Diogenidae, , de Saint Laurent and McLaughlin, 2000

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 : 113-114

Key to genera of Diogenidae

1. Well developed arthrobranchs present on arthrodial membranes at bases of cheliped and maxilliped 3; pleurobranch present on somite XI (thoracomere 5, above pereopod 2) ( Fig. 4a).......................... 2

— Reduced or vestigial arthrobranchs present on arthrodial membranes at bases of cheliped and maxilliped 3; no pleurobranch present on somite XI (thoracomere 5, above pereopod 2) ( Fig. 4c).......................................... Pseudopaguristes McLaughlin, 2002

2. 14 pairs of gills; pleurobranch present on somite XIV (thoracomere 8, above pereopod 5) ( Fig. 4a)........... 3

— 13 pairs of gills; no pleurobranch present on somite XIV (thoracomere 8, above pereopod 5) ( Fig. 4b)........ 14

3. Endopod of maxillule with well developed external lobe ( Fig. 4k)................................... 4

— Endopod of maxillule without well developed external lobe ( Fig. 4l)................................ 8

4. Ischium of maxilliped 3 with well developed crista dentata ( Figs 2b–f )............................ 5

— Ischium of maxilliped 3 without well developed crista dentata ( Fig. 2a )............................. 12

5. Chelipeds equal or unequal, each with stridulatory mechanism developed on mesial face of palm ( Fig. 6a ) 6

— Chelipeds markedly unequal, left largest; neither with stridulatory mechanism developed on mesial face of palm ( Fig. 6b )..... Allodardanus Haig and Provenzano, 1965

6. Chelipeds with acute, corneous-tipped spines on carpi and chelae; males often with pleopod 2 paired, endopod well developed, reduced or absent ( Fig. 7g )..................................... Strigopagurus Forest, 1995

— Chelipeds with tubercles or transverse striate on carpi and chelae; males without pleopod 2 paired............ 7

7. Chelipeds equal or left larger; carpus and palm with transverse striae bordered with fine setae ( Fig. 6c ); dactyls of ambulatory legs equal to or longer than propodi; females with unpaired pleopods 2–5 egg-carrying..................................... Ciliopagurus Forest, 1995

— Chelipeds equal, carpus and palm covered with generally blunt tubercles; dactyls of ambulatory legs much shorter than propodi; females with unpaired pleopod 5 non egg-carrying................ Trizopagurus Forest, 1952

8. Chelae symmetrical, together forming operculum ( Fig. 6d ); uropods symmetrical ( Fig. 8j ).................................. Cancellus H. Milne Edwards, 1836

— Chelae symmetrical or asymmetrical, together not forming operculum; uropods generally asymmetrical........ 9

9. Chelipeds unequal, right distinctly larger.................................. Petrochirus Stimpson, 1858

— Chelipeds subequal or unequal, left usually at least slightly larger............................... 10

10. Shield with prominent Y-shaped linea in posterior half ( Figs 8a, c ); posterior carapace primarily membranous; left cheliped slightly to considerably larger than right....11

— Shield without prominent Y-shaped linea ( Fig. 8b, l ); posterior carapace well calcified; chelipeds subequal....................... Tisea Morgan and Forest, 1991

11. Rostrum triangular; ocular acicles each with prominent triangular or subtriangular acicular projection ( Figs 8a, b ); chelipeds and ambulatory legs with ring-like transverse striae ( Fig. 6e ); females with well-developed brood pouch ( Fig. 7d ).................... Aniculus Dana, 1852

— Rostrum broadly rounded or obsolete ( Fig. 8c, l ); ocular acicles each with subrectangular or subquadrate acicular projection ( Fig. 8c ); chelipeds and ambulatory legs most often without ring-like transverse striae; females without brood pouch............... Dardanus Paul’son, 1875

12. Antennal flagella with microscopic setae.................................. Pseudopagurus Forest, 1952

— Antennal flagella with paired, moderate to long setae ( Fig. 2k ).................................. 13

13. Chelipeds equal or slightly subequal, similar, dactyls opening in generally horizontal plane ( Fig. 6f ).........…......................... Isocheles Stimpson, 1858

— Chelipeds unequal and dissimilar; dactyls opening in almost vertical plane ( Fig. 6g ). Loxopagurus Forest, 1964

14. Males with pleopods 1 and/or 2 paired, modified as gonopods ( Figs 7a, e, f ); females with ( Figs 2m , 7b, c ) or without pleopod 1 paired, modified.............. 15

— No paired pleopods in either sex................ 16

15. Pereopod 4 chelate ( Fig. 5t ); unpaired pleopods 3–5 occurring on either right or left side of abdomen................................ Paguropsis Henderson, 1888

— Pereopod 4 not chelate; unpaired pleopods 3–5 occurring on left side of abdomen only... Paguristes Dana, 1851

16. Chelipeds subequal ( Fig. 1f ).................... 17

— Chelipeds unequal, left appreciable larger ( Figs 1d, e, g ).… .......................................... 18

17. Ocular acicles bi or multispinose, contiguous or closely set ( Fig. 1f , 8b ); posterior margin of abdominal somite 6 unarmed.................. Clibanarius Dana, 1852

— Ocular acicles simple, widely separated ( Figs 3a, d ); posterior margin of abdominal somite 6 spinulose............................. Bathynarius Forest, 1989

18. Rostrum obsolete, roundly subtriangular or broadly round- ed, intercalary rostral process present, well developed, reduced or vestigial ( Fig. 3i )...... Diogenes Dana, 1851

— Rostrum moderate to well developed, triangular, intercalary rostral process absent... Calcinus Dana, 1851

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].