Fejervarya moodiei (Taylor, 1920)

Yodthong, Siriporn, Stuart, Bryan L. & Aowphol, Anchalee, 2019, Species delimitation of crab-eating frogs (Fejervarya cancrivora complex) clarifies taxonomy and geographic distributions in mainland Southeast Asia, ZooKeys 883, pp. 119-153 : 119

Fejervarya moodiei (Taylor, 1920)

Rana moodiei Taylor, 1920: 234

Rana cancrivora : Taylor 1962: 377

Fejervarya moodiei : Dubois and Ohler 2000: 35; Brown et al. 2013: 17

Fejervarya cancrivora : Chan-ard 2003: 107; Chuaynkern and Chuaynkern 2012: 169; Kurniawan et al. 2010: 3

Fejervarya Bangladesh mangrove type Islam et al. 2008: 1084

Fejervarya cancrivora mangrove type Kurniawan et al. 2010: 222; Kurniawan et al. 2011: 12

Fejervarya cf. cancrivora Harikrishnan & Vasudevan, 2018: 241

Diagnosis.

Fejervarya moodiei can be characterized by the following combination of characters: (1) medium to large size, SVL 42.7-62.7 mm in males, 50.0-81.8 mm in females (Table 4 ; Appendix 2, 3); (2) head length slightly greater than head width; (3) skin on dorsum and flank with spinules, and glandular warts, with irregular skin folds not arranged in series, with darker marking on dorsal surface of forelimbs and hindlimbs; (4) relative finger lengths II <IV <I <III; (5) Most individual have dermal fringe on fingers II and III; (6) prepollax indistinct; (7) palmar tubercles indistinct; (8) foot moderately webbed, with webbing formula I1 –11/2II1–2III1–2IV2– 1V; (9) dermal flap on postaxial side of Toe V; (10) Fejervaryan lines absent; (11) inner metatarsal tubercles prominent; (12) indistinct inner tarsal ridge on distal half to two-thirds of tarsus ( Fig. 6 C–D ) and (13) vocal sacs in adult males with wrinkled skin covered by triangular, very dark brown blotches on each side of throat.

Description of holotype.

Taylor (1920) described the species based on an adult female, CM 3724, from Manila, Luzon, Philippines ( Fig. 8A, B ; Appendix 3). We supplement his description of the holotype, as follows: rather large body size; head narrow, slightly longer than wide; snout tip oval in dorsal view, round in lateral view, projecting beyond lower jaw; nostril dorsolateral, oval, with small lateral flap, closer to tip of snout than eye; canthus indistinct, rounded; loreal region slightly concave and oblique [loreal region broadly sloping, not concave according to Taylor (1920)]; eye diameter about 60% snout length [eye diameter equal to snout length according to Taylor (1920)]; interorbital region flat, about half width of upper eyelid and slightly less than internarial distance; pineal body present; tympanum distinct, rounded, about 90% of eye diameter, not depressed relative to skin of temporal region, tympanic rim weakly elevated relative to tympanum, dorsoposterior margin obscured by supratympanic fold; vomerine ridge present in two strongly oblique series, very slightly closer to each other than to choanae [based on Taylor (1920), not examined by us].

Forelimbs short, rather robust; fingers rather long, slightly swollen; tips of fingers slightly rounded, terminus slightly swollen but not expanded into discs; relative finger lengths II <IV <I <III [first finger longer than second and fourth according to Taylor (1920)]; dermal fringe on fingers absent; webbing on fingers absent; subarticular tubercles distinct; supernumerary tubercles absent; prepollex indistinct, oval; palmar tubercles indistinct.

Hindlimbs moderately short, robust; tibia slightly longer than thigh, but shorter than distance from base of inner metatarsal tubercle to tip of Toe IV; toe long, stout; tips of toes rounded, not expanded into discs; relatively toe lengths I <II <III <IV, webbing moderate, deeply excised between toes, formula I1 –11/2II1–1III1–2IV2– 1V, Toe I webbed to base of distal phalanx; preaxial side of Toe II webbed to point between distal subarticular tubercle and distal phalanx, continuing as narrow fringe to base of distal phalanx; postaxial side of Toe II webbed to base of distal phalanx; preaxial side of Toe III webbed to distal subarticular tubercle, continuing as narrow fringe to base of distal phalanx, postaxial side of Toe III webbed to base of distal phalanx; preaxial side of Toe IV wedded to webbed to proximal distal subarticular tubercle, continuing as narrow fringe to base of distal phalanx, postaxial side of Toe IV wedded to webbed to proximal distal subarticular tubercle, continuing as narrow fringe to base of distal phalanx, Toe V webbed to base of distal phalanx; dermal flap well developed, extending along postaxial side of Toe V from level of inner metatarsal tubercles to distal phalanx; subarticular tubercles prominent, inner metatarsal tubercle prominent, oval, length about 30% that of Toe I; distinct dermal ridge extending along inner metatarsal tubercle to distal phalanx of Toe I; indistinct inner tarsal ridge on distal two-third of tarsus ( Fig. 7C ); outer metatarsal tubercles absent; supernumerary tubercles absent; tarsal tubercle absent.

Skin on snout and between the eyes shagreened; skin on eyelid shagreened with glandular warts; skin on dorsum shagreened with glandular warts and irregular skin folds; dorsolateral fold extending posteriorly to two-thirds length of dorsum; skin on side of head smooth; skin on flank with glandular warts; skin on cloacal region with glandular warts; forelimbs shagreened; thigh with indistinct glandular warts; tibia, tarsus, throat, chest and belly smooth.

Coloration of holotype in preservative.

Coloration mostly lost in preservative. Dorsum and side of head medium brown with a few dark brown markings; tympanum translucent brown with pale brown spot in center; flank pale brown with faint brown marbling; three wide brown vertical spots on upper lips; dorsal surfaces of forelimbs, thigh, tibia, and foot medium brown with a few dark brown spots, posterior surface of thigh with irregular pattern of indistinct dark brown marbling on light background; chin, chest, belly, and ventral surfaces of forelimb and hindlimb pale brown; ventral surfaces of hand and foot pale brown; lower lip pale brown with a few dark brown spots; vertebral and tibial stripes absent; Fejervaryan lines absent.

Coloration of referred Thai specimen in life.

Adult male ZMKU AM 01390 ( Fig. 9 A–E ) from Mueang Phang-nga District, Phang-nga Province, Thailand. SVL 44.7 mm. Dorsum and side of head light brown with indistinct olive brown marking; olive-brown band between outer margin of upper eyelids; tympanum with orange-brown blotches in center; olive-brown streak on canthus rostralis from tip of snout to eye; dark brown streak from eye along supratympanic fold to posterior rim of tympanum; flank creamy white with dark brown marbling; three wide dark brown spots on upper lips; dorsal part of limbs: forelimbs, thigh, tibia, and foot light brown with olive-brown transverse spots, posterior part of thigh with irregular pattern of dark brown marbling on creamy yellow background; ventral part of body: chin creamy white with indistinct mottled dark brown, triangular dark brown blotches and mottling on each side of throat; forelimbs, chest, belly creamy white and hindlimbs with indistinct dark brown mottling, hand brown and foot dark brown; lower lip creamy white with dark brown spots; Fejervaryan lines absent.

Variations.

Vomerine ridges slightly closer to choanae than to each other in some individuals. Most adult males have nuptial pads with small translucent spinules on dorsal and medial surface of Finger I from base of distal phalanx to base of prepollax, but some individuals have the nuptial pad extending to slightly over the base of prepollex. Most adult males have dense, fine spinules covering only the chin, but some individuals have dense, fine spinules on the chin and chest. Adult males have vocal sac present on each side of throat with wrinkled skin covered by triangular, very dark brown blotches. Adult males with larger spinules and glandular warts on dorsum, dorsal surfaces of forelimbs, flank, hindlimbs and vent region. Females are distinctly larger in size (Table 4 , Appendix 3), lack nuptial pads and vocal sacs, having fewer spinules and glandular warts on dorsal surface of body and flank than males.

Dorsal coloration in preservative varies in males and females from brown to dark brown with darker markings. Two female specimens from Trat Province, Thailand (ZMKU AM 01444 and 01451) have dark orange markings on anterior part of dorsum. Markings or transverse spots on dorsum and dorsal surfaces of forelimbs and hindlimbs usually distinct, but faint in a few individuals. Coloration on flank usually creamy white, but pale brown, with dark brown marbling, in some individuals. Ventral coloration usually creamy white, but pale brown with indistinct dark mottling on chin and chest in some individuals. Hand usually creamy white, but light brown in some individuals. Most specimens have dermal fringe on fingers II and III (males N = 21, 70%; females N = 23, 69.7%), but some individuals lack this fringe (males N = 9, 30%; females N = 10, 30.3%). One specimen from Narathiwat Province, Thailand (THNHM 19720) has a vertebral stripe.

Distribution.

Based on a combination of morphological and genetic studies of F. cancrivora mangrove type ( Kurniawan et al. 2010, 2011, 2014) and Fejervarya Bangladesh mangrove type ( Islam et al. 2008), the reported distribution of F. cancrivora ( Chan-ard 2003; Chuaynkern and Chuaynkern 2012), F. moodiei ( Brown et al. 2013), and F. cf. cancrivora ( Harikrishnan and Vasudevan 2018), and specimens studied here, F. moodiei occurs in coastal areas from eastern India, the Andaman and Nicobar Islands, and southern China, southward through Vietnam, Thailand, Myanmar, Malaysia and Luzon Island in the Philippines. In Thailand, F. moodiei was documented in all coastal regions except the extreme southeastern Gulf of Thailand coast, where it is replaced by F. cancrivora ( Fig. 1 ).

Habitat, ecology, and natural history.

In Thailand, specimens were collected at night (1900-2200 h) in a variety of coastal habitats at elevations ranging from 0-16 m asl. Most specimens were observed in marshes near slow flowing ditches, ponds, or canals in mangrove forest ( Fig. 7C ). The species was also found in man-made environments such as agricultural fields adjacent to mangroves. In Kleang District, Rayong Province, most specimens were collected in and around brackish fish ponds and ditches in mangrove areas near the mouth of the Prasae River ( Fig. 7D ). Specimens from Pak Phanang District, Nakhon Si Thammarat Province were found around brackish shrimp ponds and ditches near the mouth of the Pak Phanang River. Frogs were observed sitting on the ground, under tree roots, or in or on the bank of water bodies. When disturbed, they usually escaped into holes in the ground or jumped into brackish water. No other anuran species were found in syntopy at this locality, although F. cancrivora was sampled at a site approximately 4.5 air-km, or 5.2 km following the river course, downriver (above).

Comparisons.

Twelve species of Fejervarya are known ( Frost 2019), with nine species occurring in East and Southeast Asia ( Sanchez et al. 2018). Four species of Fejervarya occur in Thailand, including F. limnocharis (Gravenhorst, 1829), F. multistriata (Hallowell, 1861), F. orissaensis (Dutta, 1997), and F. triora Stuart et al, 2006. Three additional Fejervarya species occur in adjacent countries, including F. iskandari Vieth et al. 2001, F. sakishimensis Matsui et al., 2007, and F. kawamurai Djong et al., 2011.

Fejervarya cancrivora and F. moodiei differ from all of these species by having the following combination of characters: (1) medium to large body size (vs. small to medium, SVL about 30-40 mm in males for F. iskandari , F. kawamurai , F. limnocharis , SVL about 40-55 mm in males for F. multistriata , F. orissaensis , F. triora [ Dutta 1997; Matsui et al. 2007; Chuaynkern et al. 2009; Djong et al. 2011]; (2) webbing formula: I1 –11/2II1–2III1–2IV2– 1V (vs. I0 –1II0–11/2III0–11/2IV11/2– 0V in F. vittigera , I1 –2II1–2III1–22/3IV22/3– 11/2V in F. limnocharis , I1 –2II1–2III1–22/3IV21/3– 1V in F. iskandari , I1 –2II1–2III11/2– 22/3 IV22/3-1V in F. multistriata , I1 –2II1–21/2III11/2–3IV3– 11/2V in F. sakishimensis , I1 –2II1–21/3III11/2–3IV3– 1V in F. kawamurai ); (3) having triangular or rectangular dark brown blotches covering vocal sacs on both sides of throat (vs. black “M” shape across throat in F. kawamurai , F. limnocharis , F. sakishimensis , F. triora , F. vittigera ); (4) having prepollax indistinct (vs. distinct in F. kawamurai , F. iskandari , F. limnocharis , F. sakishimensis , F. triora ), and (6) having palmar tubercles indistinct (vs. distinct F. kawamurai , F. limnocharis , F. sakishimensis , F. triora ).

Fejervarya moodiei differs from F. cancrivora by having: (1) SVL 42.7-62.7 mm in males, 50.0-81.8 mm in females (vs. 60.2-79.8 mm in males, 85.1-107.1 mm in females of F. cancrivora , Table 4 ; Appendix 2, 3); (2) indistinct, slightly raised inner tarsal ridge on tarsus (vs. distinct, strongly raised inner tarsal ridge on distal half or two-thirds of tarsus in F. cancrivora ) ( Fig. 5 A–D ); and (3) in body proportions (Table 4 ). In Thailand, F. moodiei appears to be closely associated with brackish water in or adjacent to mangrove forest, whereas F. cancrivora also occurs in freshwater wetlands.

Figures

Figure 1. Map of sampling localities of the Fejervarya cancrivora complex, including F. cancrivora neotype (yellow pentagon), F. cancrivora sensu stricto (yellow circles), F. moodiei holotype (blue diamond), F. moodiei (blue triangles), and F. cancrivora samples that were referred to F. raja (red circles) prior to this study. Open symbols indicate molecular data only, shaded symbols indicate morphological data only, and shaded symbols with center dots indicate both molecular and morphological data were studied.

Figure 5. Plantar and metatarsal views of A adult male neotype of Fejervarya cancrivora (FMNH 256688) B adult male F. cancrivora (ZMKU AM 01426) from Khuan Khanun District, Phatthalung Province, Thailand C adult female holotype of F. moodiei holotype (CM 3724), and D adult male F. moodiei (ZMKU AM 10390) from Mueang Phang-nga District, Phang-nga Province, Thailand. The inner metatarsal ridge on the tarsus of F. cancrivora is indicated with an arrow.

Figure 6. Adult male Fejervarya cancrivora (ZMKU AM 01426) from Khuan Khanun District, Phatthalung Province, Thailand (SVL = 66. 9 mm) immediately prior to preservation in A right lateral B dorsal C ventral D right palmar, and E right plantar views. Photographs by Attapol Rujirawan.

Figure 7. Exemplar habitats in Thailand of A Fejervarya cancrivora at a wetland in Khuan Khanun District, Patthalung Province B F. cancrivora at a brackish shrimp pond near Pak Phanang river, Pak Phanang District, Nakhon Si Thammarat Province C F. moodiei at mangrove forest in Thai Mueang Distrinct, Phang-nga Province, and D F. moodiei at brackish fish ponds near mangroves at the mouth of the Prasae River, Kleang District, Rayong Province. Photograph A by Attapol Rujirawan.

Figure 8. Adult female holotype of Fejervarya moodiei (CM 3724) in preservative in A dorsal and B ventral views. Photograph B by Carnegie Museum of Natural History.

Figure 9. Adult male Fejervarya moodiei (ZMKU AM 01390) from Mueang Phang-nga District, Phang-nga Province, Thailand (SVL = 60.6 mm) immediately prior to preservation in A right lateral B dorsal C ventral D right palmar, and E right plantar views. Photographs by Attapol Rujirawan.