Integrative taxonomy reveals previously undescribed diversity within the Gloydius himalayanus complex (Squamata, Viperidae, Crotalinae) from the Himalaya and Hindu Kush Author Jablonski, Daniel 0000-0002-5394-0114 Department of Zoology, Comenius University in Bratislava, Ilkovičova 6, Mlynská dolina, 842 15, Bratislava, Slovakia Author Tillack, Frank 0000-0003-1418-6531 Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Invalidenstraße 43, 10115 Berlin, Germany Author Mahlow-Tillack, Kristin 0000-0002-0625-965X Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Invalidenstraße 43, 10115 Berlin, Germany Author Petzold, Alice 0000-0002-0743-2004 Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Invalidenstraße 43, 10115 Berlin, Germany & Institute for Biochemistry and Biology, University of Potsdam, Karl-Liebknecht-Str. 24 – 25, 14476 Potsdam, Germany Author Wilzo, Madita Institute for Biochemistry and Biology, University of Potsdam, Karl-Liebknecht-Str. 24 – 25, 14476 Potsdam, Germany Author Das, Abhijit Wildlife Institute of India, Chandrabani, Dehradun 248001, Uttarakhand, India Author Idrees, Muhammad Department of Zoology, University of Peshawar, Peshawar 25120, Khyber Pakhtunkhwa, Pakistan Author Baniya, Chitra B. 0000-0002-8746-7601 Central Department of Botany, Tribhuvan University, Kirtipur, Kathmandu, Nepal Author Masroor, Rafaqat 0000-0001-6248-546X Zoological Sciences Division, Pakistan Museum of Natural History, Shakarparian, Islamabad, Pakistan Author Hofmann, Sylvia 0000-0003-0621-9049 Museum Koenig, Leibniz Institute for the Analysis of Biodiversity Change, Adenauerallee 127, 53113 Bonn, Germany text ZooKeys 2026 2026-05-21 1280 83 153 journal article 315245 10.3897/zookeys.1280.182768 09a16018-0bd4-4559-8810-31c3fc4de7a0 451B220F-E2BB-4453-A019-39DC922D050E

Table 4. Summary statistics of diagnostic meristic and morphological characters for females of the species of the Gloydius himalayanus complex. Highlighted character states are those that differ significantly (p < 0.05) from G. himalayanus (h), G. hindukushensis sp. nov. (u), G. hazarensis sp. nov. (z), or G. nepalensis sp. nov. (n).

Females	G. himalayanus	G. hazarensis sp. nov.	G. hindukushensis sp. nov.	G. nepalensis sp. nov.
Ven
n	14	7	4	12
Range (mean ± SD)	150–164 (158.9 n, z ± 4.0)	158–165 (162.4 h, n ± 2.4)	147–159 (152.5 ± 5.5)	146–161 (152.5 h, z ± 5.1)
Ven + SubC
n	14	7	3	12
Range (mean ± SD)	195–216 (206 n ± 6.9)	201–215 (209.7 n ± 4.7)	198–204 (200.3 ± 3.2)	188–205 (195.7 h, z ± 5.7)
SubL
n	14	7	4	12
Range (mean ± SD)	8–11 (9.6 n ± 0.8)	8–10 (9.7 n ± 0.8)	8–9 (8.8 ± 0.5)	7–10 (8.8 h, z ± 0.8)
PrO
n	14	7	4	12
Range (mean ± SD)	2–3 (2.1 z ± 0.4)	3 (3 n ± 0)	3 (3 n ± 0)	2 (2 u, z ± 0)
DSR 3
n	14	7	4	12
Range (mean ± SD)	17 (17 u ± 0)	17 (17 ± 0)	17–19 (18 h, n ± 1.2)	15–17 (16.8 u ± 0.6)
PrefL (mm)
n	16	6	4	7
Range (mean ± SD)	1.5–4.6 (3.5 z ± 0.8)	2.0–3.7 (2.7 h ± 0.6)	1.8–4.16 (2.7 ± 1.0)	3.1–3.7 (3.3 ± 0.2)
PrefW (mm)
n	16	6	4	7
Range (mean ± SD)	2.2–4.5 (3.4 u, z ± 0.5)	2.1–3.3 (2.7 h, n ± 0.5)	1.9–3.31 (2.3 h, n ± 0.7)	3.1–3.8 (3.4 u, z ± 0.3)
INW (mm)
n	16	6	3	7
Range (mean ± SD)	2.3–3.9 (3.1 u, z ± 0.4)	1.3–2.3 (1.7 h ± 0.3)	1.5–1.73 (1.6 h ± 0.1)	2.3–3.2 (2.8 ± 0.2)
INL (mm)
n	16	6	3	7
Range (mean ± SD)	1.2–2.6 (1.7 u, z ± 0.3)	1.0–3.3 (2.4 h ± 1.0)	1.9–4.02 (2.6 h ± 1.2)	1.5–2.9 (2.0 ± 0.5)
INW. L (mm)
n	16	6	3	7
Range (mean ± SD)	1.4–2.8 (1.9 u, z ± 0.4)	0.5–2.3 (0.9 h ± 0.7)	0.43–0.8 (0.7 h ± 0.2)	0.8–1.8 (1.5 ± 0.4)