taxonID	type	description	language	source
03C287D3FFB0330DFD8E170E25554148.taxon	type_taxon	Type species Capnia nigra (Pictet, 1833).	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB0330DFD8E170E25554148.taxon	diagnosis	Diagnosis Additionally, the entirely setose main epiproct sclerite is unique and will easily distinguish it from all other males in the species group.	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB0330AFF48149326EA43F5.taxon	diagnosis	Diagnosis Both the male and female are macropterous, the forewing having a curved A 1 and R 1 and a quadrangular cubital cell (Fig. 3). The mesothoracic postfurcasternum is separated from the furcasternum and furcasternal pit (Murányi et al. 2014: fig. 53). The male epiproct has a large basal sclerite that is weakly divided from the smaller laterobasal sclerites. The main epiproct sclerite is laterally entire and fully divided on the ventral surface. The caudal setae are absent and the inner sclerite is weak but long. The epiproct tip lacks an eversible crest (Murányi et al. 2014: figs 11 – 12). The male paraprocts are long and widely separated at the apex (Murányi et al. 2014: fig. 26). The fusion plate is long and narrow, the small retractoral plate not fused (Murányi et al. 2014: fig. 37). The male tergum IX has a single posteromedial process that lacks scales or a sensilla basiconica. Tergum X is subdivided and posteriorly connected. A vesicle is usually present on the sternum IX and the subgenital plate is divided from sternum and tergum IX (Figs 4, 10). The female subgenital plate is large and its lateral portions are often weakly sclerotized. The posterior lobe is distinct and variable with the lateral and anterior sclerites being distinct. The inner sclerite and postgenital plate are lacking (Fig. 8).	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB0330AFF48149326EA43F5.taxon	discussion	Affinities The Cox 1 based molecular analyses well support both the monophyly of the group, and its distinction from Capnia s. str. (Murányi et al. 2014). It deserves definition as a new genus, as it will be described in the ongoing genus level revision of the Capniidae being conducted by Dávid Murányi. Morphologically, males are closely related to the genus Zwicknia Murányi, 2014, though molecular analyses do not suggest a close relationship (Murányi et al. 2014). The species of Zwicknia differ by the presence of a distinct eversible crest on the apicolaterally divided main epiproct sclerite; alternatively, the C. cordata group lacks the eversible crest. Females of the cordata group are distinctive among Palaearctic Capniidae by the distinct posterior lobe of the large but usually laterally weakly sclerotized subgenital plate, combined with the lack of an inner sclerite of the vaginal complex, a normal anal field of the hind wing, and a quadrangular cubital cell of the forewing. The larva is known only for two species (Zwick & Sivec 1980; Rehman et al. 2022); the tuft of hairs on the tip of the galea (Zwick & Sivec 1980) is a possible distinctive character for the group.	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB0330AFF48149326EA43F5.taxon	distribution	Distribution Members of the C. cordata group are distributed in High Asia, where they are the most common and diverse Capniidae. Species are known from the Tien Shan, Hindukush, Pamir, Himalayas and the eastern ranges, sloping from the Qinghai-Tibet Plateau (Fig. 1). The morphologically similar Zwicknia has a mainly West Palaearctic distribution, extending eastwards to the Central Asian high mountains. This distribution pattern elucidates the geographically vicariant relationship between Zwicknia and the C. ordata group.	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB63304FDDA130926244562.taxon	description	urn: lsid: zoobank. org: act: B 54 BC 6 B 4 - 681 D- 4 ECA- 833 B- 192955 B 1 B 8 C 7 Figs 1 – 8, 11 A – B, 12 A – C, 13 – 14	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB63304FDDA130926244562.taxon	diagnosis	Diagnosis The male of this new species differs markedly from that of most other members of the C. cordata species group by its caudally projecting posteromedial process of tergum IX ending in two acute spines.	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB63304FDDA130926244562.taxon	etymology	Etymology The specific epithet ‘ bispina ’ is derived from Latin ‘ bi ’ (meaning ‘ two’ or ‘ double’) and ‘ spina ’ (meaning ‘ spine’ or ‘ thorn’) in reference to the two apical spines of the posteromedial process of male tergum IX. Use as a noun, gender feminine.	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB63304FDDA130926244562.taxon	materials_examined	Type material Holotype CHINA • ♂; Qinghai Province, Xining City, Datong County, Niangniang Mountain; 36 ° 54 ′ 56.81 ″ N, 101 ° 36 ′ 10.51 ″ E; 2864 m a. s. l.; 17 Mar. 2021; Wei-Hai Li and Fan-Bin Kong leg.; HIST. Paratypes CHINA • 9 ♂♂, 5 ♀♀; same data as for holotype; HIST.	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB63304FDDA130926244562.taxon	description	Description Male ADULT HABITUS (Fig. 2). Body generally dark brown. Body length 9.1 – 9.3 mm. Forewing length 8.8 – 9.0 mm, hindwing length 7.0 – 7.2 mm (N = 10). Macropterous. Wing venation typical for species group. Forewing mottled, dark band in distal third, light band basal to it, other clouds of pigmentation. Head slightly wider than pronotum, with three ocelli, M-line and tentorial callosities dark brown. Antennae and palpi brown to dark brown. Pronotum trapezoid with dark rugosities, corner obtuse. Legs brown; tibiae lighter. Cerci with 17 – 18 slightly clubbed segments, each with apical whorl of long hairs. ABDOMEN (Figs 2 A, 4 – 7, 11 A – B, 12 A – B). Tergum I medially divided, terga II – IV subdivided by anteromedial membranous portion; antecosta medially divided on terga III and X, medially indent but entire on terga VI – IX (Fig. 11 B). Terga IV – IX laterally with paired longitudinal patch and pair of spots. Tergum IX strongly sclerotized, posterior edge indenting laterad to posteromedial process, process slightly raised and extending to anterior margins of tergum X, 1 / 5 × as wide as segment, falciform in shape and bilobed apically. Tergum X subdivided by evenly rounded anteromedial indentation (Figs 4 – 5). Sternum I entire and unmodified; sternum II with posterior sclerite not fused with anterior and lateral sclerites, sternum III with posterior sclerite not fused with anterior sclerites; sterna IV – VIII with anterior and lateral sclerites fused with posterior sclerite. Sternum IX fused with tergum IX anterolaterally (Fig. 11 B); subgenital plate divided from other sclerites and tongue-shaped, with darkly sclerotized lateral margins; vesicle very large and covered with dense long hairs, rounded and wider than long. Short basal stalk, extending over ½ of subgenital plate (Fig. 4 B). Paraprocts with long and wide apex, fusion plate long and narrow (Fig. 5 B). Epiproct consist of large basal sclerite fused with relatively large, triangular basolateral sclerites; main epiproct sclerite 0.35 mm in length, long and widely triangular in dorsal view, dorsal edge convex while ventral edge convex in lateral view but apical third slightly dilated then abruptly narrowed into sharp, slightly downcurved apex. Dorsal division of main epiproct sclerite straight and narrow, only slightly opens after NaOH treatment; basal portion slightly darker, with delicate keel laterobasally; basal fork relatively large, fused with laterobasal sclerites; apex sharply pointed in both lateral and dorsal aspects (Figs 5, 12 A – B); sclerite covered with dense setae on its dorsal and lateral surface, setae slightly shorter on basal portion and not similar to caudal setae of other Capniidae, sensillae occur on apex; inner sclerite long but thin and indistinct (Figs 6 – 7). Female ADULT HABITUS (Fig. 2 B). Body length 10.5 – 10.6 mm. Forewing length 9.2 – 9.3 mm, hindwing length 8.2 – 8.3 mm. Habitus generally similar to male. ABDOMEN (Figs 8, 11 A, 12 C). Terga I – VIII divided by wide median membranous area, each marked with pair of small dark spots; tergum IX with pair of anterior indentations, tergum X entire. Sternum I entire and unmodified; sterna II – VI with anterior sclerites not fused with posterior sclerite and lateral sclerites lacking, sternum VII with anterior sclerites not fused with posterior sclerite but small lateral sclerites present and fused (Figs 8 A, C, 11 A). Anterior sclerites of sternum VIII quadrangular, fused with subgenital plate. Subgenital plate wide trapezoidal, covering most of sternum (Fig. 12 C), flat in lateral view; evenly sclerotized, setation lacking from medial portion and posterior lobe; posterior lobe less wide than ⅓ of subgenital plate width, not further lobed but expanded laterad with sharp, triangular portions. Lateral sclerites bean-shaped and bald, not fused with subgenital plate nor with tergum VIII. Postgenital plate and inner vaginal sclerite lacking. Sternum IX with wavy anterior edge, not fused with tergum IX; paraprocts triangular (Fig. 8 B).	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB63304FDDA130926244562.taxon	distribution	Distribution China: Qinghai Province, Datong County. Adults were collected in mid-March, emerging from a slow to moderately fast flowing, small headwater tributary of the Datong River, its streambed consisting of large boulders and gravel (Fig. 13).	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB63304FDDA130926244562.taxon	discussion	Remarks Only C. s. lat. bilobata Chen & Song, 2019 and C. s. lat. badakhshanica Zhiltzova, 1974 share a similar tergum IX process, but in C. s. lat. bilobata, it is wider and bilobed without apical spines and in C. s. lat. badakhshanica, it is shorter and smaller. The entirely setose main epiproct sclerite is unique in the species group, also easily distinguishing the male. The female of C. s. lat. bispina Cao & Li sp. nov. appears the closest to C. s. lat. qilianshana Li & Yang, 2009 and C. s. lat. yunnana Li & Yang, 2011, sharing a similar narrow posterior lobe of the subgenital plate that is expanded lateral with triangular portions. However, the evenly sclerotized subgenital plate easily distinguishes the female from those two species.	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB8331EFDCD16D024604207.taxon	description	urn: lsid: zoobank. org: act: 91 C 439 D 8 - 907 B- 4 E 78 - 9024 - 9 F 824 A 83837 D Figs 1, 9 – 10, 11 C, 12 D – E	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB8331EFDCD16D024604207.taxon	diagnosis	Diagnosis The male of this new species differs markedly from that of most other members of the C. cordata species group by its straight process of tergum IX and the long and narrow triangular epiproct sclerite with the ventral and dorsal edges slightly but evenly convex in lateral view.	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB8331EFDCD16D024604207.taxon	etymology	Etymology The specific epithet ‘ huanglong ’ is derived from the Huanglong Scenic Spot, where the new species was found and probably restricted to these places and the nearby ranges. Used as a noun, gender neutral.	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB8331EFDCD16D024604207.taxon	materials_examined	Material examined Holotype CHINA • ♂; Sichuan Province, Aba Tibetan and Qiang Autonomous Prefecture, Songpan County, Huanglong Scenic Spot; 103 ° 49 ′ 32.52 ″ E, 32 ° 44 ′ 55.68 ″ N; 3550 m a. s. l.; 2 Aug. 2011; Si-Pei Liu and Wei-Hai Li leg.; HIST.	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB8331EFDCD16D024604207.taxon	description	Description Male ADULT HABITUS (Figs 2 – 3). Body generally dark brown. Body length 8.9 mm. Forewing length 10.3 mm, hindwing length 8.0 mm. Macropterous. Wing venation typical for species group. Head slightly wider than pronotum, with three pale and small ocelli, M-line and tentorial callosities dark brown. Antennae and palpi brown. Pronotum trapezoid with dark rugosities, corner obtuse. Legs brown; femora darker. Cerci with clubbed segments, each with apical whorl of long hairs; apical part of cerci broken in holotype. ABDOMEN (Figs 9 A, 10, 11 C, 12 D – E). Terga I – IV medially divided by membranous portion; antecosta medially divided on terga II – IV and X, medially indenting but entire on terga V – IX; short, medial longitudinal band projecting from antecosta on terga VIII – IX. Tergum IX strongly sclerotized, posterior edge nearly straight in dorsal view, process elevated and triangular in lateral view, with two small sclerotized teeth projecting into small membranous patch. Tergum X subdivided by widely rounded anteromedial indentation. Sternum I entire and unmodified; sternum II with anterior sclerites not fused with posterior sclerite, lateral sclerites lacking; sterna III – VIII with anterior and lateral sclerites, all fused with posterior sclerite (Fig. 11 C). Sternum IX fused with tergum IX anterolaterally; subgenital plate divided from other sclerites and subtriangular, with darkly sclerotized lateral margins; vesicle elliptical and medium-sized, covered with numerous long hairs, posteriorly reaching ⅓ of subgenital plate. Paraprocts with long and wide apex, fusion plate long and narrow (Fig. 10). Epiproct consists of large basal sclerite fused with small, triangular laterobasal sclerites; main epiproct sclerite 0.6 mm in length, long and narrowly triangular in dorsal view, both ventral and dorsal edge slightly but evenly convex in lateral view. Dorsal division of main epiproct sclerite slightly dilated in apical third, widely opens after NaOH treatment; basal portion darkly sclerotized with raised subtriangular structure in dorsal aspect; basal fork small, fused with laterobasal sclerites; apex pointed but its tip blunt in both lateral and dorsal aspects; sclerite bald besides few apical sensillae; inner sclerite long but thin and indistinct (Figs 10, 12 D – E). Female Unknown.	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB8331EFDCD16D024604207.taxon	distribution	Distribution China: Sichuan Province. Presently only known from Huanglong Scenic Spot.	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
03C287D3FFB8331EFDCD16D024604207.taxon	discussion	Remarks The new species is closely related to C. s. lat. manii, but can be separated by the tip of the main epiproct sclerite, which is more pointed in the latter. Furthermore, C. s. lat. manii has a vestigial ventral vesicle, in contrast to the normally developed vesicle found in the new species. The main epiproct sclerite of C. s. lat. huanglong Cao & Li sp. nov. is also different from that of other Chinese species within the C. s. lat. cordata species group: in C. s. lat. yunnana, the main epiproct sclerite has a swollen apex; in C. s. lat. qilianshana, the main epiproct sclerite has a rounded tip and its subapical area is slightly swollen; in C. s. lat. oblata, the main epiproct sclerite has a flattened apex; in C. s. lat. xiei, the main epiproct sclerite is high and has a basal bilamellar structure, while these characteristics are absent in C. s. lat. huanglong. Molecular analysis To characterize in detail the species limits within the C. cordata species group, we consider morphological characters and mitochondrial marker Cox 1, under a phylogenetic analysis. The aligned data matrix was 651 base pairs (bp) in length. The best fit substitution model TNe + I: CO 1 _ p 1, F 81 + F: CO 1 _ p 2, TN + F + G 4: CO 1 _ p 3 was selected for this concatenated data matrix. The Maximum Likelihood tree supported the morphological distinction of C. s. lat. bispina Cao & Li sp. nov. and the separation from the other species of the C. cordata species group with 100 % nodal support (Fig. 14). Interspecific divergence values for C. s. lat. bispina ranged from 5.03 – 19.44 % of genetic divergence in comparison with the other species whereas intraspecific between an associated male-female pair was only 0.31 % (Table 3). Combined, this is good evidence for the validity of C. s. lat. bispina (Hebert et al. 2003; Zhou et al. 2009). Meanwhile, from the analyses we demonstrated that C. s. lat. bispina is a member of the C. cordata species group. Unfortunately, the specimen of C. s. lat. huanglong Cao & Li sp. nov. was 14 years old, and failed to be sequenced successfully.	en	Cao, Zhishan, Cao, Jinjun, Yang, Ding, Li, Weihai, Murányi, Dávid (2025): Integrative taxonomy of the Capnia cordata species group (Plecoptera: Capniidae). European Journal of Taxonomy 1012: 81-105, DOI: 10.5852/ejt.2025.1012.3027, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3027/13591
