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FIGURE 1. A in Significance of the sexual openings and supplementary structures on the phylogeny of brachyuran crabs (Crustacea, Decapoda, Brachyura), with new nomina for higher-ranked podotreme taxa

FIGURE 1. A, Anatomical relationships of penis and gonopods in Brachyura exemplified by Chionoecetes opilio (Fabricius, 1788) (Oregoniidae) (modified from Beninger et al. 1991: fig. 1C; G1 basal setae not figured); B–D, schematic representation of the dispositions of male gonopore, penis, and ejaculatory duct (indicated by dotted line) in Brachyura: B, C, perforation of P5 coxa by the ejaculatory duct: B, coxal gonopore, with penis emerging from P5 coxa (Podotremata and Heterotremata pro parte); C, coxo-sternal condition, with coxal gonopore and penis emerging at level of thoracic sternal suture 7/8 (Heterotremata pro parte); D, E, perforation of thoracic sternum by ejaculatory duct: D, sternal gonopore at the suture 7/8 level; E, sternal gonopore far from suture 7/8, in posteriormost location in relation to sternite 8. a1, abdominal somite 1; cx5, P5 coxa; g, gonopore; G1, first gonopod; G2, second gonopod; p, penis; p5, fifth pereopod; 7, 8, thoracic sternites 7, 8; 7/8, thoracic sternal suture 7/8. Scale bar: 1 mm (A).

FIGURE 14 in Significance of the sexual openings and supplementary structures on the phylogeny of brachyuran crabs (Crustacea, Decapoda, Brachyura), with new nomina for higher-ranked podotreme taxa

FIGURE 14. Derived coxo-sternal condition showing penis sheltered in gutter and location of "supplementary" suture 7/8 at complete dorsal junction of sternites 7 and 8 (both separated here to expose penis), "true" suture 7/8 situated below. Prionoplax ciliata Smith, 1870 (Panopeidae Eucratopsinae), male 13.4 × 20.3 mm, Panamá (MNHN-B24490). cx4, cx5, P4, P5 coxae; g, gutter; p, penis; pa, papilla; 7, 8, thoracic sternites 7, 8; 6/7, 7/8, thoracic sternal sutures 6/7, 7/8. Scale bar: 2 mm.

FIGURE 27. Vestigial P5 in Significance of the sexual openings and supplementary structures on the phylogeny of brachyuran crabs (Crustacea, Decapoda, Brachyura), with new nomina for higher-ranked podotreme taxa

FIGURE 27. Vestigial P5 coxa, penis, and organisation of posterior portion of cephalothorax in Hexapodidae. A, B, Hexaplax megalops Doflein, 1904, male 13.0 × 19.0 mm, Fiji (MNHN-B29154); C,?Hexapus sexpes (Fabricius, 1798), female 10.6 × 18.0 mm, Persian Gulf (MNHN-B10209): exposed, translucent apodeme prolonging into sclerotised P4 platelet (left). a, apodeme; a.p., apodemal platelet; c, coxo-sternal condyle; cx4, P4 coxa; cx5, vestigial coxa of P5; e7, episternite 7; p, penis; p4, pereopod 4; pl7, pleurite of thoracic somite 7; 7, thoracic sternite 7; 8, reduced thoracic sternite 8; 6/7, thoracic sternal suture 6/7. Scale bar: 5 mm (C).

FIGURE 6. Complex vulvae. A in Significance of the sexual openings and supplementary structures on the phylogeny of brachyuran crabs (Crustacea, Decapoda, Brachyura), with new nomina for higher-ranked podotreme taxa

FIGURE 6. Complex vulvae. A, Cavoportunus dubius (Laurie, 1906) (Portunidae Portuninae), female 18.5 × 14.3 mm, Philippines, Bohol, Panglao I. (ZRC 2001.0627) (Photographs by S.T. Nguyen & P.K.L. Ng); B–D, Episesarma lafondii (H. Jacquinot in Hombron & H. Jacquinot 1846) (Sesarmidae), female 26.9 × 30.7 mm, Western New Guinea, Kamora River (ZRC2000.2066). B, vulvae; C, D, detailed views showing sternal vulvar cover with raised posterior wall, shorter inner wall, and folded central operculum. Photographs by P. Davie, Queensland Museum. o, operculum; r, rim; v.c., sternal vulvar cover.

FIGURE 26 in Significance of the sexual openings and supplementary structures on the phylogeny of brachyuran crabs (Crustacea, Decapoda, Brachyura), with new nomina for higher-ranked podotreme taxa

FIGURE 26. Thoracotreme condition in Heloecius cordiformis (H. Milne Edwards, 1837) (Heloeciidae). A, male 15.0 × 23.0 mm, Australia, New South Wales (MNHN-B21489): male gonopore sternal, with penis emerging close to P5 coxa; B, dissected male (same data, approximately same size): P5 (left) with penis protruding from sternite 8. c, P5 coxo-sternal condyle; cx5, P5 coxa; c.p., slightly sclerotised proximal portion of penis; pa, papilla; 8, sternite 8.

FIGURE 10 in Significance of the sexual openings and supplementary structures on the phylogeny of brachyuran crabs (Crustacea, Decapoda, Brachyura), with new nomina for higher-ranked podotreme taxa

FIGURE 10. The two patterns of P5 coxa in Dromioidea, with penis either as independent, mobile tube (A, Dromiidae Dromiinae) or enclosed in extended coxa (B, Dynomenidae). A, Takedromia cristatipes (Sakai, 1969), male 19.0 × 21.5 mm, New Caledonia (MNHN-B25400): left, lowered penis; B, Dynomene hispida Latreille in Milbert 1812, male 8.8 × 11.5 mm, New Caledonia (MNHN-B22091). cx2–cx5, coxae of P2–P5; e, extension of P5 coxa; p, penis; 6–8, thoracic sternites 6–8. Scale bar: 1 mm.

FIGURE 5 in Significance of the sexual openings and supplementary structures on the phylogeny of brachyuran crabs (Crustacea, Decapoda, Brachyura), with new nomina for higher-ranked podotreme taxa

FIGURE 5. Vulvae in Hexapodidae (A–C) and Retroplumidae (D). A,?Hexapus sexpes (Fabricius, 1798), 10.6 × 18.0 mm, Persian Gulf (MNHN-B10209). B, Hexapodidae sp., female 3.0 × 4.7 mm, Gabon (MNHN-B27907). C, Thaumastoplax anomalipes Miers, 1881, female 4.0 × 6.0 mm, Congo (R.O.C.) (MNHN-B27913). D, Bathypluma forficula Saint Laurent, 1989, female 12.0 × 14.0 mm, Indonesia, Makassar Strait (MNHN-B11229). p.b., press-button; v.c., sternal vulvar cover; 4–7, thoracic sternites 4–7; 5/6, thoracic sternal sutur 5/6. Scales bars: 5 mm (A, D), 2 mm (B, C).

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FIG. 1. Putative brachyuran phylogeny, a in The occurrence, mechanics and signiŽcance of burying behaviour in crabs (Crustacea: Brachyura)

FIG. 1. Putative brachyuran phylogeny, a composite cladogram incorporating the cladograms from Spears et al. (1992) and Bellwood (1996). 5families where burying behaviour is reported.

FIGURE 53. A–C, E, F in Brachyuran crabs (Crustacea, Decapoda) from the remote oceanic Archipelago Trindade and Martin Vaz, South Atlantic Ocean

FIGURE 53. A–C, E, F, Plagusia depressa (Fabricius, 1775). A, B, Male, cl 33 mm, cw 37 mm (MZUSP 41244). C, Male, cl

FIGURE 51. A, B in Brachyuran crabs (Crustacea, Decapoda) from the remote oceanic Archipelago Trindade and Martin Vaz, South Atlantic Ocean

FIGURE 51. A, B, Planes marinus (Rathbun, 1914), juvenile female, cl 7 mm, cw 8 mm (MZUSP 40330). C, D, Planes minutus (Linnaeus, 1758), ovigerous female, cl 12.9 mm, cw 13.5 mm (MZUSP 40257). Note in (C) setae on the upper anterior margin of the propodi of the pereopods 2–5 forming a natatory fringe (arrow); natatory fringe worn in (A). Habitus, dorsal (A, C) and ventral (B, D) aspects. Photographs by A. Pol.

FIGURE 49. A–D in Brachyuran crabs (Crustacea, Decapoda) from the remote oceanic Archipelago Trindade and Martin Vaz, South Atlantic Ocean

FIGURE 49. A–D, Grapsus grapsus (Linnaeus, 1758), Trindade Island. A, C, D, May 2012. B, June 2016. Note in (A) flattened sharp-tipped spine of the adults (arrow). Photographs by J.B. Mendonça.

FIGURE 50. A–F in Brachyuran crabs (Crustacea, Decapoda) from the remote oceanic Archipelago Trindade and Martin Vaz, South Atlantic Ocean

FIGURE 50. A–F, Pachygrapsus corrugatus (von Martens, 1872). A, B, Male, cl 14.5 mm, cw 16.1 mm (MZUSP 40237). C, D, Male, cl 15.3 mm, cw 17.5 mm (MZUSP 40243). E, F, Ovigerous female, cl 11.1 mm, cw 13.3 mm (MZUSP 40351). Habitus, dorsal (A, C, E) and ventral (B, D, F) aspects. Note in (B, E) longitudinal striae on the lateral face of the cheliped propodus (arrows). Photographs by A. Pol.

FIGURE 55. A in Brachyuran crabs (Crustacea, Decapoda) from the remote oceanic Archipelago Trindade and Martin Vaz, South Atlantic Ocean

FIGURE 55. A, Map of the South Atlantic Ocean showing major oceanic islands. B, Jaccard dissimilarity dendrogram. Note that TMV-FN-RA and AS-SH clustered into two distinct groups according to the composition of their crab fauna. C, Nonmetric multidimensional scaling (NMDS) applied to a dissimilarity matrix generated by Jaccard's (1908) coefficient. Note the species that cause segregation of the crab faunal assemblages between TMV-FN-RA and AS-SH, and the gradient in species composition between the two groups of islands formed by the amphi-Atlantic species. AS, Ascension; EA, eastern Atlantic; EP, eastern Pacific; FN, Fernando de Noronha; IWP, Indo-West Pacific; RA, Rocas Atoll; SH, Saint Helena; TMV, Trindade and Martin Vaz. WA, western Atlantic.

FIGURE 48. A–F in Brachyuran crabs (Crustacea, Decapoda) from the remote oceanic Archipelago Trindade and Martin Vaz, South Atlantic Ocean

FIGURE 48. A–F, Johngarthia lagostoma (H. Milne Edwards, 1837), Trindade Island. A, Males fighting each other to access females (21 November 2014). B, Crabs mating, female on top (19 November 2014). Note both first gonopods (arrow) inserted into the female. C, Mating female in hard-shelled condition separated from male (21 November 2014). Note genital operculum (arrow). D, Ovigerous females migrating towards the beach near sunset (4 December 2017 at 6h:30 pm). E, F, Ovigerous females spawning at night (23 November 2017 at 7h:27 pm). Photographs: (A–C) by J.B. Mendonça, (D–F) by A. Pol.

FIGURE 46. A–I in Brachyuran crabs (Crustacea, Decapoda) from the remote oceanic Archipelago Trindade and Martin Vaz, South Atlantic Ocean

FIGURE 46. A–I, Johngarthia lagostoma (H. Milne Edwards, 1837), Trindade Island. A, Pico do Desejado, around 600 m high (July 2018). B, Enseada dos Portugueses. Crabs seeking for food at night near the Brazilian Navy facilitiy (November 2017). C, Retreated to its burrow during the heat hours (December 2017). D, Preying upon a newborn sea turtle at night (May 2012). E, F,

FIGURE 45. A–F in Brachyuran crabs (Crustacea, Decapoda) from the remote oceanic Archipelago Trindade and Martin Vaz, South Atlantic Ocean

FIGURE 45. A–F, Troglocarcinus corallicola Verrill, 1908. A, B, Male, cl 2.8 mm, cw 2.3 mm (MZUSP 40210). C, D, Ovigerous female, cl 2.8 mm, cw 2.3 mm (MZUSP 40217). E, Day time, in situ photograph of a free-living male on a coral (Trindade Island, April, 2014). Note crab concealing itself by matching the color of the background. F, Day time, in situ photograph of crab inside its chamber in the coral host (Trindade Island, November, 2017). Note crescent-shaped dwelling opening. Photographs: (A–D) by A. Pol, (E, F) by J.B. Mendonça.

FIGURE 52. A–F in Brachyuran crabs (Crustacea, Decapoda) from the remote oceanic Archipelago Trindade and Martin Vaz, South Atlantic Ocean

FIGURE 52. A–F, Percnon gibbesi (H. Milne Edwards, 1853). A, B, Male, cl 16.4 mm, cw 14.8 mm (MZUSP 40168). C, D, Juvenile female, cl 11.9 mm, cw 10.5 mm (MZUSP 40166). E, Ovigerous female, cl 16.1 mm, cw 14.5 mm (MZUSP 33824). Habitus, dorsal (A, C, E) and ventral (B, D) aspects. F, Percnon gibbesi (arrow) in the borehole of the urchin Diadema ascensionis Mortensen. Trindade Island, June 2012. Photographs: (A–E) by A. Pol, (F) by J.B. Mendonça.

FIGURE 44. A, D in Brachyuran crabs (Crustacea, Decapoda) from the remote oceanic Archipelago Trindade and Martin Vaz, South Atlantic Ocean

FIGURE 44. A, D, Opecarcinus hypostegus (Shaw & Hopkins, 1977), after Kropp & Manning (1987: 11, fig. 5c; 12, fig. 6f, respectively). B, E, Kroppcarcinus siderastreicola Badaró, Neves, Castro & Johnsson, 2012, after Badaro et al. (2012: 522, fig. 3A; 523, fig. 4C, respectively). C, F, G, Troglocarcinus corallicola Verrill, 1908. C, F, after Kropp & Manning (1987: 16, fig. 7d; 17, fig. 8d, respectively). G, Female, cl 4.1 mm, cw 3 mm (USNM 1207444) paratype of T. hirsutus Badaró et al., 2012 (junior synonym of T. corallicola). A–C, Female cheliped sternite. D–F, First male gonopod. G, Semi-diagrammatic outline of the posterior region of the body, dorsal view. Note in (B) anterior margin of the sternite tuberculate (arrow) and in (G) the coxa of the fifth pereopod provided with a well-developed anterior lobe (arrow).

FIGURE 47. A–G in Brachyuran crabs (Crustacea, Decapoda) from the remote oceanic Archipelago Trindade and Martin Vaz, South Atlantic Ocean

FIGURE 47. A–G, Johngarthia lagostoma (H. Milne Edwards, 1837). A, F, Male (MZUSP 9583). B–E, G, Male (MZUSP 21430). A, F, Anterior view of body and second pereopod in lateral view, respectively. B, C, Third maxilliped in ventral and dorsal view, respectively. D, E, Second (Mxp2) and first (Mxp1) maxillipeds in dorsal view, respectively. G, Distal part of the first gonopod in pleonal view. Note in (A) Mxp3 gaping noticably, exposing the mandibles; in (B) small palp and small exopodite devoid of flagellum articulating at the ventral side of the merus and basis, respectively; in (D, E) exopodite of Mxp2 and Mxp1 devoid of flagellum; in (G) corneous unguis with subdistal opening. Scales: A, F, 10 mm. B–E, 5 mm. G, 1.5 mm.

FIGURE 41. A, B in Brachyuran crabs (Crustacea, Decapoda) from the remote oceanic Archipelago Trindade and Martin Vaz, South Atlantic Ocean

FIGURE 41. A, B, Paraliomera dispar (Stimpson, 1871), habitus dorsal and ventral views, respectively, male cl 4.9 mm, cw 6.9 mm (MZUSP 41301). Note in (B) dense pilose area on the mesial side of the major chelipede (arrow). C, D, Xanthias inornatus (Rathbun 1898), habitus dorsal and ventral views, respectively, female, cl 12 mm, cw 15.5 mm (MZUSP 39692). Photographs by A. Pol.