Uperodon mormorata ( Rao, 1937 )

Garg, Sonali, Senevirathne, Gayani, Wijayathilaka, Nayana, Phuge, Samadhan, Deuti, Kaushik, Manamendra-Arachchi, Kelum, Meegaskumbura, Madhava & Biju, Sd, 2018, An integrative taxonomic review of the South Asian microhylid genus Uperodon, Zootaxa 4384 (1), pp. 1-88 : 29-32

Uperodon mormorata ( Rao, 1937)

Mottled Globular Frog

( Figs. 1 , 2A, 2C , 3F , 5F , 7M–S , 8E–H , 9A–D ; Tables 1–4)

Original name and description. Ramanella mormorata Rao, 1937 . On some new forms of Batrachia from S. India. Proceedings of the Indian Academy of Sciences. Section B, 6: 387–427 . Neotype. By present designation, ZSI/ WGRC /V/A/958, an adult female, SVL 32.4 mm, from Sakleshpur, Karnataka state, India, collected by SD Biju on 26 June 2003. Type locality. “ Saklespur (= Sakleshpur ), Hassan District , Mysore ”, India. Current status of specific name. Valid name, as Uperodon mormorata ( Rao, 1937) .

Neotypification of Ramanella mormorata Rao, 1937 . This species was described from “Saklespur” based on a single specimen “total length 25.00 mm”, which was deposited in the Central College, Bangalore. The original name bearing type (or types) of this species are considered lost ( Dubois 1984; Das & Whitaker 1997; Biju 2001), and this nominal taxon was also presumed to be “lost” until its rediscovery from “Canacona (15°01’N 74°04’E), adjacent to Cotigao Wildlife Sanctuary (Goa, south-Western India)” based on an available museum specimen from “Malabar” at MCZ, USA ( Das & Whitaker 1997). As per the original description, Rao had several available specimens including “young”, “immature”, “mature males” and “mature females”, however, the description was based on a single specimen for which absolute measurements were included. Das & Whitaker (1997) provided a detailed description of this species based on examined specimens, however, without designating a neotype. Our new collection from the type locality Sakleshpur is comparable with the original description of Ramanella mormorata Rao, 1937 , in overall general morphology, body size and dorsal and ventral markings. Since the original name-bearing type of this species is lost, we consider designation of a neotype necessary to define this nominal taxon objectively for clarification of its taxonomic status and to establish nomenclatural stability. Hence, in accordance with Article 75 of The Code, and based on diagnostic characters found useful for differentiating Ramanella mormorata Rao, 1937 (= Uperodon mormorata ) from its congeners (as stated in the ‘comparison’ section below), we herein formally designate ZSI/WGRC/V/A/958 (an adult female, SVL 32.4 mm) from the original type locality Sakleshpur as the neotype of Ramanella mormorata Rao, 1937 . The neotype description provided subsequently also shows that the neotype is largely consistent with what is known of the former namebearing type.

Comparison. For comparison of Uperodon mormorata with U. globulosus , U. systoma , U. taprobanicus , U. anamalaiensis and U. montanus see ‘comparison’ section of those species. Uperodon mormorata differs from U. nagaoi , U. rohani sp. nov. and U. variegatus by its densely mottled ventral surface (vs. inconspicuous spots and speckles present or absent in U. nagaoi ; and complete absence of spots, blotches or mottling in U. rohani sp. nov. and U. variegatus ) and relatively larger snout-vent size, male SVL 31–40 mm, N = 9, female SVL 32–43 mm, N = 3 (vs. relatively smaller, U. nagaoi : male SVL 25–32 mm, N = 8; U. rohani sp. nov.: male SVL 26–34 mm, N = 16, female SVL 28–35 mm, N = 8; U. variegatus : male SVL 25–33 mm, N = 12, female SVL 29–36 mm, N = 7). This species could be confused with U. anamalaiensis , U. obscurus , U. palmatus and U. triangularis due to its mottled ventral skin. However, U. mormorata differs from all these species by its relatively larger snout-vent size, male SVL 31–40 mm, N = 9, female SVL 32–43 mm, N = 3 (vs. smaller, U. obscurus : male SVL 22–31 mm, N = 17, female SVL 30–38 mm, N = 5; U. palmatus : male SVL 27–31 mm, N = 4, female SVL 34–36 mm, N = 2; U. triangularis : male SVL 25–37 mm, N = 20, female SVL 33–42 mm, N = 4), its ventral surface (including throat and limbs) being densely marbled (vs. scattered spots or blotches, largely on chest and belly), and shank relatively shorter than thigh, male: SHL/TL ratio 0.8–1.1, N = 9, female: SHL/TL ratio 0.8–0.9, N = 3 (vs. nearly equal, U. obscurus : male: SHL/TL ratio 0.9–1.0, N = 17, female: SHL/TL ratio 0.9–1.1, N = 5; U. palmatus : male: SHL/TL ratio 1.0, N = 4, female: SHL/TL ratio 1.0, N = 2; U. triangularis : male: SHL/TL ratio 0.9–1.1, N = 20, female: SHL/TL ratio 0.9–1.0, N = 4).

Genetic divergence. For 16S mitochondrial gene sequences, the sampled populations of Uperodon mormorata showed an average intraspecific distance of 0.6% (range 0–1.2%, N = 11). Populations from Maharashtra and Karnataka were observed to be divergent by up to 1.2%. Genetically, U. mormorata is closely related to U. triangularis , from which it was found to differ by an average uncorrected genetic distance of 2.9% (range 2.4– 3.6%, N = 99). For interspecific genetic distances with all other members of the genus, see Table 3.

Description of neotype (measurements in mm) ( Figs. 3F , 7M–P ). Small-sized (SVL 32.4), relatively robust adult female; head small (HW 9.6, HL 8.8, IFE 4.4, IBE 8.6), less than one-third (27.2%) of body length, wider than long (HW/HL ratio 1.1); snout truncate in dorsal and ventral view, vertical in lateral view, its length (SL 3.1) shorter than horizontal diameter of eye (EL 2.6); loreal region acute with rounded canthus rostralis; interorbital space nearly twice as wide (IUE 3.0) as upper eyelid width (UEW 1.6); nostril closer to tip of snout (NS 0.6) than to eye (EN 1.5); supratympanic fold distinct, extending from posterior corner of upper eyelid to insertion of forelimb at axilla; eye diameter (EL 2.6); vomerine odontophores present on the palate, prominent, without teeth; weakly-developed neopalatinal ridges on posterior side of each choana, appear fused with the vomerine odontophores; tongue moderately large, emarginate. Forelimbs moderately long and thin; forearm (FAL 6.5) shorter than hand length (HAL 9.1); finger length formula I<II<IV<III; tips of all fingers with truncate discs, without grooves, moderately wide compared to finger width (FD I 1.5, FW I 0.3; FD II 1.5, FW II 0.4; FD III 1.8, FW III 0.5; FD IV 1.5, FW IV 0.4); subarticular tubercles prominent, oval, all present; two well-developed palmar tubercles (inner, oval, 1.6 mm long; outer, bilobed, 1.9 mm long). Hind limbs relatively long and thin, thigh length (TL 13.5) longer than shank (SHL 11.1) and foot (FOL 12.9); tips of all toes with small truncate to rounded discs, discs rather wide compared to toe width (TD I 0.7, TW I 0.4; TD II 0.8, TW II 0.6; TD III 1.0, TW III 0.7; TD IV 1.0, TW IV 0.6; TD V 0.8, TW V 0.5); foot webbing present: I2 – – 2II 2 – 3III 2 – 3IV 3 – 2V; well-developed dermal fringe present on all toes; subarticular tubercles prominent, oval; two smooth metatarsal tubercles, oval, outer slightly longer than the inner.

Skin of snout, between eyes, sides of head, anterior part of the dorsum, posterior parts of back, and upper and lower parts of flank, granular; dorsal surfaces of forelimb, thigh, tibia and tarsus, shagreened to sparsely granular ( Figs. 3F , 7M ); ventral surfaces shagreened.

Colouration. In preservation: Dorsum olive grey with scattered dark brown spots, dark brown inverted ‘V’ at the level of shoulder (directed downwards), a dark brown line between eyes, another two lines: one on snout, and another between inverted ‘V’ and line connecting the eyes; dorsal surfaces of forelimb, thigh and shank (including fingers and toes), olive grey with dark brown cross bands; groin and anterior parts of thigh, light grey with light brown mottling, and a light brown patch near cloacal region; throat, chest, belly, and ventral surfaces of forelimbs and hind limbs, light brown with prominent light grey reticulations ( Fig. 3F ). Colour in life (SDBDU 2016.3404): Dorsum dark olive grey with scattered dark brown spots especially on posterior part of dorsum, thigh and shank, faint dark brown inverted ‘V’ at the level of shoulder (directed downwards); dorsal surfaces of forelimb, thigh and shank (including fingers and toes), olive grey with dark brown cross bands; groin and anterior parts of thigh, light brown with light grey and white mottling; throat, chest, belly, and ventral surfaces of forelimb and hind limb, dark brown with prominent light greyish-blue reticulations ( Figs. 9A–C ).

Variations. Morphometric measurements for 12 specimens, including the holotype, are given in Table 4 . Overall, the skin colouration and texture (except markings), are nearly similar in all the examined specimens.

However, webbing between toes is observed to be highly variable, without being consistent even between the same sexes. Foot webbing: SDBDU 2002.567 (female): medium, I2 – – 2II 2 – 3III 2+ – 3IV 3 – 2V; SDBDU 2012.43 (male): large, I1 – 2II 1 – 1III 1 – 2IV 2 – 1V; SDBDU 2015.3082 (male): medium, I1 – 2II 1 – 2III 2 – 3IV 3 – 2V ( Figs. 7Q–S ). SDBDU 2015.3085: dorsum with more prominent blackish-brown spots.

Note. In the original description, Rao (1937) stated this species to have “No occipital fold”. However, Das & Whitaker (1997) discussed its presence based on their observation of occipital folds in both their examined samples of Uperodon mormorata (MCZ A. 116283 and MCZ A. 15421). Variable occurrence of occipital fold was earlier also reported by Parker (1934). We closely examined the presence and absence of occipital fold in members of the genus Uperodon , and observe that its presence may be an artifact of preservation caused due to stretching of loose skin of the snout towards the head.

Secondary sexual characters. Male (SDBDU 2016.3404): Vocal sac externally visible on the lower jaw; female (ZSI/WGRC/V/A/958) ova white, pigmented on pole (diameter 0.6–0.8 mm, N = 20).

Vocalization. Male of Uperodon mormorata ( SDBDU 2016.3404 ) from Goa produced a single type of call with pulsatile temporal structure. The calls had uniform intervals and were not delivered in groups. A typical male call had a duration of 182.6 ms, rise time of 108.3 ms, and fall time of 57.4 ms. The call comprised of 28 pulses that were delivered at a rate of 154.7 pulses/s. The spectrum was characterized by single broad peak with mean dominant frequency of 2.6 kHz ( Figs. 8E–H ).

Geographical distribution and habitat. In the present study, the presence of Uperodon mormorata is confirmed in the Western Ghats states of Karnataka, Maharashtra, Goa and Gujarat ( Fig. 2C ; Tables 1). For additional previous records and references thereto see supplementary Table S1. Report of this species from Assam ( Dey 2001) is doubtful. We collected this species predominantly from terrestrial habitats such as rock crevices, fallen logs and under stones, and occasionally from tree barks about two metres above the ground. It was found in and around secondary forest areas (Amba, Amboli, Bygoor, Dandeli, Londa, Netravali, Sakleshpur, Samot and Castle rock), primary forests (Shimoga), as well as marshes (coastal Goa), at elevations from sea level up to 1050 m asl.

Figures

FIGURE 1. Maximum Likelihood phylogram based on 511 bp of mitochondrial 16S rRNA gene sequences from 73 individuals representing all known members of the genus Uperodon. Three samples representing taxon under synonymy are shown inquotationmarks.Outgroup not shown for graphical purposes.BayesianPosterior Probabilities and bootstrap values of>50 are shown above and below the branches, respectively. Values towards the species labels represent mean uncorrected genetic distances between the species with range values in parentheses.

FIGURE 2. A. Geographic distribution of the genus Uperodon in Bangladesh, Bhutan, India, Nepal, Pakistan and Sri Lanka; B. Distributionof Uperodon globulosus, U. systomaand U.taprobanicus; C. Distribution of four endemic Uperodon species of the Western Ghats (excluding the widely distributed Indian member U. variegatus, shown in Fig.12A); D. Distribution of four endemic Uperdodon species of Sri Lanka. Details of localities reported in the present study are provided in Table 1. Records from literature are listed in supplementary Table S1 and indicated here based onapproximate coordinates.

FIGURE 3. Preserved type specimens of Uperodon species in dorsal and ventral view. A. Uperodon systoma, Syntype, ZMB 3551 (female); B. Uperodon globulosus, Syntype, NHM 62.12.29.4 (ex. BMNH 1947.2.10.97) (female); C. Kaloula pulchra taprobanica, Holotype, NHM 1905.3.25.138 (ex. BMNH 1947.2.10.62) (female); D. Ramanella anamalaiensis, Neotype, ZSI/ WGRC/V/A/956 (male); E. Hylaedactylus montanus, Holotype, NHM 72.4.17.212 (ex. BMNH 1947.2.11.19) (female); F. Ramanella mormorata, Neotype, ZSI/WGRC/V/A/958, (female); G. Ramanella nagaoi, Holotype, WHT 2826 (male); H. Callula obscura, Holotype, NHM 58.11.28.42 (ex. BMNH 1947.2.11.20) (female); I. Ramanella palmata, Holotype, NHM 90.11.8.62 (ex.BMNH 1947.2.11.21); J. Callula triangularis, Lectotype, NHM 74.4.29.897 (ex.BMNH 1947.2.11.15) (male); K.Hylaedactylus carnaticus, Lectotype, ZSI 2761(female); L. Uperodonrohani sp. nov., Holotype, DZ1449 (male).(Images: SD Biju and Sonali Garg).

FIGURE 5. Ventral surface of the palate in Uperodon species showing vomerine odontophores separated or fused with dermal projectionsand/or neopalatinalridgeson posterior sides ofthe choanae (indicated witharrows).A. Uperodon systoma, SDBDU 2005.2787 (female) and SDBDU 2005.2806 (male); B. Uperodon globulosus, SDBDU 2014.2673 (female) and SDBDU 2005.4762 (female); C. Uperodon taprobanicus, SDBDU 2007.6364 (female) and SDBDU 2005.4732A (male); D. Uperodon anamalaiensis, SDBDU 2013.2302 (female) and SDBDU 2002.3050 (male); E. Uperodon montanus, SDBDU 2002.24B (female) and SDBDU 2003.40115 (male); F. Uperodon mormorata, SDBDU 2011.507 (male) and SDBDU 2012.43 (male); G. Uperodon obscurus, DZ 1366 (male); H. Uperodon palmatus, DZ 1331 (male); I. Uperodon triangularis, SDBDU 2007.6400 (female) and ZSI/WGRC/V/A/959 (male); J. Uperodon variegatus, SDBDU 2005.4726 (female) and SDBDU 2005.4727 (male); K.Uperodon rohani sp. nov., DZ 1521 (male).(Images:SD Biju).

FIGURE 8. Male advertisement calls of Uperodon anamalaiensis, U. mormorata and U. triangularis. A–D. Uperodon anamalaiensis: A. ten second call segment; B. one second call segment; C. 0.1 second segment showing a single non-pulsatile call; D.spectrogram of the call shown in Figure8B; E–H. Uperodon mormorata:E.ten second call segment; F. one second call segment; G. 0.1 second segment showing a single non-pulsatile call; H. spectrogram of the call shown in Figure 8G; I–L. Uperodon triangularis: I. ten second call segment; J. one second call segment; K. 0.1 second segment showing a single nonpulsatile call; L. spectrogram of the call shown in Figure 8J.

FIGURE7. Head, hand, foot and foot webbing in Uperodon anamalaiensis, U. montanus, U. mormorata and U. nagaoi. A–E. Uperodon anamalaiensis: A–D. Neotype of Uperodon anamalaiensis (ZSI/WGRC/V/A/956, male):A. lateral view of head; B. ventral view of hand; C. ventral view of foot; D. schematic illustration of foot webbing; E. schematic illustration of foot webbing in referred specimen of U. anamalaiensis (SDBDU 2013.928, female); F–L. Uperodon montanus: F–H. Topotype of Uperodon montanus (SDBDU 2011.849, male): F. lateral view of head; G. ventral view of hand; H. ventral view of foot; I. schematic illustration of foot webbing; J–L. schematic illustration of foot webbing in referred specimens of Uperodon montanus: J. SDBDU 2015.3001, male; K. SDBDU 2003.40115, female; L. SDBDU 2005.30, female; M–S. Uperodon mormorata: M–P. Neotype of Uperodon mormorata (ZSI/WGRC/V/A/958, female): M. lateral view of head; N. ventral view of hand; O. ventral view of foot; P. schematic illustration of foot webbing; Q–S. schematic illustration of foot webbing in referred specimens of Uperodon mormorata: Q. SDBDU 2002.567, female; R. SDBDU2012.43, male; S. SDBDU 2015.3082, male.T–W. Holotype of Uperodon nagaoi (WHT 2826, male):T. lateral view ofhead;U. ventral view of hand;V. ventral view of foot; W. schematic illustration of foot webbing. (Illustrations:SD Biju).

FIGURE9. Uperodon mormorata, U. nagaoi and U.obscurus inlife.A–C. Uperodonmormorata (SDBDU 2016.3404,male): A. Dorsolateral view; B. dorsal view; C. ventral view; D. Uperodon mormorata (SDBDU 2011.835, male), dorsolateral view; E–G. Uperodon nagaoi (DZ 1367, male): E. Dorsolateral view; F. dorsal view; G. frontal view; H–K. Uperodon obscurus: H. Dorsolateral view (DZ 1366, male); I. Dorsal view (DZ 1366, male); J. dorsolateral view (DZ 1408, male); K. dorsolateral view (DZ 1369, male); L. ventral view (DZ 1369, male). (Images: SD Biju and E–G Madhava Meegaskumbura).