ISSN 0031-0301, Paleontological Journal, 2019, Vol. 53, No. 10, pp. 984–993. © Pleiades Publishing, Ltd., 2019.

A Summary of Recent Results in the Study of Taimyr Amber E. E. Perkovskya, b, * and D. V. Vasilenkob, c, ** aSchmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, Kiev, 01-030 Ukraine bBorissiak Paleontological Institute, Russian Academy of Sciences, Moscow, 117647 Russia cCherepovets State University, Cherepovets, 162600 Russia *e-mail: [email protected] **e-mail: [email protected] Received February 11, 2019; revised February 14, 2019; accepted February 14, 2019

Abstract—Two families, two tribes, 30 genera and 57 species are described as new from Taimyr amber, as well as the first new plant genus and species to be described from this amber during the 21st century. Based on the study of the Taimyr amber, two zoogeographic realms (“Baeomorpha” and “Isoptera”) are recognized.

Keywords: Taimyr amber, review, new taxa

DOI: 10.1134/S0031030119100149

INTRODUCTION Unfortunately, the technique for working with fos- sil resin (retinite) was not yet developed in the 1970s, Taimyr amber is known from a large number of and the most valuable representatives of many groups, localities (Rasnitsyn et al., 2016; Fig. 1) ranging in age including holotypes, are now significantly or irrevers- from Albian to Santonian (Perkovsky and Wegierek, ibly damaged (Makarkin and Perkovsky, 2017; Fig. 2). 2018; Gumovsky et al., 2018). New collections were made by the expedition of the The Taimyr amber forest represents a unique world Paleontological Institute as early as 2012, mainly in of its own, and the history of its study resembles a play, Yantardakh (Santonian), where D.S. Kopylov, with two acts separated by a rather long intermission E.A. Sidorchuk, and D.D. Vorontsov collected more (although not as long as in the studies, than 60 kilograms of retinite. where it lasted nearly a century). Among the new finds are the first Cretaceous As early as the 18th century, the Russian research- (later six aberrant genera and one genus with spotted ers S.P. Krasheninnikov, P.S. Pallas, and Kh.P. Laptev wings were also found in mid-Cretaceous Kachin began studying the fossil resins of Siberia, but only (Burmese) amber: Makarkin, pers. com.) and the A.F. Middendorf (1882) summarized the information northernmost representatives of spongillaflies (Neu- known at that time. However, it was only in 1970 that roptera: Sisyridae), which have an obligate associa- the expedition of the Paleontological Institute to tion with freshwater sponges and bryozoans Taimyr (for more details, see http://palaeoentomo- (Perkovsky and Makarkin, 2015; Makarkin and log.ru/Publ/amberproject.html) brought the first Perkovsky, 2016); freshwater sponges have not yet inclusions in the multi-kilogram collections of Taimyr been found in the Mesozoic of Eurasia. These Sisyri- amber presently kept at the Paleontological Institute dae are not fundamentally different from extant taxa, of the Russian Academy of Sciences (PIN), the tech- unlike most Kachin representatives (Makarkin, 2016; nical and scientific processing of which continues to Khramov et al., 2019). A new dustywing found in this day. This first expedition showed the potential of Cenomanian Nizhnyaya Agapa is assigned to a genus future collecting trips, which were held in 1971, 1973, previously known from Barremian Lebanese amber 1976 and 1977 at different Taimyr localities and in (Makarkin and Perkovsky, 2019). Previously the Yakutia (see Zherikhin, 1978, for a summary of the same genus of Lygistorrhinidae (Protognoriste) was results of those trips). These expeditions to Taimyr described from Nizhnyaya Agapa and Kachin amber resulted in the washing of many tons of tar sands, and (Blagoderov and Grimaldi, 2004); a member of Psy- more than 200 kg of resin were obtained (Sukatsheva chodidae found in Zhdanikha is also assigned to a et al., 2016). Lebanese genus (Azar et al., 2007). Another Leba-

984 A SUMMARY OF RECENT RESULTS IN THE STUDY OF TAIMYR AMBER 985

Taimyra Lake

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Kheta R. Pysina R. Polovinnoe Lake Korennoe Lake 1 Pyasinnoe Lake 30 km 2000 km

Fig. 1. Map of Taimyr amber localities: (1) Nizhnyaya Agapa, (2) Romanikha, (3) Yantardakh, (4) Isaevsky, (5) Bulun, (6) Sokolovsky, (7) Tumulda, (8) Ugolyak, (9) Kresty, (10) Gubina Gora, (11) Zhdanikha, (12) Novaya, (13) Moroskop, (14) Nikon-Yuryakh, (15) Bediey, (16) Khatangskaya Guba, (17) Logata, (18) Baikura; 1—Late Senomanian; 2–5, 7, 8—Late Santonian; 9, 11, 13–15—Albian–Late Senomanian; 17, 18—Albian–Early Senomanian; 6, 10, 12, 16—Unspecified age.

(a) (b)

Fig. 2. Types from Yantardakh damaged by placing them in castor oil: (a) Nganasania khetica Zherikhin, 1977, holotype PIN, no. 3311/45; (b) Shaposhnikovia electri Kononova, 1976, holotype PIN, no. 3311/259.

PALEONTOLOGICAL JOURNAL Vol. 53 No. 10 2019 986 PERKOVSKY, VASILENKO nese connection of Taimyr fauna appears Yantardakh nas of New Zealand and the Valdivian forests of Chile, bethylid genus in the subfamily Holopsenellinae containing many faunal elements, can Engel, Ortega et Azevedo, 2016 (only Taimyr and be attributed, in our opinion, to the low numbers and Lebanese genera were known: Azevedo et al., 2018, diversity of ants (Gumovsky et al., 2018). p. 262); third genus was described this year from Bur- A record of the oldest amber Chrysomelidae and mese amber. in particular of the oldest member of Galerucinae The first fossil record of an extant genus (new tribe Taimyralticini from Yantardakh) (Nadein Eutheia of the subfamily Scydmaeninae known to feed and Perkovsky, 2018) led to the analysis of the distri- on oribatid mites (Jałoszyński and Perkovsky, 2016) bution of Phytophaga (Chrysomeloidea and Curcu- comes from Yantardakh; previously Cretaceous mem- lionoidea) in the Late Cretaceous that revealed Cur- bers of Eutheiini were only known from the Albian Span- culionoidea not found in Taimyr and Chrysome- ish amber. A new species of the genus Iberomaimetsha loidea extremely rare (a single specimen found). The (: Maimetshidae) was described from amber faunas of the Baeomorpha realm were com- Yantardakh (Perrichot and Perkovsky, 2016); this pared for the first time with the only known Creta- genus was previously known only from two species ceous Lagerstätte not containing Phytophaga, i.e., from the Albian Spanish amber. Cretaceous Crypto- Santonian-Campanian Obeschayuschiy (Magadan phagini (Coleoptera: Cryptophagidae) also were Region). As recently established by paleobotanists, described only from Yantardakh and Spanish amber e.g., Herman (2013), Mesophytic floras with ancient (Lyubarsky and Perkovsky, 2015, 2017a, 2017b; Peris gymnosperms and ferns existed in northern Eurasia et al., 2017). until the end of the Cretaceous, and did not mix with The earliest paedogenetic gall midge from the the Cenophytic floras (with angiosperms and derived monotypic tribe Krassiloviolini (Fedotova and gymnosperms), and the distribution of Cenophytic Perkovsky, 2017), with 5-segmented tarsomeres that floras in the north of the Baeomorpha realm was often are absent in other paedogenetic gall midges, was restricted to river valleys, whereas Mesophytic floras described from Yantardakh: paedogenesis is here occupied the remaining territory. In the Late Creta- directly confirmed by the presence of two big eggs with ceous, Phytophaga were associated only with Ceno- larvae inside found next to the gall midge. phytic associations (present-day chrysomelids feed- With the description of Taimyrobryum martynov- ing on gymnosperms or ferns are exclusively rare) iorum (Ignatov et al., 2016) (first genus and species which explains their extreme rarity in the Taimyr of Taimyr amber plants), Yantardakh became the amber that formed on flat interfluves above narrow fourth locality in the world to yield Cretaceous river valleys with Cenophytic assemblages (Nadein Bryopsida. and Perkovsky, 2018). In our opinion, it was the pres- ervation of Mesophytic communities in the Baeo- The analysis of the composition of the Cretaceous morpha realm that led to a disparity between their faunas resulted in a revision of the Cretaceous faunal evolutionary and geological age (Rasnitsyn et al., elements in the northern hemisphere. Two zoogeo- 2016). graphic realms: “Baeomorpha” and “Isoptera” were recognized (Gumovsky et al., 2018). The Baeomor- The earliest known Old World amber black pha Realm, with a temperate or warm temperate cli- (Diptera: Simuliidae, Simuliini) were described from mate, apart from the presence of numerous parasitic the Santonian Taimyr amber (Perkovsky and Sukhom- wasps of the genus Baeomorpha (Rotoitidae), is char- lin, 2016; Perkovsky et al., 2018). The study of unex- acterized by very abundant aphid fossils (eight fami- pectedly abundant Ugolyak black flies showed that the lies from four superfamilies are known from this ornithuromorph birds, on which they most likely fed, realm only), few termites and no webspinners. The colonized northern Siberia as early as in the Mesozoic. Isoptera Realm, which had a warmer climate, con- In the Cretaceous of the northern hemisphere, almost tained no or very few Rotoitidae, no or few aphids everywhere that black flies are found, there are also (often with strongly reduced hind wings), whereas bird feathers (Perkovsky et al., 2018); the same applies termites and webspinners (Martynova et al., 2019) to the southern hemisphere (Australian Koonwarra). were abundant and diverse (webspinners at least in Simulioidea representative from Mongolia appears the south of the realm). Extant Rotoitidae have a rel- not black (Lukashevich et al., 2019); from Siberia ict distribution in environments with anomalously opposite birds (Enantiornithes) are unknown, so now low biocenotic pressure from ants (in New Zealand the connections between amber black flies and orni- and locally in Los Lagos Region of southern Chile, thuromorphs look even more reliable. mostly on Chiloé Island). The same is characteristic Altogether in this century, from the Taimyr of the Cretaceous assemblages with abundant Baeo- amber, two families (Shcherbakov, 2007; Popov, morpha (LaPolla et al., 2013; Perkovsky and Wegi- 2016; Fig. 3), two tribes, 30 genera, and 57 species of erek, 2018). The relict appearance of the modern fau- (including 16 genera and 29 species solely

PALEONTOLOGICAL JOURNAL Vol. 53 No. 10 2019 A SUMMARY OF RECENT RESULTS IN THE STUDY OF TAIMYR AMBER 987

(a)

(b) 1 mm (c) 1 mm

Fig. 3. Representatives of new families from Taimyr amber: (a) Cretargus emeljanovi Shcherbakov, 2007, PIN, no. 3311/561, nymph; (b, c) Taimyrocoris sukatshevae Popov, 2016, holotype PIN, no. 3311/459, (b) lateral and (c) ventral. from the material collected in the last century) are tardakh (Makarov and Perkovsky, 2019) were described as new, as well as one genus and species of recorded. The list of new taxa of plants and arthro- plants (Perkovsky, 2018; Jałoszyński and Perkovsky, pods described from Taimyr amber in the 21st cen- 2019). In addition, for the first time, for the Taimyr tury is given in Table 1. fauna, one order, four families, one subfamily, three tribes and three insect genera were recorded; this Given the inconsistency of results obtained by year additional gall midge genus from Ugolyak and comparing faunas of the Baeomorpha and Isoptera the first Cretaceous false skin beetle genus from Yan- realms (Rasnitsyn et al., 2016), a detailed inventory of

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Table 1. New species of plants and described from Taimyr amber in the 21st century Taxon Locality Citation Sphaeropsocidae: Globopsocus aquilonius Azar and Engel, 2008 Nizhnyaya Agapa Azar and Engel, 2008 Hemiptera Palaeoaphididae: Ambaraphis baikurensis Perkovsky and Wegierek, 2017 Baikura Perkovsky and Wegierek, 2017 Perforissidae: Cixitettix yangi Shcherbakov, 2007 Yantardakh Shcherbakov, 2007 Cretargus emeljanovi Shcherbakov, 2007 Yantardakh Shcherbakov, 2007 Taimyrocoridae: Taimyrocoris sukatshevae Popov, 2016 Yantardakh Popov, 2016 Coleoptera Chrysomelidae: Taimyraltica calcarata Nadein and Perkovsky, 2018 Yantardakh Nadein and Perkovsky, 2018 Cryptophagidae: Ennoticus mnemosynon Lyubarsky and Perkovsky, 2017 Yantardakh Lyubarsky and Perkovsky, 2017 Microticus khatanga Lyubarsky and Perkovsky, 2015 Yantardakh Lyubarsky and Perkovsky, 2015 Nganasania taymyrica Lyubarsky and Perkovsky, 2014 Yantardakh Lyubarsky and Perkovsky, 2014 Staphylinidae: Taimyraphes microscopicus Jałoszyński and Perkovsky, 2019 Yantardakh Jałoszyński and Perkovsky, 2019 Neuroptera Coniopterygidae: Glaesoconis popovi Makarkin and Perkovsky, 2017 Yantardakh Makarkin and Perkovsky, 2017 Libanoconis siberica Makarkin and Perkovsky, 2019 Nizhnyaya Agapa Makarkin and Perkovsky, 2019 Sisyridae: Prosisyrina sphinga Makarkin and Perkovsky, 2016 Yantardakh Makarkin and Perkovsky, 2016 Prosisyrina sukachevae Perkovsky and Makarkin, 2015 Yantardakh Perkovsky and Makarkin, 2015 Trichoptera : Taymyrodipseudon protopegasus Ivanov and Melnitsky, 2017 Yantardakh Ivanov and Melnitsky, 2017 Hydrobiosidae: Kliganigadukia taymyrensis Ivanov and Melnitsky, 2017 Yantardakh Ivanov and Melnitsky, 2017 : Siberoclea parapolaria Ivanov and Melnitsky, 2017 Yantardakh Ivanov and Melnitsky, 2017 : Archaeopolycentra yantardakh Ivanov and Melnitsky, 2017 Yantardakh Ivanov and Melnitsky, 2017 Diptera Lestremiidae: Caputmunda yantardakhica Fedotova and Perkovsky, 2016 Yantardakh Fedotova and Perkovsky, 2016 Corporesana khatanga Fedotova and Perkovsky, 2016 Yantardakh Fedotova and Perkovsky, 2016 Cretoperomyia dmitrii Fedotova and Perkovsky, 2016 Yantardakh Fedotova and Perkovsky, 2016 Cretomycophila ekaterinae Fedotova and Perkovsky, 2016 Yantardakh Fedotova and Perkovsky, 2016 Krassiloviola geniusloci Fedotova and Perkovsky, 2017 Yantardakh Fedotova and Perkovsky, 2017 Menssana norilsk Fedotova and Perkovsky, 2016 Yantardakh Fedotova and Perkovsky, 2016 Palaeostrobliella zherikhini Fedotova and Perkovsky, 2016 Yantardakh Fedotova and Perkovsky, 2016 Yantardakhiella pusilla Fedotova and Perkovsky, 2016 Yantardakh Fedotova and Perkovsky, 2016 Zherikhiniella pedicellata Fedotova and Perkovsky, 2016 Yantardakh Fedotova and Perkovsky, 2016

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Table 1. (Contd.) Taxon Locality Citation Lygistorrhinidae: Plesiognoriste zherikhini Blagoderov and Grimaldi, 2004 Yantardakh Blagoderov and Grimaldi, 2004 Protognoriste amplicauda Blagoderov and Grimaldi, 2004 Nizhnyaya Agapa Blagoderov and Grimaldi, 2004 : Apolephthisa bulunensis Blagoderov and Grimaldi, 2004 Bulun Blagoderov and Grimaldi, 2004 Ectrepesthoneura succinimontana Blagoderov and Grimaldi, 2004 Yantardakh Blagoderov and Grimaldi, 2004 Izleiina mirifica Blagoderov and Grimaldi, 2004 Yantardakh Blagoderov and Grimaldi, 2004 Nedocosia exsanguis Blagoderov and Grimaldi, 2004 Yantardakh Blagoderov and Grimaldi, 2004 Nedocosia sibirica Blagoderov and Grimaldi, 2004 Yantardakh Blagoderov and Grimaldi, 2004 Neuratelia maimecha Blagoderov and Grimaldi, 2004 Yantardakh Blagoderov and Grimaldi, 2004 incertae sedis: ?Thereotricha agapa Blagoderov and Grimaldi, 2004 Nizhnyaya Agapa Blagoderov and Grimaldi, 2004 Thereotricha sibirica Blagoderov and Grimaldi, 2004 Yantardakh Blagoderov and Grimaldi, 2004 Simuliidae: Simuliites yantardakh Perkovsky and Sukhomlin, 2016 Yantardakh Perkovsky and Sukhomlin, 2016 Ugolyakia kaluginae Perkovsky and Sukhomlin, 2018 Ugolyak Perkovsky et al., 2018 Psychodidae: Paleopsychoda zherikhini Azar, Adaymeh and Jreich, 2007 Zhdanikha Azar et al., 2007 Hymenoptera Ichneumonidae: Agapia sukatchevae Kopylov, 2012 Nizhnyaya Agapa Kopylov, 2012 Agapteron popovi Kopylov, 2012 Nizhnyaya Agapa Kopylov, 2012 Eubaeus abdominalis Kopylov, 2012 Nizhnyaya Agapa Kopylov, 2012 Urotryphon baikurensis Kopylov, 2012 Baikura Kopylov, 2012 Maimetshidae: Iberomaimetsha pallida Perrichot and Perkovsky, 2016 Yantardakh Perrichot and Perkovsky, 2016 Rotoitidae: Baeomorpha avamica Gumovsky, 2018 Yantardakh Gumovsky et al., 2018 Baeomorpha baikurensis Gumovsky, 2018 Baikura Gumovsky et al., 2018 Baeomorpha bianellus Gumovsky, 2018 Yantardakh, Romanikha Gumovsky et al., 2018 Baeomorpha caeleps Gumovsky, 2018 Yantardakh, Romanikha Gumovsky et al., 2018 Baeomorpha gracilis Gumovsky, 2018 Yantardakh Gumovsky et al., 2018 Baeomorpha ingens Gumovsky, 2018 Yantardakh Gumovsky et al., 2018 Baeomorpha popovi Gumovsky, 2018 Yantardakh Gumovsky et al., 2018 Baeomorpha quattorduo Gumovsky, 2018 Yantardakh, Romanikha Gumovsky et al., 2018 Baeomorpha quattoruno Gumovsky, 2018 Yantardakh, Ugolyak Gumovsky et al., 2018 Baeomorpha yantardakh Gumovsky, 2018 Yantardakh Gumovsky et al., 2018 Baeomorpha zherikhini Gumovsky, 2018 Baikura Gumovsky et al., 2018 Taimyromorpha pusilla Gumovsky, 2018 Yantardakh Gumovsky et al., 2018 Bryopsida Familia incertae sedis: Taimyrobryum martynoviorum Ignatov, Heinrichs, Schaf.-Verw. and Perkovsky, 2016 Yantardakh Ignatov et al., 2016

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Fig. 4. The first planidium from Taimyr, PIN, no. 3311/3243, Yantardakh, bottom view. the faunas of Taimyr amber is extremely important Perkovsky, 2019, etc.; Figs. 4, 5) would have been sim- (Olmi et al., 2019). ply technically impossible. A significant proportion of the results of recent studies of inclusions in Taimyr amber is based on the ACKNOWLEDGMENTS contribution of E.A. Sidorchuk. On her initiative and with her participation, more than a third of all amber We are grateful to A.P. Rasnitsyn and I.D. Sukatsheva samples with inclusions were collected, and without (Paleontological Institute of the Russian Academy of Sci- her participation, the description or redescription of a ences) for their constant attention to amber research, valu- able comments on the manuscript and their invaluable con- significant part of the most interesting samples (Igna- tribution to the study of the entomofauna of Taimyr amber; tov et al., 2016; Kathirithamby et al., 2017; Lyubarsky D.E. Shcherbakov (Paleontological Institute of the Russian and Perkovsky, 2017a, 2017b; Nadein and Perkovsky, Academy of Sciences) for the photographs of Hemiptera, 2018; Makarkin and Perkovsky, 2019; Jałoszyński and A.V. Gumovsky (Institute of Zoology, National Academy of

PALEONTOLOGICAL JOURNAL Vol. 53 No. 10 2019 A SUMMARY OF RECENT RESULTS IN THE STUDY OF TAIMYR AMBER 991

(a) 0.2 mm

(b) 0.2 mm

Fig. 5. Males of Baeomorpha from Yantardakh: (a) Baeomorpha caeleps Gumovsky, 2018, part of 10 gregarious male types, PIN, no. 3311/2341-2350; (b) B. popovi Gumovsky, 2018, holotype, PIN, no. 3311/1386.

Sciences of Ukraine) for a photographs of Baeomorpha, Azar, D., Adaymeh, C., and Jreich, N., Paleopsychoda R.S. Vargovich (Institute of Zoology, National Academy of zherikhini, a new Cretaceous species of moth flies from Sciences of Ukraine) for a photograph of a planidium. Taimyr amber (Diptera: Psychodidae: Psychodinae), Afr. Invertebr., 2007, vol. 48, no. 1, pp. 163–168. Azevedo, C.O., Alencar, I.D.C.C., Ramos, M.S., Bar- REFERENCES bosa, D.N., Colombo, W.D., Vargas, J.M.R., and Lim, J., Global guide of the flat wasps (Hymenoptera, Azar, D. and Engel, M.S., A sphaeropsocid bark in Bethylidae), Zootaxa, 2018, vol. 4489, no. 1, pp. 1–294. Late Cretaceous amber from Siberia (Psocoptera: Sphaeropsocidae), Trans. Kans. Acad. Sci., 2008, vol. 111, Blagoderov, V.A. and Grimaldi, D.A., Fossil Sciaroidea no. 1/2, pp. 141–146. (Diptera) in Cretaceous Ambers, Exclusive of Cecidomyiidae,

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PALEONTOLOGICAL JOURNAL Vol. 53 No. 10 2019 A SUMMARY OF RECENT RESULTS IN THE STUDY OF TAIMYR AMBER 993

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PALEONTOLOGICAL JOURNAL Vol. 53 No. 10 2019