Rhinolophus nippon Temminck, 1835

Saikia, Uttam, Chakravarty, Rohit, Csorba, Gabor, Laskar, Mostaque Ahmed & Ruedi, Manuel, 2025, Taxonomic reassessment of bats from the Western Himalayas, India and description of a new species of the Myotis frater complex (Mammalia, Chiroptera, Vespertilionidae), Zootaxa 5644 (1), pp. 1-78 : 16-18

4. Rhinolophus nippon Temminck, 1835

(Japanese greater horseshoe bat)

New material: 1 F, 31.05.2017, Barog Tunnel, Solan District , Himachal Pradesh, V /M/ERS/ 403; 1 M , 02.06.2017, Mount Karol, Solan District , Himachal Pradesh, V /M/ERS/402 .

Morphological description of specimens: Four individuals from Himachal Pradesh were examined of which two were old museum specimens. They had an average forearm length of 60.5 mm. One of the females had the largest forearm length of 62.5 mm and longest ear at 27 mm. Dorsally, the pelage was light golden brown while the belly was slightly lighter in colour. The dorsal hairs had light brown tips and dark brown roots. Ears were pointed at the tip with 8–9 ridges on the inner side. The horseshoe width was 7.2–7.3 mm, the median emargination of the horseshoe was broad. The superior connecting process of sella was rounded off when viewed laterally ( Fig. 8A ). The lancet was pointed at the tip and sides are concave. The lower lip had three mental grooves, the middle one extended forward. On average, the 3 rd metacarpal was 9.2% shorter than the 4 th. The length of first phalanges of the 3 rd metacarpal was 52.8 % of the length of the metacarpal.

The skull was robust (zygomatic width considerably exceeding mastoid width) with an average greatest length of 24.77 mm. A well–developed sagittal crest was present which bifurcated anteriorly forming a shallow nasal pit and extended posteriorly till the lambda, albeit weakly. Prominent lamboid crests were present laterally. The palate was highly emarginated, its anterior margin extending till the middle of the first molar and the posterior border resting approximately at the level of the line drawn between M 2 and M 3 ( Fig. 9B ).

DNA: We obtained 651 bp of the COI gene and 1140 bp of the CYTB from one individual captured in Mt Karol (M 2215/ V /M/ERS/402), which represents the first DNA evidence from northern Indian specimens of the Rh. ferrumequinum species complex. None of the CYTB sequences available in the GeneBank matched the Himachal Pradesh sample, the most similar sequence (at 5% K2P distance) being that of a sample from Yunnan (GB ON640696 ) and labelled as “ Rh. ferrumequinum ” ( Table S2). The COI gene also gave similar results, the closest match (at 3.4% K2P distance) being Asian samples from southern China (e.g., from Zhejiang, GB OR 467324), or a sample of Rh. nippon from Japan (GB KT779432 ) at 3.5% divergence. Western Palaearctic and some Central Asian sequences of genuine Rh. ferrumequinum s.s. were all more divergent,>5% for the COI (e.g., GB OQ 706678), or>6.5% for the CYTB (e.g., GB OQ 885419) (see Tables S2 and S 3). Thus, both molecular markers suggest that the Himachal Pradesh sample is a unique and rather distinctive lineage. However, its phylogenetic position with respect to the two major mitochondrial clades assigned respectively to ferrumequinum and nippon by Koh et al. (2014) places it as sister to the latter with strong bootstrap support ( Figs 3 and 5). Consistent with our results, Uvizl et al. (2024) also pointed that the smaller Central Asian Rh. bocharicus was also part of this species complex but distinct from other species.

Locality records and ecological notes: Himachal Pradesh: Barog Tunnel (1560 m), Lutru Cave near Arki (1550 m), Mount Karol (1890 m), Solan Town (1500 m) in Solan District ( Saikia et al. 2011; present study); Chakmoh (c. 760 m), Hamirpur District ( Ghosh 2008); Chamba (c. 1000 m), Chamba District (Chakraborty 1977); Kullu Valley, Manali (1950 m) Kullu District ( Allen 1908; Lindsay 1927); Mandi (c. 1050 m), Mandi District ( Ghosh 2008); Ghannati (c. 1640m), Shimla (2100 m), Tottu (c. 1900 m) in Shimla District ( Dodsworth 1913; Bates & Harrison 1997; Ghosh 2008). Uttarakhand: Katarmal (1380 m) and Almora (c. 1600 m) in Almora district and Mussorie (2000 m) in Dehradun district ( Bhat 1974; Bates & Harrison 1997). A female specimen was caught at about 300 m inside Barog tunnel (1530 m asl). They were seen roosting in two small groups with females carrying babies. We could record two other rhinolophids namely Rh. lepidus and Rh. perniger inside the tunnel although in lesser numbers. Another specimen was caught in a mist net at Mount Karol (1850 m) inside oak forest. A cave located nearby this forest patch held a breeding population of Rh. sinicus although no Rh. nippon could be observed there.

Taxonomic note: Traditionnally, Rh. ferrumequinum was considered as a single, polytypic species spanning the entire Palearctic region, from Europe to Japan, and parts of northern Oriental Region (e.g. Bates & Harrison 1997; Csorba et al. 2003). Bates & Harrison (1997) showed that northern India was inhabited by two distinct forms, a smaller one found in Kashmir (proximus Andersen, 1905), and a larger one ( tragatus Hodgson, 1835 ) found in Himachal Pradesh to Nepal, Arunachal Pradesh and southern China ( Csorba et al. 2003). Recent revisions based on molecular and morphological characters ( Koh et al. 2014; Uvizl et al. 2024) showed that Rh ferrumequinum is a species complex, represented in Asia by the nominal species in the western half of the continent, by the smaller Rh. bocharicus in Central Asia and by Rh. nippon further east to the Japanese Archipelago. The exact geographic extent of each species though was unclear owing to inadequate sampling from the Himalayan region. We show here that the taxon tragatus from Himachal Pradesh is genetically closely related, but morphologically slightly distinct from Rh. nippon , and thus should be considered as a distinctive subspecies of the latter (i.e., Rh. nippon tragatus ). Under this concept, Himachal Pradesh would represent the westernmost limit of distribution of Rh. nippon , while further west Rh. ferrumequinum s.s. occur. According to this suggested taxonomic arrangement, the smaller form from Kashmir (proximus) might represent the easternmost representative of Rh. ferrumequinum s.s., pending new genetic analyses to address its extact phylogenetic relationships. However, as some species of Rhinolophus may show instances of mitonchdrial introgression, blurring phylogenetic relationships ( Dool et al. 2016; Uvizl et al. 2024), taxonomic conclusions should ideally be based on several independent DNA markers, which is currently lacking for Himalayan specimens.

Figures

FIGURE 3: Neighbor-joining tree reconstruction of examined rhinolophids and hipposiderids COI sequences and based on the K2P matrix in Table S2. Branches are proportional to the number of mutational steps. Sequence names are composed of the first two letters of the genus, followed by the species name, the GenBank number (or field number for newly sequenced samples), and the region of origin. Samples from India are highlighted in bold face.

FIGURE 8. Portrait showing frontal and side views of the noseleaf structure of A) Rh. nippon (ZSI–NERC V/M/ERS/403) and B) Rh. sinicus (ZSI–NERC V/M/ERS/404) from the Western Himalayas.

FIGURE 9. Skull features of a Rh. nippon from Himachal Pradesh (ZSI–NERC V/M/ERS/402).A) Lateral view of the rostrum, B) occlusal view of palate and dentition, and C) occlusal view of the posterior portion of skull showing relative position of the glenoid cavity and tympanic bullae.