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Latin american journal of aquatic research

On-line version ISSN 0718-560X

Lat. Am. J. Aquat. Res. vol.41 no.2 Valparaíso Apr. 2013

http://dx.doi.org/10.3856/vol41-issue2-fulltext-6 

Research Article

 

Rapid assessment survey for exotic benthic species in the São Sebastião Channel, Brazil

Estudio de evaluación rápida de especies bentónicas exóticas en São Sebastião, Brasil

 

Antonio C. Marques 1, Aline dos Santos Klôh 2, Alvaro Esteves Migotto 3, Ana C. Cabral 2, Ana P. Ravedutti Rigo 2, Ariane Lima Bettim 2, Emanuel L. Razzolini 2, Helena Matthews Cascon 4, Juliana Bardi 1, Laura Pioli Kremer 2, Leandro Manzoni Vieira 1, Luis E. Arruda Bezerra 5, Maria A. Haddad 2, Ronaldo Ruy de Oliveira Filho 6, Silvia M. Millan Gutierre 7, Thaís Pires Miranda 1, Wilson Franklin Jr .6 & Rosana Moreira da Rocha 2

1 Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo R. Matão, Trav. 14, 05508-090, São Paulo, Brazil
2
Departamento de Zoologia, Universidade Federal do Paraná CP 19020, 81531-980, Curitiba, Brazil
3
Centro de Biologia Marinha, Universidade de São Paulo, Rodovia Manoel Hypólito do Rego 131,5 km, Cabelo Gordo, 11600-000, São Sebastião, Brazil
4 Departamento de Biologia, Centro de Ciências, Universidade Federal do Ceará Campus do Pici, Departamento de Biologia Bloco, 906, Pici, 60455-760, Fortaleza, Brazil
5
Departamento de Oceanografia, Universidade Federal de Pernambuco, Av. da Arquitetural s/n Cidade Universitária, 50740-550, Recife, Brazil
6
Instituto de Ciências do Mar, Universidade Federal do Ceará Av. Abolição, 3207, Meireles, 60165-081, Fortaleza, Brazil
7
Departamento de Fisiologia, Universidade Federal do Paraná CP 19031, 81531-980, Curitiba, Brazil
Corresponding author: Rosana Moreira da Rocha (rmrocha@ufpr.br)


ABSTRACT. The study of biological invasions can be roughly divided into three parts: detection, monitoring, mitigation. Here, our objectives were to describe the marine fauna of the area of the port of São Sebastião (on the northern coast of the state of São Paulo, in the São Sebastião Channel, SSC) to detect introduced species. Descriptions of the faunal community of the SSC with respect to native and allochthonous (invasive or potentially so) diversity are lacking for all invertebrate groups. Sampling was carried out by specialists within each taxonomic group, in December 2009, following the protocol of the Rapid Assessment Survey (RAS) in three areas with artificial structures as substrates. A total of 142 species were identified (61 native, 15 introduced, 62 cryptogenic, 4 not classified), of which 17 were Polychaeta (12, 1, 1, 3), 24 Ascidiacea (3, 6, 15, 0), 36 Bryozoa (17, 0, 18, 1), 27 Cmdana (2, 1, 24, 0), 20 Crustacea (11, 4, 5, 0), 2 Entoprocta (native), 16 Mollusca (13, 3, 0, 0). Twelve species are new occurrences for the SSC. Among the introduced taxa, two are new for coastal Brazil. Estimates of introduced taxa are conservative as the results of molecular studies suggest that some species previously considered cryptogenic are indeed introduced. We emphasize that the large number of cryptogenic species illustrates the need for a long-term monitoring program, especially in areas most susceptible to bioinvasion. We conclude that rapid assessment studies, even in relatively well-known regions, can be very useful for the detection of introduced species and we recommend that they be carried out on a larger scale in all ports with heavy ship traffic.

Keywords: bioinvasion, fouling, artificial structures, port, São Sebastião, Brazil, southwest Atlantic.


RESUMEN. El estudio de invasiones biológicas puede ser dividido en tres partes: detección, supervisión y atenuación. El objetivo fue describir la fauna marina del puerto de São Sebastião (costa norte del estado de São Paulo, en el canal de São Sebastião, SSC) para detectar las especies introducidas. No existen descripciones de la comunidad faunística del SSC en relación a la diversidad nativa y alóctona (invasiva o potencialmente) de todos los grupos de invertebrados. El muestreo se efectuó por especialistas de cada grupo taxonómico en diciembre de 2009, siguiendo el protocolo de Estudio de Evaluación Rápida (EER) en tres zonas con sustratos artificiales. Se identificaron 142 especies (61 nativas, 15 introducidas, 62 criptogénicas y cuatro no determinadas), de las cuales 17 correspondieron a poliquetos (12, 1, 1, 3), 24 a ascidias (3, 6, 15, 0), 36 a briozoarios (17, 0, 18, 1), 27 a cnidarios (2, 1, 24, 0), 20 a crustáceos (11, 4, 5, 0), 2 a entoproctos (nativo) y 16 a moluscos (13, 3, 0, 0). Doce especies constituyen nuevos registros para el SSC. Entre los taxa introducidos, dos son nuevos para la costa de Brasil. Las estimaciones de los taxa introducidos son conservativas dado que los resultados obtenidos en estudios moleculares sugieren que algunas especies anteriormente consideradas criptogénicas son introducidas. Se destaca que el gran número de especies criptogénicas refleja la necesidad de un programa de monitoreo a largo plazo, especialmente en las zonas más susceptibles a la invasión biológica. Se concluye que los estudios de evaluación rápida, incluso en las regiones relativamente bien conocidas, pueden ser útiles para detectar especies introducidas y se recomienda expandir este tipo de estudios en todos los puertos con tráfico marítimo.

Palabras clave: bioinvasión, incrustaciones, sustratos artificiales, puerto, São Sebastião, Brasil, Atlántico suroccidental.


 

INTRODUCTION

Bioinvasions are biological phenomena with ecological and evolutionary consequences to local biota. The study of biological invasions can be roughly divided into detection, monitoring, and remediation/mitigation, and all of them are better informed by basic (e.g., biogeographically, ecological, ecophysiological constraints) and applied knowledge (e.g., prevention, management, eradication, conservation). Biological invasions are ubiquitous, continental and marine, in all biomes and ecosystems. The introduction of species is the successful establishment of a species in a region where it did not occur before (Elton, 1958). This process can happen naturally by the expanding population of a species, but can also be caused by human activities, that occur more frequently in recent decades (Carlton, 1996).

Becoming invasive, introduced species may compete with native species (including those of commercial or cultural importance) and displace or prevail over native species, change trophic relationships in the food chain, introduce new diseases or toxic substances that affect native organisms and human populations. Thus, biological invasions have obvious negative consequences for biodiversity and public health, cultural and economical issues for the affected regions.

In the marine realm, biological invasion vectors are fundamentally associated with economic activities such as commerce and tourism, both of which are associated with ports. Consequently, records of invasive species in ports around the world are increasing (Carlton, 1989; Ruiz et al., 2000), often seriously endangering natural habitats. Transportation of potential invaders may be by hull fouling and its associated species, or ballast water (Fofonoff et al., 2003). Both have reached alarming proportions as ships become more numerous, faster and larger, thereby requiring more ballast (Cohen & Carlton, 1998). Despite of the impact of marine bioinvasions, the marine realm has historically received less attention compared to terrestrial and freshwater habitats (Carlton, 1989).

In port areas, the availability of hard artificial substrates (such as buoys, ropes, concrete walls, and marinas) provides many opportunities for settlement and metamorphosis for larvae of introduced species (Stachowicz et al., 1999; Tyrrel & Byers, 2007). After recruitment and establishment, a following stage of the invasion process comprises expansion of the geographical distribution of the species or populations. Virtually any marine taxon may include invasive lineages, such as algae (Mathieson et al., 2003) and Chordata (Castilla et al., 2004).

Detecting invasive species depends on monitoring and adequate knowledge of the native fauna, including genetic, taxonomic, biological and ecological data. Invasion and establishment of a species is frequently not recorded and very seldom observed (Carlton, 2009). Ideally, the control of exotic species will be more efficient with early detection, when populations are small and more easily eradicated or controlled (Bax et al., 2001). The lack of previous information about the species makes it difficult to determine its invasive status, in which case the species is of uncertain origin and is classified as cryptogenic (Carlton, 1996).

Strategies for monitoring species in port areas include Rapid Assessment Survey (RAS, see for instance Cohen et al., 2005; Pederson et al., 2005), a procedure with standardized field sampling effort carried out by specialists in taxonomy of target groups, in which the goal is to sample the maximum number of taxa possible in a short time period. The only previous large scale RAS to detect introduced species in Brazil was in Sepetiba Bay, with three ports, as a demonstration study of the Globallast program of the International Maritime Organization -IMO (Clarke et al., 2004). Few other surveys have been carried out and are only available for the port of Paranaguá, southern Brazil (Neves et al., 2007), Ilha Grande Bay, southeastern Brazil (Ignacio et al., 2010), and the port of Recife, northeastern Brazil (Amaral et al., 2010; Lira et al., 2010). Nonetheless, the literature on introduced species identified in specific taxonomic groups is growing, such as those for ascidians (Rocha & Kremer, 2005; Rocha et al., 2009; Marins et al., 2010); cnidarians (Calder & Mayal, 1998; Paula & Creed, 2004; Nogueira Jr. & Oliveira, 2006; Bardi & Marques, 2009); crustaceans (Tavares, 2004; Farrapeira, 2010a); mollusks (Domaneschi & Martins, 2002; Souza et al., 2003; Silveira et al., 2006; Breves et al., 2010; Lopez et al., 2010) and bryozoans (Gordon et al., 2006; Farrapeira, 2010b). General overviews were given by Lopes et al. (2009) and Ferreira et al. (2009). Unfortunately, comprehensive surveys in Brazil of the benthos, including those not focusing on bioinvasion, are also rare, even for the better known areas of the country (Migotto & Marques, 2006).

Here we attempt to detect potential bioinvasion of benthic species in the São Sebastião Channel, an important Brazilian port area, using a Rapid Assessment Survey technique.

MATERIALS AND METHODS

Study area

The São Sebastião Channel (SSC), on the northern coast of the state of São Paulo, Brazil, comprises several areas of preservation, and among the best known biodiversity in the coast of Brazil, due to the many extensive floral and faunal surveys carried out under the auspices of the Center of Marine Biology of the University of São Paulo. The SSC also includes the large Port of São Sebastião, in use for more than 50 years, and through which passes a total load of 50 million tons year-1. An important part of the port is the Terminal Marítimo Almirante Barroso (TEBAR-Transpetro/Petrobras), responsible for intense traffic of ships importing and exporting petroleum products both coastal and long-distance (http://www.transportes.gov.br/-bit/portos/ssebastiao/deposaosebastiao.htm).

Rapid Assessment Survey (RAS)

Surveys were carried out in three sites in the vicinity of the port of São Sebastião: a) Ilhabela Yacht Club (IYC, 23°46'27"S, 45°21'20"W), a floating marina located on São Sebastião Island, on 11 December 2009; b) Petrobras Pier (PP, 23o48'07''S, 45°23'27"W), a concrete structure with large columns on 12 December 2009; and c) Pontal da Cruz Pier (PCP, 23°46'53"S, 45o23'48''W), a cement structure, on 12 December 2009. IYC and PCP were examined for two hours by 17 people including specialists in Annelida Polychaeta, Ascidiacea, Bryozoa, Cnidaria, Crustacea Cirripedia, Entoprocta and Mollusca that defined the taxonomical scope of the survey. PP was surveyed to a maximum depth of 8.5 m for an hour by four scuba divers.

Benthic animals were collected by hand on artificial substrates, including floats and pier columns, during low tide at PCP and by snorkeling in shallow waters at IYC and PCP. When necessary, samples were kept separately in individualized jars in the field, and larger samples were packed to be sorted in the laboratory. Material was brought to the laboratory and maintained in vivo to be examined by specialists. Dissecting and compound microscopes were used when necessary. Samples were identified while alive to the lowest taxonomic level possible. Part of the material was anesthetized (using magnesium chloride, menthol or M-aminobenzoate ethyl) and fixed (in formalin at 4% prepared with saltwater or in ethanol 70%-100%) following standard procedures for each taxon. The material sampled was deposited in the scientific collections of the Federal University of Ceará, Federal University of Paraná, and University of São Paulo.

RESULTS

A total of 142 taxa were identified, twelve of which are new records for the SSC, but may not all be introduced (Table 1). IYC had 89 taxa, PCP had 81 and PP had 70. The taxa comprise Annelida Polychaeta (17), Ascidiacea (24), Bryozoa (36), Cnidaria (27), Crustacea (20), Entoprocta (2) and Mollusca (16). The majority of taxa were classified as either cryptogenic (69), native (53) or introduced (15), and five were not classified because they were not identified at the species level.

 

Table 1. List of species found in São Sebastião Channel (SSC) in three sites: Ilhabela Yatch Club (IYC), Petrobras Pier (PP) and Pontal da Cruz Pier (PCP) in December 2009.
 
 
 
 
 
 
 
 

 

Most taxa were found in only one site (50.7%), with 30.3% in two sites and 19.0% in all three sites (Table 2). By taxon status, half of the native and more than half of the cryptogenic species were in only one site, while only 20% of the introduced species were from one site and 40% in two or three sites (Table 3).

 

Table 2. Number of species (percentage) present in one, two or three sites in the São Sebastião Channel.
 

 

Table 3. Number of species (percentage) by status present in one, two or three sites in the São Sebastião Channel.
 

 

The fifteen introduced species are Polychaetes (Branchiomma luctuosum), Ascidiacea (Clavelina oblonga, Distaplia stylifera, Aplidiopsis sp., Ascidia sydneiensis, Styela plicata, Eusynstyela sp.), Cnidaria (Carijoa riisei), Crustacea Cirripedia (Amphibalanus amphitrite, Amphibalanus reticulatus, Megabalanus coccopoma, Balanus trigonus), and Mollusca Bivalvia (Myoforceps aristatus, Isognomon bicolor, Perna perna). Three introduced species are reported here for the first time in Brazilian waters (D. stylifera, Aplidiopsis sp., Eusynstyela sp.).

DISCUSSION

Although the SSC may have the best known marine fauna in Brazil (Migotto & Marques, 2006), at least 14 species found, 9.8% of the total, are new records (four polychaetes, two ascidians, one cnidarian, one bryozoan, one barnacle, two entoprocts and three mollusks). Clearly, more taxonomical studies are necessary even for the "well-known" regions. Of the 15 recognized introductions, only eight were previously listed in Lopes (2009), with the remaining seven either not included in that publication (C. oblonga, C. riisei), considered cryptogenic (A. amphritite, B. trigonus), or are new introductions (D. stylifera, Aplidiopsis sp., Eusystyela sp). All the 10 introductions recorded in Ilha Grande Bay (Ignacio et al., 2010) were also found in SSC, evidence of the established condition of these species in the coast of Brazil. Among them, one bryozoan species reported by Ignacio et al. (2010), Schizoporella errata, is herein identified as Schizoporella pungens. Winston (2005) suggested S. errata to be part of a species complex that includes S. pungens, described from the Caribbean, and Schizoporella isabelleana (D'Orbigny, 1842), described from Rio de Janeiro. The similarity among the species in the complex, the necessity of molecular techniques to confirm the identity of Brazilian specimens (as suggested by Tompsett et al., 2009) and the absence of additional studies on biogeography, led us to give a cryptogenic status for S. pungens. We also found Scrupocelaria aff. diadema but this is another complex of species in need of revision and without knowing which species is actually in Brazil we also prefered to give a cryptogenic status to the complex.

It is important to note that most taxa were found in only one site (Table 2). Of the introduced taxa, most were in two or three sites (80%), and only 20% in one site (Table 3). Thus, distribution patterns of the introduced taxa are quite different than those of the native and cryptogenic taxa, and may indicate that the introduced are already widespread at the SSC, in the process of successful establishment. It is therefore urgent that monitoring in the region (as well as other major ports) be swiftly carried out to better understand whether those species are also successfully colonizing natural habitats and threatening the native species elsewhere. The study in Ilha Grande Bay showed that most introduced species also occurred on natural substrata in that region (Ignacio et al., 2010).

Although the sites were not surveyed with the same effort, the number of species in each site was not very different (IYC = 89, PP = 70, PCP = 81). Subtidal areas are usually richer in species than intertidal, which is the case for IYC and PP, but effort in PP was much less then IYC and PCP. PCP survey was mainly intertidal. The fact the most species were found in only one site shows that sites were complementary in species composition and that the RAS should include different types of habitats to comprehensively survey a region.

In this study, Ascidiacea has the greatest number of introduced species, followed by Cirripedia, and Bivalvia. However, if we base ranking on the proportion of introduced, relative to the total number per taxa, the order changes to Cirripedia, Bivalvia, and Ascidiacea (Table 4). In the SSC, Cirripedia is a critical case in which most taxa are introduced, and all introduced taxa are widespread (three species in three sites, one in two sites). These introduced species have been reported elsewhere along the Brazilian coast (Farrapeira, 2010a) and some are very old introductions (Carlton et al., 2011). The taxon Cirripedia certainly deserves attention, especially because of its difficult taxonomy and the few taxonomists that address biodiversity inventories (which may have caused other introduced species to be overlooked).

 

Table 4. Number of species (percentage) by status in the São Sebastião Channel. N: native, I: Introduced, C: Cryptogenic.
 

 

The three introduced species of bivalves reported here were also recorded elsewhere in Brazil. Perna perna was probably introduced during intensive ship traffic between Africa and Brazil during the 1800 and 1900s; earlier sambaquis (coastal Indian fossil deposits) do not contain shells of this species (Souza et al., 2003). It is now established throughout the region between Espirito Santo and Santa Catarina and is cultivated for food. Myoforceps aristatus, widely distributed in the Atlantic Ocean, is a borer that lives in hard substrates and shells of other mollusks. It was first recorded in southeastern Brazil in 2006 (Simone & Gonealves, 2006). Isognomon bicolor, from the Caribbean, has been seen on rocky coasts in Brazil since the 1990s and was first recorded in 1989 in Santa Catarina (Domaneschi & Martins, 2002). It is believed that this invasive bivalve was accidentally introduced between 1970 and 1980 by petroleum platforms, boat hulls or by ballast water of ships. Their rapid population growth must have occurred during the 1990s. The species has no commercial value but competes for the same habitat with other commercially valuable species, causing economic losses. In 2002/2003 it has reached very dense populations of more than 800 individuals 100 cm-2 (Breves-Ramos et al., 2010) in Rio de Janeiro, causing serious changes to the natural hard bottom benthic community, but suffered great mortality in this region in 2006 (Lopes et al., 2009).

Ascidiacea, with the greatest number of introduced species in our study, includes well-known important and aggressive invasive species (McKindsey et al., 2007). Globally, the distributions of many introduced ascidians are restricted to artificial substrates in ports or marinas (Lambert & Lambert, 2003), but little is known about their impact on natural habitats. The cryptogenic Didemnum perlucidum, Diplosoma listerianum, Styela canopus, Microcosmus exaspe-ratus, Herdmania pallida are members of this group. Styela plicata and Ascidia sydneiensis, are well known introduced species, both invading the bivalve commercial cultures in the state of Santa Catarina (Rocha et al., 2009). Eusynstyela sp. (previously identified as Eusynstyela floridana, Rodrigues et al., 1998) may be a new species that was probably introduced in Brazil during the 1990s. Distaplia stylifera was introduced during the last ten years, while the species previously identified as D. stylifera by Rodrigues et al. (1998) is another new and also introduced species. Clavelina oblonga was classified as cryptogenic, but a recent molecular study suggested that it is introduced (Rocha et al., 2012). The genus Aplidiopsis was not found in Brazil until this RAS. We were unable to identify it to species due to the lack of reproductive structures.

Two species of polychaetes, Branchiomma (B. patriota and B. luctuosum) were previously identified as B. nigromaculatum (Baird, 1965) which was reported in coastal Brazil (Rullier & Amoureux, 1979; Duarte & Nalesso, 1996), including at SSC. Because of this erroneous identification, B. patriota was described only recently by Nogueira et al. (2006). According to these authors, B. luctuosum is an introduced species known in the SSC at least since 2009 (Amaral et al., 2010b), while its type locality is in the Red Sea and it is known to be an invasive species in Italy. The cryptogenic species of Spionidae, Polydora colonia, was first recorded in 2001 at Ilha do Mel, Paranaguá Bay, in the state of Paraná (Neves, 2006; Neves & Rocha, 2008). Its records in coastal North and Central America raise doubts as to whether it is an introduced species and hence we consider it to be cryptogenic. Many other species of polychaetes were classified as cryptogenic because of lack of knowledge about their life history and dispersal patterns. Eunice rubra has a wide distribution along the Brazilian coast, in addition to Gulf of Mexico and Caribbean Sea, what led us to consider the hypothesis that its natural dispersal area is throughout tropical and subtropical Western Atlantic, and thus it was classified as native. The other native species are those reported only to the Brazilian coast: Pseudonereis palpata, Nicolea uspiana and Branchiomma patriota.

The cnidarian Carijoa riisei was considered native to the Caribbean until a recent molecular analysis showed that this species is actually Indo-Pacific in origin and the Atlantic records are therefore introductions (Concepcion et al., 2010). It is now very widespread along the Brazilian coast and can be considered naturalized.

Decapod crustaceans, entoprocts, polychaetes and gastropods all have high proportions of native species (over 70%, see Table 4). Pachigrapsus transversus was previously known to occur on both sides of the Atlantic and in the eastern Pacific (Manning & Holthuis, 1981; Hendrickx, 1995; Melo, 1996; Poupin et al., 2005). Recently, Schubart et al. (2005), using morphological and genetic differences (16S mt DNA sequences), revalidated the species P. socius Stimpson, 1871 for the eastern Pacific, limiting the occurrence of P. transversus to the Atlantic Ocean. However, we maintained the status of the species as cryptogenic because it has been proposed that the different populations on both sides of Atlantic may be introduced in some areas. If so, to determine which populations are introduced will require further molecular studies. On the other hand, the decapod Pachycheles monilifera, native to Brazil, has been introduced in Ecuador, in the eastern Pacific (Veloso & Melo, 1993).

In this study, Bryozoa has the greatest number of native species (17). Of these species, both Amathia distans and A. brasiliensis were reported as widespread in warm tropical waters, although a recent study suggested a restricted distribution of these taxa in the western Atlantic (Fehlauer-Ale et al., 2011). In addition, the four widespread species in the western Atlantic found in our study (A. distans, A. brasiliensis, Bowerbankia maxima and Biflustra sp.) were also found in pelagic algae and may be dispersed by algal rafting, as reported for other bryozoans (Taylor & Monks, 1997; Vieira et al., 2010).

It is remarkable that while taxa with a longer history of surveys such as Crustacea Decapoda and Mollusca have a very small proportion of cryptogenic species, a large proportion of species of ascidians, bryozoans and cnidarians, are still considered cryptogenic. These taxa are typically found in small colonies that may have been overlooked in previous faunal studies, and they often comprise many species with wide geographic distribution. Thus the uncertainty of their status, also illustrating the need for periodic monitoring of areas sensitive to bioinvasion, comprehensive surveys of natural areas, and molecular studies to understand their geographical distribution. Also, some widespread bryozoans have been reported to be quite variable morphologically in disjoint areas, which suggest that, in species with short-lived larvae, a complex of cryptic species and hidden endemism may be common (Vieira et al., 2010).

With the use of molecular tools, populations of a given species have been shown to be introduced in other areas (e.g., populations of the Atlantic Clavelina lepadiformis introduced in the Mediterranean - Turon et al., 2003) and species previously considered widely distributed have been split in one or more new species with narrower geographical ranges (e.g., Thais haemastoma - Claremont et al., 2011; Pachigrapsus transversus - Schubart et al., 2005; Botryllus schlosseri - Bock et al., 2012). We suggest that this kind of genetic monitoring is also important for the study of marine bioinvasions, and such studies are being conducted by our research group for ascidians, bryozoans, and cnidarians. Previous results from these molecular studies show that some species of these groups, now considered cryptogenic, may be instead introduced species or introduced populations of haplotypes. Therefore, the estimated number of introduced taxa should be thought to be very conservative and with continued study, many more species will be shown to be introduced.

Here we demonstrate that RASs, even for reasonably well-known regions of Brazil, such as the São Sebastião Channel, are useful strategies to monitor and detect introduced species. We recommend that RASs be replicated on a large scale in all ports with moderate to heavy ship traffic.

ACKNOWLEDGMENTS

We thank the Center of Marine Biology of the University of São Paulo for hosting us during the field studies, the administration of the Ilhabela Yacht Club for allowing us to sample in the marina; Marcos Barbeitos and Carlos Rocha for their field help; Vasily Radashevsky for helping with the identification of the Spionidae Polydora colonia; James Roper for the English text review; María de Los Angeles Becerril for the Spanish abstract; Julia Beneti for information on Anthozoa. This study was an initiative of and primarily supported by the project CAPES-PROCAD 2007/150 "Benthos in port areas along the Brazilian coast: biodiversity, phylogeography and aspects of bioinvasion by biofouling". ACM, JB, and TPM have support from FAPESP (2004/09961-4; 2006/58226-0; 2010/06927-0), CAPES (Pró-Equipamentos and Prodoc projects) and Boticário Foundation. RMR received a research grant from CNPq; APRR, ASK received a master scholarship from CAPES; LMV received a doctoral scholarship from FAPESP (2008/10619-0); LEAB received a post-doctoral scholarship from PNPD/CAPES and FACEPE (BCT 0039-1.08/10). This contribution has the participation of several scientists of NP-BioMar, USP.

REFERENCES

Allman, G.J. 1888. Report on the Hydroida dredged by H.M.S. Challenger during the years 1873-1876. Part II. The Tubularinae, Corymorphinae, Campanularinae, Sertularinae, and Thalamophora. Rep. Scient. Res. Voy. Challenger Zool., 23: 1-90.         [ Links ]

Alves, T.M. 2008. Contribuição ao conhecimento taxonômico de Terebellidae e Trichobranchidae (Annelida: Polychaeta) da região sudeste-sul do Brasil. Dissertação do grau de Mestre. Instituto de Biociências, Universidade de São Paulo, 176 pp.         [ Links ]

Amaral, A.C.Z. 1977. Anelídeos poliquetos do infralitoral em duas enseadas da região de Ubatuba -Aspectos Ecológicos. Tese do grau de Doutor, Universidade de São Paulo, São Paulo, 137 pp.         [ Links ]

Amaral, A.C.Z., S.A.H. Nallin & T.M. Steiner. 2010a. Catálogo das espécies de Annelida Polychaeta do Brasil. [http://www.ib.unicamp.br/destaques\biota\bentos_marinho\prod_cien-\texto_poli.pdf]. Reviewed: 20 January 2011.         [ Links ]

Amaral, A.C.Z., M.R. Denadai, A. Turra & A.E. Rizzo. 2003. Intertidal macrofauna in Brazilian subtropical tide-dominate sandy beaches. J. Coastal Res., 35: 446-455.         [ Links ]

Amaral, A.C.Z., A.E. Migotto, A. Turra & Y. Schaeffer-Novelli. 2010b. Araçá: biodiversity, impacts and threats. Biota Neotrop., 10(1): 219-264.         [ Links ]

Amaral, F.M.D., C.M.R. Farrapeira, S.M.A. Lira & C.A.C. Ramos. 2010. Benthic macrofauna inventory of two shipwrecks from Pernambuco coast, northeastern of Brazil. Rev. Nordest. Zool., 4(1): 24-41.         [ Links ]

Bardi, J. & A.C. Marques. 2009. The invasive hydromedusa Blackfordia virginica (Cnidaria: Black-fordiidae) in southern Brazil, with comments on taxonomy and distribution of the genus Blackfordia. Zootaxa, 2198: 41-50.         [ Links ]

Bax, N., J.T. Carlton, A. Mathews-Amos, R.L. Haedrich, F.G. Howarth, J.E. Purcell, A. Riese & A. Gray. 2001. The control of biological invasions in the world's ocean. Conserv. Biol., 15(5): 1234-1246.         [ Links ]

Bock, D.G., H.J. MacIsaac & M.E. Cristescu. 2012. Multilocus genetic analyses differentiate between widespread and spatially restricted cryptic species in a model ascidian. Proc. Roy. Soc. B, DOI 10.1098/rspb.2011.2610.         [ Links ]

Bollens, S.M., J.R. Cordell & S. Avent. 2002. Zooplankton invasions: brief review plus two case studies from the northeast Pacific Ocean. Hydrobiologia, 480: 87-110.         [ Links ]

Bonnet, N.Y.K. & R.M. Rocha. 2011. The Ascidiidae (Ascidiacea, Tunicata) in coastal Brazil. Zool. Stud., 50(6): 809-825.         [ Links ]

Breves-Ramos, A., A.O.R. Junqueira, H.P. Lavrado, S.H.G. Silva & M.A.G. Ferreira-Silva. 2010. Population structure of the invasive bivalve Isognomon bicolor on rocky shores of Rio de Janeiro State (Brazil). J. Mar. Biol. Assoc. UK, 90(3): 453-459.         [ Links ]

Calder, D.R. & E.M. Mayal. 1998. Dry season distribution of hydroids in a small tropical estuary, Pernambuco, Brasil. Zool. Verhandel., 323: 69-78.         [ Links ]

Cangussu, L.C., L. Altvater, M.A. Haddad, A.C. Cabral, H.L. Heyse & R.M. Rocha. 2010. Substrate type as a selective tool against colonization by non-native sessile invertebrates. Braz. J. Oceanogr., 58(3): 219-231.         [ Links ]

Cano, G. 1889. Crostacei brachiuri ed anomuri raccolti nel viaggio de la "Vettor Pisani" in torno al globo. Boll. Soc. Natural. Napoli, 1(3): 79-268.         [ Links ]

Carlton, J.T. 1989. Man's role in changing the face of the ocean: biological invasions and implications for conservation of near-shore environments. Conserv. Biol., 3(3): 265-273.         [ Links ]

Carlton, J.T. 1996. Pattern, process and prediction in marine invasion ecology. Biol. Conserv., 78: 97-106.         [ Links ]

Carlton, J.T. 2009. Deep invasion ecology and the assembly of communities in historical time. In: G. Rilov & J.A. Crooks (eds.). Biological invasions in marine ecosystems. Springer-Verlag, Berlin, pp. 1356.         [ Links ]

Carlton, J.T., W.A. Newman & F.B. Pitombo. 2011. Barnacle invasions: introduced, cryptogenic, and range expanding Cirripedia of North and South America. In: B.S. Galil, P.F. Clark & J.T. Carlton (eds.). In the wrong place-alien marine crustaceans: distribution, biology and impacts. Springer Netherlands, Dorchester, pp. 159-213.         [ Links ]

Castilla, J.C., N.A. Lagos & M. Cerda. 2004. Marine ecosystem engineering by the alien ascidian Pyura praeputialis on a mid-intertidal rocky shore. Mar. Ecol. Prog. Ser., 168: 119-130.         [ Links ]

Christoffersen, M.L. 1980. Taxonomia e distribuição geográfica dos Alpheoidea (Crustacea, Decapoda, Natantia) do Brasil, Uruguai e norte da Argentina, incluindo considerações sobre a divisão sul do continente em províncias biogeográficas marinhas. Tese do grau de Doutor. Universidade de São Paulo, São Paulo, 467 pp.         [ Links ]

Claremont, M., S.T. Williams, T.G. Barraclough & D.G. Reid. 2011. The geographic scale of speciation in a marine snail with high dispersal potential. J. Biogeogr., 38: 1016-1032.         [ Links ]

Clarke, C., R. Hilliard, A.O.R. Junqueira, A.C.L. Neto, J. Polglaze & S. Raaymakers. 2004. Ballast water risk assessment, Port of Sepetiba, Federal Republic of Brazil. GloBallast Monog. Ser., 14: 63 pp.         [ Links ]

Coelho, P.A. 1964. Lista dos Porcellanidae (Crustacea, Decapoda, Anomura) do litoral de Pernambuco e dos estados vizinhos. Trab. Oceanogr. UFPE, 5/6: 51-68.         [ Links ]

Cohen, A.N. & J.T. Carlton. 1998. Accelerating invasion rate in a highly invaded estuary. Science, 279: 555-558.         [ Links ]

Cohen, A.N., L.H. Harris, B.L. Bingham, J.T. Carlton, J.W. Chapman, C.C. Lambert, G. Lambert, J.C. Ljubenkov, S.N. Murray, L.C. Rao, K. Reardon & E. Schwindt. 2005. Rapid Assessment Survey for exotic organisms in southern California bays and harbors, and abundance in port and non-port areas. Biol. Invas., 7: 995-1002.         [ Links ]

Concepcion, G.T., S.E. Kahng, M.W. Crepeau, E.C. Franklin, S.L. Coles & R.J. Toonen. 2010. Resolving natural ranges and marine invasions in a globally distributed octocoral (genus Carijoa). Mar. Ecol. Progr. Ser., 401: 113-127.         [ Links ]

Correa, D.D. 1964. Corallimorpharia e Actniaria do Atlântico Oeste Tropical. Tese de livre docência. Universidade de São Paulo, São Paulo, 179 pp.         [ Links ]

Dando, P.R. & A.J. Southward. 1980. A new species of Chthamalus (Crustacea Cirripedia) characterized by enzyme electrophoresis and shell morphology: with a revision of the other species of Chthamalus from the western shores of the Atlantic Ocean. J. Mar. Biol. Assoc. UK., 60: 787-831.         [ Links ]

Darwin, C.R. 1854. A monograph on the sub-class Cirripedia with figures of all the species. The Balanidae (or sessile cirripedes); the Verrucidae, etc. Roy. Soc. London, 2: 684 pp.         [ Links ]

Domaneschi, O. & C.M. Martins. 2002. Isognomon bicolor (C.B. Adams) (Bivalvia, Isognomonidae): primeiro registro para o Brasil, redescrição da espécie e considerações sobre a ocorrência e distribuição de Isognomon na costa brasileira. Rev. Bras. Zool., 19: 611-627.         [ Links ]

Duarte, L.F.L. & R.C. Nalesso. 1996. The sponge Zygomycale parishii (Bowerbank) and its endobiotic fauna. Estuar. Coast Shelf Sci., 42: 139-151.         [ Links ]

Elton, C.S. 1958. The ecology of invasions by animals and plants. Methuen, London, 181 pp.         [ Links ]

Farrapeira, C.M.R. 2006. Barnacles (Cirripedia Balanomorpha) of the estuarine region of Recife, Pernambuco, Brazil. Trop. Oceanogr., 34(2): 100-119.         [ Links ]

Farrapeira, C.M.R. 2010a. Shallow water Cirripedia of the northeastern coast of Brazil: the impact of life history and invasion on biogeography. J. Exp. Mar. Biol. Ecol., 392(1-2): 210-219.         [ Links ]

Farrapeira, C.M.R. 2010b. The introduction of the bryozoan Zoobotryon verticillatum (Della Chiaje, 1822) in northeast of Brazil: a cause for concern. Biol. Invas., 13(1): 13-16.         [ Links ]

Fausto-Filho, J. 1970. Quarta contribuição ao inventário dos crustáceos decápodos marinhos do Nordeste Brasileiro. Arq. Ciênc. Mar, 10(1): 55-60.         [ Links ]

Fehlauer-ale, K.H., L.M. Vieira & J.E. Winston. 2011. Molecular and morphological characterization of Amathia distans Busk and Amathia brasiliensis Busk (Bryozoa: Ctenostomata) from the tropical and subtropical western Atlantic. Zootaxa, 2962: 49-62.         [ Links ]

Ferreira, C.E.L., A.O.R Junqueira, M.C. Villac & R.M. Lopes. 2009. Marine bioinvasions in the Brazilian coast: brief report on history of events, vectors, ecology, impacts and management of non indigenous species. In: G. Rilov & J.A. Crooks (eds.). Biological invasions of marine ecosystems: patterns, effects and management. Springer-Verlag, Berlin, pp. 459-478.         [ Links ]

Fofonoff, P.W., G.M. Ruiz, B. Steves & J.T. Carlton. 2003. In ships or on ships? Mechanisms of transfer and invasion for nonnative species to the coasts of North America. In: G.M. Ruiz & J.T. Carlton (eds.). Invasive species: vectors and management strategies. Island Press, Washington, pp. 152-182.         [ Links ]

Gordon, D.P., L.V. Ramalho & P.D. Taylor. 2006. An unreported invasive bryozoan that can affect livelihoods - Membraniporopsis tubigera in New Zealand and Brazil. Bull. Mar. Sci., 78(2): 331-342.         [ Links ]

Hartmeyer, R. 1912. Die Ascidien der Deutschen Tiefsee-Expedition. In: Wiss. Ergebn. Dt. Tiefsee-Exped. Valdivia, 1898-1899, 16(3): 225-392.         [ Links ]

Hendrickx, M.E. 1995. Checklist of brachyuran crabs (Crustacea: Decapoda) from the eastern tropical Pacific. Bull. K. Belg. Inst. Nat. Wet., 65: 125-150.         [ Links ]

Ignacio, B.L., L.M. Julio, A.O.R. Junqueira & M.A.G. Ferreira-Silva. 2010. Bioinvasion in a Brazilian Bay: filling gaps in the knowledge of southwestern Atlantic biota. Plos One, 5: 13065.         [ Links ]

Jäderholm, E. 1903. Aussereuropäische Hydroiden im schwedischen Reichsmuseum. Ark. Zool., 1: 259-312.         [ Links ]

Laborel, J. 1969. Madréporaires et hydrocoralliaires recifaux des côtes brésiliennes. Systematique, ecologie, repartition verticale et geographie. Ann. Inst. Oceanogr., 47: 171-229.         [ Links ]

Lacombe, D. 1977. Cirripédios da Baía da Ribeira, Angra dos Reis. Publ. Inst. Pesq. Marinha, RJ., 109: 1-13.         [ Links ]

Lacombe, D. & W. Monteiro. 1974. Balanideos como indicadores de poluição na Baia de Guanabara. Rev. Bras. Biol., 34: 633-644.         [ Links ]

Lambert, C.C. & G. Lambert. 2003. Persistence and differential distribution of nonindigenous ascidians in harbors of the Southern California Bight. Mar. Ecol. Progr. Ser., 259: 145-161.         [ Links ]

Linnaeus, C. 1758. Systema Naturae. Editio Decima. Regnum Animale. Laurentii Salvii, Holmiae, 1: 824 pp.         [ Links ]

Lira, S.M.A., C.M.R. Farrapeira, F.M.D. Amaral & C.A.C. Ramos. 2010. Sessile and sedentary macro-fauna from the Pirapama shipwreck, Pernambuco, Brazil. Biota Neotrop., 10(4): 155-165.         [ Links ]

Lopes, R.M. 2009. Informe sobre as espécies exóticas invasoras marinhas no Brasil. Série Biodiversidade. Ministério do Meio Ambiente, Brasília, 33: 439 pp.         [ Links ]

López, M.S., R. Coutinho, C.E.L. Ferreira & G. Rilov. 2010. Predator-prey interactions in a bioinvasion scenario: differential predation by native predators on two exotic rocky intertidal bivalves. Mar. Ecol. Prog. Ser., 403: 101-112.         [ Links ]

Luederwaldt, H. 1929. Resultados de uma excursão científica à Ilha de São Sebastião em 1925. Rev. Mus. Paul., 16: 22-79.         [ Links ]

Manning, R.B. & L.B. Holthuis. 1981. West African brachyuran crabs (Crustacea: Decapoda). Smith. Contr. Zool., 306: 1-379.         [ Links ]

Marins, F.O., R.L.M. Novaes, R.M. Rocha & A.O.R. Junqueira. 2010. Non indigenous ascidians in port and natural environments in a tropical Brazilian bay. Zoologia, 27(2): 213-221.         [ Links ]

Marques, A.C. 2001. O gênero Eudendrium (Cnidaria, Hydrozoa, Anthomedusae) no Brasil. Pap. Avuls. Zool., 41: 329-405.         [ Links ]

Marques, A.C. & O.M.P. Oliveira. 2003. Eudendrium caraiuru sp. n. (Hydrozoa; Anthoathecata; Eudendriidae) from the southeastern ocean of Brazil. Zootaxa, 307: 1-12.         [ Links ]

Mathieson, A.C., C.J. Dawes, L.G. Harris & E.J. Hehre. 2003. Expansion of the Asiatic green alga Codium fragile subsp. tomentosoides in the Gulf of Maine. Rhodora, 105: 1-53.         [ Links ]

Melo, G.A.S. 1996. Manual de identificação dos Brachyura (caranguejos e siris) do litoral brasileiro. Plêiade, São Paulo, 603 pp.         [ Links ]

Melo, G.A.S. 1999. Manual de identificação dos Crustacea Decapoda do litoral brasileiro: Anomura, Thalassinidea, Palinuridea, Astacidea. Plêiade, São Paulo, 551 pp.         [ Links ]

Michaelsen, W. 1923. Neue und altbekannte Ascidien aus dem Reichsmuseum zu Stockholm. Mitt. Zool. Mus., 40: 1-60.         [ Links ]

Miers, E.J. 1886. Report on the Brachyura collected by H.M.S. Challenger during the years 1873-1876. In: C. Wyville-Thomson & J. Murray (eds.). Report on the scientific results of the voyage of H.M.S. Challenger during the years 1873-1876. Zoology, 17(49): 1-362.         [ Links ]

Migotto, A.E. 1996. Benthic shallow-water hydroids (Cnidaria, Hydrozoa) of the ocean of São Sebastião, Brazil, including a checklist of Brazilian hydroids. Zool. Verhandel., 306: 1-125.         [ Links ]

Migotto, A.E. & L.P. Andrade. 2000. The life cycle of Hebella furax (Cnidaria: Hydrozoa): a link between a lafoeid hydroid and a laodiceidae medusae. J. Nat. Hist., 34: 1871-1888.         [ Links ]

Migotto, A.E. & A.S. Cabral. 2005. Lafoeina amiran-tensis (Cnidaria: Hydrozoa, Campanulinoidea), the hydroid stage of the medusa Cirrholovenia tetranema (Cnidaria: Hydrozoa, Lovenelloidea). Zootaxa, 919: 1-16.         [ Links ]

Migotto, A.E. & A.C. Marques. 2006. Invertebrados marinhos. In: T. Lewinshn (org.). Avaliação do estado do conhecimento da diversidade brasileira. Série Biodiversidade. Ministério do Meio Ambiente, Brasília, 15(1-2): 147-202.         [ Links ]

Migotto, A.E. & F.L. Silveira. 1987. Hidróides (Cnidaria, Hydrozoa) do litoral sudeste e sul do Brasil: Halocordylidae, Tubulariidae e Corymorphidae. Iheringia, sér. Zool., 66: 95-115.         [ Links ]

Migotto, A.E., C.G. Tiago & A.R.M. Magalhães. 1993. Malacofauna marinha da região costeira do Canal de São Sebastião, SP, Brasil: Gastropoda, Bivalvia, Polyplacophora e Scaphopoda. Bolm Inst. Oceanogr., 41(1): 13-27.         [ Links ]

Migotto, A.E., L.M. Vieira & J.E. Winston. 2011. Bryozoa. In: A.C.Z. Amaral & S.A.H. Nallin (ed.). Biodiversidade e ecossistemas marinhos bentônicos do litoral norte de São Paulo, Sudeste do Brasil. UNICAMP/IB, Campinas, pp. 265-272.         [ Links ]

Millar, R.H. 1958. Some ascidians from Brazil. Ann. Mag. Nat. Hist., 13(1): 497-514.         [ Links ]

Millar, R.H. 1977. Ascidians (Tunicata: Ascidiacea) from the northern and north-eastern Brazilian shelf. J. Nat. Hist., 11(2): 169-223.         [ Links ]

Milne-Edwards, H. 1834. Histoire naturelle des crustacés, comprenant l'anatomie, la physiologie et la classification de ces animaux. Librairie Encyclo-pédique de Roret, Paris, Roret, (1)468, (2)532, (3)638, Atlas 1-32, Plates I-XLII.         [ Links ]

Monniot, C. 1969-1970. Ascidies phlebobranches et stolidobranches. Campagne de la Calypso au large des cotes de l'Amerique du Sud. Ann. Inst. Oceanogr., 47: 33-59.         [ Links ]

Morgado, E.H. & A.C.Z. Amaral. 1981a. Anelídeos poliquetos associados a um briozoário I: Eunicidae, Lumbrineridae, Lysaretidae e Dorvilleidae. Iheringia, sér. Zool., 60: 33-54.         [ Links ]

Morgado, E.H. & A.C.Z. Amaral. 1981b. Anelídeos poliquetos associados a um briozoário II: Palmyridae. Bolm Inst. Oceanogr., 30(1): 87-89.         [ Links ]

Morgado, E.H. & A.C.Z. Amaral. 1985. Anelídeos poliquetos associados ao briozoário Schizoporella unicornis (Johnston). V. Syllidae. Rev. Bras. Zool., 3(4): 219-227.         [ Links ]

Müller, F. 1867. Uber Balanus armatus und einen Bastard dieser Art und des Balanus improvisus var. assimilis. Darw. Arch. Naturg., 33(1): 329-356.         [ Links ]

Neves, C.S. & R.M. Rocha. 2008. Introduced and cryptogenic species and their management in Paranaguá Bay, Brazil. Braz. Arch. Biol. Tech., 51(3): 623-633.         [ Links ]

Neves, C.S., R.M. Rocha, F.B. Pitombo & J.J. Roper. 2007. Artificial substrate use by introduced and cryptogenic marine species in Paranaguá Bay, southern Brazil. Biofouling, 23: 319-330.         [ Links ]

Nogueira, C.C., P.A. Grohmann & V.M.A.P. Silva. 1997. Hydroids from the vicinity of a nuclear power plant site (CNAAA-Unidade I) at Angra-dos-Reis, Rio de Janeiro, southeastern Brazil. In: J.C. den Hartog (ed.). Proceedings of the 6th International Conference on Coelenterate Biology. National Natuurhistorisch Museum, Leiden, pp. 365-369.         [ Links ]

Nogueira, J.M.M., M.C.S. Rossi & E. López. 2006. Intertidal species of Branchiomma Kölliker and Pseudobranchiomma Jones (Polychaeta: Sabellidae: Sabellinae) occurring on rocky shores along the State of São Paulo, Southeastern Brazil. Zool. Stud., 45(4): 586-610.         [ Links ]

Nogueira, Jr., M. & J.S. Oliveira. 2006. Moerisia inkermanica Paltschikowa-Ostroumova (Hydrozoa; Moerisidae) e Blackfordia virginica Mayer (Hydrozoa; Blackfordiidae) na Baía de Antonina, Paraná, Brasil. Pan-Am. J. Aquat. Sci., 1(1): 35-42.         [ Links ]

Nonato, E.F. 1981. Contribuição ao conhecimento dos anelídeos poliquetas bentônicos da plataforma continental brasileira, entre Cabo Frio e o Arroio Chui. Tese de livre docência, Instituto Oceanográfico, Universidade de São Paulo, São Paulo, 246 pp.         [ Links ]

Nonato, E.F. & J.A.C. Luna. 1970. Anelídeos poliquetas do Nordeste do Brasil. I. Poliquetas bentônicos da costa de Alagoas e Sergipe. Bolm Inst. Oceanogr., 19: 57-130.         [ Links ]

Nutting, C.C. 1900. American hydroids. Part I. The Plumulariidae. Spec. Bull. U.S. Natl. Mus., 4: 1-285.         [ Links ]

Oliveira, L.P.H. 1941. Contribuição ao conhecimento dos crustáceos do Rio de Janeiro, subordem "Balanomorpha" (Cirripedia: Thoracica). Mem. Inst. Oswaldo Cruz, 36:1-31.         [ Links ]

Oliveira, J.S., E. Redaelli, A.J. Zaharenko, R.R. Cassulini, K. Konno, D.C. Pimenta, J.C. Freitasm, J.J. Clare & E. Wanke. 2004. Binding specificity of sea anemone toxins to Na (subscript v) 1.1-1.6 sodium channels. J. Biol. Chem., 279(32): 33323-33335.         [ Links ]

Omena, E.P. & A.C.Z. Amaral. 2003. Sandy beach morphodynamic and the Polychaete fauna in Southeast Brazil. J. Coastal Res., 35: 431-439.         [ Links ]

Paula, A.F. & J.C. Creed. 2004. Two species of the coral Tubastraea (Cnidaria, Scleractinia) in Brazil: a case of accidental introduction. Bull. Mar. Sci., 74: 175-183.         [ Links ]

Pederson, J., R. Bullock, J. Carlton, J. Dijkstra, N. Dobroski, P. Dyrynda, R. Fisher, L. Harris, N. Hobbs, G. Lambert, E. Lazo-Wasem, A. Mathieson, M.P. Miglietta, J. Smith, J.S. III & M. Tyrrell. 2005. Rapid assessment survey of non-native and native marine species of floating dock communities, August 2003. MIT Sea Grant College Program, Cambridge, 40 pp.         [ Links ]

Pilsbry, H.A. 1916. The sessile barnacles (Cirripedia) contained in the collections of the US. National Museum: including a monograph of the American species. Bull. U.S. Natl. Mus., 93: 1-366.         [ Links ]

Pires-Vanin, A.M.S., T.N. Corbisier, E. Arasaki & A.M. Mõellmann 1997. Composição e distribuição espaço-temporal da fauna bêntica no Canal de São Sebastião. Relat. Téc. Inst. Oceanogr., 41: 29-46.         [ Links ]

Poupin, J., P.J.F. Davie & J.C. Cexus. 2005. A revision of the genus Pachygrapsus Randall, 1840 (Crustacea: Decapoda: Brachyura, Grapsidae), with special reference to the southwest Pacific species. Zootaxa, 1015: 1-66.         [ Links ]

Rathbun, M.J. 1894. Notes on the crabs of the family Inachidae in the U.S. National Museum. Proc. U.S. Natl. Mus., 17: 43-75.         [ Links ]

Rathbun, M.J. 1898. The Brachyura collected by the U.S. Fish Comission Steamer Albatross on the voyage from Norfolk, Virginia, to San Francisco, California, 1887-1888. Proc. U.S. Natl. Mus., 21: 567-616.         [ Links ]

Rathbun, M.J. 1900. The decapod and stomatopod Crustacea I: Results of the Branner-Agassiz Expedition to Brazil. Proc. Wash. Acad. Sci., 2: 133-156.         [ Links ]

Rios, E.C. 1975. Brazilian marine mollusks iconography. Fundação Universidade de Rio Grande, Rio Grande, 331 pp.         [ Links ]

Ritchie, J. 1909. Supplementary report on the hydroids of the Scottish National Antarctic Expedition. Trans. Roy. Soc. Edinb., 47: 65-101.         [ Links ]

Rocha, R.M. & N.Y.K. Bonnet 2009. Levantamento de ascídias (Tunicata: Ascidiacea) introduzidas no Arquipélago de Alcatrazes, SP. Iheringia, sér. Zool., 99: 27-35.         [ Links ]

Rocha, R.M. & L.P. Kremer. 2005. Introduced ascidians in Paranaguá Bay, Paraná, southern Brazil. Rev. Bras. Zool., 22(4): 1170-1184.         [ Links ]

Rocha R.M. & F. Monniot. 1995. Taxonomic and ecological notes on some Didemnum species (Ascidiacea, Didemnidae) for São Sebastião Channel, south-east Brazil. Rev. Bras. Biol., 55(4): 639-649.         [ Links ]

Rocha, R.M., L.P. Kremer & K.H. Fehlauer-Ale. 2012. Lack of COI variation for Clavelina oblonga (Tunicata, Ascidiacea) in Brazil: evidence for its human-mediated transportation? Aquat. Invas., 7(3): 419-424.         [ Links ]

Rocha R.M., T.R. Moreno & R. Metri. 2005. Ascídias (Tunicata, Ascidiacea) da Reserva Biológica Marinha do Arvoredo, Santa Catarina, Brasil. Rev. Bras. Zool., 22(2): 461-476.         [ Links ]

Rocha, R.M., L.P. Kremer, M.S. Baptista & R. Metri. 2009. Bivalve cultures provide habitat for exotic tunicates in southern Brazil. Aquat. Invas., 4: 195-205.         [ Links ]

Rodrigues, S.A. 1962. Algumas ascídias do litoral sul do Brasil. Bolm. Fac. Filos. Ciên. Let. Univ. São Paulo, sér. Zool., 24: 193-216.         [ Links ]

Rodrigues, S.A. & R.M. Rocha. 1993. Littoral compound ascidians (Tunicata) from São Sebastião, Estado de São Paulo, Brazil. Proc. Biol. Soc. Wash., 106(4): 728-739.         [ Links ]

Rodrigues, S.A., R.M. Rocha & T.M.C. Lotufo. 1998. Guia ilustrado para identificação das ascídias do Estado de São Paulo. FAPESP, São Paulo, 190 pp.         [ Links ]

Rossi, M.C.S. 2008. Contribuição ao conhecimento taxonômico dos poliquetas Sabelideos (Polychaeta: Sabellidae) da região sudeste do Brasil. Dissertação do grau de Mestre, Instituto de Biociências, Universidade de São Paulo, Brasil, 176 pp.         [ Links ]

Ruiz, G.M., P. Fofonoff, J.T. Carlton, M.J. Wonham & A.H. Hines. 2000. Invasions of coastal marine communities in North America: apparent patterns, processes, and biases. Ann. Rev. Ecol. Syst., 31: 481-531.         [ Links ]

Rullier, F. & L. Amoureux. 1979. Annélides Polychaètes. Ann. Inst. Océanogr., 55: 145-206.         [ Links ]

Salvador, L.B., O. Domaneschi, A.C.Z. Amaral, E.H. Morgado & S.A. Henriques. 1998. Malacofauna da região entremarés de praias da ilha de São Sebastião (São Paulo, Brasil). Rev. Bras. Zool., 15(4): 1013-1035.         [ Links ]

Santos, C.S.G. & T.M. Steiner. 2006. Familia Nereididae. In: A.C.Z. Amaral, A.E. Rizzo & E.P. Arruda (Orgs.). Manual de identificação dos invertebrados marinhos da região Sudeste-Sul do Brasil. Editora da Universidade de São Paulo, São Paulo, 1: 121-133.         [ Links ]

Schubart, C.D., J.A. Cuesta & D.L. Felder. 2005. Phylogeography of Pachygrapsus transversus (Gibbes, 1850): The effect of the American continent and the Atlantic Ocean as gene flow barriers and recognition of Pachygrapsus socius Stimpson 1871 as a valid species. Nauplius, 13(2): 99-113.         [ Links ]

Silveira, F.L. 1986. Aspectos da biologia de Lophogorgia punicea (Milne-Edwards & Haime, 1857) (Gorgo-nacea, Gorgoniidae) do canal de São Sebastião, Estado de São Paulo. Doctoral Thesis, Universidade de São Paulo, São Paulo, 211 pp.         [ Links ]

Silveira, N.G., R.C.C.L. Souza, F.C. Fernandes & E.P. Silva. 2006. Occurrence of Perna perna, Modiolus carvalhoi (Mollusca, Bivalvia, Mytilidae) and Megabalanus coccopoma (Crustacea, Cirripedia) off Areia Branca, Rio Grande do Norte State, Brazil. Biociências, 14(1): 89-90.         [ Links ]

Simone, L.R.L. & E.P. Gonçalves. 2006. Anatomical study on Myoforceps aristatus, an invasive boring bivalve in S.E. Brazilian coast (Mytilidae). Pap. Avuls. Zool., 46(6): 57-65.         [ Links ]

Smith, S.I. 1869. Notice of the Crustacea collected by Prof. C.F. Hartt on the coast of Brazil in 1867. Trans. Conn. Acad. Arts Sci., 2: 1-41.         [ Links ]

Souza, R.C.C.L., F.C. Fernandes & E.P. Silva. 2003. A study on the occurrence of the brown mussel Perna perna on the sambaquis of the Brazilian coast. Rev. Mus. Arqueol. Etnol. São Paulo, 13: 3-24.         [ Links ]

Souza. R.C.R. 1989. A Fauna dos bancos de areia de Phragmatopoma lapidosa Kinberg, 1867 (Annelida, Polychaeta) da região de Ubatuba, SP. Dissertação do grau de Mestre. Instituto de Biologia, Universidade Estadual de Campinas, 83 pp.         [ Links ]

Stachowicz, J.J., R.B. Whitlatch & R.W. Osman. 1999. Species diversity and invasion resistance in a marine ecosystem. Science, 286: 1577-1579.         [ Links ]

Stechow, E. 1919. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und anderer Gebiete, nebst Angaben íiber einige Kirchenpauer'sche Typen von Plumulariden. Zool. Jb., 42: 1-172.         [ Links ]

Tavares, M.D.S. 2004. On Halicarcinus planatus (Fabricius) (Brachyura, Hymenosomatidae) transported from Chile to Brazil along with the exotic oyster Crassostrea gigas (Thunberg). Nauplius, 11(1): 45-50.         [ Links ]

Taylor, P.D. & N. Monks. 1997. A new cheilostome bryozoan genus pseudoplanktonic on mollusks and algae. Invertebr. Biol., 116: 39-51.         [ Links ]

Tompsett, S., J. Porter & P.D. Taylor. 2009. Taxonomy of the fouling cheilostome bryozoans Schizoporella unicornis (Johnston) and Schizoporella errata (Waters). J. Nat. Hist., 43 (35-36): 2227-2243.         [ Links ]

Treadwell, A.L. 1932. Annelidos Polychetos novos da Ilha de São Sebastião. Rev. Mus. Paulista, 2(17): 121.         [ Links ]

Turon, X., I. Tarjuelo, S. Duran & M. Pascual. 2003. Characterizing invasion processes with genetic data: an Atlantic clade of Clavelina lepadiformis (Asci-diacea) introduced into Mediterranean Harbours. Hydrobiologia, 503: 29-35.         [ Links ]

Tyrrel, M.C. & J.E. Byers. 2007. Do artificial substrate favor nonindigenous fouling species over native species? J. Exp. Mar. Biol. Ecol., 342: 54-60.         [ Links ]

Van Name, W.G. 1945. The North and South American ascidians. Bull. Am. Mus. Nat. Hist., 84: 1-476.         [ Links ]

Vannucci, M. 1949. Hydrozoa do Brasil. Bolm. Fac. Filos. Cienc. Let. Univ. S. Paulo, 99(14): 216-266.         [ Links ]

Vannucci, M. 1950. Resultados científicos do Cruzeiro do Baependi e do Vega a Ilha da Trindade. Hydrozoa. Bolm Inst. Oceanogr., 1(1): 81-96.         [ Links ]

Vannucci, M. 1954. Hydrozoa e Scyphozoa existentes no Instituto Oceanográfico. II. Bolm Inst. Oceanogr., 5: 95-149.         [ Links ]

Vannucci, M. & W.J. Rees. 1961. A revision of the genus Bougainvillia (Anthomedusae). Bolm Inst. Oceanogr., 11: 57-100.         [ Links ]

Vannucci-Mendes, M. 1946. Hydroida Thecaphora do Brasil. Arq. Zool. Est. Sao Paulo, 4: 535-598.         [ Links ]

Veloso, V.G. & G.A.S. Melo. 1993. Taxonomia e distribuição da família Porcellanidae (Crustacea, Decapoda, Anomura) no litoral brasileiro. Iheringia, sér. Zool., 75: 171-186.         [ Links ]

Vieira, L.M., A.E. Migotto & J.E. Winston. 2010. Shallow-water species of Beania Johnston, 1840 (Bryozoa, Cheilostomata) from the tropical and subtropical western Atlantic. Zootaxa, 2550: 1-20.         [ Links ]

Winston, J.E. 2005. Re-description and revision of Smitt's Floridan Bryozoa in the collection of the Museum of Comparative Zoology, Harvard University. Virginia Mus. Nat. Hist. Mem., 7: 1-147.         [ Links ]

Young, P.S. 1989. Establishment of an Indo-Pacific barnacle in Brazil. Crustaceana, 56: 212-214.         [ Links ]

Young, P.S. 1994. Superfamily Balanoidea Leach (Cirripedia, Balanomorpha) from the Brazilian coast. Bolm. Mus. Nac., ser Zool., 356: 1-36.         [ Links ]

Young, P.S. 1998. Maxillopoda Thecostraca. In: P.S. Young (ed.). Catalogue of Crustacea from Brazil. Serie Livros 7, Mus. Nac. Rio de Janeiro, pp. 263-285.         [ Links ]

Zullo, V.A. 1992. Balanus trigonus Darwin (Cirripedia, Balaninae) in the Atlantic basin: an introduced species? Bull. Mar. Sci., 50: 66-74.         [ Links ]

 


Received: 16 May 2011; Accepted: 22 October 2012

 

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