Paguridae, Latreille, 1802

McLaughlin, P. A., 2003, Illustrated keys to families and genera of the superfamily Paguroidea (Crustacea: Decapoda: Anomura), with diagnoses of genera of Paguridae, Memoirs of Museum Victoria 60 (1), pp. 111-144 : 115-118

Key to genera of Paguridae

1. Gill formula includes 3 well developed or reduced pleurobranchs, 1 each on somites XI–XIII (thoracomeres 5–7, above pereopods 2–4) ( Fig. 4b).................. 2

— Gill formula includes fewer than 3 pleurobranchs ( Figs 4c, d)................................ 10

2. Pleurobranchs on somites XI and XII (thoracomeres 5 and 6, above pereopods 2 and 3) reduced, rudimentary or vestigial.................................... 3

— Pleurobranchs on somites XI and XII (thoracomeres 5 and 6, above pereopods 2 and 3) well developed........ 4

3. Chelipeds markedly unequal; female with paired gonopores on coxae of pereopod 3 ( Fig. 2m )............ Propagurus McLaughlin and de Saint Laurent, 1998

— Chelipeds subequal; female with single gonopore on coxa of left pereopod 3...... Chanopagurus Lemaitre, 2003

4. No unpaired pleopods in males; tergite of abdominal somite 6 strongly calcified...................... 5

— Some unpaired pleopods in males; tergite of abdominal somite 6 not strongly calcified................... 6

5. Chela of right cheliped with large spine at base of dactyl ( Fig. 6j ); males with paired, modified pleopods 1and 2; abdominal tergite 6 operculate ( Fig. 8d )........................... Xylopagurus A. Milne-Edwards, 1880

— Chela of right cheliped without large spine at base of dactyl; males without paired, modified pleopod 1; abdominal tergite 6 not operculate.................................... Lithopagurus Provenzano, 1968

6. Males with ( Fig. 7a ) or without at least 1 pair of modified pleopods; females with or without pleopod 1 paired, modified.................................... 7

— Males with no pleopods paired, modified; females with pleopod 1 paired, modified ( Figs 7b, c )............ 9

7. Males with pleopod 2 paired, modified.................................... Tomopaguroides Balss, 1912

— Males without pleopod 2 paired, modified.......... 8

8. Right cheliped much larger than left, with massive chela ( Fig. 6k )......... Bathypaguropsis McLaughlin, 1994

— Right cheliped only slightly larger than left, chela not massive.............. Tomopaguropsis Alcock, 1905

9. Right cheliped with dactyl opening obliquely ( Fig. 6h ); pereopod 4 semichelate ( Figs 5n, p, r, s, v ); protopods of uropods without elongate spine.......................................... Pylopaguropsis Alcock, 1905

— Right cheliped with dactyl opening horizontally ( Fig. 6f ); pereopod 4 not semichelate; protopods of uropods each with elongate spine ( Fig. 5j ).................................. Munidopagurus A. Milne-Edwards, 1880

10. Pleurobranch present on somite XII (thoracomere 7, above pereopod 4) ( Fig. 4d)......................... 11

— No pleurobranch present above pereopod 4........ 70

11. Arthrobranchs well developed on maxilliped 3 ( Figs 4a, b, d) ...................................... 12

— Arthrobranchs rudimentary, vestigial or absent on maxilliped 3 ( Fig. 4c)........................ 74

12. Gill structure distally or deeply quadriserial ( Figs 4g, h)...........................................13

— Gill structure biserial ( Fig. 4f)..................21

13. Crista dentata of maxilliped 3 with 1 or more accessory teeth ( Figs 2c, f )............................ 14

— Crista dentata of maxilliped 3 without accessory tooth ( Figs 2b, d, e ).............................. 19

14. Chelipeds subequal ( Fig. 1f, q )................. 15

— Chelipeds distinctly unequal, right largest ( Figs 1i–p )... .......................................... 17

15. Females with paired, modified pleopod 1 ( Figs 7b, c )................... Michelopagurus McLaughlin, 1997

— Females without paired, modified pleopod 1....... 16

16. Rostrum triangular; ventral margins of dactyls of ambulatory legs each with row of corneous spinules.......................... Pagurodes Henderson, 1888

— Rostrum broadly rounded; ventral margins of dactyls of ambulatory legs each with row of long stiff bristles.................... Pseudopagurodes McLaughlin, 1997

17. Males with short (1–2 coxal lengths) left sexual tube ( Figs 7m –o, q ); females with paired, modified pleopod 1 ( Figs 7b, c )........ Tarrasopagurus McLaughlin, 1997

— Males with medium (>2–5 coxal lengths) to long (>5 coxal lengths) right sexual tube ( Figs 7h, j– m ); females without paired, modified pleopod 1.................... 18

18. Male right sexual tube directed across body ventrally from right to left ( Figs 7j, k, m ); female with paired gonopores ( Fig. 2m ).............. Cestopagurus Bouvier, 1897

— Male right sexual tube directed toward exterior ( Figs 7h, i, l ); female with single left gonopore............................. .. Trichopagurus de Saint Laurent, 1968

19. Chelipeds subequal, right stronger, but not appreciable longer.. Iridopagurus de Saint Laurent-Dechancé, 1966a

— Chelipeds distinctly unequal; right usually appreciably longer.................................... 20

20. Male with very short (<1 coxal length) to short (1–2 coxal lengths) left sexual tube ( Figs 7h–o, q ); female with paired, modified pleopod 1 ( Figs 7b, c ).................... Pagurojacquesia de Saint Laurent and McLaughlin, 2000

— Male with moderate (>2–5 coxal lengths) to long (>5 coxal lengths) left sexual tube ( Fig. 7p ); female without paired, modified pleopod 1...................................... Turleania McLaughlin, 1997

21. Lateral margins of shield each developed into pair of blunt or spiniform, wing-like processes ( Fig. 3k )............................. Porcellanopagurus Filhol, 1885a

— Lateral margins of shield not developed into pair of blunt or spiniform, wing-like projections.............. 22

22. Males with very short (<1 coxal length) to long sexual tube(s) (>5 coxal lengths) ( Figs 7h–q )............ 23

— Males without sexual tube(s) ( Figs 7r, s ).......... 45

23. Females with paired, modified pleopod 1 ( Figs 7b, c ). 24

— Females without paired, modified pleopod 1....... 25

24. Carpus of right cheliped strongly produced ventrally ( Fig. 6o ); uropods asymmetrical......................................... Goreopagurus McLaughlin, 1988

— Carpus of right cheliped not strongly produced ventrally; uropods symmetrical or nearly so ( Fig. 8j )............ Pylopagurus A. Milne-Edwards and Bouvier, 1891 (part)

25. Distinct male sexual tube produced from gonopore on only 1 coxa ( Fig. 7o, p ), papilla present or absent from opposite gonopore.................................. 26

— Distinct male sexual tubes produced from gonopores on both coxae ( Figs 7h–n, q )..................... 37

26. Males with left sexual tube.................... 27

— Males with right sexual tube................... 31

27. Right chela markedly larger than left............. 28

— Right chela not markedly larger than left................................. Spiropagurus Stimpson, 1858

28. Telson with transverse indentation ( Figs 5b, c, f–i ); male with paired gonopores ( Figs 2l, m ).............. 29

— Telson without transverse indentation ( Figs 5d, e, j, k ); male sometimes without right gonopore ( Fig. 7p )......................... Micropagurus McLaughlin, 1986

29. Telson with terminal margin(s) unarmed ( Figs 5e, i, j ) 30

— Telson with terminal margin(s) armed with spines ( Figs 5d, f, g, k )............... Anapagurus Henderson, 1886

30. Telson with terminal margin entire ( Figs 5d, e ); ocular peduncles with corneas strongly dilated ( Fig. 3m ).................... Forestopagurus García-Gómez, 1994

— Telson with terminal margin marked by prominent median cleft ( Figs 5b, c, f–i ); ocular peduncles with corneas reduced......... Pygmaeopagurus McLaughlin, 1986

31. Females with paired gonopores................. 32

— Females with single gonopore on coxa of left pereopod 3 Anapagrides de Saint Laurent-Dechancé, 1966b (part)

32. Males with 3 or fewer unpaired pleopods.......... 33

— Males with 4 unpaired pleopods............................ Acanthopagurus de Saint Laurent, 1968 (part)

33. Sexual tube very short (<1 coxal length) to moderate (>2–5 coxal lengths).............................. 34

— Sexual tube long (>5 coxal lengths).............. 35

34. Rostral lobe broadly rounded; pereopod 4 with single row of corneous scales in propodal rasp ( Figs 5p, q, u ); sexual tube of moderate length, directed toward exterior..................... Catapagurus A. Milne-Edwards, 1880

— Rostral lobe triangular; pereopod 4 with 2 or more rows of corneous scales in propodal rasp ( Figs 5n, r, s, v ); sexual tube short or very short, directed anteriorly or posteriorly..... Parapagurodes McLaughlin and Haig, 1973 (part)

35. Right sexual tube directed toward exterior and upward across dorsal body surface... Hemipagurus Smith, 1881

— Right sexual tube directed toward exterior, but not over dorsal body surface ( Figs 7i, l ).................. 36

36. Sexual tube terminating in elongate filament ( Fig. 7h )... Nematopaguroides Forest and de Saint Laurent, 1968 (part)

— Sexual tube not terminating in elongate filament..................... Solenopagurus de Saint Laurent, 1968

37. Abdomen reduced ( Figs 1i, l, n–p ); males without paired or unpaired pleopods; females only with unpaired uniramous pleopods 2–4............................... 38

— Abdomen well developed ( Figs 1d–g,m ); males usually with some unpaired pleopods; females with unpaired biramous pleopods 2–4, usually also with reduced pleopod 5. .... ... ... .... ... ... .... ... ... .... ... .. 39

38. Rostrum developed as prominent slender spine; pereopod 5 subchelate ( Figs 5w, x )............................... Alainopagurus Lemaitre and McLaughlin, 1995

— Rostrum broad, blunt or subacute, upturned; pereopod 5 weakly chelate ( Fig. 5y )........................................ Alainopaguroides McLaughlin, 1997

39. Females with paired, modified pleopod 1.............. Nematopagurus A. Milne-Edwards and Bouvier, 1892

— Females without paired, modified pleopod 1....... 40

40. Right sexual tube very short (<1 coxal length) to short (1–2 coxal lengths).............................. 41

— Right sexual tube long (>5 coxal lengths)............. Nematopaguroides pusillus Forest and de Saint Laurent, 1968

41. Antennal acicles each with row of spines ( Figs 8b, c, l )....................... Alloeopagurodes Komai, 1998

— Antennal acicles without row of spines........... 42

42. Propodal rasp of pereopod 4 with 2 or more rows of corneous scales ( Figs 5r, s, v ).......................... Parapagurodes McLaughlin and Haig, 1973 (part)

— Propodal rasp of pereopod 4 with 1 row of corneous scales ( Figs 5o, p, q ).............................. 43

43. Lateral projections prominently produced; telson with rounded posterior lobes, each armed with few long, slen- der, corneous spines ( Fig. 5g )........................................ Icelopagurus McLaughlin, 1997

— Lateral projections not prominently produced; telson with obtusely subtriangular posterior lobes, each armed with few minute spinules.......................... 44

44. Coxa of right pereopod 5 in males with short sexual tube, coxa of left pereopod 5 usually without papilla; females with paired gonopores................................... Acanthopagurus de Saint Laurent, 1968 (part)

— Coxa of right pereopod 5 in males with very short sexual tube, coxa of left pereopod 5 with or without papilla; females with single left gonopore.................. Anapagrides de Saint Laurent-Dechancé, 1966 (part)

45. Females with paired, modified pleopod 1.......... 46

— Females without paired, modified pleopod 1....... 60

46. Abdomen reduced; males without unpaired pleopods; females with unpaired pleopods 2–4..................... Protoniopagurus Lemaitre and McLaughlin, 1996

— Abdomen not reduced; males with some unpaired pleopods; females with unpaired pleopods 2–5..... 47

47. Right cheliped markedly elongate....................................... Ceratopagurus Yokoya, 1933

— Right cheliped not markedly elongate............ 48

48. Protopods of uropods prominently produced posteriorly ( Fig. 5j ); dorsal surface of right chela commonly with characteristic covering of mushroom-shaped tubercles.................... Agaricochirus McLaughlin, 1981

— Protopods of uropods not prominently produced posteriorly; dorsal surface of right chela usually without characteristic covering of mushroom-shaped tubercles....... 49

49. Spines on dorsal surfaces of chelae with basal rosettes ( Fig. 6m )........... Rhodochirus McLaughlin, 1981

— Spines on dorsal surfaces of chelae without basal rosettes .......................................... 50

50. Propodal rasp of pereopod 4 with more than one row of corneous scales ( Figs 5r, s, v ).................. 51

— Propodal rasp of pereopod 4 with one row of corneous scales ( Figs 5p, q, u )......................... 53

51. Left chela triangular or subtriangular in cross-section, dactyl and fixed finger not dorsoventrally flattened.. 52

— Left chela not triangular or subtriangular in cross-section, dactyl and fixed finger dorsoventrally flattened........................ Manucomplanus McLaughlin, 1981

52. Telson with lateral indentations suggesting division into anterior and posterior portions ( Figs 5b , f-i)............................ Anisopagurus McLaughlin, 1981

— Telson without lateral indentations suggesting division into anterior and posterior portions ( Figs 5d, e ).......................... Enallopaguropsis McLaughlin, 1981

53. Ocular acicles simple ( Figs 3a, m , 8a ); coxae of male pereopods 5 symmetrical...................... 54

— Ocular acicles multispinose ( Fig 8e ); coxae of male pereopods 5 asymmetrical............................. Pylopaguridum McLaughlin and Lemaitre, 2001b

54. Telson with lateral indentations suggesting division into anterior and posterior portions ( Figs 5b, f–i )....... 55

— Telson without lateral indentations suggesting division into anterior and posterior portions ( Figs 5d, e )............................. Enallopagurus McLaughlin, 1981

55. Chela of right cheliped subovate to subcircular, margins unarmed, weakly tuberculate or minutely crenulate and/or serrate, but never armed with prominent, blunt or acute spines ( Figs 6i, o )........................... 56

— Chela of right cheliped variable, margins armed with prominent, blunt or acute spines or tubercles ( Figs 6l, m ) .......................................... 57

56. Pereopod 4 with large, very prominent preungual process at base of claw ( Fig. 5p, s, u )......................................... Phimochirus McLaughlin, 1981

— Pereopod 4 without large, very prominent preungual process at base of claw ( Figs 5o, q, r )............... Pylopagurus A. Milne-Edwards and Bouvier, 1891 (part)

57. Dactyl and fixed finger of left chela excavated ventrally, spoon-shaped................................. Tomopagurus A. Milne-Edwards and Bouvier, 1893 (part)

— Dactyl and fixed finger of left chela not excavated ventrally, not spoon-shaped.................... 58

58. Right chela circumscribed by row of dorsomesial, dorsoproximal and dorsolateral marginal spines ( Fig. 6l ); left cheliped with rotation of propodal-carpal articulation 45°- 90° from horizontal plane............................. Lophopagurus (Australeremus) McLaughlin, 1981

— Right chela not circumscribed by row of dorsomesial, dorsoproximal and dorsolateral marginal spines; left cheliped with rotation of propodal-carpal articulation much less than 45° from horizontal plane......... 59

59. Left chela with midline elevated into prominent keel or crest ( Fig. 6n )...................................... Lophopagurus (Lophopagurus) McLaughlin, 1981

— Left chela with midline sometimes elevated, but not into prominent keel or crest...... Haigia McLaughlin, 1981

60. Antennal peduncle with prominent, hooked spine at laterodistal margin of segment 1 ( Fig. 8k )........................... Tomopagurus wassi McLaughlin, 1981

— Antennal peduncle without prominent, hooked spine at laterodistal margin of segment 1.................. 61

61. Ultimate segment of antennular peduncles with very long seta provided with long paired setules ( Fig. 2i ); abdomen elongate, generally straight............................. Pagurus imafukui McLaughlin and Konishi, 1994

— Ultimate segment of antennular peduncles without very long seta provided with long paired setules; abdomen variable................................... 62

62. Coxae of male pereopod 5 asymmetrical ( Figs 7s, t ). 63

— Coxae of male pereopod 5 generally symmetrical ( Figs 2m , 7r ) .................................... 64

63. Male with 3 unpaired pleopods; coxa of right pereopod 5 produced, gonopore masked by tuft of long, stiff setae ( Fig. 7s )................... Pagurixus Melin, 1939

— Male without pleopods; coxa of left pereopod 5 produced, gonopore masked by tuft of long, stiff setae ( Fig. 7t )............................ Paguridium Forest, 1961

64. Males with both coxae of pereopod 5 produced, gonopores each masked by tuft of long, stiff setae; telson with markedly concave terminal margin, outer angles acute, with extremely prominent pair of spines adjacent to median cleft ( Fig. 5h )..................................... Diacanthurus McLaughlin and Forest, 1997

— Males without coxae of pereopods 5 produced; telson without markedly concave terminal margin, outer angles variable, without extremely prominent pair of spines adjacent to median cleft....................... 65

65. Telson with distinct transverse indentation ( Figs 5b,c, f–i ) .......................................... 66

— Telson without distinct transverse indentation ( Figs 5d, e, j )............ Discorsopagurus McLaughlin, 1974

66. Posterior portion of cephalothorax, at least in part calcified ( Fig. 1l ); abdomen reduced Labidochirus Benedict, 1892

— Posterior portion of cephalothorax membranous; abdomen well developed ( Figs 1d–g, m ).................. 67

67. Left chela with pronounced counterclockwise torsion; pereopods 4 each with prominent circular “ type A P4 structure” on lateral face of dactyl ( Fig. 5v ).................................... Elassochirus Benedict, 1892

— Left chela without pronounced counterclockwise torsion; pereopod 4 without prominent circular “ type A P4 structure” on lateral face of dactyl............... 68

68. Uropods generally asymmetrical; abdomen spirally flexed......................... Pagurus Fabricius, 1775

— Uropods generally symmetrical; abdomen not spirally flexed.................................... 69

69. Males with 3 unpaired pleopods; females with 4 unpaired pleopods.............. Orthopagurus Stevens, 1927

— Males without unpaired pleopods; females with 3 unpaired pleopods................... Paguritta Melin, 1939

70. Crista dentata with 3 or 4 very large, widely-spaced spine-like teeth ( Fig. 2g )................................... Scopaeopagurus McLaughlin and Hogarth, 1998

— Crista dentata well developed or reduced, but never with only 3 or 4 widely-spaced spine-like teeth......... 71

71. Ambulatory dactyls paddle-shaped ( Fig. 1n ); females with paired gonopores; males with right sexual tube ….................... Ostraconotus A. Milne-Edwards, 1880

— Ambulatory dactyls not paddle-shaped; females with single left gonopore; males with pair of sexual tubes... 72

72. Lateral margins of shield not drawn out into 3 prominent lobes; males with some unpaired pleopods; females without paired, modified pleopod 1.................. 73

— Shield with lateral margins each drawn out into 3 prominent lobes ( Fig. 3l ); males without unpaired pleopods; females with paired, modified pleopod 1................................ Solitariopagurus Türkay, 1986

73. Males with 3 unpaired pleopods; left sexual tube partially obscured by tufts of setae........................... Catapaguroides A. Milne-Edwards and Bouvier, 1892

— Males with 4 unpaired pleopods................................... Decaphyllus de Saint Laurent, 1968

74. Rostrum strongly deflected downward, with prominent epirostral spine ( Fig. 8i )............................................ Enneophyllus McLaughlin, 1997

— Rostrum not strongly deflected downward, without prominent epirostral spine..................... 75

75. Male sexual tube with terminal fringe of dense curved setae; no preungual process at base of claw of pereopod 4.................. Enneopagurus McLaughlin, 1997

— Male sexual tube without terminal fringe of dense curved setae; preungual process developed at base of claw of pereopod 4.......... Enneobranchus García-Gómez, 1988

Figures

Figure 1. Morphological diversity among members of the Paguroidea. a, b, Coenobitidae, c, Pylochelidae; d–g, Diogenidae; h–k, Lithodidae, l–o, Paguridae, p, q, Parapaguridae: a, Birgus latro Leach; b, Coenobita clypeatus (Fabricius); c, Trizocheles spinosus (Henderson); d, Allodardanus bredini Haig and Provenzano; e, Dardanus venosus (H. Milne Edwards); f, Clibanarius arethusa De Man; g, Calcinus tibicen (Herbst); h, Cryptolithodes sitchensis Brandt; i, Hapalogaster dentata (De Haan); j, Sculptolithodes derjugini Makarov; k, Lithodes murrayi Henderson; l, Labidochirus splendescens (Owen); m, Propagurus gaudichaudi (H. Milne Edwards); n, Ostraconotus spatulipes A. Milne–Edwards; o, Porcellanopagurus edwardsi Filhol; p, Tylaspis anomala Henderson; q, Probeebei mirabilis Boone. [a, f after Alcock, 1905; b, from Chace and Hobbs, 1969; c, k, p, from Henderson, 1888; d, e, g, after Chace et al. 1985; h, from Makarov, 1938; i, j, from Vinogradov, 1950; l from McLaughlin, 1974; m, from Benedict, 1901 as Eupagurus patagonensis Benedict; n, after A. Milne-Edwards and Bouvier, 1893; o, after Forest, 1951; q, from Wolff, 1961; not to scale.]

Figure 2. Bases and paired basis-ischium of maxilliped 3: a, Coenobitidae – Coenobita clypeatus (Fabricius); b, Diogenidae – Clibanarius vittatus (Bosc); c, Pylochelidae – Mixtopagurus paradoxus A. Milne-Edwards; d, Pylojacquesidae – Pylojacquesia colemani McLaughlin and Lemaitre; e, Parapaguridae – Parapagurus pilosimanus Smith; f, Paguridae – Pagurus pollicaris Say; g, reduced teeth on crista dentata of ischium, Scopaeopagurus megalochirus McLaughlin and Hogarth. Antennular and antennal flagella. h–j, antennular flagella: h, Coenobitidae; i, Pagurus imafukui McLaughlin and Konishi; j, generalised flagella of Diogenidae, Paguridae and Parapaguridae; k, antennal flagellum with paired ventral setae. Thoracic sternites and coxae of pereopods: l, Pylojacquesia colemani McLaughlin and Lemaitre; m, generalised Paguridae; n, Lithodes aequispinus Benedict (sternites X and XI only; groove and pit of sternite XI indicate by arrow). Abbreviations: act = accessory tooth (teeth) indicat- ed by arrows; ap = anterior portion; C 1–5 = coxae of pereopods 1–5; gp = gonopore; mh = membranous hinge; pp = posterior portion. [a–f, l, from McLaughlin and Lemaitre, 2001c; g, from McLaughlin and Hogarth, 1998; h, from McLaughlin and Dworschak, 2001; i from McLaughlin and Konishi, 1994; j, from Forest et al. 2000; k, from McLaughlin and Haig, 1996b, m, adapted from McLaughlin, 1974; not to scale]

Figure 3. Basic morphology: a, diagrammatic pagurid (whole animal, dorsal view); b, diagrammatic lithodid (whole animal, dorsal view). Cephalothorax or shield, with or without cephalic appendages: c–h Pylochelidae; i, Diogenidae; j, Pylojacquesidae; k–m Paguridae. c, Pylocheles; d, Trizocheles; e, Cheiroplatea; f, Pomatocheles; g, Parapylocheles; h, Cancellocheles; i, Diogenes; j, Pylojacquesia; k, Porcellanopagurus; l, Solitariopagurus; m, Hemipagurus. Abbreviations: aa = antennal acicle; ant. = antenna; antu = antennule; c, cornea; car = carpus; cg = cervical groove; dac = dactyl; ff = fixed finger; irp = intercalary rostral process; la = linea anomurica; lf ch = left cheliped; lf ur = left uropod; lp = lateral projection; lt = linea transversalis; mer = merus; oa = ocular acicle; op = ocular peduncle; P2–5 = pereopods 2–5; pcl = posterior carapace lobe; pcme = posterior carapace median element; pl3–5 = pleopods 3–5; plm = palm; pmp = posterior median plate; pop = postocular projection; pro = propodus; r = rostrum or rostral lobe; rt ch = right cheliped; s = shield; sl1–3 = shield lobes 1–3; t6 = abdominal tergite 6; tel = telson. [a, b, adapted from Sandberg and McLaughlin, 1998; c, d from Forest et al. 2000; e–h, from Forest, 1987; i, from McLaughlin and Clark, 1997; j, from McLaughlin and Lemaitre, 2001c; k, l, from McLaughlin, 2000; m, from McLaughlin, 1997 (as Catapagurus); not to scale.]

Figure 5. Representative telsons: a, b, Pylochelidae; c–j, Paguridae; k, Parapaguridae. Sixth abdominal tergite, protopods of uropods and telson: j, Munidopagurus. Dactyl and propodus of pereopod 4: l, o, simple; m, q, subchelate; n, r, semichelate with multiple rows of corneous scales in propodal rasp and no preungual process; p, u, t, semichelate with single row of corneous scales in propodal rasp and preungual process at base of claw; s, semichelate with multiple rows of corneous scales in propodal rasp and preungual process at base of claw; t, chelate; v, semichelate with “type A” (cf. McLaughlin, 1974) sensory structure on lateral face of dactyl. Dactyl and propodus of pereopod 5: w, x subchelate; y, semichelate; z, chelate. [a, b, n, from Forest and McLaughlin, 2000; c, from Lewinsohn, 1982; d, e, from McLaughlin, 1982; f–i from McLaughlin, 1997; j, adapted from Provenzano, 1971; k, from Lemaitre, 1996; l, o–q, s, u, y, from McLaughlin, 1997; m, from McLaughlin and Lemaitre, 1997; q, w, from McLaughlin and Lemaitre, 2001c, v, from McLaughlin, 1974; x, after Lemaitre, 1998; not to scale].

Figure 6. Chelipeds: a, left chela of Ciliopagurus (mesial view) showing stridulating mechanism (indicated by arrows); b, left chela of Allodardanus (mesial view) lacking stridulating mechanism; c, left chela and carpus of Ciliopagurus (lateral view); d, chelae of Cancellus togeth- er forming operculum; e, left carpus and chela of Aniculus; f, left chela and carpus of Isocheles (dorsal view), with dactyl opening horizontally (as indicated by arrow), g, left chela and carpus of Loxopagurus (dorsolateral view) with dactyl opening vertically (as indicated by arrow); h, right chela and carpus of Paragiopagurus (dorsal view) with dactyl opening obliquely (as indicated by arrow); i, right chela of Pylojacquesia; j, right chela of Xylopagurus; k, right chela of Bathypaguropsis; l, right chela of Lophopagurus (Australeremus); m, right chela of Rhodochirus; n, left chela of Lophopagurus (Lophopagurus); o, right carpus and chela of Goreopagurus (lateral view); p, right carpus and chela of Oedignathus (mesial view); q, right carpus and chela of Dermaturus (mesial view). [a, c, from Forest, 1952; b, after Haig and Provenzano, 1965; d, after Mayo, 1973; e, from McLaughlin and Hoover, 1995; f, g, from Forest and de Saint Laurent, 1968; h, from Lemaitre, 1996; i, from McLaughlin and Lemaitre, 2001c; j, from Lemaitre, 1995; k, from McLaughlin, 1994; l, n, from McLaughlin and Gunn, 1992; m, from Williams, 1984; o, from McLaughlin and Haig, 1995; k, l, after Vinogradov, 1950; not to scale].

Figure 7. Secondary sexual appendages and structures: a, coxae of pereopods 5 and abdominal somites 1 and 2 of male with pleopods 1 and 2 paired, modified; b, c, coxae of pereopods 5 and abdominal somite 1 of female with pleopod 1 paired, modified; d, female brood pouch; e–g, male pleopod 2; h–q, male sexual tubes; r, male gonopores without sexual tube development; s, coxa of right pereopod 5 of male with gonopore masked by tuft of stiff setae; t, coxa of left pereopod 5 of male with gonopore masked by tuft of stiff setae. [a, from Forest et al. 2000; b, from McLaughlin and Haig, 1995; c, q, r, from McLaughlin and Lemaitre, 2001b; d, from McLaughlin and Provenzano, 1975; e, f, from Lemaitre, 1989; g, from Forest, 1995; i–n from McLaughlin, 1997; h, from Wang and McLaughlin, 2000; p, from McLaughlin, 1986; s, from Melin, 1939; t, from Forest, 1961; not to scale].

Figure 8. Additional morphological characters: a, c, shield with Y-shaped posterior groove; b, shield without Y-shaped posterior groove; d, last thoracic somite and abdomen of Xylopagurus (dorsal view); e, multifid ocular acicles; f, Lithodes rostral spine complex; g, dorsal and ventral rostral spines of Glyptolithodes; h, Cryptolithodes (ventral view) with carapace covering body and appendages; i, rostrum with epirostral spine (lateral view); j, symmetrical uropods and posterior portion of abdominal tergite 6, plus telson (dorsal view) k, right antennal peduncle with hooked spine (indicated by arrow) on lateral margin of segment 1; l, shield of Typhlopagurus showing spinose ocular and antennal acicles and lack of ocular peduncles; m, parapagurid epistome and labrum. Abbreviations: apr = anterior rostral process; ds = dorsal spine(s); es = epistomial spine; ls = labral spine; vs = ventral spine; 6 indicates abdominal tergite 6. [a, from McLaughlin and Hoover, 1996; b, from Forest and de Saint Laurent, 1968; c, from Forest and McLaughlin, 2000; d, from Lemaitre, 1995; e, McLaughlin and Murray, 1990; f, from Vinogradov, 1950; g, after Haig, 1974; h, from Makarov, 1938; i, from McLaughlin, 1997; j, from McLaughlin and Lemaitre, 1993; k, from McLaughlin, 1981; l, from de Saint Laurent, 1972; m, after Lemaitre, 1989; not to scale].