Brachiopods from the Cisuralian–Guadalupian of Darvaz,Tajikistan and implications for Permian stratigraphic correlations

In this paper, we describe the upper Cisuralian Safetdara and Gundara formations of the Darvaz mountains, North Pamir, which were part of the Kunlun Arc, developed along the active Eurasian margin. The Safetdara Formation comprises massive limestones (mainly cyanobacterial, Tubiphytes and Archaeolithoporella boundstones) alternating with well-bedded bioclastic and oncoidal limestones and an interval of recessive shales. The formation crops out above the Chelamchi Formation consisting of turbiditic siltstones and sandstones with bioclastic silty limestones yielding massive limestone olistoliths. The Gundara Formation consists of fine sandstones at the base, followed by well-bedded marly bioclastic, oncoidal and microbial limestones, bearing a rich silicified brachiopod fauna in life-position. Two new taxa have been identified in this association: the cemented coralliform Gundaria insolita n. gen. n. sp. and the pedicle attached Hemileurus politus n. sp. The inferred environmental setting is that of shoal deposits of warm, shallow, high energy, clear marine waters for the Safetdara Formation. The agglutinated microbial reefs to cluster reefs of the Gundara Formation were probably growing in a muddier, quieter and probably slightly deeper setting.The foraminifers of the Brevaxina Zone suggest a Bolorian age for the top of the Chelamchi Formation, the Safetdara Formation and the base of the Gundara Formation. Kungurian conodonts have been found in the lower part of the Safetdara Formation. The biostratigraphic data from the sedimentary succession of North Pamir, integrated with those already obtained from Southeast Pamir, allow to refine the correlations between the Tethyan regional scale and the International Time Scale. In particular, it seems now clear that the Bolorian and the lower part of the Kubergandian correlate to the Kungurian.     

Enigmatic dinosaur trackways from the lower jurassic (Pliensbachian) of the Sarca valley,Northeast Italy

Four large limestone blocks with dinosaur footprints have been discovered in a large landslide situated in the Sarca Valley (northeast Italy). The trampled sediments have been ascribed to the upper part (Pliensbachian) of the Lower Jurassic Calcari Grigi Formation, which in the Sarca Valley consists of oolitic‐bioclastic limestones. These carbonates were originally deposited on the western margin of the Trento Platform, a paleogeographic structural high that was located in the northern tropical belt at the westernmost end of Tethys. Several footprints form trackways of quadrupedal dinosaurs. The data suggest the presence, within the ichnoassociation, of possible prosauropods and basal Thyrephora. This inference is consistent with the chronological and geographical distribution of both prosauropods and Thyreophora.     

A new early Frasnian (Late Devonian) foraminifer from eastern Taurides (Turkey): Evolutionary and paleobiogeographic implications

A complete Devonian sequence is well exposed in the eastern Taurides, forming more than 1000 m-thick succession of carbonate and siliciclastic sediments. The carbonate succession, stratigraphically ranging from Middle Devonian to early Late Devonian and mostly comprising limestones, dolomitic limestones and reefal limestones, contains abundant and diverse assemblages of foraminifers, corals, stromatoporoids, calcareous algae, bivalves, brachiopods, ostracods, and conodonts. The limestone samples collected from a more closely sampled stratigraphic section have been investigated for their foraminiferal content. The micropalaeontological analyses carried out on these samples have revealed the presence of an early Frasnian foraminiferal assemblage including predominantly unilocular parathuramminid species and multilocular forms of the genera Nanicella, Paratikhinella and Semitextularia? and further indicated the presence of a new genus and a new species Halevikia deveciae n. gen. n. sp. which appears as an important phylogenetic and stratigraphic transitional taxon between the families Baituganellidae n. fam. and Tournayellinidae, the phylogenetic potentiality of which during the Late Devonian is currently probably underestimated.     

Mise en evidence de Campanien continental (facies provencal) a Salavas (Ardeche,S-E de la France): Description,sedimentologie, paleogeographie

The continental sediments in the Northern Alès Basin are dated as Campanian age based on gastropod Melania praelonga MATHERON. The lithology is made up by red shales, conglomerates and pedologic limestones (crusts and nodules) with microcodiums. The conglomerates are heterometric and made up by limestone (Urgonian facies) and standstone boulders of Cenomanian and Turonian age. Also sandy limestones of coniacian age are found, all belonging to the local substratum. This discovery helps to understand the evolution of the Upper Cretaceous paleogeography.     

Comparative karyotype analysis of populations in the Alstroemeria presliana Herbert (Alstroemeriaceae) complex in Chile

Alstroemeria L., one of the most diverse genera of the Chilean flora and of high floricultural value, is represented by 35 species, most of them distributed between 28–38° S in the Mediterranean zone of Central Chile. There are 24 complex-forming taxa, of which 18 have conservation problems (8 are considered “endangered” and 10 as “vulnerable”). One of these complexes is Alstroemeria presliana Herb. with two subspecies: subsp. presliana and subsp. australis Bayer. Alstroemeria presliana grows in Chile and Argentina: subsp. presliana is distributed from Reserva Nacional Siete Tazas (35°27′ S, Region of Maule) to Antuco, (37°25′ S, Region of Bío-Bío), and is also found in Neuquén, Argentina; subsp. australis is endemic to the Cordillera of Nahuelbuta. A comparative karyotype study was carried out among six populations of A. presliana subsp. presliana and five populations of A. presliana subsp. australis. The eleven populations presented an asymmetric karyotype, with 2n = 2× = 16 chromosomes but with different karyotype formulae. A. presliana subsp. presliana shows the haploid formula 2m + 2m-sat + 1sm-sat + 1st-sat + 1t + 1 t-sat, and A. preslianasubsp. australis presents a formula 1m + 2m-sat + 1sm + 2t + 2t-sat chromosomes. The architecture of the karyotype between the subspecies is very different. The scatter plot among CV_CL vs. M_CA shows different groupings between populations of the two subspecies. According to the results obtained it is possible to consider raising Alstroemeria presliana subsp. australis at species level.     

Environmental change in the Early Permian of NE Svalbard: from a warm-water carbonate platform (Gipshuken Formation) to a temperate,mixed siliciclastic-carbonate ramp (Kapp Starostin Formation)

A detailed facies study of Early Permian strata within NE Svalbard reveals a fundamental change of the depositional setting, from a restricted-marine, warm-water carbonate platform to an open-marine, temperate-water, mixed siliciclastic-carbonate ramp. The uppermost strata of the Gipshuken Formation (Templet and Sørfonna members; Sakmarian–early Artinskian?) consist of microbialites (algal mats), mudstones, bioclastic/peloidal limestones, carbonate breccias and Microcodium facies reflecting peritidal platform areas and supratidal sabkhas. A mixed heterozoan/reduced photozoan assemblage indicates temperate-water conditions within neighboring deeper, open-marine mid-platform areas, while warm-water conditions still prevailed within inner platform zones. In contrast, the lowermost strata of the overlying Kapp Starostin Formation (Vøringen Member; late Artinskian?–Kungurian) show a fully heterozoan biotic assemblage reflecting temperate water conditions within open-marine, storm-dominated, nearshore to transitional offshore areas of a mixed carbonate-siliciclastic ramp. The Vøringen Member comprises three facies associations, which form a shallowing-upward sequence subsequent to an initial transgression. The sediments reflect bryozoan bioherms in most distal areas, followed by stacked tempestites of sandy brachiopodal shell banks and Skolithos piperocks, grading into broad sand flats in most proximal areas of the inner ramp. The above environmental change is regarded as a regional event taken place across the entire shelf along the northern margin of Pangea and is attributed to paleoclimatic, paleoceanographic, as well as paleogeographic changes, possibly related to the overall northwards drift of the supercontinent. An abrupt increase in terrigenous input coinciding with this change is ascribed to the uplift of a new local source area, probably to the north or east of the investigation area.     

Late Triassic sedimentary evolution of Slovenian Basin (eastern Southern Alps): description and correlation of the Slatnik Formation

Successions of the Slovenian Basin structurally belong to the easternmost Southern Alps. During the Late Triassic, they were part of the Adriatic continental margin. Norian–Rhaetian successions of the Slovenian Basin are characterized mainly by dolomite of the Bača Dolomite Formation. However, in the northern part of the basin, Late Triassic limestone is preserved above Bača Dolomite Formation and is formalized as the Slatnik Formation. It is composed of hemipelagic limestone alternating with resedimented limestones. The succession documents an upward progradation of the slope environment composed of three high-frequency cycles. Most prominent progradation is referred to the second, i.e., Early Rhaetian cycle. The Slatnik Formation ends with thin-bedded hemipelagic limestone that records the end-Triassic productivity crisis, or rapid sea-level fall. The overlying resedimented limestones of the Early Jurassic Krikov Formation, document the recovery of production and shedding from the adjacent carbonate platform.     

The Middle Pleistocene Al Mahrúqah Formation in the Murzuq Basin,northern Sahara,Libya evidence for orbitally-forced humid episodes during the last 500,000 years

Geological and geomorphological work in the endorheic Murzuq Basin (Fezzan) in the northern Sahara Desert, SW Libya, has revealed that the Middle Pleistocene Al Mahrúqah Formation consists of at least four limestone beds at four distinct elevations. The corresponding limestone beds are named (from oldest to youngest) the Antalkhatá Member, the Brak Member, the Bi'r az Zallaf Member, and the Aqar Member. The four members are interpreted as lacustrine deposits, and they are well distinguished by their spatial extent and lithology. Twenty-five radiometric and thermal ionisation mass spectrometric (TIMS) ²³⁰Th/U ages ranging from about 490–130 ka have been determined on these lacustrine limestones. According to these radiometric ²³⁰Th/U ages, the accumulation of the four limestone members of the Al Mahrúqah Formation followed the orbitally-forced (Milankovitch) eccentricity cycle of 100 ka during at least the last 400 ka. The four members formed in humid episodes of several tens of thousands of years' duration, when large lakes might have existed. The δ¹³C and δ¹⁸O values from samples of the four members of the Al Mahrúqah Formation provide independent genetic evidence on the lacustrine origin of the deposits in the Murzuq Basin. The spatial extent of the limestone deposits, and thus the intensity of the humid episodes, decreases at least from the Brak Member to the Aqar Member from about 52,000 km² to 1400 km², respectively. In spite of the limited precision and reliability of the new ²³⁰Th/U ages, it seems possible that short humid episodes interrupted the intervening arid episodes. Other long-term palaeohydrological records from northern and southern Africa as well as the Mediterranean region support the palaeoclimatic results from the Murzuq Basin.     

Elephas reckiDietrich,, 1915 (Proboscidea,Elephantidae)). Évolution au cours du Plio-pléistocène en Afrique orientale

A revision of the presently available materialrefered to the species Elephas reckiDietrich.. 1915, collected in East african plio-pleistocene levels, leads up to distinguish a succession of five subspecies within the species: E. r. brumpti n. subsp.; E. r. shungurensis n. subsp.; E. r. atavusArambourg 1947; E. r. ileretensis n. subsp. and E. r. reckiDietrich,, 1915.The five subspecies diagnosis are proposed here. They together represent a monophyletic lineage, the study of which allows to propose new interpretations regarding the history and evolutionary trends of the genus in Africa.     

Coral-rudist bioconstructions in the Upper Cretaceous Haidach section (Gosau Group; Northern Calcareous Alps, Austria)

Summary In the area of Haidach (Northern Calcareous Alps, Austria), coral-rudist mounds, rudist biostromes, and bioclastic limestones and marls constitute an Upper Cretaceous shelf succession approximately 100 meters thick. The succession is part of the mixed siliciclasticcarbonate Gosau Group that was deposited at the northern margin of the Austroalpine microplate. In its lower part, the carbonate succession at Haidach comprises two stratal packages that each consists, from bottom to top, of a coral-rudist mound capped by a rudist biostrome which, in turn, is overlain by bioclastic limestones and, locally, marls. The coral-rudist mounds consist mainly of floatstones. The coral assemblage is dominated by Fungiina, Astreoina, Heterocoeniina andAgathelia asperella (stylinina). From the rudists, elevators (Vaccinites spp., radiolitids) and recumbents (Plagioptychus) are present. Calcareous sponges, sclerosponges, and octocorals are subordinate. The elevator rudists commonly are small; they settled on branched corals, coral heads, on rudists, and on biolastic debris. The rudists, in turn, provided settlement sites for corals. Predominantly plocoid and thamnasteroid coral growth forms indicate soft substrata and high sedimentation rates. The mounds were episodically smothered by carbonate mud. Many corals and rudists are coated by thick and diverse encrustations that indicate high nutrient level and/or turbid waters. The coral-rudist mounds are capped byVaccinites biostromes up to 5 m thick. The establishment of these biostromes may result from unfavourable environmental conditions for corals, coupled with the potential of the elevator rudists for effective substrate colonization. TheVaccinites biostromes are locally topped by a thin radiolitid biostrome. The biostromes, in turn, are overlain by bioclastic limestones; these are arranged in stratal packages that were deposited from carbonate sand bodies. Approximately midsection, an interval of marls with abundantPhelopteria is present. These marls were deposited in a quiet lagoonal area where meadows of sea grass or algae, coupled with an elevated nutrient level, triggered the mass occurrence ofPhelopteria. The upper part of the Haidach section consists of stratal packages that each is composed of a rudist biostrome overlain by bioclastic wackestones to packstones with diverse smaller benthic foraminifera and calcareous green algae. The biostromes are either built by radiolitids,Vaccinites, andPleurocora, or consist exclusively of radiolitids (mainlyRadiolites). Both the biostromes and the bioclastic limestones were deposited in a low-energy lagoonal environment that was punctuated by high-energy events.In situ-rudist fabrics typically have a matrix of mudstone to rudistclastic wackestone; other biogens (incl. smaller benthic foraminifera) are absent or very rare. The matrix of rudist fabrics that indicate episodic destruction by high-energy events contain a fossil assemblage similar to the vertically associated bioclastic limestones. Substrata colonized by rudists thus were unfavourable at least for smaller benthic foraminifera. The described succession was deposited on a gently inclined shelf segment, where coral-rudist mounds and hippuritid biostromes were separated by a belt of bioclastic sand bodies from a lagoon with radiolitid biostromes. The mounds document that corals and Late Cretaceous elevator rudists may co-occur in close association. On the scale of the entire succession, however, mainly as a result of the wide ecologic range of the rudists relative to corals, the coral-dominated mounds and the rudist biostromes are vertically separated.     

La transgression thanétienne (Paléocène supérieur) dans l’Aurès occidental (Algérie), d’après les associations de Foraminifères de la coupe d’El Kantara

In the W-Aures (Algeria), the El Kantara pass displays about 50 m of Red Marls overlying the Maastrichtian limestones with Laffitteina. The Red Marls are, in their turn, overlain by the Thanetian marly limestones of the river El Haï. The micropaleontological inventory of these Red Marls establishes the co-occurrence of subaerian (Microcodium = Paronipora), fresh-water (Charophyta) and marine microfossils (Foraminifers). The stratigraphical significance of these microfossils is discussed. The upper part of the Red Marls, yielding abundant Valvulina and scarce Glomalveolina, is of Thanetian age. The age of the lower part is less established, but the occurrence of Microcodium suggests a Thanetian age too. Located at the margin of the northern opening of the trans-Saharan epeiric sea, the El Kantara section establishes a Thanetian age for the beginning of the Paleocene transgression. Danian deposits are missing, in spite of the absence of an obvious hiatus on the field. After general emersion during the early Paleocene, the Thanetian transgression starts with the set up of lakes at the depositional area of the Red Marls. Marine influences, limited at first, become gradually prevailing, and end with the deposition of neritic marly limestones of river El Haï.     

Oldest scleractinian coral reefs on the North American craton: Upper Triassic (Carnian), northeastern British Columbia,Canada

Bioclastic accumulations composed of crinoids, brachiopods, molluscs, spongiomorphs and scleractinian corals occur within Upper Triassic strata of the lower Baldonnel Formation at Pardonet Hill in northeastern British Columbia Canada. These small buildups (∼100 to 500 m³) have planar bases and broadly convex tops. These mounds are interpreted as small patch reefs composed of packstone, bioclastic floatstone/rudstone and carbonate breccia intercalated with mixed siliciclastic carbonate sediments deposited in a shallow subtidal setting (i.e. above fairweather wave base). Amalgamated hummocky cross-stratified to current ripple-laminated, quartz-dominated sandstone beds and numerous sharp-based, normally graded bioclastic (commonly encrinitic) packstone/grainstone — quartz–sandstone couplets characterize inter-reef lithologies.Conodont biostratigraphy indicates that the Pardonet Hill patch reefs occur within strata dated as earliest Upper Carnian (lower nodosus zone). The Pardonet Hill patch reefs originated and developed during an interval of regional sea level lowstand. Strata within which these patch reefs occur represent the westernmost migration of the Triassic shoreline in western Canada. Disappearance of coral reefs in the study area may have been affected by rapid marine transgression and failure of reef faunas to recolonize the new shore zone further to the east.The Pardonet Hill locality occurred on the western margin of the North American craton during the Triassic. Prior to their discovery reef-like structures dominated by corals in the western Panthalassa were limited to allochthonous terranes (now part of the Cordillera). The Pardonet Hill patch reefs occur at approximately 30° Triassic paleolatitude. In modern settings, this is at the extreme latitudinal margin of subtropical zooxanthellate reef development. The presence of benthic faunas characteristic of low-paleolatitude settings on the northwestern coast of Pangea has significant implications in paleotectonic and paleoenvironmental reconstructions.     

塔里木板块古城4井中奥陶统一间房组的海进序列

塔里木板块中央隆起区奥陶系碳酸盐岩台地东南缘的古城4井钻遇约130m厚的中奥陶统达瑞威尔阶一间房组灰岩。该组下部为表附藻-肾形钙藻-瓶筐石礁丘群落,间夹有薄层砂屑滩,海水深度推测位于10-30m的潮下带至正常波浪带之间;上部为中-低能带藻屑砂屑组合,灰泥含量增高,大致指示正常波浪带之下到最大浪基面之间30-60m的深度。古城4井与中央隆起区西北部巴楚露头区同期沉积均显示海水逐渐加深的过程,到上奥陶统桑比阶吐木休克组沉积期两地都达到了奥陶系的最大海泛面。     

Paleobiogeographic significance of a new ozawainelloid fusulinid Pseudoacutella partoazari n. sp., from the Asselian (lowermost Permian) of Gaduk (Central Alborz,Iran)

The uppermost Carboniferous–Lower Permian Dorud Group that crops out at the Gaduk section of Central Alborz is over 44 m thick and includes thick sequences of conglomerate, quartzarenite, calcareous sandstone, oncolitic fusulinid limestone, sandy limestone, sandstone and shale. The Toyeh, Emarat and Shah Zeid formations of this Group were dated as latest Gzhelian to Early Sakmarian. A review of the Asselian fusulinids and smaller foraminifers of Iran emphasizes (1) how the foraminiferal assemblages of Alborz and central Iran differ; (2) why non-Pseudofusulina cannot be a nomen substitutum to Pseudofusulina (even if this latter was often misinterpreted in the Tethyan regions). A new smaller fusulinid Pseudoacutella partoazari n. sp. is described from the Emarat Formation (Asselian) that crops out in the Gaduk area. The palaeobiogeographic significance of Pseudoacutella is discussed, because this genus, which was cosmopolitan from Late Bashkirian to Moscovian, became rare in the Upper Pennsylvanian of the USA and the Carnic Alps, and then, very rare in Lower Permian of Texas, Arizona and Iran.     

Palaeoenvironmental changes in the Pliocene Intra-Apenninic Basin,near Bologna (Northern Italy)

Combined palaeoecologic and sedimentologic studies reveal remarkable palaeoenvironmental changes in the Pliocene Intra-Apenninic Basin (PIB) of the Bologna Apennines (Northern Italy). At the northern margin of PIB, the stratigraphic succession is approximately 1000 m thick, and displays an overall coarsening-upward trend, including a lower mudstone-dominated succession (Argille Azzurre Formation) of Zanclean age, overlain by a sandstone-rich unit (M. Adone Formation), assigned to the Zanclean-Piacenzian. The vertical succession of biofacies within the Argille Azzurre Fm. indicates an obvious shallowing-upward tendency, from slope/outer-shelf (150–300 m water depths) to inner-shelf (30–50 m) environments. A vertical cyclic pattern of facies, with alternating sandy siltstones and sandstone bodies, characterizes M. Adone Formation. Each elementary cycle, about 30–50 m thick, includes a couple of siltstone-dominated inner-shelf and prodelta deposits, passing upwards into progradational delta front, tabular sand bodies. These high-rank cycles, spanning an interval of time in the order of magnitude of 40 ky, are separated by flooding surfaces, clearly recognizable on the basis of palaeoecologic analysis, and reflect an aggradational stacking pattern of parasequences superposed to the general shallowing-upward trend.     

Species of the genus Nikanoria related to Nikanoria mongolica (Hymenoptera, Eurytomidae)

A complex of the species related to Nikanoria mongolica Zerova is discussed. This complex includes a new subspecies (N. mongolica punctulata subsp. n.) and two new species (N. kazakhstanica sp. n. and N. sugonjaevi sp. n.). Data on the distribution and trophic associations of these species are provided. The type material is deposited in the collection of the Schmalhauzen Institute of Zoology, National Ukrainian Academy of Sciences, Kiev.     

A taxonomic revision of the Antennaria rosea (Asteraceae: Inuleae: Gnaphaliinae) polyploid complex

TheAntennaria rosea species complex is circumscribed to contain four subspecific taxa. A complete synonymy and key to the subspecies ofA. rosea is presented. The following new combinations are proposed:Antennaria rosea subsp.arida (E. Nels.) R. Bayer,A. rosea subsp. confinis (Greene) R. Bayer, andA. rosea subsp. pulvinata (Greene) R. Bayer. The affinities ofA. rosea to other species ofAntennaria are discussed and a key to separateA. rosea from related polyploid complexes is provided.     

Assessment of Daucus carota L. (Apiaceae) subspecies by chemotaxonomic and DNA content analyses

For a clearer distinction between the four subspecies of Daucus carota native from Portugal (subsp. carota, subsp. maximus, subsp. gummifer and subsp. halophilus), morphological features of the fruits, DNA content analyses by flow cytometry, and chemical characterization of the essential oils were undertaken.We found chemotaxonomic evidences to consider D. carota subsp. maximus as a separate species rather than a subspecies of D. carota. This separation is based on the morphometric analysis of the fruits and in the high levels of asarone present only in the essential oil of the subsp. maximus. The remaining subspecies are difficult to distinguish from each other based on the morphology of the fruits and in DNA content. However, based on the essential oils, it was possible to distinguish the subspecies halophilus from the other two (subsp. gummifer and subsp. carota) because of its high content of elemicin, with the other two having high levels of geranyl acetate.Based on these results, the subspecies maximus is proposed as a different species (Daucus maximus Desf.) and the taxonomic status of other three subspecies (subsp. carota, subsp. gummifer and subsp. halophilus) is maintained. Still, the latter three taxa need to be further studied for a more precise taxonomic characterization.     

Cardiolinka bohemica (Barrande, 1881) – A first representative of the Late Silurian Bohemian type Bivalvia fauna from the northern Arabian Plate,Southeast Turkey

The Rheic Ocean was a major oceanic domain between Baltica, Laurentia, Perunica and Gondwana in Ordovician-Silurian times. The cosmopolitan nepiomorphian bivalves Praecardioidei Newell, 1965 and Antipleuroidei Kříž, 2007 are characteristic of the Silurian of Perunica, peri-Gondwana, and Baltica, and occur also in Laurentia and Siberia. The Bohemian-type bivalve Cardiolinka Kříž, 1981 (Nepiomorphia Kříž, 2007, Cardiolidae Hoernes, 1884), from the Late Silurian of the Bahar-1 well core, has been found for the first time in southeastern Turkey. The strata containing the species Cardiolinka bohemica (Barrande, 1881) occur in the middle part of the Dadaş Formation in the interior Petroleum District X-Siirt of the northern parts of the Arabian Plate. The cosmopolitan species C. bohemica was until now known from the Latest Ludlow to Pridoli of the Prague Basin, France, Carnic Alps, Sardinia, East European Platform (Poland), eastern Serbia, Moesian Platform, and Arctic Canada. The new surprising subsurface data on C. bohemica in Diyarbakır-Bismil area (southeastern Turkey) therefore represent another piece of evidence in favour of strong faunistic affinity between Perunica, peri-Gondwanan Europe and the northern Gondwana margin.     

Ranges and cold hardiness of two earthworm subspecies (Eisenia nordenskioldi, Lumbricidae, Oligochaeta)

The ranges of two earthworm subspecies, Eisenia nordenskioldi nordenskioldi (Eisen 1879) and E. n. pallida Malevi 1956, differ in area and partially overlap. E. n. nordenskioldi populates the entire Asian Russia and eastern regions of the Russian Plain, from the lower reaches of the Volga and Don rivers to the Arctic Ocean coasts, while E. n. pallida has not expanded to the Asian permafrost zone and does not occur in European Russia and in the Urals. These subspecies “hold the record” in cold hardiness: the worms and cocoons of the nominotypical subspecies withstand temperatures down to ?34 and ?40°C, and those of E. n. pallida, to ?28 and ?23°C, respectively. Hence, their distribution is independent of subzero temperatures, and their ability to overwinter at any phase of the life cycle makes them also independent of heat supply during the summer period. Differences in geographic range may also be due to biological features of the subspecies. The nominotypical subspecies feeds belongs to the epiendogeic morphoecological type (feeding on the ground surface), whereas E. n. pallida is a true endogeic earthworm. Both subspecies have similar requirements for soil acidity; however, conditions in coarse-humus organomineral horizons of frozen soils appear to be unfavorable for E. n. pallida, which accounts for the absence of this subspecies in the permafrost zone.