taxonID	type	format	identifier	references	title	description	created	creator	contributor	publisher	audience	source	license	rightsHolder	datasetID
BB28BA3B958B54458D7A490593DE77BB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481167	https://doi.org/10.3897/dez.72.159716.figure4	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
BB28BA3B958B54458D7A490593DE77BB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481168	https://doi.org/10.3897/dez.72.159716.figure5	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
BB28BA3B958B54458D7A490593DE77BB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481189	https://doi.org/10.3897/dez.72.159716.figure26	Figure 26. Pterostichus (Morphohaptoderus) chachi sp. nov. (A, E – F. Paratype; B – D. Holotype). A. Habitus, female; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E, F. Elytral apex. Scale bars: 1.0 mm (A – D).	Figure 26. Pterostichus (Morphohaptoderus) chachi sp. nov. (A, E – F. Paratype; B – D. Holotype). A. Habitus, female; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E, F. Elytral apex. Scale bars: 1.0 mm (A – D).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
1814CB2348C45A89BFA4B61D08F27C97.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481164	https://doi.org/10.3897/dez.72.159716.figure1	Figure 1. Characters of P. (Morphohaptoderus) spp. A. Antennomere 3 of P. wangshu sp. nov. B. Antennomere 3 of P. yaotiao sp. nov. C. Apical lamella of P. lisao sp. nov. D. Apical lamella of P. lianquan sp. nov. E. Fifth tarsomere of P. yaotiao sp. nov. F. Fifth tarsomere of P. hubeicus; G. Fifth tarsomere of P. shennongjianus; H. Elytral microsculpture of P. dentellus; I. Elytral microsculpture of P. shennongjianus. Scale bars: 0.25 mm (A – G); 0.05 mm (H – I).	Figure 1. Characters of P. (Morphohaptoderus) spp. A. Antennomere 3 of P. wangshu sp. nov. B. Antennomere 3 of P. yaotiao sp. nov. C. Apical lamella of P. lisao sp. nov. D. Apical lamella of P. lianquan sp. nov. E. Fifth tarsomere of P. yaotiao sp. nov. F. Fifth tarsomere of P. hubeicus; G. Fifth tarsomere of P. shennongjianus; H. Elytral microsculpture of P. dentellus; I. Elytral microsculpture of P. shennongjianus. Scale bars: 0.25 mm (A – G); 0.05 mm (H – I).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
1814CB2348C45A89BFA4B61D08F27C97.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481165	https://doi.org/10.3897/dez.72.159716.figure2	Figure 2. Male sternum of P. (Morphohaptoderus) spp. A, D. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; E. P. lisao sp. nov.; F. P. lingjun sp. nov. Scale bars: 1 mm.	Figure 2. Male sternum of P. (Morphohaptoderus) spp. A, D. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; E. P. lisao sp. nov.; F. P. lingjun sp. nov. Scale bars: 1 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
1814CB2348C45A89BFA4B61D08F27C97.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481166	https://doi.org/10.3897/dez.72.159716.figure3	Figure 3. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov. Scale bars: 0.5 mm.	Figure 3. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
1814CB2348C45A89BFA4B61D08F27C97.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481168	https://doi.org/10.3897/dez.72.159716.figure5	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
1814CB2348C45A89BFA4B61D08F27C97.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481169	https://doi.org/10.3897/dez.72.159716.figure6	Figure 6. Pterostichus (Morphohaptoderus) dentellus Facchini & Sciaky, from Shennongjia, Hubei. A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – D); 2.0 mm (E, F).	Figure 6. Pterostichus (Morphohaptoderus) dentellus Facchini & Sciaky, from Shennongjia, Hubei. A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – D); 2.0 mm (E, F).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
1814CB2348C45A89BFA4B61D08F27C97.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481170	https://doi.org/10.3897/dez.72.159716.figure7	Figure 7. Endophallus of P. (Morphohaptoderus) spp. A, B. P. dentellus. A. Left lateral view; B. Right lateral view. C, D. P. quyuani sp. nov. C. Left lateral view; D. Right lateral view. Abbreviations: gp gonopore, gpl gonoporal piece, bl basal lobe, lp left pre-apical lobe, la left apical lobe, ra right apical lobe. Scale bars: 0.5 mm.	Figure 7. Endophallus of P. (Morphohaptoderus) spp. A, B. P. dentellus. A. Left lateral view; B. Right lateral view. C, D. P. quyuani sp. nov. C. Left lateral view; D. Right lateral view. Abbreviations: gp gonopore, gpl gonoporal piece, bl basal lobe, lp left pre-apical lobe, la left apical lobe, ra right apical lobe. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
1814CB2348C45A89BFA4B61D08F27C97.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481175	https://doi.org/10.3897/dez.72.159716.figure12	Figure 12. Median lobe of aedeagus, left lateral view. A – C. P. dentellus; D, E. P. wangshu sp. nov.; F – I. P. quyuani sp. nov.; J. P. lingjun sp. nov. A. Non-type specimen, Shennongjia, Yanziya, 2300 m; B. Non-type specimen, Shennongjia, Jiuhuping, 1800–1950 m; C. Non-type specimen, Yintiaoling, Shizhuzi, 2137 m; D. Paratype, Wufeng, Houhe Nature Reserve, 900–1000 m; E. Holotype, Wufeng, Houhe Nature Reserve, 900–1000 m; F. Holotype, Shennongjia, Wenshui Village, 1700 m; G – I. Paratype, Shennongjia, Wenshui Village, 1700 m; J. Holotype, Shennongjia, Guanmen Mountain, 1361 m. Scale bars: 0.5 mm.	Figure 12. Median lobe of aedeagus, left lateral view. A – C. P. dentellus; D, E. P. wangshu sp. nov.; F – I. P. quyuani sp. nov.; J. P. lingjun sp. nov. A. Non-type specimen, Shennongjia, Yanziya, 2300 m; B. Non-type specimen, Shennongjia, Jiuhuping, 1800–1950 m; C. Non-type specimen, Yintiaoling, Shizhuzi, 2137 m; D. Paratype, Wufeng, Houhe Nature Reserve, 900–1000 m; E. Holotype, Wufeng, Houhe Nature Reserve, 900–1000 m; F. Holotype, Shennongjia, Wenshui Village, 1700 m; G – I. Paratype, Shennongjia, Wenshui Village, 1700 m; J. Holotype, Shennongjia, Guanmen Mountain, 1361 m. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
0B8225D7EAA950529997B36E383D65E5.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481164	https://doi.org/10.3897/dez.72.159716.figure1	Figure 1. Characters of P. (Morphohaptoderus) spp. A. Antennomere 3 of P. wangshu sp. nov. B. Antennomere 3 of P. yaotiao sp. nov. C. Apical lamella of P. lisao sp. nov. D. Apical lamella of P. lianquan sp. nov. E. Fifth tarsomere of P. yaotiao sp. nov. F. Fifth tarsomere of P. hubeicus; G. Fifth tarsomere of P. shennongjianus; H. Elytral microsculpture of P. dentellus; I. Elytral microsculpture of P. shennongjianus. Scale bars: 0.25 mm (A – G); 0.05 mm (H – I).	Figure 1. Characters of P. (Morphohaptoderus) spp. A. Antennomere 3 of P. wangshu sp. nov. B. Antennomere 3 of P. yaotiao sp. nov. C. Apical lamella of P. lisao sp. nov. D. Apical lamella of P. lianquan sp. nov. E. Fifth tarsomere of P. yaotiao sp. nov. F. Fifth tarsomere of P. hubeicus; G. Fifth tarsomere of P. shennongjianus; H. Elytral microsculpture of P. dentellus; I. Elytral microsculpture of P. shennongjianus. Scale bars: 0.25 mm (A – G); 0.05 mm (H – I).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
0B8225D7EAA950529997B36E383D65E5.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481167	https://doi.org/10.3897/dez.72.159716.figure4	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
0B8225D7EAA950529997B36E383D65E5.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481181	https://doi.org/10.3897/dez.72.159716.figure18	Figure 18. Pterostichus (Morphohaptoderus) hubeicus Facchini & Sciaky, from Shennongjia, Hubei. A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	Figure 18. Pterostichus (Morphohaptoderus) hubeicus Facchini & Sciaky, from Shennongjia, Hubei. A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
0B8225D7EAA950529997B36E383D65E5.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481182	https://doi.org/10.3897/dez.72.159716.figure19	Figure 19. Endophallus of P. (Morphohaptoderus) spp. A, B. P. hubeicus. A. Left lateral view; B. Right lateral view. C, D. P. yaotiao sp. nov. C. Left lateral view; D. Right lateral view. Abbreviations: gp gonopore, bl basal lobe, lp left pre-apical lobe, la left apical lobe, ra right apical lobe. Scale bars: 0.5 mm.	Figure 19. Endophallus of P. (Morphohaptoderus) spp. A, B. P. hubeicus. A. Left lateral view; B. Right lateral view. C, D. P. yaotiao sp. nov. C. Left lateral view; D. Right lateral view. Abbreviations: gp gonopore, bl basal lobe, lp left pre-apical lobe, la left apical lobe, ra right apical lobe. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
011E9897A6B2525F8E15B45A0192CB39.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481164	https://doi.org/10.3897/dez.72.159716.figure1	Figure 1. Characters of P. (Morphohaptoderus) spp. A. Antennomere 3 of P. wangshu sp. nov. B. Antennomere 3 of P. yaotiao sp. nov. C. Apical lamella of P. lisao sp. nov. D. Apical lamella of P. lianquan sp. nov. E. Fifth tarsomere of P. yaotiao sp. nov. F. Fifth tarsomere of P. hubeicus; G. Fifth tarsomere of P. shennongjianus; H. Elytral microsculpture of P. dentellus; I. Elytral microsculpture of P. shennongjianus. Scale bars: 0.25 mm (A – G); 0.05 mm (H – I).	Figure 1. Characters of P. (Morphohaptoderus) spp. A. Antennomere 3 of P. wangshu sp. nov. B. Antennomere 3 of P. yaotiao sp. nov. C. Apical lamella of P. lisao sp. nov. D. Apical lamella of P. lianquan sp. nov. E. Fifth tarsomere of P. yaotiao sp. nov. F. Fifth tarsomere of P. hubeicus; G. Fifth tarsomere of P. shennongjianus; H. Elytral microsculpture of P. dentellus; I. Elytral microsculpture of P. shennongjianus. Scale bars: 0.25 mm (A – G); 0.05 mm (H – I).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
011E9897A6B2525F8E15B45A0192CB39.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481167	https://doi.org/10.3897/dez.72.159716.figure4	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
011E9897A6B2525F8E15B45A0192CB39.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481168	https://doi.org/10.3897/dez.72.159716.figure5	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
011E9897A6B2525F8E15B45A0192CB39.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481184	https://doi.org/10.3897/dez.72.159716.figure21	Figure 21. Pterostichus (Morphohaptoderus) lianquan sp. nov. (A – D. Holotype; E. Paratype, Houhe Nature Reserve, Wufeng County, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, dorsal view; C. Right paramere; D. Habitus, male; E. Habitus, female. Scale bars: 0.5 mm (A – C); 2.0 mm (D, E).	Figure 21. Pterostichus (Morphohaptoderus) lianquan sp. nov. (A – D. Holotype; E. Paratype, Houhe Nature Reserve, Wufeng County, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, dorsal view; C. Right paramere; D. Habitus, male; E. Habitus, female. Scale bars: 0.5 mm (A – C); 2.0 mm (D, E).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
011E9897A6B2525F8E15B45A0192CB39.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481185	https://doi.org/10.3897/dez.72.159716.figure22	Figure 22. Pterostichus (Morphohaptoderus) zhizheng sp. nov. (A – D. Holotype; E. Paratype, Houhe Nature Reserve, Wufeng County, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, dorsal view; C. Right paramere; D. Habitus, male; E. Habitus, female. Scale bars: 0.5 mm (A – C); 2.0 mm (D, E).	Figure 22. Pterostichus (Morphohaptoderus) zhizheng sp. nov. (A – D. Holotype; E. Paratype, Houhe Nature Reserve, Wufeng County, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, dorsal view; C. Right paramere; D. Habitus, male; E. Habitus, female. Scale bars: 0.5 mm (A – C); 2.0 mm (D, E).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
518B5E183ABA51399B236AFF4868F5BD.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481165	https://doi.org/10.3897/dez.72.159716.figure2	Figure 2. Male sternum of P. (Morphohaptoderus) spp. A, D. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; E. P. lisao sp. nov.; F. P. lingjun sp. nov. Scale bars: 1 mm.	Figure 2. Male sternum of P. (Morphohaptoderus) spp. A, D. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; E. P. lisao sp. nov.; F. P. lingjun sp. nov. Scale bars: 1 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
518B5E183ABA51399B236AFF4868F5BD.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481166	https://doi.org/10.3897/dez.72.159716.figure3	Figure 3. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov. Scale bars: 0.5 mm.	Figure 3. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
518B5E183ABA51399B236AFF4868F5BD.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481168	https://doi.org/10.3897/dez.72.159716.figure5	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
518B5E183ABA51399B236AFF4868F5BD.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481175	https://doi.org/10.3897/dez.72.159716.figure12	Figure 12. Median lobe of aedeagus, left lateral view. A – C. P. dentellus; D, E. P. wangshu sp. nov.; F – I. P. quyuani sp. nov.; J. P. lingjun sp. nov. A. Non-type specimen, Shennongjia, Yanziya, 2300 m; B. Non-type specimen, Shennongjia, Jiuhuping, 1800–1950 m; C. Non-type specimen, Yintiaoling, Shizhuzi, 2137 m; D. Paratype, Wufeng, Houhe Nature Reserve, 900–1000 m; E. Holotype, Wufeng, Houhe Nature Reserve, 900–1000 m; F. Holotype, Shennongjia, Wenshui Village, 1700 m; G – I. Paratype, Shennongjia, Wenshui Village, 1700 m; J. Holotype, Shennongjia, Guanmen Mountain, 1361 m. Scale bars: 0.5 mm.	Figure 12. Median lobe of aedeagus, left lateral view. A – C. P. dentellus; D, E. P. wangshu sp. nov.; F – I. P. quyuani sp. nov.; J. P. lingjun sp. nov. A. Non-type specimen, Shennongjia, Yanziya, 2300 m; B. Non-type specimen, Shennongjia, Jiuhuping, 1800–1950 m; C. Non-type specimen, Yintiaoling, Shizhuzi, 2137 m; D. Paratype, Wufeng, Houhe Nature Reserve, 900–1000 m; E. Holotype, Wufeng, Houhe Nature Reserve, 900–1000 m; F. Holotype, Shennongjia, Wenshui Village, 1700 m; G – I. Paratype, Shennongjia, Wenshui Village, 1700 m; J. Holotype, Shennongjia, Guanmen Mountain, 1361 m. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
518B5E183ABA51399B236AFF4868F5BD.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481178	https://doi.org/10.3897/dez.72.159716.figure15	Figure 15. Pterostichus (Morphohaptoderus) lingjun sp. nov. (A – F. Holotype; G. Paratype, Guanmen Mountain, Shennongjia, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	Figure 15. Pterostichus (Morphohaptoderus) lingjun sp. nov. (A – F. Holotype; G. Paratype, Guanmen Mountain, Shennongjia, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
518B5E183ABA51399B236AFF4868F5BD.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481171	https://doi.org/10.3897/dez.72.159716.figure8	Figure 8. Pterostichus (Morphohaptoderus) quyuani sp. nov. (A – F. Holotype; G. Paratype, Wenshui Village, Shennongjia, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	Figure 8. Pterostichus (Morphohaptoderus) quyuani sp. nov. (A – F. Holotype; G. Paratype, Wenshui Village, Shennongjia, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
518B5E183ABA51399B236AFF4868F5BD.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481176	https://doi.org/10.3897/dez.72.159716.figure13	Figure 13. Median lobe of aedeagus, left lateral view. A – E. P. lisao sp. nov.; F – J. P. shennongjianus. A. Holotype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; B, C. Paratype, Yintiaoling, Shizhuzi, 2137 m; D, E. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; F. Paratype, Shennongjia, Yixiantian, 1500 m; G. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; H. Paratype, Shennongjia, Hongping Town, 2500 m; I – J. Paratype, Yintiaoling, Zhuanping Guard Station to main peak, 2021–2200 m. Scale bars: 0.5 mm.	Figure 13. Median lobe of aedeagus, left lateral view. A – E. P. lisao sp. nov.; F – J. P. shennongjianus. A. Holotype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; B, C. Paratype, Yintiaoling, Shizhuzi, 2137 m; D, E. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; F. Paratype, Shennongjia, Yixiantian, 1500 m; G. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; H. Paratype, Shennongjia, Hongping Town, 2500 m; I – J. Paratype, Yintiaoling, Zhuanping Guard Station to main peak, 2021–2200 m. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
518B5E183ABA51399B236AFF4868F5BD.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481177	https://doi.org/10.3897/dez.72.159716.figure14	Figure 14. Pterostichus (Morphohaptoderus) lisao sp. nov. (A – F. Holotype; G. Paratype, Wenshui Village, Shennongjia, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Abbreviations: rr right ridge, mr median ridge, lr left ridge. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	Figure 14. Pterostichus (Morphohaptoderus) lisao sp. nov. (A – F. Holotype; G. Paratype, Wenshui Village, Shennongjia, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Abbreviations: rr right ridge, mr median ridge, lr left ridge. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
62F01D972423564EAC81A51619912EA8.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481164	https://doi.org/10.3897/dez.72.159716.figure1	Figure 1. Characters of P. (Morphohaptoderus) spp. A. Antennomere 3 of P. wangshu sp. nov. B. Antennomere 3 of P. yaotiao sp. nov. C. Apical lamella of P. lisao sp. nov. D. Apical lamella of P. lianquan sp. nov. E. Fifth tarsomere of P. yaotiao sp. nov. F. Fifth tarsomere of P. hubeicus; G. Fifth tarsomere of P. shennongjianus; H. Elytral microsculpture of P. dentellus; I. Elytral microsculpture of P. shennongjianus. Scale bars: 0.25 mm (A – G); 0.05 mm (H – I).	Figure 1. Characters of P. (Morphohaptoderus) spp. A. Antennomere 3 of P. wangshu sp. nov. B. Antennomere 3 of P. yaotiao sp. nov. C. Apical lamella of P. lisao sp. nov. D. Apical lamella of P. lianquan sp. nov. E. Fifth tarsomere of P. yaotiao sp. nov. F. Fifth tarsomere of P. hubeicus; G. Fifth tarsomere of P. shennongjianus; H. Elytral microsculpture of P. dentellus; I. Elytral microsculpture of P. shennongjianus. Scale bars: 0.25 mm (A – G); 0.05 mm (H – I).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
62F01D972423564EAC81A51619912EA8.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481165	https://doi.org/10.3897/dez.72.159716.figure2	Figure 2. Male sternum of P. (Morphohaptoderus) spp. A, D. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; E. P. lisao sp. nov.; F. P. lingjun sp. nov. Scale bars: 1 mm.	Figure 2. Male sternum of P. (Morphohaptoderus) spp. A, D. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; E. P. lisao sp. nov.; F. P. lingjun sp. nov. Scale bars: 1 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
62F01D972423564EAC81A51619912EA8.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481166	https://doi.org/10.3897/dez.72.159716.figure3	Figure 3. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov. Scale bars: 0.5 mm.	Figure 3. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
62F01D972423564EAC81A51619912EA8.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481168	https://doi.org/10.3897/dez.72.159716.figure5	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
62F01D972423564EAC81A51619912EA8.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481174	https://doi.org/10.3897/dez.72.159716.figure11	Figure 11. Endophallus of P. (Morphohaptoderus) spp. A, B. P. shennongjianus. A. Left lateral view; B. Right lateral view. C, D. P. lisao sp. nov. C. Left lateral view; D. Right lateral view. Abbreviations: gp gonopore, bl basal lobe, lp left pre-apical lobe, la left apical lobe, ra right apical lobe. Scale bars: 0.5 mm.	Figure 11. Endophallus of P. (Morphohaptoderus) spp. A, B. P. shennongjianus. A. Left lateral view; B. Right lateral view. C, D. P. lisao sp. nov. C. Left lateral view; D. Right lateral view. Abbreviations: gp gonopore, bl basal lobe, lp left pre-apical lobe, la left apical lobe, ra right apical lobe. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
62F01D972423564EAC81A51619912EA8.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481176	https://doi.org/10.3897/dez.72.159716.figure13	Figure 13. Median lobe of aedeagus, left lateral view. A – E. P. lisao sp. nov.; F – J. P. shennongjianus. A. Holotype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; B, C. Paratype, Yintiaoling, Shizhuzi, 2137 m; D, E. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; F. Paratype, Shennongjia, Yixiantian, 1500 m; G. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; H. Paratype, Shennongjia, Hongping Town, 2500 m; I – J. Paratype, Yintiaoling, Zhuanping Guard Station to main peak, 2021–2200 m. Scale bars: 0.5 mm.	Figure 13. Median lobe of aedeagus, left lateral view. A – E. P. lisao sp. nov.; F – J. P. shennongjianus. A. Holotype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; B, C. Paratype, Yintiaoling, Shizhuzi, 2137 m; D, E. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; F. Paratype, Shennongjia, Yixiantian, 1500 m; G. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; H. Paratype, Shennongjia, Hongping Town, 2500 m; I – J. Paratype, Yintiaoling, Zhuanping Guard Station to main peak, 2021–2200 m. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
62F01D972423564EAC81A51619912EA8.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481177	https://doi.org/10.3897/dez.72.159716.figure14	Figure 14. Pterostichus (Morphohaptoderus) lisao sp. nov. (A – F. Holotype; G. Paratype, Wenshui Village, Shennongjia, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Abbreviations: rr right ridge, mr median ridge, lr left ridge. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	Figure 14. Pterostichus (Morphohaptoderus) lisao sp. nov. (A – F. Holotype; G. Paratype, Wenshui Village, Shennongjia, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Abbreviations: rr right ridge, mr median ridge, lr left ridge. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
27DCEAF8E3A25697A0064F8C6A9A0643.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481165	https://doi.org/10.3897/dez.72.159716.figure2	Figure 2. Male sternum of P. (Morphohaptoderus) spp. A, D. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; E. P. lisao sp. nov.; F. P. lingjun sp. nov. Scale bars: 1 mm.	Figure 2. Male sternum of P. (Morphohaptoderus) spp. A, D. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; E. P. lisao sp. nov.; F. P. lingjun sp. nov. Scale bars: 1 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
27DCEAF8E3A25697A0064F8C6A9A0643.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481166	https://doi.org/10.3897/dez.72.159716.figure3	Figure 3. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov. Scale bars: 0.5 mm.	Figure 3. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
27DCEAF8E3A25697A0064F8C6A9A0643.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481168	https://doi.org/10.3897/dez.72.159716.figure5	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
27DCEAF8E3A25697A0064F8C6A9A0643.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481170	https://doi.org/10.3897/dez.72.159716.figure7	Figure 7. Endophallus of P. (Morphohaptoderus) spp. A, B. P. dentellus. A. Left lateral view; B. Right lateral view. C, D. P. quyuani sp. nov. C. Left lateral view; D. Right lateral view. Abbreviations: gp gonopore, gpl gonoporal piece, bl basal lobe, lp left pre-apical lobe, la left apical lobe, ra right apical lobe. Scale bars: 0.5 mm.	Figure 7. Endophallus of P. (Morphohaptoderus) spp. A, B. P. dentellus. A. Left lateral view; B. Right lateral view. C, D. P. quyuani sp. nov. C. Left lateral view; D. Right lateral view. Abbreviations: gp gonopore, gpl gonoporal piece, bl basal lobe, lp left pre-apical lobe, la left apical lobe, ra right apical lobe. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
27DCEAF8E3A25697A0064F8C6A9A0643.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481171	https://doi.org/10.3897/dez.72.159716.figure8	Figure 8. Pterostichus (Morphohaptoderus) quyuani sp. nov. (A – F. Holotype; G. Paratype, Wenshui Village, Shennongjia, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	Figure 8. Pterostichus (Morphohaptoderus) quyuani sp. nov. (A – F. Holotype; G. Paratype, Wenshui Village, Shennongjia, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
27DCEAF8E3A25697A0064F8C6A9A0643.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481175	https://doi.org/10.3897/dez.72.159716.figure12	Figure 12. Median lobe of aedeagus, left lateral view. A – C. P. dentellus; D, E. P. wangshu sp. nov.; F – I. P. quyuani sp. nov.; J. P. lingjun sp. nov. A. Non-type specimen, Shennongjia, Yanziya, 2300 m; B. Non-type specimen, Shennongjia, Jiuhuping, 1800–1950 m; C. Non-type specimen, Yintiaoling, Shizhuzi, 2137 m; D. Paratype, Wufeng, Houhe Nature Reserve, 900–1000 m; E. Holotype, Wufeng, Houhe Nature Reserve, 900–1000 m; F. Holotype, Shennongjia, Wenshui Village, 1700 m; G – I. Paratype, Shennongjia, Wenshui Village, 1700 m; J. Holotype, Shennongjia, Guanmen Mountain, 1361 m. Scale bars: 0.5 mm.	Figure 12. Median lobe of aedeagus, left lateral view. A – C. P. dentellus; D, E. P. wangshu sp. nov.; F – I. P. quyuani sp. nov.; J. P. lingjun sp. nov. A. Non-type specimen, Shennongjia, Yanziya, 2300 m; B. Non-type specimen, Shennongjia, Jiuhuping, 1800–1950 m; C. Non-type specimen, Yintiaoling, Shizhuzi, 2137 m; D. Paratype, Wufeng, Houhe Nature Reserve, 900–1000 m; E. Holotype, Wufeng, Houhe Nature Reserve, 900–1000 m; F. Holotype, Shennongjia, Wenshui Village, 1700 m; G – I. Paratype, Shennongjia, Wenshui Village, 1700 m; J. Holotype, Shennongjia, Guanmen Mountain, 1361 m. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
27DCEAF8E3A25697A0064F8C6A9A0643.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481173	https://doi.org/10.3897/dez.72.159716.figure10	Figure 10. Pterostichus (Morphohaptoderus) shennongjianus Facchini & Sciaky, from Shennongjia, Hubei. A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F – G).	Figure 10. Pterostichus (Morphohaptoderus) shennongjianus Facchini & Sciaky, from Shennongjia, Hubei. A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F – G).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
27DCEAF8E3A25697A0064F8C6A9A0643.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481176	https://doi.org/10.3897/dez.72.159716.figure13	Figure 13. Median lobe of aedeagus, left lateral view. A – E. P. lisao sp. nov.; F – J. P. shennongjianus. A. Holotype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; B, C. Paratype, Yintiaoling, Shizhuzi, 2137 m; D, E. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; F. Paratype, Shennongjia, Yixiantian, 1500 m; G. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; H. Paratype, Shennongjia, Hongping Town, 2500 m; I – J. Paratype, Yintiaoling, Zhuanping Guard Station to main peak, 2021–2200 m. Scale bars: 0.5 mm.	Figure 13. Median lobe of aedeagus, left lateral view. A – E. P. lisao sp. nov.; F – J. P. shennongjianus. A. Holotype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; B, C. Paratype, Yintiaoling, Shizhuzi, 2137 m; D, E. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; F. Paratype, Shennongjia, Yixiantian, 1500 m; G. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; H. Paratype, Shennongjia, Hongping Town, 2500 m; I – J. Paratype, Yintiaoling, Zhuanping Guard Station to main peak, 2021–2200 m. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
27DCEAF8E3A25697A0064F8C6A9A0643.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481172	https://doi.org/10.3897/dez.72.159716.figure9	Figure 9. Habitus of Pterostichus (Morphohaptoderus) shennongjianus Facchini & Sciaky. A. Holotype; B. Median lobe of aedeagus, left lateral view, paratype; C. Paratype. Scale bars: 2 mm.	Figure 9. Habitus of Pterostichus (Morphohaptoderus) shennongjianus Facchini & Sciaky. A. Holotype; B. Median lobe of aedeagus, left lateral view, paratype; C. Paratype. Scale bars: 2 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
E01713ABC2AA542B9045BD7B3D4CB1A1.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481167	https://doi.org/10.3897/dez.72.159716.figure4	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
E01713ABC2AA542B9045BD7B3D4CB1A1.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481168	https://doi.org/10.3897/dez.72.159716.figure5	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
E01713ABC2AA542B9045BD7B3D4CB1A1.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481188	https://doi.org/10.3897/dez.72.159716.figure25	Figure 25. Pterostichus (Morphohaptoderus) rongyu sp. nov. (A – D. Holotype; E. Paratype, Huanglianchanggou, Yintiaoling, Chongqing). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, dorsal view; C. Right paramere; D. Habitus, male; E. Habitus, female. Scale bars: 0.25 mm (A – C); 1.0 mm (D, E).	Figure 25. Pterostichus (Morphohaptoderus) rongyu sp. nov. (A – D. Holotype; E. Paratype, Huanglianchanggou, Yintiaoling, Chongqing). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, dorsal view; C. Right paramere; D. Habitus, male; E. Habitus, female. Scale bars: 0.25 mm (A – C); 1.0 mm (D, E).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
E95CC9A38F355AEC8440AD5F04B0DB5C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481164	https://doi.org/10.3897/dez.72.159716.figure1	Figure 1. Characters of P. (Morphohaptoderus) spp. A. Antennomere 3 of P. wangshu sp. nov. B. Antennomere 3 of P. yaotiao sp. nov. C. Apical lamella of P. lisao sp. nov. D. Apical lamella of P. lianquan sp. nov. E. Fifth tarsomere of P. yaotiao sp. nov. F. Fifth tarsomere of P. hubeicus; G. Fifth tarsomere of P. shennongjianus; H. Elytral microsculpture of P. dentellus; I. Elytral microsculpture of P. shennongjianus. Scale bars: 0.25 mm (A – G); 0.05 mm (H – I).	Figure 1. Characters of P. (Morphohaptoderus) spp. A. Antennomere 3 of P. wangshu sp. nov. B. Antennomere 3 of P. yaotiao sp. nov. C. Apical lamella of P. lisao sp. nov. D. Apical lamella of P. lianquan sp. nov. E. Fifth tarsomere of P. yaotiao sp. nov. F. Fifth tarsomere of P. hubeicus; G. Fifth tarsomere of P. shennongjianus; H. Elytral microsculpture of P. dentellus; I. Elytral microsculpture of P. shennongjianus. Scale bars: 0.25 mm (A – G); 0.05 mm (H – I).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
E95CC9A38F355AEC8440AD5F04B0DB5C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481165	https://doi.org/10.3897/dez.72.159716.figure2	Figure 2. Male sternum of P. (Morphohaptoderus) spp. A, D. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; E. P. lisao sp. nov.; F. P. lingjun sp. nov. Scale bars: 1 mm.	Figure 2. Male sternum of P. (Morphohaptoderus) spp. A, D. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; E. P. lisao sp. nov.; F. P. lingjun sp. nov. Scale bars: 1 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
E95CC9A38F355AEC8440AD5F04B0DB5C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481166	https://doi.org/10.3897/dez.72.159716.figure3	Figure 3. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov. Scale bars: 0.5 mm.	Figure 3. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
E95CC9A38F355AEC8440AD5F04B0DB5C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481168	https://doi.org/10.3897/dez.72.159716.figure5	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
E95CC9A38F355AEC8440AD5F04B0DB5C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481172	https://doi.org/10.3897/dez.72.159716.figure9	Figure 9. Habitus of Pterostichus (Morphohaptoderus) shennongjianus Facchini & Sciaky. A. Holotype; B. Median lobe of aedeagus, left lateral view, paratype; C. Paratype. Scale bars: 2 mm.	Figure 9. Habitus of Pterostichus (Morphohaptoderus) shennongjianus Facchini & Sciaky. A. Holotype; B. Median lobe of aedeagus, left lateral view, paratype; C. Paratype. Scale bars: 2 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
E95CC9A38F355AEC8440AD5F04B0DB5C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481173	https://doi.org/10.3897/dez.72.159716.figure10	Figure 10. Pterostichus (Morphohaptoderus) shennongjianus Facchini & Sciaky, from Shennongjia, Hubei. A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F – G).	Figure 10. Pterostichus (Morphohaptoderus) shennongjianus Facchini & Sciaky, from Shennongjia, Hubei. A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F – G).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
E95CC9A38F355AEC8440AD5F04B0DB5C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481174	https://doi.org/10.3897/dez.72.159716.figure11	Figure 11. Endophallus of P. (Morphohaptoderus) spp. A, B. P. shennongjianus. A. Left lateral view; B. Right lateral view. C, D. P. lisao sp. nov. C. Left lateral view; D. Right lateral view. Abbreviations: gp gonopore, bl basal lobe, lp left pre-apical lobe, la left apical lobe, ra right apical lobe. Scale bars: 0.5 mm.	Figure 11. Endophallus of P. (Morphohaptoderus) spp. A, B. P. shennongjianus. A. Left lateral view; B. Right lateral view. C, D. P. lisao sp. nov. C. Left lateral view; D. Right lateral view. Abbreviations: gp gonopore, bl basal lobe, lp left pre-apical lobe, la left apical lobe, ra right apical lobe. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
E95CC9A38F355AEC8440AD5F04B0DB5C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481176	https://doi.org/10.3897/dez.72.159716.figure13	Figure 13. Median lobe of aedeagus, left lateral view. A – E. P. lisao sp. nov.; F – J. P. shennongjianus. A. Holotype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; B, C. Paratype, Yintiaoling, Shizhuzi, 2137 m; D, E. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; F. Paratype, Shennongjia, Yixiantian, 1500 m; G. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; H. Paratype, Shennongjia, Hongping Town, 2500 m; I – J. Paratype, Yintiaoling, Zhuanping Guard Station to main peak, 2021–2200 m. Scale bars: 0.5 mm.	Figure 13. Median lobe of aedeagus, left lateral view. A – E. P. lisao sp. nov.; F – J. P. shennongjianus. A. Holotype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; B, C. Paratype, Yintiaoling, Shizhuzi, 2137 m; D, E. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; F. Paratype, Shennongjia, Yixiantian, 1500 m; G. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; H. Paratype, Shennongjia, Hongping Town, 2500 m; I – J. Paratype, Yintiaoling, Zhuanping Guard Station to main peak, 2021–2200 m. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
F193283C18895C82A4A890A11AA5A68D.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481167	https://doi.org/10.3897/dez.72.159716.figure4	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
F193283C18895C82A4A890A11AA5A68D.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481168	https://doi.org/10.3897/dez.72.159716.figure5	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
F193283C18895C82A4A890A11AA5A68D.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481186	https://doi.org/10.3897/dez.72.159716.figure23	Figure 23. Pterostichus (Morphohaptoderus) toledanoi Facchini & Sciaky, from Shennongjia, Hubei. A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	Figure 23. Pterostichus (Morphohaptoderus) toledanoi Facchini & Sciaky, from Shennongjia, Hubei. A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481164	https://doi.org/10.3897/dez.72.159716.figure1	Figure 1. Characters of P. (Morphohaptoderus) spp. A. Antennomere 3 of P. wangshu sp. nov. B. Antennomere 3 of P. yaotiao sp. nov. C. Apical lamella of P. lisao sp. nov. D. Apical lamella of P. lianquan sp. nov. E. Fifth tarsomere of P. yaotiao sp. nov. F. Fifth tarsomere of P. hubeicus; G. Fifth tarsomere of P. shennongjianus; H. Elytral microsculpture of P. dentellus; I. Elytral microsculpture of P. shennongjianus. Scale bars: 0.25 mm (A – G); 0.05 mm (H – I).	Figure 1. Characters of P. (Morphohaptoderus) spp. A. Antennomere 3 of P. wangshu sp. nov. B. Antennomere 3 of P. yaotiao sp. nov. C. Apical lamella of P. lisao sp. nov. D. Apical lamella of P. lianquan sp. nov. E. Fifth tarsomere of P. yaotiao sp. nov. F. Fifth tarsomere of P. hubeicus; G. Fifth tarsomere of P. shennongjianus; H. Elytral microsculpture of P. dentellus; I. Elytral microsculpture of P. shennongjianus. Scale bars: 0.25 mm (A – G); 0.05 mm (H – I).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481165	https://doi.org/10.3897/dez.72.159716.figure2	Figure 2. Male sternum of P. (Morphohaptoderus) spp. A, D. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; E. P. lisao sp. nov.; F. P. lingjun sp. nov. Scale bars: 1 mm.	Figure 2. Male sternum of P. (Morphohaptoderus) spp. A, D. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; E. P. lisao sp. nov.; F. P. lingjun sp. nov. Scale bars: 1 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481193	https://doi.org/10.3897/dez.72.159716.figure30	Figure 30. Habitus of P. (Neohaptoderus) spp. A. P. yulongshanensis; B – F. P. megaloderus. A. Paratype; B. Paratype; C. Median lobe of aedeagus, left lateral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Metacoxae. Scale bars: 0.5 mm.	Figure 30. Habitus of P. (Neohaptoderus) spp. A. P. yulongshanensis; B – F. P. megaloderus. A. Paratype; B. Paratype; C. Median lobe of aedeagus, left lateral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Metacoxae. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481175	https://doi.org/10.3897/dez.72.159716.figure12	Figure 12. Median lobe of aedeagus, left lateral view. A – C. P. dentellus; D, E. P. wangshu sp. nov.; F – I. P. quyuani sp. nov.; J. P. lingjun sp. nov. A. Non-type specimen, Shennongjia, Yanziya, 2300 m; B. Non-type specimen, Shennongjia, Jiuhuping, 1800–1950 m; C. Non-type specimen, Yintiaoling, Shizhuzi, 2137 m; D. Paratype, Wufeng, Houhe Nature Reserve, 900–1000 m; E. Holotype, Wufeng, Houhe Nature Reserve, 900–1000 m; F. Holotype, Shennongjia, Wenshui Village, 1700 m; G – I. Paratype, Shennongjia, Wenshui Village, 1700 m; J. Holotype, Shennongjia, Guanmen Mountain, 1361 m. Scale bars: 0.5 mm.	Figure 12. Median lobe of aedeagus, left lateral view. A – C. P. dentellus; D, E. P. wangshu sp. nov.; F – I. P. quyuani sp. nov.; J. P. lingjun sp. nov. A. Non-type specimen, Shennongjia, Yanziya, 2300 m; B. Non-type specimen, Shennongjia, Jiuhuping, 1800–1950 m; C. Non-type specimen, Yintiaoling, Shizhuzi, 2137 m; D. Paratype, Wufeng, Houhe Nature Reserve, 900–1000 m; E. Holotype, Wufeng, Houhe Nature Reserve, 900–1000 m; F. Holotype, Shennongjia, Wenshui Village, 1700 m; G – I. Paratype, Shennongjia, Wenshui Village, 1700 m; J. Holotype, Shennongjia, Guanmen Mountain, 1361 m. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481176	https://doi.org/10.3897/dez.72.159716.figure13	Figure 13. Median lobe of aedeagus, left lateral view. A – E. P. lisao sp. nov.; F – J. P. shennongjianus. A. Holotype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; B, C. Paratype, Yintiaoling, Shizhuzi, 2137 m; D, E. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; F. Paratype, Shennongjia, Yixiantian, 1500 m; G. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; H. Paratype, Shennongjia, Hongping Town, 2500 m; I – J. Paratype, Yintiaoling, Zhuanping Guard Station to main peak, 2021–2200 m. Scale bars: 0.5 mm.	Figure 13. Median lobe of aedeagus, left lateral view. A – E. P. lisao sp. nov.; F – J. P. shennongjianus. A. Holotype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; B, C. Paratype, Yintiaoling, Shizhuzi, 2137 m; D, E. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; F. Paratype, Shennongjia, Yixiantian, 1500 m; G. Paratype, Shennongjia, Shennongding Scenic Area Entrance, 1884 m; H. Paratype, Shennongjia, Hongping Town, 2500 m; I – J. Paratype, Yintiaoling, Zhuanping Guard Station to main peak, 2021–2200 m. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481177	https://doi.org/10.3897/dez.72.159716.figure14	Figure 14. Pterostichus (Morphohaptoderus) lisao sp. nov. (A – F. Holotype; G. Paratype, Wenshui Village, Shennongjia, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Abbreviations: rr right ridge, mr median ridge, lr left ridge. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	Figure 14. Pterostichus (Morphohaptoderus) lisao sp. nov. (A – F. Holotype; G. Paratype, Wenshui Village, Shennongjia, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Abbreviations: rr right ridge, mr median ridge, lr left ridge. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481188	https://doi.org/10.3897/dez.72.159716.figure25	Figure 25. Pterostichus (Morphohaptoderus) rongyu sp. nov. (A – D. Holotype; E. Paratype, Huanglianchanggou, Yintiaoling, Chongqing). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, dorsal view; C. Right paramere; D. Habitus, male; E. Habitus, female. Scale bars: 0.25 mm (A – C); 1.0 mm (D, E).	Figure 25. Pterostichus (Morphohaptoderus) rongyu sp. nov. (A – D. Holotype; E. Paratype, Huanglianchanggou, Yintiaoling, Chongqing). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, dorsal view; C. Right paramere; D. Habitus, male; E. Habitus, female. Scale bars: 0.25 mm (A – C); 1.0 mm (D, E).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481169	https://doi.org/10.3897/dez.72.159716.figure6	Figure 6. Pterostichus (Morphohaptoderus) dentellus Facchini & Sciaky, from Shennongjia, Hubei. A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – D); 2.0 mm (E, F).	Figure 6. Pterostichus (Morphohaptoderus) dentellus Facchini & Sciaky, from Shennongjia, Hubei. A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – D); 2.0 mm (E, F).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481181	https://doi.org/10.3897/dez.72.159716.figure18	Figure 18. Pterostichus (Morphohaptoderus) hubeicus Facchini & Sciaky, from Shennongjia, Hubei. A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	Figure 18. Pterostichus (Morphohaptoderus) hubeicus Facchini & Sciaky, from Shennongjia, Hubei. A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481166	https://doi.org/10.3897/dez.72.159716.figure3	Figure 3. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov. Scale bars: 0.5 mm.	Figure 3. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481171	https://doi.org/10.3897/dez.72.159716.figure8	Figure 8. Pterostichus (Morphohaptoderus) quyuani sp. nov. (A – F. Holotype; G. Paratype, Wenshui Village, Shennongjia, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	Figure 8. Pterostichus (Morphohaptoderus) quyuani sp. nov. (A – F. Holotype; G. Paratype, Wenshui Village, Shennongjia, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481173	https://doi.org/10.3897/dez.72.159716.figure10	Figure 10. Pterostichus (Morphohaptoderus) shennongjianus Facchini & Sciaky, from Shennongjia, Hubei. A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F – G).	Figure 10. Pterostichus (Morphohaptoderus) shennongjianus Facchini & Sciaky, from Shennongjia, Hubei. A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F – G).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481178	https://doi.org/10.3897/dez.72.159716.figure15	Figure 15. Pterostichus (Morphohaptoderus) lingjun sp. nov. (A – F. Holotype; G. Paratype, Guanmen Mountain, Shennongjia, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	Figure 15. Pterostichus (Morphohaptoderus) lingjun sp. nov. (A – F. Holotype; G. Paratype, Guanmen Mountain, Shennongjia, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481168	https://doi.org/10.3897/dez.72.159716.figure5	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481167	https://doi.org/10.3897/dez.72.159716.figure4	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481183	https://doi.org/10.3897/dez.72.159716.figure20	Figure 20. Pterostichus (Morphohaptoderus) yaotiao sp. nov. (A – E. Holotype; F. Paratype, Wenshui Village, Shennongjia, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	Figure 20. Pterostichus (Morphohaptoderus) yaotiao sp. nov. (A – E. Holotype; F. Paratype, Wenshui Village, Shennongjia, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481186	https://doi.org/10.3897/dez.72.159716.figure23	Figure 23. Pterostichus (Morphohaptoderus) toledanoi Facchini & Sciaky, from Shennongjia, Hubei. A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	Figure 23. Pterostichus (Morphohaptoderus) toledanoi Facchini & Sciaky, from Shennongjia, Hubei. A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481187	https://doi.org/10.3897/dez.72.159716.figure24	Figure 24. Pterostichus (Morphohaptoderus) xuanzhang sp. nov. (A – E. Holotype; F. Paratype, Jiuhuping, Shennongjia, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	Figure 24. Pterostichus (Morphohaptoderus) xuanzhang sp. nov. (A – E. Holotype; F. Paratype, Jiuhuping, Shennongjia, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481185	https://doi.org/10.3897/dez.72.159716.figure22	Figure 22. Pterostichus (Morphohaptoderus) zhizheng sp. nov. (A – D. Holotype; E. Paratype, Houhe Nature Reserve, Wufeng County, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, dorsal view; C. Right paramere; D. Habitus, male; E. Habitus, female. Scale bars: 0.5 mm (A – C); 2.0 mm (D, E).	Figure 22. Pterostichus (Morphohaptoderus) zhizheng sp. nov. (A – D. Holotype; E. Paratype, Houhe Nature Reserve, Wufeng County, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, dorsal view; C. Right paramere; D. Habitus, male; E. Habitus, female. Scale bars: 0.5 mm (A – C); 2.0 mm (D, E).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
9661F056BB13544A849004C3B97A264C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481184	https://doi.org/10.3897/dez.72.159716.figure21	Figure 21. Pterostichus (Morphohaptoderus) lianquan sp. nov. (A – D. Holotype; E. Paratype, Houhe Nature Reserve, Wufeng County, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, dorsal view; C. Right paramere; D. Habitus, male; E. Habitus, female. Scale bars: 0.5 mm (A – C); 2.0 mm (D, E).	Figure 21. Pterostichus (Morphohaptoderus) lianquan sp. nov. (A – D. Holotype; E. Paratype, Houhe Nature Reserve, Wufeng County, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, dorsal view; C. Right paramere; D. Habitus, male; E. Habitus, female. Scale bars: 0.5 mm (A – C); 2.0 mm (D, E).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
A3980CD5DAB45CAAABA2B721331A67E5.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481164	https://doi.org/10.3897/dez.72.159716.figure1	Figure 1. Characters of P. (Morphohaptoderus) spp. A. Antennomere 3 of P. wangshu sp. nov. B. Antennomere 3 of P. yaotiao sp. nov. C. Apical lamella of P. lisao sp. nov. D. Apical lamella of P. lianquan sp. nov. E. Fifth tarsomere of P. yaotiao sp. nov. F. Fifth tarsomere of P. hubeicus; G. Fifth tarsomere of P. shennongjianus; H. Elytral microsculpture of P. dentellus; I. Elytral microsculpture of P. shennongjianus. Scale bars: 0.25 mm (A – G); 0.05 mm (H – I).	Figure 1. Characters of P. (Morphohaptoderus) spp. A. Antennomere 3 of P. wangshu sp. nov. B. Antennomere 3 of P. yaotiao sp. nov. C. Apical lamella of P. lisao sp. nov. D. Apical lamella of P. lianquan sp. nov. E. Fifth tarsomere of P. yaotiao sp. nov. F. Fifth tarsomere of P. hubeicus; G. Fifth tarsomere of P. shennongjianus; H. Elytral microsculpture of P. dentellus; I. Elytral microsculpture of P. shennongjianus. Scale bars: 0.25 mm (A – G); 0.05 mm (H – I).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
A3980CD5DAB45CAAABA2B721331A67E5.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481166	https://doi.org/10.3897/dez.72.159716.figure3	Figure 3. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov. Scale bars: 0.5 mm.	Figure 3. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
A3980CD5DAB45CAAABA2B721331A67E5.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481168	https://doi.org/10.3897/dez.72.159716.figure5	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
A3980CD5DAB45CAAABA2B721331A67E5.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481179	https://doi.org/10.3897/dez.72.159716.figure16	Figure 16. Pterostichus (Morphohaptoderus) wangshu sp. nov. (A – F. Holotype; G. Paratype, Houhe Nature Reserve, Wufeng County, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	Figure 16. Pterostichus (Morphohaptoderus) wangshu sp. nov. (A – F. Holotype; G. Paratype, Houhe Nature Reserve, Wufeng County, Hubei). A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F, G).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
A3980CD5DAB45CAAABA2B721331A67E5.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481180	https://doi.org/10.3897/dez.72.159716.figure17	Figure 17. Endophallus of P. (Morphohaptoderus) spp. A, B. P. wangshu sp. nov. A. Left lateral view; B. Right lateral view. C, D. P. zhizheng sp. nov. C. Left lateral view; D. Right lateral view. Abbreviations: gp gonopore, lp left pre-apical lobe, la left apical lobe, ra right apical lobe. Scale bars: 0.5 mm.	Figure 17. Endophallus of P. (Morphohaptoderus) spp. A, B. P. wangshu sp. nov. A. Left lateral view; B. Right lateral view. C, D. P. zhizheng sp. nov. C. Left lateral view; D. Right lateral view. Abbreviations: gp gonopore, lp left pre-apical lobe, la left apical lobe, ra right apical lobe. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
A3980CD5DAB45CAAABA2B721331A67E5.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481175	https://doi.org/10.3897/dez.72.159716.figure12	Figure 12. Median lobe of aedeagus, left lateral view. A – C. P. dentellus; D, E. P. wangshu sp. nov.; F – I. P. quyuani sp. nov.; J. P. lingjun sp. nov. A. Non-type specimen, Shennongjia, Yanziya, 2300 m; B. Non-type specimen, Shennongjia, Jiuhuping, 1800–1950 m; C. Non-type specimen, Yintiaoling, Shizhuzi, 2137 m; D. Paratype, Wufeng, Houhe Nature Reserve, 900–1000 m; E. Holotype, Wufeng, Houhe Nature Reserve, 900–1000 m; F. Holotype, Shennongjia, Wenshui Village, 1700 m; G – I. Paratype, Shennongjia, Wenshui Village, 1700 m; J. Holotype, Shennongjia, Guanmen Mountain, 1361 m. Scale bars: 0.5 mm.	Figure 12. Median lobe of aedeagus, left lateral view. A – C. P. dentellus; D, E. P. wangshu sp. nov.; F – I. P. quyuani sp. nov.; J. P. lingjun sp. nov. A. Non-type specimen, Shennongjia, Yanziya, 2300 m; B. Non-type specimen, Shennongjia, Jiuhuping, 1800–1950 m; C. Non-type specimen, Yintiaoling, Shizhuzi, 2137 m; D. Paratype, Wufeng, Houhe Nature Reserve, 900–1000 m; E. Holotype, Wufeng, Houhe Nature Reserve, 900–1000 m; F. Holotype, Shennongjia, Wenshui Village, 1700 m; G – I. Paratype, Shennongjia, Wenshui Village, 1700 m; J. Holotype, Shennongjia, Guanmen Mountain, 1361 m. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
A3980CD5DAB45CAAABA2B721331A67E5.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481173	https://doi.org/10.3897/dez.72.159716.figure10	Figure 10. Pterostichus (Morphohaptoderus) shennongjianus Facchini & Sciaky, from Shennongjia, Hubei. A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F – G).	Figure 10. Pterostichus (Morphohaptoderus) shennongjianus Facchini & Sciaky, from Shennongjia, Hubei. A. Apical lamella, dorsal-apical view; B. Median lobe of aedeagus, left lateral view; C. Median lobe of aedeagus, ventral view; D. Median lobe of aedeagus, dorsal view; E. Right paramere; F. Habitus, male; G. Habitus, female. Scale bars: 0.25 mm (A); 0.5 mm (B – E); 2.0 mm (F – G).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
A3980CD5DAB45CAAABA2B721331A67E5.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481165	https://doi.org/10.3897/dez.72.159716.figure2	Figure 2. Male sternum of P. (Morphohaptoderus) spp. A, D. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; E. P. lisao sp. nov.; F. P. lingjun sp. nov. Scale bars: 1 mm.	Figure 2. Male sternum of P. (Morphohaptoderus) spp. A, D. P. dentellus; B. P. quyuani sp. nov.; C. P. shennongjianus; E. P. lisao sp. nov.; F. P. lingjun sp. nov. Scale bars: 1 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
AC79F1DA4BC857CD8200130F086496D7.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481167	https://doi.org/10.3897/dez.72.159716.figure4	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
AC79F1DA4BC857CD8200130F086496D7.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481168	https://doi.org/10.3897/dez.72.159716.figure5	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
AC79F1DA4BC857CD8200130F086496D7.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481187	https://doi.org/10.3897/dez.72.159716.figure24	Figure 24. Pterostichus (Morphohaptoderus) xuanzhang sp. nov. (A – E. Holotype; F. Paratype, Jiuhuping, Shennongjia, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	Figure 24. Pterostichus (Morphohaptoderus) xuanzhang sp. nov. (A – E. Holotype; F. Paratype, Jiuhuping, Shennongjia, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
AC79F1DA4BC857CD8200130F086496D7.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481186	https://doi.org/10.3897/dez.72.159716.figure23	Figure 23. Pterostichus (Morphohaptoderus) toledanoi Facchini & Sciaky, from Shennongjia, Hubei. A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	Figure 23. Pterostichus (Morphohaptoderus) toledanoi Facchini & Sciaky, from Shennongjia, Hubei. A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
F1FCA1E507225325962184131A512133.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481164	https://doi.org/10.3897/dez.72.159716.figure1	Figure 1. Characters of P. (Morphohaptoderus) spp. A. Antennomere 3 of P. wangshu sp. nov. B. Antennomere 3 of P. yaotiao sp. nov. C. Apical lamella of P. lisao sp. nov. D. Apical lamella of P. lianquan sp. nov. E. Fifth tarsomere of P. yaotiao sp. nov. F. Fifth tarsomere of P. hubeicus; G. Fifth tarsomere of P. shennongjianus; H. Elytral microsculpture of P. dentellus; I. Elytral microsculpture of P. shennongjianus. Scale bars: 0.25 mm (A – G); 0.05 mm (H – I).	Figure 1. Characters of P. (Morphohaptoderus) spp. A. Antennomere 3 of P. wangshu sp. nov. B. Antennomere 3 of P. yaotiao sp. nov. C. Apical lamella of P. lisao sp. nov. D. Apical lamella of P. lianquan sp. nov. E. Fifth tarsomere of P. yaotiao sp. nov. F. Fifth tarsomere of P. hubeicus; G. Fifth tarsomere of P. shennongjianus; H. Elytral microsculpture of P. dentellus; I. Elytral microsculpture of P. shennongjianus. Scale bars: 0.25 mm (A – G); 0.05 mm (H – I).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
F1FCA1E507225325962184131A512133.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481167	https://doi.org/10.3897/dez.72.159716.figure4	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
F1FCA1E507225325962184131A512133.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481168	https://doi.org/10.3897/dez.72.159716.figure5	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
F1FCA1E507225325962184131A512133.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481182	https://doi.org/10.3897/dez.72.159716.figure19	Figure 19. Endophallus of P. (Morphohaptoderus) spp. A, B. P. hubeicus. A. Left lateral view; B. Right lateral view. C, D. P. yaotiao sp. nov. C. Left lateral view; D. Right lateral view. Abbreviations: gp gonopore, bl basal lobe, lp left pre-apical lobe, la left apical lobe, ra right apical lobe. Scale bars: 0.5 mm.	Figure 19. Endophallus of P. (Morphohaptoderus) spp. A, B. P. hubeicus. A. Left lateral view; B. Right lateral view. C, D. P. yaotiao sp. nov. C. Left lateral view; D. Right lateral view. Abbreviations: gp gonopore, bl basal lobe, lp left pre-apical lobe, la left apical lobe, ra right apical lobe. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
F1FCA1E507225325962184131A512133.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481183	https://doi.org/10.3897/dez.72.159716.figure20	Figure 20. Pterostichus (Morphohaptoderus) yaotiao sp. nov. (A – E. Holotype; F. Paratype, Wenshui Village, Shennongjia, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	Figure 20. Pterostichus (Morphohaptoderus) yaotiao sp. nov. (A – E. Holotype; F. Paratype, Wenshui Village, Shennongjia, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
F1FCA1E507225325962184131A512133.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481181	https://doi.org/10.3897/dez.72.159716.figure18	Figure 18. Pterostichus (Morphohaptoderus) hubeicus Facchini & Sciaky, from Shennongjia, Hubei. A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	Figure 18. Pterostichus (Morphohaptoderus) hubeicus Facchini & Sciaky, from Shennongjia, Hubei. A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, ventral view; C. Median lobe of aedeagus, dorsal view; D. Right paramere; E. Habitus, male; F. Habitus, female. Scale bars: 0.5 mm (A – D); 2.0 mm (E, F).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
E226522645FD532187031177305CA6E4.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481167	https://doi.org/10.3897/dez.72.159716.figure4	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	Figure 4. Pronotum posterior angle and basal area of Pterostichus (Morphohaptoderus) spp. A. P. yaotiao sp. nov.; B. P. zhizheng sp. nov.; C. P. lianquan sp. nov.; D. P. hubeicus; E. P. toledanoi; F. P. xuanzhang sp. nov.; G. P. chachi sp. nov.; H. P. rongyu sp. nov. Scale bars: 0.5 mm (A – F); 0.25 mm (G – H).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
E226522645FD532187031177305CA6E4.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481168	https://doi.org/10.3897/dez.72.159716.figure5	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	Figure 5. Gonocoxites 2 of ovipositor, dorsal view. A. P. dentellus; B. P. shennongjianus; C. P. quyuani sp. nov.; D. P. lisao sp. nov.; E. P. lingjun sp. nov.; F. P. wangshu sp. nov.; G. P. toledanoi; H. P. xuanzhang sp. nov.; I. P. hubeicus; J. P. yaotiao sp. nov.; K. P. lianquan sp. nov.; L. P. zhizheng sp. nov.; M. P. chachi sp. nov.; N. P. rongyu sp. nov. Scale bars: 0.25 mm, a for A, B, b for C – H, c for I – N.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
E226522645FD532187031177305CA6E4.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481180	https://doi.org/10.3897/dez.72.159716.figure17	Figure 17. Endophallus of P. (Morphohaptoderus) spp. A, B. P. wangshu sp. nov. A. Left lateral view; B. Right lateral view. C, D. P. zhizheng sp. nov. C. Left lateral view; D. Right lateral view. Abbreviations: gp gonopore, lp left pre-apical lobe, la left apical lobe, ra right apical lobe. Scale bars: 0.5 mm.	Figure 17. Endophallus of P. (Morphohaptoderus) spp. A, B. P. wangshu sp. nov. A. Left lateral view; B. Right lateral view. C, D. P. zhizheng sp. nov. C. Left lateral view; D. Right lateral view. Abbreviations: gp gonopore, lp left pre-apical lobe, la left apical lobe, ra right apical lobe. Scale bars: 0.5 mm.	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
E226522645FD532187031177305CA6E4.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1481185	https://doi.org/10.3897/dez.72.159716.figure22	Figure 22. Pterostichus (Morphohaptoderus) zhizheng sp. nov. (A – D. Holotype; E. Paratype, Houhe Nature Reserve, Wufeng County, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, dorsal view; C. Right paramere; D. Habitus, male; E. Habitus, female. Scale bars: 0.5 mm (A – C); 2.0 mm (D, E).	Figure 22. Pterostichus (Morphohaptoderus) zhizheng sp. nov. (A – D. Holotype; E. Paratype, Houhe Nature Reserve, Wufeng County, Hubei). A. Median lobe of aedeagus, left lateral view; B. Median lobe of aedeagus, dorsal view; C. Right paramere; D. Habitus, male; E. Habitus, female. Scale bars: 0.5 mm (A – C); 2.0 mm (D, E).	2025-12-03	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang		Zenodo	biologists	Wu, Zou-Yan;Chen, Jia-Heng;Shi, Hong-Liang			
