Contents of Proceedings of the Zoological Institute

L.I. Khosatzky. 100 years anniversary (instead of the foreword).

The life, scientific and social activity of Dr. Lev I. Khosatzky (1913–1992), a remarkable zoologist, herpetologist and paleontologist of Leningrad (Saint Petersburg) is briefly outlined. His affiliation with Leningrad (now Saint Petersburg) State University, Zoological Institute, Russian Academy of Sciences (Saint Petersburg), Paleontological Institute, Russian Academy of Sciences (Moscow), as well as the All-Union Paleontological Society (Saint Petersburg) is discussed. A wide scope of his scientific interests is analysed. The list of Khosatzky’s students contributed to zoology and paleontology is provided.

Plates.

This paper presents results of the comparative morphometric study of the Horsfield’s tortoise (Agrionemys horsfieldi) from the Ogzi-Kichik archaeological site of the Stone Age (Southern Tajikistan) and a sample of representatives of modern populations of this species. From the time of existence of the Ogzi-Kichik population to the Present, noticeable changes (decrease of the shell length, its height and convexity and changes in the proportions of some plates of the shell) have occurred in the shell morphology of this species. These changes, probably, reflect intensification of the burrowing adaptations connected with the aridization of the climate in the Southern Tajikistan during the mentioned historical period.

This paper presents a review of the chelonological research of L.I. Khosatzky (1913–1992), a well-known Soviet zoologist-herpetologist and paleontologist from Leningrad (now Saint Petersburg). It provides an annotated bibliography of his 115 publications on turtles. Khosatzky’s turtle research was devoted to the origin of turtles, systematics, phylogeny and biogeography of fossil and recent turtles, biomechanics, morphology and evolution of the turtle shell, and the ecological physiology of turtles. Khosatzky supervised numerous student works (course projects, graduate and candidate theses) on different aspects of turtle biology, the results of which were only partially published. Some of Khosatzky’s students became well-known specialists on turtles.

Methods of modeling and predicting the potential ranges to determine the limiting factors and conservation priorities are increasingly important in modern herpetological studies. In present paper we provide an analysis and forecast of the potential distribution of the Central Asian rock agamid species, – Turkestan rock agama (Paralaudakia lehmanni), and analyze the characteristics of its habitat preference using Maxent (www.cs.princeton.edu/~schapire/maxent/). We used the data from 19 bioclimatic variables (Bio 1–19) obtained from the database WorldClim (http://www.worldclim.org/current/), and the geographical coordinates of 70 localities from the whole distribution range, explored at different times. This method allows identifying the factors that have a major impact on the distribution of living organisms. The constructed model of distribution of P. lehmanni revealed enough good performance (AUC = 0.990 at variance 0.004) and predicted that the most suitable Turkestan rock agama habitats are located in the eastern Uzbekistan, western and central Tajikistan, as well as in adjacent areas of the western Kyrgyzstan and northern Afghanistan.

This is the first annotated check-list on taxonomic structure of Pseudotrapelus, a genus that was considered previously to be monotypic. The paper is based on a large collection (more than 250 specimens from the whole range of the genus), and represents the first publication about this genus in Russian. Five Pseudotrapelus species are divided into two groups: the African one with Ps. sinaitus sensu stricto and the Arabian one with Ps. neumanni, Ps. aqabensis, Ps. dhofarensis and Ps. jensvindumi. The division into two groups was caused, probably, by separation of Arabia from Africa 28–23 mya. The collision of these landmasses via the Gomphoterium land bridge, south from the Sinai Peninsula 18–15 mya caused the secondary colonization of Arabia by Pseudotrapelus (this time by Ps. sinaitus) and formation of the sympathry zone and, probably, hybridization with the Arabian species group in the Aqaba area. South-Western Arabia is the center of recent taxonomic diversity of Pseudotrapelus.

Excavations and sediment screenwashing at the Middle Jurassic Berezovsk Quarry locality in Krasnoyarsk Territory, Russia, yielded rare isolated teeth, osteoderms and fragments of cranial bones of crocodyliforms. All these remains were referred to Goniopholididae indet. on the basis of the following combination of features: a relatively narrow and long snout, contribution of splenials to the mandibular symphysis, dermal sculpturing consists of almost circular and slightly elongated oval pits, polygonal ventral osteoderms, and conical teeth with strongly striated crowns with weakly developed unserrated lateral carinae. The Berezovsk goniopholidid represents one of the oldest goniopholidid records in Asia and, geographically, the northernmost occurrence of this group in the Jurassic of Asia.

Peculiarities of the ventral color pattern variation in two cryptic species of newts of the genus Lissotriton (L. vulgaris and L. lantzi) have been examined. The belly and throat of these species have black spots on the pale-grey, yellow or orange background. 626 specimens, including 279 males and 244 females of L. vulgaris (11 samples from Russia, Ukraine, Romania, and Turkey), and 52 males and 51 females of L. lantzi (7 samples from Russia and Abkhazia) were used in this study. It was shown that, as a rule, the pattern of ventral coloration in these two species is different.

The paper presents new data on the geographic distribution and frequency of colour aberration cineracea Szczerbak, 1960 (= concolor) in Podarcis tauricus from the Crimea. It is established that the finds of individuals belonging to this aberration are grouped at the junctions of landscapes of the Crimean Mountains, where the ancient geochemical boundaries are localized. The majority of aberrant specimens were found in populations living on the Upper Jurassic conglomerates containing pebbles of crystalline rocks, and within the Middle Jurassic volcanic massif of Karadagh. The frequency of manifestation of this mutagenic type of color is almost independent of the altitude above sea level and more likely it is determined by geochemical composition of rocks prevailing in the locality which indirectly affects the synthesis of melanin pigment, rather than by climatic characteristics of the habitat. In addition, a description of the unique color aberration plumbea, discovered in a single locality in the Sudak District of the Crimea near the upper limit of the species distribution, is provided.

Some general principles of zoogeography are considered. The regionalization based on various animal groups might result in different zoogeographic schemes. Therefore, it would be quite desirable to accumulate reliable data with various taxonomic groups as a subject of comparative zoogeography. Zoogeographic regionalization should be based on animal distribution itself but not on any other external factors. The identification of realm affinities of species should include both the recent distribution of the species and its phylogenetic position (relationships) within higher taxa (species groups, genus, etc.). The distribution of amphibian species in the north, west (Atlantic archipelagos) and east of temperate Eurasia was analysed in order to identify the borders of the Palearctic Realm. Special attention was paid to the southern limits of the Palearctic in North Africa (Sahara Desert), Middle East (Sinai, Arabian Peninsula, Iran), Afghanistan, Pakistan, western Himalaya, as well as in China аnd Japan. Outlined territory of the Palearctic harbors 214 amphibian species from 47 genera, 17 families and two orders. The composition of amphibians of the Palearctic is analysed in taxonomic and zoogeographic aspects. The Palearctic share was evaluated in relation to amphibians of the World, endemism level among species, genera, and families was calculated as well. Differentiation of genera (monotypic and other, in terms of species richess and range’s size) as well as species was discussed. Attached are the description of a new toad genus, Strauchophryne gen. nov., and full list of amphibian species, genera and families of the Palearctic, with designation of realm affinity and endemism for all species.

An inventory of the insectivorous mammal fauna of the Ba Vi National Park (northern Vietnam) based on the 1993–2013 surveys was conducted. A total of five species of Eulipotyphla were found in different types of the montane broad-leafed and mixed forests and anthropogenic habitats at the altitudes 50–1000 m a.s.l. New faunistic records of the short-tailed mole Euroscaptor subanura, the Etruscan shrew Suncus etruscus and the Taiwanese gray shrew Crocidura tanakae from Vietnam are reported.

The forelimb musculature of the relict rodent Laonastes aenigmamus Jenkins et al., 2005 (Diatomyidae) was studied for the first time. Ctenodactylus gundi and Chinchilla lanigera were also first included in the morphological analysis for comparative purposes. These species belong to different infraorders, Ctenodactylomorphi and Hystricognathi, the members of which could be closely related to Laonastes according to available hypotheses. The peculiar features of the examined muscles, like the more primitive topography of the m. rhomboideus, the unusual insertion of the m. endopectoralis and absence of the m. flexor digitorum sublimis, were revealed in Laonastes. In addition, a number of common characters in the structure of the locomotor musculature were also observed in Laonastes and Ctenodactylus: a single undifferentiated condition of the m. trapezius; the muscular origin of the m. sternomastoideus; the unusual attachments of the m. latissimus dorsi and the unique origin of the m. dorsoepitrochlearis unnoted in other rodents. The majority of these features are probably indicative of more archaic organisation. The obtained results support the hypothesis of the close affinity between Diatomyidae and Ctenodactylidae proposed on the basis of the moleculargenetic and some morphological data.

Dollosaurus lutugini (Yakovlev, 1901) is the only valid species known from the territory of the former USSR. It was described from Campanian deposits of Eastern Ukraine on the basis of an incomplete skeleton. This study provides a description of an osteological material including the previously undescribed epipterygoid and squamosal. Phylogenetic analysis of a data matrix of 37 terminal taxa and 135 characters shows that D. lutugini and Prognathodon solvayi Dollo, 1889 are sister taxa. P. lutugini is distinct from P. solvayi by possession of smooth enamel surface of teeth, depression of anteriormost trunk vertebrae condyles, larger relative length of cervical vertebrae, and a distinct, horizontally interdigitating articulating surface of the splenial and angular. We conclude that D. lutugini is in Prognathodon clade; generic name of Dollosaurus Yakovlev, 1901 is a subjective junior synonym of the Prognathodon Dollo, 1889.

A new species of ichneumon-fly, Ctenochira phyllocolpae sp. nov. is described from Russia (Karelia, Transbaikalia and Yakutia). The new species was reared from three species of gall-forming sawflies of the genus Phyllocolpa. Ctenochira phyllocolpae sp. nov. and a closely related Palaearctic species C. romani (Pfankuch) and the Nearctic C. deplanata Townes et Townes are distinguished and characterized in a key. Illustrations and photos of all these species are given.

Two new species of the Palaearctic Eucnemidae: Microrhagus nuristanicus sp. n. from the Eastern Afghanistan and Entomophthalmus kirejtshuki sp. n. from the Russian Far East are described.

A new species of Pogonophryne sarmentifera sp. nov. from the Ross Sea (Southern Ocean) is described. The new species belongs to the «mentella» group of the genus Pogonophryne and is characterized by having a mental barbel long (68.0–70.0% of head length), sarment-shaped with a slender terminal expansion (5.9–8.1% of barbel length) composed of rows of short and flattened incubous processes; сanals of seismosensory system top of head have the small pores; weakly developed internasal organ, peritoneum is light.

New records of Darevskia alpina from the vicinities of Zemo Bachiani Village in South Ossetia are presented and discussed: D. brauneri from the place of the confluence of the Luhunistskale River into the Rion River in Georgia and vicinities of Ertso Lake in South Ossetia, D. pontica from the vicinities of Slavyansk-na-Kubani, Timashevsk, Novotitarovskaya Village, Poltavskaya Village and Burakovsky Village in Krasnodar Region, D. portschinskii nigrita from the vicinities of Kizilkilisa Village in Georgia, D. praticola from the vicinities of Tskhinval, Acrisheu Village, Malaya Gufta Village, Hetagurova Village and the southern slope of the Zonkar Ridge in South Ossetia. Information on an attempt to introduce D. pontica into the territory of Rostov-on-Don is provided.

Description of the genital structures of the following species of the genus Platydracus Thomson, 1858 are presented: P. stercorarius (Olivier, 1795), P fulvipes (Scopoli, 1763), P. сhalcocephalus (Fabricius, 1801), P. flavopunctatus Latreille, 1804 P. brevicornis Motschulsky, 1862 P. latebricola (Gravenhorst, 1806), P. maculosus (Erichson, 1839), P. fossator (Gravenhorst, 1802), P. ochropygus (Nordmann 1837) and P. allardi Levasseur, 1967. The morphological structures and morphogenesis of the genitalia of the genus Platydracus are discussed.

Locomotor activity of the blue mussels Mytilus edulis L. was investigated in the laboratory experiments. Movement speed of molluscs with shell length from 1 to 60 mm was analysed. Average values of motion measured during short-time intervals ranged from 8 to 12 mm/min and did not depend on the mussel size. Generally, movement of the mussels was uneven with periods of inactivity (breaks). Young mussels moved continuously with rare and short breaks. Frequency and duration of the breaks increased with mussel’s size. The rate of movement estimated for the period of experimental time changed according to the duration of breaks. Mussels with a shell length of 1–30 mm showed the maximal movement speed, while the motion activity of the bigger molluscs was significantly lower. The obtained results are discussed in view of behavioural changes in mussels during ontogenesis.

Data on the quantitative indices of occurrence, spatial-bathymetric distribution, size-weight structure of the longnose poacher Sarritor leptorhynchus (Agonidae) and composition of species accompanying in catches in the Pacific waters off the south-eastern Kamchatka and northern Kuril Islands are provided. The relationship between capture depth and the mean body weight of the longnose poacher and its diet composition are analysed. It was shown that its catch rates and occurrence in Pacific waters off the south-eastern Kamchatka and northern Kuril Islands are subjected to some interannual, seasonal, and diurnal dynamics.

The morphometric relationships among 20 geographical samples of Sorex minutissimus from the Russian part of species range were examined using geometric morphometrics and linear measurements of skull and mandible. Previous studies showed significant differences between immature and mature shrews, and no significant differences between immature males and females. In this paper we propose a method for combining small-sized samples and algorithms for checking the adequacy of such an association. We have identified two main geographical groups of shrews, “Western” and “Eastern”, distinguished by the shape of the mandible; these groups combine samples from the European (S. m. minutissimus) and East-Eurasian (S. m. barabensis, S. m. abnormis, S. m. caudata, S. m. tschuktschorum and S. m. ussurensis) parts of the range. The agreement of our results with phylogeographic data allows us to distinguish two approaches to determining the amount of intraspecific units – “phylogeographic” and “environmental”. On the basis of “environmental” approach within the Russian range we propose to consider seven subspecies: S. m. minutissimus, S. m. barabensis, S. m. stroganovi, S. m. caudata, S. m. tschuktschorum, S. m. ussurensis and S. m. tscherskii.

A new species of notothenioid fish, Pogonophryne pavlovi sp. nov., from bathyal depths of the Ross Sea, Antarctica is described. The new species belongs to the “albipinna” group of the genus Pogonophryne and is characterized by having a mental barbel with a long (61.2–65.2% of barbel length) terminal expansion composed of densely packed, brainlike convolutions.

The problem of a sable (M. zibellina) and a pine marten (M. martes) potential cohabitation in the northwest of Eastern Europe was resolved using methods of multidimensional data analysis for the first time. The skulls (n = 48) and mandibles (n = 194) of representatives of the genus Martes from ten archaeological sites of the middle Holocene (Neolithic camps) and late Holocene (Iron Age settlements) of Onega Lake region and upper reaches of Western Dvina River region were studied. 18 complete skulls and large fragments of skulls were identified morphologically as M. martes remains. 42 mandibles of M. martes from eight middle and late Holocene sites of Western Dvina River region and two mandibles of M. zibellina from two middle Holocene sites of Onega Lake region were identified using discriminant analysis methods (8 – 15 metrical characters in each model).

Type species of the genus Odontogryllus Sauss. (O. setosus Sauss.) described from a nymph is redescribed on the base of adult specimens possibly belonging to this species. Seven new species (O. proximus sp. nov., O. morona sp. nov., O. rhombicus sp. nov., O. ucayali sp. nov., O. sympatricus sp. nov., O. nimius sp. nov., and O. lacandona sp. nov.) are described from Ecuador, Peru and Mexico. Diagnostic characters and systematic position of the tribe Odontogryllini are briefly discussed.

Results of monitoring study of Ranodon sibiricus (Amphibia: Caudata: Hynobiidae) populations in the southern part of its range (Southeastern Kazakhstan) are reported in the paper. Spatial and quantitative characteristics of species distribution for 2009–2011 were compared with those obtained in the last quarter of the past century. It is suggested that the southern populations are more endangered than others. Glacier degradation due to climate warming is assumed as a key limited factor for the species in the coming decades.

Two new oribatid mite species, Epidamaeus parayunnanensis sp. nov. and Taiwanoppia (Taiwanoppia) paraflagellifera sp. nov. (Acari: Oribatida) are described from India. In India, the genera Epidamaeus and Taiwanoppia are recorded for the first time. An identification key to all known species of the nominative subgenus Taiwanoppia is provided.

Three new species of the genus Hilara are described from the Northwest Caucasus: H. arkhyziensis sp. nov., H. caucasica sp. nov., H. pseguashae sp. nov. COI barcodes are provided for the latter two species.

This paper presents a description of a new material of Adocus bostobensis, a poorly known adocid turtle from the Late Cretaceous of Kazakhstan. The new material of A. bostobensis comes from the Bostobe Formation (Santonian – early Campanian) of Shakh-Shakh II locality (northeastern Aral Sea area, Kazakhstan) and includes about 30 shell fragments and several elements of non-shell postcrania, potentially, from a single individual. This material allows us to reveal some previously unknown characters of Adocus bostobensis, improve its diagnosis and include this species in a phylogenetic analysis of Adocusia (Adocidae + Nanhsiungchelyidae) for the first time. The phylogenetic analysis places A. bostobensis in a clade with A. aksary and A. amtgai (both from the Late Cretaceous of Asia). Finally, our study demonstrates presence of at least two different lineages of Adocus (A. bostobensis and A. foveatus) in the Santonian of Western Asia.

For the first time the hair microstructure of two individuals of the bison Bison priscus found at the North East of Russia (the basins of Tirekhtyakh River in Yakutia and Rauchua River in Chukotka) have been studied using SEM in comparison with the hair structure of the recent representatives of genus Bison. The high thermo-insulating and hard-wearing characteristics of pelage and of different categories of the hair – guard hairs and wooly hairs – were revealed. The hair microstructure of ancient bison is similar to those of recent bison B. bison and aurochs B. bonasus but its pelage is more differentiated and thick.

The upper molar of a small ursid from Priozernoe locality in Moldova (the Trans-Dniester Region) incorporating remains of Moldavian faunal vertebrate complex (MN15) was found to belong to the Asian black bear (Ursus thibetanus). This discovery ascertains the time of first appearance of U. thibetanus in Europe in the early Pliocene.

Three new species of the feather mite family Pterodectinae (Protophyllodidae) are described from passerines captured in the Cat Tien National Park (Vietnam): Montesauria malacopteroni sp. nov. from Malacopteron cinereum Eyton (Pellorneidae), Proterothrix cyornis sp. nov. from Cyornis tickelliae Blyth (Muscicapidae), and P. hypothymis sp. nov. from Hypothymis azurea Boddaert (Monarchidae). Brief comments on systematics of the pterodectine genera Montesauria Oudemans, 1905 and Proterothrix Gaud, 1968 are given.

Four new species and one new genus: Leptochiton shapovalovi Sirenko sp. nov., L. liapini Sirenko sp. nov., L. dellangelloi Sirenko sp. nov., and Hoarechiton guzhovi Sirenko gen. nov. et sp. nov. are described from the Middle Russian Sea (Jurassic). L. shapovalovi is probably a xylophagous chiton because this species was found along with pieces of sunken wood and has an aesthete group very similar to some recent xylophagous species. It speculated that the genus Leptochiton was formed under rather temperate or cold conditions in the end of the Paleozoic.

Four new species of the genus Empis belonging to the nominative subgenus are described from the Northwest Caucasus: E. (E.) arkhyziensis sp. nov. (Russia: Karachay-Cherkessia), E. (E.) cherkessica sp. nov. (Russia: Karachay-Cherkessia), E. (E.) temryukiensis sp. nov. (Russia: Krasnodarskiy Territory), and E. (E.) xanthopoda sp. nov. (Russia: Krasnodarskiy Territory). Empis (E.) albopilosa de Meijere, 1935 is recorded from the Caucasus for the first time.

The results of statistical analysis of morphological characters and the study of the cytochrome b gene variability (mtDNA) of rock lizards of the Darevskia (saxicola) complex distributed in the Caucasus and the Crimea, have shown significant differentiation within this complex, assuming specific independence of D. szczerbaki (Lukina, 1963) and subspecific status of recently described D. brauneri myusserica Doronin, 2011. According to our data, within D. brauneri (Mehely, 1909) only two forms of subspecific status can be distinguished, namely D. b. brauneri and D. b. myusserica. In our opinion, Lacerta saxicola darevskii Szczerbak, 1962 (= D. brauneri darevskii) should be considered to be a junior synonym of D. b. brauneri. The possible scenarios of range formation in the Darevskia (saxicola) complex are discussed.

Eurytemora caspica sp. nov. (Crustacea: Temoridae) from the North part of the Caspian Sea and the delta of the Volga River is described. The new species belongs to the Asian clade of the Eurytemora affinis complex. This species is intermediate morphologically between the North-American and European species of the E. affinis- complex. E. caspica sp. nov. differs from nearest congeners by the shape of the mandible, absence of the seta segmentation on the caudal rami and on swimming legs, the shape of the genital double-somite and by the armament of the thoracic legs. E. caspica sp. nov. is possibly widely distributed along the North part of the Caspian Sea, as well as in the lower reaches of rivers flowing into this sea. The Japan fresh lakes are probably inhabited by undescribed species, very close morphologically to E. caspica sp. nov.

Ultrastructural organization of the foregut and intestinal caeca was investigated in the developing and mature metacercariae of Diplostomum pseudospathaceum. The features of metamorphosis were found in the development of the metacercarial caeca and oral sucker. It is suggested that metamorphic character of the development of the metacercarial gut may be related to peculiarities of the structural specialization of the free living larvae. In D. pseudospathaceum metacercariae, the functions of the gut are discussed.

In the subfamily Tryphoninae, a new species Phytodietus (Neuchorus) enriquei sp. nov. is described from Tamaulipas Province of Mexico. The new species differs from all other Neotropic and Nearctic species of the genus by the combination of white postmedian ring on antennae and pale fulvous metasoma.