Hipposideros kingstonae Wongwaiyut & Karapan & Saekong & Francis & Guillén-Servent & Senawi & Khan & Bates & Jantarit & Soisook 2023, sp. nov.

Hipposideros kingstonae sp. nov. Figs. 3–6, 9; Tables 1–2 [= H. cineraceus- B (Kingston et al., 2006; Murray et al., 2012; 2018)] Holotype. PSUZC-MM2014.164 (field number PS140903.2), adult male, body in alcohol, skull and baculum extracted, collected on 3 September 2014, by Sunate Karapan, Puchit Saekong and Pipat Soisook. Full measurements (in mm) of the holotype are as follows; FA: 38.4, HB: 40.2, E: 14.2, Tail: 25.9, TIBIA: 16.7, HF: 6.81, GTL: 16.35, SL: 16.13, CBL: 14.54, CCL:13.94, ZB: 8.20, BB: 7.66, MW: 8.50, PC: 2.75, C–M3: 5.54, C1–C1: 3.25, M3–M3: 5.83, c–m3: 5.88, ML: 9.62, ALSW: 4.18, AMSW: 2.19. Type locality. Phru To Daeng Peat Swamp Forest (= Sirindhorn Peat Swamp Forest), Princess Sirindhorn Wildlife Sanctuary, Narathiwat Province, Thailand, 6°4’ N, 101°58’ E, 170 m a.sl. The specimen was collected in a harp trap in combination with a mist net set along a boardwalk in the swamp forest. Paratypes. Thailand — PSUZC-MM2022.2 (field number PS211113.5), adult male ♁, body in alcohol, skull and baculum extracted, collected on 13 November 2021 by Sunate Karapan, Phutita Wongwaiyut and Pipat Soisook from the same area as the holotype but in a different spot, near the edge of the forest (6°4.3’ N, 101°57.8’ E). It was caught in an 18 m-long mist net along a boardwalk by the edge of the swamp. PSUZC-MM2022.1 (field number BL160219.1), adult female, body in alcohol, skull extracted, collected by Puchit Saekong and Sunate Karapan from the same locality as the holotype on 19 February 2016. Referred specimens. Thailand — PSUZC-MM2014.165 (field number PS140830.2), adult female, body in alcohol, skull extracted, collected from Border Police Base, Bang Lang Dam, Hala-Bala Wildlife Sanctuary, Yala Province, 6°4’ N, 101°17’ E, 22 m a.s.l., on 30 August 2014, by Sunate Karapan, Puchit Saekong and Pipat Soisook. Malaysia — Uncatalogued specimen field number TK020622.10, adult female, body in alcohol, skull extracted, collected from Lubuk Baung, Krau Wildlife Reserve, Pahang, peninsular Malaysia, on 22 June 2002, by Juliana Senawi. This specimen was caught in a four-bank harp trap set across a forest trail in lowland dipterocarp forest. DWNP-M- 1996-07-29 -05653 (field number TK960729.1), adult male, body in alcohol, skull extracted, collected from Kuala Lompat, Krau Wildlife Reserve, Pahang, peninsular Malaysia, on 29 July 1996 by Tigga Kingston. This specimen was caught in a four-bank harp trap in a lowland dipterocarp forest trail. DWNP-M- 1996-05-12 - 05654 (field number TK960519.1), adult male, body in alcohol, skull and baculum extracted, collected from Kuala Lompat, Krau Wildlife Reserve, Pahang, peninsular Malaysia on 19 May 1996 by Tigga Kingston. This specimen was caught in a four-bank harp trap in a lowland dipterocarp forest trail. SMF83823 (field number CMF920706- 01), immature male, collected from Krau Wildlife Reserve, Pahang, peninsular Malaysia on 5 July 1992 by Charles M. Francis. EBD 23565 (field number 960523n05), adult female, EBD 23561 (field number 960523n06) and EBD 23560 (field number 969523n33), adult males, previously identified as H. cf. cineraceus, caught together with 6 other individuals (3 female, 3 male) in a four-bank harp trap set in a lowland forest trail about ~150 m from the entrance of Madai Caves, Sabah by Antonio Guillén-Servent and Charles M. Francis. EBD 23821 (field number 960604n01), adult female, previously identified as H. cf. cineraceus, caught in a mist-net set in the understory of the lower montane forest near the park headquarters in Gunung Kinabalu National Park, Sabah by Antonio Guillén-Servent. Diagnosis. This is a small Hipposideros with a FA of 35.3–42.6 mm and GTL of 15.94–17.90 mm. The sides of the anterior part of the noseleaf are slightly concave; the anterior border has a deep V-shaped median notch and is somewhat angular in appearance. The internarial septum is large, rounded, and distinctly swollen from the middle to the top. The lateral leaflet is absent. The dorsal pelage is dark brown, with the individual hairs creamy-white from the base to the mid-part. The ventral pelage is orange-brown; the hairs are dark brown at the tip, and paler at base. The baculum is very small, 0.5 mm in length, with short but distinct distal prongs. The constant frequency (CF) element of the echolocation call is 141.0–144.0 kHz in the Thai-Malay Peninsula individuals and 132.3-141.4 kHz in the Bornean individuals. Etymology. The species is named in honour of Tigga Kingston, who as the chair and founder of the Southeast Asian Bat Conservation Research Unit (SEABCRU), spearheads the global bat research community in understanding diversity and promoting bat conservation. Description. Hipposideros kingstonae sp. nov. is a small hipposiderid with a forearm length of 35.3–42.6 mm (Table 1). The body mass is 4.9–7.0 g (n=4), with three male specimens at 4.9–5.6 g, and one female at 7.0 g. The ear is rounded with a pointed tip, and a height of 14.0– 18.7 mm; it has short brown hairs along the inner sides. The tail is relatively short (21.5–30.0 mm) in comparison to the head and body length (40.2–49.0 mm). The hindfoot is shorter than half of the tibia in length, 5.0– 7.1 mm versus 14.7–17.9 mm, respectively. The noseleaf is without a lateral supplementary leaflet (Fig. 3). The anterior leaf is dark brown, slightly concave on both lateral borders. The anterior border (bottom edges) has an angular appearance and is separated by a deep groove in the middle. The internarial septum is large, rounded, and distinctly swollen from the middle to the top. The intermediate leaf is relatively broad and sparsely haired. The male specimens have a well-defined frontal sac behind the posterior leaf. The dorsal pelage is dark brown (Fig. 3), with the base to the middle of the hairs creamy white. The ventral pelage is orange brown to dark brown at the hair tips, and paler at the bases. The baculum is very short, 0.5 mm in length (n=3) (Fig. 4a). The shaft is narrow and straight in dorsal and ventral view, with a rounded base and bifid tip. In lateral view, it is slightly curved from the middle towards the base. The skull is elongate, with a mean greatest length of the skull (GTL) of 16.29 mm (15.94–17.90 mm), a skull length (SL) of 16.13 mm (15.73–17.79 mm) and a condylocanine length (CCL) of 14.00 mm (13.56–15.94 mm) (Table 2). The mastoid width (MW) is 8.32 mm (8.07–8.50 mm). This slightly exceeds the zygomatic breadth (ZB), which is 8.01 mm (7.79–8.26 mm). In lateral view, the nasal swelling is well-developed, and the sagittal crest is clearly defined particularly on the anterior part of the braincase (Fig. 5a). The anterior median swellings are rounded, with the AMSW 3.07 mm (1.99–4.28 mm) in width. The frontal depression is shallow when viewed from either the side or the top (Fig. 5a). The postorbital constriction (PC) is 2.62 mm (2.40–2.91 mm). The zygomata are narrow with an angular process projecting upwards in the mid-part of the jugal bone (Fig. 5a). The upper canine (C1) is large, about twice the height of the second upper premolar (P4). The crown area of the P4 is about twothirds that of the C1 (Fig. 6a). The first upper premolar is very small and fully extruded, so that the C1 and P4 are in contact. The upper toothrow length (C–M3) is 5.41 mm (5.12–5.81 mm). The lower toothrow length (c–m3) is 5.67 mm (5.05–6.28 mm) and the mandible length (ML) is 9.64 mm (9.22–10.92 mm) (Table 2). The lower canine (c1) has an elongated postero-basal heel (Fig. 6a). The c1 is twice the height of the second lower premolar (p4). The p4 is rounded and subequal to that of the c 1 in crown area. The first lower premolar (p2) is short and only about half the height of (p4) (Fig. 5a). Echolocation. The echolocation call of H. kingstonae sp. nov. is a typical CF-FM signal. In three individuals recorded from Thailand, the frequency of maximum energy (FmaxE), corresponding to the frequency of the constant element (CF), is 142.4 kHz (141.0–143.4 kHz; n=3). In peninsular Malaysia, the FmaxE from Krau Wildlife Reserve is 144.0 kHz (n=1), whereas in Malaysian Borneo, the FmaxE of the bats from the Madai caves was 138.4 kHz (136.6-141.4 kHz; n=5 individuals) for males and 137.7 kHz (137.4–138.2 kHz; n=4 individuals) for females, while the female from Gunung Kinabalu had a FmaxE of 132.3 kHz. Genetics. The phylogenetic trees based on mitochondrial COI reveal that the new species clusters with H. kunzi and H. bicolor (Fig. 7), with a genetic distance of 2.3% and 4.9%, respectively (see Appendix 3). Although clustered as a subclade of the new species, the Sabah population is only 0.3% different from the specimens from the Thai-Malay Peninsula (Fig. 7; Appendix 3), and no significant difference in morphology were observed. The new species is not closely related to any of other forms currently referred to H. cineraceus. The tree topology based on ND2 and CytB is similar (Fig. 8), as previously noted by Murray et al. (2012; 2108). The ND2 and partial CytB sequences of 4 specimens from the island of Palawan deposited in the FMNH as H. ater (Esselstyn et al., 2012) belong in the clade of the new species, as sister to the Sabah specimens with less than 1% genetic distance from them. The genetic distances between the new species and H. kunzi and H. bicolor, are 8.0% and 7.3% based on CytB, and 7.3% and 10.5%, respectively, based on ND2. In addition, these two species are clearly different from the new species in terms of morphology (see comparison section below). Morphologically, the new species is similar to H. einnaythu (see comparison section below), which is also present in the region (Douangboubpha et al., 2011; Douangboubpha, 2019) but has a genetic distance of 9.6% and 10.4% based on COI and ND2, respectively. In addition, this study provides, for the first time, genetic data (as well as echolocation—see below) of H. einnaythu from Tanintharyi, Myanmar and Thailand. Besides being closely clustered with H. kunzi, H. bi

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