Eating contest between native and non-indigenous bivalve species: estimating capture efficiencies and clearance rates using natural seston

125 Eating contest between native and non-indigenous bivalve species: estimating capture efficiencies and clearance rates using natural seston Sara Cabral1, Frederico Carvalho1, Joana P. C. Cruz1, Joshua Heumüller1, Jose M. F. Babarro2, Luc A. Comeau3, Paula Chainho1,4 , Ana C. Brito1,5 1 MARE—Marine and Environmental Science Centre, ARNET—Aquatic Research Network, Faculdade de Ciências, Universidade de Lisboa, 1749-016 Lisboa, Portugal 2 Instituto de Investigaciones Marinas CSIC, Eduardo Cabello 6, 36208 Vigo, Spain 3 Fisheries and Oceans Canada, Gulf Fisheries Centre, P.O. Box 5030, Moncton, E1C 9B6 New Brunswick, Canada 4 Escola Superior de Tecnologia, Instituto Politécnico de Setúbal, 2914-508 Setúbal, Portugal 5 Departamento de Biologia Vegetal, Faculdade de Ciências, Universidade de Lisboa, 1749-016 Lisboa, Portugal Corresponding author: Sara Cabral (sdcabr[email protected]) Copyright: This is an open access article distributed under the terms of the CC0 Public Domain Dedication. Research Article Abstract Despite the burgeoning number of non-indigenous species (NIS) in worldwide coastal ecosystems, the quantification of their direct impacts on native communities remains largely unexplored. This is particularly true concerning feeding competition in sympatric filter-feeding bivalves. In this study, our aim was to fill a gap of knowledge on the potential trophic competition between native and non-indigenous bivalves, namely by focusing on three species that co-occur in Portuguese estuarine systems: the native cockle (Cerastoderma edule) and Portuguese oyster (Magallana angulata) and the non-indigenous Manila clam (Ruditapes philippinarum). The specific objectives were to i) estimate their capture efficiency (≈ particle retention efficiency; CE); ii) assess their clearance rates (CR); and iii) provide a science-based support for suitable management measures regarding NIS. Experiments were conducted in both field and laboratory conditions using the natural seston present in the seawater. The CE was higher for the larger size classes (8–14 μm) of particles measured (ranging from 4–14 μm), regardless of the species. While the individual CRs were not significantly different among species, the CR per gram of ash-free dry body tissue weight was significantly higher for the native cockle, suggesting that the NIS does not hold a competitive advantage in clearing suspended particles. However, the Manila clam might be limi ting food sources availability to the native species since there is an overlap of their ecological niches. Key words: Cerastoderma edule, Magallana angulata, Portugal, retention efficiency, Ruditapes philippinarum Introduction Biological invasions are considered one of the main threats to the marine environment, being a key driver of biodiversity change (Ruiz et al. 1999; Streftaris et al. 2005; Zorita et al. 2013; Jimenez et al. 2018). Non-indigenous species (NIS) have become ubiquitous across coastal ecosystems such as estuaries, bays, and coastal lagoons (Ruiz et al. 2009), since these are sheltered zones presenting a high level of anthropogenic pressure and activities (Ruiz et al. 2000). The adverse impacts of NIS are multiple and interdisciplinary ranging from economic (affecting fisheries, Academic editor: Tammy Robinson-Smythe Received: 30 January 2025 Accepted: 25 March 2025 Published: 7 October 2025 Citation: Cabral S, Carvalho F, Cruz JPC, Heumüller J, Babarro JMF, Comeau LA, Chainho P, Brito AC (2025) Eating contest between native and non-indigenous bivalve species: estimating capture efficiencies and clearance rates using natural seston. In: Anastácio P, Ribeiro F, Chainho P (Eds) Invasions in Aquatic Systems. NeoBiota 102: 125–150. https://doi. org/10.3897/neobiota.102.148326 NeoBiota 102: 125–150 (2025) DOI: 10.3897/neobiota.102.148326 Advancing research on alien species and biological invasions A peer-reviewed open-access journal NeoBiota 126 NeoBiota 102: 125–150 (2025), DOI: 10.3897/neobiota.102.148326 Sara Cabral et al.: Estimating capture efficiencies and clearance rates of three bivalve species aquaculture, and tourism sectors, among others; Galil et al. 2014) to ecological (including single species interactions, population declines, local extinctions, changes in community composition, and effects on entire ecosystem processes and functions; Katsanevakis et al. 2014; Anton et al. 2019). Despite the growing knowledge about NIS impacts, this information is not yet properly addressed under the European Union legal frameworks for the protection and management of marine and freshwater ecosystems. Although temporal occurrence, abundance and spatial distribution of NIS are indicators (descriptor D2) used to assess the Environmental Status (ES) of marine waters under the Marine Strategy Framework Directive (MSFD 2008), no specific consensual tools to measure the magnitude of bioinvasion impacts have been formally adopted yet. Likewise, the indices proposed to assess the Ecological Quality Status (EQS) in transitional and coastal waters under the Water Framework Directive (WFD 2000) do not address impacts of NIS (Zaiko and Daunys 2015). One of the potential ecological hazards posed by the introduction of NIS that should be considered in assessment tools and by decision makers is the displacement of native species (Truhlar et al. 2014). This can occur due to numerous reasons such as restricted mobility when competing for space availability (Shinen and Morgan 2009), direct predation (Battini et al. 2021), parasitism and diseases (Stiger-Pouvreau and Thouzeau 2015) or a high potential for food competition between native and NIS (Comeau et al. 2015; Nielsen et al. 2017; Sonier 2017; Rodrigues et al. 2023). Clearance rate measurements (CR; defined as the volume of water cleared of particles per unit of time) may offer insight into the potential trophic competition between suspension-feeding bivalves. An extensive CR literature spanning across decades (Møhlenberg and Riisgård 1978; Iglesias et al. 1998; Troost et al. 2009; Yu et al. 2017) revealed the sensitivity of CR values to several factors, such as temperature (Pernet et al. 2008; Cranford et al. 2011), salinity (Casas et al. 2018), distinctive experimental set-ups (Riisgård 2001), food quantity and quality (Navarro and Iglesias 1993), and contamination level at sampling sites (Widdows 2001). To date, these experiments have been conducted primarily under laboratory conditions (Møhlenberg and Riisgård 1978, 1979; Denis et al. 1999; Nakamura 2001; Haure et al. 2003; Troost et al. 2009; Joyce et al. 2019) or using artificial diets (Navarro and Widdows 1997; Petersen et al. 2004; Pascoe et al. 2009; Nagasoe et al. 2011; Tamayo et al. 2013). Nevertheless, it is recognized that laboratory CR results may deviate from in situ measurements (Cranford 2001; Petersen et al. 2004; Clavier and Chauvaud 2010; Rosa et al. 2018). Therefore, different species should be compared within the same trophic competition study (Troost et al. 2009), with the animals maintained under the same experimental conditions and preferably grazing on all the diversity of seston naturally available in the seawater. Notwithstanding the vast research undertaken focusing on CR, some terms (e.g. filtration rate, retention efficiency, pumping rate, capture efficiency) have been interchangeably used (Rosa et al. 2018; Steeves et al. 2022), leading to confusion. The term retention efficiency is mostly used to refer to the proportion of particles that encounter the gill filaments and are cleared from the water, being essential to determine it before measuring a bivalve’s CR (Cranford et al. 2016). Following the recommendation of Rosa et al. (2018), ‘retention efficiency’ will hereafter be denoted as capture efficiency (CE). This is because the number of particles that are retained by the gill can only be accurately determined if in vivo direct approaches are used (Shimeta and Jumars 1991; Ward et al. 1998; Rosa et al. 2018). 127 NeoBiota 102: 125–150 (2025), DOI: 10.3897/neobiota.102.148326 Sara Cabral et al.: Estimating capture efficiencies and clearance rates of three bivalve species The non-indigenous Manila clam Ruditapes philippinarum is a bivalve native to the Indo-Pacific Ocean that was firstly reported in Portugal in 1984 for aquaculture purposes (Ruano and Sobral 2000). Its presence throughout European countries has been recognized to affect biogeochemical cycles and macroalgal growth (Bartoli et al. 2001) as well as causing a reduction in both density and spatial distribution area of native bivalves (Pranovi et al. 2006, ICES 2008). Furthermore, Maia et al. (2025) suggested that R. philippinarum holds a competitive advantage when compared with native sympatric species, not only due to its rapid growth and early sexual maturity, but also because of its broad diet and flexibility in feeding on multiple food sources according to their availability in the water column. The purpose of the present study was to assess the potential trophic competition between native and non-indigenous bivalves feeding on natural particles. The specific objectives were to i) compare the CE of two native species (the cockle Cerastoderma edule and the Portuguese oyster Magallana angulata) and one non-indigenous species (the Manila clam R. philippinarum); ii) estimate their CR under field and laboratory conditions; and iii) provide a science-based support to competent authorities and regulators for suitable management measures regarding NIS. Methods Experimental design To estimate the Capture Efficiency (CE) and the Clearance Rate (CR), a total of 14 experiments (seven in the field and seven in the laboratory; Fig. 1) were conducted with three species of bivalves: the native common cockle (Cerastoderma edule) and Portuguese oyster (Magallana angulata) and the non-indigenous Manila clam (Ruditapes philippinarum). The field experiments took place between September 20th and October 11th, 2018, in an oyster aquaculture farm located in the Sado estuary (Setúbal, Portugal). This site is known for its wide variability of in-situ environmental parameters, such as temperature and salinity (AQUASADO 2022). Thus, to reduce the influence of these fluctuating variables, the experiments were also replicated in the laboratory, where conditions are inherently more stable. These experiments were conducted between the 5th and the 21st of December 2018 at the MARE-ULisboa (Lisboa, Portugal). Field experiments Wild specimens of C. edule and R. philippinarum were collected in an intertidal area of the Sado estuary (38°32.01'N, 8°46.85'W), while M. angulata individuals were obtained directly from the aquaculture farm where the experiments occurred. A set of 12 incubators (with a maximum volume of 1.2 L each) were placed on a mobile platform (Fig. 2). A single specimen of one of the selected species was placed in each incubator and three empty incubators were used as a control. Incubators were distributed randomly over the experiment platform and an air pump was attached to the wall of each incubator to homogenize the water inside them and minimize sedimentation of particles. Three replicates of each species were used per experiment. The platform was lowered to maintain the incubators permanently immersed and after 48 hours of acclimatization the experiment began by gently raising the platform with 128 NeoBiota 102: 125–150 (2025), DOI: 10.3897/neobiota.102.148326 Sara Cabral et al.: Estimating capture efficiencies and clearance rates of three bivalve species Figure 1. Diagram of the water samples (volume and timings) collected in the field and laboratory experiments for the three bivalve species (C. edule, M. angulata and R. philippinarum). Figure 2. Experimental set-up of the field experiments, conducted in the Sado estuary (Setúbal, Portugal), using a mobile platform with incubators for three bivalve species (C. edule, M. angulata and R. philippinarum) and controls. the incubators until all were isolated out of the water column. Water temperature and salinity were continuously monitored with two loggers HOBO U24 Conductivity (U24-002-C): one was positioned near the mobile platform and the other one further away at a fixed point in the water canal (Table 1). 129 NeoBiota 102: 125–150 (2025), DOI: 10.3897/neobiota.102.148326 Sara Cabral et al.: Estimating capture efficiencies and clearance rates of three bivalve species Each experiment was conducted for 90 minutes, with water samples being collected from each incubator every 15 minutes, including a sample at T0 (time of start) and another at Tf (end time). A total of 45 mL of water was collected from each incubator at each time, preserved with 1% Lugol’s Solution (Merck 109261) and subsequently stored in the dark at room temperature until further particle measurements. In addition, 15 mL of water were collected from each incubator at T0 to measure turbidity with a Lovibond infrared turbidimeter TB 210 IR (Table 1), as a proxy of the available seston load. Unpublished data measured at the same location revealed a high correlation between turbidity and suspended particulate matter (SPM; See Suppl. material 1); denoting that the average SPM during field experiments was 38.35 mg L-1. Laboratory experiments The field experiments were replicated in the laboratory using seawater collected in the Sado estuary. Similarly, specimens of C. edule and R. philippinarum were collected in an intertidal area of the Sado estuary (38°32.01'N, 8°46.85'W), while M. angulata individuals were obtained directly from the aquaculture farm where the field experiments occurred. Afterwards, all animals were transported cooled with icepacks to MARE-ULisboa, where they were maintained permanently immersed for 48 hours in an aquarium to acclimate. A continuous natural seawater flow was maintained in the aquarium to ensure adequate physico-chemical conditions and food supply. The laboratory experiments used 15 incubators (with a maximum volume of 1.2 L each) placed inside an aquarium (with 200 L of volume) with seawater (Fig. 3). Four specimens of each of the selected species (C. edule, M. angulata and R. philippinarum) were placed in four individual aerated incubators and three empty aerated incubators were used as control. Incubators were randomly distributed inside the aquarium. Water temperature and salinity (Table 1) were continuously monitored with one logger HOBO U24 Conductivity (U24-002-C), while dissolved oxygen and pH values were recorded daily with a multiparametric probe YSI EXO2. Additionally, the concentration of ammonia was also monitored by spectrophotometry to guarantee that it was always low. Samples of 45 mL were taken every 15 minutes from each incubator, for a total of 60 minutes during each experiment, including a sample at T0 (time of start) and another at Tf (end time). These samples were preserved with 1% Lugol’s Solution (Merck 109261) and stored in the dark at room temperature until subsequent particle measurements. In addition, 15 mL of water were also collected from each incubator during T0 (time of start) to measure the turbidity with a Lovibond infrared turbidimeter TB 210 IR (Table 1) with the aim of verifying the availability of particles. According to the correlation between turbidity and SPM (See Suppl. material 1), the average SPM during the laboratory experiments was 11.33 mg L-1. Table 1. Environmental conditions during field and laboratory experiments. Experiment Date Temperature (min. – max. (°C)) Salinity (min. – max.) Turbidity (mean ± s.d. (NTU)) Field September to October 2018 13–20 21–25 30.88 ± 21.35 Laboratory December 2018 15–20 22–25 6.32 ± 3.01 130 NeoBiota 102: 125–150 (2025), DOI: 10.3897/neobiota.102.148326 Sara Cabral et al.: Estimating capture efficiencies and clearance rates of three bivalve species Valvometry In both experiments, valve opening movements of the bivalves were recorded following a methodology described in Nagai et al. (2006) and Comeau et al. (2012). Each bivalve specimen had a coated Hall element sensor (HW-300a, Asahi Kasei, Japan) glued to one valve and a magnet attached to the other valve (directly in the opposite position of the Hall sensor), creating a magnetic field (Fig. 4). This magnetic field was recorded as output voltage by strain recording devices (DC 104R, Tokyo Sokki Kenkyujo Co., Japan) and afterwards converted into valve opening distance. These measurements allowed the identification of specimens that were completely and consistently closed during the experiments and thus, undoubtedly not interacting with their environment. These specimens were discarded from the experiments as they were presumably stressed by the manipulations. This approach minimized the potential underestimation of CR. Estimation of the Condition Index (CI) At the end of the experiments, all bivalves were placed in separate plastic bags, preserved in a freezer at -20 °C and stored for future analysis. Later, all specimens were processed in the laboratory: the bivalve shells were first measured and then the soft body was separated from the shell with a scalpel and tweezers, and both parts were dried at 60 °C for 72 hours, to obtain the dry weight. Lastly, the dry meat Figure 3. Experimental set-up of the laboratory experiments, conducted at the MARE-ULisboa (Lisboa, Portugal), using an aquarium with incubators for the three bivalve species (C. edule, M. angulata and R. philippinarum). 131 NeoBiota 102: 125–150 (2025), DOI: 10.3897/neobiota.102.148326 Sara Cabral et al.: Estimating capture efficiencies and clearance rates of three bivalve species was burnt in a muffle furnace until weight stabilization at 500 °C, to estimate the ashes’ weight. The Condition Index (CI) was calculated according to the following formula adapted from Walne and Mann (1975): The mean shell length, mean ash-free dry weight (AFDW) of the body tissue and mean condition index (CI) for each species used in field and laboratory experiments are summarized in Table 2. Taking into consideration that the mean ash-free dry weight (AFDW) of the body tissue of the specimens varied greatly between field and laboratory experiments, data were analysed separately for each experiment type. Estimation of Capture Efficiency (CE) and Clearance Rates (CR) The particle count and particle size distribution were determined using a PAMAS laser particle counter (model S4031 GO) containing an HCB-LD-50/50 light-scattering sensor. Particles were counted using the light blocking principle: when the cell is 100% illuminated by laser light, the particle blocks light, and casts a shadow on a photodetector. The particle size is determined by the amount of blocked light. Control incubators were also analysed to identify any reduction in particles due to bottom deposition and adjust accordingly the incubators with bivalve specimens by subtracting this decline in particle concentration. Furthermore, these controls were also used to characterize the natural seston present in each experiment. For all experiments, the PAMAS used an initial volume of 15 mL of each water sample, in which particles were counted in 1 mL of seawater. The PAMAS was initially set to determine 33 classes of sizes ranging from 1 to 150 μm to provide a very differentiated analysis of the sizes. However, the actual range monitored to determine both CEs and CRs was set between 4 and 14 μm due to the following: 1) the highest retention efficiency values for bivalves are attained for particles with 4 μm and above (Møhlenberg and Riisgård 1978; Cranford et al. 2016); and 2) most of the particles found in the seawater were within this size range. Otherwise, if the number of particles of a specific size is reduced, Figure 4. A diagram of an oyster with a Hall element sensor attached to a valve and a small magnet attached to the opposite valve B hall element sensor glued to the valve of a R. philippinarum specimen. 132 NeoBiota 102: 125–150 (2025), DOI: 10.3897/neobiota.102.148326 Sara Cabral et al.: Estimating capture efficiencies and clearance rates of three bivalve species incorrect interpretations may be drawn, since the particles might not be captured due to its size or simply because their availability in the natural seawater is too low to be effectively captured. The estimation of CE and CR for all species and both field and laboratory experiments followed the static method described by Cranford et al. (2016). A suspension feeder pumping at a constant rate and without water renewal will remove particles at a rate that declines exponentially (Coughlan 1969). Consequently, the PAMAS can be used to measure the slope of the natural logarithm of particle concentration decline over time (λ), taking into consideration that for all measurements, this decline should be linear over time in order to ensure the constant pumping rate assumption (Cranford et al. 2016). The CE was estimated for each individual and each particle size according to the following equation adapted from Cranford et al. (2016): where λsample, size represents the slope of the linear regression between the particle concentration (transformed in natural logarithm) and the elapsed time for a given particle size class in the sample; λcontrol, size is the slope of the linear regression between the particle concentration (transformed in natural logarithm) and the elapsed time for the same particle size class in the control (all controls were averaged for each sampling date); and λsample, average is the average of the λsample, size across all particle size classes of each individual. This standardization by the average value across all particle size classes allowed to minimize the effect of potential measurement errors from a single particle size class count. The resulting CE values range from 0 to 1, representing particles sizes that are not captured and captured at maximum efficiency, respectively. Nevertheless, averaging particles across size classes, rather than using the maximum, may lead to CE values exceeding 1. Suppl. material 2 depicts an example of how some components of the CE formula were estimated for each specimen. Subsequently, the mean CE for each particle size was estimated for each species, by averaging all the individuals CE. See Table 3 for the sample sizes per species and type of experiment included in the CE estimation. CR was estimated for each specimen, according to the equation adapted from Casas et al. (2018): Table 2. Mean (± SD) shell length (mm), ash-free dry body tissue weight (AFDW; g) and condition index (%) for both field and laboratory experiments of three bivalve species (C. edule, M. angulata and R. philippinarum). Experiment Species Mean shell length (mm) Mean AFDW of body (g) Mean Condition Index (%) Field C. edule 31.30 ± 1.39 0.55 ± 0.24 9.61 ± 4.92 M. angulata 85.17 ± 6.91 3.82 ± 1.55 7.60 ± 2.58 R. philippinarum 43.57 ± 2.07 1.51 ± 0.79 10.43 ± 4.25 Laboratory C. edule 29.50 ± 1.66 0.18 ± 0.02 4.12 ± 0.61 M. angulata 91.11 ± 4.93 1.41 ± 0.20 3.16 ± 0.70 R. philippinarum 47.09 ± 1.79 0.50 ± 0.06 4.59 ± 0.51 133 NeoBiota 102: 125–150 (2025), DOI: 10.3897/neobiota.102.148326 Sara Cabral et al.: Estimating capture efficiencies and clearance rates of three bivalve species where λsample represents the slope of the linear regression between the particle concentration (transformed in natural logarithm) and the elapsed time in the sample; λcontrol is the slope of the linear regression between the particle concentration (transformed in natural logarithm) and the elapsed time in the control (all controls were averaged for each sampling date); and V is the volume of seawater in each incubator (1.2 L). With the aim of reducing the weight of potential outliers on the estimates of the CR, the approach of Casas et al. (2018) was followed. This method integrates the particle concentration values recorded throughout the multiple sampling times, instead of using only data from the beginning and the end of the experiments. Additionally, only linear regressions with r2 > 0.90 were selected for the subsequent analyses. Suppl. material 3 depicts an example of how the CR was estimated for each specimen and Table 3 describes the number of bivalves per species and type of experiment included in the CR estimation. Lastly, since the studied bivalve species have discrepant sizes, CR absolute values are only representative when comparing results between individuals of the same species. Hence, following the CR estimations, these values were divided by the ash-free dry weight (AFDW) of the body tissue of the respective specimen to be comparable between the different species. Statistical analysis The Shapiro-Wilk’s and Levene’s tests were used to assess the normality of residuals and the homogeneity of variances, respectively. To test for differences in CE values between species and across particle size classes a Permutational Analysis of Variance (PERMANOVA; Anderson et al. 2008) was conducted for each experiment type (field and laboratory) with Bray-Curtis similarity matrices and 9999 permutations. This analysis was performed on PRIMER® v7 (Clarke and Gorley 2015), using a two-way fixed-effect crossed design (Factors: species; particle size). Afterwards, Kruskal–Wallis tests were used to compare the mean differences in CR between species, both for field and laboratory results. All comparisons were performed for CR (L h-1 ind-1) and CR (L h-1 g-1 AFDW). Dunn’s post hoc tests were used for pairwise comparisons between groups. Apart from the PERMANOVA, all the analyses were performed in R version 4.4.1 (RStudio version 2024.12.0). Results In spite of generally similar mean shell length across experiments (within each species), the mean AFDW of body tissue was always higher in field experiments, regardless of the species (Table 2). Considering this outcome, no direct comparisons between field and laboratory experiments were performed in this paper. At the onset of the experiments, the average concentration of particles present in the seawater was estimated for each particle size (Fig. 5). Particle concentrations in the field were on average 6 to 11 times higher than in the laboratory. The maximum average number of particles per ml were recorded for the size classes of 4–5 μm (12805.12 particles ml-1 and 2202.87 particles ml-1 for field and laboratory, respectively), whereas the minimum was observed for particles within the size class of 13–14 μm (1940.39 particles ml-1 and 190.69 particles ml-1 for field and laboratory, respectively). 140 NeoBiota 102: 125–150 (2025), DOI: 10.3897/neobiota.102.148326 Sara Cabral et al.: Estimating capture efficiencies and clearance rates of three bivalve species Consequently, governing entities and regulators should be cautious when establishing new areas for bivalve aquaculture production and ensure that data on CE and CR of both native and NIS are taken into account since there is an overlap of their ecological niches. Besides, considering that the individuals used in these experiments were representative of the exploitable populations (since their shell lengths were all above the minimum marketing size), this information should also be considered when managing NIS and native species’ stocks. Conclusion In conclusion, our results indicate that C. edule, M. angulata and R. philippinarum have similar capture efficiencies for the size classes of particles measured in this study (ranging from 4–14 μm). Moreover, R. philippinarum did not present a competitive advantage in clearing suspended particles when compared with the natives M. angulata and C. edule, with the latter revealing to be the most efficient species on a tissue weight basis. Therefore, the invasion success of R. philippinarum may be explained by other biological characteristics or advantages in ecological interactions such as reproductive rates, space competition or overlapping niche spaces. It is highly recommended that future studies focus on these aspects in order to provide the full extent of the potential impacts of NIS on native species. Notwithstanding, NIS may still jeopardize the entire ecosystem, by exerting an additional pressure on its carrying capacity. Finally, this study provides an important baseline contribution for the management of NIS by determining capture efficiencies and clearance rates estimated exclusively with natural seawater depicting a more accurate representation of the processes that occur in the environment. Acknowledgements A special thank you to Afonso Ferreira, Filipa Afonso, Inês Afonso, Inês Teodoro, Isabel Cardoso, Margarida Raposo, Marion Cottet, Pedro Coelho, Pedro Gomes, Pedro M. Oliveira, Teresa Camelo and Thomas Goulding, for assisting us during the experiments. Thanks are also due to Ramón Filgueira for the valuable suggestions. This study benefited from discussions within the Working Group on Introductions and Transfers of Marine Organisms (WGITMO) under the International Council for the Exploration of the Seas (ICES). Finally, the authors gratefully acknowledge the suggestions of the reviewers that improved the manuscript. Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. Funding This work was supported by the Portuguese Foundation for Science and Technology (FCT), through the grants SFRH/BD/145746/2019 (https://doi.org/10.54499/SFRH/BD/145746/2019), 2020.06325. BD (https://doi.org/10.54499/2020.06325.BD), UI/BD/151399/2021, CEECIND/00095/2017, 141 NeoBiota 102: 125–150 (2025), DOI: 10.3897/neobiota.102.148326 Sara Cabral et al.: Estimating capture efficiencies and clearance rates of three bivalve species 2020.01797.CEECIND, UIDB/04292/2020 (https://doi.org/10.54499/UIDB/04292/2020), and the Associate Laboratory ARNET (https://doi.org/10.54499/LA/P/0069/2020). Additionally, this work was also funded by the MAR2020 projects AQUASADO (MAR-02.01.01-FEAMP-0051) and NIPOGES (MAR-01.03.02-FEAMP-0013) and by the European Union’s Interreg Atlantic project COCKLES (EAPA_458/2016). Author contributions Conceptualization: PC, ACB. Formal analysis: SC, ACB. Funding acquisition: PC, ACB. Investigation: JH, PC, SC, FC, JPCC, JMFB, LAC, ACB. Methodology: JMFB, JPCC, FC, SC, JH, LAC, ACB. Project administration: ACB. Resources: JMFB, LAC, PC. Supervision: ACB, PC. Writing - original draft: SC. Writing - review and editing: ACB, JMFB, FC, PC, JH, JPCC, LAC. 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Aquaculture (Amsterdam, Netherlands) 479: 824–828. https://doi.org/10.1016/j.aquaculture.2017.07.030 Zaiko A, Daunys D (2015) Invasive ecosystem engineers and biotic indices: Giving a wrong impression of water quality improvement? Ecological Indicators 52: 292–299. https://doi.org/10.1016/j. ecolind.2014.12.023 Zorita I, Solaun O, Borja A, Franco J, Muxika I, Pascual M (2013) Spatial distribution and temporal trends of soft-bottom marine benthic alien species collected during the period 1989-2008 in the Nervión estuary (southeastern Bay of Biscay). Journal of Sea Research 83: 104–110. https://doi. org/10.1016/j.seares.2013.04.009 Supplementary material 1 Relationship between the Suspended Particulate Matter (SPM) and the turbidity for the field experiments site (an aquaculture farm located in the Sado estuary, Setúbal, Portugal) Authors: Sara Cabral, Frederico Carvalho, Joana P. C. Cruz, Joshua Heumüller, Jose M. F. Babarro, Luc A. Comeau, Paula Chainho, Ana C. Brito Data type: pdf Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/neobiota.102.148326.suppl1 Supplementary material 2 Representation for a single incubator with a bivalve actively filtering Authors: Sara Cabral, Frederico Carvalho, Joana P. C. Cruz, Joshua Heumüller, Jose M. F. Babarro, Luc A. Comeau, Paula Chainho, Ana C. Brito Data type: pdf Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/neobiota.102.148326.suppl2 149 NeoBiota 102: 125–150 (2025), DOI: 10.3897/neobiota.102.148326 Sara Cabral et al.: Estimating capture efficiencies and clearance rates of three bivalve species Supplementary material 3 Representation for both a sample with bivalve (green) and the control (grey) of (A) the decline in the number of particles over the experiment time and (B) the linear regression between the particle concentration (transformed in natural logarithm) and the elapsed time Authors: Sara Cabral, Frederico Carvalho, Joana P. C. Cruz, Joshua Heumüller, Jose M. F. Babarro, Luc A. Comeau, Paula Chainho, Ana C. Brito Data type: pdf Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/neobiota.102.148326.suppl3 Supplementary material 4 PERMANOVA analysis results on capture efficiency (CE) of three bivalve species (C. edule, M. angulata and R. philippinarum) Authors: Sara Cabral, Frederico Carvalho, Joana P. C. Cruz, Joshua Heumüller, Jose M. F. Babarro, Luc A. Comeau, Paula Chainho, Ana C. Brito Data type: pdf Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/neobiota.102.148326.suppl4 Supplementary material 5 PERMANOVA Pair-wise test to detect differences on CE between particle sizes for field experiments of three bivalve species (C. edule, M. angulata and R. philippinarum) Authors: Sara Cabral, Frederico Carvalho, Joana P. C. Cruz, Joshua Heumüller, Jose M. F. Babarro, Luc A. Comeau, Paula Chainho, Ana C. Brito Data type: pdf Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/neobiota.102.148326.suppl5