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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).

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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).

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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.

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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).

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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).

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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).

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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

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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).

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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).

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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).

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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.

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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.

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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.

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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

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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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FIGURE 23 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 23. Sternal male gonopores in Thoracotremata. A, Cardisoma carnifex (Herbst, 1796) (Gecarcinidae), male, Madagascar (MNHN-B29589); B, Plagusia depressa (Fabricius, 1775) (Plagusiidae), male 40.0 × 44.0 mm, Senegal (MNHN- B11693): schematic view (see Fig. 33A) (abdomen indicated by dotted line); C, Ucides cordatus (Linnaeus, 1763) (Ucididae), male 70.0 × 93.0 mm, Brazil (MNHN-B13237) (abdomen indicated by dotted line); D, Grapsus grapsus (Linnaeus, 1758) (Grapsidae), male 57.0 × 60.0 mm, Clipperton I. (MNHN-B11538): schematic representation after dissection; E, F, Ocypode cursor (Linnaeus, 1758) (Ocypodidae), male 38.0 × 49.0 m, Senegal (MNHN): E, before dissection along dotted line; F, after dissection showing ejaculatory duct perforating somite 8, instead of P5 coxa. c, P5 coxo-sternal condyle; cx5, P5 coxa; e.d., ejaculatory duct; e7, episternite 7; g, gonopore; p, penis; p5, pereopod 5; 7, 8, sternites 7, 8; 7/8, thoracic sternal suture 7/8.

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FIGURE 4 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 4. Vulvae in Goneplacidae (notice the complete thoracic sternal suture 6/7, incomplete suture 7/8, and median line line along sternites 7, 8). A, Carcinoplax longimanus (De Haan, 1833), female 29.1 × 37.8 mm, Taiwan (MNHN-B29648); B, Pycnoplax victoriensis (Rathbun, 1923), female 12.1 × 14.6 mm, Australia, Victoria (NMV J52213), remains of sperm plug; C, Thyraplax crosnieri (Guinot & Richer de Forges, 1981), female 16.7 × 21.9 mm, Philippines, Panglao I. (ZRC 2004.0720) (after Castro 2007: figs. 1, 18, 19 respectively). s.p., sperm plug; v, vulva; v.c., sternal vulvar cover; 4–8, thoracic sternites 4–8.

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FIGURE 7 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 7. Sperm plugs in Dromiidae. A, Austrodromidia octodentata (Haswell, 1882), female 44.0 × 47.0 mm, South Australia (AM P2311); B, Pseudodromia latens Stimpson, 1858, female 29.0 × 21.0 mm, South Africa (SAM A1725). g, gonopore; s, aperture of spermatheca; s.p., sperm plug; 6–8, thoracic sternites 6–8; 7/8, thoracic sternal suture 7/8. Scale bars: 5 mm (A); 2.5 mm (B).

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FIGURE 16 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 16. Coxo-sternal condition in Dorippidae. Dorippe quadridens (Fabricius, 1793), male 33.0 × 33.5 mm, Vietnam (MNHN-B11177): A, genital region; B, genital region with penis exposed (distal half of penis not shown in its normal vertical position); C, G1, penis, G2 in situ; D, G1 apex; E, G2. a1, abdominal somite 1; b, sclerotised bulb; c, coxo-sternal condyle; ca, carapace; cx4, cx5, P4, P5 coxae; G1, G2, gonopods 1, 2; en, endopodite; pa, papilla; t, sclerotised tube; 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. Scale bars: 5 mm (A–C); 1 mm (D); 2 mm (E).

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FIGURE 11 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 11. Penis in Dynomeniformia (Homolodromiidae, Dromiidae, Dynomenidae) either enclosed in extended P5 coxa (A–H) or as independent, mobile tube from P5 coxa (I–L). A, Homolodromia paradoxa A. Milne-Edwards, 1880 (Homolodromiidae), 24.0 × 15.0 mm, Bahamas (RMNH); B, Dicranodromia mahieuxii A. Milne-Edwards, 1880 (Homolodromiidae), 9.0 × 6.5 mm, holotype, Bay of Biscay (MNHN-B21681); C, Sphaerodromia lamellata Crosnier, 1994 (Dromiidae Sphaerodromiinae), 47.8 × 42.6 mm, holotype, New Caledonia (MNHN-B24724); D, S. brizops McLay & Crosnier, 1991 (Dromiidae Sphaerodromiinae), 36.4 × 36.4 mm, holotype, Seychelles (MNHN-B24560); E, Dynomene hispida (Latreille in Milbert 1812) (Dynomenidae Dynomeninae), 8.8 × 11.5 mm, New Caledonia (MNHN-B22091); F, Hirsutodynomene ursula (Stimpson, 1860) (Dynomenidae Dynomeninae), 9.6 × 11.7 mm, Galápagos Is. (MNHN-B27637); G, Metadynomene tanensis (Yokoya, 1933) (Dynomenidae Metadynomeninae), 12.5 × 13.5 mm, New Caledonia (MNHN- B25583); H, Paradynomene sp. (Dynomenidae Paradynomeninae), 11 mm width, New Caledonia (MNHN-B24780); I, Hypoconcha californiensis Bouvier, 1898 (Dromiidae Hypoconchinae), 20.3 × 19.4 mm, syntype, Gulf of California (MNHN- B22066); J, H. panamensis Smith in Verrill 1869 (Dromiidae Hypoconchinae), 20 × 21 mm, Panamá (MNHN-B21597); K, Moreiradromia antillensis (Stimpson, 1858) (Dromiidae Dromiinae), 18.0 × 18.0 mm, French Guiana (MNHN-B22030); L, Metadromia wilsoni (Fulton & Grant, 1902) (Dromiidae Dromiinae), 22.0 × 37.0 mm (MNHN-B26254). Scale bars: 1 mm (A, C, D, H–L); 0.5 mm (B, E–G).

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