Six new records of decapod crustacean species (Anomura, Brachyura) from southern Brazil

Abstract Five anomuran and one brachyuran species collected from continental shelf and slope off southern Brazil have their geographic ranges considerably extended southward: Neolithodes agassizii (Smith), Paguristes spectabilis McLaughlin and Provenzano, Pagurus heblingi Nucci and Melo, Phimochirus occlusus (Henderson), Mixtopagurus paradoxus A. Milne-Edwards, and the brachyuran Euchirograpsus antillensis Türkay. Pagurus heblingi was previously known only from its type locality (off Cabo Frio, Rio de Janeiro) and from off Espírito Santo. Paguristes spectabilis was known only from the holotype (off Trinidad, Caribbean Sea) and two additional females from off Rio de Janeiro. One female was found of the brachyuran Robertsella meridionalis Tavares and Gouvêa, a species previously known only from two males. Additionally, the occurrence of the hermit crabs Dardanus venosus (H. Milne Edwards) and Oncopagurus gracilis (Henderson), and the brachyuran Hexapanopeus paulensis Rathbun are confirmed for Santa Catarina. The record of the brachyuran Neopilumnoplax americana (Rathbun) from Santa Catarina (27°S) probably actually refers to Neopilumnoplax lipkeholthuisi Tavares and Melo, a species described from Rio Grande do Sul (33°S) and Mar del Plata (37°S).


INTRODUCTION
In 2009, the CEPSUL-ICMBio (Centro Nacional de Pesquisa e Conservação da Biodiversidade Marinha do Sudeste e Sul-Instituto Chico Mendes da Conservação da Biodiversidade) launched the project MOBIO (Monitoramento da Biodiversidade Marinha no Sul do Brasil) aimed at monitoring marine biodiversity in Santa Catarina, southern Brazil.From 2009 to 2011, as part of MOBIO, the research vessel "Soloncy Moura" was commissioned to conduct eleven oceanographic cruises along the shelf and upper slope of Santa Catarina in depths between 20 and 1000 m.The main results of the MOBIO Project concerning the decapod crustaceans were compiled by Boos et al. (2012), Misturini and Segal (2017), and references therein.
Among the decapod crustaceans amassed during the MOBIO in Santa Catarina there were five anomuran species collected between 21-500 m, which constitute new records for the region.The opportunity is taken to include one deep-water brachyuran species caught by the fishing vessel "Cordeiro de Deus I" off the coast of Rio Grande do Sul, whose geographic range is considerably extended southward.The study of this material forms the basis of this report.

MATERIAL AND METHODS
The material studied here was obtained with an otter-trawl and fishing pots (four pots per main line) baited with skipjack tuna on the research vessel "Soloncy Moura" in Santa Catarina (Fig. 1), and ottertrawl on the fishing vessel "Cordeiro de Deus I" off the coast of Rio Grande do Sul.
The specimens are deposited in the collections of the Museu de Zoologia, Universidade de São Paulo (MZUSP).Specimens from Santa Catarina and Rio Grande do Sul not previously reported in the literature are listed under "New material"; comparative specimens are listed when relevant.Remarks on morphology, taxonomy, geographic distribution and habitat are included when relevant.Updated geographic distribution are also included for all species.
Acronyms: TAAF MD55, refers to the deep-water expedition conducted off the southeastern coast of Brazil in depths down to 5,155 m (Guille and Ramos, 1987;Tavares, 1999).Guille and Ramos (1987) provided the bottom type for each location sampled by the TAAF MD55 expedition.Otherwise, information on bottom type was essentially obtained from the compilations by Melo (1996), Felder et al. (2009), and Nucci and Melo (2015).GEDIP (Grupo Executivo do Desenvolvimento da Indústria da Pesca) refers to a research program aimed at the evaluation of the fishing potential of Rio Grande do Sul (Brazil) and Maldonado (Uruguay); the oceanographic operations were conducted aboard R/V "Prof.W. Besnard" from Instituto Oceanográfico, Universidade de São Paulo (Miranda, 1971;Furtado, 1973;Miranda et al., 1973).REVIZEE (Recursos Vivos na Zona Econômica Exclusiva) was a research program carried out to evaluate the living resources in Brazil's exclusive economic zone.
Abbreviations: ARM, baited fishing trap ("pot"); cl, carapace length for Neolithodes agassizii (Smith, 1882), measured from the posterior margin of the orbit to the midpoint of the posterior margin of the carapace and given in millimeters (mm); DC, Charcot dredge; F/V, fishing vessel; R/V, research vessel; stn, sampling station."Lance" refers to a trawling operation.
Distribution and habitat.Western Atlantic: eastern coast of United States (North and South Carolina, Alabama), Gulf of Mexico, Bahamas, Caribbean Sea, Martinique, Colombia, Suriname, French Guiana, and Brazil (Rio Grande do Norte, Bahia, Espírito Santo, and Rio de Janeiro); from 332 to 2076 m (Serejo et al., 2007;Alves-Júnior et al., 2019, and references therein).This is the first record of Nl. agassizii from Rio Grande do Sul.
Remarks.Males and females of Nl. agassizii can grow as large as 167 mm and 154 mm in carapace length, respectively (Macpherson, 1988).Small specimens are known to differ from larger ones in being proportionally narrower and in having stronger development of the spinulation of carapace and appendages (Smith, 1882: plate I;Macpherson, 1988: fig. 13, plate 2C;Alves-Júnior et al., 2019: fig. 1a).The adult and juvenile males MZUSP 25992 and MZUSP 42409 (Fig. 2A, B), respectively, agree well with N. agassizii as understood by Macpherson (1988).Lianos et al. (2017) recorded an adult male of Neolithodes asperrimus Barnard, 1947 from Santa Catarina caught in depths between 250-500 m.This species was previously known from the eastern Atlantic, between 250-2000 m.Neolithodes asperrimus and Nl.agassizii are morphologically closely related but differ in some aspects including the spinulation of the walking legs and the proportions of the third walking leg relative to carapace length (see Macpherson, 1988).The adult male MZUSP 25992 can be assigned to Nl. agassizii and differs from the male of N. asperrimus reported by Lianos et al. (2017) in having: 1) the main spines on the walking legs distinctly larger (vs.spines tending to be reduced to thick granules or very short spines in Nl. asperrimus); 2) the third walking legs more than three times the carapace length (vs.third walking legs 2.5 times the carapace length in N. asperrimus); and 3) the merus of the third walking legs longer than carapace length (vs.merus of the third walking legs shorter than carapace length in males in Nl. asperrimus).

Distribution and habitat. Western Atlantic:
Caribbean Sea (Trinidad), and Brazil (off Rio de Janeiro); on soft mud bottoms; from 137 to 430 m (Lemaitre and Tavares, 2015).This is the first record of P. spectabilis from Santa Catarina.
Remarks.Paguristes spectabilis was only known from the holotype (a female caught off Trinidad Island on the northern edge of the South America mainland) and two additional females reported from off Rio de Janeiro by Lemaitre and Tavares (2015).The eight additional females reported here extend the distribution of this species much farther south.Lemaitre and Tavares (2015) noted that the specimens from off Rio de Janeiro differed from the holotype in lacking spines on the lateral margins of the anterior lobes of the telson, whereas in the holotype the telson has one to three small spines on the left and right lobes, respectively.The specimens from off Santa Catarina, while fitting well with Pt. spectabilis, also lack spines on the lateral margins of the anterior lobes of the telson.Males of this species remain unknown.
Distribution and habitat.Western Atlantic: Brazil (from Rio de Janeiro to Rio Grande do Sul), Uruguay, and Argentina; in fine sand and mud bottoms; from 10 to 50 m (Melo, 1999;Nucci and Melo, 2007).
Remarks.The female MZUSP 37644 provides the opportunity to corroborate the presence of Pr. exilis in Santa Catarina, whose record has been repeated from source to source, but no voucher specimen has been hitherto located to corroborate it (Hebling and Rieger, 1986;Rieger, 1997;1998;Melo, 1999;Nucci and Melo, 2007;Boos et al., 2012;Branco et al., 2015;Rodrigues Filho et al., 2016;Stanski et al., 2016).The specimen MZUSP 37644 match well with the diagnostic characters provided by Nucci and Melo (2007) in having the ocular peduncle (including cornea) less than 3 times the diameter of the cornea, the ultimate segment of the antennular peduncle shorter than the ocular peduncle (including cornea), and the right cheliped with the carpus covered by rounded tubercles.
Diagnosis.See Nucci and Melo (2003;2007).Distribution and habitat.Western Atlantic: Brazil (Espírito Santo, Rio de Janeiro, and Santa Catarina); in soft mud bottoms with pteropods and foraminiferans; from 300 to 600 m (Lemaitre and Tavares, 2015).This is the first record of P. heblingi from Santa Catarina.
Remarks.This species was previously known only from its type locality, off Cabo Frio, Rio de Janeiro (Nucci and Melo, 2003) and from off Espírito Santo (Lemaitre and Tavares, 2015).The present material considerably extends the species' geographic range southward.Lemaitre (2001), and Nucci and Melo (2011).

Diagnosis. McLaughlin and
Distribution and habitat.Western Atlantic: eastern coast of United States (from North Carolina to Florida), Gulf of Mexico, Caribbean Sea, Antilles, and Brazil (Amapá to Santa Catarina); in soft mud bottoms from 11 to 1020 m (Lemaitre and Tavares, 2015).
Remarks.The record of Py. discoidalis from Santa Catarina has been hitherto based on only one small, damaged specimen reported by Nucci and Melo (2011).The present new material confirms Santa Catarina as the southern limit of Py. discoidalis.
Distribution and habitat.Western Atlantic: eastern coast of United States (from off North Carolina), Gulf of Mexico, Bahamas, and Caribbean Sea to Brazil (Amapá, Pará, Ceará, Rio de Janeiro, São Paulo), from 194 to 567 m (Lemaitre and Tavares, 2015).This is the first record of M. paradoxus from Santa Catarina.
Remarks.This species was previously known from the Straits of Florida and Gulf of Mexico to Brazil and as far south as the coast of Santos, São Paulo (Lemaitre and Tavares, 2015).The present record extends the range of M. paradoxus southward to Santa Catarina.
Distribution and habitat.Western Atlantic: eastern coast of United States (North and South Carolina, Florida), Gulf of Mexico, Antilles, Colombia, Venezuela, and Brazil (São Paulo and Rio Grande do Sul), in soft and hard bottoms from 15 to 430 m (Felder et al., 2009;Melo, 2010).This first record of E. antillensis from Santa Catarina begins to fill a gap in its geographic range between São Paulo and Rio Grande do Sul.
Remarks.The diagnostic characters for E. antillensis, more easily recognized in adult specimens, are less evident in juveniles.In the juvenile from Santa Catarina the second pereopod is provided with a small subdistal spine on the merus ventrolateral margin, whereas that margin is granulated in the adults.
Distribution and habitat.Western Atlantic: South Carolina, Florida, Gulf of Mexico, and Brazil (from Pará to Santa Catarina) in sand, broken shells and rocky bottoms and in association with ascidians, bryozoans, from the intertidal to 34 m (Melo, 1996;this study).
Remarks.The records of Hexapanopeus paulensis from Santa Catarina are unclear due to lack of voucher material (Melo, 1996;Bouzon and Freire, 2007;Almeida and Coelho, 2008;Boos et al., 2012).Its occurrence in Santa Catarina in now conclusive.
Remarks.Robertsella meridionalis is known so far only from two males: the holotype caught in Santa Catarina and a young male from much farther north in Natal, Rio Grande do Norte (Tavares and Gouvêa, 2013;Alves-Júnior et al., 2021).Poupin and Corbari (2016) provisionally attributed one specimen (sex not mentioned) from Guadeloupe to R. meridionalis.Here we report on a young female and an additional young male both from Santa Catarina.Both specimens agree well with the holotype, except for the male abdominal suture 3/4 which is only evident as a lateral notch in the young male, whereas it is evident along its entire

New crustacean records from southern Brazil
Nauplius, 31: e2023015 length in the adult holotype.The young female has heterochelous chelipeds and a stridulatory apparatus similar to that described for the male by Tavares and Gouvêa (2013).Boos et al. (2012) listed 280 species of decapods from Santa Catarina based on records from the literature complemented by the examination of specimens from various collections.Of these, eight marine species were recorded for the first time from Santa Catarina.However, the authors cautioned that several records (unspecified) were unverifiable for lack of voucher material.Such is the case of the brachyuran H. paulensis, only now confirmed for Santa Catarina (see above).

DISCUSSION
The diogenid hermit crab Dardanus venosus (H.Milne Edwards, 1848), recently recorded from Santa Catarina on account of a single male by Lima et al. (2019), is here confirmed for Santa Catarina based on an additional male (MZUSP 36037) caught in the Arvoredo Island.Dardanus venosus is a wellknown species commonly found from the eastern coast of the United States, Bermuda, Florida, Antilles, northern coast of South America and Brazil (Amapá to São Paulo).It is also found in the oceanic islands of Fernando de Noronha, Rocas Atoll and Trindade and Martin Vaz and the seamounts Almirante Saldanha, Dogaressa and Minerva (Lima et al., 2019).Dardanus venosus is apparently less common southwards to São Paulo, where its presence went unrecorded and unnoticed previously (e.g., Hebling and Rieger, 1986;Bouzon and Freire, 2007;Boos et al., 2012).Santa Catarina may well be the southernmost limit to the distribution of D. venosus.
The amphi-Atlantic deep-water parapagurid Oncopagurus gracilis (Henderson, 1888) is known to occur in the eastern Atlantic in the Gulf of Guinea and off Angola.In the western Atlantic, O. gracilis has a much wider distribution ranging from the Straits of Florida and Gulf of Mexico in the northern hemisphere to off the coast of Brazil (Rio de Janeiro and Santa Catarina), where it inhabits soft mud with rubble between 146 and 900 m (Nucci et al., 2004;Lemaitre and Tavares, 2015, respectively).Nucci et al. (2004) mentioned two specimens from Santa Catarina without further specifics, a record not verified by Boos et al. (2012) for lack of voucher material.The occurrence of O. gracilis was confirmed in this study, based on 1 specimen (MZUSP 37704, Santa Catarina, 27°35.611"S47°09.351'W)caught by the R/V "Soloncy Moura", MOBIO IV, Lance 58, 02.vii.2010, between 426 and 460 m.The specimen reported here fits well with the description of O. gracilis and also agrees with the diagnostic characters provided by Lemaitre (2014).Oncopagurus gracilis and Oncopagurus bicristatus (A .Milne-Edwards, 1880) are similar and can occur sympatrically.However, they can be differentiated from each other by the shape of the right palm, armature of antennal acicles, and degree of development of the first and second gonopods in males (Lemaitre, 2014).Bouzon and Freire (2007) recorded Neopilumnoplax americana (Rathbun, 1898) from Santa Catarina (Arvoredo Island, 27°17'7''S 48°25'30''W) without further specifics.This record, uncritically accepted by Boos et al. (2012), probably actually refers to Np. lipkeholthuisi Tavares and Melo, 2010, a species described from Rio Grande do Sul (33°S) and Mar del Plata (37°S) (Tavares and Melo, 2010).

Figure 1 .
Figure 1.Sampling operations.A, The R/V "Soloncy Moura" at sea during the MOBIO Project conducted in Santa Catarina, southwestern Brazil, February 2011; B, C, Fishing operations with otter-trawl and fishing pots baited with skipjack tuna conducted by the R/V "Soloncy Moura" in December 2009.Photographs from the CEPSUL collection.

crustacean records from southern Brazil
(Nucci and Melo, 2011)at.Western Atlantic: Antilles and Brazil (from Pernambuco to São Paulo, and Santa Catarina); in Halimeda rich sand (present study); from 100 to 640 m(Nucci and Melo, 2011).This is the first record of Ph. occlusus from Santa Catarina.