Article. On new collections of deep-sea Gadiformes (Actinopterygii: Teleostei) from the Brazilian continental slope, between 11 and 23 S

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1 Zootaxa 2433: (2010) Copyright 2010 Magnolia Press Article ISSN (print edition) ZOOTAXA ISSN (online edition) On new collections of deep-sea Gadiformes (Actinopterygii: Teleostei) from the Brazilian continental slope, between 11 and 23 S MARCELO R. S. MELO 1,4, ADRIANA C. BRAGA 2, GUSTAVO W. A. NUNAN 3 & PAULO A. S. COSTA Funchess Hall, Auburn University, Auburn, AL USA, mrsmelo@hotmail.com 2 Laboratório de Dinâmica de Populações Marinhas, Universidade Federal do Estado do Rio de Janeiro UNIRIO, Av. Pasteur, 458 Urca, Rio de Janeiro, RJ Brazil. acbragaz@hotmail.com; pauloascosta@uol.com.br 3 Dept. de Vertebrados, Museu Nacional/UFRJ, Quinta da Boa Vista, Rio de Janeiro, RJ Brazil, gwanunan@acd.ufrj.br 4 Corresponding author Abstract New collections made by the French research vessel Thalassa and the Brazilian Astro Garoupa on the Brazilian continental slope, between 11 and 23 S revealed a great diversity of deep-sea gadiforms obtained between 200 and 2270 m. Of the 34 species collected, 13 (38%) are being reported for the first time in the western South Atlantic and one may represent a new species. The most species-rich family in the area is Macrouridae (25 species), followed by Moridae (4), Phycidae (2), Merlucciidae (2), and Bregmacerotidae (1). Most of the species collected is also found in the Gulf of Mexico and Caribbean and have their known ranges of distribution extended into the tropical waters of central Brazil. Four species typically occur in temperate waters and were found only in the southernmost stations. A summary comparison of the bathymetric distributions of each species on the slope is provided. Key words: Western South Atlantic; demersal; pelagic; gadiforms; taxonomy Resumo Novas coleções realizadas pelo navio oceanográfico francês Thalassa e brasileiro Astro Garoupa no talude continental brasileiro, entre 11 e 23 S, revelaram uma grande diversidade de peixes Gadiformes em águas profundas entre 200 e 2270 m. Das 34 espécies coletadas, 13 (38%) estão sendo relatadas pela primeira vez no Atlântico Sul ocidental e uma pode representar uma espécie nova. As famílias mais diversificadas na área foram Macrouridae (25 espécies), seguida de Moridae (4), Phycidae (2), Merlucciidae (2) e Bregmacerotidae (1). A maioria das espécies coletadas também é encontrada no Golfo do México e Caribe e tiveram suas áreas de distribuição conhecidas estendidas para as águas tropicais do Brasil central. Quatro espécies têm ocorrência típica em águas temperadas e foram encontradas somente nas estações mais ao sul. A comparação resumida entre a distribuição batimétrica para cada espécie é fornecida. Introduction The Gadiformes comprises a highly diversified group with more than 500 species in 75 genera and 9 families, commonly known as cods, hakes, and grenadiers. The gadiforms inhabit a great variety of marine environments, such as the tropical and temperate shallow waters and can be even found in freshwaters (genus Lota), but it is in the deep-waters of the world oceans where the greatest diversity is found. Families such as Gadidae and Merlucciidae have been well-studied taxonomically and ecologically because of their economic importance. The greatest diversity within this group, however, lies in the benthopelagic family Macrouridae with more than 350 species. Despite of the economical importance of some species, the gadiforms compromises a group that still needs considerable ecological, taxonomic and phylogenetic attention (Lloris et al. 2003; Cohen et al. 1990; Nelson, 2006). Accepted by M. Craig: 6 Mar. 2010; published: 22 Apr

2 The first records or species descriptions of gadiforms in the western South Atlantic were made by Günther (1877; 1878; 1887) based on material collected by the H.M.S. Challenger off the coasts of Brazil, Argentina and Uruguay. After a gap of more than 100 years from Günther s works, taxonomic knowledge has rapidly expanded due to a number of deep-water fisheries surveys off South America. The most remarkable records of deep-sea gadiforms were made by the Japanese Marine Fishery Resource Center (JAMARC) in the northern and southern extremes of South America (Uyeno et al. 1983; Nakamura et al. 1986), the U.S. Bureau of Commercial Fisheries, off the mouth of the Amazon River (Iwamoto & Arai 1987), and several Russian expeditions around the Falkland Islands and the Rio Grande Rise (Parin et al.1995; Brickle & Laptikhovsky 2002). Very few expeditions investigated the Brazilian slope, but those made by the French R/Vs Marion Dufresne and Thalassa, and by the Brazilian R/V Professor W. Bersnard in the past 20 years should be mentioned (Séret & Andreata 1992; Figueiredo et al. 2002; Bernardes et al. 2005; Mincarone et al. 2008; Melo et al. 2009; this contribution). These expeditions and some other random captures resulted in accumulated knowledge of 63 species in the western South Atlantic, ten of which were described from the area of study: Antimora rostrata (Günther 1878); Coryphaenoides affinis Günther 1878; C. leptolepis (Günther 1877); Coelorinchus marinii Hubbs 1934; Hymenocephalus billsam Marshall and Iwamoto 1973; Lepidion ensiferus (Günther 1887); Malacocephalus okamurai Iwamoto and Arai 1987; Merluccius hubbsi Marini 1933; Urophycis brasiliensis (Kaup 1858); and U. mystacea Ribeiro Recently, five expeditions by the French R/V Thalassa and the Brazilian R/V Astro Garoupa explored the diversity of the Brazilian slope between 11 and 23 S, in depths from 200 up to 2270 meters. These collections include 31 species from six families of Gadiformes, 13 species of which are being recorded for the area for the first time. Herein we give species accounts for the gadiforms collected during those cruises and previously listed by Costa et al. (2007), and a review of the specimens collected by the R/V Marion Dufresne which are deposited at the Universidade Santa Úrsula, Brazil. FIGURE 1. Trawl stations of the research vessels Thalassa and Astro Garoupa off eastern Brazil. Seamounts associated with the Abrolhos chain (Minerva, Rodger and Hotspur) and those linked to the Vitória-Trindade chain (Besnard, Vitória, Montagne, Jaseur, Davis, Dogaressa, Columbia). 26 Zootaxa Magnolia Press MELO ET AL.

3 Material and methods TERMS OF USE The specimens listed herein were collected during five oceanographic cruises of the French R/Vs Thalassa and Marion Dufresne, and the Brazilian R/V Astro Garoupa. The collections were made using different types of pelagic and bottom trawls, towed on the Brazilian continental slope off Bahia, Espirito Santo and Rio de Janeiro States, from S, and W, at depths from 200 to 2271 m (Fig. 1). Further details on the method of capture are given by Mincarone et al. (2008). The complete list of demersal trawls that captured gadiforms is listed in Tables 1 and 2, and Séret and Andreata (1992). All pictures of fresh specimens were taken onboard the R/V Thalassa; in some cases imaged specimens could not be tracked to the museum specimens. For each species listed below the following information is given: catalog number by lot, total number of specimens in the lot, range of specimens in total length, abbreviation for the research vessel (T for R/V Thalassa, AG for R/V Astro Garoupa, and MD for R/V Marion Dufresne), and the collecting station. Total range for each species is based on literature records; type locality follows Eschmeyer (2010) coordinates are only made available when mentioned by the author. Taxonomic classification follows Nelson (2006). Abbreviations of the lender institutions follow Leviton et al. (1985), with the addition of USU for Universidade Santa Úrsula fish collection, Rio de Janeiro, RJ, Brazil; WSA stands for western South Atlantic. TABLE 1. List of stations, which collected Gadiformes, made by the French R/V Thalassa on the Brazilian continental slope. Station Date Coordinates Depth range Initial Final (m) D /6/ 'S, 'W 'S, 'W D /6/ 'S, 'W 'S, 'W D /6/ 'S, 'W 'S, 'W D /6/ 'S, 'W 'S, 'W D /6/ 'S, 'W 'S, 'W D-538 7/7/1999 no data 'S, 'W E-495 6/6/ 'S, 'W 'S, 'W E-496 7/6/ 'S, 'W 'S, 'W E-497 7/6/ 'S, 'W 'S, 'W E-498 8/6/ 'S, 'W 'S, 'W E-499 8/6/ 'S, 'W 'S, 'W E-500 8/6/ 'S, 'W 'S, 'W E-501 9/6/ 'S, 'W 'S, 'W E-502 9/6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W continued next page DEEP-SEA GADIFORMS FROM BRAZIL Zootaxa Magnolia Press 27

4 TABLE 1. (continued) TERMS OF USE Station Date Coordinates Depth range Initial Final (m) E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E /6/ 'S, 'W 'S, 'W E-535 1/7/ 'S, 'W 'S, 'W E-536 2/7/ 'S, 'W 'S, 'W E-537 2/7/ 'S, 'W 'S, 'W E-538 2/7/ 'S, 'W 'S, 'W E-539 4/7/ 'S, 'W 'S, 'W E-540 4/7/ 'S, 'W 'S, 'W E-541 4/7/ 'S, 'W 'S, 'W E-543 5/7/ 'S, 'W 'S, 'W E-544 5/7/ 'S, 'W 'S, 'W E-545 6/7/ 'S, 'W 'S, 'W E-546 6/7/ 'S, 'W 'S, 'W E-547 6/7/ 'S, 'W 'S, 'W E-548 7/7/ 'S, 'W 'S, 'W E-549 7/7/ 'S, 'W 'S, 'W E-550 7/7/ 'S, 'W 'S, 'W E-551 8/7/ 'S, 'W 'S, 'W E-552 8/7/ 'S, 'W 'S, 'W Zootaxa Magnolia Press MELO ET AL.

5 TABLE 2. List of stations which collected Gadiformes, made by the Brazilian R/V Astro Garoupa on the Brazilian continental slope. Station Date Coordinates Depth range Initial Final (m) ASTRO /2/ 'S, 'W 'S, 'W ASTRO /2/ 'S, 'W 'S, 'W ASTRO /2/ 'S, 'W 'S, 'W ASTRO /2/ 'S, 'W 'S, 'W ASTRO /2/ 'S, 'W 'S, 'W ASTRO /2/ 'S, 'W 'S, 'W ASTRO /2/ 'S, 'W 'S, 'W ASTRO /2/ 'S, 'W 'S, 'W ASTRO /8/ 'S, 'W 'S, 'W ASTRO /8/ 'S, 'W 'S, 'W ASTRO /8/ 'S, 'W 'S, 'W ASTRO /8/ 'S, 'W 'S, 'W ASTRO /8/ 'S, 'W 'S, 'W Results Bregmacerotidae Bregmaceros sp. (Fig. 2 A) Material examined. 74 specimens: MNRJ (1, 53.0 mm), ASTRO ; MNRJ (23, mm), S, W, m, R/V Thalassa, pelagic trawl; MNRJ (4, mm), D-502; MNRJ (8, mm), D-503; MNRJ (1, 56.0 mm), E-524; MNRJ (1, 61.0 mm), E-504; MNRJ (4, mm), E-538; MNRJ (1, 61.0 mm), E-510; MNRJ (29, mm), E-504; MNRJ (2, mm), E-505. Remarks. Collected from off Bahia, Espírito Santo and Rio de Janeiro, at depths of 266 to 1762 m. Five species of Bregmaceros were assigned for the western South Atlantic: Bregmaceros atlanticus Goode and Bean 1886; B. cantori Milliken and Houde 1984; B. cayorum Nichols 1952; B. houdei Saksena and Richards 1986; and B. macclellandi Günther 1862 (Milliken & Houde 1984; Shen & Wang 1991; Menezes 2003a; Torii et al. 2004). The genus is under revision and several new species remain to be described, including a species from the area of study (A. Harold, pers. comm., 2008). Gadidae Urophycis brasiliensis (Kaup 1858) (Fig. 2 B) Material examined. One specimen: MNRJ (1, mm), T, E-539. Distribution. Collected from off Espírito Santo, at depths from 192 to 207 m. Urophycis brasiliensis was described from off Montevideo, Uruguay; it is also known from Argentina to Southern Brazil (Cousseau 1993; Bernardes et al. 2005; Menezes 2003b). DEEP-SEA GADIFORMS FROM BRAZIL Zootaxa Magnolia Press 29

6 FIGURE 2. (A) Bregmaceros sp. (MNRJ 27001, 56.0 mm); (B) Urophycis brasiliensis (MNRJ 27035, mm); (C) Urophycis mystacea (MNRJ 26978, mm); (D) Gadomus arcuatus* (MNRJ 27067, mm); (E) Gadomus capensis* (MNRJ 26921, mm); (F) Coelorinchus caribbaeus (MNRJ (, mm); (G) Coelorinchus aff. carminatus (MNRJ 26985, mm); (H) Coelorinchus marinii (MNRJ 26925, mm); (I) Coelorinchus occa (MNRJ 27095, mm); (J) Cetonurus globiceps* (MNRJ unknown, ca mm); (K) Coryphaenoides cf. leptolepis (MNRJ 26899, mm); (L) Coryphaenoides rudis (MNRJ 27070, mm). Asterisk indicates pictures taken on board of the collecting vessel, prior to the specimen fixation. 30 Zootaxa Magnolia Press MELO ET AL.

7 Urophycis mystacea Ribeiro 1903 (Fig. 2 C) Material examined. 43 specimens: MNRJ (1, mm), T, E-530; MNRJ (16, mm), T, E-539; MNRJ (5, mm), T, E-531; MNRJ (5, mm), T, E-531; MNRJ (1, mm), T, E-531; MNRJ (1, mm), T, E-545; MNRJ (2, mm), T, E-541; MNRJ (2, mm), T, E-532; MNRJ (1, mm), T, E-546; MNRJ (1, mm), T, E-541; MNRJ (1, mm), T, E-546; MNRJ (1, mm), T, E- 530; MNRJ (1, mm), T, E-541; MNRJ (1, mm), T, E-541; MNRJ (1, mm), T, E-538; MNRJ (1, mm), T, E-534. Distribution. Collected from off Espírito Santo and Rio de Janeiro, at depths from 182 to 616 m; one specimen collected at m is understood as an accidental catch during net transition since this is a well known shallow water species (Cohen et al. 1990). Urophycis mystacea was described from off Rio de Janeiro, Brazil and is generally regarded as the species from southern Brazil to Argentina. Cousseau (1993) suggest that U. mystacea is a junior synonym of U. cirrata (Goode & Bean 1896), which currently regarded as the species from Florida, Gulf of Mexico, Caribbean, and Venezuela to off the mouth of the Orinoco River (Cohen et al. 1990; Menezes 2003b). Macrouridae Gadomus arcuatus (Goode & Bean 1886) (Fig. 2 D) Material examined. One specimen: MNRJ (1, mm), T, E-522. Distribution. First record of Gadomus arcuatus in the WSA. Collected from off Bahia, at depths from 1044 to 1275 m. It was described from off Martinique and was previously known in the western North Atlantic from the Gulf of Mexico, Caribbean, Surinam and French Guyana; and in the eastern North Atlantic, from Morocco and the Canary Islands (Cohen et al. 1990; Geistdoerfer 1990; Howes & Crimen 1990). Gadomus capensis (Gilchrist & Von Bonde 1924) (Fig. 2 E) Material examined. One specimen: MNRJ (1, mm), T, E-507. Distribution. Collected from off Bahia, at depths from 1012 to 1049 m. Gadomus capensis was described from Table Bay, South Africa (33 30 S, E); in the WSA, it is also known from the eastern South Atlantic to Mozambique, western Indian, and from the Rio Grande Rise (Howes & Crimen 1990; Parin et al. 1995). Coelorinchus caribbaeus (Goode & Bean 1885) (Fig. 2 F) Material examined. 41 specimens: MNRJ (8, mm), T, E-516; MNRJ (1, mm), AG, ASTRO ; MNRJ (3, mm), T, E-533; MNRJ (6, mm), T, E-511; MNRJ (1, mm), T, E-515; MNRJ (2, mm), T, E-549; MNRJ (3, mm), T, E-521; MNRJ (5, mm), T, E-500; MNRJ (3, mm), T, E-508; MNRJ (5, mm), T, E-511; MNRJ (1, mm), T, E-549; UF (1, mm), T, E-511; USNM (2, mm), T, E-516. DEEP-SEA GADIFORMS FROM BRAZIL Zootaxa Magnolia Press 31

8 Distribution. Collected from off Bahia, Espírito Santo and Rio de Janeiro, at depths from 100 to 1721 m. Coelorinchus caribbaeus was described from the Gulf of Mexico ( N, W), and is known from the Gulf of Mexico, off the mouth of the Amazon River and southern Brazil (Marshall & Iwamoto 1973a; Cohen et al. 1990; Séret & Andreata 1992; Menezes & Figueiredo 2003a). Coelorinchus aff. carminatus (Goode 1880) (Fig. 2 G) Material examined. 35 specimens: MNRJ (3, mm), T, D-538; MNRJ (1, mm), T, D-464; MNRJ (1, mm), T, D-502; MNRJ (1, mm), T, E-521; MNRJ (1, mm), T, E-515; MNRJ (2, mm), E-534; MNRJ (3, mm), T, E-543; MNRJ (9, mm), T, E-502; MNRJ (1, mm), E-541; MNRJ (7, mm), T, E-508; MNRJ (2, mm), T, E-517; UF (2, mm), T, D-538; USNM (2, mm), T, D-538. Remarks. Collected from off Bahia, Espírito Santo and Rio de Janeiro, at depths from 379 to 801 m. Coelorinchus aff. carminatus belongs to a species complex of five species, sometimes treated as subspecies of Coelorinchus caelorinchus (Risso 1810): C. caelorinchus; C. carminatus (Goode 1880); C. geronimo Marshall and Iwamoto 1973; C. marinii Hubbs 1934; and C. polli Marshall and Iwamoto 1973 (T. Iwamoto, pers. comm. 2008). The shape of the snout plate is similar to what is described for C. carminatus; however, the pattern of spinules on the scales differs from specimens from North Atlantic and Gulf of Mexico. The type locality of C. carminatus is off New York, USA ( N, W), but holotype is lost (USNM 26001; D. Smith, person. comm., 2009). Further comparisons between our material and specimens obtained from the western and central North Atlantic are being made by the senior author. This species has probably been reported from Brazil and Argentina as C. carminatus (Uyeno 1983; Séret & Andreata 1992; Cousseau 1993; Menezes & Figueiredo 2003a). Coelorinchus marinii Hubbs 1934 (Fig. 2 H) Material examined. Three specimens: MNRJ (2, mm), T, D-464; MNRJ (1, mm), T, E-543. Distribution. Collected from off Rio de Janeiro, at depths from 617 to 776 m. Coelorinchus marinii was described from off the province of Buenos Aires, Argentina (38 52 S, W) and is known from southern Brazil to Antarctica (Cohen et al. 1990; Iwamoto 1990; Cousseau 1993; Menezes & Figueiredo 2003a). Coelorinchus occa (Goode & Bean 1885) (Fig. 2 I) Material examined. 10 specimens: MNRJ (1, mm), T, D-503; MNRJ (1, mm), T, D-503; MNRJ (1, mm), T, D-506; MNRJ (1, mm), T, D-506; MNRJ (1, mm), T, D-506; MNRJ (1, mm), T, D-506; MNRJ (1, mm), T, E-503; MNRJ (1, mm), T, E-524; MNRJ (2, mm), T, E-499. Distribution. First record of Coelorinchus occa in the WSA. Collected from off Bahia, Espírito Santo and Rio de Janeiro, at depths from 714 to 1004 m. It was described from the Gulf of Mexico (28 34 N, W), and was previously known from the Straits of Florida, Gulf of Mexico, southern part of Caribbean, and Surinam (Marshall & Iwamoto 1973a; Cohen et al. 1990). 32 Zootaxa Magnolia Press MELO ET AL.

9 Cetonurus globiceps (Vaillant 1888) (Fig. 2 J) Material examined. 41 specimens: MNRJ (1, mm), AG, ASTRO ; MNRJ (1, mm), T, D-504; MNRJ (2, mm), T, E-536; MNRJ (8, mm), T, E- 527; MNRJ (16, mm), T, E-526; MNRJ (1, mm), T, E-552; MNRJ (1, 84.0 mm), T, E-526; MNRJ (1, mm), T, E-535; MNRJ (1, mm), T, E-550; MNRJ (1, mm), T, E-544; MNRJ (1, mm), T, E-552; MNRJ (3, mm), T, E-525; MNRJ (4, mm), T, E-547. Distribution. Collected from off Espírito Santo and Rio de Janeiro, at depths from 909 to 1699 m. Cetonurus globiceps was described from off Western Sahara (23 57'N, 17 12'W), and is also known in eastern Atlantic from Mauritania, Senegal, Canary Islands, Bay of Biscal and Azores, in the western Atlantic from the Gulf of Mexico, Caribbean and Brazil, and in the Pacific from Japan and Indonesia (Iwamoto 1966; Marshall 1973a; Gaistdoerfer 1990; Séret & Andreata 1992; Menezes & Figueiredo 2003a). Coryphaenoides cf. leptolepis (Günther 1877) (Fig. 2 K) Material examined. One specimen: MNRJ (1, mm), AG, ASTRO Distribution. Collected from off Rio de Janeiro, at depths from 1071 to 1205 m. The type locality of Coryphaenoides leptolepis is Pernambuco, Brazil (BMNH ; 9 05 S, W), but specimens from other localities in the Pacific were also used: Yokohama (BMNH ), and mid-pacific (BMNH ). Günther (1887:144) restricted its distribution to Brazil, reallocating the other specimens to Macrurus liocephalus Günther, 1887, and Marshall (1973b) designated the lectotype (Eschmeyer 2010). Remarks. Coryphaenoides leptolepis is being reported from a badly preserved specimen with a broken tail, body scales lacking, and the opercular region mostly destroyed on both sides. Comparison between our specimen and data available in literature (Günther 1887; Marshall 1973b; Cohen et al. 1990) indicated a few meristic differences: pelvic-fin rays 7 (vs. 9), first dorsal-fin rays 11 (vs. 10), and length of pelvic-fin filament 60% in head length (vs. 96%). Our specimen, however, agrees with the illustration provided by Günther (1887, plate XXXI) by having the snout naked with enlarged pores. Divergence in the number of pelvic-fin rays suggests that this can be a different species from C. leptolepis, but more specimens are needed for a conclusive identification. Coryphaenoides rudis Günther 1878 (Fig. 2 L) Material examined. 15 specimens: MNRJ (1, mm), T, E-509; MNRJ (1, mm), T, E-520; MNRJ (1, mm), T, E-551; MNRJ (1, mm), T, E-509; MNRJ (1, mm), T, E-549; MNRJ (1, mm), T, E-549; MNRJ (1, mm), T, E-549; MNRJ (1, mm), T, E-537; MNRJ (1, mm), T, E-537; MNRJ (1, mm), T, E-496; MNRJ (1, mm), T, E-496; MNRJ (1, mm), T, E-540; MNRJ (1, mm), T, E-496; MNRJ (2, mm), T, E-520. Distribution. First record of Coryphaenoides rudis in the WSA. Collected from off Bahia Espírito Santo and Rio de Janeiro, at depths from 1334 to It was described from the South Pacific (28 33 S, W) and is widespread in the Indian and central Atlantic (Günther 1887; Shcherbachev & Iwamoto 1995; Iwamoto 2002). Cohen et al. (1990) referred to it as a possible junior synonym of P. paradoxus. DEEP-SEA GADIFORMS FROM BRAZIL Zootaxa Magnolia Press 33

10 FIGURE 3. (A) Coryphaenoides thelestomus* (MNRJ 27044, mm); (B) Haplomacrourus nudirostris (MNRJ 26880, mm); (C) Hymenocephalus cf. aterrimus (MNRJ 27013, 92.1 mm); (D) Hymenocephalus billsam (MNRJ mm); (E) Macrosmia phalacra (MNRJ 26891, mm); (F) Macrouroides inflaticeps* (MNRJ unknown, ca mm); (G) Malacocephalus laevis (MNRJ 30474, mm); (H) Malacocephalus occidentalis (MNRJ 30476, mm); (I) Malacocephalus okamurai (MNRJ 26988, mm); (J) Nezumia atlantica (MNRJ 26922, mm); (K) Nezumia suilla (MNRJ 26887, mm); (L) Sphagemacrurus grenadae (MNRJ 26879, mm). Asterisk indicates pictures taken on board of the collecting vessel, prior to the specimen fixation. Remarks. This species was mistakenly referred to by Costa et al. (2007) as Coryphaenoides cf. longicirrhus (Gilbert 1905). Coryphaenoides rudis can be distinguished from C. longicirrhus by the mandibulary teeth in two or more rows (vs. mandibulary teeth in a single row). 34 Zootaxa Magnolia Press MELO ET AL.

11 Coryphaenoides thelestomus Maul 1951 (Fig. 3 A) Material examined. Two specimens: MNRJ (1, mm), T, E-512; MNRJ (1, mm), T, E-505. Distribution. First record of Coryphaenoides thelestomus in the WSA. Collected from off Bahia, at depths from 1036 to 1197 m. It was described from off Madeira, western North Atlantic. It is considered to be a rare species, previously known from few specimens collected in the North Atlantic (Crabtree 1983; Gaistdoerfer 1990). Haplomacrourus nudirostris Trunov 1980 (Fig. 3 B) Material examined. Two specimens: MNRJ (1, mm), T, D-504; MNRJ (1, mm), AG, ASTRO Distribution. First record of Haplomacrourus nudirostris in the WSA. Collected from off Espírito Santo and Rio de Janeiro, at depths from 909 to 1135 m. It was described from the eastern South Atlantic, off South Africa (37 09 S, 7 40 E), and was also known from the Indo-Pacific (Iwamoto 2001; Iwamoto et al. 2004). Hymenocephalus cf. aterrimus Gilbert 1905 (Fig. 3 C) Material examined. One specimen: MNRJ (1, 92.1 mm), T, E-498. Distribution. Collected from off Bahia, at depths from 614 to 629 m. Hymenocephalus aterrimus was described from off the Kauai Island, Hawaii. It is known to have a worldwide distribution in tropical regions of the Pacific and Atlantic Oceans (Iwamoto 1999; Iwamoto 2002); in the WSA, it was previously recorded from northeastern Brazil (3 22 S, W) by Marshall and Iwamoto (1973b). Hymenocephalus billsam Marshall & Iwamoto 1973 (Fig. 3 D) Material examined. 35 specimens: MNRJ (3, mm), T, E-503; MNRJ (2, mm), T, E-510; MNRJ (2, mm), T, E-511; MNRJ (6, mm), T, E-543; MNRJ (4, mm), T, E-499; MNRJ (4, mm), T, E-517; MNRJ (4, mm), T, E-522; MNRJ (10, mm), T, D-538. Distribution. Collected from off Bahia and Rio de Janeiro, at depths from 233 to 1275 m. The type locality of Hymenocephalus billsam is off Puerto Rico, but the paratypes were collected from several parts of the Gulf of Mexico and southern Brazil (Marshall & Iwamoto 1973b; Menezes & Figueiredo 2003a). It was recorded in Argentina by Cousseau (1993). Macrosmia phalacra Merrett, Sazonov & Shcherbachev 1983 (Fig. 3 E) Material examined. 10 specimens: MNRJ (8, mm), AG, ASTRO ; MNRJ (1, mm), AG, ASTRO ; MNRJ (1, mm), T, E-537. DEEP-SEA GADIFORMS FROM BRAZIL Zootaxa Magnolia Press 35

12 Distribution. First record of Macrosmia phalacra in the WSA. Collected from off Espírito Santo and Rio de Janeiro, at depths from 1060 to 1567 m. It is rare species, known from the type locality in the Ninety East Ridge, Indian Ocean (17 04 S, E), and few other specimens from the western North Atlantic (Merrett et al. 1983). Macrouroides inflaticeps Smith & Radcliffe 1912 (Fig. 3 F) Material examined. 10 specimens: MNRJ (1, mm), T, E-549; MNRJ (1, mm), T, E-525; MNRJ (1, mm), T, E-527; MNRJ (1, mm), T, E-526; MNRJ (1, mm), T, E-552; MNRJ (5, mm), T, E-523. Distribution. First record of Macrouroides inflaticeps in the WSA. Collected from off Bahia, Espírito Santo and Rio de Janeiro, at depths from 875 to 1721.m. It was described from off the Luzon Island, Philippines ( N, E). It is known from several localities in the Atlantic, Indian, and Pacific oceans (Adam et al. 1998; Pequeño 1989; Uyeno et al. 1983) Malacocephalus laevis (Lowe 1843) (Fig. 3 G) Material examined. 42 specimens: MNRJ (1, mm), T, D-464; MNRJ (1, mm), T, D-502; MNRJ (1, mm), T, E-510; MNRJ (2, mm), T, E-502; MNRJ (1, mm), T, E-503; MNRJ (1, mm), T, E-498; MNRJ (6, mm), T, E- 541; MNRJ (1, mm), T, E-541; MNRJ (1, mm), T, E-518; MNRJ (7, mm), T, E-534; MNRJ (1, mm), T, E-517; MNRJ (1, damaged, ca mm), T, E- 517; MNRJ (3, mm), T, E-546; MNRJ (3, mm), T, E-499; MNRJ (3, mm), T, E-503; MNRJ (2, mm), T, E-503; MNRJ (4, mm), T, E-521; MNRJ (2, ), T, E-543; MNRJ (1, mm), E-503. Distribution. Collected from off Bahia, Espírito Santo and Rio de Janeiro, at depths from 457 to 822 m. Malacocephalus laevis was described from off Madeira, and is known from the tropical Atlantic, Indian and Pacific Oceans (Cohen et al. 1990; Geistdoerfer 1990; Menezes & Figueiredo 2003). Malacocephalus occidentalis Goode & Bean 1885 (Fig. 3 H) Material examined. 22 specimens: MNRJ (12, mm), T, D-464; MNRJ (2, mm), T, E-534; MNRJ (1, mm), T, E-500; MNRJ (1, mm), T, E-503; MNRJ (3, mm), T, E-541; MNRJ (3, mm), T, E-546. Distribution. Collected from off Bahia, Espírito Santo and Rio de Janeiro, at depths from 360 to 822 m. Malacocephalus occidentalis was described from off North Carolina, USA (35 44 N, W), and is known from about 40 N to 40 S in the western Atlantic, and 30 N to 20 S in the eastern Atlantic (Cohen et al. 1990; Menezes & Figueiredo 2003a). Malacocephalus okamurai Iwamoto & Arai 1987 (Fig. 3 I) Material examined. Nine specimens: MNRJ (4, mm), T, E-515; MNRJ (1, mm), T, E-508; MNRJ (1, mm), T, E-518; MNRJ (3, mm), T, E Zootaxa Magnolia Press MELO ET AL.

13 Distribution. First record of Malacocephalus okamurai in the WSA. Collected from off Bahia, at depths from 379 to 627 m. It was described from off the mouth of the Amazon River, Brazil (2 04 N, W); it is a rare species which is only known from the type locality (Iwamoto & Arai 1987). Nezumia atlantica (Parr 1946) (Fig. 3 J) Material examined. Three specimens: MNRJ (2, mm), T, E-503; MNRJ (1, mm), T, E-523. Distribution. First record of Nezumia atlantica in the WSA. Collected from off Bahia and Espírito Santo, at depths from 714 to 942 m. It was described from the Gulf of Mexico (27 48 N, W), and was previously recorded from Straits of Florida, throughout the Caribbean to 5 N in the northern part of South America (Cohen et al. 1990). Nezumia suilla Marshall & Iwamoto 1973 (Fig. 3 K) Material examined. 257 specimens: MNRJ (13, mm), T, D-502; MNRJ (18, mm), T, D-502; MNRJ (34, mm), T, D-504; MNRJ (1, mm), T, D-506; MNRJ (21, mm), T, D-506; MNRJ (1, mm), AG, ASTRO ; MNRJ (1, mm), AG, ASTRO ; MNRJ (1, mm), AG, ASTRO ; MNRJ (1, mm), AG, ASTRO ; MNRJ (1, mm), AG, ASTRO ; MNRJ (82, mm), T, D-503; MNRJ (4, mm), T, E-498; MNRJ (8, mm), T, E-503; MNRJ (3, mm), T, E- 502; MNRJ (1, mm), T, E-517; MNRJ (4, mm), T, E-507; MNRJ (2, mm), T, E-543; MNRJ (3, mm), T, E-523; MNRJ (22, mm), T, E-517; MNRJ (18, mm), T, E-499; MNRJ (2, mm), T, E-524; MNRJ (4, mm), T, E-497; MNRJ (3, mm), T, E-547; MNRJ (1, mm), T, E-506; MNRJ (1, mm), T, E-531; MNRJ (1, mm), T, E-512; MNRJ (2, mm), T, E-544; UF (2, mm), T, D-503; USNM (2, mm), T, D-503. Distribution. Collected from off Bahia, Espírito Santo and Rio de Janeiro, at depths from 182 to 1593 m. Nezumia suilla was described from the Gulf of Mexico off Panama (9 20 N, W), and is known from Gulf of Mexico, Caribbean, Guiana, Venezuela, and southern Brazil (Uyeno et al. 1983; Cohen et al. 1990; Serét & Andreata 1992; Menezes & Figueiredo 2003a). Sphagemacrurus grenadae (Parr 1946) (Fig. 3 L) Material examined. 31 specimens: MNRJ (25, mm), T, D-506; MNRJ (1, 62.0 mm), AG, ASTRO ; MNRJ (2, mm), T, E-527; MNRJ (1, mm), T, E-520; MNRJ (1, mm), T, E-512; MNRJ (1, mm), T, E-526. Distribution. Collected from off Bahia, Espírito Santo and Rio de Janeiro, at depths from 1036 to 2271 m. Sphagemacrurus grenadae was described from off Grenada, and is known from the Gulf of Mexico and Caribbean to southern Brazil (Marshall 1973c; Serét & Andreata 1992). DEEP-SEA GADIFORMS FROM BRAZIL Zootaxa Magnolia Press 37

14 Squalogadus modificatus Gilbert & Hubbs 1916 (Fig. 4 A) Material examined. Eight specimens: MNRJ, (2, mm), T, E-519; MNRJ (2, mm), T, E-519; MNRJ (1, mm), T, E-525; MNRJ (1, mm), T, E-526; MNRJ (1, mm), T, E- 527; MNRJ (1, mm), T, E-552. Distribution. First record of Squalogadus modificatus in the WSA. Collected from off Bahia, Espírito Santo and Rio de Janeiro, at depths from 1342 to 1929 m. It was described from off Kyushu, Japan (32 32 N, W), and is known to have a wide distribution throughout the North Atlantic and Pacific (Geistdoerfer 1990; Iwamoto 1999; Iwamoto 2002). Trachonurus sulcatus (Goode & Bean 1885) (Fig. 4 B) Material examined. 36 specimens: MNRJ (1, mm), T, D-506; MNRJ (1, mm), AG, ASTRO ; MNRJ (1, mm), AG, ASTRO ; MNRJ (3, mm), T, D-504; MNRJ (1, mm), T, D-506; MNRJ (1, mm), T, E-506; MNRJ (1, mm), T, E-497; MNRJ (1, mm), T, E-550; MNRJ (3, mm), T, E- 536; MNRJ (5, mm), T, E-523; MNRJ (1, mm), T, E-525; MNRJ (4, mm), T, E-547; MNRJ (1, mm), T, E-537; MNRJ (4, mm), T, E- 524; MNRJ (4, mm), T, E-540; MNRJ (4, mm), T, E-527. Distribution. First record of Trachonurus sulcatus in the WSA. Collected from off Bahia, Espírito Santo and Rio de Janeiro, at depths from 875 to 1680 m. It was described from the Gulf of Mexico (28 38 N, W), and is known from about 10 N to 40 N in both western and eastern Atlantic (Cohen et al. 1990). Ventrifossa macropogon Marshall 1973 (Fig. 4 C) Material examined. 74 specimens: MNRJ (1, mm), T, D-502; MNRJ (4, mm), T, D-502; MNRJ (1, mm), T, D-502; MNRJ (6, mm), T, D-464; MNRJ (3, mm), T, E-543; MNRJ (1, mm), T, E-517; MNRJ (3, mm), T, E-543; MNRJ (5, mm), T, E-541; MNRJ (7, mm), T, E-503; MNRJ (5, mm), T, D-538; MNRJ (2, mm), T, E-535; MNRJ (4, mm), T, E-520; MNRJ (3, mm), T, E-510; MNRJ (6, mm), T, E-517; MNRJ (1, mm), T, E-534; MNRJ (11, mm), T, E-534; MNRJ (11, mm), T, E-499. Distribution. Collected from off Bahia, Espírito Santo and Rio de Janeiro, at depths from 545 to 2271 m. Ventrifossa macropogon was described from off Jamaica (16 35 N, W), and is known from off the Atlantic coast of Florida, Gulf of Mexico and Caribbean and Brazil; in the Pacific it was recorded from New Caledonia (Cohen et al. 1990; Serét & Andreata 1992; Merrett & Iwamoto 2000). Ventrifossa mucocephalus Marshall 1973 (Fig. 4 D) Material examined. 21 specimens: MNRJ (10, mm), T, D-502; MNRJ (9, mm), T, D-502; MNRJ (1, mm), T, D-502; MNRJ (1, mm), T, E Zootaxa Magnolia Press MELO ET AL.

15 Distribution. First record of Ventrifossa mucocephalus in the WSA. Collected from off Bahia at depths from 705 to 801 m. It was described from northern Cuba ( N, W), and is known from off the Atlantic coast of Florida, Gulf of Mexico and Caribbean (Cohen et al. 1990). FIGURE 4. (A) Squalogadus modificatus* (MNRJ unknown, ca mm); (B) Trachonurus sulcatus* (MNRJ 27063, mm); (C) Ventrifossa macropogon (MNRJ 30429, mm); (D) Ventrifossa mucocephalus (MNRJ 26874, mm); (E) Merluccius hubbsi* (MNRJ unknown, ca mm); (F) Steindachneria argentea* (MNRJ unknown, ca mm); (G) Antimora rostrata* (MNRJ unknown, ca mm); (H) Gadella imberbis* (MNRJ unknown, ca mm); (I) Halargyreus johnsonii* (MNRJ unknown, ca mm); (J) Laemonema goodebeanorum* (MNRJ unknown, ca mm). The pictures of specimens indicated by an asterisk were taken on board of collecting vessel, prior to the specimen fixation. DEEP-SEA GADIFORMS FROM BRAZIL Zootaxa Magnolia Press 39

16 Merlucciidae Merluccius hubbsi Marini 1933 (Fig. 4 E) Material examined. 23 specimens: MNRJ (1, mm), T, E-546; MNRJ (4, mm), T, D-464; MNRJ (1, mm), T, E-534; MNRJ (6, mm), T, E-539; MNRJ (1, mm), T, E-545; MNRJ (6, mm), T, E-546; MNRJ (2, mm), T, E-541; MNRJ (2, mm), T, E-532. Distribution. Collected from off Espírito Santo and Rio de Janeiro, at depths from 192 to 632 m. Merluccius hubbsi was described from off Buenos Aires, Argentina, and is known from S to 49 S, southern Brazil to Argentina (Cousseau 1993; Lloris et al. 2003; Menezes 2003c). Steindachneria argentea Goode & Bean 1896 (Fig. 4 F) Material examined. 202 specimens: MNRJ (3, mm), T, E-510; MNRJ (1, mm), T, E-533; MNRJ (9, mm), T, E-516; MNRJ (8, mm), T, E-521; MNRJ (19, mm), T, E-502; MNRJ (15, mm), T, E-495; MNRJ (5, mm), T, E-511; MNRJ (5, mm), T, E-500; MNRJ (17, mm), T, E-508; MNRJ (2, mm), T, E-518; MNRJ (114, mm), T, D-538; MNRJ (2, ca , cs.), T, D-538; UF (2, mm), T, E-502. Distribution. First record of Steindachneria argentea in the WSA. Collected from off Bahia, at depths from 100 to 607. It was described from off the Mississippi delta, Gulf of Mexico (39 14 N, W), and is known from the Straits of Florida, Gulf of Mexico, Caribbean and Venezuela (Cohen et al. 1990); more recently, Hartel et al. (2008) recorded it in southern Hudson Canyon, off New England, considerably increasing the known distribution of the species. Moridae Antimora rostrata (Günther 1878) (Fig. 4 G) Material examined. 53 specimens: MNRJ (5, mm), AG, ASTRO ; MNRJ (4, mm), AG, ASTRO ; MNRJ (2, mm), AG, ASTRO ; MNRJ (1, mm), AG, ASTRO ; MNRJ (1, mm), AG, ASTRO ; MNRJ (1, mm), AG, ASTRO ; MNRJ (1, mm), AG, ASTRO ; MNRJ (1, mm), AG, ASTRO ; MNRJ (3, mm), T, E-537; MNRJ (1, mm), T, E-552; MNRJ (4, mm), T, E- 535; MNRJ (1, mm), T, E-512; MNRJ (1, mm), T, E-549; MNRJ (2, mm), T, E-540; MNRJ (2, mm), T, E-529; MNRJ (2, mm), T, E- 501; MNRJ (3, mm), T, E-547; MNRJ (2, mm), T, E-523; MNRJ (1, mm), T, E-536; MNRJ (9, mm), T, E-526; MNRJ (1, mm), T, E-544; MNRJ (1, mm), T, E-551; MNRJ (1, mm), T, E-527; MNRJ (3, mm), T, E-527. Distribution. Collected from off Bahia, Espírito Santo and Rio de Janeiro, at depths from 875 to 1721 m. Antimora rostrata was described from off Argentina, and is known from all oceans, except north of 10 N in the Pacific (Cohen et al. 1990; Cohen 1990). 40 Zootaxa Magnolia Press MELO ET AL.

17 Gadella imberbis (Vaillant 1888) (Fig. 4 H) Material examined. 98 specimens: MNRJ (4, mm), T, E-535; MNRJ (1, mm), T, E-498; MNRJ (5, mm), T, E-518; MNRJ (2, mm), T, E-543; MNRJ (10, mm), T, D-464; MNRJ (14, mm), T, E-503; MNRJ (42, mm), T, E-499; MNRJ (19, mm), T, E-502; MNRJ (1, mm), T, E-511. Distribution. Collected from off Bahia, Espírito Santo and Rio de Janeiro, at depths from 233 to 1016 m. Gadella imberbis was described from off Cape Verde Islands (16 53 N, W), and is known from the eastern and western Atlantic, from about 40 N to 30 S (Cohen et al. 1990; Menezes & Figueiredo 2003b). Halargyreus johnsonii Günther 1862 (Fig. 4 I) Material examined. Six specimens: MNRJ (1, mm), T, E-526; MNRJ (1, mm), T, E-526; MNRJ (1, mm), T, E-526; MNRJ (1, mm), T, E-526; MNRJ (1, mm), T, E-527; MNRJ (1, mm), T, E-537. Distribution. Collected from off Espírito Santo, at depths from 1342 to 1649 m. Halargyreus johnsonii was described from off Madeira, and has anti-tropical distribution in the Atlantic, Indian, Pacific and Southern Oceans, except for North Pacific (Cohen et al. 1990; Chiu et al. 1990). Laemonema goodebeanorum Meléndez & Markle 1997 (Fig. 4 J) Material examined. 20 specimens: MNRJ (11, mm), T, E-534; MNRJ (4, mm), T, E-541; MNRJ (2, mm), T, E-510; MNRJ (3, mm), T, E Distribution. Collected from off Bahia, Espírito Santo and Rio de Janeiro, at depths from 545 to 766 m. Laemonema goodebeanorum was described from the Gulf of Mexico (28 34 N, W), and is also known in the western Atlantic from Brazil to New England, USA (Meléndez & Markle 1997; Menezes & Figueiredo 2003b; Moore et al. 2003). Discussion This contribution documents a number of new records of deep-water gadiforms from the western South Atlantic, of the 34 species reported herein, 13 (38%) are new records to the WSA: Coelorinchus occa, Coryphaenoides rudis, Coryphaenoides thelestomus, Gadomus arcuatus, Haplomacrourus nudirostris, Macrosmia phalacra, Macrouroides inflaticeps, Malacocephalus okamurai, Nezumia atlantica, Steindachneria argentea, Squalogadus modificatus, Trachonurus sulcatus, and Ventrifossa mucocephalus. Among these, five species were previously known from very few specimens: Coryphaenoides leptolepis, C. thelestomus, Macrosmia phalacra, Haplomacrourus nudirostris, and Malacocephalus okamurai. Moreover, Coelorinchus aff. carminatus probably represents a new species and is under further investigation. Séret and Andreata (1990) listed one lot of Bathygadus melanobranchus Vaillant 1888 (USU 01409) among the species collected by the R/V Marion Dufresne; the lot, in fact, refers to a badly preserved specimen tentatively identified as Leucicorus (Ophidiidae: Ophiidiformes). Therefore, presence of B. melanobranchus in Brazilian waters is questionable. DEEP-SEA GADIFORMS FROM BRAZIL Zootaxa Magnolia Press 41

18 Four of the species reported herein are endemic to the western South Atlantic: Coelorinchus marinii, Merluccius hubbsi, Urophycis brasiliensis and U. mystacea there are some taxonomic questions regarding the status of the latter, see species account. These species are typically found in the southern part of South America but only north to 19 S, and the latter three are important to fisheries. Six species are known from western North Atlantic, Caribbean and Gulf of Mexico: Coryphaenoides thelestomus, Coryphaenoides leptolepis, Laemonema goodebeanorum, Malacocephalus occidentalis, Nezumia atlantica, and Sphagemacrurus grenadae. Seven species known from the Caribbean and Gulf of Mexico had their range of distribution extended considerably southwards: Coelorinchus caribbaeus, C. occa, Hymenocephalus billsam, Malacocephalus okamurai, Nezumia suilla, Ventrifossa macropogon, and V. mucocephalus. Two species previously reported only from the western North Atlantic are now being recorded from the WSA: Gadomus arcuatus and Trachonurus sulcatus. Nine species have a wide distribution, being found not only in the Atlantic but also in some regions of the Indian and/or Pacific: Cetonurus globiceps, Coryphaenoides rudis, Gadella imberbis, Halargyreus johnsonii, Hymenocephalus aterrimus, Macrosmia phalacra, Macrouroides inflaticeps, Malacocephalus laevis, and Squalogadus modificatus. The most remarkable record of these is Haplomacrourus nudirostris, which was known before from the western South Pacific, Indian and eastern Atlantic that is being recorded for the first time in the eastern Atlantic (Iwamoto & Graham 2001; Iwamoto et al. 2004). Current taxonomic knowledge regarding the species of gadiforms in the western South Atlantic indicates the existence of 74 species in the region (Nakamura et al. 1986; Cousseau 1993; Parin et al. 1995; Brickle & Laptikovsky 2002; Menezes et al. 2003; Caires et al. 2008; present contribution). The number of species for the region, however, will probably rise with additional expeditions to the deep waters. The Gadiformes is a group which is largely diversified in cold waters as the temperate and subtropical zones, and deep-waters. The great richness of species in the area of study can be explained either by the presence of widely-distributed deep-water species that had never before been recorded, or by the diversity of environments: the tropical northern South America shares several species with the Gulf of Mexico and Caribbean (e.g., Steindachneria argentea, Coelorinchus caribbaeus); the temperate central-south has some endemic species (e.g., Coelorinchus marinii and Merluccius hubbsi); and the extreme southern South America has species typical of the cold waters of Antarctica (e.g., Muraenolepis spp.). Bathymetric distribution of the species The bathymetric distribution of the gadiforms reported herein is summarized in Figure 5. Gadiformes can be divided in two groups, according to their life style: 1) Bregmaceros is pelagic, and can only be found in the water column, far from the ocean floor; 2) Macrouridae, Merlucciidae and Gadidae are benthopelagic i.e. usually live and feed near the bottom, but can sometimes be found in the pelagic zone (e.g. Haedrich 1974). The depth range of Bregmaceros sp. was not included because it is pelagic and the depth observations can be biased by the method of sampling. Along the slope, the gadiforms had a gradual transition of species, similar to other groups already studied from the area (Mincarone et al. 2008; Melo et al. 2008). Following the classification provided by Haedrich and Merrett (1988) for strata of the continental slope, seven species are found exclusively in the upper slope ( m): Urophycis brasiliensis, Steindachneria argentea, Urophycis mystacea, Malacocephalus okamurai, Hymenocephalus cf. aterrimus, Merluccius hubbsi, and Coelorinchus marinii. Urophycis spp. and Merluccius hubbsi are representatives of shallow cold waters, and are more commonly found on the continental shelf. Five species are restricted to the middle slope ( m): Gadomus capensis, Haplomacrourus nudirostris, Coryphaenoides cf. leptolepis, Coryphaenoides thelestomus, and Gadomus arcuatus. Nine species range from the upper to the middle slopes: Laemonema goodebeanorum, Ventrifossa mucocephalus, Coelorinchus aff. carminatus, Malacocephalus laevis, Malacocephalus occidentalis, Nezumia atlantica, Gadella imberbis, Coelorinchus occa, and Hymenocephalus billsam. There are no species restricted to the lower slope ( m); and ten species range from the middle to the lower slopes: Macrosmia phalacra, Halargyreus johnsonii, Trachonurus sulcatus, Cetonurus globiceps, 42 Zootaxa Magnolia Press MELO ET AL.