05_Perkovsky_01_22.indd UDC 595.74:553.99:553.555(1-18:477.82:571.511) TWO DIFFERENT CRETACEOUS WORLDS: TAIMYR AND KACHIN AMBER TRICHOPTEROFAUNAS E. E. Perkovsky Schmalhausen Institute of Zoology NAS of Ukraine, vul. B. Khmelnytskogo, 15, Kyiv, 01030 Ukraine E-mail: perkovsk@gmail.com E. Perkovsky  (https://orcid.org/0000-0002-7959-4379) Two Diff erent Cretaceous Worlds: Taimyr and Kachin Amber Trichopterofaunas. Perkovsky, E. E. — Polycentropodidae constitute 60 % of Taimyr amber caddisfl y species with known males, and only 4.8  % of caddisfl y species with known males in Kachin amber. Micro-caddisfl ies obviously dominate in Taimyr amber (Archaeopolycentra, Polycentropodidae), Kachin (Burmese) amber (Palerasnitsynus, Psychomyiidae) and New Jersey amber (Hydroptilidae); both Psychomyiidae and Hydroptilidae are absent in Taimyr amber, Polycentropodidae are absent in New Jersey amber and rare in Kachin amber. Th e domination of Polycentropodidae was proposed as a new characteristic of Baeomorpha Realm, their rarity proposed as a new characteristic of Isoptera Realm. K e y w o r d s : Insecta, Trichoptera, Polycentropodidae, Hydroptilidae, Taimyr amber, burmite, Rovno amber. Zoodiversity, 56(1): 51–56, 2022 DOI 10.15407/zoo2022.01.051 Paleontology 52 E. E. Perkovsky Introduction Santonian Taimyr amber trichopterofauna with 11 species was “on the 1st place among all Cretaceous deposits counting also the rock imprints” (Ivanov & Melnitsky, 2017, p. 131) in 2017; three species and.species were added in 2021 and 2022 (Melnitsky & Ivanov, 2021, 2022 a, b). It put Taimyr trichopterofauna on the second place aft er fauna of burmite (earliest Cenomanian Kachin amber) with 45 species versus 7 species in 2017 (Ivanov & Melnitsky, 2017, 2021; Wichard et al., 2022). Latest great advances in the burmite studies were reviewed and checklisted by A. Ross (2019, 2020, 2021); recent results in the Taimyr amber studies were sum- marized by E. E. Perkovsky & D. V. Vasilenko (2019). Th e goal of the paper was to compare some characters of the two biggest Cretaceous trichopterofaunas. Material and methods Twelve species of Taimyr amber caddisfl ies were collected in Yantardakh (Santonian) in 1971 by the expe- dition of V. V. Zherikhin and I. D. Sukacheva and in 2012 by expedition of the D. S. Kopylov, E. A. Sidorchuk, and D. D. Vorontsov that collected more than 60 kilograms of retinite (Rasnitsyn et al., 2016; Perkovsky & Vasi- lenko, 2019). All trichopterans found by the latter expedition except the holotype of Siberoclea parapolaria Iva- nov et Melnitsky, 2017 were selected by V. V. Martynov (Slavyansk), primary cutting and polishing was done by A. P. Vlaskin (Rovno). All specimens were fi nally polished by by E. A. Sidorchuk. It is very interesting that only one species appears to be common for both 1971 and 2012 Yantardakh collections (Ivanov & Melnitsky, 2017, 2021). Single assumed calamoceratid species was described from Santonian of Ugolyak from the material, collected by I. D. Sukacheva in 1973 (Botosaneanu & Wichard, 1983); assumed philopotamid was reported, but not named from Santonian of Bulun (Botosaneanu & Wichard, 1983; Perkovsky & Vasilenko, 2019). All the Taimyr material is housed in the Paleontological Institute, Moscow (PIN). Th e Kachin amber (‘Burmese amber’, or burmite in the nearly all papers published before 2020) is mined in Hukawng Valley (Kachin State, Myanmar) (Rasnitsyn et al., 2016).  Results Th irty-eight percent of the reported Taimyr species (Ivanov & Melnitsky, 2021) as well as at least 47 % of the reported specimens (Botosaneanu & Wichard, 1983; Ivanov & Melnitsky, 2017, 2021) belong to Polycentropodidae; in Yantardakh their share reaches 42  % of species and at least half of specimens; genus Archaeopolycentra Botosaneanu et Wichard, 1983 with fi ve species in Yantardakh comprises 42 % of all species and half of specimens and even 67 % of all species and 78 % of specimens in the new (2012) collection. Th ree Kachin polycentropodid species from three genera (Wichard, 2021) constitute only 6.7  % of all the named Kachin trichopterans, while Kachin Wormaldia McLachlan, 1865 (Philopotamidae) are represented by nine species as well as Palerasnitsynus Wichard, Ross et Ross, 2011 (Psychomyiidae); Palaeopsilotreta Wichard et Wang, 2017 (Odontoceridae) is represented by fi ve species and Cretapsyche Wichard et al., 2018 (Cretapsychidae) by four species; these four genera constitute 60 % of all known Kachin trichopteran diversity. Mi- cro-caddisfl ies obviously dominate both in Taimyr (Ivanov & Melnitsky, 2021) and Kachin (Wichard et al., 2018; Wichard, 2021) amber, but these micro-caddisfl ies belong to the dif- ferent families (table 1). Th e share of polycentropodid species in Yantardakh is 6.3 times higher than in Kachin amber. Comparable share of polycentropodid species is known only in Priabonian European ambers and Florissant (Ivanov et al., 2016): e. g., in the Baltic amber their share constitutes 34.8 % of species (Ivanov et al., 2016) and 80 % of specimens (Wichard, 2021). Th e ratio between polycentropodid share in equable Baltic (Archibald & Farrell, 2003; Radchenko & Perkovsky, 2021) and tropical Dominican amber (two from 32 Dominican species, 6.25 %; Wichard, 2007; Wichard & Neumann, 2021) equals 5.6, so it is nearly the same as the ratio between Yantardakh and Kachin polycentropodid share. Moreover, a male of Kachin poly- centropodid Neucentropus (Hnamadawgyia) macularis (Wang et al., 2019) is unknown, although “the description of extinct polycentropodid species embedded in amber should consider only the males” (Wichard, 2021, p. 3); Polycentropodidae comprise 4.8  % of Kachin species with known males. Four Taimyr species (both hydrobiosids, leptocerid and calamoceratid) are known only based on the holotypes which are females or specimens of unknown sex, thus, Polycentropodidae include 60 % of Taimyr species with known males. 53Two Diff erent Cretaceous Worlds: Taimyr and Kachin Amber Trichopterofaunas Table 1. Caddisfl ies from Taimyr and Kachin amber (the dagger symbol † indicates taxa known only as fossils)  Family Genera (number of species) Kachin amber Taimyr amber Philopotamidae Wormaldia (9) — Dipseudopsidae — †Taymyrodipseudon (1) Psychomyiidae †Palerasnitsynus (9) — Kambaitipsychidae †Myanpsyche (1) — Pseudoneureclipsidae †Amberclipsis (3)  — Polycentropodidae †Electrocentropus (1), Neucentropus (1), Neureclipsis (1) †Archaeopolycentra (6) Superfamily Psychomyioidea   Family incertae sedis †Protoclipsis (3) — Rhyacophilidae — Rhyacophila (1) Hydrobiosidae — †Palaeohydrobiosis (1), †Kliganigadukia (1) Hydroptilidae †Burminoptila (1), †Cretacoptila (1) — Helicopsychidae †Cretahelicopsyche (1)  — †Taymyrelectronidae — †Taymyrelectron (1) †Burmapsychidae  †Burmapsyche (2) — †Cretapsychidae  †Cretapsyche (4) — Calamoceratidae †Bipectinata (1) †Cretaganonema (1) †Calamodontus (1) Odontoceridae Psilotreta (1), †Palaeopsilotreta (5) — Leptoceridae — †Praeathripsodes (1) †Siberoclea (1) Domination of Polycentropodidae is even more evident at the level of specimens: 69 % of males belong to Polycentropodidae. Hydrobiosidae is unknown from all Cretaceous deposits, except Taimyr amber (Iva- nov & Melnitsky, 2017). Larvae of Hydrobiosidae are free-living active predators under stones in clean rapid streams (Ivanov & Melnitsky, 2017). Th eir larvae, as well as larvae of Rhyacophilidae (Ivanov & Melnitsky, 2017), are dependent on cool and fast moving water for their development, preferably that of the smaller rivers. As it was assumed by Ivanov & Melnitsky (2017), the richness of Hydrobiosidae in the Taimyr deposits might indicate at the cool climate and relatively low water temperatures in streams. Th is reason may explain the polycentropodid domination in the Taimyr trichopterofauna. Polycentropodidae is a single family known from both Kachin amber and Yantardakh (16.7  % of all families) vs. fi ve Yantardakh families (83.3  %) common with the Baltic amber; all Taimyr amber trichopterofauna has 3 from 8 families (37.5  %) common with Cenomanian Burma (table 1; third common family is Philopotamidae). Th e Taimyr and Kachin amber faunas do not have any common caddisfl y genera; one of Yantardakh genera (Rhyacophila) is known from Baltic amber. Half of the extant Kachin genera belong to Polycentropodidae. As we indicated earlier (Perkovsky et al., 2018), the right tributaries of the Kheta and left tributaries of Kotuj run mainly along the Putorana Plateau, which was present in the Cretaceous, bounded by granite canyons, creating a narrow area for the small freshwater streams surrounded by the amber forest. Th e Ugolyak (Perkovsky & Vasilenko, 2019) is the only known important retinite locality of the Kheta Formation of the river catchment area in the fl at Khatanga lowland (Perkovsky et al., 2018) where the Cretaceous river val- leys were not bounded by granite canyons. Th ere is not much space for small tributaries and lakes in the granite canyons, however, the fl at area surely allow them to exist, so the fi nding of the single assumed Taimyr calamoceratid species in Ugolyak and the absence of 54 E. E. Perkovsky polycentropodids in the taphocoenosis could be not accidental. In the Northern Hemisphere two Cretaceous zoogeographic realms (“Baeomorpha” and “Isoptera”) were recognized (Gumovsky et al., 2018). Th e Baeomorpha Realm, with a temperate or warm temperate climate, is characterized, in addition to the presence of nu- merous parasitic wasps of the genus Baeomorpha (Rotoitidae), by very abundant aphid fos- sils (eight families from four superfamilies are known from this realm only), few termites and no webspinners. Th e Isoptera Realm, which had a warmer climate, contained very few Rotoitidae, few aphids (oft en with strongly reduced hind wings), whereas termites and webspinners (Perkovsky et al., 2020) were abundant and diverse (webspinners at least in the south of the realm). Water insects were not used in the characterization of the realms; 70 species of Cretaceous amber caddisfl ies (Ivanov & Melnitsky, 2017, 2021; Wichard & Azar, 2018; Wichard et al., 2022) appear instrumental in it.  Archaeopolycentra is the only not monotypic genus of caddisfl ies in Baeomorpha Realm; at the same time all three Polycentropodidae Isoptera Realm genera are monotypic versus half of monotypic genera in all other families. It is noteworthy to mention a di- versity of Burmese Pseudoneureclipsidae which is the family closest to Polycentropodidae (Chamorro & Holzenthal, 2011). Th ere are reasons to believe that Protoclipsis Wichard et al., 2022 also belongs to Pseudoneureclipsidae, in particular because the structure of the in- ferior appendages of diff erent species of Protoclipsis is similar either to Antillopsyche Banks, 1941 or Amberclipsis Wichard et al., 2022 (Wichard et al., 2022). Nominative extant genus Pseudoneureclipsis Ulmer, 1913 inhabits large southern parts of the Old World including southeastern, eastern, and southern Asia; the Bismarck Archipelago; Central Africa and Madagascar; and the Mediterranean Region; the second extant pseudoneureclipsid genus Antillopsyche has been found only on Cuba, Hispaniola, and Puerto Rico (Malicky, 2020); extinct species of Antillopsyche reported from Miocene Dominican and Mexican amber. Some genera belonging to thermophile Psychomyiodea families were even three times more diverse in Kachin amber forest (table 1). Micro-caddisfl ies from the genus Palerasnit- synus have forewings with lengths of 1.8–2.6 mm and account at present for almost 60 % of all caddisfl ies in Kachin amber (Wichard et al., 2018). Oft en they can be found in aggrega- tions with 20 to 100 specimens in one piece of amber, thus indicating swarming activities of the adults (Wichard et al., 2018, fi g. 12). Kachin Wormaldia with wing lengths of half of the species 2.5–2.8 mm (Wichard et al., 2020; Wang et al., 2021) are “about 25 % small- er than their recent relatives and, like the psychomyiids, can be considered to be extinct “microcaddisfl ies” (Wichard, 2021, p. 6). Compared with Palerasnitsy nus the Wormaldia specimens occur sporadically in Kachin amber and do not seem to swarm (Wichard et al., 2020). Taimyr philopotamid from Bulun with wing length 5.5–5.7 mm (Botosaneanu & Wichard, 1983) is two times bigger. New Jersey and Tennessee amber species are as big as recent philopotamid species as well. Except for burmite, more than two species of cad- disfl ies of Isoptera Realm were found only in Turonian New Jersey amber, where 4 out 9 species (44.4 %) belong to Hydroptilidae; endemic subgenus Nanoagraylea Botosaneanu, 1995 (3 species) and genus Novajerseya Botosaneanu et al., 1998 are characterized by very small sizes. Hydroptilidae comprise 3.8 % of species in well-studied Baltic amber (Ivanov et al., 2016) versus 11.4 % in more southern (Mitov et al., 2021) and less studied Rovno amber (Melnitsky et al., 2021 a, b) and 25 % in tropical Dominican amber (Wichard, 2007; Wich- ard, Neumann, 2021); polycentropodid/hydroptilid species ratio equals 9.3 in Baltic amber (Ivanov et al., 2016; Melnitsky & Ivanov, 2019) and 4.5 in Rovno amber (Perkovsky, 2017; Melnitsky et al., 2021 a, b, c) versus 0.25 in Dominican amber. Eskov with co-authors as- sumed that hydroptilids “initially, from the time of their origin, were related to the tropical and subtropical regions avoiding the temperate ones” (Eskov et al., 2004, p. 43). It is quite probable that this assumption is also true with regard to the thermophile forms of other trichopteran families as well. 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