Research Article |
Corresponding author: Zhao-Chi Zeng ( zhaochizeng@outlook.com ) Corresponding author: Jian Wang ( wangj-1994@outlook.com ) Academic editor: Anthony Herrel
© 2024 Shi-Shi Lin, Hong-Hui Chen, Yuan-Hang Li, Zhao-Ning Peng, Zhao-Chi Zeng, Jian Wang.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Lin S-S, Chen H-H, Li Y-H, Peng Z-N, Zeng Z-C, Wang J (2024) A new Boulenophrys species (Anura, Megophryidae) from the coastal hills of eastern Fujian Province, China. ZooKeys 1216: 1-15. https://doi.org/10.3897/zookeys.1216.130017
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A new species of the genus Boulenophrys is described from the coastal hills of eastern Fujian Province, China. The new taxon can be distinguished from all recognized congeners by a combination of discrete morphological character state differences and genetic divergences in the combined mitochondrial 16S + CO1 genes. We also provide a map showing the distribution pattern of Boulenophrys species in Fujian and a provincial-specific key, which will aid their conservation by helping the local authorities accurately identify species during field identifications and data collection efforts.
Boulenophrys lichun sp. nov., conservation actions, distribution pattern, diversity, Horned Toads, identification, new species, provincial key, taxonomy
The Chinese Horned Toads (Boulenophrys Fei, Ye & Jiang, 2016) comprise 68 recognized species which are classified within the subfamily Megophryinae (Bonaparte, 1850) (
During recent field surveys in eastern Fujian, we collected a series of Boulenophrys specimens (Fig.
All examined specimens were fixed in 10% buffered formalin and later transferred to 70% ethanol. All studied specimens have been deposited at the
Guangdong Polytechnic of Environmental Protection Engineering (GEP), Foshan City, Guangdong, and the
Herpetological Museum, Chengdu Institute of Biology, the Chinese Academy of Sciences (
External measurements were recorded with a digital caliper (Neiko 01407A stainless steel 6-inch digital caliper) to the nearest 0.1 mm. These measurements are as follows:
SVL (snout–vent length, from tip of snout to posterior margin of vent);
HDL (head length, from tip of snout to the articulation of the jaw);
HDW (head width, head width at the commissure of the jaws);
SNT (snout length, from tip of snout to the anterior corner of the eye);
IND (internasal distance, distance between nares);
IOD (interorbital distance, minimum distance between upper eyelids);
ED (eye diameter, from the anterior corner of the eye to posterior corner of the eye);
TD (tympanum diameter, horizontal diameter of tympanum);
TED (tympanum–eye distance, from anterior edge of tympanum to posterior corner of the eye);
HND (hand length, from the proximal border of the outer palmar tubercle to the tip of digit III);
RAD (forearm or radiulna length, from the flexed elbow to the proximal border of the outer palmar tubercle);
FTL(foot length, from distal end of shank to the tip of digit IV);
TIB (crus or tibiofibula length, from the outer surface of the flexed knee to the heel).
Sex was determined by external secondary sexual characters, such as the presence of vocal sacs, nuptial pads, or spines in males and their absence in females (
Morphological characters of all 68 recognized species of the genus Boulenophrys used for comparisons were based on information available in the literature (Table
Literature for morphological characters of 68 recognized species of Boulenophrys.
Boulenophrys species | References |
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B. acuta (Wang, Li & Jin, 2014) |
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B. angka (Wu, Suwannapoom, Poyarkov, Pawangkhanant, Xu, Jin, Murphy & Che, 2019) |
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B. anlongensis (Li, Lu, Liu & Wang, 2020) |
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B. baishanzuensis (Wu, Li, Liu, Wang & Wu, 2020) |
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B. baolongensis (Ye, Fei & Xie, 2007) |
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B. binchuanensis (Ye & Fei, 1995) |
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B. binlingensis (Jiang, Fei & Ye, 2009) |
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B. boettgeri (Boulenger, 1899) |
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B. brachykolos (Inger & Romer, 1961) |
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B. caobangensis (Nguyen, Pham, Nguyen, Luong & Ziegler, 2020) |
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B. caudoprocta (Shen, 1994) |
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B. congjiangensis (Luo, Wang, Wang, Lu, Wang, Deng & Zhou, 2021) |
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B. cheni (Wang & Liu, 2014) |
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B. chishuiensis (Xu, Li, Liu, Wei & Wang, 2020) |
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B. daiyunensis (Lyu, Wang & Wang, 2021) |
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B. daoji (Lyu, Zeng, Wang & Wang, 2021) |
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B. daweimontis (Rao & Yang, 1997) |
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B. dongguanensis (Wang & Wang, 2019) |
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B. elongata Zeng, Wang, Chen, Xiao, Zhan, Li & Lin, 2024 |
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B. fengshunensis Wang, Zeng, Lyu & Wang, 2022 |
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B. fanjingmontis (Zhang, Liang, Ran & Shen, 2012) |
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B. fansipanensis (Tapley, Cutajar, Mahony, Nguyen, Dau, Luong, Le, Nguyen, Nguyen, Portway, Luong & Rowley, 2018) |
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B. frigida (Tapley, Cutaja, Nguyen, Portway, Mahony, Nguyen, Harding, Luong & Rowley, 2021) |
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B. hoanglienensis (Tapley, Cutajar, Mahony, Nguyen, Dau, Luong, Le, Nguyen, Nguyen, Portway, Luong & Rowley, 2018) |
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B. hungtai Wang, Zeng, Lyu, Xiao & Wang, 2022 |
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B. hengshanensis Qian, Hu, Mo, Gao, Zhang & Yang, 2023 |
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B. insularis (Wang, Liu, Lyu, Zeng & Wang, 2017) |
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B. jiangi (Liu, Li, Wei, Xu, Cheng, Wang & Wu, 2020) |
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B. jingdongensis (Fei & Ye, 1983) |
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B. jinggangensis (Wang, 2012) |
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B. jiulianensis (Wang, Zeng, Lyu & Wang, 2019) |
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B. kuatunensis (Pope, 1929) |
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B. leishanensis (Li, Xu, Liu, Jiang, Wei & Wang, 2018) |
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B. lushuiensis (Shi, Li, Zhu, Jiang, Jiang & Wang, 2021) |
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B. liboensis (Zhang, Li, Xiao, Li, Pan, Wang, Zhang & Zhou, 2017) |
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B. lini (Wang & Yang, 2014) |
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B. lishuiensis (Wang, Liu & Jiang, 2017) |
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B. minor (Stejneger, 1926) |
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B. mirabilis (Lyu, Wang & Zhao, 2020) |
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B. mufumontana (Wang, Lyu & Wang, 2019) |
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B. nankunensis (Wang, Zeng & Wang, 2019) |
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B. nanlingensis (Lyu, Wang, Liu & Wang, 2019) |
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B. obesa (Wang, Li & Zhao, 2014) |
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B. ombrophila (Messenger & Dahn, 2019) |
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B. omeimontis (Liu, 1950) |
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B. palpebralespinosa (Bourret, 1937) |
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B. pepe (Wang & Zeng, 2024) |
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B. puningensis Wang, Zeng, Lyu, Xiao & Wang, 2022 |
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B. qianbeinsis (Su, Shi, Wu, Li, Yao, Wang & Li, 2020) |
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B. rubrimera (Tapley, Cutajar, Mahony, Chung, Dau, Nguyen, Luong & Rowley, 2017) |
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B. sangzhiensis (Jiang, Ye & Fei, 2008) |
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B. sanmingensis (Lyu & Wang, 2021) |
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B. shimentaina (Lyu, Liu & Wang, 2020) |
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B. shuichengensis (Tian & Sun, 1995) |
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B. shunhuangensis (Wang, Deng, Liu, Wu & Liu, 2019) |
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B. spinata (Liu & Hu, 1973) |
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B. tongboensis (Wang & Lyu, 2021) |
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B. tuberogranulatus (Shen, Mo & Li, 2010) |
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B. wugongensis (Wang, Lyu & Wang, 2019) |
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B. wuliangshanensis (Ye & Fei, 1995) |
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B. wushanensis (Ye & Fei, 1995) |
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B. xiangnanensis (Lyu, Zeng & Wang, 2020) |
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B. xianjuensis (Wang, Wu, Peng, Shi, Lu & Wu, 2020) |
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B. xuefengmontis Lyu & Wang, 2023 |
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B. yangmingensis (Lyu, Zeng & Wang, 2020) |
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B. yaoshanensis Qi, Mo, Lyu, Wang & Wang, 2021 |
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B. yingdeensis Qi, Lyu, Wang & Wang, 2021 |
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B. yunkaiensis Qi, Wang, Lyu & Wang, 2021 |
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We use two partial mitochondrial genes, the 16S ribosomal RNA (16S) and the cytochrome c oxidase 1 (COI), for phylogenetic analysis. DNA extraction, PCR amplification, and sequencing protocols follow that of
We used Clustal X 2.0 (
The BI and ML phylogenetic trees resulted in essentially identical topologies, with the ML phylogenetic tree shown in Fig.
Holotype. China • ♂; Fujian Province, Ningde City, Jiaocheng District, Mt Nanji; 26.645774°N, 119.519939°E, ca. 230 m elev.; 4 Feb. 2024; Jian Wang, Zhao-Chi Zeng, Shi-Shi Lin and Yuan-Hang Li leg.; GEP a214.
Paratypes.
China • 4♂♂; same data as for holotype;
The specific name lichun is derived from Chinese Pinyin Lì Chūn, i.e. 立春 in Chinese, which means the beginning of spring, the first of the 24 solar terms (24节气) of China. The specific name refers to the breeding season of the new species which begins around this period. The song of the new species heralds the spring of a year. The type specimens of the new species were also collected on “Lichun” of the Year 2024.
(1) small size (SVL 33.5–37.0 mm in five adult males, SVL 47.1 mm in a single adult female); (2) canthus rostralis well developed, tongue not notched posteriorly; (3) tympanum distinct; (4) vomerine ridges and vomerine teeth present; (5) dorsal skin rough and highly granular, discontinuous X-shaped ridge on center of dorsum, discontinuous dorsolateral ridges present, sparse large tubercles on flanks, dorsal limbs with discontinuous transverse ridges and tubercles, ventral skin with dense raised tubercles; (6) outer margin of upper eyelid with a small horn-like prominent tubercle, supratympanic fold distinct and narrow, curving posteroventrally to above arm; (7) two metacarpal tubercles distinct, inner one observably enlarged; relative finger lengths I < II < IV < III; distinct subarticular tubercle at base of each finger; (8) heels not meeting when hindlimbs folded; tibio-tarsal articulation reaching shoulder to posterior corner of eye; (9) toes without webbing and lateral fringes, inner metatarsal tubercle long ovoid, outer one absent, relative toe length I < II < V < III < IV; (10) dorsal surface yellowish-brown with irregular dark-brown patches, and dark-brown triangular marking between eyes, dorsal limbs and digits light brown with dark-brown transverse bands; and (11) dense nuptial spines on dorsal bases of fingers I and II in breeding adult males, subgular vocal sac present in males.
Adult male. Body size small, SVL 37.0 mm. Head width larger than head length, HWD/HDL 1.04; snout rounded in dorsal view, projecting, sloping backward to mouth in profile, protruding well beyond margin of lower jaw; top of head flat; eyes moderate in size, ED 0.34 of HDL, pupil vertical, near diamond-shaped; nostril obliquely ovoid; canthus rostralis well developed; loreal region slightly oblique; internasal distance slightly larger than interorbital distance; tympanic region oblique, tympanum distinct and visible in dorsal view; tympanum moderate in size, margin clear, upper margin in contact with supratympanic fold, lower margin in contact with upper lip, TD/ED 0.55; large ovoid choanae at base of maxilla; vomerine ridge and vomerine teeth present, maxillary teeth present; margin of tongue rounded, not notched distally; presence of single subgular vocal sac.
Forearm length 0.23 of SVL, hand 0.24 of SVL; webbing absent between fingers, lateral fringes absent, relative finger length I < II < IV < III; tips of fingers slightly dilated, round; subarticular tubercles on base of fingers present, distinct; inner metacarpal tubercle observably enlarged, outer one slightly smaller; single nuptial pad bearing nuptial spines present on dorsal surface of first and second fingers, respectively. Hindlimbs short, tibio-tarsal articulation reaching forward to posterior conner of eye when hindlimb stretched along body; heels not meeting when flexed hindlimbs held at right angles to body axis; crus length 0.40 of SVL and foot length 0.58 of SVL; relative toe length I < II < V < III < IV; tips of toes round and slightly dilated; toes without lateral fringes and webbing; subarticular tubercles on base of toes present and distinct; inner metatarsal tubercle long ovoid and lacking outer metatarsal tubercle.
Dorsal skin rough and highly granular; dense large tubercles on flanks; single horn-like prominent tubercle on edge of upper eyelid; obvious supratympanic fold curving posteroventrally from posterior corner of eye to level above insertion of arm; upper lip, mandibular articulation, loreal, temporal region excluding tympanum, upper eyelid and surface around cloaca with conical tubercles; discontinuous X-shaped ridge on center of dorsum, discontinuous dorsolateral ridges present; ventral surface with dense raised tubercles; tubercles on ventral hindlimbs and around cloaca bearing tiny spines on their tips; small and distinct pectoral gland closer to axilla; single femoral gland positioned on posterior surface of thigh at midpoint between knee and cloaca.
In life, dorsal surface of body yellowish-brown with irregular dark-brown patches, dark-brown X-shaped marking on center of dorsum, dark-brown triangular marking between eyes. A vertical dark-brown band present below eye. Dorsal surface of limbs with dark-brown transverse bands. Tubercles on edge of upper eyelids orange. Supratympanic fold light brown. Surface of throat and chest yellowish-brown with irregular dark brown and white patches and white and orange dots. Center of throat with black longitudinal band. Surface of abdomen white, mottled with orange dots and black patches. Surface of ventral limbs purple brown, with white mottling and dark-brown patches. Spines on tips of tubercles on ventral hindlimbs and area around cloaca black. Digits gray white; subarticular tubercles, inner and outer metacarpal tubercles and inner metatarsal tubercle grayish-brown. Pectoral glands and femoral glands white. Iris yellowish-brown with range mottling.
In preservative, the dorsal surface of the body is dark brown, with markings and patches more distinct. Surface of chest, throat and limbs are dark brown, with dark-brown markings and patches more distinct, white patches and dots faded and orange dots absent. Color of pectoral glands and femoral glands faded.
Morphometric variation is listed in Table
Measurements (in mm) of voucher specimens of Boulenophrys lichun sp. nov.; * holotype.
Voucher |
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GEP a211 | GEP a212 | GEP a213 | GEP a214 * | GEP a215 |
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Sex | male | male | male | male | male | female |
SVL | 36.7 | 33.5 | 35.7 | 34.6 | 37.0 | 47.1 |
HDL | 13.1 | 12.7 | 13.5 | 12.9 | 13.3 | 15.3 |
HDW | 13.6 | 13.3 | 13.8 | 13.6 | 13.9 | 16.5 |
ED | 4.5 | 4.0 | 4.5 | 4.5 | 4.8 | 5.5 |
TD | 2.3 | 2.6 | 2.5 | 2.5 | 2.6 | 2.8 |
TED | 1.7 | 1.7 | 1.8 | 1.8 | 1.7 | 2.4 |
SNT | 4.5 | 4.4 | 4.5 | 4.5 | 4.6 | 4.9 |
IND | 3.6 | 3.7 | 3.8 | 3.7 | 3.9 | 4.3 |
IOD | 3.5 | 3.3 | 3.4 | 3.5 | 3.7 | 4.3 |
HDN | 8.7 | 7.8 | 8.7 | 9.1 | 9.2 | 10.2 |
RAD | 8.6 | 7.0 | 8.5 | 8.1 | 8.7 | 9.0 |
FTL | 20.7 | 18.9 | 20.7 | 20.9 | 21.4 | 24.1 |
TIB | 14.3 | 12.5 | 14.3 | 14.5 | 14.7 | 15.7 |
Boulenophrys lichun sp. nov. can easily be distinguished from the following congeners by its heels not meeting when flexed hindlimbs held at right angles to body axis: B. anlongensis, B. baishanzuensis, B. binlingensis, B. caudoprocta, B. cheni, B. chishuiensis, B. congjiangensis, B. daweimontis, B. fanjingmontis, B. fansipanensis, B. frigida, B. hoanglienensis, B. jiangi, B. jingdongensis, B. jinggangensis, B. jiulianensis, B. leishanensis, B. liboensis, B. lini, B. lushuiensis, B mirabilis, B. mufumontana, B. nanlingensis, B. omeimontis, B. palpebralespinosa, B. qianbeiensis, B. sangzhiensis, B. sanmingensis, B. shimentaina, B. shunhuangensis, B. spinata, B. sanmingensis, B. tongboensis, B. tuberogranulatus, B. wuliangshanensis, B. xianjuensis, B. yangmingensis, B. yaoshanensis, B. yingdeensis (vs. heels overlapping), from B. binchuanensis, B. elongata, B. lishuiensis, B. minor, B. xiangnanensis, B. xuefengmontis (vs. heels just meeting), and from B. angka, B. daiyunensis, B. baolongensis, B. wushanensis, B. yunkaiensis (vs. heels just meeting or slightly overlapping).
Boulenophrys lichun sp. nov. can easily be distinguished from the following congeners by its tongue not notched distally: B. brachykolos, B. insularis, B. pepe (vs. tongue notched distally). Boulenophrys lichun sp. nov. can easily be distinguished from the following congeners by its presence of vomerine teeth: B. acuta, B. boettgeri, B. caobangensis, B. daoji, B. hungtai, B. hengshanensis, B. kuatunensis, B. ombrophila, B. obesa, B. shuichengensis, B. wugongensis (vs. vomerine teeth absent).
Boulenophrys lichun sp. nov. can easily be distinguished from the following congeners by its absence of lateral fringes on webbing on toes: B. dongguanensis, B. fengshunensis, B. nankunensis, B. puningensis (vs. toes with rudimentary webbing), and from B. rubrimera (vs. toes with narrow lateral fringes).
Currently, Boulenophrys lichun sp. nov. is only known from the coastal hills of Ningde City, eastern Fujian Province, China. It inhabits flowing montane seeps and the nearby forest floor and leaf litter. The habitat is surrounded by secondary forest mixed with bamboo groves at elevations between 150–510 m. Advertisement calls of males were heard from February to May. Males were found calling in rock crevices.
The lack of follow-up surveys can pose issues in terms of endangered species listing. Boulenophrys lichun is currently only known from the coastal hills of Ningde City, eastern Fujian. The development of tourism infrastructure, stream diversion and tea leaf cultivation have gradually affected and threatened the habitats of the new species. Thus, more data (i.e., distribution, population size, potential and existing risk factors, etc.) from long-term extensive surveys are urgently required to make an assessment of their endangered status.
Boulenophrys possess limited dispersal abilities and narrow ecological niches, resulting in the restricted distribution ranges of many species (
1 | Vomerine ridges and vomerine teeth present | 2 |
– | Vomerine ridges and vomerine teeth absent | 3 |
2 | Toes with rudimentary webbing and wide lateral fringes | B. daiyunensis |
– | Toes without webbing and lateral fringes | B. lichun |
3 | Toes with rudimentary webbing and lateral fringes | 4 |
– | Toes without webbing and lateral fringes | B. ombrophila |
4 | Toes with wide lateral fringes | 5 |
– | Toes with narrow lateral fringes | B. kuatunensis |
5 | Round light patches on the shoulder present | B. boettgeri |
– | Round light patches on the shoulder absent | B. sanmingensis |
We thank Chong Xu, Xin Zhao, Qi-Feng Yan, Yu-Sen Chen, and Gen-De Yu, for their long-term monitoring of the breeding season of Boulenophrys lichun. We thank Chao Huang from the Australian Museum, Australia for proofreading the manuscript.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This work was supported by the Project of Innovation team of General Institutes of Higher Education in Guangdong Province (2024KCXTD078), the Project of Integration of resource monitoring, epidemic diseases monitoring and rescue capability of wildlife in 2023 (ZT202304111), the Project of Innovation team of General Institutes of Higher Education in Guangdong Province (2024KCXTD078) and the Project of Background survey of biosafety in Guangdong Province (STST-2021-10).
Conceptualization: Jian Wang, Zhao-Chi Zeng; Data curation: Jian Wang, Zhao-Chi Zeng; Funding acquisition: Shi-Shi Lin; Investigation: All authors; Writing – original draft, review and editing: All authors.
Shi-Shi Lin https://orcid.org/0000-0001-7709-7005
Hong-Hui Chen https://orcid.org/0000-0001-5026-2920
Yuan-Hang Li https://orcid.org/0009-0009-6543-9245
Zhao-Ning Peng https://orcid.org/0009-0008-5103-2797
Zhao-Chi Zeng https://orcid.org/0000-0003-4054-8399
Jian Wang https://orcid.org/0000-0003-4249-7767
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Localities, voucher information, and GenBank accession numbers for all samples used in this study
Data type: xlsx