Phryganea vkoivekiraz Melnitsky, Ivanov & Perkovsky, 2024

Phryganea vkoivekiraz Melnitsky, Ivanov & Perkovsky sp. n.

https://zoobank.org/ urn:lsid:zoobank.org:act:B79DED16-A1C3-4CE8-AF3D-4AD7569622A8

( Figs 5 – 8 View Figure 5 View Figure 6 View Figure 7 View Figure 8 , 9 C, D View Figure 9 )

Type material: Holotype. Male. SIZK UA –28653, Rovno Oblast, Rovno amber, late Eocene.

Description. Body length 9.0 mm; forewing length 9.5 mm. Head, antennae, legs, thorax and wings light brown. Abdomen yellowish. Wings with white hairs. Warts of head and thorax with long brown hairs. Hind wings with F 2 in the middle part two times wider than F1.

Male genitalia. Inferior appendages consist of irregularly triangular distally rounded ventral part, dorsal part absent. Two long pointed processes visible on the sides of the phallobase are associated with the segment X. The lateral appendages (Wichard 2013; lateral appendages of X segment in Wiggins 1998) strongly sclerotized and pointed. The phallobase is narrow on ventral view and very wide on lateral view.

Comparison. The new species is similar to Phryganea picea (Pictet, 1856) from Baltic amber in presence of four pointed projections in the male genitalia. The new species differs from it by the structure of the inferior appendages without deep caudal incision and in the shape of lateral appendages pointed in the new species.

Etymology. The specific name vkoivekiraz is indeclinable and must not agree in gender with its generic name; it derives from Russian and means “only once”. Distribution. Priabonian Rovno amber. Remarks. The specimen is found in the piece of amber (weight 5.0 g) obtained from Mykola Khomych (Rovno).

Syninclusions: SIZK UA–28653 - Staphylinidae : Scydmaenitae, SIZK UA–28654 –

Psychodidae, SIZK UA –28655 – Chironomidae .

Both described Rovno pieces with Phryganeidae have Scydmaenitae as syninclusions as well as dipterans from Sciara zone ( Perkovsky 2017b, this paper), i. e. both most probably originated on the lower part of the trunk ( Perkovsky et al. 2012).

Abbreviations: X – 10 th segment, aed – aedeagus, eia – flat extension of inferior appendages, ia – inferior appendages, lp – lateral appendages, pb – phallobase, pX – process of segment X, spX – spines of segment X, vpX – ventral process of segment X.

Discussion

Caddisflies are the only group of aquatic insects from Rovno amber that has been studied in detail (e. g., Zelentsov et al. 2012; Perkovsky 2013, 2017a; Baranov et al. 2014, 2016; Perkovsky & Sukhomlin 2015; Ivanov et al. 2016; Giłka et al. 2021; Martynov et al. 2022). They are not directly associated with the amber forest and among them a fairly high proportion of cryophobic insects is found ( Melnitsky et al. 2024a, 2024с). It is therefore not surprising that this fauna has a high proportion of taxa unknown from Baltic amber (see Legalov et al. 2024 a, 2024b). Our data suggest 5 genera of 21 (23.8%) and 34 species of 46 (73.9%) ( Melnitsky et al. 2024b; this paper).

The prevailing family in the Rovno fauna of Trichoptera is the family Polycentropodidae , that were dominant group of caddisflies in the microthermal regions (with Mean Annual Temperature lower than 13 °C) of Eurasia and North America at least since Upper Cretaceous until end of Eocene ( Ivanov et al. 2016; Perkovsky 2022). This family makes 54.4% of the species known from Rovno ( Melnitsky et al. 2024b, and this paper). Some 68% species of Polycentropodidae known from Rovno were not found in the Baltic amber compared to 80.5% of species belonging to all other Trichoptera families. Hence, a half of the Rovno endemics belong to this family. Among the genus Holocentropus , which is most closely associated with microthermal regions, only 6 of 11 species (54.6%) are endemic, while in the genus Plectrocnemia known in tropics since the mid-Cretaceous (Ross 2024) five of seven species (71.4%) are endemic.

Phryganeidae is an extratropical family the only of the kind that shows obvious faunal differences in the European amber faunas at the genus level ( Ivanov et al. 2016). The genus Phryganea is the first Rovno genus of its family, common with the Baltic amber. This family is represented in the Baltic amber by almost the same number of individuals (70 vs. 74), studied by Ulmer (1912), as the genus Nyctiophylax Brauer, 1865 , the most species rich genus in Baltic amber ( Melnitsky et al. 2024b). At the same time, 80% of the Baltic phryganeids examined by Ulmer were identified as Phryganea picea Pictet, 1856 ; contrary, only 29.7% of the individuals studied by Ulmer (1912) belong to abundant Nyctiophylax varians Ulmer, 1912 , also the only species of the genus known from two amber faunas. Since the Baltic amber species of Nyctiophylax were 2–5 times smaller than the Baltic Phryganeidae , the larger caddisflies were underrepresented in the fossil resins. The fore wing length in the amber species described by Ulmer was 9–19 mm, while the Priabonian fossil imprints from Florissant Formation the wing size were 19–24 mm ( Cockerell 1913, 1914), and the extant Phryganea imagines are sufficiently larger with wing length up to 32 mm. In Florissant, phryganeids make up 5% of all caddisflies, the same as psychomyids ( Ivanov et al. 2016), while they make up only 1.4% of the Baltic caddisflies studied by Ulmer and 1.6% of the Rovno caddisflies identified to family (our data). Indirect evidence of their rarity in amber is also the fact that Wichard (2013) not only did not describe new phryganeids, but also did not use new finds to redescribe already known species; phryganeids are not indicated from Bitterfeld and Danish amber ( Ivanov et al. 2016). At the same time, phryganeids are among the few families of caddisflies whose larvae are known from Baltic amber (Wichard 2013). Large, well-preserved phryganeids can only be found in large pieces of amber and are of great collection value; very few such pieces with inclusions are known from Bitterfeld and Danish amber. The only large undescribed phryganeid in a large piece of Rovno amber is kept in the private collection of Viktor Gusakov from Moscow Region ( Perkovsky 2017b).

Holocentropus vottakvot sp. n. is the fourth caddisfly species known from the Varash District. Three of them (75%) belong to the Polycentropodidae . One of the latter species, Holocentropus atratus (Pictet, 1856) , is known from Baltic amber, but is not recorded from the best-studied Klesov deposit; the other three species are known only from the Varash District. The peculiarity of the caddisfly fauna confirms the need for an in-depth study of inclusions from the Varash District, where numerous interesting new specimens are constantly being sampled ( Anisyutkin et al. 2024; Vilhelmsen et al. 2024; Melnitsky et al. 2024b, 2024с and references therein).

Earlier we supposed that the warm-adapted caddisflies represent a majority of the taxa endemic for Rovno amber ( Melnitsky et al. 2024c). However, Leptoceridae , Hydroptilidae and Helicopsychidae taken together account for only 21.7% of caddisfly species in Rovno amber. The proportion of species in these families is higher than in the Baltic amber (14.4%), and 90% of the species are endemic, but this is still not enough to absorb the majority of endemics in Rovno amber. The climate of the Rovno amber forest, although warmer than that of the Baltic, was still equable ( Jenkins Shaw et al. 2024; Chemyreva et al. 2024), and temperate taxa predominated there ( Perkovsky 2016, 2017a). Previously, the presence of groups in the fauna of European ambers that are boreal or cold adapted in the modern fauna was indicated, in particular, by species of the genus Lype McLachlan, 1878 , representatives of the families Phryganeidae , Molannidae , and Beraeidae ( Ivanov et al. 2016) . These taxa include only 7.9% of the species of Baltic amber and 10.9% of the species of Rovno amber, although Lype sericea (Pictet, 1856) is the third most abundant amber species ( Ivanov et al. 2016). The greatest number of new findings in European amber should be expected in the most abundant and species-rich genera of polycentropodids. The most promising are Holocentropus , very species rich genus in European amber with 21 species in Baltic, 11 in Rovno, and 5 endemic species in Bitterfeld fossil resin, and Plectrocnemia , the most diverse genus in European amber having 35 species: 23 in Baltic, 8 in Rovno, 6 endemic in Bitterfeld amber ( Ivanov et al. 2016; Melnitsky & Ivanov 2019, 2024; Melnitsky et al. 2024b).

Acknowledgements

We thank Alexandr P. Rasnitsyn (Paleontological Institute, Moscow, Russia) for useful discussion and anonymous reviewers for improving the overall quality of the manuscript.

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