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Ovitamon lubang sp. nov. (Figs. 2, 3 C, D) Material examined. NMCR 25009, holotype male (29.7 × 22.7 mm), Philippines, Mindoro Occidental Province, Lubang Island, Barangay Sorville, Mount Ambolong, mid-mountain creek near Philippine Air Force Radar Station, coll R. V. Sison; 27 April 2001. Paratype: ZRC (ex NMCR 25033), 1 female (34.4 × 26.4 mm), same data as the holotype. Diagnosis. Carapace with dorsal surface finely pitted or with rugosities; branchial regions gently convex; anterolateral margin strongly convex, with distinct cristae, adjacent areas with granules; epibranchial tooth small, blunt, distinctly separated from external orbital angle by small cleft; external orbital angle triangular, outer margin gently serrrated to granulated, cristate (Fig. 2 A, B); suborbital, pterygostomial regions gently rugose; posterior margin of epistome with prominent triangular median lobe with gently concave margins, about half width of margin (Fig. 2 B); chelipeds with outer surfaces distinctly rugose; fingers gaping slightly (Fig. 2 A, D); ambulatory legs rugose, dorsal margins of merus uneven, with scattered granules (Fig. 2 A, C); male sterno-abdomimal cavity reaching to imaginary line joining median part of bases of coxae of chelipeds (Fig. 2 C); male abdomen narrowly triangular (Fig. 2 C); G 1 terminal segment subcylindrical, slightly longitudinally twisted, ca. 0.5 times length of subterminal segment; almost straight, tip relatively sharp; subterminal segment subquadrate, stout (Fig. 3 C, D). Description of holotype male. Carapace transversely ovoid, dorsal surface finely pitted or with short rugosities, evenly convex; branchial regions gently convex; cervical groove shallow, broad; H-shaped median grooves distinct (Fig. 2 A). Frontal margin almost straight or slightly sinuous, cristate, bent downwards, forming low tranverse ridge (Fig. 2 B). Anterolateral margin strongly convex, cristate, adjacent areas with granules; epibranchial tooth small, blunt, distinctly separated from external orbital angle by small cleft (Fig. 2 A). Orbital margins gently sinuous, cristate, adjacent surface rugose (Fig. 2 B). Epigastric crista well defined, blunt, medially separated by short groove (Fig. 2 A). Postorbital ridge uneven, rounded, confluent with epibranchial tooth (Fig. 2 A). External orbital angle triangular, outer margin gently serrrated to granulated, gently concave, cristate (Fig. 2 A). Posterolateral margins sinuous, converging to gently convex posterior carapace margin, with some low oblique lateral ridges (Fig. 2 A). Suborbital, pterygostomial regions gently rugose (Fig. 2 B). Buccal cavern squarish; third maxilliped with distinct flagellum, shorter than width of merus; sulcus on ischium shallow (Fig. 2 B). Epistome relatively wide, surface rugose, margins cristate; posterior margin of epistome with prominent triangular median lobe with gently concave margins, about half width of margin; lateral lobes low (Fig. 2 B). Chelipeds unequal; outer surfaces rugose; merus relatively short, dorsal margin crenulated, rugose, but otherwise unarmed; carpus rounded with strong inner distal tooth, surface strongly rugose; major palm swollen, fingers gaping slightly; pollex with 2 subventral longitudinal grooves; dactylus with 3 shallow longitudinal grooves lined with pits; cutting margin with numerous denticles, low teeth (Fig. 2 A, D). Ambulatory legs slender, long, surface rugose; with scattered stiff setae on margins; dorsal margins of merus uneven, with scattered granules; dactylus relatively long (Fig. 2 A). Thoracic sternites smooth, with short setae only on margins; suture present between sternites 2, 3 gently convex towards buccal cavity; sternites 1, 2 completely fused, broadly triangular with sharp apex (Fig. 2 C). Sternoabdomimal cavity reaching to imaginary line joining median part of bases of coxae of chelipeds. Abdomen narrowly triangular; all somites free; telson longer than other somites, lateral margins almost straight, tip rounded; lateral margins of somite 3-6 gently convex to slightly concave (Fig. 2 C). G 1 subcylindrical, twisted; terminal segment ca. 0.5 times length of subterminal segment; terminal segment covered with spinules, stiff setae, almost straight, tip relatively sharp; distinctly demarcated from subterminal segment by wide cleft resembling collar, covered with very short stiff setae; subterminal segment subquadrate, stout, lateral margins lined with setae (Fig. 3 C, D). G 2 long, sinuous; flagellum about half length of basal segment, proximal part broadly dilated. Female. The female abdomen completely covers the thoracic sternum. The vulvae are relatively large, without a visible operculum. Etymology. Named after Lubang Island, the type locality. The name is used as a noun in apposition. Remarks. The G 1 structure of O. lubang sp. nov. is characteristic, with the G 1 terminal segment long and straight, with the tip sharp and bent backwards (Fig. 3 C, D). In this regard, it is perhaps most similar to the condition seen in O. arcanum from Marinduque Island much farther east. The type locality of O. lubang sp. nov. is on Lubang Island (13 ° 48 ’N 120 ° 8 'E) to the north of the island of Mindoro. The subterminal segment of O. lubang sp. nov. is nevertheless relatively straighter and the terminal segment is proportionately longer (Fig. 3 C, D) (versus gently subterminal segment sinuous in O. arcanum, Ng & Takeda 1992: fig. 1 D-G). In addition, the anterolateral margin of O. lubang sp. nov. is distinctly cristate (Fig. 2 A, B), whereas it is almost smooth and rounded in O. arcanum (Ng & Takeda 1992: fig. 1 A).

Family Potamidae, Published as part of Manuel-Santos, Marivene R. & Ng, Peter K. L., 2013, Two new species of Ovitamon Ng & Takeda, 1992 (Crustacea: Brachyura: Potamidae) from the Philippines, pp. 394-400 in Zootaxa 3619 (3) on page 394, DOI: 10.11646/zootaxa.3619.3.8, http://zenodo.org/record/222698

Ovitamon baloy sp. nov. (Figs. 1, 3 A, B) Material examined. NMCR 15007, holotype male (19.5 × 14.7 mm), Philippines, Panay island, Antique, Valderrama, Mount Baloy, 1340 m asl, coll. M. R. Manuel, October 1989. Paratypes: ZRC (ex NMCR 25034), 1 male (18.0 × 13.6 mm), NMCR 25034, 1 female (18.1 × 14.3 mm), same data as holotype. Diagnosis. Carapace with dorsal surface smooth or finely pitted; branchial regions prominently swollen; anterolateral margin strongly convex, with very low, weak cristae; epibranchial tooth low, blunt, not distinctly separated from external orbital angle by notch or fissure; external orbital angle broadly triangular, blunt, outer margin smooth, gently concave, cristate (Fig. 1 A, B); suborbital, pterygostomial regions smooth; posterior margin of epistome with triangular median lobe about one-third width of margin (Fig. 1 B); chelipeds with outer surfaces almost smooth, fingers prominently gaping in adult male (Fig. 1 A, D); ambulatory legs smooth, dorsal margin of merus almost smooth (Fig. 1 A, C); male sterno-abdomimal cavity reaching to imaginary line joining anterior edges of coxae of chelipeds; male abdomen broadly triangular (Fig. 1 C); G 1 relatively stout; terminal segment ca. 0.3 times length of subterminal segment, cylindrical, gently twisted longitudinally, gently curved, tip rounded (Fig. 3 A, B). Description of holotype male. Carapace transversely ovoid, dorsal surface smooth or finely pitted, evenly convex; branchial regions prominently swollen; cervical groove distinct, shallow, broad; median grooves Hshaped, distinct (Fig. 1 A). Frontal margin gently sinuous, cristate, bent downwards, forming low transverse ridge (Fig. 1 B). Anterolateral margin strongly convex, with very low, weak cristae; epibranchial tooth very low, blunt, almost confluent with epigastric margin, not distinctly separated from external orbital angle (Fig. 1 A, B). Orbital margins gently sinuous, cristate. Epigastric cristae not well defined, low, separated by shallow Y-shaped groove (Fig. 1 A). Postorbital cristae uneven, low, confluent with epibranchial tooth (Fig. 1 A). External orbital angle broadly triangular, blunt, outer margin smooth, gently concave, cristate (Fig. 1 A). Posterolateral margins sinuous, converging to convex posterior carapace margin, no oblique lateral ridges discernible (Fig. 1 A). Suborbital, pterygostomial regions smooth (Fig. 1 B). Buccal cavern quadrate; third maxilliped with distinct flagellum, slightly shorter than width of merus; sulcus on ischium very shallow (Fig. 1 B). Epistome relatively wide, covered with short thick setae, margins cristate; posterior margin of epistome with triangular median lobe with concave margins, about one-third width of margin; lateral lobes low (Fig. 1 B). Chelipeds unequal; outer surfaces mostly smooth; merus relatively short, dorsal margin crenulated but otherwise unarmed; carpus rounded with strong inner distal tooth; major palm swollen, fingers prominently gaping; pollex with subventral longitudinal groove; dactylus with 3 shallow longitudinal grooves lined with pits; cutting margin with small denticles to almost smooth (Fig. 1 A, D). Ambulatory legs slender, long, smooth; with some stiff setae on margins; dorsal margin of merus almost smooth; dactylus relatively long (Fig. 1 A). Thoracic sternites smooth, with setae only on margins; suture present between sternites 2, 3 gently convex towards buccal cavity; sternites 1, 2 completely fused, broadly triangular with apex long, sharp (Fig. 1 C). Sternoabdomimal cavity reaching to imaginary line joining anterior edges of coxae of chelipeds (Fig. 1 C). Abdomen broadly triangular; all somites free; telson longer than other somites, lateral margins vaguely sinuous, tip rounded; lateral margins of somite 3–6 straight to gently convex (Fig. 1 C). G 1 relatively stout; terminal segment ca. 0.3 times length of subterminal segment, cylindrical, gently twisted longitudinally, gently curved towards median part of thoracic sternum, densely covered with spinules, hooked spines, hooked stiff setae, tip rounded; collar separating terminal, subterminal segments narrow, not swollen; subterminal segment with margins setose (Fig. 3 A, B). G 2 long, sinuous; flagellum about half length of basal segment, proximal part broadly dilated. Female. The female abdomen completely covers the thoracic sternum. The vulvae are relatively large, without a visible operculum. Etymology. The name comes from Baloy, the mountain where this species was collected. The name is used as a noun in apposition. Colour in life. Branchial regions reddish-violet, margins orange; gastric region brownish purple; chelipeds purple; spines on legs orange. Remarks. Two other species of Ovitamon, O. tomaculum and O. cumingii, are known from the island of Panay in central Philippines. The most significant difference between O. tomaculum and O. baloy sp. nov. is the structure of the G 1. In O. baloy sp. nov., the G 1 is proportionately much stouter (especially the subterminal segment), with the terminal segment relatively shorter, and the tip is broadly rounded (Fig. 3 A, B) (G 1 relatively more slender overall with the terminal segment distinctly tapering in O. tomaculum, cf. Ng & Takeda 1992: fig. 3 E-H). These differences are substantial and cannot be accounted for by size-associated changes or individual variation. The holotype male of O. cumingii is dried and the important G 1 structure is not available for examination (Ng & Takeda 1993). The carapaces of the two species, however, are very different, with that of O. baloy sp. nov., relatively rounder and smoother, with the anterolateral margin rounded and almost smooth with the cristae very low (Fig. 1 A, B) (versus carapace proportionately broader, with lateral regions rugose and prominent anterolateral cristae, cf. Ng & Takeda 1993: fig. 1 A, B). In any case, the type locality of O. baloy sp. nov. (Mount Baloy, 11 ° 8 ' 56 "N 122 ° 15 ' 9 "E) is 100 km northwest of Guimaras Island (ca. 10 ° 35 ʹN 12 ° 236 ʹE), the type locality of O. cumingii.

Ovitamon Ng & Takeda, 1992 Type species. Ovitamon arcanum Ng & Takeda, 1992, by original designation; gender of genus neuter., Published as part of Manuel-Santos, Marivene R. & Ng, Peter K. L., 2013, Two new species of Ovitamon Ng & Takeda, 1992 (Crustacea: Brachyura: Potamidae) from the Philippines, pp. 394-400 in Zootaxa 3619 (3) on page 394, DOI: 10.11646/zootaxa.3619.3.8, http://zenodo.org/record/222698

Manuel-Santos, Marivene R., Ng, Peter K. L. (2013): Two new species of Ovitamon Ng & Takeda, 1992 (Crustacea: Brachyura: Potamidae) from the Philippines. Zootaxa 3619 (3): 394-400, DOI: http://dx.doi.org/10.11646/zootaxa.3619.3.8

Vultocinus anfractus, new species Figs. 1 –5, 8 A, 9 A, 10 A, 11 A, 12 A, 13 A Material examined. Holotype: male (cl 19.5 mm, cw 20.3 mm) (NMCR 19114); station M 27, Santiago, Maribojoc Bay, Panglao, Bohol, Philippines, 9 ° 0.043 'N 123 ° 0.051 'E, coll. 23 Jun 2004. Paratypes: 1 male (cl 19.9 mm, cw 22.0 mm) (ZRC), station P1, 90– 200m, Maribojoc Bay, Panglao, Bohol, Philippines, 9 ° 36.1 'N 123 °45.0'E, coll. fishermen with tangle nets, 27 Jun 2004; 1 male (cl 27.3 mm, cw 25.3 mm) (ZRC, ex NMCR 19115), Maribojoc Bay, Panglao, Bohol, Philippines, in tangle nets, ca. 200m, coll. J. Arbasto, Oct 2004 – Feb 2005; 1 male (cl 27.4 mm, cw 29.0 mm) (MNHN), Maribojoc Bay, Panglao, Bohol, Visayas, Philippines, in tangle nets, 80–140m, coll. J. Arbasto, between 2004 and 2005. Non-type: male (cl 15.6 mm, cw 16.8 mm) (ZRC), station DB 16, sandy slope with coral patches, Tutuba off Santo, Vanuatu, in tangle nets, ₁ ₅ ․ ₃ ₆ ° S 167.16 °E, 32 ̄ 40 mm depth, coll. J. Arbasto, 14 Sep 2006. Description of male holotype. Carapace subquadrate, squarish to slightly broader than long; dorsal surface relatively flattened with low convex surfaces; regions well defined, separated by deep, prominent grooves, anterior part more convex than posterior part, marked by prominent areolae, covered with numerous, densely packed very short, stiff, black setae, velvet-like, partially obscuring surface but not completely obscuring margins; 1 F divided into 2 cone-shaped lobes; 2 F complete, surface uneven; 1 M complete; 2 M with median depression; 3 M not continuous; 4 M laterally long, distinct; 1 P with small areolae; 2 P long, narrow; 2 L triangular in shape; 3 L elongated, triangular in shape; 6 L clearly demarcated, with tubercles of varying sizes; 3 R with small areolae (Figs. 1, 2, 3 A, D). Frontal region almost one-third width of carapace, with a distinct notch formed by 2 conical extensions (Figs. 2, 3 A, D). Orbits rounded, eyes fitting tightly into cavity; margins areolate with small flat granules on rim, separated into 3 parts by deep sulci (Fig. 3 A, D); supra-orbital border tuberculated, each tubercle with smaller granules on surface, with prominent inner orbital tooth and lower external orbital tooth, with 2 smaller teeth between which are separated by a deep narrow fissure, one tooth separated from inner orbital tooth by a cleft, the other separated from outer orbital tooth by a deep narrow fissure (Fig. 3 A, D); infra-orbital border more or less elevated, with 3 deep notches (Fig. 3 A, D). Anterior dorsal and distal parts of eyestalks fringed with long thick and thin setae. Antennal basal segment almost reaching small ventral prolongation of orbital hiatus; flagellum slender, long, about twice length of diameter of orbit (Fig. 3 D). Antennules folding obliquely (Fig. 3 D). Suborbital and pterygostomial surfaces tomentose but smooth underneath; subhepatic region tomentose with small scattered granules. Anterolateral margin with 4 teeth (excluding external orbital tooth), first tooth smallest, areolate, with 3 small flat granules on rim; second tooth bigger, areolate, ringed with small flat granules; third tooth longest, more conical, with 1 flat granule, directed anteriorly; fourth tooth low, rounded, ringed by small flat granules (Figs. 1, 2, 3 A). Posterolateral margin almost straight, gently converging towards posterior carapace margin, lined with rather 5 large rounded granules that form 3 granulated crests (Figs. 1, 2, 3 A). Posterior carapace margin gently convex, median part slightly concave (Figs. 3 A, 11 A). Third maxilliped almost completely covered by very short tomentum; merus quadrate, antero-external angle expanded, 3 flat granules lining mid-anteroexternal angle towards upper margin of merus; merus medially concave with a deep sulcus near lateral portion, a shallower sulcus near inner margin; ischium subquadrangular, with a deep median sulcus; exognath straight, inner subdistal angle with prominent triangular tooth, with distinct flagellum that extends beyond width of merus (Figs. 3 B, C, 5 A, 8 A); ischium and basis separated by prominent suture (Fig. 5 A). Epistome appears sunken; posterior margin divided into 4 lobes, median lobes rectangular, separated by deep narrow fissure, lateral lobes separated from median lobes by uneven cleft (Fig. 3 B, D). Endostome with long, prominent ridge running from distal edge of median lobe of posterior margin of epistome to near mandibles, interrupted subdistally, just as epistomal lobe ends. Both chelipeds large, subequal, laterally flattened, almost entirely covered with dense heavy tomentum (Figs. 2, 3 D, 4 A, B). Merus of cheliped with 2 large granules on proximal portion, 2 spinules on distal part, forming 2 granulated crests; carpus with 2 large tubercles, several nodules, granules on outer dorsal surface. Ischio-merus fused but suture still discernible. Chela somewhat flattened laterally; outer surface covered with numerous granules and tubercles of various sizes, with one median longitudinal ridge on outer surface evident, inner subventral surface with a curved row of sharp, orange-tipped tubercles; fingers shorter than palm; propodal finger with 3 nodules on inner margin, immovable finger with lower margin covered with dense thick tomentum, elevated, forming stiff fringe; dactylus with 2 deep sulci filled with short thick tomentum; dark brown pigmentation present only on distal part of fingers and cutting edges (Fig. 4 A, B). Ambulatory legs relatively long, relatively stout, thickly fringed by dense, stiff setae; P 3 longest, P 5 shortest, distinctly upturned (Figs. 1, 2). Merus of P 2 –P 4 with 2 longitudinal ridges (anterior one larger), distinctly meeting near distal segment on outer surface, margins fringed by dense stiff thick setae, margins somewhat carinate, dorso-subdistal area with deep cleft that forms prominent subdistal tooth with rounded tip and larger sharp sublamellate distal tooth which joins distinctly carinate proximal margin, area appears cup-like; merus of P 5 with 1 longitudinal bony ridge, cleft, 2 teeth, cup-like part on dorso-subdistal area less prominent (Figs. 2, 3 A, 4 C, D); ventral surface of merus of P 2 –P 4 with 2 or 3 long perpendicular spines on posterior margin, scattered granules on anterior margin; ventral surface of merus of P 5 without spines, only granules on posterior margin, scattered much smaller granules on anterior margin (Figs. 2, 3 A, 4 C, D). Carpus with prominent median ridge lined with several flat granules (Figs. 2, 3 A, 4 C, D). Propodus of P 2 –P 5 with 2 prominent median granular ridges (Figs. 2, 4 D). Dactylus long, as long as merus, covered by dense thick setae which hides tip, distal part not pigmented (Figs. 2, 4 C, D). Thoracic sternum relatively narrow transversely, entire surface covered with very short tomentum, small granules (Figs. 3 B, C, 9 A, 10 A, 12 A). Sternite 3 with shallow but distinct median longitudinal groove; sternite 4 as long as sternites 5 and 6 combined; sternite 7 very narrow longitudinally, posterior part tapering abruptly (Fig. 10 A), with posterior part of episternite 7 prolonged posteriorly to form spur which just touches coxa of P 5 and covers penis (Fig. 13 A, B); sternite 8 not exposed when abdomen closed (Fig. 10 A). Longitudinal line running through median part of sternites 7 and 8, those on sternites 4 and 6 incomplete, that of sternite 5 undiscernible (Fig. 12 A). Suture 1 / 2 absent; suture 2 / 3 complete; suture 3 / 4 distinct with median part relatively shallower; sutures 4 / 5 and 5 / 6 medially interrupted (Fig. 12 A); suture 6 / 7 appears complete but median portion very shallow (Fig. 12 A); suture 7 / 8 distinctly complete with median part deep (Fig. 12 A). Episternites 4–7 not prominently expanded laterally (Figs. 9 A, 10 A, 12 A). Sterno-abdominal cavity relatively narrow transversely, almost reaches anterior edge of sternite 4 (Figs. 9 A, 12 A). Press button of abdominal locking mechanism distinct, on posterior outer edge of sternite 5 of sterno-abdominal cavity (Fig. 12 A). Penis soft, effectively coxal, exits at base of coxa of P 5 but covered by posterior prolongation of episternite 7 (Figs. 13 A, B). Abdomen relatively narrow transversely, outer surfaces mostly covered with small low granules, very short, dense tomentum (Figs. 5 B, 9 A, 11 A). Telson triangular with gently concave lateral margins, rounded apex (Figs. 5 B, 9 A); somite 1 longitudinally and transversely broad, trapezoidal, exposed, not covered by posterior carapace margin (Fig. 11 A), somites 2–6 relatively narrow transversely, somite 3 broadest transversely; somite 2 with convex lateral margins, subrectangular in shape; somite 3 with gently convex lateral margins, trapezoidal; somites 4 larger than somite 5, both trapezoidal with gently concave lateral margins; somite 6 as long as broad, lateral margins concave (Figs. 9 A, 10 A); sutures between all somites and telson distinct, but somites 3 and 4 immovable, somites 4 and 5 slightly mobile (Figs. 5 B, 9 A, 11 A). Penis effectively coxosternal although it exits clearly from coxa, enters directly into sterno-abdominal cavity, not protected by sternal plates except for posterior part of sternite 7 which forms a spur-like structure that covers it (Figs. 13 A, B). G 1 relatively long, slender, with the rounded tip reaching suture of sternite 4, distal surfaces lined with numerous small spinules, tip open (Figs. 5 C–F). G 2 as long as G 1, very slender, long, with tip reaching beyond mid-portion of suture of sternite 5, distal segment ca. 0.2 times length of subdistal segment, base of distal segment forming cup-like structure lined with several spinules (Figs. 5 G, H). Females not known. Colour. The carapace and pereiopods are white without any distinct pigmentation, but this is barely discernible as the dark brown or orangish-brown tomentum and setae covers most of the animal (Fig. 1). The mandibles are a dark brown, like dark-coloured chitin, and this colour stays even after many months and many changes of alcohol (Fig. 8 A). The posterior margin of the epistome is lined with small orange and white granules when fresh. Some of the the larger granules on the carapace and pereiopods (especially those on the chelipeds) are orange. Etymology. The name is derived from the Latin anfractus for having many turns, alluding to the complex sculpture on the carapace. Remarks. The unusual carapace and pereiopods of V. anfractus are unique in the Goneplacoidea as presently understood. The complex labyrinth-like setal pattern is reminiscent of pilumnids like Pilumnus labyrinthicus Miers, 1884, or pseudoziids like Planopilumnus spongiosus (Balss, 1933) although they are clearly unrelated (see discussion above). Certainly none possess such a complex setal pattern and the ambulatory legs are not armed with spines. The dense carapace and pereiopod setation of V. anfractus superficially resembles those of the mathildellid, Intesius pilosus Guinot & Richer de Forges, 1981, but in the latter species, the setae are more scattered and the carapace lacks the dense setae and distinct setal arrangement of V. anfractus (see also Crosnier & Ng 2004). The holotype male of V. anfractus was collected from the rocky shoreline outside a Nipa mangrove near a river. It was crawling along on the shore in the afternoon when it was caught by the second author. We believe it had been actually caught by fishermen with tangle nets in the waters offshore as they had been observed cleaning their nets there earlier. In addition, there is a small piece of net stuck at the base of the chela of the specimen. It was almost certainly thrown onto the shore just before it was collected. The larger paratype male was collected from about 200m depth hiding inside a piece of sunken wood about 0.7 m in length and 0.3 m in diameter which was stuck in a tangle net (J. Arbasto, pers. comm.). The colour of the body and legs (dark brown) matches the wood substrate where the animal was found and it uses the spiniform ventral margins of the ambulatory meri to help anchor itself to the wood (J. Arbasto, pers. comm.). Freshly caught specimens were very dirty, being covered with detritus and mud, clearly a consequence of its setose body and pereiopods. This probably aids in camouflaging them against the wood as well. The single male specimen of Vultocinus anfractus collected from Vanuatu was from a tangle net set at about ₃ ₂ ɭΟ ɬ0 m depth. The specimen was obtained alive but we have no indication if it had been living on wood. This specimen differs from the type series of the species in having an orangish-brown body (versus dark brown), somewhat rounder carapace, and proportionately slightly longer ambulatory legs (Figs. 1 C, 2 C). It agrees with the type material, in almost all other aspects, and we prefer to regard it as conspecific with the Vultocinus anfractus material from the Philippines for the moment. Although mature, it is the smallest specimen of Vultocinus available, and the differences observed may well be due to size and habitat. A larger series of specimens from Vanuatu or other areas will be needed to ascertain this. The specimen of V. anfractus was observed alive for several hours before being preserved. It moved slowly and was not aggressive when handled, but that may have been due to it being tangled in the nets for many hours. When placed against a hard substrate, the ambulatory legs invariably clamp backwards, confirming what we believe the spines on the meri are used for. It is unfortunate that all the specimens collected were males, so key female characters could not be determined.

Vultocinus, new genus Diagnosis. Entire dorsal carapace surface covered with short, stiff setae, with underlying complex pattern of ridges and grooves, regions well marked by high tuberculated and granulated swellings (Figs. 2, 3 A). Mandibles dark coloured even after in preservatives (Fig. 8 A). Thoracic sternum relatively narrow transversely (Fig. 9 A), sternite 3 with shallow median longitudinal groove (Fig. 9 A), posterior edge of episternite 7 strongly produced posteriorly to form spur which reaches coxa of P 5 (Fig. 13 A, B). Meri of ambulatory legs with distinct median carinae, ventral margins of P 2 ���P 4 with 2 or 3 prominent perpendicular spines each (Fig. 4 C, D). Male abdomen relatively narrow transversely (Figs. 9 A, 10 A), sutures for all somites and telson clearly visible but somites 3 and 4 immovable, fused, other somites freely articulating (Fig. 5 B). G 1 relatively slender, gently curved, surface with simple spines (Fig. 5 C-F); G 2 with distal segment short, about one-sixth length of basal segment (Fig. 5 G, H). Etymology. The name is derived from the Latin vultuosus for ���full of expression���, alluding to the distinctively marked carapace features of the type species. The name is an arbitrary combination with the Latin word for crab, Carcinus. Gender masculine. Remarks. See discussion for family., Published as part of Ng, Peter K. L. & Manuel-Santos, Marivene R., 2007, Establishment of the Vultocinidae, a new family for an unusual new genus and new species of Indo-West Pacific crab (Crustacea: Decapoda: Brachyura: Goneplacoidea), with comments on the taxonomy of the Goneplacidae, pp. 39-68 in Zootaxa 1558 on page 49, DOI: 10.5281/zenodo.178296

Geosesarma lawrencei, new species (Figs. 1, 2) Material examined. Holotype, male (10.3 x 9.7 mm) (NMCR 27023), Sitio Kamantian, Barangay Samarinana [=Samareñana], Brooke’s Point, Palawan, Philippine Islands. 550 m asl, coll. L. Liao, Dec. 2001. Paratypes: Male (9.5 by 8.6 mm) (ZRC 2007.0121), female (9.9 by 9.5 mm) (NMCR 28011), same data as holotype. Diagnosis. Carapace squarish, very slightly broader than long, anterior dorsal carapace and anterolateral regions covered with groups of small sharp granules. Frontal margin strongly sinuous, bilobed, subequal to half carapace width. Postfrontal cristae distinctly separated by deep median groove, each crista bilobed with each lobe separated by shallow groove. External orbital angle acutely triangular, directed obliquely outwards, separated from first epibranchial tooth by broad, shallow but distinct V-shaped cleft. Lateral margin slightly convex posteriorly. Exopod of third maxilliped not reaching midlength of merus, with numerous long setae distally, with well-developed flagellum exceeding width of merus. Ambulatory legs relatively long, slender. Male abdomen triangular; lateral margins of fifth and sixth segments convex. G 1 slender, almost straight; distal part short, less than 0.2 times length of proximal part, bent very slightly outwards, tip rounded; proximal part with upper inner margin gently convex, outer margin gently undulating. Description of holotype. Carapace squarish, very slightly broader than long, dorsal surface covered with small sharp granules in groups of two or three; regions distinctly demarcated, with relatively deep grooves; median gastric groove deep, H-shaped; anterior grooves shallow; transverse groove separating cardiac and intestinal regions distinct (Fig. 1 A). Frontal margin strongly deflexed, strongly sinuous, bilobed, subcristate (Fig. 1 A–C). Postfrontal cristae distinct, sharp, separated by deep median groove, each crista bilobed, each lobe separated by shallow groove; surface of frontal region vertically concave (Fig. 1 A, B). External orbital angle acutely triangular, directed obliquely outwards, separated from first epibranchial tooth by broad, shallow but distinct V-shaped cleft; first epibranchial tooth smaller than external orbital angle, triangular with very shallow posterior notch delineating it from lateral margin; second epibranchial tooth separated from the first epibranchial tooth by very shallow notch, third indistinct (Fig. 1 A). Lateral margin slightly convex posteriorly; posterolateral regions with low oblique striae (Fig. 1 A). Eyes large, reaching the tip of external orbital angle (Fig. 1 A–C). Merus of third maxilliped round, with median portion of lateral margin produced as slight, blunt angle, about two times greater than proximal margin width, with strong median oblique ridge; ischium with shallow median sulcus, outer ridge distinct; exopod slender, not reaching midlength of merus, with numerous long setae distally, with well-developed flagellum exceeding width of merus (Fig. 1 B, C, 2 C). Chelipeds subequal; outer surface of palm spinulose, covered with low, scattered spinules especially on upper and lower margins; fingers subequal to length of palm, proximal part of upper margin of dactylus of left chela lined with 12 low, anteriorly-directed granules (10 granules on right chela of dactylus), cutting edge lined with numerous teeth and denticles, distal part of finger pectinated to form blade-like cutting edges; inner margin of carpus gently serrated, inner distal spine weak but distinct, with scattered low granules on dorsal surface; outer surface of merus slightly rugose, inner surface smooth, with serrated inner margin, without subdistal spine (Fig. 1 A–C). Ambulatory legs long, slender; third pair longest; merus broad, with low, truncated subdistal dorsal tooth, upper margin very gently serrated; propodus slender; dactylus shorter than propodus, straight, gently curved distally; segments lined with numerous short, stiff spiniform hairs. Abdomen triangular, lined with numerous short setae; seventh segment with rounded tip, longer than sixth, lateral margins of sixth and seventh segments convex (Fig. 1 A, C). G 1 slender, almost straight; distal part bent very slightly outwards, pectinated, straight then gently tapering to a rounded tip, tube-like with distal opening on dorsal side, laterally flattened towards ventral side; proximal part with upper inner margin gently convex; outer margin gently undulating, distally with numerous setae (Fig. 2 A, B). G 2 very short, less than 0.3 times length of G 1, lacking flagellum, expanded proximally, tapered distally, curved, with distal part directed outwards (Fig. 2 D). Paratypes. The paratype male is slightly smaller than the holotype but agrees with it in all major aspects. The paratype female is similar to the male holotype except for the external reproductive structures. Etymology. The species is named after its collector, Dr. Lawrence Liao, whose interests extend far beyond his love and dedication for his work as a phycologist. He is also an avid field collector who has contributed a wealth of new discoveries to the fauna of the Philippines, including crabs. Remarks. The slender G 1 of Geosesarma lawrencei, new species, is similar to that of G. rathbunae from Panay Island, which is also the geographically closest Philippine congener. However, G. l a w re n c e i is easily distinguished from G. rathbunae by the following characters: frontal margin strongly sinuous, distinctly bilobed when viewed dorsally (versus frontal margin gently sinuous, weakly bilobed when viewed dorsally); frontal margin subequal in length to half carapace width resulting in eyes appearing closer together (versus frontal margin longer than half carapace width, eyes appearing further apart); postfrontal cristae distinctly separated by deep median groove, with each crista bilobed and each lobe being separated by shallow groove (versus postfrontal cristae weakly separated by shallow median groove, each crista with undulating margin, not appearing bilobed); cleft separating external orbital angle from first epibranchial tooth comparatively broad and shallow (versus cleft narrower and deeper); carapace lateral margins slightly convex posteriorly (versus lateral margins straight); ambulatory legs comparatively long and slender (versus ambulatory legs shorter and stouter); G 1 almost straight with distal part bent very slightly outwards, with a rounded tip (versus G 1 with distal part bent obliquely outwards at approximately 30 degrees to the long axis, with a sharp tip); G 1 outer margin gently undulating (versus outer margin straight); pectinated distal part of G 1 proportionally shorter, about 0.17 times of proximal part (versus pectinated distal part proportionally longer, about 0.36 times of proximal part); and G 1 proximal part with upper inner margin gently convex (versus proximal part upper inner margin distinctly convex, appearing hump-like) (Figs. 1 A–C, 2 A, B; cf. Serène 1968: Figs 5, 6, Pl. 1 fig. 4). Geosesarma lawrencei can be distinguished from G. protos by the squarish carapace (versus trapezoidal carapace); the well-developed flagellum on the exopod of the third maxilliped (versus third maxilliped lacking a flagellum on the exopod); and the slender G 1 (versus stout G 1) (Figs. 1 A, 2 A–C; cf. Ng & Takeda 1992: Figs. 1 A, B, G–J). There is currently some doubt over Serène and Soh’s (1970) generic placement of Geosesarma vicentese due to its unusual morphology (Ng & Takeda 1992). In any case, G. vicentese is least likely among the Philippine species to be confused with G. lawrencei as the new species differs considerably from the former in the anterior portions of the carapace and anterolateral regions being covered with sharp granules (versus flattened granules); the postfrontal cristae being strongly separated by distinct deep grooves (versus postfrontal cristae weakly separated by gentle, shallow grooves); and the lateral margins of the fifth and sixth male abdominal segments being convex (versus concave) (Figs. 1 A–C; cf. Rathbun 1914; Ng & Takeda 1992). In addition to the known species of Geosesarma from the Philippines, two other congeners, G. sabanus Ng, 1995 a, and G. a u r a n t i u m Ng, 1995 b, both from Sabah in northern Borneo, warrant comparison by virtue of the proximity of their type localities to Palawan. Both species, however, are clearly differentiated from G. lawrencei by the absence of a flagellum on the third maxilliped and by the stout and distally bent form of their G 1 (versus flagellum present on third maxilliped and slender and almost straight G 1) (Fig. 2 A–C; cf. Ng 1995 a: Figs. 1 B, E–G, 1995 b: Figs. 1 G, 2). Distribution. Palawan, Philippines.

Vultocinidae, new family Diagnosis. Dorsal surface of carapace subquadrate, with complex pattern of ridges, grooves, dense short setae (Figs. 2, 3 A). Mandibles prominently dark coloured even after prolonged preservation (Fig. 8 A). Thoracic sternum relatively narrow transversely (Fig. 9 A); episternites 4–6 not prominently expanded laterally (Figs. 9 A, 10 A, 12 A); sternite 7 very narrow longitudinally, tapering abruptly (Fig. 10 A), posterior part of episternite 7 prominently prolonged backwards to form long spur which covers penis and just touches coxa of P 5 (Figs. 13 A, B); sutures 4 / 5 and 5 / 6 prominently interrupted medially (Fig. 12 A); suture 6 / 7 appears complete but with median portion very shallow but discernible (Fig. 12 A); suture 7 / 8 distinctly complete with median part deep (Fig. 12 A); sternite 8 not exposed when abdomen closed (Fig. 10 A). Ventral margins of meri of P 2 – P 4 with 2 or 3 strong perpendicular spines (Fig. 4 C, D). Sterno-abdominal cavity almost reaches anterior edge of sternite 4 (Figs. 9 A, 12 A); press button of male abdominal locking mechanism distinct, on posterior edge of thoracic sternite 5 (Fig. 12 A). Penis soft, effectively coxal, exits at base of coxa of P 5 but covered by posterior prolongation of episternite 7 (Figs. 13 A, B). Male abdomen relatively narrow overall (Figs. 3 B, 5 B, 9 A, 10 A), sutures for all somites and telson clearly visible; somite 1 transversely and longitudinally broad, trapezoidal in shape, exposed, not covered by posterior carapace margin (Fig. 11 A), somites 2–6 relatively narrow transversely, somite 6 as long as broad (Figs. 9 A, 10 A), somites 3, 4 immovable, fused, other somites freely articulating (Fig. 5 B). G 1 relatively slender, gently curved, surface with simple spines (Fig. 5 C-F); G 2 with short distal segment, about 0.2 times length of basal segment (Fig. 5 G, H). Type genus. Vultocinus, new genus. Remarks. Vultocinus, new genus, possesses a suite of unusual characters that make its precise affinities difficult to ascertain. Superficially, its carapace shape and setation pattern resembles species of Lophoplax Tesch, 1918, but members of this genus have slender and sinuous G 1 s with a very short sigmoid G 2, and are currently classified in the subfamily Rhizopinae Stimpson, 1858, family Pilumnidae (see Takeda 1977; Ng & Davie 1991; Ng & Huang 2002; review by Ng 1987). The general carapace shape and setose surfaces of Vultocinus also resembles members of the genus Planopilumnus Balss, 1933, of the subfamily Planopilumninae Serène, 1984, family Pseudoziidae; but members of this family have stout G 1 s and a G 2 which is only half the length of the G 1 (see Ng & Clark 2000 a, b; Ng et al. 2001; Ng & Liao 2002). The possession of male abdominal somites 2 and 3 which are immovable, of a long and relatively stout G 1 and of a long G 2 which is subequal in length to the G 1, appears to ally Vultocinus with some members of the Goneplacidae MacLeay, 1838 sensu lato. While the taxonomy of this family has been a major problem for many years (see Guinot & Breton 2006), recent studies which built on the key contributions made by Guinot (1969 a – c, 1971) have made the systematic position of the taxon much clearer. This is especially so when using the suite of many new characters pioneered by her over the years (see Guinot 1979; Guinot & Bouchard 1998). Over the years, many taxa and subfamilies have been transferred from the Goneplacidae into other families (see Castro, in press, for review) and until 2003, the Goneplacidae was believed to be composed of five subfamilies, viz. the Goneplacinae MacLeay, 1838, Carcinoplacinae H. Milne Edwards, 1852, Euryplacinae Stimpson, 1871, Chasmocarcininae Serène, 1964, and Trogloplacinae Guinot, 1986 (see Ng et al. 2001; Davie 2002). Recent studies have substantially changed the landscape of goneplacid taxonomy – Karasawa & Kato (2003 a, b), Števčić (2005), Karasawa & Schweitzer (2006) and Castro (in press); which are discussed individually below. These discussions are not always restricted to the extant fauna, a major shortcoming since there is often a need to use many important characters like male abdominal structure, as well as detailed sternal and penial features which are rarely observable in fossils. Karasawa & Kato (2003 a, b) reappraised the phylogeny of the Goneplacidae and rearranged the subfamilial system based on selected extant and fossil taxa. They synonymised the Carcinoplacinae under the Goneplacinae, and also established a new subfamily, Mathildellinae, for Beuroisia Guinot & Richer de Forges, 1981, Intesius Guinot & Richer de Forges, 1981, Mathildella Guinot & Richer de Forges, 1981, Platypilumnus Alcock, 1894, and Neopilumnoplax Serène, 1969, which had originally been placed in the Carcinoplacinae. Although Karasawa & Kato (2003) define the Mathildellinae with a suite of carapace characters, one of the most diagnostic characters (but unfortunately only usable for fresh specimens) is whether somites 3 and 4 are mobile. All known extant adult male goneplacines (and carcinoplacines) have somites 3 and 4 freely mobile (with one exception, Neommatocarcinus Takeda & Miyake, 1969, which has somites 3–5 fused, see also Castro, in press). On the other hand, all extant adult male mathildellines have abdominal somites 3 and 4 immobile although the sutures separating them are still visible (see also Ng & Chan 2000; Ng & Ho 2003; Crosnier & Ng 2004). The articulation between somites 4 and 5 is usually more difficult to ascertain. In the mathildellines we have examined, there is much more limited movement between these somites compared to other somites. In a major reappraisal, Števčić (2005) substantially reorganized the family but with minimal or no discussion. He restricted what he defined as the superfamily Goneplacoidea to just three families: Goneplacidae (with two subfamilies, Goneplacinae and Carcinoplacinae, with two tribes—Carcinoplacini, Psopheticini Števčić, 2005), Geryonidae Colosi, 1923 (with three subfamilies, Geryoninae with six tribes—Geryonini, Progeryonini Števčić, 2005, Paragalenini Števčić, 2005, Mathildellini, Intesiini Števčić, 2005, Platycheloniini Števčić, 2005; Platypilumninae Števčić, 2005; and Bathyplacinae Števčić, 2005), and Planopilumnidae (as Planopilumninae [sic]) Serène, 1984. Many other taxa which had been placed in the Goneplacidae up to 2005 were transferred to other groups or whole new families and subfamilies. His recognition of some of them as families, e.g. the Euryplacidae and Chasmocarcininae as distinct families was expected as there had been much prior discussion on their status, although his unexplained elevation of both groups to superfamily status seems unwarranted. The same applies for placing Scalopidia Stimpson, 1858, into its own family, the Scalopidiidae Števčić, 2005, in the superfamily Chasmocarcinoidea. The subfamily Trogloplacinae Guinot, 1986 (regarded as a family by Davie 2002), has clear chasmocarcinid affinities (see Guinot & Davie 1996), but this was ignored and Števčić (2005) recognised it as a distinct superfamily, separate from his Chasmocarcinoidea. Two genera which had been allied to the Chasmocarcinidae, Parapilumnus Kossmann, 1877, and Acidops Stimpson, 1871, were inexplicably referred to their own families (Parapilumnidae Števčić, 2005, and Acidopsidae Števčić, 2005, respectively) and superfamilies in two different parts of his paper, although the affinities between them are very clear and should be classified together (Ng 2002). Two other genera linked with the Chasmocarcinidae, Raoulia Ng, 1987, and Typhlocarcinodes Alcock, 1900, were also referred to their own families (Raoulidae Števčić, 2005, and Typhlocarcinodidae Števčić, 2005, respectively), and own superfamilies. One genus clearly affiliated with the Goneplacidae and Goneplacinae sensu lato, Notonyx A. Milne- Edwards, 1873, was removed to its own family (Notonycidae Števčić, 2005) and superfamily Notonycoidea Števčić, 2005 (see also Clark & Ng 2005; Castro in press). The genus Conleyus Ng & Ng, 2003, originally placed in the Carcinoplacinae, was refered to its own family (Conleyidae Števčić, 2005). Some of his actions are very difficult to rationalize, e.g. Parapilumoidea and Acidoposidea, even if both are not true chasmocarcinids as defined by Guinot & Davie (1996). The same situation prevails for Raoulia and Typhlocarcinodes. Both genera may not be chasmocarcinids, but to classify them as belonging to separate superfamilies is difficult to understand. The case with Notonyx and Notonycoidea also suggests that Stevčić (2005) placed too much emphasis on general external morphology. While Conleyus has many peculiar features, the need for a separate superfamily is also unwarranted because the genus has many affinities with goneplacids. Stevčić’s (2005) inclusion of Geryonidae and Planopilumnidae into his Goneplacoidea is also difficult to understand. This is also true for his transfer of the Mathildellinae (as a tribe) into his Geryoninae and the removal of genera previously included in it like Intesius and Platypilumnus into its own tribe and subfamily respectively. The affinities of the geryonids with portunoids has been well argued (see Ng & Manning 1998; Ng & Guinot 1999), while planopilumnids have been included in the Pseudoziidae Alcock, 1898 by Ng et al. (2001) and Ng & Liao (2002). Intesius (and Platypilumnus) is closely allied to Mathidella (see also Karasawa & Kato 2003 a; Crosnier & Ng 2004) and separating them makes little sense. The lack of any explanation or justification by Stevčić (2005) makes these taxonomic actions and transfers difficult to accept, refute or even discuss. Karasawa & Schweitzer (2006), in another reappraisal of the Goneplacidae, this time comparing it with the Xanthoidea, followed some aspects of Stevčić’s (2005) system, but argued that the Mathildellinae was actually a portunoid and recognized it as a separate family there. They regarded the Goneplacinae as synonymous with the Carcinoplacinae; and placed both the Mathildellidae (as a family) and the Geryonidae in the Portunoidea. The transfer of the Mathildellinae is difficult to contend with as with regards to the general form of the carapace, sternal, abdominal and gonopodal characters, it has many affinities with the goneplacids. Other parts of their Goneplacoidea appear to be more reasonable, with the Euryplacidae, Chasmocarcinidae, Trogloplacidae and Hexapodidae Miers, 1886. More difficult to accommodate is their transfer of the genus Progeryon Bouvier, 1922, to its own family (i.e. the Progeryonidae Števčić, 2005) and superfamily which they justified stating that the form of the male thoracic sternum is different from those in other portunoids, including mathildellids. They left the matter of Paragalene Kossmann, 1878, and the Paragaleninii Števčić, 2005, unresolved, Most recently, Castro (in press) reviewed and revised the members of the species which have been placed in Goneplacinae MacLeay, 1838, and Carcinoplacinae H. Milne Edwards, 1852, and supported Karasawa & Kato’s (2003) synonymisation of these two subfamilies. He also recognised the Mathildellinae as a separate subfamily within the Goneplacidae, and synonymised Notonycidae under the Goneplacinae. As to the status of genera like Bathyplax A. Milne-Edwards, 1880, Progeryon, Paragalene, Conleyus, etc., he argued that they were not certainly not goneplacines in the strict sense, but may perhaps be retained in the Goneplacidae sensu lato provisionally. How does Vultocinus fit into all these recent developments? The problem is that many of the subfamilies or families defined by the earlier workers have omitted key characters associated with the thoracic sternum, abdominal locking mechanism, position of the penis and associated structures. The most comprehensive study (Castro, in press) has many of the characters that are deemed useful in a modern classification but it is mostly restricted to the Goneplacinae sensu stricto, although he does briefly discuss many of the other “goneplacid allies”. To resolve the systematic position of Vultocinus, it was important that representative genera of the various supra-generic groupings discussed above be re-examined. To this effect, we examined in detail representative genera from the Goneplacinae s. str., Mathildellinae s. str., as well as two of the problematic genera, Progeryon and Conleyus, and compared them with Vultocinus. The position of the press button of the abdominal locking mechanism (see Guinot & Bouchard 1998) on or along thoracic sternite 5 is important. While all of the genera considered here have the button on thoracic sternite 5 at the lateral edges of the sterno-abdominal cavity,there are two distinct patterns. In one group with Goneplax Leach, 1814, and Carcinoplax H. Milne Edwards, 1852, the press button is on the anterior part of sternite 5, close to sternite 4 (Fig. 12 B, C). In all the other genera— Mathildella, Intesius, Platypilumnus, Progeryon, Conleyus and Vultocinus, the press button is on the posterior part of sternite 5, close to sternite 6 (Fig. 12 A, D–H). This correlates somewhat with the extent of the sterno-abdominal cavity in some cases. In Goneplax and Carcinoplax, the sterno-abdominal cavity is usually relatively deep, and extends almost to the anterior edge of thoracic sternite 4 (Figs. 9 B, C, 12 B, C). In Mathildella, Intesius and Platypilumnus, the sterno-abdominal cavity is usually relatively shallow, and extends only to the middle part of thoracic sternite 4 (Figs. 9 D–F, 12 D–F). The position of the sterno-abdominal cavity in Progeryon, however, is like that of Goneplax and Carcinoplax, almost reaching the anterior edge of sternite 4 (Fig. 9 G) but the cavity is relatively shallower (Fig. 12 G). The position of the sterno-abdominal cavity in Conleyus is intermediate between that of Goneplax, Carcinoplax and Progeryon on one hand, and Mathildella, Intesius and Platypilumnus on the other (Fig. 9 H), although the cavity itself is relatively deep (Fig. 12 H). Vultocinus stands out in that it has a deep sterno-abdominal cavity which is relatively narrow transversely (Fig. 12 A) and almost reaches the anterior edge of sternite 4 (Fig. 9 A). A note on the use of the extent of the sterno-abdominal cavity as a character is useful. In some taxa, the anterior-most point of the cavity extends beyond the tip of the telson, even when the male abdomen is fully closed and locked into position by the press button mechanism. It is most distinct in genera like Goneplax, Carcinoplax and Vultocinus (Fig. 9 A–C), and to a lesser extent by genera like Progeryon and Conleyus (Fig. 9 G, H). It is not so obvious in Mathildella, Intesius and Platypilumnus (Fig. 9 D–F) because the sterno-abdominal cavity is relatively shallower and the male abdomen always locks onto the sternum very tightly. The gap is presumably to enable the animals to defecate without moving the rest of the abdomen or even the telson (see also Guinot & Bouchard 1998). The structure of the sutures 6 / 7 and 7 / 8 are useful in understanding the relationships between the groups. In the two species of Goneplax examined in this study, suture 6 / 7 can be regarded as complete even though it becomes rather shallow medially and appears interrupted (Fig. 12 B). In Carcinoplax, the same suture also appears to be complete but there is a narrow median gap (Fig. 12 C) and so may be regarded as interrupted. In the case of Vultocinus, this suture curves anteriorly, becoming very shallow medially so it is not as easy to ascertain if this is truly complete or a case of incipient interruption (Fig. 12 A). In the Goneplacinae sensu Castro (in press), suture 6 / 7 varies between complete and interrupted. In all the other genera examined here, suture 6 / 7 is clearly complete (Fig. 12 D–H). With regards to suture 7 / 8, in Goneplax, it is prominently interrupted medially (Fig. 12 B). In Carcinoplax, the suture appears to be complete but there is a narrow median interruption (Figs. 7 E, 12 C) and so may be regarded as interrupted as well. According to Castro (in press), while most Goneplacinae are like Goneplax and Carcinoplax, there some taxa which have suture 7 / 8 complete. In all the other genera examined here (including Vultocinus), suture 7 / 8 is clearly complete (Fig. 12 A, D–H). Also of interest is the arrangement of thoracic sternite 8 in relation to the adjacent coxa of P 5 and male abdomen. Guinot (1969 b: 520) first highlighted this interesting character in her attempts to better understand goneplacoid relationships. In genera like Goneplax and Carcinoplax, part of sternite 8 is still clearly visible even after the abdomen is completely closed. In species of Goneplax sensu stricto, the exposed part of sternite 8 is quite substantial (Fig. 7 C) (see also Guinot 1969 b: Figs. 63, 64). In at least some of the Indo-West Pacific species of “ Goneplax ”, the exposed part of sternite 8 is relatively small (Figs. 10 B, 11 b), while in genera like Carcinoplax, the exposed part of sternite 8 varies from small (Figs. 10 C, 11 C) to relatively large (see Guinot 1969 b: Figs. 60 –62, 65). In genera like Mathildella, Intesius and Platypilumnus, the exposed part of sternite 8 is either completely covered by the abdomen (Figs. 10 D, F, 11 D, F) or just visible with only a small part of the edge visible (Figs. 10 E, 11 E). In Progeryon and Conleyus, the lateral part of sternite 8 completely covered by the male abdomen (Figs. 10 G, H, 11 G, H). The position of the penis is of fundamental importance (Guinot, D., pers. comm.). This character is closely associated with the shape of episternite 7 and the lateral margin of abdominal somite 3. In all cases, the exposed part of the penis is protected by abdominal somite 3 when the abdomen is closed. How this is accomplished is what distinguishes each of the groups. In the Goneplacidae sensu stricto, the penis leaves the coxa at its base, but continues under the sternum for a short distance, only exiting and becoming completely exposed inside the sterno-abdominal cavity after the coxal condyle (Fig. 13 C, D). Episternite 7 is expanded laterally to partially overlap the entire inner margin of the coxa of P 5 (Figs. 7 D, 10 B, 13 C) or at least most of the anterior part (Figs. 10 C, 12 D). The lateral edge of abdominal somi

The Viviparidae, commonly known as River Snails, is a dominant group of freshwater snails with a nearly worldwide distribution that reaches its highest taxonomic and morphological diversity in Southeast Asia. The rich fossil record is indicative of a probable Middle Jurassic origin on the Laurasian supercontinent where the group started to diversify during the Cretaceous. However, it remains uncertain when and how the biodiversity hotspot in Southeast Asia was formed. Here, we used a comprehensive genetic data set containing both mitochondrial and nuclear markers and comprising species representing 24 out of 28 genera from throughout the range of the family. To reconstruct the spatiotemporal evolution of viviparids on a global scale, we reconstructed a fossil-calibrated phylogeny. We further assessed the roles of cladogenetic and anagenetic events in range evolution. Finally, we reconstructed the evolution of shell features by estimating ancestral character states to assess whether the appearance of sculptured shell morphologies was driven by major habitat shifts. The molecular phylogeny supports the monophyly of the three subfamilies, the Bellamyinae, Lioplacinae, and Viviparinae, but challenges the currently accepted genus-level classification in several cases. The almost global distribution of River Snails has been influenced both by comparatively ancient vicariance and more recent founder events. In Southeast Asia, Miocene dispersal was a main factor in shaping the modern species distributions. A recurrent theme across different viviparid taxa is that many species living in lentic waters exhibit sculptured shells, whereas only one strongly sculptured species is known from lotic environments. We show that such shell sculpture is habitat-dependent and indeed evolved several times independently in lentic River Snails. Considerably high transition rates between shell types in lentic habitats probably caused the co-occurrence of morphologically distinct shell types in several lakes. In contrast, directional evolution toward smooth shells in lotic habitats, as identified in the present analyses, explains why sculptured shells are rarely found in these habitats. However, the specific factors that promoted changes in shell morphology require further work. [biogeographical analyses; fossil-calibrated phylogeny; fossil-constrained analyses; Southeast Asia; stochastic character mapping.].