Zootaxa 2568: 39–66 (2010) www.mapress.com / zootaxa/ ISSN 1175-5326 (print edition) Article Copyright © 2010 · Magnolia Press ZOOTAXA ISSN 1175-5334 (online edition) Biodiversity of free-living marine nematodes on the coast of Brazil: a review VIRÁG VENEKEY1,2, VERÔNICA G. FONSECA-GENEVOIS3 & PAULO J. P. SANTOS3 1 UFPA, ICB, Av. Augusto Corrêa, 01. Guamá – Belém – PA, Brazil. CEP: 66075-110. E-mail: venekey@ufpa.br UFPE, PPG Oceanografia 3 UFPE, CCB, Depto. Zoologia, R. Prof. Moraes Rêgo s/n. Cid. Universitária – Recife – PE, Brazil. CEP: 50670-920 2 Abstract The taxonomic richness of the marine Nematoda in coastal habitats of Brazil and similarities in generic composition among them are analysed. A complete faunal list is presented, containing 11 orders, 59 families, 294 genera and 231 species, among which 1 family, 10 genera and 87 species were discovered for the first time in Brazil. Seven habitats were considered (sandy beaches, estuaries, phytal, oceanic islands, beach rocks, salt works and artificial substrates): sandy beaches had the greatest generic richness (241), followed by estuaries (142) and the phytal environment (126). Taxonomic composition was similar to that of other coastal habitats sampled worldwide, with Chromadoridae and Xyalidae the most representative families. The three major habitats (beaches, rocky shores and estuaries), showed statistically significant differences in faunas. Estuaries were the most uniform in composition. Key words: nematode, composition, coastal environments, beaches, estuaries, rocky shores Introduction Phylum Nematoda has four to five thousand free-living marine species (Lorenzen 1994). This number seems very low considering the total number of known nematode species (26646) and even lower considering the estimates of up to one to one hundred million living species (Hugot et al. 2001, Lambshead 1993). Discrepancies regarding the total number of nematode species are due to recording a low number of valid species from limited locations (Coomans 2001). While interest in parasitic nematodes has been growing, attention to aquatic and terrestrial nematodes from different environments has been decreasing (Ferris 1994). Moreover, free-living nematodes are normally only a few millimetres long, making taxonomic studies difficult (Heip et al. 1982). In addition to the difficulties mentioned above, in Brazil there were no specialists working with the group until very recently. The first record of marine nematodes—a list of only three species (Alaimella truncata, Litotes minuta and Synonema brasiliense)—was made on the coast of the state of Bahia (northeastern region) by Cobb (1920). Taxonomic studies on nematodes in Brazil effectively began with the work of Dr. Sebastian Gerlach, an invited researcher to the University of São Paulo in the 1950's (Gerlach 1954; 1956a; 1956b; 1957a; 1957b). His publications, together with those for semi-limnetic environments by Meyl (1956 and 1957), were an important first step and resulted in records of 209 species, among which 106 were new to science. Taxonomic studies restarted only in the 1990s, resulting in new manuscripts (Netto & Gallucci 2003; Fonsêca-Genevois et al. 2004; Fonsêca-Genevois et al. 2006) as well as the first descriptions of new species and genera since those by Gerlach and Meyl in the 1950s (e.g. Venekey et al. 2005; Castro et al. 2006; Fonsêca-Genevois et al. 2009). Reviews of many other benthic animal groups have been carried out for the Brazilian coast: Mollusca (Rios 1994); Annelida Polychaeta (Amaral & Nonato 1996); Coral Reefs (Maida & Ferreira 1997); and Crustacea (Young 1998). However, most of the biodiversity data regarding Brazilian nematodes has remained in the gray literature: particularly theses or undergraduate papers. In this review paper we present an updated Accepted by M. Hodda: 3 Jul. 2010; published: 16 Aug. 2010 39 taxonomic list (genera and species), with revised nomenclature (modifications in species names, new taxonomic combinations, etc.) and a comparison of genus richness between different environments along the Brazilian coast. Material and methods Data collection. A uniform taxonomic list was constructed from genus and species records of nematodes from the Brazilian coast available to 2008 from PhD theses (Medeiros 1997; Netto 1999; Bezerra 2001; Castro 2003; Curvelo 2003; Da Rocha 2003; Maranhão 2003; Moellmann 2003; Pinto 2003; Venekey 2007), MSc theses (Rodrigues 2002; Nunes 2003; M.C. Silva 2004; N.R.R. Silva 2004; Lage 2005; Maria 2007; Thomas 2007; Gomes 2008), undergraduate papers (Silva 2001; Santos 2002; Fonsêca 2003; Miranda-Júnior 2006; Oliveira 2007; Pereira 2008) and other manuscripts (Cobb 1920; Gerlach 1954; Gerlach 1956a; Gerlach 1956b; Gerlach 1957a; Gerlach 1957b; Netto & Gallucci 2003; Esteves 2004; Fonsêca-Genevois et al. 2004; Fonsêca-Genevois et al. 2006). This list was also used for comparisons between different environments. Ecological studies with incomplete taxon lists (e.g. Kapusta et al. 2006; Pinto et al. 2006) were used only for their taxonomic lists. Abstracts published at scientific meetings were not considered. The studies by Meyl (1956 and 1957) were not considered either, as they were on a semi-limnetic environment (Rodrigo de Freitas Lake in Rio de Janeiro). The genus and species names were reviewed taking into account synonyms, nomenclatural changes and new combinations listed by Gerlach & Riemann 1973/1974, Abebe et al. (2007) and Deprez (2006). The taxonomic classification of De Ley & Blaxter (2004) was followed. The taxon lists were divided into seven basic habitats: sandy beaches, estuaries, phytal environment, oceanic islands, beach rocks, salt works and artificial substrates (aluminum plates and plastic plants). For statistical proposes, these were grouped into three major environments (beach, rocky shore and estuary). Studies addressing more than one environment were split between the different lists (e.g. Da Rocha 2003). The locations sampled along the Brazilian coast are presented in Fig.1 and Table 1, along with the site classification. Data analysis. The data set consisted of presence/absence data for genera. Initially, cluster analysis was carried out using the list in order to investigate similarities between different regions and habitats. Due to the limited number of investigations into certain environments, the studies were grouped into three major habitats only. The major habitats were compared with one-way analysis of similarities (one-way ANOSIM) using the Bray-Curtis index. Results were expressed in a non-metric multidimensional scaling biplot (nMDS). All tests were performed using the PRIMER (version 5.1.2) software program (Clarke & Warwick 1994), with the level of significance set at p < 0.05. Results By 2008, a total of 11 orders, 59 families, 294 genera and 231 species of nematodes were recorded for marine environments on the Brazilian coast. Among these, one family (Peresianidae), 10 genera (Assia, Bernardius, Conilia, Elzalia, Ingenia, Kosswigonema, Litotes, Manunema, Robbea and Synonema) and many species were discovered for the first time in Brazil. Of the orders, Enoplida had the highest number of families (14) and genera (63). Sandy beaches had the greatest taxon richness: 10 orders, 55 families and 241 genera, 71 found exclusively in this habitat (Fig. 2). In this environment only the order Triplonchida was not recorded from those known for the Brazilian coast (Enoplida, Triplonchida, Dorylaimida, Mononchida, Chromadorida, Desmodorida, Desmoscolecida, Monhysterida, Araeolaimida, Plectida). Estuaries were home to the second highest number of families (36) and genera (142), but not orders (7). Phytal and Oceanic Islands had 126 from 6 orders and 110 genera from 9 orders, respectively. Salt works and beach rocks had a similar number of 40 · Zootaxa 2568 © 2010 Magnolia Press VENEKEY ET AL. orders and genera. All environments had endemic genera, with the exception of artificial substrates. Only 15 of the 294 genera were common to all environments: Chromadorella, Daptonema, Desmodora, Dichromadora, Euchromadora, Marylynnia, Metachromadora, Metalinhomoeus, Microlaimus, Oncholaimus, Sabatieria, Spirinia, Terschellingia, Theristus and Viscosia. The families Chromadoridae and Xyalidae had the highest numbers of genera recorded in all environments. FIGURE 1. Map of Brazil showing locations that have been studied. Numbers refer to sites listed in Table 1. Cluster analysis revealed a group of all lists from estuarine environments with 40% similarity, but other environments did not form separate groups (Fig. 3). The one-way ANOSIM analysis detected significant differences (RGlobal = 0.339, p < 0.01) between the major environments. The pairwise comparisons also revealed differences between all environments: Beach x Rocky Shore (RGlobal = 0.291, p < 0.01), Beach x Estuary (RGlobal = 0.379, p < 0.01) and Rocky Shore x Estuary (RGlobal = 0.441, p < 0.01) (Fig. 4). Over time, there were few genera found early, then there was a pause after 1920 until new records appeared in the 1950s and then again a new pause until the 1990s (Fig. 5). About 40% of the genera were found before the 1960s. The 1990s contributed with an enlargment of 23% of the list and, in the last 8 years (since 2000), the other 37% were found. BIODIVERSITY OF MARINE NEMATODES IN BRAZIL Zootaxa 2568 © 2010 Magnolia Press · 41 TABLE 1. List of studies used. Geographical coordinates as listed by the original authors (N.I. = Not informed) and the environmental classification used here (N.C. = Not considered for environmental analysis) for the locations studied in Brazil. Study Coordinates Sampling Location Basic Habitat Major Habitat Latitude Longitude 1. Cobb (1920) N.I. N.I. Bahia Sandy Beach N.C. 2. Gerlach (1954) N.I. N.I. Lagoa Rodrigo de Freitas – RJ Estuary N.C. 3. Gerlach (1956a) N.I. N.I. Cananéia, Santos, São Sandy Beach Sebastião and Porto Fino (all in São Paulo State) Beach 4. Gerlach (1956b) (1) 7°00’S / (2) 7°45’S / (3) 9°05’S N.I. (1) Paraíba and (2,3) Pernambuco States Sandy Beach Beach 5. Gerlach (1957a) 25°00’S N.I. Cananéia – SP Estuary Estuary 6. Gerlach (1957b) (1) N.I. / (2) 20°00’S / (3) and (4) N.I. N.I. (1) Bahia, (2) Espírito Santo, (3) Rio de Janeiro and (4) São Paulo States Sandy Beach Beach 7. Medeiros (1997) 23°32’S 45°04’W Ilha Anchieta – SP Sandy Beach Beach 8. Netto (1999) 3°51’S 33°49’W Atol das Rocas – RN Oceanic Island Rocky Shore 9. Bezerra (2001) 8°02’S 34°51’W Olinda – PE Sandy Beach Beach 10. Silva (2001) 5°05’S 36°16’W Galinhos – RN N.C. N.C. 11. Rodrigues (2002) 8°04’S 34°52’W Recife – PE Estuary Estuary 12. Santos (2002) 23°44’S 42°00’W Arraial do Cabo – RJ N.C. N.C. 13. Castro (2003) 8°04’S 44°55’W Recife – PE Estuary Estuary 14. Curvelo (2003) 23°22’S 44°55’W Picinguaba Bay – SP Sandy Beach Beach 15. Fonseca (2003) 28º12’S 48º38’W Laguna – SC Estuary Estuary 16. Maranhão (2003) 8°33’S 35°00’W Porto de Galinhas - PE Beach Rocks Rocky Shore 17. Moellmann (2003) N.I. N.I. Guarujá – SP and Peruíbe – SP Sandy Beach Beach 18. Netto & Gallucci (2003) 27°29’S 48°30’W Santa Catarina Island – SC Estuary Estuary 19. Nunes (2003) 23°44’S 42°00’W Arraial do Cabo – RJ Artificial Rocky Shore 20. Pinto (2003) 32°02’S 52°07’W Rio Grande – RS Estuary Estuary 21. Da Rocha (2003) (1) 7°33’S 8°55’S / (2) 23°44’ S (1)34°49’W - (1) Pernambuco (2) Arraial do Cabo –RJ 35°11’W / (2) 42°00’W Sandy Beach / Phytal Beach / Rocky Shore 22. Esteves (2004) 22°55’S 43°05’W Coroa Grande – RJ Estuary Estuary 23. Fonsêca-Genevois et al. (2004) 23°44’S 42°00’W Arraial do Cabo – RJ Sandy Beach Beach 24. Silva (a) (2004) 7°49’S 34°50’W Igarassú – PE Estuary Estuary 25. Silva (b) (2004) 5°05’S 36°16’W Galinhos – RN Salt Work Estuary 26. Lage (2005) 23°44’S 42°00’W Arraial do Cabo – RJ Phytal Rocky Shore 27. Fonsêca-Genevois et al. (2006) 23°44’S 42°00’W Arraial do Cabo – RJ Artificial Rocky Shore 28. Miranda-Junior (2006) 0°55’N 29°21’W Saint Peter and Saint Paul Rocks Oceanic Island Rocky Shore / / Artificial Rocky Shore 29. Maria (2007) 22°24’S 42°33’W Guanabara Bay – RJ Sandy Beach Beach 30. Oliveira (2007) 8°25’S 34°55’W Cupe – PE Phytal Rocky Shore 31. Thomas (2007) 25º30’S 48º25’W Baía de Paranaguá-PR Estuary Estuary 32. Venekey (2007) 8°46’S 34°59’W Tamandaré – PE Sandy Beach Beach 33. Gomes (2008) 1º49’N 50º34’W Piratuba-AP Estuary Estuary 34. Pereira (2008) 8º32’S 35º00’W Maracaípe-PE Sandy Beach Beach 42 · Zootaxa 2568 © 2010 Magnolia Press VENEKEY ET AL. FIGURE 2. Number of orders, families, total and endemic genera found in the habitats (SB = Sandy Beaches; ES = Estuaries; PH = Phytal, OI = Oceanic Islands; BR = Beach Rocks; SW = Salt Works and AR = Artificial Substrate) . FIGURE 3. Cluster analysis of the studies. “]” represents the studies from estuaries. BIODIVERSITY OF MARINE NEMATODES IN BRAZIL Zootaxa 2568 © 2010 Magnolia Press · 43 FIGURE 4. Non-metric scaling plot of the taxonomic lists grouped in three major habitats: beaches, estuaries and rocky shores. FIGURE 5. Number of genera recorded for the Brazilian coast since 1920. Discussion Nematoda is the most abundant metazoan group in nearly all environments (McIntyre 1971). Ecological studies initially considered nematodes to be a single taxonomic or functional unit, but recently they have been 44 · Zootaxa 2568 © 2010 Magnolia Press VENEKEY ET AL. recognized as a very heterogeneous group occupying different trophic positions in marine ecosystems (Heip et al. 1982). Ecological studies including taxonomic detail only become possible as nematode taxonomy developed at the end of the 19th century throught the work of Cobb (e.g. 1890; 1891 and 1894) and De Man (e.g. 1876; 1888). The number of taxonomists increased with the evolution of microscopy and 42 manuscripts were published by 50 marine nematologists between 1971 and 1975, describing 111 new taxa each year (Gerlach 1980). From the 1980’s onward, a decline in classical taxonomy was observed for the majority of animal groups, possibly due to development of molecular techniques and taxonomy becoming more stagnant and attracting fewer scientists (Wilson 2000). The same occurred with Nematoda, mainly due to difficulties in identification using classical methods, which demand considerable experience and well-equipped laboratories (Coomans 2001). In Brazil, taxonomic research on marine nematodes began significantly only in the second half of the 1950s, with studies by Gerlach (1956a; 1956b; 1957a; 1957b). This was a very late start in comparison to other countries. The increasing records of genera and species in this country at the end of the 20th century indicate a “taxonomic flourishing” - an opposite behavior to the tendency seen in the rest of the world. This increase may be the consequence of the training of Brazilian nematologists by the Postgraduate International Nematology Course in Belgium (http://www.pinc.ugent.be) and cooperation with other groups through projects such as the “Darwin Project” (England - http://web.pml.ac.uk/nematode). In general terms, the drafting of pictorial keys, such as the “Illustrated Guide” by Tarjan (1980) and the “Free-Living Marine Nematodes”, divided into three volumes [“Enoplids” (Platt & Warwick 1983), “Chromadorids” (Platt & Warwick 1988) and “Monhysterids” (Warwick et al. 1998)] has been helpful for nematode identification worldwide. Recently, an Internet site was launched by the Marine Section of the University of Ghent, with original descriptions and data on Nematoda species: the “Nemys” database (Deprez 2006). Prior to pictorial key publications, complete international lists were drafted by Stiles & Hassal (1905), Baylis & Daubney (1926), Hope & Murphy (1972) and the best known Bremerhaven Checklist by Gerlach & Riemann (1973, 1974). Regional taxonomic lists similar to the one presented here were also published for North America (Cobb 1920; Wieser & Hopper 1967), the North Sea (Schuurmans Stekhoven 1935), the Baltic Sea (Schneider 1939), Chile (Wieser 1953; Wieser 1954; Wieser 1956) and the Antarctic continent (Allgén 1959). Knowledge on Nematoda biogeography is fragmented due to the lack of sampling in many regions. Some free-living and parasitic species seem to have worldwide distribution, but there are controversies regarding the validity of many identifications or the possibility that these species are complexes of cryptic species (Coomans 2001). Doubts persist because there are only a few studies on nematode dispersion (Gerlach 1977; Coomans et al. 1985; Winoto Suatmadji et al. 1988; Eyualem & Coomans 1995). There are about 450 marine nematode genera mentioned in the literature (Heip et al. 1982) and, according to the present study, 294 (approximately 65%) have been found in one or more locations on the coast of Brazil. This seems a significant number but is actually not high if we consider environmental heterogeneity in Brazil and the approximately 8000 km of coastline. Comparatively, on the coast of Great Britain and Ireland, corresponding to approximately 4500 km, 170 genera have been found (Warwick et al. 1998). Fleeger and Decho (1987) pointed out the influence of sampling effort on the determination of biodiversity in meiofauna groups. We may therefore suppose that our results are an underestimation of actual nematode diversity. Thus, as the majority of the coast continues without studies, the real diversity of Nematoda in Brazil remains unknown and cannot be compared with better known regions in the world. Another fact that merits attention regarding the taxonomic list of Brazil is the quantity of taxa that have been changed nomenclature (mostly changed combinations) since their first records. At species level, about 21,6% (50 of 231) were first recorded under a different name, sometimes within a different genus and, in a few cases, even within a different family. Nomenclature and description of species should be carried out in such a way that two taxonomists working in different locations could reach the same conclusion when studying the same species (Dubois 2005). Descriptions of a same species by different names are currently much rarer due to the advances in communication since the late 20th century. The exchange and checking of BIODIVERSITY OF MARINE NEMATODES IN BRAZIL Zootaxa 2568 © 2010 Magnolia Press · 45 species type material (Dubois & Nemésio 2007) have also been made easier. Nevertheless, the search for complete information available on species descriptions and records is strongly recommended, even if there are recent regional or taxonomic reviews. For Nematoda, we suggest special attention for the species described prior to the creation of the “Nemys” database (Deprez 2006) or the publication of the Bremerhaven Checklist (Gerlach & Riemann 1973, 1974). Sandy beaches were the most diverse habitats with the regard to the number of taxa (10 orders, 55 families and 241 genera). This result was expected, as the majority of studies (a total of 12) were carried out in such a habitat. Estuaries, with a similar number of studies (11), had the second highest number of genera (142) and families (36), but not orders (7). This is in agreement with many studies from very early on (Remane 1933), identifying the estuarine benthic community, as less diverse compared to truly marine or freshwater environments. Furthermore, taxonomic lists for estuarine habitats were very similar to one another, as demonstrated by the Cluster analysis, and they were also quite distinct from sandy beaches or rocky shore environments, as confirmed by ANOSIM results. The above results may be explained by the greater homogeneity of estuarine environments and the consequent lower number of microhabitats when compared with sandy beaches or rocky shores. In this regard, Flemming and Fricke (1983) point out that beach sediments are neither homogenous nor uniform in terms of distribution, even on very small scales; therefore, their configuration permits the formation of many different micro-habitats. Other studies, such as those by Wieser (1959) and Hopper and Meyer (1967), also described greater diversity in sandy environments, ascribed to more niches and heterogeneity with lower siltclay contents. All taxonomic works developed in Brazilian estuaries with nematodes sampled exclusively sediment, for this reason we can agree with the considerations of the authors cited here. Nevertheless, we should consider our results with caution as the incorporation of environments such as the phytal and artificial substrates within the rocky shore group may have made such environments more heterogeneous artifactually. It was a surprise to find oceanic islands having the second highest number of orders (9) along with a considerable number of families (29) and genera (110). This may be explained by the high sampling effort that is generally employed in locations with particularly difficult access, such as the Saint Peter and Saint Paul Rocks and the Rocas Atoll, located in the middle of the South Atlantic Ocean. The taxonomic lists for salt works and beach rocks are the results of a single study for each habitat. Taking this fact into account, the number of genera recorded (75 and 76, respectively) can be considered quite surprising, as some are endemic to these environments. The same observation may be made for phytal habitats (only 3 studies), with a list of 127 genera, including five endemic genera. Artificial substrate environments (aluminum plates and plastic plants) did not add any genus to the Brazilian taxonomic list. This result could have been expected, as the natural locations where the experiments with these artificial substrates were conducted had been studied in previous investigations. Finally, considering the genus composition in all environments, Xyalidae or Chromadoridae were always the families with the highest number of genera, which could be related to the fact that they have more described genera (Lorenzen 1994). The limited number of studies in many environments hinders comparisons of taxonomic lists. Although 15 genera were found to be common to the seven environments considered here, it would be hasty to conclude anything. No ecological or morphological pattern can be distinguished: they do not have the same buccal cavity; are from different orders; and inhabit different sediment types. Nevertheless, these 15 genera are frequently recorded in studies on marine Nematoda. Furthermore, the fact that approximately 25% of the genera were found exclusively in one study also demonstrates the small amount of information. The generic composition of each of the habitats considered was similar to that from other places with similar characteristics. Considering results from the habitats with the largest number of studies, we did not detect a single genus common to all 12 locations studied on sandy beaches. The genus Metachromadora was the most common - found in 11 studies (absent only in Bezerra 2001). This genus occurs frequently in marine benthos (e.g. Ott 1972; Blome 1983; Tita et al. 1999; Nicholas 2001). In estuaries, Anoplostoma, Daptonema, Metachromadora, Terschellingia, Theristus and Viscosia were present in all eleven studies checked. Among these, only Anoplostoma is in the group of 15 genera found in all habitats along the Brazilian coast, but it is 46 · Zootaxa 2568 © 2010 Magnolia Press VENEKEY ET AL. also frequently recorded in estuaries (i.e. Bouwman 1983; Pinto et al. 2006). All records support the opinion of Heip et al. (1985) that there are no genera that are typical of hypersaline or estuarine environments. In terms of taxonomic composition, the studies on the Brazilian coast have contributed one new family, 10 new genera and 87 new species to the list of marine nematodes. Among these, Assia, Bernardius, Conilia, Ingenia, Kosswigonema and Litotes have not been recorded in other regions around the world, whereas Elzalia, Manunema, Robbea and Synonema have been found in different environments outside Brazil (Gerlach & Riemann 1973, 1974; Deprez 2006). Considering all generic records in Brazil, 3.5% of the genera were unknown to science prior to the studies compiled here. The number of new taxa is not surprising. Knowledge on the biodiversity of nematodes worldwide is restricted mostly to Europe and North America and about 10% of all species were described for Great Britain and Ireland (Lambshead 2003). The description of new taxa as previously unknown regions are sampled, as in the case of Brazil, demonstrates the need for further studies and the importance of traditional taxonomy. The major problem for continuing descriptions of new species or higher taxon is the urgency in renewing interest in traditional taxonomy. Coomans (2001) considered the decline in the number of active taxonomists alarming and it continues to decrease. In a short time, even if interest is piqued among young researchers, there will not be anyone able to transfer the necessary knowledge. This observation has led to the recent renewal in Nematode taxonomy, particularly in Brazil. During the Thirteenth International Meiofauna Conference meeting in 2007, about 25% of all abstracts were on taxonomic studies, which is good news for this neglected scientific field. Acknowledgements We acknowledge a postgraduate grant to V.V. from CAPES and a research grant to P.J.P.S. from CNPq, both provided by the Brazilian government. We are grateful to Mr. Richard Boike for the English revision and to Viviane Lira for the help with the figures. 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PHYLUM NEMATODA CLASS ENOPLEA SUBCLASS ENOPLIA ORDER ENOPLIDA Suborder Enoplina Superfamily Enoploidea Family Enoplidae Enoplus Dujardin, 1845 Enoplus schulzi Gerlach, 1952 Fenestrolaimus Filipjev, 1927 Unidentified species Family Thoracostomopsidae Enoploides Saweljev, 1912 Enoploides longispiculosus Vitiello, 1967 Enoploides vectis (Gerlach, 1957) Wieser & Hopper, 1967* [Enoploides brunette Gerlach, 1957] Enoplolaimus De Man, 1893 Enoplolaimus connexus Wieser, 1953 Enoplolaimus distortus Gerlach, 1957* Epacanthion Wieser, 1953 Epacanthion cf. buetschli (Southern, 1914) Wieser, 1953 Mesacanthion Filipjev, 1927 Mesacanthion hirsutum Gerlach, 1953 Mesacanthion longispiculum Gerlach, 1954 Mesacanthion proximum Gerlach, 1957* Mesacanthion rigens Gerlach, 1957* Mesacanthoides Wieser, 1953 Unidentified species Oxyonchus Filipjev, 1927 Oxyonchus dentatus (Ditlevsen, 1919) Filipjev, 1927 Paramesacanthion Wieser, 1953 Unidentified species Trileptium Cobb, 1933 Trileptium stylum Gerlach, 1956 Trileptium subterraneum (Gerlach, 1952) Cobb, 1933 Family Anoplostomatidae Anoplostoma Bütschli, 1874 Anoplostoma hirtum Gerlach, 1956* Anoplostoma subulatum Gerlach, 1957* Anoplostoma viviparum (Bastian, 1865) Butschli, 1874 Chaetonema Filipjev, 1927 Chaetonema canellatum Gerlach, 1956* Family Phanodermatidae Crenopharyncx Filipjev, 1934 Unidentified species Phanoderma Bastian, 1865 BIODIVERSITY OF MARINE NEMATODES IN BRAZIL Zootaxa 2568 © 2010 Magnolia Press · 51 Phanoderma tenuicaudum Allgén, 1951 Phanodermella Kreis, 1928 Unidentified species Phanodermopsis Ditlevsen, 1926 Phanodermopsis necta Gerlach, 1957* Family Anticomidae Anticoma Bastian, 1865 Anticoma aff. eberthi Bastian, 1865 Anticomopsis Micoletzky, 1930 Unidentified species Odontanticoma Platonova, 1976 Unidentified species Suborder Trefusiina Superfamily Trefusioidea Family Trefusiidae Cytolaimium Cobb, 1920 Cytolaimium conicum Gerlach, 1957* [Trefusia conica Gerlach, 1957] Rhabdocoma Cobb, 1920 Unidentified species Trefusia De Man, 1893 Unidentified species Trefusialaimus Riemann, 1974 Unidentified species Family Lauratonematidae Lauratonema Gerlach, 1953 Lauratonema hospitum Gerlach, 1956* Lauratonema reductom Gerlach, 1953 Family Xenellidae Xenella Cobb, 1920 Unidentified species Suborder Oncholaimina Superfamily Oncholaimoidea Family Oncholaimidae Adoncholaimus Filipjev, 1918 Unidentified species Metaparoncholaimus Filipjev, 1918 Unidentified species Metoncholaimus Filipjev, 1918 Unidentified species Meyersia Hopper, 1967 Unidentified species Oncholaimellus De Man, 1890 Oncholaimellus carlbergi Allgén, 1947 Oncholaimus Dujardin, 1845 [Oncholaimium Cobb, 1930] Oncholaimus cavatus Gerlach, 1956 Oncholaimus cobbi (Kreis, 1932) Rachor, 1969 [Oncholaimium cobbi Kreis, 1932] Oncholaimus gladius Gerlach, 1956 Oncholaimus cf. longispiculosus Gerlach, 1955 Oncholaimus manilius Gerlach, 1957* Oncholaimus oxyuris Ditlevsen, 1911 52 · Zootaxa 2568 © 2010 Magnolia Press VENEKEY ET AL. Oncholaimus rapax Kreis, 1932 Pontonema Leidy, 1855 Pontonema ardens Gerlach, 1957* Prooncholaimus Micoletzky, 1924 Prooncholaimus aransas Chitwood, 1951 Viscosia De Man, 1890 Viscosia glabra (Bastian, 1865) De Man, 1890 Viscosia keiensis Kreis, 1932 Family Enchelidiidae Bathyeurystomina Lambshead & Platt, 1979 Unidentified species Belbolla Andrássy, 1973 Unidentified species Bernardius Fonsêca-Genevois, Smol, Decraemer & Venekey, 2009* Bernardius lineatus Fonsêca-Genevois, Smol, Decraemer & Venekey, 2009* Calyptronema Marion, 1870 Calyptronema cf. acuminatum Wieser, 1953 Calyptronema cf. denticulatum Micoletzky, 1930 Calyptronema pigmentatum Gerlach, 1957* Calyptronema cf. setifer (Gerlach, 1953) Wieser, 1953 Eurystomina Filipjev, 1921 Eurystomina sawayai Gerlach, 1956* Eurystomina terrícola De Man, 1907 Pareurystomina Micoletzky, 1930 Unidentified species Polygastrophora De Man, 1922 Polygastrophora septembulba Gerlach, 1954 Symplocostoma Bastian, 1865 Unidentified species Suborder Ironina Superfamily Ironoidea Family Ironidae Conilia Gerlach, 1956* Conilia divina Gerlach, 1956* Dolicholaimus De Man, 1888 Unidentified species Syringolaimus De Man, 1888 Syringolaimus cf. striatocaudatus De Man, 1888 Thalassironus De Man, 1889 Unidentified species Trissonchulus Cobb, 1920 Trissonchulus oceanus Cobb, 1920 [Dolicholaimus nudus Stekhoven, 1943] Family Leptosomatidae Barbonema Filipjev, 1927 Barbonema flagrum Gerlach, 1957* Cylicolaimus De Man, 1889 Unidentified species Leptosomatum Bastian, 1865 Unidentified species Platycoma Cobb, 1894 Platycoma curiosa (Gerlach, 1955) Gerlach, 1962 [Platycomopsis curiosus Gerlach, 1955] Platycomopsis Ditlevsen, 1926 Unidentified species Thoracostoma Marion, 1870 BIODIVERSITY OF MARINE NEMATODES IN BRAZIL Zootaxa 2568 © 2010 Magnolia Press · 53 Unidentified species Family Oxystominidae Halalaimus De Man, 1888 Halalaimus ciliocaudatus Allgén, 1932 Halalaimus gracilis De Man, 1888 Halalaimus longicaudata (Filipjev, 1927) Schneider, 1939 Halalaimus longicollis Allgén, 1932 Litinium Cobb, 1920 Unidentified species Nemanema Cobb, 1920 Unidentified species Oxystomina Filipjev, 1921 Oxystomina acuta Gerlach, 1957* Oxystomina affinis Gerlach, 1956 Oxystomina alpha Chitwood, 1936 Paroxystomina Micoletzky, 1924 Unidentified species Thalassoalaimus De Man, 1893 Thalassoalaimus brasiliensis Gerlach, 1956* Wieseria Gerlach, 1956 Unidentified species Suborder Tripyloidina Superfamily Tripyloidoidea Family Tripyloididae Bathylaimus Cobb, 1894 Bathylaimus australis Cobb, 1894 [Bathylaimus assimilis De Man, 1922] Bathylaimus capacosus Hopper, 1962 Ingenia Gerlach, 1957* Ingenia mirabilis Gerlach, 1957* Tripyloides De Man, 1886 [Nannonchus Cobb, 1913] Tripyloides amazonicus (Gerlach, 1957) Riemann, 1970* [Nannonchus amazonicus Gerlach, 1957] Tripyloides granulatus (Cobb, 1913) Wieser, 1956 [Nannonchus granulatus Cobb, 1913] ORDER TRIPLONCHIDA Suborder Tobrilina Superfamily Prismatolaimoidea Family Prismatolaimidae Prismatolaimus De Man, 1880 Unidentified species Superfamily Tobriloidea Família Rhabdodemaniidae Rhabdodemania Baylis & Daubney, 1926 Unidentified species SUBCLASS DORYLAIMIA ORDER DORYLAIMIA Suborder Dorylaimina Superfamily Tylencholaimoidea 54 · Zootaxa 2568 © 2010 Magnolia Press VENEKEY ET AL. Family Tylencholaimidae Tylencholaimus De Man, 1876 Unidentified species Family Leptonchidae Leptonchus Cobb, 1920 Unidentified species Suborder Nygolaimina Superfamily Nygolaimoidea Family Nygolaimidae Nygolaimus Cobb, 1913 Unidentified species Superfamily Dorylaimoidea Family Dorylaimidae Mesodorylaimus Andrássy, 1959 Mesodorylaimus parasubtilis (Meyl, 1957) Andrássy, 1959* [Dorylaimus parasubtilis Meyl, 1957] Mesodorylaimus paulbuchneri (Meyl, 1956) Andrássy, 1959* [Dorylaimus paulbuchneri Meyl, 1956] Mesodorylaimus sveltus (Meyl, 1957) Andrássy, 1959* [Dorylaimus sveltus Meyl, 1957] Prodorylaimus Andrássy, 1988 Prodorylaimus brasiliensis (Meyl, 1956) Andrássy, 1959* [Dorylaimus brasiliensis Meyl, 1956] Prodorylaimus rionensis (Gerlach, 1954) Andrássy, 1959* [Dorylaimus rionensis Gerlach, 1954] Family Qudsianematidae Allodorylaimus Andrássy, 1986 Allodorylaimus santosi (Meyl, 1957) Andrássy, 1986* [Dorylaimus santosi Meyl, 1957] Family Aporcelaimidae Aporcelaimellus Heyns, 1965 Aporcelaimellus gerlachi (Meyl, 1956) Heyns, 1965 [Aporcelaimus gerlachi Meyl, 1956] Aporcelaimellus paraconicaudatus (Meyl, 1956) Heyns, 1965 [Aporcelaimus paraconicaudatus Meyl, 1956] Aporcelaimellus vanderlaani (Meyl, 1957) Heyns, 1965 [Aporcelaimus vanderlaani Meyl, 1957] ORDER MONONCHIDA Suborder Mononchina Superfamily Mononchoidea Family Mononchidae Unidentified genus and species CLASS CHROMADOREA SUBCLASS CHROMADORIA ORDER CHROMADORIDA Suborder Chromadorina Superfamily Chromadoroidea Family Chromadoridae Acantholaimus Allgén, 1933 Unidentified species Actinonema Cobb, 1920 BIODIVERSITY OF MARINE NEMATODES IN BRAZIL Zootaxa 2568 © 2010 Magnolia Press · 55 Actinonema pachydermatum Lorenzen, 1972 Atrochromadora Wieser, 1959 Unidentified species Chromadora Bastian, 1865 Chromadora aff. nudicapitata Bastian, 1865 Chromadorella Filipjev, 1918 Chromadorella macris (Gerlach, 1956) Lorenzen, 1972* [Trichromadora macris Gerlach, 1956] Chromadorina Filipjev, 1918 Unidentified species Chromadorita Filipjev, 1922 [Allgéniela Strand, 1934] Chromadorita cf. leuckarti (De Man, 1876) Chromadorita tenuis (Schneider, 1906) [Allgéniela tennuis Schneider, 1906] Dichromadora Kreis, 1929 Dichromadora geophila (De Man, 1876) Kreis, 1929 Endeolophus Boucher, 1976 Unidentified species Euchromadora De Man, 1886 Unidentified species Graphonema Cobb, 1898 Unidentified species Hypodontolaimus De Man, 1886 Hypodontolaimus pumilio Gerlach, 1956* Innocuonema Inglis, 1969 Unidentified species Karkinochromadora Blome, 1982 Unidentified species Neochromadora Micoletzky, 1924 Neochromadora bonita Gerlach, 1956* Nygmatonchus Cobb, 1933 Unidentified species Parachromadorita Blome, 1974 Unidentified species Parapinnanema Inglis, 1969 Unidentified species Prochromadora Filipjev, 1922 Unidentified species Prochromadorella Micoletzky, 1924 [Trichromadora Kreis, 1929] Prochromadorella paramucrodonta (Allgén, 1929) Wieser, 1951 Prochromadorella spinosa Gerlach, 1957* Prochromadorella tenuicaudata Gerlach, 1954 Ptycholaimellus Cobb, 1920 Unidentified species Rhips Cobb, 1920 Rhips ornate Cobb, 1920 Spiliphera Bastian, 1865 Unidentified species Spilophorella Filipjev, 1917 Spilophorella meyerabichi Gerlach, 1955 Steineridora Inglis, 1969 Unidentified species Trochamus Boucher & Bovée, 1972 Unidentified species Family Ethmolaimidae Ethmolaimus De Man, 1880 Ethmolaimus cf. multipapillatus Paramonov, 1926 Paraethmolaimus Jensen, 1994 Unidentified species 56 · Zootaxa 2568 © 2010 Magnolia Press VENEKEY ET AL. Family Neotonchidae Comesa Gerlach, 1956 Unidentified species Filitonchus Platt, 1982 Unidentified species Gomphionchus Platt, 1982 Unidentified species Gomphionema Wieser & Hopper, 1966 Gomphionema compactum (Gerlach, 1957) Wieser & Hopper, 1966* [Neotonchus compactus Gerlach, 1957] Gomphionema fellatur Wieser & Hopper, 1966 Nannolaimus Cobb, 1920 Nannolaimus complicatus Gerlach, 1957* Neotonchus Cobb, 1933 Unidentified species Family Cyatholaimidae Acanthonchus Cobb, 1920 Acanthonchus viviparous Cobb, 1920 Cyatholaimus Bastian, 1865 Unidentified species Kraspedonema Gerlach, 1954 Unidentified species Longicyatholaimus Micoletzky, 1924 Unidentified species Marylynnia Hopper, 1977 Marylynnia choanolaimoides (Stekhoven, 1942) Hopper, 1977 [Longicyatholaimus cf. choanolaimoides Stekhoven, 1942] Metacyatholaimus Stekhoven, 1942 Unidentified species Minolaimus Vitiello, 1970 Unidentified species Nannolaimoides Ott, 1972 Unidentified species Paracanthonchus Micoletzky, 1924 Paracanthonchus batidus Gerlach, 1957* Paracanthonchus cochlearis Gerlach, 1957* Paracanthonchus digitatus Gerlach, 1957* Paracyatholaimoides Gerlach, 1953 Paracyatholaimoides multispiralis Gerlach, 1953 Paracyatholaimoides serpens Gerlach, 1957* Paracyatholaimus Micoletzky, 1922 Paracyatholaimus paucipapillatus Gerlach, 1955 Paracyatholaimus separatus Wieser, 1954 Paracyatholaimus ternus Wieser, 1954 Paracyatholaimus vitraeus Gerlach, 1957* Paralongicyatholaimus Stekhoven, 1942 Unidentified species Pomponema Cobb, 1917 Pomponema tautraense (Allgén, 1933) Lorenzen, 1972 [Longicyatholaimus tautraensis Allgén, 1933] Praeacanthonchus Micoletzky, 1924 Unidentified species Family Selachinematidae Cheironchus Cobb, 1917 Unidentified species Choanolaimus De Man, 1880 Unidentified species Choniolaimus Ditlevsen, 1918 Unidentified species BIODIVERSITY OF MARINE NEMATODES IN BRAZIL Zootaxa 2568 © 2010 Magnolia Press · 57 Demonema Cobb, 1894 Unidentified species Gammanema Cobb, 1920 Unidentified species Halichoanolaimus De Man, 1886 Halichoanolaimus chordiurus Gerlach, 1955 Halichoanolaimus quattordecimpapillatus Chitwood, 1951 Kosswigonema Gerlach, 1964* Kosswigonema acanthum (Gerlach, 1957) Gerlach, 1964* [Selachinema acanthum Gerlach, 1957] Latronema Wieser, 1954 Latronema botulum Gerlach, 1956* Latronema orcinum (Gerlach, 1952) Wieser, 1954 Richtersia Steiner, 1916 Richtersia imparis Gerlach, 1956* Synonchiella Cobb, 1933 Synonchiella forceps (Gerlach, 1957) Gerlach, 1964* [Trogolaimus forceps Gerlach, 1957] Synonchium Cobb, 1920 Synonchium depressum Gerlach, 1954 Synonchium obtusum Cobb, 1920 ORDER DESMODORIDA Superfamily Desmodoroidea Family Desmodoridae Acanthopharynx Marion, 1870 Unidentified species Catanema Cobb, 1920 Unidentified species Chromaspirinia Filipjev, 1918 Chromaspirinia rabosa (Gerlach, 1956) Gerlach, 1963 [Desmodora rabosa Gerlach, 1956] Croconema Cobb, 1920 Unidentified species Desmodora De Man, 1889 [Micromicron Cobb, 1920] Desmodora aff. granulata Vincx & Gourbault, 1989 Desmodorella Cobb, 1933 Echinodesmodora Blome, 1982 Unidentified species Eubostrichus Greef, 1869 Unidentified species Leptonemella Cobb, 1920 Unidentified species Metachromadora Filipjev, 1918 [Ichthyodesmodora Chitwood, 1951] Metachromadora chandleri (Chitwood, 1951) Timm, 1961 [Ichthyodesmodora chandleri Chitwood, 1951] Metachromadora clavata Gerlach, 1957* Metachromadora onyxoides Chitwood, 1936 Metachromadora pneumática Gerlach, 1956* Metachromadora aff. remanei Gerlach, 1951 Metachromadora aff. serrata Gerlach, 1963 Metachromadora spectans Gerlach, 1957* Molgolaimus Ditlevsen, 1921 Unidentified species Onyx Cobb, 1891 Unidentified species Paradesmodora Stekhoven, 1950 Paradesmodora campbelli (Allgen, 1932) Gerlach, 1963 [Spirinia campbelli Allgén, 1962] Perspiria Wieser & Hopper, 1967 Unidentified species Polysigma Cobb, 1920 58 · Zootaxa 2568 © 2010 Magnolia Press VENEKEY ET AL. Unidentified species Pseudochromadora Daday, 1889 Pseudochromadora cazca Gerlach, 1956 [Desmodora cazca Gerlach, 1956] Pseudochromadora incubans Gourbault & Vincx, 1990 Pseudochromadora luticola (Timm, 1952) Andrássy, 1959 [Micromicron cephalatum Cobb, 1920] Robbea Gerlach, 1956* Robbea caelestis Gerlach, 1956* Sigmophoranema Hope & Murphy, 1972 Unidentified species Spirinia Gerlach, 1963 Spirinia elongata Castro, Bezerra, Silva & Fonsêca-Genevois, 2006* Spirinia parasitifera Bastian, 1865 Spirinia septentrionalis Cobb, 1914 Stygodesmodora Blome, 1982 Unidentified species Family Epsilonematidae Bathyepsilonema Steiner, 1931 Bathyepsilonema bahiae Gerlach, 1957* Epsilonema Steiner, 1927 Epsilonema espeeli Verschelde & Vincx, 1994 Metepsilonema Steiner, 1927 Unidentified species Perepsilonema Lorenzen, 1973 Perepsilonema kellyae Gourbault & Decraemer, 1988 Family Draconematidae Dracognomus Allen & Noffsinger, 1978 Unidentified species Dracograllus Allen & Noffsinger, 1978 Unidentified species Draconema Cobb, 1913 [Drepanonema Cobb, 1933] Draconema brasiliensis Venekey, Lage & Fonsêca-Genevois, 2005* Draconema fluminensis Venekey, Lage & Fonsêca-Genevois, 2005* Notochaetosoma Irwin-Smith, 1918 Unidentified species Prochaetosoma Micoletzky, 1922 Prochaetosoma lugubre (Gerlach, 1957) Allen & Noffsinger, 1978* [Drepanonema lugubre Gerlach, 1957] Superfamily Microlaimoidea Family Microlaimidae Aponema Jensen, 1978 Unidentified species Bolbolaimus Cobb, 1920 Unidentified species Calomicrolaimus Lorenzen, 1971 Calomicrolaimus formosus Jensen, 1978 Microlaimus De Man, 1880 Microlaimus capillaris Gerlach, 1957* Microlaimus formosus Gerlach, 1957* Microlaimus spinosus Gerlach, 1957* Spirobolbolaimus Soeatert & Vincx, 1988 Unidentified species Family Aponchidae Synonema Cobb, 1920* Synonema braziliense Cobb, 1920* BIODIVERSITY OF MARINE NEMATODES IN BRAZIL Zootaxa 2568 © 2010 Magnolia Press · 59 Family Monoposthidae Monoposthia De Man, 1889 Unidentified species Nudora Cobb, 1920 Nudora besnardi Gerlach, 1956* [Monoposthia besnardi Gerlach, 1956] Nudora ilhabelae Gerlach, 1957* [Monoposthia ilhabelae Gerlach, 1957] Rhinema Cobb, 1920 Unidentified species ORDER DESMOSCOLECIDA Suborder Desmoscolecina Superfamily Desmoscolecoidea Family Desmoscolecidae Desmogerlachia Freudenhammer, 1975 Unidentified species Desmolorenzenia Freudenhammer, 1975 Unidentified species Desmoscolex Claparède, 1863 Unidentified species Quadricoma Filipjev, 1922 Unidentified species Tricoma Cobb, 1893 Unidentified species Family Meyliidae Boucherius Decraemer & Jensen, 1982 Unidentified species Gerlachius Andrássy, 1976 Unidentified species Family Cyartonematidae Cyartonema Cobb, 1920 Unidentified species ORDER MONHYSTERIDA Suborder Monhysterina Superfamily Monhysteroidea Family Monhysteridae Diplolaimella Allgén, 1929 Diplolaimella chitwoodi Gerlach, 1957 Diplolaimelloides Meyl, 1954 Diplolaimelloides cf. altherri Meyl, 1954 Gammarinema Kinne & Gerlach, 1953 Unidentified species Litotes Cobb, 1920* (gen. inq.) Litotes minuta Cobb, 1920* Monhystera Bastian, 1865 Unidentified species Thalassomonhystera Jacobs, 1987 Unidentified species Zygonemella Cobb, 1920 Zygonemella striata Cobb, 1920 60 · Zootaxa 2568 © 2010 Magnolia Press VENEKEY ET AL. Superfamily Sphaerolaimoidea Family Sphaerolaimidae Doliolaimus Lorenzen, 1966 Unidentified species Sphaerolaimus Bastian, 1865 Sphaerolaimus gracilis De Man, 1876 Sphaerolaimus lamasus Gerlach, 1956 Sphaerolaimus lodosus Gerlach, 1956 Sphaerolaimus cf. maeoticus Filipjev, 1922 Sphaerolaimus penicillus Gerlach, 1956 Subsphaerolaimus Lorenzen, 1978 Unidentified species Family Xyalidae Ammotheristus Lorenzen, 1977 Unidentified species Amphimonhystera Allgén, 1929 Unidentified species Amphimonhystrella Timm, 1961 Unidentified species Cobbia De Man, 1907 Cobbia cf. dentata Gerlach, 1953 Daptonema Cobb, 1920 Daptonema curvatus Gerlach, 1957 [Theristus cf. curvatus Gerlach, 1956] Daptonema setosus (Butschli, 1874) Lorenzen, 1977 [Theristus setosus Butschli, 1874] Daptonema tenuispiculum (Ditlevsen, 1918) Wieser, 1956 [Theristus tenuispiculum Ditlevsen, 1918] Daptonema trichinus Gerlach, 1956 [Theristus cf. trichinus Gerlach, 1956] Elzalia Gerlach, 1957* Elzalia floresi Gerlach, 1957* Gnomoxyala Cobb, 1920 Unidentified species Gonionchus Cobb, 1920 Unidentified species Linhystera Juario, 1974 Unidentified species Metadesmolaimus Stekhoven, 1935 Metadesmolaimus cf. labiosetosus Stekhoven, 1935 Metadesmolaimus tersus (Gerlach, 1956) Lorenzen, 1972* [Theristus tersus Gerlach, 1956] Omicronema Cobb, 1920 Omicronema clavulatum Gerlach, 1957* Omicronema litorium Cobb, 1920 Paramonohystera Steiner, 1916 [Leptogastrella Cobb, 1920] Paramonohystera stricta (Gerlach, 1956) Wieser, 1956* [Leptogastrella stricta Gerlach, 1956] Promonhystera Wieser, 1956 Unidentified species Prorhynchonema Gourbault, 1982 Unidentified species Pseudosteineria Wieser, 1956 Pseudosteineria scopae (Gerlach, 1956) Wieser, 1959 [Steineria scopae Gerlach, 1956] Retrotheristus Lorenzen, 1977 Unidentified species Rhynchonema Cobb, 1920 Rhynchonema cinctum Cobb, 1920 Scaptrella Cobb, 1917 Scaptrella cf. brevicaudata Gerlach, 1953 Scaptrella cincta Cobb, 1917 Steineria Micoletzky, 1922 Steineria ericia Gerlach, 1956 BIODIVERSITY OF MARINE NEMATODES IN BRAZIL Zootaxa 2568 © 2010 Magnolia Press · 61 Steineria marcorum Gerlach, 1956 Steineria pavo Gerlach, 1957 Steineria tripartita Gerlach, 1957* Stylotheristus Lorenzen, 1977 Unidentified species Theristus Bastian, 1865 Theristus acribus Gerlach, 1956* Theristus flevensis Stekhoven, 1935 Theristus heterospiculoides Gerlach, 1953 Theristus macroflevensis Gerlach, 1954 Theristus metaflevensis Gerlach, 1954 Theristus pertenuis Bresslau & Stekhoven in Stekhoven, 1935 Theristus aff. rhynchonemoides Hopper, 1961 Theristus stranus Gerlach, 1957* Trichotheristus Wieser, 1956 Trichotheristus heterus (Gerlach, 1957) Wieser & Hopper, 1967* [Theristus heterus Gerlach, 1957] Trichotheristus mirabilis (Stekhoven & De Conick, 1933) Wieser, 1956 [Steineria cf. mirabilis Stekhoven & De Connick, 1933] Valvaelaimus Lorenzen, 1977 Unidentified species Xenolaimus Cobb, 1920 Unidentified species Xyala Cobb, 1920 Xyala striata Cobb, 1920 Suborder Linhomoeina Superfamily Siphonolaimoidea Family Siphonolaimidae Astomonema Ott, Rieger & Enderes, 1982 Unidentified species Parastomonema Kito, 1989 Unidentified species Siphonolaimus De Man, 1893 Siphonolaimus weismanni (zur Strassen, 1904) Steiner, 1916 Family Linhomoeidae Anticyathus Cobb, 1920 Anticyathus trochus (Gerlach, 1957) Gerlach, 1963 [Prosphaerolaimus trochus Gerlach, 1957] Desmolaimus De Man, 1880 Desmolaimus calvus Gerlach, 1956* Didelta Cobb, 1920 Unidentified species Disconema Filipjev, 1918 Unidentified species Eleutherolaimus Filipjev, 1922 Eleutherolaimus obtusicaudatus Allgén, 1947 Halinema Cobb, 1920 Halinema varicans Gerlach, 1956* Linhomoeus Bastian, 1865 Unidentified species Megadesmolaimus Wieser, 1954 Unidentified species Metalinhomoeus De Man, 1907 Metalinhomoeus filiformis (De Man, 1907) Stekhoven, 1935 Metalinhomoeus typicus De Man, 1907 Paralinhomoeus De Man, 1907 62 · Zootaxa 2568 © 2010 Magnolia Press VENEKEY ET AL. Paralinhomoeus conspicuus Gerlach, 1957* Terschellingia De Man, 1888 Terschellingia communis De Man, 1888 Terschellingia longicaudata De Man, 1907 Terschellingia mora Gerlach, 1956* Terschellingia papillata Gerlach, 1955 ORDER ARAEOLAIMIDA Superfamily Axonolaimoidea Family Axonolaimidae Apodontium Cobb, 1920 Apodontium procerus Gerlach, 1957* [Axonolaimus procerus Gerlach, 1957] Ascolaimus Ditlevsen, 1919 Unidentified species Axonolaimus De Man, 1889 Axonolaimus caudostriatus Boucher, 1973 Axonolaimus demani De Conick & Stekhoven, 1933 Axonolaimus aff. steineri Timm, 1952 Axonolaimus tirrenicus Brunetti, 1941 Odontophora Bütschli, 1874 Odontophora paragranulifera Timm, 1952 Odontophora urothrix Gerlach, 1957* Odontophoroides Boucher & Helléouët, 1977 Unidentified species Parodontophora Timm, 1963 Unidentified species Pseudolella Cobb, 1920 Pseudolella intermedia Gerlach, 1957* Synodontium Cobb, 1920 Unidentified species Family Comesomatidae Actarjania Hopper, 1967 Unidentified species Cervonema Wieser, 1954 Unidentified species Comesoma Bastian, 1865 Comesoma arenae Gerlach, 1956 Comesoma sipho Gerlach, 1956* Dorylaimopsis Ditlevsen, 1918 Unidentified species Hopperia Vitiello, 1969 Unidentified species Laimella Cobb, 1920 Unidentified species Metacomesoma Wieser, 1954 Unidentified species Paracomesoma Hope & Murphy, 1972 Unidentified species Paramesonchium Hopper, 1967 Unidentified species Pierrickia Vitiello, 1970 Unidentified species Sabatieria Rouville, 1903 Sabatieria breviseta Stekhoven, 1935 [Sabatieria quadripapillata Filipjev, 1922] Sabatieria celtica Southern, 1914 [Sabatieria cupida Bresslau & Stekhoven in Stekhoven, 1935] Sabatieria mortenseni (Ditlevsen, 1921) Filipjev, 1922 BIODIVERSITY OF MARINE NEMATODES IN BRAZIL Zootaxa 2568 © 2010 Magnolia Press · 63 Sabatieria praedatrix De Man, 1907 Sabatieria pulchra (Schneider, 1906) Riemann, 1970 [Sabatieria claviculata Filipjev, 1918] Sabatieria punctata Kreis, 1924 Sabatieria rota Gerlach, 1957* Sabatieria supplicans Gerlach, 1956* Setosabatieria Platt, 1985 Unidentified species Vasostoma Wieser, 1954 Unidentified species Family Diplopeltidae Araeolaimus De Man, 1888 Unidentified species Campylaimus Cobb, 1920 Unidentified species Diplopeltis Cobb in Stiles & Hassal, 1905 Diplopeltis cirrbatus (Berth, 1863) Cobb, 1891 Diplopeltula Gerlach, 1950 Diplopeltula intermedia Gerlach, 1954 Morlaixia Vincx & Gourbault, 1988 Unidentified species Southerniella Allgén, 1932 Unidentified species ORDER PLECTIDA Superfamily Leptolaimoidea Family Leptolaimidae Alaimella Cobb, 1920* Alaimella truncata Cobb, 1920* Antomicron Cobb, 1920 Antomicron pellucidum Cobb, 1920 Assia Gerlach, 1957* Assia laureata Gerlach, 1957* Camacolaimus De Man, 1889 Unidentified species Cricolaimus Southern, 1914 Unidentified species Dagda Southern, 1914 Unidentified species Deontolaimus De Man, 1880 Unidentified species Diodontolaimus Southern, 1914 Unidentified species Halaphanolaimus Southern, 1914 Unidentified species Ionema Cobb, 1920 Ionema cobbi (Steiner, 1916) Micoletzky, 1924 [Ionema ocellatum Cobb, 1920] Leptolaimoides Vitiello, 1971 Unidentified species Leptolaimus De Man, 1876 Leptolaimus surdus Gerlach, 1957* Listia Blome, 1982 Unidentified species Onchium Cobb, 1920 Unidentified species Procamacolaimus Gerlach, 1954 Procamacolaimus cosmius Gerlach, 1956* 64 · Zootaxa 2568 © 2010 Magnolia Press VENEKEY ET AL. Stephanolaimus Ditlevsen, 1914 Unidentified species Family Aphanolaimidae Anonchus Cobb, 1913 Anonchus mangrove Gerlach, 1957 Paraphanolaimus Micoletzky, 1923 Paraphanolaimus cantor Gerlach, 1957* Family Aegialoalaimidae Aegialoalaimus De Man, 1907 Unidentified species Southernia Allgén, 1929 Unidentified species Family Diplopeltoididae Diplopeltoides Gerlach, 1962 Unidentified species Family Paramicrolaimidae Paramicrolaimus Wieser, 1954 Paramicrolaimus papillatus (Gerlach, 1956) Wieser & Hopper, 1967 [Microlaimus papillatus Gerlach, 1956] Superfamily Plectoidea Family Chronogasteridae Chronogaster Cobb, 1913 Chronogaster alatum Gerlach, 1956* Cynura Cobb, 1920 Cynura cerambus Andrássy, 1973 Cynura uniformis Cobb, 1920 Superfamily Ceramonematoidea Family Tarvaiidae Tarvaia Allgén, 1934 Unidentified species Family Ceramonematidae Ceramonema Cobb, 1920 Ceramonema filum Gerlach, 1957* Ceramonema rectum Gerlach, 1957* Dasynemella Cobb, 1933 Dasynemella cincta Gerlach, 1957* Dasynemoides Chitwood, 1936 Dasynemoides cristatus Gerlach, 1957* Dasynemoides latus Gerlach, 1957* Metadasynemella De Coninck, 1942 Unidentified species Metadasynemoides Haspeslagh, 1973 Unidentified species Pselionema Cobb, 1933 Unidentified species Pterygonema Gerlach, 1954 Unidentified species Family Tubolaimoididae Tubolaimoides Gerlach, 1963 Unidentified species BIODIVERSITY OF MARINE NEMATODES IN BRAZIL Zootaxa 2568 © 2010 Magnolia Press · 65 Superfamily Haliplectoidea Family Peresianidae* Manunema Gerlach, 1957* Manunema proboscidis Gerlach, 1957* Family Haliplectidae Haliplectus Cobb, 1913 [Aegialospirinia De Conick, 1943] Haliplectus bibulbosus (Schulz, 1935) Gerlach, 1956 [Aegialospirinia bibulbosa Gerlach, 1960] Haliplectus floridanus Cobb in Chitwood, 1956 Haliplectus schulzi (De Conick, 1943) Gerlach, 1957 Setoplectus Vitiello, 1971 Unidentified species ORDER RHABDITIDA Suborder Tylenchina Infraorder Cephalobomorpha Superfamily Cephaloboidea Family Cephalobidae Acrobeles Linstow, 1877 Unidentified species Eucephalobus Steiner, 1936 Unidentified species Heterocephalobus (Brzeski, 1960) Brzeski, 1961 Unidentified species Infraorder Tylenchomorpha Superfamily Tylenchoidea Family Hoplolaimidae Unidentified genus and species Suborder Rhabditina Infraorder Rhabditomorpha Superfamily Rhabditoidea Family Rhabditidae Paradoxorhabditis Khera, 1971 Unidentified species Rhabditis Dujardin, 1845 Rhabditis marina Bastian, 1865 66 · Zootaxa 2568 © 2010 Magnolia Press VENEKEY ET AL.