6 .April 1998 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(11:110-1 39. 1998. A revision of the freshwater crabs of the family Pseudothelphusidae (Decapoda: Brachyura) from

Gilberto Rodnguez and Richard von Sternberg

(GR) Cenlro de l-cologia, Institute Venezolano de Investigaciones Cienifficas, Apart ado 21827, Caracas 1020 A. Venezuela; (RvS) Center for Intelligent Systems, SUNY-Binghamton. New York. 13902-6000, U.S.A.

Abstract*—Revised diagnoses and illustrations or" the species of pseudo- thelphusid crabs previously known to occur in Ecuador are provided and five new species. Hypoiobocera esmeraldensis, H. konstanzae, H. muisnensis, H. mindonensis and Lindacatalina sumacensis, are described. The validity of pen- tanomial names proposed in the literature for some Ecuadorian species is re­ vised in the light of series of specimens collected in their distributional areas. The genera Hypoiobocera* Moritschus and Lindacatalina are redefined and several species are reassigned among these genera.

The Systematics of Neotropical freshwa­ example, what he called Hypoiobocera ter crabs of the family Pseudothelphusidae (Hypoiobocera) aequatorialis aequatoriaiis is relatively complex due to lack of de­ in 1972, became Hypoiobocera (Hypoio­ pendable laxonomic differences in their car­ bocera ) [aequatorialis'] aequatorialis ae­ apace and appendages. The male first gon- quatorialis in his later contributions. The opods provide the most reliable characters erection of these infraspecific categories of diagnostic value, whereas potential so­ was based usually on one or two specimens. matic characters, such as the dentition of In some cases two infraespecific categories the lateral border form and position of the of the same species were reported from the cervical grooves and postfrontal lobes, same localities (see for instance Hypoio­ sculpturing of the front, and proportions of bocera {Hypoiobocera) [peruviana] henrici the third maxilliped, display very slight in­ henrici and Hypoiobocera {Hypoiobocera) terspecific differences. {peruviana] henrici nora). This treatment of In almost all cases it is not possible to the Ecuadorian species has resulted in con­ establish infraspecific categories that will siderable confusion and serious difficulties meet the two criteria accepted for other an­ for the identification of binomial taxa. imal groups, viz., that the differences are In the present contribution the validity of slight, but constant through large series of some of these infraspecific taxa is revised specimens (Mayr 1964), and that no overlap in the light of series of specimens collected occurs in the geographical distribution of in the same areas as Pretzmann*s materials the supposed subspecies (Mayr et al. 1953). (Pretzmann & Radda 1978). All new or In his revision of the Pseudothelphusi­ critical species reported here are fully illus­ dae, Pretzmann (1972) proposed the orga­ trated. For other species only figures oi the nization of the taxa into a tetranomial first male gonopods are given, together with scheme (genus, subgenus, species and sub­ references to adequate illustrations in the species). In further contributions he ar­ literature (see ^Additional illustrations" ranged some crabs from Ecuador into a under each species). Two species, Hypoio­ more elaborate pentanomial nomenclature bocera conradi (NobilL 1897) and Linda­ (Pretzmann 1978, 1983a, 1983b). Thus, for catalina hauserae Pretzmann, 1977b, are VOLLAU-; 1 I 1, NLMBfcK I 111 not illustrated for lack of material. Termi­ Tahlt 1.—Curapiicf breadth of largesl males nology for gonopod morphology follows recorded and proportions of the cxogEiaih lo ischium of cndogiuilh of third muxdlipedv in Hcuadoriaji Pseu­ Smalley < 1964). dothelphusidae. Abbreviations used are cl. for carapace length and cb. for carapace breadth. The E\(i^r.ilh.: fh (mm} materials recorded are deposited in the Ref­ erence Collection of the Instituto Venezo- H\plati 65 1 0.30 the Museum of Natural History of Tulane li. i-smrrahiensts v^.S 0.40 University, New Orleans (TU). the National fi. t.xtfiti MS 0.30 Museum of Natural History, Smithsonian H, i!iui\titfuiU-ti\i\ 44.1 0 30 J Institution, Washington, D. C (USNMl, the H kfni.\Uinztit. 56.8 0.30 H mhuitmt.'itsi\ 27.1 0.30 Museum nationale d'Histoire naturelle, Par­ it, muisni'nsis 51 A 0.35 is (MNHN), the Strasbourg Museum (SM) H, titrtwi 2.V5 0.30 and the Nuturmuseum und Forschunginsti- //. ntthhttui 45.0 0.35 tut Senckenberg, Frankfurt am Main UtniantUilitia hnf\ •ipvttit 27.5 0.65 (SMF). L. htutst'mv 25.0 0.50 /,. UltifWtlii 55.7 0.50 L. t*rit'tthrfis 2N.0 0.65 Systematics /.. pu\t HMi 32.3 0.45 A. Mtrudu•«.%/.* 35.6 0.45 Family Pseudothelphusidae Ralhbun, 1893 \ft>tft

1. Lateral margin of hrsi gonopods pro­ duced into a delined. although some­ times reduced, lateral lobe 2 distal view, with round papilla near sper­ -. Lateral margin widening progressively matic channel (Fig, IB). towards the apex which extends consid­ Type species.—Potamia chilensis H. erably laterally, giving the apex in cau­ Milne Edwards & Lucas, 1844. dal view a characteristic rriangular-elon- Distribution.—Panama. Colombia. Ven­ gated appearance Mohtschits ezuela. Ecuador and Peru. 2. Lateral lobe densely covered with spi- nulev Lxognaih of third maxilliped usu­ ally more than 0.45 length of ischium of Key to Species from Ecuador endognath Lmdtwtitalinct -. Lateral lobe naked or with a few sparse 1. Lateral lobe o\ lirsl gonopods reduced spinules and short hairs. Exognath of or obsolescent (Figs. IE, 4A) 2 third maxilliped usually less than 0.45 -. Lateral lobe well developed 6 length of ischium of endognath ..... 2. Lateral lobe with small scattered papil­ Hvff'ttthocerti lae (Fig. 7A> H. tnindoneusis - Lateral lobe with smooth surface or with scattered short hairs 3 L Hxpolttbocera Ortmann. IS >7 3. One or two prominent tubercles on apex of tirst gonopods 4 Diagnosis.—Exognalh of third maxilli­ -. No prominent tubercles on apex of first ped usually less than 0.45 length of ischium gonopods 5 of endognath (Table 1}. First male gono­ 4. One prominent tubercle: apex produced pods with strong longitudinal ridge on cau­ laterally into extraordinarily long, ob­ dal surface, and well defined (although tuse lobe H. exiwa sometimes reduced} lateral lobe (Fig. 1A); -. Two prominent tubercles on apex of apex truncated, either circular or oblong in gonopods . • < • ... , . . H orcv.si PROCEEDINGS OK THL BIOLOGICAL SOCIETY Oh WASHINGTON

lii:. I. J; L i s. I left i:onopod of Ecuadorian PseudodielphLisidae: A, B, Hvpolebui eta aafuufor'taUs lOrtmann, 1S1>7). hololypc I rum Ecuador (SMK C. D. H. cn/nttii iNobiiL 1901). from Rio Quevedo (IVIC 62S): L. P. //. raifihuni Prel/inauu. IWS. from Kit) Peripa. between Aurora en Puerto Limun lIVIC 631): G. H. \hntr\chny i'cututtn-t'n.\h tRaihlnm. I S97 L lioiu UCM of Gualea (BM 9l8.L31.Lh: L J, M. Iwnriri [NohUL 1X97}, ham Ecuador (IVIC 615); KM. Limhtcuntihui hrwiptnis (Rodritiue/ & DKI/}. 39NL from Ecuador llVlC 606): N P. A. lot i penis iPi-iM/mann, 1968). from Ecuador (IVIC 62 h. A. C. L. Ci. L K. N. caudal: M. O. lateral: B. D. F. II. J. L. P. apex, distal: lo, lateral lobe; a\ caudal ridge: nl. supplementary lobe, Scales = 2 mm.

5- With large tubercle on external surface 6. Lateral lobe of iirsl gonopods long, ob- o( palm H. conradi long, wilh proximal angle rounded and -. Without a large tubercle on external distal angle sloping gently to apex ... 7 surface of palm H, rathbutu -. Lateral lobe triangular or subiriangular. . . 8 VOLUME 111. NUMBER I 113

7. A large tubercle on external surface of Hypolohocera {Hypolohocera) [aequatori­ palm H. delsolari alis] aequatorialis nigra.—Pretzmann, ~. Without a large tubercle on external 1983b:352, figs. 3, 17, 25, 35, 52. 55, 72. surface of palm H. aequatorialis 8. Apex of first gonopods in caudal view Material.—Ecuador: Leg. Reiss, 1 male forms very elongated spine, projected holotype of Potamocarcinus aequatorialis laterally and dislally (Figs. 1C, 3A) . . 9 Ortmann, 1897 (SM).—Arroyo de Arrayan, -. Apex in caudal view with lateral border N of Banos, Parroquia de Chirgua, TUngur- obtuse or with short spine (Figs. ahua Province, 1750 m alt., 7 Nov 1980, 5A.6A.8A) 10 9. Border of lateral lobe straight or slight­ leg. H. Diaz, 3 males cl. 31.5, 31.0 and 21.8 ly convex distally H. caputii mm, cb. 49.8, 48.8 and 33.7 mm, 2 mature -. Border of lateral lobe rounded distally females cl. 35.0 and 34.1 mm, cb. 57.0 and H. esmeraldensis 53.5 mm, 1 immature female cl. 27.8 mm, 10. Apex in distal view with lateral margin cb. 43.5 mm (IVIC 590).—Banos, Tungur- acute or ending in a short point directed ahua Province, Dec 1984, leg. Ferro, 1 laterally 11 male, I immature female (IVIC 972).— -. Apex in distal view with lateral margin Quebrada Punsan, Pueblo de Alba, E of rounded H. muisnensis Banos, , 1600 m alt., 11. Border of lateral lobe expanded and 7 Nov 1980, leg. H. Diaz, 1 male cl. 27.8 rounded distally H. guayaquilensis -. Border of lateral lobe narrow and trans­ mm, cb. 43.4 mm, I immature female cl. verse distally H. konstanzoe 31.4 mm, cb. 49.5 mm (IVIC 591).—Rio Villa. Ponce, 44 km N Machala, . 50 m alt., II Nov 1980, leg. H. Hypolohocera aequatorialis Diaz, 70 males, the largest cl. 40.8 mm, cb. (Ortmann 1897) 66.8 mm, 57 females, the largest cl. 32.5 Figs. 1A, B mm, cb. 51.6 mm (IVIC 624).—Canton San Pseudothelphiisa dentata.—Ortmann, 1893: Miguel, 5 km N of Balsapamba, roadside 493 (pro parte ex. b, c). stream feeding into Rio Cristal, Bolivar Potamocarcinus aequatorialis Ortmann, Province, 20 May 1996, leg. R. von Stern­ 1897:317, 319, pi. 17, fig. 5. berg, 7 males, the largest cl. 19.4 mm, cb. Pseudothelphiisa aequatorialis.—Rathbun, 30.5 mm, I juvenile (IVIC 940).—Town of 1898:532. 537.—Young, 1900:213.— Pullatanga. Province. 15 Feb Nobili, 1901:38.—Rathbun, 1905:285.— 1996, leg. R. von Sternberg. 6 males, 2 fe­ Colosi, 1920:18.—Coifmann. 1939:106, males (TV1C 969).—Village of Ocana. Cafi- Strengeria (Strengeria) aequatorerorialis ar Province, 8 Jun 1996, leg. F. von Stern­ |sicj.—Pretzmann, 1965:7. berg, 1 male, 10 juveniles (IVIC 970). Hypolohocera (Hypolohocera) aequatori­ Additional illustrations.—Rodriguez alis aequatorialis.—Bott, 1967:368, fig. (1982, figs. 19k; 22d,i; 23f; 33a-f.) 3a-c—Pretzmann. 1971:17; 1972:43, Diagnosis.—Carapace with upper frontal figs. 186-189, 265-267. margin angled, with faintly indicated papil­ Hypolohocera aequatorialis.—Rodriguez, lae and deep notch at middle. Larger chela 1982:61 (pro parte and fig. 33e, f). with oblong, but not well developed, dark Hypolohocera (Hypolohocera) [aequatori­ protuberance near articulation of fingers alis | aequatorialis aequatorialis.—Pretz• and smaller dark tubercle above it. Lateral mann, 1983b:351, figs. 4. 18. 26. 39. 54. lobe of first male gonopods prominent, 56, 71. square in outline: apex in caudal view fun­ Hypolohocera [Hypolohocera) aequatori­ nel-shaped; in distal view elongated later­ alis nigra Pretzmann. 1968:6; 1972:44. ally and ending in spine directed distally figs. 167-169.262-264; 1971:17. and transversely to main axis of appendage. 1 14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Remarks.—Ortmann (1893) identified as cera) [aequatorialis] delsolari delsolari is Pseudothelphusa dentata three lots of crabs considered as a separate species and Hy­ from South America, Later he (Ortmann polobocera (Hypolobocera) [aequatorialis\ 1897) separated lots b and c. from Rio delsolari Isabella a junior synonym of this. UcuyallL Peril, and the Eastern Cordillera Pretzmann's (1968) original material of of Ecuador, respectively, under his new spe­ Hypolobocera (Hypolobocera) [aequatori­ cies Poramocarcinus aequatohalis, but he alis] aequatohalis nigra comprised 1 male used as types only the specimens from the holotype, 1 male paratype and 3 females, second locality. The first male gonopods of collected by Cayan in 1883 at an undeter­ these specimens (Rodriguez 1982, fig. 33e, mined locality in Ecuador. The first gono­ f) closely correspond with those of the pods of the holotype was illustrated in specimens recorded above from the vicinity Pretzmann 1972. Subsequently, he (Pretz­ of Bailos and from Ponce, near Machala, mann 1977b) recorded the distribution of but the specimens recorded by Rodriguez his taxon as 4"Westrand der Anden nord- (1982, fig. 33a—d) from Rio Jubones belong westlich Machala". However, this distribu­ in Hypolobocera delsolari. Bott (1967) re­ tion must refer to two lots of crabs recorded corded the species from Paramba, on the later (Pretzmann 1983b) from 20 and 35 km headwaters of Rio Mira, 15 km from Tul- NE of Machala in the Rio Jubones basin. cau. Imbambura Province. In the material There are no clear cut characters that sep­ reported above from Ponce, near Machala, arate our specimens collected around Mach­ the first gonopods exactly correspond with ala from the typical H. aequatohalis, even those of the type material and with those of in the coloration of the specimens which our specimens from Bahos: the only differ­ was given as one of the diagnostic charac­ ence is that in the largest male (cl. 40.8 ters. Tn specimens from a single locality mm) from the first loclity the spine of the preserved in alcohol, some specimens are apical lobe is directed laterally and perpen­ dark brown, almost black, on anterior por­ dicularly to main axis of the appendage. In tion, including cervical grooves, while oth­ the specimens trom Ponce the papillae on er have cervical grooves and cardiac aniero-lateral margins of carapace are more regions olive. On the other hand. H. del­ clearly defined. According to these records, solari, H. ftiuistiensis and H. orcesi also dis­ H. aequatohalis occupies widely separated play this last pattern of coloration, with the areas on the eastern and western slopes of cervical grooves and gastric regions of a the Eastern Cordillera of Ecuador. lighter shade than the dorsal surface of car­ Pretzmann used the specific name Hy­ apace. polobocera aequatohalis in several contri­ butions (Pretzmann 1968. 1977b, 1983b). Hypolobocera caputii (Nobili 1901) In his most recent one (Pretzmann 1983b) Fig. 1C, D he grouped under this species the following forms: Hypolobocera {Hypolobocera) [ae- Pseudothelphusa caputii Nobili, 1901: quatorialis] aequatorialis aequatorialis. 38.—Rathbun. 1905:299.—Colosi. 1920: Hypolobocera {Hypolobocera) [aequatori­ 20.—Coifmann. 1939:107.—Rodriguez. alis] aequatorialis nigra, Hypolobocera 1982:190. ( Hypolobocera) [ aequatorialis \ delsolari Strengeria (Strengeria) caput! | sic ].— delsolari. Hypolobocera {Hypolobocera) Pretzmann. 1965:7 (pro parte). [aequatorialis] delsolari isabellcL It is not Strengeria (Strengeria) caputii.—Pretz• possible to discern from this pentanomial mann. 1972:40: 1983b:353. nomenclature whether the author assigned a Hypolobocera (Hypolobocera) caputii ca- subspecific rank to these forms. In the pres­ putii.—Pretzmann, 1971:17: 1972:40 ent contribution Hypolobocera {Hypolobo­ (pro parte) figs. 254, 255. not figs. 270- VOI.UMK I ! I. NUMBER 1 U.S

272, 302, 303 [=Hypoiocera chilensis the holotype, thus validating his report of (H. Milne Edwards & Lucas, 1844)); this species. 1983b:353, tigs. 2, 22. 29, 38, 48, 59. 65. NobilTs species has been recorded in the Hypolobocera {Hypo I oboe era) \chilensis\ literature from Rio Peripa (Nobili 1901, caputii.—Pretzmann, 1977b:436. Pretzmann 1983b), 42 km from Quevedo Hypolobocera quevedensis Rodriguez & (Rodriguez & Diaz 1981. holotype of Hy­ Diaz. 1981:308, figs. 2, 6. 7. polobocera quevedensis): Quevedo and Mindo (Pretzmann 1983b), The latter au­ Material.—Ecuador: Rfo Quevedo, 36 thor gives as the general distribution of Hy­ km N of Quevedo, Province, 24 polobocera caputii the basins of the Daule Jun 1976, leg. H. Diaz, I male holotype of and Vincens rivers. H. quevedensis, cL 26.8 mm, cb. 41.9 mm (IVIC 628).—Puerto Rico. Quevedo, Los Hypolobocera conradi (Nobili 1897) Rfos Province, 3 males cl. 21.5, 17.9 and 14.0 mm, cb. 33.4, 27,8 and 21,2 mm (TU Pseudotheiphusa coti radi NobiI i, 1897:3: 94-100-1, USNM 273521). 1901:38.—Rathbun, 1898:533, 537 (pro Additional illustrations.—Rodriguez & parte): 1905:298, fig. 90a, d (pro parte, Diaz (1981, figs. 2, 6, 7). not materia] from Peru and fig. 90b, c),— Diagnosis.—Carapace with upper frontal Young. 1900:21 7.—Colosi. 1920:19.— margin well defined although not projected, Coifmann, 1939:107.—Rodriguez, 1982: with some tubercles faintly indicated and 63. deep notch at middle. Larger chela with Strengeria (Strengeria) conradi,—Pretz• small swelling on outer surface, at articu­ mann, 1965:7. lation of dactylus. First male gonopods with Potamocarcinus (Hypolobocera) conra­ lateral lobe well developed, long, subtrian- di.—Bow, 1967:367, fig. 2a, b. c. gular, with distal margin angled, advanced; Hypo 1 obocera {Hypolobocera) conradi apex with conspicuous lanceolate lobe di­ conradi.—Pretzmann, 1971:17; 1972:41, rected distally. fig. 273, 274: 1977b:430. tig. 1: 1983b: Remarks.—Nobili (1901) in his original 356, tigs. 74, 79-83. description of Pseudothelphusa caputii did Pseudothelphusa dubia Colosi, 1920:19.— not give an illustration of the male gono­ Coifmann. 1939:107. pods, and they were only vaguely described Hypolobocera (Hypolobocera) dubia.— as "lunghe e robuste, troncate e svasate ob- Pretzmann, 1972:48 (pro parte, not the liquamente ad'apice." Since the holotype material from Colombia and figs. 224- and only specimen recorded could not be 226, 230-232, 236, 237). located at the Museo Zoologico di Torino, Material.—Ecuador: Sabanilla, 18 km where it was presumably to be deposited. NNE from Loja, headwaters of Rfo Zamo- Rodriguez (1982) considered this species ra. Zamora Province, Sep 1985, leg. L. Col- incertae sedis. These circumstances also Jed oma, 2 males cl. 20,7 and 19.1 mm. cb. Rodriguez & Diaz (1981) to erroneously 31.5 and 29.2 mm (IVIC 946). describe their material from Quevedo under Additional illustrations.—Bott (1967, fig. a new species. H. quevedensis. Pretzmann 2a, b, c). (1965. 1971 and 1972) recorded Hypolo­ Diagnosis.—Carapace with upper frontal bocera caputii on several occasions, but margin angled, with flat papillae and deep never stated that he had examined the ho­ notch at middle. Large flat tubercle on the lotype. although subsequently he (Pretz­ insertion of the dactylus of the larger chela, mann 1983b) illustrated the gonopod, car­ fingers conspicuously high. First male gon­ apace, orbital area and third maxilliped of opods slender, with lateral lobe long, re- 116 PROCLHD1NGS OF THF, BIOLOGICAL SOCIETY OF WASHINGTON

Ri!. 2. Hvptifffrtn cm ihhuhin Pretzmann. l^7X. male Ironi Quebrada Celala. A/uay Province, hcuador (IVIC 4W)i: A, iirst lelt uunopod. caudal; B. apex, distal: C. chela ot' largest cheliped. external wevv: IX dtusal view nt" rijihi side til" canipace. K, third maxiHiped- Scale-; : ~S inni. traded; apex with conspicuous lateral ex­ Hypolobocera delsolah Pretzmann. 1978 pansion, in distal view triangular. Fig. 2 Remarks.—The status of NobilPs type Hypolobocera iHypolobocera) [aeyuarori- material was discussed by Rodriguez ( 1982). alis] delsohiri del solan Pretzmann who considered the male recorded by Bolt 1977b:436 inomen nudum); 1978:163, (1970) from Rio Santiago as the neotype of fig. I; I983a:304, rigs. IK 12: 1983b: the species. Our specimens were collected in 350, fig. 58.—Rodriguez, 1982:210. a locality within this basin. Since we have Hypolobocera {Hypohtbocera) [aequatori- examined only the two small males men­ alix] delsolah Isabella Pretzmann 1977b: tioned above, it is not possible to revise the 436 (nomen nudum); 1978:163, fig. 2; description of the species or to present ad­ 1983a:304, tigs. 13. 14;1983b:35.—Rio of carapace smooth and polished, with Chorro, affluent of Rio San Vicente in the regions only slightly indicated. Lateral bor­ Rio Jubones basin, near Giron, Azuay Prov­ der of carapace with shallow postorbital ince, 1 I Nov 1980, leg. H. Diaz, I mule cl. notch, without teeth or papillae up to level 37.6 mm, cb, 60.6 mm, I immature female of cervical grooves; rest of border with ap­ cl. 29.0 mm, cb. 45.3 mm (IVIC 959).— proximately 12 distinct triangular teeth Quera. military checkpoint, border of which diminishes in size progressively and Azuay and El Oro provinces, stream feed­ end at middle of border. Cervical grooves ing into Rio Jubones, 22 May 1996, leg. R. straight and deep, not reaching margins of von Sternberg, chela and portions of cara­ carapace. Postfrontal lobes absent, its place pace of a specimen cl. 36.5 mm. cb. 57.5 marked only by I or 2 punctae: median mm, 4 juveniles UV1C 942).—Village of groove absent. Upper margin of front al­ Tics Banderas. Azuay Province, roadside most straight or slightly convex in dorsal ditch. Rio Jubones adjacent. 22 May 1996, view, without median notch and devoid of leg+ R. von Sternberg. 2 young males, the tubercles. Lower margin sinuous in frontal largest cl. 17.6 mm. cb. 26.1 mm, 2 juvenile view; space between both margins narrow. females (IVIC 941). Palm of larger cheliped (left) moderately Diagnosis.—Carapace with upper frontal inflated, fingers slightly gaping. Exognath margin angled, devoid of papillae, with of third maxilliped 0.4 length of ischium of deep notch at middle. Larger chela with endognalh. large rounded tubercle below articulation oi' First male gonopods strongly arcuate in daetylus. First male gonopods with lateral lateral view; caudal ridge curved, becoming lobe long, oblong, wider proximally; apex indistinct distally; lateral lobe narrow, ex­ wilh conspicuous lanceolate lobe directed cavated and more prominent distally: apex distally. in distal view with strong curved point pro­ Remarks.—The apex of the first gono­ jected laterally and distally, in caudal view- pods in this species resembles that o\ Hx­ very elongated. polohocera caputa (Nobili, 1901) in the lat­ Erymoltrgy.—The species is named after eral apical process, but differs in the shape the where the species of the lateral lobe. was collected.

Hxpolohocera esmeraldensis* new species Hxpolohocera cxnca Pretzmann. 1977b Fig. 3 Fig. 4 Hxpolohocera tHxpolohocera) \ conradi] Mate rial—Ecuador: Esmeraldas Prov­ exuca Pretznumn. 1977b:437. fig. 8: ince, leg. Juan Carlos. I male holotype cl. 1983b:357, figs. 91-94. 20.6 mm. cb. 33.8 mm, 1 immature female Hxpolohocera riven' Rodriguez. 1980:891; cl. 20.9 mm, cb. 33.6 mm (TU 94-100- 1982:49. rigs. 19b. 20e. j. 23b. 25a-e. 2).—Chone River. Manabf Province, 1 ma­ ture male, broken carapace (TU 94-100-3). Materia!.—Ecuador: 10 km N of La Diagnosis.—Carapace with upper frontal TroncaL on Rio Culebras/Taura. Caiiar margin devoid of median notch and tuber­ Province, in a concrete storm drainage cles. First gonopods with caudal ridge ob­ ditch, 6 Jun 1996. leg. R, von Sternberg. 1 solescent distally: lateral lobe narrow, more male cl, 37.6 mm. cb. 59.3 mm. I female ] IK PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. }. Hyfutlohtneta vsmerukdvnsis. new species, holotype male from Esmeralda Province, Ecuador fTLJ 94-100-2): A, firs? left gonopod. cauda); B. apex, distal: C, chela of largest cheliped. external view; D. dorsal view of riizht side of carapace: E, third maxilliped. Scales - 3 mm. cl. 21.1 mm, cb. 33.2 mm, 9 juveniles shaped, with strong elongated projection (IV1C 949).—Between La Troncat and Ma­ ending in truncated tip; in distal view nuel J. Calles, 65 km SE of , strongly expanded cephalically, with con­ Canar Province, 100 m alt., JO Nov 1980, spicuous ridge on caudal side of expansion; leg. H. Diaz. 1 immature male cl. 18.2 mm, prominent subtriangular papilla on caudal cb. 27.2 mm (IVIC 627).—Town of Las side of gonopore; distinct subapical ridge Pampas, on Rio Toachi, Province, on mesial side. Oct 1988, leg. G. Onore, 1 male cl.32.4 Remarks.—Pretzmann (1977b) gave as mm, cb. 50.5 mm (IVIC 950).—Ecuador, his type locality 'Cordillere". The present without other data, leg. P. Rivet, 2 males cl. records fix the area of distribution of the 40.8 and 23.6 mm, cb. 61.8 and 37.6 mm, species between the provinces of Canar and holotype and paratype respectively of Hy- Cotopaxi. polobocera riveti Rodriguez, 1980 (MNHN B-5087). Hypolobocera guayaquilensis Bott, 1967 Diagnosis.—Carapace with upper frontal Fig. 5 margin angled, with small papillae and deep notch at middle. First male gonopods with Potamacarcinus (Hypolobocera) aequato- lateral lobe absent, replaced by wide de­ rialis guayaquilensis Bott, 1967:368, pression; apex in lateral view funnel- figs. 4a, b, c. YOLL'Mk I 11. NtMBKR I 119

Hii 4. Ih-paltihtH-era t'\mn Fret/mann, I l>77h. A-G. male from ID km N of La TYoncal. Omar Province. licuaJor lIVR" LMCJ); A. lirsi left gonopod. caudal; B. lateral; C. apex. cando-distal: D. apex, distal: H. chela vA' laiLicsi cheliped. external view. K dorsal view of right side of carapace: G. third niuxittiped. hL specimen t'mm l.as Pampas Colopaxi Province. Kcuador ([VIC *J?0|. first left gonopod. apex, distal. Scales = 3 mm.

Hypoiobocera {Hypoiobocera) caputii gu- tubercles, with notch at middle. First male ayaquilensis.—Pretzmann. 1971:17 (by gonopods strongly arcuate in lateral view; inference). caudal ridge prominent, moderately curved Hypoiobocera (Hypoiobocera) gitayaqni- proximally, straight distally; lateral lobe tri­ lensis.—Pretzmann. 1972:42, rigs. 173- angular, increasing in width distally. with 175. textrig 10. Hypoiobocera {Hypoio­ distal margin rounded: apex in distal view bocera) laequatf>rialis\ guayaqui lea­ elongated along meso-lateral axis, ending in sts.— Pretzmann, 1983b:353, rigs. 5, 16. short lateral point directed laterally. 2X. 37, 53, 57. 69. Remarks.—The type locality is Babahoyo. Hypoiobocera gitayaquilensis.—Rodriguez, on the Daule-Vincens basin. The specimens 1982:64. reported above come from a locality 140 km NNW of Babahoyo. on the coastal plain, but Material—Ecuador: Chone. Estero Don- the water divide between both basins, on the de, Manabi Province, 9 Aug 1967. 3 males Conguillo Mountains, is less than 300 m el. 26.2. IS.3 and 14,7 mm, cb. 44.1, 29.3 above sea level in some places, and 23.5 mm, 1 female with young under the abdomen cl. 34.5 mm. cb. 56.2 mm (TL1 Hypoiobocera konstanzae. new species 6374). Fig. 6 Diagnosis.—Carapace with upper frontal Material.—Ecuador: Estero San Agustin. 4 margin angulated and devoid of conspicuous km S from the bridge. Rio Banchal. Manabt PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

hi^, 5. HvpulohtKcm t>uaytufitilcrixi\ Butl. 1967. male I'roin Chone. Manubi Province:, Ecuador (Tl i 6374); A. tirsi left gnuopoil. cuudjil: R. upc.v distal: C. chela of largest chclipcd. exlemal vicu: D. dorsal view of right Mile ot carapace: H. third mnxilliped. Scales = 3 nun.

Province. 6 Jun 1977, leg. H. Dfaz, 1 male Description of holotype.—Carapace nar­ holotype cl. 37.5 nun. cb. 56.8 mm (IVIC row (cb/cl = 1.51), surface smooth. Antero­ 593).—Village of Caseol. stream adjacent to lateral margins without shallow notch behind Rio BanchaL lower Rio Daule basin, Manabi external orbital angles, margin behind it and Province, approx. 1° 40' S, 80° 30' W, 4 Jun up to level of cervical grooves smooth, al­ 19%. leg. R. von Sternberg, 2 young males most straight: approximately 8 small papilli- paratypes (the largest soft shell) el. 21.8 and form teeth behind level of cervical grooves, IS.9 mm, cb. 33.5 and 29.1 mm respectively, progressively less prominent posteriorly. Cer­ 3 juvenile males, 3 immature females, the vical grooves almost straight, deep, not reach­ largest cl. 22.3 mm. cb. 34.3 mm {1V1C 951). ing margins of carapace, Postfrontal lobes Diagnosis.—Carapace with upper frontal well delined, transverse, with anterior margin margin angled, devoid of papillae, with deep rounded; median groove well defined, very notch at middle. First male gonopods with narrow anteriorly, making deep incision on lateral lobe reduced, subtriangular. more ex­ upper frontal margin of carapace. Upper fron­ panded distally; apex in caudal view runnel- tal margin bilobed in dorsal view; angled, de­ shaped, moderately elongated laterally, not void of papillae; lower margin strongly sin­ ending in defined spine; in distal view rough­ uous; space between upper and lower mar ly obtuse-angled, lateral expansion rounded. gins narrow, voi IMI J 11. NLMBE:K I 1:1

Fig r> flvpnitifai<-i'f\t konsttwztit'. neu species. Iu>lnt\pc male fmm Rio Banchiil. Manabi Prmincu. Ecuador il\'K" 5^M' A. tirst let'i iionopod, caudal; H, ape\. distal; C. chela of largest ehehjvd. external view; D. dorsal \ic« of ne!u MIII1 ol carapace; h. rhird ma\il1iped. Scales - _i mm

Exi)gnath of third maxilliped 0.30 length terolateral margins possess 10-12 small of ischium of endognath. Chelipeds elongat­ well defined teeth behind level of cervical ed. Fingers gapping, covered by punctae and grooves, which are progressively less prom­ few inconspicuous papillae. inent posteriorly; the cervical grooves are First male gonopods with caudal ridge straight, and reach the margins of carapace: proximally strong and curved to follow stran­ the upper frontal margin is covered with ru­ gled shape ol basal portion: distally progres­ dimentary papillae; the lower margin is sively indistinct: lateral lobe reduced, sub- moderately sinuous. triangular, more expanded distally: apex in Etymology.—The species is dedicated to caudal view funnel-shaped, moderately elon­ Miss Konstanza von Sternberg, for her as­ gated laterally, not ending in defined spine: sistance in the collection of crabs in Ecua­ in distal view roughly obtuse-angled, lateral dor, expansion rounded; papilla on caudal side of gonopore replaced by semicircular ridge. Hxpolobocera mitulonensis* new species Remarks.—The paratypes display the fol­ Fig. 7 lowing differences in regard to holotype: The carapace surface has small pores and Materia!,—Ecuador: Confluence of Rio papillae not visible to naked eye; the an­ Salaya and Rio Mindo, Pichincha Prov- 122 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fie. 7. HypofolwH'i'ru nihulotwtisix* new species, holotype male from the confluence of Rfo Salaya and Rfo Mindo. Ecuador (TU 44-100-4): A, rirst Lett gonopod. caudal: B. laieral: C, apex, caudo-distal: D. apex, distal: b. chela of largest ehehped. external vilew : K dorsal view of right side of carapace: G, third maxilliped. Scales = 3 mm. ince. between 1000 and 1200 m altitude, 23.0 mm, with 13 and 18 eggs respective­ Sep 1968, leg. M. Olalla, 1 male holotype ly, 10 mature female paratypes cl. 13.6— cl. 11.3 mm, cb. 27.1 mm, 14 male para- 22.3 mm, cb, 20.5-33.4 mm, 6 immature lypescl. 16.9, 14.7, 14.7, 14.4, 14.4, 14.2, females, the largest cl. 17.0 mm, cb. 26.2 14,2, 13.7, 13.6, 12.9, 12.0, I 1.5, 11.0, mm (TU 94-100-4). and 10.9 mm, cb. 25.2, 23.9, 23.2, 22.2, Diagnosis.—Carapace with upper fron­ 23.7, 23.3, 23.1, 21.4, 21.0, 19.4, 18.1, tal margin rounded, with small papillae 17.5, 16.5, and 16.4 mm, 2 ovigerous fe­ and deep notch at middle. First male gon- males cl. 15.5 and 15.1 mm, cb. 24.9 and opods with lateral lobe narrow and regu- WM.UME in. M-Mbhk i 123 larly rounded in outline, covered with 952).—Estero E) Canero, La Concordia, minute flattened papillae; apex in caudal Esmeraldas Province, I Oct 1988, leg, J. view funnel-shaped; elongated in distal C. Vieira, I male cl. 20.7 mm, cb. 31.9 view, strong triangular tooth on lateral mm (1VIC 953).—Same data, 8 Oct 1988, corner and conical elevated tubercle on I male cl. 24.2 mm. cb. 39.5 mm HV1C caudal side of gonopore. 954).—Estero Moneauve, Recinto Mon- Description of hoIotype.—Cervical cauve, Esmeraldas Province. 5 Jan 1988. grooves deep and wide proximally, shal­ leg. J. C. Vieira, 1 male cl. 24.2 mm, cb. low and straight distally. not reaching 39.5 mm (IVIC 955). margins of carapace, Antero-lateral mar­ Diagnosis.—Carapace with upper fron­ gins with well defined incision behind tal margin well defined, angled, without outer orbital angle and several ill-defined conspicuous papillae, with deep notch at papillae behind it; tooth at level of cer­ middle. First male gonopods with lateral vical grooves and 8-10 very small teeth lobe triangular, increasing in width distal­ over rest of margin. Posttrontal lobes al­ ly. with distal margin excavated; apex most obsolete, its place indicated by 2 elongate along meso-lateral axis in distal slight swellings; median groove shallow. view, cephalic margin rounded, horseshoe wide. Upper margin of front rounded, shaped. slightly bilobed in dorsal view, with small Description of holotype.—Lateral bor­ papillae which are obsolescent towards der of carapace with shallow notch behind sides, and deep notch at middle: lower outer orbital angle, followed by short un­ margin moderately sinuous in frontal dulated segment; 12-16 very small trian­ view; space between upper and lower gular teeth behind level of cervical margins narrow. grooves, regularly-spaced, subequal in Exognath of third maxilliped 0.30 size except for last 3-4 which are squam> length of ischium of endognath. Larger iform. Cervical grooves deep and wide, cheliped with palm inflated, smooth; fin­ slightly arched, reaching margin of cara­ gers slightly gapping, with brown-black pace. Posttrontal lobes well marked, with punctae arranged in parallel lines. distal margin transverse; median groove First male gonopods strongly constrict­ well defined, forming deep incision at up­ ed at middle; caudal ridge curved, strong­ per border of front: this border well de­ ly contoured proximally, becoming indis­ fined, angled, without conspicuous papil­ tinct distally; lateral lobe narrow and reg­ lae; lower margin thick, strongly sinuous, ularly rounded in outline, with minute advanced in relation to upper margin; flattened papillae; apex funnel shaped in space between upper and lower margins caudal view; elongated along latero-me- narrow. sial axis, with strong triangular tooth on Exognath of third maxilliped 0.35 lateral corner in distal view; conical ele­ length of ischium of endognath. Chelipeds vated papilla on caudal side of gonopore. heavy, ringers gaping, with rows of min­ Etymology.—The species is named af­ ute dark points over outer surface. ter Rio Mindo, where it was collected. First male gonopods strongly arcuate in lateral view; caudal ridge prominent, Hypolobocera muisnensis, new species. curving proximally, straight distally; lat­ Fig. 8 eral lobe triangular, increasing in width distally, with distal margin excavated; Mate rial.—Ecuador: Estero Lojca apex in distal view elongated along meso- More, Muisne Salima, Esmeraldas Prov­ lateral axis, cephalic margin rounded. ince, 21 Oct 1988, leg. J. C. Vieira, 1 male horseshoe shaped. holotypc cl. 31.1 mm, cb. 51.6 mm (1VIC Etymology.—The specific epiteth is de- 124 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Hg,. S. Hvfwlohoctrti mi4t\ni'n.\i.\, new species, liolorype male from MuiMie Sidima. Bcuiirior (IV1C 9^21: A. tirsl left jionopoik IL apex. dist.il: C. chela of largest eheliped, external view: D. dorsal view of right side of carapace; E. thud maxilliped. Scales = 3 mm. rived from part of the locality's name Diagnosis,—Carapace with upper frontal where the species was collected, margin rounded, devoid of defined papillae. First male gonopods with caudal ridge Hypolohocera orcesi Pretzmann, 1978 strongly geniculated proximally, indistinct Fig. 9 dis tally; lateral lobe undifferentiated in caudal view, forming thin ridge in lateral view; apex Hypolohocera iLindacatalina) [plana] or­ funnel-shaped in caudal view, projected ce- cesi Pretzmann, 1978:166, fig. 6: 1983b: phalicaily in lateral view, subtriangular in dis­ 361. tal view, with lateral margin rounded; 2 Hypolohocera (Lindacatalina) orcesi,— prominent papillae near gonopore and on ce­ Pretzmann, 1983a:303, pi. 7, 8. phalic expansion. Material.—Ecuador: Valley of Rfo Mindo, Remarks.—The distal angle of the lateral 5 km from Mindo, , 28 lobe of the gonopod is more squarish in May 1996, leg. R. von Sternberg, 1 male ch Pretzmann's (1978, fig. 6). Otherwise our 14.2 mm. cb. 23,5 mm. 1 male soft shell cl. materia] closely corresponds with the original 11.1 mm, ! juvenile male. 1 mature female description and with the supplementary char­ cl. 14,8 mm, cb. 24.3 mm (IV1C 956), acters given by Pretzmann (1983a. b). The VOLLML I II. M'MBbK I 125

Fig. 4. Ihpttlabttvvra urcvsi Pret/mann, I lJ7K. male from valley of Rio Mirulo. Feuador (I VIC* 456): A. tirsl left iionopml. caiHlul: B. lateral: C eephalie. D. apev distal: H. chela o\~ largest eheUpcd. external view; F. dorsal \ieu of nt: In Mile of" carapace. G. third inavilliped Scales = 3 mm.

rvvo prottuneni papillae, one near the gono- Hypolohocera (Hypolohocera) guayaqui- pore and another on the cephalic expansion lensis rathhuni.—Pretzmanru 1972:42. of the apex, are characteristic of this species. rigs, 287-289. Hypolohocera {Hypolohocera) caput is rath­ Hypolohocera rathhuni Pretzmann. 1968 huni.—Pretzmann, 1983b: 354, figs. 1, Fig. 1E-F 23, 30, 40, 51, 60, 64. Hypolohocera {Hypolohocera) rathhuni Hypolohocera rathhuni.—Rodriguez* 1982: Pretzmann. 1968:5. 63. rig. 19p: 22b. g: 23e: 34a-c. I2h PROCEEDINGS OE THE BIOLOGICAL SOCIETY OF WASHINGTON

Material.—Ecuador: Santo Domingo, Pi- type which was not observed in the rest of chincha Province, 490 m alt., 1 male neo- the material listed above. type (Rodriguez 1982) cl. 23.7 mm, cb. 37.5 mm IBM 1918. 1.31.12).— Rio Peripa, Lindacatalina Pretzmann, 1977b between Aurora and Puerto Limon, SW of Santo Domingo de los Colorados, Pichin- Diagnosis.—Exognath of third maxilli- cha Province, 29 Nov 1980, leg. H. Diaz. ped usually more than 0.45 length of ischi­ 18 males, the largest cl. 27.7 mm, cb. 45.0 um of endognath (Table 1). First male gon­ mm. 1 mature female cl. 22.2 mm, cb. 35.9 opods with strong longitudinal ridge on mm, 12 immature females, the largest cl. caudal surface: well developed lateral and 19.6 mm, cb. 31.1 mm (IVIC 631).—Rio supplementary cephalic lobes (this last rare­ Peripa, Puerto Limon, SW of Santo Domin­ ly absent), both covered by minute spinules; apex truncated, circular in distal view, with go de los Colorados. Pichincha Province, two flat papillae near spermatic channel 200 m alt,, 29 Nov 1980, leg. H. Diaz, 1 (Fig. IK, L, M). male cl. 18.2 mm, cb. 28.6 mm, 1 spent Type species.—Hypolobocera (Lindaca­ female cl. 29.3 mm. cb. 46.4 mm. 1 im­ talina) hauserae Pretzmann. 1977b. mature female cl. 17.0 mm, cb. 26.7 mm, Distribution.—Southern Colombia and 2 juveniles (IVIC 629).—Rio Peripa, San Ecuador. Miguel, 5 kin from Aurora, SW of Santo Remarks.—We use Pretzmann's genus to Domingo de los Colorados, Pichincha Prov­ group all Hypolobocerini with the lateral ince, 29 Nov 1980, leg. H. Diaz, 1 imma­ lobe of gonopods densely covered by spi­ ture male cl. 14.3 mm. cb. 22.1 mm, 1 im­ nules and frequently possessing a supple­ mature female, 3 juveniles (IVIC 630). mentary lobe, equally spinulous, on the ce­ Additional illustrations.—Rodriguez phalic side. We exclude from this genus two (1982, figs. 19p: 22b, g: 2^: 34a-c). species that were included by Pretzmann Diagnosis.—Carapace with upper frontal (1977b), Hypolobocera orcesi which has a margin well defined by row of distinct pa­ few sparse spinules on the lateral lobe, but pillae on each side and deep notch at mid­ not a continuous covering of this process, dle. First male gonopods with caudal ridge and H. nobili whose holotype is a female, prominent and curved proximally, obsoles­ and consequently their gonopods are not cent on distal half: lateral lobe long, nar­ known. We place in this genus Hypolobo­ row, slightly expanded distally, with outer cera brevipenis Rodriguez & Diaz, 1981. border sinuous; apex in caudal view trans­ and a new species, L. sumacensis. Thus de­ verse, ending laterally in long spine: in dis­ fined, the genus consist of an homogeneous tal view narrow, very elongated laterally, group of species restricted to a small area ending laterally in long acuminate spine: on the Amazonian drainage of Southern flat digitiform papilla on caudal side of gon- Colombia and Ecuador. opore. Remarks.—There are slight differences Key to Species of Lindacatalina between the neotype from Santo Domingo de los Colorados (Rodriguez 1982) and the \. Lateral lobe of first gonopods with sup­ specimens from Rio Peripa. The relation­ plementary cephalic lobe 2 ship between the length of the exognath and -. Lateral lobe without supplementary lobe 5 the ischium of endognath in the third max- 2, Lateral and supplementary lobes fused illiped of the neotype is 0.28, whereas in distally 3 the others specimens range between 0.37 -. Lateral and supplementary lobes distinct and 0.39. The flat papilla on the apex of 4 gonopods has a minute denticle in the neo­ 3. Supplementary lobe in cephalic view al- VOl.UMT- III. NUMBER 1 127

mosl pyramidal ur strongly excavated on Diagnosis.—Carapace without upper the mesial side , L. pttyensis frontal margin. First male gonopods slen­ -. Supplementary lobe in cephalic view der widening progressively proximally; globular L. latipenis apex oval-elongated in distal view, wider 4, Lateral lobe wide, rounded, distinct; sup- laterally than mesially; lateral lobe wide. plemcntary lobe iriangular, not expanded placed in line with main axis of appendage; distally - L. brevipenis -. Lateral lobe narrow, partially fused to caudal ridge slightly rounded. caudal ridge; supplementary lobe round­ Remarks.—We have not seen material of ed, expanded distally L. sttmarensi.s this species, which is the type species of the 5. Lateral lobe square; wide apical portion genus Lindacataliua, The diagnosis given almost reaching apex L. hauserae above was derived from Pretzmann's -. Lateral lobe rounded, not reaching apex (1977b, 1983a, b) diagnoses and keys. The L. orientails only specimens known, six males, three fe­ males, and six juveniles came from 2 km Uudacatalina brevipenis (Rodriguez & East of Mendez, Morona-Santiago Prov­ Diaz 1981) ince. Figs. 1K-M: 12F, G Lindacataliua latipenis (Pretzmann 1968) Hypolobocera brevipenls Rodriguez & Figs. 1N-P; 12A-C Dfaz, 1981:309, figs. 3, 8, 9. Hypolobocera {Hypolobocera) latipenis Material.—Ecuador: without other data, Pretzmann, 1968:8. leg. M. Olalla, 1 male holotype cl. 15.9 Hypolobocera {Hypolobocera) conradi la­ mm, cb. 27.7 (SMF 9140), 1 male paratype tipenis.—Pretzmann, 1971:17; 1972;41, cl. 12.0 mm, cb. 20.3 mm fIVIC 606). figs. 281-283. Additional illustrations.—Rodriguez & Hypolobocera (Lindacatalina) latipenis la­ Diaz (19X1, tigs. 3, 8, 9). tipenis.—Pretzmann, 1977b:432. figs. 5. Diagnosis.—Carapace with upper frontal 6, 1 I. margin rounded, devoid of tubercles. First Hypolobocera (Lindacataliua) [latipenis] male gonopods very short and stout; caudal latipenis latipenis.—Pretzmann, 1983b: ridge strong, geniculated and wrinkled at 357, figs. 12, 21, 34, 45, 49, 63, 68, 77. middle; lateral lobe auriculariform. wrin­ 78, 90. kled; supplementary cephalic lobe forming Hypolobocera latipenis.—Rodriguez, 1982: winged triangular expansion, distinct from 54, figs. 19n; 20a, f; 23a; 8a-e. lateral lobe, both lobes covered by minute spinules; apex oval and expanded laterally Material.—Ecuador: Faldas del Monte in distal view. Sumaeo, Loreto, , 450 m alt., Remarks.—This species can be easily Jun 1968, leg. M. Olalla, 2 males, 1 female distinguished from other within the genus {TU 94-100-5).—Ecuador, leg. M. Olalla, 1 because the lateral and supplementary lobes male cL 33.8, cb. 55.7 mm (1VIC 621). are distinct and resemble each other in Additional illustrations.—Rodriguez shape (Fig. 121"). (1982:54, tigs, 19n; 20a, f; 23a; 8a-e). Diagnosis.—Carapace with upper frontal margin well marked, with scattered tuber­ Lindacataliua hauserae Pretzmann, 1977b cles. First male gonopods with caudal ridge Hypolobocera (Lindacataliua) hauserae distinct and strongly geniculated; lateral Pretzmann, 1977b:437, fig, 1.0; 1983a: lobe large, wide, rounded; supplementary 301, pis. 1,2. cephalic lobe forms large digitiform process Hypolobocera (Lindacataliua) [nobili] hau­ transversely directed and fused distally to serae.—Pretzmann, 1983b:358, fig. 10. lateral lobe; both lobes covered by wrinkles I:N PROCEEDINGS OF TIIF, BIOLOGICAL SOCIETY OF WASHINGTON and spinules, apex in distal view circular, 1994, leg. J. Garces, 1 male cl. 16.2 mm, with wide Mat papilla near gonopore. cb. 28.0 mm (TVIC 957). Remarks. — Pret/.rnann (1972, 1983b) Diagnosis.—Carapace with upper frontal gives as the original citation of this species margin rounded, devoid of defined papillae. Strengcria (Strengeria) iatipenis Pretz­ First male gonopods with caudal ridge mann, 1965. However, the specific name strongly geniculated at middle, progressive­ did not appear for the first time in Pretz­ ly tapering to end near apex; lateral lobe mann ( 1965), but latter, in Pretzmann very broad, extending from middle of ap­ ( 1968) as Hypolobocera (Hypolobocera) pendage to near apex, rounded, covered by Iatipenis. minute spinules on lateral surface; apex truncated in caudal view, oblong, expanded Lindacatalina orientalis (Pretzmann 1968) laterally into rounded projection in distal Fig. 10 view. Remarks.—The type material of Pseu- Hypolobocera {Hypolobocera) plana orien­ dothelphusa plana Smith, 1870. consisted tals Pretzmann, 1968:2: 1971:17: 1972: of 2 females (cl 16,6 and 13,6 mm. cb, 27.7 60, figs. 162-164, 214-221, and 22.4 mm) from . Peru, in the Mu­ Hypolobocera (Lindacatalina) [plana] seum of Yale College, collected by Prof. plana oriental is.—Pretzmann, 1983 b: James Orton. Smith (1870) description of 360, figs. 8. 15. 33. 36, 46, 73. carapace and appendages, although de­ Hypolobocera oriental is.—Rodriguez, tailed, are generic for many species of Pseu- 1982:52, tigs, 19d, 20c. h, 26a-c. dothelphusidae and he did not include il­ Hypolobocera [Hypolobocera) plana plana lustrations of the gonopods. The species Pretzmann, 1972:49, figs. 275-277, 304- was latter cited in the literature by Pocock 307. Not Pseadothelphusa plana Smith, (1889), Nobili (1897), Rathbun (1898, 1870:146, 147.—Pocock, 1889:10.—No- 1905), Young (1900), Coif man n (1939). bili, 1897:3, 5—Ralhbun, 1898:535, and Ortmann (1897). but any of these au­ 537—Young. 1900:211—Rathbun, 1905: thors examined materials of the species. 278—Coifmann, 1939:109—Rodriguez, The types that, according to Smith 1982:192. Not Potatnocarcinits plan­ (1870), were "rather badly preserved spec­ us.—Orimann, 1897:318 (see Remarks). imens," deteriorated further, and latter, Hypolobocera {Lindacatalina) [plana] Pretzmann (1972) stated that they were no plana plana.—Pretzmann, 1983b:359, longer available. Consequently he renamed tigs. 9, 14, 31, 41, 70. the species as Hypolobocera (Hypolobo­ Material,—Ecuador: Oriental Cordillera, cera) plana plana* proposed as neotype a Ecuador, 1874, leg, Reiss, 1 male holotype male from Ecuador in the USNM (labelled cl. 14.3 mm, eh. 22.3 mm, 7 males para- as follows: Mindo, Pichincha Province, Ec­ types cl. 12,8, 12.6. 12.1, I 1.9, I 1.5. 10.1 uador. 1919 Irwing Expedition, leg. C, N, and 9.9 mm. cb. 20.7. 19.6. 19.2. 18,9, Eigenmann, cl. 7.9 mm. cb. 13.9 mm. 17.3, 15.4 and 15.0 mm, 2 immature fe­ USNM 68558), and also included under males cl. 9.9 and 9.7 mm. cb. 14.7 and 14.3 this species 10 males and 1 I females from mm

Fit! 10. i.tfhituuhilitni tmt'nniHf iPrct/m;mn. L468i: AG. male from ID km S of Mindo. Ecuador (IVK"

<->^Ki \+ first loft gonorunJ. caudal; B, lateral; C. cephalic. D, ape\, disial; ti, chela of largest chehped, external ucw; L lUnvil view of ndn side ci carapace: G. third nia\illiped. H. male from Mindo ilV'JC *->?7i. ;tpe\ of hrsl ^nnopotJ. disial. Scales = 3 mm. theiphusa plana as incertae sedis on the tie his species to the material examined by grounds that the original type locality (Pai- Pretzmann in the USNM, ta) was isolated by a desert (Tumbez), and The gonopods of our specimens from the neotype locality (Mindo) was 600 km Mindo recorded above, and those of Pretz- to the north, on an entirely different river mann's (1972) neotype (USNM 68558) are basin. Furthermore, there is no diagnostic identical to the illustrations of the gonopods character in Smith's original description to of Hypolobocera iHypolobocera) plana or- Un PROCEEDINGS Ol THE BIOLOGICAL SOCIETY OF WASHINGTON tentaiis Pretzmann. 1968 (see Pretzmann and of L. latipenis come from localities 1972, 1983b). within a small area between Puyo and Tena, Another related taxon erected by Pretz­ and north of Tena. The specimens reported mann (1977). Hypolobocera {Lmdacatali- above are from localities 22 km and ^5 km na) [plana] plana olallaL cannot be differ­ from the type locality of L. puyensis. The entiated from Hypolobocera {Hypolobo­ type locality of L. latipenis is 40 km ENE cera) \plana\ plana plana i=Lindacatalina from Archidona. All these localities are lo­ oriental is) from the diagnosis or the cated in a few small river basins draining sketchy illustration of the gonopod given by to the upper reaches of the Napo River Pretzmann (1978. 1983b), We have found The first gonopods of the two species no specimens that could he attributed to Hy­ closely resemble each other, but in L. puy­ polobocera {Limlacatalina) [plana] plana ensis the supplementary lobe in cephalic olallai in our collections from the type lo­ view is oblong, almost pyramidal, and cality (Aurora, Rio Peripa) of this taxon. strongly excavated on the mesial side (Fig. I 2D. E). whereas in L. latipenis it is clearly Limlacatalina puyensis Pretzmann. 1978 globular (Fig. 12A, B. C). The carapace Fig. 12 D, E breadth of all male specimens attributed to L. puyensis (3 type specimens and present Hypolobocera (Lindacatalina) laiipenis records) is less than 35 mm, while L. lati­ puyensis Pretzmann, I977b:438 (nomen penis attains a carapace breadth of at least nudum); 1978:165, rig. 7; 1983a:302. pi. 58 mm. It is possible that the male speci­ 5, 6. mens attributed to L. puyensis are younger Hypolobocera {Limlacatalina) [latipenis] specimens of L. latipenis, or populations of laiipenis puyensis.—Pretzmann, 1983b: dwarf individuals, a phenomenon that has KX. lig. 11. been described in other Brachyura (Conde Material.—Ecuador: Teniente Ortiz, 10 et al. 1989). This situation could be solved km N of Puyo, Rio Rebadeneira, affluent of only by the discovery of mature females of Rio A raj u no. Pasta/a Province. 980 m alt., small size. Pretzmann (1983a. b) recorded 5 Nov 1980, leg. H. Diaz, 1 male cl. 19.8 small females, but did not state whether mm. cb. 32.3 mm (IVIC 626).—Archidona. they were mature. We are keeping both spe­ Cuevas de San Bernardo. Napo Province. cies distinct until more information is forth­ 12 Jun 1986. leg. P Villamar, 3 males cl. coming. 16.8, 10.8 and 9.9 mm. cb. 29,0. 17.4 and 16.5 mm, 1 juvenile female, carapace bro­ Limlacatalina sumacensis. new species ken, cb. aprox. 9 mm tlVIC 947).—Village Figs. II, I2H, I of Talac. Napo Province, 1500 m. 28 Dec 1993, leg. S. Baez, 1 male cl. 18.0 mm, cb. Material.—Ecuador: Faldas del Monte 30.4 mm (IVIC 962). Sumaco. Loreto, Napo Province, 450 m Diagnosis.—Carapace with upper frontal all., Jun 1968, leg. M. Olalla. 1 male ho- margin well marked. First male gonopods lotype cl. 21.9 mm, cb. 35.6 mm, 10 with caudal ridge distinct and strongly ge- males paratypes, 27 females, 7 of the larg­ nicuhited; lateral lobe large, wide, rounded: est mature females as follows: cl. 22.8. supplementary cephalic lobe forms large 20.9,20.6,20.5, 19.1. 19.1. and 18.4 mm, conical process transversely directed and cb. 35,3. 32.9. 33.8. 34.9. 30.5. 29.3. and fused distully to lateral lobe; both lobes 29.1 mm

J \ ( i II, ^N 1% P •• )

Fiji. 11 I.inJtutitiiiitHi \itmncrn.\i\. new species. hoUitype male from Monte Sumaco. Napo Province. F.c- uador

margin angled, without conspicuous papil­ Cervical grooves wide and shallow, becom­ lae, with inconspicuous notch at middle. ing indistinct toward margins of carapace. First gonopods with caudal ridge straight, Postfrontal lobes small, but well defined. tused to lateral lobe, covered by minute placed transversely in relation to middle transverse wrinkles; lateral lobe rounded, axis of carapace; median groove wide and expanded distally; supplementary cephalic shallow. Upper margin of front rounded in lobe rounded, thick, covered by minute spi- dorsal view, with inconspicuous notch at nules; apex oval in distal view. middle, angled in frontal view, without con­ Description of holotype*—Antero-lateral spicuous papillae; the lower margin strong­ margins with wide notch after external or­ ly sinuous; space between both margins bital angles and another at level of cervical very narrow. Upper surface of carapace grooves; between these two notches and for smooth and polished, covered by closely short space behind second one, border de­ placed papillae not visible to naked eye. void of teeth or papillae; towards the mid­ Chelipeds strongly unequal; paim of dle of border begins series of approximately larger one (right) inflated; fingers strongly \5 small but well defined triangular teeth. gaping; movable ringer strongly arched. Ex- 132 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 12. First male gonopods of Undacatalina, A. B. C, L, tatipenis (Pretzmann. 1968). from Ecuador (IVIC 621); D.E, L. puyensis Pretzmann. 1978. from Teniente Ortiz (IVIC 6261; EG. L. brevipenis (Rodriguez & Diaz. 1981). from Ecuador (IVIC 606); H. I. L. sumacensis new species, hulolype male from Monte Sumaco (TU 94- 100-6). (A.D.EH. cephalic; C. disto-cephalic: B.E.G.I. distal). \Ol.r\U- l I L NUMBER I m ognaih of third maxilliped 0.46 length of small closely set spinules; caudal end ischium of endognath. produced in sirong tingerdike process di­ First male gonopods slender, strongly rected proximally; elongate process over arched doj so-vent rally: caudal ridge field of spines formed by one papilla, displaced towards lateral expansion, its straight, fused with lateral lobe, covered by distal margin bordered by minute spi­ minute transverse wrinkles; lateral lobe nules M. narinnensis (Colombia) rounded, expanded distally; supplementary 2. Llongated process over field of spines cephalic lobe rounded, thick, covered by placed near opening of spermatic chan­ minute spines; outline of apex oval in distal nel. Adult specimens very large (more view. than 6 cm eh.* . A/, henrici Etymology.—This species is named after -. Llongaied process over field of spines Monte Sumaeo, where it was collected, displaced towards lateral expansion. Adult specimens very small leb less than Moritschus Prel/mann. 1965 .1 cm) M. ccutuiorensis Diagnosis.—Exognath of third maxilli­ Moritschus ecuadorensis (Rathbun 1897) ped usually more than 0.30 length of ischi­ Fig. 1G. H um of endognath (Table I K First male gon­ opods with strong longitudinal ridge on Pseuditthelphusa ecuadorensis Rathbun. caudal surface; lateral expansion continuos 1897:59; 1898:534, 537; 1905:279, fig. 7, with apex of appendage; apex truncated. pL 13, rig. 8.—Young, 1900:210— No­ very elongated in distal view, with two flat bili. 1901:38.—Colosi, 1920:17.—Coif* papillae on side or in front of spermatic inunn, 1939:107. channel (Fig. II. J). Guinotia {Moritschus) ecuadorensis.— Type species.—Pseudothelphusa ecu­ Prelzmann. 1965:3. adorensis Rathbun, 1897. Potamocarcinus (Hypolobocera) ecuador­ Distribution.—Southern Colombia, Ec­ ensis.—Ron, 1967:370, fig. 5a-c. uador and northern Peru. Hypolobocera (Moritschus) ecuadoren­ Remarks.—Pseudothelphusa ecuadoren­ sis.—Pretzmann, 1971:18; I983b:348, sis Rathbun. 1897. P. henrici Nobili, 1897, 363 and Moritschus nariunesis Campos & Rod­ Hypolobocera i Moritschus) ecuadoriensis riguez, 1988 (from southern Colombia) dis­ (sic).—Prelzmann. 1972:52. tigs. 249, play the same lateral elongation of the gon­ 250. 316-318. opods that results in the distal migration of Moritschus ecuadorensis. — Rodriguez, the lateral lobe and the consequent narrow­ 1982:68, fig, 37a-d. ing and elongation o\ the apex. These char­ acters show that the three species are close­ Material.—Ecuador: Alluriquin, affluent ly related, although their sizes are extreme­ of Rio Touch i* SE of Santo Domingo de los ly different. Colorado*, Piehincha Province, 31 Nov 1980, leg. PL Diaz, 12 males, the largest cl. Key to Species of Moritschus 13.9 mm. cb. 22.0 mm (IVIC 651).—West of Guulea, 880 m alt., leg. O. Thomas. I 1. Lateral margin of tirsi gonopinl's apex male el. 13.2 mm. cb- 25.5 mm (BM without Npinules; caudal end produced in 1918.1.331.11). shorn beak; elongate process over field of Additional illustrations, — Rodnguez spines with 2 rudimentary papillae di­ rected laterally, placed near opening of (1982, rig. 37a-d). spermatic channel or displaced towards Diagnosis.—Carapace with upper frontal lateral expansion, its distal margin entire margin absent. First male gonopods long 2 and slenden with lateral margin widening -. Lateral margin of apex covered with progressively towards apex, this extending 134 PROCHfcDINGS OK THt BIOLOGICAL SOCIETY OF WASHINGTON considerably laterally and produced in short dia, affluent of Rfo Ansu, Rio Napo basin, beak; elongate process over field of spines Santa Clara, , 550 m alt., displaced towards lateral expansion. 4 Nov 1980. leg. H. Diaz, 3 males cl. 18.5, 12.5 and 9.9 mm, cb. 28.7, 19.5 and 13.4 Morilschus henrici (Nobili 1897) mm, 1 immature female cl. 10,2 mm, cb. Fig. II, J 15.3 mm (IV1C 618).—Road Mera-Banos, 4 km from Mera, Rio Pastaza basin, Pastaza Pseudothelphusa henrici Nobili, 1 897:1; Province, 1100 m alt., 6 Nov 1980, leg, H. 1901:40.—Rathbun, 1898:534, 537; Diaz, 1 mature female cl. 48.4 mm, cb. 76.5 1905:302.—Young, 1900:219.—Coiosi, mm (IVIC 617). 1920:40.—Coif mann, 1939:108. Additional illustrations.—Rodri'guez Strengeria (Strengeria) henrici.—Pretz• (1982:66, figs. !9o; 22a, f; 23d; 36a, d). mann, 1965:7. Diagnosis.—Carapace with upper frontal Hypolohocera (Hypolohocera) henrici hen­ margin angled, advanced, with ill-defined rici,— Pretzmann, 1971:17; 1972:39, figs. papillae and deep notch at middle. Third 260, 26 L 294, 296. abdominal tergites with unusually deep cav­ Hypolohocera henrici.—Pretzmann & ities to receive apex of first gonopods, al­ Mayta, 1980:139, figs. 5, 6.—Rodriguez, ready present in juveniles. Propodous of 1982:66, figs. 19o; 22a, f; 23d; 36a, d. fifth pereiopods wide, with row of plumose Hypolohocera {Hypolohocera) [peruviana] setae on infero-posterior margin. First male henrici henrici.—Pretzmann, 1983b;355, gonopods extraordinarily large, lateral mar­ figs. 7, 19, 34, 42, 50, 62. 66, 75. gin widening progressively towards apex Hypolohocera {Hypolohocera) \henrici\ which extends considerably laterally, giving henrici nora Pretzmann. 1977b:436 (no- apex in caudal view triangular-elongated men nudum); 1978:164, fig. 3. appearance; elongate process over field of Hypolohocera {Hypolohocera) [peruviana] spines with 2 rudimentary papillae directed henrici nora.—Pretzmann, 1983b:356, laterally, placed near opening of spermatic figs. 6, 20, 32, 43, 47, 61, 67, 84-88. channel. Material.—Ecuador: Leg. M. Olalla, 38 Remarks.—The material of Hypoloho­ males, the largest cl. 54.8, cb. 91.1 mm, 59 cera (Hypolohocera) [henrici] henrici nora females, the largest cl. 35.4 mm, cb. 56.7 Pretzmann, 1978, consist of two male spec­ mm (TVIC 615).—Monte Sumaco, Lore to, imens, collected by Pretzmann at two lo­ Napo Province, 450 m, Jun 1968, leg. M. calities, Mendez and Rfo Arajuno respec­ Olalla, 1 male, 1 immature female (TU 94 tively, widely separated from each other in 100-7).—Rio Latas, affluent of Rio Napo, the basins of Rio Napo and Rio Santiago, between Tena and Puerto MisuahallL Napo Pretzmann also collected specimens of the Province, 3 Nov 1980, leg, H. Diaz, 3 males typical form of H, henrici from the same cl. 24.5, 16.2 and 8.3 mm. cb. 38.6. 24.8 localities (Pretzmann 1978). The wide dis­ and 12.9 mm {IVIC 616).—Cuevas de Ju- tribution of both subspecies, and their over­ mundf, 5 km N of Arehidona, near Tena, in lap at two localities, suggest that they can­ affluent of Rio Napo, Napo Province, 3 not be .separated as distinct subspecies. On Nov 1980. 1 immature male (1VIC 968).— the other hand it is not possible to give spe Rio Rebadeneira, affluent of Rio Arajuno, cific rank to Hypolohocera {Hypolohocera) Rio Napo basin, Teniente Ortiz, 18 km N henrici [henrici] nora, since the characters of Puyo, Pastaza Province, 980 m alt., 5 are part of the variability of H. henrici. The Nov 1980, leg. H. Diaz, 1 male cl. 21.3 round distal margin of the apex in the first mm, cb. ?>5.5 mm (TVfC 619).—Puyo, Pas­ gonopods, mentioned by Pretzmann (1978) taza Province, 820 m alt., 10 Jan 1986, leg. for H. nora ("Oberrand der Kronchens, in Lilian Real, 1 male (IVIC 939).—Rio Pla- seitlicher Ansicht, starker geknickt"), is ob- VOU'MF I I I. NUMBER 1 135 served in all our immature specimens. The extension and topographic complexity of lateral lobe of rirst gonopods ("Aupenrand these basins, but possibly also to a more der Lateral lobe lateral stark ausgebaucht") intensive collecting effort due to the vicin­ displays a similar variation in our series. ity of large cities. Several species are Another character mentioned by Pretz- known from single localities, underlining mann (1978) is the relative wideness of car­ our imperfect knowledge of the fauna; these apace in both subspecies. The relationship are: Hypolobocera orcesi, H. muisnensis, eb/cl in H. henhci is stated to be more than H. mindimensis, Lindacatalina hauserae, L. 1.64 ("Cpx. breit, Index liber 1.64," Pretz- sumacensis. The most interesting cases are mami 1977b), whereas in the subspecies those of trans basin distribution. Two ex­ nora it is described as wider, with the radius treme examples are Hypolobocera aequa- of curvature of lateral borders larger torialis and Lindacatalina. orientals* ( "VSR-Krummungradius grop . . . Cpx which involve distribution across the main breiter." Pretzmann 1978). The length of Andean water divide. H. esmeraldensis, H. The radius mentioned is of course a function guayaquilensis, H. rat h bun i, H. exited, and of the carapace width. In a series of 26 Moritschus henhci* exhibit similar trans-ba­ specimens from one locality we examined, sin distributions. According to the present the relationship cb/cl varies between 1.51 records and others available in the litera­ and 1.66 according to the size of the spec­ ture, the area of distribution of H. henhci imens. covers the basins of four of the largest ef­ fluents of the Amazon: Rio Napo (Pretz­ mann 1978), Rio Pastaza guavaquilen- ter the estuary of Guayas lies the basin of sis. H, konstanzoe and //. muisnensis^ have the Rio Jubones and a few minor littoral been found along the coastal plain of the streams. The basin of the Daule-Vinces riv­ Esmeraldas and Manabi provinces, between ers, enclosed between the Andes and the 150 and 200 m, but H. guayaquilensis has Manabf mountains, is the most extensive been also recorded inland in the Daule val­ valley on the Pacific drainage. The Ama­ ley (Fig. 13). The other species on the Pa­ zonian drainage comprises to the North, the cific side are inland dwellers: H. caputii and basin of the Napo River, and to the South //. rathbuni, found at 200 m and between the basins of the Pastaza and Santiago riv­ 200 and 450 m, respectively; H. mindonen- ers. These last two drain the internal valleys sis, H. orcesi. and Moritschus ecitadorensis of the Oriental and Central Cordilleras, and between 950 and 1200 m; and H. e.xuca at discharge into the Maranon River. 100 and 2000 m. On the Amazonian side The distribution of species among these the species range from 400 to 1500 m: H. eight basins is unequal. The largest num­ conradi between 900 and 1500 m; Linda- bers occur in the Esmeraldas and Napo ba­ catatina orientalis and L, puyensis between sins, with ten and seven species respective­ 1000 and 1200 m; and L. hauserae, L. la- ly. This abundance is related mainly to the tipenis and L. sumacensis\ between 400 and IM<;u \l. MK'IL ! ^ Of W -\SMINul t >N

I 78° W

L-l SOUTH AMERICA

/. I" ' I-

/ /

/-'-'.< -, - V__ ;• v -"•.5->.. !> '\ »''..w'_--•''EX. /"T V..' ° ^ HE.'

< -.v / • /HE)

• J An •!•' --/ 'fv-- -. -* '1/ ':!: -*t ' I V' />;•*! •2CS-, /f; -?<> «• >>. rnEXi /^ \- • 1/ *

• - * v ;• \

JUBONE&V

•' iw T

\5 • v

v v ---WATER DIVIDES \\ ;' t ;--o TRANS-BASIN * \ •'/ vO \ ,- A DISTRIBUTIONS,*'.„•-' ' \ \ • ' v /. * \

: 1; ••!.; Hinr -n- -1 H. :A:L.\S. -i ) ~-,-.i 1. .1- ,1 /i' ..V-v',: | 1 - // .,": .v. -..•• 1 ^ \ \ = u. .- ^.\.-- ... . M!

H\ = / - ;.::>-. ,.• . V VOLUME 111. NUMBER I 137

Table 2.—Altitudes reported for Ecuadorian Pseu- Fitzpatrick, Jr. for entrusting to the senior dolhelphusids

500 m; Moritschus henrici between 450 and Boa, R. 1967. FluPkrahbcn aus dem wesllichen Sii- 1100 m. damerika (Crust.. Decapod.).—Senckenbergi- ana Biologica 48<5/6>:365-372. Hypolobocera aequatorialis has been . 1970, Belrachtungen Liber die Emwicklungs- collected at 50 m on the Pacific side, and geschichtc und Verbreitung dcr Sujisswasser- from 700 to 1750 m on the Amazonian Krahbcn nach der SanimJung des Naturhisto- side. This vertical distribution, together rischen Museums in Gcnf/Schwci/,—Revue with the trans-basin distribution mentioned Suisse dc Zoologie 77. fascicule 2<24>:327-244. Campos. M.. & G. Rodriguez. 1988. Notes on the above, is rather peculiar for a species of freshwater crabs of the genus Moritschus Pretz- Pseudothelphusidae. However, we were un­ mann. 1965

McGraw-Hill Book Company, Inc. New York, den.—Sit/ungsberichtcn der Osterreiehischcn 328 pp. Akademie der Wissensehaften, Mathematisch Milne Edwards, H. & H Lucas. |842-t844_ Crusta- Naturwisscnschaftlichc Klasse il) 187:163- ces. In: A, d'Orbigny* Voyage dans TAmcnqnc 170. mendionale dans Ic cours des annes 1826— 1833 . 1983a. Ergebnisse cinigcr Sammelrcisen in 6li):]-39. Atlas S);pL 1-17. Siidamerika 1. Tcil: Ncue Pseudothelphusi­ Nobili. G. 1847. Viaggio del Dr. Eurico Festa nctla dae-—Annalen der Naturhistorisches Museum Republica delFEeuador e regioni vieine. 1 De- (Wien) 84/B:301-305. capodi terrestri e d'acqua dolce.—Bollettino dei . 1983b. Die Pseudothelphusidae von Ecua­ Musei di Zoologia ed Anatomia Comparata del- dor.—Annalen der Naturhistorisches Museum la R Univcrsita di Torino 12(275): 1-6. (Wien) 84/B:347^-368. . 1901, Deeapodi raccolti dal Dr Filipo SiL . & R, Mayta. 1980. Uber einige Supwasser- vestri ncll' America meridionals—Bollettino krabben aus Peni.—Anzeiger der Mathematisch dei Musei di Zoologia ed Anatomia Comparata Natui wissenschaftliehe Klasse der Osterrei- del la R. Universita di Torino 16<402 >: 1-16. chischen Akademie der Wissenschaften 9:137- Ortmann, A. I W^3. Die DekapodeivKrebse des Strass- 144. burger Museums, mit besonderer Beriicksichti- . & A. Radda. 1978. Berich iiber zoologisehe gung der von Herrn Dr. Doderlein bei Japan und Studien- und Sammeireisen in Peru and Ecua­ bci den Liu-Kiu-lnseln gesammelten und zur dor 1976/77.—Annalen der Naturhistorisches Zeit in Stiassburger Museum aufbewahrten For- Museum (Wien) 81:589-595. men. VII, Theil. Ahtheilung: Brachyunt Rathbiin. M. J. 1893, Descriptions of new species of (Braehyiira genuina Boas) II Uuterabtbeilung: American freshwater crabs. — Proceedings of Cancroidea. 2 Section: Cancrinea. 1. Gruppe: the United States National Museum lfi(959): Cyclometopa. —Zoologisehe JahrbLieher. Abth- 649-661. eilung tin Systematik. Geographie und Biologic . 1897. Descriptions de nouvelles especes de der Tfuere 7:41 1-49?. Crabes d'eau douce appartenant aux collections . 1847. Careinologisehe Studien.—Zoologis­ du Museum d'Histoire naturelle de Paris.—Bul­ ehe JahrbLicher. Ahtheilung fur Systematik. letin du Museum nationale d'Histoire naturelle Geographic und Biologic der Thiere 10:258- (Paris) 3(2t:58-61. 372. . 1897. Descriptions de nouvelles especes de Pocock. R. 1. 1889. Contributions to our knowledge Crabes d'eau douce appartenant aux collections of the Crustacea ot Dominica.—Annals and du Museum d'Histoire naturelle de Paris.—Bul­ Magazine ot Natural History (6)3:6—22. letin du Museum nationale d'Histoire naturelle Pret/.mami. G. J^ft.V Vorlauhger Bencht tiber die (Paris) 3(2):58-6L Famihe Pseudothelphusidae.— 'Anzeigee der . 1898. A contribution to a knowledge of the Maihetuatiseh Naturwissensehaftliehe Klasse freshwater crabs of America. The Pseudotbel- der OsieiTeichischen Akademie der Wissen- phusinae.—Proceedings of die United States schaltcn (1)1:1-10, National Museum 21(1 l58);507-537. , I9OS, Ncue Siidamerikanische Siifiwasscr- , 1905, Le^ crabes d'eau douce (Potamoni- krabben.—Entomologisehes Nachrichtenblatt dae).— Nouvelles Archives du Museum 77u, Noli/en /ur Biologic der Supwasser- Famille des Pseudothelphusidae,—Faune Trop- krabhen.—Anzeiger der Mathematisch Nalur- icalc 22:1-223. wissen^cliatlhche Klasse der Osterreiehischcn . & H. Diaz. 1981. New Species of Freshwater Akadenne der Wissenschatten 7:87-8^. Crabs from the Andes (Ciustacea. Deeapoda. 1477b, Zur Taxonomie, Chorolo^ie und Sys- Pseudothelphusidae).—Senckenbergiana Biolo- loniiUik der miuelandischen Hypoloboeerini.— gica 61:305-312. Sit/amgsberichtcn tier Oslcrrekhischen Akade- , & M. Campos, 1998. A cladistic revision of mic tier Wisscnschattcn. Mathematisch Natur- the genus h'redius (Crustacea: Deeapoda: Pseu­ wissenschafthche Klasse tl) 186:429-439. dothelphusidae) and its significance to ihe bio- . 1978. Neue Su|iwasserkiaben aus den An- geography of the Guianan lowlands of South VOLUME 111. NUMBF.R 1 139

America. —Journal of Natural History (In of the American river crabs.—Systematic Zo press). ology 13:28-31. , & G. Pereira, 1992. New species, cludistic Smith, S. I. 1870. Notes on American Crustacea. I. relationships and biogeography of the genus Ocypodoidea.—Transactions of the Connecticut Fredius (Crustacea: Decapoda: Pseudothelphu- Academy of Arts and Sciences 2:1 13-176. sidae} from South Amenca.—Journal of Crus­ Young, C. G. 19(X). The stalk-eyed Crustacea of the tacean Biology 12:298-311. British Guiana, West Indies and Bermuda. Lon­ Smalley. A. E. 1964. A terminology for the gonopods don. 514 pp.