Embiodea, Kusnezov, 1903

Anisyutkin, Leonid N., Legalov, Andrei A. & Perkovsky, Evgeny E., 2024, New species and new record of embiids (Insecta: Embiodea) from upper Eocene of Europa, Ecologica Montenegrina 79, pp. 16-28 : 21-28

Embiodea gen. sp.

( Figs 9 – 13 )

Material. Female or last instar larva of female. SIZK LKV- 1005. SIZK LKV- 1005/A, Rovno amber, Priabonian.

Description of inclusion. Amber piece ( Fig. 9 ) is cut into two parts, so the inclusion is divided between two unequal pieces of amber. Total weight is 3.43 (2.1 + 1.33) g. The inclusion chip passes through the specimen, so that the upper part of the insect lies in the larger and the lower part in the smaller fragment of inclusion. The interior of the specimen is empty, with traces of charred cuticle. First, larger, piece in shape of irregular elongated prism, approximately 32 × 8.5 × 2.5 mm, only partly transparent, is including only the upper part of webspinner. Second, smaller, part of inclusion in shape of shorter prism, approximately 22.5 × 11.0 × 2.0 mm, transparent, is including the lower part of webspinner and elongated fragment of plant debris. The specimen is located in a curved position, the head and prothorax are higher than the abdomen ( Figs 10, 11 ).

Description. General color, as far as can be judged from the inclusions, yellowish-brown, distal segments of antennae probably slightly darker ( Figs 10, 11 ). Surfaces, as far as can be judged from the inclusion, smooth. Pubescence well developed, more expressed on antennae, legs, lower side of abdomen and cerci; lower side of tarsi with dense brush of chaetae; abdomen from below and cerci with long recumbent chaetae. Head large and elongated ( Figs 10–13 ), epicranial sutures or carinas absent; posterocaudal angles rounded; eyes small, about twice shorter then postocular space, not prominent; labrum medially slightly emarginated. Antennae about (as visible from the inclusion) 13–15 segmented (right and left antenna, respectively); scapus short and massive, about as long as wide; pedicellum subquadrate, smaller than scapus; 3rd segment about as long as scapus; 4th about as long as pedicellum; other segments of about equal length, slightly elongated, with exception of slightly longer 3 penultimate ones. Mouthparts indiscernible with exception of maxillary palps and distal part of mentum; maxillary palps with 1st-4th segments short and subequal in length, ultimate segment longer; prementum of usual for webspinners structure (similar to fig. 11 in Ross 2000), anterior angles of submentum rounded, not produced. Pronotum transverse, with swollen prescutum; mesonotum large, about as long as wide, with distinct acrotergite. Fore and hind wings completely absent. Metanotum transverse, without visible acrotergite; median grooves absent. Prosternum small; mesosternum strongly deformed; metasternum transverse, constricted and widely rounded caudally. Fore probasitarsus (1st tarsal segment of fore leg) widened, about 2 times long as wide (about 0.6 mm x 0.35 mm); 2nd tarsal segment about as long as wide (~ 0.15 mm), with rounded distolateral margins; 3rd tarsal segments very short (~ 0.1 mm). Fore femora moderately widened. Mid probasitarsus elongated, with single apical pulvilla, length of tarsal segments about 0.4, 0.1, 0.25 mm. Hind femora strongly widened, hind probasitarsus stout, with single apical pulvilla, length of tarsal segments about 0.3, 0.1, 0.25 mm. All claws simple and symmetrical, arolia absent. Abdomen elongated, widest at V–VII abdominal segments. Terminalia nearly symmetrical, Xth abdominal tergum simple, subtrapezoidal in shape, with caudal margin slightly asymmetrical; cerci symmetrical, consist of two fusiform segments, proximal segment slightly shorter and wider than distal one; IX sternum without visible specializations; two symmetrical triangular paraprocts located behind IX sternum.

Measurements (mm). Length of head 1.1; width of head 0.8; length of pronotum 0.6; width of pronotum 0.8; length of mesonotum 1.25; width of mesonotum 1.1; length of metanotum 0.7; width of metanotum 1.0; length of fore femur 0.8; length of midtibia 0.4; length of hind femur 1.0; length of hind tibia 0.7; width of Xth abdominal tergite 0.75. All measurements approximate due to inclusion distortion and specimen deformation.

Distribution. Rovno amber, Priabonian.

Note. Unfortunately, the present material, a single female or even a larva, does not allow us to determine the webspinner from Rovno amber even to the family level. However, the species from Rovno amber is obviously not conspecific with Electroembia species, readily differs by the presence of only one pulvilla on the hind basitarsus and no protruding anterior angles of submentum.

Discussion

At the present time, fossil Cenozoic webspinners are known from the early and late Eocene of Europe (Oise amber, Baltic amber, Rovno amber), early Eocene of South Asia ( Cambay amber), late Eocene of North America (Florissant) and Miocene of Mesoamerica ( Ross 2000, Chiapas amber) and Caribbean (Dominican amber) ( Engel et al. 2011; Falières et al. 2021). Apparently, embiids were numerous in the warm and humid climate of the first half of the Cenozoic.

The genus Electroembia is apparently close to Galloembia Falières, Engel & Nel, 2021 in the structure of the male terminalia. The latter genus was described from Oise amber of France dating to the earliest Eocene ( Falières et al. 2021).

Unfortunately, descriptions of late Eocene (Florissant) Lithembia florissantensis ( Cockerell, 1908) are too scanty. The abdominal apex is not well preserved in the known specimens of L. florissantensis , and other characters are described too schematically ( Cockerell 1908; Ross 1944, 1984). This species was described in the genus Embia Latreille, 1829 ( Cockerell 1908), later it was putatively ascribed to Clothoda Enderlein, 1909 ("Evidence for placing it in the genus Clothoda is very inconclusive" – Ross 1944, p. 406) and finally erected in monotypical genus Lithembia Ross, 1984 . The diagnosis of this genus is unsuitable and therefore the name Lithembia is nomen nudum ( Engel & Grimaldi 2006; Miller 2009).

It is noteworthy that Electroembia is similar to extant genera from Africa and the New World ( Ross 1966). This may serve as confirmation of the existence at the end of the Mesozoic – the beginning of the Cenozoic of a more or less uniform fauna of Polyneoptera from the Caribbean basin to the Far East ( Shelford 1910; Vršanský et al. 2011, 2014; Anisyutkin 2020).

Edward Ross (1956, p. 80) supposed "Apterism in male Embioptera is characteristic of modern species which live in ecologically marginal life zones of the world. The apterous condition is almost universal in regions experiencing prolonged dry seasons. Thus, male apterism in Electroembia strongly suggests that the early Tertiary environment of the species suffered a long dry season in the summer months (the season of sexual maturity in modern North-temperate species)".

Still all that known about Baltic amber fauna contradicts this supposition (Rasnitsyn, pers. com.). The absence of winged Embioptera males, if confirmed, may support the hypothesis of greater dryness of the Volhynian land in the late Eocene vs. Russo-Scandia (see also Lyubarsky & Perkovsky 2012, 2019; Dietrich & Perkovsky 2020 and references therein).

Webspinners are generally perceived as thermophiles. At the same time, at least some representatives of the order are cryophobes (see Archibald et al. 2023, Jenkins Shaw et al. 2024a).

Extant web-spinners were found in the microthermal locations, i. e. Kazantip in Crimea (MAT +11.4 °C) and Aktau (former Shevchenko) in Kazakhstan (MAT +12,6 °C) ( Nazarenko et al. 2009; Temreshev 2023). It is even possible that these insects were introduced to the northernmost locations of their range by humans ( Ross 2000; Nozaki et al. 2018), although for Florissant Lagerstätte MAT is 11.6 ± 3.3°C ( Allen et al. 2020). Even more noteworthy looks a strong presence of extant webspinners on Abrau Peninsula (MAT +13.2 °C, Korobushkin et al. 2016), where these insects are dominant in shrubland ( Gongalsky et al. 2006). Moreover, the high abundance of embiopterans on the burned areas (27.8% of all large soil invertebrates in the area 7 years after fire) probably indicates that they are well adapted to fires ( Gongalsky et al. 2011).

The most likely reason for the abundance of webspinners in shrubland is the weakness of the ant pressure. The ants in the shrubland are represented by small species that are unable to control webspinners (Dubovikoff, pers. com., Perkovsky, unpublished). We have already cited the weakness of the ant (and bird) pressure as one of the reasons for the abundance and diversity of webspinners in the Kachin amber forest ( Perkovsky et al. 2020; Anisyutkin et al. 2022).

Adaptation to fire could be very important webspinners’ adaptation to sub-Mediterranian climate ( Gongalsky et al. 2011). If it is really connected with a loss of wings by the males, the ratio of the winged and wingless males could be an important source of information about the ecology areas, neighbouring with European amber forests.

Webspinners apparently ranged much further north in the Cretaceous than in the late Eocene, as suggested by the discovery of Sclerogibbidae (obligate parasitoids of web-spinners) in Aptian of Kanto Region, Japan (Choshi amber) ( Perkovsky et al. 2020). At the same time, while webspinners or Sclerogibbidae are known from all ambers of Isoptera Realm except Spanish and New Jersey ones ( Perkovsky et al. 2020) in the Cretaceous, Baeomorpha Realm looks too cold for them ( Jenkins Shaw et al. 2024b; Perkovsky 2022 and references therein).

Until now, more than one webspinner species was known from only two fossil resins: 15 species from Kachin amber ( Ross 2024; Liu et al. 2024) and 5 species (including 4 formally described) from Dominican amber ( Martynova et al. 2019, Supplemental Table 2). In this paper, they are supplemented with Baltic amber. Rovno amber is significantly richer in cryophobic species than Baltic amber ( Yamamoto et al. 2022; Belokobylskij et al. 2023; Kirichenko-Babko & Perkovsky 2023; Legalov et al. 2023 a, 2023b; Telnov et al. 2023; Chemyreva et al. 2024; Melnitsky et al. 2024b etc.), so further study of the Rovno amber fauna is very likely to reveal more than one species of webspinner.

Acknowledgements

The authors thank Carsten Gröhn ( Glinde , Germany) who made available for study unique Baltic amber inclusions, Mykola Khomich (Rivne, Ukraine) for donating Rovno amber inclusion to SIZK and Alexander P. Rasnitsyn for useful and fruitful discussions .

The authors thank anonymous reviewers for the valuable comments that improved the manuscript.

L.N. Anisyutkin’s research was performed in the frames of the state research project 122031100272-3 (Systematics, morphology, ecophysiology and evolution of insects).

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Figures

Figures 9–11. Embiodea gen. sp. 9 – General view of inclusion; 10, 11 – webspinner from above (10) and below (11). Scale bar for 1 = 10 mm, 10, 11 – out of scale.

Figures 12–13. Embiodea gen. sp. General view from above (12) and below (13). Scale bar 1 mm.