264 species, with eight species frequently occurred (i.e., A. lessonii , A. lobifera , E. craticulatum , H. depressa , N. calcar, P. pertusus , P. planatus , S. orbiculus ). The assemblages consisted of a variety of symbiont-bearing taxa dominated by A. lessonii , A. radiata , N. calcar , and O. ammonoides , including a new species that was recently described ( Peneroplis hoheneggeri nov. sp.). The genera Ammonia and Elphidium were commonly found as the opportunistic taxa and mainly represented by A. beccarii , Ammonia convexa , Ammonia parkinsoniana , Ammonia supera , A. tepida , E. advena , E. craticulatum , E. crispum , E. excavatum , E. fichtelianum , Elphidium gerthi , E. hispidulum , Elphidium neosimplex , and Elphidium striatopunctatum . 3.2.5 Oceanian assemblages The largest coral reef system, the Great Barrier Reef, has been extensively studied (Uthicke & Nobes, 2008 ). Other locations, such as Moreton Bay (Narayan & Pandolfi, 2010 ), Polynesia (Oron et al., 2024 ), Bora Bora (Parker & Gischler, 2024 ), New Caledonia (Debenay, 2012 ), and Papua New Guinea (Langer & Lipps, 2003), have also been investigated. The highest number of species recorded in these regions, with Australia exhibiting the greatest diversity (n = 135 species), followed by Bora Bora (n = 118 species), New Caledonia (n = 69 species) and Polynesia (n = 55 species). Five species frequently occurred, including Hauerina pacifica, Miliolinella oceanica, Quinqueloculina parkeri , and Sorites orbiculus . The region also recorded high numbers of symbiont-bearing taxa (n = 34 species), dominated with taxa such as A. lessonii , A. papillosa , A. radiata , N. calcar , Calcarina hispida , M. vertebralis , and S. orbiculus . Assemblages from Bora Bora were notable for a high number of small heterotrophic taxa (n = 110 species), contributing to a total of 319 taxa recorded in the Oceania region, including the identification of a new species, Textularia boraboraensis nov. sp. 3.3 Pattern of similarity/dissimilarity 3.3.1 Comparison between continents The β index values revealed substantial species turnover in foraminiferal assemblages across the continents (Table 2 ). The global β w calculated was 2.5537, reflecting average dissimilarity among regions. The highest dissimilarities were observed between Europe and Africa (β = 0.92), followed closely by Europe vs. Americas and Europe vs. Asia (β = 0.89), suggesting minimal species overlap and high community distinctiveness. Europe vs. Oceania also showed considerable dissimilarity (β = 0.88), reinforcing Europe’s unique biogeographic identity and environmental characteristics. Among the remaining continents, Americas vs. Africa (β = 0.81), Americas vs. Oceania (β = 0.81), and Americas vs. Asia (β = 0.79) reflected moderate dissimilarity. The lowest β values occurred between Africa vs. Oceania (β = 0.60), Africa vs. Asia (β = 0.55), and Asia vs. Oceania (β = 0.54), indicating relatively greater species overlap and compositional similarity within these tropical or subtropical regions. Table 2 Values of β diversity and \(\:\stackrel{-}{D}\) J for each continental region. β w = 2.5537 Europe Americas Africa Asia Oceania Europe - Americas 0.89 - Africa 0.92 0.81 - Asia 0.89 0.79 0.55 - Oceania 0.88 0.81 0.60 0.54 - \(\:\stackrel{-}{\varvec{D}}\) J =0.86 Europe Americas Africa Asia Oceania Europe - Americas 0.94 Africa 0.96 0.89 Asia 0.94 0.89 0.71 Oceania 0.93 0.89 0.75 0.70 - β w = Whittaker global diversity \(\:\stackrel{-}{D}\) J = Average Jaccard dissimilarity Complementing these findings, the D J mirrored similar trends. The global Jaccard dissimilarity ( \(\:\stackrel{-}{D}\) J = 0.86) suggested that foraminiferal communities are relatively similar overall. Europe vs. Africa again exhibited the highest dissimilarity ( D J = 0.96), with only 23 shared species, followed by Europe vs. Americas ( D J = 0.94), Europe vs. Asia ( D J = 0.94), and Europe vs. Oceania ( D J = 0.93). The Americas assemblages showed high dissimilarity with other continents ( D J = 0.89). Comparisons among Africa, Asia, and Oceania yielded the lowest dissimilarity scores, i.e., Africa vs. Oceania ( D J = 0.75), Africa vs. Asia ( D J = 0.71), and Asia vs. Oceania ( D J = 0.70), consistent with shared biogeographical patterns across the Indo-Pacific realm. Both indices consistently suggest that Europe harbors a distinct foraminiferal assemblage, while Africa, Asia, and Oceania show greater ecological and compositional continuity, potentially reflecting their geographic proximity and environmental connectivity. 3.3.2 Comparison between oceans Similarly, the comparison between the main ocean body followed similar trends with β w value slightly lower (β w = 1.3526) (Table 3 ). The highest dissimilarity values were observed between the Atlantic and Pacific Oceans (β = 0.83) and Atlantic and Indian Oceans (β = 0.81), suggesting distinct foraminiferal assemblages with limited species overlap in these regions. In contrast, the Pacific and Indian Oceans exhibited a notably lower β value of 0.56, reflecting greater similarity and shared species between these two adjacent oceanic regions. Table 3 Values of β diversity and \(\:\stackrel{-}{D}\) J for oceanic body. β w = 1.3526 Atlantic Indian Pacific Atlantic - Indian 0.81 - Pacific 0.83 0.56 - \(\:\stackrel{-}{\varvec{D}}\) J =0.84 Atlantic Indian Pacific Atlantic - Indian 0.90 - Pacific 0.91 0.72 - β w = Whittaker global diversity \(\:\stackrel{-}{D}\) J = Average Jaccard dissimilarity These patterns are supported by the D J , which similarly showed high dissimilarity between the Atlantic and Indian Oceans ( D J = 0.90) and the Atlantic and Pacific Oceans ( D J = 0.91), reinforcing the distinctness of the Atlantic fauna compared to the Indo-Pacific realms. The Pacific vs . Indian Ocean comparison again showed the lowest dissimilarity ( D J = 0.72), consistent with a higher degree of faunal overlap, likely due to the more direct connectivity and environmental continuity between these two ocean basins. 3.3.3 Cosmopolitan distribution Three opportunistic species were recorded across oceanic regions (i.e., A. beccarii , A. tepida , E. advena ), with five symbiont-bearing (i.e., A. hemprichii , B. pulchra , B. schlumbergeri , H. depressa , P. pertusus ) and 48 small heterotrophic species. Among the small heterotrophic species recorded in all major oceans worldwide were A. beccarii, A. tepida, Articulina pacifica, B. striatula, Bolivina variabilis, Cancris auricula, Cibicides refulgens, Cornuspira planorbis, Cycloforina granulocostata, Cymbaloporetta squamosa, Lobatula lobatula, Melonis affinis, Pyrgo oblonga, Quinqueloculina agglutinans, Quinqueloculina bicarinata, Quinqueloculina bosciana, Q. lamarckiana, Q. parkeri, Quinqueloculina philippinensis, Quinqueloculina poeyana, Quinqueloculina seminulum, Rosalina globularis, Rotorbis auberii, Spiroloculina antillarum, Spiroloculina communis, Spiroloculina corrugata, Textularia agglutinans, Textularia candeiana, Textularia pseudogramen, Triloculina rotunda, Triloculina tricarinata , and T. trigonula . 4. Discussion 4.1 Biogeographic patterns of foraminifera in coral reefs The global distribution of benthic foraminiferal assemblages in coral reefs reveals distinct regional patterns in terms of species richness, dominant taxa, and ecological strategies (i.e., heterotrophic, symbiont-bearing, and opportunistic). The highest overall species richness is recorded in Europe (γ = 391) and Oceania (γ = 319), followed by America (γ = 356), Asia (γ = 264), and Africa (γ = 191). However, these figures are strongly influenced by environmental settings, sampling effort and ecological diversity within study sites. European cold-water coral reefs (e.g., Rockall Bank, Porcupine Seabight) are dominated by small heterotrophic taxa, as symbiont-bearing forms are absent at bathyal depths. This pattern reflects the oligotrophic, deep-water environments where cold-water corals prevail, limiting light availability necessary for photosymbionts (Chen & Lin, 2017 ; Avnaim-Katav et al., 2020 ; Vicente et al., 2021 ). In contrast, tropical reefs in America, Africa, Asia, and Oceania exhibit mixed assemblages with varying proportions of symbiont-bearing and opportunistic taxa, consistent with warmer, shallower reef habitats. In American reefs, although the number of shared species among sites was low, the dominance of miliolid genera such as Quinqueloculina, Triloculina , and Spiroloculina reflects a robust heterotrophic community. Several larger symbiont-bearing taxa, including A. angulatus and H. depressa , suggest healthy carbonate platforms and reef systems. African reefs, particularly from the Zanzibar and Bazaruto Archipelagos, show comparable proportions of heterotrophic, symbiont-bearing, and opportunistic taxa. The occurrences of both Amphistegina and Elphidium species points to transitional reef conditions, potentially affected by both coral reef development and environmental disturbances (Murray, 2006 ; Girard et al., 2021 ; O’Brien et al., 2021 ). Asian assemblages, being the most geographically diverse in the study, harbor a relatively high diversity of both symbiont-bearing and opportunistic foraminifera. The consistent presence of taxa such as A. lessonii, N. calcar , and P. planatus across multiple sites underlines their resilience and ecological success in Indo-Pacific reefs. In Oceania, particularly in the Great Barrier Reef and Bora Bora, a high diversity of both small heterotrophic and symbiont-bearing taxa is observed. The identification of new species (e.g., T. boraboraensis ) underscores the region’s importance as a hotspot for foraminiferal diversity. The overlap of species such as S. orbiculus and P. acervalis across multiple sites indicates broad ecological tolerances and potential for wide biogeographic dispersal (Alve & Goldstein, 2003 ; Murray, 2013 ; Prazeres et al., 2020 ). 4.2 Patterns of similarity and turnover Beta diversity analyses reveal clear biogeographic segregation among regions. The higher similarity of benthic foraminiferal assemblages in certain areas can be attributed to the lack of recent and past geographic or environmental separation. Such limited separation reduces habitat heterogeneity and environmental gradients, leading to lower beta diversity and greater faunal overlap among sites (Renema, 2006 ; Uthicke et al., 2009 ; Diaz et al., 2024 ). Europe exhibits the highest species turnover, when compared to other continents, that can be ascribed to the peculiar environmental setting hosting cold-water reef ecosystems. In contrast, Asia, Africa, and Oceania exhibit a lower turnover (β ≈ 0.54–0.60), suggesting regional continuity and biogeographic connectivity in the Indo-Pacific (Förderer et al., 2018 ). Oceanic comparisons further reinforce this pattern. The Atlantic Ocean stands out as the most distinct, exhibiting the least similarity with both the Indian and Pacific Oceans ( D J ≈ 0.90–0.91), likely due to historical isolation and contrasting environmental conditions. Conversely, Pacific and Indian Ocean reefs share a high degree of similarity ( D J = 0.72), reflecting the contiguous reef systems across Southeast Asia and Melanesia. The pronounced dissimilarity between Atlantic and Pacific benthic foraminiferal assemblages likely reflects the closure of the Isthmus of Panama (~ 3 Ma), which terminated faunal exchange between the two oceanic realms, reorganized ocean circulation, and created distinct environmental regimes that promoted biogeographic divergence (Collins et al., 1996 ; McDougall, 1996 ; Bornmalm, 1997 ; Groeneveld et al., 2014 ). 4.3 Cosmopolitan taxa and ecological indicators Despite regional differences, several cosmopolitan species are recorded across all oceanic regions, indicating their ecological behaviors. Notably, three opportunistic taxa ( A. beccarii, A. tepida , and E. advena ) and five symbiont-bearing species ( A. hemprichii, B. pulchra, B. schlumbergeri, H. depressa, P. pertusus ) are found globally. These taxa are commonly associated with both healthy reef conditions (in the case of larger symbiont-bearers) and environments under anthropogenic stress (in the case of opportunists). The compiled dataset (n = 1,054 species) indicates that the coral reefs is the hotspot for biodiversity, as this number is higher than previous compilation from other location such as Gulf of Mexico (n = 987 species) (Gupta & Smith, 2010 ), Sahul Shelf and Timor Sea (n = 946 species) (Loeblich & Tappan, 1994 ), (n = 304 species) Korean Peninsula ( Kim et al., 2016) and Aegen Sea, Mediterranean (n = 267 species). However, it is important to note that these studies were not exclusively based on carbonate platform environments, where foraminifera typically reach their highest diversity. A slightly higher species diversity has been reported by Dorst et al. (2013) from continental shelf of North-East Atlantic with 1,486 species. This highlights the ecological adaptability of benthic foraminifera, which can thrive across a wider range of substrates than previously emphasized. By contrast, studies restricted to larger benthic foraminifera (n = 105 species) have identified peak richness in the Western Coral Triangle and Sahul Shelf provinces of the Central Indo-Pacific, reflecting the well-established role of these provinces as global biodiversity hotspots (Förderer et al., 2022). Interestingly, even in the oligotrophic Arabian Gulf, more than 492 benthic foraminiferal species have been recorded, and these assemblages extend beyond reef environments into adjacent habitats (Amao et al., 2025). Comparable findings have also been reported in the Mediterranean Sea, where benthic foraminiferal assemblages associated with cold-water coral ecosystems, vermetid reef platforms, and algal-dominated hard substrates display unexpectedly high diversity beyond classic reefal systems (Stalder et al., 2021 ; Rossbach et al., 2022; Manda et al., 2024 ). This highlights that high species richness is not confined to classic reefal systems but can also emerge in marginal and environmentally extreme settings. Nonetheless, it is important to acknowledge that these sensitive coral environments remain poorly studied, and many taxa have not yet been identified to the species level, which likely underestimates true diversity. Moreover, none of the existing studies accounted for allogromids (i.e., naked forams). This suggests that the actual number of taxa is expected to be much higher than currently reported. Foraminiferal assemblages in coral reef environments are predominantly composed of a mixture of calcareous hyaline and porcelaneous forms, particularly among the small heterotrophic species, consistent with the observations of Murray ( 2006 ). Although symbiont-bearing foraminifera are ecologically dominant in these habitats, the most numerically abundant group consists of small heterotrophic species, with a minor proportion of opportunistic taxa. The extensive list of small heterotrophic species with global distributions (n = 38) further highlights the ecological flexibility and dispersal capability of these taxa. Some of these, such as L. lobatula , Q. philippinensis , and S. communis , are well-known for their euryhaline and eurytopic nature. By 2023, the projection that nearly 90% of global reefs will face degradation was alarming (Narayan et al., 2021 ). The ocean warming, ocean acidification, and habitat loss threaten coral reefs where increasing coral bleaching event frequency occurred worldwide since 1998 (Eakin et al., 2019 ; Thangadurai et al., 2024 ). Symbiont-bearing taxa such as Amphistegina , Sorites and Peneroplis are highly sensitive to thermal stress and bleaching (Hallock, 2000 ; Schmidt et al., 2016 ; Narayan et al., 2021 ), while acidification reduces calcification and test integrity (Prazeres et al., 2015 ; Guamán-Guevara et al., 2019 ). As coral cover declines, the loss of reef habitat further reduces assemblage diversity (Doo et al., 2020 ), with potential consequences for carbonate production and reef resilience. Consequently, climate-driven reef degradation poses a critical risk to benthic foraminiferal diversity, particularly for symbiont-bearing species that serve as key bioindicators of reef health. 4.4 Limitation Despite the inclusion of a high number of publications, one of the key limitations of this study is the presence of incomplete or inconsistent species names in the original sources, which may have constrained the comprehensiveness and accuracy of the species database. Although we carefully selected and validated species lists from each continent and ocean basin, the reliance on published records introduces potential biases due to taxonomic discrepancies, synonymy, or unidentified taxa. Furthermore, while the analysis captures broad-scale biogeographic patterns, the generalizations derived from this dataset may overlook finer ecological or regional variations. Additionally, several other limitations should be noted: (a) naked foraminifera were not accounted for in the reviewed studies; (b) sampling efforts in some regions remain comparatively lower than in others; (c) many species are still left in open nomenclature, which further obscures true diversity; and (d) differences in sampling strategies across studies may introduce inconsistencies in diversity estimates. As such, caution is advised in interpreting the findings as fully representative of global foraminiferal diversity. Future efforts should aim to integrate molecular data, high-resolution taxonomic revisions, and standardized sampling protocols to enhance the reliability of beta diversity assessments. 5. Conclusion This global synthesis of benthic foraminiferal assemblages from coral reefs reveals significant regional differences driven by environmental conditions, reef type (cold-water vs. tropical), and biogeographic history. High β-diversity values, particularly between Europe and other continents, reflect strong species turnover and endemism. Conversely, the Indo-Pacific region demonstrates high compositional similarity, suggesting ecological connectivity and potential for dispersal among reef systems. The consistent presence of specific symbiont-bearing and opportunistic taxa across regions suggests their potential as universal bioindicators for reef health and environmental change. Moreover, the discovery of new taxa highlights the need for continued exploration and taxonomic resolution, particularly in under-studied regions. Future studies should focus on integrating molecular approaches to complement traditional taxonomy, assess cryptic diversity, and explore population connectivity across coral reef systems. Additionally, monitoring shifts in assemblage composition in response to climate change and anthropogenic impacts will be crucial for understanding reef ecosystem resilience and guiding conservation strategies. Declarations Conflict of interest The authors declare that they have no conflict of interest. Consent to participate The authors are agreeing to be accountable for all aspects of the work in ensuring that questions related to the accuracy or integrity of any part of the work are appropriately investigated and resolved. Consent to publish The authors provides consent to publish the data in this manuscript. 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Sedimentology 69(1):121–161. https://doi.org/10.1111/sed.12858 O’Brien PaJ, Asteman IP, Bouchet VMP (2021) Benthic Foraminiferal Indices and Environmental Quality Assessment of Transitional Waters: A review of current challenges and future research perspectives. Water 13(14):1898. https://doi.org/10.3390/w13141898 Oron S, Friedlander AM, Sala E, Goodman-Tchernov BN (2024) Shallow water foraminifera from Niue and Beveridge Reef (South Pacific): insights into ecological significance and ecosystem integrity. Royal Soc Open Sci 11(1). https://doi.org/10.1098/rsos.230997 Parker JH, Gischler E (2010) Modern foraminiferal distribution and diversity in two atolls from the Maldives. Indian Ocean Marine Micropaleontology 78(1–2):30–49. https://doi.org/10.1016/j.marmicro.2010.09.007 Parker JH, Gischler E (2024) Taxonomy and systematics of shallow-water tropical benthic foraminifera from the lagoon environments at Bora Bora, Society Islands, French Polynesia. Micropaleontology 70(4):301–404. https://doi.org/10.47894/mpal.70.4.01 Pinheiro HT, Shepherd B, Castillo C, Abesamis RA, Copus JM, Pyle RL, Greene BD, Coleman RR, Whitton RK, Thillainath E, Bucol AA, Birt M, Catania D, Bell MV, Rocha LA (2019) Deep reef fishes in the world’s epicenter of marine biodiversity. Coral Reefs 38(5):985–995. https://doi.org/10.1007/s00338-019-01825-5 Prazeres M, Uthicke S, Pandolfi JM (2015) Ocean acidification induces biochemical and morphological changes in the calcification process of large benthic foraminifera. Proceedings of the Royal Society B: Biological Sciences , 282(1803), 20142782. https://doi.org/10.1098/rspb.2014.2782 Prazeres M, Martínez-Colón M, Hallock P (2019) Foraminifera as bioindicators of water quality: The FoRAM Index revisited. Environ Pollut 257:113612. https://doi.org/10.1016/j.envpol.2019.113612 Prazeres M, Morard R, Roberts TE, Doo SS, Jompa J, Schmidt C, Stuhr M, Renema W, Kucera M (2020) High dispersal capacity and biogeographic breaks shape the genetic diversity of a globally distributed reef-dwelling calcifier. Ecol Evol 10(12):5976–5989. https://doi.org/10.1002/ece3.6335 Ranjan D, Chandravanshi S, Verma P, Singh MB, Verma DK, Maurya P, Upadhyay AK, Raghunath N, Tiwari AK, Sahu KK (2023) Effects of Coral Reef Destruction on Humans and the Environment. Int J Environ Clim Change 13(10):716–725. https://doi.org/10.9734/ijecc/2023/v13i102708 Renema W (2003) Larger foraminifera on reefs around Bali (Indonesia). Zoölogische Verhandelingen 345:337–366. https://www.narcis.nl/publication/RecordID/oai%3Arepository.naturalis.nl%3A220323 Renema W (2006) Habitat variables determining the occurrence of large benthic foraminifera in the Berau area (East Kalimantan, Indonesia). Coral Reefs 25(3). https://doi.org/10.1007/s00338-006-0119-4 Renema W (2008) Habitat selective factors influencing the distribution of larger benthic foraminiferal assemblages over the Kepulauan Seribu. Mar Micropaleontol 68(3–4):286–298. https://doi.org/10.1016/j.marmicro.2008.06.002 Rodríguez L, Acosta A, González-Zapata FL, Gómez-Corrales M, Marrugo M, Alvarado-Ch EM, Dueñas LF, Andrade J, Gutierrez-Cala L, Sánchez JA (2023) Conservation at the edge: connectivity and opportunities from non-protected coral reefs close to a National Park in the Colombian Caribbean. Biodivers Conserv 32(5):1493–1522. https://doi.org/10.1007/s10531-022-02539-x Rossbach FI, Merk B, Wild C (2021) High diversity and abundance of foraminifera associated with Mediterranean benthic red algae mats. Diversity 14(1):21. https://doi.org/10.3390/d14010021 Schmidt C, Titelboim D, Brandt J, Herut B, Abramovich S, Almogi-Labin A, Kucera M (2016) Extremely heat tolerant photo-symbiosis in a shallow marine benthic foraminifera. Sci Rep 6(1). https://doi.org/10.1038/srep30930 Sous D, Chevalier C, Devenon J, Blanchot J, Pagano M (2017) Circulation patterns in a channel reef-lagoon system, Ouano lagoon, New Caledonia. Estuar Coastal Shelf Sci 196:315–330. https://doi.org/10.1016/j.ecss.2017.07.015 Spezzaferri S, Ruggeberg A, Stalder C, Margreth S (2013) Benthic Foraminifer Assemblages from Norwegian Cold-Water Coral Reefs. J Foraminifer Res 43(1):21–39. https://doi.org/10.2113/gsjfr.43.1.21 Stalder C, ElKateb A, Spangenberg JE, Terhzaz L, Vertino A, Spezzaferri S (2021) Living benthic foraminifera from cold-water coral ecosystems in the eastern Alboran Sea, Western Mediterranean. Heliyon 7(9):e07880. https://doi.org/10.1016/j.heliyon.2021.e07880 Thangadurai T, De K, Murugesan S, Peter PS, Pasiyappazham R, Selvin R, Jose J, P. A., Bellantuono A (2024) History of recurrent short- and long-term coral bleaching events in Indian coral reefs: a systematic review of contrasting bleaching patterns, lessons learned, and future directions. Estuar Coastal Shelf Sci 109112. https://doi.org/10.1016/j.ecss.2024.109112 Uthicke S, Nobes K (2008) Benthic Foraminifera as ecological indicators for water quality on the Great Barrier Reef. Estuar Coastal Shelf Sci 78(4):763–773. https://doi.org/10.1016/j.ecss.2008.02.014 Uthicke S, Thompson A, Schaffelke B (2009) Effectiveness of benthic foraminiferal and coral assemblages as water quality indicators on inshore reefs of the Great Barrier Reef, Australia. Coral Reefs 29(1):209–225. https://doi.org/10.1007/s00338-009-0574-9 Vicente TM, Yamashita C, De Mello E, Sousa SH, Ciotti AM (2021) Evaluation of the relationship between biomass of living (stained) benthic foraminifera and particulate organic matter vertical flux in an oligotrophic region, Campos Basin, southeastern Brazilian continental margin. J Sea Res 176:102110. https://doi.org/10.1016/j.seares.2021.102110 Wooldridge SA (2017) Instability and breakdown of the coral–algae symbiosis upon exceedence of the interglacial pCO2 threshold (> 260 ppmv): the missing Earth-System feedback mechanism. Coral Reefs 36(4):1025–1037. https://doi.org/10.1007/s00338-017-1594-5 World Register of Marine Species (WoRMS) Accessed at https://www.marinespecies.org on [24 October 2024]. https://doi.org/10.14284/170 Supplementary Table S1: Sources used for the global compilation of foraminiferal data in coral reefs Additional Declarations The authors declare no competing interests. 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Malek","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA6klEQVRIiWNgGAWjYFACHhDBDEKMD6BCBkRrYTZgSCBJCwMDmwRRWnRn5B78dKPGOo+/nTutmveHXT4De/M2Ccacwzi1mN3IS5bOOZZeLHGYd9ttnoRkywaeY2USjNvwackxkM5hO5zYANEC9I9EjhkhLca/c/4dTpwP1FLMk1BvwCD/hqAWM+nctsOJG4BamHkSDgNt4SGg5cwbM+vcvvTEjYd5N0vOSTtuwMaTVmyRuC0dt5bjOca3c75ZJ847f3bjhzc21Qb87Ic33vi4zRqnFkzABiISGJpJ0AIFdaRrGQWjYBSMguEKAB1XUVVYWlObAAAAAElFTkSuQmCC","orcid":"https://orcid.org/0000-0002-3730-8389","institution":"Universiti Tunku Abdul Rahman, Malaysia","correspondingAuthor":true,"prefix":"","firstName":"Muhamad","middleName":"Naim Bin Abd","lastName":"Malek","suffix":""},{"id":532782828,"identity":"0f37e37c-c8ae-434c-982f-de8156204d12","order_by":1,"name":"Fabrizio Frontalini","email":"","orcid":"https://orcid.org/0000-0002-0425-9306","institution":"University of Urbino, 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04:21:40","extension":"xml","order_by":9,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":565202,"visible":true,"origin":"","legend":"","description":"","filename":"rs78634290structuring.xml","url":"https://assets-eu.researchsquare.com/files/rs-7863429/v1/3a1b5e1dbd83b903ffb83711.xml"},{"id":94056691,"identity":"c8eb6b79-924b-4e43-8ffb-e2c01d900803","added_by":"auto","created_at":"2025-10-22 04:21:40","extension":"html","order_by":10,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":618163,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-7863429/v1/801c32a874ff58094e1befb9.html"},{"id":94056694,"identity":"14a6304e-0c1a-43a9-af3e-94b0a14114fb","added_by":"auto","created_at":"2025-10-22 04:21:40","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":1183751,"visible":true,"origin":"","legend":"\u003cp\u003eNumber of documents and citations from 1981 to 2024. Blue histograms represent the number of publications, whereas the pink diamonds and lines define the trend of publication related to benthic foraminifera studies in coral reefs. The dashed line separates the three phases of the trend.\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-7863429/v1/048fb4ceee3de8b37790633d.png"},{"id":94056886,"identity":"a59f4738-08d3-4131-9cf5-e0499f935d50","added_by":"auto","created_at":"2025-10-22 04:29:39","extension":"jpeg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":143929,"visible":true,"origin":"","legend":"\u003cp\u003e(A) Worldwide corals distribution modified from Allen Coral Atlas (2022). Red shaded colour showed cold-water corals while green shaded colour indicates the warm-water corals. (B) Map showing location of studied benthic foraminifera from coral reefs worldwide: Malaysia: \u003cstrong\u003e1\u003c/strong\u003e-Tioman Island, \u003cstrong\u003e2\u003c/strong\u003e- Bidong Island, \u003cstrong\u003e3\u003c/strong\u003e-\u003cem\u003e \u003c/em\u003eRedang Island; Philippines: \u003cstrong\u003e4\u003c/strong\u003e- Palawan, \u003cstrong\u003e5\u003c/strong\u003e- Nogas Island; Indonesia: \u003cstrong\u003e6\u003c/strong\u003e- Tambelan Island, \u003cstrong\u003e7\u003c/strong\u003e- Kepulauan Seribu, \u003cstrong\u003e8\u003c/strong\u003e- Bali; Australia: \u003cstrong\u003e9\u003c/strong\u003e- Great Barrier Reef, \u003cstrong\u003e10\u003c/strong\u003e- Moreton Bay; \u003cstrong\u003e11\u003c/strong\u003e- New Caledonia; Polynesia: \u003cstrong\u003e12\u003c/strong\u003e- Niue; French Polynesia: \u003cstrong\u003e13\u003c/strong\u003e- Bora bora; \u003cstrong\u003e14\u003c/strong\u003e- Galapagos Island; Panama: \u003cstrong\u003e15\u003c/strong\u003e- Punta Galeta; Colombia: \u003cstrong\u003e16\u003c/strong\u003e- Isla Barú; Puerto Rico: \u003cstrong\u003e17\u003c/strong\u003e- Jobos Bay; Brazil: \u003cstrong\u003e18\u003c/strong\u003e- Pirangi and Maracajaú; Ireland: \u003cstrong\u003e19\u003c/strong\u003e- Rockall Bank; Norway: \u003cstrong\u003e20\u003c/strong\u003e- Norway Margin; \u003cstrong\u003e21- \u003c/strong\u003eAlboran Sea; \u003cstrong\u003e22\u003c/strong\u003e- Qeshm Island; Maldives:\u003cstrong\u003e 23\u003c/strong\u003e- Ari Atoll, \u003cstrong\u003e24\u003c/strong\u003e- Rasdhoo Atoll; Tanzania: \u003cstrong\u003e25\u003c/strong\u003e- Zanzibar; Ecuador: \u003cstrong\u003e26\u003c/strong\u003e- Galapagos Archipelagos.\u003c/p\u003e","description":"","filename":"floatimage1.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7863429/v1/a8b19ebba95fb2a66bfdcc93.jpeg"},{"id":94057136,"identity":"6558546a-7f50-4a5d-bda7-8ce3b6910aa4","added_by":"auto","created_at":"2025-10-22 04:45:41","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2610406,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7863429/v1/f634ac65-853e-46b7-9ba6-e6e4711dfcb6.pdf"},{"id":94056683,"identity":"e1952270-fb74-4dfa-9d36-b77f908c5334","added_by":"auto","created_at":"2025-10-22 04:21:39","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":18385,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementaryTableS1.docx","url":"https://assets-eu.researchsquare.com/files/rs-7863429/v1/31c54d54303ff9935e386a6f.docx"}],"financialInterests":"The authors declare no competing interests.","formattedTitle":"\u003cp\u003e\u003cstrong\u003eAssessing the diversity of benthic foraminifera in coral reefs: a global perspective\u003c/strong\u003e\u003c/p\u003e","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eCoral reefs are among the most biologically diverse and ecologically important ecosystems on Earth, commonly referred to as the \"rainforests of the sea\" (Wooldridge, \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). These complex bioconstructions, primarily composed of calcium carbonate laid down by corals and calcareous algae, support an immense array of marine life, housing an estimated 30% of all known marine species despite occupying less than 0.5% of the ocean floor (Bravo et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). They are primarily distributed in shallow tropical and subtropical waters, typically between latitudes 30\u0026deg;N and 30\u0026deg;S, with major reef systems located in the Indo-Pacific, Caribbean, and Red Sea regions (Ranjan et al., \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The Coral Triangle, located in the western Pacific Ocean, is commonly regarded as the global epicentre of marine biodiversity, which host a variety of species and endemisms (Pinheiro et al., \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Mills et al., 2023). Approximately 845 species of reef-building corals are distributed globally (DeVantier et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). These ecosystems, which offer ecosystem services like the availability of renewable resources like fisheries, shoreline protection from erosive processes, biogeochemical cycles like nitrogen fixation, and valuable cultural and recreational tourism opportunities, are essential to humankind (Elliff \u0026amp; Silva, \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eDespite their importance, these vital ecosystems are facing unprecedented threats from both localized anthropogenic pressures and global climate change, threatening their structural integrity (Sous et al., \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). The primary threats include ocean warming due to climate change, which leads to coral bleaching, ocean acidification, overfishing, coastal development, and pollution from land-based sources such as agricultural runoff and plastic debris. These stressors have contributed to widespread degradation of reef systems, with reports indicating that more than 50% of coral reefs globally have experienced significant decline over the past few decades (Aswani et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Ranjan et al., \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The cumulative effect of these disturbances has pushed many coral reef systems to the brink of ecological collapse, which triggers urgent attention for comprehensive conservation and management strategies to enhance their resilience against future impacts (Chung et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Coral cover and reef resilience have significantly decreased due to the increased frequency and severity of mass bleaching events, particularly those linked to strong El Ni\u0026ntilde;o phenomena (Heenan et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Furthermore, reef habitats are still being harmed by destructive fishing methods like blast and cyanide fishing, especially in areas with weak enforcement capabilities (Carneiro \u0026amp; Martins, \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eTo mitigate this situation, several international and regional bodies are actively monitoring coral reef health and promote sustainable management. Some of the key organizations include the International Coral Reef Initiative (ICRI), the Global Coral Reef Monitoring Network (GCRMN), and the United Nations Environment Programme (UNEP), alongside regional programs such as the Coral Triangle Initiative (CTI). These governing bodies collaborate with national governments, NGOs, and research institutions in order to assess reef status, implement marine protected areas, and foster community-based conservation. However, effective management remains constrained due to limited funding, inconsistent enforcement, and the accelerating impacts of global climate change, urgent the need for coordinated international action to safeguard these vital ecosystems.\u003c/p\u003e\u003cp\u003eCoral reefs are host to diverse assemblages of microorganisms, including coral-associated benthic foraminifera, a single-celled eukaryotes with calcareous shells that play a crucial role in reef sediment formation and biogeochemical cycling (Dawson et al., \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Langlet et al., \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Narayan et al., \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). These foraminifera are particularly abundant in warm, shallow reef environments and have been commonly used as bioindicators of reef health due to their sensitivity to environmental changes such as water temperature, salinity, pH, and pollution levels (Hallock et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; A\u0026rsquo;ziz et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Mar\u0026iacute;n, \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Belart et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Foraminiferal assemblages can reflect both natural variability and anthropogenic disturbances, making them essential in long-term reef monitoring programs (Prazeres et al., \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). A decline in symbiont-bearing large benthic foraminifera, such as \u003cem\u003eAmphistegina\u003c/em\u003e spp. and \u003cem\u003eMarginopora\u003c/em\u003e spp., commonly parallels with coral bleaching events, signalling broader ecosystem stress (Girard et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Inferring foraminiferal diversity in coral reef ecosystems enhances our understanding of micro- to macroscale ecological shifts and provides an additional layer of insight for conservation and restoration efforts (Hallock et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Prazeres et al., \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Mar\u0026iacute;n, \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e\u003cp\u003ePreviously, biogeographical distribution of benthic foraminifera has been carried out from various settings (e.g., Culver \u0026amp; Buzas, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Murray, \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2001\u003c/span\u003e, \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Gooday \u0026amp; Jorissen, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Camacho et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Frontalini et al., \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Kim et al., 2016; F\u0026ouml;rderer et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Malek et al., \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Amao et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2019\u003c/span\u003e, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Mamo et al., \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). However, despite these efforts, the compilation and synthesis of species-level data remain ambiguous, particularly due to regional biases, inconsistent taxonomic resolution, and limited integration of datasets across biogeographical provinces.\u003c/p\u003e\u003cp\u003eThe present study aims to bridge this gap by compiling and analyzing species-level occurrence records from a wide range of coral reef habitats across major ocean basins, thereby offering new insights into the global taxonomical diversity of benthic foraminifera in these sensitive ecosystems. This work is intended to provide a list of benthic foraminiferal species from coral reefs worldwide in order to assess the overall biodiversity and identify geographical pattern.\u003c/p\u003e"},{"header":"2. Materials and methods","content":"\u003cp\u003eCompiling species-level data is a challenging and time-consuming task that is commonly hindered by inconsistencies in the reporting of species names in published studies. Scopus databases are used for collecting publication by using keywords such as \u0026ldquo;foraminifera\u0026rdquo;, \u0026ldquo;benthic foraminifera\u0026rdquo;, and \u0026ldquo;coral reefs\u0026rdquo;. The time frame was selected to include publication from the beginning of the year these keywords emerged. From the initial 428 documents retrieved, we excluded publications not directly related to benthic foraminifera in coral reef environments. To ensure consistency and relevance, we restricted the dataset to articles that specifically addressed species diversity and were published in the English language. The search strings used are shown below:\u003c/p\u003e\u003cp\u003e(TITLE-ABS-KEY (benthic foraminifera) AND TITLE-ABS-KEY (coral reef)) AND PUBYEAR\u0026thinsp;\u0026gt;\u0026thinsp;1980 AND PUBYEAR\u0026thinsp;\u0026lt;\u0026thinsp;2025 AND (LIMIT-TO (DOCTYPE, \"ar\")) AND (LIMIT-TO (EXACTKEYWORD, \"Foraminifera\") OR LIMIT-TO (EXACTKEYWORD, \"Coral Reef\") OR LIMIT-TO (EXACTKEYWORD, \"Benthic Foraminifera\") OR LIMIT-TO (EXACTKEYWORD, \"Coral Reefs\") OR LIMIT-TO (EXACTKEYWORD, \"Reefs\") OR LIMIT-TO (EXACTKEYWORD, \"Foraminifer\") OR LIMIT-TO (EXACTKEYWORD, \"Coral\") OR LIMIT-TO (EXACTKEYWORD, \"Reef\") OR LIMIT-TO (EXACTKEYWORD, \"Benthic Foraminifers\") OR LIMIT-TO (EXACTKEYWORD, \"Foraminifers\") OR LIMIT-TO (EXACTKEYWORD, \"Corals\")) AND (LIMIT-TO (LANGUAGE, \"English\")).\u003c/p\u003e\u003cp\u003eIn some cases, authors chose not to include detailed species identifications in their work, either due to a focus on higher taxonomic levels or limitations in available taxonomic resources. This lack of uniformity complicates efforts to create comprehensive datasets and may lead to gaps in our understanding of the overall species diversity and distribution. To address these challenges, we selected publications for taxonomic review based on the following criteria:\u003c/p\u003e\u003cp\u003e\u003col\u003e\u003cspan\u003e\u003cli\u003e\u003cp\u003epublications that include species names;\u003c/p\u003e\u003c/li\u003e\u003c/span\u003e\u003cspan\u003e\u003cli\u003e\u003cp\u003epublications that specify the sampling area as coral reefs;\u003c/p\u003e\u003c/li\u003e\u003c/span\u003e\u003c/ol\u003e\u003c/p\u003e\u003cp\u003eIn addition, a compilation of species from cold-water reefs, as reported in the dissertation of Margreth (2010), was included.\u003c/p\u003e\u003cp\u003eSystematic classification of the benthic foraminiferal species, genera, and the suprageneric taxonomic categories are adopted from the World Register of Marine Species (WoRMS, 2025). Each species and genus were verified against WoRMS, using only the accepted name. Open nomenclature species were not considered in the species lists compilation.\u003c/p\u003e\u003cp\u003eThe presence-absence list of taxa was compiled and used for the β diversity (differentiation) and γ diversity (regional) calculation (Al-Enezi et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The β diversity was calculated using the PAlaeontological STatistics (PAST) data analysis software (version 4.02) (Hammer et al., \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2001\u003c/span\u003e) and expressed as the global β diversity (βw) (Whittaker, 1960), while γ diversity is computed as the total species richness. The βw represents the variation in species composition, whereby the highest difference is indicated with the high value of βw (Al-Enezi et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In addition, Jaccard dissimilarity index (\u003cem\u003eD\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e) was calculated, which ranges from 0 to 1, and provided the degree of overlap between the datasets, with higher values suggesting greater similarity (Jaccard, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1912\u003c/span\u003e).\u003c/p\u003e"},{"header":"3. Results","content":"\u003cdiv id=\"Sec4\" class=\"Section2\"\u003e\u003ch2\u003e3.1 Data compilation\u003c/h2\u003e\u003cp\u003eA total of 317 documents were retrieved using the search strings, and the publication and citation trends from 1981 to 2024, are presented in Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. The trend fluctuates annually but shows an overall general upward pattern in both publications and citations. The trends are tentatively divided into three phases. The first phase, from 1981 to 2000 represents the formative years of research output, during which only a small number of papers were published annually, commonly fewer than five per year. This period reflects the early exploration of the field, with publications appearing sporadically and a relatively low cumulative citation count. The second phase between 2001 and 2012 marked as a period of steady growth. During this time, the annual publication rate gradually increased, consistently exceeding five papers per year, with several peaks reaching 10 to 15 publications. In the recent phase (i.e., 2013 to 2024), the period is characterized by higher productivity, with annual publications frequently ranging between 15 to 25 documents. The cumulative citation curve continued to climb steeply, surpassing 9,000 citations by 2024, reflecting both the maturity of the field and its wider impact. Despite some fluctuations in annual outputs, the overall trend shows sustained and intensive research interest during this phase.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eFollowing the search criteria, 24 publications of benthic foraminifera from coral reefs were selected comprising seven locations from Atlantic Ocean (i.e., Norway, Puerto Rico, Ireland, Alboran Sea, Brazil, Panama, and Columbia), seven from Pacific Ocean (i.e., Malaysia, New Caledonia, Australia, Philippines, Ecuador, Polynesia, and Bora bora) and five from Indian Ocean (i.e., Iran, Indonesia, Tanzania, Maldives and Mozambique) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e2\u003c/span\u003e; Table S1). Overall, the data collection recorded 1,054 species (γ diversity) of benthic foraminifera that are represented by 371 genera, 123 families, and 13 orders (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e; Table S2). The most diverse order belonged to Rotaliida, with a total of 65 families, followed by Lituolida (15 families) and Miliolida (13 families). Moderate diversification was observed in orders like Astrorhizida (nine families), Textulariida (six families), and Nodosariida (four families). The Order Loftusiida and Spirillinida was less common, with only two families, while the orders Robertinida, Allogromiida, Carterinida, and Vaginulinida were represented by a single family.\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\u003eBenthic foraminiferal taxa recognized in coral reefs worldwide.\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=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eOrder\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eFamily\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eGenus\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eSpecies\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAllogromiida\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eAllogromiidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAstrorhizida\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eAstrorhizoidea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eBotellinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eHyperamminidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eSaccamminidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e5\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\u003cp\u003eStannomidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eRhabdamminidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e4\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\u003cp\u003eHippocrepinellidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eHyperamminidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003ePsammosphaeridae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCarterinida\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCarterinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eLoftusiida\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHaddoniidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eGlobotextulariidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eLituolida\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eAdercotrymidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\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\u003cp\u003eAmmosphaeroidinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e6\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\u003cp\u003eTrilocularenidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eDiscamminidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\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\u003cp\u003eHaplophragmoididae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e5\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\u003cp\u003eHormosinellidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eHormosinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e7\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\u003cp\u003eLituolidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e3\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\u003cp\u003eNouriidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003ePlacopsilinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eProlixoplectidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eRemaneicidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\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\u003cp\u003eSpiroplectamminidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e3\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\u003cp\u003eTrochamminidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e19\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\u003cp\u003eVerneuilinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMiliolida\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003eAlveolinidae\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e5\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\u003cp\u003eCornuspiridae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e4\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\u003cp\u003eCribrolinoididae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e4\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\u003cp\u003eFischerinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e10\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\u003cp\u003eHauerinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e47\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e268\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\u003cp\u003eMiliamminidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\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\u003cp\u003eMiliolidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eNubeculariidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e4\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\u003cp\u003eOphthalmidiidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e3\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\u003cp\u003e\u003cb\u003ePeneroplidae\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e23\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\u003cp\u003eRiveroinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e5\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\u003cp\u003e\u003cb\u003eSoritidae\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e10\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\u003cp\u003eSpiroloculinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e38\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNodosariida\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eAmmolagenidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eLagenidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e16\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\u003cp\u003eNodosariidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e9\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\u003cp\u003eReophacidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePolymorphinida\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eEllipsolagenidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e42\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\u003cp\u003eGlandulinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e3\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\u003cp\u003ePolymorphinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e12\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eRobertinida\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRobertinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eRotaliida\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eAcervulinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e8\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\u003cp\u003eAlabaminidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e3\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\u003cp\u003eAlmaenidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eAmmoniidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e17\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\u003cp\u003eAnomalinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e10\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\u003cp\u003e\u003cb\u003eAmphisteginidae\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e6\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\u003cp\u003eAsterigerinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eAsterigerinatidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\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\u003cp\u003eAstrononionidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\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\u003cp\u003eBaggininae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eBolivinitidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e32\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\u003cp\u003eBolivinellidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eBuliminidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e8\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\u003cp\u003eBuliminoididae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003e\u003cb\u003eCalcarinidae\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e18\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\u003cp\u003eCancrisidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e10\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\u003cp\u003eCassidulinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e15\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\u003cp\u003eCibicididae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e28\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\u003cp\u003eChilostomellidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eCymbaloporidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e5\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\u003cp\u003eDiscorbidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e4\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\u003cp\u003eDiscorbinellidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e6\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\u003cp\u003eElphidiellidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\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\u003cp\u003eElphidiidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e41\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\u003cp\u003eEpistomariidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e5\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\u003cp\u003eEponididae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e14\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\u003cp\u003eGavelinellidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e8\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\u003cp\u003eGlabratellidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e3\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\u003cp\u003eGlobigerinitidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\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\u003cp\u003eGlobobuliminidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e3\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\u003cp\u003eHeronalleniidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eHomotrematidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e3\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\u003cp\u003eHaynesinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e4\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\u003cp\u003eMelonidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e3\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\u003cp\u003eMurrayinellidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\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\u003cp\u003eMississippinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eNonionidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e16\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\u003cp\u003eNotorotaliidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003e\u003cb\u003eNummulitidae\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e13\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\u003cp\u003ePavoninidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e4\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\u003cp\u003ePegidiidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003ePlacentulinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\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\u003cp\u003ePlanorbulinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e4\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\u003cp\u003ePlanulinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003ePseudoparrellidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e3\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\u003cp\u003ePulleniidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e6\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\u003cp\u003eReussellidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e4\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\u003cp\u003eRosalinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e42\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\u003cp\u003eRotaliidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e4\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\u003cp\u003eSiphoninidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e8\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\u003cp\u003eSiphogenerinoididae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e4\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\u003cp\u003eSphaeroidinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e3\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\u003cp\u003eStainforthiidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e3\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\u003cp\u003eSvratkinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\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\u003cp\u003eTortoplectellidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\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\u003cp\u003eTosaiidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eTrimosinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eTurrilinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\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\u003cp\u003eUngulatellidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eUvigerinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e9\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\u003cp\u003eVictoriellidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\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\u003cp\u003eEpistominidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\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\u003cp\u003eAmmodiscidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\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\u003cp\u003ePatellinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003ePlanispirillinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSpirillinida\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eAmmodiscidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eSpirillinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTextulariida\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eEggerellidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e33\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\u003cp\u003eKaminskiidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003eOlgiidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\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\u003cp\u003ePseudogaudryinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e8\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\u003cp\u003eTextulariidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e9\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\u003cp\u003eValvulinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eVaginulinida\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eVaginulinidae\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e17\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eTOTAL\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003e371\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003e1,054\u003c/b\u003e\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\u003e\u003cb\u003eBold\u003c/b\u003e families indicate symbiont-bearing type.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eFigure \u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e2\u003c/span\u003e: (A) Worldwide corals distribution modified from Allen Coral Atlas (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Red shaded colour showed cold-water corals while green shaded colour indicates the warm-water corals. (B) Map showing location of studied benthic foraminifera from coral reefs worldwide: Malaysia: \u003cb\u003e1\u003c/b\u003e-Tioman Island, \u003cb\u003e2\u003c/b\u003e- Bidong Island, \u003cb\u003e3\u003c/b\u003e- Redang Island; Philippines: \u003cb\u003e4\u003c/b\u003e- Palawan, \u003cb\u003e5\u003c/b\u003e- Nogas Island; Indonesia: \u003cb\u003e6\u003c/b\u003e- Tambelan Island, \u003cb\u003e7\u003c/b\u003e- Kepulauan Seribu, \u003cb\u003e8\u003c/b\u003e- Bali; Australia: \u003cb\u003e9\u003c/b\u003e- Great Barrier Reef, \u003cb\u003e10\u003c/b\u003e- Moreton Bay; \u003cb\u003e11\u003c/b\u003e- New Caledonia; Polynesia: \u003cb\u003e12\u003c/b\u003e- Niue; French Polynesia: \u003cb\u003e13\u003c/b\u003e- Bora bora; \u003cb\u003e14\u003c/b\u003e- Galapagos Island; Panama: \u003cb\u003e15\u003c/b\u003e- Punta Galeta; Colombia: \u003cb\u003e16\u003c/b\u003e- Isla Bar\u0026uacute;; Puerto Rico: \u003cb\u003e17\u003c/b\u003e- Jobos Bay; Brazil: \u003cb\u003e18\u003c/b\u003e- Pirangi and Maracaja\u0026uacute;; Ireland: \u003cb\u003e19\u003c/b\u003e- Rockall Bank; Norway: \u003cb\u003e20\u003c/b\u003e- Norway Margin; \u003cb\u003e21-\u003c/b\u003e Alboran Sea; \u003cb\u003e22\u003c/b\u003e- Qeshm Island; Maldives: \u003cb\u003e23\u003c/b\u003e- Ari Atoll, \u003cb\u003e24\u003c/b\u003e- Rasdhoo Atoll; Tanzania: \u003cb\u003e25\u003c/b\u003e- Zanzibar; Ecuador: \u003cb\u003e26\u003c/b\u003e- Galapagos Archipelagos.\u003c/p\u003e\u003cp\u003eThe family Hauerinidae represented the most taxonomically diverse group, comprising 47 genera, the highest recorded among benthic foraminifera family. Notably, it encompassed a wide range of genera, including \u003cem\u003eQuinqueloculina\u003c/em\u003e (109 species), \u003cem\u003eTriloculina\u003c/em\u003e (35 species), \u003cem\u003ePyrgo\u003c/em\u003e (17 species), and \u003cem\u003eMiliolinella\u003c/em\u003e (14 species). Other diverse genera were \u003cem\u003eElphidium\u003c/em\u003e with 31 species and \u003cem\u003eTextularia\u003c/em\u003e with 27 species as well as \u003cem\u003eRosalina\u003c/em\u003e (16 species), \u003cem\u003eBolivina\u003c/em\u003e (17 species), and \u003cem\u003eCibicides\u003c/em\u003e (12 species). Despite the occurrence of symbiont-bearing foraminifera in coral reefs, these families only represented a minor part of the species. Among them were reported Alveolinidae (i.e., \u003cem\u003eAlveolinella\u003c/em\u003e, \u003cem\u003eBorelis\u003c/em\u003e), Peneroplidae (i.e., \u003cem\u003eCoscinospira\u003c/em\u003e, \u003cem\u003eDendritina\u003c/em\u003e, \u003cem\u003eEuthymonacha\u003c/em\u003e, \u003cem\u003eLaevipeneroplis\u003c/em\u003e, \u003cem\u003eMonalysidium\u003c/em\u003e, \u003cem\u003ePeneroplis\u003c/em\u003e, \u003cem\u003eParasorites\u003c/em\u003e), Amphisteginidae (i.e., \u003cem\u003eAmphistegina\u003c/em\u003e), Nummulitidae (i.e., \u003cem\u003eHeterostegina\u003c/em\u003e, \u003cem\u003eNummulites\u003c/em\u003e, \u003cem\u003ePlanostegina\u003c/em\u003e, \u003cem\u003eOperculina\u003c/em\u003e, \u003cem\u003eOperculinella\u003c/em\u003e, \u003cem\u003eNeoassillina\u003c/em\u003e, \u003cem\u003ePlanoperculina\u003c/em\u003e), Calcarinidae (i.e., \u003cem\u003eBaculogypsina\u003c/em\u003e, \u003cem\u003eBaculogypsinoides\u003c/em\u003e, \u003cem\u003eCalcarina\u003c/em\u003e, \u003cem\u003eNeorotalia\u003c/em\u003e, \u003cem\u003ePararotalia\u003c/em\u003e, \u003cem\u003eSchlumbergerella\u003c/em\u003e), and Soritidae (i.e., \u003cem\u003eArchaias\u003c/em\u003e, \u003cem\u003eAmphisorus\u003c/em\u003e, \u003cem\u003eCycloputeolina\u003c/em\u003e, \u003cem\u003eCyclorbiculina\u003c/em\u003e, \u003cem\u003eMarginopora\u003c/em\u003e, \u003cem\u003eSorites\u003c/em\u003e).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec5\" class=\"Section2\"\u003e\u003ch2\u003e3.2 Distribution of species\u003c/h2\u003e\u003cdiv id=\"Sec6\" class=\"Section3\"\u003e\u003ch2\u003e3.2.1 European assemblages\u003c/h2\u003e\u003cp\u003eThe benthic foraminiferal assemblages on European coral reefs were studied at three locations in Norwegian margin reefs (Margreth, 2010; Spezzaferri et al., \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), Rockall Bank, Ireland (Margreth, 2010; Morigi et al., \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), and Alboran Sea (Margreth, 2010; Stalder et al., \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). These studies site consisted of cold-water coral reefs assemblages. The total number of species were high with 391 species: 180 species in Rockall Bank, 278 species in Porcupine Seabight and 173 species in Alboran Sea. Dominant species in Rockall Bank were \u003cem\u003eEpistominella exigua\u003c/em\u003e, \u003cem\u003eGlobocassidulina subglobosa\u003c/em\u003e, \u003cem\u003eCassidulina carinata\u003c/em\u003e, \u003cem\u003eCassidulina laevigata\u003c/em\u003e, \u003cem\u003eDiscanomalina coronata\u003c/em\u003e, while Porcupine Seabight with \u003cem\u003eAlabaminella weddellensis\u003c/em\u003e, \u003cem\u003eAdercotryma wrighti\u003c/em\u003e, \u003cem\u003eEhrenbergina carinata\u003c/em\u003e, \u003cem\u003eParatrochammina challenger\u003c/em\u003e, \u003cem\u003ePlanulina ariminensis\u003c/em\u003e, \u003cem\u003eSpirillina vivipara\u003c/em\u003e, and \u003cem\u003eTrochammina inflata\u003c/em\u003e. Meanwhile, in Alboran Sea, deep-water species such as \u003cem\u003eBulimina marginata, Bolivina dilatata, Bolivina striatula\u003c/em\u003e, \u003cem\u003eBolivina alata\u003c/em\u003e, \u003cem\u003eCassidulina crassa\u003c/em\u003e, \u003cem\u003eC. laevigata\u003c/em\u003e, \u003cem\u003eRectuvigerina elongatastriata\u003c/em\u003e, \u003cem\u003eUvigerina peregrina\u003c/em\u003e, and \u003cem\u003eGlobobulimina doliolum\u003c/em\u003e were dominant. The cold-water coral assemblages mainly consisted of small heterotrophic and opportunistic species.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec7\" class=\"Section3\"\u003e\u003ch2\u003e3.2.2 America assemblages\u003c/h2\u003e\u003cp\u003eFour locations of reefs have been studied, Jobos Bay, Puerto Rico (Marin et al., 2024), Pirangi and Maracaja\u0026uacute;, Brazil (Eichler \u0026amp; de Moura, \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), Punta Galeta, Panama (Elsa et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) and Galapagos Island, Ecuador (Humphreys et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Overall, 356 species has been identified with Ecuador recorded highest species (n\u0026thinsp;=\u0026thinsp;142 species), followed by Panama (n\u0026thinsp;=\u0026thinsp;88 species), Puerto Rico (n\u0026thinsp;=\u0026thinsp;73), and Brazil (n\u0026thinsp;=\u0026thinsp;53 species). Only three species were found at all locations (i.e., \u003cem\u003eQuinqueloculina laevigata\u003c/em\u003e, \u003cem\u003eQuinqueloculina lamarckiana\u003c/em\u003e, and \u003cem\u003eTriloculina trigonula\u003c/em\u003e). The heterotrophic assemblages in American reefs were the most diverse dominated with genera like \u003cem\u003eQuinqueloculina\u003c/em\u003e, \u003cem\u003eSpiroloculina\u003c/em\u003e, and \u003cem\u003eTriloculina\u003c/em\u003e, while symbiont-bearing taxa recorded such as \u003cem\u003eAmphisorus hemprichii\u003c/em\u003e, \u003cem\u003eAmphistegina gibbosa\u003c/em\u003e, \u003cem\u003eArchaias angulatus\u003c/em\u003e, \u003cem\u003eBorelis pulchra\u003c/em\u003e, \u003cem\u003eBorelis schlumbergeri\u003c/em\u003e, \u003cem\u003eBorelis clarionensis\u003c/em\u003e, \u003cem\u003eHeterostegina antillarum\u003c/em\u003e, \u003cem\u003eHeterostegina depressa\u003c/em\u003e, \u003cem\u003eHeterostegina curva\u003c/em\u003e, \u003cem\u003eLaevipeneroplis proteus\u003c/em\u003e, \u003cem\u003ePeneroplis carinatus\u003c/em\u003e, \u003cem\u003ePeneroplis pertusus\u003c/em\u003e, \u003cem\u003eSorites marginalis\u003c/em\u003e, \u003cem\u003eSorites marginata\u003c/em\u003e, \u003cem\u003ePlanogypsina acervalis\u003c/em\u003e, and \u003cem\u003eVertebrasigmoilina mexicana\u003c/em\u003e.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec8\" class=\"Section3\"\u003e\u003ch2\u003e3.2.3 African assemblages\u003c/h2\u003e\u003cp\u003eTwo sites were studied in the African region: the Zanzibar Archipelago (Narayan et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) and Bazaruto Archipelago, Mozambique (Langer et al., \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). A total of 124 species of benthic foraminifera were found in Zanzibar, belonging to 56 genera while slightly lower species in Mozambique with 95 species belonging to 63 genera. Both \u003cem\u003eAmphistegina lessonii\u003c/em\u003e and \u003cem\u003eAmphistegina lobifera\u003c/em\u003e were dominant in the assemblage. Although a large number of small heterotrophic species was found, the number of symbiont-bearing and opportunistic taxa were almost similar. In Zanzibar, symbiont-bearing taxa were \u003cem\u003eAlveolinella quoyi\u003c/em\u003e, \u003cem\u003eA. hemprichii\u003c/em\u003e, \u003cem\u003eA. lessonii\u003c/em\u003e, \u003cem\u003eA. lobifera\u003c/em\u003e, \u003cem\u003eAmphistegina papillosa\u003c/em\u003e, \u003cem\u003eAmphistegina radiata\u003c/em\u003e, \u003cem\u003eCoscinospira hemprichii\u003c/em\u003e, \u003cem\u003eH. depressa\u003c/em\u003e, \u003cem\u003eMarginopora vertebralis\u003c/em\u003e, \u003cem\u003eNeorotalia calcar\u003c/em\u003e, \u003cem\u003eOperculina ammonoides\u003c/em\u003e, \u003cem\u003eP. pertusus\u003c/em\u003e, \u003cem\u003ePeneroplis planatus\u003c/em\u003e, and \u003cem\u003eSorites orbiculus\u003c/em\u003e, while eight species were found in Mozambique (i.e., \u003cem\u003eA. hemprichii\u003c/em\u003e, \u003cem\u003eC. hemprichii\u003c/em\u003e, \u003cem\u003eH. depressa\u003c/em\u003e, \u003cem\u003eN. calcar\u003c/em\u003e, \u003cem\u003eP. planatus\u003c/em\u003e, \u003cem\u003eP. acervalis\u003c/em\u003e, \u003cem\u003eS. orbiculus\u003c/em\u003e, and \u003cem\u003eSorites variabilis\u003c/em\u003e). Opportunistic taxa were \u003cem\u003eAmmonia beccarii\u003c/em\u003e, \u003cem\u003eAmmonia tepida\u003c/em\u003e, \u003cem\u003eElphidium advena\u003c/em\u003e, \u003cem\u003eElphidium craticulatum\u003c/em\u003e, \u003cem\u003eElphidium crispum\u003c/em\u003e, \u003cem\u003eElphidium fichtelianum\u003c/em\u003e, \u003cem\u003eElphidium limbatum\u003c/em\u003e, \u003cem\u003eElphidium striatopunctatum\u003c/em\u003e, \u003cem\u003eElphidium macellum\u003c/em\u003e, \u003cem\u003eElphidium milletti\u003c/em\u003e, \u003cem\u003eEuloxostomum pseudobeyrichi\u003c/em\u003e, \u003cem\u003eElphidium hispidulum\u003c/em\u003e, \u003cem\u003eReusella spinulosa\u003c/em\u003e, \u003cem\u003eSagrina zanzibarica\u003c/em\u003e, \u003cem\u003eSagrinella durrandii\u003c/em\u003e, and \u003cem\u003eSagrinella jugosa.\u003c/em\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec9\" class=\"Section3\"\u003e\u003ch2\u003e3.2.4 Asian assemblages\u003c/h2\u003e\u003cp\u003eThe Asian region has the highest number of location studied for coral reefs foraminifera, including three coral reefs in Malaysia (Bidong Island (Husain et al., \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), Tioman Island (A'ziz et al., 2021), Redang Island (Minhat et al., \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), three in Indonesia (i.e., Riau (Junita et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), Kepulauan Seribu (Renema, \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2008\u003c/span\u003e) and Bali (Renema, \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2003\u003c/span\u003e), two in the Philippines (Palawan (F\u0026ouml;rderer \u0026amp; Langer, \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) and Nogas Island (Gonzales et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), two at Maldives atolls (Parker \u0026amp; Gischler, 2011; Giraldo-Gomez et al, 2024), and one at each Iran (Maghsoudlou et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and Egypt (Badr El Din \u0026amp; Hallock, 2024). It corresponded of total 264 species, with eight species frequently occurred (i.e., \u003cem\u003eA. lessonii\u003c/em\u003e, \u003cem\u003eA. lobifera\u003c/em\u003e, E. \u003cem\u003ecraticulatum\u003c/em\u003e, \u003cem\u003eH. depressa\u003c/em\u003e, \u003cem\u003eN. calcar, P. pertusus\u003c/em\u003e, \u003cem\u003eP. planatus\u003c/em\u003e, \u003cem\u003eS. orbiculus\u003c/em\u003e). The assemblages consisted of a variety of symbiont-bearing taxa dominated by \u003cem\u003eA. lessonii\u003c/em\u003e, \u003cem\u003eA. radiata\u003c/em\u003e, \u003cem\u003eN. calcar\u003c/em\u003e, and \u003cem\u003eO. ammonoides\u003c/em\u003e, including a new species that was recently described (\u003cem\u003ePeneroplis hoheneggeri\u003c/em\u003e nov. sp.). The genera \u003cem\u003eAmmonia\u003c/em\u003e and \u003cem\u003eElphidium\u003c/em\u003e were commonly found as the opportunistic taxa and mainly represented by \u003cem\u003eA. beccarii\u003c/em\u003e, \u003cem\u003eAmmonia convexa\u003c/em\u003e, \u003cem\u003eAmmonia parkinsoniana\u003c/em\u003e, \u003cem\u003eAmmonia supera\u003c/em\u003e, \u003cem\u003eA. tepida\u003c/em\u003e, \u003cem\u003eE. advena\u003c/em\u003e, \u003cem\u003eE. craticulatum\u003c/em\u003e, \u003cem\u003eE. crispum\u003c/em\u003e, \u003cem\u003eE. excavatum\u003c/em\u003e, \u003cem\u003eE. fichtelianum\u003c/em\u003e, \u003cem\u003eElphidium gerthi\u003c/em\u003e, \u003cem\u003eE. hispidulum\u003c/em\u003e, \u003cem\u003eElphidium neosimplex\u003c/em\u003e, and \u003cem\u003eElphidium striatopunctatum\u003c/em\u003e.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec10\" class=\"Section3\"\u003e\u003ch2\u003e3.2.5 Oceanian assemblages\u003c/h2\u003e\u003cp\u003eThe largest coral reef system, the Great Barrier Reef, has been extensively studied (Uthicke \u0026amp; Nobes, \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Other locations, such as Moreton Bay (Narayan \u0026amp; Pandolfi, \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), Polynesia (Oron et al., \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), Bora Bora (Parker \u0026amp; Gischler, \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), New Caledonia (Debenay, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), and Papua New Guinea (Langer \u0026amp; Lipps, 2003), have also been investigated. The highest number of species recorded in these regions, with Australia exhibiting the greatest diversity (n\u0026thinsp;=\u0026thinsp;135 species), followed by Bora Bora (n\u0026thinsp;=\u0026thinsp;118 species), New Caledonia (n\u0026thinsp;=\u0026thinsp;69 species) and Polynesia (n\u0026thinsp;=\u0026thinsp;55 species). Five species frequently occurred, including \u003cem\u003eHauerina pacifica, Miliolinella oceanica, Quinqueloculina parkeri\u003c/em\u003e, and \u003cem\u003eSorites orbiculus\u003c/em\u003e. The region also recorded high numbers of symbiont-bearing taxa (n\u0026thinsp;=\u0026thinsp;34 species), dominated with taxa such as \u003cem\u003eA. lessonii\u003c/em\u003e, \u003cem\u003eA. papillosa\u003c/em\u003e, \u003cem\u003eA. radiata\u003c/em\u003e, \u003cem\u003eN. calcar\u003c/em\u003e, \u003cem\u003eCalcarina hispida\u003c/em\u003e, \u003cem\u003eM. vertebralis\u003c/em\u003e, and \u003cem\u003eS. orbiculus\u003c/em\u003e. Assemblages from Bora Bora were notable for a high number of small heterotrophic taxa (n\u0026thinsp;=\u0026thinsp;110 species), contributing to a total of 319 taxa recorded in the Oceania region, including the identification of a new species, \u003cem\u003eTextularia boraboraensis\u003c/em\u003e nov. sp.\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e\u003ch2\u003e3.3 Pattern of similarity/dissimilarity\u003c/h2\u003e\u003cdiv id=\"Sec12\" class=\"Section3\"\u003e\u003ch2\u003e3.3.1 Comparison between continents\u003c/h2\u003e\u003cp\u003eThe β index values revealed substantial species turnover in foraminiferal assemblages across the continents (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The global β\u003csub\u003ew\u003c/sub\u003e calculated was 2.5537, reflecting average dissimilarity among regions. The highest dissimilarities were observed between Europe and Africa (β\u0026thinsp;=\u0026thinsp;0.92), followed closely by Europe \u003cem\u003evs.\u003c/em\u003e Americas and Europe \u003cem\u003evs.\u003c/em\u003e Asia (β\u0026thinsp;=\u0026thinsp;0.89), suggesting minimal species overlap and high community distinctiveness. Europe \u003cem\u003evs.\u003c/em\u003e Oceania also showed considerable dissimilarity (β\u0026thinsp;=\u0026thinsp;0.88), reinforcing Europe\u0026rsquo;s unique biogeographic identity and environmental characteristics. Among the remaining continents, Americas \u003cem\u003evs.\u003c/em\u003e Africa (β\u0026thinsp;=\u0026thinsp;0.81), Americas \u003cem\u003evs.\u003c/em\u003e Oceania (β\u0026thinsp;=\u0026thinsp;0.81), and Americas \u003cem\u003evs.\u003c/em\u003e Asia (β\u0026thinsp;=\u0026thinsp;0.79) reflected moderate dissimilarity. The lowest β values occurred between Africa \u003cem\u003evs.\u003c/em\u003e Oceania (β\u0026thinsp;=\u0026thinsp;0.60), Africa \u003cem\u003evs.\u003c/em\u003e Asia (β\u0026thinsp;=\u0026thinsp;0.55), and Asia \u003cem\u003evs.\u003c/em\u003e Oceania (β\u0026thinsp;=\u0026thinsp;0.54), indicating relatively greater species overlap and compositional similarity within these tropical or subtropical regions.\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\u003eValues of β diversity and \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\stackrel{-}{D}\\)\u003c/span\u003e\u003c/span\u003e\u003csub\u003eJ\u003c/sub\u003e for each continental region.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"6\"\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\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eβ\u003csub\u003ew\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;2.5537\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eEurope\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eAmericas\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAfrica\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eAsia\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eOceania\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEurope\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAmericas\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.89\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAfrica\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.92\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.81\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAsia\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.89\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.79\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eOceania\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.88\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.81\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.60\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.54\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\stackrel{-}{\\varvec{D}}\\)\u003c/span\u003e\u003c/span\u003e\u003csub\u003eJ\u003c/sub\u003e=0.86\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003eEurope\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003eAmericas\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003eAfrica\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cb\u003eAsia\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cb\u003eOceania\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEurope\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAmericas\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.94\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAfrica\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.96\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.89\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAsia\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.94\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.89\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.71\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eOceania\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.93\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.89\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.75\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003ctfoot\u003e\u003ctr\u003e\u003ctd colspan=\"6\"\u003eβ\u003csub\u003ew\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;Whittaker global diversity\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd colspan=\"6\"\u003e\u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\stackrel{-}{D}\\)\u003c/span\u003e\u003c/span\u003e\u003csub\u003eJ\u003c/sub\u003e = Average Jaccard dissimilarity\u003c/td\u003e\u003c/tr\u003e\u003c/tfoot\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eComplementing these findings, the \u003cem\u003eD\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e mirrored similar trends. The global Jaccard dissimilarity (\u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\stackrel{-}{D}\\)\u003c/span\u003e\u003c/span\u003e\u003csub\u003eJ\u003c/sub\u003e = 0.86) suggested that foraminiferal communities are relatively similar overall. Europe \u003cem\u003evs.\u003c/em\u003e Africa again exhibited the highest dissimilarity (\u003cem\u003eD\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e = 0.96), with only 23 shared species, followed by Europe \u003cem\u003evs.\u003c/em\u003e Americas (\u003cem\u003eD\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e = 0.94), Europe \u003cem\u003evs.\u003c/em\u003e Asia (\u003cem\u003eD\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e = 0.94), and Europe \u003cem\u003evs.\u003c/em\u003e Oceania (\u003cem\u003eD\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e = 0.93). The Americas assemblages showed high dissimilarity with other continents (\u003cem\u003eD\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e = 0.89). Comparisons among Africa, Asia, and Oceania yielded the lowest dissimilarity scores, i.e., Africa \u003cem\u003evs.\u003c/em\u003e Oceania (\u003cem\u003eD\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e = 0.75), Africa \u003cem\u003evs.\u003c/em\u003e Asia (\u003cem\u003eD\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e = 0.71), and Asia \u003cem\u003evs.\u003c/em\u003e Oceania (\u003cem\u003eD\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e = 0.70), consistent with shared biogeographical patterns across the Indo-Pacific realm.\u003c/p\u003e\u003cp\u003eBoth indices consistently suggest that Europe harbors a distinct foraminiferal assemblage, while Africa, Asia, and Oceania show greater ecological and compositional continuity, potentially reflecting their geographic proximity and environmental connectivity.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec13\" class=\"Section3\"\u003e\u003ch2\u003e3.3.2 Comparison between oceans\u003c/h2\u003e\u003cp\u003eSimilarly, the comparison between the main ocean body followed similar trends with β\u003csub\u003ew\u003c/sub\u003e value slightly lower (β\u003csub\u003ew\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;1.3526) (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The highest dissimilarity values were observed between the Atlantic and Pacific Oceans (β\u0026thinsp;=\u0026thinsp;0.83) and Atlantic and Indian Oceans (β\u0026thinsp;=\u0026thinsp;0.81), suggesting distinct foraminiferal assemblages with limited species overlap in these regions. In contrast, the Pacific and Indian Oceans exhibited a notably lower β value of 0.56, reflecting greater similarity and shared species between these two adjacent oceanic regions.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eValues of β diversity and \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\stackrel{-}{D}\\)\u003c/span\u003e\u003c/span\u003e\u003csub\u003e\u003cem\u003eJ\u003c/em\u003e\u003c/sub\u003e for oceanic body.\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\u003cp\u003eβ\u003csub\u003ew\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;1.3526\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eAtlantic\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eIndian\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePacific\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAtlantic\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eIndian\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.81\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePacific\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.83\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.56\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\stackrel{-}{\\varvec{D}}\\)\u003c/span\u003e\u003c/span\u003e\u003csub\u003eJ\u003c/sub\u003e=0.84\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003eAtlantic\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003eIndian\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003ePacific\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAtlantic\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eIndian\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.90\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePacific\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.91\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.72\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003ctfoot\u003e\u003ctr\u003e\u003ctd colspan=\"4\"\u003eβ\u003csub\u003ew\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;Whittaker global diversity\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd colspan=\"4\"\u003e\u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\stackrel{-}{D}\\)\u003c/span\u003e\u003c/span\u003e\u003csub\u003eJ\u003c/sub\u003e = Average Jaccard dissimilarity\u003c/td\u003e\u003c/tr\u003e\u003c/tfoot\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eThese patterns are supported by the \u003cem\u003eD\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e, which similarly showed high dissimilarity between the Atlantic and Indian Oceans (\u003cem\u003eD\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e = 0.90) and the Atlantic and Pacific Oceans (\u003cem\u003eD\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e = 0.91), reinforcing the distinctness of the Atlantic fauna compared to the Indo-Pacific realms. The Pacific \u003cem\u003evs\u003c/em\u003e. Indian Ocean comparison again showed the lowest dissimilarity (\u003cem\u003eD\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e = 0.72), consistent with a higher degree of faunal overlap, likely due to the more direct connectivity and environmental continuity between these two ocean basins.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec14\" class=\"Section3\"\u003e\u003ch2\u003e3.3.3 Cosmopolitan distribution\u003c/h2\u003e\u003cp\u003eThree opportunistic species were recorded across oceanic regions (i.e., \u003cem\u003eA. beccarii\u003c/em\u003e, \u003cem\u003eA. tepida\u003c/em\u003e, \u003cem\u003eE. advena\u003c/em\u003e), with five symbiont-bearing (i.e., \u003cem\u003eA. hemprichii\u003c/em\u003e, \u003cem\u003eB. pulchra\u003c/em\u003e, \u003cem\u003eB. schlumbergeri\u003c/em\u003e, \u003cem\u003eH. depressa\u003c/em\u003e, \u003cem\u003eP. pertusus\u003c/em\u003e) and 48 small heterotrophic species. Among the small heterotrophic species recorded in all major oceans worldwide were \u003cem\u003eA. beccarii, A. tepida, Articulina pacifica, B. striatula, Bolivina variabilis, Cancris auricula, Cibicides refulgens, Cornuspira planorbis, Cycloforina granulocostata, Cymbaloporetta squamosa, Lobatula lobatula, Melonis affinis, Pyrgo oblonga, Quinqueloculina agglutinans, Quinqueloculina bicarinata, Quinqueloculina bosciana, Q. lamarckiana, Q. parkeri, Quinqueloculina philippinensis, Quinqueloculina poeyana, Quinqueloculina seminulum, Rosalina globularis, Rotorbis auberii, Spiroloculina antillarum, Spiroloculina communis, Spiroloculina corrugata, Textularia agglutinans, Textularia candeiana, Textularia pseudogramen, Triloculina rotunda, Triloculina tricarinata\u003c/em\u003e, and \u003cem\u003eT. trigonula\u003c/em\u003e.\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e"},{"header":"4. Discussion","content":"\u003cdiv id=\"Sec16\" class=\"Section2\"\u003e\u003ch2\u003e4.1 Biogeographic patterns of foraminifera in coral reefs\u003c/h2\u003e\u003cp\u003eThe global distribution of benthic foraminiferal assemblages in coral reefs reveals distinct regional patterns in terms of species richness, dominant taxa, and ecological strategies (i.e., heterotrophic, symbiont-bearing, and opportunistic). The highest overall species richness is recorded in Europe (γ\u0026thinsp;=\u0026thinsp;391) and Oceania (γ\u0026thinsp;=\u0026thinsp;319), followed by America (γ\u0026thinsp;=\u0026thinsp;356), Asia (γ\u0026thinsp;=\u0026thinsp;264), and Africa (γ\u0026thinsp;=\u0026thinsp;191). However, these figures are strongly influenced by environmental settings, sampling effort and ecological diversity within study sites.\u003c/p\u003e\u003cp\u003eEuropean cold-water coral reefs (e.g., Rockall Bank, Porcupine Seabight) are dominated by small heterotrophic taxa, as symbiont-bearing forms are absent at bathyal depths. This pattern reflects the oligotrophic, deep-water environments where cold-water corals prevail, limiting light availability necessary for photosymbionts (Chen \u0026amp; Lin, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Avnaim-Katav et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Vicente et al., \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In contrast, tropical reefs in America, Africa, Asia, and Oceania exhibit mixed assemblages with varying proportions of symbiont-bearing and opportunistic taxa, consistent with warmer, shallower reef habitats.\u003c/p\u003e\u003cp\u003eIn American reefs, although the number of shared species among sites was low, the dominance of miliolid genera such as \u003cem\u003eQuinqueloculina, Triloculina\u003c/em\u003e, and \u003cem\u003eSpiroloculina\u003c/em\u003e reflects a robust heterotrophic community. Several larger symbiont-bearing taxa, including \u003cem\u003eA. angulatus\u003c/em\u003e and \u003cem\u003eH. depressa\u003c/em\u003e, suggest healthy carbonate platforms and reef systems. African reefs, particularly from the Zanzibar and Bazaruto Archipelagos, show comparable proportions of heterotrophic, symbiont-bearing, and opportunistic taxa. The occurrences of both \u003cem\u003eAmphistegina\u003c/em\u003e and \u003cem\u003eElphidium\u003c/em\u003e species points to transitional reef conditions, potentially affected by both coral reef development and environmental disturbances (Murray, \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Girard et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; O\u0026rsquo;Brien et al., \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Asian assemblages, being the most geographically diverse in the study, harbor a relatively high diversity of both symbiont-bearing and opportunistic foraminifera. The consistent presence of taxa such as \u003cem\u003eA. lessonii, N. calcar\u003c/em\u003e, and \u003cem\u003eP. planatus\u003c/em\u003e across multiple sites underlines their resilience and ecological success in Indo-Pacific reefs. In Oceania, particularly in the Great Barrier Reef and Bora Bora, a high diversity of both small heterotrophic and symbiont-bearing taxa is observed. The identification of new species (e.g., \u003cem\u003eT. boraboraensis\u003c/em\u003e) underscores the region\u0026rsquo;s importance as a hotspot for foraminiferal diversity. The overlap of species such as \u003cem\u003eS. orbiculus\u003c/em\u003e and \u003cem\u003eP. acervalis\u003c/em\u003e across multiple sites indicates broad ecological tolerances and potential for wide biogeographic dispersal (Alve \u0026amp; Goldstein, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Murray, \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Prazeres et al., \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec17\" class=\"Section2\"\u003e\u003ch2\u003e4.2 Patterns of similarity and turnover\u003c/h2\u003e\u003cp\u003eBeta diversity analyses reveal clear biogeographic segregation among regions. The higher similarity of benthic foraminiferal assemblages in certain areas can be attributed to the lack of recent and past geographic or environmental separation. Such limited separation reduces habitat heterogeneity and environmental gradients, leading to lower beta diversity and greater faunal overlap among sites (Renema, \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Uthicke et al., \u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Diaz et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eEurope exhibits the highest species turnover, when compared to other continents, that can be ascribed to the peculiar environmental setting hosting cold-water reef ecosystems. In contrast, Asia, Africa, and Oceania exhibit a lower turnover (β\u0026thinsp;\u0026asymp;\u0026thinsp;0.54\u0026ndash;0.60), suggesting regional continuity and biogeographic connectivity in the Indo-Pacific (F\u0026ouml;rderer et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Oceanic comparisons further reinforce this pattern. The Atlantic Ocean stands out as the most distinct, exhibiting the least similarity with both the Indian and Pacific Oceans (\u003cem\u003eD\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e \u0026asymp; 0.90\u0026ndash;0.91), likely due to historical isolation and contrasting environmental conditions. Conversely, Pacific and Indian Ocean reefs share a high degree of similarity (\u003cem\u003eD\u003c/em\u003e\u003csub\u003eJ\u003c/sub\u003e = 0.72), reflecting the contiguous reef systems across Southeast Asia and Melanesia. The pronounced dissimilarity between Atlantic and Pacific benthic foraminiferal assemblages likely reflects the closure of the Isthmus of Panama (~\u0026thinsp;3 Ma), which terminated faunal exchange between the two oceanic realms, reorganized ocean circulation, and created distinct environmental regimes that promoted biogeographic divergence (Collins et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; McDougall, \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Bornmalm, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Groeneveld et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec18\" class=\"Section2\"\u003e\u003ch2\u003e4.3 Cosmopolitan taxa and ecological indicators\u003c/h2\u003e\u003cp\u003eDespite regional differences, several cosmopolitan species are recorded across all oceanic regions, indicating their ecological behaviors. Notably, three opportunistic taxa (\u003cem\u003eA. beccarii, A. tepida\u003c/em\u003e, and \u003cem\u003eE. advena\u003c/em\u003e) and five symbiont-bearing species (\u003cem\u003eA. hemprichii, B. pulchra, B. schlumbergeri, H. depressa, P. pertusus\u003c/em\u003e) are found globally. These taxa are commonly associated with both healthy reef conditions (in the case of larger symbiont-bearers) and environments under anthropogenic stress (in the case of opportunists).\u003c/p\u003e\u003cp\u003eThe compiled dataset (n\u0026thinsp;=\u0026thinsp;1,054 species) indicates that the coral reefs is the hotspot for biodiversity, as this number is higher than previous compilation from other location such as Gulf of Mexico (n\u0026thinsp;=\u0026thinsp;987 species) (Gupta \u0026amp; Smith, \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), Sahul Shelf and Timor Sea (n\u0026thinsp;=\u0026thinsp;946 species) (Loeblich \u0026amp; Tappan, \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e1994\u003c/span\u003e), (n\u0026thinsp;=\u0026thinsp;304 species) Korean Peninsula ( Kim et al., 2016) and Aegen Sea, Mediterranean (n\u0026thinsp;=\u0026thinsp;267 species). However, it is important to note that these studies were not exclusively based on carbonate platform environments, where foraminifera typically reach their highest diversity. A slightly higher species diversity has been reported by Dorst et al. (2013) from continental shelf of North-East Atlantic with 1,486 species. This highlights the ecological adaptability of benthic foraminifera, which can thrive across a wider range of substrates than previously emphasized. By contrast, studies restricted to larger benthic foraminifera (n\u0026thinsp;=\u0026thinsp;105 species) have identified peak richness in the Western Coral Triangle and Sahul Shelf provinces of the Central Indo-Pacific, reflecting the well-established role of these provinces as global biodiversity hotspots (F\u0026ouml;rderer et al., 2022). Interestingly, even in the oligotrophic Arabian Gulf, more than 492 benthic foraminiferal species have been recorded, and these assemblages extend beyond reef environments into adjacent habitats (Amao et al., 2025). Comparable findings have also been reported in the Mediterranean Sea, where benthic foraminiferal assemblages associated with cold-water coral ecosystems, vermetid reef platforms, and algal-dominated hard substrates display unexpectedly high diversity beyond classic reefal systems (Stalder et al., \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Rossbach et al., 2022; Manda et al., \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). This highlights that high species richness is not confined to classic reefal systems but can also emerge in marginal and environmentally extreme settings. Nonetheless, it is important to acknowledge that these sensitive coral environments remain poorly studied, and many taxa have not yet been identified to the species level, which likely underestimates true diversity. Moreover, none of the existing studies accounted for allogromids (i.e., naked forams). This suggests that the actual number of taxa is expected to be much higher than currently reported.\u003c/p\u003e\u003cp\u003eForaminiferal assemblages in coral reef environments are predominantly composed of a mixture of calcareous hyaline and porcelaneous forms, particularly among the small heterotrophic species, consistent with the observations of Murray (\u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). Although symbiont-bearing foraminifera are ecologically dominant in these habitats, the most numerically abundant group consists of small heterotrophic species, with a minor proportion of opportunistic taxa. The extensive list of small heterotrophic species with global distributions (n\u0026thinsp;=\u0026thinsp;38) further highlights the ecological flexibility and dispersal capability of these taxa. Some of these, such as \u003cem\u003eL. lobatula\u003c/em\u003e, \u003cem\u003eQ. philippinensis\u003c/em\u003e, and \u003cem\u003eS. communis\u003c/em\u003e, are well-known for their euryhaline and eurytopic nature.\u003c/p\u003e\u003cp\u003eBy 2023, the projection that nearly 90% of global reefs will face degradation was alarming (Narayan et al., \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The ocean warming, ocean acidification, and habitat loss threaten coral reefs where increasing coral bleaching event frequency occurred worldwide since 1998 (Eakin et al., \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Thangadurai et al., \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Symbiont-bearing taxa such as \u003cem\u003eAmphistegina\u003c/em\u003e, \u003cem\u003eSorites\u003c/em\u003e and \u003cem\u003ePeneroplis\u003c/em\u003e are highly sensitive to thermal stress and bleaching (Hallock, \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Schmidt et al., \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Narayan et al., \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), while acidification reduces calcification and test integrity (Prazeres et al., \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Guam\u0026aacute;n-Guevara et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). As coral cover declines, the loss of reef habitat further reduces assemblage diversity (Doo et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), with potential consequences for carbonate production and reef resilience. Consequently, climate-driven reef degradation poses a critical risk to benthic foraminiferal diversity, particularly for symbiont-bearing species that serve as key bioindicators of reef health.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec19\" class=\"Section2\"\u003e\u003ch2\u003e4.4 Limitation\u003c/h2\u003e\u003cp\u003eDespite the inclusion of a high number of publications, one of the key limitations of this study is the presence of incomplete or inconsistent species names in the original sources, which may have constrained the comprehensiveness and accuracy of the species database. Although we carefully selected and validated species lists from each continent and ocean basin, the reliance on published records introduces potential biases due to taxonomic discrepancies, synonymy, or unidentified taxa. Furthermore, while the analysis captures broad-scale biogeographic patterns, the generalizations derived from this dataset may overlook finer ecological or regional variations. Additionally, several other limitations should be noted: (a) naked foraminifera were not accounted for in the reviewed studies; (b) sampling efforts in some regions remain comparatively lower than in others; (c) many species are still left in open nomenclature, which further obscures true diversity; and (d) differences in sampling strategies across studies may introduce inconsistencies in diversity estimates.\u003c/p\u003e\u003cp\u003eAs such, caution is advised in interpreting the findings as fully representative of global foraminiferal diversity. Future efforts should aim to integrate molecular data, high-resolution taxonomic revisions, and standardized sampling protocols to enhance the reliability of beta diversity assessments.\u003c/p\u003e\u003c/div\u003e"},{"header":"5. Conclusion","content":"\u003cp\u003eThis global synthesis of benthic foraminiferal assemblages from coral reefs reveals significant regional differences driven by environmental conditions, reef type (cold-water \u003cem\u003evs.\u003c/em\u003e tropical), and biogeographic history. High β-diversity values, particularly between Europe and other continents, reflect strong species turnover and endemism. Conversely, the Indo-Pacific region demonstrates high compositional similarity, suggesting ecological connectivity and potential for dispersal among reef systems. The consistent presence of specific symbiont-bearing and opportunistic taxa across regions suggests their potential as universal bioindicators for reef health and environmental change. Moreover, the discovery of new taxa highlights the need for continued exploration and taxonomic resolution, particularly in under-studied regions. Future studies should focus on integrating molecular approaches to complement traditional taxonomy, assess cryptic diversity, and explore population connectivity across coral reef systems. Additionally, monitoring shifts in assemblage composition in response to climate change and anthropogenic impacts will be crucial for understanding reef ecosystem resilience and guiding conservation strategies.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eConflict of interest\u003c/strong\u003e\u003cp\u003eThe authors declare that they have no conflict of interest.\u003c/p\u003e\u003c/p\u003e\u003cp\u003e\u003cstrong\u003eConsent to participate\u003c/strong\u003e\u003cp\u003eThe authors are agreeing to be accountable for all aspects of the work in ensuring that questions related to the accuracy or integrity of any part of the work are appropriately investigated and resolved.\u003c/p\u003e\u003c/p\u003e\u003cp\u003e\u003cstrong\u003eConsent to publish\u003c/strong\u003e\u003cp\u003eThe authors provides consent to publish the data in this manuscript.\u003c/p\u003e\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAl-Enezi E, Khader S, Balassi E, Frontalini F (2020) Modern Benthic Foraminiferal Diversity: An Initial Insight into the Total Foraminiferal Diversity along the Kuwait. 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Coral Reefs 29(1):209\u0026ndash;225. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s00338-009-0574-9\u003c/span\u003e\u003cspan address=\"10.1007/s00338-009-0574-9\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eVicente TM, Yamashita C, De Mello E, Sousa SH, Ciotti AM (2021) Evaluation of the relationship between biomass of living (stained) benthic foraminifera and particulate organic matter vertical flux in an oligotrophic region, Campos Basin, southeastern Brazilian continental margin. 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Coral Reefs 36(4):1025\u0026ndash;1037. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s00338-017-1594-5\u003c/span\u003e\u003cspan address=\"10.1007/s00338-017-1594-5\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eWorld Register of Marine Species (WoRMS) Accessed at \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.marinespecies.org\u003c/span\u003e\u003cspan address=\"https://www.marinespecies.org\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e on [24 October 2024]. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.14284/170\u003c/span\u003e\u003cspan address=\"10.14284/170\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eSupplementary Table S1: Sources used for the global compilation of foraminiferal data in coral reefs\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"highlight":"","institution":"Universiti Tunku Abdul Rahman","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":"bleaching event, symbiont-bearing, carbonate, biodiversity, Indo-Pacific, Atlantic","lastPublishedDoi":"10.21203/rs.3.rs-7863429/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7863429/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eUtilizing a comprehensive dataset of species records from the Indo-Pacific and Atlantic regions, this work explores the diversity, distribution, and paleoenvironmental significance of benthic foraminifera across different coral reef ecosystems. Overall, 1,054 species of benthic foraminifera from coral ecosystems are recognized in this compilation, belonging to 371 genera, 123 families, and 13 orders. Although symbiont-bearing taxa such as \u003cem\u003eAmphistegina\u003c/em\u003e, \u003cem\u003eHeterostegina\u003c/em\u003e, and \u003cem\u003eCalcarina\u003c/em\u003e are typical of reef environments, the dataset is dominated by small heterotrophic species. This compilation identifies several cosmopolitan taxa, including three opportunistic species (\u003cem\u003eAmmonia beccarii\u003c/em\u003e, \u003cem\u003eAmmonia tepida\u003c/em\u003e, \u003cem\u003eElphidium advena\u003c/em\u003e), with five symbiont-bearing (\u003cem\u003eAmphisorus hemprichii\u003c/em\u003e, \u003cem\u003eBorelis pulchra\u003c/em\u003e, \u003cem\u003eBorelis schlumbergeri\u003c/em\u003e, \u003cem\u003eHeterostegina depressa\u003c/em\u003e, \u003cem\u003ePeneroplis pertusus\u003c/em\u003e) and 38 small heterotrophic species. European and American assemblages recorded the highest species diversity, with 391 and 356 species, respectively. The highest β-diversity was observed between the Atlantic and Pacific oceans (β\u0026thinsp;=\u0026thinsp;0.83), followed by the Atlantic and Indian oceans (β\u0026thinsp;=\u0026thinsp;0.81), whereas the Indian and Pacific oceans shared more species, with a lower β-diversity (β\u0026thinsp;=\u0026thinsp;0.56). The findings highlight significant species turnover across continents and ocean basins, indicating high foraminiferal diversity and the influence of regional environmental gradients on assemblage composition in coral reef settings.\u003c/p\u003e","manuscriptTitle":"Assessing the diversity of benthic foraminifera in coral reefs: a global perspective","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-10-22 04:21:34","doi":"10.21203/rs.3.rs-7863429/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":"fbae5bc4-05fd-4c88-8c98-e491835e4f4f","owner":[],"postedDate":"October 22nd, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":56311352,"name":"Marine and Freshwater Ecology"}],"tags":[],"updatedAt":"2025-10-22T04:21:34+00:00","versionOfRecord":[],"versionCreatedAt":"2025-10-22 04:21:34","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-7863429","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7863429","identity":"rs-7863429","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}