Print
Revision of the deep-water spider crab genus, Scyramathia A. Milne-Edwards, 1880, with the description of a new species from the Mediterranean and notes on Rochinia A. Milne-Edwards, 1875, and Anamathia Smith, 1885 (Crustacea, Decapoda, Brachyura, Epialtidae)
expand article infoBee Yan Lee, Bertrand Richer De Forges§, Peter K. L. Ng
‡ National University of Singapore, Singapore, Singapore
§ Muséum National d’Histoire Naturelle, Paris, France
Open Access

Abstract

The taxonomy of the deep-water spider crabs of the genus Scyramathia A. Milne-Edwards, 1880, is revised and four extant species are recognised from the Atlantic and western Indian Ocean: S. carpenteri (Norman, in Thomson 1873) (type species), S. umbonata (Stimpson, 1871), S. hertwigi Doflein, in Chun 1900, and S. tenuipes sp. nov. Scyramathia tenuipes sp. nov. from the Mediterreanean is easily distinguished from its congeners by its slender and elongate ambulatory legs. All species are diagnosed and figured. The taxonomy of two allied genera from the Atlantic and Mediterranean, Rochinia A. Milne-Edwards, 1875, and Anamathia Smith, 1885, are also treated and their type species redescribed and figured.

Key Words

deep-sea, Epialtidae, new species, revision, spider crabs, taxonomy

Introduction

The deep-water epialtid spider crab genus, Scyramathia A. Milne-Edwards, 1880, was recently reinstated by Tavares and Santana (2018) who recognised two extant valid species, S. carpenteri (Norman, in Thomson 1873), and S. umbonata (Stimpson, 1871), as well as one fossil taxon, S. boschii (Casadío, Feldmann, Parras & Schweitzer, 2005). Scyramathia had previously been in the synonymy of Rochinia A. Milne-Edwards, 1875, for many years (see Griffin and Tranter 1986, Ng et al. 2008). When A. Milne-Edwards (1880a) described Scyramathia, he did not provide the distinguishing morphological characters despite placing two species in it and this has caused problems in defining it. Although Tavares and Santana (2018) recognised Scyramathia, they focused more on the differences between it and Rochinia s. str.

With a good series of Scyramathia specimens from various parts of Europe and Africa, including types, the genus is herein revised and diagnosed. Four species are here recognised, including a new species from the Mediterranean. These species are diagnosed and figured. To provide context and necessary comparisons, the genera Rochinia A. Milne-Edwards, 1875, and Anamathia Smith, 1885, are also diagnosed, and their type species figured.

Material and methods

The material examined are deposited in the Muséum national d’Histoire naturelle (MNHN), Paris, France; The Natural History Museum (NHM), London, United Kingdom; U.S. National Museum of Natural History (USNM), Smithsonian Institution, Washington D.C., U.S.A; Naturalis Biodiversity Centre (ex. Rijksmuseum van Natuurlijke Historie, RMNH), Leiden, Netherlands; The Australian Museum (AM), Sydney, Australia; Museum of Comparative Zoology (MCZ), Harvard University, U.S.A.; Biological Reference Collection, Institute of Marine Sciences (ICMD), Barcelona, Spain; National Museum of Science and Technology (NSMT), Tokyo, Japan; Senckenberg Museum Frankfurt (SMF), Frankfurt, Germany; Museum für Naturkunde (ZMB), Berlin, Germany; South African Museum (Iziko Museums of South Africa) (SAM), Cape Town, South Africa; National Oceanography Centre, Southampton, U.K.; and Zoological Reference Collection (ZRC), Lee Kong Chian Natural History Museum, National University of Singapore, Singapore.

Measurements provided, in millimetres, are of the carapace length (excluding pseudorostral spine) and carapace width (to the base of the lateral spine). The maximum length and width of the merus of the ambulatory legs are measured following Maenosono and Naruse (2016). The abbreviations used are as follow: G1 and G2 = male first and second gonopods, respectively; P2 and P5 = second and fifth ambulatory legs, respectively; and stn = station.

With regard to the taxa named by Sir Charles Wyville Thomson, his family name has been cited as “Thomson” (see examples: Conan et al. 1981; MacPherson 1983; De Mol et al. 2002; Pawson and Pawson 2013), “Wyville Thomson” (see examples: D’Udekem d’Acoz 1999; Ng and Richer de Forges 2007; Ng et al. 2008) or “Wyville-Thomson” (see examples: Alvarado et al. 2008; O’Hara et al. 2018; Gan and Li 2019). His family name is Thomson and his given name is Charles Wyville. “Wyville” came from his maternal grandfather, Dr Wyville Smith, who had been a hospital inspector. He clearly preferred using Wyville to Charles as his first name, and tended to sign his name “C. Wyville Thomson”, even when writing to his son. Thomson was knighted and was addressed frequently as Sir Wyville. In the United Kingdom, when a person is knighted, the title “Sir” is conferred and usually used together with the given name and not the family name; and that was why in the narrative for the Challenger Expedition reports, John Murray referred to him as “Sir Wyville”. The author’s name is here written as “Thomson”.

Systematic account

Superfamily Majoidea Samouelle, 1819

Family Epialtidae MacLeay, 1838

Rochinia A. Milne-Edwards, 1875

Rochinia A. Milne-Edwards 1875 [in 1873–1880]: 86 (footnote); Stebbing 1910: 289 (list); Rathbun 1925: 204, 210; Garth 1958: 282, 283; Serène and Lohavanijaya 1973: 54; Ingle 1980: 140; Williams 1984: 315 (key), 322; Griffin and Tranter 1986: 174; Tavares 1991: 161, 162; Richer de Forges 1995: 45; De Melo 1996: 251 (key); D’Udekem d’Acoz 1999: 194; Davie 2002: 329; Ng and Richer de Forges 2007: 61–63; Ng et al. 2008: 105; Richer de Forges and Poore 2008: 64; Richer de Forges and Ng 2009: 2; Ng and Richer de Forges 2013: 362, 363; Richer de Forges and Ng 2013: 468; Pettan 2013: 10.

Rachinia [sic]: Alcock 1895: 165 (list); A. Milne-Edwards and Bouvier 1900: 132 (list); Stebbing 1910: 289 (list).

Type species

Rochinia gracilipes A. Milne-Edwards, 1875, by monotypy; gender feminine. Monotypic.

Remarks

The study by Tavares and Santana (2018: 222) suggests that Rochinia sensu stricto “should be restricted to its type species, R. gracilipes A. Milne-Edwards, 1875 with perhaps a few additional closely related species”. To this effect, Tavares and Santana (2018: 223) left the rest of the other species, mainly from the Indo-West Pacific region, in Rochinia sensu lato. Ng and Richer de Forges (2013) had discussed the problems with Rochinia at length and suggested that more genera may need to be recognised. The recent work on the epialtid spider crabs from Papua New Guinea by Lee et al. (2019) described two species from Rochinia sensu lato and transferred four Rochinia sensu lato species to Crocydocinus Lee, Richer de Forges & Ng, 2019. Lee et al. (2019) noted that a revision for Rochinia is in preparation (see also Lee et al. 2017). In this study, one species, S. hertwigi Doflein, in Chun 1900, is transferred out of Rochinia sensu lato and back to Scyramathia A. Milne-Edwards, 1880.

Rochinia gracilipes A. Milne-Edwards, 1875

Figs 1A–C, 2A–F

Rochinia gracilipes A. Milne-Edwards 1875 [in 1873–1880]: 86 (footnote) (type locality: Cape Corrientes, Argentina); Griffin and Tranter 1986: 174 (list); Rathbun 1925: 210 (key), 218, pl. 229 figs 1–4; Monod 1956: 32 (list), 516–517 (in part); Garth 1958: 282 (list), 283 (list); Manning and Holthuis 1981: 253 (list), 254, 255; Tavares 1991: 160, 161 (list), 164, figs 3A, 5B; De Melo 1996: 266 (key), 268, unnumbered in-text fig.; Tavares and De Melo 2004: 130; Ng and Richer de Forges 2007: 61, 63 (list); Ng et al. 2008: 105 (list); Ng and Richer de Forges 2013: 362, fig. 5A; Pettan 2013: 2 (list), 3 (list), 5 (list), 6 (list), 10 (list), 11 (list), 12, 14, 16, 20, 26, 41, 47, 86 (key), figs 10a, b, 16d, 23a, b; Griffiths et al. 2014: 185, map 8; Pettan and Tavares 2014: 306 (list); Ceccon and De Angeli 2018: 151 (list); Tavares and Santana 2018: 201, 204, 223 (list), figs 2, 9D, 10A, B, 11A, 12A, 13I–J; Spivak et al. 2019: 95, fig. 51.

Rachinia [sic] gracilipes: A. Milne-Edwards 1875 [in 1873–1880]: pl. 18 fig. 1–1d; Lagerberg 1905: 22–24.

Materials examined

Paralectotypes : 1 ♂ [dry] (20.0 × 15.0 mm, 1 ♀ (10.4 × 8.5 mm) (MNHN-IU-2000-4460 [= MNHN-B4460]), Cap. Corrientes, coll. A Milne-Edwards. Other material: 3 ovigerous ♀♀ (15.1 × 11.1 mm, 14.9 × 10.8 mm, 14.5 × 10.1 mm), 3 ♀♀ (15.2 × 11.1 mm, 14.1 × 9.7 mm, 12.5 × 9.2 mm) (ZRC 2019.1634), Mar del Plata Habour, Argentina, 38°02'S, 57°31'30"W, coll. N Farias, 9 October 2014; 1 ovigerous ♀ (16.0 × 11.5 mm) (USNM 1277591), Mar del Plata, Buenos Aires Province, Argentina, South Atlantic Ocean, coll. M Doella-Jurado, 7 August 1911; 2 ♂ (15.2 × 10.6 mm, 11.9 × 9.8 mm) (USNM 1150573), Mar del Plata, Buenos Aires, Argentina, South Atlantic Ocean, coll. Franceschi and Lelois, February 1924; 1 ♂ (17.2 × 12.4 mm) (USNM 1150574), Mar del Plata, Buenos Aires, Argentina, South Atlantic Ocean, coll. M Doella-Jurado, 1914; 1 ♂ (10.5 × 7.1 mm), 1 ovigerous ♀ (14.0 × 10.1 mm) (USNM 55118), 40°22'S, 60°35'W, 55 m, coll. Hassler Expedition, 1951.

Figure 1. 

Rochinia gracilipes A. Milne-Edwards, 1875, paralectotype male [dry] (20.0 × 15.0 mm) (MNHN-IU-2000-4460 [= MNHN-B4460]), Cap. Corrientes. A. overall dorsal view; B. overall ventral view; C. lateral view of carapace.

Remarks

No type specimen was designated when this species was described by A. Milne-Edwards (1875: 86 [footnote]) although various specimens from “la collection du Muséum” (i.e., the Muséum national d’Histoire naturelle (MNHN), Paris) as well as specimens from Hassler expedition were mentioned. There is also a figure of a male specimen provided in the same paper, but the name was incorrectly spelled as “Rachinia gracilipes” in the caption (see A. Milne-Edwards 1875: pl. 18 fig. 1). The male specimen that is found in the MNHN (MNHN-IU-2000-4460 [= MNHN-B4460]) matches the figure provided by A. Milne-Edwards (1875: pl. 18 fig. 1–1d). The same male specimen was used by Tavares (1991: figs 3A, 5B) and was considered a syntype by the author. This was, however, not listed by Tavares and Santana (2018) in their study to redefine Rochinia, when they designated another specimen as the lectotype for this species. It was mentioned by Tavares and Santana (2018) that the type material needs clarification and that the male specimen that was illustrated by A. Milne-Edwards (1875: pl. 18, fig. 1–1d) and deposited in the MNHN was clearly not the holotype as had been identified by Rathbun (1925). Rathbun’s (1925) recognition of the “holotype” is incorrect as no type was selected in the original description by A. Milne-Edwards (1875) and all the material must be considered syntypes. Tavares and Santana (2018) designated one male specimen (MCZ 1950) as the lectotype for this species and five other female specimens as the paralectotypes, all collected from the Hassler expedition. Based on the original description by A. Milne-Edwards (1875: 86) and the figure (A. Milne-Edwards 1875: pl. 18 fig. 1–1d), the specimens from MNHN are also part of the type series. The G1 and G2 of the paralectotype male (MNHN-IU-2000-4460 [= MNHN-B4460]) is illustrated here (Fig. 2A–F). The type locality of this species is Cape Corrientes (= Cabo Corrientes), Argentina, and specimens are found in relatively shallow waters (about 55 m depth) (see also Tavares & Santana, 2018: 221).

Rochinia gracilipes was recorded from the coast of Africa near Gabon by Monod (1956). The male specimen figured (Monod 1956: figs 706–708) was compared with the MNHN type specimen and it does resemble the present concept of R. gracilipes. Monod (1956: 517), however, was not certain of the origin of the specimen and noted that its presence in African waters needs to be confirmed. Monod (1956)’s dubious African record is not included in the synonymy for this species for the time being.

Distribution

Currently only known from its type locality, Cape Corrientes (= Cabo Corrientes), Mar del Plata, Argentina (A. Milne-Edwards 1875), southeastern Brazilian coast (Spivak et al. 2019), South Shetland Islands (Griffiths et al. 2014); Antarctic Peninsula (De Melo, 1996; Tavares and Melo 2004).

Figure 2. 

A–D. Rochinia gracilipes A. Milne-Edwards, 1875, paralectotype male (20.0 × 15.0 mm) (MNHN-IU-2000-4460 [= MNHN=B4460]), Cap. Corrientes, left G1; A. ventral view; B. ventral view of distal portion; C. dorsal view; D. dorsal view of distal portion; E–F. R. gracilipes A. Milne-Edwards, 1875, male (15.2 × 10.6 mm) (USNM 1150573), Argentina, left G2; E. ventral view of G2; F. dorsal view of G2. Scale bars: 1 mm.

Scyramathia A. Milne-Edwards, 1880

Scyramathia A. Milne-Edwards 1880a: 356; A. Milne-Edwards 1880b: 277; Sars 1885: 5, 6; Smith 1886: 625[21]; A. Milne-Edwards and Bouvier 1894: 12, 13; Alcock 1895: 201; A. Milne-Edwards and Bouvier 1900: 131; Rathbun 1901: 61; Stebbing 1902: 5; Doflein 1904: 80, 81; Stebbing 1905: 25; Stebbing 1910: 289; A. Milne-Edwards and Bouvier 1923: 379; Barnard 1950: 48 (key), 49; Tavares 1991: 161, 162; Tavares and Santana 2018: 204, 208.

? Scyramathia: Alcock 1895: 165 (list).

Type species

Amathia carpenteri Norman, in Thomson 1873, subsequent designation by Rathbun 1925; gender feminine.

Diagnosis

Carapace pyriform, with strong raised plate-like structures in large adult specimens (Figs 3, 4A, 5A, 7A, C, 9A–I, 10A, 11A, 12, 13A); weak plates on smaller specimens (see Tavares et al. 2015: Fig. 1A–C). Pseudorostral spines long, slender, diverging at approximately 45° angle or less. Supraorbital eave truncated with sharp preorbital angle; strong postorbital lobe fused with hepatic spine, round anterior margin (Figs 3, 4A, 5A, 7A, C, 9A–I, 10A, 11A, 12, 13A). Mesogastric and cardiac regions with plates; long lateral branchial spine pointing outwards, more plate-like in large specimens (usually exceeding 20 mm carapace width) (Figs 3, 4A, 5A, 7A, 9A–I, 10A, 11A, 12, 13A). Antennal flagellum short, about half pseudorostral spine length. Basal antennal article longer than broad, with straight to gently convex outer margin. Distal angle of buccal frame blunt, slightly raised. Pterygostomial region with large, flattened granules on outer margin (Figs 4B, 5B, 6C, 7B, D, 10B, 11B, E, 13B). Chelipeds slender with slight carinate margins; propodus slender, longer than fingers; carpus with carinate margins; merus triangular in cross-section with carinate margins. Ambulatory legs long with cylindrical articles, rounded margins; merus with blunt distal angle; P2 longest (Figs 3, 4A, 5A, 10A, 11A, 12, 13A). Male thoracic sternum transversely broad, deeply groove; sternites 3, 4 relatively wide with slightly constricted lateral margin. Male pleon rectangular, telson triangular; surface of somites smooth (Figs 4B, 5B, 6D, F, 10B, 11B, E, 13B). G1 straight, distal part elongated, with sharp angle at tip (Figs 8C–F, 15A–H); G2 with wide distal tip forming C-shape lateral margin (Fig. 8G, H).

Figure 3. 

Illustration of Scyramathia carpenteri (Norman, in Thomson 1873), “sandy chalk of Holtenia ground”, overall dorsal view. (After Norman, in Thomson 1873: fig. 35).

Figure 4. 

Scyramathia carpenteri (Norman, in Thomson 1873), lectotype male (21.6 × 15.9 mm) (NHM 1907.8.28.3), “Holtenia ground” (= between north Scotland and the Faeroe Islands). A. overall dorsal view; B. overall ventral view; C. lateral view of carapace.

Figure 5. 

Scyramathia carpenteri (Norman, in Thomson 1873), male (41.3 × 34.0 mm) (NHM 1983.449), North Sea. A. overall dorsal view; B. overall ventral view; C. lateral view of carapace.

Remarks

When establishing Scyramathia A. Milne-Edwards, 1880, A. Milne-Edwards (1880a: 356) transferred Scyra umbonata Stimpson, 1871, and Amathia carpenteri Norman, in Thomson 1873, to it, commenting that both species do not belong to the genus they were described in: “Je ferai remarquer que l’Amathia Carpenteri n’appartient pas au genre Amathia et que la prétendue Scyra umbonata n’est certainement pas une Scyra” [I will point out that Amathia carpenteri does not belong to the Amathia genus and that alleged Scyra umbonata is certainly not Scyra]. He did not elaborate on the reasons why the two genera were different, nor did he compare it to Rochinia A. Milne-Edwards, 1875, which he described only seven years earlier. There were no distinguishing or diagnostic morphological characters provided for Scyramathia A. Milne-Edwards, 1880. No type species for the genus was proposed, Rathbun (1925) subsequently selected Amathia carpenteri Norman, in Thomson 1873, as the type species for Scyramathia.

Scyramathia carpenteri (Norman, in Thomson 1873) was described in Amathia Roux, 1828, but this genus name is preoccupied by Amathia Lamouroux, 1812 (Bryozoa), and as such, a new name Anamathia, was proposed by Smith (1885). Alcock (1895: 201) re-described Scyramathia as he understood it and remarked that it is distinct from Anamathia, with more affinities to Hyastenus White, 1847, and Pugettia Dana, 1851. Alcock (1895), however, had based his understanding of Scyramathia on the Indo-West Pacific species. In selecting S. carpenteri (Norman, in Thomson 1873) as the type species of Scyramathia, Rathbun (1925) also synonymised it under Rochinia, but without any explanation. Although Garth (1958: 283) listed Scyramathia A. Milne-Edwards, 1880, and Anamathia Smith, 1885, in the synonymy of Rochinia, he commented that the genus is not monophyletic due to the varying forms of the G1. Scyramathia has nevertheless remained under the synonymy of Rochinia since. Serène and Lohavanijaya (1973: 55) discussed the history of Rochinia, and noted that some authors (e.g. Barnard 1950) still considered Scyramathia a valid genus.

The species in Scyramathia s. str. have characteristic carapace plates that become more pronounced and lamelliform with age (see Tavares et al. 2016; Tavares and Santana 2018). Our studies confirm the findings of Tavares and Santana (2018). Tavares and Santana (2018) noted that Scyramathia includes the following species: S. carpenteri Norman, in Thompson, 1873, S. umbonata Stimpson, 1871, and a fossil species, S. boschii (Casadío, Feldmann, Parras & Schweitzer, 2005) (see Casadío et al. 2005). Tavares and Santana (2018), however, retained one species, Scyramathia hertwigi Doflein, in Chun 1900, from Africa in Rochinia sensu lato. It is clearly a Scyramathia species (present study).

Scyramathia s. str. is herein re-defined. With the transfer of S. hertwigi, as well as the description of a new species, S. tenuipes sp. nov., from the Mediterranean Sea, there are now four extant species and one fossil species in this genus.

Key to the extant species of Scyramathia A. Milne-Edwards, 1880 (based on adult male specimens)

1 Pseudorostral spines long, slightly longer than or equal to half carapace length; basal antennal article with straight to gently convex outer margin 2
Pseudorostral spines short, shorter than half carapace length; basal antennal article with convex outer margin 3
2 With weak or relatively distinct plates on carapace; lateral branchial spine sharp, pointing laterally, slightly upwards; pterygostomial region with plate-like granules on outer margins; P2 merus length 11.8–14.0 times width; P5 merus length 5.5–7.3 times width Scyramathia carpenteri (Norman, in Thomson 1873)
With weak plates on carapace; lateral branchial spines sharp, pointing outwards; pterygostomial region with distinct granules on outer margins; P2 merus length 19.5–20.5 times width; P5 merus length 8.0–9.6 times width………….. Scyramathia tenuipes sp. nov.
3 With distinct, slightly raised plates on carapace; fused L-shaped postorbital and hepatic plates directed upwards; lateral branchial plate slightly curved, pointed upwards in large males Scyramathia hertwigi Doflein, in Chun 1900
With distinct, raised leaf-like plates on carapace; fused L-shaped postorbital and hepatic plates curved upwards; lateral branchial plate curved upwards in large males Scyramathia umbonata (Stimpson, 1871)

Scyramathia carpenteri (Norman, in Thomson, 1873)

Figs 3, 4A–C, 5A–C, 6A–F, 7A–D, 8A–H, 9A–I, 14A, B

Amathia carpenteri Norman, in Thomson 1873: 175 (fig.), 176, fig. 35 (type locality: “sandy chalk of the Holtenia ground” [= between North Scotland and the Faeroe Islands]).

Scyramathia Carpenteri: A. Milne-Edwards 1880a: 356 [new combination]; A. Milne-Edwards 1880b: 277; Filhol 1885: 123, 154, fig. 38; Sars 1885: 6–11, 274, pl. 1 figs 1–7; Perrier 1886: 208, fig. 217; Smith 1886: 625[21], 626[22]; Bourne 1890: 308; A. Milne-Edwards and Bouvier 1894: 13, 14 (in part); A. Milne-Edwards and Bouvier 1899: 43 (in part); Doflein 1904: 81 (list); A. Milne-Edwards and Bouvier 1900: 133–138, pl. 20 fig. 1–10; Hansen 1908: 12; Bouvier 1922: 81 (in part); Grieg 1927: 44–45.

Anamathia Carpenteri: Smith 1886: 626[22]; Pocock 1889: 425, 426; Bourne 1890: 314, 315.

Anamathia carpenteri: Faxon 1895: 10.

Rochinia carpenteri: Rathbun 1925: 204 (list) [new combination]; Miranda y Rivera 1933: 41; Sivertsen and Holthuis 1956: 49; Allen 1967: 35 (list), 64 (key), 97 (fig.); Sankarankutty 1968: 51, figs 5D–F; Christiansen 1969: 20, 122–124, 123 (map), fig. 50; Ingle 1979: 47–53, 59, figs 1–7; Ingle 1980: 15 (list), 19 (table), 20 (table), 140, fig. 108, pl. 33 fig. b; Manning and Holthuis 1981: 254; Shelton and Dooley 1982: 109; Clark 1986: 190, 191 (map); Griffin and Tranter 1986: 180 (list); Rice 1990: 10, fig. 7; Tavares 1991: 161 (list), 172; d’Udekem d’Acoz 1999: 194 (in part); Casadío et al. 2005: 158 (list); Ng and Richer de Forges 2007: 63 (list); Ng et al. 2008: 105 (list); Ng and Richer de Forges 2013: 362, fig. 5B.

Rochinia Carpenteri: Bouvier 1940: 344, 345 (in part).

Scyramathia carpenteri: Tavares and Santana 2018: 204 (in part).

Material examined

Lectotype : ♂ (21.6 × 15.9 mm) (NHM 1907.8.28.3), stn 47, ‘Holtenia ground’, 59°34'N, 7°18'W, 958 m, coll. H.M.S. “Porcupine”, 1869. Paralectotypes: 1 ♀ (13.0 × 8.4 mm) (NHM 1911.11.8.377), same locality and collection data as lectotype; 1 damaged juvenile ♀ (5.4 × 3.0 mm) (NHM 1910.2.4.213), stn 48, ‘Holtenia ground’ 59°32'N, 6°59'W, coll. H.M.S. “Porcupine”, 1869. Other material: Scotland/North Sea • 1 ♂ (35.0 × 27.1 mm) (NHM 1981.108), stn 18, Sula Sgeir, 59°33.7'N, 06°49'W to 59°29.5'N, 06°43.3'W, 915–880 m, coll. A Wheeler, 16 July 1973; 2 ♂♂ (41.3 × 34.0 mm, 38.8 × 31.6 mm) (NHM 1983.449), 56°23'N, 09°18.2'W, 1010–1030 m, coll. W Wales, 20 October 1977; 7 ♂♂ (37.1 × 30.5 mm, 35.0 × 27.6 mm, 30.5 × 23.0 mm, 28.2 × 22.7 mm, 24.6 × 17.2 mm), 3 ♀♀ (37.1 × 30.6 mm, 33.5 × 27.5 mm, 22.4 × 17.2 mm), 1 ovigerous ♀ (35.8 × 29.0 mm) (NHM 1978.99), Atlantic Ocean, 56°33'N, 09°13'W to 56°30'N, 09°14'W, 750 m, coll. RW Ingle, 23 October 1977. Porcupine Seabight • 3 ♂♂ (25.0 × 18.9 mm, 15.5 × 11.1 mm, 9.0 × 6.2 mm), 1 ♀ (34.8 × 28.2 mm), 1 ♀ (with bopyrid; 15.6 × 11.6 mm) (National Oceanography Centre), stn 50601, 51°19.2'N, 11°41.1'W51°21.1'N, 11°42.9'W, 927–770 m, coll. H.M.S. “Discovery” Collection, 1 July 1979; 1 ♂ (23.9 × 18.4 mm), 3 ♀♀ (17.1 × 13.8 mm, 13.6 × 10.6 mm, 12.5 × 9.4 mm) (National Oceanography Centre), stn 9752 #1, 51°16.3'N, 11°42.5'W51°18.6'N, 11°42.8'W, 1010–1045 m, coll. H.M.S. “Discovery” Collection, 7 April 1978.

Diagnosis

Carapace pyriform, strong diverging pseudorostral spines. Postorbital and hepatic spine fused, forming distinct L-shape plate on larger specimens. Carapace with 1 oblong mesogastric plate, 1 oblong cardiac plate, 2 metabranchial plates, 2 horseshoe-shaped epibranchial plates (Figs 3, 4A, 5A, 7A, C, 9A–I). Lateral branchial spine plate-like on specimens with strong carapace plates, present as strong spine on specimens with weak carapace plates (Figs 3, 4A, 5A, 7A, C, 9A–I). Male thoracic sternum deeply concave with distinct grooves (Figs 4B, 5B, 6D). G1 straight, with flattened, sharp tip; straight top margin (Fig. 8C–F).

Figure 6. 

A. Scyramathia carpenteri (Norman, in Thomson 1873), male (35.0 × 27.1 mm) (NHM 1981.108), North Sea; B–F. S. carpenteri (Norman, in Thomson 1873), male (41.3 × 34.0 mm) (NHM 1983.449), North Sea. A, B. lateral view of carapace; C. ventral frontal surface of carapace; D. male thoracic sternum view; E. dorsal surface of male left chela; F. posterior ventral view of male pleon.

Description

Carapace covered by a thick tomentum of setae; with several plates and spines. Pseudorostral spines sharp, straight, diverging, slightly deflected. Supraorbital eave with sharp preorbital spine; sharp angle preorbital spine on juvenile specimens. Hepatic spine pointing upwards, fused with postorbital plate; proportionally longer on juvenile specimens (Figs 4C, 5C, 6A, B, 8A, B). Carapace with plate-like structures: 1 oblong mesogastric plate; 1 oblong cardiac plate; 2 metabranchial plates; 2 horseshoe-shaped epibranchial plates (Figs 3, 4A, 5A, 7A, C, 9A–I). One long sharp branchial spine pointing laterally; proportionately longer in juveniles and specimens with weak carapace. One large intestinal granule on posterior border of carapace (Figs 3, 4A, 5A, 7A, C, 9A–I).

Figure 7. 

Illustration of Scyramathia carpenteri (Norman, in Thomson 1873). A, B. lectotype male (21.6 × 15.9 mm) (NHM 1907.8.28.3), “Holtenia ground” (= between north Scotland and the Faeroe Islands); C, D. paralectotype female (13.0 × 8.4 mm) (NHM 1911.11.8.377), same locality as lectotype. A, C. dorsal view of carapace; B, D. ventral frontal surface of carapace.

Antennal flagellum shorter than pseudorostral spines, about half or nearly equal in length. Basal antennal article fused to carapace; longer than broad; straight to gently convex outer margin; blunt distal angle of article. Pterygostomial region with several granules on outer margin (Figs 4B, 5B, 6C, 7B, D). Buccal frame square, covered by third maxiliped; strong, blunt distal angle of buccal frame (Figs 4B, 5B, 6C, 7B, D).

Chelipeds long, slender; propodus longer than dactylus; dactylus curved, serrulated along entire length (Fig. 6E); carpus short with granules on margin, dorsal margin carinated; merus with 1 tooth on distal upper border. Ambulatory legs slender; merus with sharp angle on distal end; male P2 merus length between 0.6–1.3 times carapace length, female P2 merus length between 0.8–1.0 times carapace length for female, male P2 merus length 8.2–14.0 times width, female P2 merus length 10.5–13.4 times width; male P5 merus length 0.4–0.6 times carapace length, female P5 merus length 0.3–0.5 times carapace length, male P5 merus length 4.2–7.3 times width, female P5 merus length 4.6–5.8 times width (Figs 3, 4A, 5A, 14A, B).

Male thoracic sternum with sternites 1–4 deep concave; sternites 1, 2 wider than long; sternites 3, 4 widest at base, lateral margin slightly constricted; sternites strongly defined (Figs 4B, 5B, 6D, F,). Male pleon with triangular telson and 6 somites, all free; widest on second and third somites. Adult female with round pleon, with all somites distinct. G1 straight, with flattened, sharp distal tip; straight top margin; slightly constricted at distal region (Fig. 8C–F).

Figure 8. 

Illustration of Scyramathia carpenteri (Norman, in Thomson 1873). A, C–H. lectotype male (21.6 × 15.9 mm) (NHM 1907.8.28.3), “Holtenia ground” (= between north Scotland and the Faeroe Islands); B. paralectotype female (13.0 × 8.4 mm) (NHM 1911.11.8.377), same locality as lectotype. A, B. lateral view of carapace; C. ventral view of G1; D. ventral view of distal portion of G1; E. dorsal view of G1; F. dorsal view of distal portion of G1; G. ventral view of G1; H. dorsal view of G2. Scale bars: 1 mm.

Remarks

The identity of Amathia carpenteri Norman, in Thomson 1873 s. str., is critical to the status of Scyramathia as it is the type species of the genus. Although Tavares and Santana (2018) clarified the identity of the genus and had material of the species, they did not examine the types for this species. The species was named by Thomson in 1873 in his famous book about the exploration of the deep Atlantic by the H.M.S. Porcupine, “The Depths of the Seas” (p. 175: fig. 35; reproduced here as Fig. 3), under the name “Amathia Carpenteri” to honour his colleague Dr. W.B. Carpenter. There was no clear indication of number of specimens examined, with the author only commenting that “Another handsome new species, Amathia carpenteri, Norman (fig. 35), was common in the sandy chalkmud of the Holtenia ground.” (Thomson 1873: 176). The figure shows that the cardiac region of the carapace appeared to be damaged (Thomson 1873: fig. 35; reproduced here as Fig. 3). In the NHM are three specimens that are labelled as syntypes of A. carpenteri, all obtained by the H.M.S. Porcupine. None of these specimens, however, match the measurements of the type figure, which was “once and a half the natural size” (Thomson 1873: fig. 35; reproduced here as Fig. 3). Of these, one male specimen (NHM 1907.8.28.3) agrees relatively well with the figure by Thomson (1873: fig. 35; Fig. 3), and the specimen label mentions that it was figured in Thomson (1873). This specimen, with catalogue number, NHM 1907.8.28.3, was noted by Christiansen (1969: 122) to be the lectotype for this species, and in an unpublished note, was also selected by Isabella Gordon (also unpublished data) to be the lectotype. The same specimen was also mentioned by Rice (1990: 10), noting that it is similar to the figure by Norman, in Thomson (1873: fig. 35). The type locality of this species was stated to be on “sandy chalk of the Holtenia ground”. Holtenia Ground is an area between North Scotland and the Faeroe Islands that covers an area of approximately 500 km2, with a depth of 820 to 1000 m (Carpenter et al. 1870; Reiswig and Champagne 1995). Specimen NHM1907.8.28.3 is here formally designated as the lectotype of Amathia carpenteri Norman, in Thomson 1873, to stabilise the taxonomy of the species and the genus.

The three type specimens differ markedly from each other in many ways although they are all from Holtenia Ground. The lectotype male (21.6 × 15.9 mm, NHM 1907.8.28.3) selected here differs markedly from the other two paralectotypes in having a smoother carapace, without the distinct raised plate-like structures on the carapace and the branchial regions are also proportionately more inflated (Figs 4A, 7A). The next largest type is a subadult female (13.0 × 8.4 mm, NHM 1911.11.8.377), and its carapace more closely resembles typical S. carpenteri, with the plates more developed. It is almost certainly the same specimen figured by Norman, in Thomson (1873: fig. 35; Fig. 3), with the shape and structures of the spines and plates agreeing very well, althought the left pseudorostral spine of the specimen is now broken (Fig. 7C). The third type specimen is a small and poorly preserved juvenile female (5.4 × 3.0 mm, NHM 1910.2.4.213) in which the dorsal surface of the carapace as well as the third maxillipeds are covered with setae but without any obvious plates. The surfaces of the third maxillipeds of the other two type specimens are finely granulated or smooth (Figs 4B, 7B, D).

Scyramathia carpenteri exhibits strong variation on the plate-like structures on the carapace of various sized specimens showed variation. D’Udekem d’Acoz (1999) observed that “L’ornementation de la carapace de R. carpenteri est extrêmement variable” but did not mention if it is due to the size of the specimens. Similar sized specimens showed either weak or strong plate-like structures (Fig. 9A–I). These differences are also seen in the type specimens where the larger sized male lectotype specimen has weak plate-like structures on the carapace whereas the same structures on the smaller sized female paralectotype specimens are relatively stronger. The variation of the plate-like structures on the carapace are therefore, not always size dependent. Other morphological characters that show variation on S. carpenteri includes the divergence of the pseudorostral spines, from relatively divergent as seen on the lectotype (Fig. 4A) to subparallel as seen on the male specimen, with catalogue number, NHM 1983.449 (Fig. 5A); the hepatic region having a hepatic spine or plate (Figs 4A, 5A, 7A, C, 9A–I); and the lateral branchial spine on the carapace, from relatively straight, slightly curved or plate-like (Figs 4A, 5A, 7A, C, 9A–I).

Figure 9. 

Variation of plate-like structures on carapace on Scyramathia carpenteri (Norman, in Thomson 1873). A. lectotype male (21.6 × 15.9 mm) (NHM 1907.8.28.3), “Holtenia ground”; B, G. female (22.4 × 17.2 mm, 35.8 × 29.0 mm, respectively) (NHM 1978.99), North Sea; C. male (41.3 × 34.0 mm) (NHM 1983.449), North Sea; D, E, H, I. male (28.2 × 22.7 mm, 37.1 × 30.5 mm, 24.6 × 17.2 mm, 33.0 × 25.7 mm, respectively) (NHM 1978.99), North Sea; F. male (35.0 × 27.1 mm) (NHM 1981.108), North Sea.

Superficially, S. carpenteri resembles S. umbonata (Stimpson, 1871) from the western Atlantic, which also appears to grow to a much larger size (57 mm maximum carapace length; see Tavares et al. 2015). There are, however, consistent differences, with S. carpenteri, which has relatively longer pseudorostral spines, and the outer margin of the basal antennal article is always straight or only slightly convex. The pseudorostral spines of S. umbonata are always proportionately shorter, even when similar sized specimens are compared, and the outer margin of the basal antennal article is always distinctly convex (cf. Tavares et al. 2015: figs 1, 2, 4, 5). These differences are reliable for specimens of similar sizes. As such both species are here recognised as separate taxa.

It is interesting to note that some studies (A. Milne-Edwards and Bouvier 1899, 1900; Bouvier 1922, 1940) mentioned that S. carpenteri does not occur in the Mediterranean, which is the region where the species, S. tenuipes sp. nov., is now found (see remarks for S. tenuipes sp. nov.). Alphonse Milne-Edwards and Bouvier (1899, 1900) and Bouvier (1940) had material they referred to “S. carpenteri” from the Azores, and up north of São Jorge, and the Sahara beach. Based on their figures (see A. Milne-Edwards and Bouvier 1899: pl. 1 fig. 4; A. Milne-Edwards and Bouvier 1900: pl. 20 Figs 1, 6; Bouvier 1940: pl. 14 fig. 1), however, the specimens they figured are more similar to S. tenuipes sp. nov., with the ambulatory legs long and slender, and these records are hence referred there for now. The plates on the carapace as figured (see A. Milne-Edwards and Bouvier 1899: pl. 1 fig. 4; A. Milne-Edwards and Bouvier 1900: pl. 20 Figs 1, 6; Bouvier 1940: pl. 14 fig. 1) appear relatively stronger than what has been observed in S. tenuipes sp. nov., but as discussed earlier, this character is known to vary in S. carpenteri s. str. in any case.

Distribution

Scyramathia carpenteri is known from the type locality, “sandy chalk of the Holtenia ground” [= between North Scotland and the Faeroe Islands] (Carpenter et al. 1870; Norman, in Thomson 1873; Reiswig and Champagne 1995), South Iceland (Hansen 1908), Southwest Faeroe Islands, South of Norway, West of British Isles (Clark 1986).

Scyramathia hertwigi Doflein, in Chun, 1900

Figs 10A–C, 14C, D, 15A–D

Scyramathia Hertwigi: Doflein, in Chun 1900: 172 (fig.); Doflein, in Chun 1902–1903: 172 (fig.).

Scyramathia Hertwigi Doflein, 1904: 81–84, pl. 27 fig. 3, pl. 28 fig. 1 (type locality: off Cape Point and on Agulhas Bank).

Scyramathia hertwigi: Stebbing 1902: 7, pl. 6; Stebbing 1905: 25; Stebbing 1910: 289; Barnard 1950: 50, 51, fig. 11b, c; Low et al. 2017: 116, fig. 2A, tables 2, 3.

Rochinia carpenteri: Monod 1956: 32 (list), 515. [not Amathia carpenteri Norman, in Thomson 1873].

Rochinia hertwigi: Serène and Lohavanijaya 1973: 56 (key); MacPherson 1983: 32–35, figs 19, 20a–c; Griffin and Tranter 1986: 175 (key), 180; Casadío et al. 2005: 159 (list); Ng and Richer de Forges 2007: 62 (list); Ng et al. 2008: 105 (list); Tavares and Santana 2018: 223 (list); Griffiths et al. 2018: 199.

Materials examined

Lectotype : ♂ (33.7 × 27.0 mm) (ZMB 13699), Atlantic, South Atlantic Ocean, Agulhasstrom [= Agulhas current], coll. Deutsche Tiefsee Expedition, 2 November 1898. Paralectotypes: 3 ♂♂ (24.7 × 19.0 mm, 23.6 × 17.7 mm, 18.7 × 13.7 mm), 2 ♀♀ (28.3 × 22.6 mm, 26.4 × 20.2 mm) (ZMB 13700), stn 103, Agulhasstrom [= Agulhas current], 500 m, coll. Deutsche Tiefsee Expedition, 2 November 1898; 8 ♂♂ (2 largest: 22.8 × 17.9 mm, 21.6 × 16.5 mm; 2 smallest: 16.2 × 12.3 mm, 16.0 × 11.9 mm), 2 ovigerous ♀♀ (25.9 × 20.5 mm, 24.3 × 19.2 mm), 8 ♀♀ (2 largest: 25.1 × 19.4 mm, 22.6 × 17.3 mm; 2 smallest: 20.1 × 15.2 mm, 14.3 × 10.3 mm) (SMF 23085), stn 103, Atlantic, South Atlantic, Agulhasstrom [= Agulhas current], 35°10'S, 23°2'E, 500 m, coll. FS ‘Valdivia”, 2 November 1898. Other material: South Africa • 1 ♂ (29.2 × 22.9 mm) (SAM-A39580), west grounds, I + J trawls, coll. UCT Ecological Survey, 22 November 1946; 2 ♂♂ (33.2 × 27.7 mm, 31.9 × 27.1 mm) (SAM-A47887), west coast, 333 m, coll. 17 January 2007; 1 ♂ (19.0 × 14.2 mm), 1 ovigerous ♀ (34.4 × 27.1 mm) (SAM-A12149), West of Slangkop, Cape Peninsula, 457 m, coll. P Henry, 16 February 1965; 1 ♂ (47.0 × 38.0 mm) (SAM-A40027), stn A10121-082-035-4197, 34°36.6'S, 25°15.1'E, 480 m, coll. R.S. Africana South Coast Hare Biomass Survey, 31 May 1990; 1 ♂ (23.7 × 18.5 mm), 1 ovigerous ♀ (36.0 × 28.5 mm), 1 ♀ (24.5 × 18.4 mm) (SAM-A47872), stn NAN401 T027, 34°20.2'S, 18°12.7'E, 290 m, coll. L Atkinson, Nansen West Coast Survey, 15 January 2007; 1 ♂ (40.7 × 31.1 mm) (AM P34653), off Cape Town, 34°21'S, 17°57'E, coll. 18 December 1929; 3 ♂♂ (34.6 × 27.3 mm, 29.3 × 23.1 mm, 22.0 × 17.0 mm) (NHM 1924.7.19.1–3), stn 103, Cape, no other data. SOUTH ATLANTIC • 1 ♀ (21.5 × 18.9 mm) (ZMB 26204), stn 153/1968, 28°22'S, 14°24'E, 470–460 m, coll. E Haeckel, 9 May 1968.

Diagnosis

Carapace pyriform, covered with short setae. Pseudorostral spine short, diverging. Supraorbital eave with sharp preorbital spine; postorbital lobe fused to hepatic plate on large size specimens. Carapace with plates: 1 hepatic plate pointing upwards, 1 short mesogastric spine above 1 oblong mesogastric plate, 1 large protogastric granule, 1 oblong cardiac plate, 1 large epibranchial plate, 1 mesobranchial plate, 1 posterior blunt spine, 1 lateral branchial plate curved upwards (Fig. 10A). Plates less distinct on juvenile specimens. Antennal flagellum equal to or slightly longer than pseudorostral spines. Basal antennal article with convex outer margin, with blunt distal angle of article. Buccal frame covered by third maxilliped, distal angle of buccal frame raised, distinct. Pterygostomial region plate-like with 3 or 4 granules on outer margin (Fig. 10B). Chelipeds slender, propodus longer than fingers, slightly carinate; carpus with granules on carina margin; merus triangular in cross-section, granulated on edge. Ambulatory legs slender; merus cylindrical; male P2 merus length 0.8–1.2 times carapace length, female P2 merus length 0.4–0.8 times carapace length, male P2 merus length 10.9–14.9 times width, female P2 merus length about 8.5 times width; male P5 merus length 0.4–0.6 times carapace length, female P5 merus length 0.4–0.5 times carapace length, male P5 merus length 4–6.8 times width, female P5 merus length 4.6–5.8 times width (Figs 10A, 14C, D). Male thoracic sternum wide, concave on sternites 1–4; regions well defined; sternites 3, 4 widest, lateral margins slightly constricted medially. Male pleon with triangular telson; surface of somites smooth (Fig. 10B). G1 straight with flattened sharp tip (Fig. 15A–D).

Figure 10. 

Scyramathia hertwigi Doflein, in Chun 1900, lectotype male (33.7 × 27.0 mm) (ZMB 13699), South Atlantic Ocean. A. overall dorsal view; B. overall ventral view; C. lateral view of carapace.

Remarks

The author and year of publication for Scyramathia hertwigi has traditionally been cited as Doflein (1904). Doflein (1904: 81), however, cited a figure of this species in Chun (1900). Hence, the authorship and date of publication for S. hertwigi should be “Doflein in Chun 1900” (see Low et al. 2017). The figure by Doflein that is in Chun (1900) was stated to be drawn to scale (Low et al. 2017). Doflein (1904), when describing the species, did not specify the number of specimens examined, but mentioned that many male and female specimens were examined. The specimen that was figured by Doflein, in Chun (1900), is currently deposited in ZMB. The measurement of the male specimen deposited in the Museum für Naturkunde, Berlin (ZMB) is very similar to that drawn by Doflein, in Chun (1900) and we are confident they are the same specimen. The location where this specimen was collected matches that listed in Doflein (1904). Since no type was designated in Chun (1900) or Doflein (1904), this male specimen (ZMB 13699) is herein designated as the lectotype of S. hertwigi.

The Atlas by Doflein (1904: pl. 27 figs 1–7) has figures that show a series of sizes of the carapace with plates, in particular the horse-shoe shaped epibranchial plate. In this species, the pseudorostral spines are always proportionately shorter than those in S. carpenteri. The differences between S. carpenteri and S. hertwigi were discussed by MacPherson (1983: 35), and we agree with his interpretation. There are also slight variations amongst the specimens from South Atlantic (SMF 23085), which tend to have less distinct plates on the carapace, and slightly shorter and stouter ambulatory legs while a large specimen (AM P34653) from South Africa has distinct plates on carapace, and somewhat longer and slender ambulatory legs. These differences, however, are not substantial and should not be a reason to treat them as different.

The specimens of “Scyramathia carpenteri” from off Cape Bajador, West Africa (= off Boujdour, Western Sahara) reported by Tavares and Santana (2018: 204) may not be that species. This location overlaps with the distribution range of S. tenuipes sp. nov., and it is possible their specimens belong to this species instead. Their material will need to be re-examined.

Distribution

Scyramathia hertwigi is known from its type locality, off Cape Point and on Agulhas Bank, South Africa, as well as parts of the south Atlantic along the African coast. It is the only species that enters the southwestern-most edge of the Indian Ocean.

Scyramathia umbonata (Stimpson, 1871)

Figs 11A–C, 14E

Scyra umbonata Stimpson 1871: 115 (type locality: off Sand Key, Florida).

Scyra umbonata: A. Milne-Edwards 1875 [in 1873–1880]: 87–88; A. Milne-Edwards 1880 [in 1873–1880]: pl. 31A fig. 5–5b; A. Milne Edwards 1880c: 2; Sars, 1885: 6 (list), 7 (list), 274 (list); Smith 1886: 626[22 on separate]

Scyramathia umbonata: A. Milne-Edwards 1880a: 356; A. Milne-Edwards 1880b: 277; A. Milne-Edwards and Bouvier 1923: 381; Tavares and Santana 2018: 204 (list), Figs 1, 12C, 13A, B.

Anamathia umbonata: Rathbun 1894: 61–62, pl. 1 Figs 13; Faxon 1895: 10.

Rochinia umbonata: Rathbun 1925: 210 (key), 222, 223, text-fig. 85, pl. 72, pl. 73 fig. 1; Bullis and Thompson 1965: 12 (list); Williams et al. 1968: 61, fig. 16; Takeda 1983: 135; Williams 1984: 322 (key), 323–325, figs 258, 260c; Abele and Kim 1986: 42, 172 (key); Griffin and Tranter 1986: 175 (list); Soto 1991: 628 (table), 632 (list), 633 (list), 634, 636 (list); Tavares 1991: 161 (list), 164; Poupin 1994: 43, 44, pl. 4 fig. g; Nizinski 2003: 128; McLaughlin et al. 2005: 253 (list); Wicksten and Packard 2005: 1762 (list); Casadío et al. 2005: 159 (list), 160; Ng and Richer de Forges 2007: 63 (list); Ng et al. 2008: 106 (list); Felder et al. 2009: 1078 (list); Pettan 2013: 72–80, 87 (key), figs 14a, b, 17h, 18b, 19b, 20b, 28a–d, 29a–c; Pettan and Tavares 2014: 306 (list), fig. 1; Tavares et al. 2016: 1065–1069, figs 1–6.

Rochinia confusa Tavares 1991: 161 (list), 162–165, fig. 1A–C, 5A, pl. 1 A–D (type locality: off Rio de Janeiro, 23°46'S, 42°09'W, Brazil); De Melo 1996: 266 (key), 267, unnumbered in-text fig.; Casadío et al. 2005: 158 (list); Ng and Richer de Forges 2007: 63 (list); Ng et al. 2008: 105 (list); Pettan 2013: 21–26, 87 (key), figs 7a, b, 16a, 18a, 19a, 20a; Pettan and Tavares 2014: 305 (list), fig. 1. (For the remaining Neotropics literature, see Tavares et al. 2016).

Material examined

Paratype of Rochinia confusa Tavares, 1991: 1 ♂ (8.0 × 5.2 mm) (MNHN-IU-2014-19836 [= MNHN-B24570]), stn 54, Brazil, 19°36'S, 38°53'W, coll. TAAF/MD55, 2 June 1987. Other material: 1 ♀ (36.7 × 27.9 mm) (NSMT-Cr7178), off Suriname, 7°47'N, 54°08'W, 60 m, coll. 1980.

Diagnosis

Carapace pyriform, covered with short setae. Pseudorostral spine short, diverging. Supraorbital eave fused to carapace with sharp preorbital spine; postorbital lobe and hepatic spine fused, forming a L-shape plate-like structure, pointing upwards; plate-like structure not formed on juvenile specimens. Carapace with raised plates: 1 hepatic plate curved, pointing upwards, 1 short mesogastric granule above 1 oblong mesogastric plate, 1 oblong cardiac plate, 1 large epibranchial plate, 1 mesobranchial plate, 1 posterior blunt spine, 1 lateral branchial plate curved upwards (Fig. 11A). Plates less distinct on juvenile specimens. Antennal flagellum equal to or slightly longer than pseudorostral spines. Basal antennal article with convex outer margin, with blunt distal angle of article. Buccal frame covered by third maxilliped, distal angle of buccal frame raised, distinct. Pterygostomial region plate-like with 3 or 4 granules on outer margin (Fig. 11B). Chelipeds slender, propodus longer than fingers, slightly carinate. Ambulatory legs slender; merus cylindrical; male P2 merus length approximately 0.7 times carapace length, female P2 merus length 0.6–1.0 times carapace length, male P2 merus length approximately 9.0 times width, female P2 merus length 9.5–12.3 times width; male P5 merus length approximately 04 times carapace length, female P5 merus length 04–0.6 times carapace length, male P5 merus length approximately 4.4 times width, female P5 merus length 4.1–6.9 times width (cf. Rathbun 1925: pl. 72, 73 fig. 1; Figs 11A, 14E). Male thoracic sternum wide, concave on sternites 1–4; regions well defined; sternites 3, 4 widest, lateral margins slightly constricted. Male pleon, telson triangular; surface of somites smooth (Fig. 11E). G1 straight with flattened sharp tip (cf. Tavares and Santana 2018: fig. 13A, B).

Figure 11. 

A–C. Scyramathia umbonata (Stimpson, 1871), female (36.7 × 27.9 mm) (NSMT-Cr7178), off Suriname; D–F. Rochinia confusa Tavares, 1991, paratype male (8.0 × 5.2 mm) (MNHN-IU-2014-19836 [= MNHN-B24570]), Brazil. A, C. overall dorsal view; B, E. overall ventral view; C, F. lateral view of carapace.

Remarks

Scyramathia umbonata was originally described as a species of Scyra by Stimpson (1871) from off Sand Keys in Florida. Stimpson (1871) provided measurements for one male specimen but did not indicate if he had more material. It was transferred to Scyramathia by A. Milne-Edwards (1880a), and Rathbun (1894) subsequently referred it to Anamathia. Rathbun (1925) later transferred the species to Rochinia when she synonymised the genera. The type material for the S. umbonata is probably no longer extant (Tavares et al. 2016).

Tavares et al. (2016) discussed the taxonomy of S. umbonata at length (see also Tavares and Santana 2018) and synonymised Rochinia confusa Tavares, 1991, under S. umbonata, stating that R. umbonata undergoes drastic changes in morphology as it grows, with more distinct plates being present on larger sized specimens. This is the same pattern as seen in S. carpenteri and S. hertwigi. Tavares and Santana (2018) later transferred it back to Scyramathia. There is no need to redescribe or figure this species as the detailed study and figures of this species by Tavares et al. (2016) and Tavares and Santana (2018) are quite sufficient. For these reasons, a neotype is also not needed for this species for the time being.

Distribution

Scyramathia umbonata is recorded from the North Carolina to Gulf of Mexico (United States of America), Nicaragua, Caribbean Island, and Brazil (Tavares et al. 2016).

Scyramathia tenuipes sp. nov.

Figs 12, 13A–C, 14F–J, 15E–H

Scyramathia Carpenteri: A. Milne-Edwards and Bouvier 1894: 13, 14 (in part); A. Milne-Edwards and Bouvier 1899: 43 (in part), pl. 1 fig. 4; Bouvier 1922: 81 (in part). [not Amathia carpenteri Norman, in Thomson 1873].

Rochinia Carpenteri: Bouvier 1940: 344, 345 (in part), fig. 209, pl. 14 figs 1–3. [not Amathia carpenteri Norman, in Thomson 1873].

Rochinia (Amathia) carpenteri: Dieuzeide 1955: 53–55, fig. 9.

Rochinia (Scyramathia) carpenteri: Dieuzeide and Roland 1958: 21 (list), 62 (list). [not Amathia carpenteri Norman, in Thomson 1873].

Rochinia carpenteri: Dieuzeide 1960: 76 (list); Fransen 1991: 6, (list), 7 (list), 181 (list); García Raso 1996: 741 (in part); D’Udekem d’Acoz 1999: 194 (in part); Abelló et al. 2002: 187 (table), 189 (list), 191 (table), 193 (table), 195 (table); Araújo et al. 2014: 38, fig. 1F. [not Amathia carpenteri Norman, in Thomson 1873].

Material examined

Holotype : ♂ (33.9 × 26.3 mm) (ICMD288), Mediterranean Sea, coll. 1994. Paratypes: 1 ovigerous ♀ (31.2 × 24.6 mm) (ICMD000708), Málaga, Mediterranean Sea, 36°31'19"N, 03°59'18"W, 650 m, coll. 1 May 2016; 1 ♀ (27.7 × 20.5 mm) (ICMD000712), Castell de Ferro, Mediterranean Sea, 36°39'19"N, 03°16'47"W, 638 m, coll. 5 May 2016; 1 ovigerous ♀ (32.2 × 25.8 mm) (ZRC 2019.1635, ex. ICMD000696), Adra, Mediterranean Sea, 36°33'51"N, 03°03'52"W, 572 m, coll 29 April 2016; 1 ♂ (32.0 × 25.4 mm) (ZRC 2019.1636, ex. ICMD000713), Málaga, Mediterranean Sea, 36°18'44"N, 04°20'25"W, 770 m, coll. 24 April 2016; 1 ♂ (29.6 × 22.9 mm) (ICMD000700), Alborán Island, Mediterranean Sea, 35°33'51"N, 03°03'52"W, 819 m, coll. 6 May 2016; 1 ♂ (29.7 × 22.0 mm) (ICMD000707), Málaga, Mediterranean Sea, 35°31'7"N, 03°59'18"W, 650 m, coll. 1 May 2016; 1 ovigerous ♀ (30.6 × 23.7 mm) (ICMB000711), Castell de Ferro, Mediterranean Sea, 36°39'19"N, 03°16'47"W, 638 m, coll. 5 May 2016; 1 ♂ (14.9 × 10.5 mm) (ZRC 2019.1637, ex. ICMD000706), Málaga, Mediterranean Sea, 36°31'19"N, 03°59'18"W, 650 m, coll. 1 May 2016; 1 ♀ (16.3 × 12.5 mm) (ZRC 2019.1638, ex. ICMB000705), Málaga, Mediterranean Sea, 36°31'19"N, 03°59'18"W, 650 m, coll. 1 May 2016.

Diagnosis

Carapace pyriform. Pseudorostral spines straight, almost half of carapace length. Supraorbital eave fused with carapace, with blunt preorbital spine; postorbital lobe fused with hepatic spine. Carapace with plates: 1 hepatic spine, 1 small granule above 1 oblong mesogastric plate, 1 protogastric granule, 1 epibranchial plate, 1 oblong cardiac plate, 1 metabranchial granule, 1 lateral branchial spine, 1 blunt posterior spine (Figs 12, 13A). Antennal flagellum shorter than pseudorostral spines, about half of pseudorostral length. Basal antennal article longer than broad, straight outer margin with blunt roundish distal angle of article (Fig. 13B). Buccal frame with distal angle of buccal frame slightly raised with round edge. Pterygostomial region with 3 or 4 granules on outer margins (Fig. 13B). Chelipeds slender, propodus longer than fingers, slightly carinate margin; carpus with granules; merus smooth, triangular in cross-section. Ambulatory legs slender; merus with distal angle blunt; male P2 merus length 1.0–1.4 times carapace length, female P2 merus length 1.0–1.3 times carapace length, male P2 merus length 17.3–20.5 times width, female P2 merus length 14.8–21.5 times width; male P5 merus length 0.4–0.7 times carapace length, female P5 merus length 0.5–0.6 times carapace length, male P5 merus length 7.1–9.5 times width, female P5 merus length 7.9–8.8 times width (Figs 12, 13A, 14F–J). Male thoracic sternum flat, slightly concave; sternites 3, 4 widest. Male pleon with triangular telson and all 6 somites free; somites 2, 3 widest; surface of somites smooth (Fig. 13B). G1 straight, distal part relatively shorter, sharp tip (Fig. 15E–H).

Figure 12. 

Colour in life of Scyramathia tenuipes sp. nov., holotype male (33.9 × 26.3 mm) (ICMD288), Mediterranean Sea (Credits: MacPherson E & Abelló P).

Etymology

The term “tenuis”, which means thin in Latin, is used for the slender ambulatory legs. The name is used as a noun in apposition.

Remarks

Scyramathia tenuipes sp. nov. is superficially similar to S. carpenteri, but there are several important differences between the two species. Firstly, all the adult specimens of S. tenuipes sp. nov. examined have relatively less distinct plates on the carapace (Figs 12, 13A) (versus adult S. carpenteri usually have more prominent plates on the carapace; Figs 3, 4A, 5A, 7A, C, 9A–I). More significantly, it also has distinctly longer and more slender ambulatory legs, with the P2 meri length of all the male specimens longer or subequal in length (1.0–1.4 times) to the carapace length, and the P5 meri length 7.1–9.5 times longer than the width (Figs 12, 13A, 14F–J) (versus S. carpenteri has relatively shorter and less slender ambulatory legs, with the P2 meri of all male specimens typically subequal in length (0.7–1.1 times) to the carapace length, and the P5 meri length 4.2–7.3 times the width; Figs 3, 4A, 5A, 14A, B). This character is consistent in all the adult male and female specimens examined. The pterygostomial region of adult S. tenuipes sp. nov. has relatively more distinct granules (Fig. 13B) compared to the more plate-like granules on that of adult S. carpenteri (Figs 4B, 5B, 6C, 7B, D); and the distal part of the G1 of S. tenuipes sp. nov. is always relatively shorter (Fig. 15E–H) compared to the longer ones of S. carpenteri (Fig. 8C–F).

Figure 13. 

Scyramathia tenuipes sp. nov., holotype male (33.9 × 26.3 mm) (ICMD288), Mediterranean Sea. A. overall dorsal view; B. overall ventral view; C. lateral view of carapace.

An unpublished genetic study by the first author comparing S. tenuipes sp. nov. with S. carpenteri, S. hertwigi, S. umbonata and other Rochinia species shows small but consistent differences that indicate we are dealing with a recent but separate species. A total of five genes, three mitochondrial genes: COI, 12S, 16S, and two nuclear genes: 18S and H3, were used for the molecular analysis in this unpublished work. The cytochrome oxidase I (COI) dendrogram for S. tenuipes sp. nov. and S. carpenteri shows a consistent 0.3% difference between them whereas the differences between the other Scyramathia and Rochinia species ranged from 1.3–10.9 %. Significantly, the phylogenetic tree from the Maximum Likelihood analysis shows an 87% support for S. tenuipes sp. nov. and S. carpenteri as separate clades.

The type locality of S. tenuipes sp. nov. is an interesting area in the Mediterranean. The Alboran Sea is at the western narrow part of Mediterranean that ends at the Strait of Gibraltar. The complex circulation of the waters through this very narrow strait of approximately 14 km, is known to play a key role in regulating the gene flow for a number of benthic species (Palero et al. 2011). The peculiar hydrographic features of the Strait of Gibraltar were established since the end of the Messinian (upper Miocene) about 5.4 million years ago, after a period of very low sea level of 500,000 years where the strait was dry, and the Mediterrannean Sea was completely closed. Between Spain and Morocco, the very narrow strait has several choke points not deeper than 350 m that seriously affects the water currents. The surface Atlantic waters are entering at speeds varying from 1 to 2 knots, whereas the outflow of the Mediterranean waters nearer the sea bottom flows at a speed of 1.1 to 0.9 knots (Millot 2005). The physical characteristics of these two opposite currents are also very different: the Atlantic surface water has a salinity of 36 g/l while the Mediterranean waters are denser at 38 g/l of salt. As a result of such conditions, the Alboran Sea has more influx from the Altantic Ocean than the rest of the Mediterranean basin (Pascual et al. 2016). These peculiar oceanographical conditions could explain why S. tenuipes sp. nov. is most common in the Alboran Sea and not in the rest of Mediterranean or even in the main Atlantic waters adjacent. In the Mediterranean, S. tenuipes sp. nov. is known only from the western part of Mediterranean along the coast of Spain (Marco-Herero et al. 2015; Palero pers. comm.). The unusual oceanographic features in the Alboran Sea are what the species prefers. A study by Tortonese (1964) on the Mediterranean fish fauna have shown that the fish fauna differs in the different regions in the Mediterranean, for example the scorpionfish, Trachyscorpia cristulata echinata (Köhler, 1896) (Sebastidae), is found only in the western part of Mediterranean, which is also observed by submersible during the Cyanalboran cruise (Zibrowius pers. comm.). It has also been discussed by d’Udekem d’Acoz (1999) that there are crustacean species that showed variations in morphological differences between the Atlantic and Mediterranean populations with unknown, continuous or absence of intermediate populations in between (see d’Udekem d’Acoz 1999: table 1). These morphological differences due to the geographical distribution are linked to the different physical characteristics of the Atlantic and Mediterranean waters (d’Udekem d’Acoz 1999). However, it is likely that these species might be separate ones, as in this case, which requires more study.

There are various records of S. tenuipes sp. nov. as “S. carpenteri”, and while A. Milne-Edwards and Bouvier (1899, 1900) and Bouvier (1922, 1940) state that S. carpenteri was not known from the Mediterranean, d’Udekem d’Acoz (1999) noted that a Mediterranean specimen was figured by Dieuzeide (1955). The figure of the Mediterranean specimen by Dieuzeide (1955: fig. 9), shows an animal with long and slender ambulatory legs, weak plates on carapace, and relatively straight and slightly divergent pseudorostral spines, diagnostic characters of S. tenuipes sp. nov. (Figs 12, 13A). Subsequently, Dieuzeide (1960) commented that the species was commonly found among the bamboo coral, Isidella elongata (Esper, 1788) (Isididae: Alcyonacea) in the Mediterranean. Some of the specimens of S. tenuipes sp. nov. that were examined in this study were found with unidentified deep-sea anemones attached on their carapace.

Distribution

Scyramathia tenuipes sp. nov. is known from its type locality, the western Mediterranean Sea, with possible records from Algeria (Dieuzeide 1955), Azores, north of Sao Jorge in Portugal, and Sahara beach (A. Milne-Edwards and Bouvier 1894, 1899, 1900; Bouvier 1940), Morocco (A. Milne-Edwards and Bouvier 1900; Sivertsen and Holthuis 1956; Fransen 1991), and Mauritania (A. Milne-Edwards and Bouvier 1900).

Figure 14. 

Dorsal view of P5. A, B. Scyramathia carpenteri (Norman, in Thomson 1873); A. male (41.3 × 34.0 mm) (NHM 1983.449), North Sea; B. ovigerous female (35.8 × 29.0 mm) (NHM 1978.99), Atlantic Ocean; C, D. S. hertwigi Doflein, in Chun 1900; C. male (34.6 × 27.3 mm) (NHM 1924.7.19.1–3), South Africa; D. female (24.5 × 18.4 mm) (SAM-A47872), South Africa; E, S. umbonata (Stimpson, 1871), female (36.7 × 27.9 mm) (NSMT-Cr7178), off Suriname; F–J. S. tenuipes sp. nov.; F. holotype male (33.9 × 26.3 mm) (ICMD288), Mediterranean Sea; G. paratype male (29.7 × 22.0 mm) (ICMD000707), Mediterranean Sea; H. paratype male (32.0 × 25.4 mm) (ZRC 2019.1636), Mediterranean Sea,; I. paratype ovigerous female (31.2 × 24.6 mm) (ICMD000708), Mediterranean Sea,; J. paratype ovigerous female (32.2 × 25.8 mm) (ZRC 2019.1635), Mediterranean Sea.

Figure 15. 

A–D. Scyramathia hertwigi Doflein, in Chun 1900, paralectotype male (24.7 × 19.0 mm) (ZMB 13700), Agulhasstrom [= Agulhas current], left G1; E–H. Scyramathia tenuipes sp. nov., holotype male (33.9 × 26.3 mm) (ICMD288), Mediterranean Sea, left G1. A, E. ventral view; B, F. ventral view of distal portion; C, G. dorsal view; D, H. dorsal view of distal portion. Scale bars: 1 mm.

Anamathia Smith, 1885

Amathia Roux 1828: 8, 11, 12, pl. 3; H. Milne Edwards 1834a: 285, 286.

Pisa (Amathia) De Haan 1839: 78 (key), 84.

Anamathia Smith 1885: 493 (replacement name for Amathia Roux, 1828); Miers 1886: 25 (in part); Alcock 1895: 165 (list); Faxon 1895: 8; Bouvier 1940: 345; Ng et al. 2008: 102 (list); Tavares and Santana 2018: 202, 208.

Type species

Amathia rissoana Roux, 1828, by monotypy, gender feminine.

Diagnosis

Carapace pyriform with spines. Pseudorostral spines relatively long, straight, stout, cylindrical, slight diverging at approximately 20° angle or less. Supraorbital eave with blunt preorbital angle; weak postorbital lobe small, round anterior margin. Carapace with strong spines; 3 spines medially: metagastric, cardiac, intestinal; strong lateral branchial spines pointed outwards and downwards (Figs 16A, 17A). Antennal flagellum shorter than pseudorostral spines (Fig. 17B). Basal antennal article longer than broad, distal angle blunt, relatively straight outer margin. Distal angle of buccal frame elongated, forming strong blunt angle. Pterygostomial region with granules on outer margin (Fig. 17B). Chelipeds slender, articles with rounded margins; propodus slender, longer than fingers; carpus with spines on outer margin; merus with spine on distal angle, rounded margins. Ambulatory legs slender, articles with rounded margins; merus with blunt distal angle; P2 longest (Figs 16A, 17A). Male thoracic sternum concave anteriorly, constricted between sternites 1, 2 and 3, 4; sternites 3, 4 with lateral margin slightly constricted. Male pleon triangular, telson triangular, margin round; with raised granules on somites 2–5 (Fig. 17B). G1 straight with flattened sharp tip (Fig. 18A–D); G2 with distal tip round (Fig. 18E, F).

Remarks

The genus was first described as Amathia by Roux (1828). The name was changed to Anamathia by Smith (1885), as the earlier name was preoccupied for a bryozoan (Lamouroux 1812). This genus was synonymised by Rathbun (1925) under Rochinia A. Milne-Edwards, 1875, with no detailed explanation. Ng et al. (2008) listed the genus as valid as well but again with no explanation. Tavares and Santana (2018) discussed the matter and considered this genus as valid, listing the morphological differences from Rochinia and Scyramathia. Anamathia hystrix (Stimpson, 1871) was also transferred to this genus by Tavares and Santana (2018) without explanation. Although the external features of this species resemble A. rissoana, R. hystrix has a strong preorbital spine and a pronounced mesogastric spine in the middle of the carapace; and on the basis of this as well as unpublished genetic data, we are of the opinion that it should not be placed in this genus. As such, R. hystrix together with the rest of the Atlantic and East Pacific Rochinia sensu lato species will be dealt with separately by Lee et al. (in prep).

Anamathia rissoana (Roux, 1828)

Figs 16A–C, 17A–C, 18A–F

Amathia rissoana Roux, 1828: 2 unnumbered paes, pl. 3 (type locality: Mediterranean).

Amathia Rissoana: H. Milne Edwards 1834a: 286; H. Milne Edwards 1840: pl. 34 bis fig. 2; Heller 1863: 29, 30, pl. 1 figs 4, 5.

Anamathia rissoana: Smith 1885: 493 (list); A. Milne-Edwards and Bouvier 1899: 43, 44, pl. 1 fig. 5; A. Milne-Edwards and Bouvier 1900: 138, pl. 20 fig. 11; Pesta 1918: 348, fig. 112; Bouvier 1922: 80; Rathbun 1925: 204 (list); Bouvier 1940: 345–347, fig. 210, pl. 14 fig. 4; Kramp 1947: 3, fig. 1; Dieuzeide 1950: 52; Dieuzeide 1955: 20 (list), 57, fig. 10; Holthuis 1962: 242 (list); Vervoort 1966: 383, fig. 6; Zariquiey Álvarez 1968: 465, figs 7c, 153b, 154g, h; Holthuis 1977: 74; Manning and Holthuis 1981: 253 (list), 254; Relini et al. 1986: 145 (table); Guerao and Abelló 1996: 245–250, Figs 13; Mura et al. 2005: 110–115, figs 1–8; Ng et al. 2008: 102 (list); Maynou and Cartes 2012: table 1; Araújo et al. 2014: 38 (list); Bo et al. 2015: 14, fig. 4P; Mastrototaro et al. 2016: 244–246, fig. 2A–H; Mastrototaro et al. 2017: 214, 216 (list), 217, 222, figs 5j, k, 6j–n, table 3; Taviani et al. 2017: 65, fig. 6B, table 3; Ceccon and De Angeli 2018: 151, fig. 2.1; Tavares and Santana 2018: 208 (list), figs 3, 12B, 13C, D.

Rochinia rissoana: Garth 1958: 283, 289; Clark 1986: 192, 193 (map); d’Udekem d’Acoz 1999: 194; González-Gordillo et al. 2001: 280 (list); Mura and Corda 2011: 679 (table).

Rochinia (Anamathia) rissoana: Cartes et al. 2013: 62.

Material examined

North Atlantic Ocean • 1 ♂ (22.9 × 16.1 mm), 1 ♀ (23.5 × 16.5 mm) (SMF 4420), stn 9C-189, 30°5.1'N, 28°38.4'W, 340–305 m, coll. Meteor Expedition, 26 July 1967; 4 ♂♂ (26.2 × 17.5 mm, 24.0 × 15.2 mm, 19.9 × 12.8 mm, 13.1 × 8.2 mm), 4 ovigerous ♀♀ (27.5 × 17.2 mm, 25.1 × 16.2 mm, 23.1 × 14.6 mm, 21.0 × 13.4 mm, 19.8 × 12.5 mm), (SMF 4410), stn 9c-130, 36°41.1'N, 14°14.8'W, 216–225 m, coll. Meteor Expedition, 3 July 1967; 1 ♂ (16.5 × 11.2 mm), 1 ♀ (24.0 × 15.4 mm), 2 juveniles (SMF 5542), stn 19–131, 30°8'N, 28°38.5'W, 269 m, coll. Meteor Expedition, 17 February 1970.

Figure 16. 

Illustration of Anamathia rissoana (Roux, 1828), Mediterranean. A. overall dorsal view; B. male pleon; C. female pleon. (After Roux 1828: pl. 3).

Description

Carapace pyriform, covered with layer of setae, smooth when denuded (Figs 16A, 17A). Supraorbital eave fused with carapace with blunt preorbital angle; postorbital lobe cup-like, small (Fig. 17B).

Carapace with 13 spines: 1 hepatic spine, 1 protogastric spine, 1 metagastric spine, 1 lateral epibranchial spine, 1 lateral branchial spine, 1 cardiac spine, 1 metabranchial spine, 1 strong intestinal spine near posterior of carapace (Figs 16A, 17A).

Antennal flagellum shorter than pseudorostral spines. Basal antennal article longer than broad, narrow; straight outer margin with blunt distal angle. Presence of granule at base of article. Buccal frame covered by third maxilliped, distal angle of buccal frame distinct, highly protruded, forming blunt angle. Pterygostomial region with 2 or 3 granules on outer margin; second granule biggest (Fig. 17B).

Chelipeds slender, palm longer than dactylus; carpus with 2 spines on distal edge; merus with sharp spine on distal angle. Ambulatory legs slender; distal angle of merus blunt; P2 longest (Figs 16A, 17A).

Male thoracic sternum slightly concave, constricted between sternites 1, 2 and 3, 4; sternites 3, 4 widest (Fig. 17B). Male pleon with semi circular telson and 6 somites; somites 2, 3 widest; large granule on middle of second to fifth somites (Fig. 16B, 17B). Adult female with round pleon, with all somites free (Fig. 16C). G1 straight, with flattened, sharp tip (Fig. 18A–D).

Figure 17. 

Anamathia rissoana (Roux, 1828), male (22.9 × 16.1 mm) (SMF 4420), North Atlantic Ocean. A. overall dorsal view; B. overall ventral view; C. lateral view of carapace.

Remarks

Roux (1828) described Amathia rissoana from the Mediterranean, and it was noted by Roux (1828) that the rare crab is found 20 metres deep amongst “les algues et les focus”. Roux (1828) did not indicate the number of specimens that was examined, but he had at least one male and one female which were figured (Roux 1828: pl. 3; reproduced here as Fig. 16A–C). These specimens are therefore syntypes; but the whereabouts of Roux’s specimens are not known. His figures, however, are relatively detailed and leave no doubt regarding the species identity.

The morphology of this species is distinct from species of Rochinia in having distinct carapace spines (Figs 16A, 17A) (versus smooth carapace in Rochinia; Fig. 1A), and buccal frame with highly protruded blunt angle (Fig. 17B) (versus lack of protruded angle on buccal frame in Rochinia; Fig. 1B).

There are studies that report the prezoea and first zoea stage morphology for A. rissoana (Guerao and Abelló 1996), as well as the size of the species at sexual maturity (Mura et al. 2005). This species has been noted to be associated with bamboo coral, Isidella elongata (Esper, 1788) (see Maynou and Cartes 2012; Cartes et al. 2013; Mastrototaro et al. 2017), and black coral, L. glaberrima (see Bo et al. 2015), and typically with the hydroid epibiont, Rosalinda incrustans (Kramp, 1947), on its carapace (Kramp 1947; Vervoort 1966; Mastrototaro et al. 2016). This species was also recently observed to have R. incrustans on its pseudorostral spines and climbing on colonies of I. elongata to catch small prey (Mastrototaro et al. 2017: 217, figs 5j, k, 6j–n).

A new fossil species, Lessiniamathia bolcense Ceccon & De Angeli, 2018, was recently described (see Ceccon and De Angeli 2018).

Distribution

Madère (= Madeira), Açores (= Azores), Méditerranée (= Mediterranean) (Bouvier 1922), NE Atlantic, SE Atlantic (Azores, Madeira), Mediterranean Spain, Baleares Islands, Italy, and Adriatic (Clark 1986).

Figure 18. 

Anamathia rissoana (Roux, 1828), male (26.2 × 17.5 mm) (SMF 4410), North Atlantic Ocean, left G1 and G2. A. ventral view; B. ventral view of distal portion; C. dorsal view; D. dorsal view of distal portion; E. ventral view of G2; F. dorsal view of G2. Scale bars: 1 mm.

Acknowledgements

The authors would like to thank many colleagues and friends who have helped in making important specimens available for this study: Enrique MacPherson and Pere Abelló (ICMD); Laure Corbari, Paula Martin-Lefèvre and Sébastien Soubzmaigne (MNHN); Paul Clark (NHM); Karen Reed and Rafael Lemaitre (USNM); Karen van Dorp and Charles Fransen (RMNH); Stephen Keable and Shane Ahyong (AM); Adam J. Baldinger (MCZ); Oliver Coleman (ZMB); Candice Untiedt and Elizabeth Hoensen (SAM); Bianca Trautwein and the late Professor Michael Türkay (SMF); Tammy Horton (National Oceanography Centre, University of Southampton); and J.C.E. Mendoza, Mohd. Dzaki and Tan Siong Kiat (LKCNHM). Our colleagues, Helmut Zibrowius and Ferran Palero, offered interesting comments about the biogeography and paleogeography of the Alboran Sea. The authors would also like to thank the captains and crews of research cruises RRS Discovery D92 and RRS Challenger CH9/79, as well as the long-term curation of specimens by the Discovery Collections, National Oceanography Centre, Southampton, U.K. that was funded by the UK Natural Environment Research Council (NERC). For discussions, we thank Marcos Tavares for his many helpful ideas, and the late Professor Michael Türkay for his comments on S. hertwigi. The drawings from Figs 7 and 8A–F are from Jean- Jean-François Dejouannet. We are also grateful to Cédric d'Udekem d'Acoz (Institut Royal des Sciences naturelles) for help with hard to get literature. Lastly, we are grateful to the editor, Sammy De Grave, and the reviewers, Marcos Tavares and Shane Ahyong, for their comments and suggestions which helped improve this manuscript.

References

  • Abele LG, Kim W (1986) An illustrated guide to the marine decapod crustaceans of Florida. Department of Environmental Regulation, State of Florida, Technical Series, 8(1), xvii + 225 pp.
  • Abelló P, Carbonell A, Torres P (2002) Biogeography of epibenthic crustaceans on the shelf and upper slope off the Iberian Peninsula Mediterranean coasts: implications for the establishment of natural management areas. In: Abelló P, Bertrand JA, Gil de Sola L, Papaconstantinou C, Relini G, Souplet A (Eds) Mediterranean marine demersal resourves: the medits international trawl survey (1994–1999). Scientia Marina 66 (supplement 2), 183–198. https://doi.org/10.3989/scimar.2002.66s2183
  • Alcock A (1895) Materials for a carcinological fauna of India. No. 1. The Brachyura Oxyrhyncha. Journal of the Asiatic Society of Bengal 44 (part 2, no. 2): 157–291. [pls. 3–5] https://doi.org/10.5962/bhl.title.16033
  • Allen JA (1967) Crustacea: Euphausiacea and Decapoda with an illustrated key to the Bristish species. The Fauna of the Clyde Sea Area, Bell and Bain Ltd., Glasgow, 1–116 pp.
  • Alvarado JJ, Solís-Marín FA, Ahearn C (2008) Equinodermos (Echinodermata) del Caribe Centroamericano. Revista de Biología Tropical 56(3): 37–55.
  • Anonymous (1881) Reports on the results of dredging, under the supervision of A. Agassiz, in the Caribbean Sea, 1878–79, along the Atlantic coast of the United States, during the summer of 1880, by the United States Coast Survey steamer ‘Blake,’ Commander J.E. Bartlett, U.S.N., commanding. Two reports have recently been published (Dec., 1880) in vol. viii of the Bulletin of the Museum of Comparative Zoölogy of Harvard College: Études préliminaires sur les Crustacés, par M. Alph. Milne-Edwards, 1re partie, pp. 1–68, with 2 plates; the other, A Preliminary Report on the Echini, by A. Agassiz, pp. 69–84. American Journal of Science (series 3) 21 (124), 338.
  • Araújo R, Biscoito M, González JA (2014) New records of decapod crustaceans from off the Archipelago of Madeira (Northeastern Atlantic). Boletim do Museu de História Natural do Funchal 64(339): 35–41.
  • Barnard KH (1950) Descriptive Catalogue of South African Decapod Crustacea (Crabs and Shrimps). Annals of the South African Museum 38: 1–837.
  • Bo M, Bavestrello G, Angiolillo M, Calcagnile L, Canese S, Cannas R, Cau A, D’Elia M, D’Oriano F, Follesa MC, Quarta G, Cau A (2015) Persistence of pristine deep-sea coral gardens in the Mediterranean Sea (SW Sardinia). PLoS ONE 10 (3): e0119393 [1–21]. https://doi.org/10.1371/journal.pone.0119393
  • Bourne GC (1890) Report of a trawling cruise in H.M.S. “Research” off the South-west coast of Ireland. Journal of the Marine Biological Association of the United Kingdom 1(3): 306–323. https://doi.org/10.1017/S0025315400072167
  • Bouvier E-L (1922) Observations complémentaires sur les Crustacés décapodes (Abstraction faite des Carides) provenant des Campagnes de S.A.S. le Prince de Monaco. In: Résultats des campagnes scientifiques accomplies sur son yacht par Albert Ier, Prince Souverain de Monaco 62: 106 + [1] pp. [pls. 1–4] [In French]
  • Bouvier E-L (1940) Décapodes Marcheurs. Faune de France 37: 1–404. [pls. 1–14]
  • Bullis HR, Thompson JR (1965) Collections by the Exploratory Fishing Vessels Oregon, Silver Bay, Combat, and Pelican made during 1956 to 1960 in the Southwestern North Atlantic. United States Fish and Wildlife Service, Special Scientific Report, Fisheries 510: 1–130.
  • Carpenter WB, Gwyn Jeffreys J, Thomson CW (1870) Preliminary report of the scientific exploration of the deep sea in H.M. Surveying-vessel ‘Porcupine,’ during the summer of 1869. Proceedings of the Royal Society of London 18: 397–492. https://doi.org/10.1098/rspl.1869.0084
  • Cartes JE, LoIacono C, Mamouridis V, López-Pérez C, Rodríguez P (2013) Geomorphological, trophic and human influences on the bamboo coral Isidella elongate assemblages in the deep Mediterranean: To what extent does Isidella form habitat for fish and invertebrates? Deep-Sea Research I 76: 52–65. https://doi.org/10.1016/j.dsr.2013.01.006
  • Ceccon L, De Angeli A (2018) Lessiniamathia bolcense n. gen., n. sp., a new Epialtidae crustacean from the early Eocene of the eastern Lessini Mounths (Verona, NE Italy). Lavori-Società Veneziana di Scienze Naturali 43: 147–154. [in Italian]
  • Christiansen ME (1969) Crustacea Decapoda Brachyura. Marine Invertebrates of Scandinacia number 2. Universitetsforlaget Oslo, Denmark, 143 pp.
  • Chun C (1900) Aus den Tiefen des Weltmeeres. Schilderungen von der Deutschen Tiefsee-Expedition. Mit 6 Chromolithographieen, 8 Heliogravüren, 32 als Tafeln gedruckten Vollbildern, 2 Karten und 390 Abbildungen im Text. Gustav Fischer, Jena, vi + 549 pp. [46 pls., 2 maps]
  • Chun C (1902–1903) Aus den Tiefen des Weltmeeres. Schilderungen von der Deutschen Tiefsee-Expedition. Mit 6 Chromolithographien, 8 Heliogravüren, 32 als Tafeln gedruckten Vollbildern, 3 Karten und 482 Abbildungen im Text. Zweite umgearbeitete und stark vermehrte Auflage. Gustav Fischer, Jena, ix + 592 pp. [46 pls., 3 maps]
  • Conan G, Roux M, Sibuet M (1981) A photographic survey of a population of the stalked crinoid Diplocrinus (Annacrinus) wyvillethomsoni (Echinodermata) from the bathyal slope of the Bay of Biscay. Deep-Sea Research 28A(5): 441–453. https://doi.org/10.1016/0198-0149(81)90136-9
  • D’Udekem d’Acoz C (1999) Inventaire et distribution des crustacés décapodes de l’Atlantique nord-oriental, de la Méditerranée et des eaux continentales adjacentes au nord de 25°N. Patrimoines naturels, Muséum national d’Histoire naturelle, Paris 40: i–x, 1–394.
  • Dana JD (1851) Conspectus Crustaceorum quae in Orbis Terrarum circumnavigatione, Carolo Wilkes e Classe Reipublicae Foederatae Duce. Pars VI. American Journal of Science and Arts (Series 2) 11(32): 268–274.
  • Davie PJF (2002) Crustacea: Malacostraca: Eucarida (Part 2): DecapodaAnomura, Brachyura. In: Wells A, Houston WWK (Eds) Zoological Catalogues of Australia. Volume 19.3B. CSIRO Publishing, Melbourne, Australia, xi + 641 pp.
  • De Haan W (1833–1850) Crustacea. In: Siebold PF von (Ed.) Fauna Japonica sive Descriptio Animalium, quae in Itinere per Japoniam, Jussu et Auspiciis Superiorum, qui Summum in India Batava Imperium Tenent, Suscepto, Annis 1823–1830 Collegit, Notis, Observationibus et Adumbrationibus Illustravit, xvi + xxi + xvii + 243 pp. [72 pls] [For dates of publication, see Holthuis (1953)]
  • De Melo GAS (1996) Manual de Identificação dos Brachyura (Caranguejos e Siris) do Litoral Brasileiro. Plêiade/FAPESP, São Paulo, Brazil, 603 pp.
  • De Mol B, Van Rensbergen P, Pillen S, Van Herreweghe K, Van Rooij D, McDonnell A, Huvenne V, Ivanov M, Swennen R, Henriet JP (2002) Large deep-water coral banks in the Porcupine Basin, southwest of Ireland. Marine Geology 188: 193–231. https://doi.org/10.1016/S0025-3227(02)00281-5
  • Dieuzeide R (1950) La faune du fond chalutable de la Baie de Castiglione. Bulletin des Travaux Publiés par la Station d’Aquiculture et de Pêche de Castiglione (Nouvelle Série 2): 11–86.
  • Dieuzeide R (1955) Recherches sur les fonds chalutables de la region d’Alger. I. Introduction: Draguages et chalutages. Notes faunistiques sur la zone meso-abyssale. Bulletin des Travaux Publiés par la Station d’Aquiculture et de Pêche de Castiglione (Nouvelle Série 7): 9–86.
  • Dieuzeide R (1960) Le fond chalutable à 600 mètres par le travers de Castiglione. Le faciès à Isidella elongata Esper. Bulletin des Travaux Publiés par la Station d’Aquiculture et de Pêche de Castiglione (Nouvelle Série 10): 61–106. [+ pl. 1]
  • Dieuzeide R, Roland J (1958) Prospections des fonds chalutables de côtes algériennes. Recherches de Nouvelles zones (Années 1956–1957). Bulletin des Travaux Publiés par la Station d’Aquiculture et de Pêche de Castiglione (Nouvelle Série 9): 11–69.
  • Doflein F (1904) Brachyura. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem Dampfer “Valdivia” 1898–1899. Gustav Fischer, Jena, volume 6: 1–6, i–xiv, 1–314. [figs 1–58]
  • Esper EJC (1788–1830) Die Pflanzenthiere in Abbildungen nach der Natur mit Farben erleuchtet nebst Beschreibungen. [Theilen 1–3 (Lieferungen 1–17)]. Raspe, Nürnberg. [completed by F.L. Hammer after 1809; for full collation, see Ott (1989, 1995) and Grasshoff and Scheer (1991).]
  • Faxon W (1895) The Stalk-Eyed Crustacea. In: Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands, in charge of Alexander Agassiz, by the U.S. Fish Commission steamer “Albatross,” during 1891, Lieut.-Commander Z. L. Tanner, U.S.N. commanding. XV. Memoirs of the Museum of Comparative Zoölogy at Harvard College XVIII: 1–292. [pls. A–K + I–LVII] [Apr.1895].
  • Felder DL, Álvarez F, Goy JW, Lemaitre R (2009) Decapoda (Crustacea) of the Gulf of Mexico, with comments on the Amphionidacea. In: Felder DL, Camp DK (Eds) Gulf of Mexico–Origins, Waters, and Biota. Biodiversity. Texas A&M University Press, College Station, Texas, 1019–1104.
  • Filhol H (1885) La vie au fond des mers. Les explorations sous-marines et les voyages du Travailleur et du Talisman. G. Masson, Paris, viii + 303 pp. [pls. 1–8]
  • Fransen CHJM (1991) Preliminary report on Crustacea collected in the eastern part of the North Atlantic during the CANCAP and MAURITANIA Expeditions of the former Rijksmuseum van Natuurlijke Historie, Leiden. Nationaal Natuurhistorisch Museum, Leiden, vi + 200 pp.
  • Gan Z, Li X (2019) Report on four deep-water barnacles (Cirripedia, Thoracica) from the north west Pacific, with remarks on Trianguloscalpellum regium (Wyville-Thomson, 1873). Zootaxa 4565(2): 201–212. https://doi.org/10.11646/zootaxa.4565.2.4
  • García Raso JE (1996) Crustacea Decapoda (excl. Sergestidae) from Ibero-Moroccan waters. Results of Balgim-84 Expedition. Bulletin of Marine Science 58(3): 730–752.
  • Garth JS (1958) Brachyura of the Pacific coast of America: Oxyrhyncha. Allan Hancock Pacific Expeditions 21 (1–2): i–xii + 1–874, figs 1–9, pls. A–Z + Z1–Z4 + 1–55, tables 1–107. [25 Jul.1958] [Note: Part 1 contains pp. i–xii, 1–499, and figs 1–9. Part 2 contains pp. 500–854, pls. A–Z + Z1–Z4 + 1–55, and tables 1–107).]
  • González-Gordillo JI, Dos Santos A, Rodríguez A (2001) Checklist and annotated bibliography of decapod crustacean larvae from the Southwestern European coast (Gibraltar Strait area). Scientia Marina 65(4): 275–305. https://doi.org/10.3989/scimar.2001.65n4275
  • Grieg JA (1927) Decapoda Crustacea from the West Coast of Norway and the North Atlantic. Bergens Museums Aarbok Naturvidenskabelig række 7: 1–53.
  • Griffin DJG, Tranter HA (1986) The Decapoda Brachyura of the Siboga expedition. Part VIII: Majidae. Siboga Expeditie Monografie 39: [i]–[vii], 1–335.
  • Griffiths CL, Landschoff J, Atkinson LJ (2018) Phylum Arthropoda. In: Atkinson LJ, Sink KJ (Eds) Field Guide to the Offshore Marine Invertebrates of South Africa. Malachite Marketing and Media, Pretoria, 133–226.
  • Guerao G, Abelló P (1996) Morphology of the prezoea and first zoea of the deep-sea spider crab Anamathia rissoana (Brachyura, Majidae, Pisinae) Scientia Marina 60 (2–3): 245–251.
  • Hansen HJ (1908) Crustacea Malacostraca. I. Danish Ingolf-Expedition 3(2): 1–120. [pls. 1–5]
  • Heller C (1863) Die Crustaceen des südlichen Europa. Crustacea Podophthalmia. Mit einer Übersicht über die horizontale Verbreitung sämmtlicher europäischer Arten. Wien: Wilhem Braumüller, 336 pp. https://doi.org/10.5962/bhl.title.13110
  • Holthuis LB (1962) Forty-seven genera of Decapoda (Crustacea); Proposed addition to the official list. Z.N.(S.) 1499. Bulletin of Zoological Nomenclature 19(4): 232–253.
  • Holthuis LB (1977) The Mediterranean decapod and stomatopod Crustacea in A. Risso’s published works and manuscripts. Annales du Muséum d’Histoire Naturelle de Nice Tome 5: 37–88.
  • Ingle RW (1979) The larval development of the spider crab Rochinia carpenteri (Thomson) [Oxyrhyncha: Majidae] with a review of majid subfamilial larval features. Bulletin of British Museum of Natural History (Zoology) 37(1): 47–66.
  • Ingle RW (1980) British Crabs. British Museum (Natural History), London, [i] + 222 pp.
  • Köhler R (1896) Poissons. Résultats scientifiques de la campagne du “cudan” dans le golfe de Gascogne, Août-Septembre, 1895. Annales de l’Université de Lyon (volume 26): 475–526. [2 pls.] https://doi.org/10.5962/bhl.title.65730
  • Kramp PL (1947) Hydroids collected by the “Skagerak” Expedition in the eastern Atlantic 1946. Göteborgs Kungliga Vetenskaps- och Vitterhets-Samhälles Handlingar 6. Följden Serie B 5(8): 1–16.
  • Lagerberg T (1905) Anomura und Brachyura der schwedischen Südpolar-Expedition. Wissenschafteliche Ergebnisse der Schwedischen Südpolar-Expedition (1901–1903) 5(7): 1–39. [pl. 1]
  • Lamouroux JVF (1812) Extrait d’un mémoire sur la classification des polypiers coralligènes non entièrement pierreux. Nouveau Bulletin des Sciences, par la Société Philomatique, Paris 63: 181–188.
  • Lee BY, Richer de Forges B, Ng PKL (2017) Deep-sea spider crabs of the families Epialtidae MacLeay, 1838 and Inachidae MacLeay, 1838, from the South China Sea, with description of two new species (Decapoda, Brachyura, Majoidea). European Journal of Taxonomy 358: 1–37. https://doi.org/10.5852/ejt.2017.358
  • Lee BY, Richer de Forges B, Ng PKL (2019) Deep-sea spider crabs of the families Epialtidae MacLeay, 1838, from Papua New Guinea, with a redefinition of Tunepugettia Ng, Komai, Sato, 2017, and descriptions of two new genera (Crustacea: Decapoda: Brachyura: Majoidea). Zootaxa 4619(1): 001–044. https://doi.org/10.11646/zootaxa.4619.1.1
  • Low MEY, Lee BY, Ng PKL (2017) The Brachyura (Crustacea: Decapoda) of the Deutschen Tiefsee-Expedition auf dem Dampfer ‘Valdivia’ 1898–1899: dates of publications and authorship. Zootaxa 4341(1): 113–123. https://doi.org/10.11646/zootaxa.4341.1.10
  • MacLeay WS (1838) On the brachyurous decapod Crustacea. Brought from the Cape by Dr. Smith. In: Smith A (Ed.) Illustrations of the Zoology of South Africa; consisting chiefly of Figures and Descriptions of the Objects of Natural History Collected During an Expedition into the Interior of South Africa, in the Years 1834, 1835, and 1836; Fitted Out by ‘The Cape of Good Hope Association for Exploring Central Africa’. Published under the Authority of the Lords Commissioners of Her Majesty’s Treasury, London, i–iv + 53–71. [2 pls.]
  • MacPherson E (1983) Crustáceos Decápodos capturados en las costas de Namibia. Resultados Expeditiones Científicas 11: 3–79.
  • Maenosono T, Naruse T (2016) New records of two sesarmid crabs (Crustacea: Decapoda: Brachyura) from Ishigakijima Island, Ryukyu Archipelago, Japan. Fauna Ryukyuana 28: 5–22.
  • Marco-Herrero E, Abelló P, Drake P, García-Raso JE, González-Gordillo JI, Guerao G, Palero F, Cuesta JA (2015) Annotated checklist of brachyuran crabs (Crustacea: Decapoda) of the Iberian Peninsula (SW Europe). Scientia Marina 79(2): 243–256. https://doi.org/10.3989/scimar.04161.27A
  • Mastrototaro F, Aguilar R, Chimienti G, Gravili C, Boero F (2016) The rediscovery of Rosalinda incrustans (Cnidaria: Hydrozoa) in the Mediterranean Sea. Italian Journal of Zoology 83(2): 244–247. https://doi.org/10.1080/11250003.2016.1181800
  • Mastrototaro F, Chimienti G, Acosta J, Blanco J, Garcia S, Rivera J, Aguilar R (2017) Isidella elongate (Cnidaria: Alcyonacea) facies in the western Mediterranean Sea: visual surveys and descriptions of its ecological role. The European Zoological Journal 84(1): 208–225. https://doi.org/10.1080/24750263.2017.1315745
  • Maynou F, Cartes JE (2012) Effects of trawling on fish and invertebrates from deep-sea coral facies of Isidella elongate in the western Mediterranean. Journal of Marine Biological Association of the United Kingdom 92(7), 1501–1507. https://doi.org/10.1017/S0025315411001603
  • McLaughlin PA, Camp DK, Angel MV, Bousfield EL, Brunel P, Brusca RC, Cadien D, Cohen AC, Conlan K, Eldredge LG, Felder DL, Goy JW, Haney T, Hann B, Heard RW, Hendrycks EA, Hobbs HH, Holsinger JR, Kensley B, Laubitz DR, LeCroy SE, Lemaitre R, Maddocks RF, Martin JW, Mikkelsen P, Nelson E, Newman WA, Overstreet RM, Poly WJ, Price WW, Reid JW, Robertson A, Rogers DC, Ross A, Schotte M, Schram FR, Shih C-T, Watling L, Wilson GDF, Turgeon DD (2005) Common and Scientific Names of Aquatic Invertebrates from the United States and Canada: Crustaceans. American Fisheries Society, Special Publication, Volume 31. Bethesda, Maryland, 1–533.
  • Miers EJ (1886) Report on the Brachyura collected by H.M.S. Challenger during the years 1873–1876. In: Thomson CW, Murray J (Eds) Report on the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873–1876 under the command of Captain George S. Nares, R.N., F.R.S. and the late Captain Frank Tourle Thomson, R.N. prepared under the Superintendence of the late Sir C. Wyville Thomson, Knt., F.R.S. &c. Regius Professor of Natural History in the University of Edinburgh Director of the civilian scientific staff on board and now of John Murray, LL.D., Ph.D., &c. one of the naturalists of the Expedition. Zoology. Published by Order of Her Majesty’s Government, London, Edinburgh and Dublin, volume 17(49): l [= 50] + 362 pp. [pls. 1–29]
  • Milne-Edwards A (1873–1880) Études sur les xiphosures et les crustacés de la région mexicaine. Mission scientifique au Mexique et dans l’Amérique centrale, ouvrage publié par ordre du Ministre de l’Instruction publique. Recherches Zoologiques pour servir à l’histoire de la faune de l’Amérique centrale et du Mexique, publiées sous la direction de M. H. Milne Edwards, membre de l’Institut. Imprimerie nationale, Paris. Cinquième partie. Tome premier. [8 unpaginated] + 1–368 + [63 unpaginated plate legends] pp., pls. 1–61 + 5A + 31A [= 63 plates]. {Dates of publication—Livraison 1: [6 unpaginated] + 1–24 pp., pls. 1–7: 20 Sep.1873; Livraison 2: 25–56 pp., pls. 8–14: 28 Oct.1873; Livraison 3: 57–120 pp., pls. 15–20: 4 Dec.1875; Livraison 4: 121–184 pp., pls. 21–27, 29, 30: 1878; Livraison 5: 185–224 pp., pls. 31–39: 1879; Livraison 6: 225–264 pp., pls. 40–43, 45–48, 5A: 1879; Livraison 7: 265–312 pp., pls. 31A, 44, 49–54: 1880; Livraison 8: 313–368 + [8 unpaginated] pp., pls. 55–61: 1880. For dates of publication, see Monod 1956: 642, Manning and Holthuis 1981: 368, and Crosnier and Clark 1998: 93. Though there appears to be some disagreement between the dates in these publications, we defer to Crosnier and Clark 1998.}
  • Milne-Edwards A (1880a) Compte rendu sommaire d’une exploration zoologique faite dans le golfe de Gascogne à bord du navire de l’Etat le Travailleur. Comptes rendus hebdomadaires des Séances de l’Académie des Sciences, Paris 91(7): 355–360. [23 August 1880 (p. 424 of issue 8 of same volume implies that issue 7 had already been published)].
  • Milne-Edwards A (1880b) Rapport sur l’exploration scientifique du Travailleur. Le Naturaliste, Journal des échanges et des nouvelles 2(35): 275–277. [1 September 1880 (p. 273 of same issue)].
  • Milne-Edwards A (1880c) Études préliminaires sur les crustacés. Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico, and in the Caribbean Sea, 1877, ‘78, ‘79, by the United States Coast Survey Steamer “Blake,” Lieut.-Commander C.D. Sigsbee, U.S.N., and Commander J.R. Bartlett, U.S.N., commanding. VIII. Bulletin of the Museum of Comparative Zoölogy, Harvard College 8(1): 1–68. [pls. 1, 2] [December 1880 (Anonymous, 1881: 338)]
  • Milne-Edwards A, Bouvier E-L (1894) Crustacés décapodes provenant des campagnes du yacht l’Hirondelle (1886, 1887, 1888). I. Brachyures et Anomoures. Résultats des Campagnes Scientifiques accomplies sur son Yacht par Albert Ier Prince Souverain de Monaco 7: 3–112. [pls 1–11]
  • Milne-Edwards A, Bouvier E-L (1899) Crustacés décapodes provenant des campagnes de l’Hirondelle (supplément) et de la Princesse-Alice (1891–1897). Résultats des Campagnes Scientifiques accomplies sur son Yacht par Albert Ier Prince Souverain de Monaco 13: 1–106.
  • Milne-Edwards A, Bouvier E-L (1900) Crustacés décapodes. Première partie. Brachyures et Anomoures. In: Milne-Edwards A (Ed.) Expéditions scientifiques du Travailleur et du Talisman pendant les années 1880, 1881, 1882, 1883. Masson, Paris, 1–396. [32 pls] https://doi.org/10.5962/bhl.title.11668
  • Milne-Edwards A, Bouvier E-L (1923) Les Porcellanides et des Brachyures. In: Reports on the results of dredging. Under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877–78), in the Caribbean Sea (1878–79), and along the Atlantic coast of the United States (1880), by the U.S. Coast Survey steamer “Blake.” Lieut.-Com. C. D. Sigsbee, U.S.N., and Commander J.R. Bartlett, U.S.N., commanding. XLVII. Memoirs of the Museum of Comparative Zoology at Harvard College 47(4): 283–395.
  • Milne Edwards H (1834) Histoire naturelle des Crustacés, comprenant l’anatomie, la physiologie et la classification de ces animaux, Librairie Encyclopédique de Roret, volume 1, Paris, 468 pp. https://doi.org/10.5962/bhl.title.39738
  • Milne Edwards H (1836–1844) Le régne animal distribué d’après son organisation, pour servir de base à l’histoire naturelle des animaux, et d’introduction à l’anatomie comparée, par Georges Cuvier. Edition accompagnée de planches gravées, représentant les types de tous le genres, les caractères distinctifs des divers groupes et les modifications de structure sur lesquelles repose cette classification; par une réunion de disciples de Cuvier. Les Crustacés. Fortin, Masson et Cie, Paris, [vi] + [87] pp. [87 pls.]
  • Miranda v A de (1933) Ensayo de un catálogo de los crustáceos decápodos marinos de España y Marruecos Español. Notas y Resumenes. Boletín Instituto Español de Oceanografía (series 2) 67: 1–72.
  • Monod T (1956) Hippidea et Brachyura ouest-africains. Mémoires de l’Institut Français d’Afrique Noire (IFAN) 45: 1–674.
  • Mura M, Orrù F, Cau A (2005) Size at sexual maturity of the spider crab Anamathia rissoana (Decapoda: Majoidea) from the Sardinian sea. Journal of Crustacean Biology 25(1): 110–115. https://doi.org/10.1651/C-2520
  • Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum: part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology 17 (supplement): 1–286.
  • Ng PKL, Richer de Forges B (2007) A new species of deep-water spider crab of the genus Rochinia A. Milne-Edwards, 1875, from Guam (Crustacea: Brachyura: Majidae). Zootaxa 1610: 61–68. https://doi.org/10.11646/zootaxa.1610.1.5
  • Ng PKL, Richer de Forges B (2013) Samadinia longispina, a new genus and species of deep-sea spider crab from the western Pacific, and a new species of Rochinia A. Milne-Edwards, 1875, from Papua New Guinea (Crustacea: Brachyura: Majoidea: Epialtidae). Zootaxa 3718(4): 357–366. https://doi.org/10.11646/zootaxa.3718.4.5
  • Nizinski MS (2003) Annotated checklist of decapod crustaceans of Atlantic coastal and continental shelf waters of the United States. Proceedings of the Biological Society of Washington 116(1): 96–157.
  • O’Hara TD, Stöhr S, Hugall AF, Thuy B, Martynov A (2018) Morphological diagnoses of higher taxa in Ophiuroidea (Echinodermata) in support of a new classification. European Journal of Taxonomy 416: 1–35. https://doi.org/10.5852/ejt.2018.416
  • Palero F, Abelló P, Macpherson E, Beaumont M, Pascual M (2011) Effect of oceanographic barriers and overfishing on the population genetic structure of the European spiny lobster (Palinurus elephas). Biological Journal of the Linnean Society 104: 407–418. https://doi.org/10.1111/j.1095-8312.2011.01728.x
  • Pascual M, Palero F, García-Merchán VH, Macpherson E, Robainas-Barcia A, Mestres F, Roda T, Abelló P (2016) Temporal and spatial genetic differentiation in the crab Liocarcinus depurator across the Atlantic-Mediterranean transition. Scientific Reports 6, 29892.https://doi.org/10.1038/srep29892
  • Pawson DL, Pawson DJ (2013) Bathyal sea urchins of the Bahamas, with notes on covering behavior in deep sea echinoids (Echinodermata: Echinoidea). Deep-Sea Research II 92: 207–213. https://doi.org/10.1016/j.dsr2.2013.01.023
  • Perrier E (1886) Les explorations sous-marines. Hachette, Paris, 352 pp.
  • Pesta O (1918) Die Decapodenfauna der Adria. Versuch Einer Monographie, Veröffentlicht Mit UnterStützung Aus der Nowakstiftung der Kaiserlichen Akademie der Wissenschaften in Wien, x + 500 pp. https://doi.org/10.5962/bhl.title.16144
  • Pettan RB (2013) Revisão taxonômica preliminar das espécies americanas do gênero Rochinia A. Milne-Edwards, 1875 (Crustacea: Brachyura: Epialtidae). Unpublished M.Sc. thesis, Departamento de Zoologia, Instituto de Biociências da Universidade de São Paulo, 143 pp.
  • Pettan RB, Tavares M (2014) Rochinia hystrix (Stimpson, 1871) in the Southwestern Atlantic, with morphological comparison to the partially sympatric congener Rochinia tanneri (Smith, 1883) (Brachyura, Epialtidae). Crustaceana 87(3): 305–311. https://doi.org/10.1163/15685403-00003292
  • Pocock RI (1899) Crustacea. LVI. Report of a Deep-sea Trawling Cruise off the S.W. Coast of Ireland, under the Direction of Rev. W. Spotswood Green, M.A., F.R.G.S. The Annals and Magazine of Natural History (series 6) 4: 425–431. https://doi.org/10.1080/00222938909460559
  • Poupin J (1994) Faune marine profonde des Antilles françaises. Récoltes du navire Polka faites en 1993: 1–79. (ORSTOM Éditions, Collection Études et Thèses, Paris).
  • Rathbun MJ (1901) The Brachyura and Macrura of Porto Rico. Bulletin of the United States Fish Commission for 1900 2(2): 3–127.
  • Reiswig HM, Champagne P (1995) The NE Atlantic glass sponges Pheronema carpenteri (Thomson) and P. grayi Kent (Porifera: Hexactinellida) are synonyms. Zoological Journal of the Linnean Society 115: 373–384. https://doi.org/10.1111/j.1096-3642.1995.tb01430.x
  • Relini G, Peirano A, Tunesi L (1986) Osservazioni sulle comunità dei fondi strascicabili del Mar Ligure Centro-Orientale. Notes on the trawlable bottom communities in the central and eastern part of the Ligurian Sea. Bollettino dei Musei e degli Istituti Biologici dell’Università di Genova 52 (supplement): 139–161.
  • Rice AL (1990) The nomenclature of crabs collected during the cruises of HMS Porcupine in 1869 and 1870, with notes on the Norman collection of invertebrata and its acquisition by the British Museum (Natural History). Bulletin of the British Museum of Natural History (History Series) 18(1): 1–23.
  • Richer de Forges B (1995) Nouvelles récoltes et nouvelles espèces de Majidae de profondeur du genre Oxypleurodon Miers, 1886 (New findings and new species of deep-sea majids of the genus Oxypleurodon Miers, 1886). Crustaceana 68(1): 43–60. https://doi.org/10.1163/156854095X01150
  • Richer de Forges B, Ng PKL (2009) New genera, new species and new records of Indo-West Pacific spider crabs (Crustacea: Brachyura: Epialtidae: Majoidea). Zootaxa 2025: 1–20. https://doi.org/10.11646/zootaxa.2025.1.1
  • Richer de Forges B, Ng PKL (2013) On a collection of spider crabs of the genera Rochinia A. Milne-Edwards, 1875 and Naxioides A. Milne-Edwards, 1865 (Crustacea, Brachyura, Majoidea, Epialtidae) from Mozambique Channel, Solomon, Vanuatu and Philippine Islands, with description of a new species of Rochinia. In: Ahyong ST, Chan TY, Corbari L, Ng PKL (Eds) Tropical Deep-sea Benthos 27. Muséum national d’Histoire naturelle, Paris, 467–483.
  • Richer de Forges B, Poore GB (2008) Deep-sea majoid crabs of the genera Oxypleurodon and Rochinia (Crustacea: Decapoda: Brachyura: Epialtidae) mostly from the continental margin of Western Australia. Memoirs of Museum Victoria 65: 63–70. https://doi.org/10.24199/j.mmv.2008.65.6
  • Roux P (1828–1830) Crustacés de la Mediterranée et de son littoral. Levrault, Paris, [iv] + [176] pp., pls. 1–45. [Note: Pages not numbered] [Published in 9 parts. Parts 1, 2 (pls. 1–10): 1828; Part 3 (pls. 11–15): 1829; Parts 4–9 (pls. 16–45): 1830; see Manning and Holthuis (1981: 374) and Monod (1956: 646).]
  • Samouelle G (1819) The Entomologist’s Useful Compendium, or an Introduction to the Knowledge of the British Insects, Comprising the Best Means of Obtaining and Preserving Them, and a Description of the Apparatus Generally Used; Together with the Genera of Linné, and the Modern Method of Arranging the Classes Crustacea, Myriapoda, Spiders, Mites and Insects, From Their Affinities and Structure, According to the Views of Dr. Leach. Also an Explanation of the Terms used in Entomology; a Calendar of the Times of Appearance and Usual Situations of 3,000 Species of British Insects; with Instructions for Collecting and Fitting up Objects for the Microscope. Thomas Boys, London, [v] + 6–496 pp. [12 pls.] https://doi.org/10.5962/bhl.title.120094
  • Sars GO (1885) Crustacea I. Den Norske Nordhavs-Expedition (= The Norwegian North-Atlantic Expedition) 1876–1878. Zoologi 6: 1–210. [pls. 1–21, 1 map]
  • Serène R, Lohavanijaya P (1973) The Brachyura (Crustacea: Decapoda) collected by the Naga Expedition, including a review of the Homolidae. Naga Report. Scientific Results of Marine Investigations of the South China Sea and the Gulf of Thailand 4(4): 1–186. [pls. 1–21]
  • Shelton RGJ, Dooley HD (1982) New records of the geryonid crab, Geryon affinis Milne Edwards & Bouvier, and other deep water Brachyura from the North East Atlantic. Crustaceana 42(1): 108–110. https://doi.org/10.1163/156854082X00768
  • Sivertsen E, Holthuis LB (1956) Crustacea Decapoda (The Penaeidae and Stenopodidae excepted). Report on the Scientific results of the “Michael Sars” North Atlantic Deep-Sea Expedition (1910) 5(12): 1–54. [pls. 1–4]
  • Smith SI (1885) On some new or little known decapod Crustacea, from recent fish commission dredgings off the east coast of the United Sates. Proceedings of the United States National Museum VII (31): 493–496. [26 Jan.1885]. https://doi.org/10.5479/si.00963801.455.493
  • Smith SI (1886) Report on the decapod Crustacea of the Albatross dredgings off the east coast of the United States during the summer and autumn of 1884. Annual Report of the Commissioner of Fish and Fisheries for 1885. Report of the Commission on Fish and Fisheries 13: 605–698 [1–94]. [pls. 1–6]
  • Soto LA (1991) Faunal zonation of the deep-water brachyuran crabs in the Straits of Florida. Bulletin of Marine Science 49(1–2): 623–637.
  • Stebbing TRR (1902) South African Crustacea. Part II. Marine Investigations in South Africa, Cape of Good Hope, Department of Agriculture, Cape Town 2: 1–92. [pls. [Crustacea] 5–16]
  • Stebbing TRR (1905) South African Crustacea. Part III. Marine Investigations in South Africa, Cape of Good Hope, Department of Agriculture, Cape Town 4: 21–123. [pls. [Crustacea] 17–21, 21 bis, 22–26]
  • Stebbing TRR (1910) General Catalogue of South African Crustacea (Part V. of S. A. Crustacea, for the Marine Investigations in South Africa). Annals of the South African Museum 6(4): 281–593. [pls. 15–21] https://doi.org/10.5962/bhl.part.15558
  • Stimpson W (1871) Brachyura. Preliminary Report on the Crustacea dredged in the Gulf Stream in the Straits of Florida, by L. F. de Pourtales, Assist. U.S. Coast Survey. Bulletin of the Museum of Comparative Zoology at Harvard College 2(2): 109–160. [1871] [Imprint is 1 Dec.1870] [For date of publication, see Manning and Holthuis 1981: 377.] [Note: The name of the “Assistant of the U.S. Coast Survey” is spelt as “Pourtalès” in three instances in the Contents but as “Pourtales” on title pages of the relevant articles.].
  • Takeda M (1983) Crustaceans. In: Takeda M, Okutani T (Eds) Crustaceans and Mollusks trawled off Suriname and French Guiana. Japan Marine Fishery Resource Research Center, Tokyo, Japan: 19–177., 178–180 [Index in Japanese], 181–185 [Index in English].
  • Tavares MS (1991) Redéfinition des genres Rochinia A. Milne Edwards, Sphenocarcinus A. Milne Edwards et Oxypleurodon Miers, et établissement du genre Nasutocarcinus gen. nov. (Crustacea, Brachyura, Majidae). Bulletin de Muséum national d’Histoire naturelle, Section A (Zoology) (Series 4) 13 (1–2): 159–179. [figs 1–6, pl. 1] [In French with and English abstract].
  • Tavares MS, De Melo GAS (2004) Discovery of the first known benthic invasive species in the Southern Ocean: the North Atlantic spider crab Hyas araneus found in the Antarctic Peninsula. Antarctic Science 16(2): 129–131. https://doi.org/10.1017/S0954102004001877
  • Tavares MS, Santana W (2018) Refining the genus Rochinia A. Milne-Edwards, 1875: reinstatement of Scyramathia A. Milne-Edwards, 1880 and Anamathia Smith, 1885, and a new genus for Amathia crassa A. Milne-Edwards, 1879, with notes on its ontogeny (Crustacea: Brachyura: Epialtidae). Zootaxa 4418(3): 201–227. https://doi.org/10.11646/zootaxa.4418.3.1
  • Tavares MS, Santana W, Pettan R (2016) Rochinia confusa, a junior synonym of R. umbonata (Crustacea: Brachyura: Epialtidae) as revealed by ontogenetic changes. Journal of the Marine Biological Association of the United Kingdom 96(5): 1065–1071. https://doi.org/10.1017/S0025315415001587 [published online on 6 October 2015, no LSID; hardcopy printed in August 2016]
  • Taviani M, Angeletti L, Canese S, Cannas R, Cardone F, Cau A, Cau AB, Follesa MC, Marchese F, Montagna P, Tessarolo C (2017) The “Sardinian cold-water coral province” in the context of the Mediterranean coral ecosystems. Deep-Sea Research II 145: 61–78. https://doi.org/10.1016/j.dsr2.2015.12.008
  • Thomson CW (1873) The depths of the sea. An account of the general results of the dredging cruises of H.M.SS. “Porcupine” and “Lightning” during the summers of 1868, 1869, and 1870, under the scientific direction of Dr. Carpenter, F.R.S., J. Gwyn Jeffreys, F.R.S., and Dr. Wyville Thomson, F.R.S. Macmillan and Co., London: i–xxi + 1–527. [pls. 1–8] https://doi.org/10.5962/bhl.title.16330
  • Tortonese E (1964) The maine biogeographical features and problems of the Mediterranean fish fauna. COPEIA 1964(1): 98–107. https://doi.org/10.2307/1440837
  • Vervoort W (1966) Skeletal structure in the Solanderiidae and its bearing on hydroid classification. In: Rees WJ (Ed.) The Cnidaria and their evolution. Symposia of the Zoological Society of London (number 16), 373–396.
  • White A (1847a) List of Specimens of Crustacea in the Collection of the British Museum. By Order of the Trustees [of the British Museum], London, viii + 143 pp.
  • White A (1847b) Descriptions of new Crustacea from the Eastern Seas. In: April 27, 1847. William Tarrell, Esq., Vice-President, in the Chair. Proceedings of the Zoological Society of London 15(172): 56–58.
  • White A (1847c) Descriptions of new Crustacea from the Eastern Seas. In: Proceedings of Learned Societies. Zoological Society. April 13, 1847.– Wm. Yarrell, Esq., Vice-President, in the Chair. Annals and Magazine of Natural History 20(130): 61–63. https://doi.org/10.1080/037454809496014
  • Wicksten MK, Packard JM (2005) A qualitative zoogeographic analysis of decapod crustaceans of the continental slopes and abyssal plain of the Gulf of Mexico. Deep-Sea Research I 52: 1745–1765. https://doi.org/10.1016/j.dsr.2005.04.006
  • Williams AB (1984) Shrimps, Lobsters, and Crabs of the Atlantic coast of the Eastern United States, Maine to Florida. Smithsonian Institution Press, Washington D.C., xviii + 550 pp.
  • Williams AB, McCloskey LR, Gray IE (1968) New records of brachyuran decapod crustaceans from the continental shelf off North Carolina, U.S.A. Crustaceana 15(1): 41–66. https://doi.org/10.1163/156854068X00755
  • Zariquiey Alvarez R (1968) Crustáceos Decápodos Ibéricos. Investigación Pesquera, Barcelona 32: xv + 510 pp1., figs 1–164. [In Spanish].