Animalia Squamata Gekkonidae LuuVinh QuangNguyenThuong HuyenDoQuyen HanhPhamCuong TheHoangTuoi ThiNguyenTruong QuangLeMinh DucZieglerThomasGrismerJesse L.GrismerL. LeeA new species of Hemiphyllodactylus (Squamata, Gekkonidae) from Ha Giang Province, VietnamZookeys22062023116735338210.3897/zookeys.1167.103713 64879394-8DB8-5685-9534-C5A76EAF82B1 Hemiphyllodactylus lungcuensis https://zoobank.org/7E96A770-35D0-4B50-82B5-F8C7A4A817FD sp. nov.Figs 7, 8 Verbatim name. Lungcu Slender GeckoMaterial examined.

Holotype.VNUF R.2021.01 (Field no. LC17.01), adult male, collected by H.M. Ly on 25 August 2017 from Lung Cu Commune, Dong Van District, Ha Giang Province (23°21'N, 105°19'E, at an elevation of 1391 m a.s.l.). Paratypes.IEBR R.5151 (Field no. LC17.02), VNUF R.2021.03 (Field no. LC17.03), adult males, IEBR R.5152 (Field no. ST17.01), VNUF R.2021.08 (Field no. HG17.08), adult females, collected at the same locality as holotype on 25 August 2017; two adult females, VNUF R.2023.01 (Field no. LC22.01) and VNUF R.2023.02 (Field no. LC22.02), collected at the same locality as the holotype on 5 August 2022.

Diagnosis.

Hemiphyllodactyluslungcuensis sp. nov. can be distinguished from its congeners by a combination of the following characters: a maximum SVL of 44.2 mm; trunk not particularly elongate (AG/SVL ratio 0.52); chin scales 8–10, distinct enlarged postmentals; circumnasal scales two or three; supralabial scales 11 or 12; infralabial scales 10 or 11; ventral scale rows 6–9; dorsal scale rows 12–16; digital lamellae formula 4(3)–4–5(4)–4 (forefoot) and 4–5–5–5 (hindfoot); precloacal pores 17–25 and continuous series in males, 0–22 pitted precloacal scales in females; cloacal spur single in both sexes; enlarged subcaudal scales absent; dorsal surface brown sand color, with irregular dark brown streaks; a distinct dark postorbital stripe extending to at least the base of the neck, uneven dark streaks running along the flanks and ending at the base of tail, a cream-colored V-shaped postsacral mark with anteriorly projecting arms; a pale yellow ventral view of the body and limbs; and unpigmented caecum and gonads.

10.3897/zookeys.1167.103713.figure720E380F7-77BD-597E-B357-14BF327AF5E7

Dorsal views of Hemiphyllodactyluslungcuensis sp. nov. in life from Ha Giang Province, northeastern Vietnam A adult male holotype VNUF R.2021.01 B adult female paratype VNUF R.2021.08.

https://binary.pensoft.net/fig/867700
10.3897/zookeys.1167.103713.figure8C0CAE215-E671-596A-B734-578452D25608

Type series of Hemiphyllodactyluslungcuensis sp. nov. in preservative from Ha Giang Province, northeastern Vietnam. A from right to left: Adult male holotype (VNUF R.2021.01), adult female paratype (IEBR R.5251), adult female paratype (VNUF R.2021.08), adult male paratype (IEBR R.5151), adult male paratype (VNUF R.2021.3). B From right to left: Adult female paratype (VNUF R.2023.02), and adult female paratype (VNUF R.2023.01).

https://binary.pensoft.net/fig/867701
Description of the holotype.

Adult male, SVL 39.2 mm; tail length (TaL 37.3 mm); body compressed (TrunkL 20.3 mm); head triangular in dorsal profile, depressed, distinct from neck, and head longer than wide (HeadL 11.0 mm, HeadW 8.0 mm); eye large, pupils vertical (EyeD 1.8 mm); snout-eye distance (SnEye 4.3 mm); snout moderate, round in dorsal profile, longer than eye diameter (EyeD/SnEye 0.42); nare-eye distance (NarEye 3.2 mm); internarial distance (SnW 1.5 mm); ear opening oblique (EarD 0.7 mm).

Proportions: EarD/EyeD 0.39, TrunkL/SVL 0.52, HeadL/SVL 0.28, HeadW/SVL 0.20, HeadW/HeadL 0.73, SnEye/HeadL 0.39, NarEye/HeadL 0.29, EyeD/HeadL 0.16, SnW/HeadL 0.14, EyeD/NarEye 0.56, SnW/HeadW 0.19.

Scalation: Rostral large, wider than high, partially divided dorsally, bordered posteriorly by two large supranasals and two internasals (SnS); nares in contact with rostral anteriorly, first supralabial ventrally, supranasal dorsally, and three nasals posteriorly on each side; supralabial scales 12/12 (left/right), infralabial scales 11/10 (left/right), gradually decreasing in size towards angle of jaw; scales on head small, round, largest on rostrum; mental triangular, bordered by first infralabials and posteriorly by two enlarged postmentals, eight chin scales touching internal edge of infralabials and mental between the juncture of the second and third infralabials on each side of the head, anterior pair enlarged; ventral scales smooth and flat, much larger than dorsal scales, subimbricate, in seven rows (contained within one eye diameter); dorsal scales small, granular in 13 rows at midbody on dorsum (contained within one eye diameter), enlarged tubercles absent; fore-limbs relatively short, covered dorsally with granular, subimbricate scales, smaller smooth scales ventrally; 25 pore bearing femoral and precloacal sacles, in a continuous row; terminal two phalanges free, claws absent on first finger and on first toe,large triangular lamellae present on the second to fifth digits of both the forefoot and hindfoot pads, with digital formula 4–4–5–4 (forefoot) and 4–5–5–5 (hindfoot); four on first fingers, four on first toes; cloacal spurs (CloacS) one on each side; semi-regenerated tail; caudal scales flat, not forming distinct caudal segments; caecum and gonads unpigmented.

Coloration in life.

Dorsal surface of head, body, and limbs brown sand color, with irregularly shaped dark-brown streaks; a dark postorbital stripe, distinct, extending to at least base of neck; uneven dark streaks running along the flanks and ending at the base of tail; two dark dots on dorsal surface of tail; dorsal surface of tail has dark bands evenly spaced from base of tail to tip of tail; postsacral mark cream-colored and V-shaped, bearing anteriorly projecting arms; ventral surface of body and limbs pale yellow; subcaudal region pale peach.

Sexual dimorphism and variation.

The scale counts vary among the type series: chin scales 9 or 10; scales between supranasals one or three; supralabials 11; infralabials 10; ventral scale rows seven or nine; 0–22 pitted precloacal scales in females (distinct pore-bearing scales in males). Dorsal surface of the tail has dark bands evenly spaced from the base of the tail to the tip of the tail in females. Subcaudal region of original part of tail of the adult male paratype (IEBR R.5151) is reddish-orange, while the regenerated portion of the tail is uniformly dark grey. For other morphological characters of the paratypes are given in Table 8.

Morphological characters of Hemiphyllodactyluslungcuensis sp. nov. (measurements in mm, * = regenerated or broken, ?= missing data, Min = minimum, Max = maximum, other abbreviations defined in the text).

VNUF R.2021.01IEBR R.5151VNUF R.2021.03VNUF R.2023.01VNUF R.2023.02IEBR R.5152VNUF R.2021.08Min-Max
HolotypeParatypeParatypeParatypeParatypeParatypeParatype
Sex malemalemalefemalefemalefemalefemale
SVL 39.24344.235.337.439.243.535.3–44.2
TaL 37.325*42.730.4*27.3*4.3*34.734.7–42.7
TrunkL 20.323.023.117.019.319.022.317.0–23.1
HeadL 11.011.611.18.98.58.510.38.5–11.6
HeadW 8.08.07.06.97.07.38.06.9–8.0
EyeD 1.82.12.41.81.82.52.11.8–2.5
SnEye 4.34.55.03.63.94.14.23.6–5.0
NarEye 3.22.93.63.13.03.23.32.9–3.6
SnW 1.52.01.71.41.41.92.11.4–2.1
EarD 0.70.90.80.60.50.60.80.5–0.9
EarD/EyeD 0.390.430.330.330.280.240.380.24–0.43
TrunkL/SVL 0.520.530.520.480.520.480.510.48–0.53
HeadL/SVL 0.280.270.250.250.230.220.240.22–0.28
HeadW/SVL 0.200.190.160.200.190.190.180.16–0.20
HeadW/HeadL 0.730.690.630.780.820.860.780.63–0.86
SnEye/HeadL 0.390.390.450.400.460.480.410.39–0.48
NarEye/HeadL 0.290.250.320.350.350.380.320.25–0.38
EyeD/HeadL 0.160.180.220.200.210.290.200.21–0.29
SnW/HeadL 0.140.170.150.160.160.220.200.14–0.22
EyeD/NarEye 0.560.720.670.580.600.780.640.56–0.78
SnW/HeadW 0.190.250.240.200.200.260.260.19–0.26
Chin 88?10109108–10
CN 33333232–3
SnS 22221131–3
SL (left/right) 12/1211/1112/1212/1111/1111/1111/1111–12
IL (left/right) 11/1011/1110/1010/910/910/1010/109–11
VS 7671011976–11
DS 1313151716161212–17
Lamellae formula of forelimbs II–V (left) 4454*44??44544454444434544(3)45(4)4
Lamellae formula of hind limbs II–V (left) 45554555455545554555455545554555
SL1F 44?44433–4
SL1T 44444444
Total femoroprecloacal pores 25172316002217–25 (in males)
CloacS on each side 11121111–2
Precloacal and femoral pore series separate (1) or continuous (0) 000000
Subcaudals enlarged, plate-like 00000000
Distribution.

Hemiphyllodactyluslungcuensis sp. nov. is currently known only from the type locality in Lung Cu Commune, Dong Van District, Ha Giang Province, northeastern Vietnam (Fig. 1).

Etymology.

Specific epithet lungcuensis is a toponym in reference to the type locality of the species. For the common names, we suggest Lungcu Slender Gecko (English) and Thạch sùng dẹp lũng cú (Vietnamese).

Natural history.

The species is known from a karstic habitat. The type series were found on karst cliffs between 19:00 and 20:30, approximately 1–1.5 m above the ground. The surrounding habitat was a disturbed karst forest with small hardwood and shrub trees, and was located near residential areas and road systems (Fig. 9). The type locality of the new species is located in an unprotected area, and thus we need to clarify the impact and threats of the localized disturbance so as to put in place effective conservation measures.

10.3897/zookeys.1167.103713.figure9A8570F1B-56DC-5177-A056-F2FCE8C31E23

Habitat of Hemiphyllodactyluslungcuensis sp. nov. in Lung Cu Commune, Dong Van District, Ha Giang Province, northeastern Vietnam.

https://binary.pensoft.net/fig/867702
Comparisons.

The molecular analyses indicate that Hemiphyllodactyluslungcuensis sp. nov. is a phylogenetically distinct lineage of the clade 6 that is not embedded within any lineage of the other 13 species of clade 6 (Agung et al. 2022) and bears an uncorrected pairwise sequence divergence of 4.6–20.2% from them (Fig. 1, Table 2). Hemiphyllodactyluslungcuensis sp. nov. is the sister species to a clade composed of H.huishuiensis and H.yanshanensis (Fig. 1) from which it bears an uncorrected pairwise sequence divergence of 5.3% from H.huishuiensis and 6.9% from H.yanshanensis (Table 2). Hemiphyllodactyluslungcuensis sp. nov. differs significantly from H.huishuiensis in mean values of EyeD (0.31 vs. 0.41, p = 0.001) and SnW (0.23 vs. 0.08, p = 0.001) and from H.yanshanensis in mean values of TrunkL (1.31 vs. 1.29, p = 0.03) and EyeD (0.31 vs. 0.38, p = 0.002). In addition, Hemiphyllodactyluslungcuensis sp. nov. differs significantly from H.nahangensis in mean values of EyeD (0.31 vs. 0.42, p = 0.001), SnW (0.23 vs. 0.05, p = 0.001), DS (14.75 vs. 21, p = 0.001), and SL1T (4 vs. 3, p = 0.001) (Tables 6, 7). Statistically significant differences of morphometric and meristic traits between Hemiphyllodactyluslungcuensis sp. nov. and all other species in clade 6 are presented in Tables 6, 7. Due to the unavailability of individual specimens of H.dushanensis for statistical analysis, we obtained integrated data of this species from Do et al. (2020: table 4) for comparison, the new species can be distinguished from H.dushanensis by the presence of dark postorbital stripe, dark dorsolateral stripe, and dark dorsal transverse blotches (vs. absent in H.dushanensis).

10.3897/zookeys.1167.103713.figure1A7805DAA-8DBA-51EE-B7D0-3793CDCBAE9C

Distribution of the species of clade 6 of the genus Hemiphyllodactylus.

https://binary.pensoft.net/fig/867694

Uncorrected genetic P-distances (%) in ND2 gene of Hemiphyllodactylus. Intraspecific p-distance are in bold font, n/a = data not applicable.

banaensis bonkowskii dupanglingensis dushanensis harterti hongkongensis huishuiensis kiziriani lungcuensis sp. nov. nahangensis ngocsonensis serpispecus typus yanshanensis zugi
banaensis n/a
bonkowskii 13.0 0.01
dupanglingensis 16.915.3 0.00
dushanensis 17.314.75.1 0.00
harterti 30.426.527.829.1 0.04
hongkongensis 16.314.67.97.127.5 0.01
huishuiensis 15.715.47.16.428.97.9 0.01
kiziriani 22.720.020.320.529.620.020.5 0.00
lungcuensis sp. nov. 15.813.86.24.827.27.05.320.2 0.02
nahangensis 15.513.65.85.525.37.86.320.14.6 0.01
ngocsonensis 14.512.87.67.927.09.27.819.36.67.0 0.02
serpispecus 12.27.516.616.025.716.415.421.414.413.212.6 n/a
typus 15.214.67.26.327.98.97.421.25.64.18.815.6 n/a
yanshanensis 17.115.28.68.326.09.44.422.26.87.38.514.78.2 0.00
zugi 16.014.04.85.827.07.47.019.96.86.27.515.07.38.1 0.00

Summary statistics of morphometric and meristic characters among the Hymiphyllodactylus species in clade 6.

Species SVL TrunkL HeadL HeadW SnEye NarEye EyeD SnW Chin CN SL IL VS DS SL1F SL1T
banaensis (n = 10)
Mean1.691.361.050.870.640.520.410.226.2310.810.310.118.554.9
SD0.0170.0140.0120.0090.0270.0210.0310.0190.42200.9190.6740.8761.17900.316
Lower1.661.341.030.860.600.480.360.19639991754
Upper1.711.391.070.890.690.560.450.26731211122055
bonkowskii (n = 6)
Mean1.641.341.010.930.670.520.450.316.539.589.2514.3325.1754.83
SD0.0430.0200.0110.0120.0190.0200.0270.0230.83700.4910.8800.8160.51600.408
Lower1.561.321.000.910.630.480.420.2753981324.554
Upper1.681.371.030.950.690.540.490.33731010.5152655
dupanglingensis (n = 6)
Mean1.611.281.030.870.650.500.410.2110.333.3311.679.8310.6714.834.834.67
SD0.0200.0110.0210.0130.0150.0130.0110.0380.820.5161.2111.1690.5160.7530.4080.516
Lower1.601.270.990.850.630.490.400.1693119101444
Upper1.651.301.050.880.670.520.430.261141412111655
hongkongensis (n = 8)
Mean1.581.291.000.850.550.440.360.085.633.1311.0610.139.3813.54.135
SD0.0440.0120.0120.0610.0220.0240.0210.0220.5180.350.7760.3530.5180.9260.6410
Lower1.521.270.980.720.510.410.320.0553109.591235
Upper1.631.301.010.910.580.470.390.12641210.5101555
huishuiensis (n = 8)
Mean1.641.331.010.890.610.470.410.089.5310.139.948.1313.6333
SD0.0580.0110.0440.0160.0270.0240.0200.0360.9300.6410.6780.8350.74400
Lower1.551.320.970.870.580.440.370.03839971333
Upper1.711.351.110.920.660.520.440.13103111191533
kiziriani (n = 10)
Mean1.581.280.820.840.590.480.410.157.7410.559.812.922.855
SD0.0250.0240.0190.0190.0210.0210.0240.0311.33700.4380.6321.1252.31200
Lower1.551.250.780.810.550.440.370.10064109111855
Upper1.611.320.850.870.630.510.450.22941110.514.52655
lungcuensis (n = 7)
Mean1.601.310.100.870.630.500.310.239.142.8611.3610.078.1414.573.864
SD0.0370.0130.0390.0260.0220.0280.0490.0460.9000.3780.4760.5351.8641.9020.3780
Lower1.551.290.940.830.590.450.270.1682119.561234
Upper1.651.331.060.910.660.540.410.301031211111744
nahangensis (n = 3)
Mean1.631.331.010.850.580.480.420.058.332.8311.1710.1710.332133
SD0.0124.15E-050.0080.0070.0130.0070.0060.0340.5770.2890.2890.5772.3092.64600
Lower1.621.331.000.840.570.470.410.01782.5119.591833
Upper1.641.331.020.860.590.490.420.099311.510.5132333
ngocsonensis (n = 4)
Mean1.661.371.010.930.660.530.440.247.5310.139.6314204.755.25
SD0.0060.0040.0080.0100.0160.0290.0100.007100.250.4790.8160.5770.50.5
Lower1.661.361.000.910.650.500.430.23631091319.545
Upper1.671.371.020.940.690.570.450.258310.5101520.556
serpispecus (n = 1)
Mean1.621.331.010.880.590.510.380.1773119102644
SDNANANANANANANANANANANANANANANANA
Lower1.621.331.010.880.590.510.380.1773119102644
Upper1.621.331.010.880.590.510.380.1773119102644
typus (n = 4)
Mean1.501.260.900.690.490.380.26-0.0312.755.2511.3610.75111544.25
SD0.0960.0260.0280.0160.0270.0150.0330.1152.6300.50.6290.51.4140.81600.5
Lower1.351.220.880.680.460.360.21-0.1910510.51091444
Upper1.561.280.940.720.520.400.290.071561211121645
yanshanensis (n = 13)
Mean1.611.290.870.900.620.500.380.188.775.07710.7710.158.2315.384.695.08
SD0.0350.0120.0210.0170.0180.0240.0390.0191.0130.2770.6960.5911.7872.9590.4800.277
Lower1.531.270.840.870.590.460.310.16859.59.571145
Upper1.671.310.930.940.650.550.450.211161211132256
zugi (n = 3)
Mean1.611.330.970.860.640.460.440.19102.511.3310.8315.3321.174.674.83
SD0.0340.0100.0120.0010.0130.0310.0010.0091.7320.51.0410.2890.5770.5770.5770.289
Lower1.581.320.960.850.620.420.4360.1859210.510.51520.544.5
Upper1.651.340.980.860.650.480.4380.20212312.5111621.555

Summary statistics of the principal component analysis for Hemiphyllodactylus species.

PC1 PC2 PC3 PC4 PC5 PC6 PC7 PC8
Standard deviation 2.3540410571.6938967421.2748197881.1676318951.0357726990.8585830050.7925727690.700362564
Proportion of Variance 0.346340.179330.101570.085210.067050.046070.039260.03066
Cumulative Proportion 0.346340.525670.627250.712460.779510.825580.864840.8955
eigen 5.5415092962.8692861741.6251654911.3633642411.0728250850.7371647770.6281715930.49050772
SVL -0.2756849390.230671227-0.1832688560.035839657-0.1181105030.272234498-0.3535480730.532231802
TrunkL -0.3080280870.2265779260.1131820.165672749-0.098377540.213756620.231486740.352374067
HeadL -0.1822060730.4016792470.1110644770.286694098-0.082466549-0.2705669180.230170931-0.024756545
HeadW -0.3162348790.087646379-0.266341502-0.2275935830.0115930530.056636999-0.034411085-0.551718699
SnEye -0.3540534410.028071822-0.1742721850.0287377970.370906143-0.0057905570.061744748-0.038483117
NarEye -0.3432024250.013213311-0.227705587-0.0974141990.248586294-0.037810729-0.0505989060.110097139
EyeD -0.316496337-0.0570540940.203121932-0.0540967720.112788940.198377934-0.488753315-0.270887117
SnW -0.331682586-0.118626-0.2200994510.0781237660.10976895-0.0790251680.348298246-0.089036833
Chin 0.2095989840.0411245150.0555045020.0157689540.7627550380.1269367540.2529757250.142123111
CN 0.219334589-0.306220475-0.340275868-0.2382457220.1065135280.269505437-0.0867748890.202680044
SL 0.1508236150.072482948-0.1415262310.5695829870.281130944-0.34518498-0.525400396-0.03369833
IL 0.1480197240.047098158-0.1604486410.539731081-0.0782045470.694783630.119615798-0.305781505
VS -0.159093977-0.324270770.5072084260.1799126330.1072842330.0467599870.085474597-0.042903745
DS -0.216885816-0.3166655930.4159172220.0136703450.0488852590.15087592-0.1397509760.074486639
SL1F -0.160449559-0.454450117-0.1642154080.230860029-0.051722119-0.1312815780.0733696090.169471363
SL1T -0.088298003-0.43959919-0.2766771580.243587056-0.224574637-0.1546361550.0811866950.024901385
PC9 PC10 PC11 PC12 PC13 PC14 PC15 PC16
Standard deviation 0.6458376460.5288067310.49495050.4608694020.4228360180.3923449470.3487834810.252014728
Proportion of Variance 0.026070.017480.015310.013280.011170.009620.00760.00397
Cumulative Proportion 0.921570.939050.954360.967630.978810.988430.996031
eigen 0.4171062650.2796365590.2449759980.2124006060.1787902980.1539345580.1216499160.063511423
SVL -0.0076537970.5048652840.0230548570.048923188-0.104515394-0.1349550960.216496623-0.075553468
TrunkL -0.155806654-0.4026687910.169935171-0.501088590.0302087210.261221952-0.1058883670.183446071
HeadL -0.196823028-0.0429285890.3089090570.583413488-0.2354825690.083561907-0.07755344-0.18027682
HeadW -0.0573559340.0187951520.420271-0.230081521-0.1699181090.0207324740.4451174820.004175194
SnEye 0.006655124-0.054733740.125911056-0.0813370710.26677457-0.615512405-0.423767342-0.211307545
NarEye 0.209982569-0.475190325-0.4612928990.279036274-0.29657524-0.0400456040.2118999660.218583377
EyeD -0.4676943930.020586761-0.2012463650.1509904540.1463310730.329187066-0.2742452120.024651913
SnW 0.3736297340.470794413-0.10280995-0.0133932820.0311196970.456870887-0.2725192770.133686181
Chin -0.2931975690.139059923-0.038969703-0.051313747-0.0527184310.180261940.28186905-0.211579231
CN -0.036235154-0.1354005180.4857355920.236559781-0.2762039180.132252018-0.3502331890.158104094
SL 0.189523962-0.0599002510.171907156-0.215076887-0.0770033570.119789393-0.029041250.125441842
IL 0.112573099-0.066456621-0.1171630240.1490199890.026731134-0.0822138230.038834193-0.012764391
VS -0.0601170230.2245847180.102501327-0.019925796-0.380657113-0.3336678220.0323689520.484590421
DS 0.471295976-0.1693348130.2071805160.049104744-0.0281956290.157392670.109826908-0.543616511
SL1F -0.192515088-0.0713054290.1828546050.252200640.5655969310.0287933690.3778982550.17530195
SL1T -0.364000904-0.005236656-0.226318873-0.228651157-0.413553347-0.023541078-0.058595931-0.412884904
AgungAPChorneliaAGrismerLLGrismerJLQuahESHLuJTomlinsonKWHughesAC (2022) Description of two new species of Hemiphyllodactylus (Reptilia: Gekkonidae) from karst landscapes in Yunnan, China, highlights complex conservation needs.Zoological Research43(5): 767786. https://doi.org/10.24272/j.issn.2095-8137.2022.105DoQHPhamCTPhanTQLeMDZieglerTNguyenTQ (2020) A new species of Hemiphyllodactylus (Squamata: Gekkonidae) from Tuyen Quang Province, Vietnam.Zootaxa4821(3): 511532. https://doi.org/10.11646/zootaxa.4821.3.5