PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 119(1):58–66. 2006. Foza raimundi, a new and of potamonautid freshwater (Crustacea: : ) from northern Madagascar

Sadie K. Reed and Neil Cumberlidge* (SKR) Department of Biology, Program in Evolution, Ecology, and Organismal Biology, ASEC 185, University of Akron, Akron, Ohio 44325-3908, U.S.A., e-mail: [email protected]; (NC) Department of Biology, Northern Michigan University, Marquette, Michigan 49855-5301, U.S.A., e-mail: [email protected]

Abstract.—Foza raimundi, a new genus and species of from AntsirananaProvince,northernMadagascar,isdescribedfromahigh-altitude locality in the isolated Marojejy mountain range. The unusual combinationof characters of the new species warrants the establishment of a new genus to accommodate this taxon. The new taxon is compared to the other species of freshwater occurring in Madagascar. The Malagasy freshwater crabs belong to the , a family found exclusively in the Afrotropical zoogeographic region. The new taxon is endemic to Madagascar, as are the other six genera and 12 species of freshwater crabs found there.

The most recent revision of the fresh- Sternberg 2002, 2003; Cumberlidge et al. water crabs of Madagascar (Cumberlidge 2004); there are also a number of reports & Sternberg 2002) recognized 12 species of their presence in phytotelmic habitats in six genera. The distribution of these in rainforest ecosystems (Cumberlidge et endemic was described in al. 2002, Cumberlidge et al. 2005). detail by Cumberlidge & Sternberg We report here on the discovery of (2002) and by Cumberlidge et al. (2004); a new genus and species of freshwater crab the latter authors also assessed the from a high altitude locality in an isolated conservation status of each taxon. Of mountain range in northern Madagascar. concern is the fact that ten Malagasy The specimens described here are distinct- species were judged to be either threat- ly different from all known taxa in ened or extremely rare/endangered, and a number of important morphological only two taxa (Hydrothelphusa agilis A. taxonomic characters including those of Milne-Edwards, 1872 and H. madagasa- the mandible, gonopods, carapace, ster- cariensis (A. Milne-Edwards, 1872)) were num, epistome, chelipeds, walking legs, considered to be secure (Cumberlidge et and third maxilliped. This unusual com- al. 2004). The high degree of endemism bination of characters is sufficient to shown by the Malagasy freshwater crabs warrant the recognition of a new mono- is a characteristic that they share with typic genus to accommodate this species. other terrestrial and freshwater organisms Foza raimundi, new genus, new species, is from this tropical . These crabs are compared to the other genera and species found in the lakes, streams, rivers, and of freshwater crabs from Madagascar. adjacent terrestrial habitats throughout The new taxon is assigned to the Potamo- the island (Bott 1960, 1965: Ng & Takeda nautidae sensu Cumberlidge (1999) and 1994; Cumberlidge 1999; Cumberlidge & Cumberlidge & Sternberg (2002). Abbreviations.—Terminology is adapt- * Corresponding author. ed from Cumberlidge (1999), and the VOLUME 119, NUMBER 1 59 classification used here follows that ter crab. Foza is treated as a feminine of Martin & Davis (2001). Abbrevia- noun. The spelling of the word ‘‘fosa’’ has tions: cw, distance across the carapace been modified because of the potential for at the widest point; cl, carapace length confusion that may arise from the simi- measured along the median line, from larity of the spelling and pronunciation of the frontal margin to the posterior ‘‘fosa’’ to the generic name and common margin; ch, carapace height, the maxi- name of two common Malagasy carni- mum height of the cephalothorax); vores. For example, the scientific name of fw, front width measured along the the Malagasy civet or striped civet is frontal margin; s, thoracic sternite; s4/s5, Fossa fossana (Viverridae), while ‘‘fossa’’, s5/s6, s6/s7, s7/s8, sternal sulci between also spelled ‘‘fosa’’, is the common name adjacent thoracic sternites; e, thoracic for the well-known Malagasy carnivore episternite; s4/e4, s5/e5, s6/e6, s7/e7, Cryptoprocta ferox (Cryptoproctidae) episternal sulci between adjacent thoracic (Dollar 2000a, 2000b). sternites and episternites; a, abdominal Remarks.—Foza, like all Malagasy segment; a6/a5, a7/a6, sutures between freshwater crab taxa, is assigned to the abdominal segments; p1–p5, pereopods Potamonautidae Bott, 1970, because of 1–5; asl, above sea level. All measure- the common possession of a number of ments are given in mm. FMNH 5 Field characters, including a two-segmented Museum of Natural History, Chicago, mandibular palp, a triangular abdomen, Illinois, U.S.A. and a first gonopod with either a short or medium length terminal article (Cumber- Systematics lidge 1999). Foza can be distinguished from the other Malagasy freshwater crab Subphylum Crustacea Bru¨nnich, 1772 genera (Hydrothelphusa A. Milne-Ed- Order Decapoda Latreille, 1802 wards, 1872, Madagapotamon Bott, Infraorder Brachyura Latreille, 1802 1965, Skelosophusa Ng & Takeda, 1994, Superfamily Potamoidea Ortmann, 1896 Marojejy Cumberlidge, Boyko, & Har- Family Potamonautidae Bott, 1970 vey, 2002, Malagasya Cumberlidge & Foza,n.gen. Sternberg, 2002, and Boreas Cumberlidge Type species.—Foza raimundi,n.sp. & Sternberg, 2002) as follows: the first Diagnosis.—Frontal margin of cara- gonopod of Foza has a wide-tipped, pace relatively narrow (fw/cw 0.25), broadly conical terminal article; the sec- sharply deflexed. Pterygostomial region ond gonopod has an extremely elongated of carapace sidewall with broad area of and inwardly curved flagellum-like termi- dense setae. Terminal article of gonopod nal article; the pterygostomial region has 1 short, wide-tipped, broadly conical. a dense field of setae, and the postfrontal Terminal article of gonopod 2 extremely crest is faint and incomplete with the long, flagellum-like, reaching anterior weak epigastric crests positioned so far margin of sterno-abdominal cavity; fla- forward on the front that they almost gellum of gonopod 2 curving inward touch the frontal margin. Interestingly, distally, tip sharply curved inward, over- this discovery means that the Marojejy all outline of flagellum forming ‘‘question mountains are home to four species of mark’’ shape; distal parts of flagellae of freshwater crabs belonging to four gen- both second gonopods overlapping one era, the other three taxa being Marojejy another, sometimes protruding from un- longimerus Cumberlidge, Boyko & Har- der closed abdominal telson. vey, 2002, Skelosophusa eumeces Ng & Etymology.—Foza is derived from the Takeda, 1994, and Hydrothelphsua Malagasy name ‘‘fosa’’ meaning freshwa- madagascariensis. 60 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Foza is morphologically close to Mala- matically reduced short stub in Madaga- gasya because both taxa share a bilobed potamon). mandibular palp, elongated walking legs, a narrow deflexed front, and two pointed Foza raimundi, new species carpal teeth on the carpus of the cheliped. Figs. 1 and 2 However, Foza can be distinguished from Malagasya by the anterolateral margin of Holotype.—FMNH 7435: adult male the carapace, which is granular in Foza, (cw 22.9, cl 17, ch 10.9, fw 5.7 mm), but conspicuously toothed in Malagasya. Madagasacar, Antsiranana Province, Foza can be distinguished from Hydro- Parc National de Marojejy (5 Marojezy), thelphusa and Marojejy, the other Mala- 13 km SE Doany, on ridge between the gasy genera with a bilobed mandibular Andranomazava and Antsahaberaoka palp, as follows: the space between the Rivers (14u26.29S, 49u37.29E), caught by exorbital and epibranchial teeth in Foza is hand in undisturbed montane forest on very narrow as the result of the advanced slope with slightly wet ground, collected position of the epibranchial tooth (which by Achille Ruselimanana, 29 Oct 2001. is so far forward that it almost touches Paratypes.—FMNH 7436: adult ovi- the exorbital tooth), whereas in both gerous female (cw 25.2, cl 18.6, ch Hydrothelphusa and Marojejy there is 12.4, fw 6.1 mm), Antsiranana Province, a wide space between these teeth. The Parc National de Marojejy (5 Marojezy), long slender walking legs of Foza further 13 km SE Doany, on ridge between the distinguish this genus from Hydrothel- Andranomazava and Antsahaberaoka phusa whose walking legs are shorter, and Rivers (14u26.29S, 49u37.29E), 1070 m not unusually elongated. The eyestalks asl, found 20 cm below surface of soil, and corneas of Foza areofnormallength collected by S. M. Goodman, 29 Oct and size, which distinguishes this genus 2001; FMNH 7437: adult female oviger- from Marojejy whose eyestalks taper ous (cw 22.2, cl 17.4, ch 11.5, fw 5.7 mm), distally and whose corneas are very Antsiranana Province, Parc National reduced. de Marojejy (5 Marojezy) (14u26.29S, Foza is distinguished from Skeloso- 49u37.29E), 875 m asl, pitfall trap, col- phusa and Boreas by the form of the lected by Voahangy & Bomoina, 11 Feb terminal segment of the mandibular palp 2002; FMNH 7438: adult ovigerous (which has an enlarged anterior process female (cw 22.8, cl 15.7, ch 10.2, fw about 0.5 times as large as the posterior 6.6 mm), Antsiranana Province, Parc lobe in Foza, but only a small ledge-like National de Marojejy (5 Marojezy) (14u anterior process in Skelosophusa and 26.29S, 49u37.29E), 850 m asl, pitfall trap, Boreas). The long slender walking legs collected by M. Raheriasena & S. M. of Foza distinguish this genus externally Goodman, 6 Feb 2002; FMNH 7439: from Boreas whose walking legs are juvenile (cw 10.9, cl 8.9, ch 5.6, fw shorter, and not unusually elongated. 2.6 mm), Antsiranana Province, Parc Foza is distinguished from Madagapota- National de Marojejy (5 Marojezy), NW mon by the terminal segment of the (14u26.29S, 49u37.29E), 810 m asl, pitfall mandibular palp (which has a distinct trap, collected by V. Soarimalala & D. lobe-shaped anterior process in Foza,but Raheriasena, 10 Feb 2002; FMNH 7440: which is simple and lacking all evidence of juvenile (cw 11.3, cl 9.3, ch 5.8, fw an anterior process in Madagapotamon) 2.9 mm), Antsiranana Province, Parc and by the length of the flagellum of the National de Marojejy (5 Marojezy) NW exopod of the third maxilliped (which is (14u26.29S, 49u37.29E), 810 m asl, pitfall a shortened flagellum in Foza,butadra- trap, collected by V. Soarimalala & D. VOLUME 119, NUMBER 1 61

Fig. 1. Foza raimundi, from Madagascar. Male holotype (FMNH 7435), cw 22.9 mm. A, cephalothorax, frontal aspect; B, cephalothorax, dorsal aspect; C, carpus and merus of right cheliped, inferior view; D, carpus and merus of right cheliped, superior view; E, right cheliped, frontal aspect; F, left cheliped, frontal aspect; G, right pereopod 4; H, right pereopod 5; I, sternum. Scale bar 5 8 mm. 62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Foza raimundi, from Madagascar. Male holotype (FMNH 7435), cw 22.9 mm. A, left third maxilliped; B, right gonopod 1, dorsal aspect; C, right gonopod 1, medial aspect, turned to show longitudinal groove; D, right gonopod 1, ventral aspect; E, right gonopod 2, ventral aspect. Female adult, paratype (FMNH 7437), cw 22.2 mm. F, sternum. Scale bar 5 4 mm (B–E), 8 mm (A, F).

Rahotomalala, 11 Feb 2002; FMNH trap, collected by V. Soarimalala & D. 7441: juvenile (cw 13.8, cl 10.5, ch 6.6, Rahotomalala, 11 Feb 2002. fw 3.6 mm) Antsiranana Province, Parc Diagnosis.—As for the genus. National de Marojejy (5 Marojezy), NW Description.—Based on holotype, an (14u26.29S, 49u37.29E), 810 m asl, pitfall adult male. Carapace outline transversely VOLUME 119, NUMBER 1 63 oval, very high (ch/fw 2.0); front narrow separated forming wide space, folds (fw/cw 0.25), deflexed; epibranchial tooth meeting midway along ventral face of small, pointed, extremely advanced in article to form longitudinal groove that position, almost touching exorbital tooth; continues to tip of article; groove not anterolateral margin evenly curved out- visible on dorsal face; subterminal seg- ward, lined by small granules, continuous ment of gonopod 1 with distinct raised with posterolateral margin, latter margin rounded shoulder on external margin crossed by carinae; postfrontal crest faint near junction with terminal article. Junc- to absent, postorbital crests lacking, tion between terminal article and sub- epigastric crests faint, positioned forward terminal segment of gonopod 1 not clear on front almost touching frontal margin; on ventral side. Terminal article, sub- deep mid-groove between epigastric crests terminal segment separated on dorsal side forked posteriorly; cardiac, urogastric by broad, subtriangular dorsal mem- grooves deep, cervical grooves deep, brane; superior margin of dorsal mem- short. Suborbital region of carapace side- brane formed by diagonal basal margin of wall smooth, subhepatic region smooth terminal article, inferior margin of mem- except for few carinae near posterolateral brane formed by diagonal slightly s- margin, pterygostomial region with broad shaped distal edge of subterminal seg- area of dense setae, vertical sulcus on ment; lateral margin of dorsal membrane carapace sidewall curved, granular, run- broad, medial margin of membrane nar- ning from base of epibranchial tooth to row, forming medial junction between epimeral sulcus. subterminal segment and terminal article. Epistomial tooth triangular, deflexed, Subterminal segments of gonopods 1 & edges smooth. Exopod of third maxilliped 2 of equal length, but terminal article of reaching to lower half of merus, flagellum gonopod 2 much longer than terminal of exopod with short flagellum, ischium article of gonopod 1. Terminal article of with faint vertical groove, curving distally gonopod 2 flagellum-like, same length as toward medial margin. Sternal sulcus s1/ subterminal segment of gonopod 2, reach- s2 short, very faint; sternal sulcus s2/s3 ing anterior margin of sterno-abdominal completely crossing sternum; sternal sul- cavity; flagellum curving inward distally, cus s3/s4 a v-shaped groove, deep at edges, tip sharply curved inward, overall outline faint in middle; anterior sterno-abdominal of flagellum forming ‘‘question mark’’ cavity lined with short setae. Episternal shape; distal parts of flagellae of both sulci s4/e4, s5/e5, s6/e6, s7/e7 absent, second gonopods overlapping, sometimes smooth. Male abdomen slim, triangular, protruding from under closed abdominal tapered, widest at a3 narrowest at a7 telson. (telson); telson outline forming straight- Dactylus of both chelipeds relatively sided triangle with broad base, rounded slender, approximately one-third height apex. Sternal groove s4/s5 meeting abdo- of palm, cutting edge lined by small even men at abdominal groove between a7/a6; teeth; upper margin of dactylus smooth; sternal groove s6/s7 meeting a6 one lower margin of propodus slightly in- quarter of segment length from a6/a5. dented. Fixed finger of propodus of Terminal article of gonopod 1 very major (right) cheliped slender with large short (ratio of length of terminal article to fused molar in proximal region followed subterminal segment 0.2), cone-shaped, by series of small pointed teeth. First directed slightly outward, straight, bris- carpal tooth on inner margin of carpus of tled, apical opening wide; lateral, medial cheliped large, pointed; second carpal folds on ventral terminal article of gono- tooth smaller, pointed, followed by series pod 1 equal in height, width, folds basally of very small teeth. Medial, lateral 64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON margins of inferior face of merus of Marojejy massif is a forested mountain cheliped distinctly toothed, inferior face range with a series of peaks some of with pointed, granulated distal meral which are more than 2000 m asl, situated tooth; superior margin and superior face between the Tsaratanana mountains to of merus of cheliped roughened by the northwest and the Masoala peninsula granules and short carinae; granules on to the southeast. The climate of Marojejy medial margin of merus continuous with is sub-humid and warm and receives granules on medial margin of ischium of heavy rain from mid-October to April cheliped, inferior margin of ischium (25uC average) and lighter rain from May rounded, smooth. Walking legs (p2–p5) to October (19uC average). The river elongated (ratio of merus length of p5 to Manantenina drains the Lokoho valley cw 0.4), slender, inner margins of propodi from west to east and empties into the of p2 to p5 smooth. , whereas the river Andra- Variation.—Anterior sternum of female nomadiobe flows south from these moun- with heavy field of setae, sterno-abdom- tains and the river Antongodriha flows inal cavity broad, shallow. Adult female north. abdomen broad, shield-shaped, lateral Ecology.—Crabs were caught in pitfall margins covering coxa of walking legs, traps set in a partially disturbed mixed telson (a7) outline broad-based triangle dry deciduous and humid forest at with rounded apex. Chelipeds subequal, altitudes between 810 and 1070 m asl. lacking enlargement of propodus palm, Interestingly, in one case a crab was lacking large molar teeth on edges of found burrowed 20 cm below the surface fingers seen in adult male. of the soil. These observations imply that Size.—The largest known specimen is this species spends at least some of its the male holotype, cw 22.9 mm. Adults as time on land, and indicates a degree of judged by size at pubertal molt beginning independence from permanent water around cw 22 mm. sources. Furthermore, the branchial Live coloration.—Carapace uniformly chambers of F. raimundi each house two deep purple, chelipeds and walking legs different respiratory organs (both a dorsal light brown, sternum and undersides of pseudolung and ventral gills) that is pereopods p1–p5 pale purple-yellow. noteworthy because pseudolungs are not Distribution.—Foza raimundi is known widespread in potamonautids (Sternberg only from a single locality in the Parc & Cumberlidge 2001). Foza shares this National de Marojejy (formerly the Re´- adaptation of its respiratory system with serve Naturelle Inte´grale de Marojejy), in a select number of terrestrial Afrotropical Antsiranana Province in northern Mada- freshwater land crabs such as gascar. macropus Rathbun, 1898 from West Remarks.—This species is distinguished Africa (Cumberlidge 1991, 1999); Deck- from other Malagasy species by a unique enia imitatrix Hilgendorf, 1868 from East combination of gonopod and carapace Africa; Madagapotamon humberti Bott, features (see Remarks under genus 1965 from Madagascar; and above). The discovery that the Marojejy alluaudi (A. Milne-Edwards & Bouvier, mountain range supports three mono- 1893) from the (Sternberg & typic genera of freshwater crabs (Foza, Cumberlidge 2001). Pseudolungs are Marojejy,andSkelosophusa) plus the widespread in neotropical pseudothelphu- widespread species H. madagascariensis sids (Rodriguez 1986), and were described (Cumberlidge et al. 2004) makes this in detail for several semi-terrestrial species isolated mountain range an area of great by Diaz & Rodriguez (1977). Foza shares interest in terms of its biodiversity. The a superficially similar morphology with VOLUME 119, NUMBER 1 65 other highly terrestrial true freshwater [5Copenhagen and Leipzig]: Apud Frider. crabs (Potamidae, Potamiscinae) from Christ. Pelt., 254 pp. [not seen; as cited in Holthuis, 1991]. Southeast Asia such as Thaipotamon Ng Dollar, L. 2000a. Cryptoprocta ferox. In IUCN & Naiyanetr, 1993, Pudaengon Ng & 2004. 2004 IUCN Red List of Threatened Naiyanetr, 1995, and Terrapotamon Ng Species. 5www.redlist.org4. & Naiyanetr, 1998, no doubt through Dollar, L. 2000b. Fossa fossana. In IUCN 2004. convergence (Ng & Naiyanetr 1993, 1995, 2004 IUCN Red List of Threatened Species. 5www.redlist.org4. 1998). Clearly, F. raimundi,asanair- Cumberlidge, N. 1991. The respiratory system of breathing highly terrestrial species, is Globonautes macropus (Rathbun 1898), a ter- worthy of comparison with land crabs restrial fresh-water crab from Liberia (Para- (including members of the predominantly thelphusoidea, ).—Crusta- marine families Ocypodidae and Grapsi- ceana 61(1):69–80. ———. 1999. The freshwater crabs of West Africa, dae) that are highly adapted to life out of family Potamonautidae.—Faune et Flore water. Tropicales, No. 35, IRD, Paris, 382 pp. Etymology.—The new species is named ———, & R. v. Sternberg. 2002. A taxonomic for the first author’s father, Raymond revision of the freshwater crabs of Madagas- Paul Reed II, nicknamed ‘‘Raimundo,’’ to car (Decapoda: Potamoidea: Potamonauti- dae).—Zoosystema 24(1):41–79. thank him for his extensive love and ———, & ———. 2003. The freshwater crabs of support throughout her graduate career Madagascar. Pp. 612–617 in S. Goodman and during which time this species was dis- J. Benstead, eds., Madagascan Natural His- covered. tory. University of Chicago Press. ———, C. B. Boyko, & A. W. Harvey. 2002. A new genus and species of freshwater crab (Dec- Acknowledgments apoda, Crustacea, Potamoidea) from north- ern Madagascar, and a second new species We thank Steven M. Goodman (De- associated with Pandanus leaf axils.—Journal of Natural History 36:65–77. partment of Zoology, FMNH) for his ———, S. K. Reed, & C. B. Boyko. 2004. unparalleled field collecting skills and for Distribution patterns of the Malagasy fresh- making the specimens available to the water crabs (Crustacea: Decapoda: Bra- authors, and we thank Janet Voight and chyura).—Journal of Natural History 38: Martin Pryzdia of the FMNH for hosting 1133–1157. visits to the museum by both authors. The ———, D. B. Fenolio, M. E. Walvoord, & J. Stout. 2005. Tree-climbing crabs (Potamonautidae National Science Foundation (Grant and Sesarmidae) from phytotelmic microhab- no. 0075614) is thanked for its support. itats in rainforest canopy in Madagascar.— Journal of Biology 25(2):302–308. Diaz, H., & G. Rodriguez. 1977. The branchial Literature Cited chamber of some terrestrial and semiterres- trial crabs.—Biological Bulletin 153:485– Bott, R. 1960. Crustacea (Decapoda): Potamonidae. 504. Pp. 13–18 in B. Hansstro¨m and others, eds., Hilgendorf, F. 1868. Ueber eine neue Gattung der South African life. Results of the kurzschwanzigen Krebse aus den Sammlun- Lund University Expedition in 1950–1952. gen des Baron von der Decken, ———. 1965. Die Su¨ßwasserkrabben von Mada- imitatrix.—Sitzungsberichte der Gesellschaft gaskar.—Bulletin du Muse´um national d’His- naturforschender Freunde zu Berlin 1869:2. toire naturelle Paris 37(2):335–350. Holthuis, L. B. 1991. Marine lobsters of the world. ———. 1970. Die Su¨ßwasserkrabben von Europa, An annotated and illustrated catalogue of Asien, Australien und ihre Stammes- species of interest to fisheries known to geschichte.—Abhandlungen der Senckenber- date.—FAO species catalogue, vol. 13, and gischen Naturforschenden Gesellschaft FAO Fisheries Synopsis no. 125. Rome: FAO Deutsch 526:1–338. i–vii, Pp 292. Bru¨nnich, M. Th. 1772. Zoologiae fundamenta Latreille, P. A. 1802. Histoire Naturelle ge´ne´rale et praelectionibus academicis accomodata. particulie`re, des Crustace´s et des Insectes, Grunde I Dyrelaeren. Hafniae et Lipsiae III:1–467. 66 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Martin, J. W., & G. E. Davis. 2001. An updated ———, & ———. 1998. Terrapotamon palian sp. classification of the recent Crustacea.—Nat- nov., a second member of the terrestrial crab ural History Museum of Los Angeles County genus Terrapotamon (Decapoda, Brachyura, Contributions in Science 39:1–124. Potamidae) from southern Thailand.—Crus- Milne-Edwards, A. 1872. Note sur les crabes d’eau taceana 71(5):487–492. douce de Madagascar.—Bibliographie e´cole ———, & M. Takeda. 1994. Skelosophusa (Crusta- Hautes e´tudes (Se´ction Sciences naturelles) cea, Decapoda, Brachyura), a new genus of Paris 5(8):1–3. potamonautid freshwater crab from Mada- ———, & E. Bouvier. 1893. Sur une espe`ce nouvelle gascar, with descriptions of two new spe- du genre Deckenia (Hilgendorf) recueilli par cies.—Bulletin of the National Science Muse- M. Alluaud aux Seychelles, Indian Ocean.— um, Tokyo, series A (Zoology) 20(4):161–172. Annales des Sciences Naturelles (Zoologie), Ortmann, A. 1896. Das system der Decapoden- Paris 15:325–336. Krebse.—Zoologische Jahrbuchner (Systema- Ng, P. K. L., & P. Naiyanetr. 1993. New and tics) 9:409–453. recently described freshwater crabs (Crusta- Rathbun, M. J. 1898. Descriptions of three new cea: Decapoda: Brachyura: Potamidae, Ge- species of fresh-water crabs of the genus carcinucidae and Parathelphusidae) from Potamon.—Proceedings of the Biological So- Thailand.—Zoologische Verhandeingen 284: ciety of Washington 12:27–30. 1–117, figs. 1–68. Rodriguez, G. 1986. Centers of Distribution of ———, & ———. 1995. Pudaengon, a new genus of Neotropical Freshwater Crabs. Pp. 51–67 in terrestrial crabs (Crustacea: Decapoda: Bra- R. H. Gore, & K. L. Heck, eds., Bio- chyura: Potamidae) from Thailand and Laos, geography of the Crustacea.—Crustacean with descriptions of seven new species.— Issues 3, figs. 1–13. Raffles Bulletin of Zoology, Singapore Sternberg, R. v., & N. Cumberlidge. 2001. Notes on 43(2):355–376. the position of the true freshwater crabs ———, & ———. 1997. A revision of the genus within the brachyrhynchan Eubrachyura Siamthelphusa Bott, 1968 (Crustacea: Deca- (Crustacea: Decapoda: Brachyura).—Hydro- poda: Brachyura: Parathelphusidae), with biologia 449(1/3):21–39. descriptions of five new species from Thai- land.—Journal of Natural History 31(12): 1751–1784. Associate Editor: Christopher B. Boyko