# Projecting expected growth period of bivalves in a coastal temperate sea

Petra Zemunik Selak, Cléa Denamiel, Melita Peharda, Bernd Schöne, Julien

Thébault, Hana Uvanović, Krešimir Markulin, Ivica Vilibić

► To cite this version:

Petra Zemunik Selak, Cléa Denamiel, Melita Peharda, Bernd Schöne, Julien Thébault, et al. Pro-
jecting expected growth period of bivalves in a coastal temperate sea. Limnology and Oceanography
Letters, 2024, 10.1002/lo12.10393 . hal-04567185

HAL Id: hal-04567185

https://hal.univ-brest.fr/hal-04567185

Submitted on 6 May 2024

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# Projecting expected growth period of bivalves in a coastal temperate sea

Petra Zemunik Selak, Cléa Denamiel, Melita Peharda, Bernd Schöne, Julien

Thébault, Hana Uvanović, Krešimir Markulin, Ivica Vilibić

► To cite this version:

Petra Zemunik Selak, Cléa Denamiel, Melita Peharda, Bernd Schöne, Julien Thébault, et al. Pro-
jecting expected growth period of bivalves in a coastal temperate sea. Limnology and Oceanography
Letters, 2024, 10.1002/lo12.10393 . hal-04567185

HAL Id: hal-04567185

https://hal.univ-brest.fr/hal-04567185v1

Submitted on 6 May 2024

HAL is a multi-disciplinary open access
archive for the deposit and dissemination of sci-
entific research documents, whether they are pub-
lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers.

L'archive ouverte pluridisciplinaire HAL, est
destinée au dépôt et à la diffusion de documents
scientifiques de niveau recherche, publiés ou non,
émanant des établissements d'enseignement et de
recherche français ou étrangers, des laboratoires
publics ou privés.

<figure><img src="image_1.png" /></figure>

Distributed under a Creative Commons Attribution 4.0 International License
## LETTER

# Projecting expected growth period of bivalves in a coastal temperate sea

Petra Zemunik Selak,¹ Cléa Denamiel ID,²,³ Melita Peharda,¹ Bernd R. Schöne,⁴ Julien Thébault ID,⁵ Hana Uvanović,¹ Krešimir Markulin,¹ Ivica Vilibić ID²,³*

¹Institute of Oceanography and Fisheries, Split, Croatia; ²Division for Marine and Environmental Research, Ruđer Bošković Institute, Zagreb, Croatia; ³Institute for Adriatic Crops and Karst Reclamation, Split, Croatia; ⁴Institute of Geosciences, University of Mainz, Mainz, Germany; ⁵Univ Brest, CNRS, IRD, Ifremer, LEMAR, Plouzane, France

### Scientific Significance Statement

Climate change is transforming coastal ecosystems; however the extent of its impact is not fully understood. This study offers insights into the growth of bivalve species at the end of this century by integrating the sclerochronological approach provided by shell measurements with the climatological aspect presented by projections of the state-of-the-art climate model. This innovative method introduces a promising strategy for forecasting future trends in bivalve growth phenology, which may be extended to other marine species across various coastal environments.

### Abstract

The impact of climate warming on coastal benthic fauna is already observed, but forecasting their long-term fate remains challenging. This study uses $\delta^{18}$O$_{shell}$ data of specimens of five bivalve species collected at six locations and results from kilometer-scale atmosphere-ocean climate model for the time intervals of 1987-2017 and 2070-2100, to estimate changes in bivalve growth phenology. All species will benefit from climate warming during winter, experiencing a longer growing season than currently. The growth of *Aequipecten opercularis*, *Flexopecten glaber*, and *Pecten jacobaeus* will decrease in summer, resulting in up to 3 months of reduced growth per year. *Glycymeris pilosa* and *Venus verrucosa* in the southern Adriatic Sea will be more affected than those in the north, with up to 4 months longer annual growth. These findings can inform adaptation plans for bivalve management in the Adriatic Sea but also in areas where the studied species are present.

<footnote>*Correspondence: ivilibic@irb.hr</footnote>

<footnote>Associate editor: Ming-Tsung Chung</footnote>

<footnote>**Author Contribution Statement:** P.Z.S. and C.D. co-led the entire manuscript effort and contributed equally. C.D., M.P., and I.V. came up with the research question and designed the study approach. C.D. designed and implemented the atmosphere-ocean model. M.P., B.R.S., J.T., H.U., and K.M. obtained shell isotope data. P.Z.S. and C.D. conducted the statistical analyses and prepared visuals. P.Z.S., C.D., and I.V. wrote the manuscript. M.P., B.R.S., J.T., H.U., and K.M. reviewed and edited the manuscript. P.Z.S. and C.D. contributed equally to this work.</footnote>

<footnote>Petra Zemunik Selak and Cléa Denamiel contributed equally and are listed as co-first authors.</footnote>

<footnote>**Data Availability Statement:** All the data (bivalve stable oxygen isotope data, eastern Adriatic Sea temperature data, Adriatic Sea and Coast (AdriSC) model results) are archived in the Open Science Framework public repository and are available at: https://osf.io/ecqu3 (Denamiel et al., 2023).</footnote>

<footnote>Additional Supporting Information may be found in the online version of this article.</footnote>

<footnote>This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.</footnote>
## LETTER

# Projecting expected growth period of bivalves in a coastal temperate sea

Petra Zemunik Selak,¹ Cléa Denamiel ID,²,³ Melita Peharda,¹ Bernd R. Schöne,⁴ Julien Thébault ID,⁵ Hana Uvanović,¹ Krešimir Markulin,¹ Ivica Vilibić ID²,³*

¹Institute of Oceanography and Fisheries, Split, Croatia; ²Division for Marine and Environmental Research, Ruđer Bošković Institute, Zagreb, Croatia; ³Institute for Adriatic Crops and Karst Reclamation, Split, Croatia; ⁴Institute of Geosciences, University of Mainz, Mainz, Germany; ⁵Univ Brest, CNRS, IRD, Ifremer, LEMAR, Plouzane, France

### Scientific Significance Statement

Climate change is transforming coastal ecosystems; however the extent of its impact is not fully understood. This study offers insights into the growth of bivalve species at the end of this century by integrating the sclerochronological approach provided by shell measurements with the climatological aspect presented by projections of the state-of-the-art climate model. This innovative method introduces a promising strategy for forecasting future trends in bivalve growth phenology, which may be extended to other marine species across various coastal environments.

### Abstract

The impact of climate warming on coastal benthic fauna is already observed, but forecasting their long-term fate remains challenging. This study uses δ¹⁸O₃ₐ₉ₐ₆ data of specimens of five bivalve species collected at six locations and results from kilometer-scale atmosphere-ocean climate model for the time intervals of 1987-2017 and 2070-2100, to estimate changes in bivalve growth phenology. All species will benefit from climate warming during winter, experiencing a longer growing season than currently. The growth of *Aequipecten opercularis*, *Flexopecten glaber*, and *Pecten jacobaeus* will decrease in summer, resulting in up to 3 months of reduced growth per year. *Glycymeris pilosa* and *Venus verrucosa* in the southern Adriatic Sea will be more affected than those in the north, with up to 4 months longer annual growth. These findings can inform adaptation plans for bivalve management in the Adriatic Sea but also in areas where the studied species are present.

**Author Contribution Statement:** P.Z.S. and C.D. co-led the entire manuscript effort and contributed equally. C.D., M.P., and I.V. came up with the research question and designed the study approach. C.D. designed and implemented the atmosphere-ocean model. M.P., B.R.S., J.T., H.U., and K.M. obtained shell isotope data. P.Z.S. and C.D. conducted the statistical analyses and prepared visuals. P.Z.S., C.D., and I.V. wrote the manuscript. M.P., B.R.S., J.T., H.U., and K.M. reviewed and edited the manuscript. P.Z.S. and C.D. contributed equally to this work.

Petra Zemunik Selak and Cléa Denamiel contributed equally and are listed as co-first authors.

**Data Availability Statement:** All the data (bivalve stable oxygen isotope data, eastern Adriatic Sea temperature data, Adriatic Sea and Coast (AdriSC) model results) are archived in the Open Science Framework public repository and are available at: https://osf.io/ecqu3 (Denamiel et al., 2023).

Additional Supporting Information may be found in the online version of this article.

This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

<footnote>*Correspondence: ivilibic@irb.hr</footnote>

<footnote>Associate editor: Ming-Tsung Chung</footnote>
Understanding the impact of climate warming on fisheries and aquaculture systems has been highlighted as a priority in the latest report of the Intergovernmental Panel on Climate Change (https://www.ipcc.ch/assessment-report/ar6/, last accessed: 22 January 2024). However, predicting the impact of climate change on bivalve species remains challenging. Recent studies (Froehlich et al. 2018; Cubillo et al. 2021; Tan et al. 2022; Gallagher and Albano 2023) have attempted to forecast the global effects of climate change on coastal benthic fauna, yet they relied on regional and global models which often fail to accurately reproduce the thermohaline and biogeochemical conditions in coastal environments (Holt et al. 2009; Reale et al. 2020). Bivalves record temperature and salinity changes in their shells in the form of stable oxygen isotopes (δ¹⁸O<sub>shell</sub>; e.g., Goodwin et al. 2003; Vihtakari et al. 2016; Kubota et al. 2017). In settings characterized by stronger variations in ocean temperature than salinity δ¹⁸O<sub>shell</sub> data can thus provide reliable temperature estimates (Mook and Vogel 1968; Schöne et al. 2004; Wanamaker et al. 2007). Consequently, a sclerochronological approach can provide insight into bivalve growth seasonality (e.g., Vihtakari et al. 2016; Branscombe et al. 2021; Ezgeta-Balić et al. 2022).

The Adriatic Sea, which presents a complex coastal topography along its eastern coast (Fig. 1) is a particularly interesting region for the reproduction and projection of the growing season duration of bivalves. The Adriatic Sea and Coast (AdriSC; Denamiel et al. 2019) kilometer-scale climate model,

with resolutions an order of magnitude higher than the available regional climate models (e.g., Med-CORDEX ensemble; Soto-Navarro et al. 2020), has been specifically designed to accurately represent the Adriatic coastal dynamics under historical (1987–2017) and extreme warming (2070–2100) time interval for a representative concentration pathways (RCP) 8.5 greenhouse scenario conditions. Previous research has indeed demonstrated that the horizontal resolutions of 3-km in the atmosphere and 1-km in the ocean used in the AdriSC model are essential to reproduce the bora wind (Grisogono and Belušić 2009) driving the Adriatic dense water dynamics (Denamiel et al. 2021a) and the Adriatic thermohaline circulation (Pranić et al. 2023). Recent studies (Peharda et al., 2019a,b; Johnson et al., 2021; Uvanović et al. 2021; Ezgeta-Balić et al. 2022) have demonstrated that five bivalve species living in the Adriatic Sea (i.e., *Aequipecten opercularis*, *Flexopecten glaber*, *Pecten jacobaeus*, *Glycymeris pilosa*, and *Venus verrucosa*) form their shells near oxygen isotopic equilibrium with the ambient seawater. These species were chosen because of their longevity (*G. pilosa*) and commercial importance (other four species). Consequently, their δ¹⁸O<sub>shell</sub> values can be used for robust water temperature reconstructions and, combined with the AdriSC climate model results, trustable reproduction and projection of the expected growth of these five species under historical and extreme warming conditions can be achieved, as demonstrated in studies by Abe (2021) in Japan and Steeves et al. (2018) along the Atlantic Canadian coast.

<figure><img src="image_2.png" /><figcaption>**Fig. 1.** Geographical location of the eastern Adriatic coast, locations of the bivalve sampling sites and description of the two datasets depending on both sampling site locations and bivalve species.</figcaption></figure>
Understanding the impact of climate warming on fisheries and aquaculture systems has been highlighted as a priority in the latest report of the Intergovernmental Panel on Climate Change (https://www.ipcc.ch/assessment-report/ar6/, last accessed: 22 January 2024). However, predicting the impact of climate change on bivalve species remains challenging. Recent studies (Froehlich et al. 2018; Cubillo et al. 2021; Tan et al. 2022; Gallagher and Albano 2023) have attempted to forecast the global effects of climate change on coastal benthic fauna, yet they relied on regional and global models which often fail to accurately reproduce the thermohaline and biogeochemical conditions in coastal environments (Holt et al. 2009; Reale et al. 2020). Bivalves record temperature and salinity changes in their shells in the form of stable oxygen isotopes (δ¹⁸O<sub>shell</sub>; e.g., Goodwin et al. 2003; Vihtakari et al. 2016; Kubota et al. 2017). In settings characterized by stronger variations in ocean temperature than salinity δ¹⁸O<sub>shell</sub> data can thus provide reliable temperature estimates (Mook and Vogel 1968; Schöne et al. 2004; Wanamaker et al. 2007). Consequently, a sclerochronological approach can provide insight into bivalve growth seasonality (e.g., Vihtakari et al. 2016; Branscombe et al. 2021; Ezgeta-Balić et al. 2022).

The Adriatic Sea, which presents a complex coastal topography along its eastern coast (Fig. 1) is a particularly interesting region for the reproduction and projection of the growing season duration of bivalves. The Adriatic Sea and Coast (AdriSC; Denamiel et al. 2019) kilometer-scale climate model,

with resolutions an order of magnitude higher than the available regional climate models (e.g., Med-CORDEX ensemble; Soto-Navarro et al. 2020), has been specifically designed to accurately represent the Adriatic coastal dynamics under historical (1987–2017) and extreme warming (2070–2100) time interval for a representative concentration pathways (RCP) 8.5 greenhouse scenario conditions. Previous research has indeed demonstrated that the horizontal resolutions of 3-km in the atmosphere and 1-km in the ocean used in the AdriSC model are essential to reproduce the bora wind (Grisogono and Belušić 2009) driving the Adriatic dense water dynamics (Denamiel et al. 2021a) and the Adriatic thermohaline circulation (Pranić et al. 2023). Recent studies (Peharda et al., 2019a,b; Johnson et al., 2021; Uvanović et al. 2021; Ezgeta-Balić et al. 2022) have demonstrated that five bivalve species living in the Adriatic Sea (i.e., *Aequipecten opercularis*, *Flexopecten glaber*, *Pecten jacobaeus*, *Glycymeris pilosa*, and *Venus verrucosa*) form their shells near oxygen isotopic equilibrium with the ambient seawater. These species were chosen because of their longevity (*G. pilosa*) and commercial importance (other four species). Consequently, their δ¹⁸O<sub>shell</sub> values can be used for robust water temperature reconstructions and, combined with the AdriSC climate model results, trustable reproduction and projection of the expected growth of these five species under historical and extreme warming conditions can be achieved, as demonstrated in studies by Abe (2021) in Japan and Steeves et al. (2018) along the Atlantic Canadian coast.

<figure><img src="image_2.png" /><figcaption>**Fig. 1.** Geographical location of the eastern Adriatic coast, locations of the bivalve sampling sites and description of the two datasets depending on both sampling site locations and bivalve species.</figcaption></figure>
In this study, we propose a more comprehensive assessment of the impact of extreme warming on the growth of five bivalve species along the eastern Adriatic coast, based on a methodology that may be generalized to other marine species, coastal seas, and climate scenarios. Our primary objective is to provide a preliminary assessment of the locations and species that will potentially either benefit from climate change or not be suitable for fisheries and aquaculture. Based on these results, we will further discuss the potential adaption plans for the commercial exploitation of bivalves along the eastern Adriatic Sea.

# Material and methods

## AdriSC climate model

The AdriSC climate model (Denamiel et al. 2019) is the first atmosphere-ocean kilometer-scale climate model implemented in the world. It is based on the coupling between the weather research and forecasting (Skamarock et al. 2005) model at up to 3-km horizontal resolution (using 58 vertical levels) and the regional ocean modeling system (Shchepetkin and McWilliams 2009) at up to 1-km horizontal resolution (using 35 vertical levels). Two different AdriSC climate simulations were performed.

The historical simulation was forced with reanalysis products and covers the time interval 1987–2017 (Denamiel et al. 2021b; Pranić et al. 2021). The evaluation of the model demonstrated that the AdriSC 1-km ocean results could reproduce key processes driving the Adriatic thermohaline circulation (Denamiel et al. 2022; Pranić et al. 2023). Additionally, in this study, the evaluation was extended by using the observations collected at the bivalve sampling sites (Supporting Information Table S1; Fig. S1) and, overall, confirmed the skills of the AdriSC model in reproducing the temperatures along the eastern Adriatic coast.

The extreme warming simulation, covering the time interval 2070–2100, was performed under the RCP 8.5 greenhouse emission scenario using the pseudo-global warming (PGW) methodology to downscale the LMDZ4-NEMOMED8 regional climate model (Hourdin et al. 2006; Beuvier et al. 2010) from the Med-CORDEX ensemble (Soto-Navarro et al. 2020). The PGW methodology (Schär et al. 1996; Denamiel et al. 2020) consists in calculating the climatological changes based on the differences between future scenario (here for 2070–2100) and historical (here for 1987–2017) results from global or regional climate model and imposing it, as forcing, on top of the reanalysis products. Consequently, the main drawback of the PGW methodology is that it might miss potential changes in intra-annual and inter-annual variability. However, its main advantage consists in projecting the impact of climate change on the atmosphere-ocean dynamics at an unprecedented spatial resolution compared with traditional global and regional models, and for a reduced numerical cost.

## Stable oxygen isotope data

Along the eastern Adriatic coast, about 1650 carbonate samples from 32 shells collected at six different sampling sites were previously extracted and analyzed for five bivalve species (Peharda et al. 2019a, 2019b; Uvanović et al. 2021; Ezgeta-Balić et al. 2022) in order to analyze their stable oxygen isotope composition (δ¹⁸O<sub>shell</sub>, reported on the Vienna Pee Dee Belemnite scale as ‰ V-PDB). These species include three pectinids in Dataset 1—the Mediterranean scallop, *P. jacobaeus*, the queen scallop, *A. opercularis*, and the smooth scallop *F. glaber*—as well as, in Dataset 2—the long-lived hairy bittersweet (Dataset 2.1), *G. pilosa*, and the commercially important warty venus (Dataset 2.2), *V. verrucosa*. The sampling sites, depths, gears, periods and number of specimens, as well as the previously published studies are fully described in Fig. 1 and Table 1 and in section S1 of the Supporting Information.

In this study, seawater salinity and temperature time-series reproduced by the AdriSC model at different sampling sites were employed to compute the expected δ¹⁸O<sub>shell</sub> chronology (hereafter δ¹⁸O<sub>exp</sub>) for the time intervals 1987–2017 and 2070–2100. For Dataset 1, the AdriSC results were extracted and averaged over the entire region (off Istra; see Fig. 1) and depth ranges covered by the beam trawl, while, for Dataset 2, the values were extracted at the model grid points with the nearest location and depth of the sampling sites. Validation of the AdriSC data on in situ temperature loggers is provided in section S2 of the Supporting Information. Model and δ¹⁸O<sub>shell</sub> data are available in the Open Science Framework repository (Denamiel et al. 2023).

As the mineralogy of the bivalve species differs (i.e., calcitic and aragonitic for respectively Datasets 1 and 2), different paleothermometry equations were used. For Dataset 1, the Friedman and O'Neil (1977) formula derived from the O'Neil et al. (1969) equation was used to compute the expected δ¹⁸O<sub>shell</sub> (δ¹⁸O<sub>exp</sub>) chronology with the AdriSC temperature (T in °C):

$$ \delta^{18}O_{\text{exp}} = (1000 + \delta^{18}O_{\text{water}}) \times e^{[2780 \times (T+273.15)^{-2} - 0.00289]} - 1000 \quad (1) $$

Because the oxygen isotope composition of the water was not monitored over the lifespan of the studied specimens, δ¹⁸O<sub>water</sub> values (reported relative to V-SMOW) were reconstructed from the AdriSC salinity (S, at transect marked in Fig. 1, Peharda et al. 2019a) using the equation derived from Stenni et al. (1995):

$$ \delta^{18}O_{\text{water}} = 0.366S - 12.568, \quad r^2 = 0.964 \quad (2) $$

The conversion from the V-SMOW scale to V-PDB was done with the equation of Coplen et al. (1983):

$$ \delta^{18}O_{\text{exp-V-PDB}} = 0.97002 \delta^{18}O_{\text{exp-V-SMOW}} - 29.98 \quad (3) $$
In this study, we propose a more comprehensive assessment of the impact of extreme warming on the growth of five bivalve species along the eastern Adriatic coast, based on a methodology that may be generalized to other marine species, coastal seas, and climate scenarios. Our primary objective is to provide a preliminary assessment of the locations and species that will potentially either benefit from climate change or not be suitable for fisheries and aquaculture. Based on these results, we will further discuss the potential adaption plans for the commercial exploitation of bivalves along the eastern Adriatic Sea.

# Material and methods

## AdriSC climate model

The AdriSC climate model (Denamiel et al. 2019) is the first atmosphere-ocean kilometer-scale climate model implemented in the world. It is based on the coupling between the weather research and forecasting (Skamarock et al. 2005) model at up to 3-km horizontal resolution (using 58 vertical levels) and the regional ocean modeling system (Shchepetkin and McWilliams 2009) at up to 1-km horizontal resolution (using 35 vertical levels). Two different AdriSC climate simulations were performed.

The historical simulation was forced with reanalysis products and covers the time interval 1987–2017 (Denamiel et al. 2021b; Pranić et al. 2021). The evaluation of the model demonstrated that the AdriSC 1-km ocean results could reproduce key processes driving the Adriatic thermohaline circulation (Denamiel et al. 2022; Pranić et al. 2023). Additionally, in this study, the evaluation was extended by using the observations collected at the bivalve sampling sites (Supporting Information Table S1; Fig. S1) and, overall, confirmed the skills of the AdriSC model in reproducing the temperatures along the eastern Adriatic coast.

The extreme warming simulation, covering the time interval 2070–2100, was performed under the RCP 8.5 greenhouse emission scenario using the pseudo-global warming (PGW) methodology to downscale the LMDZ4-NEMOMED8 regional climate model (Hourdin et al. 2006; Beuvier et al. 2010) from the Med-CORDEX ensemble (Soto-Navarro et al. 2020). The PGW methodology (Schär et al. 1996; Denamiel et al. 2020) consists in calculating the climatological changes based on the differences between future scenario (here for 2070–2100) and historical (here for 1987–2017) results from global or regional climate model and imposing it, as forcing, on top of the reanalysis products. Consequently, the main drawback of the PGW methodology is that it might miss potential changes in intra-annual and inter-annual variability. However, its main advantage consists in projecting the impact of climate change on the atmosphere-ocean dynamics at an unprecedented spatial resolution compared with traditional global and regional models, and for a reduced numerical cost.

## Stable oxygen isotope data

Along the eastern Adriatic coast, about 1650 carbonate samples from 32 shells collected at six different sampling sites were previously extracted and analyzed for five bivalve species (Peharda et al. 2019a, 2019b; Uvanović et al. 2021; Ezgeta-Balić et al. 2022) in order to analyze their stable oxygen isotope composition (δ¹⁸O<sub>shell</sub>, reported on the Vienna Pee Dee Belemnite scale as ‰ V-PDB). These species include three pectinids in Dataset 1—the Mediterranean scallop, *P. jacobaeus*, the queen scallop, *A. opercularis*, and the smooth scallop *F. glaber*—as well as, in Dataset 2—the long-lived hairy bittersweet (Dataset 2.1), *G. pilosa*, and the commercially important warty venus (Dataset 2.2), *V. verrucosa*. The sampling sites, depths, gears, periods and number of specimens, as well as the previously published studies are fully described in Fig. 1 and Table 1 and in section S1 of the Supporting Information.

In this study, seawater salinity and temperature time-series reproduced by the AdriSC model at different sampling sites were employed to compute the expected δ¹⁸O<sub>shell</sub> chronology (hereafter δ¹⁸O<sub>exp</sub>) for the time intervals 1987–2017 and 2070–2100. For Dataset 1, the AdriSC results were extracted and averaged over the entire region (off Istra; see Fig. 1) and depth ranges covered by the beam trawl, while, for Dataset 2, the values were extracted at the model grid points with the nearest location and depth of the sampling sites. Validation of the AdriSC data on in situ temperature loggers is provided in section S2 of the Supporting Information. Model and δ¹⁸O<sub>shell</sub> data are available in the Open Science Framework repository (Denamiel et al. 2023).

As the mineralogy of the bivalve species differs (i.e., calcitic and aragonitic for respectively Datasets 1 and 2), different paleothermometry equations were used. For Dataset 1, the Friedman and O'Neil (1977) formula derived from the O'Neil et al. (1969) equation was used to compute the expected δ¹⁸O<sub>shell</sub> (δ¹⁸O<sub>exp</sub>) chronology with the AdriSC temperature (T in °C):

$$ \delta^{18}O_{\text{exp}} = (1000 + \delta^{18}O_{\text{water}}) \times e^{[2780 \times (T+273.15)^{-2} - 0.00289]} - 1000 \quad (1) $$

Because the oxygen isotope composition of the water was not monitored over the lifespan of the studied specimens, δ¹⁸O<sub>water</sub> values (reported relative to V-SMOW) were reconstructed from the AdriSC salinity (S, at transect marked in Fig. 1, Peharda et al. 2019a) using the equation derived from Stenni et al. (1995):

$$ \delta^{18}O_{\text{water}} = 0.366S - 12.568, \quad r^2 = 0.964 \quad (2) $$

The conversion from the V-SMOW scale to V-PDB was done with the equation of Coplen et al. (1983):

$$ \delta^{18}O_{\text{exp-V-PDB}} = 0.97002 \delta^{18}O_{\text{exp-V-SMOW}} - 29.98 \quad (3) $$
<figure><img src="image_3.png" /><figcaption>**Fig. 2.** Impact of climate change on shell growth of the bivalves species, *Aequipecten opercularis* (A.O.), *Flexopecten glaber* (F.G.), and *Pecten jacobaeus* (P.J.) (= Dataset 1 species) along Istria (IS) for conditions during 1987–2017 (= “Historical,” blue) and for conditions during 2070–2100 (= “Extreme warming,” red). (A) Distributions of the reconstructed shell oxygen stable isotope data (δ¹⁸Oexp values) with conditions favorable to growth derived from observed δ¹⁸Oshell (shaded), (B) duration of the annual growing season, and (C) duration of the monthly growing season.</figcaption></figure>

conditions, shells deposited almost no material during this
period of the year. The growth of Dataset 1 species will,
however, substantially decrease during the warmer season

(from May to October). This will likely far more affect the
growth of *A. opercularis* and *P. jacobaeus*, with favorable con-
ditions occurring throughout the entire year during
<figure><img src="image_3.png" /><figcaption>**Fig. 2.** Impact of climate change on shell growth of the bivalves species, *Aequipecten opercularis* (A.O.), *Flexopecten glaber* (F.G.), and *Pecten jacobaeus* (P.J.) (= Dataset 1 species) along Istria (IS) for conditions during 1987–2017 (= “Historical,” blue) and for conditions during 2070–2100 (= “Extreme warming,” red). (A) Distributions of the reconstructed shell oxygen stable isotope data (δ¹⁸Oexp values) with conditions favorable to growth derived from observed δ¹⁸Oshell (shaded), (B) duration of the annual growing season, and (C) duration of the monthly growing season.</figcaption></figure>

conditions, shells deposited almost no material during this
period of the year. The growth of Dataset 1 species will,
however, substantially decrease during the warmer season

(from May to October). This will likely far more affect the
growth of *A. opercularis* and *P. jacobaeus*, with favorable con-
ditions occurring throughout the entire year during
historical times, than *F. glaber* that was restricted to the annual thermal minimum. Consequently, *A. opercularis* and *P. jacobaeus* will potentially have an up to 3 months shorter annual growing season.

In Dataset 2, δ¹⁸O<sub>shell</sub> values for both *G. pilosa* (Fig. 3) and *V. verrucosa* (Fig. 4) range from negative to positive. This is in contrast with Dataset 1 species which displayed only positive values. Consequently, Dataset 2 species can thrive in warm

<figure><img src="image_4.png" /><figcaption>**Fig. 3.** Dataset 2.1: *Glycymeris pilosa* (G.P.) at Barbariga (BA), Pag Bay (PB), Pašman Channel (PC), and Drače (DR). For explanation see caption of Fig. 2.</figcaption></figure>
historical times, than *F. glaber* that was restricted to the annual thermal minimum. Consequently, *A. opercularis* and *P. jacobaeus* will potentially have an up to 3 months shorter annual growing season.

In Dataset 2, δ¹⁸O<sub>shell</sub> values for both *G. pilosa* (Fig. 3) and *V. verrucosa* (Fig. 4) range from negative to positive. This is in contrast with Dataset 1 species which displayed only positive values. Consequently, Dataset 2 species can thrive in warm

<figure><img src="image_4.png" /><figcaption>**Fig. 3.** Dataset 2.1: *Glycymeris pilosa* (G.P.) at Barbariga (BA), Pag Bay (PB), Pašman Channel (PC), and Drače (DR). For explanation see caption of Fig. 2.</figcaption></figure>
<figure><img src="image_5.png" /><figcaption>**Fig. 4.** Dataset 2.2: *Venus verrucosa* (V.V.) at Barbariga (BA), Pag Bay (PB), and Kaštela Bay (KB). For explanation see caption of Fig. 2.</figcaption></figure>

waters and are expected to have a longer growing season under extreme warming. Further, Dataset 2 species in the southern Adriatic (e.g., Kaštela Bay and Drače) are likely to benefit more from extreme warming than those living in the

north. Indeed, the temperatures along the eastern Adriatic coast will likely be warmer by up to 2°C, and about 0.4°C on average, in the south than in the north (Supporting Information Table S2; Fig. S2). Additionally, the warming is likely to
<figure><img src="image_5.png" /><figcaption>**Fig. 4.** Dataset 2.2: *Venus verrucosa* (V.V.) at Barbariga (BA), Pag Bay (PB), and Kaštela Bay (KB). For explanation see caption of Fig. 2.</figcaption></figure>

waters and are expected to have a longer growing season under extreme warming. Further, Dataset 2 species in the southern Adriatic (e.g., Kaštela Bay and Drače) are likely to benefit more from extreme warming than those living in the

north. Indeed, the temperatures along the eastern Adriatic coast will likely be warmer by up to 2°C, and about 0.4°C on average, in the south than in the north (Supporting Information Table S2; Fig. S2). Additionally, the warming is likely to
be stronger at shallower than at deeper sites. This will poten-
tially result in an up to 4 months longer annual growing sea-
son for *V. verrucosa* in Kaštela Bay than at Barbariga under
extreme warming. Furthermore, for *G. pilosa* (Fig. 3), δ¹⁸Oexp
distributions in historical and extreme warming conditions at
Barbariga and Pag Bay show nearly equal peaks in summer
and winter, while a more distinct peak during the winter sea-
son is present at Pašman Channel and Drače. This indicates
both different historical thermohaline conditions and varying
impacts of climate change at each location. Additionally, the
effect of extreme warming will be the strongest in Pašman
Channel, with a more than 3 months longer growing season.
The growth of this species was restricted from May to
November in historical times and it will likely be extended
to the entire year, especially in the central and southern
Adriatic (Pašman Channel and Drače) while, in the north
(Barbariga and Pag Bay), the shell growth will still be relatively
limited during winter months. *V. verrucosa* has fairly similar
growth conditions as *G. pilosa* (Fig. 4). In Kaštela Bay, where
conditions were most optimal for growth during the past,
with 244 d of shell growth per year, this species will presum-
ably have the ability to grow throughout the entire year. The
increase is also evident in the northern locations (from 174 to
240 growth days at Barbariga and from 172 to 257 growth
days at Pag Bay), although less pronounced as for *G. pilosa*.
Consequently, species from Dataset 2 would clearly benefit
from climate warming, which predicts an extended growing
season and favorable thermal conditions for growth.

Discussion

Our preliminary study is based on the unique combination
of δ¹⁸Oshell data of five bivalve species and the AdriSC
kilometer-scale atmosphere-ocean climate model results for
historical and extreme warming conditions. Overall, it reveals
a substantial influence of extreme warming on the expected
duration of all bivalve growing seasons that can be summa-
rized as follows:

* At all sites the five studied species will likely benefit from climate warming during the colder season, i.e., they will grow longer than currently between November and April.

* The expected growing season of scallops (Dataset 1 species) will likely substantially decrease during summer, i.e., between May and October, resulting in an up to 3 months shorter growing season.

* *G. pilosa* and *V. verrucosa* (Dataset 2 species) in the south and at shallower habitats will likely be more impacted by climate warming than those living in the north and deeper habitats, resulting in an up to 4 months' longer growth season.

However, this study presents several limitations. First, the
evaluation of the AdriSC temperatures (Table S1; Fig. S1)

shows that the results should be interpreted with some cau-
tion, particularly at Drače, where the model results deviate the
most from the observations. Second, the climate uncertainty
derived from ensembles of simulations forced by multiple
global climate models under multiple warming scenarios
(Semenov and Stratonovitch 2010) is not considered. Third,
the δ¹⁸Oshell thresholds defining the conditions favorable for
bivalve growth are derived from a small number of specimens,
particularly for Dataset 1 (e.g., only two specimens of
A. opercularis and F. glaber; Table 1), which might impact their
accuracy. Fourth, in contrast to the study by Abe (2021), our
results do not account for the impact of extreme warming on
the food availability (Norkko et al. 2005) derived from either
the primary production (e.g., Bonitz et al. 2018) or the acidifi-
cation (e.g., Tan et al. 2019). Lastly, no adaption of the inves-
tigated species to the new climate is assumed as no
information is available, although some bivalve species are
found to better adapt if inhabiting zones prone to higher tem-
perature oscillations (Zhang et al. 2020). Despite the crucial
need to reproduce these processes in complex coastal environ-
ments (Mishra et al. 2023), no kilometer-scale biogeochemical
climate model has yet been implemented (Ani and
Robson 2021). Therefore, our analyses could only use the
reconstructed δ¹⁸Oexp values. The reconstructed δ¹⁸Oexp values
are, however, in a good agreement with the measured δ¹⁸Oshell
values, as demonstrated in previous research (Peharda
et al. 2019a,b; Uvanović et al. 2021; Ezgeta-Balić et al. 2022)
that employed the alignment technique described in the
Methods. Although different climate models were used in
these studies, the AdriSC model was successfully employed
in Ezgeta-Balić et al. (2022).

In terms of bivalve management along the eastern Adriatic
Sea, *P. jacobaeus* is commercially the most important scallop
species in the Adriatic (Mattei and Pellizzato 1996; Peharda
et al., 2019a; Ezgeta-Balić et al., 2022). In the last 40 years,
due to a combination of overfishing, decline in eutrophica-
tion (Iveša et al. 2016) and positive temperature trends
(Vilibić et al. 2019), its biomass has already declined
(Mazzoldi et al. 2014). As its main growing season could fur-
ther be reduced by 30%, its accelerated decline or eventual
vanishing could occur by the end of this century in the north-
ern Adriatic. Owing to a high nutritional quality, *A. opercularis*
has been proposed for human consumption (Kovačić
et al. 2023), with winter and spring being the most suitable
period of the year for the harvest. However, our results indi-
cate a substantial change in its growing season, posing a ques-
tion if and how this species can adapt to that. *F. glaber*, which
grows particularly fast and is free of diseases, could also be
farmed (Marčeta et al. 2016). However, due to its sensitivity to
warming and heavy metal pollution (Nardi et al. 2018;
Telahigue et al. 2022) and the projected shortening of its
growing season by 3 months, its commercial exploitation may
never be possible. Our results indicate that, among the three
be stronger at shallower than at deeper sites. This will poten-
tially result in an up to 4 months longer annual growing sea-
son for *V. verrucosa* in Kaštela Bay than at Barbariga under
extreme warming. Furthermore, for *G. pilosa* (Fig. 3), δ¹⁸Oexp
distributions in historical and extreme warming conditions at
Barbariga and Pag Bay show nearly equal peaks in summer
and winter, while a more distinct peak during the winter sea-
son is present at Pašman Channel and Drače. This indicates
both different historical thermohaline conditions and varying
impacts of climate change at each location. Additionally, the
effect of extreme warming will be the strongest in Pašman
Channel, with a more than 3 months longer growing season.
The growth of this species was restricted from May to
November in historical times and it will likely be extended
to the entire year, especially in the central and southern
Adriatic (Pašman Channel and Drače) while, in the north
(Barbariga and Pag Bay), the shell growth will still be relatively
limited during winter months. *V. verrucosa* has fairly similar
growth conditions as *G. pilosa* (Fig. 4). In Kaštela Bay, where
conditions were most optimal for growth during the past,
with 244 d of shell growth per year, this species will presum-
ably have the ability to grow throughout the entire year. The
increase is also evident in the northern locations (from 174 to
240 growth days at Barbariga and from 172 to 257 growth
days at Pag Bay), although less pronounced as for *G. pilosa*.
Consequently, species from Dataset 2 would clearly benefit
from climate warming, which predicts an extended growing
season and favorable thermal conditions for growth.

Discussion

Our preliminary study is based on the unique combination
of δ¹⁸Oshell data of five bivalve species and the AdriSC
kilometer-scale atmosphere-ocean climate model results for
historical and extreme warming conditions. Overall, it reveals
a substantial influence of extreme warming on the expected
duration of all bivalve growing seasons that can be summa-
rized as follows:

* At all sites the five studied species will likely benefit from climate warming during the colder season, i.e., they will grow longer than currently between November and April.

* The expected growing season of scallops (Dataset 1 species) will likely substantially decrease during summer, i.e., between May and October, resulting in an up to 3 months shorter growing season.

* *G. pilosa* and *V. verrucosa* (Dataset 2 species) in the south and at shallower habitats will likely be more impacted by climate warming than those living in the north and deeper habitats, resulting in an up to 4 months' longer growth season.

However, this study presents several limitations. First, the
evaluation of the AdriSC temperatures (Table S1; Fig. S1)

shows that the results should be interpreted with some cau-
tion, particularly at Drače, where the model results deviate the
most from the observations. Second, the climate uncertainty
derived from ensembles of simulations forced by multiple
global climate models under multiple warming scenarios
(Semenov and Stratonovitch 2010) is not considered. Third,
the δ¹⁸Oshell thresholds defining the conditions favorable for
bivalve growth are derived from a small number of specimens,
particularly for Dataset 1 (e.g., only two specimens of
A. opercularis and F. glaber; Table 1), which might impact their
accuracy. Fourth, in contrast to the study by Abe (2021), our
results do not account for the impact of extreme warming on
the food availability (Norkko et al. 2005) derived from either
the primary production (e.g., Bonitz et al. 2018) or the acidifi-
cation (e.g., Tan et al. 2019). Lastly, no adaption of the inves-
tigated species to the new climate is assumed as no
information is available, although some bivalve species are
found to better adapt if inhabiting zones prone to higher tem-
perature oscillations (Zhang et al. 2020). Despite the crucial
need to reproduce these processes in complex coastal environ-
ments (Mishra et al. 2023), no kilometer-scale biogeochemical
climate model has yet been implemented (Ani and
Robson 2021). Therefore, our analyses could only use the
reconstructed δ¹⁸Oexp values. The reconstructed δ¹⁸Oexp values
are, however, in a good agreement with the measured δ¹⁸Oshell
values, as demonstrated in previous research (Peharda
et al. 2019a,b; Uvanović et al. 2021; Ezgeta-Balić et al. 2022)
that employed the alignment technique described in the
Methods. Although different climate models were used in
these studies, the AdriSC model was successfully employed
in Ezgeta-Balić et al. (2022).

In terms of bivalve management along the eastern Adriatic
Sea, *P. jacobaeus* is commercially the most important scallop
species in the Adriatic (Mattei and Pellizzato 1996; Peharda
et al., 2019a; Ezgeta-Balić et al., 2022). In the last 40 years,
due to a combination of overfishing, decline in eutrophica-
tion (Iveša et al. 2016) and positive temperature trends
(Vilibić et al. 2019), its biomass has already declined
(Mazzoldi et al. 2014). As its main growing season could fur-
ther be reduced by 30%, its accelerated decline or eventual
vanishing could occur by the end of this century in the north-
ern Adriatic. Owing to a high nutritional quality, *A. opercularis*
has been proposed for human consumption (Kovačić
et al. 2023), with winter and spring being the most suitable
period of the year for the harvest. However, our results indi-
cate a substantial change in its growing season, posing a ques-
tion if and how this species can adapt to that. *F. glaber*, which
grows particularly fast and is free of diseases, could also be
farmed (Marčeta et al. 2016). However, due to its sensitivity to
warming and heavy metal pollution (Nardi et al. 2018;
Telahigue et al. 2022) and the projected shortening of its
growing season by 3 months, its commercial exploitation may
never be possible. Our results indicate that, among the three
studied scallop species, *F. glaber* is expected to be most impacted by climate warming scenarios. In contrast, *G. pilosa* which, due to its longevity is regarded as an archive for environmental change in the Mediterranean (Peharda et al. 2016), and *V. verrucosa* which has a high commercial value and occurs in the entire Mediterranean Sea (Arneri et al. 1998; Poppe and Goto 2000), will benefit from extreme warming with an up to 45% longer growing season along the southeastern Adriatic coast, where *V. verrucosa* could successfully be exploited in the future.

To the date, it has not been assessed how the growing season of the target species vary across latitudes or (paleo)climates. However, for *Mercenaria mercenaria* it has been found that variations in growth rate over different latitudes do not affect the shell δ¹⁸O values (Elliot et al. 2003), while latitudinal changes in temperature were the major factor in shaping growth rate (Palmer et al. 2021). Similarly, *P. maximus* exhibits a latitudinal reduction of the growth due to a shorter phenology (Chauvaud et al. 2012). Assuming the same for the species investigated here, a migration to higher latitudes may be a way for preserving bivalves for which a substantial decline in growth in the future climate is envisaged. Changes in growth dynamics, for these, as well as other bivalve species, will potentially have impact on the marine food webs. Bivalve predators belong to different taxa, including echinoderms, crustaceans, gastropods, cephalopods, and fish (see Gosling 2015). Thereby, it can be expected that a wide range of marine organisms will be impacted by changes in seawater temperature.

If generalized, this study might lead to better adaptation plans (e.g., Roman et al. 1999; Papa et al. 2021; Marčeta et al. 2022) for possible cultivation of the studied bivalve species, along the eastern Adriatic Sea, as well as in other areas where these species live. Furthermore, similar scenarios of changes in growth patterns may occur for other bivalve species in temperate coastal areas around the world, which may constitute a growing portion of the food production in the future (Jennings et al. 2016; Tan et al. 2020; Willer et al. 2021).

## References

* Abe, H. 2021. Climate warming promotes Pacific oyster (*Magallana gigas*) production in a subarctic lagoon and bay, Japan: Projection of future trends using a three dimensional physical-ecosystem coupled model. Reg. Stud. Mar. Sci. **47**: 101968. doi:10.1016/j.rsma.2021.101968

* Ani, C. J., and B. Robson. 2021. Responses of marine ecosystems to climate change impacts and their treatment in biogeochemical ecosystem models. Mar. Pollut. Bull. **166**: 112233. doi:10.1016/j.marpolbul.2021.112223

* Arneri, A., G. Giannetti, and B. Antolini. 1998. Age determination and growth of *Venus verrucosa* L. (Bivalvia: Veneridae)

in the southern Adriatic and the Aegean Sea. Fish. Res. **38**: 193-198. doi:10.1016/S0165-7836(98)00146-5

* Beuvier, J., F. Sevault, M. Herrmann, H. Kontoyiannis, W. Ludwig, M. Rixen, E. Stanev, K. Béranger, and S. Somot. 2010. Modeling the Mediterranean Sea interannual variability during 1961-2000: Focus on the eastern Mediterranean transient. J. Geophys. Res. Atmos. **115**: C08017. doi: 10.1029/2009JC005950

* Bonitz, F. G. W., C. Andersson, T. Trofimova, and H. Hatum. 2018. Links between phytoplankton dynamics and shell growth of *Arctica islandica* on the Faroe shelf. J. Mar. Syst. **179**: 72-87. doi:10.1016/j.jmarsys.2017.11.005

* Branscombe, T., R. Schulting, J. Le-Thorp, and M. J. Leng. 2021. The potential of marine bivalve *Spisula sachalinensis* as a marine temperature record. Palaeogeogr. Palaeoclimatol. Palaeoecol. **582**: 110634. doi:10.1016/j.palaeo.2021.110634

* Chauvaud, L., and others. 2012. Variation in size and growth of the great scallop *Pecten maximus* along a latitudinal gradient. PloS One **7**: e37717. doi:10.1371/journal.pone.0037717

* Coplen, T. B., C. Kendall, and J. Hopple. 1983. Comparison of stable isotope reference samples. Nature **302**: 236-238. doi: 10.1038/302236a0

* Cubillo, A. M., J. G. Ferreira, J. Lencart-Silva, N. G. H. Taylor, A. Kennerley, J. Guilder, S. Kay, and P. Kamermans. 2021. Direct effects of climate change on productivity of European aquaculture. Aquac. Int. **29**: 1561-1590. doi:10.1007/s10499-021-00694-6

* Denamiel, C., J. Šepić, D. Ivanković, and I. Vilibić. 2019. The Adriatic Sea and coast modelling suite: Evaluation of the meteotsunami forecast component. Ocean Model. **135**: 71-93. doi:10.1016/j.ocemod.2019.02.003

* Denamiel, C., P. Pranić, F. Quentin, H. Mihanović, and I. Vilibić. 2020. Pseudo-global warming projections of extreme wave storms in complex coastal regions: The case of the Adriatic Sea. Climate Dynam. **55**: 2483-2509. doi: 10.1007/s00382-020-05397-x

* Denamiel, C., I. Tojčić, and I. Vilibić. 2021a. Balancing accuracy and efficiency of atmospheric models in the northern Adriatic during severe bora events. J. Geophys. Res. Atmos. **126**: e2020JD033516. doi:10.1029/2020JD033516

* Denamiel, C., P. Pranić, D. Ivanković, I. Tojčić, and I. Vilibić. 2021b. Performance of the Adriatic Sea and coast (AdriSC) climate component – A COAWST V3.3-based coupled atmosphere-ocean modelling suite: Atmospheric dataset. Geosci. Model Dev. **14**: 3995-4017. doi:10.5194/gmd-14-3995-2021

* Denamiel, C., I. Tojčić, P. Pranić, and I. Vilibić. 2022. Modes of the BiOS-driven Adriatic Sea thermohaline variability. Climate Dynam. **59**: 1097-1113. doi:10.1007/s00382-022-06178-4

* Denamiel, C. M., M. Peharda, and B. Schöne. 2023. *Projecting expected growth period of bivalves [Dataset]*. Open Science Framework. doi:10.17605/osf.io/ecqu3
studied scallop species, *F. glaber* is expected to be most impacted by climate warming scenarios. In contrast, *G. pilosa* which, due to its longevity is regarded as an archive for environmental change in the Mediterranean (Peharda et al. 2016), and *V. verrucosa* which has a high commercial value and occurs in the entire Mediterranean Sea (Arneri et al. 1998; Poppe and Goto 2000), will benefit from extreme warming with an up to 45% longer growing season along the southeastern Adriatic coast, where *V. verrucosa* could successfully be exploited in the future.

To the date, it has not been assessed how the growing season of the target species vary across latitudes or (paleo)climates. However, for *Mercenaria mercenaria* it has been found that variations in growth rate over different latitudes do not affect the shell δ¹⁸O values (Elliot et al. 2003), while latitudinal changes in temperature were the major factor in shaping growth rate (Palmer et al. 2021). Similarly, *P. maximus* exhibits a latitudinal reduction of the growth due to a shorter phenology (Chauvaud et al. 2012). Assuming the same for the species investigated here, a migration to higher latitudes may be a way for preserving bivalves for which a substantial decline in growth in the future climate is envisaged. Changes in growth dynamics, for these, as well as other bivalve species, will potentially have impact on the marine food webs. Bivalve predators belong to different taxa, including echinoderms, crustaceans, gastropods, cephalopods, and fish (see Gosling 2015). Thereby, it can be expected that a wide range of marine organisms will be impacted by changes in seawater temperature.

If generalized, this study might lead to better adaptation plans (e.g., Roman et al. 1999; Papa et al. 2021; Marčeta et al. 2022) for possible cultivation of the studied bivalve species, along the eastern Adriatic Sea, as well as in other areas where these species live. Furthermore, similar scenarios of changes in growth patterns may occur for other bivalve species in temperate coastal areas around the world, which may constitute a growing portion of the food production in the future (Jennings et al. 2016; Tan et al. 2020; Willer et al. 2021).

## References

* Abe, H. 2021. Climate warming promotes Pacific oyster (*Magallana gigas*) production in a subarctic lagoon and bay, Japan: Projection of future trends using a three dimensional physical-ecosystem coupled model. Reg. Stud. Mar. Sci. **47**: 101968. doi:10.1016/j.rsma.2021.101968

* Ani, C. J., and B. Robson. 2021. Responses of marine ecosystems to climate change impacts and their treatment in biogeochemical ecosystem models. Mar. Pollut. Bull. **166**: 112233. doi:10.1016/j.marpolbul.2021.112223

* Arneri, A., G. Giannetti, and B. Antolini. 1998. Age determination and growth of *Venus verrucosa* L. (Bivalvia: Veneridae)

in the southern Adriatic and the Aegean Sea. Fish. Res. **38**: 193-198. doi:10.1016/S0165-7836(98)00146-5

* Beuvier, J., F. Sevault, M. Herrmann, H. Kontoyiannis, W. Ludwig, M. Rixen, E. Stanev, K. Béranger, and S. Somot. 2010. Modeling the Mediterranean Sea interannual variability during 1961-2000: Focus on the eastern Mediterranean transient. J. Geophys. Res. Atmos. **115**: C08017. doi: 10.1029/2009JC005950

* Bonitz, F. G. W., C. Andersson, T. Trofimova, and H. Hatum. 2018. Links between phytoplankton dynamics and shell growth of *Arctica islandica* on the Faroe shelf. J. Mar. Syst. **179**: 72-87. doi:10.1016/j.jmarsys.2017.11.005

* Branscombe, T., R. Schulting, J. Le-Thorp, and M. J. Leng. 2021. The potential of marine bivalve *Spisula sachalinensis* as a marine temperature record. Palaeogeogr. Palaeoclimatol. Palaeoecol. **582**: 110634. doi:10.1016/j.palaeo.2021.110634

* Chauvaud, L., and others. 2012. Variation in size and growth of the great scallop *Pecten maximus* along a latitudinal gradient. PloS One **7**: e37717. doi:10.1371/journal.pone.0037717

* Coplen, T. B., C. Kendall, and J. Hopple. 1983. Comparison of stable isotope reference samples. Nature **302**: 236-238. doi: 10.1038/302236a0

* Cubillo, A. M., J. G. Ferreira, J. Lencart-Silva, N. G. H. Taylor, A. Kennerley, J. Guilder, S. Kay, and P. Kamermans. 2021. Direct effects of climate change on productivity of European aquaculture. Aquac. Int. **29**: 1561-1590. doi:10.1007/s10499-021-00694-6

* Denamiel, C., J. Šepić, D. Ivanković, and I. Vilibić. 2019. The Adriatic Sea and coast modelling suite: Evaluation of the meteotsunami forecast component. Ocean Model. **135**: 71-93. doi:10.1016/j.ocemod.2019.02.003

* Denamiel, C., P. Pranić, F. Quentin, H. Mihanović, and I. Vilibić. 2020. Pseudo-global warming projections of extreme wave storms in complex coastal regions: The case of the Adriatic Sea. Climate Dynam. **55**: 2483-2509. doi: 10.1007/s00382-020-05397-x

* Denamiel, C., I. Tojčić, and I. Vilibić. 2021a. Balancing accuracy and efficiency of atmospheric models in the northern Adriatic during severe bora events. J. Geophys. Res. Atmos. **126**: e2020JD033516. doi:10.1029/2020JD033516

* Denamiel, C., P. Pranić, D. Ivanković, I. Tojčić, and I. Vilibić. 2021b. Performance of the Adriatic Sea and coast (AdriSC) climate component – A COAWST V3.3-based coupled atmosphere-ocean modelling suite: Atmospheric dataset. Geosci. Model Dev. **14**: 3995-4017. doi:10.5194/gmd-14-3995-2021

* Denamiel, C., I. Tojčić, P. Pranić, and I. Vilibić. 2022. Modes of the BiOS-driven Adriatic Sea thermohaline variability. Climate Dynam. **59**: 1097-1113. doi:10.1007/s00382-022-06178-4

* Denamiel, C. M., M. Peharda, and B. Schöne. 2023. *Projecting expected growth period of bivalves [Dataset]*. Open Science Framework. doi:10.17605/osf.io/ecqu3
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Acknowledgments

Comments and suggestions provided by the editor and two anonymous
reviewers greatly improved the manuscript. This work has been supported
by the project "Bivalves: Archives of environmental change in the coastal
marine ecosystems" (BivACME, Croatian Science Foundation Grant IP-
2019-2104-8542). AdriSC climate simulations have been enabled by the
European Centre for Middle-range Weather Forecast (ECMWF) through
Special Projects "Using stochastic surrogate methods for advancing
towards reliable meteotsunami early warning systems" and "Numerical
modelling of the Adriatic-Ionian decadal and inter-annual oscillations:
From realistic to process-oriented simulations."

Submitted 03 November 2023

Revised 21 March 2024

Accepted 01 April 2024
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Acknowledgments

Comments and suggestions provided by the editor and two anonymous
reviewers greatly improved the manuscript. This work has been supported
by the project "Bivalves: Archives of environmental change in the coastal
marine ecosystems" (BivACME, Croatian Science Foundation Grant IP-
2019-2104-8542). AdriSC climate simulations have been enabled by the
European Centre for Middle-range Weather Forecast (ECMWF) through
Special Projects "Using stochastic surrogate methods for advancing
towards reliable meteotsunami early warning systems" and "Numerical
modelling of the Adriatic-Ionian decadal and inter-annual oscillations:
From realistic to process-oriented simulations."

Submitted 03 November 2023

Revised 21 March 2024

Accepted 01 April 2024