Diversity and Distributions, (Diversity Distrib.) (2015) 21, 1075–1086

BIODIVERSITY What guides invasion success? Ecological RESEARCH correlates of arrival, establishment and spread of Red Sea bivalves in the Mediterranean Sea Rafał Nawrot1*, Devapriya Chattopadhyay2 and Martin Zuschin1

1Department of Palaeontology, University of ABSTRACT Vienna, Althanstrasse 14, Vienna 1090, Aim The opening of the Suez Canal in 1869 re-established the direct link Austria, 2Department of Earth Sciences, between long-separated biogeographic realms, allowing hundreds of marine Indian Institute of Science Education and Research (IISER) Kolkata, Mohanpur, species to spread from the Red Sea to the Mediterranean. We use marine bival- WB-741246, India ves to relate species-level attributes to successful transition through successive stages of the invasion process.

Location Mediterranean and Red Sea.

Methods We compiled data on taxonomic composition, body size, life habit A Journal of Conservation Biogeography and geographic distribution of the Red Sea bivalve fauna from published litera- ture, museum collections and our own field surveys. Using multimodel infer- ence, we examined selectivity of the Lessepsian invasion and identify traits that distinguish successful species at three major stages of invasion: arrival, estab- lishment and spread.

Results The upper limit of bathymetric range and occurrence outside the trop- ical zone in other regions are the strongest predictors of successful transition through the Suez Canal. Establishment in the Mediterranean is positively corre- lated with earlier arrival and association with hard-bottom habitats. Preference for hard substrates together with large body size is the primary factor distin- guishing invasive aliens representing a significant threat to recipient ecosystems from other established species.

Main conclusions The relative strength of abiotic and biotic filters changes along the course of the invasion: environmental affinity and climate match con- strain the pool of potential invaders, while the establishment in the new region and invasive status depend on the habitat preferences and life history traits of aliens, affecting their interactions with resident species. Our results together with previous studies suggest that the eastern Mediterranean rocky shores are more susceptible to the establishment of Lessepsian species, many of which may induce strong pressure on recipient communities as ecosystems engineers and competitors of native species. *Correspondence: Rafał Nawrot, Department

and Distributions Keywords of Palaeontology, University of Vienna, Althanstrasse 14, 1090 Vienna, Austria. Alien species, biological invasions, biotic interchange, invasive species, Lessep- E-mail: [email protected] sian migration, marine invasions, Mollusca.

into a global hotspot of marine bioinvasions (Rilov & Galil, INTRODUCTION 2009; Por, 2010; Edelist et al., 2013). This so-called Lessep- The opening of the Suez Canal in 1869 re-established the sian or Erythrean invasion (Por, 1971; Rilov & Galil, 2009; direct link between biogeographic realms separated since the Safriel, 2013) has led to profound changes in marine com-

Diversity Middle Miocene and allowed hundreds of tropical species to munities of the eastern Mediterranean, which are now often spread from the Red Sea to the Mediterranean, turning it dominated by Red Sea taxa (Galil, 2007; Edelist et al., 2013;

DOI: 10.1111/ddi.12348 ª 2015 John Wiley & Sons Ltd http://wileyonlinelibrary.com/journal/ddi 1075 R. Nawrot et al.

Rilov, 2013). In contrast to other human-mediated invasions, We use information on taxonomic composition, bathy- the Lessepsian invasion encompasses whole complexes of metric and geographic distribution, body size and life habits species that are sympatric in their native range (Por, 2010) of the Red Sea bivalve fauna to quantitatively examine the and maintain high levels of gene flow between their newly biological selectivity of the consecutive stages of the Lessep- established and parent populations (Bernardi et al., 2010). sian invasion. We attempt to determine which characteristics Although the introduction of some Red Sea species is of the Red Sea species predispose them to enter the Mediter- undoubtedly facilitated by maritime transport (Shefer et al., ranean Sea and thrive there, and how particular traits of suc- 2004), for most of them, the natural dispersal through the cessful immigrants affect their post-establishment spread and Suez Canal and along the Mediterranean coasts remains the impact on native communities. primary means of range expansion (Gofas & Zenetos, 2003; Ben Rais Lasram et al., 2008; Tzomos et al., 2012). METHODS The Lessepsian invasion can be regarded as a unique biogeographic experiment (Por, 2010) that shares many simi- Data collection larities with large-scale biotic interchanges in the geological past (Vermeij, 2005). It can thus serve as a model system for We compiled a database on body size and ecological charac- comparing marine invasions in natural and anthropogenical- teristics of 394 bivalve species occurring at continental shelf ly altered ecosystems. Moreover, identification of factors depths (<200 m) in the Red Sea (see Appendix S1 in Sup- contributing to the invasion process may help to alleviate the porting Information). Data were collected from primary lit- threats it poses to native biodiversity and human economy erature, major monographs and museum collections (see (Streftaris & Zenetos, 2006; Galil, 2007; Katsanevakis et al., Appendix S2). Substantial information was derived from 2014). Oliver (1992), Dekker & Orlin (2000), Huber (2010) and Molluscs, with over 200 alien species, are currently the from our own field studies (Zuschin & Oliver, 2003, 2005; most species-rich animal phylum among non-native taxa in Zuschin & Ebner, 2015). Taxonomic nomenclature was pri- the Mediterranean Sea (Zenetos et al., 2012). In this study, marily based on the World Register of Marine Species we focus on marine bivalves, which are routinely used as a (WoRMS Editorial Board, 2014). Teredinid bivalves (12 spe- model clade in macroecological and macroevolutionary cies) were excluded from the analyses. This group is poorly studies (e.g. Roy et al., 2000; Vermeij et al., 2008; Berke studied in the Red Sea and may include cryptogenic species. et al., 2013), and encompass some of the most notorious Species of Red Sea origin that are non-indigenous to the invasive species (Streftaris & Zenetos, 2006; Katsanevakis Mediterranean fauna were identified based on the inventories et al., 2014). Taxonomic identity and distribution of Lessep- of Gofas & Zenetos (2003) and Zenetos et al. (2010, 2012), sian bivalves in the Mediterranean Sea is well documented updated with the most recent records (Appendix S2). We (Gofas & Zenetos, 2003; Zenetos et al., 2010; Tzomos et al., did not attempt to differentiate between species spreading 2012), and our knowledge about the ecology and biology of through the Suez Canal by natural means of dispersal, that is selected species is continuously expanding (e.g. Rilov et al., ‘true’ Lessepsian migrants sensu Por (1971), and those pas- 2004; Zurel et al., 2012). Nonetheless, the strong focus of sively introduced by ships. For many species, both vectors previous studies on just a few of the most conspicuous may operate simultaneously, especially during their second- immigrants (reviewed by Safriel, 2013) is hindering our ary spread within the Mediterranean Sea (e.g. Shefer et al., ability to understand the dynamics of the entire invasion 2004; Galil, 2008). We use the term Lessepsian species in a process. We therefore address the biological correlates of the broad sense to denote all Red Sea species that have pene- Lessepsian invasion in the whole pool of potential bivalve trated through the Suez Canal. The three cosmopolitan spe- immigrants. cies Perna perna (Linnaeus, 1758), Irus irus (Linnaeus, 1758) The invasion is a multistage process (Blackburn et al., and Martesia striata (Linnaeus, 1758) were considered native 2011) in which the effect of any particular trait on the spe- to both regions and excluded from the source pool of cies’ success may change depending on the stage at which it invaders. is evaluated (Cassey et al., 2004; Jeschke & Strayer, 2006). The information on the acclimatization status of Lessep- Accordingly, proper delineation of the species pool and an sian species was taken from Zenetos et al. (2010, 2012) rep- appropriate control group at each successive stage is a prere- resenting the most comprehensive overview of marine alien quisite for the identification of factors promoting or imped- species in the Mediterranean Sea currently available (see also ing success of alien species (Cassey et al., 2004; Miller & Appendix S2 for supplementary sources). Their classification Ruiz, 2009). Three stages of the Lessepsian invasion can be of alien species can be easily incorporated into the multistage distinguished following the modified framework of Black- framework for biological invasions of Blackburn et al. burn et al. (2011): (1) arrival of a Red Sea species to the (2011). We considered all Red Sea bivalves with confirmed Mediterranean Sea, (2) establishment of a viable alien popu- records in the Mediterranean Sea as alien (i.e., Lessepsian) lation, and (3) population growth and spread, imposing a species that successfully completed the arrival stage. Depend- considerable impact on native communities (see also discus- ing on the successful establishment of free-living and self- sion in Safriel, 2013). sustaining populations in the new region, they can be

1076 Diversity and Distributions, 21, 1075–1086, ª 2015 John Wiley & Sons Ltd Invasion success in marine bivalves divided into casual (non-established) and established aliens. widely used in previous studies on large-scale body-size pat- Species recorded at least twice from different localities and at terns in marine bivalves (e.g. Roy et al., 2000; Berke et al., different times were also included in the latter group (Zene- 2013). tos et al., 2010). Those among established aliens that were identified as invasive by Zenetos et al. (2010) were regarded Life habits as the ones that reached the spread stage. Such species not only are able to disperse away from the place of initial intro- Species were assigned to soft or hard substrate and classified duction and reproduce in new areas, but also have noticeable into five trophic guilds (suspension feeders, deposit feeders, ecological or economic impacts in the recipient ecosystems chemoautotrophs, carnivores and zooxanthellae bearers) and (Streftaris & Zenetos, 2006; Zenetos et al., 2010). In sum- five life habits (infaunal, endobysate, epifaunal, boring and mary, from the source pool of 391 Red Sea species, 52 were commensal). Taxa inhabiting mixed bottoms were included identified as Lessepsian, including 22 casual, 23 established in the hard-substrate group; the alternative assignment of non-invasive and 7 invasive species. We analysed their char- such forms yielded qualitatively similar results. acteristics separately for each stage by comparing successful and failed immigrants within an appropriate species pool (all Year of introduction Red Sea species, all alien species or only established aliens). Information on life history and reproductive traits is lack- Species that entered a new region earlier have a greater ing for most marine invertebrates (Tyler et al., 2012). We chance to become established and eventually invasive at the therefore restricted our analyses to a suite of traits that time of the most recent survey. We used the year of the potentially facilitate invasion (e.g. Roy et al., 2001; Miller first Mediterranean record taken from Gofas & Zenetos et al., 2007; Ben Rais Lasram et al., 2008) and can be easily (2003) and Tzomos et al. (2012) as a proxy for the arrival obtained for a large pool of Red Sea species: bathymetric and of alien species and considered it as a correlate of estab- geographic distribution, body size, prefered substrate type lishment and spread. Whenever available, the date of col- and trophic guild (see Appendix S2). lection of the actual specimens was preferred over the publication year. Because the first detection of a species can post-date its arrival and establishment by several years Minimum depth or even decades, such data provide only the minimum, Shallow-water dwellers are more likely to pass through the and thus very conservative, estimate of the actual timing of barrier of the Suez Canal and are also expected to have the introduction. greater potential to establish in a new region (Harley et al., 2003). The minimum depth of occurrence of species’ post- Data analyses larval stages was measured in metres. Intertidal species were assigned a value of zero. When no quantitative estimate To assess the relative role of species-level traits in determin- could be obtained (6.8% of the species), the minimum depth ing the invasion success, we used generalized linear models was approximated by the depth of the shallowest environ- with binomial error structure and logit link function (i.e. mental zone occupied by a species. Four such zones were logistic regression). The outcome of transition through each distinguished: intertidal (0–1 m depth), the shallow sublit- stage was coded as a binary response. As most of the life toral (1–30 m), the deep sublittoral (30–200 m) and the habit categories were represented by only few species, we deep sea (>200 m). simplified this classification by dividing all species into two feeding guilds (suspension and non-suspension feeding) and two substrate groups (hard- and soft-substrate-associated Extratropical occurrence species). Additional analyses were performed using more Occurrence outside the tropical zone can serve as a coarse- detailed classification of life habits and employing Firth’s scale indicator of the climate match between the native range bias-reduced logistic regression to account for statistical sep- of Red Sea species and the highly seasonal, warm-temperate aration (Heinze & Schemper, 2002) (see Appendix S3). To Mediterranean Sea (Ben Rais Lasram et al., 2008). Species facilitate comparison of parameter estimates for binary and are identified as extratropical if their geographic range continuous variables, the latter were standardized to a mean extends beyond the tropical marine realms as defined in of 0 and standard deviation of 0.5 (Gelman, 2008). Spalding et al. (2007). We employed a sample size-corrected form of the Akaike information criterion (AICc) to rank alternative models based on the trade-off between the model fit and its com- Body size plexity (Burnham & Anderson, 2002; Grueber et al., 2011).

The log2-transformed geometric mean of the shell length and For each invasion stage, we generated a set of models with height of the largest specimen reported from the Red Sea all possible combinations of predictors and scored them was used as a proxy for body size of a species (see Chatto- according to their relative support as measured by AICc. To padhyay et al., 2014 for details). This measure has been account for the uncertainty in model selection, we performed

Diversity and Distributions, 21, 1075–1086, ª 2015 John Wiley & Sons Ltd 1077 R. Nawrot et al. model averaging, applying a cut-off criterion of ΔAICc ≤ 2 establishment and spread stage, by repeatedly drawing 30 to define the confidence set of models. To identify the fac- species from the pool of 52 aliens and seven species from the tors that have the strongest effect on invasion success, the pool of 30 established aliens, respectively. All statistical analy- parameter estimates were calculated by averaging over all ses were performed in R 3.1 (R Core Team, 2014). selected models, that is using the zero method (Burnham & Anderson, 2002; Grueber et al., 2011). The relative impor- RESULTS tance of each predictor was computed as the sum of AICc weights of top-scoring models that included a given variable, Arrival with the value of 1 indicating that the predictor was present in all of them. The area under the receiver operating charac- Compared to 339 species that have not succeeded at crossing teristic curve (AUC) was used as a measure of predictive the Suez Canal, the 52 Lessepsian species tended to be larger, power of the models (Fielding & Bell, 1997). The value of occurred at shallower depths and extended into higher lati- the AUC varies between 0.5 and 1.0 and, in the context of tudes in their native range (Fig. 1). Although the share of our study, can be interpreted as the probability that when hard-bottom and epifaunal forms was also higher among successful and unsuccessful invaders are selected at random, them (Fig. 1 and Fig. S1 in Appendix S3), this difference was the former will have greater probability of success estimated not greater than expected by a random sampling from the by a model. source species pool (Fig. 2a and Fig. S2a in Appendix S3). In To aid the interpretation of the model-selection results, we turn, suspension feeders were significantly over-represented constructed a randomization-based null model (cf. Blackburn among successful species (Fig. 2b). None of the commensal, & Cassey, 2007) predicting the number of species from each carnivorous or zooxanthellae-bearing bivalves entered the life habit expected to invade assuming the lack of ecological Mediterranean Sea; this apparent selectivity may be an arte- selectivity. From the original source pool of 391 Red Sea spe- fact of the small number of species in these groups (16, six cies, we sampled at random and without replacement as and three species, respectively), as suggested by the null many species as there were aliens (i.e. 52 species) and model (Fig. S2 in Appendix S3). The minimum water depth counted the number of species belonging to each feeding and extratropical status were the most important predictors guild and substrate group. This simulation was repeated of the transition through the first invasion stage (Fig. 3a, 10,000 times to obtain the median and 95% confidence Table 1). Feeding mode emerged as a significant correlate of intervals for the expected number of species. A similar proce- the invasion success among simplified life habits, but not dure was used to determine the ecological selectivity of the when more detailed guild subdivision was used (see

Body size Minimum depth Year of introduction Arrival Establishment Spread Arrival Establishment Spread Establishment Spread 8 (size) 2 Year Log Depth [m] 246 1880 1920 1960 2000 100.0 10.0 1.0 0.1 N-AlnAln Cas Est N-Inv Inv N-AlnAln Cas Est N-Inv Inv Cas Est N-Inv Inv

Substrate type Feeding mode Extratropical status Arrival Establishment Spread Arrival Establishment Spread Arrival Establishment Spread % of species % of species % of species 0.0 0.2 0.4 0.6 0.8 1.0 0.0 0.2N-Aln 0.4 0.6Aln 0.8 1.0 Cas Est N-Inv Inv N-AlnAln Cas Est N-Inv Inv 0.0 0.2 N-Aln 0.4 0.6Aln 0.8 1.0 Cas Est N-Inv Inv Soft Hard Suspension Non-suspension Absent Present

Figure 1 Comparison of the ecological characteristics of Red Sea bivalve species that succeeded and failed at different stages of the Lessepsian invasion. Arrival stage: non-alien (N-Aln; n = 339) and alien Red Sea species (Aln; n = 52); establishment stage: casual (Cas; n = 22) and established aliens (Est; n = 30); spread stage: non-invasive (N-Inv; n = 23) and invasive established aliens (Inv; n = 7).

1078 Diversity and Distributions, 21, 1075–1086, ª 2015 John Wiley & Sons Ltd Invasion success in marine bivalves

(a) Arrival Establishment Spread Arrival 50 1 (a) Relative Expected importance AUC = 0.73 Observed - 40 0.8 Intercept

Minimum depth 1 30 - 0.6 - Extratropical 1 20 - - 0.4 % of species

Number of species - Suspension feeding 1 10 0.2

- Hard substrate 0.44 0 P = 0.21 P = 0.02 P = 0.09 0 –8 –4 0 4 8 NofInv NofAln NofEst %ofInv %ofAln %ofEst Establishment Relative (b) importance AUC = 0.83 (b) Arrival Establishment Spread - 50 - 1 Intercept - Introduction year 1 40 - - 0.8

Hard substrate 1 30 0.6 - Extratropical 0.36 20 0.4 % of species Suspension feeding 0.35 Number of species 10 0.2 - Size 0.12 0 P = 0.01 P =0.1 P = 0.77 0 –8 –4 0 4 8

Spread Relative NofInv %ofInv NofAln NofEst %ofAln %ofEst (c) importance AUC = 0.85

Figure 2 Number and proportion of (a) hard-substrate and (b) Intercept - suspension-feeding species expected to succeed at each stage of the Lessepsian invasion under the null model of random Hard substrate 0.8 sampling from the source species pool. The median number of species (Æ95% confidence intervals) expected to represent a Size 0.79 given life habit was obtained from 10,000 resampling iterations. The P-values (permutation test) correspond to the proportion of Extratropical 0.19 randomizations that produced a value equal or more extreme than the observed number of species. Aln, alien species; Est, established aliens; Inv, invasive aliens. –8 –4 0 4 8 Effect size (logit scale)

Appendix S3: Fig. S3, Table S1). One of the top models con- Figure 3 Results of model averaging based on the best- tained the substrate, but model-averaged 95% confidence supported subset of multivariate logistic regression models relating species characteristics of Red Sea bivalves to success at intervals for this variable overlapped zero (Fig. 3a). the (a) arrival, (b) establishment and (c) spread stage of the Lessepsian invasion. Average effect sizes with corresponding Establishment standard errors (thick bars) and 95% confidence intervals are ordered vertically according to the relative importance of the The set of established aliens was characterized by a larger predictors. median size, greater proportion of extratropical species and significant over-representation of hard-substrate forms models for this stage, the year of introduction and preferred (Figs 1 and 2). The first Mediterranean records of established substrate type were consistently included in all top models aliens were much older compared to those of the majority of and had a significant effect on the probability of establish- casual species. Despite a rather large set of best-supported ment. Extratropical status, feeding mode and body size had

Diversity and Distributions, 21, 1075–1086, ª 2015 John Wiley & Sons Ltd 1079 R. Nawrot et al.

Table 1 Highest ranked logistic models (ΔAICc ≤ 2) investigating ecological correlates of the successful transition of Red Sea bivalves through the consecutive stages of the Lessepsian invasion.

Stage Model AICc ΔAICc wAICc R2

Arrival EX + MD + FD 283.44 0.00 0.31 0.141 EX + MD + FD + SB 284.96 0.52 0.24 0.148 Establishment YI + SB 58.50 0.00 0.17 0.409 YI + SB + EX 59.07 0.58 0.13 0.440 YI + SB + FD 59.12 0.63 0.13 0.439 YI + SB + EX + FD 60.19 1.69 0.08 0.463 YI + SB + SZ 60.41 1.92 0.07 0.417 Spread ST + SZ 32.38 0.00 0.11 0.320 ST 33.57 1.19 0.06 0.165 SZ 33.67 1.30 0.06 0.160 ST + SZ + EX 33.80 1.42 0.06 0.368

ΔAICc, difference in AICc from the best-supported model; wAICc, model weights; R2, Nagelkerke’s R2. Predictors: EX, extratropical occurrence; FD, feeding mode; YI, year of introduction; MD, minimum depth; SB, substrate; SZ, size. only minor effects (Fig. 3b). Similar results were obtained Arrival using the alternative classification of life habits (see Appendix S3: Figs S2 and S3, Table S1). Factors related to the environmental and geographic distri- bution of a species were the most important correlates of the Lessepsian status among Red Sea bivalves. The Suez Canal is Spread a shallow waterway, with depths initially not exceeding 8 m Only seven species, representing <2% of the original Red Sea and only recently increased to 24 m after several phases of species pool, became invasive aliens. The number of epifau- progressive enlargement (Rilov & Galil, 2009; Suez Canal nal forms among them (Figs S1 and S2a in Appendix 3) and Authority, 2015). Consequently, the minimum depth of spe- their median size (Fig. 1) were significantly larger than cies’ occurrence was identified as the strongest determinant expected by a random sampling from the pool of established of the successful passage. A similar over-representation of species (P = 0.041 and 0.001, respectively; 10,000 iterations). shallow-water species is evident among alien fishes (Rilov & Substrate type and body size were the strongest predictors of Galil, 2009). This bathymetric filter acting already at the arri- the invasive status (Fig. 3c and Table 1), although only the val stage limits the pool of invaders exclusively to species latter trait received similar support in analyses based on the inhabiting highly variable intertidal and upper sublittoral alternative guild classification (see Appendix 3: Fig. S3 and zones. Their greater environmental tolerance may facilitate Table S1). The 95% confidence intervals for the model-aver- subsequent establishment and range expansion (Harley et al., aged coefficient estimates, however, were visibly large and 2003). included zero. This lack of precision likely reflects the very Occurrence outside the tropical zone in other regions, small number of invasive species. indicative of a broad thermal tolerance of a species, was the second significant predictor of the completion of the arrival stage. The increase in the proportion of extratropical species DISCUSSION is also evident at later invasion stages. Low winter seawater Our study is apparently the first to search for ecological cor- temperatures in the eastern Mediterranean are one of the relates of successful transition across a complete series of main factors limiting the survival and reproductive success steps in the invasion process in a large assemblage of marine of Red Sea immigrants (Rilov & Galil, 2009; Zurel et al., invertebrates. Some of the analysed variables show consistent 2012), although high summer extremes may be detrimental patterns: median body size, proportion of hard-substrate, as well (Belmaker et al., 2013). Similarly, fish species experi- suspension-feeding and extratropical species increased at encing a broader range of temperatures in their native range each stage (Fig. 1). However, the relative importance and are more likely to enter the Mediterranean Sea (Belmaker strength of the effect of any particular predictor differed et al., 2013) and have greater rates of subsequent dispersal between stages, pointing to shifts in the nature of filters, and (Ben Rais Lasram et al., 2008). The importance of climate consequently biological mechanisms, operating at each phase similarity between the native range and the locality of intro- of the process. A similar idiosyncratic pattern of invasion has duction has also been documented for terrestrial vertebrates been also observed among terrestrial and aquatic vertebrates (e.g. Duncan et al., 2001; Rago et al., 2012). On the other (Cassey et al., 2004; Marchetti et al., 2004; Jeschke & Strayer, hand, recent evidence for the widespread climatic niche 2006). expansion among Lessepsian fishes (Parravicini et al., 2015)

1080 Diversity and Distributions, 21, 1075–1086, ª 2015 John Wiley & Sons Ltd Invasion success in marine bivalves highlights the uncertainty involved in predicting invasion be more prone to invasion and/or are exposed to greater risk based on the climatic factors alone. propagule pressure. Recent surveys of benthic fauna on rocky The strong bathymetric and climatic constraints on the reefs along the Israeli cost have revealed the strong domina- pool of potential invaders affect the distribution of other tion of bivalve, gastropod and fish assemblages by Indo-Paci- biological attributes among immigrants. The relative paucity fic species (Rilov, 2013). Analogous differences in invasion of non-suspension-feeding aliens can be largely explained by levels between soft and hard substrates have been detected non-random patterns in the environmental distribution of both in modern marine introductions in California (Wasson different trophic guilds: mostly deep-water occurrence of car- et al., 2005) and in the Pliocene trans-Arctic interchange nivorous bivalves, and predominately tropical affinity of between northern Pacific and Atlantic biotas (Vermeij, chemosymbiont-bearing lucinids, deposit-feeding tellinids 1991). and zooxanthellae-bearing tridacnids. Relatively greater availability of suitable hard bottoms – The duration of the planktonic larval stage and, conse- both natural and artificial ones – in shallow coastal waters quently, larval-development mode may affect the passage farther away from the Canal may be responsible for the pre- through the Suez Canal (Gofas & Zenetos, 2003; Safriel, dominance of epifaunal forms among established aliens. On 2013), but information on these traits is not available for the the other hand, only one fourth of approximately 180 species great majority of the Red Sea bivalves. While larval plankto- of bivalves native to the south-eastern Mediterranean Sea is trophy, typically associated with prolonged larval stage, may associated with hard substrates (Barash & Danin, 1992 and facilitate invasion, the conditions in the Suez Canal can be our unpublished data). The smaller pool of incumbent spe- deleterious for plankton-feeding larvae (Por, 1971). We cies in Levantine rocky habitats may indicate lower biotic expect therefore that the larval-development mode is not a resistance (Olyarnik et al., 2009), which – combined with a key factor controlling this invasion, as already suggested for wealth of artificial hard substrates providing space for alien Lessepsian gastropods (Gofas & Zenetos, 2003), which are recruitment (Glasby et al., 2007) – should greatly facilitate dominated by non-planktotrophic species (Chemello & the establishment of immigrants. At the same time, the rates Oliverio, 2001). Unfortunately, thorough tests of this of introduction and establishment of cemented and bysally- hypothesis have to wait until more life history data are col- attached epifauna are likely accelerated by the dispersal as lected. Adult traits have been shown to be more important ship-fouling adults (Rilov & Galil, 2009). The high pheno- than the larval-dispersal potential in determining natural typic plasticity typical for sessile, substrate-attached species range expansions of marine bivalves and fishes (Roy et al., (Bayne, 2004) may further promote their survival and expan- 2001; Luiz et al., 2012). sion in the new region. Finally, among suspension-feeding bivalves, epifaunal species tend to have higher clearance rates and greater feeding efficiency under low seston loads (Bacon Establishment et al., 1998; Velasco & Navarro, 2002). This would provide The time since the first Mediterranean record (1–140 years, an important advantage in the highly oligotrophic conditions Fig. 1) had the strongest effect on the probability of estab- of the Levantine basin. lishment of Lessepsian bivalves. This likely reflects the inher- ent lag in the growth of incipient populations related to Spread early colonization dynamics (Crooks, 2005), but also lags in alien species detection, as the establishment status of non- The small number of invasive species among Lessepsian biv- native marine invertebrates is typically inferred from the alves calls for caution when interpreting the result for the number and spatio-temporal distribution of their records in last invasion stage. Association with hard substrate and large the invaded region. Delayed establishment of early immi- body size were the strongest predictors of the invasive status. grants may be also related to the general warming trend Both factors may favour greater detectability of conspicuous, (Raitsos et al., 2010) and progressive environmental changes large epifaunal species. However, several lines of evidence in the eastern Mediterranean (Rilov et al., 2004; Rilov & suggest that the patterns observed at the establishment and Galil, 2009). No significant differences were found in the spread stage cannot be entirely attributed to a sampling bias. date of the first Mediterranean record between non-invasive First, such bias should be strongest during the detection of and invasive alien molluscs. This confirms the markedly var- newly arrived aliens, but only a minor shift in median body ied dynamics of the spread of Lessepsian species (Rilov et al., size and proportion of hard-substrate species can be observed 2004; Ben Rais Lasram et al., 2008; Galil, 2008) and under- at the arrival stage (Fig. 1). Second, among 42 species of Les- scores the difficulty in predicting the outcome of any given sepsian bivalves that are known from the Mediterranean introduction event. coast of Israel, undoubtedly the best-sampled sector of the Despite the paucity of natural rocky shores in the Suez Levantine shelf (e.g. Barash & Danin, 1992), hard-substrate Canal and on its both ends (Por, 1971), the ratio between forms predominate among established and invasive species hard- and soft-bottom species systematically increases along just like in the total pool of aliens. Third, in identifying spe- the course of invasion, especially during the establishment cies that reached the spread stage, we followed Zenetos et al. stage. This suggests that Mediterranean hard substrates may (2010), who defined invasive aliens based on their significant

Diversity and Distributions, 21, 1075–1086, ª 2015 John Wiley & Sons Ltd 1081 R. Nawrot et al. effects on native communities, rather than just a large geo- graphic range in the Mediterranean Sea. Over-representation of epifaunal species among invasive 8 aliens may be related to anthropogenic alteration of recipient hard-substrate communities, which can increase the impact ● of newcomers by putting native species in competitive disad- ● ● vantage (Byers, 2002a). A recent shift in habitat conditions ● ● 6 ● along the Levantine rocky shores has been identified as the ● ● main driver of the rapid population increase in Brachidontes ● ● ● ● pharaonis (Fischer, 1870) (Rilov et al., 2004) and may (size) 2 ● enhance the success of other epifaunal aliens. Several Lessep- ● ●

Log ● sians are acting as ecosystems engineers, increasing the struc- 4 ● tural complexity of hard-bottom habitats and affecting the diversity and composition of local communities. Examples ● ● include dense mussel beds formed by B. pharaonis and reefs constructed by Spondylus spinosus Schreibers, 1793, and Cha- ma pacifica Broderip, 1835, that are colonized by a variety of 2 native and alien epibionts (Mienis et al., 1993; Streftaris & ● Invasive Zenetos, 2006). ● Established Positive association between body size and invasion success ● Casual has been suggested in the context of natural (Roy et al., 2001; Vermeij et al., 2008) and human-assisted range expansions of marine bivalves (Roy et al., 2002; but see Miller et al., 2002). Mytilidae Pteriidae Cardiidae This relationship, however, was important only at the last Ostreidae Chamidae stage of the Lessepsian invasion in our study. Comparison of Spondylidae alien species with those that failed to enter the Mediterranean Figure 4 Body size of Lessepsian species within bivalve families Sea indicates that that passage through the Suez Canal (i.e. containing at least one invasive alien. Invasive species are always the arrival stage) is not associated with any consistent pattern the largest aliens within a given family, with Mytilidae being the of size selectivity within individual bivalve lineages (Fig. S4 in only exception. Appendix S3). In contrast, the association of large body size with the invasive status is evident in the pool of species that crossed the Canal: with the exception of mytilids, high- if they are able to overcome the limits imposed by competi- impact invaders are always the largest among alien species tion and predation from the resident species (Bishop & Pet- belonging to a given family (Fig. 4). erson, 2006; Duyck et al., 2007), which in turn may depend Large size provides a refuge from predation and is linked on physical attributes of the habitat (Safriel & Sasson-Frostig, to greater competitive abilities (Vermeij et al., 2008). This is 1988; Byers, 2002b; Krassoi et al., 2008). In contrast to other congruent with the decline of native Mediterranean bivalves invasive bivalves that are mostly confined to subtidal habi- in response to the spread of larger-bodied Red Sea species tats, B. pharaonis is spreading primarily in the intertidal (Safriel & Sasson-Frostig, 1988; Galil, 2007; Crocetta et al., zone, where predation and competition levels are lower. 2013). The key role of interactions with resident species in B. pharaonis is a better competitor and a larger species than mediating post-establishment expansion of alien populations the Mediterranean mytilid Mytilaster minimus (Poli, 1795), in marine ecosystems (Byers, 2002a,b; Bishop & Peterson, and at the same time a better colonist (but worse competi- 2006) is also supported by the non-random patterns of mor- tor) than M. auriculatus (Safriel & Sasson-Frostig, 1988; Saf- phospace occupation among the dominant Lessepsian fishes riel, 2013). In accordance with this competitive hierarchy (Azzurro et al., 2014). and colonization–competition trade-off, B. pharaonis is pro- Mytilids represent an exception among Lessepsian families gressively displacing the smaller native species from the inter- in terms of body size of invasive species (Fig. 4). Previous tidal habitats of the eastern Mediterranean, where it is a studies on the population biology of the mussel B. pharaonis more successful invader than the larger M. auriculatus (reviewed by Safriel, 2013) have shown that this invader is (Safriel, 2013). characterized by a more opportunistic life history strategy and smaller body size compared to the closely related but CONCLUSIONS much less widespread Lessepsian Modiolus auriculatus (Kraus, 1848). Life history theory predicts trade-offs between Patterns of selectivity among Lessepsian bivalves indicate that colonization abilities and attributes related to competitive the relative strengths of abiotic and biotic filters change at performance and antipredatory defence (Pianka, 1970). Con- different steps of the invasion process. Occurrence outside sequently, r-selected invaders should be more successful only the tropical zone, pointing to the climate match between the

1082 Diversity and Distributions, 21, 1075–1086, ª 2015 John Wiley & Sons Ltd Invasion success in marine bivalves native and alien range of immigrants, together with the shal- Barash, A. & Danin, Z. (1992) Fauna Palaestina: Mollusca I. low depths of the invasion corridor defines the pool of spe- Annotated list of Mediterranean molluscs of Israel and Sinai. cies potentially capable of reaching and surviving in the The Israel Academy of Sciences and Humanities, Jerusa- Mediterranean Sea. In turn, substrate relationship and large lem. body size were the strongest predictors of establishment and Bayne, B.L. (2004) Phenotypic flexibility and physiological spread of alien species, suggesting that their subsequent suc- tradeoffs in the feeding and growth of marine bivalve mol- cess is largely determined by the habitat-specific strength of luscs. Integrative and Comparative Biology, 44, 425–432. biotic resistance set by the interactions with native species. Belmaker, J., Parravicini, V. & Kulbicki, M. (2013) Ecological The strong evidence for the role of bathymetric and cli- traits and environmental affinity explain Red Sea fish intro- matic filters in limiting invasion of ecologically disparate duction into the Mediterranean. Global Change Biology, 19, groups such as marine bivalves and fishes suggests that the 1373–1382. planned enlargement of the Suez Canal (Galil et al., 2015), Ben Rais Lasram, F., Tomasini, J.A., Guilhaumon, F., Rom- combined with the continuing warming of the eastern Medi- dhane, M.S., Do Chi, T. & Mouillot, D. (2008) Ecological terranean Sea (Raitsos et al., 2010), may have unprecedented correlates of dispersal success of Lessepsian fishes. Marine consequences for the rates of inflow of Lessepsian immi- Ecology Progress Series, 363, 273–286. grants. If species associated with hard substrates have greater Berke, S.K., Jablonski, D., Krug, A.Z., Roy, K. & Toma- probability of becoming established and invasive, as illus- sovych, A. (2013) Beyond Bergmann’s rule: size–latitude trated by our results for alien bivalves, effects of these relationships in marine Bivalvia world-wide. Global Ecology changes will be especially devastating for rocky shore ecosys- and Biogeography, 22, 173–183. tems. Rising sea surface temperatures are likely responsible Bernardi, G., Golani, D. & Azzurro, E. (2010) The genetics for the recent population collapses of several ecologically of Lessepsian bioinvasions. Fish invasions of the Mediterra- important native species on the eastern Mediterranean rocky nean Sea: change and renewal (ed. by D. Golani and B. Ap- reefs (Rilov, 2013). Both the shift in environmental condi- pelbaum-Golani), pp. 71–84. Pensoft Publishers, Sofia- tions and related decline of resident species may facilitate Moscow. establishment of aliens. At the same time, many epifaunal Bishop, M.J. & Peterson, C.H. (2006) When r-selection may invaders represent potential competitors for space and habi- not predict introduced-species proliferation: predation of a tat-forming species further altering the structure and func- nonnative oyster. Ecological Applications, 16, 718–730. tioning of the recipient ecosystems. Blackburn, T.M. & Cassey, P. (2007) Patterns of non-ran- domness in the exotic avifauna of Florida. Diversity and Distributions, 13, 519–526. ACKNOWLEDGEMENTS Blackburn, T.M., Pysek, P., Bacher, S., Carlton, J.T., Duncan, We thank A. Eschner (Vienna) and H. K. Mienis (Jerusalem R.P., Jarosık, V., Wilson, J.R.U. & Richardson, D.M. and Tel Aviv) for help in accessing museum collections in (2011) A proposed unified framework for biological inva- their care. K. Kleemann and P. G. Oliver provided informa- sions. Trends in Ecology & Evolution, 26, 333–339. tion on the taxonomy and ecology of Red Sea bivalves. We Burnham, K.P. & Anderson, D.R. (2002) Model selection and are grateful to P. Albano, M. Stachowitsch, A. Tomasovych multimodel inference: a practical information-theoretic and three anonymous reviewers for their comments, which approach. Springer, Berlin. greatly improved the manuscript. This project was funded by Byers, J.E. (2002a) Impact of non-indigenous species on Ernst Mach grant and KWA grant from the University of natives enhanced by anthropogenic alteration of selection Vienna. R.N. and D.C. are supported by the uni:doc fellow- regimes. Oikos, 97, 449–458. ship of the University of Vienna and the Academic Research Byers, J.E. (2002b) Physical habitat attribute mediates biotic Grant of IISER Kolkata, respectively. resistance to non-indigenous species invasion. Oecologia, 130, 146–156. Cassey, P., Blackburn, T.M., Jones, K.E. & Lockwood, J.L. 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flat and shallow subtidal at the northern Red Sea. Facies BIOSKETCHES (in press), doi: 10.1007/s10347-015-0428-6. Zuschin, M. & Oliver, P.G. (2003) Bivalves and bivalve habi- Rafał Nawrot is a PhD student at the University of Vienna. tats in the northern Red Sea. The northern Bay of Safaga His work focuses on the effects of regional extinctions and (Red Sea, Egypt): an actuopalaeontological approach. VI. Na- biotic invasions on diversity of marine communities in the turhistorisches Museum, Wien. Neogene of the circum-Mediterranean region. Zuschin, M. & Oliver, P.G. (2005) Diversity patterns of biv- alves in a coral dominated shallow-water bay in the north- Devapriya Chattopadhyay is interested in molluscan pal- – ern Red Sea – high species richness on a local scale. aeoecology and predator prey dynamics in Recent and fossil Marine Biology Research, 1, 396–410. assemblages. She is currently working on the effect of climatic variation on diversity of Miocene marine molluscs of India.

SUPPORTING INFORMATION Martin Zuschin is interested in evolutionary palaeoecology and diversity dynamics of the Cenozoic marine faunas. He is Additional Supporting Information may be found in the currently working on diversity, conservation palaeobiology online version of this article: and historical ecology in the Adriatic Sea and Indian Ocean. Appendix S1 List of Red Sea bivalve species used in this Author contributions: R.N. and D.C. conceived the ideas; study. R.N., D.C. and M.Z. collected the data; R.N. analysed the Appendix S2 Literature sources used to compile species list data; and R.N. and M.Z. led the writing. and ecological data. Appendix S3 Supplementary figures and tables. Editor: Hugh MacIsaac

1086 Diversity and Distributions, 21, 1075–1086, ª 2015 John Wiley & Sons Ltd Appendix S1 List of Red Sea bivalve species used in this study.

Supraspecific classification and taxonomic nomenclature was based primarily on Carter et al. (2011) and the World Register of Marine Species (WoRMS Editorial Board, 2014), and the acclimatization status in the Mediterranean Sea on Zenetos et al. , 2010. See Appendix S2 for the complete list of literature sources. N-Aln, non-alien species; Cas, casual alien; Est, established non-invasive alien; Inv, established invasive alien.

Order Family Species Important synonyms Invasive status in the Mediterranean Nuculida Nuculidae Nucula inconspicua H. Adams, 1871 N-Aln Solemyida Nucinellidae Huxleyia diabolica (Jousseaume, 1897) N-Aln Solemyida Solemyidae Solemya sp. N-Aln Nuculanida Nuculanidae Nuculana sculpta (Issel, 1869) N-Aln Mytilida Mytilidae Brachidontes pharaonis (P. Fischer, 1870) B. variabilis (Krauss, 1948) Inv Mytilida Mytilidae Septifer cumingii Récluz, 1849 S. forskali Dunker, 1855 Inv Mytilida Mytilidae Septifer excisus (Wiegmann, 1837) N-Aln Mytilida Mytilidae Perna perna (Linnaeus, 1758) P. picta (Born, 1778) N-Aln Mytilida Mytilidae Musculus viridulus (H. Adams, 1871) N-Aln Mytilida Mytilidae Musculus coenobitus (Vaillant, 1865) N-Aln Mytilida Mytilidae Musculus cumingianus (Reeve, 1857) N-Aln Mytilida Mytilidae Arcuatula senhousia (Benson in Cantor, 1842) A. arcuatula sensu auct. N-Aln 1) Mytilida Mytilidae Arcuatula perfragilis (Dunker, 1857) Est Mytilida Mytilidae Solamen striatissima (G. B. Sowerby III, 1904) N-Aln Mytilida Mytilidae Solamen persicum (E. A. Smith, 1906) S. adamsianum (Melvill & Standen, 1907) N-Aln Mytilida Mytilidae Rhomboidella vaillanti (Issel, 1869) N-Aln Mytilida Mytilidae Fungiacava eilatensis Goreau et al., 1968 N-Aln Mytilida Mytilidae Gregariella ehrenbergi (Issel, 1869) N-Aln Mytilida Mytilidae Modiolus auriculatus (Krauss, 1848) Est Mytilida Mytilidae Lioberus ligneus (Reeve, 1858) Cas Mytilida Mytilidae Jolya rhomboidea (Reeve, 1857) Modiolus sirahensis (Jousseaume, 1891) N-Aln Mytilida Mytilidae Modiolus philippinarum (Hanley, 1843) N-Aln Mytilida Mytilidae Amygdalum striatum (Hutton, 1873) A. cf. dendriticum sensu auct. N-Aln Mytilida Mytilidae Botula cinnamomea (Gmelin, 1791) N-Aln Mytilida Mytilidae Lithophaga teres (Philippi, 1846) N-Aln Mytilida Mytilidae Lithophaga robusta Jousseaume in Lamy, 1919 N-Aln Mytilida Mytilidae Lithophaga laevigata (Quoy & Gaimard, 1835) N-Aln Mytilida Mytilidae Lithophaga malaccana (Reeve, 1857) N-Aln Mytilida Mytilidae Lithophaga purpurea Kleemann, 1980 N-Aln Mytilida Mytilidae Lithophaga parapurpurea Kleemann, 2008 N-Aln Mytilida Mytilidae Lithophaga dahabensis Kleemann, 2008 N-Aln Mytilida Mytilidae Lithophaga simplex Iredale, 1939 N-Aln Mytilida Mytilidae Lithophaga obesa (Philippi, 1847) N-Aln Mytilida Mytilidae Lithophaga pulchra Jousseaume, 1919 N-Aln Mytilida Mytilidae Lithophaga lima (Jousseaume in Lamy, 1919) N-Aln Mytilida Mytilidae Lithophaga lessepsiana (Vaillant, 1865) N-Aln Mytilida Mytilidae Lithophaga hanleyana (Reeve, 1857) N-Aln Mytilida Mytilidae Lithophaga tripartita (Jousseaume, 1894) N-Aln Arcida Arcidae Arca acuminata Krauss, 1848 N-Aln Arcida Arcidae Arca patriarchalis Röding, 1798 A. avellana Lamarck, 1819 N-Aln Arcida Arcidae Arca ventricosa Lamarck, 1819 N-Aln Arcida Arcidae Arca navicularis Bruguière, 1789 N-Aln Arcida Arcidae Acar plicata (Dillwyn, 1817) Cas Arcida Arcidae Vitracar sulcata (Lamarck, 1819) Barbatia reticulata sensu auct. N-Aln Arcida Arcidae Samacar strabo (Hedley, 1915) Barbatia acupicta (Viader, 1951) N-Aln Arcida Arcidae Barbatia cometa (Reeve, 1844) N-Aln Arcida Arcidae Barbatia foliata (Forsskål in Niebuhr, 1775) N-Aln Arcida Arcidae Barbatia trapezina (Lamarck, 1819) B. decussata (Sowerby, 1833) N-Aln Arcida Arcidae Barbatia perinesa Oliver & Chesney, 1994 N-Aln Arcida Arcidae Barbatia amygdalumtostum Röding, 1798 B. fusca (Bruguière, 1789) N-Aln Arcida Arcidae Barbatia setigera (Reeve, 1844) N-Aln Arcida Arcidae Barbatia parva (G.B. Sowerby I, 1833) N-Aln Arcida Arcidae Calloarca tenella (Reeve, 1844) N-Aln Arcida Arcidae Trisidos tortuosa (Linnaeus, 1758) N-Aln Arcida Arcidae Anadara antiquata (Linnaeus, 1758) N-Aln Arcida Arcidae Anadara uropigimelana (Bory de Saint-Vincent, 1827) N-Aln Arcida Arcidae Anadara vellicata (Reeve, 1844) A. birleyana (Melvill & Standen, 1907) N-Aln Arcida Arcidae Anadara ehrenbergi (Dunker, 1868) N-Aln Arcida Arcidae Anadara auriculata (Lamarck, 1819) N-Aln Arcida Arcidae Mosambicarca erythraeonensis (Jonas in Philippi, 1851) N-Aln Arcida Arcidae Anadara pygmaea (H. Adams, 1872) N-Aln Arcida Arcidae Anadara natalensis (Krauss, 1848) Est Arcida Arcidae Bathyarca anaclima (Dall, 1981) N-Aln Arcida Noetiidae Ribriarca polycymoides (Thiele & Jaeckel, 1931) Arcopsis ornata (Viader, 1951) N-Aln Arcida Noetiidae Striarca erythraea (Issel, 1869) N-Aln Arcida Noetiidae Sheldonella lateralis (Reeve, 1844) N-Aln Arcida Cucullaeidae Cucullaea labiata (Lightfoot, 1786) C. cucullata (Röding, 1798) N-Aln Arcida Limopsidae Limopsis multistriata (Forsskål in Niebuhr, 1775) Cas Arcida Limopsidae Limopsis marerubra Oliver & Zuschin, 2000 N-Aln Arcida Limopsidae Limopsis elachista Sturany, 1899 N-Aln Arcida Glycymerididae Tucetona audouini (Jousseaume in Lamy, 1916) N-Aln Arcida Glycymerididae Glycymeris livida (Reeve, 1844) N-Aln Arcida Glycymerididae Glycymeris arabica (H. Adams, 1871) Cas Ostreida Pinnidae Pinna muricata Linnaeus, 1758 N-Aln Ostreida Pinnidae Pinna bicolor Gmelin, 1791 N-Aln Ostreida Pinnidae Atrina pectinata (Linnaeus, 1767) N-Aln Ostreida Pinnidae Atrina vexillum (Born, 1778) N-Aln Ostreida Pinnidae Streptopinna saccata (Linnaeus, 1758) N-Aln Ostreida Malleidae Malleus regula Forsskål, 1775 Est Ostreida Malleidae Malleus anatinus (Gmelin, 1791) M. normalis Lamarck, 1819 N-Aln Ostreida Malleidae Malleus albus Lamarck, 1819 M. savignyi Jousseaume in Lamy, 1919 N-Aln Ostreida Pteriidae Pteria aegyptiaca (Dillwyn, 1817) N-Aln Ostreida Pteriidae Pteria avicular (Holten, 1802) P. producta (Reeve, 1857) N-Aln Ostreida Pteriidae Pteria penguin (Röding, 1798) P. macroptera (Lamarck, 1819) N-Aln Ostreida Pteriidae Pteria tortirostris (Dunker, 1848) N-Aln Ostreida Pteriidae Electroma alacorvi (Dillwyn, 1817) N-Aln Ostreida Pteriidae Electroma vexillum (Reeve, 1857) Est Ostreida Pteriidae Pteria physoides (Lamarck, 1819) Electroma zebra (Reeve, 1857) N-Aln Ostreida Pteriidae Pinctada margaritifera (Linnaeus, 1758) N-Aln 2) Ostreida Pteriidae Pinctada imbricata radiata (Leach, 1814) P. vulgaris (Schumacher, 1817) Inv Ostreida Pteriidae Vulsella fornicata (Forsskål in Niebuhr, 1775) N-Aln Ostreida Pteriidae Vulsella vulsella (Linnaeus, 1758) N-Aln Ostreida Pteriidae Isognomon nucleus (Lamarck, 1819) N-Aln Ostreida Pteriidae Isognomon legumen (Gmelin, 1791) N-Aln Ostreida Pteriidae Isognomon isognomum (Linnaeus, 1758) N-Aln Ostreida Pteriidae Crenatula picta (Gmelin, 1791) N-Aln Ostreida Ostreidae Saccostrea cucullata (Born, 1778) Est Ostreida Ostreidae Lopha cristagalli (Linnaeus, 1758) N-Aln Ostreida Ostreidae Nanostrea fluctigera (Jousseaume in Lamy, 1925) N. exigua Harry, 1985; Est Ostrea deformis Lamarck, 1819 Ostreida Ostreidae Dendostrea crenulifera (Sowerby, 1871) Est Ostreida Ostreidae Alectryonella plicatula (Gmelin, 1791) Cas Ostreida Ostreidae Dendostrea folium (Linnaeus, 1758) D. frons sensu auct. Inv Ostreida Ostreidae Booneostrea subucula (Jousseaume in Lamy, 1925) N-Aln Ostreida Gryphaeidae Hyotissa hyotis (Linnaeus, 1758) N-Aln Ostreida Gryphaeidae Parahyotissa numisma (Lamarck, 1819) N-Aln Pectinida Pectinidae Pecten erythraeensis G.B. Sowerby II, 1842 N-Aln Pectinida Pectinidae Gloripallium maculosum (Forsskål, 1775) N-Aln Pectinida Pectinidae Gloripallium pallium (Linnaeus, 1758) N-Aln Pectinida Pectinidae Excellichlamys spectabilis (Reeve, 1853) N-Aln Pectinida Pectinidae Decatopecten plica (Linnaeus, 1758) N-Aln Pectinida Pectinidae Decatopecten amiculum Philippi, 1851 N-Aln Pectinida Pectinidae Mirapecten yaroni Dijkstra & Knudsen, 1998 N-Aln Pectinida Pectinidae Mirapecten tuberosus Djikstra & Kilburn, 2001 M. rastellum sensu auct. N-Aln Pectinida Pectinidae Juxtamusium maldivense (E.A. Smith, 1903) N-Aln Pectinida Pectinidae Semipallium crouchi (E.A. Smith, 1903) N-Aln Pectinida Pectinidae Laevichlamys rubromaculata (Sowerby II, 1842) N-Aln Pectinida Pectinidae Laevichlamys superficialis (Forsskål, 1775) N-Aln Pectinida Pectinidae Laevichlamys andamanica (Preston, 1908) N-Aln Pectinida Pectinidae Pedum spondyloideum (Gmelin, 1791) N-Aln Pectinida Pectinidae Mimachlamys sanguinea (Linnaeus, 1758) M. senatoria (Gmelin, 1791) Cas Pectinida Pectinidae Cryptopecten nux (Reeve, 1853) N-Aln Pectinida Pectinidae Delectopecten musorstomi Poutiers, 1981 N-Aln Pectinida Pectinidae Coralichlamys madreporarum (G. B. Sowerby II, 1842) N-Aln Pectinida Propeamussiidae Parvamussium formosum (Melvill in Melvill & Standen, N-Aln 1907) Pectinida Propeamussiidae Parvamussium siebenrocki (Sturany, 1901) N-Aln Pectinida Propeamussiidae Parvamussium thyrideus (Melvill in Melvill & Standen, N-Aln 1907) Pectinida Propeamussiidae Similipecten eous (Melvill in Melvill & Standen, 1907) N-Aln Pectinida Spondylidae Spondylus groschi Lamprell & Kilburn, 1995 S. marisrubri sensu auct. N-Aln 2) Pectinida Spondylidae Spondylus darwini Jousseaume, 1882 S. pickeringae Lamprell, 1998; N-Aln S. crassisquamatus sensu auct. Pectinida Spondylidae Spondylus spinosus Schreibers, 1793 S. marisrubri Röding, 1798 Inv Pectinida Spondylidae Spondylus nicobaricus Schreibers, 1793 S. hystrix Röding, 1798 Cas Pectinida Spondylidae Spondylus candidus Lamarck, 1819 N-Aln Pectinida Spondylidae Spondylus fauroti Jousseaume, 1888 S. smytheae Lamprell, 1998 N-Aln Pectinida Spondylidae Spondylus layardi Reeve, 1856 N-Aln Pectinida Spondylidae Spondylus avramsingeri Kovalis, 2010 N-Aln Pectinida Plicatulidae Plicatula plicata (Linnaeus, 1767) N-Aln Pectinida Plicatulidae Plicatula australis Lamarck, 1918 N-Aln Pectinida Limidae Lima paucicostata G. B. Sowerby II, 1843 N-Aln Pectinida Limidae Lima vulgaris (Link, 1807) N-Aln Pectinida Limidae Limaria viali (Jousseaume in Lamy, 1916) N-Aln Pectinida Limidae Ctenoides annulatus (Lamarck, 1819) N-Aln Pectinida Limidae Ctenoides concentricus (G. B. Sowerby III, 1888) N-Aln Pectinida Limidae Limatula pusilla (H. Adams, 1871) N-Aln Pectinida Limidae Limea pectinata H. Adams, 1870 N-Aln Pectinida Limidae Limaria fragilis (Gmelin, 1791) N-Aln Pectinida Anomiidae Anomia achaeus Gray, 1850 N-Aln Pectinida Anomiidae Monia nobilis (Reeve, 1859) N-Aln Pectinida Placunidae Placuna placenta (Linnaeus, 1758) N-Aln Carditida Carditidae Cardita variegata Bruguière, 1792 N-Aln Carditida Carditidae Cardites rufus (Deshayes in Laborde & Linant, 1830) N-Aln Carditida Carditidae Centrocardita akabana (Sturany, 1899) Cas Carditida Carditidae Cardita ffinchi (Melvill, 1898) N-Aln Carditida Carditidae Beguina semiorbiculata (Linnaeus, 1758) N-Aln Carditida Carditidae Beguina gubernaculum (Reeve, 1844) N-Aln Carditida Crassatellidae Bathytormus jousseaumei (Lamy, 1919) N-Aln Lucinida Lucinidae Pegophysema kora (Taylor & Glover, 2005) Anodontia edentula sensus auct. N-Aln Lucinida Lucinidae Pegophysema philippiana (Reeve, 1850) N-Aln Lucinida Lucinidae Euanodontia ovum (Reeve, 1850) N-Aln Lucinida Lucinidae Cryptophysema vesicula (Gould, 1850) N-Aln Lucinida Lucinidae Cryptophysema ovulum (Reeve, 1850) N-Aln Lucinida Lucinidae Leucosphaera salamensis (Thiele & Jaeckel, 1931) N-Aln Lucinida Lucinidae Cardiolucina semperiana (Issel, 1869) N-Aln Lucinida Lucinidae Lamellolucina dentifera (Jonas, 1846) N-Aln Lucinida Lucinidae Lamellolucina oliveri Taylor & Glover 2002 N-Aln Lucinida Lucinidae Rasta lamyi (Abrard, 1942) N-Aln Lucinida Lucinidae Codakia tigerina (Linnaeus, 1758) N-Aln Lucinida Lucinidae Codakia paytenorum (Iradale, 1937) N-Aln Lucinida Lucinidae Ctena divergens (Philippi, 1850) N-Aln Lucinida Lucinidae Cavilucina fieldingi (H. Adams, 1871) N-Aln Lucinida Lucinidae Chavania erythraea (Issel, 1869) N-Aln Lucinida Lucinidae Pillucina vietnamica Zorina, 1978 P. fischeriana (Issel, 1869); N-Aln P. concinna (Adams, 1871) Lucinida Lucinidae Divaricella macandrewae (H. Adams, 1871) N-Aln Lucinida Lucinidae Divalinga arabica Dekker & Goud, 1994 Cas Lucinida Lucinidae Liralucina sperabilis (Hedley, 1909) N-Aln Lucinida Lucinidae Funafutia crosseana (Issel, 1869) N-Aln Lucinida Lucinidae Barbierella scitula Oliver & Abou-Zeid, 1986 N-Aln Lucinida Thyasiridae Thyasira sp. N-Aln Pholadida Myidae Cryptomya elliptica (A. Adams, 1851) N-Aln Pholadida Myidae Tugonia nobilis H. Adams & A. Adams, 1856 N-Aln Pholadida Myidae Tugonia decurtata (A. Adams, 1851) N-Aln Pholadida Myidae Sphenia rueppellii A. Adams, 1851 Est Pholadida Corbulidae Corbula sulculosa H. Adams, 1870 N-Aln Pholadida Corbulidae Corbula erythraeensis H. Adams, 1871 N-Aln Pholadida Corbulidae Corbula taitensis Lamarck, 1818 N-Aln Pholadida Pholadidae Barnea manilensis (Philippi, 1847) B. erythraea (Gray, 1851) N-Aln Pholadida Pholadidae Pholadidea fauroti Jousseaume, 1888 N-Aln Pholadida Pholadidae Martesia striata (Linnaeus, 1758) N-Aln Cardiida Ungulinidae Diplodonta subrotunda (Issel, 1869) N-Aln Cardiida Ungulinidae Diplodonta globosa (Forsskål, 1775) N-Aln Cardiida Ungulinidae Diplodonta moolenbeeki van Aartsen & Goud, 2006 N-Aln Cardiida Ungulinidae Diplodonta bogii van Aartsen, 2004 Est Cardiida Ungulinidae Diplodonta genethlia (Melvill, 1898) D. raveyensis sensu auct. N-Aln Cardiida Lasaeidae Sagamikellia khoroica Oliver & Chesney, 1997 N-Aln Cardiida Galeommatidae Levanderia erythraeensis Sturany, 1905 N-Aln Cardiida Galeommatidae Amphilepida aurantia (Dashayes, 1835) Scintilla obockensis Jousseaume, 1888 N-Aln Cardiida Galeommatidae Amphilepida faba (Dashayes, 1856) N-Aln Cardiida Galeommatidae Lionelita denticulata (Dashayes, 1856) N-Aln Cardiida Galeommatidae Scintillula scintillula Jousseaume, 1888 S. sulphurea Sturany, 1899 N-Aln Cardiida Galeommatidae Scintillula variabilis (Sturany, 1899) N-Aln Cardiida Galeommatidae Issina issina Jousseaume, 1898 N-Aln Cardiida Lasaeidae Kellia cycladiformis (Dashayes, 1851) N-Aln Cardiida Lasaeidae Marikellia pustula (Dashayes, 1863) N-Aln Cardiida Basterotiidae Basterotia borbonica (Deshayes in Maillard, 1863) N-Aln Cardiida Basterotiidae Basterotia angulata (H. Adams, 1871) N-Aln Cardiida Basterotiidae Basterotia arcula Melvill, 1898 Eucharis spaldingi Jousseaume in Lamy, 1925 N-Aln Cardiida Chamidae Chama lazarus Linnaeus, 1758 N-Aln Cardiida Chamidae Chama asperella (Lamarck, 1819) Cas Cardiida Chamidae Chama limbula (Lamarck, 1819) N-Aln Cardiida Chamidae Chama brassica (Reeve, 1847) C. brassica elatensis Delsaerdt, 1986 N-Aln Cardiida Chamidae Chama croceata Lamarck, 1819 C. imbricata Broderip, 1835; N-Aln C. savignyi Jousseaume in Lamy, 1921; C. plinthota Cox, 1927 Cardiida Chamidae Chama pacifica Broderip, 1834 C. reflexa Reeve, 1846 Inv Cardiida Chamidae Chama aspersa Reeve, 1846 Est Cardiida Chamidae Chama yaroni Delsaerdt, 1986 N-Aln Cardiida Chamidae Pseudochama corbierei (Jonas, 1846) Est Cardiida Chamidae Pseudochama rianae Delsaerdt, 1986 N-Aln Cardiida Cardiidae Fragum sueziense (Issel, 1869) N-Aln Cardiida Cardiidae Afrocardium richardi (Audouin, 1826) Est Cardiida Cardiidae Ctenocardia fornicata (Sowerby II, 1840) N-Aln Cardiida Cardiidae Fragum nivale (Reeve, 1845) N-Aln Cardiida Cardiidae Lunulicardia hemicardium (Linnaeus, 1758) N-Aln Cardiida Cardiidae Fulvia fragilis (Forsskål in Niebuhr, 1775) Inv Cardiida Cardiidae Fulvia australis (Sowerby II, 1834) Cas Cardiida Cardiidae Lyrocardium anaxium Oliver & Chesney, 1997 N-Aln Cardiida Cardiidae Vasticardium pectiniforme (Born, 1780) N-Aln Cardiida Cardiidae Vasticardium marerubrum (Voskuil & Onverwagt, 1991) N-Aln Cardiida Cardiidae Acrosterigma maculosum (W. Wood, 1815) Vasticardium arenicolum (Reeve, 1845) N-Aln Cardiida Cardiidae Laevicardium biradiatum (Bruguière, 1789) N-Aln Cardiida Cardiidae Laevicardium attenuatum (G. B. Sowerby II, 1841) N-Aln Cardiida Cardiidae Frigidocardium torresi (E. A. Smith, 1885) N-Aln Cardiida Cardiidae Maoricardium pseudolima (Lamarck, 1819) N-Aln Cardiida Cardiidae Vasticardium assimile (Reeve, 1844) N-Aln Cardiida Cardiidae Lunulicardia auricula (Niebuhr, 1775) N-Aln Cardiida Cardiidae Lunulicardia orlini Mienis, 2009 L. auricula sensu auct. N-Aln Cardiida Cardiidae Tridacna maxima (Röding, 1798) N-Aln Cardiida Cardiidae Tridacna squamosa Lamarck, 1819 N-Aln Cardiida Cardiidae Tridacna squamosina Sturany, 1899 T. costata Richter et al. 2008 N-Aln Cardiida Mactridae Mactra achatina Holten, 1802 N-Aln Cardiida Mactridae Mactra lilacea Lamarck, 1818 Est Cardiida Mactridae Mactrotoma ovalina (Lamarck, 1819) N-Aln Cardiida Mactridae Mactra olorina Philippi, 1846 Est Cardiida Mactridae Mactrinula tryphera Melvill, 1899 N-Aln Cardiida Mactridae Lutraria turneri Jousseaume, 1891 L. australis sensu auct. N-Aln Cardiida Mactridae Meropesta nicobarica (Gmelin, 1791) N-Aln Cardiida Mactridae Eastonia solanderi (Gray, 1837) N-Aln Cardiida Mactridae Raeta pellicula (Dashayes, 1855) N-Aln Cardiida Mesodesmatidae Atactodea striata (Gmelin, 1791) A. glabrata (Gmelin, 1791) Cas Cardiida Mesodesmatidae Coecella geratensis Morris & Morris, 1993 C. cf. zebuensis sensu auct. N-Aln Cardiida Tellinidae Angulus vernalis (Hanley, 1844) N-Aln Cardiida Tellinidae Angulus flaccus (Römer, 1871) Cas Cardiida Tellinidae Exotica triradiata (H. Adams, 1871) N-Aln Cardiida Tellinidae Moerella lactea (H. Adams, 1871) N-Aln Cardiida Tellinidae Semelangulus mesodesmoides Oliver & Zuschin, 2000 N-Aln Cardiida Tellinidae Tellinella staurella (Lamarck, 1818) N-Aln Cardiida Tellinidae Tellinella philippii (Philippi, 1844) T. rastellum Hanley, 1844 N-Aln Cardiida Tellinidae Pharaonella perna (Spengler, 1798) N-Aln Cardiida Tellinidae Pharaonella pharaonis (Spengler, 1798) N-Aln Cardiida Tellinidae Tellinella asperrima (Hanley, 1844) N-Aln Cardiida Tellinidae Pristipagia adamsii (Bertin, 1878) N-Aln Cardiida Tellinidae Tellinella virgata (Linnaeus, 1758) N-Aln Cardiida Tellinidae Tellinella crucigera (Lamarck, 1818) N-Aln Cardiida Tellinidae Serratina sulcata (Wood, 1815) N-Aln Cardiida Tellinidae Serratina capsoides (Lamarck, 1818) N-Aln Cardiida Tellinidae Quidnipagus palatum (Iradale, 1929) N-Aln Cardiida Tellinidae Obtellina sericata (Melvill, 1898) N-Aln Cardiida Tellinidae Scutarcopagia scobinata (Linnaeus, 1758) N-Aln Cardiida Tellinidae Tellidora lamellosa (Issel, 1869) N-Aln Cardiida Tellinidae Arcopella isseli (H. Adams, 1871) N-Aln Cardiida Tellinidae Arcopaginula inflata (Gmelin, 1791) N-Aln Cardiida Tellinidae Herouvalia pulchella (H. Adams, 1870) Gari granulifera Lamy, 1938 N-Aln Cardiida Tellinidae Pinguitellina pinguis Hanley, 1844 N-Aln Cardiida Tellinidae Phylloda foliacea (Linnaeus, 1758) N-Aln Cardiida Tellinidae Tellinides ovalis (Sowerby I, 1825) N-Aln Cardiida Tellinidae Moerella philippinarum (Hanley, 1844) N-Aln Cardiida Tellinidae Angulus bertini (Jousseaume, 1895) N-Aln Cardiida Tellinidae Angulus arsinoensis (Issel, 1869) N-Aln Cardiida Tellinidae Nitidotellina valtonis (Hanley, 1844) Est Cardiida Tellinidae Psammotreta praerupta (Salisbury, 1934) Cas Cardiida Tellinidae Psammotreta edentula (Spengler, 1797) N-Aln Cardiida Tellinidae Florimetis coarctata (Philippi, 1845) N-Aln Cardiida Tellinidae Loxoglypta secunda (Bertin, 1878) N-Aln Cardiida Tellinidae Jactellina clathrata (Dashayes, 1835) N-Aln Cardiida Tellinidae Loxoglypta subpallida (E. A. Smith, 1891) N-Aln Cardiida Semelidae Abra fragillima (Issel, 1869) N-Aln Cardiida Semelidae Semele carnicolor (Hanley, 1847) N-Aln Cardiida Semelidae Semele cordiformis (Holten, 1802) S. sinensis Adams, 1854 N-Aln Cardiida Semelidae Semele lamellosa (Sowerby, 1830) N-Aln Cardiida Semelidae Iacra seychellarum (A. Adams, 1856) N-Aln Cardiida Semelidae Iacra kallima (A. E. Salisbury, 1934) I. speciosa (Dashayes, 1856) N-Aln Cardiida Semelidae Leptomya cochlearis (Hinds, 1844) N-Aln Cardiida Semelidae Leptomya rostrata (H. Adams, 1868) L. subrostrata (Issel, 1869) N-Aln Cardiida Semelidae Leptomyaria etesiaca (Hedley, 1909) N-Aln Cardiida Semelidae Abra aegyptiaca Oliver & Zuschin, 2000 N-Aln Cardiida Semelidae Ervilia scaliola Issel, 1869 N-Aln Cardiida Semelidae Ervilia purpurea (Smith, 1906) N-Aln Cardiida Semelidae Rochefortina sandwichensis (E.A. Smith, 1885) N-Aln Cardiida Semelidae Semele striata (Reeve, 1853) N-Aln Cardiida Psammobiidae Gari pallida (Dashayes, 1855) N-Aln Cardiida Psammobiidae Gari insignis (Dashayes, 1855) N-Aln Cardiida Psammobiidae Gari maculosa (Lamarck, 1818) N-Aln Cardiida Psammobiidae Gari pulcherrima (Deshayes, 1855) N-Aln Cardiida Psammobiidae Gari occidens (Gmelin, 1791) N-Aln Cardiida Psammobiidae Heteroglypta contraria (Deshayes in Maillard, 1863) N-Aln Cardiida Psammobiidae Gari pennata (Dashayes, 1855) N-Aln Cardiida Psammobiidae Gari sharabatiae Rusmore-Villaume, 2005 N-Aln Cardiida Psammobiidae Asaphis violascens (Forsskål, 1775) N-Aln Cardiida Psammobiidae Hiatula rosea (Gmelin, 1791) Soletellina ruppelliana (Reeve, 1857) Cas Cardiida Solecurtidae Solecurtus subcandidus Sturany, 1899 N-Aln Cardiida Solecurtidae Azorinus coarctatus (Gmelin, 1791) N-Aln Cardiida Donacidae Donax erythraeensis Bertin, 1881 N-Aln Cardiida Donacidae Donax clathratus Dashayes, 1855 N-Aln Cardiida Donacidae Donax trifasciatus (Linnaeus, 1758) D. veneriformis Lamarck, 1818; N-Aln D. biradiatus (Forsskal, 1775) Cardiida Donacidae Donax scalpellum Gray, 1825 N-Aln Cardiida Trapezidae Trapezium oblongum (Linnaeus, 1758) N-Aln 2) Cardiida Trapezidae Trapezium bicarinatum (Schumacher, 1817) N-Aln Cardiida Trapezidae Glossocardia obesa (Reeve, 1843) N-Aln Cardiida Trapezidae Coralliophaga coralliophaga (Gmelin, 1791) N-Aln Cardiida Kelliellidae Alveinus miliaceus (Issel, 1869) N-Aln Cardiida Glossidae Meiocardia vulgaris (Reeve, 1845) M. moltkiana sensu auct. N-Aln Cardiida Veneridae Antigona lamellaris Schumacher, 1817 Cas Cardiida Veneridae Periglypta reticulata (Linnaeus, 1758) P. puerpera sensu auct. N-Aln Cardiida Veneridae Periglypta crispata (Dashayes, 1853) N-Aln Cardiida Veneridae Globivenus orientalis (Cox, 1930) Ventricolaria toreuma sensu auct. N-Aln Cardiida Veneridae Circe scripta (Linnaeus, 1758) Cas Cardiida Veneridae Circe rugifera (Lamarck, 1818) N-Aln Cardiida Veneridae Circe crocea (Gray, 1838) N-Aln Cardiida Veneridae Circe intermedia (Reeve, 1863) N-Aln Cardiida Veneridae Redicirce sulcata (Gray, 1838) N-Aln 2) Cardiida Veneridae Gafrarium pectinatum (Linnaeus, 1758) Est Cardiida Veneridae Gafrarium dispar (Holten, 1802) N-Aln Cardiida Veneridae Sunetta effossa (Hanley, 1843) N-Aln Cardiida Veneridae Sunetta sunettina (Jousseaume, 1891) S. contempta Smith, 1891 N-Aln Cardiida Veneridae Tivela stefaninii (Nardini, 1933) T. ponderosa (Koch in Philippi, 1844) N-Aln Cardiida Veneridae Dorisca amica (E.A. Smith, 1885) D. nana (Melvill, 1898) N-Aln Cardiida Veneridae Circenita callipyga (Born, 1778) Cas Cardiida Veneridae Gouldiopa dilecta (Gould, 1861) N-Aln Cardiida Veneridae Gouldiopa consternans (Oliver & Zuschin, 2001) Pauliella miliacea sensu auct. Cas Cardiida Veneridae Callista florida (Lamarck, 1818) Cas Cardiida Veneridae Callista erycina (Linnaeus, 1758) N-Aln Cardiida Veneridae Lioconcha philippinarum (Hanley, 1844) N-Aln Cardiida Veneridae Lioconcha ornata (Dillwyn, 1817) N-Aln Cardiida Veneridae Lioconcha arabaya van der Meij, Moolenbeek & Dekker, L. castrensis sensu auct. N-Aln 2010 Cardiida Veneridae Lioconcha polita (Röding, 1798) N-Aln Cardiida Veneridae Pitar hebraeus (Lamarck, 1818) N-Aln Cardiida Veneridae Aphrodora yerburyi (E.A. Smith, 1891) N-Aln Cardiida Veneridae Hyphantosoma spoori Lamprell & Whitehead 1990 N-Aln Cardiida Veneridae Costellipitar chordatus (Römer, 1869) N-Aln Cardiida Veneridae Samarangia lewinsohni Mienis, 2011 N-Aln Cardiida Veneridae Clementia papyracea (Gray, 1825) Est Cardiida Veneridae Dosinia caelata (Reeve, 1851) N-Aln Cardiida Veneridae Dosinia histrio (Gmelin, 1791) N-Aln Cardiida Veneridae Dosinia pubescens (Philippi, 1847) N-Aln Cardiida Veneridae Dosinia erythraea Römer, 1860 Est Cardiida Veneridae Dosinia hepatica (Lamarck, 1818) N-Aln Cardiida Veneridae Dosinia alta (Dunker, 1848) N-Aln Cardiida Veneridae Timoclea marica (Linnaeus, 1758) Cas Cardiida Veneridae Timoclea roemeriana (Issel, 1869) N-Aln Cardiida Veneridae Timoclea hypopta (Sturany, 1899) N-Aln Cardiida Veneridae Placamen foliaceum (Philippi, 1846) N-Aln Cardiida Veneridae Tapes deshayesii (Hanley, 1844) N-Aln Cardiida Veneridae Tapes literatus (Linnaeus, 1758) N-Aln Cardiida Veneridae Tapes sulcarius (Lamarck, 1818) N-Aln Cardiida Veneridae Venerupis rugosa (G.B. Sowerby II, 1854) N-Aln Cardiida Veneridae Paphia textile (Gmelin, 1791) Est Cardiida Veneridae Protapes ziczac (Linnaeus, 1758) P. sinuosa sensu auct. N-Aln Cardiida Veneridae Marcia opima (Gmelin, 1791) N-Aln Cardiida Veneridae Marcia cordata (Forsskål in Niebuhr, 1775) M. flammea (Gmelin, 1791) N-Aln Cardiida Veneridae Irus irus (Linnaeus, 1758) I. macrophylla (Deshayes, 1853) N-Aln Cardiida Veneridae Aphrodora pudicissima (E. A. Smith, 1894) N-Aln Cardiida Veneridae Kyrina kyrina Jousseaume, 1894 N-Aln Cardiida Veneridae Lajonkairia elegans (H. Adams, 1870) N-Aln Cardiida Veneridae Petricola fabagella Lamarck, 1818 P. hemprichii Issel, 1869 Est Cardiida Veneridae Petricola lapicida (Gmelin, 1791) N-Aln Cardiida Veneridae Asaphinoides madreporicus Jousseaume, 1895 N-Aln Cardiida Gastrochaenidae Gastrochaena cuneiformis Spengler, 1783 G. gigantea Deshayes, 1830; N-Aln G. ruppellii Dashayes, 1855; G. savignyi Pallary, 1926 Cardiida Gastrochaenidae Dufoichaena dentifera (Dufo, 1840) N-Aln Cardiida Gastrochaenidae Dufoichaena pexiphora (Sturany, 1899) N-Aln Cardiida Gastrochaenidae Lamychaena weinkauffi (Sturany, 1899) L. inaequistriata (Jousseaume in Lamy, 1923) N-Aln Cardiida Gastrochaenidae Spengleria mytiloides (Lamarck, 1818) S. plicatilis (Deshayes, 1855) N-Aln Cardiida Gastrochaenidae Cucurbitula cymbium (Spengler, 1783) Est Poromyida Cuspidariidae Cardiomya singaporensis (Hinds, 1843) C. pulchella Adams, 1871 N-Aln Poromyida Cuspidariidae Cardiomya alcocki (E. A. Smith, 1894) N-Aln Pandorida Penicillidae Brechites attrahens (Lightfoot, 1786) N-Aln Pandorida Clavagellidae Bryopa lata (Broderip, 1834) N-Aln Pandorida Pandoridae Pandora flexuosa G.B. Sowerby I, 1822 N-Aln Pandorida Laternulidae Laternula anatina (Linnaeus, 1758) Est Thraciida Periplomatidae Offadesma sp. N-Aln Thraciida Thraciidae Thracia adenensis Melvill, 1898 N-Aln Solenida Pharidae Ensiculus cultellus (Linnaeus, 1758) Cas Solenida Pharidae Siliqua polita (W. Wood, 1815) N-Aln Solenida Solenidae Solen ceylonensis Leach, 1814 N-Aln Solenida Solenidae Solen digitalis Jousseaume, 1891 N-Aln Solenida Solenidae Solen lischkeanus Dunker, 1865 N-Aln Solenida Solenidae Solen cylindraceus Hanley, 1843 N-Aln Solenida Solenidae Solen roseomaculatus Pilsbry, 1901 N-Aln

1) Arcuatula senhousia is an invasive alien in the Mediterranean Sea, where it was accidentally introduced with mariculture from the Western Pacific (Zenetos et al. , 2010). This species was, therefore, excluded from the list of Lessepsian bivalves.

2) Species identified as questionable aliens by Zenetos et al. (2010) were excluded from the list of Lessepsian species.

References

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Zuschin M. & Oliver P.G. (2003) Bivalves and bivalve habitats in the northern Red Sea. The northern Bay of Safaga (Red Sea, Egypt): an actuopalaeontological approach. VI. Appendix S3 Supplementary figures and tables.

Figure S1. Comparison of life habits of Red Sea bivalve species that succeed and failed at different stages of the Lessepsian invasion. Arrival stage: non-alien (N-Aln; n = 339) and alien Red Sea species (Aln; n = 52); establishment stage: casual (Cas; n = 22) and established aliens (Est; n = 30); spread stage, non-invasive (N-Inv; n = 23) and invasive established aliens (Inv; n = 7). Com, commensal; Bor, boring; Epi, epifaunal; End, endobysate; Inf, infaunal; Zoo, zooxanthellae bearers; Car, carnivores; Che, chemoautotrophs; Dep, deposit feeders; Sus, suspension feeders. Figure S2. Number species belonging to different (a) substrate groups and (b) feeding guilds expected to succeed at each invasion stage under the null model of random sampling from the source species pool. The median number of species (± 95 % confidence intervals) expected to represent a given life habit was obtained from 10,000 resampling iterations. The P-values correspond to the proportion of randomizations that produced a value as low or lower than the observed (for lower tail of the null distribution), or greater than or equal to the observed (for the upper tail). Filled points denote observed numbers of species that are significantly larger or smaller than expected from the null model. See Fig. S1 for explanations of abbreviations. Figure S3. Results of model averaging obtained using the alternative classification of life habits. Model-averaged parameter estimates (± 1 standard error and 95 % confidence intervals) are based on the best-supported subset of multivariate logistic regression models (Table S1) linking attributes of Red Sea species with the successful transition through different stages of the Lessepsian invasion. Estimates for substrate guilds express the difference in the probability of success relative to infaunal bivalves. None of the top models included feeding mode. See Fig. S1 for explanations of variable abbreviations. Figure S4. Body size of Red Sea species that succeed (Alien) and failed (Non-Alien) in crossing the Suez Canal within bivalve families containing at least one alien species. Families are ordered according to increasing mean size of all species. Thick horizontal bars represent the median size per group. Note that the direction of change in the median size between the non-Lessepsian and Lessepsian species can be different in individual lineages. Table S1. Top-scoring models (∆AICc<2) liking attributes of Red Sea bivalves to their success at the consecutive stages of the Lessepsian invasion. The best-supported models were selected from a set generated using the alternative classification of life habits. The AICc score, differences from the best model (∆AICc) and model weights (wAICc) are shown. Predictors: EX, extratropical occurrence; YI, year of introduction; MD, minimum depth; SB, substrate; SZ, size.

Model AICc ∆AICc wAICc a) Arrival EX+MD+SZ+SB 280.75 0.00 0.26 EX+MD+SB 281.04 0.29 0.23 EX+MD+SZ 281.84 1.09 0.15 EX+MD 281.86 1.12 0.15 b) Establishment YI+SB 62.08 0.00 0.15 YI+SB+EX 62.64 0.56 0.11 YI+EX 63.11 1.03 0.09 YI 63.49 1.41 0.07 YI+SB+SZ 63.79 1.70 0.06 YI+SB+MD 63.86 1.78 0.06 c) Spread SZ 32.36 0.00 0.20 SZ+MD 33.90 1.53 0.09 SZ+EX 33.92 1.55 0.09 EX 34.36 2.00 0.07