PSEUDOTHELPHUSOIDEA, ORTMANN, 1893

Álvarez, Fernando, Ojeda, Juan Carlos, Souza-Carvalho, Edvanda, Villalobos, José Luis, Magalhães, Célio, Wehrtmann, Ingo S. & Mantelatto, Fernando L., 2020, Revision of the higher taxonomy of Neotropical freshwater crabs of the family Pseudothelphusidae, based on multigene and morphological analyses, Zoological Journal of the Linnean Society 193, pp. 973-1001 : 995-996

KEY TO THE FAMILIES AND SUBFAMILIES OF THE SUPERFAMILY PSEUDOTHELPHUSOIDEA

1. Carapace transversely oval, dorsal surface slightly convex or flat, cervical groove incipient or shallowly marked. In frontal view, suborbital internal tooth high, forming slender, grooved plate reaching frontal margin, closing orbital hiatus completely; branchial efferent channel with tooth near the aperture (except in Epilobocera wetherbeei Rodríguez & Williams, 1995 ). Third maxilliped with exopod longer than lateral border of ischium, without flagellum [except in Epilobocera sinuatifrons (A. Milne-Edwards, 1866) ]. Male G1 long, cylindrical, moderately slender, without lateral lobes on principal axis; apical portion club-shaped, laterodistally directed, without lobules, projections or strong plates, armed with short, stout, curved spines ........................................................................................... Family Epiloboceridae (one genus and 11 species)

- Carapace transversely oval, dorsal surface flat or slightly to moderately convex, cervical groove wide and shallow or narrow and deep, well marked. In frontal view, suborbital internal tooth low, triangular or as a grooved plate, never reaching frontal margin or closing orbital hiatus completely; branchial efferent channel without tooth near aperture. Third maxilliped with exopod generally shorter or almost as long as lateral border of ischium, occasionally a little longer than ischium ( Strengeriana spp. ), and without flagellum (except in Strengeriana flagellatta Campos & Rodríguez, 1993 ). Male G1 long, moderately slender to strong, with or without lateral lobes on principal axis; apical portion variously shaped, distally, laterodistally, or laterally directed, apical elements in the form of plates, projections, processes or lobules; smooth or armed with short, stout, curved spinules or spines ......Family Pseudothelphusidae (8 subfamilies, 48 genera and 293 species) 2

2. Male G1 slender to moderately robust. In mesial view, principal axis with one well developed lateral lobe, sometimes divided forming an accessory lobe ( Neostrengeria spp. ), or reduced [ Martiana clausa ( Rathbun, 1915) ]; apical portion truncated, directed distally and not complexly ornamented with plates, projections, processes or lobules, and rarely ornamented with spines, spinules or granules ( Lindacatalina spp. and Moritschus spp. could present spinules); mesial process reduced or well developed; marginal plate forming caudal crest ........................................................... Subfamily Hypolobocerinae (five genera and 84 species)

- Male G1 long, moderately slender to robust. In mesial view, without lateral lobe on principal axis [except for Ptychophallus spp. and Spirothelphusa verticalis ( Rathbun, 1893) ]; apical portion variously shaped, directed distally, laterodistally or laterally [ Kunziana irengis ( Pretzmann, 1971) ]; apical elements in the form of plates, projections, processes or lobules, arranged in simple or complex structures, reduced or well developed, smooth or armed with short, stout, curved spinules or spines; mesial process in various sizes and forms; marginal plate reaching apical cavity forming part of its border or extended supra-apically ........................................ 3

3. Male G1 with apical cavity in vertical or approximately vertical position relative to main axis of gonopod ........................................................................................................................................................................... 4

- Male G1 with apical cavity transversal or in moderately oblique position relative to main axis of gonopod ........................................................................................................................................................................... 5

4. Male G1 with elongated supra-apical process, formed by distal extension of mesial and caudal surfaces, apical cavity and field of spines in vertical position relative to main axis of gonopod, opening cephalically, laterally or proximally due to strong deflexed apical complex [ Kunziana irengis ( Pretzmann, 1971) ] ......................................................................................... Subfamily Kingsleyinae (14 genera and 63 species)

- Male G1 with elongated supra-apical process, formed by distal extension of lateral surface produced into a spiraliform process turning laterally; apical cavity and field of spines oriented vertically and partially transversally, situated at base of internal surface of spiraliform process ....................................................... ......................................................................................... Subfamily Guinotinae (one genus and one species)

5. Male G1 with marginal plate and caudal surface fused, twisted mesially to end apically in caudomarginal projection, not extending beyond apical cavity, produced cephalically, ending in one to three lobes. Marginal suture fading away on distal third, not reaching apex. Apical cavity transversal or slightly oblique ( Disparithelphusa pecki Smalley & Adkison, 1984 ) to principal axis of gonopod, U-shaped, rounded or exceptionally as narrow slit [ Ehecatusa chiapensis ( Rodríguez & Smalley, 1972) ]; spermatic channel opening in caudal position. Mesial process as triangular or subrectangular plate ( Pseudothelphusa galloi Alvarez & Villalobos, 1990 ), or as axe-shaped laminar expansion, commonly in transversal position relative to principal axis of gonopod, closing apical cavity on cephalic surface. Lateral process absent (except in Epithelphusa chiapensis and Epithelphusa mixtepensis ). ............ Subfamily Pseudothelphusinae (five genera, 40 species)

- Male G1 with marginal plate not fused to caudal surface, ending distally at or beyond apex. Marginal suture straight and visible throughout principal axis of G1. Apical cavity transversal to principal axis of G1, rounded, oval or narrow and elongated caudocephalically. Mesial process in various sizes, forms and positions ........................................................................................................................................................... 6

6. Male G1 with marginal plate extending throughout main axis over caudal and mesial surfaces, ending distally beyond apical cavity, over or aside apical lobe of mesial surface. The latter projected distally forming hood, field of spines on its basal portion. Apical cavity usually oblong; spermatic channel opening in mesocaudal position. Mesial process as strong, conical, acute or triangular spine, cephalically directed ............................. ......................................................................................... Subfamily Potamocarcininae (two genera, 14 species)

- Male G1 with marginal plate extending throughout main axis over caudal and mesial surfaces, becoming distally an edge, in caudal or mesocaudal position, limiting apical cavity, with rounded, oval or narrow shape, elongated caudocephalically; spermatic channel opening in mesocaudal or cephalic position. Mesial process varies in position, as strong, conical, acute or triangular spine, cephalically directed ...................... .......................................................................................................................................................................... 7

7. Apical cavity of male G1 next to caudal surface, oval shaped, elongated caudocephalically, limited by marginal plate and distal crest of lateral surface, with narrow caudal notch. Aperture of spermatic channel in cephalic position. Marginal suture straight, ending distally in triangular marginal process, rarely surpassing apical cavity. Mesial process subapical, generally well developed, triangular or axe-shaped [ Achlidon puntarenas (Hobbs, 1991) ]. Lateral lobe on principal axis present ( Ptychophallus spp. , Camptophallus botti Smalley, 1965 ) or absent ( Achlidon spp. , Allacanthos spp ) ........................................... ...................................................................................... Subfamily Ptychophallinae (four genera, 19 species)

- Apical cavity of male G1 variable in shape, usually as narrow channel, oriented caudocephalically, limited by distal crests of mesial and lateral surfaces, caudally by distal border of marginal plate. Aperture of spermatic channel generally in mesocaudal position. Marginal suture straight, well marked over caudal and mesial surfaces. Apical elements following two patterns, extended distally beyond apical cavity or expanded and projected cephalically forming complex of triangular or dentated processes with accessory spines ............. 8

8. Third maxilliped with exopod as long or longer than ischium (except in Chaceus spp. slightly shorter than ischium); aperture of efferent branchial channel closed or partially closed by jugal angle spine and lateral lobe of epistome. Male G1 with apical elements evidently expanded and projected cephalically forming a complex of processes (except in Chaceus spp. in which apical elements are distally directed), triangular or dentate, with accessory spines ( Strengeriana spp. ), or divided into two lobes, projected cephalically (Pallangothelphusa spp.), or as conical, acute spines, projected cephaloproximally [ Eidocamtophallus chacei (Pretzmann, 1967) ] ....................................................... Subfamily Strengerianinae (four genera, 32 species)

- Third maxilliped with exopod shorter than ischium; aperture of efferent branchial channel open. Male G1 simple, straight, strong, apex moderately ornamented or complex, twisted along main axis [ Spirothelphusa verticalis ( Rathbun, 1893) ], with apical elements variously shaped, directed distally or laterodistally, as plates, projections, processes or lobules. Apical elements arranged in simple or complex ways, reduced or well developed, smooth or armed with short, stout, curved spinules or spines; apical cavity elongated, narrow or oval shaped, limited by mesial and lateral distal crests, directed distally, caudodistally, laterally or even proximally ( Phrygiopilus spp. ). Mesial process usually as strong, conical or triangular spine, cephalad or proximally directed, or subquadrate, pentagonal or rounded ( Raddaus spp. ). Lateral apical surface smooth, with conical spine in different positions or with strong spinate crest arising from lateral border of apical cavity [ Villalobosius lopezformenti (Alvarez & Villalobos, 1991) ]. Marginal plate reaching apical cavity forming apical crest or extending distally beyond it (except in Spirothelphusa verticalis ) ...... ............................................................................................. Subfamily Raddausinae (11 genera, 38 species)