A. Northeast to southwest view of Pagan Island. B. Ash and gas plume emitted from Mount Pagan on July 14, 2010. C,D. High cyanobacterial cover on hard-bottom reef habitats.

A. Northeast to southwest view of Pagan Island. B. Ash and gas plume emitted from Mount Pagan on July 14, 2010. C,D. High cyanobacterial cover on hard-bottom reef habitats.

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Volcanically active islands abound in the tropical Pacific and harbor complex coral communities. Whereas lava streams and deep ash deposits are well-known to devastate coral communities through burial and smothering, little is known about the effect of moderate amounts of small particulate ash deposits on reef communities. Volcanic ash contains a d...

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... The cumulative impact of disturbance events have caused rapid and pronounced changes on coral reefs, many of which do not completely recover and instead undergo a transition (phase shift) to an alternative stable state [31][32][33]. The best-known phase shifts on coral reefs are transitions from scleractinian-dominated communities to a dominance of fleshy macroalgae [33], though other transitions to communities dominated by coralline or peyssonnelioid algae [34,35], sponges [36,37], corallimorphs [38], or other zoantharians [39,40] have also been reported. Shifts in community composition and overall reef degradation could drive local or regional extinctions of reef-associated species, emphasizing the need to examine reef dynamics at a local scale [41]. ...
... Despite the significant decline in live coral cover, the structural complexity of reefs at Lafac Bay remained high (average rugosity of 2.81 and slope of 44 degrees) [115], indicating that the reef had not 'flattened' or eroded yet [28]. Examples of coral recovery following acute or recurrent disturbances have been reported for the tropical Pacific [37,99]. Reefs like those at Lafac Bay, which experience limited anthropogenic impacts and are primarily disturbed by natural events, have been observed to return to a coral-dominated state following a stage of macroalgal dominance [116]. ...
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The island of Guam in the west Pacific has seen a significant decrease in coral cover since 2013. Lafac Bay, a marine protected area in northeast Guam, served as a reference site for benthic communities typical of forereefs on the windward side of the island. The staghorn coral Acropora abrotanoides is a dominant and characteristic ecosystem engineer of forereef communities on exposed shorelines. Photoquadrat surveys were conducted in 2015, 2017, and 2019, and a diver-operated hyperspectral imager (i.e., DiveRay) was used to survey the same transects in 2019. Machine learning algorithms were used to develop an automated pipeline to assess the benthic cover of 10 biotic and abiotic categories in 2019 based on hyperspectral imagery. The cover of scleractinian corals did not differ between 2015 and 2017 despite being subjected to a series of environmental disturbances in these years. Surveys in 2019 documented the almost complete decline of the habitat-defining staghorn coral Acropora abrotanoides (a practically complete disappearance from about 10% cover), a significant decrease (~75%) in the cover of other scleractinian corals, and a significant increase (~55%) in the combined cover of bare substrate, turf algae, and cyanobacteria. The drastic change in community composition suggests that the reef at Lafac Bay is transitioning to a turf algae-dominated community. However, the capacity of this reef to recover from previous disturbances suggests that this transition could be reversed, making Lafac Bay an excellent candidate for long-term monitoring. Community analyses showed no significant difference between automatically classified benthic cover estimates derived from the hyperspectral scans in 2019 and those derived from photoquadrats. These findings suggest that underwater hyperspectral imagers can be efficient and effective tools for fast, frequent, and accurate monitoring of dynamic reef communities.
... Volcanic eruptions are the most common natural disaster affecting marine ecosystems, with dozens of them occurring every year (Global Volcanism Program, 2023), and depending on their intensity and type of materials released, they totally or partially affect the ecosystems (Crisafulli et al., 2015). Thus, pyroclasts and ash can partially or even completely cover an area, significantly reducing or altering ecosystem biodiversity (Vroom and Zgliczynski, 2011;Schils, 2012;Hart et al., 2022), as incandescent lava flows obliterate all forms of life on the territory (Thorton, 2007). Despite their catastrophic effects, lava flows are suitable for studying key ecological processes because they are free habitats. ...
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We studied the primary succession of benthic communities in the lava flows of the Tajogaite volcano at 2, 4.5, and 7 months after the eruption ended. The lava from the Tajogaite created several lava flows and sterile rocky reefs that were monitored in both intertidal and subtidal areas up to 20 m depth. Sampling included macroinvertebrates and algae in the intertidal, and fishes, macroinvertebrates, and algae in the subtidal. A control zone was selected to compare the early colonisation of lava flows with that of a mature ecosystem. Colonisation of the lava flow was swift, with numerous species arriving and proliferating soon after the eruption ended. After 7 months, the total number of species recorded in the lava flows was 70, representing 64% of those found in the control zone. thus, communities were gradually becoming increasingly complex owing to the continuous incorporation of species. The number of fishes, and macroinvertebrates in both the intertidal and subtidal, lava flows increased progressively, approaching the values of the control zone. However, algae, in terms of total cover, presented values similar to the control zone from the beginning of the monitoring. All the communities have followed the same trajectory to converge towards communities like those in the control zone, although the rate at which they have changed with time differs. After seven months, differences in fishes between lava flows and the control zone were small, but they were still large with respect to macroinvertebrates and algae. Thus, according to each community of organisms, the benthic ecosystem of the lava flows was found at different stages of succession.
... Las coladas de lava incandescente son los materiales emitidos que a priori provocan mayor impacto puesto que tienen efectos devastadores sobre el territorio, aniquilando todas las formas de vida a su paso (Thorton, 2007). Pero también, los piroclastos y las cenizas pueden afectar a una zona cubriendo total o parcialmente la superficie del terreno, diezmando o modificando la biodiversidad de los ecosistemas, aunque en este caso sin provocar la mortalidad de todas las formas de vida (Vrom & Zgliczynski, 2011;Schils, 2012;Hart et al., 2022). ...
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La erupción del volcán Tajogaite ha creado un nuevo ecosistema, un laboratorio natural ideal para el estudio de procesos ecológicos, muchos de los cuales no pueden reproducirse en condiciones controladas de laboratorio. La sucesión primaria se ha podido estudiar de forma temprana, apenas dos meses después del final del proceso eruptivo, y ha incluido diferentes grupos de organismos, lo que nos ha permitido tener una visión amplia de la formación de un ecosistema. Algunos organismos colonizaron las coladas de lava más rápido que otros, la capacidad de movilidad de los peces fue una ventaja que les permitió llegar primero y evolucionar rápidamente hacia comunidades clímax. El resto de comunidades de organismos estudiadas, invertebrados y algas, han evolucionado más lentamente. Sólo en las algas encontramos unreemplazo de especies a través de la sucesión, mientras que en peces e invertebrados, las especies que colonizaron primero las coladas de lava fueron esencialmente las mismas que estaban presentes en la zona de control. Describimos un ecosistema donde aún las interacciones entre organismos son incipientes, esto ha dado lugar al asentamiento e insólita proliferación de algunas especies. Esta contribución, como las futuras en las que estamos trabajando, nos permitirán aportar conocimiento de las variaciones espacio-temporales de las comunidades y de las dinámicas a largo plazo en territorios volcánicos de nueva formación, tan necesarios para una correcta gestión de la biodiversidad.
... Bruno et al., 2009), other organisms have also been reported to increase in abundance after coral mortality (Norström et al., 2009;Bell et al., 2021). Coral decline has corresponded with increased sponge abundance at some locations in the Caribbean (Diaz and Rutzler, 2001;McMurray et al., 2010), Brazil (Kelmo et al., 2013), the Indo-Pacific (Biggerstaff et al., 2017) and the Central Pacific (Schils, 2012;Knapp et al., 2016). In addition, experimental evidence indicates that many tropical sponge species, although vulnerable to high temperature, may be more tolerant than corals to ocean warming and acidification (Bennett et al., 2017). ...
... Sponges have many important ecological roles, including removal of organic matter from the water column, bioerosion of the reef framework, nutrient cycling, competition for space, and provision of food for predators (reviewed in Bell, 2008). Sponges can also grow into complex three-dimensional structures (Boury-Esnault and Rützler, 1997;Schönberg, 2021), and potentially contribute to reef structural complexity, especially when they are present at high density and diversity (Maldonado et al., 2016;Harris et al., 2021). Sponges can host diverse endobiotic communities of invertebrates in their internal system of aquiferous canals (reviewed in Wulff, 2006a). ...
... High levels of tephra deposition can result in water turbidity (cloudiness) (Lee, 1996), is linked to a low surface water pH (Blong, 1984;Wall-Palmer et al., 2011), and can directly lead to the smothering of organisms (Eldredge and Kropp, 1985;Ono et al., 2002;Vroom and Zgliczynski, 2011;Wu et al., 2018). Existing studies that investigate the interaction of volcanic ash deposition on coral reefs, all of which are exclusively based on posteruptive field studies, commonly report mass coral mortality (Heikoop et al., 1996;Vroom and Zgliczynski, 2011;Wu et al., 2018) through smothering, bleaching or coral overgrowth by macroalgae (Schils, 2012). However, if the physical effects of ash fallout do not result in direct smothering, tephra input may have a beneficial role on benthic communities (Frenzel, 1983;Duggen et al., 2007;Pearson et al., 2009), with reefs exhibiting high rates of recovery following exposure to volcanic ash (Smallhorn-West et al., 2020). ...
... It is, to our knowledge, the first time that exposure to volcanic ash deposition has been observed to positively impact coral physiology. This is likely because the majority of studies are field-based and focused on proximal locations to volcanic eruptions, where smothering can lead to coral mortality (Heikoop et al., 1996;Vroom and Zgliczynski, 2011;Schils, 2012;Wu et al., 2018). Nevertheless, the energy gain from photosynthesis we have observed in our experiments, which results from metal leaching in ambient seawater or in situ within the organism after ingestion, could be excreted in the form of coral mucus (Xu et al., 2022) as it is a protective mechanism against sedimentation stress (reviewed in Brown and Bythell, 2005). ...
... Explosive volcanic eruptions are capable of generating large volumes of tephra (10 11 -10 15 kg of released magma (Self, 2006)), disrupting all ecosystems in their vicinity and influencing biogeochemical cycles globally. Studies reporting on the effect of volcanic ash deposition on coral reefs are scarce and commonly report mass mortality (Heikoop et al., 1996;Vroom and Zgliczynski, 2011;Schils, 2012;Wu et al., 2018) of large portions of the coral reef close to the volcano. Information about the physiological response of corals to volcanic ash has not been gathered yet. ...
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Coral reefs, which are among the most productive ecosystems on earth, are in global decline due to rapid climate change. Volcanic activity also results in extreme environmental changes at local to global scales, and may have significant impacts on coral reefs compared to other natural disturbances. During explosive eruptions, large amounts of volcanic ash are generated, significantly disrupting ecosystems close to a volcano, and depositing ash over distal areas (10s - 1000s of km depending on i.a. eruption size and wind direction). Once volcanic ash interacts with seawater, the dissolution of metals leads to a rapid change in the geochemical properties of the seawater column. Here, we report the first known effects of volcanic ash on the physiology and elemental cycling of a symbiotic scleractinian coral under laboratory conditions. Nubbins of the branching coral Stylophora pistillata were reared in aquaria under controlled conditions (insolation, temperature, and pH), while environmental parameters, effective quantum yield, and skeletal growth rate were monitored. Half the aquaria were exposed to volcanic ash every other day for 6 weeks (250 mg L-1 week-1), which induced significant changes in the fluorescence-derived photochemical parameters (ΦPSII, Fv/Fm, NPQ, rETR), directly enhanced the efficiency of symbiont photosynthesis (Pg, Pn), and lead to increased biomineralization rates. Enhancement of symbiont photosynthesis is induced by the supply of essential metals (Fe and Mn), derived from volcanic ash leaching in ambient seawater or within the organism following ingestion. The beneficial role of volcanic ash as an important micronutrient source is supported by the fact that neither photophysiological stress nor signs of lipid peroxidation were detected. Subaerial volcanism affects micronutrient cycling in the coral ecosystem, but the implication for coral ecophysiology on a reef scale remains to be tested. Nevertheless, exposure to volcanic ash can improve coral health and thus influence resilience to external stressors.
... Beyond fish, VA deposition can impact other marine organisms. For instance, iron enrichment can favor phytoplankton species that are detrimental to coral reefs (Schils, 2012). Thick ash deposition can also directly obliterate coral reefs (Vroom and Zgliczynski, 2011) or lead to the loss of shallow water habitat (Zimmermann et al., 2018). ...
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Volcanic eruptions can be catastrophic events, particularly when they occur in inhabited coastal environments. They also play important roles in climate and biogeochemical cycles, including through nutrient deposition in the ocean. Volcanic ash studies in the ocean have focused on the phytoplankton response, generally quantifying changes in chlorophyll-a concentration. Many gaps remain in addressing fundamental questions regarding why volcanic ash deposition may enhance or limit both phytoplankton growth and/or drive community composition shifts. Here we outline a wide, multidisciplinary vision for monitoring volcanic eruptions near ocean ecosystems from satellites, including considerations for characteristics of airborne volcanic ash and ash geochemistry in seawater. Ultimately, observations beyond chlorophyll-a are needed to quantify phytoplankton communities (including harmful algal blooms) and possible impacts across higher trophic levels. We synthesize relevant research from volcanic studies as well as atmospheric and ocean sciences to identify the 'known unknowns' in ash-ecosystem studies. Our goal is to move toward an improved understanding of how real-time and near-real-time monitoring of volcanic eruptions can help address societally relevant questions.
... Even if categories like geniculate coralline algae and crustose coralline algae are employed in benthic surveys, branching Lithophyllum taxa are not a proper fit with either of these categories, resulting in high observer bias. As such, reliable baseline data on the abundance and persistence of branching Lithophyllum taxa are usually non-existent and therefore it is unknown if the observed community transitions are a short-term phenomenon [16] or represent a new type of phase shift on tropical Pacific reefs [17]. This question is relevant because certain species of non-geniculate coralline algae promote coral recovery as they serve as preferred settlement and recruitment substrates for various invertebrate larvae, including scleractinian corals [18]. ...
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A unique shift in benthic community composition, where scleractinian corals are replaced by coralline algae, has been observed on coral reefs in Guam in the western Pacific. Guam's reefs have been subjected to intense fishing pressure and impaired water quality for decades. Since 2013, heat stress has emerged as an additional major threat to the island's coral reefs. After a severe coral bleaching and mortality event in 2017, branching coralline algae of the genus Lithophyllum rapidly overgrew dead coral skeletons of the ecosystem engineer Acropora abrotanoides and have since become major components of forereef communities over a broad depth range. By now, the persistence of increased Lithophyllum cover meets the temporal criterium of phase shifts, but accurate estimates on the degree of dominance over appropriate spatial scales are lacking due to the absence of reliable baseline data. The ecological impacts of coral reef transitions towards increased coralline cover are unclear. Whereas carbonate budgets and reef growth could remain positive in the long term, the downstream effect of changes in structural complexity, (micro)habitat diversity, and benthic community composition on ecological processes and reef-associated faunal assemblages is unknown.
... Various studies on the Caribbean and Great Barrier Reef sponges showed high tolerance to thermal stress (for instance, up to 36°C for larvae of sponge R. odorabileis and 32°C for adults) (Webster et al. 2008Pantile and Webster 2011;Fan et al. 2013). Due to differential tolerance capacity of sponges and corals, reports from different environmental habitats (such as Caribbean reef, Atlantic coral reef in Brazil, Wakatobi Marine National Park in Indonesia, Palmyra Atoll lagoon in Central PaciBc and the Northern Mariana Islands) predicted the shifts towards sponge-dominated systems from coral-dominated systems (Aronson et al. 2002;Kelmo 2002;L opez-Victoria and Zea 2005;Ward-Paige et al. 2005;Maliao et al. 2008;Pawlik 2011;Schils 2012;Knapp et al. 2013). These community shifts are attributed to the ability of sponges to occupy newly available substrates and thrive under elevated temperatures, highly turbid and polluted environments . ...
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Marine intertidal organisms face extreme environmental fluctuations due to tidal cycles. To investigate the impact of environmental changes (high salinity, sea surface temperature, and anthropogenic pollution) on the health of the coastal area, we selected the dominant intertidal organisms: sponge (Cinachyrella cf. cavernosa) and zoanthid (Zoanthus sansibaricus) as our model system from a rocky beach Anjuna, Goa, India. Sponges and zoanthids were transplanted from their natural habitat (mid-tide pools) to new habitat (upper-tide pools). The study was carried out for 15 days when these pools experienced flushing during the first and last 5 days (high tide was >1.8 m) and no flushing during the middle 5 days (high tide was <1.8 m). The indices of physiological response (such as in budding activity and oscular openings) in the sponge and (chlorophyll estimation and symbiotic zooxanthellae count) in zoanthid were estimated at natural and transplanted habitats. Upper intertidal pools experienced intense temperature (29.9–36.15°C), salinity (35.03–46.7 ppt) and variability in dissolved oxygen (3.2–9.5 mg/l). The transplanted sponge did show the closure of oscula (90.62%) and reduction in buds (40–94.9%), but they could survive beyond 15 days. While transplanted zoanthid showed bleaching and did not survive beyond 72 h due to reduced chlorophyll content (up to 44.21%) and zooxanthellae count (34.54%). Our results depicted the differential tolerance capacity of sessile organisms like sponge and zoanthid to environmental changes, which govern the vertical zonation in the rocky intertidal region. It highlights the role of these sessile invertebrates as early indicators of environmental changes.
... Brazilian coast, Brazil n/a n/a n/a n/a Large 1998-2012 Invasive species [88] Curacao, Netherlands Antilles n/a n/a n/a n/a Small 2014-2016 n/a [89] Hon Сau Island, Vietnam n/a n/a n/a 14-54 Small 2013 n/a [33] b Tiao-Shi Reef, Taiwan n/a n/a n/a 32 Small 1992-2005 Storm, overfishing, pollution [71,72] Todos os Santos Bay, Brazil n/a 13 n/a 12 Medium 2015-2017 Invasive species [90] Sponges Bahia, Brazil n/a n/a n/a n/a Medium n/a El Niño Southern Oscillation (ENSO) [73,74] Curacao and Bonaire, Netherlands Antilles 30 10 1 2 Small 1973-2013 Coral bleaching events [75] Fort Lauderdale, Florida n/a n/a n/a n/a Medium 2000-2015 Coral bleaching events [76] Gulf of Mannar, India n/a n/a n/a n/a Small 2020 Coral bleaching events [91] Komodo Park, Indonesia n/a n/a n/a n/a Small 2016 n/a [77] b La Parguera, Puerto Rico n/a <1 n/a 11 Small 1975Small -1992 Coral bleaching, white-band disease [78,79] Mariana Islands n/a n/a n/a n/a Medium 2007-2011 Volcanic eruptions [21] Mauritius 33 26 10 13 Small 2014-2015 n/a [92] Okinawa, Japan n/a 4 n/a 24 Small 2010-2014 n/a [25,80] Palmyra Atoll, US Line Islands n/a n/a n/a 28 Large 2008 ...
... However, from most of these reports, the exact extent of the change is currently unclear. The reports from the Mariana islands [21] and Okinawa [22] are particularly interesting since large increases in the abundances of the coral-killing sponge Terpios hoshinota have been reported. This sponge covered up to 24% in Okinawa, and although the sponge had virtually disappeared within four years following a severe typhoon, it then recovered nearly half its area occupied within only 3 years. ...
Article
Despite the global focus on the occurrence of regime shifts on shallow-water tropical coral reefs over the last two decades, most of this research continues to focus on changes to algal-dominated states. Here, we review recent reports (in approximately the last decade) of regime shifts to states dominated by animal groups other than zooxanthellate Scleractinian corals. We found that while there have been new reports of regime shifts to reefs dominated by Ascidacea, Porifera, Octocorallia, Zoantharia, Actiniaria and azooxanthellate Scleractinian corals, some of these changes occurred many decades ago, but have only just been reported in the literature. In most cases, these reports are over small to medium spatial scales (<4 × 104 m2 and 4 × 104 to 2 × 106 m2, respectively). Importantly, from the few studies where we were able to collect information on the persistence of the regime shifts, we determined that these non-scleractinian states are generally unstable, with further changes since the original regime shift. However, these changes were not generally back to coral dominance. While there has been some research to understand how sponge- and octocoral-dominated systems may function, there is still limited information on what ecosystem services have been disrupted or lost as a result of these shifts. Given that many coral reefs across the world are on the edge of tipping points due to increasing anthropogenic stress, we urgently need to understand the consequences of non-algal coral reef regime shifts.
... The onset of meso-to eutrophic conditions will see a community shift from coralgal and foramol assemblages to bryomol and crinoid assemblages with lower calcification rates (Weissert & Erba 2004). Under fully eutrophic conditions, mesotrophic and oligotrophic carbonate producing communities are at danger of being overgrown by rapidlygrowing eutrophic organisms (Hallock & Schlager 1986;Reuter & Piller 2011) including cyanobacterial mats (Schils 2012), fleshy algae, (Hallock 1988), coralline algae (Littler et al. 1991;Wilson & Lokier 2002;Reuter & Piller 2011) and sponges (Reuter & Piller 2011;Schils 2012). Eutrophication will also ...
... The onset of meso-to eutrophic conditions will see a community shift from coralgal and foramol assemblages to bryomol and crinoid assemblages with lower calcification rates (Weissert & Erba 2004). Under fully eutrophic conditions, mesotrophic and oligotrophic carbonate producing communities are at danger of being overgrown by rapidlygrowing eutrophic organisms (Hallock & Schlager 1986;Reuter & Piller 2011) including cyanobacterial mats (Schils 2012), fleshy algae, (Hallock 1988), coralline algae (Littler et al. 1991;Wilson & Lokier 2002;Reuter & Piller 2011) and sponges (Reuter & Piller 2011;Schils 2012). Eutrophication will also ...
... Although some coral assemblages are able to tolerate eutrophic conditions (Mitchell 2002), increased nutrient levels will severely hinder growth (Hallock 2001a). Eutrophication will also fuel the growth of fleshy macroalgae and sponges that will rapidly colonise the substrate and further inhibit coral recruitment and development (Hall-Spencer et al. 2008;Schils 2012;Enochs et al. 2015;Enochs et al. 2016). ...
Article
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Carbonate sediments have been produced and deposited in areas of active volcanism since, at least, the Paleoarchean. Despite early recognition of a significant relationship between volcanism and marine carbonate systems, research in this field has been largely neglected. With increasing recognition of the accelerating effects and significance of anthropogenically-driven climate change on the ocean-atmosphere system, the time is ripe for studying volcanism-influenced carbonates as a natural analogue for future environmental scenarios. We undertake a detailed assessment of the state-of-the-science in our understanding of these systems. We identify significant bilateral division in approaches, with the geological and biological communities rarely interacting. The study of ancient volcanic-carbonate systems, in particular, appears to have ‘fallen-between-two-stools’ with both the volcanic and sedimentological communities shying away from studying these cross-disciplinary systems. Observations of recent volcanic-carbonate interactions are challenging, long periods of volcanic quiescence are punctuated by brief episodes of activity. Recent developments in robust remotely deployable instrumentation offer an opportunity to safely undertake sustained monitoring of these systems before, during and after eruptions. Informed assessment of the likely responses of carbonate ecosystems to future climatic challenges requires the initiation of an integrated, collaborative, cross-disciplinary approach to studying the complex interactions within these challenging mixed depositional systems.