taxonID	type	description	language	source
0397FD10BB653D3FFF5B50B0FB64FE20.taxon	type_taxon	Type genus. Whetwhetaksa Archibald & Cannings, 2021	en	Wang, Yehao, Nel, André, Cai, Chenyang, Fu, Yanzhe, Liu, Qi, Zeng, Guangchun, Lian, Xinneng, Huang, Diying (2025): New insect assemblage from the early Oligocene in Ningming Basin, Guangxi, China. Zootaxa 5739 (1): 21-35, DOI: 10.11646/zootaxa.5739.1.6, URL: https://doi.org/10.11646/zootaxa.5739.1.6
0397FD10BB653D3FFF5B50B0FB64FE20.taxon	discussion	Remarks. Archibald et al. (2021) tentatively included the Whetwhetaksidae in the suborder Cephalozygoptera Archibald, Cannings & Erickson, 2021, mainly based on head characters, which are unknown in this family. Later Nel & Zheng (2021) suggested that this order is poorly defined on the basis of deformations of the heads of the concerned fossils. Archibald & Cannings (2021) denied the values of Nel & Zheng (2021) ’ arguments, but Nel & Jouault (2022) proposed new arguments to confirm their opinion. More recently, Ware et al. (2025: figs 3 – 4) proposed a phylogenetic analysis of the Cephalozygoptera, without defined outgroup (s), and thus also without strong proof of the monophyly of this set of taxa. Thus, the definition and value of the Cephalozygoptera remain dubious and we prefer to consider the Whetwhetaksidae as a family of ‘ Zygoptera’ sensu lato (sister group to the Epiproctophora). A new analysis of these enigmatic damselflies is under study by Jouault et al.	en	Wang, Yehao, Nel, André, Cai, Chenyang, Fu, Yanzhe, Liu, Qi, Zeng, Guangchun, Lian, Xinneng, Huang, Diying (2025): New insect assemblage from the early Oligocene in Ningming Basin, Guangxi, China. Zootaxa 5739 (1): 21-35, DOI: 10.11646/zootaxa.5739.1.6, URL: https://doi.org/10.11646/zootaxa.5739.1.6
0397FD10BB643D3FFF5B5565FADBFC88.taxon	description	urn: lsid: zoobank. org: act: 987426 A 9 - 4 B 33 - 4 EAD-BCEA- 6 A 134753 B 5 F 8	en	Wang, Yehao, Nel, André, Cai, Chenyang, Fu, Yanzhe, Liu, Qi, Zeng, Guangchun, Lian, Xinneng, Huang, Diying (2025): New insect assemblage from the early Oligocene in Ningming Basin, Guangxi, China. Zootaxa 5739 (1): 21-35, DOI: 10.11646/zootaxa.5739.1.6, URL: https://doi.org/10.11646/zootaxa.5739.1.6
0397FD10BB643D3FFF5B5565FADBFC88.taxon	type_taxon	Type species. Sinowhetaksa incompleta Nel sp. nov.	en	Wang, Yehao, Nel, André, Cai, Chenyang, Fu, Yanzhe, Liu, Qi, Zeng, Guangchun, Lian, Xinneng, Huang, Diying (2025): New insect assemblage from the early Oligocene in Ningming Basin, Guangxi, China. Zootaxa 5739 (1): 21-35, DOI: 10.11646/zootaxa.5739.1.6, URL: https://doi.org/10.11646/zootaxa.5739.1.6
0397FD10BB643D3FFF5B5565FADBFC88.taxon	etymology	Etymology. Named after ‘ Sinica’, Latin name for China and the suffix ‘ whetaksa’, employed for the genera of Whetwhetaksidae. Gender feminine.	en	Wang, Yehao, Nel, André, Cai, Chenyang, Fu, Yanzhe, Liu, Qi, Zeng, Guangchun, Lian, Xinneng, Huang, Diying (2025): New insect assemblage from the early Oligocene in Ningming Basin, Guangxi, China. Zootaxa 5739 (1): 21-35, DOI: 10.11646/zootaxa.5739.1.6, URL: https://doi.org/10.11646/zootaxa.5739.1.6
0397FD10BB643D3FFF5B5565FADBFC88.taxon	diagnosis	Diagnosis. Wing venation characters only. Base of RP 2 five cells distad subnodus; 19 postnodal crossveins; base of IR 2 one cell basad nodus, one row of cells in area between C and RA just distad pterostigma.	en	Wang, Yehao, Nel, André, Cai, Chenyang, Fu, Yanzhe, Liu, Qi, Zeng, Guangchun, Lian, Xinneng, Huang, Diying (2025): New insect assemblage from the early Oligocene in Ningming Basin, Guangxi, China. Zootaxa 5739 (1): 21-35, DOI: 10.11646/zootaxa.5739.1.6, URL: https://doi.org/10.11646/zootaxa.5739.1.6
0397FD10BB643D32FF5B54A9FE6BFE78.taxon	description	urn: lsid: zoobank. org: act: 5500 D 7 E 4 - A 836 - 4 A 4 D- 821 F- 317199603 CFC	en	Wang, Yehao, Nel, André, Cai, Chenyang, Fu, Yanzhe, Liu, Qi, Zeng, Guangchun, Lian, Xinneng, Huang, Diying (2025): New insect assemblage from the early Oligocene in Ningming Basin, Guangxi, China. Zootaxa 5739 (1): 21-35, DOI: 10.11646/zootaxa.5739.1.6, URL: https://doi.org/10.11646/zootaxa.5739.1.6
0397FD10BB643D32FF5B54A9FE6BFE78.taxon	materials_examined	Material. Holotype, a fragmented wing attached to thorax (Fig. 7), NIGP 200692, stored at NIGPAS.	en	Wang, Yehao, Nel, André, Cai, Chenyang, Fu, Yanzhe, Liu, Qi, Zeng, Guangchun, Lian, Xinneng, Huang, Diying (2025): New insect assemblage from the early Oligocene in Ningming Basin, Guangxi, China. Zootaxa 5739 (1): 21-35, DOI: 10.11646/zootaxa.5739.1.6, URL: https://doi.org/10.11646/zootaxa.5739.1.6
0397FD10BB643D32FF5B54A9FE6BFE78.taxon	etymology	Etymology. Named after the incomplete state of preservation of the holotype.	en	Wang, Yehao, Nel, André, Cai, Chenyang, Fu, Yanzhe, Liu, Qi, Zeng, Guangchun, Lian, Xinneng, Huang, Diying (2025): New insect assemblage from the early Oligocene in Ningming Basin, Guangxi, China. Zootaxa 5739 (1): 21-35, DOI: 10.11646/zootaxa.5739.1.6, URL: https://doi.org/10.11646/zootaxa.5739.1.6
0397FD10BB643D32FF5B54A9FE6BFE78.taxon	diagnosis	Diagnosis. As for the genus.	en	Wang, Yehao, Nel, André, Cai, Chenyang, Fu, Yanzhe, Liu, Qi, Zeng, Guangchun, Lian, Xinneng, Huang, Diying (2025): New insect assemblage from the early Oligocene in Ningming Basin, Guangxi, China. Zootaxa 5739 (1): 21-35, DOI: 10.11646/zootaxa.5739.1.6, URL: https://doi.org/10.11646/zootaxa.5739.1.6
0397FD10BB643D32FF5B54A9FE6BFE78.taxon	materials_examined	Locality and horizon. Ningming Formation; along the Luoyue Avenue, Chengzhong Town, Ningming County, Chongzuo City, Guangxi, China; early Oligocene (Rupelian).	en	Wang, Yehao, Nel, André, Cai, Chenyang, Fu, Yanzhe, Liu, Qi, Zeng, Guangchun, Lian, Xinneng, Huang, Diying (2025): New insect assemblage from the early Oligocene in Ningming Basin, Guangxi, China. Zootaxa 5739 (1): 21-35, DOI: 10.11646/zootaxa.5739.1.6, URL: https://doi.org/10.11646/zootaxa.5739.1.6
0397FD10BB643D32FF5B54A9FE6BFE78.taxon	description	Description. Wing hyaline with brown pterostigma; wing ca. 27.0 mm long, ca. 7.7 mm wide; petiole rather short, 2.9 mm long; CuP just basad base of AA; anal area narrow below subdiscoidal cell, with one row of cells; distance from base to nodus 6.6 mm; from nodus to pterostigma 14.2 mm; from pterostigma to wing apex ca. 1.9 mm; pterostigma elongate, 3.3 mm long, width between 0.6 mm and 0.8 mm (the vein RA along its posterior margin compressed and deformed so that the pterostigma seems to be wider than it should be); basal side of pterostigma very oblique; distal side even more oblique and very long, with a distinct ‘ prolongation’ along costal margin, 0.3 mm long; one row of cells in area between C and RA just distad pterostigma but two rows distally; no oblique pterostigmal brace; 19 postnodal crossveins, basal-most ones being well aligned with postsubnodal crossveins and distal ones not aligned; nodal crossvein and subnodus well aligned and strongly oblique; base of RP 2 five cells distad subnodus; base of IR 1 ca. six cells distally; base of IR 2 one cell basad nodus; CuA terminating at margin over half wing length, well distad nodus; MA close to, subparallel with RP 3 - 4 to posterior margin; MA and MP diverging to posterior margin; MP and CuA subparallel to posterior margin.	en	Wang, Yehao, Nel, André, Cai, Chenyang, Fu, Yanzhe, Liu, Qi, Zeng, Guangchun, Lian, Xinneng, Huang, Diying (2025): New insect assemblage from the early Oligocene in Ningming Basin, Guangxi, China. Zootaxa 5739 (1): 21-35, DOI: 10.11646/zootaxa.5739.1.6, URL: https://doi.org/10.11646/zootaxa.5739.1.6
0397FD10BB643D32FF5B54A9FE6BFE78.taxon	discussion	Remarks. Although fragmentary, this wing is attributed to Zygoptera sensu lato because of the presence of a petiole, shape of subdiscoidal cell, and very long postnodal area. It does not fit in the epiproctophoran Isophlebioptera Bechly, 1996 that have also these characters, because of the narrower anal area and subdiscoidal cell. This last clade was still present during the Eocene through the family Pseudostenolestidae (Garrouste & Nel, 2015). There are rather few damselfly families with such broad wings, elongate pterostigma, long postnodal area compared to antenodal one, and so numerous postnodal crossveins, viz. the Dysagrionidae, Sieblosiidae, Whetwhetaksidae, Shundeagrionidae, Menatlestidae, Latibasaliidae, Thaumatoneuridae, and Heliocharidae. Affinities with the Heliocharidae are excluded because of the longer petiole, and base of IR 2 close to nodus; affinities to Thaumatoneuridae are excluded because the base of RP 2 is only five cells distad subnodus (Münz, 1919). Affinities with the Sieblosiidae are excluded because the subnodus is in straight line with the nodal crossvein and of strong ‘ normal’ obliquity, and the base of IR 2 is one cell basad the nodus. Affinities with the Shundeagrionidae, Menatlestidae, and Latibasaliidae are excluded because the base of IR 2 is one cell basad nodus vs midway between nodus and arculus, and distal side of the pterostigma is strongly oblique vs much less in the latter groups (Petrulevičius & Nel, 2007; Nel & Jouault, 2022; Huang et al., 2025). Affinities with the Dysagrionidae are excluded because the subnodus is much more oblique than in this family (with a weak ‘ normal’ obliquity to perpendicular to RP in the Dysagrionidae) (Archibald et al., 2021: figs 67 - 67; Simonsen et al., 2022: figs 1 A, 2 A). Also the distal side of the pterostigma of the new fossil has a very particular shape, very oblique with a short but distinct ‘ prolongation’ along costal margin. This character is present in the Whetwhetaksidae, but not in the other families, even if some Dysagrionidae (e. g., Okanagrion, Dysagrion) also have an oblique distal side of the pterostigma, but less than in the new fossil and the Whetwhetaksidae, and no costal ‘ prolongation’. This character is not listed in the diagnoses proposed by Archibald et al. (2021: 110), Simonsen et al. (2022: 587), and Ware et al. (2025) for the Whetwhetaksidae, but it is a putative synapomorphy for this family, present in the type genus Whetwhetaksa and in Danowhetaksa. The preserved characters of the new fossil fit well with those listed in the diagnoses of this family, viz. distinctively long pterostigma [Simonsen et al. indicated ‘ length at least seven times width’, while it is five to eight times, depending if we count the width of the vein RA along its posterior margin in the new fossil, but this character can greatly vary in representatives of the same family, e. g., see the different situations in the Dysagrionidae]; nodus to base of pterostigma 55 – 60 % of nodus to apex [75 % in the new fossil, due to the slightly shorter pterostigma]; arculus just distal to Ax 1, closer to it than to Ax 2 [unknown in the new fossil]; oblique vein O absent [probably shared by the new fossil]; nodal, subnodal crossveins of ‘ normal’ obliquity [shared by the new fossil]; base of RP 2 close to nodus [shared by the new fossil]; CuA terminates at margin over half wing length [shared by the new fossil]; MA close to, subparallel with RP 3 - 4 to margin [shared by the new fossil]; MA and MP diverge to margin [shared by the new fossil]; MP and CuA remain subparallel to margin [shared by the new fossil]. In conclusion, the new fossil can be attributed to the Whetwhetaksidae, especially because of the specialized shape of the distal margin of the pterostigma. Its base of RP 2 is five cells distad subnodus, vs. only two to four cells in Whetwhetaksa, and eight cells in Danowhetaksa. It also has only 19 postnodal crossveins, vs. ca. 20 in Whetwhetaksa, and ca. more than 25 in Danowhetaksa. Its base of IR 2 is one cell basad the nodus, vs. in a well basal position in Whetwhetaksa and two small cells basad in Danowhetaksa. Lastly, there is only one row of cells in area between C and RA just distad the pterostigma, vs. two rows in Whetwhetaksa and Danowhetaksa, but this last character is more of species level than the preceding. It is not very surprising to find a representative of the Whetwhetaksidae in China as this family is present in the Paleogene of Eastern North America and Western Europe. This group is either quite ancient, Cretaceous or probably could ‘ migrate’ between Eurasia and North America through the ‘ Beringian Bridge’ as for some other Odonata (Garrouste & Nel, 2019).	en	Wang, Yehao, Nel, André, Cai, Chenyang, Fu, Yanzhe, Liu, Qi, Zeng, Guangchun, Lian, Xinneng, Huang, Diying (2025): New insect assemblage from the early Oligocene in Ningming Basin, Guangxi, China. Zootaxa 5739 (1): 21-35, DOI: 10.11646/zootaxa.5739.1.6, URL: https://doi.org/10.11646/zootaxa.5739.1.6
