Justification
This widely distributed species is rare. Global level, species-specific population data are limited; however, coral reefs have declined globally and are expected to continue rapidly declining due to increasing severe bleaching conditions under temperature stress caused by climate change as well as a variety of other threats. Our species-specific vulnerability traits analysis indicates this species is moderately susceptible to major threats related to coral reef degradation (e.g., disease and bleaching). We applied two analytical approaches involving two different global coral datasets and the species’ distribution map as proxies to infer population decline. Based on global coral cover monitoring data, this species experienced a suspected decline of less than 25% over the past three generations, or since 1989. Based on the projected onset of annual severe bleaching (ASB) conditions via both SSP2-4.5 and SSP5-8.5 scenarios of global climate model data, in combination with the species’ depth range, distribution and bleaching vulnerability, this species is suspected to decline by less than 25% over the next three generations, or by 2050. It is listed as Least Concern. The change in status from the previous assessment reflects updated declines calculated from improved data on modeled coral cover loss and projected date of annual severe bleaching, along with improved knowledge of species traits.
Geographic Range Information
This species is distributed from eastern Sri Lanka, Myanmar, Thailand, Southeast Asia (DeVantier et al. 2009), Vietnam, South China Sea (Huang et al. 2016), Japan, southern Papua New Guinea, eastern Australia (Roelofs 2018), the Torres Strait (Sweatman et al. 2015), New Caledonia (Osborne et al. 2013), Fiji, Samoa (Fenner 2013), Phoenix Island, Line Islands (Kenyon et al. 2010), Hawaii (Kenyon and Aeby 2009), and Johnston Atoll. It has also been confirmed from Palau (DeVantier and Turak 2017).
The depth range is 3-20 m.
Population Information
This species is rare (DeVantier and Turak 2017). It was found within all of 20 surveyed sites in West Hawaii during 2015 (Maynard et al. 2016).
Species-specific, global level population information is limited. However, coral reefs are experiencing severe global level declines due to increasing water temperatures caused by climate change (Hoegh-Guldberg et al. 2017, Hughes et al. 2018, Donovan et al. 2021). For the purposes of this Red List assessment, we used species-specific vulnerability traits and two analytical approaches based on two global coral datasets to infer past (GCRMN 2021) and future (UNEP 2020) population trends.
Approach 1: Future population trend
The projected onset of annual severe bleaching (ASB) was applied as a proxy to estimate global level population decline. ASB represents the date at which a coral reef will likely experience severe bleaching conditions annually, and beyond which the species will experience a greater than 80% decline as it is not expected to recover (van Hooidonk et al. 2014). ASB is defined as at least eight Degree Heating Weeks (DHW) occurring over a three-month period within a year, and where a DHW occurs when the sea surface temperature is at least 1°C above the maximum monthly mean (van Hooidonk et al. 2014; 2015). We defined the onset of ASB as corresponding to 80% or more decline, however, this is conservative as other studies have found that coral populations may experience near complete mortality and are unlikely to recover with just two incidences of ASB per decade (Obura et al. 2022).
To calculate ASB for each species we applied spatial data made publicly available via a United Nations Environment Programme report (UNEP 2020) that used the 2019 IPCC CMIP6 global climate models to estimate the projected onset of ASB for the years 2015-2100 on a 27 km x 27 km grid according to the 2018 WCMC-UNEP global coral reef distribution map, which has a resolution to 30 m depth. These data are available via two scenarios of Shared Socioeconomic Pathways (SSP), with SSP5-8.5 representing current global emissions and SSP2-4.5 representing a future reduction in emissions (UNEP 2020). We applied SSP5-8.5 since it follows the precautionary approach recommended by the IUCN Red List methodology and SSP2-4.5 since it represents a more moderate climate change scenario that better tracks current policy projections (Roelfsema et al. 2020, Obura et al. 2022). To acknowledge varying levels of coral adaptation to thermal stress, both of these spatial data layers are available for all quarter degree intervals between 0° and 2°C (UNEP 2020); however, coral adaptation in general is little understood and varies by species and locality (Bay et al. 2017, Matz et al. 2020, Logan et al. 2021). To account for adaptation, we calculated two estimates of ASB onset for both the SSP5-8.5 and the SSP2-4.5, where the first estimate assumes the species has no level of adaptation (0°C) and the second assumes a capacity for 1°C of adaptation. We clipped each of these four UNEP (2020) spatial data layers to the species’ distribution and calculated the average year of ASB onset across all overlapping grid cells.
Based on this spatial analysis, the onset of ASB across this species’ range is projected to occur on average by the year 2034 for SSP5-8.5 and by 2038 for SSP2-4.5 assuming no level of adaptation and by the year 2062 for SSP5-8.5 and by 2071 for SSP2-4.5 assuming 1°C of adaptation. For species where the onset of ASB occurs within 3-generation lengths, the 3-generation reduction is calculated as 80% multiplied by two proportions: (i) the proportion of the species' depth range that is in 0-30 m range, and (ii) for widespread species, the proportion of cells within the species' range that are expected to experience ASB under SSP2-4.5 before 2050 (three generation lengths). We inferred that the uncertainty associated with the estimate of population decline based on 1°C of adaptation is lower given this species is primarily restricted to depths shallower than 30 m and is moderately susceptible to bleaching. For widespread species, the final estimate of decline was further adjusted by excluding the proportion of cells within its range that were expected to experience ASB under SSP2-4.5 after 2050 (three generation lengths), in order to account for the potential resilience of species to the asynchronous variability of bleaching events that occur across the Indo-Pacific. The relative vulnerability to bleaching (i.e., highly susceptible, moderately susceptible, or more resilient) is primarily based on scientific species expert knowledge. The application of the species’ depth range as a vulnerability factor is based on the assumption that a coral species with shallow depth preferences is more frequently exposed to extreme temperatures and might decline at a faster rate in some places than species that also occur in deeper, cooler waters (Riegl and Piller 2003), although this is not always the case (e.g., Smith et al. 2016, Frade et al. 2018). Ocean acidification, which is measured by aragonite saturation, is also considered a major threat to corals due to the impacts of climate change, however, the impacts are expected to be more severe in cooler and/or deeper waters (Couce et al. 2013, van Hooidonk et al. 2014, Hoegh-Guldberg et al. 2017). Although the exact threshold of aragonite saturation that is expected to cause significant decline is not well-known, in the Pacific, changes in aragonite saturation are expected to be most severe in high-latitude reefs (van Hooidonk et al. 2014). Therefore, this species is suspected to experience a projected global level decline of less than 25% by the year 2050, regardless of the SSP2-4.5 or SSP5-8.5 scenario.
Approach 2: Past population trend
Coral reef monitoring data were also applied as a proxy to estimate global level population decline. The Global Coral Reef Monitoring Network (GCRMN) compiled data related to the status and trends of coral reefs in 10 regions from 1978-2019 via the scientific monitoring observations of more than 300 network members located throughout the world. We applied the publicly available data on estimations of the percent of live hard coral cover loss at the 20%, 50% and 80% confidence intervals in the 37 subregions of the Indo-Pacific (GCRMN 2021) to estimate species population decline over the past three generations (1989-2019). The proportion of the species’ range that overlapped with each of the subregions was estimated using the Red List distribution map. The sum of the proportion of the subregional species distribution multiplied by the percent of coral cover loss in each subregion was then used to calculate the 20%, 50% and 80% estimates of coral loss across this species’ range.
To inform the choice of the best (i.e., lowest level of uncertainty) out of the three percentile declines, we considered 11 species-specific traits related to vulnerability to coral cover loss. Given this species’ depth range is 3-20 m and is predominately found at depths greater than 10 m, generalized abundance is considered rare, overall population is not restricted or highly fragmented, does occur off-reef, is highly susceptible to disease, has a moderate recovery potential from bleaching or disease, has a low susceptibility to crown-of-thorns starfish, is moderately susceptible to bleaching, has an unknown susceptibility to the impacts of ocean acidification (Kornder et al. 2018), did not have >10,000 pieces exported annually in the aquarium trade between 2010-2019, it is overall suspected to be moderately susceptible to threats related to reef degradation. Therefore, past decline was inferred from the 50% percentile of estimated coral cover loss, resulting in a suspected global level decline of less than 25% since 1989, or over the past three generations.
Habitat and Ecology Information
This species occurs in shallow, tropical reef environments on sheltered reefs. Sparse colonies are found from 12-15 m depth in the South China Sea and Gulf of Siam (Titlyanov and Titlyanova 2002). It is found on subtidal rock and rocky reefs, on the back and foreslope of the reef, and in lagoons.
The age at first maturity of most reef-building corals is typically three to eight years (Wallace 1999). Based on this, we infer that the average age of mature individuals of this species is greater than eight years. Based on average sizes and growth rates, we also infer that the average length of one generation is 10 years. Longevity is not known, but is likely to be greater than 10 years. Therefore, any population decline rates estimated for the purposes of this Red List assessment are measured over a time period of 30 years.
Threats Information
The major threat to corals is global climate change, resulting in an increasing frequency, duration and intensity of marine heatwaves (Frölicher et al. 2018). There is a high likelihood that these temperature extremes will lead to prolonged or repeated coral bleaching events in future, similar to the pan-tropical 2014 to 2017 event. This event affected 75% of well studied reefs across the globe, resulting in declines in live coral cover of up to 90% (Hughes et al. 2018). Climate change presents a range of additional threats to the long term persistence of coral reefs. Tropical cyclones will become increasingly intense under climate change (Cheal et al. 2017), likely resulting in more extensive physical damage to reefs. Sea-level rise threatens coral reefs if rates of increase exceed the ability of reefs to accrete calcium carbonate and keep pace with future sea-level (Perry et al. 2018).
Ocean acidification threatens coral reefs due to an altered seawater chemistry from increasing atmospheric carbon dioxide concentrations (Hoegh-Guldberg et al. 2007). The synergistic impact of global warming and ocean acidification is likely to result in a loss of coral reef calcium carbonate framework by 2100, if carbon dioxide emissions continue on their present trajectory (Dove et al. 2020).
Outbreaks of coral disease continue to threaten coral reefs worldwide, supporting initial correlations of increased disease incidence with higher ocean temperatures (Boyett et al. 2007). Recent evidence finds that coral disease prevalence tracks ENSO climate cycles, and suggests that disease outbreaks may become more frequent as ENSO cycles become shorter with climate change (Randall and van Woesik 2017).
Localized threats to corals include fisheries, human development (industry, settlement, tourism, and transportation), changes in native species dynamics (competitors, predators, pathogens and parasites), invasive species (competitors, predators, pathogens and parasites), dynamite fishing, chemical fishing, pollution from agriculture and industry, domestic pollution, sedimentation, and human recreation and tourism activities. These local threats interact with global climate-related threats resulting in either “chain” or “modification” effects (Franca et al. 2020). Chain effects occur where a local threat amplifies the magnitude of a globalised climate related threat. Alternatively, modification effects can result in additive, antagonistic or synergistic interactions. For example, an antagonistic interaction occurs where reefs with high turbidity and sediment load are less likely to bleach due to reduced light stress. In contrast, reefs with high nutrient loads or overfishing are likely to be those which experience the greatest declines and slowest recovery following bleaching events.
Use and Trade Information
Conservation Actions Information
All stony corals are listed on CITES Appendix II. All stony corals (Scleractinia) fall under Annex B of the European Union Wildlife Trade Regulations, and have done so since 1997. Furthermore, several countries (India, Israel, Somalia, Djibouti, Solomon Islands and the Philippines) at various stages have banned either the trade or export of CITES II listed species, which includes all stony corals, since 1999.
Recommended measures for conserving this species include research in taxonomy, population, abundance and trends, ecology and habitat status, threats and resilience to threats, restoration action; identification, establishment and management of new protected areas; expansion of protected areas; recovery management; and disease, pathogen and parasite management. Artificial propagation and techniques such as cryo-preservation of gametes may become important for conserving coral biodiversity.
Recommended conservation measures include the promulgation of a network of MPAs to monitor the effects of collecting for the aquarium trade, especially in Indonesia.
The Convention on Biological Diversity adopted an updated Strategic Plan for Biodiversity 2011–2020, which now includes Aichi Biodiversity Target 11, calling for 10% of coastal and marine areas to be conserved by 2020. And in 2016, the IUCN World Conservation Congress agreed upon a target of >30% global marine protection by 2030.
It is crucial that global warming is constrained well below 2°C (the goals of the Paris Agreement).