Gibbes 1850: 189 Clibanarius tricolor Stimpson 1858: 234 Benedict 1901: 142 Schmitt 1924: 94 1935: 200 Boone 1927: 77 Provenzano 1959: 366 Coelho 1965: 309 Sánchez & Campos 1978: 25 Rieger 1998: 422 Melo 1999: 54 Coelho et al . 2007: 9 McLaughlin et al. 2010: 20 Nucci & Melo 2015: 333 Lemaitre & Tavares 2015: 453 Verrill, 1908: 450 Bouvier, 1918: 9 Forest & de Saint Laurent, 1968: 102 Coelho & Ramos 1972: 169 Coelho & Santos 1980: 143 Clibanarius tricolor Clibanarius antillensis Stimpson, 1869 Clibanarius tricolor ( Gibbes, 1850 ) Clibanarius aff. tricolor ( Gibbes, 1850 ) Paguroids (Decapoda: Anomura: Diogenidae and Paguridae) of the remote oceanic Archipelago Trindade and Martin Vaz, off southeast Brazil, with new records, description of three new species and zoogeographical notes Lima, Daniel Tavares, Marcos Jr, Joel Braga De Mendonça Zootaxa 2019 2019-11-05 4694 1 1 63  Nucci & Melo 2015: 333  fe04ce3e-2749-4d42-8323-ecd6f94d8fe9 (Gibbes, 1850) Lima & Tavares & Jr 2019 Gibbes 1850 [151,629,903,930] Malacostraca Diogenidae Clibanarius Animalia Decapoda 10 11 Arthropoda species tricolor  ( Fig. 4A–C)     Pagurus tricolor  Gibbes 1850: 189.    Clibanarius tricolor.—  Stimpson 1858: 234;  Benedict 1901: 142, pl. 6, fig. 2; Verril 1908: 447, figs. 61–63 [in part];  Schmitt 1924: 94;  1935: 200, fig. 61; 1939: 28;  Boone 1927: 77;  Provenzano 1959: 366, fig. 5a; 1960: 119; 1961: 153;  Coelho 1965: 309; 1969: tab. 1;  Sánchez & Campos 1978: 25, fig. 6; Coelho & Ramos-Porto: 1986: 51;  Rieger 1998: 422;  Melo 1999: 54, fig. 12 a–d;  Coelho et al. 2007: 9, tab. 4;  McLaughlin et al.2010: 20;  Nucci & Melo 2015: 333, figs. 2c, 6a. 25, fig. 6;  Lemaitre & Tavares 2015: 453, tab. 1.    Clibanarius hebes  Verrill, 1908: 450, figs. 65, 66 [in part].    Clibanarius brachyops  Bouvier, 1918: 9, fig. 3.    Clibanarius tricolor tricolor  Forest & de Saint Laurent, 1968: 102, fig. 61, tab. II;  Coelho & Ramos 1972: 169;  Coelho & Santos 1980: 143.   Clibanarius tricolor.— Rieger 1997: 114; Masunari et al. 1998: 224, 225; Nucci & Melo 2015: 333[in part, Espirito Santoonly]; Sousa et al.2015: 341, table 2, 3. [=  Clibanarius antillensisStimpson, 1869, not  Clibanarius tricolor( Gibbes, 1850)] Trindade specimens.  Clibanariusaff. tricolor( Gibbes, 1850):  6 males, 6 females, 2 juveniles, + 88 specimens( MZUSP 36151), Brazil, Espírito Santo, Trindade Island, Crista do Galo, 20°29’22.1”S, 29°20’03.1”W, J.B. Mendonçacoll.,  12.vi.2012, tide pool; 8 males, 8 females, 7 juveniles, + 128 specimens( MZUSP 36142), idem, Crista do Galo, stn tide pool, 20°29’22.1”S, 29°20’03.1”W, J.B. Mendonçacoll.,  17.vi.2012, snorkeling,  2.5 m; 3 males, 9 females, 3 juveniles( MZUSP 36148), idem, Crista do Galo, stn tide pool, 20°29’22.1”S, 29°20’03.1”W, J.B. Mendonçacoll.,  18.vii.2015, snorkeling,  1.5 m; 5 males, 1 female, 6 juveniles, + 52 specimens( MZUSP 36149), idem, Crista do Galo, stn tide pool, 20°29’22.1”S, 29°20’03.1”W, J.B. Mendonçacoll.,  17.vi.2012;  1 male( MZUSP 36146), idem, Enseada dos Portugueses, stn Farol, 20°29’52.3”S, 29°19’15.6”W, J.B. Mendonçacoll.,  10.vii.2012,  SCUBA,  14.6 m; 21 males, 3 females( MZUSP 36156), idem, Praia da Calheta, 20°30’37.6”S, 29°18’29.1”W, J.B. Mendonçacoll.,  12.vi.2012, tide pool; 33 males, 2 females( MZUSP 36155), idem, Praia da Calheta, 20°30’37.6”S- 29°18’28.1”W, J.B. Mendonçacoll.,  11.vi.2012, tide pool; 4 males, 6 females, 1 juvenile( MZUSP 36144), idem, Praia dos Andradas, 20°30’45.7”S, 29°18’21.9”W, J.B. Mendonçacoll.,  21.vii.2013, tide pool; 3 males( MZUSP 36147), idem, Praia dos Andradas, 20°30’45.7”S, 29°18’21.9”W, J.B. Mendonçacoll.,  05.viii.2013, tide pool; 4 males, + 37 specimens( MZUSP 36141), idem, Ponta dos Andradas, 20°30’45.7”S, 29°18’21.9”W, J.B. Mendonçacoll.,  21.vii.2013, tide pool; 8 males, 2 females, 7 juveniles( MZUSP 36158), idem, Praia das Tartarugas, 20°30’54.1”S, 29°18’18.2”W, J.B. Mendonçacoll., 05.vii.  2013m tide pool and rock shore; 1 male( MZUSP 36161), idem, Parcel das Tartarugas, stn tide pool, 20°30’10.4”S, 29°17’58.3”W, J.B. Mendonçacoll.,  19.vi.2016, intertidal and snorkeling, 1.0 m; 1 male, 1 female, 2 juveniles( MZUSP 36139), idem, Praia das Tartarugas, 20°30’54.1”S, 29°18’18.2”W, J.B. Mendonçacoll.,  05.vii.2013, tide poll and rock shore; 1 juvenile( MZUSP 36159), idem, Parcel das Tartarugas, stn tide pool, 20°31’10.4”S, 29°17’58.4”W, J.B. Mendonçacoll.,  19.vi.2016, intertidal e snorkeling, 1.0 m; 1 male( MZUSP 36160), idem, Parcel das Tartarugas, stn tide pool, 20°31’10.4”S, 29°17’58.4”W, J.B. Mendonçacoll.,  19.vi.2016, intertidal and snorkeling, 1.0 m; 1 male, 2 juvenile( MZUSP 36179), idem, Praia das Tartarugas, 20°31’10.4”S, 29°17’58.4”W, J.B. Mendonçacoll.,  12.iv.2014, tide pool; 1 male, 1 female( MZUSP 36145), idem, Parcel das Tartarugas, stn Piscinal Natural, 20°31’10.4”S, 29°17’58.4”W, J.B. Mendonçacoll.,  19.vi.2016, intertidal and snorkeling, 1.0 m; 1 male( MZUSP 36154), idem, Praia das Tartarugas, 20°31’10.4”S, 29°17’54.4”W, J.B. Mendonçacoll.,  12.iv.2014, tide pool; 6 males, 6 females, 2 juveniles( MZUSP 36379), idem, Parcel das Tartarugas, stn tide pool, 20°31’10.4”S, 29°17’58.4”W, J.B. Mendonçacoll.,  19.vi.2016, intertidal and snorkeling; 3 males, 4 females, 3 juveniles, 2 megalopa ( MZUSP 36153), idem, Parcel, stn tide pool, 20°31’11.4”S, 29°18’00.6”W, J.B. Mendonçacoll.,  15.vii.2012, snorkeling; 3 males, 3 females, 6 juveniles, 2 megalopa, + 118 specimens( MZUSP 36150), idem, Parcel, stn tide pool, 20°31’11.4”S, 29°18’00.6”W, J.B. Mendonçacoll.,  15.vii.2012, snorkeling; 2 males, 5 females, 4 ovigerous females ( MZUSP 36143), idem, Praia do Lixo, 20°31’29.8”S, 29°19’43.9”W, J.B. Mendonçacoll.,  25.x.2014,  SCUBA,  13.7 m; 1 female( MZUSP 36324), idem, Enseadado Príncipe, stn Paredão, 20°31’22.4”S, 29°19’52.0”W, J.B. Mendonçacoll.,  09.vii.2013,  SCUBA,  19.5 m; 1 male( MZUSP 36152), idem, Ponta do Túnel, 20°31’33.0”S, 29°17’59.4”W, J.B. Mendonçacoll.,  11.vi.2012.   Comparative material examined.  Clibanarius tricolor( Gibbes, 1850):  2 males, 2 females( MZUSP 36967), Brazil, Pernambuco, Fernando de Noronha, east of Praia do Porto, 03°50’05.1”S, 32°24’04.1”W, L.R.L. Simonecoll.,  30.iv.2005;  2 males, 2 females( MZUSP 36968), Fernando de Noronha, west of Praia do Porto, 03°50’01.8”S, 32°24’01.1”W, L.R.L. Simonecoll.,  01.v.2005;  14 males, 8 ovigerous females, 4 females, 1 juvenile( MZUSP 36969), Fernando de Noronha, Praia do Atalaia, 03°51’22.6”S, 32°24’37.2”W, L.R.L. Simonecoll.,  04.v.2005.   Typelocality. Key West, Florida, USA.   Distribution. Bermuda, Florida, Gulf of Mexico, Antilles, Colombiaand Brazil, where it is restricted to oceanic islands so far (Rocas Atoll, Fernando de Noronha and Trindade ( Coelho 1965; Paiva et al. 2007; Nucci & Melo 2015; present study).   Remarks.The specimens of  Clibanariusfrom Trindade are herein provisionally assigned to  C. tricolor( Gibbes, 1850). The specimens from Trindade share with  C. tricolor s. str.the same pattern of transverse orange and blue bands on the articulations of P2 and P3. However, in  C. aff. tricolorfrom Trindade the meri, carpi and propodi of P2 and P3 have whitish or pale blueish backgrounds ( Fig. 4A–C), instead of the bright blue backgrounds of the adults of  C. tricolor s. str.( Fig. 4E). In adults of both,  C. tricolor s. str.and  C. aff. tricolorfrom Trindade, the dark color on the chelae usually extends to the tips of the fingers ( Fig. 4B, E), whereas in juveniles the cheliped fingers are usually lighter than the palm ( Fig. 4D; Table 2). Juveniles of  C. tricolor s. str.can have P2 and P3 with pale blueish backgrounds, cheliped palm dark, with whitish fingers ( Fig. 4D). In some species of Clibararius, such as  C. virescens( Krauss, 1843), there are considerable intraspecific differences in color between specimens from different localities or even within the same local population (e.g. Morgan 1985; Yoshikawa et al.2018). At this point, however, it is unclear whether the differences in size and color patterns between populations from northern localities and Trindade correspond with an intraspecific color variation of  C. tricolor s. str.or with a yet undescribed species. Ongoing molecular and morphological studies of populations of  C. tricolor s. l.from throughout its presumed distributional range are expected to add to our understanding of the boundaries between  C. tricolor s. str.and  C. aff. tricolorfrom Trindade.   Clibanarius tricoloris morphologically closely related to  C. antillensis Stimpson, 1859. Previous authors have found it difficult to separate between preserved specimens of  C. tricolorand  C. antillensisin which the color in life has faded away (e.g. Schmitt 1935; Provenzano 1959), and indeed we know of no realiable characters to date to distinguish between  C. tricolorand  C. antillensisother than the striking differences in color patterns. Specimens of  C. tricolorand  C. antillensiswith color in life information can be easily distinguished from each other, for instance, by the presence of a transverse orange and blue bands on the articulations of P2 and P3, Fig. 4A–C ( vs.P2 and P3 pereopods with a broad, longitudinal, light stripe on a dark background at their lateral sides in  C. antillensis, Fig. 4F).  Coelho (1965)recorded  C. tricolorfor the first time from south of the Caribbean Sea (Rocas Atoll, some 263 kmoff the coast of Brazil). A few years later, Forest & de Saint Laurent (1968)assigned 133 specimenscollected by J. Forest in the oceanic island of Fernando de Noronha (FN) to  Clibanarius tricolor(FN is located about 152 kmeast of the Rocas Atoll). Forest & de Saint Laurent (1968: 103)stressed that the specimens from FN are morphologically inseparable from  C. chapini Schmitt, 1926(known from the Cape Vert islands, and from Senegalto Angolacoastal waters): “Aucune différence morphologique ne sépare les formes est- et ouest-atlantiques [  C. chapiniand  C. tricolor, respectively] et seule une différence de coloration, minime mais constante, nous interdit de les identifier”. To the best of our knowledge, there are only a few, scattered references to the colors of the chelipeds and ambulatory legs for  C. chapini( Schmitt 1926; Forest & de Saint Laurent 1968; Manning & Chace 1990), and our attempts to obtain color photographs of  C. chapinifailed. However, the brief color description by Forest & de Saint Laurent (1968: 99, key to  Clibanariusspecies; 103, Table 2) for the FN material closely matches the color of the specimens from Trindade, but differs from  C. tricolor s. str.(see above and also Table 2).   FIGURE 4. (A–C)  Clibanariusaff. tricolor( Gibbes, 1850). (A–C) males from Trindade Island. (A) sl 2.0 mm (MZUSP 38885); (B) sl 1.7 mm (MZUSP 36160); (C) sl 1.6 mm (MZUSP 38883). (D, E) specimens from Guadalupe: aspect of juvenile and photograph of an in situspecimen, respectively. (F)  Clibanarius antillensis Stimpson, 1859from Bahia, Brazil. Photographs by A. Pol (A–C), J. Poupin and L. Corbari (D, E), and D. Lima (F).   Clibanarius rosewateri, a species being small in size (sl. ranging from 0.4–2.4 mm) so far endemic to Ascension Island, is also morphologically related to  C. tricolor s. str.and  C. chapini. However,  C. rosewateriis unique in having a distinctive distal blue spot on the mesial and lateral surfaces of the propodus of P2 and P3, a character found neither in  C. tricolor s. str.nor in  C. chapini(viz. Manning & Chace 1990), or  C. aff. tricolorfrom Trindade. It is worth noting that in the original description of  C. rosewateri, Manning & Chace (1990)involuntarily also used three times the manuscript name  Clibanarius ascensionis, which becomes a nomen nudumand hence is unavailable under the ICNZ (1999: Article 13). In the Table 1we summarize the distinctive color characters between  Clibanariusaff. tricolorfrom Trindade Island,  C. tricolor s. str.,  C. chapini, and  C. rosewateri. In addition to the oceanic island records, there are three records of  C. tricolorfrom the Brazilian continental shores. Masunari et al.(1998)recorded  C. tricolorfor the first time from the coast of Paraná, based on a single specimen caught during a 1-year survey at a rocky shore in Matinhos, with no further informations. Surprisingly,  C. antillensiswas not found during their survey, despite the species being long known from along the coast of Paraná.  Nucci & Melo (2015)referred three males and three females (two ovigerous) from Camburí beach, Espírito Santo, southeastern Brazil(MZUSP 8867) to  C. tricolor. These specimens were actually caught in April 1960by Henrique Rodrigues da Costa (see Rodrigues da Costa 1964) and were in his collections before they went to MZUSP in 1987, with no color informations. However, Rodrigues’ da Costa (1964)report on his field trip to Espírito Santoin April 1960should have given rise to caution as actually only two  Clibanariuswere recorded from Camburí, Tubarão Port:  C. sclopetarius(Herbst, 1796)and  C. antillensis. The third record of  C. tricoloris an odd one from a Brazilian mangrove under strong influence of Amazonian waters in São Marcos Bay, Maranhão. This record was based upon a single specimen taken over muddy sediment with scattered vegetation ( Sousa et al. 2015). We strongly suspect that the above records of  C. tricolor( Masunari et al. 1998; Nucci & Melo 2015; Sousa et al. 2015) from the Brazilian shores actually refer to  C. antillensisas, in absence of adequate color pattern informations, this species is easily mistaken for  C. tricolor. 2465041977 2012-06-12 MZUSP J. B. Mendonca Brazil -20.489473 Crista do Galo 1 -29.334194 Trindade Island 10 11 MZUSP 36151 102 6 2 6 Espirito Santo 2446841303 2012-06-17 MZUSP J. B. Mendonca -20.489473 Crista do Galo 1 -29.334194 10 11 MZUSP 36142 151 8 7 8 2446841524 2015-07-18 MZUSP J. B. Mendonca 3 -20.489473 Crista do Galo 1 -29.334194 10 11 MZUSP 36148 15 9 3 3 2446840933 2012-06-17 MZUSP J. B. Mendonca 2 -20.489473 Crista do Galo 1 -29.334194 10 11 MZUSP 36149 64 1 6 5 2446840952 2012-07-10 MZUSP Enseada dos Portugueses & J. B. Mendonca 2 -20.497862 Farol 1 -29.321001 10 11 MZUSP 36146 1 1 2446840541 2012-06-12 MZUSP J. B. Mendonca 15 -20.510445 Praia da Calheta 1 -29.308083 10 11 MZUSP 36156 24 3 21 2446840290 2012-06-11 MZUSP J. B. Mendonca 15 -20.510445 Praia da Calheta 1 -29.307804 10 11 MZUSP 36155 35 2 33 2446841458 2013-07-21 MZUSP J. B. Mendonca 15 -20.512695 Praia dos Andradas 1 -29.306082 10 11 MZUSP 36144 11 6 1 4 2446840993 2013-08-05 MZUSP J. B. Mendonca 15 -20.512695 Praia dos Andradas 1 -29.306082 10 11 MZUSP 36147 3 3 2446840999 2013-07-21 MZUSP Ponta dos Andradas & J. B. Mendonca 15 -20.512695 Praia dos Andradas 1 -29.306082 10 11 MZUSP 36141 41 4 2465041978 2013-07-21 MZUSP J. B. Mendonca 15 -20.515028 Praia das Tartarugas 1 -29.305056 10 11 MZUSP 36158 17 2 7 8 2465041976 2013-07-21 MZUSP J. B. Mendonca 15 -20.515028 Praia das Tartarugas 1 -29.305056 10 11 MZUSP 36158 17 2 7 8 2446841520 2013-07-05 MZUSP J. B. Mendonca 2013 11 12 -20.515028 Praia das Tartarugas 1 -29.305056 10 11 MZUSP 36139 4 1 2 1 2446841383 2016-06-19 MZUSP Parcel das Tartarugas & J. B. Mendonca 2013 -20.519556 Praia das Tartarugas 1 -29.299555 11 12 MZUSP 36159 1 1 2446841105 2016-06-19 MZUSP Parcel das Tartarugas & J. B. Mendonca 2013 -20.519556 Praia das Tartarugas 1 -29.299555 11 12 MZUSP 36160 1 1 2446840996 2014-04-12 MZUSP J. B. Mendonca 2013 -20.519556 Praia das Tartarugas 1 -29.299555 11 12 MZUSP 36179 3 2 1 2446840895 2016-06-19 MZUSP Parcel das Tartarugas & J. B. Mendonca 2013 -20.519556 Piscinal Natural 1 -29.299555 11 12 MZUSP 36145 2 1 1 2446841403 2014-04-12 MZUSP J. B. Mendonca 2013 -20.519556 Praia das Tartarugas 1 -29.298445 11 12 MZUSP 36154 1 1 2446841516 2016-06-19 MZUSP Parcel das Tartarugas & J. B. Mendonca 2013 -20.519556 Praia das Tartarugas 1 -29.299555 11 12 MZUSP 36379 14 6 2 6 2446841414 2012-07-15 MZUSP J. B. Mendonca 2013 -20.519833 Parcel 1 -29.300165 11 12 MZUSP 36153 10 4 3 3 2446841171 2012-07-15 MZUSP J. B. Mendonca 2013 -20.519833 Parcel 1 -29.300165 11 12 MZUSP 36150 130 3 6 3 2446840407 2014-10-25 MZUSP J. B. Mendonca 2013 -20.524944 Praia do Lixo 1 -29.328861 11 12 MZUSP 36143 7 5 2 2446841514 2013-07-09 MZUSP J. B. Mendonca Sao Tome and Principe 14 -20.522888 Enseada 1 -29.331112 Paredao 11 12 MZUSP 36324 1 1 2446841265 2012-06-11 MZUSP J. B. Mendonca Sao Tome and Principe 20 -20.525833 Ponta do Tunel 1 -29.299833 Paredao 11 12 MZUSP 36152 1 1 2446841421 2005-04-30 MZUSP L. R. L. Simone Brazil -3.83475 Praia do Porto 1 -32.401142 Fernando de Noronha 11 12 MZUSP 36967 4 2 2 Pernambuco 2446841519 2005-05-01 MZUSP L. R. L. Simone Brazil -3.8338332 Praia do Porto 1 -32.400307 Fernando de Noronha 11 12 MZUSP 36968 4 2 2 Pernambuco 2465041979 2005-05-04 MZUSP L. R. L. Simone Brazil -3.8562777 Praia do Atalaia 1 -32.410336 Fernando de Noronha 11 12 MZUSP 36969 27 4 1 14 Pernambuco