Contents of Proceedings of the Zoological Institute

Volume 325 (2), 2021

Date of publication — June 25, 2021

Benthic fauna of the Kara Sea: new results and study prospects. Foreword

New data on the fauna and the biogeographic structure of the Hydrozoa, Scyphozoa, Staurozoa and Alcyonacea (Anthozoa) of the Kara Sea

The paper considers the species composition of the fauna of several cnidarian groups of the Kara Sea. The author presents a list of species of the studied groups and indicates the types of habitat for each species. The analysis was based on the literature data, the collections of the Zoological Institute of the Russian Academy of Sciences and material collected in the Kara Sea during the expedition to the R/V Professor Multanovsky in 2019. In total, 87 species of Hydrozoa, 3 species of Scyphozoa, 4 species of Staurozoa, and 5 species of the order Alcyonacea from the class Anthozoa were recorded for the fauna of the Kara Sea, based on the new material obtained by the author and published literature data. The report presents the biogeographic structure of the discussed cnidarian groups. According to the types of biogeographic ranges, the fauna of the above-mentioned cnidarian groups in the Kara Sea mostly consists of representatives of the Boreal-Arctic type of habitat (63%), the Boreal and Amphiboreal biogeographic groups each containing 12% of the total number of described species, and the Panoceanic and Arctic groups together accounting for only 9% and 4% of the fauna of the Kara Sea. Two species new for the Kara Sea, Neoturris pileata (Forsskål, 1775) and Neoturris pileata (Forsskål, 1775), are described. Neoturris pileata is an element of the warm-water Atlantic fauna that penetrated into the Kara Sea with waters of Atlantic genesis. Nausithoe werneri is an element of the cold-water Arctic fauna that penetrated into the Novaya Zemlya Trough of the Kara Sea from the north-western side from the St. Anna Trough, which was open to the Polar Basin.

The history of study, the taxonomic composition and the origin of the sea anemone fauna of the Kara Sea (Actiniaria, Anthozoa, Cnidaria)

Based on the study of new collection material and literature data, the composition of the sea anemone fauna of the Kara Sea is described. It includes 13 species of 7 families. Among them are 10 species that have previously been known for this region: Urticina crassicornis (O.F. Muller, 1776), Aulactinia stella (Verrill, 1864), Hormathia digitata (O.F. Muller, 1776), Allantactis parasitica Danielssen, 1890, Actinostola callosa (Verrill, 1882), A. spetsbergensis (Carlgren, 1893), Anthosactis janmayeni Danielssen, 1890, Halcampa arctica Carlgren, 1893, Edwardsia arctica Carlgren, 1921, Ptychodactis patula Appellof, 1893, and 3 species reported for the Kara Sea for the first time: Cactosoma abyssorum Danielssen, 1890, Haliactis arctica Carlgren, 1921, and Edwardsia vitrea Danielssen, 1890. The ranges of the species are described. A detailed consideration of the species’ geographical distribution, determined by the temperature, depth and salinity, allows the anemones to be subdivided into 3 groups: a) inhabiting the shelf and associated with high salinity; b) eurybathic, associated with the high latitudes, high salinity and a narrow interval of low temperatures; c) the most widespread, withstanding desalination and low temperatures. The geographic ranges of epibenthic polyps of the infraorder Thenaria have now been thoroughly studied, but the distribution of the representatives of the infraorder Athenaria in Arctic is known only from several finds. The distribution of the only Arctic representative of the suborder Ptychodacteae, Ptychodactis patula, is known from a fairly large number of finds, but its habitat requirements are little known. Therefore, our reasoning about the sources of the formation of the Kara Sea fauna is based mainly on the data derived from the thenarians. Our analysis has confirmed the participation of the species of Atlantic origin, most resistant to low temperatures, in the formation of the Kara Sea fauna, but no convincing evidence has been found for the participation of Pacific elements in the settlement of this region.

Polychaetes (Annelida: Polychaeta) of the Kara Sea

As a result of processing materials from the expeditions of the Saint-Petersburg State University in 2012–2013 and the Transarctic expedition of 2019, 131 polychaete taxa were recorded, of which 124 were identified to species. On the basis of data from previous and recent collections, the general list of polychaetes in the Kara Sea was updated; it includes 209 species belonging to 98 genera, 37 families, and 15 orders. The following species are reported for the first time from the sea: Abyssoninoe hibernica (McIntosh, 1903), Ampharete falcata Eliason, 1955, Brada rugosa Hansen, 1882, Microclymene acirrata Arwidsson, 1906; Nephtys pente Rainer, 1984; Ophelina abranchiata Stop-Bowitz, 1948, Scolelepis burkovskii Sikorski, 1994, and Travisia forbesii Johnson, 1840. Polychaetes were found at all stations; the number of species per station ranged from 1 to 28. Shannon’s indices (H'SpA and H'spB) showed that at most stations the diversity of species was high and resistance to stress (DE') was quite satisfactory. Only in a few cases some tension existed in the ecological situation, estimated by the index of difference in evenness and this applies, to an overwhelming degree, to the shallow water stations in the area of the outlet from the Ob and Yenisei bays. New data show a more pronounced Arctic character of the Kara Sea fauna in comparison with the polychaete fauna of neighboring seas. The influence of the Pacific and Atlantic oceans both on the polychaete fauna of the Kara Sea and on the polychaete fauna of the neighboring seas is noticeable and natural, which is confirmed by changes in the proportion of species of Pacific and Atlantic origin. The trophic structure of polychaetes is represented by four main groups; the distribution of trophic groups is closely related to environmental conditions.

Shell-bearing mollusks of the order Cephalaspidea (Gastropoda: Heterobranchia) of the Kara Sea: fauna and ecology

The survey of shell-bearing heterobranchs of the order Cephalaspidea of the Kara Sea has been done based on unpublished and literature data. The studied material includes that collected during expeditions aboard R/V Professor Multanovskiy in August–September 2019 and R/V Dalnie Zelentsy in August 2012. Additionally, the material of collections of the Zoological Institute of the Russian Academy of Sciences (Saint Petersburg) has been studied. The fauna of heterobranch molluscs of the order Cephalaspidea in the Kara Sea includes 18 described and 1 undescribed species that belong to 10 genera and 6 families. It has been specified that the type locality of Retusa semen Reeve, 1855 (Port Refuge) is located at Port Refuge National Historic Site of Canada, Devon Island, Nunavut, not in the Kara Sea. Retusa semen is considered here as species inquirenda because of the brief original description, significant differences in drawings of the shell of this species made by different authors, and the absence of type specimens. The presence of Diaphana minuta Brown, 1827, Laona quadrata (Wood, 1839), Philine sinuata Stimpson, 1851 and Retusa semen in the Kara Sea needs confirmation. Species numbers of heterobranch molluscs of the order Cephalaspidea decrease from the Barents Sea (21) through the Kara Sea (19) to the Laptev Sea (15) because of the decreasing proportion of boreal-arctic species. Among cephalaspids of the Kara Sea, the highest average abundance (individuals/m2) was observed for Cylichnoides densistriatus (Leche, 1878), and the highest average biomass for Cylichna corticata (Møller, 1842). The proportion of live individuals in the populations does not exceed 18% in late August – early September. The taxonomic structure of the cephalaspid fauna of the Kara Sea has been studied based exclusively on morphological characters. Further study of the fauna is needed based on molecular analyses of the material fixed in 96˚ alcohol.

Bryozoans of the Kara Sea: estimation of species diversity and patterns of biogeographic composition

Until now, information on the bryozoan fauna of the Kara Sea was as the most unordered in comparison to other Arctic seas of Russia. The information in literature is mainly based on the data collected more than 80 years ago. Collections carried out over the last 30 years made it possible to expand the understanding not only of the species richness of this group, but also to study its spatial variation within the sea. At the moment, the species list of bryozoans in the Kara Sea includes 230 names. Of these, 42 species were recorded for the first time in this area of the Arctic. The fauna richness varies with depth and geographic location of sampling sites. Comparison of the species composition of bryozoans within 6 sectors defined on the basis of differences in environmental parameters indicates the presence of a single faunistic complex. The biogeographic composition of the fauna is characterized by the predominance of the boreal-arctic species (67%) over the arctic (30%); the share of boreal species is only 3%. Comparison of bryozoans from the Kara Sea with faunas from other areas of the Arctic region indicates a closer similarity of its fauna with the fauna of the Barents Sea than with the fauna of bryozoans from the Laptev Sea.

Biodiversity and distribution of brittle stars (Echinodermata, Ophiuroidea) in the Kara Sea

Brittle stars are one of the leading components of the benthic communities in the Kara Sea. The fauna of the Kara Sea brittle stars is represented by 12 species. Ophiocten sericeum (Forbes, 1852), Ophiopleura borealis Danielssen et Koren, 1877, Ophiacantha bidentata (Bruzelius, 1805), and Ophioscolex glacialis Müller et Troschel, 1842 are most widespread within the sea. Based on the available data, distribution maps for each species were constructed. A comparative analysis of the Barents Sea, the Kara Sea and the Laptev Sea fauna was carried out. It was found that during evolution the fauna of Kara Sea brittle stars at the genus level was under balanced influence of autochthonous and allochthons processes; at the species level the autochthonous processes were predominant. The obtained value of the taxonomic uniqueness index characterizes the fauna of the Kara Sea brittle stars as quite isolated at all taxonomic levels. Six biogeographic groups were distinguished in the biogeographic structure of the fauna of the sea, of which the boreal-Arctic and high-boreal-Arctic forms are the most represented. The use of the Jaccard species similarity coefficients and Pearson correlation showed that the greatest similarity at the species level is observed between the Kara and the Laptev seas.

Notes on the fishes of the Severnaya Zemlya archipelago and the spawning area of polar cod Boreogadus saida (Gadidae)

Data on the fishes of the high-latitude Severnaya Zemlya archipelago (the North Land) is presented. The archipelago is located in the Arctic on the border between the Kara Sea and the Laptev Sea. The ichthyofauna of the archipelago has not been studied; therefore, even small collections are of interest. Fish samples were obtained during the expedition “Open Ocean: Arctic Archipelagos – 2019: Severnaya Zemlya”. In addition, the samples from this area in the collections of the Zoological Institute (ZIN) were studied, which have been received from polar expeditions to the Kara and Laptev seas during the entire era of polar research. The most significant fact is the discovery of mass accumulation of polar cod Boreogadus saida (Lepechin, 1774) larvae in Mikoyan Bay (Bolshevik Island), which gives evidence of important spawning grounds near Severnaya Zemlya. Indirect evidence of this can be found in the publications of polar explorers who overwintered on Severnaya Zemlya in the 1930s–1950s and have reported that the polar cod approaches the shores for spawning in August, in huge schools. The waters of Severnaya Zemlya represent the spawning area of polar cod in the central part of the Eurasian shelf, which is not mentioned in current literature. In addition to polar cod, a few more species are registered in samples from the coastal waters of the archipelago (depths to 38 m), rough hookear sculpin Artediellus scaber Knipowitsch, 1907, twohorn sculpin Icelus bicornis (Reinhardt, 1840) (family Cottidae), Liparis tunicatus Reinhardt, 1836, black-bellied snailfish L. cf. fabricii (Liparidae), Knipowich eelpout Gymnelus knipowitschi Chernova, 1999 (Zoarcidae) and three-spined stickleback Gasterosteus aculeatus (Linnaeus, 1758) (Gasterosteidae). In the deepwater straits, snailfish Careproctus sp. (174–234 m) and pale eelpout Lycodes pallidus Collett, 1879 (105–348 m) were found. The Arctic charr Salvelinus alpinus (Linnaeus, 1758) (Salmonidae) inhabits some lakes of the archipelago. This is the first finding of a three-spined stickleback in the east of the Kara Sea.

Volume 325 (1), 2021

Date of publication — March 25, 2021

The taxonomic status of the vipers of the Pelias (kaznakovi) complex in the middle-flow of the Kura River basin in the East Transcaucasia

Recently, the Caucasian viper was found in a number of localities in South Ossetia (Tuniyev et al. 2017a,b; Tuniyev et al. 2019) and in the lower gate of the Borzhom Gorge of Eastern Georgia. Animals from the new localities are compared to Pelias kaznakovi from the Western Caucasus – northeastern Turkey and P. dinniki. The canonical discriminate analysis (CDA) showed that the snakes from the left-bank basin of the middle flow of the Kura River in East Georgia and South Ossetia differ significantly from Pelias kaznakovi from Krasnodar Territory and Abkhazia in the mean values of several plastic and meristic characters. Thus, there is discrimination of the groups allocated according to geographic location and gender. Cluster analysis based on meristic characters of pholidosis showed the distance between P. dinniki and the remaining “kaznakovi” complex forms. Based on the results of the analysis of the mitochondrial cytochrome b gene, the snakes from the left-bank basin of the middle flow of the Kura River form a separate cluster, not sister to Pelias kaznakovi. The results obtained on morphology, genetics and ecology of the studied vipers suggest the taxonomic validity of the vipers from the left bank basin of the middle flow of the Kura River in Eastern Georgia and South Ossetia. The species is named after Boris Tuniyev, who made a significant contribution to the study of the Caucasian herpetofauna and taxonomy of the shield-head viper snakes. Among the representatives of the “kaznakovi” complex, males of the new species are characterized by the minimal values of body length, pileus length, head width, number of ventral and subcaudal shields, as well as the shields around the eyes; differences are also noted in the structure of the hemipenis; the females have the minimum values of body length, pileus length, head width, number of shields around the eyes and loreals, fewer wings of zigzag. All known finds are located in the left bank basin of the middle flow of the Kura River from the Borzhom Gorge (east slope of Meskheti Ridge) in Georgia to the mid-altitude mountain districts of South Ossetia (east foothills of Likhsky Ridge and south spurs of Central Caucasus). Biotopes of the new species are character- ized by more xero-mesophilic traits and a moderately warm mezo-climate than the typical mesophilic biotopes of North-Colchian (Krasnodar Territory – Abkhazia) and, especially, Adzharo-Lazistanian (Georgia – Turkey) P. kaznakovi living in the humid subtropical climate.

A new species of Gymnelus (Perciformes, Zoarcidae) from Greenland, similar to G. viridis

Zoarcid fishes of the genus Gymnelus Reinhardt inhabit the shelves of the North Pacific Ocean and the Arctic. A new species, G. pseudosquamatus sp. nov., is described from trawl samples taken at depths off South-West Greenland. It is most similar to the type species of the genus, Common Fish Doctor G. viridis, inhabiting the coastal waters of Greenland. The characters of the latter are specified on materials from the type locality, including the neotype and specimens of J.C.H. Reinhardt. Both species are in the group of Gymnelus with an interrupted supratemporal commissura, two supratemporal sensory pores (1+0+1), and a dorsal fin originating above the pectoral fin. The two species differ in a complex of characters, including habitus. In G. viridis, the trunk is roundish in cross section, highest above the beginning of the anal fin; the anterior rays of the dorsal fin are shortened, and covered with a thick and somewhat fleshy skin. In G. pseudosquamatus, the body is compressed and deeper anteriorly, the dorsal-fin rays are evenly elongated, and the fin membrane is thin. Differences in counts and measurements are statistically significant, including the number of vertebrae and rays in the dorsal, anal and pectoral fins, the number of teeth on the jaws, as well as the length and depth of the head, predorsal length, the length of pectoral fins, eye diameter and length of the gill slit. The color of G. pseudosquamatus, with 8–16 wide brown mottled bands, is also unusual, as the skin is dotted with light speckles that create the illusion of tiny scales, which is the reason for the name “pseudosquamatus”, the Deceptive Fish Doctor. While G. viridis is found inshore in a zone of macroalgae, the new species is found in deeper waters (100–457 m) along the shelf edge of South-West Greenland. The name Ophidium stigma Lay et Bennett, 1839 (=Gymnelus stigma) should be excluded from the synonymy of G. viridis, since the original description mentions the presence of scales on the body, which are absent in Gymnelus.

Phylogeography of a Darevskia (caucasica) complex (Lacertidae: Sauria) based on the cytochrome b mitochondrial gene analysis

The range of superspecific complex of rock lizards Darevskia (caucasica) (D. alpina, D. c. caucasica, D. c. vedenica, D. daghestanica) includes the Main Caucasian ridge and its spurs on the southern and northern macroslopes within heights from 500 m to 3200 m above sea level. A fragment of the mitochondrial cytochrome b gene (1096 bp) was analyzed in 48 specimens of these lizards. Specimens were collected in 23 localities. We have found 39 unique haplotypes. The minimum genetic diversity was found in D. c. vedenica, maximum in D. c. caucasica. All taxa of the complex, except for D. c. vedenica, demonstrate genetic heterogeneity. Four clades can be distinguished on the dendrogram of phylogenetic relationships. They correspond to D. alpina, D. c. caucasica, D. c. vedenica, D. daghestanica. D. c. caucasica forms two distinct haplogroups. On the phylogenetic tree, the sample of D. c. vedenica formed an independent clade close to D. c. caucasica. The Darevskia (caucasica) complex is polyphyletic according to the analysis of the mitochondrial marker. Nucleotide sequences of D. alpina formed a single clade with Darevskia (saxicola) complex representatives and combined with the sequences of D. brauneri into subclade. On the example of the population of D. daghestanica, found in a forest biotope without rocky outcrops and genetically not differing from individuals from the adjacent territory, the ability of rocky lizards to master biotopes unusual for them in a short period of time was demonstrated. This example demonstrates the ability of rock lizards to adapt to unusual biotopes in a short period of time. Genetic distance between D. c. vedenica and D. c. caucasica is comparable to the distance between D. c. caucasica and D. daghestanica. We assume the existence of an independent form of D. caucasica ssp. inhabiting South Ossetia and adjacent territories.

The Planthopper family Achilidae (Hemiptera: Auchenorrhyncha: Fulgoroidea) in the United Arab Emirates

This is first review of the planthopper family Achilidae of the United Arab Emirates. Two achilid species are reported from the country – Akotropis fulgens Linnavuori, 1973 and Deferunda incompta Dlabola, 1961. The finding of D. incompta in the UAE is the most western point of the genus Deferunda Distant, 1912 distribution and the first record of this genus from the Arabian Peninsula. External view photos of A. fulgens and D. incompta are provided and the latter is illustrated for the first time.

Remains of large mammals from the Epigravettian site of Yudinovo

The Epigravettian site of Yudinovo (Bryansk oblast, Russia) was discovered in 1930 by K.M. Polikarpovich. It is located in the Sudost’ river valley and has a unique stratigraphy. This article presents the results of the study of the remains of large mammals from Yudinovo and also discusses their significance in revising the former interpretation of the existence of the site during the very end of the final part of the Late Pleistocene. In total, 38 268 mammalian bones were identified from cultural layers excavated between 1947–2019. The faunal assemblage is relatively small with a dominance of woolly mammoth and arctic fox, typical of a cold and dry tundra-steppe environment. We undertook stable isotopic tracking from samples of bones. Our analyses confirm the hunting of both adult and juvenile larger mammals by ancient humans. Based on the eruption sequence and wear of the milk teeth from young animals, we were able to clarify the season of their death. It seems that these individuals were hunted during the late spring or early autumn. Traces of gnawing by dogs were recorded on a few bones. We also present in this article the results of the study of so-called “dwellings”, constructed by stacking up body parts and bones that were extracted from carcasses of freshly killed mammoths. We interpret these structures as middens representing the remains of ritually deposited hunted game. It can be assumed that these “dwellings” were probably an important part of the socio-symbolic system of the peoples, who created them.

Faunal complex of the Early Pleistocene Muhkai 2 locality

The Early Pleistocene locality Muhkai 2 was discovered in Central Dagestan, northeastern Caucasus, Russia, in 2006. Archaeological investigations of the site have been directed by Corresponding member of RAS H.A. Amirkhanov (Institute of Archaeology RAS, Moscow). The article presents the results of the study of osteological material from the excavations of 2006–2017. The cultural layers with stone tools and mammal bones are found within stratum of brownish-grey petrified loams with reversed magnetization. We identified 14 large species of mammals, which are indicators for the dating of the faunal assemblage and crucial in identifying the specific characteristics of the natural environment at the time of site formation. Stenon horse and Etruscan wolf dominate the spectrum of species at Muhkai 2. All large mammal species from the site are inhabitants of open and semi-open landscapes and animals indicative of closed biotopes are absent. The biotope appears to have been a dry savannah-steppe with small areas of forest vegetation. The climate at this time was warm and quite arid. Most likely, the animals died here due to natural causes and their carcasses were buried at the bottom of an ancient, slightly saline and shallow, temporary body of water. Data from the theriofauna place the age of the site Muhkai 2 within the chronological range of 2.1–1.77 Mya.

Parameters of ontogeny and population dynamics modeling of Panagrolaimus detritophagus (Nematoda: Rhabditida) in vitro

The parameters of individual development and population cycle in in vitro nematodes Panagrolaimus detritophagus were revealed. The nematodes are bacterial feeders and commensals of the cerambycid Monochamus galloprovincialis from the pine Pinus sylvestris; nematodes use beetles as vectors. Mean development time (T) from egg to juvenile is 1–2 days for J2, 3–4 days for J3, and 4–7 days for J4; to adults (G, generation) 7 (6–8) days. In vitro the population cycle is equal to 4 generations and ends with 90% of survival juveniles (J3, day 34). In the growth phase of the population, the proportion of eggs exceeds the proportion of other stages of the developmental cycle: 39±11% for 7 days; 53±10% for 21 days. The average oviposition rate of females is 4.5±1.3/day and only 56±12% of eggs proceed to immediate development (hatching and molting of juveniles). The remaining mass of eggs enter development only after 27 days (4 individual generations). This feature may be considered as a form of delay or a brief diapause at the egg stage. Individual females may accumulate up to 4 synchronous eggs in the body and lay them simultaneously. The average life span of an adult female is 13–20 days. Formulas for the exponential growth of the number of females and the total nematode population have been developed.

Variability of thermoregulation parameters of the common viper

The results of processing the data of continuous recording of body temperature using loggers obtained with the enclosure keeping 23 individuals of the common viper Vipera berus L. in Karelia (62.068310°N, 33.958824°E) are presented. Nine temperature characteristics, subdivided into 2 groups, were quantitatively evaluated – thermoecological, describing the dependence of the course of the viper’s body temperatures on the heat flux of the medium, and thermophysiological, stable parameters of thermoregulation. The factors that determine the variability of temperature characteristics are considered: time of day, weather conditions, morphometric indicators of an individual. A wide individual variability of thermoecological indicators and low variability of thermophysiological indicators were revealed, which was not associated with significant differences in different individuals. The parameter “maximum voluntary temperature”, which characterizes body temperature while avoiding overheating, is the closest to the physiological parameter of thermoregulation; obtaining such data is a very laborious process associated with deciphering the snake’s behavior from video recordings. An indicator that gives close values to the maximum voluntary temperature, which has a very simple calculation algorithm – “median active temperature (for a sample of temperatures above 30°C)”; however, it requires a volumetric series of continuously recorded temperature records for analysis. An accurate, statistically robust “maximum typical temperature” parameter gives an idea of the true maximum temperature characteristic of a species, but has a complex calculation method. These thermophysiological parameters are recommended to be used for intrapopulation and interspecies comparisons.

On the circumstances of the transfer of the Lepidoptera collection of Grand Duke Nikolai Mikhailovich to the Zoological Museum in St. Petersburg

The work examines the circumstances of the transfer of the largest collection of butterflies, collected by Grand Duke Nikolai Romanov for 26 years, to the Zoological Museum of the Imperial Academy of Sciences in St. Petersburg in 1899–1900. The gift was made on the condition that the collection would retain the name of the donor; that, because of its considerable size, there will be a special custodian; that it would remain untouchable and without resupply, except for the species that may come from the Russian Empire; that it will be available for the work of scientists and professionals interested in the field. Two requirements were added a little bit later. The collection should have been kept in the same cabinets as it was at the Grand Duke’s palace. Otto Hertz was to be left the custodian with the position of senior zoologist. The main sources of the article are the minutes of the meetings of the Physics and Mathematics Department of the Imperial Academy of Sciences, museum reports, books of receipts from the Scientific Archives of the Zoological Institute of the Russian Academy of Sciences, materials of the St. Petersburg branch of the Archive of RAS and the Russian State Historical Archives.



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Last modified: June 25, 2021