A new subspecies of the Asiatic softshell turtle Amyda ornata (Gray, 1861) from its northern distribution range Author Praschag, Peter Turtle Island, Am Katzelbach 98, 8054 Graz, Austria ppraschag@turtle-island.at Author Gemel, Richard Natural History Museum Vienna, 1. Zoological Department, Herpetological Collection, Burgring 7, 1010 Vienna, Austria text Herpetozoa 2022 2022-05-25 35 81 93 http://dx.doi.org/10.3897/herpetozoa.35.e86055 journal article http://dx.doi.org/10.3897/herpetozoa.35.e86055 2682-955X-35-81 066DFFD046A44E588F982AF034E299E6 0789ED2026475723A7980113BCC27431 Amyda ornata jongli ssp. nov. Holotype. NHMW 40287 (Natural History Museum Vienna, herpetological collection): subadult specimen; skull and lower jaw; incomplete postcranial skeleton: parts of discus, shoulder girdle, pelvic girdle and extremities (Figs 2 - 5 ). Measurements: bony disc length 10 cm, bony disc width 13,6 cm. Plastron: Straight width 14,9 cm. Skull: Straight length 6,6 cm, straight width 3,5 cm. Figure 2. Holotype (NHMW 40287) of A. o. jongli ssp. nov. Bony elements of carapace (dorsal) and plastron (dorsal/visceral). Some neurals, 1st pleural right, left epiplastron and xiphiplastra missing. Photo: Natural History Museum Vienna, Alice Schumacher. Terra typica. Sangu River near Dohazari, Chittagong hill tracts, south-eastern Bangladesh, don. P. Praschag 3/2016. Paratypes. The paratypes consist of five living voucher specimens, one male and four females in the facility of "Turtle Island", NHMW 41463: 1-5 (Fig. 6a-j ). Location of origin for all five paratypes: Sangu River near Dohazari (terra typica), Chittagong hill tracts, south-eastern Bangladesh (same location as holotype-specimen). The specimens will be transferred post mortem to the herpetological collection of the Natural History Museum Vienna. For measurements see Table 1 . Figure 3. Holotype (NHMW 40287) of A. o. jongli ssp. nov. Lower jaw (dorsal with alveolar surface of mandible) and skull (dorsal). Photo: Natural History Museum Vienna, Alice Schumacher. Derivatio nominis. The subspecies name Amyda ornata jongli (noun in apposition) refers to the special term of local fishermen in Bangladesh who call this softshell turtle " Amyda ornata jongli " in Bengali, deriving from "jangala" in Sanskrit. It means wild and ferocious, as the animals are powerful and struggle when captured. Simultaneously it means also "... coming from rivers of jungle woodlands". Rahman et al. (2015) mention also the name "Tui-lip" , used by the Mro tribe. Suggested common names: English: "Northern Asiatic softshell turtle". German: "Noerdliche Knorpel-Weichschildkroete" . Taxonomic status. The Maximum Likelihood Tree for Amyda and related softshell turtles shows five distinct clades, divided into three species. Clades 4 ("Pale coloured, large nuchal and back tubercles" now herein described as A. o. jongli ssp. nov.), 5 (" Trionyx phayrei " / " Trionyx ephippium ") and 6 (" Trionyx ornatus ") show similar levels of genetic differentiation from each other, and constitute the three identified subspecies from Mainland Southeast Asia ( Fritz et al. 2014 ). Figure 4. Holotype (NHMW 40287) of A. o. jongli ssp. nov. Skull in ventral view. Photo: Natural History Museum Vienna, Alice Schumacher. Diagnosis on morphological base. The heads in subadults and adults are stout (see Figs 5 , 7a ), with a dome rising up steeply from the top of the proboscis to the top of the head, in the region of prefrontals and frontals (see Fig. 9 ). Compared to the two other subspecies of A. ornata , a broader distance between the eyes is seen in A. o. jongli (see Figs 3 , 7b ), although this relative distance varies depending on the ontogenetic stage of the specimen considered. Rather uniform pale and greyish colouration, carapace is olive greyish, bright yellow pigmentation missing (Fig. 6a-j ). Yellow-greyish dots, and vermiculation on head and neck is faded and not distinctly marked, ochre-coloured irises, ring-shaped skin around eyes with indistinct light dots, no pronounced "zebra striped" pattern; carapace with (or sometimes without, but less common) distinct black strokes and blotches or irregular saddle-shaped dark colouration on carapace (see Fig. 9 ). Black pigmentation mostly in the form of square spots on the rear edge of the carapace and two or three stripes running sideways from the center line of the carapace (compare Fig. 6a-j of the paratypes). Distinct tubercles in nuchal and back region. Nuchal tubercles protrude in a round or pointed shape along the front edge of the carapace in subadult and adult specimens (Fig. 8 ). Juveniles with tubercles on carapace running concentric, not longitudinal. Plastron white (in juveniles) changing to yellowish-grey during growth, mottled with clouds of black pigments especially on the parts of underlying bony skeleton. Figure 5. Holotype (NHMW 40287) of A. o. jongli ssp. nov. Skull in lateral view. Note the convex structure of frontal, prefrontal and maxillary. Photo: Natural History Museum Vienna, Alice Schumacher. Figure 6. Paratypes, NHMW 41463:1 ( a, b ); NHMW 41463:2 ( c, d ); NHMW 41463:3 ( e, f ); NHMW 41463:4 ( g, h ); NHMW 41463:5 ( i, j ), living specimens of A. o. jongli ssp. nov. in the holdings of "Turtle Island", dorsal and ventral view. Photos: Peter Praschag. Diagnostic comparisons. The genus Amyda is distinguished from all other trionychids by the following suite of characteristics: Carapace with black markings, sometimes black and yellow or bright dots, but no ocelli; skull: snout longer than the diameter of the orbit (more in juveniles than in adults); alveolar surface of the mandible with a longitudinal ridge at the symphysis; surface of jaws light; first and second neural fused; epiplastra in contact or narrowly separated, front part of epiplastra are elongated (see Fig. 7c ); further detailed skull morphology is given by Van Dijk (1992) . Figure 7. (a - c) Computer Tomography of A. o. jongli ssp. nov, NHMW 41463:1. Photos: Thomas Hildebrandt and G. Fritsch, Department of Reproduction Management, Leibnitz Institute for Zoo and Wildlife Research. A. ornata differs from A. cartilaginea (sensu stricto) and Amyda sp. (= candidate species A, sensu Fritz et al. 2014 ) by the stouter head versus narrow head tapering lengthwise with a longer proboscis. Amyda o. ornata differs from the other two subspecies by its distinct yellow pigments on carapace, head and neck; the presence of a "crown" : two or three converging black lines forming an arrow on the upper side of the head; tubercles predominantly in the nuchal and front region of the carapace; saddle-shaped dark colouration on carapace uncommon. Distribution: Vietnam, Laos, Cambodia (Chaco Phraya, Irrawaddy and Salween River systems). Amyda o. phayrei differs from the other two subspecies by its dull brown carapace, yellow pigments present forming distinct yellow dots on the head and neck; carapace with or without distinct saddle-shaped dark colouration; with eyes open, the eyelids have distinct yellow spots alternating with grey "zebra striped" colouration; tubercles in the nuchal and back region of the carapace but not as distinct as in A. o. jongli ssp. nov.; head in upper side view more slender and narrowed to the proboscis. Distribution: Thailand, Myanmar (delta region of the Irrawaddy (Ayeyarwady) and possibly neighbouring Salween (Thanlwin) river, NE India (Mizoram) and Yunnan ( Kuchling 1995 ). Distribution data of Amyda ornata jongli ssp. nov. (Fig. 1 , Table 2 ). Bangladesh : Chittagong hill tracts: Sangu River system [=Sangpo river, Sankho river, Shankha River] and tributaries, Matamuhuri river; Karnaphuli river system. Sangu River: Dohazari (Terra typica) (Fig. 1 , pt 1); four Mro villages along upstream of Sangu River in Sangu-Matamuhuri Reserve Forest (villages Nr. 23, 24, 25 and 26 in Fig. 1 , Rahman et al. 2015 ) (Fig. 1 , pt 2); Thanci Upazila [=Thanchi]: "rocky hill stream Remakri Khal" in Bandarban and Mro village "Nepew Para" ( Khan (2013 "2012" ) ), (Fig. 1 , pt 3); village on the Matamuhuri River (village Nr. 1 in Fig. 1 , Rahman et al. 2015 ) (Fig. 1 , pt 4); Sangu River at Remarki, border to Myanmar Fatik Chhari (pers. comm. Scott Trageser, Comitee Creative Conservation Alliance) (Fig. 1 , pt 6); Durung Khal river, mouth of the Halda river (pers. comm. Sureshchandra Das) (Fig. 1 , pt 7). Table 2. Waypoints of known localities of A. o. jongli ssp. nov.

Point (pt)	Locality	Coordinates (WGS 84) (Latitude, Longitude)
1	Dohazari (Terra typica), Bangladesh	22.163958 , 92.065016
2	Sangu River - CHT, Sangu-Matamuhuri Reserve Forest, Bangladesh ( Rahman et al. 2015 )	21.3455827 , 92.6069887
3	Thanchi upazila, Bandarban, Bangladesh ( Khan (2013 "2012" ) )	21.8217887 , 92.4377149
4	Matamuhri river, Bangladesh (village Nr. 1 in Fig. 1 , Rahman et al. 2015 )	21.8830881 , 92.2497277
5	Ngengpui River, Ngengpui wildlife sanctuary, Lawngtlai, Mizoram, India (central point selected for map)	22.43134 , 92.7867
6	Sangu River at Remarki, Fatik Chhari, border to Myanmar, Bangladesh	21.3030315 , 92.6511433
7	Durung Khal River, mouth of the Halda river, Bangladesh	22.7612597 , 91.755951
8	Tuirial River drainage, Aizawl District, India ( Hmar et al. 2020 )	23.555 , 92.779
9	Narichera Stream, tributary of Khowai River, North District, Tripura, India ( Das et al. 2016 )	23.7804 , 91.8430
10	Dhalchera, Phaisen Hills, Cachar district, India ( Nath et al. 2018 )	24.264611 , 92.415256
11	Guwahati, Assam - Fish market, India ( Anonymous 2020 )	26.16945 , 91.76448

India : Mizoram: Ngengpui River, Ngenpui Wildlife Sanctuary, Lawngtlai (Pawar and Choudhuri 2000): bony disc deposited in the holdings of the Museum of Bombay Natural History Society under BNHM 1446 as confirmed by Rahul Khot, Curator of National History Collection department of the Bombay Natural History Society (pers. comm. 7th, April 2016) (Fig. 1 , pt 5); Tuirial River drainage, Aizawl District ( Hmar et al. 2020 ) (Fig. 1 , pt 8) (additionally occurring in most of the major river systems of the northern part and documented here from the rivers Teirei, Tut, Tlawng, Tuirini, Tuirial, Tuivawl & Tuipuibari, pers. comm., 6th March 2016 Tlau Vanlal Hrima, not considered in Fig. 1 ); Tripura : Narichera Stream, North District ( Das et al. 2016 ) (Fig. 1 , pt 9); Assam : Dhalchera river, Phaisen Hills, Cachar District ( Nath et al. 2018 ) (Fig. 1 , pt 10), fish market, Guwahati ( Anonymous 2020 ) (Fig. 1 , pt 11). The area registered in the distribution records comprises two main river systems separated by a watershed running through the centre part of Mizoram: the southern region in southern Mizoram and Chittagong hill tracts containing Sangu River as a part of the "Kolodyne Basin" and draining in the Bay of Bengal. The northern part is situated in northern Mizoram Assam and Tripura and is dominated by the Barak River system ("Barak Basin") draining into the Brahmaputra; in the recent past a specimen was even reported from the fish market in Guwahati, Assam (Fig. 1 , pt 11).