taxonID	type	format	identifier	references	title	description	created	creator	contributor	publisher	audience	source	license	rightsHolder	datasetID
A92D04029B6C5BEAB7D58D671D1BE6A7.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652561	https://doi.org/10.3897/zookeys.1280.182768.figure10	Figure 10. Skull of the adult male of Gloydius chambensis BMNH 1898.5. 17.4: A. Dorsal; B. Ventral (lower jaws virtually extracted); C. Lateral view; D. Pterygoid virtually isolated, ventral view; E. Lower jaw virtually isolated, medial view. Scale bars: 4.0 mm.	Figure 10. Skull of the adult male of Gloydius chambensis BMNH 1898.5. 17.4: A. Dorsal; B. Ventral (lower jaws virtually extracted); C. Lateral view; D. Pterygoid virtually isolated, ventral view; E. Lower jaw virtually isolated, medial view. Scale bars: 4.0 mm.	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
A92D04029B6C5BEAB7D58D671D1BE6A7.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652562	https://doi.org/10.3897/zookeys.1280.182768.figure11	Figure 11. The colour and pattern in Gloydius chambensis and its typical habitat. A. The specimens (uncollected) from Teppa, Chamba District, Himachal Pradesh State, India, 2543 m elevation (photo by Girish Choure); B. Gloydius chambensis (uncollected) from Teppa, Chamba District, Himachal Pradesh State, India, 2543 m elevation (photo by Girish Choure); C. Specimens from Chamba Valley (photo by Rakeshwar Kapoor, https://www.inaturalist.org/observations/52247726); D. General view of the Chamba Valley (https://commons.wikimedia.org/wiki/File%3AChamba.jpg).	Figure 11. The colour and pattern in Gloydius chambensis and its typical habitat. A. The specimens (uncollected) from Teppa, Chamba District, Himachal Pradesh State, India, 2543 m elevation (photo by Girish Choure); B. Gloydius chambensis (uncollected) from Teppa, Chamba District, Himachal Pradesh State, India, 2543 m elevation (photo by Girish Choure); C. Specimens from Chamba Valley (photo by Rakeshwar Kapoor, https://www.inaturalist.org/observations/52247726); D. General view of the Chamba Valley (https://commons.wikimedia.org/wiki/File%3AChamba.jpg).	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
A92D04029B6C5BEAB7D58D671D1BE6A7.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652560	https://doi.org/10.3897/zookeys.1280.182768.figure9	Figure 9. A. Geographic localities of species within the Gloydius himalayanus complex, compiled from literature sources, museum records, citizen science platforms (iNaturalist 2025), and data obtained in this study (see Gazetteer Suppl. material 1: table SS 3 for details). Grey circles indicate ambiguous records or populations of uncertain taxonomic status, where evidence was insufficient for reliable identification (" transition zone "). Dashed lines delineate hypothesised biogeographic boundaries between major lineages, i. e., the Upper Indus Valley, Kashmir Valley, Ravi River Valley, and Karnali River Valley, landscape features likely acting as geographic barriers influencing species distribution; B. Inset map of the Kashmir region (Western Himalaya) showing the distributions and contact zones of the taxa defined here, highlighting a key area for future research. Question marks highlight regions that remain poorly explored or unsampled. The map was generated using QGIS 3.28 (https://qgis.org/) with the WGS 84 coordinate reference system.	Figure 9. A. Geographic localities of species within the Gloydius himalayanus complex, compiled from literature sources, museum records, citizen science platforms (iNaturalist 2025), and data obtained in this study (see Gazetteer Suppl. material 1: table SS 3 for details). Grey circles indicate ambiguous records or populations of uncertain taxonomic status, where evidence was insufficient for reliable identification (" transition zone "). Dashed lines delineate hypothesised biogeographic boundaries between major lineages, i. e., the Upper Indus Valley, Kashmir Valley, Ravi River Valley, and Karnali River Valley, landscape features likely acting as geographic barriers influencing species distribution; B. Inset map of the Kashmir region (Western Himalaya) showing the distributions and contact zones of the taxa defined here, highlighting a key area for future research. Question marks highlight regions that remain poorly explored or unsampled. The map was generated using QGIS 3.28 (https://qgis.org/) with the WGS 84 coordinate reference system.	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
B6B9CCFF76035A5D80AD6567D371F49E.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652569	https://doi.org/10.3897/zookeys.1280.182768.figure18	Figure 18. The adult female holotype of Gloydius hazarensis sp. nov. (UF 70652). A – D. Lateral, dorsal and ventral views of the head; (E, F) dorsal and ventral views of the body.	Figure 18. The adult female holotype of Gloydius hazarensis sp. nov. (UF 70652). A – D. Lateral, dorsal and ventral views of the head; (E, F) dorsal and ventral views of the body.	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
B6B9CCFF76035A5D80AD6567D371F49E.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652570	https://doi.org/10.3897/zookeys.1280.182768.figure19	Figure 19. Skull of the adult female holotype of Gloydius hazarensis sp. nov., UF 70652: A. Dorsal, B. Ventral (lower jaws virtually extracted) and C. Lateral view; D. Pterygoid virtually isolated, ventral view; E. Lower jaw virtually isolated, medial view. Scale bar: 2.0 mm.	Figure 19. Skull of the adult female holotype of Gloydius hazarensis sp. nov., UF 70652: A. Dorsal, B. Ventral (lower jaws virtually extracted) and C. Lateral view; D. Pterygoid virtually isolated, ventral view; E. Lower jaw virtually isolated, medial view. Scale bar: 2.0 mm.	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
B6B9CCFF76035A5D80AD6567D371F49E.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652571	https://doi.org/10.3897/zookeys.1280.182768.figure20	Figure 20. The colour and pattern in Gloydius hazarensis sp. nov. and its typical habitat. A – E. Adult specimens from Naran Valley, Khyber Pakhtunkhwa, Pakistan (photos by Rafaqat Masroor) and the habitat of the type locality of the species near Naran town (F; photo by Daniel Jablonski).	Figure 20. The colour and pattern in Gloydius hazarensis sp. nov. and its typical habitat. A – E. Adult specimens from Naran Valley, Khyber Pakhtunkhwa, Pakistan (photos by Rafaqat Masroor) and the habitat of the type locality of the species near Naran town (F; photo by Daniel Jablonski).	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
B6B9CCFF76035A5D80AD6567D371F49E.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652560	https://doi.org/10.3897/zookeys.1280.182768.figure9	Figure 9. A. Geographic localities of species within the Gloydius himalayanus complex, compiled from literature sources, museum records, citizen science platforms (iNaturalist 2025), and data obtained in this study (see Gazetteer Suppl. material 1: table SS 3 for details). Grey circles indicate ambiguous records or populations of uncertain taxonomic status, where evidence was insufficient for reliable identification (" transition zone "). Dashed lines delineate hypothesised biogeographic boundaries between major lineages, i. e., the Upper Indus Valley, Kashmir Valley, Ravi River Valley, and Karnali River Valley, landscape features likely acting as geographic barriers influencing species distribution; B. Inset map of the Kashmir region (Western Himalaya) showing the distributions and contact zones of the taxa defined here, highlighting a key area for future research. Question marks highlight regions that remain poorly explored or unsampled. The map was generated using QGIS 3.28 (https://qgis.org/) with the WGS 84 coordinate reference system.	Figure 9. A. Geographic localities of species within the Gloydius himalayanus complex, compiled from literature sources, museum records, citizen science platforms (iNaturalist 2025), and data obtained in this study (see Gazetteer Suppl. material 1: table SS 3 for details). Grey circles indicate ambiguous records or populations of uncertain taxonomic status, where evidence was insufficient for reliable identification (" transition zone "). Dashed lines delineate hypothesised biogeographic boundaries between major lineages, i. e., the Upper Indus Valley, Kashmir Valley, Ravi River Valley, and Karnali River Valley, landscape features likely acting as geographic barriers influencing species distribution; B. Inset map of the Kashmir region (Western Himalaya) showing the distributions and contact zones of the taxa defined here, highlighting a key area for future research. Question marks highlight regions that remain poorly explored or unsampled. The map was generated using QGIS 3.28 (https://qgis.org/) with the WGS 84 coordinate reference system.	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
198FFC36425756CBA28203A75C34EC93.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652552	https://doi.org/10.3897/zookeys.1280.182768.figure1	Figure 1. A. Phylogenetic placement of the G. himalayanus complex within the genus Gloydius (for details see Fig. 2 A); B. Geographic distribution (dark shading) of recognised lineages within the G. himalayanus complex, along with the sampling localities of genetically investigated specimens (see Table 1, Suppl. material 1: table SS 1). The position of the locality for the lectotype BMNH 1946.1. 19.64 is only for map purposes (see remarks on the lectotype in the species account). Due to incomplete locality data in Kuttalam et al. (2022) (see Fig. 2 A), some sample locations are not depicted in panel (B); an inset shows the approximate extent of the major mountain ranges in the region, following Spicer et al. (2021, 2025). Question marks indicate uncertain or potential occurrences in eastern Afghanistan. The map was generated using QGIS 3.28 (https://qgis.org/) with the WGS 84 coordinate reference system. Sketch shows a syntype of Halys himalayanus, taken from Günther (1864). This individual corresponds to the specimen BMNH 1946.1. 19.64, which is designated here as the lectotype for Gloydius himalayanus.	Figure 1. A. Phylogenetic placement of the G. himalayanus complex within the genus Gloydius (for details see Fig. 2 A); B. Geographic distribution (dark shading) of recognised lineages within the G. himalayanus complex, along with the sampling localities of genetically investigated specimens (see Table 1, Suppl. material 1: table SS 1). The position of the locality for the lectotype BMNH 1946.1. 19.64 is only for map purposes (see remarks on the lectotype in the species account). Due to incomplete locality data in Kuttalam et al. (2022) (see Fig. 2 A), some sample locations are not depicted in panel (B); an inset shows the approximate extent of the major mountain ranges in the region, following Spicer et al. (2021, 2025). Question marks indicate uncertain or potential occurrences in eastern Afghanistan. The map was generated using QGIS 3.28 (https://qgis.org/) with the WGS 84 coordinate reference system. Sketch shows a syntype of Halys himalayanus, taken from Günther (1864). This individual corresponds to the specimen BMNH 1946.1. 19.64, which is designated here as the lectotype for Gloydius himalayanus.	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
198FFC36425756CBA28203A75C34EC93.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652563	https://doi.org/10.3897/zookeys.1280.182768.figure12	Figure 12. The adult male lectotype of Gloydius himalayanus (BMNH 1946.1. 19.64). A – D. Lateral, dorsal and ventral views of the head; (E, F) dorsal and ventral views of the body.	Figure 12. The adult male lectotype of Gloydius himalayanus (BMNH 1946.1. 19.64). A – D. Lateral, dorsal and ventral views of the head; (E, F) dorsal and ventral views of the body.	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
198FFC36425756CBA28203A75C34EC93.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652557	https://doi.org/10.3897/zookeys.1280.182768.figure6	Figure 6. Dorsolateral scalation and pattern in four species of the Gloydius himalayanus complex: A. Gloydius himalayanus (BMNH 1946.1. 19.64, lectotype); B. G. hazarensis sp. nov. (UF 70652, holotype); C. G. hindukushensis sp. nov. (NHMW 41993, holotype); D. G. nepalensis sp. nov. (ZMB 65613, holotype).	Figure 6. Dorsolateral scalation and pattern in four species of the Gloydius himalayanus complex: A. Gloydius himalayanus (BMNH 1946.1. 19.64, lectotype); B. G. hazarensis sp. nov. (UF 70652, holotype); C. G. hindukushensis sp. nov. (NHMW 41993, holotype); D. G. nepalensis sp. nov. (ZMB 65613, holotype).	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
198FFC36425756CBA28203A75C34EC93.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652564	https://doi.org/10.3897/zookeys.1280.182768.figure13	Figure 13. Skull of the adult male lectotype of Gloydius himalayanus BMNH 1946.1. 19.64: A. Dorsal, B. Ventral (lower jaws virtually extracted) and C. Lateral view; D. Pterygoid virtually isolated, ventral view; E. Lower jaw virtually isolated, medial view. Scale bars: 4.0 mm.	Figure 13. Skull of the adult male lectotype of Gloydius himalayanus BMNH 1946.1. 19.64: A. Dorsal, B. Ventral (lower jaws virtually extracted) and C. Lateral view; D. Pterygoid virtually isolated, ventral view; E. Lower jaw virtually isolated, medial view. Scale bars: 4.0 mm.	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
198FFC36425756CBA28203A75C34EC93.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652565	https://doi.org/10.3897/zookeys.1280.182768.figure14	Figure 14. The colour and pattern in Gloydius himalayanus and its typical habitat. (A, B) Adult male (uncollected) from between Bhebra camp site and Manjhi (Baharat Range), Uttarkashi District, Uttarakhand State, India, 2400 m elevation (photo by Frank Tillack); (C, D) An uncollected specimen from Khaliya [Khulia] Danda, Pithoragarh District, Uttarakhand State, India, 3433 m elevation (photo by Ashok Captain and Emmanuel Theophilus); (E, F) An uncollected specimen from Dungair Dhar, Pithoragarh District, Uttarakhand State, India, 3535 m elevation (photo by Ashok Captain and Emmanuel Theophilus); G. Juvenile (uncollected) from Dhapa Bend, Pithoragarh District, Uttarakhand State, India, 1794 m elevation (photo by Ashok Captain and Emmanuel Theophilus); H. General view of the habitat of Gloydius himalayanus at Khaliya [Khulia] Danda, Pithoragarh District, Uttarakhand State, India, 3433 m elevation (photo by Ashok Captain and Emmanuel Theophilus); I. Microhabitat of G. himalayanus between Bhebra camp site and Manjhi (Baharat Range), Uttarkashi District, Uttarakhand State, India, 2400 m elevation; J. General view of the habitat of G. himalayanus between Bhebra camp site and Manjhi (Baharat Range), Uttarkashi District, Uttarakhand State, India, 2400 m elevation; K. Habitat of G. himalayanus at Dungair Dhar, Pithoragarh District, Uttarakhand State, India, 3535 m elevation (photo by Ashok Captain and Emmanuel Theophilus).	Figure 14. The colour and pattern in Gloydius himalayanus and its typical habitat. (A, B) Adult male (uncollected) from between Bhebra camp site and Manjhi (Baharat Range), Uttarkashi District, Uttarakhand State, India, 2400 m elevation (photo by Frank Tillack); (C, D) An uncollected specimen from Khaliya [Khulia] Danda, Pithoragarh District, Uttarakhand State, India, 3433 m elevation (photo by Ashok Captain and Emmanuel Theophilus); (E, F) An uncollected specimen from Dungair Dhar, Pithoragarh District, Uttarakhand State, India, 3535 m elevation (photo by Ashok Captain and Emmanuel Theophilus); G. Juvenile (uncollected) from Dhapa Bend, Pithoragarh District, Uttarakhand State, India, 1794 m elevation (photo by Ashok Captain and Emmanuel Theophilus); H. General view of the habitat of Gloydius himalayanus at Khaliya [Khulia] Danda, Pithoragarh District, Uttarakhand State, India, 3433 m elevation (photo by Ashok Captain and Emmanuel Theophilus); I. Microhabitat of G. himalayanus between Bhebra camp site and Manjhi (Baharat Range), Uttarkashi District, Uttarakhand State, India, 2400 m elevation; J. General view of the habitat of G. himalayanus between Bhebra camp site and Manjhi (Baharat Range), Uttarkashi District, Uttarakhand State, India, 2400 m elevation; K. Habitat of G. himalayanus at Dungair Dhar, Pithoragarh District, Uttarakhand State, India, 3535 m elevation (photo by Ashok Captain and Emmanuel Theophilus).	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
198FFC36425756CBA28203A75C34EC93.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652560	https://doi.org/10.3897/zookeys.1280.182768.figure9	Figure 9. A. Geographic localities of species within the Gloydius himalayanus complex, compiled from literature sources, museum records, citizen science platforms (iNaturalist 2025), and data obtained in this study (see Gazetteer Suppl. material 1: table SS 3 for details). Grey circles indicate ambiguous records or populations of uncertain taxonomic status, where evidence was insufficient for reliable identification (" transition zone "). Dashed lines delineate hypothesised biogeographic boundaries between major lineages, i. e., the Upper Indus Valley, Kashmir Valley, Ravi River Valley, and Karnali River Valley, landscape features likely acting as geographic barriers influencing species distribution; B. Inset map of the Kashmir region (Western Himalaya) showing the distributions and contact zones of the taxa defined here, highlighting a key area for future research. Question marks highlight regions that remain poorly explored or unsampled. The map was generated using QGIS 3.28 (https://qgis.org/) with the WGS 84 coordinate reference system.	Figure 9. A. Geographic localities of species within the Gloydius himalayanus complex, compiled from literature sources, museum records, citizen science platforms (iNaturalist 2025), and data obtained in this study (see Gazetteer Suppl. material 1: table SS 3 for details). Grey circles indicate ambiguous records or populations of uncertain taxonomic status, where evidence was insufficient for reliable identification (" transition zone "). Dashed lines delineate hypothesised biogeographic boundaries between major lineages, i. e., the Upper Indus Valley, Kashmir Valley, Ravi River Valley, and Karnali River Valley, landscape features likely acting as geographic barriers influencing species distribution; B. Inset map of the Kashmir region (Western Himalaya) showing the distributions and contact zones of the taxa defined here, highlighting a key area for future research. Question marks highlight regions that remain poorly explored or unsampled. The map was generated using QGIS 3.28 (https://qgis.org/) with the WGS 84 coordinate reference system.	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
78DE78E2577A5E2F95AFF27D63FED0F2.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652566	https://doi.org/10.3897/zookeys.1280.182768.figure15	Figure 15. The adult female holotype of Gloydius hindukushensis sp. nov. (NHMW 41993). A – D. Lateral, dorsal and ventral views of the head; (E, F) dorsal and ventral views of the body.	Figure 15. The adult female holotype of Gloydius hindukushensis sp. nov. (NHMW 41993). A – D. Lateral, dorsal and ventral views of the head; (E, F) dorsal and ventral views of the body.	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
78DE78E2577A5E2F95AFF27D63FED0F2.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652567	https://doi.org/10.3897/zookeys.1280.182768.figure16	Figure 16. Skull of the adult female holotype of Gloydius hindukushensis sp. nov., NHMW 41993: A. Dorsal, B. Ventral (lower jaws virtually extracted) and C. Lateral view; D. Pterygoid virtually isolated, ventral view; E. Lower jaw virtually isolated, medial view. Scale bar: 4.0 mm.	Figure 16. Skull of the adult female holotype of Gloydius hindukushensis sp. nov., NHMW 41993: A. Dorsal, B. Ventral (lower jaws virtually extracted) and C. Lateral view; D. Pterygoid virtually isolated, ventral view; E. Lower jaw virtually isolated, medial view. Scale bar: 4.0 mm.	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
78DE78E2577A5E2F95AFF27D63FED0F2.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652568	https://doi.org/10.3897/zookeys.1280.182768.figure17	Figure 17. The colour and pattern in Gloydius hindukushensis sp. nov. and its typical habitat. A. Freshly killed holotype (NHMW 41993) from Kumrat Valley, Khyber Pakhtunkhwa, Pakistan; (B, C) The paratype (CUHC 10088) from Kumrat Valley, Khyber Pakhtunkhwa, Pakistan, all photos by Daniel Jablonski; (D, E) The specimen PMNH 5150 collected on 30 April 2023 from Lal Qilla, Lower Dir, Khyber Pakhtunkhwa, Pakistan; (F, G) The specimen PMNH 5149 collected on 16 October 2023 from Lal Qilla, Lower Dir, Khyber Pakhtunkhwa, Pakistan, photos by Muhammad Idrees; H. The specimens (uncollected) observed on 28 May 2025 from Broon area, Bomborate, Kalash Valley, Chitral, Khyber Pakhtunkhwa, Pakistan, photo by Abdul Basit; I. The habitat of the type locality of G. hindukushensis sp. nov. in Kumrat Valley, photo by Daniel Jablonski; J. The habitat in Lal Qilla, photo by Muhammad Idrees; K. The habitat of the species in Chitral Gol National Park, Chitral, Pakistan, photo by Daniel Jablonski.	Figure 17. The colour and pattern in Gloydius hindukushensis sp. nov. and its typical habitat. A. Freshly killed holotype (NHMW 41993) from Kumrat Valley, Khyber Pakhtunkhwa, Pakistan; (B, C) The paratype (CUHC 10088) from Kumrat Valley, Khyber Pakhtunkhwa, Pakistan, all photos by Daniel Jablonski; (D, E) The specimen PMNH 5150 collected on 30 April 2023 from Lal Qilla, Lower Dir, Khyber Pakhtunkhwa, Pakistan; (F, G) The specimen PMNH 5149 collected on 16 October 2023 from Lal Qilla, Lower Dir, Khyber Pakhtunkhwa, Pakistan, photos by Muhammad Idrees; H. The specimens (uncollected) observed on 28 May 2025 from Broon area, Bomborate, Kalash Valley, Chitral, Khyber Pakhtunkhwa, Pakistan, photo by Abdul Basit; I. The habitat of the type locality of G. hindukushensis sp. nov. in Kumrat Valley, photo by Daniel Jablonski; J. The habitat in Lal Qilla, photo by Muhammad Idrees; K. The habitat of the species in Chitral Gol National Park, Chitral, Pakistan, photo by Daniel Jablonski.	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
78DE78E2577A5E2F95AFF27D63FED0F2.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652560	https://doi.org/10.3897/zookeys.1280.182768.figure9	Figure 9. A. Geographic localities of species within the Gloydius himalayanus complex, compiled from literature sources, museum records, citizen science platforms (iNaturalist 2025), and data obtained in this study (see Gazetteer Suppl. material 1: table SS 3 for details). Grey circles indicate ambiguous records or populations of uncertain taxonomic status, where evidence was insufficient for reliable identification (" transition zone "). Dashed lines delineate hypothesised biogeographic boundaries between major lineages, i. e., the Upper Indus Valley, Kashmir Valley, Ravi River Valley, and Karnali River Valley, landscape features likely acting as geographic barriers influencing species distribution; B. Inset map of the Kashmir region (Western Himalaya) showing the distributions and contact zones of the taxa defined here, highlighting a key area for future research. Question marks highlight regions that remain poorly explored or unsampled. The map was generated using QGIS 3.28 (https://qgis.org/) with the WGS 84 coordinate reference system.	Figure 9. A. Geographic localities of species within the Gloydius himalayanus complex, compiled from literature sources, museum records, citizen science platforms (iNaturalist 2025), and data obtained in this study (see Gazetteer Suppl. material 1: table SS 3 for details). Grey circles indicate ambiguous records or populations of uncertain taxonomic status, where evidence was insufficient for reliable identification (" transition zone "). Dashed lines delineate hypothesised biogeographic boundaries between major lineages, i. e., the Upper Indus Valley, Kashmir Valley, Ravi River Valley, and Karnali River Valley, landscape features likely acting as geographic barriers influencing species distribution; B. Inset map of the Kashmir region (Western Himalaya) showing the distributions and contact zones of the taxa defined here, highlighting a key area for future research. Question marks highlight regions that remain poorly explored or unsampled. The map was generated using QGIS 3.28 (https://qgis.org/) with the WGS 84 coordinate reference system.	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
5E547ECEBB9E52728C7B311678FF74DE.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652572	https://doi.org/10.3897/zookeys.1280.182768.figure21	Figure 21. The adult male holotype of Gloydius nepalensis sp. nov. (ZMB 65613). A – D. Lateral, dorsal and ventral views of the head; (E, F) dorsal and ventral views of the body; G. Hemipenis.	Figure 21. The adult male holotype of Gloydius nepalensis sp. nov. (ZMB 65613). A – D. Lateral, dorsal and ventral views of the head; (E, F) dorsal and ventral views of the body; G. Hemipenis.	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
5E547ECEBB9E52728C7B311678FF74DE.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652573	https://doi.org/10.3897/zookeys.1280.182768.figure22	Figure 22. Skull of the adult male holotype of Gloydius nepalensis sp. nov., ZMB 65613: A. Dorsal, B. Ventral (lower jaws virtually extracted) and C. Lateral view; D. Pterygoid virtually isolated, ventral view; E. Lower jaw virtually isolated, medial view. Scale bar: 3.0 mm.	Figure 22. Skull of the adult male holotype of Gloydius nepalensis sp. nov., ZMB 65613: A. Dorsal, B. Ventral (lower jaws virtually extracted) and C. Lateral view; D. Pterygoid virtually isolated, ventral view; E. Lower jaw virtually isolated, medial view. Scale bar: 3.0 mm.	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
5E547ECEBB9E52728C7B311678FF74DE.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652574	https://doi.org/10.3897/zookeys.1280.182768.figure23	Figure 23. The colour and pattern in Gloydius nepalensis sp. nov. and its habitat. (A, B) The holotype (ZMB 65613, adult male) from Kalopani village, Mustang District, Gandaki Province, Nepal, 2500 m elevation; C – E. The paratype (ZMB 65612, adult female) from Syang village, Mustang District, Gandaki Province, Nepal, 2700 m elevation; F. The paratype (NME R 0544 / 07), adult male from Kermi, 14 km northwest of Simikot, Humla District, Karnali Province, Nepal (photo by Ulrich Scheidt); G. Neonate of the blotched phase (ZMB 65607, female, mother from Syang village, Mustang District, Gandaki Province, Nepal, 2700 m elevation); H. Neonate of the striped phase (ZMB 65606, male; mother from Syang village, Mustang District, Gandaki Province, Nepal, 2700 m elevation); I. Representative habitat of G. nepalensis sp. nov. in the vicinity of Marpha, Mustang District, Gandaki Province, Nepal, courtesy National Trust for Nature Conservation Annapurna Conservation Area Project, Jomsom, Mustang, Nepal.	Figure 23. The colour and pattern in Gloydius nepalensis sp. nov. and its habitat. (A, B) The holotype (ZMB 65613, adult male) from Kalopani village, Mustang District, Gandaki Province, Nepal, 2500 m elevation; C – E. The paratype (ZMB 65612, adult female) from Syang village, Mustang District, Gandaki Province, Nepal, 2700 m elevation; F. The paratype (NME R 0544 / 07), adult male from Kermi, 14 km northwest of Simikot, Humla District, Karnali Province, Nepal (photo by Ulrich Scheidt); G. Neonate of the blotched phase (ZMB 65607, female, mother from Syang village, Mustang District, Gandaki Province, Nepal, 2700 m elevation); H. Neonate of the striped phase (ZMB 65606, male; mother from Syang village, Mustang District, Gandaki Province, Nepal, 2700 m elevation); I. Representative habitat of G. nepalensis sp. nov. in the vicinity of Marpha, Mustang District, Gandaki Province, Nepal, courtesy National Trust for Nature Conservation Annapurna Conservation Area Project, Jomsom, Mustang, Nepal.	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
5E547ECEBB9E52728C7B311678FF74DE.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1652560	https://doi.org/10.3897/zookeys.1280.182768.figure9	Figure 9. A. Geographic localities of species within the Gloydius himalayanus complex, compiled from literature sources, museum records, citizen science platforms (iNaturalist 2025), and data obtained in this study (see Gazetteer Suppl. material 1: table SS 3 for details). Grey circles indicate ambiguous records or populations of uncertain taxonomic status, where evidence was insufficient for reliable identification (" transition zone "). Dashed lines delineate hypothesised biogeographic boundaries between major lineages, i. e., the Upper Indus Valley, Kashmir Valley, Ravi River Valley, and Karnali River Valley, landscape features likely acting as geographic barriers influencing species distribution; B. Inset map of the Kashmir region (Western Himalaya) showing the distributions and contact zones of the taxa defined here, highlighting a key area for future research. Question marks highlight regions that remain poorly explored or unsampled. The map was generated using QGIS 3.28 (https://qgis.org/) with the WGS 84 coordinate reference system.	Figure 9. A. Geographic localities of species within the Gloydius himalayanus complex, compiled from literature sources, museum records, citizen science platforms (iNaturalist 2025), and data obtained in this study (see Gazetteer Suppl. material 1: table SS 3 for details). Grey circles indicate ambiguous records or populations of uncertain taxonomic status, where evidence was insufficient for reliable identification (" transition zone "). Dashed lines delineate hypothesised biogeographic boundaries between major lineages, i. e., the Upper Indus Valley, Kashmir Valley, Ravi River Valley, and Karnali River Valley, landscape features likely acting as geographic barriers influencing species distribution; B. Inset map of the Kashmir region (Western Himalaya) showing the distributions and contact zones of the taxa defined here, highlighting a key area for future research. Question marks highlight regions that remain poorly explored or unsampled. The map was generated using QGIS 3.28 (https://qgis.org/) with the WGS 84 coordinate reference system.	2026-05-21	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia		Zenodo	biologists	Jablonski, Daniel;Tillack, Frank;Mahlow-Tillack, Kristin;Petzold, Alice;Wilzo, Madita;Das, Abhijit;Idrees, Muhammad;Baniya, Chitra B.;Masroor, Rafaqat;Hofmann, Sylvia			
