Taxonomy of the freshwater crabs of Costa Rica, with a revision of the genus Ptychophallus Smalley, 1964 (Crustacea: Decapoda: Pseudothelphusidae) Author Magalhães, Célio Author Wehrtmann, Ingo S. Author Lara, Luis Rólier Author Mantelatto, Fernando L. text Zootaxa 2015 3905 3 301 344 journal article 42408 10.5281/zenodo.235127 16b434af-bed4-43fd-bdfc-0f29bcaec7bb 1175-5326 235127 4DA803F1-30C6-4FC3-8EEF-9690A89822A6 Ptychophallus paraxanthusi ( Bott, 1968 ) ( Figs. 44–53 , 82 ) Pseudothelphusa colombiana — Rathbun 1898 : 533 (part).— Young 1900 : 219 (part).— Rathbun 1905 : 302 (part). Pseudothelphusa xantusi — Rathbun 1898 : 536 (part).— Young 1900 : 208 (part).— Rathbun 1905 : 292 (part), fig. 85b.— Rathbun 1906 : 501 (in list). Pseudothelphusa ( Ptychophallus ) xantusi — Smalley 1964b : 11 , fig. 15. Pseudothelphusa ( Ptychophallus ) paraxanthusi Bott, 1968 : 43 , fig. 8. Ptychophallus ( Semiptychophallus ) columbianus paraxanthusi [sic]— Pretzmann 1971 : 21 . Ptychophallus ( Semiptychophallus ) colombianus paraxanthusi — Pretzmann 1972 : 89 , figs. 488–491, 516–518. Ptychophallus paraxanthusi — Rodriguez 1982 : 82 , figs. 49a, d, j, 50a–d.— Campos & Lemaitre 1999 : 560 (map).— Rodríguez & Hedström 2000 : 424 .— Ng et al. 2008 : 177 (in list).— Villalobos Hiriart & Álvarez 2008 : 297 (in list).— Lara et al. 2013 : 689 (table), 691, fig. 3 (map). Ptychophallus osaensis Rodríguez, 2001 : 440 , fig. 3; Ng et al. 2008 : 177 (in list).— Villalobos Hiriart & Álvarez 2008 : 297 (in list). Material examined . COSTA RICA . Locality not specified: 3 males , 1 female , UCR-MZ 678, Los Reyes Creek, Iglesia Los Reyes, 22.vii.1972 , V. Juárez leg.—San José Province, Pacific drainage: 1 male , UCR-MZ 646, La Palma, La Bonita Creek, 6.5 km from San Isidro, 5.xii.1971 , V. Juárez leg.; 6 males , UCR-MZ 673, Pérez Zeledón Canton, Convento River, 38 km E San Isidro de General, 31.i.1972 , V. Juárez leg.; 11 males , 6 females , UCR-MZ 640, Pérez Zeledón Canton, Cajón River, on Panamerican Highway, 24 km from San Isidro del General, 28.ii.1971 , V. Juárez, C. Villalobos & R.A. Cruz leg.; 1 male , UCR-MZ 657, same data; 3 males , 2 females , UCR-MZ 689, Pérez Zeledón Canton, San Pablo, Platanares River, 15.vii.1972 , V. Juárez leg.; 2 males , 1 immat. male, 2 females , UCR-MZ 697, Las Bonitas Creek, San Rafael de Platanares, 15.vii.1972 , V. Juárez leg.; 2 males , UCR-MZ 672, Platanares River, 75 m N from bridge, Paso de Pejivalle [= Pejibaye] to Achotales, 22.vii.1972 , V. Juárez leg.; 1 male , 1 female , UCR-MZ 674, Ignacio Mata Creek, 2 km N from Pejivalle [= Pejibaye], direction to San Isidro, 22.vii.1972 , V. Juárez leg.; 2 males , 1 female , UCR-MZ 692, Pérez Zeledón Canton, creek 100 m N from plaza de Pejivalle [= Pejibaye], 22.vii.1972 , V. Juárez leg.— Puntarenas Province, Pacific drainage: 4 males , 1 immat. male, 1 immat. female, UCR-M 694, Ceibo River, 35 km E San Isidro, 19.ii.1972 , V. Juárez leg.; 6 males , 1 female , INPA 1854, Cola de Gallo River, tributary of Cabagra River, 9°01’29.748”N 83°10’25.680”W , 17.v.2007 , L.R. Lara leg.; 2 males , UCR-MZ 676, Sabanilla River, Paso de Sabanillas, 12.iii.1972 , V. Juárez leg.; 3 males , UCR-MZ 677, Escuadra River, 2.1 km S Paso Real, 5.iii.1972 , V. Juárez leg.; 1 male , UCR-MZ 679, Curré River, 10.5 km S Paso Real, 3.iii.1972 , V. Juárez leg.; 2 males , UCR-MZ 691, Puerto Nuevo River, 16 km S Paso Real, 5.iii.1972 , V. Juárez leg.; 2 males , UCR-MZ 675, Disciplina River, 20 km S Paso Real, 5.iii.1972 , V. Juárez leg.; 1 male , 1 female , INPA 1855, Veraguas Creek, tributary General River, 09°03’26.00"N 83°28’47.02"W , 15.v.2007 , L.R. Lara leg.; 1 male (cw 26.0, cl 16.3) 1 female (cw 21.4, cl 13.5) 1 juv. , USNM 20331, Boruca, undated, H. Pittier leg.; right G1, holotype , MNHN-B 5328, Boruca, undated, H. Pittier leg.; 4 male , UCR-MZ 683, La Guácima Creek, on road to San Vito, 12.iii.1972 , V. Juárez leg.; 5 males , 2 females , UCR-MZ 2650-01, Cacao Creek, tributary Fresco River, Grande de Térraba River basin, 22.v.2007 , L.R. Lara leg.; 2 ad. males, 1 juv. male, UCR- MZ 3217-05, road to Ciudad Neily-San Vito, below bridge of Agua Buena, 8°42’45.45”N 82°54’33.33”W , 1117 m , 27.i.2014 , I.S. Wehrtmann, C. Magalhães & E. Souza-Carvalho leg.; 6 males 2 females , UCR-MZ 696, Catarata River, 7 km S of bridge over El Brujo, Panamerican Highway, 19.ii.1972 , V. Juárez leg.; 5 m , UCR-MZ 681, La Vieja River, 7.8 km S from Paso Real, 5.iii.1972 , V. Juárez leg.; 1 male , UCR-MZ 2660-01, Catarata River, 10 km NW intersection of Interamericana highway, road to San Vito, 14.vi.1986 , Smalley, Campo, Kersting & Kersting leg.; 5 males immat., 11 females immat., 2 juv. , INPA 1856, Limón River (arriba), 8°49’26.724”N 83°01’41.520”W , 8.v.2007 , R.L. Lara leg.; 1 male , UCR-MZ 2652-01, Batambal Creek (Desfogue), Palmar Norte, 8.ix.2007 , L.R. Lara leg.; 1 male , 1 female , UCR-MZ 2653-01, Escuadra Creek, Cataratas, Buenos Aires, 15.v.2007 , L.R. Lara leg.; 2 males , 1 female , UCR-MZ,2658-01, El Silencio Creek, Palmar Norte, 27.vi.2007 , L.R. Lara leg.; 3 males , UCR-MZ 687, Honda Creek, Palmar Norte, 7.vii.1972 , V. Juárez leg.; 2 males , UCR-MZ 682, Zapote River, Palmar Norte, 7.vii.1972 , V. Juárez leg.; 5 right and 2 left G1, INPA 1657, Camaronal River, Grande de Térraba River basin, 08°57’52.02"N 83°28’14.14"W , 30.v.2007 , L.R. Lara leg.; 2 males , UCR-MZ 690, Osa Canton, creek 100 m N from Zapote River, 7.vii.1972 , V. Juárez leg.; 1 male , 1 female , UCR-MZ 684, Zapotico Creek, Palmar Norte, 7.vi.1972 , V. Juárez leg.; 1 male , UCR-MZ 695, Salama Viejo River, 30 km S from bridge over Grande de Térraba River, Palmar Norte, 5.vii.1972 , V. Juárez leg.; 1 male , INPA 1655, Grande de Térraba River basin, 9.v.2007 , L.R. Lara leg.; 4 males , 1 female , INPA 1852, Térraba Creek, Grande de Térraba River basin, 9°03’56.952”N 83°17’28.824”W , 9.v.2007 , L.R. Lara leg.; 1 male , INPA 1833, near Sierpe, Estero Seco, 08°47.182’N 82°55.666’W , 16.xi.2009 , L.R. Lara leg.; 2 males , holotype (cw 47.9, cl 28.5) and paratype (cw 31.9, cl 19.9) of Ptychophallus osaensis Rodríguez, 2001 , TMNH 6271, Rincón de Osa,1971, leg. unknown; 8 males , 1 female , 4 juv. , TMNH 6251, Rincón de Osa, vii.1972 , R. Zeledón leg.; 3 males , 1 immat. male, 2 immat. females, UCR-MZ 680, Rincón de Osa, creek 2 km W from Barracas OTS, 8.v.1971 , C. Villalobos & R. Barraotes leg.; 8 males , 10 females , UCR-MZ 653, Rincón de Osa, creek behind Estación OTS, 8.v.1971 , C. Villalobos leg.; 1 male , 7 juv. females, UCR-MZ 693, Rincón de Osa, creek behind Estación OTS, 7.v.1971 , C. Villalobos leg.; 3 males , 2 females , UCR-MZ 724, Rincón de Osa, creek behind Estación OTS, 9.v.1971 , C. Villalobos leg.; 4 males , 1 female , UCR-MZ 724, Rincón de Osa, creek behind Estación OTS, 9.v.1971 , C. Villalobos leg.; 2 males , 6 females , 1 juv. , UCR-MZ 749, Rincón de Osa, Agua Buena Creek, behind Estación Experimental, 2.vi.1972 , E. Solis & G. Rojas leg.; 2 males , 1 female , INPA 1853, same data; 1 juv. male, UCR-MZ 3213-01, peninsula de Osa, road to Rincón a Drake, creek after Rancho Quemado, 8°41’26.26”N 83°34’36.36”W , 191 m , 26.i.2104 , I.S. Wehrtmann, C. Magalhães & V. Souza-Carvalho leg. Locality not specified: 1 male (cw 43.0, cl 24.1), USNM 20035, “ Mexico , 300 m ”, undated, leg. unknown. PANAMA . Chiriquí Province, Pacific drainage: 1 male (cw 30.8, cl 19.0), USNM 1089231, Cerro Punta, 1- 3.iii.1963 , E.J. Morris leg.; 3 males , 5 females , 4 juv. , USNM 1089234, vicinities La Concepción, probably Mula River, iv.1964 , leg. local children. Description of gonopod . Straight in mesiocaudal view. Marginal suture on mesial side, straight; marginal process broadly rounded, not produced beyond distal border of apex. Mesial process usually broadly rounded to subretangular, transversally projected (not downturned). Lateral process lengthy, bilobed, approximately 60% of stem length (from proximal opening to caudal border of apex), with shallow depression in middle; lateral border with usually shallow median incision; proximal lobe broader, wider than distal one, as wide as or slightly wider than lateral end of apex (mesiocaudal view); distal lobe weakly to moderately developed, obliquously directed towards caudal side, usually narrower than lateral end of apex in mesiocaudal view. Distocaudal ridge rather long (nearly reaching level of median depression of lateral process), narrow, separated from distal border of lateral process by shallow depression. Apex peduncule bent approximately 50–60° towards laterocephalic side; oblong, rather wide in distal view; caudal border of apex slightly convex, with notch near lateral end. Field of apical spines well developed, facing towards cephalolateral side. Holotype and type locality . Male, MNHN-B 5328 (only G1 examined). Costa Rica , Puntarenas Province, Boruca (Pacific drainage). Distribution . Pacific drainage of eastern Costa Rica (provinces of San José and Puntarenas ) and western Panama (province of Chiriquí) ( Fig. 82 ). FIGURES 44–60 . Ptychophallus paraxanthusi (Bott, 1968) : 44, left first gonopod, holotype, MNHN B-5328, caudo-mesial view; 45, same data, latero-cephalic view of distal part; 46, right first gonopod, holotype of Ptychophallus osaensis Rodríguez, 2001 , TMNH 6271, caudo-mesial view; 47–48, outline of right first gonopod, two males from INPA 1853, caudo-mesial view; 49, same data, INPA 1657; 50, same data, INPA 1855; 51–52, outline of left first gonopod, two males from INPA 1854, caudomesial view; 53, same data, INPA 1855. Ptychophallus tumimanus (Rathbun, 1898) : 54, left first gonopod, holotype, USNM 19484, latero-cephalic view of distal part; 55, same data, caudo-mesial view; 56, same data, holotype of Ptychophallus ( Ptychophallus ) tumimanus ingae Pretzmann, 1978 , NHMW 4009, caudo-mesial view; 57, outline of left first gonopod, INPA 1832, caudo-mesial view; 58, same data, USNM 230099; 59–60, same data, two males from INPA 1835. Scale bars = 1 mm. Remarks . The holotypes of P. paraxantusi and P. osaensis were collected from localities (Boruca and Rincón, respectively) approximately 40 km apart in southern Puntarenas Province, Costa Rica . The slight morphological differences seen in their gonopods are not consistent to treat them as separate taxa and therefore they are hereby synonymyzed. Both lobes of the lateral process are less developed in the G1 of the holotype of P. paraxantusi ( Figs. 44, 45 ) than those of the G1of the holotype of P. osaensis ( Fig. 46 ) and bears some minute setae along the lateral border, besides the distal lobe being somewhat irregular in the former. The lobes are slightly more prominent in the G1 of the holotype of P. osaensis , with distal lobe knob-like and bearing a smooth border ( Fig. 46 ). Such differences, however, could be attributed to the size of the specimens ( holotype of P. paraxantusi is subadult with cw 26 mm ; holotype of P. osaensis is a fully grown adult with cw 47.9 mm ) as well as to intraspecific variability as intermediate conditions have been observed in the numerous specimens examined from the region. In both topotypic males in INPA 1853 the gonopods have a distal lobe that is narrow and with smooth border, but with a wide proximal lobe ( Figs. 47, 48 ), whereas the male collected approximately midway from the type localities of both species (Camaronal River, INPA 1657) has the distal lobe bearing an irregular border and wide proximal lobe ( Fig. 49 ). Although the proximal lobe is usually larger than the distal lobe, some variability may occur in their degree of development (see Figs. 47–53 ). In some specimens the lobes can have nearly the same size (as in some males in INPA 1854, 1855; Figs. 52–53 ), and in a few specimens the distal lobe can be even slightly larger (e.g. male in INPA 1854; Fig. 51 ). The mesial process can also be slightly variable, with rounded (e.g. TU 6271; Fig. 46 ), subtriangular (e.g. MNHN-B 5328, INPA 1657, 1855, 1854; Figs. 44, 49, 52 , respectively), or rarely subretangular shape (e.g. INPA 1853; Fig. 47 ) shape. These conditions are illustrated in Figs. 44–53 . The specimens from USNM 20331 (Boruca, Costa Rica ) were identified by Rathbun (1898 ; 1905 ) as Pseudothelphusa xantusi , a species listed as incertae sedis by Rodriguez (1982) . Rathbun (1905) stated that the lot had three males and one young male, which was sent to MNHN. The USNM specimens consist of one male, one female, and one young specimen; the male has both G1 missing, but a small label stating “app. fig.” may indicate that it was the one used by Rathbun (1905: 292, fig. 85b) to illustrate the G1. The lot USNM 20035 with a male identified as Pseudothelphusa colombiana by Rathbun (1898: 533) has a label stating “ Mexico , 300 m , British Museum, Acc. 31482”, and a second with “ Ptychophallus tristani Rathbun, Det. Smalley, Jun 1966 . Strongly suspect an incorrect label”. The specimen was examined by the senior author, who identified it as P. paraxanthusi ; therefore the Mexican locality is highly doubtful.