Introduction
The odonate genus Labandeiraia Petrulevičius et al. (Cephalozygoptera, Eodichromatidae, Eodichromatinae) is known from the earliest Ypresian of Denmark (L. europae Petrulevičius et al. and undescribed species, unpublished data) and the Ypresian Green River and Wind River Formations of Wyoming and Colorado, United States of America (L. americaborealis Petrulevičius et al., L. riveri Bechly et al., and L. browni Bechly et al.; Petrulevičius et al. Reference Petrulevičius, Nel, Rust, Bechly and Kohls2007; Bechly et al. Reference Bechly, Garrouste, Aase, Karr, Grande and Nel2020). A single species of Eodichromatinae has been described from the Okanagan Highlands at Republic, Washington, United States of America, Republica weatbrooki Archibald and Cannings. At that time, Eodichromatidae was considered a subfamily of the zygopteran family Epallagidae (= Euphaeidae) (Archibald and Cannings Reference Archibald and Cannings2021b). Here, we describe two Labandeiraia wings from the Okanagan Highlands (Archibald et al. Reference Archibald, Greenwood, Smith, Mathewes and Basinger2011a) Allenby Formation of the southern interior of British Columbia, Canada, near the village of Princeton.
Material and methods
We examined the part (A side) and counterpart (B side) of two fossil odonate wings, BBM P014397A, B and BBM P014398A, B, not joined but in close association. One is hyaline, and the other is darkly infuscate. They were found in Allenby Formation Vermilion Bluffs Unit lacustrine shale (see Read Reference Read2000 for the geology) about 4 km southwest of Princeton, British Columbia, Canada. The “Billy’s Family Restaurant” exposure of Vermilion Bluffs shale in Princeton has an estimated Ypresian maximum likelihood age of 51.85 ± 0.85 Ma (Rubino et al. Reference Rubino, Leier, Cassel, Archibald, Foster-Baril and Barbeau2021).
We follow the systematics of the suborder Cephalozygoptera of Ware et al. (Reference Ware, Simonsen, Rasmussen and Archibald2025) and the morphological terminology of Bechly (Reference Bechly1998) and Garrison et al. (Reference Garrison, von Ellenrieder and Louton2010): aa, accessory antenodal crossveins; Ax0, Ax1, and Ax2, primary antenodal crossveins; C, costa; CuA, anterior branch of the cubitus; discoidal bracket, the thickened distal sides of the quadrangle (MAb) and subquadrangle (basal CuA), aligned with reverse obliquity (red; Fig. 2); hs, hyperstigmal crossvein(s) between the pterostigma and C; IR1 and IR2, intercalary veins; MA and MP, anterior and posterior branches of the media; n, nodus; pt, pterostigma; RA, anterior branch of the radius; and RP1, RP2, RP3-4, branches of the posterior radius.
These wings are housed in the collections of the Beaty Biodiversity Museum (BBM), University of British Columbia, Vancouver, British Columbia, Canada.
The part and counterpart of the infuscate wing mostly preserve different portions, with a small, shared portion that we used to join them graphically in Adobe Illustrator (Adobe, San Jose, California, United States of America) when the counterpart portion is flipped left to right (Figs. 1 and 2).
Figure 1. A, Photograph of the holotype hyaline wing of Labandeiraia burlingameae n. sp. part BBM P014397A (left) and Labandeiraia sp. A (right), the basal portion part (BBM P014398A) graphically joined with the distal portion counterpart (BBM P014398B), flipped left–right. B, Drawings of these wings with information from their parts and counterparts. Arrow indicates direction of extension shown in the Figure 2 drawings. Scalebar = 1 cm.
Figure 2. Drawings from the part and counterparts of A, the Labandeiraia burlingameae n. sp. holotype (BBM P014397A, B), and B, Labandeiraia sp. A (BBM P014398A, B), bottom, as in Figure 1B, but corrected for distortion (see the text and grey arrow in Fig. 1). The quadrangle is blue, the subquadrangle is green, and the discoidal bracket is red. These presumed undistorted wing shapes are speculative, so no scale is indicated.
These wings were deformed by geologic forces, common at this exposure; see discussion and illustration of a stretched March fly, Plecia Wiedemann (Diptera, Bibionidae) in Archibald and Cannings (Reference Archibald and Cannings2021a, fig. 3) and of the giant ant Titanomyrma Archibald et al. (Hymenoptera, Formicidae, Formiciinae) in Archibald et al. (Reference Archibald, Mathewes and Aase2023, fig. 2). The wings are at a right angle to each other, so graphically stretching them together on a plane perpendicular to the length of the hyaline wing until their nodus-to-apex lengths are the same likely closely reproduces their life shape. Because of uncertainty, however, measurements are not reported. The same shape could be generated by compressing the fossil at 90° to this along the plane of the length of the hyaline wing, but the wings in that reconstruction would be unreasonably small.
We use the family, subfamily, and tribe names Epallagidae rather than Euphaeidae, Eodichromatinae rather than Eodichrominae, and Eodichromatini rather than Eodichromini (Bechly Reference Bechly1998, Reference Bechly1999).
We use the mean annual temperature categories of Wolfe (Reference Wolfe1975): microthermal, colder than or equal to 13 °C; mesothermal, warmer than 13 °C, colder than 20 °C; megathermal, at least 20 °C.
Systematic paleontology of Labandeiraia burlingameae and Labandeiraia sp. A
Order Odonata Fabricius
Suborder Cephalozygoptera Archibald et al .
Superfamily Eodichromatoidea Ware et al .
Family Eodichromatidae Cockerell
Subfamily Eodichromatinae Cockerell
Tribe Eodichromatini Cockerell
Genus Labandeiraia Petrulevičius et al .
Labandeiraia burlingameae n. sp.
Zoobank Nomenclature Act: urn:lsid:zoobank.org:act:8C8B50D3-6C78-40DB-8B20-ED0D3091D333
Type material. HOLOTYPE: BBM P014397A, B, part and counterpart. An almost complete hyaline forewing. Collected by Beverley Burlingame, 14 October 2024, and housed in the BBM collections.
Diagnosis. The L. burlingameae forewing can be separated from fore- and hind wings of all other Labandeiraia species by lack of colouration, from all others except L. americaborealis by extensive distal undulate curvature towards and away from each other of long veins and supplementary sectors between them. Forewing is further separated from L. americaborealis by having at least 14 more postnodal (C-RA space) crossveins (53 preserved, likely more than 60 total) than the L. americaborealis hind wing (44 preserved, likely 46 total). Labandeiraia europae and L. riveri have seven more in forewing than in hind wing; L. browni’s forewing is not known.
Description. HOLOTYPE: BBM P014397A, B – as in diagnosis and elongate-oval, posterior margin rather smoothly curved from apex to base. With numerous supplementary sectors between long veins; dense crossvenation throughout clearly present, especially when wetted with ethanol, but not preserved well enough to clearly photograph or accurately draw other than 53 postnodal crossveins, probably approximately 63 total, none branched. Basal-most portion where Ax0, Ax1, and Ax2 might be not preserved; three accessory antenodal crossveins. Pterostigma long, with basal, distal sides strongly oblique. At least one hyperstigmal crossvein. Region where brace vein might occur is poorly preserved. Discoidal bracket present. RP1-2 does not closely approach RA. Subnodus not preserved, but RP2 origin posterior to nodus where subnodus confidently would be; IR1 origin closely distad. No veins zigzagged. MA, MP, CuA, and many supplementary sectors smoothly undulate as above; MP, CuA with subparallel curves; at least two supplementary sectors between MP and CuA. The CuA space expanded with many supplementary sectors. Cells about twice as high as wide in basal-posterior region, mostly posterior to subquadrangle, some immediately distad this.
Etymology. The specific epithet is a patronymic formed from the surname of Beverley Burlingame, recognising her discovery of these wings, as well as her making them available to us, her donating them to the Beaty Biodiversity Museum, and her many other contributions to the paleontology of the Allenby Formation over the years.
Range and age. The Late Ypresian Vermilion Bluffs Member of the Allenby Formation near Princeton, British Columbia.
Labandeiraia sp. A
Material. A darkly infuscate forewing BBM P014398A, B, preserved in close proximity on the same rock as the L. burlingameae holotype. About the basal quarter of the wing is missing. Collected by Beverley Burlingame, as above, and housed in the BBM collections.
Description. BBM P014398A, B. – wing darkly infuscate; main veins and supplementary sectors between them in preserved portions like those of the L. burlingameae holotype; crossvenation seen in places to be dense, but almost none preserved clearly enough to accurately draw.
Systematics of Labandeiraia burlingameae and Labandeiraia sp. A
Ware et al. (Reference Ware, Simonsen, Rasmussen and Archibald2025) raised the Eodichromatinae (as Zygoptera, Epallagidae) to family status and transferred it to the Cephalozygoptera. We assign L. burlingameae to the Eodichromatidae and so to the Cephalozygoptera by the diagnostic combination of its eodichromatoid quadrangle (see Ware et al. Reference Ware, Simonsen, Rasmussen and Archibald2025: rectangular or subrectangular, with distal end slightly if at all broader than basal end, anterior side about same length as posterior, corner angles 90° or close to it, length twice or more the maximum height, may be curved); discoidal bracket; accessory antenodal crossveins; expanded cubito-anal area with supplementary sectors; RP1-2 not closely approaching RA near its base; cells about twice as high as wide in the basal-posterior region; and undulate long veins. Other character states considered diagnostic of the Eodichromatidae by Bechly (Reference Bechly1998) and Petrulevičius et al. (Reference Petrulevičius, Nel, Rust, Bechly and Kohls2007) as a subfamily of Euphaeidae, Zygoptera, and by Ware et al. (Reference Ware, Simonsen, Rasmussen and Archibald2025) as a family of Cephalozygoptera cannot be evaluated due to the specimens’ preservation.
Within the Eodichromatidae, the Allenby wings are most like that of Labandeiraia americaborealis, which is known only by a hind wing. They possess numerous long veins, with the intercalated veins curving away from and towards each other in their distal portion in a similar fashion. In the hind wings of some other Labandeiraia species, a few veins may be curved this way but less than is seen in L. americaborealis and the new Allenby wings or in any other eodichromatid forewing known to us.
In the subfamily Eodichromatinae, hind wings are wider than forewings and are triangular, with a rounded angle point on the posterior margin (e.g., undescribed Eodichromatinae from the Fur Formation of Denmark, unpublished data, and see Petrulevičius et al. Reference Petrulevičius, Nel, Rust, Bechly and Kohls2007, fig. 2 – Labandeiraia europae Petrulevičius et al., and Bechly et al. Reference Bechly, Garrouste, Aase, Karr, Grande and Nel2020, fig. 5 – Labandeiraia riveri Bechly et al. and fig. 6 – Eodysphaea magnifica Bechly et al.). Both incomplete Allenby forewings are reconstructed as not unusually wide, elongate–oval with the preserved posterior margin rather evenly curved – that is, without a distinct angle – consistent with eodichromatine forewings (Fig. 2).
Because one wing is uniformly infuscate and the other is uniformly hyaline, they are preserved about 1.5 cm apart, and because a difference in colouration as an artefact of preservation is not known to us in any Allenby Formation fossil, we are confident that their preserved colour difference represents a true difference. We can’t tell what colour the preserved portion of the infuscate wing was in life (its hue), but we are confident that it was dark (its value).
Although the chances seem small for reasons discussed below, the hyaline forewing and the dark L. americaborealis hind wing might be conspecific, with this colouration resulting from sexual dimorphism, polyphenism, or a forewing–hind wing difference. All described species and undescribed specimens of Labandeiraia known to us have dark wings. Treating this forewing as a new species is further supported by the number of its postnodal crossveins relative to that of the L. americaborealis hind wing. Both the L. europae and L. riveri forewings have about seven more postnodal crossveins than their hind wings do. If this pattern is stable within the genus, the hyaline wing is excluded from L. americaborealis by having at least 14 more (53 preserved, likely more than 60 total) than the L. americaborealis hind wing (44 preserved, likely 46 total).
The new infuscate wing is dark throughout like the L. americaborealis hind wing and has similar venation to it and to that of L. burlingameae. It cannot be compared with these species by the number of its postnodal crossveins because these are not preserved distinctly enough to accurately count. The L. americaborealis diagnosis of Petrulevičius et al. (Reference Petrulevičius, Nel, Rust, Bechly and Kohls2007) separates it from L. europae (all that was necessary at the time) and includes character states of the hind wing, the basal region, and other morphology that cannot be evaluated in the present specimen by preservation but might be resolved in future specimens.
Labandeiraia americaborealis is from the Parachute Creek Member of the Green River Formation in Colorado (Petrulevičius et al. Reference Petrulevičius, Nel, Rust, Bechly and Kohls2007), which is, like the Allenby Formation, in the second half of the Ypresian (Smith et al. Reference Smith, Singer and Carroll2003, Reference Smith, Chamberlain, Singer and Carroll2010; Rubino et al. Reference Rubino, Leier, Cassel, Archibald, Foster-Baril and Barbeau2021). The Green River Formation is located about 1400 km southeast of the Allenby and preserved life in a hot, upper mesothermal to megathermal climate (Archibald et al. Reference Archibald, Johnson, Mathewes and Greenwood2011b), as opposed to the Allenby Formation, which was deposited in a considerably cooler microthermal montane setting (Greenwood et al. Reference Greenwood, Archibald, Mathewes and Moss2005).
We find it most conservative to designate the hyaline wing as the holotype of L. burlingameae and treat the infuscate wing as Labandeiraia sp. A because it could belong to L. burlingameae, L. americaborealis, or its own new species.
Acknowledgements
The authors thank Beverley Burlingame of Princeton for finding the specimen and donating it to the Beaty Biodiversity Museum; Emilia Silvestre for photography in the fossil lab; Karen Needham, Curator of Entomology at the Beaty Biodiversity Museum, for use of her lab’s microscope with digital camera; and Aliye Sasmaz for photography. S.B.A. thanks Rolf Mathewes of Simon Fraser University, Burnaby.
Financial support
S.B.A. received support from Natural Sciences and Engineering Grant 2020-05026 to Rolf Mathewes (Simon Fraser University, Burnaby, British Columbia).
Competing interests
The authors declare that they have no competing interests.
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