When resilience is not enough: 2022 extreme marine heatwave threatens climatic refugia for a habitat-forming Mediterranean octocoral.

Climate change is impacting ecosystems worldwide, and the Mediterranean Sea is no exception. Extreme climatic events, such as marine heat waves (MHWs), are increasing in frequency, extent and intensity during the last decades, which has been associated with an increase in mass mortality events for multiple species. Coralligenous assemblages, where the octocoral Paramuricea clavata lives, are strongly affected by MHWs. The Medes Islands Marine Reserve (NW Mediterranean) was considered a climate refugia for P. clavata, as their populations were showing some resilience to these changing conditions. In this study, we assessed the impacts of the MHWs that occurred between 2016 and 2022 in seven shallow populations of the octocoral P. clavata from a Mediterranean Marine Protected Area. The years that the mortality rates increased significantly were associated with the ones with strong MHWs, 2022 being the one with higher mortalities. In 2022, with 50 MHW days, the proportion of total affected colonies was almost 70%, with a proportion of the injured surface of almost 40%, reaching levels never attained in our study site since the monitoring was started. We also found spatial variability between the monitored populations. Whereas few of them showed low levels of mortality, others lost around 75% of their biomass. The significant impacts documented here raise concerns about the future of shallow P. clavata populations across the Mediterranean, suggesting that the resilience of this species may not be maintained to sustain these populations face the ongoing warming trends.

El canvi climatic està impactant els ecosistemes arreu del planeta, i el Mar Mediterrani no n'és una excepció. Els esdeveniments climàtics extrems, com ara les onades de calor marines, estan augmentant en freqüència, extensió i intensitat en les darreres dècades, i estan sent associades a un increment dels esdeveniments de mortalitat massiva de múltiples espècies. El coral·ligen, on hi viu l'octocorall Paramuricea clavata, està altament afectat per les onades de calor marines. La Reserva Marina de les Illes Medes (NO del Mediterrani) es considerava un refugi climàtic per aquesta espècie, degut a que les seves poblacions mostraven certa resiliència a les condicions canviants. En aquest estudi hem avaluat els impactes de les onades de calor marines succeïdes entre els anys 2016 i 2022 a set poblacions someres de l'octocorall P. clavata, en una Àrea Marina Protegida del Mediterrani. Els anys en els quals les taxes de mortalitat van incrementar significativament s'associen amb els anys amb fortes onades de calor marines, sent el 2022 l'any amb la mortalitat més elevada. Al 2022, amb 50 dies d'onada de calor, la proporció total de colònies afectades va ser prop del 70%, amb un percentatge de superfície afectada de gairebé el 40%, arribant a valors mai observats en el lloc d'estudi des de que es va iniciar el seguiment d'aquestes poblacions. També hem observat variabilitat espacial entre les poblacions mostrejades. Mentre que algunes d'elles han mostrat poca mortalitat, altres han perdut al voltant del 75% de la seva biomassa. Els impactes documentats en aquest estudi mostren un futur preocupant de les poblacions someres de P. clavata arreu del Mediterrani, i això suggereix que la resiliència d'aquesta espècie podria no ser suficient per mantenir les seves poblacions en l'escenari d'escalfament que es preveu.

Marine heatwaves drive recurrent mass mortalities in the Mediterranean Sea.

Climate change is causing an increase in the frequency and intensity of marine heatwaves (MHWs) and mass mortality events (MMEs) of marine organisms are one of their main ecological impacts. Here, we show that during the 2015-2019 period, the Mediterranean Sea has experienced exceptional thermal conditions resulting in the onset of five consecutive years of widespread MMEs across the basin. These MMEs affected thousands of kilometers of coastline from the surface to 45 m, across a range of marine habitats and taxa (50 taxa across 8 phyla). Significant relationships were found between the incidence of MMEs and the heat exposure associated with MHWs observed both at the surface and across depths. Our findings reveal that the Mediterranean Sea is experiencing an acceleration of the ecological impacts of MHWs which poses an unprecedented threat to its ecosystems' health and functioning. Overall, we show that increasing the resolution of empirical observation is critical to enhancing our ability to more effectively understand and manage the consequences of climate change.

Ecological traits, genetic diversity and regional distribution of the macroalga Treptacantha elegans along the Catalan coast (NW Mediterranean Sea).

The widespread decline of canopy-forming macroalgal assemblages has been documented in many regions during the last decades. This pattern is often followed by the replacement of structurally complex algal canopies by more simplified habitats (e.g., turfs or sea urchin barren grounds). Against all odds, the fucoid Treptacantha elegans, a large Mediterranean brown macroalga, broadened its depth range to deeper and exposed environments and displayed an unexpected range expansion along the northern coast of Catalonia over the last two decades. Here, we reconstruct the spread of T. elegans in time and space and unravel ecological and demographic traits such as population dynamics and genetic patterns to provide a comprehensive and integrated view of the current status and geographical expansion for this species. Fast-growing dynamics, early fertile maturity, and high turnover rate are the main competitive advantages that allow the exposed populations of T. elegans to colonize available substrata and maintain dense and patchy populations. We also provided evidence that the deeper and exposed populations of T. elegans constitute a single group across the Catalan coast, with little genetic differentiation among populations. This seems to support the hypothesis of a unique source of spread in the last decades from the Medes Islands No-Take Zone towards both southern and northern waters.

Long-term monitoring of temperate macroalgal assemblages inside and outside a No take marine reserve.

Macroalgal communities have an essential role in the shallow benthic habitats of temperate seas, where changes in their composition can resonate through entire coastal ecosystems. As all major ecosystems on Earth, algal beds have already been affected by multiple disturbances. Passive conservation tools, such as marine protected areas or No-take zones, have the potential to reduce some of the anthropogenic impacts by limiting human activity. However, without a good knowledge of the natural community dynamics, it is not easy to discern between changes fruit of the intrinsic variability of biological communities and the ones caused by human-related stressors. In this study, we evaluated the natural variability of macroalgal communities' composition inside and outside a Mediterranean No-Take marine reserve during 15 years. We described their temporal dynamics considering their main drivers and we tested the effect of protection in seaweed beds. We did not find differences either in the composition of the macroalgal assemblages or the total algal cover between protected and non-protected locations over the fifteen years of study. Nevertheless, we observed a positive effect of the protection increasing the cover of some specific species, such as the canopy-forming Treptacantha elegans. Our results highlight the importance of obtaining long-term data in ecological studies to better understand the natural variability of marine communities. Accordingly, a robust understanding of the community dynamics would help us to avoid misinterpretations between 'impacted' or 'in-recovery' communities when recovery times are longer than the study periods.

Response diversity in Mediterranean coralligenous assemblages facing climate change: Insights from a multispecific thermotolerance experiment.

Climate change threatens coastal benthic communities on a global scale. However, the potential effects of ongoing warming on mesophotic temperate reefs at the community level remain poorly understood. Investigating how different members of these communities will respond to the future expected environmental conditions is, therefore, key to anticipating their future trajectories and developing specific management and conservation strategies. Here, we examined the responses of some of the main components of the highly diverse Mediterranean coralligenous assemblages to thermal stress. We performed thermotolerance experiments with different temperature treatments (from 26 to 29°C) with 10 species from different phyla (three anthozoans, six sponges and one ascidian) and different structural roles. Overall, we observed species-specific contrasting responses to warming regardless of phyla or growth form. Moreover, the responses ranged from highly resistant species to sensitive species and were mostly in agreement with previous field observations from mass mortality events (MMEs) linked to Mediterranean marine heat waves. Our results unravel the diversity of responses to warming in coralligenous outcrops and suggest the presence of potential winners and losers in the face of climate change. Finally, this study highlights the importance of accounting for species-specific vulnerabilities and response diversity when forecasting the future trajectories of temperate benthic communities in a warming ocean.

No-take marine reserves control the recovery of sea urchin populations after mass mortality events.

Understanding how no-take zones (NTZs) shape the population dynamics of key herbivores is crucial for the conservation and management of temperate benthic communities. Here, we examine the recovery patterns of sea urchin populations following a high-intensity storm under contrasting protection regimes in the NW Mediterranean Sea. We found significant differences in the recovery trends of Paracentrotus lividus abundance and biomass in the five years following the storm. The P. lividus populations outside the NTZ recovered faster than the populations inside the NTZ, revealing that predation was the main factor controlling the sea urchin populations inside the NTZ during the study period. Arbacia lixula reached the highest abundance and biomass values ever observed outside the NTZ in 2016. Our findings reveal that predation can control the establishment of new sea urchin populations and emphasize top-down control in NTZs, confirming the important role of fully protected areas in the structure of benthic communities.

Divergent responses to warming of two common co-occurring Mediterranean bryozoans.

Climate change threatens the structure and function of marine ecosystems, highlighting the importance of understanding the response of species to changing environmental conditions. However, thermal tolerance determining the vulnerability to warming of many abundant marine species is still poorly understood. In this study, we quantified in the field the effects of a temperature anomaly recorded in the Mediterranean Sea during the summer of 2015 on populations of two common sympatric bryozoans, Myriapora truncata and Pentapora fascialis. Then, we experimentally assessed their thermal tolerances in aquaria as well as different sublethal responses to warming. Differences between species were found in survival patterns in natural populations, P. fascialis showing significantly lower survival rates than M. truncata. The thermotolerance experiments supported field observations: P. fascialis started to show signs of necrosis when the temperature was raised to 25-26 °C and completely died between 28-29 °C, coinciding with the temperature when we observed first signs of necrosis in M. truncata. The results from this study reflect different responses to warming between these two co-occurring species, highlighting the importance of combining multiple approaches to assess the vulnerability of benthic species in a changing climate world.