Abavorana luctuosa (Peters, 1871)

Quah, Evan S. H., Grismer, L. Lee, Wood Jr., Perry L., Lim, Kelvin K. P., Imbun, Paul Y. & Anuar, M. S. Shahrul, 2021, An investigation into the taxonomy of Abavorana luctuosa (Peters, 1871) (Anura, Ranidae) and the resurrection of Rana decorata Mocquard, 1890 from Borneo, Vertebrate Zoology 71, pp. 75-99 : 75

Abavorana luctuosa (Peters, 1871)

Common Mahogany Frog Figs 3 , 4 , 7E-H , 8A-F , 9A-B

Limnodytes luctuosus : Peters 1871:579, 1872:43 & pl. 6; Capocaccia 1957:208 & 215.

Rana luctuosa (in part): Boulenger 1882:68, 1891:341, 1912:238, 1920:183 & 184; Flower 1896:904, 1899:896 & 897; Hanitsch 1900:73; Butler 1904:199 & 200; Van Kampen 1923:196 & 197; Smith 1930:94 & 103, 1931:16; Capocaccia 1957: 215; Taylor & Elbel 1958:1040; Taylor 1962:451-453; Inger 1954:250, 1966:206-208, 1978:312, 1985:iii, 5, 38, 40, 56, 57, 76; Grandison 1972:66 & 67; Berry 1975:74 & 75; Dring 1979:200; Inger & Stuebing 1992:42, 2005:19, 27, 33, 149 & 150; Inger & Tan 1996:563; Inger et al. 1996:363; Manthey & Grossmann 1997:111 & 112; Khonsue & Thirakhupt 2001:73; Malkmus et al. 2002: 163-165; Das 2006a:5; Jaafar et al. 2008:39.

Rana (Limnodytes) luctuosa : Mocquard 1890:122.

Rana (Hylorana) luctuosa (in part): Boulenger 1920:126.

Rana (Hylarana) luctuosa (in part): Van Kampen 1923:196 & 197; Bourret 1942:360; Dubois, 1987: 42.

Hylorana luctuosa (in part): Deckert 1938:144.

Rana (Pulchrana) luctuosa (in part): Dubois 1992:326.

Pulchrana luctuosa (in part): Frost et al. 2006:369; Fei et al. 2010:33.

Hylarana luctuosa (in part): Manthey 1983:22; Che et al. 2007: 1-13 (by implication); Das et al. 2007:159, 160, 167, 170, fig. 7d; Imbun 2014:99.

Abavorana luctuosa : Oliver et al. 2015:186 (in part); Quah et al. 2017: 272-288 (in part); Zainudin et al. 2017:876-891 (in part); Chan et al. 2019:1057, 1062.

Hylarana (Abavorana) luctuosa (in part): Inger et al. 2017:147 & 148.

Holotype.

Adult female ( MSNG 29344) collected by Giacomo Doria and Odoardo Beccari from “Sarawak”, (Borneo) Malaysia ( Capocaccia 1957) (Fig. 3 ).

Diagnosis.

Body robust, medium-sized; head moderate; snout short, rounded, canthus rostralis smoothly rounded; interorbital space broader than the upper eyelid; tympanum distinct, not quite two-thirds the size of the eye with no pale colouration on the margins of the tympanum; no vocal sacs in males; vomerine teeth in two small oblique patches on a level with the posterior edge of the choanae; length of 1st finger greater than 2nd finger; disc width to finger width ratios of finger 3 and toe 4 is 1-1.5; dorsolateral fold indistinct or absent; the humeral gland in males is prominent, raised and centrally positioned on the ventral surface of the upper arm; a weak or absent rictal ridge; outer metatarsal tubercle weak or absent; subarticular tubercles moderate; skin of dorsum smooth or finely shagreened; throat, abdomen, and flanks smooth; posterior section of venter and back of the thigh rugose; dorsum reddish-orange to chocolate-brown, encircled by a white or cream coloured dorsolateral line that encircles the snout, canthus rostralis, outer edge of the upper eyelids, and dorsum along the dorsolateral fold to the vent; lower flanks dark-brown or black below the dorsolateral line grading into a paler venter; dorsal colouration of the limbs same as the flanks with whitish or light-grey speckles or stripes. Abavorana luctuosa can be easily differentiated from its congeners on the basis of its ventral colour pattern which is usually immaculate or with only very faint, sparse, light speckling (Figs 3 , 4 , 9A-B ). Adult males with SVL 35.05-44.27 mm, adult females with SVL 37.05-51.06 mm; adult males with SW 7.11-8.62 mm, adult females with SW 7.65-9.38 mm; adult males with IOD 3.25-5.14 mm, adult females with IOD 3.80-5.64 mm; adult males with TD 2.50-3.83 mm, adult females with TD 2.95-4.24 mm; adult males with FL 17.81-22.04 mm, adult females with FL 19.13-26.37 mm (Table 5 ).

Distribution.

Its distribution spans southern Peninsular Thailand, Peninsular Malaysia, Borneo, and Sumatra (Manthey & Grossmann 1997; Oliver et al. 2015; Quah et al. 2017). Within Borneo it is confirmed from Sabah state: Imbak Canyon and Sipitang; Sarawak state: Bukit Kana, Samarakan and Tubau in Bintulu Division and Kubah National Park where it is sympatric with Abavorana decorata (Fig. 10 ). The species is expected to range across Borneo.

Figures

Figure 3. Holotype of Abavorana luctuosa; A: Original illustration of the holotype of Abavorana luctuosa from Peters (1872). B: Dorsum of the holotype of A. luctuosa (MSNG 29344). C: Venter of the holotype of A. luctuosa (MSNG 29344). Photographs by Giuliano Doria.

Figure 4. Bornean Abavorana luctuosa; Dorsal (A & C) and ventral (B & D) pattern of Abavorana luctuosa from Borneo. A & B: Top row (from left to right); FMNH 234972, FMNH 234973, FMNH 234975 and FMNH 234977 (all specimens from Sipitang District, Sabah). Bottom row (from left to right); FMNH 234980 (Sipitang District, Sabah), FMNH 269142, FMNH 273216 and FMNH 273217 (specimens from Bintulu Division, Sarawak). C & D (from left to right): ZRC 1.11938, ZRC 1.11943 (both from Kubah, Sarawak) and ZRC 1.2270 (unspecified location in Borneo).

Figure 7. Living specimens of Abavorana decorata and Bornean A. luctuosa; A: Abavorana decorata from Tawau Hills, Sabah, East Malaysia. B: A. decorata from Maliau Basin, Sabah, East Malaysia. C: A. decorata from Kubah National Park, Sarawak, East Malaysia. D: A. decorata from Melawi Regency, West Kalimantan Province, Indonesia. E & F: A. luctuosa from Kubah National Park, Sarawak, East Malaysia. G: A. luctuosa (FMNH 238720) from Sipitang, Sabah, East Malaysia. H: A. luctuosa (FMNH 242883) from Sipitang, Sabah, East Malaysia. (Photographs by Alexander Haas [A, C & E], Paul Bertner [B], Muhammad Aini [D], Chien C. Lee [F], and Robert F. Inger [G & H], Figs G & H © Field Museum of Natural History. Created by Field Museum of Natural History, Amphibian and Reptile Collection and licensed under CC-BY-SA 4.0).

Figure 8. Peninsular Malaysian, Thai and Sumatran Abavorana luctuosa, and A. nazgul; A: A. luctuosa from Sungai Durian Forest Reserve, Kelantan, Peninsular Malaysia. B: A. luctuosa from Janda Baik, Pahang, Peninsular Malaysia. C: A. luctuosa from Semenyih, Selangor, Peninsular Malaysia. D: A. luctuosa from Fraser's Hill, Pahang, Peninsular Malaysia. E: A. luctuosa from Hala-Bala Wildlife Sanctuary, Thailand. F: A. luctuosa from Sumatra, Indonesia. G: A. nazgul (USMHC 2572) from Gunung Jerai, Kedah, Peninsular Malaysia. H. A. cf. nazgul from Thale Ban National Park, Satun Province, Thailand. (Photographs by Zaharil Dzulkafly [A], Steven Wong [B], Kurt H. P. Guek [C], Evan Quah [D & G], Ian Dugdale [E], Hellen Kurniati [F] & Bunyarit Dechochai [H]).

Figure 9. Colour pattern comparison among species of the genus Abavorana; Dorsum (A, C & E) and venter (B, D & F) in life of A. luctuosa (USMHC 2576) (A & B) from Penang Hill, Penang Island, Malaysia, A. decorata from Maliau Basin, Sabah, East Malaysia, Borneo (C & D) and A. nazgul (USMHC 2233) (E & F) from Gunung Jerai, Kedah, West Malaysia. (Photographs by Evan Quah [A, B, E & F] and Stefan T. Hertwig [C & D]).

Figure 10. Distribution map of Abavorana decorata and A. luctuosa in Borneo; Known distribution of Abavorana decorata (red) and A. luctuosa (blue) in Borneo. Star = type locality.