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FIGURE 36 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 36. Sternal penis protection in Varunidae. Varuna litterata (Fabricius, 1798), male 41.2 × 46.0 mm, New Caledonia (MNHN-B-22720): penis emerging from large orifice on thoracic sternite 8. A, invaginated papilla; B, extended papilla. o, sclerotised sheath acting as an operculum; pa, papilla. Scale bar: 1 mm.
FIGURE 53 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 53. Carrying behaviour in Dromiidae (A, B), Homolidae (C, D), and Cyclodorippidae (E). A, B, Tumidodromia dormia (Linnaeus, 1763), Mayotte, carrying sponges; C, Homola orientalis Henderson 1888, French Polynesia, with the carried sponge (detached); D, Lamoha inflata (Guinot & Richer de Forges 1981), French Polynesia, carrying a sea anemone; E, Deilocerus planus (Rathbun, 1900) carrying piece of shell (see Wicksten 1982). A, B, photographs by J. Dumas; C, D, by J. Poupin; E, sketch by M. Wicksten.
FIGURE 40 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 40. Evolution of the abdominal-locking mechanism among the Gymnopleura: ancestral condition in extinct †Palaeocorystoidea (F–I) and derived condition in extant Raninoidea of the family Lyreididae (A–E). A, Lyreidus tridentatus De Haan, 1841, ovig. female 38.0 × 23.0 mm, Australia, New South Wales (MNHN-B13316); B, L. brevifrons Sakai, 1937, male 49.4 × 23.4 mm, New Caledonia (MNHN-B24319): ventral surface, abdomen in locking position (A), abdomen removed (B); C, D, Lysirude channeri (Wood-Mason, 1885), Philippines (MNHN-B11562): sexual dimorphism of episternite 5 hook: strong teeth in male (C, 25.5 × 17.4 mm), blunt, eventually lost, and hook setose in large female (D, 33.7 × 20.5 mm); E, Lyreidus stenops Wood- Mason, 1887, male 33 × 18 mm, Philippines (MNHN-B13367): hook with 2 strong teeth; F, G, †Notopocorystes stokesii (Mantell, 1844), Middle Albian, indeterminate sex, 35 × 22 mm (overall), France (MAB k. 2870): F, ventral surface without abdomen; G, detail of double peg on episternite 5; H, I, †Eucorystes iserbyti Van Bakel, Guinot, Artal, Fraaije & Jagt, 2012, Middle Albian, indeterminate sex, height of image 6.0 mm, France, Marne (MAB k. 2871): H, thoracic sternum without abdomen; I, detail of double peg on salient episternite 5, notice the two recurved teeth. c.s., crescent-shaped suture; d.p., double peg; cx2, cx3, P2, P3 coxae; e4, e5, episternites 4, 5; g4, gynglyme for P1 on thoracic sternite 4; g5, gynglyme for P2 on thoracic sternite 5; G1, first gonopod; h, episternite 5 hook; to, tooth; 4, 5, thoracic sternites 4, 5. C–E, scanning electron micrographs. Scale bars: 100 µm (C, E), 1 mm (D). (C–E, courtesy of J.-M. Bouchard; F–I, courtesy of B. van Bakel).
FIGURE 56 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 56. Diagrammatic representation of patterns of thoracic sternal sutures 4/5–7/8 and modalities of median line (median plate not taken into account) in Eubrachyura. A, pattern 1: sutures 4/5–7/8 complete (uninterrupted), median line along sternites 5–8, may extend on sternite 4; B, pattern 2: only suture 4/5 interrupted, median line along sternites 6–8; C, pattern 3: sutures 4/5, 5/6 interrupted, median line along sternites 7, 8; D, pattern 4: sutures 4/5–6/7 interrupted, median line on sternite 8; E, generalised pattern 5: sutures 4/5–7/8 incomplete, median line variable among subpatterns a-f; F, pattern 6: sutures 4/5–7/8 interrupted, suture 6/7 complete, median line along sternites 7, 8; G, subpattern 5a: sutures 4/5–7/8 interrupted, median line absent; H, subpattern 5b: sutures 4/5–7/8 interrupted, median line along sternites 3–8; I, subpattern 5c: sutures 4/5–7/8 interrupted, median line along sternites 5–8, sometimes partially on sternite 4; J, subpattern 5d: sutures 4/5–7/8 interrupted, median line along sternites 5–7; K, subpattern 5e: sutures 4/5–7/8 interrupted, median line along sternites 7, 8; L, subpattern 5f: sutures 4/5–7/8 interrupted, median line on sternite 4. Each thoracic sternum with same width; all sutures represented parallel, equidistant, and similar. Some additional patterns are not figured.
FIGURE 31. A–D 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 31. A–D. Condylar protection: penis within P5 coxo-sternal condyle. A, Orithyia sinica (Linnaeus, 1771) (Orithyiidae), male 80.0 × 74.0 mm, North China Sea (MNHN-B11612); B, Sayamia germaini (Rathbun, 1902) (Parathelphusidae), male 35.9 × 44.0 mm, holotype of Potamon (Parathelphusa) germaini, Vietnam (MNHN-B5162): penis emerging from extremity of long condyle; C, Leurocyclus tuberculosus (H. Milne Edwards & Lucas, 1842) (Inachoididae), male 33.6 × 36.8 mm, Urugay (MNHN-B9366); D, Stenorhynchus lanceolatus (Brullé, 1837) (Inachoididae), male 34.0 × 16.0 mm, West Africa (MNHN-B21437). E. Non-condylar protection in Dilocarcinus pagei pagei Stimpson, 1861 (Trichodactylidae), male 40.0 × 50.0 mm, Bolivia (MNHN-B12813): penis, with thick sheath, emerging from oblique superior border of condyle. F. Vestigial pleopods on male abdominal somite 5 of Orithyia sinica, same data as in A. a5, a6, abdominal somites 5, 6; c, coxo-sternal condyle; cx5, coxa of P5; e7, episternite 7; G1, first gonopod; I, prominence of sternite 8; m.l., median line; p, penis; pa, papilla; t, sclerotised sheath; te, telson, v, vestigial pleopod; 6–8, thoracic sternites 6–8; 7/8, thoracic sternal suture 7/8. Scale bars: 5 mm (A, F); 1 mm (partial enlargement of A).
FIGURE 32 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 32. Coxo-sternal condition in Palicidae (A, B, E), Crossotonotidae (C, D), schematic representation. A, B, Palicus caronii (Roux, 1830), male 9.0 × 11.0 mm, Canary Is. (MNHN-B16335). A, thoracic sternum and abdomen in situ; B, detail of A. C, D, Crossotonotus spinipes (de Man, 1888), New Caledonia (MNHN-B26811): detail of sternites 7 and 8. C, male 36.0 × 44.0 mm; D, female 29.0 × 32.0 mm. E, Palicus zonatus (Rathbun, 1893), male 10.0 × 13.0 mm, Gulf of California (MNHN- B26745): ventral view, abdomen removed; invaginated sternite 8 has been partially cut to expose underlying penis (left). a1, abdominal somite 1; b, press-button; cx4, cx5, P4, P5 coxae; e3–e7, episternites 3–7; l, invagination line of sternite 8; m.l., median line; p, penis (schematically represented too thin, straight in A and B, correctly represented thick and at an angle in Fig. E); p5, pereopod 5; 6, 7, thoracic sternites 6, 7; 8, thoracic sternite 8 in the form of double plate. Scale bar: 3 mm (E).
FIGURE 33. Thoracotreme disposition. 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 33. Thoracotreme disposition. A, Plagusia squamosa (Herbst, 1790) (Plagusiidae), male cw 55.0 mm (dissected), French Polynesia (MNHN-B8889): trajectory of vas deferens directly connecting gonad to sternal gonopore. B, Ostracotheres tridacnae (Rüppell, 1830) (Pinnotheroidea), male, 9.0 × 8.6 mm, Egypt, Suez (MNHN-B10578): thoracic sternum; C, Pinnotheres pisum (Linnaeus, 1767) (Pinnotheroidea), male 4.7 × 4.6 mm, Bay of Biscay, La Rochelle (MNHN-B10604): male gonopore (B, C, after Guinot 1979a: figs. 24D, 53G respectively). cx5, P5 coxa; g, sternal gonopore; j, lateral extension of sternite 4 resulting in incomplete thoracic sternum/pterygostome junction; m.l., median line; mxp3, external maxillipeds; p, penis; t, telson; v.d., vas deferens; 3–8, thoracic sternites 3–8; 4/5–7/8, thoracic sternal sutures 4/5–7/8. Scale bar: 1 mm (A).
FIGURE 30 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 30. Coxal to coxo-sternal condition in Leucosiidae, penis enclosed inside P5 coxo-sternal condyle (abdomen removed). A, Ebalia nux A. Milne-Edwards, 1883, male, Bay of Biscay (MNHN-B20067); B, Pseudomyra mbizi Capart, 1951, male 25.0 × 20.0 mm, Congo (R.O.C.) (MNHN-B17570); C, Philyra pisum De Haan, 1841, male 21.0 × 20.5 mm, Japan (MNHN-B16982). D–F, schematic representation of A–C, portion of P5 coxa covered by episternites 7 and 8 (dotted line in C and F). a8, p8, anterior and posterior expansions of sternite 8; c, P5 coxo-sternal condyle; cx4, cx5, P4, P5 coxae; e7, e8, episternites 7, 8; g, gonopore; 7, 8, thoracic sternites 7, 8; 6/7, 7/8, thoracic sternal sutures 6/7, 7/8.
FIGURE 28 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 28. Coxal condition in Hexapodidae. A, Paeduma cylindraceum (Bell, 1859), female, 9.4 × 14.4 mm, west coast of Mexico (USNM 170897): posterior view of cephalothorax, with carapace slightly displaced to expose pleurite of thoracic somite 7 (pleurite 7). B, C, Hexaplax megalops Doflein, 1904, male 12.0 × 17.0 mm, New Caledonia (MNHN-B19244): schematic representation of posterior part of body, abdomen removed. a1–a3, abdominal somites 1–3; c, P5 coxo-sternal condyle; cx4, P4 coxa; cx5, vestigial coxa of P5; e7, episternite 7; p, penis; pl7, pleurite of thoracic somite 7; s8, exposed, visible dorsally portion of thoracic sternite 8; 6–8, thoracic sternites 6–8; 6/7, thoracic sternal suture 6/7. Scale bar: 5 mm (A).
FIGURE 29 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 29. Male gonopore (A, B) and dorsal platelets of the abdomen (C–E) in Hymenosomatidae. A, Cancrocaeca xenomorpha Ng, 1991, male 3.9 × 4.6 mm, paratype, Indonesia, Sulawesi (MNHN-B24450); B, Odiomaris pilosus (A. Milne- Edwards, 1873) (Odiomarinae), male 20.0 × 22.6 mm, New Caledonia (MNHN-B26146); C–E, same data as B: abdomen in dorsal and ventral views; notice on pleotelson (somite 6 fused to telson) the vestigial uropod as intercalated, dorsal platelet, with socket situated beneath, and uropod (D) with electron microscope (all after Guinot & Richer de Forges 1997: figs. 5B–D, 9A, B). a.c., sterno-abdominal cavity; cx5, P5 coxa; g, gonopore; G1, first gonopod; p, uropod as dorsal platelet; pt, pleotelson; s, socket; 6–8, thoracic sternites 6–8; 6/7, 7/8, thoracic sternal sutures 6/7, 7/8. Scale bar: 100µm (E).
FIGURE 25. Litocheira bispinosa Kinahan, 1856 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 25. Litocheira bispinosa Kinahan, 1856 (Litocheiridae), male 8.2 × 10.0 mm, female 8.5 × 11.0 mm, Australia, Tasmania (MNHN-B17754). A–C, G1, two different views and apex; D, G2; E, male abdomen; F, female thoracic sternum, with large vulvae; G, penis lying in depression posterior to suture 7/8; notice expanded episternite 7. cx5, P5 coxa; p, penis; v, vulva. 4–8, thoracic sternites 4–8; 4/5–7/8, thoracic sternal sutures 4/5–7/8. Scale bars: 2 mm (A–C, G); 1 mm (D); 3 mm (E); 5 mm (F).
FIGURE 22 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 22. Coxo-sternal condition in Ethusidae. Ethusina abyssicola Smith, 1884, male 12.6 × 11.0 mm, Massachusetts (MNHN-B24048): A, genital region; B, genital region with penis exposed by dissection; C, P5 coxa, penis, G1, and G2 in situ; D, G1 apex. a1, a2, abdominal somites 1, 2; c, coxo-sternal condyle; ca, carapace; cx4, cx5, P4, P5 coxae; en, endopodite; G1, G2, gonopods 1, 2; p, penis; pa, papilla; 7, 8, thoracic sternites 7, 8; 6/7, 7/8, thoracic sternal sutures 6/7, 7/8. Outline of
FIGURE 24 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 24. Coxo-sternal condition in Chasmocarcinidae. A, B, Trogloplax joliveti Guinot, 1986 (Trogloplacinae), male 20.0 × 27.0 mm, paratype, Papua New Guinea, New Britain (MNHN-B12794). A, thoracic sternum (abdomen removed); B, details of sternites 7 and 8, portion of penis (dotted line) enclosed in the "double plate". C, Chasmocarcinus typicus Rathbun, 1898 (Chasmocarcininae), male 9.0 × 12.0 mm, Brazil (MNHN-B29900): penis (proximal portion dorsally visible) enclosed in invagination of sternite 8 after perforating P5 coxa, and emerging as papilla (left); P5 coxa removed to expose ejaculatory duct (right). c, coxo-sternal condyle; cx4, cx5, P4, P5 coxae; e.d., ejaculatory duct; e.p., exposed part of penis; g.i., gutter for carapace-thoracic sternum interlocking mechanism; l, invagination line of sternite 8; m.l., median line; p, penis; pa, papilla; p.p., proximal portion of penis; s.c., sterno-abdominal cavity; 4–8, sternites 4–8; 4/5–7/8, thoracic sternal sutures 4/5–7/8. Scale bar: 5 mm (C).
FIGURE 18 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 18. Coxo-sternal condition in Dorippidae. Dorippoides facchino (Herbst, 1785), male 25.0 × 31.0 mm, India, Pondichery (MNHN-B11178): A, genital region; B, genital region with penis exposed; C, penis, G1, G2 in situ; D, G1 apex. a1, a2, abdominal somites 1, 2; b, sclerotised bulb; ba, basipodite; c, coxo-sternal condyle; ca, carapace; co, coxopodite; cx4, cx5, P4, P5 coxae; en, endopodite; e7, episternite 7; G1, G2, gonopods 1, 2; pa, papilla; td, distal portion of penial tube; tp, inclined proximal portion of penial tube; 7, 8, thoracic sternites 7, 8; 6/7, 7/8, sternal sutures 6/7, 7/8. Outline of the abdomen indicated by dotted line. Scale bars: 5 mm (A); 2 mm (B); 3 mm (C); 1 mm (D).
FIGURE 17 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 17. Coxo-sternal condition in Dorippidae. A, B, Dorippe tenuipes Chen, 1980. A, male 21.0 × 25.0 mm, South China Sea (MNHN-B8937). B, male 19.0 × 20.0 mm, holotype of Dorippe miersi Serène, 1982, Vietnam (MNHN-B7279): A, genital region covered by abdomen; B, genital region uncovered by abdomen, penis covered by sternite 8. C, Paradorippe granulata (De Haan, 1841), male 11.0 × 11.0 mm, Japan (MNHN-B11181): genital region. D, Philippidorippe philippinensis Chen, 1985, male 25.0 × 30.0 mm, Philippines (MNHN-B18913): genital region covered by abdomen. a1–a3, abdominal somites 1–3; b, sclerotised bulb; c, coxo-sternal condyle; ca, carapace; cx4, cx5, P4, P5 coxae; e7, episternite 7; pa, papilla; td, distal portion of penial tube; tp, inclined proximal portion of penial tube; 6–8, thoracic sternites 6–8; 6/7, 7/8, sternal sutures 6/7, 7/8. Scale bars: 5 mm (A, D); 3 mm (B, C).
FIGURE 13 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 13. Modalities of penis protection in Gymnopleura (A–E) and Cyclodorippiformia (F, G). A, Ranina ranina (Linnaeus, 1758), 98.0 × 89.0 mm, Réunion (MNHN-B17715): abdominal margin as dotted line; B, Raninoides laevis (Latreille, 1825), French Guiana (MNHN-B16169); C, Cyrtorhina granulosa Monod, 1956, 38.0 × 33.0 mm, Gulf of Guinea, Principe I. (MNHN-B16181); D, E, Symethis corallica Davie, 1989, 18.3 × 12.2 mm, paratype, Coral Sea, Chesterfield Is. (MNHN-B24039): dorsal (D) and ventral (E) views. F, Cymopolus asper A. Milne-Edwards, 1880, 8.5 × 6.5 mm, paratype, West Indies, Montserrat (MNHN-B13479); G, Cymonomus quadratus A. Milne-Edwards, 1880, male, cw approximately 7.0 mm (USNM). Scale bars: 5 mm (C); 2.5 mm (D, E). cx5, P5 coxa; p, penis.
FIGURE 12 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 12. Penis in Homoliformia, located in demarcated excavation, covered by row of setae. Homola ranunculus Guinot & Richer de Forges, 1995 (Homolidae), male 26.4 × 21.6 mm, New Caledonia (MNHN-B19871). cx4, cx5, P4, P5 coxae; g, gonopore; m.l., median line; p, penis; 7, 8, thoracic sternites 7, 8; 7/8, thoracic sternal suture 7/8. Scale bar: 5 mm.
FIGURE 9 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 9. Penes and two pairs of gonopods acting together as a copulatory complex in Brachyura. Callinectes sapidus Rathbun, 1896 (Portunidae, Portuninae). Copyright ©2003 Steven C. Zinski.
FIGURE 21 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 21. Coxo-sternal condition in Ethusidae. A, Ethusina talismani A. Milne-Edwards & Bouvier, 1897, male 7.0 × 6.0 mm, lectotype, Azores (MNHN-B24049): genital region (proximal portion of penis not shown). B, Ethusa rosacea A. Milne- Edwards & Bouvier, 1897, male 12.0 × 11.0 mm, syntype, off Canary Is. (MNHN-B13545): genital region. C, Ethusa dilatidens Chen, 1997, male 18.0 × 18.0 mm, Philippines (MNHN-B18930): penis with papilla protruding from funnel-like structure. a1, abdominal somite 1; c, coxo-sternal condyle; cx4, cx5, P4, P5 coxae; f, funnel; pa, papilla; t, sclerotised tube; 6– 8, thoracic sternites 6–8; 6/7, 7/8, thoracic sternal sutures 6/7, 7/8. Scale bars: 1 mm (A); 3 mm (B); 1 mm (C). 6–8, thoracic sternites 6–8
FIGURE 20 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 20. Coxo-sternal condition in Ethusidae. Ethusa mascarone (Herbst, 1785), male 11.0 × 9.8 mm, Senegal (MNHN- B20933). A, genital region; B, P5 coxa, penis, G1, G2 in situ; C, G1 apex; D, G2. a1, abdominal somite 1; ba, basipodite; c, coxo-sternal condyle; ca, carapace; co, coxopodite; cx4, cx5, P4, P5 coxae; en, endopodite; G1, G2, gonopods 1, 2; p, penis; pa, papilla; 6–8, thoracic sternites 6–8; 6/7, 7/8, thoracic sternal sutures 6/7, 7/8. Outline of the abdomen indicated by dotted line. Scale bars: 2 mm (A, B, D); 1 mm (C).
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