Raddaus bocourti (A. Milne-Edwards, 1866 )

Magalhães, Célio & Wehrtmann, Ingo S., 2025, Taxonomic revision and distribution of the freshwater crabs of the family Pseudothelphusidae (Decapoda, Brachyura) from Guatemala, with the descriptions of two new species and keys to the species of all genera, Zoosystema 47 (26), pp. 581-615 : 596-600

Raddaus bocourti (A. Milne-Edwards, 1866)

( Figs 6 A-C; 7; 16)

Boscia bocourti A. Milne-Edwards, 1866: 203 . — Smith 1870: 146 (in list).

Potamocarcinus bocourti – Ortmann 1897: 317 (in list).

Pseudothelphusa bocourti – Pocock 1889: 10 (in list). — Rathbun 1898: 513, 533, 537 (in list); 1904: 242 (in list); 1905: 294, pl. 15 fig. 1; 1906: 502 (in list). — Young 1900: 220. — Coifmann 1939: 106 (in list).

Pseudothelphusa similis Rathbun, 1904: 242 ( nomen nudum).

Pseudothelphusa similis Rathbun, 1905: 295 , text-fig. 87, pl. 15, fig. 4; 1906: 502 (in list). — Coifmann 1939: 109 (in list).

Pseudothelphusa cobanensis Rathbun, 1904: 242 ( nomen nudum).

Pseudothelphusa cobanensis Rathbun, 1905: 296 , text-fig. 88, pl. 15, fig. 5; 1906: 502 (in list). — Coifmann 1939: 107 (in list). — Pretzmann 1965: 3 (in list). — Smalley 1964: 28, 29, fig. 1A, C, E.

Pseudothelphusa grallator Rathbun, 1904: 242 ( nomen nudum).

Pseudothelphusa grallator Rathbun, 1905: 297 , text-fig. 89, pl. 15, fig. 6; 1906: 502 (in list). — Coifmann 1939: 108 (in list). — Pretzmann 1965: 3 (in list).

Pseudothelphusa magna – Rathbun 1904: 242 (in list); 1905: 299 (part,? ♀ from Pansos Palochie, Haute Vera Paz); 1906: 502 (in list).

Pseudothelphusa cobanensis cobanensis – Bott 1956: 230 (part, typus).

Pseudothelphusa ( Anaphyrmus) bocourti – Smalley 1965: 651 (in list).

Pseudothelphusa ( Anaphyrmus) similis – Smalley 1965: 651 (in list).

Pseudothelphusa ( Anaphyrmus) cobanensis – Smalley 1965: 651 (in list).

Potamocarcinus ( Raddaus) similis – Pretzmann 1965: 3 (in list).

Potamocarcinus ( Raddaus) grallator – Pretzmann 1965: 3 (in list).

Potamocarcinus ( Raddaus) cobanensis – Pretzmann 1965: 3 (in list). — Smalley 1970: 102, fig. 14.

Potamocarcinus ( Raddaus) boccourti [sic] – Pretzmann 1965: 3 (in list).

Potamocarcinus ( Raddaus) bocourti bocourti – Pretzmann 1971: 20 (in list); 1972: 76, text-figs 16, 17, figs 442, 443, 453, 698, 699; 1980: 656.

Potamocarcinus ( Raddaus) bocourti cobanensis – Pretzmann 1971: 20 (in list).

Potamocarcinus ( Raddaus) bocourti similis – Pretzmann 1971: 20 (in list); 1972: 76, figs 447-449, 453-456; 1980: 656.

Potamocarcinus ( Raddaus) bocourti grallator – Pretzmann 1971: 20 (in list); 1972: 76, figs 444-446, 472-474.

Raddaus bocourti – Rodríguez 1982: 98, figs 64b, e, j, k, 65a-g; 2001: 442. — Villalobos F. 1982: 220 (in list). — Rodríguez & Magalhães 2005: 360 (in table). — Villalobos Hiriart & Álvarez 2008: 297 (in list). — Ng et al. 2008: 177 (in list). — Wehrtmann et al. 2014: 123, fig. 1A, B; 2016: 781 (in list). — Acevedo-Alonso & Cumberlidge 2022: 583 (Appendix 2, in list).

TYPE MATERIAL EXAMINED. — Lectotype (by inference of “type” by Rathbun 1905: 294, 295). Guatemala • 1♀ (damaged, 76.5:48.5 according to Rodríguez 1982), dry; Alta Verapaz, environs de Cobán [ 15°27’55.74”N, 90°22’20.10”W; 1303 m elev.]; M. Bocourt leg.; Expédition du Mexique, 1864-1866; MNHN Entry 190-1866; MNHN-IU-2000-4052 (= MNHN-B22814 ) .

Paralectotype (by inference of “cotype” by Rathbun 1905: 295). Guatemala • 1♀ (cw 74.3 mm × cl 48.4 mm, according to Rodríguez 1982), dry; Alta Verapaz, environs de Cobán [ 15°27’55.74”N, 90°22’20.10”W; 1303 m elev.]; M. Bocourt leg.; Expédition du Mexique, 1864-1866; MNHN-IU-2000-4626 (= MNHN-B22814 ) . Holotype of Pseudothelphusa cobanensis Rathbun, 1905 (by original designation). Guatemala • 1♀ (62.5:40.5), dry,; Alta Verapaz, environs de Cobán; M. Bocourt leg.; Expédition du Mexique, 1864-1866; MNHN Entry 190-1866; MNHN-IU-2000-4625 (= MNHN-B4625 , ex B4053) .

Paratype of Pseudothelphusa cobanensis Rathbun, 1905 . Guatemala • 1 ♂ (carapace damaged), dry,; Alta Verapaz, environs de Cobán; date unknown; M. Bocourt leg.; Expédition du Mexique, 1864-1866; MNHN-IU-2000-4628 (= MNHN-B4628 , ex B4053) . Holotype of Pseudothelphusa similis Rathbun, 1905 . Guatemala • 1♂ (70.2:42.5), dry; Alta Verapaz, rivière de Pansos-Palochie [= Polochic river, municipality of Panzós; 15°23’20.74”N, 89°37’30.23”W; 10 m elev.]; XII.1865; M. Bocourt leg.; Expédition du Mexique, 1864-1866; MNHN Entry 190-1866; MNHN-IU-2000-4624 (= MNHN-B4624 , ex-B4058) .

Holotype of Pseudothelphusa grallator Rathbun, 1905 (by original designation. Guatemala • 1♀ (61.3:38.9), dry, Totonicapán, montagnes des Altos, environs de Totonecapan [? San Francisco El Alto ; 14°55’53.07”N, 91°26’42.10”W; 2351 m elev.], surrondings of Totonicapán]; M. Bocourt leg.; Expédition du Mexique, 1864- 1866; MNHN-IU-2000-4627 (= MNHN-B4627 , ex B4055).

Paratypes of Pseudothelphusa grallator Rathbun, 1905 . Guatemala • 1 ♀, dry, 1 ♂ (remnants), dry; same data as in type; MNHN-IU- 2000-4055 (= MNHN-B4627 ) .

OTHER MATERIAL EXAMINED. — Guatemala • 1♂; collecting data unknown; UVGCR 46 • 1 ♂; collecting data unknown; UVGCR 209 • 1♂, UVGCR 510 , collecting data unknown; 1 ♂; Costa Sur ; 28.X.1993; K. Ness M. leg.; UVGCR 97 • 1 ♂, Costa Sur; XI.1993; C. Cardona leg.; UVGCR 367 • 1 juv.♂ (24.5:16.6); Petén, Poptún, Finca Ixobel , cerca a riachuelo; 16°18’14”N, 89°25’13”W; [ 537 m elev.]; 11.I.2014; I. S. Wehrtmann & M. Orozco leg.; UCR-MZ 3236-01 • 2 juv. ♂ (12.4:8.6; 19.8:13.3); Petén, Dolores, Río Sanicté ; 16°36’55”N, 89°32’37”W; 270 m elev.; 12.01.2014; I. S. Wehrtmann & M. Orozco leg. UCR-MZ 3233 • 1 juv. ♂ (21.8:14.5) 1 juv. ♀; Quiché, Sacapulas, quebrada El Tesoro, Pacan , 15°18’11”N, 91°04’49”W; 1954 m elev.; 20.I.2014; I. S. Wehrtmann, M. Orozco, C. Magalhães & M. W. Dix leg.; UCR-MZ 3232-01 • 2♂ spec., Alta Verapaz, Río Chilax, in the surroundings of San Juan Chamelco ; [ 15°26’15.30”N, 90°20’00.89”W; 1368 m elev.]; 1975; Böttger leg.; SMF 8866 • 2♂ (31.3:20.4; 48.3:30.4) 2♀ (22.7:14.7; 42.6:27.1); Izabal, Puerto Barrios, Río Las Escobas ; [ 15°41’09.92”N, 88°38’40.91”W; 93 m elev.]; 18.X.1986; leg. unknown; UVGCR 430 • 1 ♂; Izabal, Río Dulce [ 15°39’26”N, 88°59’58”W; 4 m elev.]; 20.X.1997; F. Bolañoz leg.; UVGCR 173 • 1 juv. ♂; Izabal, Puerto Santo Tomás [ 15°41’37”N, 88°37’32”W; 22 m elev.], Playa Ramonastes ; 29.V.1994; M. L. Priaz & W. Dio leg.; UVGCR 243 • 1 immat. ♂; Izabal, Cerro San Gil, Río Las Escobas [ 15°39’56.40”N, 88°46’54.72”W; 1113 m elev.]; 21.X.2001; Ecología Avanzada – grupo 2 leg.; UVGCR 454 • 2 ♂ (32.8:21.1; 38.8:24.0) 1 ♀ (58.1:35.2); Baja Verapaz, Salamá [ 15°06’01.88”N, 90°18’51.60”W; 940 m elev.]; 20.IV.1934; E. R. Blake leg.; FMNH 1829 • 1 ♂ (33.5:21.4) 3 ♀ (18.8:12.8-27.7:17.5); Zacapa, Villa Rocosa, Río El Jutillo, near Usumatlán ; 15°00’45”N, 89°49’58”W; [ 717 m elev.]; 21.VII.2014; I. S. Wehrtmann & M. Orozco leg.; UCR-MZ uncat . • 1♂; Chimaltenango, Yepocapa [ 14°30’N, 90°57’W]; 1400 m elev.; 6.VI.1949; leg. unknown; USNM 230092 • 1 ♂ 2 ♀ 1 juv., Chimaltenango, Yepocapa; 1460 m elev.; XI.1949; leg. unknown; USNM 230093 • 1♂; Chimaltenango, Yepocapa , 13-14. VII.1949, H. T. Dalmat leg.; USNM 90364 • 1♂ (77.4:48.3) 1 ♀ (80.9:49.6), Chimaltenango, stream in area San Pedro Yepocapa ; XII.1948; H. T. Dalmat leg.; USNM 88721 , • 1♂; Guatemala, Lago de Amatitlán, Tacatón ; [ 14°26’27.35”N, 90°33’33.02”W; 1195 m elev.]; 31.X.1993; C. MacVean leg.; UVGCR 135 • 1 ♂; Guatemala, Amatitlán ; [ 14°28’29.74”N, 90°34’38.94”W; 1190 m elev.]; 6.IV.1991; J. Ruiz R. leg.; UVGCR 148 • 1 ♂; Guatemala, Ciudad de Guatemala, mercado La Plazita [probably captured in Lago Amatitlán ]; 14.XI.2000; F. Peloéz leg.; UVGCR 58 • 1♂ (74.8:46.6); Guatemala, Santa Catarina Pinula, Puerta Parada [ 14°33’21.21”N, 90°27’09.28”W; 1837 m elev.], Río Acatán ; 2.III.1987; W. Dix leg.; UVGCR 523 • 1♂; Guatemala, Santa Catarina Pinula, Puerta Parada ; 1978; M. Dix leg.; UVGCR 438 • 1♂ (83.2:52.3); Guatemala, carretera to El Salvador, km 16 ; [ 14°32’47.82”N, 90°27’06.33”W; 1877 m elev.]; date unknown; S. Rodas leg.; UVGCR 499 • 1♂ (33.4:21.7) 1 ♀ (45.5:29.1); Solalá, boca del río Panajachel [ 14°44’04.96”N, 91°09’26.20”W; 1565 m elev.]; 13.ii.1970; W. Bussing leg.; UCR-MZ 2671-01 • 1 ♂ (70.7:45.8) 1 ♀ (61.1:42.3); Solalá, Lake Atitlán [ 14°42’N, 91°12’W; 1558 m elev.]; date unknown; G. A. Artamonoff leg.; FMNH 12288 • 1 ♂; Solalá, Lago Atitlán ; V.1977; G. Martínez leg.; UVGCR 111 • 1 ♂, Solalá, Panajachel [ 14°44’29”N, 91°09’48”W]; 6.X.1997; J. Cruz leg.; UVGCR 172 • 1 ♂, Solalá, Panajachel, cerca del muelle ; [ 14°44’29.25”N, 91°09’48.20”W; 1563 m elev.]; data unknown; J. Cruz leg.; UVGCR 146 • 1♂ (65.0:40.0); Solalá, Panajachel (bought in the market, captured at Lago Atitlán) ; 21.VII.2013; I. S. Wehrtmann leg.; UCR-MZ 3114-03 • 1 ovig. ♀, Solalá, Panajachel (bought in the market, captured at Lago Atitlán) ; 21.VII.2013; I. S. Wehrtmann leg.; UCR-MZ 3114-01 • 4 ♂ (53.4:33.9-59.0:36.6) 1 ♀ (61.8:38.8); Solalá, Panajachel (bought in the market, captured at lake Atitlán) ; I.2014; C. Magalhães leg.; INPA 2020 • 1 ♂, Sacatepéquez, Municipalidad Mixco, Finca La Esmeralda [? Finca La Esmeralda, in San Lucas Sacatepéquez, Department Sacatepéquez : 14°34’53.88”N, 90°37’03.71”W; 1637 m elev.]; 17.VIII.1999; C. Cardona leg.; UVGCR 16 • 1♂ (23.8:16.1); Suchitepéquez, Patutul, Finca Los Tarrales , stream inside the farm; 14°31’28”N, 91°08’07”W; 875 m elev.; 19.I.2014; I. S. Wehrtmann, M. Orozco, C. Magalhães & M. W. Dix leg.; UCR-MZ 3231-02 • 15 spec.; Santa Rosa, laguna de Ayarza ; [ 14°25’07”N, 90°07’33”W; 1412 m elev.]; 10-11.V.1947; R. R. Miller leg.; USNM 1089230 • 1 ♂; Santa Rosa, Barbarena ; [ 14°18’34”N, 90°21’40”W; 1225 m elev]; 9.XI.2000; J. Castillo leg.; UVGCR 171 • 1 ♂; Santa Rosa, Taxisco ; [ 14°04’02”N, 90°28’04”W; 179 m elev.]; V.1994; J. R. Valdéz leg.; UVGCR 112 • 2 juv. ♂, Santa Rosa, Taxisco-Monterrico ; [ 13°59’38.63”N, 90°27’52.75”W; 30 elev.]; 20.X.2000; leg. unknown.; UVGCR 456 • 1 ♂; Escuintla, Puerto San José ; [ 13°55’45”N, 90°49’13”W; 5 m elev.]; 7.XI.2000; D. Maza leg.; UVGCR 390 • 1 ♂; Escuintla, Puerto San José ; 21.IX.1997; A. Turcois leg.; UVGCR 159 • 1 ♂; Escuintla, aeropuerto Las Lisas [ 13°55’43”N, 90°49’53”W; 12 m elev.], municipalidad San José; 7.X.1976; M. W. Dix leg.; UVGCR 435 .

DISTRIBUTION. — Guatemala ( Alta Verapaz, Baja Verapaz, Chimaltenango, Chiquimula, Escuintla, Guatemala, Izabal, Petén, Quiché, Santa Rosa, Solalá, Suchitepéquez, Totonicapán, Zacapa) ( Fig. 16), Mexico, Belize, Honduras, and El Salvador ( Pretzmann 1980; Rodríguez 1982, 2001; Villalobos Hiriart & Álvarez 2008; Wehrtmann et al. 2016; this study).

OTHER RECORDS IN GUATEMALA. — Pretzmann (1980): Petén, Mopán, at Río Melchor Mencos, border with Belize [ 17°03’27”N, 89°09’06”W; 92 m elev.]; Petén, near Poptún [ 16°21’N, 89°26’W; 544 m elev]; Alta Verapaz, tributary of Río Colochie [= Polochic], NE Tucura [= Tucurú, 15°17’51.32”N, 90°06’58.78”W; 434 m elev.]; Chiquimula, Río Apantes [? Aldea Apantes, 14°32’45.73”N, 89°24’28.64”W; 984 m elev., municipality of Concepción Las Minas], Melen, NW Esquipulas; Guatemala, 14 km S Guatemala City, N San José Pinula [ 14°32’46”N, 90°24’41”W; 1748 m elev.]. Rodríguez (2001): Solalá, Lago de Atitlán, 4 miles E of Panajachel [ 14°41’50.43”N, 91°07’05.52”W; 1604 m elev.]; Guatemala, Río Piscaya [= Pixcaya, Department Chimaltenango, 14°51’24.85”N, 90°39’44.99”W; 759 m elev.], 2 km N junction National RT5-RT4, small tributary of Río Piscaya, at 40 km N from Guatemala; Guatemala, km 56 from Guatemala City on route 5 to Salamá, small tributary of Río Grande [ El Quiché, 14°53’48.53”N, 90°40’20.42”W; 733 m elev.]; Guatemala, km 71 from Guatemala City on route 5 to Salamá, small tributary of Río Grande [ Baja Verapaz, 14°54’48.73”N, 90°31’24.78”W; 967 m elev.].

DESCRIPTION OF G1 ( Fig. 6 A-C)

Straight in mesial view, narrow in the proximal 2/3, wide in the distal third, moderately compressed mesolaterally; caudal margin moderately convex subdistally in mesial view; cephalic margin somewhat irregular, with few short setae more widely spaced medianly, close together subdistally at base of widened mesial process. Marginal suture on mesial surface, straight up to apex, with row of large marginal setae along proximal portion partially extending to marginal plate as row of short setae. Marginal process somewhat narrow, slightly rounded, extended as long as mesial margin of apex. Lateral suture incomplete, marked by shallow sulcus on caudal surface. Mesial process widened, nearly as wide as main stem, roughly trapezoidal, directed cephalad, bilobed, with proximal lobe larger, broadly rounded; distal lobe narrow, rounded, situated obliquely in relation to proximal lobe, slightly longer in mesial view; cephalic surface of mesial process distinctly concave in lateral view, convex in mesial view. Apex oblong, situated apically, open, mesial and lateral margins of same height; field of apical spines well developed, with short, small spines; spermatic channel opening on apex cavity, situated towards caudal end. Cephalic surface with strong triangular, subdistal cephalic spine, pointing cephalad.

REMARKS

The correct status of the type material of this species and three other synonymous taxa is somewhat unclear. Alphonse Milne-Edwards (1866) offered a very brief description of apparently a single specimen from a river of Coban (Alta Verapaz) for which he provided measurement (cw 75 mm × cl 47 mm) but did not inform the sex. However, it is not explicitly mentioned in the text that the specimen would be the type (in the sense of the name-bearing specimen). Although it may seem that A. Milne-Edwards (1866) used a single specimen, this cannot be unequivocally assumed because Rathbun (1905) mentioned a slightly smaller female with the same characters in the MNHN collection that she treated as “cotype”. Furthermore, when visiting the MNHN in September 2015 for examining the Guatemalan material, the senior author found a box containing two dry females of R. bocourti ( MNHN entry 190-1866 in Bocourt’s “Expédition du Mexique ” [catalogued as MNHN-B22814]) with indications of “type” and “co-type”, one of which partially disarticulated and with the carapace badly damaged ( Fig. 7A). Rathbun (1905) mentioned that “Le type du P. Bocourti , c’est-à-dire l’individu que M. A. Milne-Edwards a mesuré, est une grande femelle” [The type of P. Bocourti , that is to say the individual that Mr. A. Milne-Edwards measured, is a large female] and that “Le femelle cotype est un peu plus petite que le type” [The female cotype is slightly smaller than the type]. By this action, Rathbun (1905) was the first author before the year 2000 to have “unambiguously selected a par - ticular syntype to act as the unique name-bearing type of the taxon” as stated in the Article 74.5 (ICZN 1999), which can be taken as a valid lectotype designation. Later, Rodríguez (1982) measured the larger female with cw 76.5 mm × cl 48.5 mm, the smaller female with cw 74.3 mm × cl 48.4 mm, and remarked that the carapace of the former was broken; he treated them as “type” and “ paratype ”, respectively. Thus, the larger, partially damaged female of MNHN-B22814 ( Fig. 7A) must be considered a valid lectotype designation by Rathbun (1905) and, assuming that A. Milne-Edwards (1866) may have had another specimen at his disposal, the smaller female treated as “cotype” by Rathbun (1905) and kept under the same catalogue number should be treated as paralectotype.

Three Pseudothelphusa species from Guatemara were described by Rathbun (1905) based on female specimens: Pseudothelphusa similis , Ps. cobanensis , and Ps. grallator , which have been considered to be junior synomyms of R. bocourti (see Rodríguez 1982; Ng et al. 2008). Rathbun (1905) founded Ps. similis on a single male specimen but also mentioned 17 juvenile specimens from the same locality and attributed a slight morphological difference between them to age. As the male, to which she provided measurements, is unequivocally termed “type”, this is the holotype fixed by original designation (ICZN 1999, Art. 73.1.1). The holotype of Ps. similis (MNHN-B4624) is a dry male with a longitudinal cleft along the middle and on the right side of the dorsal region of the carapace ( Fig. 7B).

The type specimens of Ps. cobanensis , a well-preserved female ( Fig. 7C) and a male in very bad condition ( Fig. 7D), are also dry. Rathbun’s (1905: 296, 297) action, by clearly treating only the female as “type”, can be interpreted as she has set it as the unique name-bearing type of this taxon (Art. 73.1.1, ICZN 1999), therefore fixing the holotype by original designation. The male was designated as “co-type” ( Rathbun, 1905: 297, text-fig. 88), which can then be considered as a reference to paratype. This was also Rodríguez’s (1982: 101) interpretation when treated the female as the holotype and the male as paratype.

Rathbun (1905: 297) based the description of Pseudothelphusa grallator on at least two specimens: a female (cw 61.3 mm × cl 38.9 mm) and a male (cw 59.2 mm × cl 37.8 mm) and, in “Dimensions”, provided a table with their measurements. In this table, she indicated the female as “Type”. In the section “Distribution”, she listed eight specimens from the same locality: three females (including the “type”, numbered 623d in the Bocourt’s material from the “Expédition du Mexique ”), one male (no. 624d in Bocourt’s catalogue), and additionally one female and three young specimens. Since the female with the number 623d (MNHN-B4627) is clearly marked as “Type” by Rathbun (1905), this specimen is the holotype fixed by original designation (Art. 73.1.1, ICZN 1999). Rodríguez (1982) remarked that the situation of these specimens in the Paris Museum is somewhat confusing but, even taking into account the poor condition of the material, he concluded that the reasonably well-preserved female measuring cw 60.5 mm × cl 39 mm should be the one Rathbun considered as the “Type”. During a visit to the MNHN in September 2015, the senior author found that the condition of some of these specimens had worsened but this dry female was still in reasonably good condition and preserved in a box with the catalogue number 4627 and with a label stating “Bocourt (Expédition du Mexique) (no. 623 d de son catalogue)”; this box also had the indicative sign of holotype (the red circle label) attached to it ( Fig. 7E). This specimen should be the one considered by Rodríguez (1982) as the holotype. Another box, bearing a handwritten label and stating “ Pseudothelphusa grallator Type 4627”, contained one dry female with the dorsal region of the carapace severely damaged and almost entirely disarticulated as well as the remnants of a male specimen with the abdomen partially broken and the sternopleonal cavity glued to the box (this male has the number “624d” written on the anterior portion of the sternum) ( Fig. 7F, G). Although these lots bear a label with the word “TYPE” written in red, these specimens should be considered as paratypes; Rodríguez (1982) considered the male as being an allotype.