ISSN 1015-3233
COMISIÓN TÉCNICA MIXTA DEL FRENTE MARÍTIMO Impreso en 2019 en PRONTOGRÁFICA Cerro Largo 850 - Tel.: 2902 3172 Montevideo Uruguay E-mail: [email protected] ii ISSN 1015-3233
COMISIÓN TÉCNICA MIXTA DEL FRENTE MARÍTIMO www.ctmfm.org
FRENTE MARÍTIMO VOLUMEN 26 - ABRIL 2019
iii Index
INTRODUCTION 3
STUDY AREA 4
MATERIALS AND METHODS 6
RESULTS 8
SYSTEMATIC ACCOUNT 12
List of species by systematic order. Symbols preceding species denotes: ►listed by Boschi et al. (1992); ● listed by Zolessi and Philippi (1995); † very dubious or incorrect reports on both lists
● Aristaeopsis edwardsiana 12
Plesiopenaeus armatus 12
Pseudaristeus speciosus 13
Benthesicymus brasiliensis 13
Gennadas brevirostris 13
Gennadas gilchristi 13
Gennadas tinayrei 13
Gennadas valens 14
►● Artemesia longinaris 14
Funchalia villosa 15
● Funchalia woodwardi 15
● Parapenaeus americanus 15
Penaeopsis serrata 16
†● Penaeus brasiliensis 16
● Penaeus notialis 16
►● Penaeus paulensis 17
Farfantepenaeus paulensis 17
● Penaeus schmitti 17 iv ● Xiphopenaeus kroyeri 17
►● Pleoticus muelleri 18
Solenocera necopina 19
● Belzebub faxoni 19
Allosergestes pectinatus 19
Allosergestes sargassi 20
Deosergestes corniculum 20
Deosergestes curvatus 20
Deosergestes disjunctus 20
►● Eusergestes antarcticus 20
Gardinerosergia splendens 21
Neosergestes edwardsii 21
Parasergestes armatus 21
Parasergestes vigilax 22
►● Peisos petrunkevitchi 22
Petalidium foliaceum 23
►● Phorcosergia potens 24
Prehensilosergia prehensilis 25
● Sergestes atlanticus 25
Sergia Stimpson 25
Sergia laminata 25
►● Pasiphaea acutifrons 26
Pasiphaea barnardi 26
Pasiphaea dofleini 26
Pasiphaea rathbunae 26
Pasiphaea scotiae 26
v ► Acanthephyra pelagica 27
Acanthephyra quadrispinosa 28
Systellaspis debilis 28
Nematocarcinus lanceopes 29
Nematocarcinus longirostris 30
Nematocarcinus tenuipes 31
Campylonotus arntzianus 32
► Campylonotus capensis 33
►● Campylonotus semistriatus 34
► Campylonotus vagans 35
Palaemon macrodactylus 36 northropi 37
►● Alpheus puapeba 37
Alpheus pouang 38
► Betaeus lilianae 38
► Betaeus truncatus 39
► Chorismus antarcticus 39
► Chorismus tuberculatus 40
Exhippolysmata oplophoroides 40
►● Latreutes parvulus 40
►● Merhippolyte americana 40
► Nauticaris magellanica 41
Eualus amandae 42
Eualus dozei 42
Eualus kinzeri 42
Lebbeus antarcticus 42
vi Processa guyanae 43
►● Processa hemphilli 43
Processa profunda 43
►● Austropandalus grayi 44
►● Pandalopsis ampla 45
Pantomus parvulus 46
Stylopandalus richardi 46
►● Aegaeon boschii 47
● Lissosabinea cf tridentata 48
► Notocrangon antarcticus 48
● Philoceras gorei 48
►● Metanephrops rubellus 49
►● Thymops birsteini 50
Thymops takedai 50
Thymopsis nilenta 51
►Anacalliax argentinensis 51
► Notiax brachyophthalma 51
Notiax santarita 51
● Poti gaucho 52
►● Sergio mirim 52
Upogebia australis 52
► Scyllarides deceptor 53
Pentacheles validus 54
Stereomastis suhmi 55
► Munida gregaria 56
● Munida iris 57
vii † Munida flinti 57
† Munida forceps 57
† Munida irrasa 57
► Munida spinosa 58
Liopetrolisthes patagonicus 58
►● Pachycheles chubutensis 59
►● Pachycheles laevidactylus 60
● Polyonyx gibbesi 61
● Porcellana sayana 61
►● Blepharipoda doelloi 62
►† Emerita analoga 63
►● Emerita brasiliensis 64
► Lithodes confundens 65
►● Lithodes santolla 66
► Lithodes turkayi 67
Neolithodes diomedeae 68
Neolithodes yaldwyni 68
► Paralomis anamerae 68
Paralomis birsteini 68
►● Paralomis formosa 69
► Paralomis granulosa 70
Paralomis longidactylus 70
► Paralomis spinosissima 71
Paralomis tuberipes 72
Paralomis zealandica 72
►● Loxopagurus loxochelis 73
viii ► Paguristes foresti 73
►● Paguristes robustus 74
Paguristes scarabinoi 74
►● Pagurus comptus 74
►● Pagurus criniticornis 75
►● Pagurus exilis 75
► Pagurus forceps 75
● Pagurus leptonyx 75
†● Pagurus provenzanoi 76
● Pagurus trichocerus 76
►● Propagurus gaudichaudii 77
► Sympagurus dimorphus 78
● Homola minima 79
● Latreilla willamsi 79
● Hepatus pudibundus 80
► Acanthocyclus albatrossis 80
►● Corystoides abbreviatus 81
● Acanthocarpus alexandri 82
Acanthocarpus meridionalis 82
►● Peltarion spinosulum 83
Metacarcinus edwardsii 84
Ethusina 85
Ethusina abyssicola 85
►● Danielethus crenulatus 86
►● Danielethus patagonicus 87
● Chasmocarcinus typicus Rathbun 88
ix ● Neopilumnoplax lipkeholthuisi 88
► Ebalia rotundata 88
● Myropsis quinquespinosa 89
● Persephona mediterranea 89
►† Leucosia planata 89
►● Leucippa pentagona 90
●† Taliepus dentatus and T. marginatus 91
Lepteces ornatus 91
►● Libidoclea granaria 92
● Libinia ferreirae 93
►● Libinia spinosa 93
►● Pelia rotunda 94
►● Rochinia gracilipes 95
● Stenocionops furcatus 96
● Anisonotus atlanticus 96
► Eurypodius latreillii 97
Eurypodius longirostris 98
● Stenorhynchus seticornis 98
►● Collodes rostratus 99
►● Leurocyclus tuberculosus 100
Pyromaia tuberculata 101
Amphithrax besnardi 102
Hyas araneus 102
► Halicarcinus planatus 103
● Mesorhoea sexspinosa 104
● Piloslambrus guerinii 104
x ● Spinolambrus fraterculus 104
● Spinolambrus meridionalis 104
►● Pilumnus reticulatus 105
Carcinus maenas 106
►● Ovalipes trimaculatus 107
►● Chaceon notialis 108
►● Coenophthalmus tridentatus 109
● Arenaeus cribrarius 110
►● Callinectes sapidus 110
● Callinectes danae 111
● Achelous spinicarpus 111
● Achelous spinimanus 111
►● Pilumnoides hassleri 112
● Cyrtoplax spinidentata 113
►● Acantholobulus schmitti 113
● Eurypanopeus depressus 114
● Hexapanopeus paulensis 115
►● Panopeus meridionalis 115
● Tetraxanthus rathbunae 116
Speocarcinus dentatus 116
● Actaea acantha 116
● Pachygrapsus transversus 117
● Planes major 117
Planes marinus 117
►● Armases rubripes 118
●† Armases miersii 118
xi ►● Neohelice granulata 119
►● Cyrtograpsus angulatus 120
►● Cyrtograpsus altimanus 121
►● Cyrtograpsus affinis 122
● Ocypode quadrata 122
►● Leptuca uruguayensis 123
►● Austinixa patagoniensis 124
► Pinnixa brevipollex 124
● Pinnixa chaetopterana 124
►● Pinnixa rapax 125
● Pinnixa sayana 125
Pinnixa valdiviensis 125
► Calyptraeotheres garthi 126
► Dissodactylus crinitichelis 127
► Fabia emiliai 127
► Pinnaxodes chilensis 127
►● Tumidotheres maculatus 128
DISCUSSION 129
SUPPLEMENTARY ONLINE FILES 137
ACKNOWLEDGEMENTS 137
REFERENCES 138
xii Prólogo
Entre los días 21 y 23 de noviembre de 2017 se llevó a cabo en Tigre, Provincia de Buenos Aires, el 18º Simposio Científico de la Comisión Técnica Mixta del Frente Marítimo. En esa oportunidad el Simposio estuvo dedicado a conmemorar el 40º aniversario de la creación de la Comisión Técnica Mixta del Frente Marítimo (CTMFM) que tuvo lugar el 16 de agosto de 1976 y de su primera Sesión Plenaria, que tuvo lugar el 4 de febrero de 1977.
El volumen 25 de FRENTE MARÍTIMO publicado en 2018 contiene, con la única excepción de la obra que aquí prologamos, todos los trabajos presentados en el Simposio que fueran oportunamente enviados al Comité Editor y aceptados para su publicación.
El presente volumen se destina entonces, en exclusividad, a publicar la mencionada obra: el Catalogo anotado y bibliografía de los camarones, langostas, cangrejos y formas afines, estuarinas y marinas (Crustacea: Decapoda), de Argentina y Uruguay (Océano Atlántico Sudoccidental).
Hace 27 años la CTMFM publicó el Catálogo ilustrado de los crustáceos estomatópodos y decápodos marinos de Argentina (Boschi et al., 1992) registrados en aguas costeras, de plataforma y de profundidad, que incluía 93 especies de decápodos. Poco después se publicó una lista de los decápodos de Uruguay (Zolessi y Philippi, 1995) con 107 especies. Desde entonces, el advenimiento de técnicas moleculares, la revalorización de la taxonomía y el incremento de las invasiones biológicas debido al intercambio comercial y al cambio climático global, hacen necesaria la revisión de los trabajos anteriores. Este Catálogo incluye la adición y remoción de especies, una actualización de la taxonomía y sistemática de los grupos involucrados, además de brindar listas bibliográficas sobre los grupos y especies más importantes.
No dudamos que será un valiosísimo material de referencia para todos los científicos, técnicos y público interesados en la biogeografía y sistemática de crustáceos.
xiii Editorial Committee
- Dr. Marcos Domingos Siqueira Tavares
- Prof. Fabrizio Scarabino
xv Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Eduardo D. Spivak1, Nahuel E. Farías1, Emiliano H. Ocampo1, Gustavo A. Lovrich2 y Tomás A. Luppi1 1Instituto de Investigaciones Marinas y Costeras (IIMyC), Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET)- Universidad Nacional de Mar del Plata (UNMdP). [email protected] 2Centro Austral de Investigaciones Científicas (CADIC) Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET)
SUMMARY: Twenty seven years ago, the Co- shelf and slope, the contiguous abysal plains, misión Técnica Mixta del Frente Marítimo pu- the Malvinas, Georgias del Sur and Sandwich del blished a catalogue of marine decapod and Sur Islands and the Scotia Sea. stomatopod crustaceans registered in coasts, continental shelf and slope, and deep waters of A total of 211 species of decapod crustaceans Argentina (Boschi et al., 1992), that included 93 are listed in this catalogue: 78 shrimps and species of decapods. Later, a list of decapods of prawns (Dendrobranchiata and Caridea), 13 Uruguay appeared with 107 marine and estua- lobsters and ghost shrimps, (Astacidea, Axiidea, rine species (Zolessi and Philippi, 1995). Since Gebiidea, Achelata, Polychelidae), 40 king crabs, then, the acknowledgement of the importance hermit crabs and other “not true” crabs (Anomu- of biodiversity for the human kind and the many ra) and 80 true crabs (Brachyura). Three families processes affecting it, such as the growing rates of Dendrobranchiata and 20 of Pleocyemata of biological invasions and global climate chan- (4 of Caridea, and 16 of Brachyura) not cited ge, make necessary to review previous knowled- by Boschi et al. (1992) have been registered, as ge on local biodiversity. Moreover, the incresing well as 2 infra-orders of Pleocyemata (Gebiidea use of molecular techniques in taxonomic revi- and Polychelidae). The additions to Boschi et al. sions have unveiled the existence of misidenti- (1992) list are of species: 1) only cited previously fications, cryptic species and species complex in waters of Uruguay, 2) from deep and/or in numbers that significantly alter our previous southern waters, 3) coastal invaders and 4) new, considerations on species richness at the local described for the first time in the southern tip of and regional scales. The present catalogue in- America. Eight species cited in previous papers cludes the addition and removal of species and have been excluded due to synonymy or mis- an actualization of the published bibliography takes in geographic location, and 14 have been related with them. The study area comprises es- renamed to date. tuaries and marine waters of the Southwestern Atlantic Ocean in front of Uruguay and Argen- Key words: shrimps, lobsters, crabs, Crustacea, tina (33-60ºS), including the coasts, continental Decapoda, Southwestern Atlantic Ocean
1 FRENTE MARÍTIMO Publicación de la Comisión Técnica Mixta del Frente Marítimo
RESUMEN: Catálogo anotado y bibliografía de Océano Atlántico Sudoccidental frente a Uru- los camarones, langostas, cangrejos, y formas guay y Argentina (33-55ºS), incluyendo la corres- afines (Crustacea: Decapoda) de Argentina y pondiente plataforma, talud y fondos abisales Uruguay (Océano Atlántico Sudoccidental). adyacentes, las islas Malvinas, Georgias del Sur y Hace 27 años la Comisión Técnica Mixta del Fren- Sándwich del Sur, y el mar de Escocia. te Marítimo publicó el “Catálogo Ilustrado de los Crustáceos Estomatópodos Decápodos Marinos” Un total de 211 especies de crustáceos decápo- (Boschi et al., 1992) registrados en aguas coste- dos integran la presente lista: 78 de langostinos ras, de plataforma y de profundidad de Argenti- y camarones (Dendrobranchiata y Caridea), 13 na, que incluía 93 especies de decápodos. Poco de langostas y camarones fantasmas (Astacidea, después apareció una lista de los decápodos de Axiidea, Gebiidea, Achelata, Polychelidae), 40 de Uruguay (Zolessi y Philippi, 1995) con 107 espe- centollas, ermitaños y afines (Anomura) y 80 de cies marinas y estuariales. Desde entonces, el re- cangrejos verdaderos (Brachyura). Se registraron conocimiento de la biodiversidad como capital 3 familias de Dendrobranchiata y 20 de Pleocye- de la humanidad y de los numerosos procesos mata (4 de Caridea, y 16 de Brachyura) no citadas que la afectan, como las crecientes tasas de in- por Boschi et al. (1992), así como 2 infra-órdenes vasiones biológicas y el cambio climático glo- más de Pleocyemata (Gebiidea y Polychelidae). bal, hacen necesario revisar los conocimientos Las adiciones a la lista de Boschi et al. (1992) pro- previos sobre la biodiversidad local. Además, el vienen de especies: 1) solo citadas en aguas de creciente uso de técnicas moleculares en las re- Uruguay, 2) de aguas profundas y/o australes, 3) visiones taxonómicas ha revelado la existencia invasoras costeras y 4) nuevas, descriptas en el de identificaciones erróneas, especies crípticas extremo sur de América. Ocho especies citadas y complejos de especies en números que alte- en los trabajos previos han sido excluidas debido ran significativamente nuestras consideraciones a sinonimias o errores en la ubicación geográfica previas sobre la riqueza faunisitica a escala local y 14 han sido renombradas a la fecha. y regional. Este catálogo incluye adiciones y re- mociones de especies y una actualización biblio- Palabras clave: camarones, langostas, cangre- gráfica del conocimiento de las mismas. El área jos, Crustacea, Decapoda, Océano Atlántico Su- de estudio abarca estuarios y costas marinas del doccidental
2 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
INTRODUCTION
Twenty seven years have passed since the publication of the “Catálogo Ilustrado de los Crustáceos Estomatópodos y Decápodos Marinos” (Illustrated catalogue of stomatopod and decapod crusta- ceans of the marine littoral of Argentina, Boschi et al., 1992). This seminal work for the Southwestern Atlantic crustacean diversity was leaded by Enrique Eduardo Boschi and included 93 species of De- capoda (two of them dismissed due to error of location and fusion of two species) and 3 of Stomato- poda, registered in coastal, shelf and deep waters along the ca. 6500 km of coastline washed by the Southwestern Atlantic Ocean, from the mouth of the Río de la Plata to the Canal Beagle. The studied area also comprised the Malvinas, Georgias del Sur and Sandwich del Sur Islands. These waters har- bor cold-temperate and warm-temperate biotas, usually assimilated to the Magellanic and Argen- tine Biogeographic provinces, respectively, corresponding to the so-called Patagonian Shelf Large Marine Ecosystem (PSLME; Boschi, 2000; Orenszanz et al., 2002 and references thereof). Three years later, a cheklist of decapods from Uruguay was published (Zolessi and Philippi, 1995), and partially reviewed a decade after by Scarabino (2006). In 2000, the first electronic checklist of the Chilean decapods appeared (Retamal, 2000), thus completing the catalogues for the southern tip of South America. In addition, Antarctic and sub-Antarctic Decapods have been reviewed by Basher and Cos- tello (2014) and Griffiths et al. (2014); more recently, a checklist of shrimps of the Southern Ocean has been published, including many species found in the Southwestern Atlantic area studied here (D’Udekem D’Acoz and De Grave, 2018).
Due to large inter-individual and ontogenic variation in diagnostic features, the existence of cases of morphotypes, evolutionary convergence and/or morphological stasis, the identification of decapods based solely in morphological features (traditional taxonomy) can be problematic in many cases, even for experts. During the last decade the irruption of DNA taxonomy and barcoding has proved to be effective tools to aid species identification and discovery, contributing to perform biodiversity assessments in a timeframe that is in line with the current accelerating threats to biodiversity such as climate change and biological invasions. However, both DNA barcoding and traditional taxonomy have virtues and drawbacks, so they become more effective when used synergistically, in what is named an “integrative taxonomy” (Lipscomb et al., 2003; Dayrat, 2005; Will et al., 2005).
In the last 27 years our knowledge on the taxonomic composition of the decapod crustacean fauna experienced several changes, due not only to the new taxonomic tools mentioned above, but also to new classifications (Martin and Davis, 2001; Ng et al., 2008; De Grave et al., 2009, 2011), changes in taxonomy and systematic placement, new findings and, also, arrival of exotic species. All these changes motivated an update of monographic works dealing with large geographic areas such as the temperate eastern United States studied, firstly by Williams (1984) and then revised by Nizinski (2003), or the Peninsula Ibérica (Zariquiey Álvarez, 1968) recently revisited by Marco-Herrero et al. (2015).
The main goal of this study was to generate a revised and up-to-date compendium of the marine decapod species reported in cold to temperate waters of Argentina and Uruguay and the adjacent ocean waters. To that, we have perfomed a review of the available literature occurrence of decapod crustaceans within the study area. The result is resulting in a comprehensive list of the species regis- tered for this region, with an updated classification and the addition and removal of several species.
3 FRENTE MARÍTIMO Publicación de la Comisión Técnica Mixta del Frente Marítimo
Thus, this study serves as the baseline and guide for further review of our knowledge of the decapod diversity under an integrative taxonomy view and, we hope, will foster and help to complete and curate extant biological collections. STUDY AREA
The geographic scope of this study can be broadly defined as the area encompassing the Patagonian Shelf Large Marine Ecosystem (PSLME) and the Scotia Sea, including the Malvinas Islands and the archipelagos conforming the Scotia arc.
The Patagonian Shelf Large Marine Ecosystem extends from the border of Uruguay and Brazil (on the coast: 33°44’S; ca. 35ºS on the shelf border) to the south of Burdwood Bank, in Argentina (55°S) and has a total area of about 1,200,000 km2. The continental shelf extending seawards from southeastern Brazil, Uruguay and Argentina widens progressively to the south, where it reaches a maximum width of some 850 km; it is one of the world’s most productive and complex marine systems (Bisbal, 1995; Acha et al., 2004; Heileman, 2009). The coastline is relatively straight from 23°S through South Brazil and Uruguay, to the Río de la Plata. From the mouth of this huge river to Tierra del Fuego, there are several major coastal geographic features: Bahia Blanca, Golfo San Matías, Golfo Nuevo and Golfo San Jorge, and Península Valdés (Fig. 1).
13º45´S 27º07´W 26º20´S 42º29´W Uruguay Am Ro Pa Ml Ma Ca Pi Argentina Mo Ce RGN BA Pb RN Brazil Pe Al Se Ba Ch South Western ES RJ Atlantic SP Pa SC SC
RGS Malvinas Uruguay TF Argentina Georgias del sur
Sandwich del sur 60ºS Southern 63º01´S 96º39´W 63º43´S 77º08´W Ocean Figure 1. Geographic references in the study area and adjacent areas. Abbreviatures are included in Table 1. Modified from Google Earth.
4 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Table 1. Political divisions in the study area used to describe geographical distribution of species in the text, with its approximate latitudinal limits.
Northern Southern Country States/Departments/Provinces Abbreviature limit limit Amapá AP 4ºN 1ºN Pará PA 1ºN 1ºS Maranhão MA 1ºS 2ºS Piauí PI 3ºS 3ºS Ceará CE 3ºS 4º50’S Rio Grande do Norte RN 4º50’S 6º30’S Paraíba PB 6º30’S 7º30’S Pernambuco PE 7º30’S 9ºS Brazil Alagoas AL 9ºS 10º30’S Sergipe SE 10º30’S 11º30’S Bahia BA 11º30’S 18º20’S Espírito Santo ES 18º20’S 21º20’S Rio de Janeiro RJ 21º20’S 23º25’S São Paulo SP 23º25’S 25º15’S Paraná PR 25º15’S 26ºS Santa Catarina SC 26ºS 29º20’S Rio Grande do Sul RS 29º20’S 33º50’S Rocha RO 33º48’S 34º49’S Maldonado ML 34º49’S 34º48’S Uruguay Canelones CA 34º48’S 34º54’S Montevideo MO 34º54’S 34º55’S Buenos Aires BA 35º40’S 41ºS Río Negro RN 41ºS 42ºS Argentina Chubut CH 42ºS 46ºS Santa Cruz SC 46ºS 52º30’S Tierra del Fuego TF 52º30’S 55º10’S
The northern limit of this catalogue was established at the latitude of Chuy lighthouse (the coastal boundary between Uruguay and Brazil: 33º44’S; 53º22’W). However, from this point the maritime boundary follows a direction nearly perpendicular to the general line of the coast (running on a bearing of 128 sexagesimal degrees, counting from true north) (Fig. 2). The species (especially oceanic) found in front of Urugayan coasts, in the area between latitude 33º44’S and the political boundary (Brazilian and international waters) were included with the corresponding explanation.
5 FRENTE MARÍTIMO Publicación de la Comisión Técnica Mixta del Frente Marítimo
30º33´S 43º10´W
33º44´S
Maritime boundary
38º46´S 62º17´W Figure 2. Maritime boundary between Uruguay and Brazil.
To the east, the study area is delimited by the Scotia arc in its southern part, and along the PSLME, by the subantarctic Malvinas current that flows northwards over the continental shelf break; this wind-driven current meets the opposite flow of the Brazil subtropical current at 36°S (Brazil-Malvinas Confluence) where both turn offshore in a series of ample meanders (Fig. 1). PSLME is also influenced by low salinity coastal waters and the upwelling of cold Antarctic waters (Acha et al., 2004; Heileman, 2009).
Lastly, the southern limit of this catalogue was established at 60ºS (following The Antarctic Treaty, https://www.ats.aq/e/ats.htm) which is generally used to set the political division for the Southern or Antarctic Ocean, and coincides with the natural bio-oceanographic boundary delimited by the Antarctic convergence within the longitudinal range of this study. MATERIALS AND METHODS
The present list includes marine (from deep water to intertidal), and brackish water (estuaries, costal lagoons, marshes) species and is based on a review of publications dealing on the systematics and distribution of marine decapods of Argentina, Uruguay and the Southern Ocean near the southern tip of America (Antarctic waters), with emphasis on those that were published after Boschi et al. (1992). The review of Uruguayan decapods by Zolessi and Philippi (1995) has several mistakes and, consequently, the original literature has been revisited whenever available. In addition, the likely southern extension of the distributional range of species of Southern Brazil to Uruguayan waters has been evaluated on the basis of the Catalogue of Crustacea of Brazil edited by Young (1998) and the contributions of Bordin (1987); Cardoso (2009, 2013); Christoffersen (1979, 1982b, 1988); Coelho (1997a, 2005); Cunha et al. (2015); D’Incao (1997, 1998, 1999a); D’Incao and Martins (2000);
6 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Lemaitre and Tavares (2015); Magalhaes et al. (2016); Mantelatto et al. (2006, 2007, 2009b, 2014); Martins and D’Incao (1996); Melo (1996, 2006, 2010), Melo et al. (2003); Nucci and Melo (2007, 2011, 2015); Prado and Melo (2002); Rodrigues (1971); Scarabino (2006); Serejo et al. (2007); Soledade and Almeida (2013); Tavares and Serejo (2007); Veloso and Melo (1993) and Vezzani (2007). Data on temperate pelagic shrimps has been completed with Judkins (2014) and the online database of the Invertebrate Zoology Collection, National Museum of Natural History, Smithsonian Institution, denoted as “USNMdb” (http://collections.nmnh.si.edu/search/iz/). The following databases were also used to complete or verify the information: World Register of Marine Species, WoRMSdb (http:// www.marinespecies.org), Ocean Biogeographic Information System, Obisdb (http://www.iobis.org), Global Biodiversity Information Facility, GBIFdb (http://www.gbif.org) and National Institute of Water and Atmospheric Research of New Zealand, NIWAdb (https://www.niwa.co.nz).
The systematic classification essentially follows Nget al. (2008); Chan (2010); McLaughlin et al. (2010); Okawa and McLaughlin (2010); De Grave and Fransen (2011); but also considers later changes in particular taxa (Vereshchaka, 2000; Guinot et al., 2013; Vereshchaka et al., 2014, 2016; De Grave et al., 2014; Davie et al. 2015; Shih et al., 2016). As in Marco-Herrero et al. (2015) paper, superfamilies are listed by systematic order following the Sections and Subsections as currently accepted, and in alphabetical order within them. Families, genera and species are also listed in alphabetical order within their respective superfamilies and families. A list of references of all taxonomic names is included in the Supplementary Online File 1. Symbols preceding species denotes: ►listed by Boschi et al. (1992); ● listed by Zolessi and Philippi (1995); † very dubious or incorrect reports on both lists. Vernacular names are included at familiar, generic or specific when corresponds; Dendrobranchata according to Pérez Farfante (1988).
Specimens collected opportunistically during different campaigns and cruises were used to add information on molecular taxonomy. After preliminary (morphology based) species identification, some specimens were photographed alive to show living colour, and muscle tissue samples taken from up to five individuals per species for further genetic analyses. Then, genomic DNA was extracted and the partial fragment of the mitochondrial gene cytochrome c oxidase subunit I (COI), used as the standard for DNA barcode of animals, was amplified using the universal primers HCO2198 (5-TAA ACTTCA GGG TGA CCA AAA AAT CA-3), and LCO1490 (5-GGT CAA CAA ATC ATA AAGTATTGG-3). Finally, the amplified fragments were bi-directionally sequenced. All steps of the genetic analyses (DNA extraction, amplification, purification and sequencing) were conducted according tothe protocols of the Canadian Centre for DNA Barcoding (Ivanova et al., 2006, CCDB protocols http:// ccdb.ca/resources.php). Specimens collected during this study and their respective tissue samples were all kept in the collection of the Invertebrates Laboratory of the Universidad Nacional de Mar del Plata, or deposited at the ‘Museo Argentino de Ciencias Naturales Bernardino Rivadavia’, Argentina, preserved in ethanol 96%. The resulting COI sequences and any other specimen information, such as the voucher id’s, individual pictures, collection dates and location, etc., was made publicly available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org). Analyses of the genetic data performed to solve taxonomic problems that arose during this study were carried out using the online analytical tools provided by the BOLD system (http://www.boldsystems.org/) or alternatively using the standard nucleotide BLAST® (http://blast.ncbi.nlm.nih.gov/Blast.cgi) for search on GenBank databases (www.ncbi.nlm.nih.gov/GenBank). A brief note was added to the remarks for those species for which barcode is available, regardless of whether the specimens from whom the
7 FRENTE MARÍTIMO Publicación de la Comisión Técnica Mixta del Frente Marítimo sequence originates were within the geographical range of our study. When barcode sequences were generated for the present work, the corresponding identification code is provided to facilitate its finding on BOLD system.
The distribution of each species has been summarized by means of the following geographic areas: AO (Arctic Ocean), NWAO (northwestern Atlantic Ocean), NEAO (northeastern Atlantic Ocean, including the Mediterranean and Black Sea), NEPO (northeastern Pacific Ocean), NWPO (northwestern Pacific Ocean), SWAO (southwestern Atlantic Ocean), SEAO (southeastern Atlantic Ocean), SEPO (southeastern Pacific Ocean), SWPO (southwestern Pacific Ocean), SO (Southern Ocean), EIO (eastern Indic Ocean) and WIO (western Indic Ocean).
The detailed distribution of each species in SWAO has been described using the administrative divisions of Argentina (Provinces), Uruguay (Departments) and Brazil (States) coasts (Table 1). The words Province/s, Department/s and State/s have been omitted. Species that are endemic of the SWAO were indicated.
The bathymetric distribution of benthic species has been summarized using the classic terminology: supralittoral zone, intertidal (= eulittoral) zone, infralittoral zone (ca. 5 m below the low water mark), circalittoral zone (from ca. 5 m depth to the edge of the continental shelf, ca. 200 meters), bathyal zone (from ca. 200 to 4000 m) and abyssal zone (> 4000 m). That of pelagic species, as follows: epipelagic (from the surface to 200 m depth), mesopelagic (from 200 to 1000 m), bathypelagic (1000 to 4000 m and abyssopelagic (more than 4000 m).
The published morphological studies on larval development of each species have been mentioned, indicating if they were based on laboratory reared or plankton collected larvae and if they were completely of partially described. A brief qualitative summary of the information on the knowledge of species biology, as well as its economic or scientific importance (if any) has been added when available. Detailed lists of references supporting that information have been included in Supplementary Online Files 2-7.
It is important to note that, with exception made of species present in our own collection, we do not revisit the records or deposited specimens cited in the bibliography. Therefore when a location or identification was dubious to us, this was commented in the correspondent remarks. RESULTS
The present catalogue includes 211 species of decapod crustaceans, belonging to 63 families that have been reported in the literature for coastal and oceanic waters of the Southwestern Atlantic Ocean from 33º45’S (boundary between Brazil and Uruguay) to 55ºS (Beagle Channel, the southernmost boundary between Argentina and Chile), the Malvinas islands, the Southern Ocean islands (the Georgias del Sur and Sandwich del Sur), and Antarctic waters. The list consists of 36 Dendrobranchiata and 175 Pleocyemata (42 Caridea, 4 Astacidea, 5 Axiidea, 1 Gebiidea, 1 Achelata, 2 Polychelida, 40 Anomura and 80 Brachyura). The number of species per family ranges from 1 to 16; the most speciose families were Sergestidae (16), Lithodidae (14), Pinnotheridae (11) and Penaeidae (9) (Table 2).
8 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Table 2. Number of species and families of decapod crustacea reported for the study area and a comparison of the number of marine and estuarine species with A) Boschi et al. (1992) and B) Zolessi and Philippi (1995).
Families Species A B Dendrobranchiata Aristeidae 3 0 1 Benthesicymidae 5 0 0 Penaeoidea Penaeidae 9 2 8 Solenoceridae 2 1 1 Luciferidae 1 0 1 Sergestoidea Sergestidae 16 3 4 Total 6 36 6 15 Pleocyemata Dendrobranchiata Pasiphaeoidea Pasiphaeidae 5 1 1 Acanthephyridae 2 1 0 Oplophoroidea Oplophoridae 1 0 0 Nematocarcinoidea Nematocarcinidae 3 0 0 Campylonotoidea Campylonotidae 4 3 1 Palaemonoidea Palaemonidae 2 0 1 Caridea Alpheidae 4 3 1 Alpheoidea Hippolytidae 6 5 1 Thoridae 4 0 0 Processoidea Processidae 3 1 1 Pandaloidea Pandalidae 4 2 2 Crangonoidea Crangonidae 4 2 3 Astacidea Nephropoidea Nephropidae 4 2 2 Axiidea Callianassidae 5 3 1 Gebiidea Upogebiidae 1 0 0 Achelata Scyllaridae 1 1 0 Polychelida Polychelidae 2 0 0 Galatheidae 6 3 1 Galatheoidea Porcellanidae 5 2 4 Blepharipodidae 1 1 1 Hippoidea Hippidae 1 2 1 Anomura Lithodoidea Lithodidae 14 7 2 Diogenidae 5 4 3 Paguroidea Paguridae 7 5 6 Parapaguridae 1 1 0 Homolidae 1 0 1 Brachyura Dromiacea Homoloidea Latreillidae 1 0 1
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Aethroidea Aethridae 1 0 1 Bellioidea Bellidae 2 2 1 Bythograeoidea Calappidae 2 0 1 Atelecyclidae 1 1 1 Cancroidea Cancridae 1 0 0 Dorippoidea Ethusidae 1 0 0 Eriphioidea Platyxanthidae 2 2 2 Goneplacoidea Chasmocarcinidae 1 0 1 Mathildellidae 1 0 0 Leucosioidea Leucosiidae 3 2 2 Epialtidae 8 5 8 Inachidae 3 1 1 Brachyura Eubrachyura Majoidea Inachoididae 4 2 3 Heterotremata Mithracidae 1 0 1 Oregonidae 1 0 0 Hymenosomatoidea Hymenosomatidae 1 1 0 Parthenopoidea Parthenopidae 5 0 4 Pilumnoidea Galenidae 1 1 1 Carcinidae 2 1 1 Geryonidae 1 1 1 Portunoidea Macropipidae 1 1 1 Portunidae 5 1 5 Pseudozioidea Pilumnoididae 1 1 1 Panopeidae 6 2 6 Xanthoidea Pseudorhombilidae 1 0 0 Xanthidae 1 0 1 Grapsidae 3 0 2 Grapsoidea Sesarmidae 1 1 2 Brachyura Eubrachyura Varunidae 4 4 4 Thoracotremata Ocypodoidea Ocypodidae 2 1 2 Pinnotheroidea Pinnotheridae 11 8 5 Total Pleocyemata 57 175 87 92 Total Decapoda 63 211 93 107
From this 211 species, 91 are from the SWAO and adjacent areas: SO, SEPO and subantarctic WIO. Forty nine have been found only in the SWAO: 4 Dendrobranchiata (Artemesia longinaris, Farfantepe- naeus paulensis, Pleoticus muelleri and Peisos petrunkevitchi), 2 Caridea (Alpheus puapeba and Betaeus lilianae), 1 Astacidea (Metanephrops rubellus), 4 Axiidea (Anacalliax argentinensis, Notiax santarita, Poti gaucho and Sergio mirim), 1 Gebiidea (Upogebia australis), 1 Achelata (Scyllarides deceptor), 10 Ano- mura (Pachycheles chubutensis, Pachycheles laevidactylus, Blepharipoda doelloi, Lithodes confundens,
10 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Loxopagurus loxochelis, Paguristes foresti, Paguristes robustus, Paguristes robustus scarabinoi, Pagurus exilis and Pagurus leptonyx) and 26 Brachyura (Latreilla willamsi, Corystoides abbreviatus, Neopilumno- plax lipkeholthuisi, Danielethus crenulatus, Danielethus patagonicus, Ebalia rotundata, Libinia spinosa, Pelia rotunda, Anisonotus atlanticus, Collodes rostratus, Leurocyclus tuberculosis –although its presence in SEPO should be corroborated-, Amphithrax besnardi, Spinolambrus meridionalis, Chaceon notialis, Coenophthalmus tridentatus, Acantholobulus schmitti, Panopeus meridionalis, Speocarcinus dentatus, Neohelice granulata, Cyrtograpsus altimanus, Cyrtograpsus affinis, Leptuca uruguayensis, Austinixa pa- tagoniensis, Pinnixa rapax, Calyptraeotheres garthi and Fabia emiliai). Two species have been found only in the SO: 1 Caridea (Eualus kinzeri) and 1 Anomura (Neolithodes yaldwyni).
In addition, 11 species have been found both in the SWAO and the SO: 7 Caridea (Pasiphaea rathbu- nae, Pasiphaea scotiae, Campylonotus arntzianus, Chorismus antarcticus, Eualus amandae, Aegaeon boschii and Notocrangon antarcticus), 2 Astacidea (Thymops takedai and Thymopsis nilenta), 1 Ano- mura (Paralomis formosa) and 1 Brachyura (Rochinia gracilipes; one of the 2 crab species that live in Antarctic waters, together with Halicarcinus planatus, present in SWAO but also extensely distributed in shallow waters associated with the Antarctic Circumpolar Current). Three species have been found in both the SWAO and the subantarctic WIO (Caridea: Campylonotus capensis and Chorismus tuber- culatus; Anomura: Paralomis anamerae), one in SO and the subantarctic WIO (Anomura: Paralomis birsteini) and one in the three areas (Anomura: Munida spinosa).
Twenty three species have been found both in the SWAO and the SEPO: 6 Caridea (Pasiphaea doflei- ni, Campylonotus vagans, Campylonotus semistriatus, Nauticaris magellanica, Eualus dozei and Austro- pandalus grayi), 1 Axiidea (Notiax brachyophthalma), 1 Polychelida (Stereomastis suhmi), 6 Anomura (Lithodes santolla, Lithodes turkayi, Paralomis granulosa -, a single record in New Zealand was repor- ted as P. cf granulosa-, Pagurus comptus, Pagurus forceps and Propagurus gaudichaudii) and 9 Bra- chyura (Acanthocyclus albatrossis, Peltarion spinosulum, Metacarcinus edwardsii, Libidoclea granaria, Eurypodius longirostris, Pilumnoides hassleri, Cyrtograpsus angulatus, Pinnixa brevipollex and Pinnixa valdiviensis). Finally, 4 species have been found in the SWAO, the SEPO and the SO: 2 Caridea (Nema- tocarcinus lanceopes and Lebbeus antarcticus) one Astacidea (Thymops birsteini) and one Anomura (Paralomis tuberipes).
The catalogue of Boschi et al. (1992) consisted of 93 species that inhabits Argentine coastal and oce- anic waters from 36ºS to 55ºS, also including the Malvinas and Southern Ocean islands. Dendro- branchiata were represented by only 6 species and Pleocyemata by 85 (18 Caridea, 2 Astacidea, 3 Axiidea, 1 Achelata, 25 Anomura and 38 Brachyura). The most speciose families were Pinnotheridae (8), Lithodidae (7), Epialtidae (5) and Paguridae (5) (Table 2).
The list of Zolessi and Philippi (1995) consisted of 107 species that inhabits coastal and oceanic waters of Uruguay from 33º45’S to 36ºS. Dendrobranchiata were represented by 15 species and Pleo- cyemata by 92 (11 Caridea, 2 Astacidea, 1 Axiidea, 18 Anomura and 60 Brachyura). The most speciose families were Epialtidae (8), Penaeidae (8), Paguridae (6), Panopeidae (6), Pinnotheridae (5) and Por- tunidae (5) (Table 2).
The families Aristeidae, Luciferidae, Palaemonidae, Homolidae, Latreillidae, Aethridae, Calappidae, Chasmocarcinidae, Majidae, Parthenopidae, Xanthidae and Grapsidae, had been listed by Zolessi and Philippi (1995), but not by Boschi et al. (1992). Moreover, Benthesicymidae, Oplophoridae, Thori-
11 FRENTE MARÍTIMO Publicación de la Comisión Técnica Mixta del Frente Marítimo dae, Nematocarcinidae, Upogebiidae, Polychelidae, Mithracidae, Oregonidae, Cancridae, Ethusidae, Mathildellidae and Pseudorhombilidae were reported neither by Boschi et al. (1992) nor by Zolessi and Philippi (1995) (Table 2).
At the time of finishing this manuscript, from the 211 species included herein, 100 had publicly avai- lable genetic barcodes. Of those species, 61 barcodes were previously available and 39 have been sequenced and published in BOLD system for first time as result of this study. A brief note pointing that the species have available barcodes was added when applicable.
SYSTEMATIC ACCOUNT
ORDER DECAPODA Latreille, 1802 SUBORDER DENDROBRANCHIATA Spence Bate, 1888 SUPERFAMILY PENAEOIDEA Rafinesque, 1815
Family Aristeidae Wood-Mason in Wood-Mason and Alcock, 1891
Aristaeopsis Wood-Mason in Wood-Mason and Alcock, 1891
● Aristaeopsis edwardsiana (Johnson, 1868) (scarlet shrimp, carabinero)
Geographic distribution: NWAO, NEAO, NWPO, SWAO, SEAO, SWPO, WIO, EIO. SWAO: Brazil (Amapá, Pará, Bahia, Espiritu Santo and Santa Catarina); off Uruguay (Itusarry, 1983, quoted by Zolessi and Philippi, 1995; D’Incao, 1998; Tavares and Serejo, 2007). Bathymetric distribution: meso and bathypelagic (Pérez Farfante and Kensley 1997).
Remarks: reported by Itusarry (1983) and listed by Zolessi and Philippi (1995) as Plesiopenaeus edwardsianus Johnson, 1868. D’Incao (1998) reported this species from two sites off Uruguay: 35º34’S; 52º32’W and 35°44’S; 52°05’W. A fishery has been developed in northern and southeastern Brazil since 2002 (Pezzuto et al., 2006). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Plesiopenaeus Spence Bate, 1881
Plesiopenaeus armatus (Spence Bate, 1881)
Geographic distribution: NWAO, NEAO, NWPO, SWAO, SWPO, WIO. SWAO: Brazil (from Bahia to Espírito Santo) (Pérez Farfante and Kensley, 1997; Tavares and Serejo, 2007); off Buenos Aires, Argentina (Spence Bate, 1888). Bathymetric distribution: meso and bathypelagic (Pérez Farfante and Kensley 1997). Larval development undescribed (dos Santos and Lindley, 2001). However, phylogenetic analysis found Cerataspis monstrosa, originally described by Gray, 1828 as a “monstrous and misshapen animal” and placed within a new genus and species of Leptostraca, as identical to P. armatus; so it is considered a larval stage of the latter (Bracken-Grissom et al., 2012).
12 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Pseudaristeus Crosnier, 1978
Pseudaristeus speciosus (Spence Bate, 1881)
Known geographic distribution: SWAO: off Buenos Aires, Argentina at 4847 m. Bathymetric distribution: abyssopelagic (Spence Bate, 1888; Pérez Farfante, 1987).
Remarks: only collected once, at 4847 m depth (lat. long: Spence Bate, 1888).
Familiy Benthesicymidae Wood-Mason in Wood-Mason and Alcock, 1891
Benthesicymus Spence Bate, 1881
Benthesicymus brasiliensis Spence Bate, 1881
Geographical distribution: NWAO, NEAO, SWAO, SEAO, SWPO. SWAO: Argentina, off Buenos Aires (Spence Bate, 1881; Zariquiey Álvarez, 1968; d’Udekem d’Acoz, 1999; Pérez Farfante and Kensley, 1997). Bathymetric distribution: meso, bathy and abyssopelagic (Tiefenbacher, 2001).
Gennadas Spence Bate, 1881
Gennadas brevirostris Bouvier, 1905
Geographical distribution: NWAO, NEAO, SEAO. SWAO: Brazil, from Alagoas to Rio Grande do Sul), Uruguay, 35°16’S; 49°26’W (D’Incao, 1998). Bathymetric distribution: meso and bathypelagic (D’Incao, 1998).
Gennadas gilchristi Calman, 1925
Geographical distribution: SWAO, SEAO, SWPO. SWAO: international waters off Uruguay and off Buenos Aires Province, Argentina. Bathymetric distribution: meso and bathypelagic (Judkins, 2014; USNMdb).
Remarks: 60 specimens collected from 2 stations located in brazilian and international waters (Fig. 2), at 34º43´S; 49º28´W and 34º06´S; 42º43´W, respectively (Atlantis R/V), were identified by D. Judkins; 2 specimens from a station located at 40º18´S; 39º12´W (Walther Herwig R/V) were identified by B. Kensley (USNM 161579; USNM 164254).
Gennadas tinayrei Bouvier, 1906
Geographical distribution: NWAO, NEAO, NWPO, NEPO, SWAO, SEAO. SWAO: off Sao Paul, Brazil, off Uruguay. Bathymetric distribution: epi, meso and bathypelagic (Judkins, 2014; USNMdb).
Remarks: 20 specimens collected from 2 stations located in international and brazilian waters (Fig. 2), respectively at 35ºS; 48º37´W and 34º06´S; 42º49´W (Atlantis R/V) were identified by D. Judkins (USNM 164231; USNM 164246).
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Gennadas valens (Smith, 1884)
Geographical distribution: NWAO, NEAO, SWAO, SEAO. SWAO: off Rio Grande do Sul, Brazil, Uruguay and Buenos Aires, Argentina. Bathymetric distribution: epi, meso and bathypelagic (Judkins, 2014; USNMdb)
Remarks: 18 specimens collected from 2 stations located in international and Uruguayan waters, (Fig. 2), at 34º43´S; 49º28´W and 36º45´S; 53º06´W, respectively, (Atlantis R/V) were identified by D. Judkins (USNM 161576; USNM 161594).
Family Penaeidae Rafinesque, 1815
Artemesia Spence Bate, 1888
►● Artemesia longinaris Spence Bate, 1888
(Argentine stiletto shrimp, camarón, camarón siete barbas, camarão argentino, camarão serrinha, camarão ferrinho, camarão barba branca, camarão barba-ruça)
Figure 3. Artemesia longinaris Spence Bate, 1888. Live color. Photography of the body in lateral view. The top-right miniature shows the antennae length relative to the body. Photo by Nahuel Farias.
14 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Geographical distribution: endemic of SWAO: from Atafona, Rio de Janeiro (21°37’S) to Chubut, Argentina (43°30’S). Bathymetric distribution: infra and circalittoral (Boschi et al., 1992; D’Incao, 1999a; Carvalho-Batista et al., 2011). Larval development completely described from laboratory reared larvae (Boschi and Scelzo, 1977)
Remarks: Artemesia is a monotypic genus; the species is target of artisanal fisheries along its entire distribution and industrial fisheries in southern Brazil and Argentina; in both countries its biology was extensively studied (recently reviewed by Gavio and Boschi, 2016; Scelzo, 2016; these articles included a detailed bibliography: papers, books and technical reports). Artemesia longinaris plays an important role in the marine trophic chain of the southwestern Atlantic, as food for different species of fish and cephalopods. Phenotypic variations among populations exist through the geographical range that seems to be due to different environmental conditions among localities since there is a continuous gene flow among them (Supplementary Online File 2; only selected references). The Barcode is available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org under code: PRIMB104-14, sampling site: 38°13’S; 57°24’W).
Funchalia Johnson, 1868
Funchalia villosa (Bouvier, 1905)
Geographical distribution: NWAO, NEAO, SWAO, SEAO. SWAO: Brazil, from Fernando de Noronha to Rio Grande do Sul, off Uruguay and Buenos Aires, Argentina. Bathymetric distribution: epi and mesopelagic (D’Incao, 1999b; Judkins, 2014; USNMdb).
Remarks: 2 specimens collected from 2 stations located in Uruguayan an international waters (Fig. 2) at 34º09´S; 42º52´W and 36º45´S; 53º06´W, respectively, (Atlantis R/V) were identified by D. Judkins (USNM 161597; USNM 164240). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
● Funchalia woodwardi Johnson, 1868
Geographical distribution: NWAO, NEAO, SWAO, SEAO, SWPO, EIO. SWAO: Brazil, Rio Grande do Sul and Uruguay. Bathymetric distribution: epi and mesopelagic (Itusarry, 1984, quoted by Zolessi and Philippi, 1995; D’Incao, 1999b; USNMdb).
Parapenaeus Smith, 1885
● Parapenaeus americanus Rathbun, 1901
Geographical distribution: NWAO, SWAO. SWAO: Brazil, from Rio de Janeiro to Rio Grande do Sul, off Uruguay; off Buenos Aires, Argentina. Bathymetric distribution: epi and mesopelagic, demersal (D’Incao, 1999a).
Remarks: a specimen collected in international waters (Fig. 2) at 36°42´S; 49°36´W by the R/V Calypso was identified by Isabel Pérez Farfante (USNM 140929). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
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Penaeopsis
Penaeopsis serrata Spence Bate, 1881 (pinkspeckled shrimp)
Geographical distribution: NWAO, NEAO, SWAO. SWAO: from Paraná to Rio Grande do Sul, Brazil; Uruguay. Bathymetric distribution: mesopelagic (D’Incao, 1999; USNMdb).
Remarks: The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Penaeus Fabricius, 1798
Remarks: the genus Penaeus sensu lato included 29 species and some of them are the most commercially important decapods worldwide. This genus has been split into six genera by Pérez Farfante and Kensley (1997): Litopenaeus Pérez Farfante, 1969, Farfantepenaeus Burukovsky, 1997, Fenneropenaeus Pérez Farfante, 1969, Marsupenaeus Timizi, 1971, Melicertus Rafinesque, 1814 and Penaeus sensu stricto Fabricius, 1798. However, some authors have refused this change and proposed the restoration of the old Penaeus genus (Ma et al., 2011 and references thereof; WoRMS database, 2014). Here both old and new names are reported
†● Penaeus brasiliensis Latreille, 1817 = Farfantepenaeus brasiliensis (Latreille, 1817) (pinkspot shrimp, camarão rosa)
Geographical distribution: NWAO, SWAO. SWAO: from Amapá to Rio Grande do Sul, Brazil. Bathymetric distribution: infra and circalittoral (D’Incao, 1999; USNMdb). Larval development described from plankton specimens collected in Venezuela (Lares, 1974).
Remarks: the only record from Uruguay was that of Barattini and Ureta (1960, quoted by Zolessi and Philippi, 1995) but attributed to P. paulensis by Demicheli and Scarabino (2006).
● Penaeus notialis Pérez Farfante, 1967 = FarfantePenaeus notialis (Pérez Farfante, 1967) (southern pink shrimp, camarão rosa)
Geographical distribution: NWAO, SWAO, SEAO. SWAO: from Bahia to Rio de Janeiro, Brazil; Maldonado and Rocha, Uruguay. Bathymetric distribution: infra and circalittoral (Pérez Farfante, 1967; Zolessi and Philippi, 1995; WoRMSdb; USNMdb).
Remarks: reported as Penaeus duoradum Burkenroad, 1939 by Boschi (1963) and Zolessi and Philippi (1995).
16 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
►● Penaeus paulensis Pérez Farfante, 1967 = Farfantepenaeus paulensis (Pérez Farfante, 1967) (São Paulo shrimp, langostino rojo, camarão rojo, camarão rosa)
Geographical distribution: endemic of SWAO: from Ilheus, Bahía, Brazil, along the continental shelves of Brazil and Uruguay, to Mar del Plata, Buenos Aires, Argentina, although its presence in the latter is certainly occasional. Bathymetric distribution: infra and circa littoral (Pérez Farfante, 1969; Boschi et al., 1992; Santana et al., 2015). Larval development completely described from laboratory reared larvae (Iwal, 1978).
Remarks: cited by Boschi et al. (1992) as Penaeus (Farfantepenaeus) paulensis Pérez Farfante, 1967. Listed by Zolessi and Philippi (1995) as Penaeus aztecus Ives 1891 (as was done Boschi, 1963) and P. paulensis. It is commercially exploited in Braziland; artisanal fishery in Uruguay. Juvenile development takes place in estuaries and coastal lagoons. Artificially reared and subject of physiological and ecological research on larvae and juveniles (Supplementary Online File 2).
● Penaeus schmitti Burkenroad, 1936 = Litopenaeus schmitti (Burkenroad, 1939) (Caribbean, Cuban or southern white shrimp, camarão branco, camarón blanco)
Geographical distribution: NWAO, SWAO. SWAO: from Amapá to Rio Grande do Sul, Brazil; Rocha, Uruguay. Bathymetric distribution: infra and circalittoral (Zolessi and Philippi, 1995; D’Incao, 1999; USNMdb). Larval development described from larvae collected in plankton samples from Venezuela (García-Pinto and Ewald, 1974).
Xiphopenaeus Smith, 1869
● Xiphopenaeus kroyeri (Heller, 1862) (seabob, camarão sete barba, camarón siete barbas)
Geographical distribution: NWAO, SWAO. SWAO: from Amapá to Rio Grande do Sul, Brazil, Rocha, Uruguay. Bathymetric distribution: infra and circalittoral (Zolessi and Philippi, 1995; USNMdb). Larval development was partially described from plankton samples collected in southern Brazilian waters (Calazans, 1993).
Remarks: the Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
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Family Solenoceridae Wood-Mason in Wood-Mason and Alcock, 1891
Pleoticus Spence Bate, 1888
►● Pleoticus muelleri Spence Bate 1888 (Argentine red shrimp, langostino, langostin, lagostinho da Argentina, camarão de Santana, camarão vermelho)
Figure 4. Pleoticus muelleri Spence Bate 1888. Live color. Photography of the body in lateral view. Photo by Nahuel Farias.
Geographical distribution: endemic of SWAO: from Rio de Janeiro, Brazil (23°S) to Santa Cruz, Argentina (50°S). Bathymetric distribution: epi and mesopelagic, demersal, more frequently found between 9 and 130 m (D’Incao, 1999a) and between 3 and 90 m (Boschi et al., 1992). Larval development completely described from laboratory reared larvae (Iorio et al., 1990; Calazans and Ingle, 1998).
Remarks: biology, fisheries and technology have been extensively studied, both in Argentina and in Brazil (recently reviewed by Moriondo Danovaro et al., 2016; Scelzo, 2016; these articles included a detailed bibliography: papers, books and technical reports). Its fishery is one of the most important in Argentina, mainly between 43°S and 47°S. The largest catches come from San Jorge Gulf and adjacent areas; in the coastal waters off Buenos Aires province (38°-40°S) there is also a small scale fishery. Only in 2005 the catch of this species in Argentine was of 7510 metric tons. In Brazil the fishery is mainly developed in San Paulo coast and in Uruguay the species is fished by an artisanal fleet (Supplementary Online File 2). The Barcode is now available online via the Barcode of Life Data System (BOLD; www. barcodinglife.org under code: PRIMB180-14, sampling site: 38°19.6’S, 57°30.4’W).
18 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Solenocera Lucas, 1849
Solenocera necopina Burkenroad, 1939 (deepwater humpback shrimp)
Geographic distribution: NWAO, SWAO. SWAO: off Uruguay. Bathymetric distribution: epipelagic (Williams, 1984).
Remarks: 4 specimens collected from 1 station located in Brazilian waters (Fig. 2) (Professor Besnard R/V: 34º26’S; 51º47’W) were identified by I. Pérez Farfante (USNM 254072, reported from “off Punta del Palmar, Rocha, Uruguay”).
SUPERFAMILY SERGESTOIDEA Dana, 1852a
Family Luciferidae De Haan, 1849 [in De Haan, 1833–1850] (planktonic shrimps)
Belzebub Vereshchaka, Olesen and Lunina, 2016
● Belzebub faxoni (Borradaile, 1915)
Geographical distribution: NWAO, SWAO. SWAO: from Canada to Rio Grande do Sul State, Brazil, and Montevideo, Uruguay. Bathymetric distribution: epipelagic; Luciferidae are one of the few holoplanktonic decapod crustaceans. B. faxoni occurs from 6 to 55 meters deep, usually in coastal waters (D’Incao, 1997; Teodoro et al., 2012). Larval development described from plankton samples (Brooks, 1882).
Remarks: listed by Zolessi and Philippi (1995) as Lucifer faxoni Borradaile, 1915. Ecological and reproductive aspects described in Brazilian waters (Supplementary Online File 2).
Family Sergestidae Dana, 1852a
General remarks: the systematic and nomenclature of sergestid shrimps has been reviewed and modified by Vereshchaka (2000); Vereshchaka and Lunina (2015) and Vereshchaka et al. (2014; 2016).
Allosergestes Judkins and Kensley, 2008
Allosergestes pectinatus (Sund, 1920)
Geographical distribution: NWAO, NEAO, NEPO, NWPO, SWAO, SEAO. SWAO: off Uruguay and Buenos Aires, Argentina. Bathymetric distribution: epi and mesopelagic (Judkins, 2014; USNMdb).
Remarks: 127 specimens collected from 7 stations located between 34º09´-36º45´S and between 42º43´-53º06´W, mainly in international waters (Atlantis R/V) were identified by D. Judkins (USNM 161601; 164227; 164234; 161586; 164244; 164249; 164263).
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Allosergestes sargassi (Ortmann, 1893a)
Geographical distribution: NWAO, NEAO, NEPO, NWPO, SWAO, SEAO. SWAO: off Uruguay and Buenos Aires province, Argentina. Bathymetric distribution: epi and mesopelagic (Judkins, 2014; USNMdb).
Remarks: 57 specimens collected from 7 stations located between 34º06´-38º37´S and between 42º43´-53º06´W (Walther Herwig R/V and Atlantis R/V), mainly in Brazilian, Uruguayan, Argentine and international waters (Fig. 2), were identified by D. Judkins and B. Kensley (USNM 258480; 161603; 164237; 161582; 164242; 164251; 164258).
Deosergestes Judkins and Kensley, 2008
Deosergestes corniculum (Krøyer, 1855)
Geographical distribution: NWAO, NEAO, SWAO, SEAO. SWAO: off Uruguay and Buenos Aires, Argentina. Bathymetric distribution: meso and bathy pelagic (Judkins, 2014; USNMdb).
Remarks: 3 specimens collected from 2 stations located in Brazilian and Uruguayan waters (Fig. 2) at 34º43´S; 49º28´W and 36º45´S; 53º06´W, respectively (Atlantis R/V) were identified by D. Judkins (USNM 161598; 161592). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Deosergestes curvatus (Crosnier and Forest, 1973)
Geographical distribution: SWAO, SEAO, Santa Helena Island. SWAO: off Buenos Aires province, Argentina. Bathymetric distribution: meso and bathypelagic (USNMdb).
Remarks: one specimen collected from 1 station located 39º46´S; 43º30´W (Walther Herwig R/V) was identified by B. Kensley (USNM 258438).
Deosergestes disjunctus (Burkenroad, 1940)
Geographical distribution: SWAO, SEAO, SWPO, WIO. SWAO: off Uruguay and Buenos Aires province, Argentina. Bathymetric distribution: meso and bathypelagic (USNMdb).
Remarks: 10 specimens collected from 2 stations located in Uruguayan and Brazilian waters (Fig. 2) at 34º43´S; 49º28´W and at 36º45´S; 53º06´W, respectively (Atlantis R/V) were identified by D. Judkins (USNM 161600; 161585).
Eusergestes Judkins and Kensley, 2008
►● Eusergestes antarcticus (Vereshchaka, 2009)
Geographical distribution: SWAO, SEAO, SEPO, SWPO, SO, Drake Passage; Western Antarctic region. SWAO: off Uruguay and Buenos Aires, Argentina; northwest of the Georgias del Sur Islands (USNM 161578, 164256, 258389, 258423, 258474) identified as Sergestes arcticus should be considered
20 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean) representative of E. antarcticus in the study area. Bathymetric distribution: epi and mesopelagic, 100- 330 m at night (most individuals at 100 m) and 500-1000 m during the day (most individuals at 670 m) (Vereshchaka, 2009). Larval development undescribed, that of E. similis, a closely related species, described from laboratory reared specimens (Knight and Omori, 1982).
Remarks: listed by Boschi et al. (1992) and Zolessi and Philippi (1995) as Sergestes arcticus Kroyer, 1865; the nomenclature and systematic placement were modified by Vereshchakaet al. (2014). Before 2009, but also today, the Southern Atlantic species has been denoted as Sergestes arcticus (Judkins, 2014). The specimens collected in waters off Uruguay and Buenos Aires, Argentina (USNM 161578, 164256, 258389, 258423, 258474) identified as Sergestes arcticus should be considered representative of E. antarcticus in the study area. All three members of Eusergestes (E. antarcticus, E. arcticus and E. similis) live at similar depths and are similar in diurnal vertical migrations and biology. They are parapatric, with E. arcticus occurring only in the North Atlantic, E. similis in the North Pacific and E. antarcticus in the Southern Ocean (Vereshchaka, 2009) although this author still had named them as “the S. arcticus species group”. E. similis Hansen (1903) was cited for Uruguayan waters by Barattini and Ureta (1960), quoted by Zolessi and Philippi (1995), and should be considered E. antarcticus.
Gardinerosergia Vereshchaka A. L., J. Olesen and A. A. Lunina, 2014
Gardinerosergia splendens (Sund, 1920)
Geographical distribution: NWAO, NEAO, SWAO, SEAO. SWAO: off Uruguay and Buenos Aires, Argentina. Bathymetric distribution: epi, meso and bathypelagic (Judkins, 2014; USNMdb).
Remarks: 5 specimens collected from 3 stations located in Brazilian and international waters (Fig. 2) at 34º43´S; 49º28´W, 35ºS; 48º37´W, 38º37´S; 51º55´W (Atlantis R/V and Walther Herwig) were identified by D. Judkins and B. Kensley (USNM 161584; 164232; 258482).
Neosergestes Judkins and Kensley, 2008
Neosergestes edwardsii (Krøyer, 1855)
Geographical distribution: NWAO, NEAO, SWAO, SEAO. SWAO: off Uruguay and Buenos Aires province, Argentina. Bathymetric distribution: epi, meso and bathypelagic (Judkins, 2014; USNMdb).
Remarks: 2 specimens collected from 1 station located in international waters (Fig. 2) at 35º01´S; 48º46´W (Atlantis R/V) were identified by D. Judkins (USNM 164229).
Parasergestes Judkins and Kensley, 2008
Parasergestes armatus (Krøyer, 1855)
Geographical distribution: NWAO, NEAO, NWPO, SWAO, SEAO, WIO. SWAO: off Uruguay and Buenos Aires province, Argentina. Bathymetric distribution: epi and mesopelagic (USNMdb).
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Remarks: 34 specimens collected from 4 stations located in Brazilian, Uruguayan, Argentine and international waters (Fig. 2) at 35º01´-39º46´S; 43º37´-52º09´W (Walther Herwig R/V and Atlantis R/V) were identified by B. Kensley and D. Judkins (USNM 161595; 164230; 161577; 164255).
Parasergestes vigilax (Stimpson, 1860a)
Geographical distribution: NWAO, NEAO, SWAO, SEAO. SWAO: off Uruguay and Buenos Aires province, Argentina. Bathymetric distribution: epi and mesopelagic (USNMdb). Larval development partially described from plankton samples (Gurney and Lebour, 1940).
Remarks: 49 specimens collected from 5 stations located in international waters (Fig. 2) at 34º06´- 35º01´S; 42º43´-48º46´W (Atlantis R/V) were identified by D. Judkins (USNM 164228; 164233; 164245; 164247; 164264).
Peisos Burkenroad, 1945
►● Peisos petrunkevitchi Burkenroad, 1945
Figure 5. Peisos petrunkevitchi Burkenroad, 1945. Photography of the body in lateral view. he miniature showing a living specimen is from http://cifonauta.cebimar.usp.br/photo/5311/ with credit to Alvaro E. Migotto licensed under a Creative Commons Attribution-NonCommercial-ShareAlike 3.0 Unported License. Photo by Nahuel Farias.
Geographical distribution: endemic of SWAO: from Rio de Janeiro, Brazil, to Uruguay and then to Playa Unión, Chubut, Argentina (44°S). Bathymetric distribution: epipelagic (Boschi et al., 1992; D’Incao and Martins, 2000). Larval development completely described from laboratory reared larvae (Mallo, 1986).
22 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Remarks: listed by Boschi et al. (1992) and Zolessi and Philippi (1995) as Peisos petrunkevitchi Burkenroad, 1945, the nomenclature had changed to Acetes petrunkevitchi (Burkenroad, 1945) according to Vereshchaka et al. (2016). However, this taxonomic decision was rejected by D’Udekem D’Acoz and De Grave (2018). This species has an important role in trophic webs, as food of fishes (Supplementary Online File 2). The Barcode is available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org under code: INVMP091-13, sampling site: 38°1.8’S; 57°31.4’W).
Petalidium Spence Bate 1881
Petalidium foliaceum Spence Bate 1881
Geographical distribution: SEAO, SO, WIO, EIO: south of the Polar Front, in Western Antarctica, Prydz Bay, and the Scotia and Weddell Seas. SWAO: off Uruguay; the Georgias del Sur Islands. Bathymetric distribution: meso and bathypelagic (Clarke and Holmes, 1986; Vereshchaka and Lunina, 2015; USNMdb). Larval development partially described form plankton collected samples (Vera and Bacardit, 1987).
Remarks: Vereshchaka and Lunina (2015) stated that “The absence of Petalidium in the South-western Atlantic region may be a result of the poor sampling efforts in this area (no stations during the ‘Dana’ and other great pelagic expeditions)”. However, 8 specimens collected from a station located at 34º43´S; 49º28´W (Atlantis R/V) in Brazilian waters (Fig. 2) were identified by D. Judkins (USNM 161575) and the presence of larvae has been reported (Vera and Bacardit, 1987).
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Phorcosergia Vereshchaka A. L., J. Olesen and A. A. Lunina, 2014
►● Phorcosergia potens (Burkenroad, 1940)
Figure 6. Phorcosergia potens (Burkenroad, 1940). Live color. Photography of the body in lateral view. Top-right: detail of the characteristic short and laminar rostrum, with a small terminal spine at the anterosuperior margin. . Photo by Nahuel Farias.
Geographical distribution: SWAO, SEAO, SEPO, SWPO, WIO, EIO. SWAO: waters off Uruguay and Buenos Aires province, Argentina. Bathymetric distribution: meso and bathypelagic, most specimens live at 200-300 m at night and 1000-2000 m during the day (Vereshchaka, 2000; USNMdb).
Remarks: listed by Boschi et al. (1992) and Zolessi and Philippi (1995) as Sergia potens; the nomenclature and systematic placement were modified by Vereshchaka et al. (2000). However, it was still named as Sergia potens by D’Udekem D’Acoz and De Grave (2018). P. potens is parapatric to P. grandis, P. bisulcata, P. phorca, and P. filicta, which live north of 10-15°S in the Atlantic, Indian and Pacific Oceans, while P. potens occurs south of 10-15°S in the same oceans (Vereshchaka, 2000). Fifteen specimens collected from 2 station located in Brazilian and international waters (Fig. 2) at 34º43´S; 49º28´W and 34º06´S; 42º43´W, respectively (Atlantis R/V) were identified by D. Judkins (USNM 161583; 164262).
24 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Prehensilosergia Vereshchaka A. L., J. Olesen and A. A. Lunina, 2014
Prehensilosergia prehensilis (Spence Bate, 1881)
Geographical distribution: SWAO, SEAO, NWPO, SWPO. SWAO: off Uruguay and Buenos Aires province, Argentina. Bathymetric distribution: epi and mesopelagic (USNMdb).
Remarks: 35 specimens collected from stations located in Brazilian, Uruguayan, Argentine and international waters (Fig. 2) at 34º06´-39º46´S; 42º43´-53º06´W (Atlantis R/V and Walther Herwig R/V) were identified by D. Judkins and B. Kensley (USNM 161574; 161602; 164235; 164248; 164260; 258437; 258441; 258475).
Sergestes H. Milne Edwards, 1830
● Sergestes atlanticus H. Milne Edwards, 1830
Geographical distribution: NWAO, NEAO, NWPO, SWAO, SEAO, SWPO. SWAO: off Uruguay and Buenos Aires province, Argentina. Bathymetric distribution: epi and mesopelagic (USNMdb). Larval development partially described from plankton samples (Gurney and Lebour, 1940).
Remarks: 2 specimens collected from 2 stations located in international and Brazilian waters (Fig. 2) at 35ºS; 48º37´W and 34º43´S; 49º28´W, respectively, (Atlantis R/V) were identified by D. Judkins (USNM 164238; 161588).
Sergia Stimpson, 1860a
Sergia laminata (Burkenroad, 1940)
Geographical distribution: SWAO, SEAO, EIO. SWAO: off Uruguay. Bathymetric distribution: mesopelagic (USNMdb).
Remarks: one specimen collected from a station located in international waters (Fig. 2) at 34º06´S- 42º43´W (Atlantis R/V) was identified by D. Judkins as Sergestes guineensis (USNM 164261).
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SUBORDER PLEOCYEMATA Burkenroad, 1963 INFRAORDER CARIDEA Dana, 1852b SUPERFAMILY PASIPHAEOIDEA Dana, 1852b
Family Pasiphaeidae Dana, 1852
Pasiphaea Savigny, 1816
►● Pasiphaea acutifrons Spence Bate, 1888
Geographical distribution: NWPO, SWAO, SEPO (from 41°S to Magellan Strait), SO. SWAO: off Uruguay and off Argentina, from 40°36’S to 54°4’S; the Malvinas Islands; Canal Beagle. Bathymetric distribution: meso and bathypelagic (Holthuis, 1952a; Takeda and Hatanaka, 1984; Boschi et al., 1992; Zolessi and Philippi, 1995; Arntz et al., 1999).
Pasiphaea barnardi Yaldwyn, 1971
Geographical distribution: SWAO, SEAO, SEPO, SWPO, SO. SWAO: off the Malvinas Islands. Bathymetric distribution: epi, meso and bathypelagic (Guzman, 2014; USNMdb).
Remarks: 2 specimens collected from one station located at 51º59’S; 56º38’W (Eltanin R/V) were identified by G. Guzmán (USNM 1277426).
Pasiphaea dofleini Schmitt, 1932
Geographic distribution: SWAO, SEPO, from 20º to 53º22’S: a single record at the eastern entrance of the Estrecho de Magallanes. Bathymetric distribution: epi-mesopelagic (Retamal, 1973; Guzmán and Wicksten, 1998).
Pasiphaea rathbunae (Stebbing, 1914)
Geographical distribution: SWAO, SO. SWAO: the Malvinas and Georgias del Sur Islands. Bathymetric distribution: meso and bathypelagic (Clarke and Holmes, 1986; Burukovsky and Romensky, 1987, cited by Guzmán, 2014; USNMdb).
Remarks: 3 specimens collected from 2 stations located at South Georgia Islands (56º57´S; 33º46´W and -58º10´S; 37º47´W, Eltanin R/V) were identified by G. Guzmán (USNM 1277390; 1277388). The specimens from the Malvinas Islands have been cited as P. balssi Burukovsky and Romensky, 1987, recently synonymyzed as P. rathbunae (WoRMSdb).
Pasiphaea scotiae (Stebbing, 1914)
Geographical distribution: SWAO, SO. SWAO: Georgias del Sur Islands. Bathymetric distribution: meso and bathypelagic (Clarke and Holmes, 1986; Gorny, 1999; USNMdb).
26 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Remarks: 30 specimens collected from 7 stations located at 53º06´-58º51´S; 33º46´-38º03´W (Eltanin R/V) were identified by G. Guzmán (USNM 1277388; 1277390; 1277389; 1277391; 1277392; 1277394; 1277395). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
SUPERFAMILY OPLOPHOROIDEA Dana, 1852
Family Acanthephyridae Spence Bate, 1888
Acanthephyra A. Milne-Edwards, 1881
► Acanthephyra pelagica (Risso, 1816)
Figure 7. Acanthephyra pelagica (Risso, 1816). Live color. Photography of the body in lateral view. Photo by Nahuel Farias.
Geographical distribution: NEAO, SEAO, SEPO, SWPO, SO, EIO, WIO. SWAO: Drake Passage. SWAO: from 26 to 54°S; Georgias del Sur Islands, Scotia Sea. Bathymetric distribution: meso and bathypelagic; it migrates between both areas (Takeda and Hatanaka, 1984; Clarke and Holmes, 1986; Wasmer, 1986, 1993; Boschi et al., 1992; Burukovsky and Andreeva, 2010; Judkins, 2014; USNMdb).
Remarks: important item of the Patagonian toothfishDissostichus eleginoides diet after its ontogenetic descent to the lower part of the continental slope (500–1000 m depths) of the Malvinas Islands (Supplementary Online File 3). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: TALII001-14, sampling site: 38°6.2’S, 54°20.9’W).
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Acanthephyra quadrispinosa Kemp, 1939
Geographical distribution: SWAO, SEAO, SEPO, SWPO, EIO, WIO. SWAO: off Uruguay. Bathymetric distribution: meso and bathypelagic (Judkins, 2014; USNMdb).
Remarks: 3 specimens collected from 2 stations located in Brazilian and international waters (Fig. 2) at 34º43´S; 49º28´W and 34º06´S-42º43´W, respectively (Atlantis R/V) were identified by D. Judkins (USNM 161590; USNM 164265).
Family Oplophoridae Dana, 1852
Systellaspis Spence Bate, 1888
Systellaspis debilis (A. Milne-Edwards, 1881)
Geographical distribution: NWAO, NEAO, NEPO (Hawai), SWAO, SEAO, SWPO, EIO, WIO. SWAO: off Uruguay. Bathymetric distribution: meso and bathypelagic (Judkins, 2014; USNMdb).
Remarks: 12 specimens collected from 3 stations located in international waters (Fig. 2) at 34º09´S; 42º52´W, 34º07´S; 42º49´W and 34º06´S; 42º43´W (Atlantis R/V) were identified by D. Judkins (USNM 164241; 164252; 164257). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
28 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
SUPERFAMILY NEMATOCARCINOIDEA Smith, 1884
Family Nematocarcinidae Smith, 1884
Nematocarcinus A. Milne-Edwards, 1881
Nematocarcinus lanceopes Spence Bate, 1888
Figure 8. Nematocarcinus lanceopes Spence Bate, 1888. Live color. Photography of the body in lateral view. Photo by Martin Rauschert .
Geographical distribution: SEPO up to Chiloé (42°S), SO: Weddell Sea. SWAO: Scotia Seas (south of 55°30’S); occasionally found on the continental slope south off Tierra del Fuego, at the southernmost tip of South America. Bathymetric distribution: bathyal, 840-4000 m (Thatje et al., 2005 and references thereof; Raupach et al., 2010; Arntz and Gorny, 1991; Gutt et al., 1991; Guzmán and Quiroga, 2005; Basher et al., 2014; Basher and Costello, 2016; Dambach et al., 2016). Larval development described from plankton samples (Thatje et al., 2005).
Remarks: model species for studies on physiological and reproductive adaptations to Antarctic waters (Supplementary Online File 3).
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Nematocarcinus longirostris Spence Bate, 1888
Figure 9. Nematocarcinus longirostris Spence Bate, 1888. Live color. Photography of the body in lateral view. Photo by Martin Rauschert.
Geographical distribution: NWPO, SWAO, SEPO, WIO, the Marion and Crozet Islands, SO: Indian Ocean sector of the Southern Ocean and parts of the Ross Sea. SWAO: off the Georgias del Sur and Sandwich del Sur Islands, in the Scotia Sea. Bathymetric distribution: bathyal (Thatje et al., 2005 and references thereof). Larval development described from plankton samples (Thatje et al., 2005).
Remarks: considered a synonym for N. proximatus Spence Bate, 1888 (Zarenkov, 1968; Tiefenbacher, 1990; Thatje et al., 2005), it has been listed as N. proximatus by D’Udekem D’Akoz and De Grave (2018).
30 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Nematocarcinus tenuipes Spence Bate, 1888
Figure 10. Nematocarcinus tenuipes Spence Bate, 1888. Live color. Close-up photography of the body in lateral view. The top-right miniature shows the length of pereiopods and antennae relative to the body. Photo by Nahuel Farias.
Geographical distribution: NWPO and Hawai, SWAO. SWAO: off Brazil, from 4º to 32ºS and off Buenos Aires, Argentina. Bathymetric distribution: bathyal (Cardoso and Burukovsky, 2014).
Remarks: the specimens found in Mar del Plata Canyon, off Buenos Aires, Argentina, are morphologically indistinguishable from those of NWPO. However, the genetic distance that separates them from Taiwan specimens and the geographical isolation of both clusters could indicate the presence of different species raises doubts on species identity (Farías, unpublished). The Barcode is available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: TALII 035-14, sampling site: 37°54.5’S 54°0.7’W).
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SUPERFAMILY CAMPYLONOTOIDEA Sollaud, 1913
Family Campylonotidae Sollaud, 1913
Campylonotus Spence Bate, 1888
Campylonotus arntzianus Thatje, 2003
Figure 11. Campylonotus arntzianus Thatje, 2003. Live color. Photography of the body in lateral view. Photo by Sven Tathje
Geographical distribution: SO: Antarctica. SWAO: The Sandwich del Sur Islands, Scotia Sea. Bathymetric distribution: bathyal (Thatje, 2003).
32 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
► Campylonotus capensis Spence Bate, 1888
Figure 12. Campylonotus capensis Spence Bate, 1888. Live color. Photography of the body in lateral view. Photo by Nahuel Farias.
Geographical distribution: WIO: Marion Island. SWAO: along the continental slope of Argentine north to the continental slope off Pernanbuco, Brazil; the Malvinas Islands (Torti and Boschi, 1973; Thatje et al., 2001). Bathymetric distribution: bathyal (Thatje, 2003 and cites thereof). Larval development partially described from plankton samples (Thatje et al., 2001).
Remarks: evidences of protandric hermaphroditism (Torti and Boschi, 1973). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: TALII040-14, sampling site: 38°4.1’S; 54°28.4’W).
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►● Campylonotus semistriatus Spence Bate, 1888
Figure 13. Campylonotus semistriatus Spence Bate, 1888. Photography of the body in lateral view. Photo by Nahuel Farias.
Geographical distribution: SEPO: from Canal Beagle and fjord system of the Estrecho de Magallanes to 32°S. SWAO: off Tierra del Fuego and Santa Cruz, Argentina. Bathymetric distribution: circalittoral and bathyal, (Takeda and Hatanaka, 1984; Thatje, 2003 and cites thereof; Arana and Ahumada, 2006). Larval development partially described from plankton samples (Thatje et al., 2001).
Remarks: type species of Campylonotus.
34 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
► Campylonotus vagans Spence Bate, 1888
Figure 14. Campylonotus vagans Spence Bate, 1888. Live color. Photography of the body in lateral view. Photo by Héctor Monsalve.
Geographical distribution: SEPO: Chilean coast from about 41 to 56°S. SO. SWAO: from Buenos Aires (about 35°S) to Tierra del Fuego, Argentina; the Georgias del Sur Islands. Bathymetric distribution: circalittoral (Boschi, 1963; Torti and Boschi, 1973; Takeda and Hatanaka, 1984; Boschi et al., 1992; Gorny, 1999; Thatje et al., 2001; Thatje, 2003 and cites thereof). Larval development abbreviated (i.e. fewer stages than related species) and early juvenile development described from laboratory reared larvae (Thatje et al., 2001, Thatje and Lovrich 2003).
Remarks: evidences of protandric hermaphroditism (Torti and Boschi, 1973; Thatje and Lovrich, 2003). Some life history traits studied (Supplementary Online File 3). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
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SUPERFAMILY PALAEMONOIDEA Rafinesque, 1815
Family Palaemonidae Rafinesque, 1815
Palaemon Weber, 1795
Palaemon macrodactylus Rathbun, 1902 (oriental shrimp)
Figure 15. Palaemon macrodactylus Rathbun, 1902. Live color. Photography of the body in lateral view. Photo by Nahuel Farias.
Geographical distribution: cosmopolitan and invader. NEPO, NWPO, SWAO, NEAO, SWPO. SWAO: from Río de la Plata to Río Negro, Argentina. Bathymetric distribution: intertidal and infralittoral (Ashelby et al., 2013 and all references thereof; Bonel et al., 2013; Martorelli et al., 2012; Spivak et al., 2006). Larval development completely described from laboratory reared larvae (Little, 1969); the number of stages varies with season and environmental conditions (salinity) (Vázquez et al., 2015).
Remarks: native from Japan and China, it is a recent invader in Buenos Aires Province, Argentina, first detected in Mar del Plata Harbor (Spivak et al., 2006).The life history of this species was studied in the native area and in Mar del Plata; in both habitats the species showed seasonal variability in reproduction and development. The egg development and its infection by bacteria and fungus have been studied (Supplementary Online File 3). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: INVMP011-13, sampling site: 38°2.5’S; 57°32.3’W).
36 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
● Palaemon northropi (Rankin, 1898)
Geographical distribution: NWAO, SWAO. SWAO: distribution, from Ceará, Brazil, to Piriápolis, Uruguay. Bathymetric distribution: intertidal in sand and mudflat pools, on reef pools during ebb tide, and in the lower portions of estuaries, near mangrove areas (Holthuis, 1952b; Pralon and Negreiros- Fransozo, 2006). Larval development described from laboratory reared larvae (Moura et al., 1990).
Remarks: population biology studied in Brazil, where is used as model species to study physiology of larvae and adults, mainly osmoregulation (Supplementary Online File 3). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
SUPERFAMILY ALPHEOIDEA Rafinesque, 1815
Family Alpheidae Rafinesque, 1815 (snapping shrimps)
Alpheus Fabricius, 1798
►● Alpheus puapeba Christoffersen, 1979
Figure 16. Alpheus puapeba Christoffersen, 1979. Live color. Photography of the body in lateral view. Photo by Nahuel Farias.
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Geographical distribution: endemic of SWAO: Espírito Santo to Rio Grande do Sul, Brazil, Buenos Aires Province, Argentina. Bathymetric distribution: circalittoral (Boschi, 1988; Christoffersen, 1979, 1998; Zolessi and Philippi, 1995; Soledade and Almeida, 2013). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: PRIMB189-14, sampling site: 38°19.6’S; 57°30.4’W).
Alpheus pouang Christoffersen, 1979
Geographical distribution: endemic of SWAO: São Paulo to Rio Grande do Sul, Brazil, and Uruguay. Bathymetric distribution: circalittoral (Christoffersen, 1979, 1998; Soledade and Almeida, 2013; Cunha et al., 2015).
Betaeus Dana, 1852
► Betaeus lilianae Boschi, 1966
Figure 17. Betaeus lilianae Boschi, 1966. Live color. Photography of the body in lateral view of female (up) and male (down). Photo by Nahuel Farias.
Geographical distribution: endemic of SWAO: from Uruguay (34°18’S to San Matías Gulf, Río Negro, Argentina (41°S). Bathymetric distribution: intertidal from San Matías to Mar del Plata (38°S) and from 60 to 160 m in Uruguay (Christoffersen, 1982a).
Remarks: morphometry and ecological relationships studied (Supplementary Online File 3).
38 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
► Betaeus truncatus Dana, 1852
Geographical distribution: SEPO: from Perú (12°S) to Southern Chile (Magellan Strait and Cape Horn). SWAO: from Tierra del Fuego to 41°S; the Georgias del Sur Islands. Bathymetric distribution: intertidal and circalittoral (Lardies and Wehrtmann, 2001; Barria et al., 2014; Marin and Anosov, 2014). Larval development partially described from plankton samples (Albornoz and Wehrtmann, 1997; Thatje and Baccardit, 2001).
Remarks: effects of latitude on life history traits studied in Chile (Supplementary Online File 3). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www. barcodinglife.org).
Family Hippolytidae Spence Bate, 1888
Chorismus Spence Bate, 1888
► Chorismus antarcticus (Pfeffer, 1887)
Figure 18. Chorismus antarcticus (Pfeffer, 1887). Live color. Photography of the body in lateral view. Photo by Nahuel Farias.
Geographical distribution: SWAO, SO: Ross Sea, South Orkney Islands: up 36°30’S; Scotia Sea. Bathymetric distribution: circalittoral and bathyal (Boschi et al., 1992; Raupach et al., 2010; Basher et al., 2014). Larval development abbreviated described from plankton samples (Gurney, 1937).
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Remarks: only protandrous hermaphrodite decapod species found on the Antarctic continental shelves (Mascetti et al., 1997). Used as model species for studies on physiological and reproductive adaptations to Antarctic waters, lipofuscin as age marker, microscopic anatomy and ultrastructure, and presence of trace metals in Antarctic crustaceans (Supplementary Online File 3). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
► Chorismus tuberculatus Spence Bate, 1988
Geographical distribution: WIO: Marion and Kerguelen Island, in the sub-antarctic Indian Ocean. SWAO: off Chubut, Santa Cruz and Tierra del Fuego, Argentina; the Malvinas and Georgias del Sur Islands. Bathymetric distribution: bathyal (Boschi et al., 1992 Gorny, 1999, Thatje and Bacardit, 2000a). Larval development partially described from plankton samples (Thatje and Baccardit, 2000a).
Exhippolysmata Stebbing, 1915
Exhippolysmata oplophoroides Holthuis, 1948
Geographical distribution: NEAO, SWAO. SWAO: from Guyana to Uruguay (33°46’S; 53°17’W). Bathymetric distribution: circalittoral (Christoffersen, 1979). Larval development partially described (Negreiros Fransozo et al., 2002).
Remarks: protandric simultaneous hermaphrodite (Supplementary Online File 3). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife. org).
Latreutes Stimpson, 1860a
►● Latreutes parvulus (Stimpson, 1871a)
Geographical distribution: NWAO, NEAO, SWAO. SWAO: from Piauí to Rio Grande do Sul, Brazil; to Monte Hermoso, Buenos Aires, Argentina. Bathymetric distribution: intertidal, infra and circalittoral, 0-124 m (Boschi, 1988; Christoffersen, 1982b; 1998).
Remarks: The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Merhippolyte Spence Bate, 1888
►● Merhippolyte americana Holthuis, 1961.
Geographical distribution: NWAO, SWAO. SWAO: from Ceará to Rio Grande, Brazil, Uruguay. Bathymetric distribution: circalittoral and bathyal, 80-275 m (Christoffersen 1979; Pachelle et al., 2016).
Remarks: the southernmost report by Christoffersen (1979) was 35º10’S; 52º46’W, erroneously assigned to Buenos Aires province, Argentina; in fact, it corresponds to Uruguayan waters. Since then, the mistake remained in cheklists (Boschi et al., 1992; Zolessi and Philippi, 1995).
40 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Nauticaris Spence Bate, 1888
► Nauticaris magellanica (A. Milne-Edwards, 1891)
Figure 19. Nauticaris magellanica (A. Milne-Edwards, 1891). Live color. Photography of the body in lateral view. Photo by Arthur Anker
Geographical distribution: SEAO: from Cape Horn to 30°S. SWAO: Uruguay; Argentina from Buenos Aires Province to Canal Beagle; the Malvinas Islands. Bathymetric distribution: circalittoral (Holthuis, 1952a; Boschi et al., 1992; Christophersen, 1982a). Larval development described from larvae reared in the laboratory reared and collected in the field (Albornoz and Wehrtmann, 1997; Thatje and Bacardit, 2000c; Wehrtmann and Albornoz, 1998, 2003).
Remarks: used in Chile as model species for the study of latitudinal variation in reproductive biology (Supplementary Online File 3). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
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Family Thoridae Kingsley, 1879
Eualus Thallwitz, 1892
Eualus amandae Nye, Copley and Linse, 2013
Geographical distribution: SO: only from non-hydrothermal areas in the Kemp Caldera. SWAO: East Scotia Ridge, Scotia Sea. Bathymetric distribution: bathyal (Nye et al., 2013).
Eualus dozei (A.Milne-Edwards, 1891)
Geographical distribution: SEPO and Southern tip of South America. Bathymetric distribution: circalittoral (Wehrtmann and Carvacho, 1997). Larval development partially described from plankton samples (Albornoz and Wehrtmann, 1997).
Eualus kinzeri Tiefenbacher, 1990
Geographical distribution: SO: Eastern Weddell Sea. Bathymetric distribution: bathyal (Arntz and Gorny, 1991; Gorny et al., 1992).
Lebbeus White, 1847a
Lebbeus antarcticus (Hale, 1941)
Geographical distribution: SWAO, SEPO, SO, Weddell and Lazarev Seas. SWAO: Tierra del Fuego, Argentina; Scotia Sea. Bathymetric distribution: bathyal (Arntz and Gorny, 1991; D’Udekem D’Akoz and De Grave, 2018; Gorny, 1999; Guzmán and Quiroga, 2005; Nye et. al., 2013).
Remarks: largest egg size reported for an Antarctic caridean (Supplementary Online File 3).
42 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
SUPERFAMILY PROCESSOIDEA Ortmann, 1896
Family Processidae Ortmann, 1896
Processa Leach, 1815 [in Leach, 1815–1875]
Processa guyanae Holthuis, 1959
Geographical distribution: NWAO, SWAO. SWAO: from Paraiba to Rio Grande do Sul, Brazil, and Uruguay. Bathymetric distribution: circalittoral (Christoffersen, 1979; Williams, 1984).
►● Processa hemphilli Manning and Chace, 1971
Geographical distribution: NWAO, SWAO. SWAO: from Rio de Janeiro to Rio Grande do Sul, Brazil, Uruguay. Bathymetric distribution: intertidal, infralittoral and circalittoral (Christoffersen, 1979).
Remarks: the southernmost report by Christoffersen (1979) was 35º18’S; 54º13’W, erroneously assigned to Buenos Aires province, Argentina; in fact, it corresponds to Uruguayan waters. Since then, the mistake remained in cheklists (Williams, 1984; Boschi, 1988; Boschi et al., 1992; Zolessi and Philippi, 1995).
Processa profunda Manning and Chace, 1971
Geographical distribution: NWAO, SWAO. SWAO: from São Paulo (Brazil) to Uruguay. Bathymetric distribution: circalittoral (Christoffersen, 1982b; Williams, 1984).
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SUPERFAMILY PANDALOIDEA Haworth, 1825
Family Pandalidae Haworth, 1825
Austropandalus Holthuis, 1952
►● Austropandalus grayi (Cunningham, 1871)
Figure 20. Austropandalus grayi (Cunningham, 1871). Female; ovigerous. Photography of the body in lateral view. Photo by Nahuel Farias.
Geographical distribution: SEPO: from Chiloé to Cape Horn. SWAO: off Buenos Aires to Tierra del Fuego, Argentina; Canal Beagle. Bathymetric distribution: circalittoral (Boschi et al., 1992; Arntz et al., 1999; Pérez-Barros et al., 2004). Larval development partially described from plankton samples and laboratory hatching (Thatje and Bacardit, 2000b).
Remarks: reproductive and developmental traits described in Chile (Supplementary Online File 3). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife. org, under code: MEJII037-14, sampling site: 39°12.4’S; 58°21.1’W).
44 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Pandalopsis Spence Bate, 1888
►● Pandalopsis ampla Spence Bate, 1888 (deepwater bigeye)
Figure 21. Pandalopsis ampla Spence Bate, 1888. Live color. Photography of the body in lateral view. Bottom-right: detail of the rostrum. Photo by Nahuel Farias.
Geographical distribution: NEPO, SWAO, SEPO, SO. SWAO: off Uruguay and off Argentina from 41 to 55°S; the Malvinas Islands. Bathymetric distribution: bathyal (Rathbun, 1904; Boschi, 1973; Takeda and Hatanaka, 1984; Zolessi and Philippi, 1995; Gorny, 1999; Guzmán and Quiroga, 2005).
Remarks: type species of Pandalopsis, listed by Zolessi and Philippi as P. amplus. Takeda and Hatanaka (1984) suspected that the Atlantic and Pacific populations may be separate species.
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Pantomus A. Milne-Edwards, 1883
Pantomus parvulus A. Milne-Edwards, 1883
Geographical distribution: NWAO, SWAO. SWAO: Rio Grande do Sul, Brazil, and Uruguay (34°28’S, 51°50’W). Bathymetric distribution: circalittoral and bathyal (Williams, 1984; Christoffersen, 1989).
Remarks: type species of Pantomus.
Stylopandalus Coutière, 1905
Stylopandalus richardi (Coutière, 1905)
Geographical distribution: NWAO, NEAO, SWAO, SEAO, SEPO, SWPO, WIO, EIO. SWAO: Rio de Janeiro, Brazil, and Uruguay. Bathymetric distribution: epi, meso and bathypelagic (Guzmán and Rivera, 2002; Cardoso, 2009, Judkins, 2014).
Remarks: its presence off Uruguay is suggested by Judkins illustration (2014, fig. 4) and should be corroborated. The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
46 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
SUPERFAMILY CRANGONOIDEA Haworth, 1825
Family Crangonidae Haworth, 1825
Aegaeon Agassiz, 1846 [in Agassiz, 1842–1846]
►● Aegaeon boschii (Christoffersen, 1988)
Figure 22. Aegaeon boschii (Christoffersen, 1988). Live color. Photography of the body in typical posture in dorsal (up) and lateral (down) views. Photo by Nahuel Farias.
Geographical distribution: SWAO, SO. SWAO: distribution, from off Rio de Janeiro, Brazil, (23°S) to Chubut, Argentina (44°S). Bathymetric distribution: circalittoral (Boschi et al., 1992; Gorny, 1999; Cardoso, 2013).
Remarks: listed by Boschi et al. (1992) and Zolessi and Philippi (1995) as Pontocaris boschii Christoffersen, 1988. The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: PRIMB130-14, sampling site: 38°19.6’S; 57°30.4’W).
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Lissosabinea Christoffersen, 1988
● Lissosabinea cf tridentata (Pequegnat, 1970)
Two specimens from off Uruguay (one was an exuvia and the other was damaged) were assigned by Christoffersen (1988) to L. cf. tridentata, and this record was listed by Spivak (1997) (Komai, 2006). However, Komai (2006) suggested that “Christoffersen (1988) was actually reporting a species other than L. tridentata, although it remains unknown what species was represented”.
Notocrangon Coutière, 1900
► Notocrangon antarcticus (Pfeffer, 1887)
Geographical distribution: SO: circum-Antarctic. SWAO: Estrecho de Magallanes; the Georgias del Sur Island. Bathymetric distribution: circalittoral and bathyal (Arntz and Gorny, 1991; Gutt et al., 1991; Boschi et al., 1992; Basher et al., 2014). Larval development described by Makarov (1973).
Remarks: model species for studies on physiological and reproductive adaptations to Antarctic waters, the potential use of lipofuscin as age marker, the microscopic anatomy and ultrastructure of the digestive system, and the presence of trace metals in Antarctic crustaceans (Supplementary Online File 3). Type species of Notograngon. The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Philocheras Stebbing, 1900
● Philoceras gorei (Dardeau, 1980)
Geographical distribution: NWAO, SWAO. SWAO: from Rio de Janeiro (Brazil) to off Uruguay. Bathymetric distribution: circalittoral (Christoffersen, 1988).
Remarks: the material studied by Christoffersen (1988) included a sample from 34º06’S; 51º33’W that corresponds to Brazilian waters. Fig. 2)
48 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
INFRAORDER ASTACIDEA Latreille, 1802 SUPERFAMILY NEPHROPOIDEA Dana, 1852a
Family Nephropidae Dana, 1852
Metanephrops Jenkins, 1972
►● Metanephrops rubellus (Moreira, 1903). (scampi, cigala, Uruguayan lobster)
Figure 23. Metanephrops rubellus (Moreira, 1903). Live color. Photography of the body in dorsal view. Photo by Fabrizio Scarabino.
Geographical distribution: endemic of the SWAO: from 23°S (off Rio de Janeiro State, Brazil) to 38°S (off Buenos Aires Province, Argentina). Bathymetric distribution: 50-270 m (Holthuis, 1991; Tavarez, 1998; Melo, 1999; Severino Rodríguez et al., 2007).
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Thymops Holthuis, 1974
►● Thymops birsteini (Zarenkov and Semenov, 1972) (langosta de profundidad)
Figure 24. Thymops birsteini (Zarenkov and Semenov, 1972). Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographical distribution: SWAO, SEPO, SO. SWAO: off Uruguay and Argentina to 53°S; the Malvinas Islands; the Georgias del Sur Islands, Scotia Sea. Bathymetric distribution: circalittoral and bathyal, (Holthuis, 1974, 1991; Itusarry, 1984, cited by Zolessi and Philippi, 1995; Laptikhovsky and Reyes, 2009).
Remarks: reproductive biology studied (Supplementary Online File 4). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Thymops takedai Ahyong, Webber and Chan, 2012.
Geographical distribution: SWAO, SO. SWAO: the Georgias del Sur Islands, Scotia Sea. Bathymetric distribution: bathyal (Ahyong et al., 2012).
50 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Thymopsis Holthuis, 1974
Thymopsis nilenta Holthuis, 1974
Geographical distribution: SWAO, SO. SWAO: south east of the Malvinas Islands; south of the San Pedro Island. Bathymetric distribution: bathyal (Holthuis, 1974; Ahyong et al., 2012).
INFRAORDER AXIIDEA de Saint Laurent, 1979
Family Callianassidae Dana, 1852 (ghost shrimp, camarón fantasma, corrupto)
Subfamily Anacalliacinae Manning and Felder, 1991
Anacalliax de Saint Laurent, 1973
► Anacalliax argentinensis (Biffar, 1971)
Geographical distribution: endemic of the SWAO: from San Blas, Buenos Aires, to Puerto Deseado, Santa Cruz, Argentina (Ferrari, 1981). Bathymetric distribution: intertidal to circalittoral (Ferrari, 1981).
Remarks: type species of Anacalliax.
Subfamily Callianassinae Dana, 1852
Notiax Manning and Felder, 1991
► Notiax brachyophthalma (A. Milne-Edwards, 1870).
Geographical distribution: SEPO, from Chiloé (Chile) to Bahía Independencia (Perú). SWAO: from Golfo San Jorge to Puerto Deseado, Santa Cruz, Argentina. Bathymetric distribution: intertidal (Boschi et al., 1992).
Remarks: listed by Boschi et al. (1992) as Callianassa brachyophthalma.
Notiax santarita Thatje, 2000
Geographical distribution: reported only for Canal Beagle, Chile-Argentina. Bathymetric distribution: circalittoral (Thatje, 2000).
Remarks: until this report, it has been assumed that Thalassinidea are absent in the extreme of South America, as well as in Antarctic waters (Thatje, 2000).
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Poti Rodrigues and Manning, 1992
● Poti gaucho Rodrigues and Manning, 1992
Geographical distribution: endemic of SWAO, found only in Brazilian waters near the Uruguayan border. Bathymetric distribution: circalittoral (Rodrigues and Manning, 1992).
Remarks: reported from “off Chui, near the border between Brazil and Uruguay, 33°43’S; 51°13’W, depth 150 m” (Rodrigues and Manning, 1992).
Subfamily Callichirinae Manning and Felder, 1991
Sergio Manning and Lemaitre, 1994
►● Sergio mirim (Rodrigues, 1966)
Geographical distribution: endemic of the SWAO: from Bahia to Rio Grande do Sul, Brazil, and from Mar del Plata to Bahía Blanca, Buenos Aires, Argentina. Bathymetric distribution: infralittoral (Ferrari, 1981, Rodrigues and Shimizu, 1998). Larval development completely described from laboratory reared larvae (Rodrigues, 1984).
Remarks: listed by Boschi et al. (1992) as Callianasa mirim and by Zolessi and Philippi (1995) as Callichirus mirim. The morphology anatomy, burrowing behavior, population dynamics and symbiotic relationships studied in Brazil (Supplementary Online File 4).
INFRAORDER GEBIIDEA de Saint Laurent, 1979
Family Upogebiidae Borradaile, 1903 (ghost shrimp, camarón fantasma)
Upogebia Leach, 1814 [in Leach, 1813–1815]
Upogebia australis Thatje and Gerdes, 2000
Geographical distribution: reported only for Canal Beagle, Chile-Argentina. Bathymetric distribution: circalittoral (Thatje and Gerdes, 2000).
Remarks: the first species of the Upogebiidae reported for Chile and Argentina; only two upogebiids from southern coastal waters of Brazil had represented the former southernmost distribution of this family in South America (Thatje and Gerdes, 2000).
52 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
INFRAORDER ACHELATA Scholtz and Richter, 1995
Family Scyllaridae Latreille, 1825
Scyllarides Gill, 1898
► Scyllarides deceptor Holthuis, 1963 (hooded slipper lobster, lagosta sapateira)
Geographical distribution: endemic of the SWAO: off Brazil in Pernambuco and from Rio de Janeiro to Rio Grande do Sul; off Uruguay and Buenos Aires Province, Argentina. Bathymetric distribution: circalittoral (Boschi et al., 1992; Oliveira et al., 2008; Tavares et al., 2009; Scarabino et al., 2017, and references thereof).
Remarks: in southeastern Brazil S. deceptor and S. brasiliensis are caught by fleets fishing for pink shrimp (Farfantepenaeus brasiliensis and F. paulensis) and octopuses (Octopus vulgaris); population and reproductive biology studied (Supplementary Online File 4).
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INFRAORDER POLYCHELIDA Scholtz and Richter, 1995
Family Polychelidae Wood-Mason, 1875
Pentacheles Spence Bate, 1878
Pentacheles validus A. Milne Edwards, 1880a (deep-sea blind lobster)
Figure 25. Pentacheles validus A. Milne Edwards, 1880. Preserved specimen. Photography of the body in dorsal (up) and lateral (down) views. Photo by Nahuel Farias.
Geographic distribution: NWAO, NEAO, SWAO, SEPO, SWPO, SO. Mid-Atlantic Ridge, EIO, WIO. SWAO: from off Bahia to Rio de Janeiro, Brazil, off Uruguay; the Georgias del Sur Islands. Bathymetric distribution: bathyal (Serejo et al., 2007; Chang et al., 2013; Farías and Scarabino, in press).
Remarks: the Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
54 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Stereomastis Spence Bate, 1878
Stereomastis suhmi (Spence Bate, 1878)
Figure 26. Stereomastis suhmi (Spence Bate, 1878). Live color. Photography of the body in dorsal (up) and lateral (down) views. Photo by Nahuel Farias.
Geographical distribution: SEPO: off Chile and archipelago of Tierra del Fuego. SWAO: Mar del Plata Canyon, off Buenos Aires, Argentina, 38°72´S; 54°12´W). Bathymetric distribution: bathyal (Spence Bate, 1888; Arntz et al., 1999; Farías et al., 2015).
Remarks: the Barcode is now available online via the Barcode of Life Data System (BOLD; www. barcodinglife.org, under code: INVMP081-13, sampling site: 37°52.0’S; 53°54.0’W).
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INFRAORDER ANOMURA MacLeay, 1838 SUPERFAMILY GALATHEOIDEA Samouelle, 1819
Family Munididae Ahyong, Baba, Macpherson and Poore, 2010
Munida Leach, 1820 (squat lobster, langostilla, bogavante, piojo de mar, langostino de los canales)
► Munida gregaria (Fabricius, 1793)
Figure 27. Munida gregaria (Fabricius, 1793). Live color. Photography of the body in dorsal view of the benthic gregaria (left) and pelagic subrugosa (right) morphotypes. Photo by Gustavo Lovrich
Geographical distribution: SEPO: from 42ºS to 55ºS, including the Straits of Magellan. SWPO: off New Zealand and eastern Australia between 36ºS and 55ºS. SO: a single record at 59º15´S; 54º25´W (GBIFdb). SWAO: from 37º17´S to 55ºS, and over all continental shelf and shelf break, including the Malvinas Islands and Burdwood Bank. Bathymetric distribution: epi and mesopelagic. Larval development: described from plankton samples and experimental cultures (Roberts, 1973; Pérez- Barros et al., 2007); megalopae of both morphotypes clearly distinguishable from plankton samples (Varisco, 2013).
Remarks: two morphotypes clearly distinguishable (Tapella and Lovrich, 2006), which correspond to the descriptions of the former M. subrugosa and M. gregaria listed by Boschi et al. (1992) and Zolessi and Philippi (1995). Both species were synonymized by Baba et al. (2008) after genetic evidences
56 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean) of identity (Pérez-Barros et al., 2008). The “gregaria” morphotype has a pelagic phase that can be found in the water column further the sexual maturity (Diez et al., 2016a; 2016b); the “subrugosa” morphotype is exclusively benthic from first juvenile stages. Ecology studied (reviewed by Diez et al., 2016c; Supplementary Online File 4). Reported by Henderson (1888) as M. spinosa for “Off Montevideo” (37°17’S; 53°52’W). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
● Munida iris A. Milne Edwards, 1880a
Known geographical distribution: NEAO, NWAO. SWAO: from Alagoas to Rio Grande do Sul, Brazil, and Uruguay. Known bathymetric distribution: circalittoral and bathyal (Melo-Filho and Melo, 2001; Melo, 2006).
Remarks: the Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
† Munida flinti Benedict, 1902
Known geographical distribution: NWAO. SWAO: from Gulf of Mexico, Mexico to Rio Grande do Sul, Brazil, and Uruguay. Known bathymetric distribution: circalittoral and bathyal (Melo-Filho and Melo, 2001; Melo, 2006).
† Munida forceps A. Milne-Edwards, 1880
Known geographical distribution: NEAO, NWAO. SWAO: from Virginia, USA to Rio Grande do Sul, Brazil, and Uruguay. Known bathymetric distribution: circalittoral and bathyal (Melo-Filho and Melo, 2001; Melo, 2006).
† Munida irrasa A. Milne-Edwards, 1880
Known geographical distribution: NEAO, NWAO. SWAO: from North Carolina, USA to Rio Grande do Sul, Brazil, and Uruguay. Known bathymetric distribution: circalittoral and bathyal (Melo-Filho and Melo, 2001; Melo, 2006).
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► Munida spinosa Henderson, 1885
Figure 28. Munida spinosa Henderson, 1885. Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographical distribution: SWPO: off Crozet Islands (OBISdb, 2017), SO: Drake Passage (GBIFdb). SWAO: continental shelf and shelf break, off Rio de la Plata (36ºS) to Banco Burdwood. Bathymetric distribution: epi to bathypelagic (Baba et al., 2008; Lovrich et al., 2005; Petraroia, 2014).
Remarks: reported by Henderson (1888) as M. spinosa for “Off Montevideo” (37°17’S; 53°52’W). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: TALII030-14, sampling site: 38°5.5’S; 54°39.5’W).
Family Porcellanidae Haworth, 1825
Liopetrolisthes Haig, 1960
Liopetrolisthes patagonicus (Cunningham, 1871)
Geographic distribution: SEPO: from San Lorenzo (Perú, 12ºS) to the Estrecho de Magallanes and near Cabo de Hornos (55º42´S). Bathymetric distribution: intertidal to circalittoral (Ceseña et al., 2016; Retamal, 2000).
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Remarks: likely present at the eastern entrance of the Estrecho de Magallanes. Morphologically similar to Liopetrolisthes mitra (Dana, 1852) but it taxonomic status has been clarified by Urbina (1991) and Weber and Galelguillos (1991). L. patagonicus is symbiotic to sea-urchins and sea-stars (Ceseña et al., 2016).
Pachycheles Stimpson, 1858
►● Pachycheles chubutensis Boschi, 1963
Figure 29. Pachycheles chubutensis Boschi, 1963. Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographical distribution: endemic of the SWAO: Santa Catarina and Rio Grande do Sul, Brazil, Cabo Polonio, Rocha, Uruguay, Buenos Aires, Río Negro and Chubut, Argentina. Bathymetric distribution: intertidal and circalittoral (Haig, 1966, Boschi et al., 1992; Zolessi and Philippi, 1995; Veloso, 1998). Larval development partially described (González-Pisani et al., 2006).
Remarks: morphologically similar, and related, to P. laevidactylus (Miranda et al., 2014). Embryonic development described (Supplementary Online File 4). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: INVMP031-13, sampling site: 38°2.2’S; 57°31.4’W).
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►● Pachycheles laevidactylus Ortmann, 1892
Figure 30. Pachycheles laevidactylus Ortmann, 1892. Live color. Photography of the body in dorsal (left) and ventral (right) views. Photo by Nahuel Farias.
Geographical distribution: endemic of the SWAO: from Pernanbuco (Brazil) to Monte Hermoso, Buenos Aires (Argentina). Bathymetric distribution: intertidal and circalittoral, 0-30 m (Harvey and De Santo, 1996). Larval development completely described from laboratory reared larvae (Boschi et al., 1967).
Remarks: listed by Boschi et al. (1992), Zolessi and Philippi (1995) and Veloso (1998) as P. haigae Rodrigues da Costa and reassigned to P. laevidactylus by Harvey and De Santo (1996). Morphologically similar, and related, to P. chubutensis (Miranda et al., 2014). Life history studied (Supplementary Online File 4). The Barcode is now available online via the Barcode of Life Data System (BOLD; www. barcodinglife.org, under code: ESTA016-15, sampling site: 38°10.0’S; 57°38.0’W).
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Polyonyx Stimpson, 1858
● Polyonyx gibbesi Haig, 1956
Geographic distribution: NWAO, SWAO. SWAO: from Sergipe (Brazil) to Buenos Aires, Argentina. Bathymetric distribution: infralittoral (Veloso, 1998; Rodríguez et al., 2005; Hirose et al., 2012; Belleggia et al., 2010).
Remarks: the previous reported southern limit of its geographical distribution was La Paloma, Uruguay (Haig, 1966). It is the only species of the genus that occurs in the western Atlantic, and is a known commensal of the polychaete worm Chaetopterus variopedatus (Rodríguez et al., 2005). The symbiotic relationships of this crab and the worm have been studied (Supplementary Online File 4). In Argentinian waters, specimens were found in stomachs of the narrownose smoothhound shark Mustelus schmitti and the absence of chaetopterids or rest of its tubes in those stomachs has suggests that crabs could live free or in another host (Belleggia et al., 2010). However, one specimen of P. gibbesi was found in a Chaetopterus tube in the mouth of Rio de la Plata in 2009 (Ocampo, unpublished report). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Porcellana Lamarck, 1801
● Porcellana sayana (Leach, 1820)
Geographic distribution: NWAO, SWAO. SWAO: from Amapa to Rio Grande do Sul, Brazil; Uruguay. Bathymetric distribution: intertidal (Veloso and Melo, 1993; Veloso, 1998; Ferreira, 2010). Larval development completely described from laboratory reared larvae (Hernández et al., 1998).
Remarks: symbiotic relationship with the hermit crab Dardanus insignis (Anomura, Diogenidae) studied (Supplementary Online File 4).
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SUPERFAMILY HIPPOIDEA Latreille, 1825
Family Blepharipodidae Boyko, 2002
Blepharipoda Randall, 1840
►● Blepharipoda doelloi Schmitt, 1942
Figure 31. Blepharipoda doelloi Schmitt, 1942. Live color. Photography of the body in dorsal (left) and lateral (up right) and ventral (down right) views. Photo by Nahuel Farias.
Geographic distribution: endemic of the SWAO. from Rio de Janeiro, Brazil, to San Matías Gulf, Rio Negro, Argentina. Bathymetric distribution: intertidal to circalittoral (Boschi et al., 1992; Boyko, 2002). Larval development completely described from larvae reared in the laboratory (Boschi et al., 1968).
Remarks: the presence B. occidentalis mentioned by Zolessi and Philippi (1995) is erroneous (Boyko, 2002).
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Family Hippidae Latreille, 1825
Emerita Scopoli, 1777 (mole crab, cangrejo topo)
►† Emerita analoga (Stimpson, 1857)
Geographic distribution: NEPO and SEPO: from Kodiak Islands, Alaska, USA, to Puerto Bueno, Magallanes, Chile. Bathymetric distribution: intertidal (Efford, 1976). Larval development described from plankton samples (Johnson and Lewis, 1942; Mujica et al., 2015).
Remarks: Efford (1976) erroneously attributed the specimen deposited in the Museum of Comparative Zoology (Harvard University) with the number MCZ 13235 and collected in False Bay, California, USA to a locality of Tierra del Fuego Province, Argentina, known as “Caleta Falsa”. This mistake was reproduced by Boschi et al. (1992).
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►● Emerita brasiliensis Schmitt, 1935 (tatucito)
Figure 32. Emerita brasiliensis Schmitt, 1935. Preserved specimen. Photography of the body in dorsal (left), lateral (up right) and ventral (down right) views. Photo by Nahuel Farias.
Geographic distribution: NWAO, SWAO. SWAO: from Sao Francisco to Santa Catarina States, Brazil; Uruguay. Bathymetric distribution: intertidal and infralittoral (Efford, 1976; Boschiet al., 1992). Larval development described from plankton samples by Veloso and Calazans (1992); larvae were reared in the laboratory by Otegui and Soares-Gomes (2007).
Remarks: Boschi et al., 1992 stated that this species has been also found in “Argentina (norte y centro de la Provincia de Buenos Aires) (Efford, 1976)”. However, Efford (1976) had written: “In the southern part of its range E. brasiliensis extends to Uruguay but has not been recorded from the beaches of neighboring Argentina (in litt. Boschi)”. Later, Orensanz et al. (2002) included E. brasiliensis reports in Buenos Aires province, Argentina, as an example of ‘extralimital records’. Life history, ecology and physiology studied in Uruguay and southern Brazil (Supplementary Online File 4). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife. org).
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SUPERFAMILY LITHODOIDEA Samouelle, 1819
Family Lithodidae Samouelle, 1819
Lithodes Latreille, 1806
► Lithodes confundens Macpherson, 1988 (centolla, southern king crab)
Figure 33. Lithodes confundens Macpherson, 1988a. Live color. Photography of the body in dorsal view. Photo by Gustavo Lovrich
Geographic distribution: SWAO: off Buenos Aires at 36ºS; coastal waters off the Eastern entrance of the Estrecho de Magallanes (50º-54ºS); Burdwood Bank; the Malvinas Islands; extending eastwards over the Northern Scotia Ridge up to Shag Rocks (53°39´S; 40°44´W). Bathymetric distribution: intertidal during mating (spring - summer) and then circatidal in southern coastal waters; bathyal on North Scotia ridge. (Lovrich et al., 2002; Wittmann et al., 2010; Sotelano et al., 2013; Anosov et al., 2014).
Remarks: it needs to be verified whether it is a morphotype of L. santolla (Pérez-Barros et al., 2015). Punta Arenas in the Straits of Magellan, reported by Macpherson (1988a) is a dubious location because is the disembarking port for an extensive fishing area and specimens could have come from the Atlantic entrance of Estrecho de Magallanes (Lovrich et al., 2002). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
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Lithodes ferox Filhol, 1885
Geographic distribution: SEAO: off western Africa between 22ºN and 28ºS. SWAO: only one record at 36º29´S; 53º47´W (Uruguayan slope, not off Brazil as otherwise stated). Bathymetric distribution: bathyal (Macpherson, 1988a).
Remarks: the Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
►● Lithodes santolla (Molina, 1782) (centolla, centolla patagónica, southern king crab, crabe royal de Patagonie)
Figure 34. Lithodes santolla (Molina, 1782). Live color. Photography of the body in dorsal view. Photo by Gustavo Lovrich
Geographic distribution: SEPO: off 41ºS to Cabo de Hornos (Retamal, 2000). SWAO: apparently disjoint; continental slope from Uruguay (ca. 36º30´S) to Cabo de Hornos; most of the continental shelf south 45ºS (Macpherson, 1988a). Bathymetric distribution: infralittoral (Canal Beagle and Estrecho de Magallanes) and bathyal northwards 40ºS, on the continental slope (Vinuesa, 1991; Boschi et al., 1992; Sotelano et al., 2013). Larval development completely described from laboratory reared larvae (Campodónico, 1971; McLaughlin et al., 2001).
Remarks: very abundant and subject of artisanal and industrial fisheries at Golfo San Jorge, Estrcho de Magallanes and Cabal de Beagle, from which almost all biological information proceeds, and recently reviewed (Lovrich and Vinuesa; 2016,Wyngaard et al., 2016; also: Supplementary Online
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File 4). Previously known as Lithodes antarctica Hombron and Jacquinot, 1842. Apparently with two morphotypes, the other being L. confundens, though not synonymized yet (Pérez-Barros et al., 2015). Cited from Uruguayan outer shelf as Pseudolithodes zenkevitchi (see Birstein & Vinogradov, 1972 and Macpherson, 1988a). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
► Lithodes turkayi Macpherson, 1988 (centolla)
Figure 35. Lithodes turkayi Macpherson, 1988a. Preserved specimen. Photography of the body in dorsal view. Photo by Gustavo Lovrich
Geographic distribution: SEPO from 32ºS to Tierra del Fuego. SWAO: Canal Beagle; the Malvinas Islands; Northern Scotia Ridge. Bathymetric distribution: circalittoral to bathyal (Campodónico and Guzmán, 1972a; Macpherson, 1988a; 2004; Vinuesa et al., 1999; Retamal, 2000; Anosov et al., 2014).
Remarks: records of L. murrayi in Estrecho de Magallanes correspond to L. turkayi (Campodónico and Guzmán, 1972a; Nakamura et al., 1986; Vinuesa et al., 1999). Presence of L. turkayi, previously misidentified as L. murrayi, is uncertain in Antarctica (Anosov et al., 2014; García Raso et al., 2005; Klages et al., 1995). There is some information on life history (Supplementary Online File 4). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www. barcodinglife.org).
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Neolithodes A. Milne Edwards and Bouvier, 1894
Neolithodes diomedeae (Benedict, 1895) (centolla de la Isla Grande de Chiloé, centolla patache, king crab, stone crab)
Geographic distribution: NEPO and SEPO: from 36ºN (off California, USA) to 45ºS (off Chiloé, Chile), and a single record at the western entrance of Estrecho de magallanes. SO: apparently misidentified off the Antarctic Peninsula. SWAO: Northern Scotia Ridge, off the Georgias del Sur Islands. Bathymetric distribution: bathyal, associated to hydrothermal vents (Báez et al., 1986; Macpherson, 1988a; Chevaldonne and Olu, 1996; Drumm et al., 2013; Anosov et al., 2014; GBIFdb).
Remarks: some information on natural history (Supplementary Online File 4). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Neolithodes yaldwyni Ahyong and Dawson, 2006 (king crab)
Geographic distribution: SO: off western Antarctic Peninsula and in Ross Sea. Bathymetric distribution: bathyal (Ahyong and Dawson, 2006; Smith et al., 2011).
Remarks: some information on ecology (Supplementary Online File 4). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Paralomis White, 1856
► Paralomis anamerae Macpherson, 1988
Geographic distribution: WIO: off Crozet and Kergelen Islands. SWAO: continental shelf off Argentina, the Malvinas and Georgias del Sur Islands. Bathymetric distribution: circalittoral and bathyal (Macpherson, 1988a; 2004; Purves et al., 2003).
Remarks: the Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Paralomis birsteini Macpherson, 1988 (king crab)
Geographic distribution: WIO: off Crozet and Kergelen Islands. SO: off western Antarctic Peninsula at Bellinghausen Sea; Ross Sea. Bathymetric distribution: bathyal (Macpherson, 1988b; Ahyong and Dawson, 2006; Thatje et al., 2008; Aronson et al., 2015; GBIFdb).
Remarks: the specimens reported by Arana and Retamal (1999) at Scotia Sea need confirmation (Anosov et al., 2014). There is some information on ecology and natural history (Supplementary Online File 4). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
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►● Paralomis formosa Henderson, 1888 (stone crab, centollón)
Figure 36. Paralomis formosa Henderson, 1888. Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: SO, South Orkney Islands. SWAO: off Spirito Santo, Brasil, off Uruguay and Argentine slope, from 18ºS; 39ºW to 47ºS; 59º30´W; along the Northern Scotia Ridge, off the Georgias del Sur Islands and Shag rocks. Bathymetric distribution bathyal. (Tavares and Albuquerque 1990, López-Abellán and Balguerias, 1994; Purves et al., 2003; Anosov et al., 2014).
Remarks: listed by Zolessi and Philippi as P. formosus. An intermittent fishery for this species and P. spinosissima off Georgias del Sur Island; some information on life history (Supplementary Online File 4). Reported by Henderson (1888) as M. spinosa for “Off Montevideo” (37°17’S.; 53°52’W). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: TALII026-14, sampling site: 37°58.7’S; 54°7.1’W).
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► Paralomis granulosa (Hombron and Jacquinot, 1846) (centollón, falsa centolla, false king crab, softshell king crab, crabe royal hérisson, snow crab in Chile)
Figure 37. Paralomis granulosa (Hombron and Jacquinot, 1846). Live color. Photography of the body in dorsal view. Photo by Gustavo Lovrich.
Geographic distribution: SEPO: from Chiloe to Cabo de Hornos and Estrecho de magallanes. SWPO: a single record in Morgue Seamount, New Zealand, 43ºS, 79º54´W at 1000 m depth reported as P. cf granulosa. SWAO: Canal Beagle, the Malvinas Islands, Banco Burdwood. Bathymetric distribution: infralittoral to circalittoral (Boschi et al., 1992; Retamal, 2000; GBIFdb). Larval development completely described from laboratory reared larvae (Campodónico and Guzmán, 1981; McLaughlin et al., 2003).
Remarks: part of the mixed fishery withLithodes santolla in Estrecho de Magallanes and Canal Beagle; information on life history and fisheries recently reviewed (Lovrich and Vinuesa; 2016, Wyngaard et al., 2016; also: Supplementary Online File 4). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Paralomis longidactylus Birstein and Vinogradov, 1972
Known geographic distribution: only one specimen was caught off Uruguay (35º34´S; 52º40´W). Known bathymetric distribution: bathyal (Birstein and Vinogradov, 1972; Macpherson, 1988b).
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► Paralomis spinosissima Birstein and Vinogradov, 1972
Figure 38. Paralomis spinosissima Birstein and Vinogradov, 1972. Preserved specimen. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: SWAO, SO. SWAO: slope off Mar del Plata; slope off E and SE of Tierra del Fuego and Cabo de Hornos, Burdwood Bank; off Georgia del sur Islands. Bathymetric distribution: bathyal (Macpherson, 1988a; López-Abellán and Balguerias, 1994; Anosov et al., 2014; Olguín et al., 2015.). Larval development completely described from laboratory reared larvae (Watts et al., 2006).
Remarks: intermittent fishery for this species and P. formosa off Georgias del Sur Islands; some information on life history (Supplementary Online File 4). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
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Paralomis tuberipes Macpherson, 1988
Geographic distribution: SEPO: only one record at the type locality off Puerto Aguirre (45º10´S; 75º33´W). SO: off western Antarctic Peninsula. SWAO: reported at 48ºS; 65ºW. Bathymetric distribution: not recorded (Macpherson, 2004; Thatje and Arntz, 2004; GBIFdb).
Paralomis zealandica Dawson and Yaldwin, 1971 (prickly King Crab)
Geographic distribution: SWPO: off eastern New Zealand (41ºS-53ºS). SWAO: Bromley Plateau (ca. 31º13´S; 34º49´W). Bathymetric distribution: bathyal (Macpherson, 1988a; GBIFdb; NIWAdb).
Remarks: the Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
SUPERFAMILY PAGUROIDEA Latreille, 1802 (hermit crab, ermitão, ermitaño)
Family Diogenidae Ortmann, 1892
Dardanus Paul’son, 1875
►● Dardanus insignis (de Saussure, 1858) (red brocade hermit crab)
Geographic distribution: NWAO, SWAO. SWAO: from Rio de Janeiro to Rio Grande do Sul, Brazil, Uruguay, from Buenos Aires to Chubut, Argentina. Bathymetric distribution: infralittoral to bathyal (Boschi et al., 1992; Rieger, 1998; Carranza and Horta, 2008). Larval development completely described by Hebling and Mansur (1995) from laboratory reared larvae.
Remarks: listed by Zolessi and Philippi (1995) as D. arrosor insignis. Population and reproductive biology, shell occupation, and symbiosis with Porcellana sayana studied in Brazil and Uruguay (Supplementary Online File 4).
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Loxopagurus Forest, 1964
►● Loxopagurus loxochelis (Moreira, 1901)
Figure 39. Loxopagurus loxochelis (Moreira, 1901). Photography of a living specimen within a shell of the sea snail Olivancillaria sp. Photo by Nahuel Farias.
Geographic distribution: endemic of the SWAO: from Bahia to Rio Grande do Sul, Brazil, Uruguay and Buenos Aires, Argentina. Bathymetric distribution: intertidal to infralittoral (Forest and de Saint Laurent, 1968; Boschi et al., 1992; Rieger, 1998; Nucci and Melo, 2015). Larval development completely described from laboratory reared larvae (Scelzo; 1976; see also Mantelatto et al., 2006) and references thereof.
Remarks: the hypothetical synonymy between the genus Loxopagurus and Isocheles has been discussed by Mantelatto et al. (2006) and Nucci and Melo (2015). Population and reproductive biology, shell occupation and epibiosis, and spermatophore morphology, studied in Brazil (Supplementary Online File 4).
Paguristes Dana, 1851a
► Paguristes foresti Scelzo, 1971
Geographic distribution: endemic of the SWAO: Buenos Aires, Argentina (Boschi et al., 1992) and Uruguay (Scelzo, 1973). Bathymetric distribution: circalittoral, 200 m (Scelzo, 1971; Boschi et al., 1992).
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►● Paguristes robustus Forest and de Saint Laurent, 1968
Geographic distribution: endemic of the SWAO: from São Paulo to Rio Grande, Brazil, through Uruguay, to Buenos Aires, Argentina. Bathymetric distribution: infra and circalittoral (Forest and de Saint Laurent, 1968; Boschi et al., 1992; Rieger, 1998; Fransozo et al., 2011).
Paguristes scarabinoi Lima and Santana, 2017
Geographic distribution: endemic of the SWAO: Uruguay. Bathymetric distribution: circalittoral (Lima and Santana, 2017)
Remarks: only reported from 35°41´S; 52°35´W and 34°43´S; 52°37´W (Lima and Santana, 2017).
Family Paguridae Latreille, 1802
Pagurus Fabricius, 1775
General remark: the highly diverse and widely distributed genus Pagurus has been sub-divided in several informal groups of species with similar morphology; some of these groups were validated by molecular analysis by Olguín and Mantelatto, 2013. However, these authors concluded that the taxonomic status of the species assigned to Pagurus, but also to Propagurus, “needs significant revision which should accord with phylogenetic relationships, and at least the proposal of new genera or new subgenera for some groups of species”. The 3 informal groups reported for the study area are the following: provenzanoi (P. provenzanoi and P. criniticornis), comptus (P. comptus and P. forceps) and exilis (P. exilis) (Olguín and Mantelatto, 2013).
►● Pagurus comptus White, 1847b
Geographic distribution: SEPO: from Coquimbo to Magellan region, Chile. SWAO: from Uruguay, to Rio Negro, Argentina; Canal Beagle; the Malvinas Islands, Scotia Sea. Bathymetric distribution: infra to circalittoral (Boschi et al., 1992; Arntz et al., 1999; Pérez Barros et al., 2004; Mantelatto et al., 2009a).
Remarks: according to Mantelatto et al. (2009a), P. forceps and P. comptus are “phylogenetically close but different species, with no genetic justification to support their synonymization”. Reproductive and populational biology, shell occupation and parasites were studied in Chile and Southern Argentina (Supplementary Online File 4). Scarabino (personal communication) stated that the report from Montevideo reproduced by Forest and de Saint Laurent (1968) is erroneous. The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
►● Pagurus criniticornis (Dana, 1852c)
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(white spotwrist hermit crab)
Geographic distribution: NWAO, SWAO. SWAO: from Pernambuco to Rio Grande do Sul, Brazil, from Buenos Aires to Rio Negro, Argentina. Bathymetric distribution: intertidal to circalittoral (Boschi et al., 1992; Rieger, 1998; Nucci and Melo, 2007). Larval development completely, and juvenile development, described from laboratory reared larvae (Hebling and Brossi-García, 1981; Brossi-García and Hebling, 2006).
Remarks: model species for the study of behavior and shell occupation in Brazil; populational and reproductive biology, as well as effects of temperature and salinity on larval development, were also studied in this country (Supplementary Online File 4). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
►● Pagurus exilis (Benedict, 1892)
Geographic distribution: endemic of SWAO: from Rio de Janeiro to Rio Grande do Sul, Brazil, through Uruguay to Buenos Aires, Argentina. Bathymetric distribution: intertidal to circalittoral (Forest and de Saint Laurent, 1968; Boschi et al., 1992; Rieger, 1998; Cervellini, 2001; Nucci and Melo, 2007). Larval development completely described from laboratory reared larvae (Scelzo and Boschi, 1969).
Remarks: population and reproductive biology, shell occupation and reproductive morphology studied in Brazil (Supplementary Online File 4).
► Pagurus forceps H. Milne Edwards, 1836
Geographic distribution: SEPO: from Antofagasta to Tierra del Fuego, Chile. SO: its presence in the Antarctic requires confirmation. SWAO: from 38°S to Tierra del Fuego, Argentina; the Malvinas Islands. Bathymetric distribution: intertidal to bathyal (Mantelatto et al. 2009a; USNM, Antarctic Invertebrates: http://invertebrates.si.edu/antiz/taxon_view.cfm?taxon=4772).
Remarks: P. forceps and P. comptus are “phylogenetically close but different species, with no genetic justification to support their synonymization” (Mantelatto et al., 2009a).
● Pagurus leptonyx Forest and de Saint Laurent, 1968
Geographic distribution: endemic of SWAO: from Rio de Janeiro, Brazil to Maldonado, Uruguay. Bathymetric distribution: circalittoral (Forest and de Saint Laurent, 1968; Zolessi and Philippi, 1995).
†● Pagurus provenzanoi Forest and de Saint Laurent, 1968
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Geographic distribution: NWAO, SWAO. SWAO: from Pernambuco to Rio Grande do Sul, Brazil, and Uruguay. Bathymetric distribution: infra and circalittoral (Rieger, 1998; Nucci and Melo, 2007).
Remarks: the identification of Uruguayan specimen is dubious (Forest and de Saint Laurent, 1968). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www. barcodinglife.org).
● Pagurus trichocerus Forest and de Saint Laurent, 1968
Geographic distribution: endemic of SWAO: only found near Cabo Polonio, Uruguay, not Rio de la Plata as stated in the official page of the Museum d´Histoire Naturelle de Paris (https://science.mnhn. fr/institution/mnhn/collection/iu/item/2008-15162). Bathymetric distribution: infralittoral (Forest and de Saint Laurent, 1968)
Remarks: described on the basis of a single male specimen from the type locality.
Propagurus McLaughlin and de Saint Laurent, 1998
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►● Propagurus gaudichaudii (H. Milne Edwards, 1836)
Figure 40. Propagurus gaudichaudii (H. Milne Edwards, 1836). Preserved specimen. Photography of the body in dorsal
view. Photo by Nahuel Farias.
Geographic distribution: SEPO: Magallanes, southern Chile. SWAO: Rio Grande do Sul, Brazil, Uruguay, from Buenos Aires to Estrecho de Magallanes, Argentina. Bathymetric distribution: infralittoral and circalittoral (Boschi et al., 1992; McLaughlin and de Saint Laurent, 1998; Rieger, 1998; Carranza and Horta, 2008; Nucci and Melo, 2011).
Remarks: listed by Boschi et al. (1992) and Zolessi and Philippi (1995) as Pagurus gaudichaudi. Type species of Propagurus (McLaughlin and de Saint Laurent, 1998). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: PRIMB168-14, sampling site: 38°27.5’S; 58°19.5’W).
Family Parapaguridae Smith, 1882
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Sympagurus Smith, 1883
► Sympagurus dimorphus (Studer, 1883)
Figure 41. Sympagurus dimorphus (Studer, 1883). Live color. Photography of the body in dorsal view. Photo by Nahuel
Farias.
Geographic distribution: SWAO, SEAO, SEPO, SWPO. SWAO: from Buenos Aires to Tierra del Fuego, Argentina. Bathymetric distribution: circalittoral and bathyal (Boschi et al., 1992; Forest et al., 2000; Lemaitre, 2004; Olguin et al., 2014). Larval development: only megalopae collected in the field described (Lemaitre and McLaughlin, 1992).
Remarks: in SWAO integrates the benthic community dominated by the Patagonian scallop Zygochlamys patagonica (King, 1832), ca. 100 m depth. It is found in gastropod shells but also associated with zoanthid colonies, especially Epizoanthus paguricola; spermatophore morphology described (Supplementary Online File 4). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: MEJII044-14, sampling site: 39°5.8’S; 55°41.2’W).
INFRAORDER BRACHYURA Linnaeus, 1758
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SECTION PODOTREMATA Guinot, 1977 SUPERFAMILY HOMOLOIDEA H. Milne Edwards, 1837
Family Homolidae H. Milne Edwards, 1837
Homola Leach, 1815
● Homola minima Guinot and Richer de Forges, 1995
Geographic distribution: NWAO from Virgina (USA) to northern Brazil, NEAO, SWAO. SWAO: from Rio Grande do Norte to Rio Grande do Sul, Brazil, Uruguay. Bathymetric distribution: circalittoral and bathyal (Tavares and Lemaitre, 2014). Larval development described from laboratory reared larvae (Rice and Provenzano, 1970).
Remarks: denoted as Thelxiope barbata by Rathbun (1937), Bordin (1987) and Zolessi and Philippi (1995) and H. barbata by Williams, 1984; Melo 1999 and references thereof and Nizinski, 2003. The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www. barcodinglife.org).
Family Latreilliidae Stimpson, 1858
Latreillia Roux, 1830 [in Roux, 1828–1830]
● Latreillia willamsi Melo, 1990 (spider crab, cangrejo araña)
Geographic distribution: endemic of SWAO: off southern Brazil coasts from Rio de Janeiro to the Uruguay border. Bathymetric distribution: circalittoral, 130-290 m (Melo, 1996; Castro et al., 2003).
Remarks: reported as L. elegans (Bordin, 1987; Zolessi and Philippi, 1995).
SECTION EUBRACHYURA de Saint Laurent, 1980
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SUBSECTION HETEROTREMATA Guinot, 1977 SUPERFAMILY AETHROIDEA Dana, 1851b
Family Aethridae Dana, 1851b
Hepatus Latreille, 1802
● Hepatus pudibundus (Herbst, 1785) (flecked box crab, caranguejo-baú, cofrecito)
Geographic distribution: NWAO and SWAO from Georgia (USA) to Rio Grande do Sul (Brazil) and Maldonado, Uruguay. Bathymetric distribution: intertidal to circalittoral (Juanicó, 1978; Williams, 1984; Bordin, 1987; Martínez et al., 2009; Nizinski, 2003). Larval development completely described from laboratory reared larvae, as well as a series of juvenile stages (Rieger and Hebling, 1993; Hebling and Rieger, 2003); the megalopa and early juvenile development from neuston samples (Negreiros- Franzoso et al., 2008).
Remarks: it has been reported as “the most abundant crab on the southeastern Brazilian coast” and there are detailed studies on its biology and ecology in this country; model species for investigate osmoregulatory physiology (Supplementary Online File 5). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
SUPERFAMILY BELLIOIDEA Dana, 1852c
Family Belliidae Dana, 1852
Acanthocyclus Lucas, 1844 [in H. Milne Edwards and Lucas, 1842–1844]
► Acanthocyclus albatrossis Rathbun, 1898a
Geographic distribution: SEPO: Southern Chile from 36 to 49ºS. SWAO: Canal Beagle, the Malvinas Islands. Bathymetric distribution: intertidal (Boschi et al., 1992; Vinuesa et al., 1999; Lovrich, 1999; Meyer et al., 2013).
Remarks: its biology has been studied in Chile; model species for the study of magnesium regulation in polar areas (Supplementary Online File 5). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Corystoides Lucas, 1844 [in H. Milne Edwards and Lucas, 1842–1844]
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►● Corystoides abbreviatus A. Milne Edwards, 1880
Figure 42. Corystoides abbreviatus A. Milne Edwards, 1880. Live color. Photography of the body in dorsal view. Photo by
Nahuel Farias.
Geographic distribution: endemic of the SWAO: from Rio de Janeiro (Brazil) to Buenos Aires Province (Argentine). Bathymetric distribution: intertidal to circalittoral (Scelzo et al., 2005). Larval development described from laboratory reared larvae (Boschi and Scelzo, 1970).
Remarks: listed by Boschi et al. (1992) and Zolessi and Philippi (1995) as C. chilensis. The hypothesis of the existence of two species (C. chilensis for the Pacific andC. abbreviatus for the Atlantic, as registered in WoRMS) instead of one as proposed by Rathbun (1930), is still unresolved (Garth, 1957; Boschi, 1964, p. 68; Guinot, 1976, WoRMSdb). Morphometry and relative growth of the Atlantic form studied (Supplementary Online File 5).
SUPERFAMILY CALAPPOIDEA De Haan, 1833 [in De Haan, 1833–1850]
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Family Calappidae De Haan, 1833 [in De Haan, 1833–1850]
Acanthocarpus Stimpson, 1871b
General remark: the specific identity of Southwestern Atlantic Acanthocarpus should be revised since Bordin (1987), Zolessi and Philippi (1995) and Melo (2010) did not take into account the study of Mañé-Garzón (1979).
● Acanthocarpus alexandri Stimpson, 1871b
Geographic distribution: NWAO, SWAO. SWAO: from Rio de Janeiro to Rio Grande do Sul, Brazil and Maldonado, Uruguay. Bathymetric distribution: circalittoral and bathyal (Bordin, 1987; Melo, 2010, and cites thereof).
Acanthocarpus meridionalis Mañé-Garzón, 1979
Geographic distribution: SWAO: restricted to Uruguay. Bathymetric distribution: circalittoral (Mañé- Garzón, 1979).
Remarks: only one adult male (type specimen) collected in Pozo del Fango, off Rocha, Uruguay (located at 34°47’S; 53°45’W, although it was erroneously reported at 38°24’S; 55°26W). The report of Mañé-Garzón (1979) was not mentioned by Scarabino (2006).
SUPERFAMILY CANCROIDEA Latreille, 1802
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Family Atelecyclidae Ortmann, 1893b
Peltarion Hombron and Jacquinot, 1846 [in Hombron and Jacquinot, 1842–1854]
►● Peltarion spinosulum (White, 1843)
Figure 43. Peltarion spinosulum (White, 1843). Male. Dorsal view. Photo by Nahuel Farias.
Geographic distribution: SWAO, SEPO. SWAO: from Uruguay to Beagle Channel through Buenos Aires, Rio Negro, Chubut, Santa Cruz and Tierra del Fuego, Argentina. Bathymetric distribution: circalittoral (Boschi et al., 1992; Arntz et al., 1997; Lovrich, 1997). Larval development described from plankton samples (Iorio, 1983).
Remarks: larval distribution reported; recognized as food of several fish species (Supplementary Online File 5). The Barcode is now available online via the Barcode of Life Data System (BOLD; www. barcodinglife.org, under code: PRIMB124-14, sampling site: 38°19.6’S; 57°30.4’W).
Family Cancridae Latreille, 1802
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Metacarcinus A. Milne-Edwards, 1862
Metacarcinus edwardsii (Bell, 1835) (jaiba mármola, jaiba mola, mole crab)
Geographic distribution: SEPO: from Ecuador to Canal Beagle, mostly associated to estuarine waters. Bathymetric distribution: infra and circalittoral (Vinuesa et al. 1999; Retamal, 2000; Stotz et al., 2007). Larval development described from laboratory reared larvae (Quintana, 1983).
Remarks: listed as Cancer edwarsii by Retamal (2000). Ecological studies in Chile (Supplementary Online File 5). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
SUPERFAMILY DORIPPOIDEA MacLeay, 1838
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Family Ethusidae Guinot, 1977
Ethusina Smith, 1884
Ethusina abyssicola Smith, 1884 (abyssal sumo crab) Figure 44. Ethusina abyssicola Smith, 1884. Preserved specimen. Photography of female in dorsal (up) and ventral (down)
view. Note the white sac beneath the abdomen which is the externa of an unidentified rhizocephalan barnacle. Photo by Emiliano Ocampo.
Geographic distribution: NWAO and SWAO: from New England, USA to Cabo Frio, Brazil, and in the deep and under-explored Mar del Plata Canyon (38°01´S; 53°39´W), Argentina. Bathymetric distribution: bathyal (Ocampo et al., 2014).
Remarks: “one of the best adapted brachyuran crabs to inhabit deep water” (Ocampo et al., 2014).
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Family Platyxanthidae Guinot, 1977
Danielethus Thoma, Ng and Felder, 2012
►● Danielethus crenulatus (A. Milne-Edwards, 1879) (purple stone crab, cangrejo de las rocas)
Figure 45. Danielethus crenulatus (A. Milne-Edwards, 1879). Live color. Photography of the body in dorsal view. Photo by
Nahuel Farias.
Geographic distribution: endemic of SWAO: from Rio Grande do Sul, Brazil, through Uruguay, to Golfo San Matías, Rio Negro, Argentina. Bathymetric distribution: intertidal and infralittoral (Bordin, 1987; Boschi et al., 1992; Melo, 1998). Larval development completely described from laboratory reared larvae (Menu-Marque, 1970).
Remarks: listed by Boschi et al. 1992 and Zolessi and Philippi (1995) as Platyxanthus crenulatus; it presumably has an important ecological role as a predator in benthic communities; functional morphology of female reproductive system described (Supplementary Online File 5). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: INVMP057-13, sampling site: 38°2.2’S; 57°31.5’W).
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►● Danielethus patagonicus (A. Milne-Edwards, 1879) (Patagonian stone crab, red stone crab, cangrejo de las rocas)
Figure 46. Danielethus patagonicus (A. Milne-Edwards, 1879). Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: endemic of SWAO: from off Rio Grande do Sul, through Uruguay, to Chubut, Argentina. Bathymetric distribution: circalittoral (Bordin, 1987; Boschi et al., 1992; Melo, 1998). Larval development completely described from laboratory reared larvae (Iorio and Boschi, 1986).
Remarks: listed by Boschi et al., 1992 and Zolessi and Philippi (1995) as Platyxanthus patagonicus; edible crab; life history studied in Patagonia (Supplementary Online File 5). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: PRIMB133-14, sampling site: 38°19.6’S; 57°30.4’W).
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SUPERFAMILY GONEPLACOIDEA MacLeay, 1838
Family Chasmocarcinidae Serène, 1964
Chasmocarcinus Rathbun, 1898b
● Chasmocarcinus typicus Rathbun, 1898b
Geographic distribution: NWAO, SWAO. SWAO: from Bahia to Rio Grande do Sul, Brazil, and Uruguay. Bathymetric distribution: circalittoral (Vezzani, 2007).
Remarks: Bordin (1987) and Zolessi and Philippi (1995) reported C. rathbuni Bouvier, 1917 from Maldonado (Uruguay) but Coelho (1997b) considered that the studied specimens were confused with juvenile C. typicus. However, all specimens studied by Vezzani (2007) were collected in Brazilian waters (Fig. 2).
Family Mathildellidae Karasawa & Kato, 2003
Neopilumnoplax Serène, 1969
Neopilumnoplax lipkeholthuisi Tavares & Melo, 2010
Geographic distribution: endemic of SWAO. The species was described and is known only from the southwestern Atlantic Ocean. The few records to date are distributed from the mouth of Rio de la Plata to south Brazil, and at 126 to 207 m depth (Tavares and Melo, 2010).
Remarks: the species resembles Neopilumnoplax americana (Rathbun,1898) but can be easily separated from its congeners by a suite of carapace and appendage characters (Tavares and Melo, 2010)
SUPERFAMILY LEUCOSIOIDEA Samouelle, 1819
Family Leucosiidae Samouelle, 1819
Subfamily Ebaliinae Stimpson, 1871b
Ebalia Leach, 1817
► Ebalia rotundata (A. Milne-Edwards, 1880b)
Geographic distribution: endemic of the SWAO: from Rio Grande do Sul, Brazil, to Santa Cruz, Argentina. Bathymetric distribution: circalittoral (Boschi et al., 1992; Torres and Melo, 2001; Giberto and Bremec, 2003).
Remarks: its geographical range had been restricted to Patagonian waters until 2000, on the basis of two specimens of the first description and numerous specimens from Golfo San Matías, Rio Negro,
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Argentina. Later, its occurrence off Argentina, Uruguay and Rio Grande (Brazil) has been reported, extending its geographical distribution more than 700 km northwards, (Torres and Melo, 2001; Giberto and Bremec, 2003). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: PRIMB081-14, sampling site: 41°9.0’S; 62°20.0’W).
Myropsis Stimpson, 1871
● Myropsis quinquespinosa Stimpson, 1871 (fivespine purse crab)
Geographic distribution: NWAO, SWAO. SWAO: from Rio de Janeiro to Rio Grande do Sul, Brazil, Uruguay (Maldonado), Argentina. Bathymetric distribution: circalittoral and bathyal (Bordin, 1987; Souza, 1997; Melo, 2010).
Remarks: the reference to “Argentina” did not inform about the collecting site.
Persephona Leach, 1817b
● Persephona mediterranea (Herbst, 1794) (mottled purse crab)
Geographic distribution: NWAO, SWAO. SWAO: from Amapa to Rio Grande do Sul, Brazil, off Uruguay from Brazilian border to Punta del Este. Bathymetric distribution: intertidal to circalittoral (Martínez et al., 2009; Magalhaes et al., 2016 and references thereof). Larval development completely described from laboratory reared larvae (Negreiros Fransozo et al., 1989; Hirose et al., 2012).
Remarks: ecology and reproductive biology studied in Brazil (Supplementary Online File 5). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www. barcodinglife.org).
Subfamily Leucosiinae Samouelle, 1819
Leucosia Weber, 1795
►† Leucosia planata (Fabricius, 1793)
Geographic distribution: SWAO: Tierra del Fuego Province, Argentina. Bathymetric distribution undescribed.
Remarks: the type specimen is the only one that has been described up to the present, but no “with enough detail to enable one to place it with certainty” (Rathbun, 1937).
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SUPERFAMILY MAJOIDEA Samouelle, 1819 (spider crab, cangrejo araña, caranguejo-aranha)
Family Epialtidae MacLeay, 1838
Subfamily Epialtinae MacLeay, 1838
Leucippa H. Milne Edwards, 1834 [in H. Milne Edwards, 1834–1840]
►● Leucippa pentagona H. Milne Edwards, 1834
Figure 47. Leucippa pentagona H. Milne Edwards, 1834. Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: NEPO and SEPO: from Baja California (Mexico) to Estrecho de Magallanes (Chile), with apparent separate populations in northern and southern hemisphere. SWAO: Ubatuba, São Paulo, Brazil; Uruguay; from Buenos Aires to Chubut, Argentina. Bathymetric distribution: infralittoral and circalittoral (Boschi et al. 1992¸ Pohle and Marques, 2003). Larval development completely described from laboratory reared larvae (Pohle and Marques, 2003).
Remarks: ecologic and reproductive biology studied (Supplementary Online File 5). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: PRIMB030-14, sampling site: 40°6.1’S; 61°55.0’W).
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Taliepus A. Milne-Edwards, 1878
●† Taliepus dentatus and T. marginatus
General remark: the presence of both species from the SEPO in Uruguay, and even Argentina and Brazil, reported by Barattini and Ureta (1960, cited by Zolessi and Philippi, 1995) is very dubious. However, Taliepus dentatus has been cited as a rare alien species in Brazil (Tavares, 2011).
Remarks: the Barcode of T. dentatus has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org)
Subfamily Pisinae Dana, 1851d
Lepteces Rathbun, 1893
Lepteces ornatus Rathbun, 1903
Geographic distribution: NWAO. SWAO: Uruguay. Bathymetric distribution: circalittoral (Melo and Crivelaro, 2002).
Remarks: one specimen collected at 34°45’ S; 52°05’W, previously reported only in the type locality, in Yucatan, Mexico (Melo and Crivelaro, 2002).
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Libidoclaea H. Milne Edwards and Lucas, 1842 [in H. Milne Edwards and Lucas, 1842–1844]
►● Libidoclea granaria H. Milne Edwards and Lucas, 1842
Figure 48. Libidoclea granaria H. Milne Edwards and Lucas, 1842. Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: SEPO: from Valparaiso to Canal Beagle. SWAO: Rio Grande do Sul, Brazil; Uruguay; from Buenos Aires to Tierra del Fuego, Argentina. Bathymetric distribution: infralittoral and circalittoral (Barattini and Ureta, 1960 quoted by Zolessi and Philippi, 1995; Boschi et al., 1992; Vinuesa et al., 1999; Melo, 2010). Larval development completely described from laboratory reared larvae (Fagetti, 1969).
Remarks: larval distribution, epibiosis, morphometry and reproductive traits were studied in Chile and Argentina (Supplementary Online File 5). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: PRIMB056-14, sampling site: 40°34.9’S; 59°10.1’W).
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Libinia Leach, 1815
● Libinia ferreirae Brito Capello, 1871
Geographic distribution: NWAO, SWAO. SWAO: from Amapá to Rio Grande do Sul, Brazil, and Uruguay. Bathymetric distribution: intertidal to circalittoral (Tavares and Santana, 2012 and references thereof). Larval development completely described from laboratory reared larvae (Bakker et al., 1990).
Remarks: it inhabits very similar environments with Libinia spinosa and their geographic and bathymetric distributions overlap for about 3000 km along the southwestern Atlantic coast (from Espírito Santo, Brazil to Uruguay); both species are morphologically very similar and their differences were revised by Tavares and Santana (2012). Larval physiology, reproductive biology and symbiotic associations studied (Supplementary Online File 5).
►● Libinia spinosa H. Milne Edwards in Guérin, 1832
Figure 49. Libinia spinosa H. Milne Edwards in Guérin, 1832. Female. Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: endemic of SWAO: from Espírito Santo, Brazil, through Uruguay to Chubut, Argentina. Bathymetric distribution: intertidal to circalittoral (Tavares and Santana, 2012 and references thereof). Larval development completely described from laboratory reared larvae (Boschi and Scelzo, 1968; Clark et al., 1998).
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Remarks: listed by Boschi et al. (1992) as Libinia spinosa H. Milne Edwards, 1834; the authorship now modified by Tavares and Santana (2012). Not distributed in Chile, as reported by Boschi et al. (1992). Ecology and reproductive biology, including behavior, has been described in detail; symbiotic associations studied (Supplementary Online File 5). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: PRIMB117-14, sampling site: 38°19.6’S; 57°30.4’W).
Pelia Bell, 1836
►● Pelia rotunda A. Milne-Edwards, 1875 [in A. Milne-Edwards, 1873–1880] (southern teardrop crab)
Figure 50. Pelia rotunda A. Milne-Edwards, 1875. Male. Preserved specimen. Photography of the body in dorsal view after being cleared of epibionts. The top-left miniature shows the usual aspect of living specimens covered mostly by sea sponges. Photo by Nahuel Farias.
Geographic distribution: endemic of SWAO: from Pará to Rio Grande do Sul, Brazil, Uruguay, and from Buenos Aires to Rio Negro, Argentina. Bathymetric distribution: intertidal to circalittoral (Boschi et al., 1992; Melo, 1998).
Remarks: the Barcode is now available online via the Barcode of Life Data System (BOLD; www. barcodinglife.org, under code: INVMP029-13, sampling site: 38°2.2’S; 57°31.4’W).
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Rochinia A. Milne-Edwards, 1875 [in A. Milne-Edwards, 1873–1880]
►● Rochinia gracilipes A. Milne-Edwards, 1875 [in A. Milne-Edwards, 1873–1880]
Figure 51. Rochinia gracilipes A. Milne-Edwards, 1875. Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: SO: South Shetland Island. Endemic of SWAO: from Cabo Frio, Rio de Janeiro, Brazil, to Cago de Hornos, Argentina. Bathymetric distribution: infra and circalittoral (Rathbun, 1925; Boschi et al., 1992; Melo, 1998; Griffiths et al. 2014; Pettan and Tavares, 2014). Larval development completely described from laboratory reared larvae (Luppi and Spivak, 2016).
Remarks: type species of Rochinia. The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: PRIMB126-14, sampling site: 38°19.6’S; 57°30.4’W).
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Stenocionops A. G. Desmarest, 1823
● Stenocionops furcatus (Olivier, 1791) (decorator crab)
Geographic distribution: NWAO, SWAO. SWAO: from Ceará to Rio Grande do Sul, Brazil and Maldonado, Uruguay. Bathymetric distribution: intertidal to circalittoral (Bordin, 1987; Melo, 2010). Larval development completely described from laboratory reared larvae (Santana et al., 2004).
Remarks: listed by Zolessi and Philippi (1995) as S. furcata. The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Family Inachidae MacLeay, 1838
Anisonotus A. Milne-Edwards, 1879 [in A. Milne-Edwards, 1873–1880]
● Anisonotus atlanticus (Coelho, 1997a)
Geographic distribution: endemic of SWAO: Santa Catarina and Rio Grande do Sul, Brazil, and in Brazilian waters off Uruguay. Bathymetric distribution: circalittoral (Coelho, 2005; Melo, 2010).
Remarks: reported as Podochela algicola by Bordin (1987) and by Zolessi and Philippi (1995). Renamed by Coelho (2005).
96 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Eurypodius Guérin, 1828
► Eurypodius latreillii Guérin, 1828
Figure 52. Eurypodius latreillii Guérin, 1828. Preserved specimen. Photography of the body in dorsal view after being cleared of epibionts. The top-left miniature shows the usual aspect of living specimens. Photo by Nahuel Farias.
Geographic distribution: SEPO. Bathymetric distribution: circalittoral (Rathbun, 1925; Boschi et al., 1992).
Remarks: reports from SWAO, off Buenos Aires, off Golfo San Matías, Rio Negro, Tierra del Fuego and Canal Beagle, Argentina, and the Malvinas Islands by Boschi et al. (1992) apparently correspond to E. longirostris.on the basis of morphology and genetics (Farías et al., unpublished). There are reports from Uruguay (erroneously Rio Grande do Sul) by Melo (2010) that should be verified. Reports from shallow waters of the east coast of Tierra del Fuego, Argentina (López Gappa and Sueiro, 2007) should be verified. The Barcode is now available online via the Barcode of Life Data System (BOLD; www. barcodinglife.org, under code: PRIMB052-14, sampling site: 40°35.6’S, 59°44.0’W).
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Eurypodius longirostris Miers, 1885
Geographic distribution: SEPO: from Quintero (32.8ºS) to the Estrecho de Magallanes. SWAO: Canal Beagle, the Malvinas Islands, Banco Burdwood, Argentina; apparently also off Buenos Aires, off Golfo San Matías, Rio Negro, Tierra del Fuego (Arntz et al., 1999; Lovrich, 1999; Pérez Barros et al., 2004; Farías et al., unpublished). Bathymetric distribution: bathyal, typically deeper than E. latreillii.
Remarks: morphologically similar to E. latreillii and frequently mis-identified. Genetically different from its relative (Meyer et al., 2013). According to these authors, both species were found in Chile and have a distinct geographical distribution pattern: E. latreillii between 36º and 43ºS and E. longirostris between 43ºS and 53ºS. Consequently, the larval development completely of E. latreillii described from laboratory reared larvae (Campodónico and Guzmán, 1972b), as well as the studies of population and reproductive biology carried on in southern Chile, and of magnesium regulation related to life in polar waters, should correspond to E. longirostris (Supplementary Online File 5). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Family Inachoididae Dana, 1851d
Stenorhynchus Lamarck, 1818
● Stenorhynchus seticornis (Herbst, 1788) (yellowline arrow crab, arrow crab)
Geographic distribution: NWAO, SWAO. SWAO: from Amapá to Rio Grande do Sul, Brazil, including Brazilian waters off Uruguay and Argentina (with no indication of locality). Bathymetric distribution: intertidal to bathyal (Rathbun, 1925; Melo, 2010; Guinot, 2012 and references thereof). Larval development completely described (Yang, 1976).
Remarks: its presence in Argentina needs to be validated. It is commonly associated with sponges, stony corals, soft corals, gorgonians, anemones and echinoderms and it is considered an “ornamental species”. Reproductive traits were studied in Brazil as well as larval eco-physiology, symbiotic associations and sperm ultrastructure (Supplementary Online File 5). It represents “a rare case in Brachyura, where ovigerous females occur throughout the year and of copulation frequently involving ovigerous females; reproductive strategy is also found in Hymenosomatidae, considered closely related to Inachoididae” (Guinot 2012). In fact, Guinot (2012) transferred the genus Stenorhynchus from Inachidae to Inachoididae on the basis of adult and larval morphology and behavior, and resurrected the Subfamily Stenorhynchinae. The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
98 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Collodes Stimpson, 1860b
►● Collodes rostratus A. Milne-Edwards, 1879
Figure 53. Collodes rostratus A. Milne-Edwards, 1879). Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: endemic of SWAO: from Rio de Janeiro to Rio Grande do Sul, Brazil, Uruguay and from Buenos Aires to Rio Negro, Argentina. Bathymetric distribution: circalittoral (Boschi et al., 1992; Melo, 2010).
Remarks: the Barcode is now available online via the Barcode of Life Data System (BOLD; www. barcodinglife.org, under code: PRIMB151-14, sampling site: 38°19.6’S; 57°30.4’W).
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Leurocyclus Rathbun, 1897a
►● Leurocyclus tuberculosus (H. Milne Edwards and Lucas, 1842) (knobbed spider crab)
Figure 54. Leurocyclus tuberculosus (H. Milne Edwards and Lucas, 1842). Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: SWAO: from Rio de Janeiro to Rio Grande do Sul, Brazil, Uruguay and from Buenos Aires to, at least, Chubut, Argentina. Bathymetric distribution: circalittoral (Boschi et al., 1992; Melo, 2010. Its presence in SEPO mentioned by these authors should be corroborated; the available information consists on the holotype (from Chile, but without certainty) and a specimen from this country described by Brito Capello (1871). Larval development completely described from laboratory reared larvae (Santana and Marques, 2009).
Remarks: according to Guinot (1984) L. gracilipes (listed by Boschi, 1964 and Zolessi and Philippi, 1995) are, in fact, juveniles and females of L. tuberculosus. The reproductive biology of this species has been studied in Argentina and aspects of its biology in Brazil (Supplementary Online File 5). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: PRIMB146-14, sampling site: 38°19.6’S; 57°30.4’W).
100 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Pyromaia Stimpson, 1871b
Pyromaia tuberculata (Lockington, 1877)
Figure 55. Pyromaia tuberculata (Lockington, 1877). Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: native to Northeastern Pacific Ocean coasts, from México to Colombia (Rathbun, 1925; Garth, 1958), it is now successfully established in several distant regions (Guinot, 2012 and references thereof). SWAO: introduced, from Rio de Janeiro to Paraná, Brazil, and in Buenos Aires, Argentina. Bathymetric distribution: infralittoral and circalittoral (Melo et al., 1989; Schejter et al., 2002). Larval development: zoeal morphology described from Pacific specimens (Webber and Wear, 1981; Terada, 1983); the complete larval development as well as juvenile morphology and growth were described from laboratory reared larvae (Fransozo and Negreiros-Fransozo, 1997; Luppi and Spivak, 2003; Oh and Ko, 2010).
Remarks: it is an invasive species found for the first time in coastal waters of Argentina in 2000, probably transported from Brazil (Schejter et al., 2002; Tavares, 2011). Life history studied in Japan; sperm ultrastructure described (Supplementary Online File 5). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: MEJII019-14, sampling site: 36°55.1’S; 55°18.2’W).
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Family Mithracidae MacLeay, 1838
Amphithrax Windsor & Felder, 2017
Amphithrax besnardi (Melo, 1990)
Geographic distribution: SWAO, Uruguay. Bathymetric distribution: bathial (Melo, 2010).
Remarks: originally named Mithrax besnardi (Melo, 1990) on the basis of “The only existing specimens (that) were on a mud bottom and a depth of 2-100 meters” and erroneously reported from Rio Grande do Sul, Brazil (Melo, 2010).
Family Oregoniidae Garth, 1958
Hyas Leach, 1814
Hyas araneus (Linnaeus, 1758)
Geographic distribution: AO, NEAO, SO: a male and a female were collected in the South Shetland Island (61°05’93”S, 55°47’07”W, 92 m) in 1986, but the species has not been re-collected in that region (Tavares and Melo, 2004). Bathymetric distribution: circalittoral. Larval development completely described from laboratory reared larvae (Christiansen, 1973).
Remarks: Tavares and Melo (2004) stated that “Whether the introduction of H. araneus has failed or been successful but actually gone unnoticed, must await acquisition of new data”. Model species for studies on larval physiology in the Northern Sea (e.g. Schiffer et al., 2013 and references thereof). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www. barcodinglife.org).
102 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
SUPERFAMILY HYMENOSOMATOIDEA MacLeay, 1838
Family Hymenosomatidae MacLeay, 1838
Halicarcinus White 1846
► Halicarcinus planatus (Fabricius, 1775) (flattened crab, cangrejito aplanado)
Figure 56. Halicarcinus planatus (Fabricius, 1775). Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: southern tip of South America and in shallow waters associated with the Antarctic Circumpolar Current. SWPO: New Zealand, Macquarie, Campbell and Auckland islands. SEPO, from Bahía de Taltal (35°S) to Cape Horn, Chile. IO: Prince Edward, Crozet and Kerguelen Islands. SWAO: from Mar del Plata, Buenos Aires Province (36°S) to Tierra del Fuego Province, Argentina, the Malvinas Islands. Bathymetric distribution: intertidal and infralittoral (Melrose, 1975; Boschi et al., 1992; Arntz et al., 1999). Larval development described from laboratory reared specimens (Boschi et al., 1969) and comprises three zoeae not followed by a megalopa stage. This peculiar pattern is typical of marine Hymenosomatidae (Lucas, 1980).
Remarks: populational, reproductive and larval biology studied in Argentina; model species for the study of magnesium regulation in polar areas. Occasionally found in Brazil, and considered alien species (Tavares, 2011). Hypothesized as a potential invader of shelf Antarctic environments and, in
103 FRENTE MARÍTIMO Publicación de la Comisión Técnica Mixta del Frente Marítimo fact, it was recently found in the shallow infralittoral zone at Deception Island, an emergent volcanic cone located off the Western Antarctic Peninsula. (Supplementary Online File 5). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: INVMP063-13, sampling site: 38°2.5’S; 57°32.3’W).
SUPERFAMILY PARTHENOPOIDEA MacLeay, 1838
Family Parthenopidae MacLeay, 1838
Subfamily Parthenopinae MacLeay, 1838 (elbow crab)
Mesorhoea Stimpson, 1871b
● Mesorhoea sexspinosa Stimpson, 1871b (sixspine elbow crab)
Geographic distribution: NWAO, SWAO. SWAO: from Pará to Rio Grande do Sul, Brazil, and Brazilian waters off Uruguay Bathymetric distribution: intertidal to circalittoral (Bordin, 1987; Melo, 2010).
Piloslambrus S. H. Tan and Ng, 2007
● Piloslambrus guerinii (Brito Capello, 1871)
Geographic distribution: NWAO, SWAO. SWAO: from Rio Grande do Norte to Rio Grande do Sul, Brazil, and Cabo Polonio, Uruguay. Bathymetric distribution: circalittoral (Bordin, 1987).
Remarks: mentioned as Parthenope (Platylambrus) guerinii by Bordin (1987) and by Zolessi and Philippi (1995).
Spinolambrus S. H. Tan and Ng, 2007
● Spinolambrus fraterculus (Stimpson, 1871b)
Geographic distribution: NWAO, SWAO. SWAO: from Amapá to Rio Grande do Sul, Brazil, Uruguay and, Argentina. Bathymetric distribution: infralittoral and circalittoral (Melo, 1990; 2010).
Remarks: listed by Zolessi and Philippi (1995) as Parthenope (Platylambrus) aylthoni. Reported also as Parthenope (Platylambrus) aylthoni for Uruguay and Argentina (38º05’S; 56º50’W) (Melo, 1990).
● Spinolambrus meridionalis (Boschi, 1965)
Geographic distribution: endemic of SWAO: Rio Grande do Sul, Brazil, and Rocha, Uruguay. Bathymetric distribution: circalittoral (Boschi, 1965; Bordin, 1987).
Remarks: listed by Zolessi and Philippi (1995) as Lambrus meridionalis.
104 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
SUPERFAMILY PILUMNOIDEA Samouelle, 1819
Family Galenidae Alcock, 1898
Subfamily Pilumninae Samouelle, 1819
Pilumnus Leach, 1816
►● Pilumnus reticulatus Stimpson, 1860b (hairy crab)
Figure 57. Pilumnus reticulatus Stimpson, 1860. Male. Dorsal and ventral view. Photo by Nahuel Farias.
Geographic distribution: NWAO, NEPO, SWAO. SWAO: from Pernambuco to Rio Grande do Sul, Brazil, Uruguay, from Buenos Aires to Rio Negro, Argentina. Bathymetric distribution: circalittoral (Rathbun, 1930, Bordin, 1987, Boschi et al., 1992). Larval development described from laboratory reared larvae (Montú et al., 1989; Spivak and Rodríguez, 2002).
Remarks: Rathbun (1930) notes that there is considerable morphological variation, and for convenience recognizes two forms, besides the “typical” one, “forma fragosa” and “forma tessellata = meridionalis”. South American specimens belong to “tessellata” (Boschi, 1964).
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SUPERFAMILY PORTUNOIDEA Rafinesque, 1815
Family Carcinidae MacLeay, 1838 Subfamily Carcininae MacLeay, 1838
Carcinus Leach, 1814 [in Leach, 1813–1815]
Carcinus maenas (Linnaeus, 1758) (shore crab, European green crab, cangrejo verde europeo)
Figure 58. Carcinus maenas (Linnaeus, 1758). Live color. Photography of the body in dorsal view. Photo by Nicolás Battini
Geographic distribution: it is a highly invasive species native from the northeast Atlantic, from North Africa to Norway (70°N). It was introduced to the east coast of North America, the northeastern Pacific, southern Australia, Tasmania, Japan, South Africa and Indian Ocean. SWAO: Chubut Province, Argentine and off Pernambuco, Brazil. Bathymetric distribution: intertidal to circalittoral (Rathbun, 1930; Hidalgo et al., 2005 and references thereof). Larval development described by Rice and Ingle (1975) from laboratory reared larvae.
Remarks: in Argentine coasts it had been discovered for the first time in 2003 in Bahía Camarones (44°79’S) and in 2015 it was found more to the north in Golfo Nuevo (42°46’S) (Hidalgo et al., 2005; Torres and González Pisani, 2016). Important model species for ecological and physiological studies of adults and larvae and, according to the Scopus database, 1067 articles published between 1895 and 2016 included the name of this species in their titles. A review of the biological studies of C. maenas has been published in 2007; growth and reproduction in Patagonia were also studied (Supplementary Online File 5). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
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Subfamily Polybiinae Ortmann, 1893b
Ovalipes Rathbun, 1898a
►● Ovalipes trimaculatus (De Haan, 1833) (paddle crab, southern surf crab, cangrejo nadador, jaiba nadadora)
Figure 59. Ovalipes trimaculatus (De Haan, 1833). Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: NWPO, SWAO, SEAO, SEPO, SWPO, WIO, EIO. SWAO: from São Paulo to Rio Grande do Sul, Brazil, Uruguay, from Buenos Aires to Chubut, Argentina. Bathymetric distribution: infralittoral and circalittoral (Bordin, 1989; Boschi et al., 1992; Melo, 1998; Martelli et al., 2016 and references thereof). Larval development (zoeal stages) described from laboratory reared larvae (Schoeman and Cockcroft, 1996).
Remarks: caught in Argentina, mostly as by-catch in artisanal bottom trawl fisheries. Annual landings have never historically exceeded 800 Tons (Boschi, 1997). Reproductive biology, development, growth, molt, regeneration, effects of pollution and behavior studied (Wyngaardet al., 2016; Supplementary Online File 5). The reports of O. catharus and O. punctatus by Zolessi and Philippi are erroneous. The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: PRIMB100-14, sampling site: 38°12.7’S; 57°25.0’W).
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Family Geryonidae Colosi, 1923
Chaceon Manning and Holthuis, 1989
►● Chaceon notialis Manning and Holthuis, 1989 (red crab, deep sea red crab, cangrejo rojo)
Figure 60. Chaceon notialis Manning and Holthuis, 1989. Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: endemic of the SWAO: from off Rio Grande, Brazil, through off Uruguay to off Buenos Aires, Argentina. Bathymetric distribution: bathyal (Boschi et al., 1992; Pezzuto et al., 2002).
Remarks: Mantelatto et al. (2014) detected the existence of at least two different groups within this species that “might indicate the existence of cryptic species”. Reproductive biology and specially fisheries of this crab studied in Brazil and Uruguay (Supplementary Online File 5). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife. org).
108 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Family Macropipidae Stephenson and Campbell, 1960
Coenophthalmus A. Milne-Edwards, 1879 [in A. Milne-Edwards, 1873–1880]
►● Coenophthalmus tridentatus A. Milne-Edwards, 1879
Figure 61. Coenophthalmus tridentatus A. Milne-Edwards, 1879. Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: endemic of SWAO: from Rio de Janeiro to Rio Grande do Sul, Brazil, Uruguay, and from Buenos Aires to Chubut, Argentina. Bathymetric distribution: infralittoral/circalittoral (Boschi et al., 1992; Melo, 1998). Larval development (first zoea) described from laboratory hatched larvae (Spivak and Luppi, 2004).
Remarks: the Barcode is now available online via the Barcode of Life Data System (BOLD; www. barcodinglife.org, under code: PRIMB138-14, sampling site: 38°19.6’S; 57°30.4’W).
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Family Portunidae Rafinesque, 1815
Subfamily Portuninae Rafinesque, 1815
Arenaeus Dana, 1851
● Arenaeus cribrarius (Lamarck, 1818) (speckled swimming crab, siri chita, cangrejo nadador)
Geographic distribution: NWAO, SWAO. SWAO: from Ceará to Rio Grande do Sul, Brazil, Buenos Aires, Argentina. Bathymetric distribution: intertidal to circalittoral (Rathbun, 1930; Melo, 1998; Scelzo, 2001). Larval development completely described from laboratory reared larvae (Stuck and Truesdale, 1988).
Remarks: reported for the first time in Argentina in Mar del Plata, Buenos Aires, in 2000 (Scelzo, 2001); mentioned as an example of extraliminal distribution by Orensanz et al. (2002). Population and reproductive biology, epibionts, symbionts and parasites, studied in Brazil (Supplementary Online File 5). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Callinectes Stimpson, 1860b (blue crab, cangrejo azul, cangrejo nadador, siri)
►● Callinectes sapidus Rathbun, 1896
Geographic distribution: World wide distributed as invader in Norteastern Atlantic coasts (Europe) including Mediterranean and Black Seas as well as in Northeastern Pacific coasts, Hawaian Islands and Japan (see Kennedy and Cronin, 2007). NEAO and SWAO: occurs naturally along the Atlantic coast of America, from Nova Scotia (Canada) to Rio de la Plata (Argentina-Uruguay). Bathymetric distribution: infralittoral, including brackish water of estuaries and occasionally fresh water, to 37 m (Rathbun, 1930; Ringuelet, 1963; Cesar et al., 2003). Larval development described from laboratory reared larvae (Costlow and Bookhout, 1959; Kennedy, 2007).
Remarks: important commercial crab and the second most studied species of the Infraorder Brachyura (after Carcinus maenas); the Scopus database records 820 papers which included the name of the species in the title, published between 1914 and 2016. Its morphology, anatomy, biology, physiology, ecology, symbiotic relationships, development, life history and fisheries has been deeply reviewed in a book edited by Kennedy and Cronin (2007) with contributions of many authors. Recent research on C. sapidus in the Southwestern Atlantic shores of Brazil included studies on population and reproductive biology, age determination, physiology and toxicology (Supplementary Online File 5). Adults live in shallow estuaries, and females migrate to the sea to lay their eggs. After the eggs have hatched, development took place in the sea and the young crabs migrate to estuaries; regional examples of this type of migration are described in Uruguayan coastal lakes (Verdi and Delgado, 2001; Uscudún Melinkov, 2014). Zolessi and Philippi (1995) reported also C. acutidens as a different species but is a synonymous. The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
110 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
● Callinectes danae Smith, 1869b
Geographic distribution: NWAO, SWAO. SWAO: from Paraíba to Rio Grande do Sul, Brazil; Uruguay. Introduced to Northeastern Atlantic waters including Mediterranean Sea (Rathbun, 1930; Santana, 1987; Melo, 1998). Bathymetric distribution intertidal to circalittoral (including brackish to hypersaline waters, in mangroves and estuaries) (Melo, 1996). Larval development (first zoea) described from laboratory hatched larvae (Sankarankutty et al., 1999).
Remarks: deeply studied in Brazil. The Scopus database includes 51 articles, from 1969 to 2016, dealing with populational and reproductive biology, physiology, toxicology, symbiosis and parasitism with Callinectes danae in the title (Supplementary Online File 5). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Achelous De Haan, 1833 (swimming crab, siri, cangrejo nadador)
● Achelous spinicarpus Stimpson, 1871b
Geographic distribution: NWAO, SWAO. SWAO: from Amapá to Rio Grande do Sul States, Brazil, and Maldonado, Uruguay. Bathymetric distribution: infralittoral to bathyal in NWAO, 9-550 m, intertidal and circalittoral, 0-20 m in SWAO (Williams, 1984; Bordin, 1987; Melo, 1998). Larval development completely described from laboratory reared larvae (Bookhout and Costlow, 1974).
Remarks: listed as Portunus (Achelous) spinicarpus by Zolessi and Philippi (1995). It is one of several American species of Portunus (Weber, 1795) transferred to the Achelous (Mantelatto et al., 2007, 2009b). Relative growth and reproduction studied in Brazilian waters and in the Gulf of Mexico (Supplementary Online File 5). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
● Achelous spinimanus (Latreille, 1819)
Geographic distribution: NWAO, SWAO. SWAO: from Pernambuco to Rio Grande do Sul States, Brazil, and Maldonado, Uruguay. Bathymetric distribution: circalittoral (Williams, 1984; Bordin, 1987; Melo, 1998). Larval development: megalopa described (Negreiros-Fransozo et al., 2007).
Remarks: listed as Portunus (Achelous) spinimanus by Zolessi and Philippi (1995). It is one of several American species of Portunus (Weber, 1795) transferred to the Achelous (Mantelatto et al., 2007, 2009b). Juvenile morphology and reproductive biology studied in Brazilian waters where it is commercially sold for human consumption (Supplementary Online File 5). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
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SUPERFAMILY PSEUDOZIOIDEA Alcock, 1898
Family Pilumnoididae Guinot and Macpherson, 1987
Pilumnoides Lucas, 1844 [in H. Milne Edwards and Lucas, 1842–1844]
►● Pilumnoides hassleri A. Milne-Edwards, 1880b
Figure 62. Pilumnoides hassleri A. Milne-Edwards, 1880b. Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: SEPO: Estrecho de Magallanes (Chile). SWAO: from Rio de Janeiro to Rio Grande do Sul, Brazil, Uruguay, and from Buenos Aires to Santa Cruz, Argentina. Bathymetric distribution: intertidal (Rathbun, 1930; Bordin, 1987; Boschi et al., 1992; Melo, 1998). Larval development described from laboratory reared larvae (Luppi et al., 2010).
Remarks: the Barcode is now available online via the Barcode of Life Data System (BOLD; www. barcodinglife.org, under code: PRIMB162-14, sampling site: 38°24.6’S; 57°36.1’W).
112 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
SUPERFAMILY XANTHOIDEA MacLeay, 1838
Family Panopeidae Ortmann, 1893b Subfamily Eucratopsinae Stimpson, 1871b
Cyrtoplax Rathbun, 1914
● Cyrtoplax spinidentata (Benedict, 1892b)
Geographic distribution: NWAO, SWAO. SWAO: from Ceará to Rio Grande do Sul, Brazil, Maldonado, Uruguay. Bathymetric distribution: infralittoral and circalittoral (Bordin, 1987; Melo, 1998, Bezerra et al., 2006).
Subfamily Panopeinae Ortmann, 1893b (mud crab, cangrejo del barro)
Acantholobulus Felder and J. W. Martin, 2003
►● Acantholobulus schmitti (Rathbun, 1930)
Figure 63. Acantholobulus schmitti (Rathbun, 1930). Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: endemic of SWAO: from Ceará to Rio Grande do Sul, Brazil, through Uruguay, to Mar del Plata, Buenos Aires, Argentina. Bathymetric distribution: intertidal and infralittoral (Boschi et al., 1992; Vieira et al., 1998; Melo, 1998). Larval development described from laboratory hatched larvae (Bakker et al., 1989; Rodríguez and Spivak (2001) as Hexapanopeus schmitti Rathbun, 1930 and Panopeus margentus Williams and Boschi, 1990, respectively.
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Remarks: listed by Boschi et al. (1992) as P. margentus and by Zolessi and Philippi (1995) as H. schmitti. Felder and Martin (2003) assigned H. schmitti to the new genus Acantholobulus and concluded that P. margentus is a junior synonym of A. schmitti.
Eurypanopeus A. Milne-Edwards, 1880 [in A. Milne-Edwards, 1873–1880]
● Eurypanopeus depressus Smith, 1869b (flatback mud crab)
Figure 64. Eurypanopeus depressus Smith, 1869. Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: NWAO, SWAO. SWAO: Uruguay, from Montevideo to Piriápolis, and Mar Chiquita Lagoon, Buenos Aires, Argentina. Bathymetric distribution: intertidal (Juanicó, 1978; Nizinski, 2003; Spivak and Luppi, 2006). Larval development described from laboratory hatched larvae (Costlow and Bookhout, 1961).
Remarks: the most common mud crab inhabiting oyster reefs along the east coast of the United States, co- occurring with Panopeus herbsti (McDonald, 1982; Meyer, 1994; Tolley et al., 2013 and references thereof). Since it was not found in Brazil (Melo, 1996) it is likely an invader (Tavares, 2011). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
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Hexapanopeus Rathbun, 1898b
● Hexapanopeus paulensis Rathbun, 1930
Geographic distribution: NWAO, SWAO. SWAO: from Ceará to Rio Grande do Sul, Brazil, to Uruguay. Bathymetric distribution: intertidal and infralittoral (Milstein et al., 1976; Williams, 1984; Melo, 1998). Larval development described from laboratory hatched larvae (Fransozo et al., 1990).
Remarks: according to Williams (1984), “reported from only a few widely separated areas”.
Panopeus H. Milne Edwards, 1834 [in H. Milne Edwards, 1834–1840]
►● Panopeus meridionalis Williams, 1984
Figure 65. Panopeus meridionalis Williams, 1984. Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: endemic of SWAO: Its presence has been only reported from Montevideo, Uruguay; “Rio de la Plata hasta Mar del Plata” and Mar Chiquita lagoon, Buenos Aires Province, Argentina. Bathymetric distribution: intertidal (Williams, 1984; Boschi et al., 1992; Spivak and Luppi, 2006). Larval development described from laboratory hatched larvae (Rodríguez et al., 2003)
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Remarks: Panopeus herbstii H. Milne Edwards 1834 sensu lato, has been considered a species complex that included 4 forms (Rathbun, 1930). However, Williams (1984) divided the complex into 6 species on the basis of morphometry, color pattern and hemocyanin analysis: P. herbsti sensu stricto, P. lacustris, P. obesus, P. simpsoni Rathbun, P. austrobesus, and P. meridionalis, the southernmost species of this group. P. austrobesus has been reported from Cabo Polonio, Rocha, Uruguay (Peluffo, 2004). However, the material has been only compared with literature and “some pictures from Brazilian P. austrobesus specimens from São Paulo”, Brazil. The morphological similitude among species of this complex made necessary a comparison with P. meridionalis. Consequently, this report should be considered dubious.
Tetraxanthus Rathbun, 1898b
● Tetraxanthus rathbunae Chace, 1939
Geographic distribution: NWAO, SWAO. SWAO: from Paraíba to Rio Grande do Sul, Brazil, Maldonado, Uruguay. Bathymetric distribution: circalittoral (Bordin, 1987; Melo, 1998).
Remarks: listed by Zolessi and Philippi (1995) as T. bidentatus (Milne Edwards, 1880).
Family Pseudorhombilidae Alcock, 1900
Speocarcinus Stimpson, 1859
Speocarcinus dentatus Brandão, Coelho-Filho & Tavares, 2012
Geographic distribution: SWAO: endemic, from Alagoas to Rio Grande do Sul, Brazil; Uruguay. Bathymetric distribution: circalittoral (Brandão, Coelho-Filho & Tavares, 2012).
Remarks: collected at 35°33’S; 53°48’W (Brandão, Coelho-Filho & Tavares, 2012). Previously reported as Speocarcinus carolinensis (Vezzani, 2007).
Family Xanthidae MacLeay, 1838
Subfamily Actaeinae Alcock, 1898
Actaea De Haan, 1833 [in De Haan, 1833–1850]
● Actaea acantha (H. Milne Edwards, 1834) (spinose rubble crab)
Geographic distribution: NWAO, SWAO. SWAO: from Amapá to Rio Grande do Sul, and oceanic islands, Brazil; Rocha and Maldonado, Uruguay. Bathymetric distribution: circalittoral (Rathbun, 1930; Bordin, 1987; Melo, 1998; Zolessi and Philippi, 1995).
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SUBSECTION THORACOTREMATA Guinot, 1977 SUPERFAMILY GRAPSOIDEA MacLeay, 1838
Family Grapsidae MacLeay, 1838
Pachygrapsus Randall, 1840
● Pachygrapsus transversus (Gibbes, 1850) (mottled shore crab)
Geographic distribution: NWAO, NEAO, NEPO, SWAO, SEAO, SEPO, SWPO. SWAO from Ceará to Rio Grande do Sul, Brazil; Montevideo, Uruguay. Bathymetric distribution: intertidal (Rathbun, 1918; Williams, 1984; Melo, 1998; Poupin, et al., 2005). Larval development: zoeal stages described from laboratory reared larvae (Hines, 1986; Cuesta and Rodríguez, 1994; Brossi-García and Rodrigues, 1997), megalopa described by Flores et al. (1998) and Arruda and Abrunhosa (2011).
Remarks: some characters used to identify this species vary according to size and/or sex (Poupin et al., 2005). The morphological and molecular variation of P. transversus from eastern, western Atlantic, and eastern Pacific populations found sufficient differentiation among these three populations that taxonomic separation can be considered and the reinstatement of P. socius Stimpson, 1871 for the east Pacific populations, based on colour pattern, adult morphometry, and DNA analysis (Cuesta and Schubart, 1998; Schubart et al., 2005). Population and reproductive biology, and foraging, have been studied in as well as physiology (Supplementary Online File 6). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Planes Bowdich, 1825 (Columbus crab, flotsam crab, oceanic crab, cangrejo de Colón)
● Planes major (MacLeay, 1838)
Geographic distribution: NWAO, NEAO, NEPO, SWAO, SWPO, WIO. SWAO: off Brazil and Uruguay. Bathymetric distribution: epipelagic, as symbiont of turtles or living on flotsam (Chace, 1951; Juanicó, 1976; Manning and Holthuis, 1981; Prado and Melo, 2002; Carranza et al., 2003; Pons et al., 2011).
Remarks: listed by Zolessi and Philippi (1995) as P. cyaneus Dana, 1851. Feeding, behavior and reproductive biology studied (Supplementary Online File 6).
Planes marinus Rathbun, 1914
Geographic distribution: NEPO, NWPO, SWAO, SWPO, Santa Helena Island. SWAO: Brazil, Mar Chiquita, Buenos Aires, Argentina. Bathymetric distribution: epipelagic, on flotsam and turtles (Dell, 1963; 1964; Spivak and Bas, 1999; Kepel et al., 2002; Prado and Melo, 2002). Larval development: first zoea described by Wear (1970).
Remarks: feeding and behavior studied (Supplementary Online File 6).
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Family Sesarmidae Dana, 1851f
Armases Abele, 1992
►● Armases rubripes (Rathbun, 1897b)
Geographic distribution: NWAO, SWAO. SWAO: from Ceará to Rio Grande do Sul, Brazil, Montevideo, Uruguay, and “the Rio de la Plata”. Bathymetric distribution intertidal (Rathbun, 1918). Larval development described from laboratory reared larvae hatched form females of Maracaibo estuary, Venezuela (Díaz and Ewald, 1968). Later, Montú et al. (1990) described an alternative way of the larval development, with five zoeae, in larvae from Pontal do Sul, Brazil. The larvae of both localities also showed some morphological differences, suggesting the existence of geographical races along the wide range of distribution of this species. Early juvenile development, morphology of the megalopa and first crab described (Negreiros-Fransozo et al., 2011).
Remarks: listed by Boschi et al. (1992) and Zolessi and Philippi (1995) as Metasesarma rubripes; these authors stated that is seldom found in Argentine waters. Luppi et al. (2003) argued that the Rio de la Plata seems to function as a biogeographic barrier to A. rubripes since its larvae did not tolerate reduced salinities. Ecology and reproductive biology studied in Brazil (Supplementary Online File 6).
●† Armases miersii Rathbun, 1897
Zolessi and Philippi (1995) listed this crab as Sesarma miersii as found in Isla de Ratas, Montevideo, Uruguay, but it is an erroneous report (Scarabino, 2006 and references thereof). These authors stated that this site is the type locality of this tropical species when, in fact, it is Bahamas (Rathbun, 1897). Rathbun (1897; 1918) included Montevideo in the geographical range of A. miersii on the basis of a the identification “with considerable hesitation” of “a male example collected at Isla Rata, Montevideo, with Chasmognathus granulatus” as Sesarma angustipes (Miers, 1881)
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Family Varunidae H. Milne Edwards, 1853
Subfamily Cyclograpsinae H. Milne Edwards, 1853
Neohelice Sakai, Türkay and Yang, 2006
►● Neohelice granulata (Dana, 1851). (cangrejo excavador, cangrejo estuarial, caranguejo, catanhão, gatanhão)
Figure 66. Neohelice granulata (Dana, 1851). Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: endemic of temperate SWAO: from Lagoa Araruama, Rio de Janeiro, Brazil (22°57’S; 42°50’W), through Uruguay, to Golfo San José, Chubut, Argentina (42°25’ S; 64°36’ W). Bathymetric distribution: intertidal (Spivak, 2010). Larval development: four or five zoeae (alternative developmental pathways), megalopae and first juvenile described from laboratory reared larvae (Boschi et al., 1967); Pestana and Ostrenski, 1995; Cuesta et al., 2001; 2002)
Remarks: listed by Boschi et al. (1992) and Zolessi and Philippi (1995) as Chasmagnathus granulata. Also known as C. granulatus. It is a key species in South American salt marshes, mudflats and estuaries; through this 20° latitudinal gradient, its habitats vary from mangroves in the north to the salt marshes of Northern Patagonia, in the south (Spivak, 2010). It is an important animal model species: “Beginning in the late 1989s, an explosion of publications appeared in international journals dealing with its ecology, physiology, toxicology and behavior. A bibliometric analysis using the Scopus database allowed detecting 309 papers that deal with this species during the period 1986–
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2009” (Spivak, 2010). Since then, 118 new papers have been added to this database. Supplementary Online File 7 contains an almost complete list of references about this. The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: PRIMB088-14, sampling site: 37°44.8’S; 57°26.2’W).
Subfamily Varuninae H. Milne Edwards, 1853
Cyrtograpsus Dana, 1851
►● Cyrtograpsus angulatus Dana, 1851
Figure 67. Cyrtograpsus angulatus Dana, 1851. Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: SEPO: San Lorenzo Island, Peru. SWAO: from Rio de Janeiro to Rio Grande do Sul, Brazil, trough Uruguay, to Santa Cruz, Argentina. Bathymetric distribution: intertidal and infralittoral (Rathbun, 1918; Boschi, 1964; Boschi, et al. 1992). Larval development completely described, and juvenile development partially described, from laboratory hatched larvae (Rieger and Vieira, 1997; Rieger and Beltrao, 2000; Luppi and Spivak, 2007).
Remarks: is an important species in coastal ecosystems of the region. There are many studies on life history, development, populational and reproductive biology, effects of pollution, morphology and histology, ecological relationships, physiology and parasitology, mainly in Argentina but also in Southern Brazil (Supplementary Online File 6). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: INVMP061-13, sampling site: 38°2.5’S; 57°32.3’W).
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►● Cyrtograpsus altimanus Rathbun, 1918
Figure 68. Cyrtograpsus altimanus Rathbun, 1918. Live color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: endemic of temperate waters of the Soutwestern Atlantic Ocean from Rio Grande do Sul State (Brazil) to Golfo San Matías (Rio Negro Province, Argentine). Bathymetric distribution: intertidal (Rathbun, 1918; Bordin, 1987; Boschi et al., 1992). Larval development completely described from laboratory hatched larvae (Szcelzo and Lichstchein de Bastida, 1978).
Remarks: morphometric comparisons between C. altimanus and C. affinis showed a continuous gradient rather than discrete differences and comparison of mitochondrial DNA sequences did not reveal diagnostic differences between them. It has been suggested a very close relationship and the possibility that the two forms represent a single species with a high ecological and phenotypic plasticity (Spivak and Schubart, 2003). Life history and physiology studied in Argentina (Supplementary Online File 6). The Barcode is now available online via the Barcode of Life Data System (BOLD; www. barcodinglife.org, under code: INVMP021-13, sampling site: 38°2.5’S; 57°32.3’W).
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►● Cyrtograpsus affinis (Dana, 1851)
Geographic distribution: endemic of temperate waters of SWAO: from Cabo San Roque, Rio Grande do Norte, Brazil, to Golfo Nuevo, Chubut, Argentina (Rathbun, 1918; Lezcano et al., 2012). However, the northern record corresponds to only one specimen studied by Rathbun (1918) and most specimens were collected from Río de la Plata to Nuevo Gulf. Bathymetric distribution: intertidal and infralittoral (Rathbun, 1918; Spivak and Cuesta, 2000; Spivak and Schubart, 2003; Lezcano et al., 2012). Larval development described from laboratory hatched larvae (Spivak and Cuesta, 2000).
Remarks: morphometric comparisons between C. altimanus and C. affinis showed a continuous gradient rather than discrete differences and comparison of mitochondrial DNA sequences did not reveal diagnostic differences between them. It has been suggested a very close relationship and the possibility that the two forms represent a single species with a high ecological and phenotypic plasticity (Spivak and Schubart, 2003; Lezcano et al., 2012). An association with the scyphozoan medusae Lychnorhiza lucerna was described (Supplementary Online File 6). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org, under code: PRIMB036-14, sampling site: 40°6’S; 61°54’W).
SUPERFAMILY OCYPODOIDEA Rafinesque, 1815
Family Ocypodidae Rafinesque, 1815
Subfamily Ocypodinae Rafinesque, 1815
Ocypode Weber, 1795
● Ocypode quadrata (Fabricius, 1787) (Atlantic ghost crab, cangrejo fantasma, caranguejo maria-farinha)
Geographic distribution: NWAO, SWAO. SWAO: from Para to Rio Grande do Sul, Brazil (Melo, 1998) and Uruguay (Scarabino, 2006 and references therof). Bathymetric distribution: supralittoral, in sandy beaches (Lucrezi and Schlacher, 2014). Larval development described from laboratory hatched larvae (Díaz and Costlow, 1972).
Remarks: many studies have been performed on this species in several countries including comparative morphology, ecology and behavior, physiology and its role as indicator of anthropic effects on sandy beaches (Lucrezi and Schlacher, 2014), several of them in Brazil (Supplementary Online File 6). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
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Subfamily Gelasiminae Miers, 1886
Leptuca Bott, 1973 (fiddler crab, cangrejo violinista, caranguejo-violinista, chama-maré)
►● Leptuca uruguayensis (Nobili, 1901)
Figure 69. Leptuca uruguayensis (Nobili, 1901). Male. Up-right miniature shows the living color. Photography of the body in dorsal view. Photo by Nahuel Farias.
Geographic distribution: endemic of SWAO: from Rio de Janeiro to Rio Grande do Sul in Brazil, Uruguay and from Bahía Samborombón to Quequén, Buenos Aires, Argentina. Bathymetric distribution: intertidal (Boschi et al., 1992; Melo 1998). Larval development described from laboratory hatched larvae (Rieger, 1996; Armendáriz, 2005).
Remarks: listed by Boschi et al. (1992) and Zolessi and Philippi (1995) as Uca uruguayensis. This species has been widely studied, mainly on its ecology and behavior, but also on its physiology and parasitic infections (Supplementary Online File 6). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
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SUPERFAMILY PINNOTHEROIDEA De Haan, 1833 [in De Haan, 1833–1850]
Family Pinnotheridae De Haan, 1833 [in De Haan, 1833–1850] (pea crab) Subfamily Pinnothereliinae Alcock, 1900
Austinixa Heard and Manning, 1997
►● Austinixa patagoniensis (Rathbun, 1918)
Geographic distribution: endemic of SWAO: from Amapá to Rio Grande do Sul, Brazil, Uruguay, and from Mar del Plata, Buenos Aires, to Golfo San Matías, Río Negro, Argentina. Bathymetric distribution: infralittoral and circalittoral related with its hosts (Fenucci, 1975; Manning and Felder, 1989; Melo, 1998).
Remarks: listed by Boschi et al. (1992) and Zolessi and Philippi (1995) as Pinnixa patagoniensis. Lives typically in burrows of the callianasid Sergio mirim and the polychaete Arenicola sp., in few cases has been sampled off their hosts. (Supplementary Online File 6). The Barcode is available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
Pinnixa White, 1846
► Pinnixa brevipollex Rathbun, 1898
Geographic distribution: SEPO: southern Chile. SWAO: from Rio de Janeiro to Rio Grande do Sul, Brazil and from Buenos Aires to Rio Negro, Argentina. Bathymetric distribution: intertidal and infralittoral (Fenucci, 1975; Martins and D´Incao, 1996; Bezerra et al., 2005; Ceseña et al., 2016).
Remarks: found in tubes of Chaetopterus variopedatus or off its host (Fenucci, 1975).
● Pinnixa chaetopterana Stimpson, 1860b
Geographic distribution: NWAO, SWAO. SWAO: from Pernambuco to Rio Grande do Sul States, Brazil, and from Maldonado to Chuy, Uruguay. Bathymetric distribution: intertidal and infralittoral (Bordin, 1987; Melo, 1998; McDermott, 2005). Larval development: first zoea described and illustrated by Hyman (1925).
Remarks: found in tubes of Chaetopterus variopedatus and other polychaetes. Its biology was studied in the USA (McDermott, 2005 and cites thereof).
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►● Pinnixa rapax Bouvier, 1917
Geographic distribution: endemic of SWAO: from Rio de Janeiro to Rio Grande do Sul, Uruguay and Mar del Plata, Buenos Aires, Argentina. Bathymetric distribution: circalittoral (Fenucci, 1975; Melo 1998).
Remarks: host unknown, sampled only free (Fenucci, 1975).
● Pinnixa sayana Stimpson, 1860b
Geographic distribution: NWAO, SWAO. SWAO: from Amapá to Rio Grande do Sul, Brazil, and Maldonado, Uruguay. Bathymetric distribution: infra to circalittoral (Williams, 1984; Bordin, 1987; Melo, 1998).
Remarks: live in burrows of Arenicola but also sampled off its host (Melo, 1998).
Pinnixa valdiviensis Rathbun, 1907
Geographic distribution: SEPO: from Chincha Islands, Peru (13°S) to southern Chile (55°S). SWAO: Bahía San Julián, Santa Cruz, Argentina. Bathymetric distribution: intertidal and infralittoral (Torres, 2006; Baeza and Hernáez, 2015; Ceseña et al., 2016).
Remarks: found in tubes of Chaetopterus variopedatus and Callichirus garthi, and as commensal of the echiurid Pinuca chilensis (Torres, 2006; Baeza and Hernáez, 2015). Populational and reproductive biology studied in Chile (Supplementary Online File 6). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
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Subfamily Pinnotherinae De Haan, 1833 [in De Haan, 1833–1850]
Calyptraeotheres Campos, 1990
► Calyptraeotheres garthi (Fenucci, 1975)
Figure 70. Calyptraeotheres garthi (Fenucci, 1975). Live color. Photography of a female within one of the slipper limpet host species (right), and dorsal views of the body of a female (up right) and the smaller male (down right). Photo by Emiliano Ocampo
Geographic distribution: endemic of the SWAO: from Mar del Plata, Buenos Aires, to Golfo San Matías, Rio Negro. Bathymetric distribution: intertidal and infralittoral (Fenucci, 1975; Ocampo et al., 2016). Larval development completely described from larvae hatched in the laboratory as well as juvenile development (Ocampo et al., 2011; Ocampo et al., 2016).
Remarks: listed by Boschi et al., (1992) as Pinnotheres garthi. Included in the genus Calyptraeotheres by Campos (1989a). It exclusively inhabits gastropods pertaining to the family Calyptraeidae (limpet snails), including Crepidula cachimilla, C. argentina, C. unguiformis, C protea and Bostrycapulus odites, among others (Ocampo et al.; 2014). Populational and reproductive biology studied in Argentina (Supplementary Online File 6). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
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Dissodactylus Smith, 1870
► Dissodactylus crinitichelis Moreira, 1901
Geographic distribution: NWAO, SWAO. SWAO: from Pará to Rio Grande do Sul, Brazil, and Mar del Plata, Buenos Aires Province, Argentina. Bathymetric distribution: intertidal to circalittoral (Fenucci, 1975; Williams, 1984; Melo 1998). Larval development described from laboratory hatched larvae (Pohle and Telford, 1981a; b).
Remarks: associated with echinoids Clypeasteroida (Encope emarginata, E. michelini, Clypeaster subdepressus, Leodia sexiesperforata, Mellita quinquiesperforata) and Spatangoida (Meoma ventricosa) and asteroids (Luidia sp.); both sexes leave their host for mate searching behavior. Host crab relationships were deeply, studied, in part in Brazil (Supplementary Online File 6). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www.barcodinglife. org).
Fabia Dana, 1851b
► Fabia emiliai (Melo, 1971)
Geographic distribution: endemic of SWAO: from Rio de Janeiro to Rio Grande do Sul, Brazil, and Mar del Plata, Buenos Aires, Argentina. Bathymetric distribution: infralittoral (Fenucci, 1975; Melo 1998).
Remarks: F. emiliai is the combination of P. emiliai Melo, 1971 and F. insularis Melo, 1971 (Fenucci, 1975; Campos, 1996; 2013); it is associated with the bivalves Glycymeris longior and Anadara brasiliana (Fenucci, 1975).
Pinnaxodes Heller, 1865
► Pinnaxodes chilensis (H. Milne Edwards, 1837 in H. Milne Edwards, 1834-1840).
Geographic distribution: SEPO: its presence in Argentine waters of Tierra del Fuego has not been confirmed. Bathymetric distribution: infralittoral zone (Fenucci, 1975; Lardies and Castilla, 2001).
Remarks: not found in Argentine waters of Tierra del Fuego (Beagle Channel) but the distribution of its host, the echinoid Loxechinus albus, suggested that it should live there (Fenucci, 1975). The Barcode has been previously available online via the Barcode of Life Data System (BOLD; www. barcodinglife.org).
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Tumidotheres Campos, 1989b
►● Tumidotheres maculatus (Say, 1818) in Say (1817-1818)
Figure 71. Tumidotheres maculatus (Say, 1818). Live color. Photography of a female within one its bivalve host Atrina seminuda (left), and dorsal views of the body of a female (down left) and the smaller male (up left). Photo by Emiliano Ocampo
Geographic distribution: NWAO, SWAO. SWAO: from Alagoas to São Paulo, Brazil, Uruguay and from Mar del Plata, Buenos Aires to Golfo San Matías, Rio Negro, Argentina, Bathymetric distribution: infralittoral (Fenucci, 1975; Melo, 1998). Larval development described from larvae hatched in the laboratory (Costlow and Bookhout, 1961).
Remarks: it was included in the genus Tumidotheres by Campos, 1989b. It was found in different hosts, mainly bivalves, but also in polychaete tubes; its biology, including the relationships with its hosts, has been studied in USA but also in Argentina (Supplementary Online File 6). The Barcode is now available online via the Barcode of Life Data System (BOLD; www.barcodinglife.org).
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DISCUSSION
A total of 211 decapod crustacean species have been found in coastal and oceanic waters of the Soutwestern Atlantic Ocean in front of Uruguay and Argentina (34ºS to 60ºS), and of the Southern Ocean (66ºW to 26ºW). The previous reviews of Argentine (Boschi et al., 1992) and Uruguayan (Zolessi and Philippi, 1995) decapods included 93 and 107 marine and estuarine species, respectively. Among the most represented taxa, shrimps (Dendrobranchiata and Caridea) showed the higher increase, from 23 (Argentina) and 26 (Uruguay) to 78 species. King crabs, hermit crabs and their allies (Anomura) increased less, from 28 (Argentina) and 18 (Uruguay) to a total of 40 species. The number of species of true crabs (Brachyura) twofold the Argentine previous record (38 to 80) and represent an almost 25% increment in species number respect to the Uruguayan list (60 to 80) (Table 2).
Three families of Dendrobranchiata (The Penaoidea Aristeidae and Benthesicymidae and the Sergestoidea Luciferidae), as well as 20 of Pleocyemata, that were not included in Boschi et al. (1992) are reported in this new catalogue (Table 2). The later suborder includes now several higher order taxa previously absent: 2 new infraorders, Gebiidea (family Upogebidae) and Polychelida (family Polychelidae), 4 families of Caridea (Oplophoridae, Nematocarcinidae, Thoridae and Palaemonidae), 2 of Brachyura Dromiacea (Homoloidea: Homolidae and Latreillidae) and 10 of Brachyura Eubrachyura Heterotremata (Aethridae, Calappidae, Cancridae, Ethusidae, Chasmocarcinidae, Oregonidae, Parthenopidae, Portunoidea Carcinidae, Portunoidea Macropipidae, Pseudorhombilidae, Mathildellidae and Xanthidae sensu stricto).
The additions to Boschi et al. (1992) list came from different sources: 1) Species recorded for Uruguayan coastal waters that were not found in Argentina: 11 shrimps (Dendrobranchiata, mainly Penaeidae), 1 Axiidea, 8 anomurans (squat lobsters, porcellanid and hermit crabs) and 30 Brachyura, mainly Majoidea, Portunidae and Panopeidae (Tables 2 and 3). These data agree with the hypothesis of the Rio de la Plata (34º–36º20’S; 55º–58º30’W) as a significant biogeographical barrier of the southwestern Atlantic coastal ecosystems (Briggs, 1974; Bisbal, 1995; Luppi et al., 2003 and references thereof). 2) Recent reports of several species of shrimps from the relatively understudied oceanic waters off Argentina and Uruguay (Table 3). Although older reports also contribute, such as Plesiopenaeus armatus, Pseudaristeus speciosus which have been described early, by Spence Bate (1881; 1888). All these shrimps belong to Dendrobranchiata (3 Aristeidae, 5 Benthesicymidae, 4 Penaeidae, and especially 13 Sergestidae) and Pleocyemata (3 Pasiphaeidae, 1 Acanthephyridae, 1 Oplophoridae, 2 Pandalidae, 1 Crangonidae (Table 3); 3) Several species of Caridean shrimps (2 Nematocarcinidae, 1 Campylonotidae and 4 Thoridae), 2 Nephropidae and 7 Lithodidae that have been recently found in Southern Ocean waters (Table 3). 4) Few alien species, a caridean shrimp and two crabs which have successfully colonized Argentine coastal waters after 1992. They are the global invaders Palaemon macrodactylus (Palaemonidae), Pyromaia tuberculata (Majoidea, Inachoididae) and Carcinus maenas (Portunoidea, Carcinidae). Hyas arenaeus invaded Antarctic waters in 1986, but the establishment of a successful population has not been demonstrated. 5) Five new species that have been described from the Southern tip of South America, the Southwestern Atlantic Ocean Islands and the Southern Ocean, one from Uruguay and Argentina, and one hermit crab from Uruguay (Table 4).
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Several species previously reported by Boschi et al. (1992) or Zolessi and Philippi (1995) have not been included in the list due to synonymy and/or having detected erroneous geographical localization or report: Penaeus brasiliensis, Munida subrugosa (synonymized with M. gregaria), Emerita analoga (erroneously reported for Tierra del Fuego, Argentina), Taliepus dentatus and T. marginatus (likely erroneous report of this SEPO species for Uruguay and Brazil), Leucosia planata (the type specimen is the only one described, but without detail) and Armases miersii (a clear erroneous report for Montevideo,Uruguay).
The following 7 species have been reported on the basis of few specimens: ● Pseudaristeus speciosus, only two specimens (male and female) collected off Buenos Aires, Argentina, ● Lissosabinea cf tridentata, two specimens from off Uruguay (one was an exuvia and the other was damaged), ● Lithodes ferox, only one record in the SWAO (36º29´S; 53º47´W); this species is reported from the SEAO, off western Africa between 22ºN and 28ºS, ● Paralomis longidactylus, only one specimen was caught near the mouth of the Rio de la Plata (35º34´S; 52º43´W), ● Pagurus trichocerus, was described on the basis of a single male specimen from the type locality, near Cabo Polonio, Uruguay, ● Acanthocarpus meridionalis, was described on the basis of a single male specimen from the type locality, Pozo de Fango, in Rocha, Uruguay (geographic collection place under discussion), ● Hyas araneus, only a male and a female of this NEAO species were collected in the South Shetland Island (61°05’93”S; 55°47’07”W, 92 m) in 1986.
The presence of the following 2 species has been hypothesized: ● Liopetrolisthes patagonicus, it is likely present at the eastern entrance of the Estrecho de Magallanes ● Pinnaxodes chilensis, it has not been found in Argentine waters of Tierra del Fuego (Canal Beagle) but the distribution of its host, the echinoid Loxechinus albus, suggested that it should live there
In addition, the reports of Pagurus provenzanoi and Panopeus austrobesus for Uruguayan waters are dubious and they need to be confirmed.
The following unresolved conflicts have been detected: ● Nematocarcinus tenuipes. Genetic differences suggest that the specific identity between SWAO and NWPO is doubtful. ● Lithodes confundens. It has been suggested that L. confundens could be a morphotype of L. santolla. ● Eurypodius latreillii and E. longirostris. Morphological and molecular evidence (yet unpublished) suggest that the species that inhabits the SWAO is E. longirostris. ● Cyrtograpsis affinis and C. altimanus. It has been suggested that both could represent a single species with a high ecological and phenotypic plasticity
Table 3. Species not registered by Boschi et al. (1992).
130 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Species Status Habitat Reported for Uruguay Argentina Dendrobranchiata Aristeidae Aristaeopsis edwardsiana Oceanic Meso-bathypelagic 37 No Plesiopenaeus armatus Oceanic Meso-bathypelagic No 26 Pseudaristeus speciosus Oceanic Abyssopelagic No 26 Benthesicymidae Benthesicymus brasiliensis Oceanic Meso-abyssopelagic No 25 Gennadas brevirostris Oceanic Meso-abyssopelagic 12 No Gennadas gilchristi Oceanic Meso-bathypelagic 17; 20 17; 20 Gennadas tinayrei Oceanic Epi-bathypelagic 17; 20 No Gennadas valens Oceanic Epi-bathypelagic 17; 20 17; 20 Penaeidae Funchalia villosa Oceanic Epi-mesopelagic 17; 20 17; 20 Funchalia woodwardi Oceanic Epi-mesopelagic 37 No P arapenaeus americanus Oceanic Epi-mesopelagic 37 20 Penaeopsis serrata Oceanic Mesopelagic 12 No Penaeus notialis Coastal Epipelagic 37 No Penaeus schmitti Coastal Epipelagic 37 No Xiphopenaeus kroyeri Coastal Epipelagic 37 No Solenoceridae Solenocera necopina Oceanic Epipelagic 20 No Luciferidae Belzebub faxoni Coastal Epipelagic 37 No Sergestidae Allosergestes pectinatus Oceanic Epi-mesopelagic 17; 20 17; 20 Allosergestes sargassi Oceanic Epi-mesopelagic 17; 20 17; 20 Deosergestes corniculum Oceanic Meso-bathypelagic 17; 20 17; 20 Deosergestes curvatus Oceanic Meso-bathypelagic No 17; 20 Deosergestes disjunctus Oceanic Meso-bathypelagic 17; 20 17; 20 Gardinosergia splendens Oceanic Epi-bathypelagic 17; 20 17; 20 Neosergestes edwarsii Oceanic Epi-bathypelagic 17; 20 17; 20 Parasergestes armatus Oceanic Epi-mesopelagic 17; 20 17; 20 Parasergestes vigilax Oceanic Epi-mesopelagic 17; 20 17; 20 Petalidium foliaceum Oceanic Meso-bathypelagic 17; 20 No Prehensilosergia prehensilis Oceanic Epi-mesopelagic 17; 20 17; 20 Sergestes atlanticus Oceanic Epi-mesopelagic 17; 20 17; 20 Sergia laminata Oceanic Mesopelagic 17; 20 17; 20 Pleocyemata Caridea Pasiphaeidae Pasiphaea barnardi Oceanic Meso-bathypelagic No 20 Pasiphaea dofleini Coastal Epi-mesopelagic No Not published Pasiphaea rathbunae Oceanic Epi-bathypelagic No 11; 20 Pasiphaea scotia Oceanic Meso-bathypelagic No 11; 20 Acanthephyridae Acanthephyra quadrispinosa Oceanic Meso-bathypelagic 17; 20 No Oplophoridae Systellaspis debilis Oceanic Meso-bathypelagic 17; 20 No Nematocarcinidae Nematocarcinus lanceopes Oceanic Bathyal No 32
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Nematocarcinus longirostris Oceanic Bathyal No 32 Nematocarcinus tenuipes Campylonotidae Campylonotus arntzianus n.s. Oceanic Bathyal No 30 Palaemonidae Palaemon macrodactylus * Coastal Intertidal, infralittoral No 28 Palaemon northropi Coastal Intertidal 37 No Alpheidae Alpheus pouang Coastal Circalittoral 10 No Hippolytidae Exhippolysmata Coastal Circalittoral 5 No oplophoroides Thoridae Eualus amandae Oceanic Bathyal No 21 Eualus dozei Circalittoral/bathyal No 36 Eualus kinzeri Oceanic Bathyal No 3; 15 Lebbeus antarcticus Oceanic Bathyal No 3; 21 Processidae Processa guyanae Coastal Circalittoral 10 No Processa profunda Coastal Circalittoral 9 No Pandalidae Pantomus parvulus Circalittoral/bathyal 8 No Stylopandalus richardi Oceanic Epi-bathypelagic 17 No Crangonidae Lissosabinea cf. tridentata Oceanic ------7 No Philocheras gorei Coastal Circalittoral 7 No Pleocyemata Astacidea Nephropidae Thymopsis nilenta Oceanic Bathyal No 2 Thymops takedai n.s. Oceanic Bathyal No 2 Pleocyemata Axiidea Callianassidae Poti gaucho Coastal Circalittoral 37 No Callianasidae Notiax santarita n.s. Circalittoral No 29 Pleocyemata Gebiidea Upogebiidae Upogebia australis n.s. Coastal Circalittoral No 31 Pleocyemata Polychelida Polychelidae Pentacheles validus Oceanic Bathyal 14 No Stereomastis sunhni Oceanic Bathyal No 13 Pleocyemata Anomura Galatheidae Munida iris Circalittoral/bathyal 37 No Munida flinti Circalittoral/bathyal 38 No Munida forceps Circalittoral/bathyal 38 NO Munida irrisa Circalittoral/bathyal 38 NO No Not Porcellanidae Liopetrolisthes patagonicus Coastal Intertidal/circalittoral published Polyonyx gibbesi Coastal Infralittoral 37 4 Porcellana sayana Coastal Intertidal 37 No Lithodidae Lithodes ferox Oceanic Bathyal No No Neolithodes diomedeae Oceanic Bathyal No No Neolithodes yaldwini n.s. Oceanic Bathyal No 1 Paralomis birsteini Oceanic Bathyal No No Paralomis longidactylus Oceanic Bathyal No No
132 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Paralomis tuberipes Oceanic ---- No No Paralomis zealandica Oceanic Bathyal No No Diogenidae Paguristes scarabinoi n.s Coastal Circalittoral 18 No Paguridae Pagurus leptonyx Coastal Circalittoral 37 No Pagurus trichocerus Coastal Infralittoral 37 No Pleocyemata Brachyura Homolidae Homola barbata Coastal Circalittoral 37 No Latreilliidae Latreilla willamsi Coastal Circalittoral 37 No Aethridae Hepatus pudibundus Coastal Intertidal/circalittoral 37 No Calappidae Acanthocarpus alexandri Coastal Circalittoral/bathyal 37 No Acanthocarpus meridionalis Coastal Circalittoral 19 No Cancridae Metacarcinus edwardsii Coastal Infra/circalittoral 35 No Ethusidae Ethusina abyssicola Oceanic Bathyal No 21 Chasmocarcinidae Chasmocarcinus typicus Coastal Circalittoral 34 No Mathildillidae Neopilumnoplax lipkeholthuisi Coastal Circalittoral/bathyal No 39 Leucosiidae Myropsis quinquespinosa Coastal Circalittoral/bathyal 37 No Persephona mediterranea Coastal Intertidal/circalittoral 37 No Epialtidae Libinia ferreira Coastal Intertidal/circalittoral 37 No Stenocionops furcatus Coastal Intertidal/circalittoral 37 No Inachidae Anisonotus atlanticus Coastal Circalittoral 37 No Not Eurypodius longirostris Oceanic Bathyal No published Inachoididae Pyromaia tuberculata * Coastal Infra/circalittoral No 24 Stenorhynchus seticornis Coastal Intertidal/bathyal 37 No Mithracidae Amphithrax besnardi Oregonidae Hyas areaneus * Coastal Circalittoral No No Not Parthenopidae Leiolambrus nitidus Coastal Circalittoral No published Mesorhoea sexspinosa Coastal Intertidal/circalittoral 37 No Piloslambrus guerinii Coastal Circalittoral 37 No Spinolambrus fraterculus Coastal Infra/circalittoral 37 No Spinolambrus meridionalis Coastal Circalittoral 37 No Carcinidae Carcinus maenas * Coastal Intertidal/circalittoral No 16 Portunidae Arenaeus cribarius Coastal Intertidal/circalittoral 37 23 Callineces danae Coastal Intertidal/circalittoral 37 No Achelous spinicarpus Coastal Intertidal/circalittoral 37 No Achelous spinimanus Coastal Circalittoral 37 No Panopeidae Cyrtoplax spinidentata Coastal Infra/circalittoral 37 No Eurypanopeus depressus Coastal Intertidal 37 No Hexapanopeus paulensis Coastal Intertidal/infralittoral 37 No Tetraxanthus rathbunae Coastal Circalittoral 37 No Actaea acantha Coastal Intertidal/infralittoral 37 No Grapsidae Pachygrapsus transversus Coastal Intertidal 37 No Pseudorhombilidae Speocarcinus dentatus Coastal Circalittoral 5 No
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Planes major Oceanic Epipelagic 37 No Planes minutus Oceanic Epipelagic No 26 Ocypodidae Ocypode quadrata Coastal Supralittoral 37 No Pinnotheridae Pinnixa chaetopterana Coastal Intertidal/infralittoral 37 No Pinnixa sayana Coastal Infra/circalittoral 37 No Pinnixa valdiviensis Coastal Intertidal/infralittoral No 33
References: * = invader, n.s. = new species; 1) Ahyongand Dawson, 2006; 2) Ahyong, Webber and Chan, 2012; 3) Arntz and Gorny, 1991; 4) Belleggia et al., 2010;5) Brandão, Coelho-Filho & Tavares, 2012; 6) Christoffersen, 1979; 7) Christoffersen, 1988; 8) Christoffersen, 1989; 9) Christoffersen, 1982b; 10) Christoffersen, 1998; 11) Clarke and Holmes, 1986; 12) D’Incao, 1998, 1999; 13) Farías et al., 2015; 14) Farías and Scarabino, in press; 15) Gorny et al., 1992; 16) Hidalgo et al., 2005; 17) Judkins, 2014; 18) Lima and Santana, 2017; 19) Mañé-Garzón, 1979; 20) USNMdb; 21) Nye et al., 2013; 22) Ocampo et al., 2014; 23) Scelzo, 2001; 24) Schejter et al., 2002; 25) Spence Bate, 1881; 26) Spence Bate, 1888; 27) Spivak and Bas, 1999; 28) Spivak et al., 2006; 29) Thatje, 2000; 30) Thatje, 2003; 31) Thatje and Gerdes, 2000; 32) Thatje et al., 2005; 33) Torres, 2006; 34) Vezzani, 2007; 35) Vinuesa et al., 1999; 36) Wehrtmann and Carvacho, 1997; 37) Zolessi and Philippi, 1995; 38) Melo Filho and Melo, 2001 ; 39) Tavares and Melo, 2010.
Table 4. New species of Pleocyemata Decapod Crustacea described after Boschi et al. (1992).
Infraorder Family Species Sites Campylonotus The Sandwich del Sur Caridea Campylonotidae Thatje, 2003 arntzianus Islands, Scotia Sea The Georgias del Sur Ahyong, Webber and Astacidea Nephropidae Thymops takedai Islands, Scotia Sea Chan, 2012 Axiidea Callianasidae Notiax santarita Canal Beagle Thatje, 2000 Gebiidea Upogebiidae Upogebia australis Canal Beagle Thatje an dGerdes, 2000 Ahyong and Dawson, Anomura Lithodidae Neolithodes yaldwini Antarctic 2006 Diogenidae Paguristes scarabinoi Uruguay Lima and Santana, 2017 Neopilumnoplax North Argentina, Brachyura Mathildellidae Tavares and Melo, 2010 lipkeholthuisi South Brasil
Only 15 species have been renamed between 1992 and the present (Table 5). Ten have been assigned to the newly established genera Belzebub, Notiax, Sergio, Propagurus, Danielethus, Leptuca, Neohelice, Austinixa and Calyptraeotheres and 5 species have been synonymyzed (one of them assigned to the new genus Acantholobulus). The genus Penaeus has been grouped into the subgenera Litopenaeus, Farfantepenaeus, Fenneropenaeus, Marsupenaeus, Melicertus and Penaeus sensu stricto by Pérez Farfante and Kensley (1997) though some authors have refused this change and proposed the restoration of the old Penaeus genus (Ma et al., 2011).
Table 5.Changes in nomenclature and taxonomy after Boschi et al. (1992).
134 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Old name New name Dendrobranchiata Luciferidae Lucifer faxoni Belzebub faxoni Pleocyemata Axiidea Callianassa brachyophthalma Notiax brachyophthalma Callianassidae Callianasa mirim/Callichirus mirim Sergio mirim Pleocyemata Anomura Galatheidae Pachycheles haigae Pachycheles laevidactylus Propaguridae Pagurus gaudichaudi Propagurus gaudichaudii Pleocyemata Brachyura Podotremata Latreillidae Latreilla elegans Latreilla willamsi Homola barbata Homola minima Pleocyemata Brachyura Eubrachyura Platyxanthus crenulatus Danielethus crenulatus Platyxanthidae Platyxanthus patagonicus Danielethus patagonicus Inachidae Podochela algicola Anisonotus atlanticus Panopeidae Panopeus margentus Acantholobulus schmitti Varunidae Chasmagnathus granulata/granulatus Neohelice granulata Ocypodidae Uca uruguayensis Leptuca uruguayensis Pinnixa patagoniensis Austinixa patagoniensis Pinnotehridae Pinnotheres garthi Calyptraeotheres garthi
Table 6. Changes in higher level systematics and nomenclature of Pleocyemata included in this catalogue:
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De Grave et al., 2009; 2014 Boschi 1992 Infraorder Superfamily Family Family Superfamily Infraorder Pasiphaeoidea Pasiphaeidae No change Atyoidea Caridea Oplophoroidea Acanthephyridae Oplophoridae Oplophoridae Campylonotoidea Campylonotidae No change Palaemonoidea Caridea Palaemonoidea Palaemonidae New Alpheidae No change Alpheoidea Alpheoidea Hippolytidae No change Thoridae No change Processoidea Processidae No change Axiidea ------Callianassidae No change Thalassinoidea Thalassinidea Achelata ------Scyllaridae No change Palinuroidea Palinura Galatheoidea Galatheidae No change Galatheoidea Anomura Porcellanidae No change Hippoidea Blepharipodidae Albuneidae Hippidae No change Anomura Lithodoidea Lithodidae No change Paguroidea Paguroidea Diogenidae No change Coenobitoidea Paguridae No change Paguroidea Parapaguridae No change Paguroidea Epialtidae Majidae Majoidea Brachyura Inachidae Majoidea Inachoididae Pseudorhombilidae Oregonidae Brachyura Eriphioidea Platyxanthidae No change Xanthoidea Eubrachyura Pilumnoidea Galenidae Xanthidae Hetero-tremata Pseudozioidea Pilumnoididae Xanthoidea Panopeidae Brachyura Eubrachyura Grapsoidea Grapsidae Grapsidae Grapsoidea Thoraco-tremata
Regarding higher taxonomic categories, the catalogue of Boschi et al. (1992) was organized on the basis of a classification of recent Crustacea published by Bowman and Abele (1982) and changes in classification and nomenclature have been accumulated since then and reviewed by De Grave et al. (2009). On one hand, the following superfamilies listed by De Grave et al. (2009) have not been mentioned by Boschi et al. (1992): Pasiphaeoidea, Oplophoroidea, Campylonotoidea, Processoidea, Hippoidea, Lithodoidea, Eriphioidea, Pilumnoidea and Pseudozioidea; this change is related with modifications in other superfamilies: Atyoidea, Palaemonoidea, Alpheoidea, Galatheoidea,
136 Annotated catalogue and bibliography of marine and estuarine shrimps, lobsters, crabs and their allies (Crustacea: Decapoda) of Argentina and Uruguay (Southwestern Atlantic Ocean)
Coenobitoidea and Paguroidea (Table 6). On the other hand, the infraorders Thalassinidae and Palinura have been renamed as Axiidea and Achelata, respectively. Other changes occurred at the familial level: Acanthephyridae has been extracted from Oplophoridae, Thoridae from Hippolytidae (De Grave et al., 2014), Blepharipodidae from Albuneidae and Majidae, Xanthidae and Grapsidae have been subdivided in several families (Table 6).
Summarizing, the present knowledge of the decapods crustacean fauna of the Southwestern Atlantic Ocean in front of Uruguay and Argentina is not uniform. Coastal species form a well-known assemblage, and the most important addition to it are alien species. The information on oceanic species, however, has increased only in the northern areas, especially off Rio de la Plata, and in the southernmost areas, including Southern Ocean. Consequently, additional samplings are necessary to improve the understanding of regional diversity.
SUPPLEMENTARY ONLINE FILES
These files are posted in the web page of the CTMFM: www.ctmfm.org
Supplementary Online File 1. References, Taxonomy Format Supplementary Online File 2. References, Dendrobranchiata Supplementary Online File 3. References, Pleocyemata Caridea Supplementary Online File 4. References, Pleocyemata, Axiidea, Achelata, Anomura Supplementary Online File 5. References, Pleocyemata Brachyura, Heterotremata Supplementary Online File 6. References, Pleocyemata Brachyura, Thoracotremata Supplementary Online File 7. References, Pleocyemata Brachyura, Neohelice granulata
ACKNOWLEDGEMENTS
The authors thank the Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET) PIP 830/12, Agencia Nacional de Promoción Científica y Tecnológica (ANPCyT) PICT 763/13 and Universidad Nacional de Mar del Plata (UNMdP) EXA 805/16, for the continuous support. Part of the work related to DNA barcoding was financed by IBOL PIP 11220110100830 3404/14. We are grateful for the support provided by the Government of Canada through Genome Canada and the Ontario Genomics Institute to the International Barcode of Life Project. This funding enabled the Canadian Centre for DNA Barcoding (University of Guelph) to carry out the sequence analysis on our specimens. We want to aknowledge Marcos Tavares and Fabrizio Scarabino for the enormous job done in the review of this paper and for their helpful suggestions and comments. Lastly, we would like to thank Magdalena Perez, Secretaría Técnica Comisión Técnica Mixta del Frente Marítimo, for her support and constant help throughout the process of preparing this work.
This paper is dedicated to Prof. Dr. Enrique Eduardo Boschi on the occasion of his 90th birthday.
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