A new species of Chinese horned toad (Anura, Megophryidae, Boulenophrys) from western Guangdong Province, China, with recommendations for extinction risk assessments for several related congeners

A new species o Chinese ho ned oad (Anu a,
Megoph yidae, Boulenoph ys) om wes e n Guangdong
P o ince, China, wi h ecommenda ions o ex inc ion
isk assessmen s o se e al ela ed congene s
Yi-Yang Zhao1*, Shi-Shi Lin2*, Yuan-Hang Li2, Zhi-Tong Lyu3, Ke Zheng4, Zhao-Chi Zeng2, Amaël Bo zée5, Jian Wang2
1 Guangdong Wildli e Moni o ing Rescue and Conse a ion Cen e , Guangzhou 510520, Guangdong, China
2 Guangdong Poly echnic o En i onmen al P o ec ion Enginee ing, Foshan 528216, Guangdong, China
3 Moun ain Ecological Res o a ion and Biodi e si y Conse a ion Key Labo a o y o Sichuan P o ince, Chengdu Ins i u e o Biology, Chinese
Academy o Sciences, Chengdu 610213, Sichuan, China
4 GuangdongYangchunEhuangzhangP o incialNa u eRese eManagemen O ice,Yangjiang529631,Guangdong,China
5 Labo a o yo AnimalBeha iou andConse a ion,Collegeo Li eSciences,NanjingFo es yUni e si y,Nanjing210037,Jiangsu,China
h ps://zoobank.o g/1888DFB8-1A61-45E4-8C8A-BD607671E666
Co esponding au ho s: Jian Wang ([email p o ec ed]); Amaël Bo zée ([email p o ec ed]); Zhao-Chi Zeng ([email p o ec ed])
* These au ho s con ibu ed equally o his wo k.
Academic edi o : Gün e Gollmann ♦
Recei ed
9 Sep embe 2025 ♦
Accep ed
17 No embe 2025 ♦
Published
24 No embe 2025
Abs ac
The genus Boulenoph ys is p edominan ly dis ibu ed in China and exhibi s high species di e si y. Howe e , accu a e conse a ion
assessmen s o his genus a e hinde ed by i s un esol ed axonomy, which is linked o mo phological conse a ism, sympa ic
dis ibu ion pa e ns, and limi ed ield su ey da a. In his s udy, we desc ibe a new species, namely Boulenoph ys yangchunensis
sp. no ., and epo new locali y eco ds o B. acu a (Wang, Li & Jin, 2014) and B. yunkaiensis Qi, Wang, Lyu & Wang, 2021 om
wes e n Guangdong P o ince, China. Addi ionally, we p opose downlis ing B. acu a, cu en ly lis ed as C i ically Endange ed (CR)
on he IUCN Red Lis o Th ea ened Species, o a lowe h ea ca ego y, and we ecommend an ex inc ion isk ca ego y upda e o
Boulenoph ys yangchunensis sp. no ., B. yunkaiensis, and B. gaolanensis Song, Wang, Qi, Wang & Wang, 2024. We u he discuss
key conse a ion challenges and p o ide a ge ed managemen ecommenda ions o add ess hese challenges.
Key Wo ds
conse a ion assessmen , managemen sugges ions, molecula phylogeny, mo phological conse a ism, ange ex ension, eassess-
men , sympa y, sys ema ics
In oduc ion
The speciose genus Boulenoph ys Fei, Ye & Jiang, 2016,
known as he Chinese ho ned oads and cu en ly com-
p ising 78 ecognized species, is widesp ead in he sub-
opical and opical a eas o mainland Eas Asia (F os
2025). Speci ically, he genus is p incipally dis ibu ed in
sou he n China and sou hwa ds in o no he nmos Vie -
nam, Laos, Myanma , and Thailand (Fei and Ye 2016;
Lyu e al. 2023; F os 2025). Boulenoph ys possess lim-
i ed dispe sal abili ies and na ow ecological niches, e-
sul ing in es ic ed dis ibu ion anges o many species
He pe ozoa 38: 345–357 (2025)
DOI 10.3897/he pe ozoa.38.e171400
Copy igh Yi-Yang Zhao e al. This is an open access a icle dis ibu ed unde he e ms o he C ea i e Commons A ibu ion License
(CC BY 4.0), which pe mi s un es ic ed use, dis ibu ion, and ep oduc ion in any medium, p o ided he o iginal au ho and sou ce
a e c edi ed.
he pe ozoa.penso .ne
Yi-Yang Zhao e al.: A new species o Boulenoph ys346
(Wang e al. 2019a; Lyu e al. 2023). Due o he combined
e ec o a ious ac o s, conse a ion e o s o his ge-
nus ha e encoun e ed signi ican challenges. Besides,
hei high le els o mo phological conse a ism and he
lack o ollow-up su eys ha e led o inadequa e p o ec-
ion due o misiden i ica ions and missing da a (Lin e al.
2024). The e o e, accu a e assessmen s o he ex inc ion
isk o Boulenoph ys species s ill equi e a signi ican
amoun o ield su eys and con inuous da a upda es.
As he esea ch p og essed, Guangdong P o ince,
China, g adually became one o he p o inces wi h ex-
emely high di e si y o he genus Boulenoph ys (20 o
77 species). Cu en ly, he ype locali y o 15 species is
loca ed wi hin he ju isdic ion o Guangdong P o ince,
i.e., B. acu a (Wang, Li & Jin, 2014), B. dongguanensis
(Wang & Wang, 2019), B. elonga a Zeng, Wang, Chen,
Xiao, Zhan, Li & Lin, 2024, B. engshunensis Wang,
Zeng, Lyu & Wang, 2022, B. gaolanensis Song, Wang,
Qi, Wang & Wang, 2024, B. hung ai Wang, Zeng, Lyu,
Xiao & Wang, 2022, B. insula is (Wang, Liu, Lyu, Zeng
& Wang, 2017), B. nankunensis (Wang, Zeng & Wang,
2019), B. nanlingensis (Lyu, Wang, Liu & Wang, 2019),
B. obesa (Wang, Li & Zhao, 2014), B. pepe Wang &
Zeng, 2024, B. puningensis Wang, Zeng, Lyu, Xiao &
Wang, 2022, B. shimen aina (Lyu, Liu & Wang, 2020),
B. yingdeensis Qi, Lyu, Wang & Wang, 2021, and
B. yunkaiensis Qi, Wang, Lyu & Wang, 2021 (Lyu e
al. 2023; Song e al. 2024; F os 2025). Among hem,
he Fengkai ho ned oad, Boulenoph ys acu a, was mos
ecen ly assessed o he IUCN Red Lis o Th ea -
ened Species in 2019 and lis ed as C i ically Endan-
ge ed (CR) unde c i e ia B1ab(iii), due o i s limi ed
dis ibu ion ange wi hin Heishiding Na u e Rese e in
Fengkai Coun y, wes e n Guangdong P o ince, China
(Li e al. 2014). Based on molecula da a, Song e al.
(2024) p o ided a ange ex ension by epo ing a new
popula ion om M . Yunkai in Xinyi Ci y, wes e n
Guangdong P o ince, China, app oxima ely 155 km
away in s aigh -line dis ance. Du ing ou ield su eys
in Ehuangzhang NR in Yangchun Ci y, wes e n Guang-
dong P o ince, China, we collec ed a se ies o sympa -
ic adpole samples and adul specimens o he genus
Boulenoph ys (Fig. 1). Subsequen molecula analysis
e ealed ha wo o he samples (GEP asp062 and GEP
asp064; En i onmen al P o ec ion Enginee ing o he
abb e ia ion GEP and amphibian issue sample o he
abb e ia ion asp) clus e ed wi h he ypical lineage o
B. acu a wi h shallow gene ic di e gence, and ou sam-
ples (GEP asp061, GEP asp063, and GEP asp065–066)
clus e ed wi h he ypical lineage o B. yunkaiensis wi h
shallow gene ic di e gence. These clus e s ep esen
new geog aphical subpopula ions o he wo species, e-
spec i ely (Fig. 2). Addi ionally, i e specimens (GEP
a168–170, SYS a009518, and CIB 121751; Chengdu
Figu e 1. Map showing he ype locali y o Boulenoph ys yangchunensis sp. no . and he new dis ibu ion si es o B. acu a and
B. yunkaiensis (1); he ype locali y o B. acu a (2); he ype locali y o B. gaolanensis (3); and he ype locali y o B. yunkaiensis and
he new dis ibu ion si e o B. acu a (4).
He pe ozoa 38: 345–357 (2025)
he pe ozoa.penso .ne
347
Ins i u e o Biology, Chinese Academy o Sciences o
he abb e ia ion CIB) exhibi ed dis inc mo phologi-
cal di e ences and signi ican gene ic di e gence om
all known congene s wi hin he genus. The e o e, we
desc ibe a new species om Ehuangzhang NR, epo
ange ex ensions o B. acu a (app oxima ely 188 km in
s aigh -line dis ance om i s ype locali y in Heishid-
ing NR) and o B. yunkaiensis (app oxima ely 52 km
om i s ype locali y in M . Yunkai), and also p o ide
a eassessmen o he endange ed s a us o B. acu a and
assessmen s o B. yunkaiensis and B. gaolanensis.
Ma e ials and me hods
Mo phology
Fi e adul male specimens (GEP a168–170; CIB 121751;
SYS a009518) o he genus Boulenoph ys we e col-
lec ed on 11 Augus 2023 om Guangdong Yangchun
Ehuangzhang P o incial Na u e Rese e (21.851912°N,
111.421553°E; ca. 1,060 m a.s.l.) in Yangchun Ci y,
Guangdong P o ince, China. All examined specimens
we e ixed in 10% bu e ed o malin and la e ans e ed
o 70% e hanol. All s udied specimens ha e been deposi -
ed a he Guangdong Poly echnic o En i onmen al P o-
ec ion Enginee ing (GEP) acili y in Foshan, Guangdong
P o ince; he Museum o Biology, Sun Ya -sen Uni e -
si y in Guangzhou, Guangdong P o ince, China (SYS);
and he He pe ological Museum a he Chengdu Ins i-
u e o Biology, Chinese Academy o Sciences (CIB) in
Chengdu, Sichuan P o ince, China.
Ex e nal measu emen s we e eco ded wi h a digi al
calipe (Neiko 01407A s ainless s eel 6-inch digi al cal-
ipe ) o he nea es 0.1 mm. These measu emen s a e as
ollows: SVL (snou – en leng h, om ip o snou o
pos e io ma gin o en ); HDL (head leng h, om ip o
snou o he a icula ion o he jaw); HDW (head wid h,
head wid h a he commissu e o he jaws); SNT (snou
leng h, om ip o snou o he an e io co ne o he
eye); IND (in e nasal dis ance, dis ance be ween na es);
IOD (in e o bi al dis ance, minimum dis ance be ween
Figu e 2. Maximum likelihood ee esul s om pa ial DNA sequences o he mi ochond ial 16S RNA and COI gene. Numbe s
be o e slashes indica e boo s ap suppo (BS), and numbe s a e slashes indica e Bayesian pos e io p obabili ies (BPP).
B. yunkaiensis
B. gaolanensis
B. yangchunensis sp. no .
B. acu a
0.05 B. shunhuangensis
B. xianjuensis
GEP asp061 ˙
B. boege i
B. palpeb alespinosa
B. mino
B. lichun
GEP asp066 ˙
B. daiyunensis
B. baolongensis
B. nanlingensis
B. baishanzuensis
B. cheni
SYS a002278 ˚
B. ansipanensis
B. yingdeensis
GEP asp064 ˙
B. binlingensis
B. daweimons
B. yaoshanensis
B. engshunensis
B. insula is
B. dalaolingensis
SYS a009518 ˙
SYS a009225 ˜
B. elonga a
SYS a004694 ˜
B. jinggangensis
B. ongboensis
B. jiulianensis
X. mangshanensis
GEP asp065 ˙
B. changyangensis
B. qianbeiensis
B. jiangi
B. ube og anula a
B. liboensis
B. angka
B. anlongensis
B. mi abilis
B. congjiangensis
SYS a009113 ˜
B. jingdongensis
B. caudop oc a
B. mu umon ana
B. hengshanensis
B. dupanglingensis
B. binchuanensis
GEP asp062 ˙
SYS a002276 ˚
GEP a169 ˙
SYS a002266 ˚
SYS a009236 ˜
B. spina a
B. anjingmons
B. hoanglienensis
SYS a009235 ˜
B. xue engmons
B. sanmingensis
B. leishanensis
B. lini
B. wushanensis
B. kua unensis
B. ub ime a
B. wugongensis
B. b achykolos
B. omb ophila
B. daoji
B. obesa
B. chishuiensis
GEP a170 ˙
B. caobangensis
B. xiangnanensis
B. hung ai
B. lushuiensis
B. daxuemons
B. dongguanensis
B. shimen aina
B. huangniushiensis
B. gu u
B. omeimons
B. sangzhiensis
B. wuliangshanensis
X. glandulosa
B. puningensis
B. yangmingensis
B. nankunensis
CIB 121751
˙
B. pepe
GEP a168 ˙
GEP asp063 ˙
B. lishuiensis
B. igida
ʒEhuangzhang Na u e Rese e, Yangchun
ʓHeishiding Na u e Rese e, Fengkai
ʔGaolan Island, Zhuhai
ʕM Yunkai, Xinyi
BS≥70 & BPP≥0.90
BS≥70 & BPP<0.90
BS<70 & BPP≥0.90
BS<70 & BPP<0.90
Boulenoph ys
Xenoph ys
100/1.00
100/1.00
91/1.00
95/1.00
72/0.95
96/0.99
he pe ozoa.penso .ne
Yi-Yang Zhao e al.: A new species o Boulenoph ys348
uppe eyelids); ED (eye diame e , om he an e io co -
ne o he eye o he pos e io co ne o he eye); TD
( ympanum diame e , ho izon al diame e o ympa-
num); TED ( ympanum–eye dis ance, om an e io
edge o ympanum o pos e io co ne o he eye); HND
(hand leng h, om he p oximal bo de o he ou e pal-
ma ube cle o he ip o digi III); RAD ( o ea m o
adioulna leng h, om he lexed elbow o he p oximal
bo de o he ou e palma ube cle); FTL ( oo leng h,
om dis al end o shank o he ip o digi IV); TIB (c us
o ibio ibula leng h, om he ou e su ace o he lexed
knee o he heel). Sex was de e mined by ex e nal sec-
onda y sexual cha ac e s, such as he p esence o ocal
sacs, nup ial pads, o spines in males and hei absence
in emales (Fei e al. 2009). Mo phological cha ac e s o
all ecognized species o he genus Boulenoph ys used
o compa isons we e based on an examina ion o mu-
seum specimens lis ed in Appendix I and in o ma ion
a ailable in he li e a u e (Table 1).
Phylogeny
Fi e issue samples we e de i ed om he i e
adul specimens (GEP a168–170; CIB 121751; SYS
a009518). Ano he six issue samples (GEP asp061–
066) we e de i ed om he ail ins o di e en adpoles
om Guangdong Yangchun Ehuangzhang P o incial
Na u e Rese e (21.851912°N, 111.421553°E; ca.
1,060 m a.s.l.), and he adpoles we e no collec ed. In
o al, 94 samples we e used o he phylogene ic analy-
ses, encompassing 11 newly sequenced indi iduals and
o he s downloaded om GenBank (h ps://www.ncbi.
nlm.nih.go /genbank/). Two samples o he genus Xe-
noph ys Gün he , 1864, we e used as ou g oups (Suppl.
ma e ial 1). Two pa ial mi ochond ial genes, he 16S
ibosomal RNA (16S) and he cy och ome c oxidase 1
(COI), we e used o phylogene ic analysis. DNA ex-
ac ion, PCR ampli ica ion, and sequencing p o ocols
ollowed Liu e al. (2018).
We used Clus al X 2.0 (Thompson e al. 1997) o align
he sequences using de aul pa ame e s. We hen used
Pa i ionFinde (Lan ea e al. 2012) o de e mine he op-
imal pa i ioning schemes, and we de e mined ha pa -
i ioning by gene was op imal o 16S, while pa i ioning
by codon posi ion was op imal o COI, wi h GTR+I+G
iden i ied as he bes - i nucleo ide subs i u ion model o
all pa i ions. We cons uc ed he phylogene ic ees using
maximum likelihood (ML) implemen ed in RAxMLGUI
.1.3 (Sil es o and Michalak 2012) and Bayesian in e -
ence (BI) using M Bayes .3.2.4 (Ronquis e al. 2012).
Fo he ML analysis, an op imal ee was ob ained, and
b anch suppo s we e e alua ed wi h 1000 apid boo -
s apping eplica es. Fo he BI analysis, wo independen
uns we e conduc ed, each unning o 10,000,000 gen-
e a ions and sampled e e y 1000 gene a ions, wi h he
i s 25% o samples disca ded as bu n-in, esul ing in a
po en ial scale educ ion ac o (PSRF) o < 0.01. Then,
he emaining ees we e used o c ea e a consensus ee.
Nodes ha ing ML boo s ap alues (BS) ≥ 70 and BI pos-
e io p obabili ies (BPP) ≥ 0.90 we e conside ed well
suppo ed. Gene ic dis ances among all Boulenoph ys
samples we e calcula ed in MEGA 11 using he unco -
ec ed p-dis ance model based on he 16S.
Th ea assessmen s
Among he ocal species in his wo k, B. acu a has
been assessed as C i ically Endange ed (ASG 2021),
bu B. yunkaiensis and B. gaolanensis ha e no been as-
sessed. He e, he h ea assessmen o B. gaolanensis is
ele an , as i is closely ela ed o he axa desc ibed in
his manusc ip . The geog aphic a ea o his s udy is cha -
ac e ized by low-ele a ion hills and moun ains in e up -
ed by low-ele a ion ag icul u al plains and la ge u ban
a eas. As a esul , he popula ions a e disconnec ed bo h
because o he landscape and human ac i i ies. To sug-
ges an ex inc ion isk assessmen o hese ou species,
we ollowed he IUCN Red Lis ca ego ies and c i e ia
(www.iucn edlis .o g/ esou ces/ edlis guidelines) due
o hei obus ness, especially o species wi h es ic ed
anges (Maes e al. 2015), and because e alua ions a e
conduc ed agains quan i a i e h esholds o i e c i e ia
(IUCN 2012). Howe e , we could no use c i e ia A (pop-
ula ion size educ ion), C (small and declining popula ion
size), and E (quan i a i e analysis o ex inc ion isk) due
o he absence o da a on popula ion sizes. Ins ead, we
elied on c i e ia B (geog aphic ange size) and D ( e y
small popula ion and/o es ic ed dis ibu ion) o he as-
sessmen s. To de e mine he ex en o occu ence (he e-
a e EOO) and he a ea o occupancy (he ea e AOO)
o each species, we used he online pla o m GeoCAT
(www.geoca .iucn edlis .o g).
Resul s
Molecula phylogene ic analysis
The BI and ML phylogene ic ees esul ed in essen ially
iden ical opologies (ML phylogene ic opology in Fig.
2). In he phylogene ic esul , he Boulenoph ys samples
om Ehuangzhang NR, Yangchun, o med h ee indepen-
den clades. Among hem, samples (GEP asp062 and GEP
asp064) om wo uncap u ed adpoles clus e ed wi h he
ypical se ies o B. acu a om Heishiding NR, Fengkai,
and ha om M . Yunkai (Song e al. 2024), Xinyi, wi h
good suppo (BS = 74; BPP = 0.95). In addi ion, samples
(GEP asp061, GEP asp063, and GEP asp065–066) om
ou uncap u ed adpoles clus e ed wi h he ypical se ies
o B. yunkaiensis om M . Yunkai, Xinyi, wi h s ong
suppo (BS = 97; BPP = 0.99), and showed e y shal-
low di e gence. Thus, B. acu a and B. yunkaiensis we e
He pe ozoa 38: 345–357 (2025)
he pe ozoa.penso .ne
349
Table 1. Li e a u e o mo phological cha ac e s o 78 ecognized species o Boulenoph ys.
ID Boulenoph ys spp. Re e ences
1B. acu a (Wang, Li & Jin, 2014) Lyu e al. 2023
2B. angka (Wu, Suwannapoom, Poya ko , Pawangkhanan , Xu, Jin, Mu phy & Che, 2019) Wu e al. 2019
3B. anlongensis (Li, Lu, Liu & Wang, 2020) Li e al. 2020; Lyu e al. 2023
4B. baishanzuensis (Wu, Li, Liu, Wang & Wu, 2020) Wu e al. 2020
5B. baolongensis (Ye, Fei & Xie, 2007) Ye e al. 2007; Fei and Ye 2016
6B. binchuanensis (Ye & Fei, 1995) Lyu e al. 2023
7B. binlingensis (Jiang, Fei & Ye, 2009) Fei e al. 2009; Lyu e al. 2023
8B. boe ge i (Boulenge , 1899) Lyu e al. 2023
9B. b achykolos (Inge & Rome , 1961) Lyu e al. 2023
10 B. caobangensis (Nguyen, Pham, Nguyen, Luong & Ziegle , 2020) Nguyen e al. 2020
11 B. caudop oc a (Shen, 1994) Shen 1994; Lyu e al. 2023
12 B. congjiangensis(Luo, Wang, Wang, Lu, Wang, Deng & Zhou, 2021) Lyu e al. 2023
13 B. changyangensis Li, Shi, Liu, Zhao, Gao & Wang, 2025 Li e al. 2025
14 B. cheni (Wang & Liu, 2014) Lyu e al. 2023
15 B. chishuiensis (Xu, Li, Liu, Wei & Wang, 2020) Lyu e al. 2023
16 B. daiyunensis (Lyu, Wang & Wang, 2021) Lyu e al. 2023
17 B. dalaolingensis Wang, Chen & Liu, 2025 Wang e al. 2025a
18 B.daoji(Lyu, Zeng, Wang & Wang, 2021) Lyu e al. 2023
19 B. daweimon is (Rao & Yang, 1997) Rao and Yang 1997
20 B. daxuemon is Liu, Li, Cheng, Wei, Wang & Cheng, 2025 Liu e al. 2025
21 B. dongguanensis (Wang & Wang, 2019) Lyu e al. 2023
22 B. dupanglingensis Xiao & Mo, 2025 Xiao e al. 2025
23 B. elonga a Zeng, Wang, Chen, Xiao, Zhan, Li & Lin, 2024 Zeng e al. 2024
24 B. engshunensis Wang, Zeng, Lyu & Wang, 2022 Lyu e al. 2023
25 B. anjingmon is(Zhang, Liang, Ran & Shen, 2012) Lyu e al. 2023
26 B. ansipanensis (Tapley, Cu aja , Mahony, Nguyen, Dau, Luong, Le, Nguyen, Nguyen, Po way, Luong &
Rowley, 2018)
Tapley e al. 2018a
27 B. igida (Tapley, Cu aja, Nguyen, Po way, Mahony, Nguyen, Ha ding, Luong & Rowley, 2021) Tapley e al. 2021
28 B. gaolanensis Song, Wang, Qi, Wang & Wang, 2024 Song e al. 2024
29 B. gu u Kuang, Wei & Shi, 2025 Kuang e al. 2025
30 B. hoanglienensis (Tapley, Cu aja , Mahony, Nguyen, Dau, Luong, Le, Nguyen, Nguyen, Po way, Luong &
Rowley, 2018)
Tapley e al. 2018a
31 B. huangniushiensis Wang, Wu, Song, Huang, Zhuo & Wang, 2025 Wang e al. 2025b
32 B. hung ai Wang, Zeng, Lyu, Xiao & Wang, 2022 Lyu e al. 2023
33 B. hengshanensis Qian, Hu, Mo, Gao, Zhang & Yang, 2023 Qian e al. 2023
34 B. insula is (Wang, Liu, Lyu, Zeng & Wang, 2017) Lyu e al. 2023
35 B. jiangi(Liu, Li, Wei, Xu, Cheng, Wang & Wu, 2020) Lyu e al. 2023
36 B. jingdongensis (Fei & Ye, 1983) Fei e al. 1983; Lyu e al. 2023
37 B. jinggangensis(Wang, 2012) Lyu e al. 2023
38 B.jiulianensis (Wang, Zeng, Lyu & Wang, 2019) Lyu e al. 2023
39 B. kua unensis (Pope, 1929) Lyu e al. 2023
40 B. leishanensis (Li, Xu, Liu, Jiang, Wei & Wang, 2018) Lyu e al. 2023
41 B. lushanensis Shen, Zhou & Li, 2025 Shen e al. 2025
42 B. lushuiensis (Shi, Li, Zhu, Jiang, Jiang & Wang, 2021) Lyu e al. 2023
43 B. lichun Lin, Chen, Li, Peng, Zeng & Wang, 2024 Lin e al. 2024
44 B. liboensis (Zhang, Li, Xiao, Li, Pan, Wang, Zhang & Zhou, 2017) Zhang e al. 2017
45 B. lini (Wang & Yang, 2014) Lyu e al. 2023
46 B. lishuiensis (Wang, Liu & Jiang, 2017) Lyu e al. 2023
47 B. mino (S ejnege , 1926) Lyu e al. 2023
48 B. mi abilis (Lyu, Wang & Zhao, 2020) Lyu e al. 2023
49 B. mu umon ana (Wang, Lyu & Wang, 2019) Lyu e al. 2023
50 B. nankunensis (Wang, Zeng & Wang, 2019) Lyu e al. 2023
51 B. nanlingensis (Lyu, Wang, Liu & Wang, 2019) Lyu e al. 2023
52 B. obesa (Wang, Li & Zhao, 2014) Lyu e al. 2023
53 B. omb ophila (Messenge & Dahn, 2019) Lyu e al. 2023
54 B. omeimon is (Liu, 1950) Lyu e al. 2023
55 B. palpeb alespinosa (Bou e , 1937) Fei e al. 2009; Lyu e al. 2023
56 B. pepe (Wang & Zeng, 2024) Wang e al. 2024
57 B. puningensis Wang, Zeng, Lyu, Xiao & Wang, 2022 Lyu e al. 2023
58 B. qianbeinsis (Su, Shi, Wu, Li, Yao, Wang & Li, 2020) Lyu e al. 2023
59 B. ub ime a (Tapley, Cu aja , Mahony, Chung, Dau, Nguyen, Luong & Rowley, 2017) Tapley e al. 2017, 2018b
60 B. sangzhiensis (Jiang, Ye & Fei, 2008) Lyu e al. 2023
61 B. sanmingensis (Lyu & Wang, 2021) Lyu e al. 2023
62 B. shimen aina (Lyu, Liu & Wang, 2020) Lyu e al. 2023
63 B. shuichengensis (Tian & Sun, 1995) Tian and Sun 1995; Tian e al. 2000; Fei and Ye 2016
64 B. shunhuangensis (Wang, Deng, Liu, Wu & Liu, 2019) Lyu e al. 2023

he pe ozoa.penso .ne
Yi-Yang Zhao e al.: A new species o Boulenoph ys350
e i ied o be dis ibu ed in Ehuangzhang Na u e Rese e
based on molecula da a, which also con i ms he expan-
sion o he dis ibu ion ange o he wo species. As he
dis ibu ion ange o he c i ically endange ed B. acu a
has been excep ionally unde es ima ed, we sugges an up-
da e o he ex inc ion isk assessmen below.
Addi ionally, samples aken om i e cap u ed adul
specimens (GEP a168–170, SYS a009518, and CIB
121751) o med a monophyle ic lineage wi h s ong
suppo (BS = 100; BPP = 1.00). This new lineage is a
sis e axon o B. acu a wi h s ong suppo (BS = 97;
BPP = 1.00) and signi ican di e gence wi h a mean
p-dis ance o 6.5% (Suppl. ma e ial 2). Fu he mo pho-
logical examina ion also showed ha he co esponding
specimens o he new lineage can be dis inguished om
all ecognized congene s. Thus, we desc ibe he co e-
sponding specimens as a new species, which is sympa ic
wi h B. acu a and B. yunkaiensis.
Taxonomic accoun
Boulenoph ys yangchunensis sp. no .
h ps://zoobank.o g/E1D9D803-9BC7-4933-AB38-13B26681B7DC
Figs 3, 4
English name: Yangchun Ho ned Toad
Chinese name: yáng chūn jiǎo chán (阳春角蟾)
Type ma e ials. Holo ype • ♂.GEP a170, collec ed by
Jian Wang, Zhao-Chi Zeng, Hong-Hui Chen, Yuan-Hang
Li, and Ke Zheng on 11 Augus 2023 om Guangdong
Yangchun Ehuangzhang P o incial Na u e Rese e
(21.851912°N, 111.421553°E; ca. 1,060 m a.s.l.) in
Yangchun Ci y, Guangdong P o ince, China.
Pa a ypes. •4 ♂: GEP a168–169, SYS a009518 ( ield
numbe : GEP a172), and CIB 121751 ( ield numbe : GEP
a171), same collec ion da a as holo ype.
E ymology. The speci ic epi he yangchunensis is in
e e ence o he ype locali y, Yangchun Ci y.
Diagnosis. (1) small size (SVL 27.6–28.7 mm in i e
adul males); (2) snou sha pened in do sal iew, can hus
os alis well de eloped, ongue no no ched dis ally; (3)
ympanum dis inc ; (4) ome ine idges and ome ine
ee h absen ; (5) do sal skin ough and highly g anula ,
discon inuous X-shaped idge on cen e o do sum, dis-
con inuous do sola e al idges p esen , spa se la ge ube -
cles on lanks, do sal limbs wi h discon inuous ans e se
idges and ube cles; (6) ou e ma gin o uppe eyelid
wi h a small ho n-like p ominen ube cle, sup a ym-
panic old dis inc and na ow, cu ing pos e o en al-
ly o abo e a m; (7) wo me aca pal ube cles dis inc ,
inne one obse ably enla ged, ela i e inge leng hs
I < II < IV < III, dis inc suba icula ube cle a base o
each inge ; (8) heels jus mee ing when hindlimbs old-
ed, ibio- a sal a icula ion eaching middle o an e io
co ne o eye; (9) oes wi hou webbing and la e al ing-
es, inne me a a sal ube cle long o oid, ou e one absen ,
ela i e oe leng h I < II < V < III < IV; (10) do sal su ace
yellowish-b own o da k b own, wi h indis inc i egula
da k-b own pa ches and small o ange do s; da k-b own
iangula ma king be ween eyes; do sal limbs and digi s
ligh b own wi h da k-b own ans e se bands; and (11) a
single nup ial pad bea ing dense nup ial spines on do sal
bases o inge s I and II in b eeding adul males, subgula
ocal sac p esen in males.
Compa isons. Boulenoph ys yangchunensis sp. no .
is phylogene ically mos closely ela ed o B. acu a and
B. gaolanensis (Fig. 2). Howe e , he new species dis-
inc i ely di e s om B. acu a by i s heels mee ing when
hindlimbs a e olded ( s. heels no mee ing), absence o
ome ine idge ( s. p esence o weak ome ine idge), ab-
sence o webbing and la e al inges on oes ( s. p esence
o udimen a y webbing and na ow la e al inges on oes),
p esence o nup ial pads bea ing dense nup ial spines on
do sal bases o inge s I and II in b eeding adul males ( s.
p esence o weak nup ial pads no bea ing nup ial spines
on do sal bases o inge s I and II in b eeding adul males).
The new species u he di e s om B. gaolanensis by i s
heels mee ing when hindlimbs olded ( s. heels no mee -
ing), sha pened snou in do sal iew ( s. ounded snou in
do sal iew), p esence o nup ial pad bea ing dense nup ial
spines on do sal bases o inge s I and II in b eeding adul
males ( s. nup ial pad and nup ial spines in isible).
Boulenoph ys yangchunensis sp. no . can be easily
dis inguished om B. b achykolos, B. daoji, B. dawei-
mon is, B. dongguanensis, B. engshunensis, B. igida,
B. hung ai, B. hengshanensis, B. kua unensis, B. lichun,
B. lushanensis, B. nankunensis, B. puningensis, B. pepe,
ID Boulenoph ys spp. Re e ences
65 B. spina a (Liu & Hu, 1973) Hu e al. 1973; Lyu e al. 2023
66 B. ongboensis (Wang & Lyu, 2021) Lyu e al. 2023
67 B. ube og anula a (Shen, Mo & Li, 2010) Mo e al. 2010; Fei and Ye 2016; Lyu e al. 2023
68 B. wugongensis (Wang, Lyu & Wang, 2019) Lyu e al. 2023
69 B. wuliangshanensis (Ye & Fei, 1995) Lyu e al. 2023
70 B. wushanensis (Ye & Fei, 1995) Ye and Fei 1995; Fei and Ye 2016; Lyu e al. 2023
71 B. xiangnanensis (Lyu, Zeng & Wang, 2020) Lyu e al. 2023
72 B. xianjuensis(Wang, Wu, Peng, Shi, Lu & Wu, 2020) Lyu e al. 2023
73 B. xue engmon is Lyu & Wang, 2023 Lyu e al. 2023
74 B. yangmingensis (Lyu, Zeng & Wang, 2020) Lyu e al. 2023
75 B. yaoshanensis Qi, Mo, Lyu, Wang & Wang, 2021 Lyu e al. 2023
76 B. yezhongensis Liu, Feng, Shen, Li, Cheng, Wei, Wang & Su, 2025 Liu e al. 2025
77 B. yingdeensis Qi, Lyu, Wang & Wang, 2021 Lyu e al. 2023
78 B. yunkaiensis Qi, Wang, Lyu & Wang, 2021 Lyu e al. 2023
He pe ozoa 38: 345–357 (2025)
he pe ozoa.penso .ne
351
B. insula is, B. obesa, B. omb ophila, and B. wugongen-
sis, by i s heels mee ing when hindlimbs olded ( s. heels
no mee ing); and om B. anlongensis, B. baishanzuensis,
B. binlingensis, B. caudop oc a, B. cheni, B. changyangen-
sis,B.congjiangensis, B. changyangensis, B. chishuiensis,
B. daxuemon is, B. dalaolingensis, B. dupanglingensis,
B. anjingmon is, B. jiangi, B. jiulianensis, B. jingdon-
gensis,B.jinggangensis, B. leishanensis, B. lushuiensis,
B. liboensis, B. lini, B. lishuiensis, B. mi abilis, B. mu u-
mon ana, B. nanlingensis, B. omeimon is, B. palpeb alespi-
nosa, B. qianbeiensis, B. sangzhiensis, B. shunhuangensis,
B. sanmingensis, B. shimen aina, B. spina a, B. ongboen-
sis, B. ube og anula a, B. xianjuensis, B. yangmingen-
sis, B. yezhongensis, B. yingdeensis, B. yunkaiensis, and
B. wuliangshanensis ( s. heels o e lapping).
Boulenoph ys yangchunensis sp. no . u he di -
e s om B. angka, B. anlongensis, B. baishanzuensis,
B. binlingensis, B. elonga a, B. ansipanensis, B. hoan-
glienensis, B. mino , B. ub ime a, B. shuichengensis,
B. xiangnanensis, B. yaoshanensis, and B. yezhongensis
by i s absence o ome ine idge ( s. p esence o om-
e ine idge); om B. daiyunensis, B. elonga a, B. an-
sipanensis, B. hoanglienensis, and B. ub ime a by i s
absence o ome ine ee h ( s. p esence o ome ine
ee h); and om B. baolongensis and B. boe ge i by i s
ongue no no ched dis ally ( s. ongue no ched). The
new species di e s om B. wushanensis and B. gu u
by i s absence o la e al inges on oes ( s. p esence
o la e al inges on oes); om B. yaoshanensis and
B. caobangensis by i s absence o webbing on oes ( s.
p esence o webbing on oes); om B. xue engmon is
by i s ibio a sal a icula ion eaching egion be ween
middle o eye o an e io co ne o eye when legs a e
s e ched o wa d ( s. be ween pos e io ma gin o ym-
panum and pos e io co ne o eye), and he absence o
g anules on en al skin ( s. p esence o se e al g an-
ules on ches ); om B. huangniushiensis by i s p esence
o nup ial pads bea ing nup ial spines in males ( s. ab-
sence o nup ial pads o nup ial spines).
Desc ip ion o holo ype. Adul male. Body size small,
SVL 27.9 mm. Head leng h almos equal o head wid h,
HDW/HDL 1.00; snou sha pened in do sal iew, p o-
jec ing, sloping backwa d o mou h in p o ile, p o uding
well beyond ma gin o lowe jaw; op o head la ; eyes
mode a e in size, ED 0.40 o HDL, pupil e ical, nea
diamond-shaped; nos il obliquely o oid; can hus os alis
well de eloped; lo eal egion sligh ly oblique; in e nasal
dis ance sligh ly la ge han in e o bi al dis ance; ympan-
ic egion oblique, ympanum dis inc and isible in do sal
iew; ympanum mode a e in size, ma gin clea , uppe
ma gin in con ac wi h sup a ympanic old, lowe ma gin
in con ac wi h uppe lip, TD/ED 0.50; la ge o oid choa-
nae a base o maxilla; ome ine idge and ome ine ee h
absen , maxilla y ee h p esen ; ma gin o ongue ounded,
no no ched dis ally; p esence o single subgula ocal sac.
Fo ea m leng h 0.20 o SVL, hand 0.30 o SVL;
webbing absen be ween inge s, la e al inges absen ,
ela i e inge leng h I < II < IV < III; ips o inge s
sligh ly dila ed, ound; suba icula ube cles on base
o inge s p esen , dis inc ; inne me aca pal ube cle
Figu e 3. The holo ype (GEP a170) o Boulenoph ys yangchunensis sp. no . in p ese a i e: do sal iew (A); en al iew (B);
en al iew o hand (C); en al iew o oo (D). Pho os by Jian Wang.
he pe ozoa.penso .ne
Yi-Yang Zhao e al.: A new species o Boulenoph ys352
obse ably enla ged, ou e one sligh ly smalle ; sin-
gle nup ial pad bea ing dense nup ial spines p esen on
do sal su ace o i s and second inge s, espec i ely.
Hindlimbs mode a e in leng h, ibio- a sal a icula ion
eaching o wa d o an e io co ne o eye when hind-
limb s e ched along body; heels jus mee ing when
lexed hindlimbs held a igh angles o body axis; c us
leng h 0.50 o SVL and oo leng h 0.70 o SVL; ela-
i e oe leng h I < II < V < III < IV; ips o oes ound
and sligh ly dila ed; oes wi hou la e al inges and
webbing; suba icula ube cles on base o oes p esen
and dis inc ; inne me a a sal ube cle long o oid and
lacking ou e me a a sal ube cle.
Do sal skin ough and highly g anula ; dense la ge
ube cles on lanks; single ho n-like p ominen ube cle
on edge o uppe eyelid; ob ious sup a ympanic old
cu ing pos e o en ally om pos e io co ne o eye o
le el abo e inse ion o a m; discon inuous X-shaped
idge on cen e o do sum, discon inuous do sola e al
idges p esen ; do sal limbs wi h discon inuous ans-
e se idges and ube cles; en al su ace smoo h;
small and dis inc pec o al gland close o axilla; single
emo al gland posi ioned on pos e io su ace o high
a midpoin be ween knee and cloaca.
Colo a ion o holo ype. In li e, do sal su ace o body
yellowish-b own wi h indis inc i egula da k-b own pa ch-
es and small o ange do s, da k-b own iangula ma king be-
ween eyes. A e ical da k-b own band p esen below eye.
Tympanum ligh b own. Do sal su ace o limbs wi h da k-
b own ans e se bands. Tube cles on edge o uppe eyelids
o ange. Sup a ympanic old g ey whi e. I is coppe y b own.
In p ese a i e, do sal su ace o body g eyish-b own,
wi h ma kings and pa ches mo e dis inc . Su ace o
ches , h oa and limbs a e yellowish b own, wi h dis inc
da k-b own ma kings and g eyish-whi e pa ches. Pec o-
al glands and emo al glands g eyish-whi e.
Va ia ion. Mo phome ic a ia ions a e lis ed in
Table 2. Mos o he pa a ypes a e simila o he holo ype
in mo phology and colo pa e n, excep o he ollowing:
ibio- a sal a icula ion eaching o wa d o middle o eye
when hindlimb s e ched along body in he pa a ype GEP
a171; head wid h la ge han head leng h in he pa a ype
SYS a009518; do sal colo a ion yellowish-b own, g ey-
ish-b own o da k b own (Fig. 4).
Figu e 4. Li e aspec o Boulenoph ys yangchunensis sp. no .: holo ype GEP a170 (A, B); pa a ype GEP a168 (C); pa a ype GEP
a172 (D). Pho os by Jian Wang.
He pe ozoa 38: 345–357 (2025)
he pe ozoa.penso .ne
353
Dis ibu ion and na u al his o y. Cu en ly, Boule-
noph ys yangchunensis sp. no . is only known om he
ype locali y, Guangdong Yangchun Ehuangzhang P o-
incial Na u e Rese e, wes e n Guangdong, China. I
inhabi s lowing mon ane s eams and he nea by o es
loo and lea li e . The habi a is su ounded by e e -
g een b oad-lea ed o es mixed wi h bamboo g o es a
ele a ions be ween 900 and 1,100 m (Fig. 5). Ad e ise-
men calls o males we e hea d om Feb ua y o May.
Males we e ound calling while pe ching on lea es ha
a e 10–30 cm abo e he g ound.
Conse a ion s a us o ele an
Boulenoph ys species in Guangdong
Recommenda ion o ex inc ion isk assessmen o
Boulenoph ys acu a
Dis ibu ion: Heishiding Na u e Rese e, Feng-
kai Coun y, Zhaoqing, Guangdong P o ince, China
(Li e al. 2014; Lyu e al. 2023); M . Yunkai, Xinyi
Ci y, Guangdong P o ince, China (Song e al. 2024);
Guangdong Yangchun Ehuangzhang P o incial Na-
u e Rese e, Yangchun Ci y, Guangdong P o ince,
China ( his s udy).
Ex inc ion isk assessmen : based on he analysis
on GeoCAT, we de e mined an AOO o 3,170 km2 and
an AOO o 12 km2. As he species is ound a h ee in-
dependen loca ions ( o assess agains c i e ia B2a), and
he e is a con inued obse ed decline in he (iii) quali y
o he habi a , we ecommend he species be lis ed as En-
dange ed ollowing he c i e ia B1ab(iii) and B2ab(iii).
I elying on he c i e ia D ( e y small popula ion and/
o es ic ed dis ibu ion) o he ecommenda ion o as-
sessmen , he species would be lis ed as Vulne able D2
because he AOO is smalle han 20 km² and he species
occu s a ewe han i e loca ions. This ecommenda-
ion o assessmen is a non-genuine downlis ing (ASG
Table 2. Measu emen s (in mm) o ouche specimens o Bou-
lenoph ys yangchunensis sp. no ., * holo ype.
Vouche GEP a168 GEP a169 GEP a170* CIB 121751 SYS a009518
Sex ♂ ♂ ♂ ♂ ♂
SVL 28.6 27.6 27.9 28.6 28.7
HDL 10.4 10.1 11.1 10.6 10.4
HDW 10.3 9.7 11.0 10.4 10.6
ED 3.7 3.6 3.8 4.0 3.7
TD 3.5 3.1 3.7 3.5 3.6
TED 3.1 3.0 3.2 3.4 3.3
SNT 3.8 3.7 4.1 3.6 3.5
IND 2.1 2.0 2.0 2.2 2.0
IOD 1.2 1.3 1.1 1.2 1.2
HND 7.8 7.8 7.9 8.1 7.6
RAD 6.0 6.1 6.5 5.9 5.9
FTL 19.9 20.1 20.7 19.7 19.7
TIB 14.0 13.8 14.5 13.3 14.1
Figu e 5. Mic ohabi a o Boulenoph ys yangchunensis sp. no . in Guangdong Yangchun Ehuangzhang P o incial Na u e Rese e,
Yangchun Ci y, Guangdong P o ince, China. Pho os by Bin-Bin Zhan.