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Research Article
Four new tube-nosed bat species of the genus Murina (Chiroptera, Vespertilionidae) from Xizang Autonomous Region, China, based on morphological and molecular data
expand article infoTao Luo§|, Ming-Le Mao§, Chang-Ting Lan§, Zi-Fa Zhao§, Zhong-Lian Wang§, Jing Yu§, Jia-Jia Wang§, Chen-Rui Yan§, Ning Xiao, Jiang Zhou§
‡ Yunnan University, Kunming, China
§ Guizhou Normal University, Guiyang, China
| Southwest United Graduate School, Kunming, China
¶ Guiyang Healthcare Vocational University, Guiyang, China
Open Access

Abstract

The genus Murina Gray, 1842, recently had four new species discovered in China over the last four years, suggesting its diversity may have been previously underestimated. Herein, we describe four new species—Murina beibengensis sp. nov., Murina medogensis sp. nov., Murina milinensis sp. nov., and Murina yadongensis sp. nov.—based on morphological and genetic evidence from specimens collected during bat diversity surveys conducted in the Xizang Autonomous Region of China over the past three years. Each of these four new species forms an independent lineage on a phylogenetic tree reconstructed using the mitochondrial COI and Cyt b genes, and each is genetically distinct from its congeners. Morphologically, the new species can be distinguished from the 43 recognized congeners by features including forearm length, hair color, and skull morphology. We elevated M. huttoni rubella from a subspecies of M. huttoni to a species based on morphological and genetic evidence. The new species discussed herein increase the number of species in the genus Murina to four worldwide and from 23 to 28 in China. This study not only enriches our understanding of bat species diversity but also underscores the importance of conducting bat surveys in the specialized highland habitats of the Himalayas.

Key Words

Diversity, Himalaya, morphology, Murina, taxonomy

Introduction

The family Vespertilionidae has an extremely high species diversity with over 500 species distributed worldwide (Moratelli et al. 2019). Within this family, species of the subfamily Murininae share a single prominent characteristic, namely, tube-shaped nostrils. There are three recognized genera in this subfamily: Murina Gray, 1842; Harpiocephalus Gray, 1842; and Harpiola Thomas, 1915 (Kuo et al. 2006; Son et al. 2015; Moratelli et al. 2019; Mou et al. 2024). For a long time, although these genera have distinguishable morphological characteristics, their phylogenetic relationships have been controversial (Tate 1941; Bhattacharyya 2002; Wilson and Reeder 2005; Kuo et al. 2006). For example, phylogenetic analyses based on the mitochondrial cytochrome c oxidase subunit I (COI) gene revealed that Harpiocephalus and Harpiola are phylogenetically embedded within Murina (Ruedi et al. 2012; Soisook 2013; Son et al. 2015; Tu et al. 2015; Soisook et al. 2017; Mou et al. 2024; Mou et al. 2025). However, recent phylogenomic studies have provided robust support for the taxonomic validity of these genera, recovering Murina as the sister clade to Harpiocephalus and Harpiola, with the phylogenetic conflicts among genera being attributable to ancient introgression events (Wang et al. 2024b).

Asian tube-nosed bats, genus Murina Gray, 1842, are small bats with a forearm length of less than 44 mm. These bats are largely distributed in the forests of East, South, and Southeast Asia, as well as in parts of North Asia (Moratelli et al. 2019) (Fig. 1). Based on the relative size of the crown area of the canines and first and second upper premolars, as well as the position of the incisors, the genus Murina has been divided into two groups (Corbet and Hill 1992). In the suilla group, the first upper incisor (I2) is situated anterior to the second (I3), I2 is clearly visible in the lateral view, and the crown area of the first upper premolar (P2) is half or less that of the second (P4); meanwhile, in the cyclotis group, I3 is adjacent to I2, such that I2 is essentially obscured by I3 when viewed laterally, and the crown area of P2 is 2/3 or more that of P3 (Csorba and Bates 2005). However, these two morphogroups do not represent monophyletic groups and were therefore later revised as “suilla-type” and “cyclotis-type” (Son et al. 2015; Soisook et al. 2017; Yu et al. 2020; Mou et al. 2024; Wang et al. 2024a; Mou et al. 2025). Currently, 43 species of the genus Murina have been recognized (Dobson 1872; Peters 1872; Scully 1881; Thomas 1891; Sowerby 1922; Hill 1964; Hill and Francis 1984; Maeda and Matsumura 1998; Csorba and Bates 2005; Kruskop and Eger 2008; Furey et al. 2009; Kuo et al. 2009; Csorba et al. 2011; Eger and Lim 2011; Francis and Eger 2012; Soisook et al. 2013a; Soisook et al. 2013b; He et al. 2015; Son et al. 2015; Chen et al. 2017; Soisook et al. 2017; Zeng et al. 2018; Moratelli et al. 2019; Yu et al. 2020; Mou et al. 2024; Mou et al. 2025) (Table 1), with 24 species occurring in China (Mou et al. 2025; Wei et al. 2025), several of which have been discovered in the last 10 years, i.e., M. fanjingshanensis He, Xiao & Zhou, 2015 (He et al. 2015), M. rongjiangensis Chen, Liu, Deng, Xiao & Zhou, 2017 (Chen et al. 2017), M. liboensis Zeng, Chen, Deng, Xiao & Zhou, 2018 (Zeng et al. 2018), M. jinchui Yu, Csorba & Wu, 2020 (Yu et al. 2020), M. yuanyang Mou, Qian, Li, Li, Luo & Li, 2024 (Mou et al. 2024), M. yushuensis Han, Csorba & Wu, 2024 (Wang et al. 2024a), and M. lvchun Mou & Li, 2025 (Mou et al. 2025). These findings show that the species diversity of the genus may have been seriously underestimated.

Table 1.

Species catalogue, dentition types, molecular markers, and references for the genus Murina (modified from Moratelli et al. 2019).

ID Species Dentition Molecular data Literature obtained
1 Murina aenea Hill, 1964 Cyclotis-type Present Hill 1964; Moratelli et al. 2019
2 Murina annamitica Francis & Eger, 2012 Cyclotis-type Present Francis and Eger 2012
3 Murina cyclotis Dobson, 1872 Cyclotis-type Present Dobson 1872; Moratelli et al. 2019
4 Murina fionae Francis & Eger, 2012 Cyclotis-type Present Francis and Eger 2012
5 Murina guilleni Soisoo, Karapan, Satasook, Thong, Khan, Maryanto, Csorba, Furey, Aul & Bates, 2013 Cyclotis-type Present Soisook et al. 2013a
6 Murina harrisoni Csorba & Bates, 2005 Cyclotis-type Present Csorba and Bates 2005
7 Murina huttoni (Peters, 1872) Cyclotis-type Present Peters 1872; Moratelli et al. 2019
8 Murina lvchun Mou & Song Li, 2025 Cyclotis-type Present Mou et al. 2025
9 Murina peninsularis Hill, 1964 Cyclotis-type Present Hill 1964; Moratelli et al. 2019
10 Murina pluvialis Ruedi, Biswas & Csorba, 2012 Cyclotis-type Present Ruedi et al. 2012
11 Murina puta Kishida, 1924 Cyclotis-type Present Moratelli et al. 2019
12 Murina recondita Kuo, Fang, Csorba & Lee, 2009 Cyclotis-type Present Kuo et al. 2009
13 Murina rozendaali Hill & Francis, 1984 Cyclotis-type Present Hill and Francis 1984; Moratelli et al. 2019
14 Murina bicolor Kuo, Fang, Csorba & Lee, 2009 Suilla-type Present Kuo et al. 2009
15 Murina aurata Milne-Edwards, 1872 Suilla-type Present Moratelli et al. 2019
16 Murina balaensis Soisook, Karapan, Satasook & Bates, 2013 Suilla-type Present Soisook et al. 2013b
17 Murina beelzebub Son, Furey, Csorba & 2011 Suilla-type Absent Csorba et al. 2011
18 Murina chrysochaetes Eger & Lim, 2011 Suilla-type Present Eger and Lim 2011
19 Murina eleryi Furey, Thong, Bates, Csorba & Son, 2009 Suilla-type Present Furey et al. 2009
20 Murina fanjingshanensis He, Xiao & Zhou, 2015 Suilla-type Present He et al. 2015
21 Murina feae (Thomas, 1891) Suilla-type Present Thomas 1891; Moratelli et al. 2019
22 Murina florium Thomas, 1908 ? Present Moratelli et al. 2019
23 Murina fusca Sowerby, 1922 Suilla-type Absent Sowerby 1922
24 Murina gracilis Kuo, Fang, Csorba & Lee, 2009 Suilla-type Present Kuo et al. 2009
25 Murina harpioloides Kruskop & Eger, 2008 Suilla-type Present Kruskop and Eger 2008
26 Murina hilgendorfi Peters, 1880 Suilla-type Present Moratelli et al. 2019
27 Murina hkakaboraziensis Soisook, Thaw, Kyaw, Oo, Pimsai, Suarez-Rubio & Renner, 2017 Suilla-type Present Soisook et al. 2017
28 Murina jaintiana Ruedi, Biswas & Csorba, 2012 Suilla-type Present Ruedi et al. 2012
29 Murina jinchui Yu, Csorba & Wu, 2020 Suilla-type Present Yu et al. 2020
30 Murina kontumensis Son, Csorba, Tu & Motokawa, 2015 Suilla-type Present Son et al. 2015
31 Murina leucogaster Milne-Edwards, 1872 Suilla-type Present Moratelli et al. 2019
32 Murina liboensis Zeng, Chen, Deng, Xiao & Zhou, 2018 Suilla-type Present Zeng et al. 2018
33 Murina lorelieae Eger & Lim, 2011 Suilla-type Present Eger and Lim 2011
34 Murina rongjiangensis Chen, Liu, Deng, Xiao & Zhou, 2017 Suilla-type Present Chen et al. 2017
35 Murina ryukyuana Maeda & Matsumura, 1998 Suilla-type Absent Maeda and Matsumura 1998
36 Murina shuipuensis Eger & Lim, 2011 Suilla-type Present Eger and Lim 2011
37 Murina suilla Temminck, 1840 Suilla-type Present Moratelli et al. 2019
38 Murina tenebrosa Yoshiyuki, 1970 Suilla-type Absent Moratelli et al. 2019
39 Murina tubinaris (Scully, 1881) Suilla-type Present Scully 1881; Moratelli et al. 2019
40 Murina ussuriensis Ognev, 1913 Suilla-type Present Moratelli et al. 2019
41 Murina walstoni Furey, Csorba & Son, 2011 Suilla-type Present Csorba et al. 2011
42 Murina yuanyang Mou, Qian, Li, Li, Luo & Li, 2024 Suilla-type Present Mou et al. 2024
43 Murina yushuensis Han, Csorba & Wu, 2024 Suilla-type Present Wang et al. 2024a
Figure 1. 

Geographical distribution of recognized species and the four new species of the genus Murina in Asia. The base maps are from the Standard Map Service website (http://bzdt.ch.mnr.gov.cn/index.html; Map Approval No. GS (2020) 4619).

Several new bat specimens were collected during biodiversity surveys conducted between July 2023 and August 2024 in China’s Xizang Autonomous Region and Gansu Province. These specimens were identified as belonging to the genus Murina based on the following diagnostic characteristics: tube-shaped nose, forearm length less than 44 mm (vs. usually greater than 44 mm in Harpiocephalus), third upper molars (M3) are reduced, with parastyle, paracone, and protocone (vs. highly reduced in Harpiocephalus, only parastyle), heights of the inner (I2) and outer (I3) upper incisors each measuring half the height of the upper canines (C1) (vs. 2/3 in Harpiola), and the height and crown area of the first premolar (P2) are significantly smaller than those of the lower canine (C1) (vs. similar in Harpiola) (Kuo et al. 2006; Mou et al. 2024). Subsequent detailed morphological comparisons and molecular analyses indicated that these specimens could be distinguished from currently recognized species. Based on these differences, we formally describe the specimens as four new species of the genus Murina: Murina beibengensis sp. nov., Murina medogensis sp. nov., Murina milinensis sp. nov., and Murina yadongensis sp. nov.

Materials and methods

Taxon sampling and morphological analyses

A total of 55 specimens from the genus Murina were collected between July 2016 and August 2024 in a bat survey that included areas in China’s Xizang, Xinjiang, Qinghai, Gansu, Guizhou, Sichuan, and Shaanxi Provinces. The experimental animals used in this study were treated in accordance with Chinese animal welfare laws (GB/T 35892–2018). All specimens used for morphological studies were fixed in 95% ethanol and stored at the Animal Ecology Laboratory of Guizhou Normal University in the city of Guiyang, Guizhou Province, China. All molecular samples were stored in a refrigerator kept at –80 °C.

Morphometric data were collected from 25 well-preserved specimens of the genus Murina (Suppl. material 2). A total of 25 measurements were recorded to the nearest 0.1 mm using digital calipers following the protocol of Wang et al. (2024a). For external measurements of the body, including HB = head-body length (from tip of snout to anus); TL = tail length (from anus to tip of tail); EL = ear length (from lower edge of external auditory meatus to tip of pinna); EW = ear width (maximum width of the external auditory meatus); TRL = tragus length (from base to tip of tragus); TRW = tragus width (maximum width of the tragus); HFL = hind foot length (from extremity of heel to tip of longest toe, not including claws); FL = forearm length (from elbow to carpus with wings folded); TIB = tibia length (from knee joint to ankle). Measurements of the cranial and dental from extracted and cleaned skull including: GTL = greatest length of skull (from anterior aspect of I2 to most prominent point of occipital region); CCL = condylocanine length (from exoccipital condyle to most anterior part of C1); BCW = braincase width (greatest width of braincase); BCH = braincase height (from horizontal plane to highest point of cranium); ZYW = zygomatic width (greatest width of skull across zygomatic arches); MAW = mastoid width (greatest distance across mastoid region); IOW = interorbital width (least width of interorbital constriction); CM3L = upper canine-molar length (from anterior of C1 to posterior of M3 crown); C1C1W = upper canine width (greatest width of outer borders of C1); M3M3W = upper molar greatest width (greatest width of outer borders of M3); CM3L = lower canine-molar length (from anterior of C1 to posterior of third lower molar (M3) crown); ML = mandible length (from anterior rim of alveolus of I2 to most posterior part of condyle); MDL = greatest length of mandible (from anterior point of first lower incisor (I2) to most posterior part of condyle); CPH = coronoid process height (least distance from apex of coronoid process to indentation of lower border of ramus mandibula).

Principal component analysis (PCA) was employed to investigate the relative contributions of specific variables to morphometric variation. Subsequently, cluster analysis was performed to group species based on the extracted principal components. For closely related species for which complete data were not available from the previously published literature, mean values were used for the subsequent analyses. All statistical analyses were conducted using SPSS v21.0 (SPSS, Inc., Chicago, IL, USA), and differences were considered statistically significant at a p-value of < 0.05.

DNA extraction, sequencing, and phylogenetic reconstruction

Genomic DNA was extracted from muscle tissue using a DNA extraction kit purchased from Tiangen Biotech (Beijing) Co., Ltd. In this study, we amplified and sequenced the mitochondrial COI and cytochrome b (Cyt b) genes from each DNA sample using the primers COI-F (5’-CCTACTCRGCCATTTTACCTATG-3’), COI-R (5’-ATCTCTGGGTGTCCAAAGAATCA-3’), Cytb-F (5’-ATGATATGAAAAACCATCGTTG-3’), and Cytb-R (5’-TTTCCNTTTCTGGTTTACAAGAC-3’). PCR amplification was performed with a 25-μl reaction volume under the following cycling conditions: an initial denaturing step at 95 °C for 5 min; 10 cycles of denaturing at 94 °C for 60 s, annealing at 46 °C for 30 s, and extension at 72 °C for 1 min; 25 cycles of denaturing at 94 °C for 60 s, annealing at 50 °C for 40 s, and extension at 72 °C for 1 min; 35 cycles of denaturing at 94 °C for 60 s, annealing at 54 °C for 40 s, and extension at 72 °C for 1 min; and a final extension at 72 °C for 10 min. The purified PCR products were sequenced using both forward and reverse primers with a BigDye Terminator Cycle Sequencing Kit according to the manufacturer’s instructions. Sequencing was performed on an ABI Prism 3730 automated DNA sequencer manufactured by Chengdu TSING KE Biological Technology Co., Ltd. (Chengdu, China). All of the relevant sequences have been deposited in GenBank (Suppl. material 1: tables S2, S3).

Mitochondrial sequences were aligned in MEGA v7.0 (Kumar et al. 2016) using the MUSCLE algorithm (Edgar 2004) with default parameters. Phylogenetic trees were constructed using both maximum likelihood (ML) and Bayesian inference (BI) methods. The ML analysis was performed using IQ-tree v2.0.4 (Nguyen et al. 2015a) based on the best-fit model with 2000 ultrafast bootstrap (UFB) replicates (Hoang et al. 2018) and continued until a correlation coefficient of at least 0.99 was achieved. The BI analysis was performed using MrBayes v3.2.1 (Ronquist et al. 2012). Each BI analysis was run independently using four Markov Chain Monte Carlo chains (with three heated chains and one cold chain) starting with a random tree; each chain was then run for 6 million generations, and sampling was conducted every 1000 generations. Data run convergence was confirmed when the average standard deviation of split frequencies was less than 0.01. Nodes in the trees were considered well-supported when the Bayesian posterior probabilities were ≥ 0.95 (Ronquist et al. 2012) and the ML UFB value was ≥ 95% (Hoang et al. 2018). The best-fit model for the BI and ML analyses was obtained based on the Bayesian information criterion computed with PartitionFinder v2.1.1 (Lanfear et al. 2017).

In the phylogenetic analysis, to elucidate the phylogenetic position of the new species within the genus Murina and the lineage relationships of the M. huttoni complex, we constructed three datasets: two COI datasets (Murina_1 and Murina_2) and one Cyt b dataset (Murina_3). Murina_1 comprises 92 sequences (657 bp), with 288 variable sites and 264 parsimony-informative sites. The evolutionary models for the first, second, and third codon positions were TIM+I+G, HKY+I, and TRN+I+G, respectively. Murina_2 includes 43 sequences (657 bp), with 188 variable sites and 142 parsimony-informative sites. The evolutionary models for the first, second, and third codon positions were TRNEF+G, F81+I, and TRN, respectively. Murina_3 consists of 46 sequences (1140 bp), with 506 variable sites and 448 parsimony-informative sites. The evolutionary models for the first, second, and third codon positions were K80+I+G, HKY+I+G, and TRN+I+G, respectively.

Species delimitation and genetic distance

To rapidly assess the boundaries of species currently recognized by taxonomists, we used a tree-based phylogenetic approach known as the multi-rate Poisson Tree Processes (mPTP) (Kapli et al. 2017). This method is comparatively more stable and widely used than other tree-based methods (e.g., GMYC and bPTP) (Blair and Bryson 2017). We used the ML tree generated by the COI sequences as input and performed the analysis on the mPTP web server (http://mptp.h.its.org) using the default parameters. Additionally, genetic distances were calculated based on COI and Cyt b sequences using the uncorrected p-distance model with 1,000 bootstrap replications in MEGA v7.0 (Kumar et al. 2016).

Results

Phylogeny and genetic divergence

Reconstructing the mitochondrial tree topology of Vespertilionidae based on the mitochondrial COI gene using both methods showed discordance (Fig. 2A and Suppl. material 1: fig. S1). The BI tree included monophyletic clades of the subfamilies Myotinae, Kerivoulinae, and Murininae, except for Vespertilioninae (Fig. 2A and Suppl. material 1: fig. S1), although the groups were not highly resolved. The monophyly of three genera of the subfamily Murininae, Harpiocephalus, Harpiola, and Murina, was not supported (Fig. 2A and Suppl. material 1: fig. S1), but could be further divided into four clades, Clades I–IV. Clade I consisted of only the single species M. jinchui; Clade II comprised M. fanjingshanensis, M. rongjiangensis, M. shuipuensis, M. leucogaster, and M. hilgendorfi; Clade III contained the genus Harpiola, M. harrisoni, M. ussuriensis, and M. huttoni; and Clade IV comprised the remaining species, including four subclades, IV1–4, that were not highly supported (Fig. 2A and Suppl. material 1: fig. S1). However, phylogenetic analysis of mitochondrial Cyt b revealed Harpiocephalus, Harpiola, and Murina as independent monophyletic clades (Fig. 2B and Suppl. material 1: fig. S2).

Figure 2. 

Phylogeny and species delimitation. A. BI tree and mPTP species delimitation based on mitochondrial COI; B. BI tree and mPTP species delimitation based on mitochondrial Cyt b; C. Bayesian tree and mPTP species delimitation in the M. huttoni complex. Scale bars represent 0.1/0.04 nucleotide substitutions per site. Taxa marked with “merger” indicate proposed merging schemes based on mPTP species delimitation results, while unmarked taxa are suggested as valid species.

Within subclade IV-1, a single specimen from Yadong County, Shigatse City, Xizang Autonomous Region, and three samples from Miling County, Nyingchi City, Xizang Autonomous Region, China, clustered together to become the sister clade of (M. yushuensis + (M. harpioloides + (M. yuanyang + M. chrysochaetes))) (Fig. 2A and Suppl. material 1: fig. S1). These two populations can be distinguished from the known species of the genus Murina and from the lineages undescribed in this study by morphological characteristics (see subsequent comparisons) and molecular data, with p-distances of 12.2% (vs. M. harpioloides) and 10.8% (vs. M. yushuensis) (Suppl. material 3). These values are greater than the genetic distances between recognized species, e.g., 4.8% (M. gracilis vs. M. recondita) and 7.6% (M. yushuensis vs. M. harpioloides) (Suppl. material 3). Similarly, these two undescribed species formed independent lineages in the phylogeny reconstructed using Cyt b (Fig. 2B and Suppl. material 1: fig. S2), with the minimum genetic distances to congeners being 11.8% and 12.5%, respectively (Suppl. material 4). Thus, the populations at these two localities represent two independently evolved lineages, which are described below as two new species: Murina yadongensis sp. nov. and Murina milinensis sp. nov.

Within subclade IV-3, four samples from Beibeng Township, Medog County, Nyingchi City, and Xizang Autonomous Region, China, clustered together to form a sister clade to (M. guilleni + M. cf. cyclotis 1) and were highly supported (BPP/UFB = 0.95/95) (Fig. 2A and Suppl. material 1: fig. S1). This population can be distinguished from known species of the genus Murina and from lineages undescribed in this study by several morphological characters (see subsequent comparisons) and molecular differences, with a high p-distance of 12.0% (vs. M. guilleni) (Suppl. material 3). This is much greater than the genetic distances between recognized species, e.g., 9.7% (M. hkakaboraziensis vs. M. kontumensis) and 7.6% (M. yushuensis vs. M. harpioloides) (Suppl. material 3). In the phylogeny reconstructed using Cyt b, this population was resolved as a basal clade to M. cyclotis and M. liboensis (Fig. 2B and Suppl. material 1: fig. S2), with the minimum genetic distance to congeners being 11.8% (vs. M. liboensis) (Suppl. material 4). Thus, the population at this locality can be considered an independently evolved lineage and is described below as a new species, Murina beibengensis sp. nov.

Within subclade IV-4, three samples from Beibeng Township, Medog County, Nyingchi City, and Xizang Autonomous Region formed a distinct cluster and were highly supported as the sister clade to M. jaintiana (BPP/UFB = 0.95/99) (Fig. 2B and Suppl. material 1: fig. S1). This population can be distinguished from known species of the genus Murina and from lineages undescribed in this study by morphological characters (see subsequent comparisons) and molecular data, with a p-distance of 7.0% (vs. M. jaintiana) (Suppl. material 3). This value is greater than the genetic distances between recognized sister species, e.g., 4.5% (M. feae vs. M. tubinaris) (Suppl. material 1: table S4). In the phylogeny reconstructed using Cyt b, this population was resolved as the sister species to M. jaintiana (Fig. 2B and Suppl. material 1: fig. S2), with a minimum genetic distance of 6.1% between them (Suppl. material 4). Thus, the population at this locality represents an independently evolved lineage and is described below as a new species, Murina medogensis sp. nov.

Further phylogenetic analysis of expanded samples of the M. huttoni complex recovered seven clades (Fig. 2C), corresponding to those shown in Figure 2A, and published sequences of the subspecies M. huttoni rubella (Zhang et al. 2016) were nested within Clade IV (Fig. 2C). Within the M. huttoni complex, genetic distances ranged from 6.2% to 6.9%, which are much greater than the genetic distances between recognized species as previously described, e.g., 2.9% (M. yuanyang vs. M. chrysochaetes) (Suppl. material 1: table S7).

Species delimitation based on mitochondrial sequences

The mPTP species delimitation resolved the most recognized species and was generally consistent with the phylogenetic results. However, mPTP also suggested that five sets of recognized and undetermined species should be merged into a single species for each set: (1) M. feae + M. tubinaris; (2) M. cf. eleryi + M. eleryi + M. gracilis + M. recondita + M. aurata + M. balaensis; (3) M. chrysochaetes + M. yuanyang; (4) M. rongjiangensis + M. shuipuensis + M. leucogaster; and (5) M. cf. huttoni 1 + M. cf. huttoni 2 + M. cf. huttoni 3 + M. cf. huttoni 5 + M. cf. huttoni 4 + M. puta + M. huttoni (Fig. 2A). Species delimitation using mPTP based on Cyt b supported the results from COI-based delimitation but also suggested the merging of several species: (1) M. feae and M. tubinaris; (2) M. cyclotis and M. liboensis; (3) M. gracilis and M. recondita; (4) M. fanjingshanensis and M. leucogaster; (5) M. rongjiangensis and M. shuipuensis; and (6) the M. huttoni complex (Fig. 2B). However, since the sequences used were not entirely derived from type specimens, these results require further validation with additional evidence. In summary, the mPTP analyses supported the identification of the new species in this study as independent phylogenetic lineages.

The mPTP analysis suggested that M. cf. huttoni from central Laos, Vietnam, China (Hunan, Xizang, Guangdong, and Jiangxi), and central Vietnam be merged into a single species as the sister species of M. puta (Fig. 2C). Therefore, based on genetic and morphological differences, we propose to elevate its subspecies M. h. rubella to the species level. Detailed morphological comparisons are given below.

Morphological analyses

We measured 24 morphological characters from 17 specimens for PCA analysis. The number of morphological characters used for PCA analysis was not entirely consistent because data from the literature were incomplete. Overall, four undescribed new species were separated from closely related species in the PCA plots.

For M. beibengensis sp. nov., M. guilleni, M. cyclotis, and M. peninsularis, a total of three principal components were extracted from the 20 traits. The first three principal components explained 90.16% of the total variation (Suppl. material 1: table S8). In the PCA plot, the cluster formed by a single sample of M. beibengensis sp. nov. was clearly separated from the remaining three species along the PC1 axis, which was loaded with all traits (Fig. 3A).

Figure 3. 

Plots of principal component analysis for the four new species and closely related species, based on morphological characters.

For M. medogensis sp. nov., M. jaintiana, M. feae, and M. tubinaris, three principal components were extracted from the 15 traits. PC1 and PC2 explained 18.85% and 16.53% of the total variance, respectively (Suppl. material 1: table S9). In the PCA plot, M. medogensis sp. nov. showed slight overlap with M. jaintiana and M. feae but was separated along the PC1 axis, forming a distinct cluster comprising a single sample (Fig. 3B).

For M. milinensis sp. nov., M. yadongensis sp. nov., M. chrysochaetes, Murina yuanyang, M. harpioloides, and M. yushuensis, a total of eight principal components were extracted. The first two principal components explained 50.14% of the total variance (Suppl. material 1: table S10). In the PCA plot, M. milinensis sp. nov. and M. yadongensis sp. nov. formed two distinct and independent clusters, which were separated from the remaining species along the PC2 axis (Fig. 3C). Traits loaded on the PC2 axis included FA, TIB, skull, and ear measurements (Suppl. material 1: table S10).

Based on mitochondrial and morphological differences, the four populations of Murina from the Xizang Autonomous Region, China, represent four new undescribed species. We formally describe them in what follows.

Taxonomic account

Murina beibengensis Luo, Mao & Zhou, sp. nov.

Figs 4, 5, Table 2, Suppl. material 2

Holotype.

• Adult male, field number XZ2024037 (Figs 4, 5, Table 2), collected by Tao Luo, Ming-Le Mao, Chang-Ting Lan, Zi-Fa Zhao, and Zhong-Lian Wang on 15 August 2024, from Beibeng Township, Medog County, Nyingchi City, Xizang Autonomous Region, China (29.22926454°N, 95.15008271°E; ca. 864 m. a.s.l.; Fig. 1).

Table 2.

External and craniodental measurements (in mm) of M. beibengensis sp. nov. and three closely related species. NA indicates that the data is not available.

Species M. beibengensis sp. nov. M. guilleni (Soisook et al. 2013a) M. cyclotis (Soisook et al. 2013a) M. peninsularis (Soisook et al. 2013a)
Holotype (Female) Holotype (Male) Male (n = 6) Female (n = 3) Male (n = 36) Female (n = 40) Male (n = 23) Female (n = 19)
BW 6.20 6.50 NA NA NA NA NA NA
HB 43.25 48.00 43.2–51.6 (48.0 ± 3.2) 48.0–48.3 (48.2 ± 0.2) 38.7–46.4 (42.3 ± 2.3) 41.1–50.0 (45.1 ± 2.8) 39.9–50.1 (46.6 ± 3.0) 42.0–55.1 (49.9 ± 3.7)
TL 32.69 35.20 28.1–39.2 (35.9 ± 4.0) 35.9–42.0 (39.1 ± 3.1) 26.2–39.0 (34.9 ± 3.9) 32.0–41.1 (37.4 ± 2.5) 32.4–42.8 (38.7 ± 3.3) 38.5–46.0 (42.3 ± 1.9)
EL 16.18 12.60 11.4–15.2 (13.7 ± 1.4) 13.5–15.1 (14.4 ± 0.8) 12.0–17.6 (14.0 ± 1.3) 12.7–16.0 (14.5 ± 1.0) 11.9–18.8 (14.2 ± 1.7) 13.0–17.0 (15.1 ± 1.2)
EW 8.61 NA NA NA NA NA NA NA
TRL 8.67 NA 8.30–9.20 (8.6 ± 0.5) NA 6.80–9.30 (8.1 ± 0.6) 5.80–9.20 (7.9 ± 0.9) 7.40–10.30 (8.4 ± 0.9) 7.60–9.00 (8.3 ± 0.5)
TRW 2.29 NA NA NA NA NA NA NA
HFL 8.64 8.10 7.7–9.4 (8.5 ± 0.6) 8.0–8.4 (8.2 ± 0.2) 6.5–8.8 (7.9 ± 0.6) 7.0–9.7 (8.3 ± 0.8) 5.6–9.1 (7.8 ± 0.9) 7.1–10.0 (9.0 ± 0.8)
FL 31.94 34.00 31.9–34.0 (33.2 ± 0.7) 35.0–35.9 (35.4 ± 0.5) 29.4–33.0 (30.7 ± 0.9) 31.6–36.8 (33.9 ± 1.0) 33.8–38.1 (35.7 ± 1.3) 34.5–39.4 (37.7 ± 1.2)
TIB 18.42 19.60 17.7–19.7 (18.4 ± 0.8) 18.1–19.7 (18.9 ± 0.8) 14.5–19.3 (17.6 ± 1.1) 17.3–20.3 (17.6 ± 1.1) 18.2–21.6 (19.5 ± 1.0) 11.1–21.0 (19.3 ± 2.5)
GTL 15.68 17.03 16.40–17.54 (17.10 ± 0.43) 17.12–18.10 (17.65 ± 0.50) 15.86–17.08 (16.47 ± 0.34) 16.6–18.18 (17.21 ± 0.47) 17.39–18.52 (17.79 ± 0.31) 17.59–19.33 (18.70 ± 0.52)
CCL 12.50 14.88 14.47–15.19 (14.85 ± 0.26) 15.09–15.76 (15.43 ± 0.34) 13.60–15.12 (14.45 ± 0.34) 14.34–16.17 (15.22 ± 0.41) 14.90–16.41 (15.52 ± 0.41) 15.53–16.89 (16.40 ± 0.36)
BCW 7.57 7.74 7.65–7.82 (7.76 ± 0.06) 7.53–7.74 (7.64 ± 0.11) 7.16–8.10 (7.64 ± 0.22) 7.40–8.17 (7.71 ± 0.19) 7.72–8.48 (8.12 ± 0.19) 7.70–8.58 (8.22 ± 0.22)
BCH 7.87 6.57 6.57–6.91 (6.68 ± 0.12) 6.78–7.10 (6.94 ± 0.16) 6.08–7.22 (6.49 ± 0.30) 6.10–7.21 (6.5 ± 0.24) 6.79–8.22 (7.32 ± 0.32) 7.10–8.37 (7.48 ± 0.34)
ZYW 8.20 9.72 9.29–9.93 (9.60 ± 0.22) 9.75–10.02 (9.92 ± 0.15) 8.78–10.05 (9.36 ± 0.31) 9.33–10.43 (9.84 ± 0.29) 9.76–11.31 (10.36 ± 0.38) 10.12–11.22 (10.80 ± 0.27)
MAW 7.58 8.15 8.00–8.28 (8.12 + 0.11) 8.18–8.43 (8.34 ± 0.14) 7.11–8.48 (7.90 ± 0.26) 7.64–8.58 (8.20 ± 0.21) 8.32–9.39 (8.74 ± 0.27) 8.08–9.62 (9.02 ± 0.34)
IOW 4.24 4.31 4.24–4.44 (4.34 ± 0.07) 4.20–4.44 (4.32 ± 0.12) 3.92–4.48 (4.17 ± 0.12) 3.99–4.52 (4.25 ± 0.13) 4.31–4.97 (4.57 ± 0.19) 4.46–4.88 (4.68 ± 0.12)
CM3L 4.98 5.44 5.44–5.72 (5.58 ± 0.12) 5.50–5.91 (5.67 ± 0.21) 5.12–5.68 (5.41 ± 0.15) 5.06–6.00 (5.61 ± 0.19) 5.52–6.09 (5.76 ± 0.18) 5.68–6.39 (6.07 ± 0.18)
C1C1W 3.65 4.16 4.12–4.31 (4.19 ± 0.07) 4.28–4.44 (4.34 ± 0.09) 3.73–4.27 (4.00 ± 0.14) 4.00–4.67 (4.25 ± 0.14) 4.28–5.28 (4.66 ± 0.25) 4.46–5.26 (4.97 ± 0.17)
M3M3W 4.61 5.80 5.36–5.80 (5.54 ± 0.16) 5.68–5.87 (5.79 ± 0.10) 5.07–5.79 (5.39 ± 0.18) 5.18–6.05 (5.57 ± 0.19) 5.45–6.22 (5.72 ± 0.18) 5.69–6.20 (5.94 ± 0.15)
RCM 0.79 0.72 NA NA NA NA NA NA
CM3L 5.92 6.00 5.83–6.12 (6.00 ± 0.09) 6.01–6.43 (6.18 ± 0.22) 5.57–6.18 (5.84 ± 0.14) 5.75–6.49 (6.11 ± 0.16) 5.94–8.02 (6.31 ± 0.42) 6.28–6.94 (6.55 ± 0.21)
ML 11.52 11.43 11.13–11.74 (11.38 ± 0.21) 11.95–12.34 (12.11 ± 0.21) 10.52–11.68 (11.17 ± 0.30) 11.32–12.78 (11.86 ± 0.35) 11.25–12.92 (11.92+0.39) 12.09–13.59 (12.75 ± 0.41)
MDL 11.98 NA NA NA NA NA NA NA
CPH 4.38 4.78 4.33–4.93 (4.63 ± 0.21) 5.05–5.15 (5.10 ± 0.05) 3.77–4.60 (4.14 ± 0.21) 4.16–5.30 (4.71 ± 0.27) 4.30–5.33 (4.86 ± 0.29) 4.72–6.08 (5.51 ± 0.39)
Figure 4. 

External morphological characteristics of Murina beibengensis sp. nov. in living holotype, XZ2024037. A. Lateral view; B. Ventral view; C. Dorsal view of the body and uropatagium; D. Ventral hairs; E. Dorsal hairs; F. Hindfoot ventral view; G. Hindfoot dorsal view.

Figure 5. 

Micro-CT reconstruction of the skull of Murina beibengensis sp. nov. holotype, XZ2024037. A. Lateral view; B. Dorsal view; C. Ventral view; D. Frontal view of mandible.

Measurements (in mm) and body mass (in g) of the holotype.

HB: 43.25, EL: 16.18, EW: 8.61, TRL: 8.67, TRW: 2.29, HFL: 8.64, FL: 31.94, TIB: 18.42, GTL: 15.68, CCL: 12.50, BCW: 7.57, BCH: 7.78, ZYW: 8.20, MAW: 7.58, IOW: 4.24, CM3L: 4.98, C1C1W: 3.65, M3M3W: 4.61, RCM: 0.79, CM3L: 5.92, ML: 11.52, MDL: 11.98, CPH: 4.38; BW: 6.20.

Etymology.

The specific epithet beibengensis refers to the type locality of the new species: Beibeng Township, Medog County, Nyingchi City, Xizang Autonomous Region, China. We propose the common English name “Beibeng Tube-nosed Bat” and the Chinese name “Bèi Bēng Guǎn Bí Fú (背崩管鼻蝠)”.

Diagnosis.

Murina beibengensis sp. nov. can be distinguished from all other congeners by the following combination of characters: (1) small-size Murina, FL 31.94 mm, GTL 15.68 mm; (2) dorsal fur orangish-yellow overall, grey at the base, gradually transitioning to orange-yellow tips halfway from the base; (3) ventral fur silvery-gray overall, dark grey at the base, with silvery-gray tips; (4) ears broadly rounded, with smoothly convex anterior margins, no notch on posterior margins; (5) forearm and wrists covered with sparse hairs; (6) wing attachment point located at 1/3 from base of claw to base of toe; (7) sagittal and lambdoid crests well developed; (8) I2 is situated laterally anterior to I3 and partially visible in the lateral view, and I2 slightly taller than I3; (9) mesostyles of M1 and M2 are reduced; (10) C1 taller than P4, P2 smaller than P4 in height, and crown area of P2 larger than 2/3 that of P4; (11) P2 approximately equal to P4 in height, with a basal area 2/3 that of P4; (12) C1 larger than P4 in height and basal area; (13) mandibular foramina clearly visible, situated below P2.

Description.

Morphometric data of Murina beibengensis sp. nov. are provided in Table 2 and Suppl. material 2. Small-size Murina, HB 43.25 mm, FL 31.94 mm, EL 16.18 mm, HFL 8.64 mm, and BW 6.20 g. Nostrils tubular, open sideways, and slightly short. Ears small, short, oval, blunt at tips, with smoothly convex anterior margins, not notched on posterior margin. Tragus short, narrow, and tapering toward a pointed tip, with slightly convex anterior margin, concave posterior margin, and basal notch, and slightly curved outward, about half as long as ear. Body covered with thick and fluffy hair. Dorsal hairs orangish-yellow overall (bicolored, grayish black at the base, gradually transitioning to orange-yellow tips halfway from base). Dorsal hairs extend onto bases of wings; uropatagium, thumbs, forearm, wrist, tibia, and feet, with well-developed fringe of hairs around margin of uropatagium. Densely furred anterior 1/3 of the dorsal uropatagium, posterior 2/3 covered with sparse hairs. Ventral hairs are silvery-gray overall, tricolored, gray-white at the base, central dark gray, with silvery-gray tips. Flesh purple around the eyes, muzzle, and lower forehead, and the face is hairy except for the long, protuberant nostrils that are naked. TL 32.69 mm, tip of the tail extending significantly past the rear edge of the uropatagium, tip slightly free. Plagiopatagium attachment point located at the middle of the claw of the first toe.

Skull robust and domed, relatively small, GTL 15.68 mm. Rostrum long, deep, gradually ascending to forehead; prominent median depression present. Sagittal and lambdoid crests well developed. In dorsal view, braincase nearly rounded; zygomatic arches weak and slender, gradually widening posteriorly, widest at the base of the zygomatic arches; posterior margin of skull not protruding; middle from snout to frontal region distinctly concave downward. In lateral view, skull slightly elongated, with elongated oval braincase; height gradually rising from snout to parietal, with slightly increasing slope from snout to frontal and decreasing slope from frontal to parietal; slight depression between snout and frontal, without distinct prominence at frontal; zygomatic arches gradually rising from anterior to posterior, highest at the middle of zygomatic arches. In ventral view, palatine wide and nearly flat, ending at midpoint of C1; basisphenoid pits slightly shallowly teardrop-shaped, extending posteriorly to anterior third of cochlea. Mandible length 11.52 mm, inverted L-shaped. Line between coronoid process and condyle nearly flat; distinct inward depression between condyle and angle; angle slightly long and wide; mandibular foramina clearly visible, situated below P2.

Dental morphology: Dental formula is I2/3, C1/1, P2/2, M3/3=34 (Fig. 5). In the maxilla, I2 is situated laterally anterior to I3 and partially visible in the lateral view; the crown area of P2 larger than 2/3 that of P4 and smaller than that of C1. Based on these characters, the species belongs to the “cyclotis-type”. Maxillary dentition converges slightly anteriorly (RCM = 0.79). I2 and I3 bicuspid, smaller secondary cusp situated posterior to primary cusp; I2 is slightly taller than I3, with a crown area half that of I3; distinct gap between posterior of I3 and C1, not in contact, about 1/3 of height of C1. C1 taller than P4, slightly elongated and lacking secondary cusps, crown area approximately equal to P4; P2 is slightly smaller, delicate and pointed, about 4/5 as high as P4 and 2/3 as high as C1, crown area of P2 is 2/3 that of P4. Mesostyles of M1 and M2 are reduced, but retaining distinct cusps; paracone, protocone, metacone, and parastyle well developed. M3 are reduced, with only parastyle, paracone, and protocone. In the mandible, the first, second, and third lower incisors (I1, I2, and I3) smaller, tricuspid, nearly equal in height and width; slight overlap of outer cusps of I1, I2, and I3 with gradual increase in height from I1 to C1. C1 contains pointed cusp on anterior inner margin, in contact with I3 outer cusp, exceeding P2 and P4 in height and basal area; C1 larger than P4 in height and basal area. P2 nearly equal to P4 in height, and the basal and crown area of P2 approximately 2/3 that of P4. In lateral view, trigonids of M1, M2, and M3, metaconid, and paraconid approximately 2/3 as high as protoconid. Talonids of M1 and M2 are approximately half the size of trigonids; entoconid and hypoconid distinctly separated from trigonid, nearly equal to metaconid and paraconid in height. M1 and M2 are nyctalodont types, with well-developed entoconids. M3 reduced, talonid approximately 1/3 as long as trigonid, paraconid, protoconid, and metaconid complete and well developed.

Morphological comparisons with congeneric species.

Based on its dentition, I2 is situated laterally anterior to I3, and I2 is partially visible in the lateral view, and crown area of P2 larger than 2/3 that of P4, Murina beibengensis sp. nov. belongs to the “cyclotis-type” a character that distinguishes 30 species belonging to the “suilla-type”, I2 is anterior to I3, I2 is clearly visible in the lateral view, and the crown area of P2 is half or less than that of P4, including M. rubella, M. aurata, M. balaensis, M. beelzebub, M. bicolor, M. chrysochaetes, M. eleryi, M. fanjingshanensis, M. feae, M. fusca, M. gracilis, M. harpioloides, M. hilgendorfi, M. hkakaboraziensis, M. jaintiana, M. jinchui, M. kontumensis, M. leucogaster, M. liboensis, M. lorelieae, M. rongjiangensis, M. ryukyuana, M. shuipuensis, M. suilla, M. tenebrosa, M. tubinaris, M. ussuriensis, M. walstoni, M. yuanyang, and M. yushuensis. Detailed morphological differences between the new species and congeners are listed in Suppl. material 5 and illustrated in Suppl. material 1: fig. S3.

Within the “cyclotis-type”, Murina beibengensis sp. nov. can be distinguished from M. aenea, M. fionae, M. harrisoni, M. huttoni, M. peninsularis, and M. puta by the forearm length 31.94 mm (vs. forearm length over 33 mm); from M. annamitica, M. pluvialis, and M. recondita by lacking off-white circumferential band around the neck (vs. present); from M. cyclotis and M. guilleni by the very well-developed sagittal and lambdoid crests (vs. poorly developed); and from M. rozendaali by ventral fur silvery-gray overall (vs. yellowish white) and mesostyles of M1 and M2 moderately reduced (vs. well-developed).

M. florium, which is not assigned to either the “suilla-type” or “cyclotis-type”, can be distinguished by the following morphological characters: P2 is 2/3 the height of P4 (vs. P2 equal to P4), and the mesostyles of M1 and M2 are reduced (vs. well-developed).

Habitat and ecology.

The new species is only found in two areas of Xizang, China. The type specimen was captured in a broad-leaved evergreen forest near Beibeng Township, Medog County, Nyingchi City, Xizang Autonomous Region, China (about 2.8 km from the town center) using a harp net. The type locality has a subtropical climate with a mean annual temperature of 16 °C and mean annual precipitation of 2500–3900 mm. The site is surrounded by several cash crops (e.g., tea trees and bananas). In addition, two Rhinolophus sp., six Hipposideros armiger, three H. pratti, and one Cynopterus sphinx were captured here and in the surrounding area. Another distribution site in a cave near Riduo Township, Maizhokunggar County, Lasa City, and Xizang Autonomous Region yielded two carcasses. This site has a highland temperate semi-arid monsoon climate, with a mean annual temperature of 0.8 °C, mean annual precipitation of 350 mm, and sparse surrounding vegetation. Accordingly, we hypothesize that this new species may have a wider distribution in the Qinghai-Xizang Plateau.

Remark.

Two genetic samples, field numbers XZ2024B05 and XZ2024B12, gender unknown, collected from a cave near Riduo Township, Maizhokunggar County, Lasa City, and Xizang Autonomous Region, China (29.69274086°N, 92.24386096°E, 4414 m) on 12 August 2024. These two samples occurred as dry and decomposed carcasses; no morphological data could be obtained, and only muscle tissue was preserved for sequencing.

Murina medogensis Mao, Lan & Zhou, sp. nov.

Figs 6, 7, 8, 9, Table 3, Suppl. material 2

Holotype.

• Adult male, field number XZ2024038 (Figs 6, 7), collected by Ming-Le Mao, Tao Luo, Chang-Ting Lan, Zi-Fa Zhao, and Zhong-Lian Wang on 16 August 2024, from Medog Town, Medog County, Nyingchi City, Xizang Autonomous Region, China (29.29867524°N, 95.28785706°E; ca. 813 m. a.s.l.; Fig. 1).

Figure 6. 

External morphological characteristics of Murina medogensis sp. nov. in living holotype, XZ2024038. A. Lateral view; B. Ventral view; C. Dorsal view of the body and uropatagium; D. Ventral hairs; E. Dorsal hairs; F. Hindfoot ventral view; G. Hindfoot dorsal view.

Figure 7. 

Micro-CT reconstruction of the skull of Murina medogensis sp. nov. holotype, XZ2024038. A. Lateral view; B. Dorsal view; C. Ventral view; D. Frontal view of mandible.

Paratype.

• Field number XZ2024006 (Figs 8, 9), adult female, collected near Beibeng Township, Medog County, Nyingchi City, Xizang Autonomous Region, China (29.2297514°N, 95.14752388°E; ca. 816 m) on 14 August 2024.

Figure 8. 

Skull and dentition of the Murina medogensis sp. nov. paratype, XZ2024006. A. Lateral view; B. Dorsal view; C. Ventral view; D. Frontal view of mandible.

Figure 9. 

External morphological characteristics of Murina medogensis sp. nov. in living paratype, XZ2024006. A. Lateral view; B. Ventral view; C. Dorsal view of the body and uropatagium; D. Ventral hairs; E. Dorsal hairs; F. Hindfoot ventral view; G. Hindfoot dorsal view.

Measurements (in mm) and body weight (in g) of the holotype.

HB: 38.46, EL: 11.89, EW: 6.79, TRL: 6.19, TRW: 1.94, HFL: 6.94, FL: 29.75, TIB: 15.47, GTL: 15.01, CCL: 14.09, BCW: 7.55, BCH: 6.71, ZYW: 8.74, MAW: 7.11, IOW: 4.39, CM3L: 4.82, C1C1W: 3.61, M3M3W: 5.07, RCM: 0.712, CM3L: 5.86, ML: 9.1, MDL: 10.21, CPH: 3.42; BW: 4.20.

Etymology.

The specific epithet medogensis refers to the type locality of the new species: Medog Town, Medog County, Nyingchi City, Xizang Autonomous Region, China. We propose the common English name “Medog Tube-nosed Bat” and the Chinese name “Mò Tuō Guǎn Bí Fú (墨脱管鼻蝠)”.

Diagnosis.

Murina medogensis sp. nov. can be distinguished from all of the other congeners by the following combination of characters: (1) small-size Murina, FL 29.75–31.24 mm, GTL 15.01–18.74 mm; (2) dorsal fur dark grayish overall, dark grey at the base, gradually transitioning to gray-brown/white tips 2/3 from the base; (3) ventral fur silvery-gray overall, dark grey at the base, with silvery-gray tips; (4) ears broadly rounded, with smoothly convex anterior margins, no notch on posterior margins; (5) forearm and wrists without covered sparse hairs; (6) wing attachment point located at 1/3 from base of claw to base of toe; (7) sagittal crest absent, lambdoid crest present and poorly developed; (8) I2 is situated laterally anterior to I3 and partially visible in the lateral view, I2 equal to I3 in height; (9) mesostyles of M1 and M2 are well developed; (10) C1 slightly lower than P4 in height and crown area of P2 larger than 2/3 that of P4; (11) P2 is approximately equal to P4 in height, with a basal area 2/3 that of P4; (12) C1 taller than P4 in height, crown area equal to P2 and 2/3 of P4; (13) mandibular foramina clearly visible, situated below P4; (14) without distinct prominence at frontal aspect of skull.

Description.

Morphometric data of Murina medogensis sp. nov. are provided in Table 3 and Suppl. material 2. Small-size Murina, HB 37.68–38.46 mm, FL 29.75–31.24 mm, EL 11.89–14.06 mm, HFL 6.94–8.52 mm, and BW 4.20 g. Nostrils tubular, open sideways, and slightly short. Ears small, short, oval, blunt at tips, with smoothly convex anterior margins, not notched on posterior margin. Tragus short, narrow, and tapering toward pointed tips, with slightly convex anterior margin, concave posterior margin, and basal notch, and it curves outwards slightly, about half as long as ear. Body covered with thick and fluffy hair. Dorsal hairs dark grayish overall (bicolored, dark grey at the base, gradually transitioning to gray-brown/white tips 2/3 from base). Dorsal hairs extend onto bases of wings, uropatagium, thumbs, tibia, and feet, with slight-developed fringe of hairs around margin of uropatagium. Densely furred anterior 1/3 of the dorsal uropatagium, posterior 2/3 covered with sparse hairs. Ventral hairs are silvery-gray overall, bicolored, dark grey at the base, with silvery-gray tips. Flesh-purple around the eyes, muzzle, and lower forehead, and the face is hairy except for the long, protuberant nostrils that are naked. TL 34.54–36.37 mm, slightly shorter than head-body length, tip of the tail extending significantly past the rear edge of the uropatagium, tip slightly free. Plagiopatagium attachment point located at about 1/3 from base of claw to base of toe, near base of claw (Fig. 6F, G).

Table 3.

External and craniodental measurements (in mm) of M. medogensis sp. nov. and three closely related species. NA indicates that the data is not available.

Species M. medogensis sp. nov. M. jaintiana (Ruedi et al. 2012) M. feae (Son et al. 2015) M. tubinaris (Csorba et al. 2011)
XZ2024038 (Male) XZ2024006 (Female) Holotype (Male) Range (n = 5) Male Female Range
BW 4.20 4.20 NA 3.5–5.3 3.9–4.4 NA
HB 38.46 37.68 40.00 NA 33.7–43.0 32.8–43.5 NA
TL 36.37 34.54 33.00 NA 31.4–39.5 30.0–41.6 NA
EL 11.89 14.06 13.90 NA 11.5–14.3 11.0–15.0 NA
EW 6.79 6.50 NA NA NA NA NA
TRL 6.19 7.70 6.30 NA NA NA NA
TRW 1.94 2.07 NA NA NA NA NA
HFL 6.94 8.52 6.80 NA 6.0–8.4 6.1–8.0 NA
FL 29.75 31.24 29.10 29.10–31.10 27.5–33.4 28.1–34.3 31.00–32.94
TIB 15.47 17.45 17.10 NA 15.7–18.1 15.6–17.8 NA
GTL 15.01 18.74 NA 14.75–15.25 14.91–16.30 15.13–16.11 14.92–15.70
CCL 14.09 16.37 NA 13.36–13.61 NA NA 13.08–13.89
BCW 7.55 9.09 NA NA NA NA 7.01–7.30
BCH 6.71 7.54 NA 5.96–6.17 NA NA 5.55–5.86
ZYW 8.74 10.85 NA 8.26–8.68 8.50–8.90 8.9 8.07–8.66
MAW 7.11 9.32 NA 7.23–7.43 7.30–7.80 7.5 7.27–7.51
IOW 4.39 5.49 NA 4.02–4.36 NA NA 4.28–4.51
CM3L 4.82 7.54 NA 4.81–5.09 5.00–5.30 5.1 4.88–5.19
C1C1W 3.61 4.85 NA 3.52–3.74 3.70–4.00 3.9–4.4 3.59–3.78
M3M3W 5.07 6.72 NA 5.04–5.20 NA NA 4.97–5.32
RCM 0.712 0.72 NA NA NA NA NA
CM3L 5.86 7.25 NA 5.18–5.46 4.62–5.25 4.89–5.47 5.37–5.69
ML 9.1 12.54 NA 9.85–10.28 9.63–10.64 9.80–10.86 10.07–10.62
MDL 10.21 13.23 NA NA NA NA NA
CPH 3.42 2.97 NA 3.20–3.40 NA NA 3.04–3.30

Skull robust and domed, relatively small, GTL 15.01–18.74 mm. Rostrum long, deep, gradually ascending to forehead; prominent median depression present. Sagittal crest absent, lambdoid crest present, poorly developed. In dorsal view, braincase nearly rounded; zygomatic arches weak and slender, gradually widening posteriorly, widest at the base of the zygomatic arches; posterior margin of skull slightly protruding; middle from snout to frontal region distinctly concave downward. In lateral view, skull slightly elongated, with elongated oval braincase; height gradually rising from snout to parietal, with slightly increasing slope from snout to frontal and decreasing slope from frontal to parietal; slight depression between snout and frontal, without distinct prominence at frontal; zygomatic arches gradually rising from anterior to posterior. In ventral view, palatine wide and nearly flat, ending at posterior margin of C1; basisphenoid pits slightly shallowly teardrop-shaped, extending posteriorly to anterior third of cochlea. Mandible length 9.1 mm, inverted L-shaped. Line between coronoid process and condyle nearly flat; distinct inward depression between condyle and angle; angle slightly long and wide; mandibular foramina clearly visible, situated below P4.

Dental morphology: Dental formula is I2/3, C1/1, P2/2, M3/3=34 (Figs 7, 8). In the maxilla, I2 is situated laterally anterior to I3 and partially visible in the lateral view; crown area of P2 larger than 2/3 that of P4 and smaller than that of C1. Based on these characters, the species belongs to the “cyclotis-type”. Maxillary dentition converges slightly anteriorly (RCM = 0.71). I2 and I3 bicuspid, smaller secondary cusp situated posterior to primary cusp; I2 equal to I3 in height, crown area of I2 less than half that of I3; distinct gap between posterior surface of I3 and C1, not in contact, about half of height of C1. C1 slightly lower than P4 in height, slightly elongated and lacking secondary cusps, crown area 2/3 that of P4; P2 slightly smaller, delicate and pointed, about half as high as P4 and 2/3 as high as C1, crown area 2/3 that of P4. Mesostyles of M1 and M2 are reduced, but retaining distinct cusps; paracone, protocone, metacone, and parastyle well developed. M3 reduced, with only parastyle, paracone, and protocone. In the mandible, I1, I2, and I3 smaller, tricuspid, almost equal in height and width; slight overlap of outer cusps of I1, I2, and I3; with gradual increase in height from I1 to C1. C1 contains pointed cusp on anterior inner margin, in contact with I3 outer cusp, exceeding P2 and P4 in height; C1 equal to P2 in basal area but smaller than P4. P2 slightly smaller than P4 in height, and the basal and crown area of P2 approximately 2/3 that of P4. In lateral view, trigonids of M1, M2, and M3, metaconid, and paraconid approximately 2/3 as high as protoconid in height. Talonids of M1 and M2 are slightly half the size of trigonids; entoconids and hypoconids distinctly separated from trigonids, lower than metaconid and paraconid. M1 and M2 are nyctalodont types, with well-developed entoconids. M3 reduced, talonid approximately 1/3 as long as trigonid, paraconid, protoconid, and metaconid complete and well developed.

Morphological comparisons with congeneric species.

Based on its dentition, I2 is situated laterally anterior to I3, and I2 is partially visible in the lateral view, and crown area of P2 larger than 2/3 that of P4, Murina medogensis sp. nov. belongs to the “cyclotis-type” a character that distinguishes 30 species belonging to the “suilla-type”, I2 is anterior to I3, I2 is clearly visible in the lateral view, and the crown area of P2 is half or less than that of P4, including M. rubella, M. aurata, M. balaensis, M. beelzebub, M. bicolor, M. chrysochaetes, M. eleryi, M. fanjingshanensis, M. feae, M. fusca, M. gracilis, M. harpioloides, M. hilgendorfi, M. hkakaboraziensis, M. jaintiana, M. jinchui, M. kontumensis, M. leucogaster, M. liboensis, M. lorelieae, M. rongjiangensis, M. ryukyuana, M. shuipuensis, M. suilla, M. tenebrosa, M. tubinaris, M. ussuriensis, M. walstoni, M. yuanyang, and M. yushuensis. Detailed morphological differences between the new species and congeners are listed in Suppl. material 5 and illustrated in Suppl. material 1: fig. S3.

Murina medogensis sp. nov. can be distinguished from Murina beibengensis sp. nov. by dorsal fur dark grayish overall (vs. orangish-yellow), forearm and wrists without covered sparse hairs (vs. covered with sparse hairs), lambdoid crest well developed (vs. absent), C1 slightly less than P4 in height (vs. C1 taller than P4), P2 slightly less than P4 in height (vs. P2 equal to P4), and mandibular foramina clearly visible, situated below P4 (vs. situated below P2).

Within the “cyclotis-type”, Murina beibengensis sp. nov. different from M. aenea, M. fionae, M. harrisoni, M. huttoni, and M. pluvialis by the forearm length 29.75–31.24 mm (vs. forearm length over 34 mm). By dorsal hairs dark grayish, Murina medogensis sp. nov. can be further distinguished from M. aenea (vs. dark brown), M. fionae (vs. orange), M. harrisoni (vs. orangish brown), M. huttoni (vs. rusty brown), and M. pluvialis (vs. reddish brown).

Murina beibengensis sp. nov. different from M. annamitica, M. cyclotis, M. guilleni, M. peninsularis, and M. recondita by the sagittal crest absence (vs. presence) and dorsal hairs dark grayish (vs. orangish brown or yellowish brown). Murina beibengensis sp. nov. different from M. puta by the forearm length 29.75–31.24 mm (vs. 33.0–39.0 mm), dorsal fur (bicolored, with dark grey at the base and gray-brown tips vs. tricolored, with black at the base, tan in the middle, and reddish tips), and ventral fur silvery-gray (vs. paler). Murina beibengensis sp. nov. different from M. rozendaali by dorsal fur dark grayish (vs. shiny yellowish brown), ventral fur silvery-gray (vs. yellowish white), mesostyles of M1 and M2 reduced (vs. well-developed).

Murina medogensis sp. nov. is morphologically similar to M. jaintiana but can be distinctly distinguished from it by combining the following morphological characters: dorsal fur bicolored, dark grey at the base, gradually transitioning to gray-brown/white tips from 2/3 from base (vs. dorsal fur has three distinct bands: basal half dark grey, almost black, middle part dirty white, distal end brownish-grey), I2 equal to I3 in height (vs. I2 smaller than I3), C1 slightly less than P4 in height (vs. C1 taller than P4), and without distinct prominence at frontal aspect of skull (vs. with distinct prominence at frontal) (Ruedi et al. 2012).

For M. florium not assigned to “suilla-type” and “cyclotis-type”, can be distinguished by the combination of the following morphological characters: dorsal fur dark grayish overall (vs. gray-brown to orange rufous brown), absence of notch at the posterior margin of the ear (vs. distinct notch on posterior margin), and P2 less than P4 in height (vs. P2 equal to P4).

Habitat and ecology.

Currently, this new species is known only from Medog County, Nyingchi City, in the Tibet Autonomous Region, China, where two specimens were captured using a harp-shaped trap in broad-leaved evergreen forests near Medog Town, Medog County, on 16 August 2024, at two locations approximately 8 km apart. The type locality exhibits a subtropical climate, characterized by a mean annual temperature of 16 °C and mean annual precipitation ranging from 2500 to 3900 mm. The area is also surrounded by various cash crops, such as tea and banana plants. Additionally, species such as Harpiola isodon, Sphaerias blanfordi, Myotis sp., and Kerivoula kachinensis were also captured in the same region.

Murina milinensis Luo, Mao & Zhou, sp. nov.

Figs 10, 11, 12, Table 4, Suppl. material 2

Holotype.

• Adult male, field number XZ2023010 (Figs 10, 11), collected by Ming-Le Mao, Chang-Ting Lan, Zi-Fa Zhao, and Zhong-Lian Wang on 14 August 2023, from Nanyi Lhoba Township, Milin County, Nyingchi City, Xizang Autonomous Region, China (29.1598922°N, 94.20712670°E; ca. 3020 m. a.s.l.; Fig. 1).

Figure 10. 

External morphological characteristics of Murina milinensis sp. nov. in living holotype, XZ2023010. A. Lateral view; B. Ventral view; C. Dorsal view of the body and uropatagium; D. Ventral hairs; E. Dorsal hairs; F. Hindfoot ventral view; G. Hindfoot dorsal view. These photographs were acquired under incandescent lighting conditions using a Canon EOS 6D Mark II camera ISO: 6400; focal length: 105 mm at approximately 22:41 on 5 August 2023.

Figure 11. 

Micro-CT graph of the skull of Murina milinensis sp. nov. paratype, XZ2024066. A. Lateral view; B. Dorsal view; C. Ventral view; D. Frontal view of mandible.

Paratypes.

• Eight specimens from the same locality as the holotype. Five males, field numbers XZ2024066, XZ2024065, XZ2024067, XZ2024074, and XZ2024075. Three females, field numbers XZ202384, XZ2024072, and XZ2024076. Female XZ2024102 is from Zari Township, Longzi County, Shannan City, Xizang Autonomous Region, China (28.68318987°N, 93.34190369°E; ca. 3450 m).

Measurements (in mm) and body weight (in g) of the holotype.

HB: 34.68, EL: 12.74, EW: 7.89, TRL: 7.81, TRW: 5.35, HFL: 5.31, FL: 30.02, TIB: 12.75, GTL: 14.07, CCL: 12.01, BCW: 7.03, BCH: 6.49, ZYW: 8.07, MAW: 6.68, IOW: 4.40, CM3L: 4.44, C1C1W: 2.21, M3M3W: 4.97, RCM: 0.44, CM3L: 4.53, ML: 9.06, MDL: 9.46, CPH: 3.20; BW: 3.36.

Etymology.

The specific epithet milinensis refers to the type locality of the new species: Nanyi Lhoba Township, Milin County, Nyingchi City, Xizang Autonomous Region, China. We propose the common English name “Milin Tube-nosed Bat” and the Chinese name “Mǐ Lín Guǎn Bí Fú (米林管鼻蝠)”.

Diagnosis.

Murina milinensis sp. nov. can be distinguished from all of the other congeners by the following combination of characters: (1) small-size Murina, FL 28.84–34.01 mm, GTL 14.07–14.27 mm; (2) dorsal fur brown-gold overall, black at the base, gradually transitioning to brown-gold tips 2/3 from the base; (3) ventral fur pale overall, dark black at the base, with grayish white at the tips; (4) ears narrow and oval, without smoothly convex anterior margins, no notch on posterior margins; (5) forearm and wrists covered with sparse hairs; (6) wing attachment point located at 1/3 from base of claw to base of toe; (7) sagittal crest absent, lambdoid crest present and poorly developed; (8) I2 is situated anterior to I3 and clearly visible in the lateral view, I2 equal to I3 in height; (9) mesostyles of M1 and M2 are slightly reduced; (10) C1 less than P4 in height, crown is about 2/3 of P4; (11) P2 approximately half of P4 in height, crown area of P2 larger than 2/3 that of P4; (12) C1 equal to P4 in height and crown area; (13) mandibular foramina clearly visible, situated below anterior margin of P4.

Description.

Morphometric data of Murina milinensis sp. nov. are provided in Table 4 and Suppl. material 2. Small-size Murina, HB 29.23–39.29 mm, FL 28.84–34.01 mm, EL 12.48–14.58 mm, HFL 5.31–9.18 mm, and BW 3.15–7.50 g. Nostrils tubular, open sideways, and slightly longer. Ears small, short, and narrow, oval, blunt at tips, with smoothly convex anterior margins, not notched on posterior margin. Tragus short, narrow, and tapering toward pointed tip, with slightly convex anterior margin, concave posterior margin, and basal notch, and it curves outwards slightly, about half as long as ear. Body covered with thick and fluffy hair. Dorsal hairs brown-gold overall (bicolored, black at the base, gradually transitioning to brown-gold tips from 2/3 from base). Dorsal hairs extend onto bases of wings, uropatagium, thumbs, forearm, wrist, tibia, and feet, with slight-developed fringe of hairs around margin of uropatagium. Densely furred anterior 1/3 of the dorsal uropatagium, posterior 2/3 covered with sparse hairs. Ventral hairs are pale overall, bicolored, bicolored, dark black at the base and grayish white at the tips. Dark flesh-purple around the eyes, muzzle, and lower forehead, and the face is hairy except for the long, protuberant nostrils that are naked. TL 24.51–30.32 mm, slightly shorter than head-body length, tip of the tail extending significantly past the rear edge of the uropatagium, tip distinctly free. Plagiopatagium attachment point located at 1/3 from base of claw to base of toe, near base of claw (Fig. 10F).

Table 4.

External and craniodental measurements (in mm) of M. milinensis sp. nov., M. yadongensis sp. nov., and four closely related species. MC: M. chrysochaetes (Eger et al. 2011); MH: M. harpioloides (Kruskop and Eger 2008); MY: M. yushuensis (Wang et al. 2024a). NA indicates that the data is not available.

Species M. yadongensis sp. nov. M. milinensis sp. nov. MC MH MY M. yuanyang (Mou et al. 2024)
Male (n = 3) Female (n = 1) Male (n = 6) Female (n = 4) Holotype (male) Holotype (male) Holotype (male) Holotype (male) Male (n = 2) Female (n = 5)
BW 5.68–7.00 (6.52 ± 0.73) 6.86 3.15–3.49 (3.37 ± 0.12) 3.35–7.50 (4.73 ± 1.88) NA NA NA 4.20 3.4–3.6 3.8–4.7 (4.3 ± 0.3)
HB 28.68–35.54 (32.21 ± 3.43) 33.88 33.70–37.89 (35.78 ± 1.57) 29.23–39.29 (34.30 ± 5.60) NA 35.00 30.44 35.05 31.37–35.94 32.11–36.34 (34.82 ± 1.52)
TL 28.32–30.46 (29.52 ± 1.09) 32.12 25.54–30.32 (28.88 ± 1.69) 24.51–28.98 (26.66 ± 2.29) 28.00 30.50 28.08 31.58 23.89–27.25 26.92–31.58 (28.39 ± 1.64)
EL 11.40–14.02 (12.85 ± 1.33) 14.2 12.48–14.58 (13.46 ± 0.82) 13.12–13.95 (13.55 ± 0.34) 11.00 12.30 11.08 12.99 12.77–13.4 12.27–14.77 (13.25 ± 0.82)
EW 6.20–7.06 (6.54 ± 0.46) 8.04 6.92–8.47 (7.70 ± 0.52) 6.76–8.58 (7.38 ± 0.84) NA NA NA NA NA NA
TRL 7.15–7.49 (7.37 ± 0.19) 7.81 6.59–7.82 (7.31 ± 0.51) 6.28–8.35 (7.22 ± 1.04) NA NA NA NA NA NA
TRW 2.14–2.15 (2.14 ± 0.01) 2.13 1.51–5.35 (2.49 ± 1.42) 1.76–2.34 (2.03 ± 0.28) NA NA NA NA NA NA
HFL 6.07–8.55 (7.15 ± 1.27) 7.06 5.31–8.89 (7.37 ± 1.32) 6.82–9.18 (7.94 ± 0.97) 7.00 NA 8.46 7.88 6.14–6.83 6.25–8.52 (7.66 ± 0.75)
FL 28.81–30.29 (29.60 ± 0.75) 31.84 28.84–30.43 (29.82 ± 0.60) 30.83–34.01 (32.12 ± 1.35) 26.35 29.70 31.34 29.80 NA NA
TIB 13.57–14.27 (13.82 ± 0.39) 14.61 12.64–13.95 (13.42 ± 0.60) 13.32–14.37 (13.94 ± 0.44) 10.92 NA 15.22 13.48 12.56–13.16 12.37–13.57 (13.15 ± 0.43)
GTL 13.73–14.11 (13.92 ± 0.27) NA 14.07–14.27 (14.17 ± 0.14) 14.16 14.05 NA 14.14 14.13 13.44–14 13.95–14.16 (14.09 ± 0.07)
CCL 12.61–12.89 (12.75 ± 0.20) NA 12.01–13.29 (12.65 ± 0.91) 12.26 12.45 12.34 13.85 12.12 11.58–11.93 11.94–12.22 (12.10 ± 0.09)
BCW 6.36–6.70 (6.53 ± 0.24) NA 6.81–7.03 (6.92 ± 0.16) 6.92 6.98 7.21 7.23 6.95 6.91–6.93 6.93–7.15 (7.00 ± 0.08)
BCH 5.66–6.53 (6.10 ± 0.62) NA 6.25–6.49 (6.37 ± 0.17) 6.64 NA 5.81 6.05 6.78 6.59–6.98 6.51–6.78 (6.62 ± 0.09)
ZYW 7.76–8.72 (8.24 ± 0.68) NA 7.63–8.07 (7.85 ± 0.31) 7.38 7.85 NA 7.94 7.73 7.41–7.52 7.44–7.81 (7.69 ± 0.13)
MAW 7.03–7.04 (7.04 ± 0.01) NA 6.13–6.68 (6.41 ± 0.39) 6.06 7.07 7.42 6.89 7.21 7.18–7.23 7.12–7.38 (7.31 ± 0.10)
IOW 3.51–4.39 (3.95 ± 0.62) NA 3.98–4.40 (4.19 ± 0.30) 3.82 NA 4.09 4.31 3.84 3.67–3.78 3.68–3.86 (3.79 ± 0.06)
CM3L 4.46–5.31 (4.89 ± 0.60) NA 4.32–4.44 (4.38 ± 0.08) 4.66 4.36 4.68 4.42 4.51 4.38–4.43 4.45–4.63 (4.53 ± 0.06)
C1C1W 2.19–2.45 (2.32 ± 0.18) NA 2.21–3.32 (2.77 ± 0.78) 2.07 NA NA NA 3.22 3.01–3.21 3.06–3.28 (3.21 ± 0.08)
M3M3W 3.30–3.80 (3.55 ± 0.35) NA 4.74–4.97 (4.86 ± 0.16) 3.38 4.36 4.88 4.83 4.74 4.59–4.61 4.62–4.74 (4.67 ± 0.04)
RCM 0.64–0.66 (0.65 ± 0.01) NA 0.44–0.70 (0.57 ± 0.18) 0.61 0.73 0.69 0.67 0.68 NA NA
CM3L 4.91–5.74 (5.33 ± 0.59) NA 4.53–5.11 (4.82 ± 0.41) 5.08 4.36 5.13 4.09 5.03 4.71–4.81 4.88–5.03 (4.93 ± 0.05)
ML 8.85–9.22 (9.04 ± 0.26) NA 8.77–9.06 (8.92 ± 0.21) 9.42 8.42 9.31 9.08 8.41 8.09–8.15 8.34–8.69 (8.49 ± 0.13)
MDL 8.72–9.49 (9.11 ± 0.54) NA 9.24–9.46 (9.35 ± 0.16) 9.82 8.42 NA NA 8.76 8.21–8.43 8.51–8.91 (8.75 ± 0.13)
CPH 3.30–3.61 (3.46 ± 0.22) NA 2.99–3.20 (3.10 ± 0.15) 3.31 3.06 3.32 2.96 2.86 2.52–2.76 2.69–2.93 (2.83 ± 0.09)

Skull robust, nearly oval, relatively small, GTL 14.07–14.27 mm. Rostrum long, deep, gradually ascending to forehead; prominent median depression present. Sagittal crest absent, lambdoid crest present, poorly developed. In dorsal view, braincase nearly domed; zygomatic arches weak and slender, gradually widening posteriorly, widest at the base of the zygomatic arches; posterior margin of skull slightly protruding; middle from snout to frontal region distinctly concave downward. In lateral view, skull slightly elongated, with elongated oval braincase; height gradually rising from snout to parietal, with slightly increasing slope from snout to frontal and decreasing slope from frontal to parietal; slight depression between snout and frontal, with slight distinct prominence at frontal; zygomatic arches gradually rising from anterior to posterior. In ventral view, palatine wide and nearly flat, ending at posterior margin of C1; basisphenoid pits slightly shallowly tadpole-shaped, extending posteriorly to anterior half of cochlea. Mandible length 8.77–9.42 mm, inverted L-shaped in lateral view. Line between coronoid process and condyle nearly flat; distinct inward depression between condyle and angle; angle short and wide; mandibular foramina clearly visible, situated below anterior margin of P4.

Dental morphology: Dental formula is I2/3, C1/1, P2/2, M3/3=34 (Figs 11, 12). In the maxilla, I2 is situated anterior to I3, and I2 clearly visible laterally; crown area of P2 approximately 1/3 that of P4 and slightly smaller than C1. Based on these characters, the species belongs to the “suilla-type”. Maxillary dentition converges slightly anteriorly (RCM = 0.44–0.70). I2 and I3 bicuspid, smaller secondary cusp situated posterior to primary cusp; I2 almost equal to I3 in height, crown area of I2 half that of I3; distinct gap between posterior surface of I3 and C1, not in contact, about half of height of C1. C1 less than P4 in height, slightly elongated and lacking secondary cusps, crown area half that of P4; P2 slightly smaller, delicate and pointed, about half as high as P4 and 2/3 as high as C1, and crown area of P2 is half that of P4 and slightly smaller than C1. Mesostyles of M1 and M2 are reduced, but retaining distinct cusps; paracone, protocone, metacone, and parastyle well developed. M3 reduced, with only parastyle, paracone, and protocone. In the mandible, I1, I2, and I3 smaller, tricuspid, almost equal in height and width; slight overlap of outer cusps of I1, I2, and I3; with gradual increase in height from I1 to C1. C1 contains pointed cusp on anterior inner margin, in contact with I3 outer cusp. C1 taller than P2 and P4 in height, with a basal area larger than P2 but smaller than P4. P2 equal to P4 in height, and the basal and crown areas are about half of P4. In lateral view, trigonids of M1, M2, and M3 clearly bicuspid, metaconid, and paraconid approximately 2/3 as high as protoconid in height. Talonid of M1 and M2 is slightly half the size of trigonid; entoconid and hypoconid distinctly separated from trigonid, lower than metaconid and paraconid, nearly equal to metaconid and paraconid in height. M1 and M2 are nyctalodont types, with well-developed entoconids. M3 reduced, talonid approximately 1/3 as long as trigonid, paraconid, protoconid, and metaconid complete and well developed.

Figure 12. 

Skull and dentition of the Murina milinensis sp. nov. holotype, XZ2023010. A. Lateral view; B. Dorsal view; C. Ventral view; D. Frontal view of mandible.

Morphological comparisons with congeneric species.

Based on its dentition, I2 situated anterior to I3, and the crown area of P2 is less than half that of P4 and smaller than that of C1, Murina beibengensis sp. nov. belongs to the “suilla-type”, a character that distinguishes 12 species belonging to the “cyclotis-type,” including M. aenea, M. annamitica, M. cyclotis, M. fionae, M. guilleni, M. harrisoni, M. huttoni, M. peninsularis, M. pluvialis, M. puta, M. recondita, and M. rozendaali. Detailed morphological differences between the new species and congeners are listed in Suppl. material 5 and illustrated in Suppl. material 1: fig. S3.

Murina milinensis sp. nov. can be distinguished from Murina beibengensis sp. nov. by dorsal fur dark grayish overall (vs. brown-gold), ventral fur black at the base (vs. dark grey at the base), sagittal crest absent (vs. well-developed), I2 is situated laterally anterior to I3, and I2 is partially visible in the lateral view (vs. I2 is anterior to I3, I2 is clearly visible in the lateral view), I2 equal to I3 in height (vs. I2 taller than I3), and C1 slightly less than P4 in height (vs. C1 taller than P4).

Murina milinensis sp. nov. can be distinguished from Murina medogensis sp. nov. by dorsal fur brown-gold (s. dark grayish), ventral fur pale (vs. silvery-gray), dorsal hairs extend onto forearm and wrist (vs. hairless forearms and wrists), I2 is situated laterally anterior to I3, and I2 is partially visible in the lateral view (vs. I2 is anterior to I3, I2 is clearly visible in the lateral view), and I2 equal to I3 in height (vs. I2 taller than I3).

Murina milinensis sp. nov. can be distinguished from M. fanjingshanensis, M. fusca, M. hilgendorfi, and M. leucogaster by the small size, forearm length 28.84–34.01 mm, and greatest length of skull 14.07–14.27 mm (vs. forearm length over 37 mm and greatest length of skull over 16 mm in the latter).

Murina milinensis sp. nov. can be distinguished from M. aurata by dorsal fur brown-gold, bicolored, black at the base, gradually transitioning to brown-gold tips from 2/3 from base (vs. golden brown), from M. balaensis (vs. orange-reddish), from M. eleryi (vs. coppery brown), from M. feae (vs. dark grayish), from M. jaintiana (vs. medium gray with brownish tinge), from M. jinchui (vs. brownish gray), from M. liboensis (vs. yellowish brown), from M. lorelieae (vs. reddish brown), from M. walstoni (vs. brownish gray), from M. walstoni (vs. warm brown, whitish at the base, orangish brown at tips), and from M. yuanyang (vs. bark gold). M. milinensis sp. nov. can be further distinguished from M. bicolor by ventral fur is pale overall, bicolored, dark at the base, pale at tips (vs. uniformly yellow), from M. rongjiangensis (vs. bright yellowish orange), and from M. shuipuensis (vs. bright orange yellow).

Murina milinensis sp. nov. can be distinguished from M. beelzebub, M. gracilis, M. harpioloides, M. hkakaboraziensis, M. kontumensis, and M. suilla by the ears without smooth convex anterior margins and no notch on posterior margin (vs. with smoothly convex anterior margins, distinct notch on posterior margin); from M. chrysochaetes, M. ryukyuana, M. tenebrosa, M. ussuriensis, and M. yushuensis (vs. with smoothly convex anterior margins). By lambdoid crest absent and I2 equal to I3 in height, Murina milinensis sp. nov. can be further distinguished from M. rubella and M. chrysochaetes (vs. present, I2 less than I3); from M. rongjiangensis (vs. present, I2 taller than I3), and M. yushuensis (vs. present). Murina milinensis sp. nov. can be distinguished from M. bicolor by sagittal crest absent (vs. present), C1 slightly less than P4 in height (vs. C1 taller than P4), P2 less than P4 in height (vs. P2 taller than P4), and C1 taller than P4 in height (vs. C1 equal to P4).

Murina milinensis sp. nov. can be distinguished from M. tubinaris by dorsal hairs brown-gold (vs. light gray), P2 less than P4 in height (vs. P2 taller than P4), ears without smooth convex anterior margins and no notch on posterior margin (vs. with smoothly convex anterior margins and small notch on posterior margins), and basal area of P2 is larger than one-half that of P4 (vs. less than one-half that of P4).

For M. florium not assigned to “suilla-type” and “cyclotis-type”, can be distinguished by the combination of the following morphological characters: absence of notch at the posterior margin of the ear (vs. distinct notch on posterior margin), I2 almost equal to I3 in height (vs. I2 taller than I3), and P2 slightly less than P4 in height and crown area (vs. equally).

Habitat and ecology.

Currently, this new species is known only from two areas in the Xizang Autonomous Region, China. The type specimen was captured using a harp trap in a water channel near Nanyi Lhoba Township, Miling County, Nyingchi City, on 14 August 2023. A single additional specimen of the same species (Specimen No. XZ2023084) was also captured at the same site. The water channel was approximately 4 m wide and 3 m deep, with a shallow flow of water (approximately 25 cm deep). Dense thickets grew along both sides of the channel. Nanyi Lhoba Township, situated at an elevation of about 3020 m, experiences a plateau temperate semi-humid monsoon climate, with an average annual temperature of approximately 9.3 °C and an annual rainfall of around 600 mm. The collection site is located in a river valley, surrounded by mixed coniferous and broadleaf forests as well as agricultural land, where wheat, highland barley, and oilseed rape are the primary crops. In August of the following year, we captured seven specimens of the same species at the same site. Additionally, the presence of Myotis sp. was recorded in this area.

Another distribution site is located in the mixed coniferous and broadleaf forest near Zari Township, Longzi County, Shannan City, where one specimen was captured using a harp-shaped trap. This site has a plateau temperate continental monsoon climate, with an average annual temperature of 10.3 °C and annual precipitation ranging from 350 to 550 mm. The collection site is also situated in a river valley with dense vegetation. In the vicinity, we recorded Rhinolophus sp. and Myotis sp.

Murina yadongensis Mao, Zhao & Zhou, sp. nov.

Figs 13, 14, Table 4, Suppl. material 2

Holotype.

• Adult male, field number XZ2023082 (Figs 13, 14), collected by Ming-Le Mao, Qing-Qing He, and Qin Yang, on 6 August 2023, at Boluoka, Xiayadong Township, Yadong County, Shigatse City, Xizang Autonomous Region, China (27.36420685°N, 88.97613108°E; ca. 2775 m. a.s.l.; Fig. 1).

Figure 13. 

External morphological characteristics of the Murina yadongensis sp. nov. in living holotype XZ2023082. A. Lateral view; B. Ventral view; C. Dorsal view of the body and uropatagium; D. Ventral hairs; E. Dorsal hairs; F. Hindfoot ventral view; G. Hindfoot dorsal view.

Figure 14. 

Skull and dentition of the Murina yadongensis sp. nov. holotype, XZ2023082. A. Lateral view; B. Dorsal view; C. Ventral view; D. Frontal view of mandible.

Paratypes.

• Three specimens from the same locality as the holotype. Two males, field numbers XZ2023036 and XZ2023016. One female, field numbers XZ2023062. XZ2023082 and XZ2023062 have been dissected for sampling and are incomplete specimens.

Measurements (in mm) and body weight (in g) of the holotype.

HB: 35.54, EL: 14.02, EW: 6.20, TRL: 7.48, TRW: 2.14, HFL: 6.07, FL: 30.29, TIB: 13.57, GTL: 13.73, CCL: 12.61, BCW: 6.36, BCH: 5.66, ZYW: 8.72, MAW: 7.04, IOW: 3.51, CM3L: 5.31, C1C1W: 2.45, M3M3W: 3.80, RCM: 0.64, CM3L: 5.74, ML: 9.22, MDL: 9.49, CPH: 3.61; BW: 5.68.

Etymology.

The specific epithet yadongensis refers to the type locality of the new species: Xiayadong Township, Yadong County, Shigatse City, Xizang Autonomous Region, China. We propose the common English name “Yadong Tube-nosed Bat” and the Chinese name “Yà Dōng Guǎn Bí Fú (亚东管鼻蝠)”.

Diagnosis.

Murina yadongensis sp. nov. can be distinguished from all of the other congeners by the following combination of characters: (1) small-size Murina, FL 28.81–31.84 mm, GTL 13.73–14.11 mm; (2) dorsal fur brown-gold, black at the base, gradually transitioning to brown-gold tips 2/3 from base; (3) ventral fur grayish brown, black at the base, gray-white at tips; (4) ears narrow and oval, without smoothly convex anterior margins, no notch on posterior margins; (5) forearm and wrists covered with sparse hairs; (6) wing attachment point located at 1/3 from base of claw to base of toe; (7) an off-white circumferential band around the neck; (8) sagittal crest absent, lambdoid crest present, poorly developed; (9) I2 is situated anterior to I3 and clearly visible in lateral view, I2 equal to I3 in height; (10) mesostyles of M1 and M2 are slightly reduced; (11) C1 taller than P4, crown is about 2/3 of P4; (12) P2 exceeded 2/3 of P4 in height, and the crown area is about 2/3 of P4; (13) C1 equal to P4 in height and crown area. and crown area of P2 larger than 2/3 that of P4.

Description.

Morphometric data of Murina yadongensis sp. nov. are provided in Table 4 and Suppl. material 2. Small-size Murina, HB 28.68–35.54 mm, FL 28.81–31.84 mm, EL 11.40–14.20 mm, HFL 6.07–8.55 mm, and BW 5.68–7.00 g. Nostrils tubular, open sideways, and slightly longer. Ears small, short, and narrow, oval, blunt at tips, without smoothly convex anterior margins, not notched on posterior margin. Tragus short, narrow, and tapering toward pointed tip, with slightly convex anterior margin, concave posterior margin, and basal notch, and it curves outwards slightly, about half as long as ear. Body covered with thick and fluffy hair. Dorsal hairs brown-gold (bicolored, black at the base, gradually transitioning to brown-gold tips from 2/3 from base). Dorsal hairs extend onto bases of wings, uropatagium, thumbs, forearm, wrist, tibia, and feet, with slight-developed fringe of hairs around margin of uropatagium. Densely furred anterior 1/3 of the dorsal uropatagium, posterior 2/3 covered with sparse hairs. Ventral hairs grayish brown overall, bicolored, black at the base, gray-white at tips. An off-white circumferential band around the neck. Dark flesh-purple around the eyes, muzzle, and lower forehead, and the face is hairy except for the long, protuberant nostrils that are naked. TL 28.32–32.12 mm, slightly shorter than head-body length, tip of the tail extending significantly past the rear edge of the uropatagium, tip distinctly free. Plagiopatagium attachment point located at 1/3 from base of claw to base of toe, near base of claw (Fig. 13E, F).

Skull robust, nearly oval, relatively small, GTL 13.73–14.11 mm. Rostrum long, deep, gradually ascending to forehead; prominent median depression present. Sagittal and lambdoid crests absent. In dorsal view, braincase domed; zygomatic arches weak and slender, gradually widening posteriorly, widest at the base of the zygomatic arches; posterior margin of skull slightly protruding; middle from snout to frontal region distinctly concave downward. In lateral view, skull slightly elongated, with elongated oval braincase; height gradually rising from snout to parietal, with slightly increasing slope from snout to frontal and decreasing slope from frontal to parietal; slight depression between snout and frontal, with slight distinct prominence at frontal; zygomatic arches gradually rising from anterior to posterior. In ventral view, palatine wide and nearly flat, ending at posterior margin of C1; basisphenoid pits slightly shallowly tadpole-shaped, extending posteriorly to anterior half of cochlea. Mandible length 8.85–9.22 mm, inverted L-shaped in lateral view. Line between coronoid process and condyle nearly flat; distinct inward depression between condyle and angle; angle short and wide; mandibular foramina clearly visible, situated below anterior margin of P4.

Dental morphology: Dental formula is I2/3, C1/1, P2/2, M3/3=34 (Fig. 14). In the maxilla, I2 is situated anterior to I3, and I2 clearly visible laterally; crown area of P2 approximately half that of P4 and slightly smaller than C1. Based on these characters, the species belongs to the “suilla-type”. Maxillary dentition converges slightly anteriorly (RCM = 0.64–0.66). I2 and I3 bicuspid, smaller secondary cusp situated posterior to primary cusp; I2 almost equal to I3 in height, with crown area approximately half that of I3; distinct gap between posterior surface of I3 and C1, not in contact, about half of height of C1. C1 smaller than P4 in height, slightly elongated and lacking secondary cusps, crown area 2/3 that of P4; P2 slightly smaller, delicate and pointed, about half as tall as P4 and C1, with crown area of P2 is half that of P4 and slightly smaller than C1. Mesostyles of M1 and M2 are reduced, but retaining distinct cusps; paracone, protocone, metacone, and parastyle well developed. M3 reduced, with only parastyle, paracone, and protocone. In the mandible, I1, I2, and I3 smaller, tricuspid, almost equal in height and width; overlap of outer cusps of I1, I2, and I3; with rapid increase in height from I3 to C1. C1 without pointed cusp on anterior inner margin, not in contact with I3 outer cusp. C1 almost equal to P4 in height and basal area, about twice as much as P2. P2 exceeded 4/5 of P4 in height, and the basal and crown areas are about 2/3 of P4. In lateral view, trigonids of M1, M2, and M3, metaconidhe slightly equal to protoconid in height, paraconid about 1/3 of protoconid. Talonid of M1 and M2 almost the same size as trigonid; entoconid and hypoconid distinctly separated from trigonid, lower than metaconid and paraconid, nearly equal to metaconid and paraconid in height. M1 and M2 are nyctalodont types, with well-developed entoconids. M3 reduced, talonid approximately 1/3 as long as trigonid, paraconid, protoconid, and metaconid complete and well developed.

Morphological comparisons with congeneric species.

Based on its dentition, I2 situated anterior to I3, and the crown area of P2 is less than half that of P4 and smaller than that of C1, Murina beibengensis sp. nov. belongs to the “suilla-type,” a character that distinguishes 12 species belonging to the “cyclotis-type,” including M. aenea, M. annamitica, M. cyclotis, M. fionae, M. guilleni, M. harrisoni, M. huttoni, M. peninsularis, M. pluvialis, M. puta, M. recondita, and M. rozendaali. Detailed morphological differences between the new species and congeners are listed in Suppl. material 5 and illustrated in Suppl. material 1: fig. S3.

Murina yadongensis sp. nov. can be distinguished from Murina beibengensis sp. nov. by dorsal fur brown-gold (vs. brown-gold), sagittal crest absent (vs. well-developed), I2 is situated laterally anterior to I3, and I2 is partially visible in the lateral view (vs. I2 is anterior to I3, I2 is clearly visible in the lateral view), I2 almost equal to I3 in height (vs. I2 taller than I3), and C1 taller than P4 in height (vs. C1 almost equal to P4).

Murina yadongensis sp. nov. can be distinguished from Murina medogensis sp. nov. by brown-gold dorsal fur (vs. dark grayish), dorsal hairs extend onto forearm and wrist (vs. hairless forearms and wrists), I2 is situated laterally anterior to I3, and I2 is partially visible in the lateral view (vs. I2 is anterior to I3, I2 is clearly visible in the lateral view), I2 almost equal to I3 in height (vs. I2 higher than I3), C1 taller than P4 (vs. C1 almost equal to P4), and an off-white circumferential band around the neck (vs. lacking); from Murina milinensis sp. nov. by lambdoid crest absent (vs. poorly developed), C1 taller than P4 (vs. C1 lower than P4), C1 taller than P4 (vs. C1 equal to P4), and an off-white circumferential band around the neck (vs. lacking).

Murina yadongensis sp. nov. can be distinguished from M. bicolor, M. fanjingshanensis, M. fusca, M. hilgendorfi, M. jinchui, M. leucogaster, and M. ryukyuana by the small size, forearm length 28.81–31.84 mm (vs. forearm length over 35 mm).

Murina yadongensis sp. nov. can be distinguished from M. aurata, M. balaensis, M. beelzebub, M. bicolor, M. chrysochaetes, M. eleryi, M. fanjingshanensis, M. feae, M. florium, M. hilgendorfi, M. hkakaboraziensis, M. jaintiana, M. jinchui, M. leucogaster, M. liboensis, M. lorelieae, M. rongjiangensis, M. ryukyuana, M. shuipuensis, M. suilla, M. tenebrosa, M. tubinaris, M. ussuriensis, M. walstoni, M. yuanyang, and M. yushuensis by an off-white circumferential band around the neck (vs. absent). By dorsal fur brown-gold (bicolored, black at the base, gradually transitioning to brown-gold tips from 2/3 from base), Murina yadongensis sp. nov. can be further distinguished from M. aurata (vs. golden brown), M. balaensis (vs. orange-reddish), M. chrysochaetes (vs. golden brown), M. eleryi (vs. coppery brown), M. feae (vs. dark grayish), M. jaintiana (vs. medium gray with brownish tinge), M. jinchui (vs. brownish gray), M. liboensis (vs. yellowish brown), M. lorelieae (vs. reddish brown), M. walstoni (vs. brownish gray), M. walstoni (vs. warm brown, whitish at the base, orangish brown at tips), and M. yuanyang (vs. bark gold). By ventral fur is grayish brown overall (bicolored, dark at the base, pale at tips), M. milinensis sp. nov. can be further distinguished from M. bicolor (vs. uniformly yellow), M. rongjiangensis (vs. bright yellowish orange), and M. shuipuensis (vs. bright orange yellow).

By the ears without smooth convex anterior margins and no notch on posterior margin, Murina yadongensis sp. nov. can be distinguished from M. gracilis, M. harpioloides, and M. kontumensis (vs. with smoothly convex anterior margins and distinct notch on posterior margin). By the lambdoid crest absent, I2 almost equal to I3 in height, C1 taller than P4, C1 equal to P4 in height and crown area, and Murina yadongensis sp. nov. can be further distinguished from M. gracilis (vs. lambdoid crest very weak), M. harpioloides (vs. I2 lower than I3 in height, C1 less than P4 in height, and C1 taller than P4), and M. kontumensis (vs. lambdoid crest present, and C1 larger than P4 in height and crown area).

Habitat and ecology.

The type locality experiences a subtropical semi-humid monsoon climate, characterized by mild and humid conditions. The average temperature in July is approximately 14.4 °C, and the mean annual precipitation is about 800 mm. The perennial influence of the monsoon, along with the mild and humid climate, contributes to the richness of forest resources in the southern part of the county. The specimen was captured in a harp trap on 6 August 2023 in a mixed coniferous forest in Xiayadong Township, Yadong County. The area is surrounded by dense thickets and bamboo forests, with rivers and primary forests situated approximately 3 kilometers away. In this forest, we also collected five species from four genera: Plecotus homochrous, Rhinolophus sinicus, Rhinolophus ferrumequinum, Myotis sp., and Submyotodon moupinensis.

Murina rubella Thomas, 1914

Figs 15, 16, 17, Table 5, Suppl. material 2

Chresonymy.

Murina huttoni rubella: Zhang 1997 (Guangxi and Fujian, China); Wang 2003 (Fujian, Jiangxi, and Guangxi, China); Csorba et al. 2007 (China); Francis and Eger 2012 (Fokien, China); Jiang 2015, 2021 (Fujian, Jiangxi, and Guangxi, China); Zhang et al. 2016 (Jinggangshan National Natural Reserve, Jiangxi, China); Moratelli et al. 2019 (Hunan, Jiangxi, Fujian, Guangdong, and Guangxi, China; Myanmar, Thailand, Laos, Vietnam, Peninsular Malaysia). Murina huttoni: Zhou et al. 2011 (Guangdong, China); Jiang 2015, 2021 (Fujian, Jiangxi, and Guangxi, China); Huang et al. 2018 (Huibei, Zhejiang, China); Wei 2022 (Xizang, Hubei, Fujian, Jiangxi, Zhejiang, Guangdong, Guangxi, China); Wei et al. 2022 (China); Qin et al. 2023 (Anhui, China); and Liao et al. 2023 (Hunan, China).

Holotype.

BMNH 1908.8.11.6., adult male.

Type locality.

Kuatun, Fokien, China (Kuatun, Wuyishan City, Fujian Province, China).

Measurements (in mm) holotype.

FL: 37.50, GTL: 18.2 (Thomas 1914).

Material examined.

• Four male specimens (XZ202387, XZ202399, XZ2023102, and XZ2023104) and three female specimens (XZ2024041, XZ2023101, and XZ2023103) were collected from Xiachayu Town, Zayu County, Nyingchi City, Xizang Autonomous Region, China. • One female specimen (GS20240031, Figu.15) was collected from Yaodu Town, Wen County, Longnan City, Gansu Province, China.

Etymology.

The specific epithet “rubella” comes from the Latin word rubellus, meaning “slightly reddish” or “reddish”. The common English name “Fujian Tube-nosed Bat” and the Chinese name “Fú Jiàn Guǎn Bí Fú (福建管鼻蝠)”.

Diagnosis.

Murina rubella can be distinguished from all of the other congeners by the following combination of characters: (1) larger body size, FL 34.24–38.45 mm, GTL 17.08–18.51 mm; (2) dorsal fur tan overall, pale brown at the base, gradually transitioning to dark tan tips from 2/3 from base; (3) ventral fur uniformly pale brown overall, black at the base, gray-white at tips; (4) ears narrow and oval, with smoothly convex anterior margins, no notch on posterior margins; (5) forearm and wrists covered with sparse hairs; (6) wing attachment point located at 1/3 from base of claw to base of toe; (7) without off-white circumferential band around the neck; (8) sagittal and lambdoid crests absent; (9) I2 is situated anterior to I3 and clearly visible in lateral view, I2 slightly smaller than I3 in height; (10) mesostyles of M1 and M2 are slightly reduced; (11) C1 slightly smaller than P4 in height, crown is about 2/3 of P4; (12) P2 approximately half of P4 in height, crown area 2/3 that of P4; (13) C1 equal to P4 in height and crown area.

Redescription.

Morphometric data of M. rubella are provided in Table 5 and Suppl. material 2. Medium-size Murina, HB 37.29–48.73 mm, FL 34.24–38.45 mm, EL 16.15–17.68 mm, HFL 7.58–9.49 mm, and BW 4.00–21.00 g. Nostrils tubular, open sideways, and short. Ears slightly large, short, and broadly rounded, blunt at tips, with smoothly convex anterior margins, not notched on posterior margin. Tragus short, narrow, and tapering toward pointed tip, with slightly convex anterior margin, concave posterior margin, and basal notch, and it curves outwards slightly, about half as long as ear. Body covered with thick and fluffy hair. Dorsal hairs tan, similar to rust color (pale brown at the base, gradually transitioning to dark tan tips from 2/3 from base). Dorsal hairs extend onto bases of wings, uropatagium, thumbs, forearm, and feet, with slight-developed fringe of hairs around margin of uropatagium. Densely furred anterior 1/3 of the dorsal uropatagium, posterior 2/3 covered with sparse hairs. Ventral hairs uniformly pale brown overall. Without off-white circumferential band around the neck. Dark reddish-brown around the eyes, muzzle, and lower forehead, and the face is hairy except for the long, protuberant nostrils that are naked. TL 36.53–40.69 mm, slightly shorter than head-body length, tip of the tail extending significantly past the rear edge of the uropatagium, tip slight free. Plagiopatagium attachment point located at 1/3 from base of claw to base of toe, near base of claw (Fig. 15G).

Table 5.

External and craniodental measurements (in mm) of M. rubella and a closely related species. NA indicates that the data is not available.

Species M. rubella M. huttoni (Chakravarty et al. 2020) M. puta (Moratelli et al. 2019)
Male (n = 4) Female (n = 4) Range Range
BW 4.00–21.00 (12.25 ± 9.54) 6.00–19.00 (10.10 ± 6.08) 8–11 5.4–9.0
HB 37.29–39.26 (38.43 ± 0.82) 42.57–48.73 (44.48 ± 2.88) NA 59.0–61.0
TL 36.54–40.53 (38.25 ± 1.71) 36.53–40.69 (38.51 ± 1.93) 31.0–37.9 32.0–36.0
EL 16.15–17.68 (17.03 ± 0.66) 16.89–17.51 (17.09 ± 0.29) 14.9–20.9 17.0–19.0
EW 9.09–9.67 (9.33 ± 0.25) 8.14–10.67 (9.51 ± 1.04) NA NA
TRL 8.66–9.96 (9.08 ± 0.61) 6.80–9.76 (8.22 ± 1.22) NA NA
TRW 2.40–2.61 (2.49 ± 0.09) 1.84–2.63 (2.31 ± 0.35) NA NA
HFL 8.27–9.49 (8.83 ± 0.61) 7.58–8.96 (8.47 ± 0.62) 8.0–10.0 10.0–11.0
FL 34.24–36.07 (35.59 ± 0.90) 36.30–38.45 (37.04 ± 1.01) 34.4–36.5 33.0–39.0
TIB 16.98–17.68 (17.37 ± 0.29) 16.55–17.74 (17.28 ± 0.55) 13.0–17.93 NA
GTL 17.08–18.28 (17.61 ± 0.51) 17.13–18.51 (17.82 ± 0.59) NA NA
CCL 15.45–16.54 (16.20 ± 0.50) 15.94–17.15 (16.68 ± 0.53) NA NA
BCW 7.65–8.07 (7.82 ± 0.20) 7.37–8.21 (7.88 ± 0.37) NA NA
BCH 6.94–7.76 (7.44 ± 0.36) 6.70–7.53 (7.19 ± 0.38) NA NA
ZYW 9.05–9.76 (9.47 ± 0.35) 9.71–10.20 (9.95 ± 0.25) NA NA
MAW 7.64–8.54 (8.02 ± 0.42) 8.21–8.66 (8.37 ± 0.20) NA NA
IOW 4.40–4.61 (4.52 ± 0.10) 4.56–4.99 (4.68 ± 0.21) NA NA
CM3L 5.68–6.03 (5.91 ± 0.17) 4.02–6.12 (5.46 ± 0.97) NA NA
C1C1W 2.81–2.98 (2.87 ± 0.08) 2.93–4.19 (3.55 ± 0.62) NA NA
M3M3W 3.73–4.47 (4.03 ± 0.32) 4.23–6.02 (5.13 ± 0.95) NA NA
RCM 0.63–0.80 (0.72 ± 0.07) 0.66–0.74 (0.69 ± 0.04) NA NA
CM3L 6.61–7.12 (6.83 ± 0.21) 4.63–6.99 (6.17 ± 1.05) NA NA
ML 11.03–12.49 (11.46 ± 0.69) 11.43–12.40 (11.85 ± 0.45) NA NA
MDL 11.53–14.42 (12.29 ± 1.42) 11.93–12.90 (12.44 ± 0.42) NA NA
CPH 4.17–4.61 (4.39 ± 0.18) 4.14–4.64 (4.45 ± 0.22) NA NA
Figure 15. 

External morphological characteristics of the M. rubella in living specimen, GS2024031. A. Lateral view; B. Ventral view; C. Dorsal view of the body and uropatagium; D. Ventral hairs; E. Dorsal hairs; F. Hindfoot ventral view; G. Hindfoot dorsal view.

Skull robust, nearly oval, relatively small, GTL 17.08–18.51 mm. Rostrum long, deep, gradually ascending to forehead; prominent median depression present. Sagittal and lambdoid crests absent. In dorsal view, braincase domed; zygomatic arches weak and slender, gradually widening posteriorly, widest at the base of the zygomatic arches; posterior margin of skull slightly protruding; middle from snout to frontal region distinctly concave downward. In lateral view, skull slightly elongated, with elongated oval braincase; height gradually rising from snout to parietal, with slightly increasing slope from snout to frontal and decreasing slope from frontal to parietal; slight depression between snout and frontal, with distinct prominence at frontal; zygomatic arches gradually rising from anterior to posterior. In ventral view, palatine wide and nearly flat, ending at posterior margin of C1; basisphenoid pits slightly shallowly teardrop-shaped, extending posteriorly to anterior half of cochlea. Mandible length 11.03–12.49 mm, inverted L-shaped in lateral view. Line between coronoid process and condyle nearly flat; distinct inward depression between condyle and angle; angle short and wide; mandibular foramina clearly visible, situated below anterior margin of P2.

Dental morphology: Dental formula is I2/3, C1/1, P2/2, M3/3=34 (Fig. 16). In the maxilla, I2 is situated anterior to I3, and I2 clearly visible laterally; crown area of P2 approximately half that of P4 and slightly smaller than C1. Based on these characters, the M. rubella belongs to the “suilla-type”. Maxillary dentition converges slightly anteriorly (RCM = 0.63–0.80). I2 and I3 bicuspid, smaller secondary cusp situated posterior to primary cusp; I2 slightly smaller than I3 in height, crown area of I2 2/3 that of I3; distinct gap between posterior surface of I3 and C1, not in contact, about half of height of C1. C1 slightly smaller than P4 in height, slightly elongated and lacking secondary cusps, crown area 2/3 that of P4. P2 smaller, delicate and pointed, about half of P4 and 2/3 of C1 in height, and crown area of P2 is half that of P4 and slightly smaller than C1. Mesostyles of M1 and M2 are reduced, but retaining distinct cusps; paracone, protocone, metacone, and parastyle well developed. M3 reduced, with only parastyle, paracone, and protocone. In the mandible, I1, I2, and I3 smaller, tricuspid, almost equal in height and width; slight overlap of outer cusps of I1, I2, and I3; with gradual increase in height from I1 to C1. C1 without pointed cusp on anterior inner margin, in contact with I3 outer cusp. C1 taller than P4 and equal in basal area to P4. P2 exceeded four-fifths of P4 in height, and the basal and crown areas are about 2/3 of P4. In lateral view, trigonids of M1, M2, and M3, metaconidhe slightly equal to protoconid in height, paraconid about 1/3 of protoconid. Talonid of M1 and M2 almost the same size as trigonid; entoconid and hypoconid distinctly separated from trigonid, lower than metaconid and paraconid, nearly equal to metaconid and paraconid in height. M1 and M2 are nyctalodont types, with well-developed entoconids. M3 reduced, talonid approximately 1/3 as long as trigonid, paraconid, protoconid, and metaconid complete and well developed.

Figure 16. 

Skull and dentition of the M. rubella. A. Lateral view; B. Dorsal view; C. Ventral view; D. Frontal view of mandible.

Figure 17. 

External morphological characteristics of the M. rubella in living specimen, XZ2024041. A. Lateral view; B. Ventral view; C. Dorsal view of the body and uropatagium; D. Ventral hairs; E. Dorsal hairs; F. Hindfoot ventral view; G. Hindfoot dorsal view.

Morphological comparisons with congeneric species.

Based on its dentition, I2 is situated anterior to I3 and crown area of P2 approximately half that of P4 and slightly smaller than C1, M. rubella belongs to the “suilla-type”, a character that distinguishes 14 species belonging to the “cyclotis-type”, including M. aenea, M. annamitica, M. cyclotis, M. fionae, M. guilleni, M. harrisoni, M. huttoni, M. peninsularis, M. pluvialis, M. puta, M. recondita, and M. rozendaali. Detailed morphological differences between the M. rubella and congeners are shown in Suppl. material 5 and Suppl. material 1: fig. S3.

Murina rubella can be distinguished from M. beibengensis sp. nov. by the dorsal hairs dark tan (vs. orangish-yellow overall), from M. medogensis sp. nov. (vs. dark grayish), from M. yadongensis sp. nov. (vs. brown-gold), M. milinensis sp. nov. (vs. brown-gold), from M. aurata (vs. golden brown with golden tips), from M. balaensis (vs. golden orangish brown, with orange reddish brown tips), from M. harpioloides (vs. orangish brown with orange gold tips), from M. hilgendorfi (vs. silvery brownish gray), from M. jaintiana (vs. medium gray with brownish tinge), from M. jinchui (vs. brownish gray), from M. liboensis (vs. yellowish brown), from M. shuipuensis (vs. golden grayish brown), from M. suilla (vs. orangish brown), from M. tubinaris (vs. light grayish brown), and from M. yushuensis (vs. brown-gold).

Murina rubella can be distinguished from M. chrysochaetes, M. gracilis, M. kontumensis, and M. yuanyang by without off-white circumferential band around the neck (vs. absent). By the ears with smooth convex anterior margins, but without notch on posterior margin, M. rubella can be distinguished from M. beelzebub, M. bicolor, M. eleryi, M. fanjingshanensis, M. hkakaboraziensis, M. leucogaster, and M. rongjiangensis (vs. with distinct notch on posterior margin).

By I2 less than I3 in height, M. rubella can be distinguished from M. lorelieae (vs. I2 higher than I3), M. ryukyuana (vs. I2 equal to I3), and M. florium (vs. I2 higher than I3). Murina rubella different from M. walstoni by sagittal and lambdoid crests absent (vs. present) and ventral hairs uniformly pale brown (vs. pure white); from M. tenebrosa and M. florium by ventral hairs uniformly pale brown (vs. paler).

Habitat and ecology.

Specimen GS2023031 was captured on 23 April 2024, using a harp trap in a mixed coniferous and broadleaf forest located at the border between Longnan City, Gansu Province, and Guangyuan City, Sichuan Province, China. The area, situated in the western part of the Qinling Mountains, is surrounded by small-scale agricultural land and tea plantations. Rich in forest resources, the region has a subtropical, mild, and humid climate, with a multi-year average temperature of 16.5 °C and an average annual precipitation of approximately 600 mm. Only one specimen was captured during the survey, and no other bats were observed or captured in subsequent surveys. We speculate that the bats in this area may have been in a post-hibernation stage and were not yet active.

Remark.

Murina rubella was described on the basis of specimens from the Guadun, Fujian, China, as a subspecies of M. huttoni and is widely accepted (Moratelli et al. 2019). Subsequently, M. huttoni has been widely recorded from southern provinces of China, including Xizang, Hubei, Fujian, Jiangxi, Zhejiang, Guangdong, Guangxi, Hunan, and Anhui (Zhang 1997; Wang 2003; Zhou et al. 2011; Jiang 2015; Huang et al. 2018; Jiang 2021; Liao et al. 2023; Qin et al. 2023; Wei et al. 2025). Recently, mitochondrial markers of the near-type locality of M. h. rubella and M. huttoni were published (Zhang et al. 2016; Chakravarty et al. 2020), offering the possibility of resolving the phylogenetic relationships of these two species. In our phylogenetic tree reconstructed on the basis of mitochondrial COI, the M. huttoni complex from different regions formed seven clades: Clade i included M. huttoni from the near type locality; Clade ii included the species M. puta; Clade iii was from central Vietnam; and Clade iv was from southern China (Guangdong, Hunan, Jiangxi), including the near-type locality M. rubella; Clades v and vi are from Xizang, China; and Clade vii is from China (Hunan), Vietnam, and central Laos. From the results of the mPTP species delimitation, Clades i, ii, and iii-vii are proposed as separate species, with genetic distances between them ranging from 6.2% to 6.9%. Based on this evidence, and the validity of M. huttoni and M. puta is recognized, then the proposal to elevate the subspecies M. h. rubella in China to species status should be accepted. For M. huttoni, we speculate that its distribution in China may be limited to the territory of Xizang Autonomous Region, along the Himalaya.

Geographic distribution.

Based on this study, phylogenetic evidence, and previous literature, M. rubella can be definitively recorded as distributed in China (Anhui, Fujian, Guangxi, Guangdong, Hubei, Hunan, Jiangxi, Zhejiang, Xizang) (Zhang 1997; Wang 2003; Francis et al. 2010; Zhou et al. 2011; Jiang 2015; Zhang et al. 2016; Huang et al. 2018; Moratelli et al. 2019; Yu et al. 2020; Jiang 2021; Liao et al. 2023; Qin et al. 2023), northern Laos (Francis et al. 2010; Francis and Eger 2012; Nguyen et al. 2015b), and Vietnam (Francis and Eger 2012; Tu et al. 2015; Tu et al. 2016). Although M. huttoni has also been recorded in northern Thailand, northern Myanmar, and Peninsular Malaysia (Moratelli et al. 2019), the authenticity of these records (Francis and Eger 2012) requires further evidence due to the lack of available genetic data. The M. huttoni sensu stricto may be confined to the periphery of the Himalayas (Moratelli et al. 2019).

Discussion

Asian tube-nosed bats, genus Murina, are the most frequently described taxon among bats in China in recent years, and seven species have been described, i.e., M. fanjingshanensis (He et al. 2015), M. rongjiangensis (Chen et al. 2017), M. liboensis (Zeng et al. 2018), M. jinchui (Yu et al. 2020), M. yuanyang (Mou et al. 2024), M. yushuensis (Wang et al. 2024a), and M. lvchun (Mou et al. 2025). We describe here four new species, named Murina beibengensis sp. nov., Murina medogensis sp. nov., Murina milinensis sp. nov., and Murina yadongensis sp. nov., based on fieldwork in Xizang, China, in the last three years, combined with morphological (refer to previous species descriptions), mitochondrial COI, and Cyt b. Morphologically, these four new species can be distinguished from conspecifics based on differences in body fur color, forearm length, dentition, and skull (Suppl. material 1: table S9). In the phylogeny, the four new species formed distinct lineages, with mitochondrial COI and Cyt b divergences from congeners ranging from 7.0% to 20.3% and 6.1% to 17.2%, respectively. These divergences are significantly greater than those observed among currently recognized species (Suppl. material 1: tables S3, S4). Thus, morphological and genetic evidence supports the validity of these four new species. The discovery of these four new species indicates that the diversity of the genus Murina is significantly underestimated, particularly in the Himalayan and Qinghai-Tibetan Plateau regions. This finding underscores the importance and urgency of conducting comprehensive surveys in these areas.

Due to the sampling constraints in evergreen broad-leaved forests where only harp traps can be effectively deployed, we obtained only one to two specimens each of Murina beibengensis sp. nov. and Murina medogensis sp. nov. Despite the limited sample size, multiple lines of evidence support their validity as distinct species. Similar situations with small type series, e.g., M. lvchun with two specimens (Mou et al. 2025), M. yushuensis with one specimen (Wang et al. 2024a), and M. hkakaboraziensis with one specimen (Soisook et al. 2017), are not uncommon in chiropteran taxonomy, particularly for rare forest-dwelling species. While we acknowledge that future collections may expand the known morphological variation ranges, the current diagnostic characters, including both discrete morphological features and genetic divergence, provide robust support for their taxonomic recognition. The congruence between morphological distinctiveness and genetic differentiation observed in our analyses strongly suggests these represent valid species rather than intraspecific variants.

Most species of the genus Murina have been found in montane forests at elevations ranging from 110 m to 2600 m (Moratelli et al. 2019) with the exception of M. yushuensis (3770 m) (Wang et al. 2024a). The four new species described here are distributed across low to mid-altitude regions of the Himalayas, though they may also inhabit higher elevations. For example, Murina beibengensis sp. nov. has been recorded at elevations ranging from 864 m to 4414 m. This remarkable altitudinal variation suggests that Murina beibengensis sp. nov. exhibits strong locomotor capabilities and may possess unique adaptations to cope with rapid environmental changes during flight, such as high temperatures and low oxygen levels. Future genomic studies of this species will provide valuable insights into the genetic mechanisms underlying its adaptation to high-altitude environments. This study describes four new species of Murina and elevates the subspecies M. huttoni rubella to species rank, increasing the total number of recognized species in the genus from 43 to 48. Given the low dispersal capability and relatively small natural population densities characteristic of Murina bats, additional conservation measures are warranted to ensure their long-term survival.

Acknowledgments

This study was supported by the program of the Survey of Wildlife Resources in Key Areas of Tibet (Phase II), Survey of Wildlife Resources in Key Areas of Tibet (ZL202203601), and Diversity and Distribution Survey of Chiroptera species in China (2021FY100302). We thank Xin-Rui Zhao, Peng-Fei Luo, Qing-Qing He, and Qin Yang for their help during the sample collection. We thank Dr. Jing-Song Shi for his help with the CT scan of the bones.

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Tao Luo and Ming-Le Mao contributed equally to this paper.

Supplementary materials

Supplementary material 1 

Additional figures and tables

Tao Luo, Ming-Le Mao, Chang-Ting Lan, Zi-Fa Zhao, Zhong-Lian Wang, Jing Yu, Jia-Jia Wang, Chen-Rui Yan, Ning Xiao, Jiang Zhou

Data type: pdf

Explanation note: fig. S1. For the genus Murina, a phylogenetic tree (ML) was reconstructed based on the mitochondrial COI gene. fig. S2. For the genus Murina, a phylogenetic tree (ML) was reconstructed based on the mitochondrial Cyt b gene. fig. S3. External morphology and skull features of species of the genus Murina, from literature and web pages. table S2. For the genus Murina, locality, voucher information, and mitochondrial COI GenBank accession numbers are provided for all samples used. table S3. For the genus Murina, locality, voucher information, and mitochondrial Cyt b GenBank accession numbers are provided for all samples used. table S4. For the M. huttoni complex, locality, voucher information, and mitochondrial COI GenBank accession numbers are provided for all samples used. table S7. Uncorrected p-distance (%) between species of the M. huttoni complex based on mitochondrial COI. table S8. Results and percentage of variance explained by principal component analysis for M. beibengensis sp. nov. and three closely related species. table S9. Results and percentage of variance explained by principal component analysis for M. medogensis sp. nov. and three closely related species. table S10. Results and percentage of variance explained by principal component analysis for M. milinensis sp. nov., M. yadongensis sp. nov., and four closely related species.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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Supplementary material 2 

Raw measurements of species were measured and collected in this study

Tao Luo, Ming-Le Mao, Chang-Ting Lan, Zi-Fa Zhao, Zhong-Lian Wang, Jing Yu, Jia-Jia Wang, Chen-Rui Yan, Ning Xiao, Jiang Zhou

Data type: xlsx

Explanation note: For closely related species, mean values were also considered.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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Supplementary material 3 

Uncorrected p-distance (%) between species of the genus Murina based on mitochondrial COI

Tao Luo, Ming-Le Mao, Chang-Ting Lan, Zi-Fa Zhao, Zhong-Lian Wang, Jing Yu, Jia-Jia Wang, Chen-Rui Yan, Ning Xiao, Jiang Zhou

Data type: xlsx

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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Supplementary material 4 

Uncorrected p-distance (%) between species of the genus Murina based on mitochondrial Cyt b

Tao Luo, Ming-Le Mao, Chang-Ting Lan, Zi-Fa Zhao, Zhong-Lian Wang, Jing Yu, Jia-Jia Wang, Chen-Rui Yan, Ning Xiao, Jiang Zhou

Data type: xlsx

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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Supplementary material 5 

Comparison of the diagnostic features of the four new species described here with those selected for the 43 recognized species of the genus Murina

Tao Luo, Ming-Le Mao, Chang-Ting Lan, Zi-Fa Zhao, Zhong-Lian Wang, Jing Yu, Jia-Jia Wang, Chen-Rui Yan, Ning Xiao, Jiang Zhou

Data type: xlsx

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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