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European glacial relict snails and plants: environmental context of theirmodern refugial occurrence in southern Siberia

MICHAL HORS�AK, MILAN CHYTR�Y, PETRA H�AJKOV�A, MICHAL H�AJEK, JI�R�I DANIHELKA, VERONIKAHORS�AKOV�A, NIKOLAI ERMAKOV, DMITRY A. GERMAN, MARTIN KO�C�I, PAVEL LUSTYK, JEFFREY C. NEKOLA,ZDENKA PREISLEROV�A AND MILAN VALACHOVI�C

Hors�ak, M., Chytr�y, M., H�ajkov�a, P., H�ajek, M., Danihelka, J., Hors�akov�a, V., Ermakov, N., German, D. A., Ko�c�ı,M., Lustyk, P., Nekola, J. C., Preislerov�a, Z. & Valachovi�c, M. 2015 (October): European glacial relict snails andplants: environmental context of their modern refugial occurrence in southern Siberia. Boreas, Vol. 44, pp. 638–657.10.1111/bor.12133. ISSN 0300-9483.

Knowledge of present-day communities and ecosystems resembling those reconstructed from the fossil record canhelp improve our understanding of historical distribution patterns and species composition of past communities.Here, we use a unique data set of 570 plots explored for vascular plant and 315 for land-snail assemblages locatedalong a 650-km-long transect running across a steep climatic gradient in the Russian Altai Mountains and theirfoothills in southern Siberia. We analysed climatic and habitat requirements of modern populations for eightland-snail and 16 vascular plant species that are considered characteristic of the full-glacial environment of centralEurope based on (i) fossil evidence from loess deposits (snails) or (ii) refugial patterns of their modern distribu-tions (plants). The analysis yielded consistent predictions of the full-glacial central European climate derived fromboth snail and plant populations. We found that the distribution of these 24 species was limited to the areas withmean annual temperature varying from �6.7 to 3.4 °C (median �2.5 °C) and with total annual precipitation vary-ing from 137 to 593 mm (median 283 mm). In both groups there were species limited to areas with colder anddrier macroclimates (e.g. snails Columella columella and Pupilla loessica, and plants Kobresia myosuroides andKrascheninnikovia ceratoides), whereas other species preferred areas with relatively warmer and/or moistermacroclimates (e.g. snails Pupilla turcmenica and P. alpicola, and plants Artemisia laciniata and Carex capillaris).Analysis of climatic conditions also indicated that distributional shifts of the studied species during the Pleisto-cene/Holocene transition were closely related to their climatic tolerances. Our results suggest that the habitatrequirements of southern Siberian populations can provide realistic insights into the reconstruction of Eurasian,especially central European, glacial environments. Data obtained from modern populations also highlight theimportance of wet habitats as refugia in the generally dry full-glacial landscape.

Michal Hors�ak ([email protected]), Milan Chytr�y, Petra H�ajkov�a, Michal H�ajek, Ji�r�ı Danihelka, VeronikaHors�akov�a, Martin Ko�c�ı, Pavel Lustyk and Zdenka Preislerov�a, Department of Botany and Zoology, MasarykUniversity, Kotl�a�rsk�a 2, CZ-611 37 Brno, Czech Republic; Petra H�ajkov�a, Ji�r�ı Danihelka, Department of VegetationEcology, Institute of Botany, Czech Academy of Sciences, Lidick�a 25/27, CZ-602 00, Brno, Czech Republic; NikolaiErmakov, Central Siberian Botanical Garden, Russian Academy of Sciences, Zolotodolinskaya 101, Novosibirsk630090, Russia; Dmitry A. German, South-Siberian Botanical Garden, Altai State University, Lenina 61, 656049Barnaul, Russia; Jeffrey C. Nekola, Biology Department, University of New Mexico, Albuquerque, NM 87131,USA; Milan Valachovi�c, Institute of Botany, Slovak Academy of Sciences, D�ubravsk�a cesta 9, SK-845 23 Bratislava,Slovakia; received 30th March 2015, accepted 3rd June 2015.

Historical distribution patterns and species composi-tions of past communities are usually assessed basedon direct fossil evidence from the target area and a par-ticular time period, but these assessments can beimproved by knowledge of modern analogues, i.e. pres-ent-day communities and ecosystems resembling thoseknown from the fossil record (e.g. Jackson & Williams2004). Modern analogues never provide a perfectmatch with past environments, but enable an examina-tion of real ecological processes across multiple habi-tats, which may be hidden in the fossil record. Theyalso help us deal with various biases existing in the fos-sil record caused by the varying capacity of the envi-ronment to preserve the fossils, mixing of multiplesediment beds and differential preservation amongsttaxa (Kidwell & Holland 2002; Jackson & Williams2004).

Recent research has provided plausible evidence thatplant and animal communities, and whole ecosystemsof the Altai-Sayan region in southern Siberia represent

possibly the closest modern analogue to those offull-glacial central Europe. Several land-snail speciestypical of central European full-glacial loess sediments,some currently extinct in Europe, have been found topossess extant populations in this part of central Eur-asia and to exhibit community assemblages that aretypical of the European full-glacial fossil record (Meng2008; Meng & Hoffmann 2009; Hors�ak et al. 2010;Hoffmann et al. 2011). Likewise, present mammaliancommunities of the Altai-Sayan region display amarked similarity to the Last Glacial mammalianfauna of the same area (Agadjanian & Serdyuk 2005),harbouring more surviving Pleistocene full-glacialmammals than any other region of the Palaearctic(Pavelkov�a �Ri�c�ankov�a et al. 2014). Comparisons of thefull- and late-glacial fossil pollen spectra from centralEurope with modern pollen assemblages of variousvegetation types from southern Siberia have also con-firmed their considerable similarity in dominant plantspecies and vegetation types (Kune�s et al. 2008;

DOI 10.1111/bor.12133 © 2015 Collegium Boreas. Published by John Wiley & Sons Ltd

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Magyari et al. 2014a; see also Jankovsk�a & Pokorn�y2008; Pel�ankov�a & Chytr�y 2009). The Altai-Sayanregion can thus be regarded as an important refugiumfor full-glacial fauna and flora, harbouring a uniquemosaic of cold steppe (e.g. snails Pupilla loessica andVallonia tenuilabris, mammals Lagurus lagurus andMarmota bobak, plants Artemisia spp. and Chenopodi-aceae) and tundra/taiga species (e.g. snails Columellacolumella and Vertigo parcedentata, mammals Lepustimidus and Microtus gregalis, and plants Betula rotun-difolia (B. nana s. lat.) and Kobresia myosuroides),occurring at the same or closeby sites (Kune�s et al.2008; Hors�ak et al. 2010; Pavelkov�a �Ri�c�ankov�a et al.2014). Therefore, data from this region can consider-ably enhance our knowledge of the structure and habi-tat heterogeneity of the European and northern AsianLast Glacial landscapes, and the autecology of glacialrelict species. As several mollusc-based transfer func-tions have been developed to reconstruct past tempera-ture and precipitation during the last glaciation(Rousseau 1991; Moine et al. 2002), the integration ofthese modern data can also help to improve the reli-ability of the reconstructions.

In this paper we examine climatic and habitat affini-ties of selected species of land snails and vascularplants in their continental refugium in the Altai Mts ofsouthern Siberia with the aim of contributing to thebiogeographical interpretation of the current isolatedoccurrences of these species in Europe, where they maybe of relict origin, and to palaeoenvironmental recon-structions. We assume that species persisting in thesouthern Siberian refugium have retained their habitatrequirements since the Last Glacial period (Hors�aket al. 2010). For land snails, relict status is directlyascertained from their known presence in the glacialloess sediments of central Europe (e.g. Lo�zek 1954,1964, 2001; S€umegi 2005; Mania 2006). For most spe-cies of vascular plants, however, relict status can onlybe indirectly inferred based on their modern distribu-tion ranges, as identifications of most herbaceousplants to the species level are difficult or impossiblebased on fossil pollen (Bennett & Willis 2001) and ma-crofossils are generally rare. Therefore, only plant spe-cies with rare isolated occurrences in Europe and amajor continuous distribution range in the continentalcentral part of Asia, or those for which a more contin-uous distribution range in Europe during the Last Gla-cial is indicated by fossil records, were considered inthis study as probable glacial relicts. We included notonly species of zonal habitats, but also wetland plantspecies.

We assessed and compared climatic and habitatrequirements of relict snails and (presumably) relictplants in parallel, using original data collected along a650-km-long transect across the Russian Altai Mts,which follows the strongest climatic gradient in thisarea. Both macroclimatic and habitat requirements of

the target relict populations are summarized anddiscussed in the broader context of central Europeanglacial-environment reconstructions. We aimed toreview the modern distribution and fossil records ofthese species and put them into the context of the cli-matic and habitat affinities of their extant relict popu-lations in southern Siberia. We believe that originalfield data on the ecology of these species that are oftenused as proxies for past environmental conditions canbe very useful for interpretation of European full-gla-cial environments. Of prime importance are those spe-cies that are today extinct or extremely rare in centralEurope, as data on their climatic and habitat affinitiesare very limited or completely missing. Such data mayalso help to understand the causes of their rarity inEurope with consequences not only for historical bio-geography, but also for biodiversity conservation.

Material and methods

Study area

In order to define the climatic ranges and habitatrequirements for modern populations of the targetEuropean full-glacial species, we sampled along an~650-km-long NNW–SSE transect across a steep cli-matic gradient running across the northern Altaifoothills and the Russian part of the Altai Mts insouthern Siberia, from the Salairskii Kryazh range inthe north to the Ukok plateau in the south (Fig. 1).The transect covered an elevational range from 192to 2924 m a.s.l. and included three main climatic andvegetation zones (Kuminova 1960; Walter 1974),which differ greatly in average elevation, temperatureand precipitation. Owing to increasing elevation,temperature sharply decreases from NNW to SSE.Whereas mean annual precipitation locally exceeds800 mm in the northern Altai foothills, it decreasessharply to the south and south-east (e.g. Polikarpovet al. 1986), resulting in very dry conditions for thesoutheastern part of the Russian Altai, where inter-montane basins receive <180 mm of annual precipita-tion (Beresneva 2006). At the scale of the wholeRussian Altai, precipitation therefore decreases withthe decrease in temperature. As a result, the climaticsettings as well as the vegetation of the NNW partof the transect resemble those of modern centralEurope except for longer winters with much lowertemperatures and several months of continuous frost,in contrast to the SSE areas, which closely match theclimate and biota reconstructed for the central Euro-pean full-glacial (Hors�ak et al. 2010).

The low mountains in the northern part of the areaare mainly forested, with Abies sibirica, Betula pendula,Picea obovata, Pinus sibirica, P. sylvestris and Populustremula being the dominant trees. The central part is aforest-steppe and forest-tundra with woodland patches

BOREAS European glacial relict snails and plants 639

dominated by Larix sibirica, Picea obovata and Pinussibirica. The southern part is predominantly an openlandscape with steppe and tundra grasslands, locallywith small patches of Larix sibirica woodland. For fur-ther details on the environment of the Russian part ofthe Altai Mts see Hors�ak et al. (2010).

Species sampling

Vascular plants were recorded from 570 quadrats of100 m2 in area that represented locally predominanthabitat types along the transect (Fig. 1). All vascularplant species were recorded in each plot. Species thatcould not be identified in the field were collected andidentified subsequently using the Flora of Siberia(Krasnoborov 1987 et seq.) and a comparative herbar-ium collection. Plant specimens are stored at the Her-barium of Masaryk University (BRNU). In general, thetaxonomy and nomenclature of vascular plants followthe Flora of Siberia as summarized by Baikov (2005).Wherever necessary, information is given on the taxo-nomic concept accepted for the purpose of this study.

Land snails were sampled from 315 of the 570 vascu-lar plant quadrats using habitat-specific sampling tech-niques that allowed for optimal encounter rates andmaximal inventory of the snail assemblage from eachplot. In non-wetland habitats snails were carefullysearched for by eye from all microhabitats. Sampling

continued until no new species were found over a 15-min period and until all species were represented by atleast three individuals. Using this rule, species-poor sites(e.g. dry loess steppes) were never sampled for <30 min,whereas at richer sites (herb-rich forest plots or calcare-ous rock outcrops) sampling generally took no morethan an hour to complete. In damper habitat types (i.e.mires, fens, wet tundra and alluvial vegetation), a 12-litre sample of the soil organic horizon including topsoil,litter, bryophytes and herbaceous vegetation was col-lected and processed in the field using the wet-sievingmethod (Hors�ak 2003). All recorded shells and liveshelled snails were collected and kept dry; samples wereidentified under a dissecting microscope in the labora-tory using the available literature (Meng 2009; Sysoev &Schileyko 2009; Hors�ak et al. 2010, 2013a) and the firstauthor’s comparative collection of Siberian snail speci-mens. All samples are stored in the collection of MichalHors�ak (Brno). Nomenclature and taxonomy generallyfollow Sysoev & Schileyko (2009), with Nekola et al.(2015) used for the genus Pupilla.

Environmental variables and data analysis

For each site with a record of any of the target rel-ict species, total annual precipitation and mean Jan-uary and July temperatures were obtained from theWorldClim database (Hijmans et al. 2005,

A B C

D

Fig. 1. Map of the study area with the position of sampling sites. Variation in numbers of recorded relict species of land snails (A) and vascu-lar plants (B) discussed in this study is expressed by different symbols. Distributions of mean January temperature (C) and total annual precip-itation (D) are shown across the study area.

640 Michal Hors�ak et al. BOREAS

www.worldclim.org) using the ArcGIS 8.3 program(www.esri.com). We chose then mean temperature ofthe coldest month as it has repeatedly emerged asthe most important thermal parameter that mayconstrain the geographical distribution of land snails(e.g. Moine et al. 2002; more details also below). Bycontrast, total annual precipitation is amongst themost important predictors for vascular plants incontinental areas (Adler & Levine 2007; Chytr�yet al. 2007; Yang et al. 2010), although their directeffect on snails is not so obvious. Because of strongcorrelations amongst the sums of annual, summerand winter precipitation (rho > 0.87), we used onlythe total annual precipitation, which appeared to bethe most informative measure of precipitation in ourprevious study of the distribution of glacial snails inthe study area (Hors�ak et al. 2010). As both taxacan also be influenced by summer temperatures, wecompleted the data by mean temperature of thewarmest month. All above mentioned climatic char-acteristics were used, along with the annual tempera-ture, to show the climate affinities of the targetspecies and to provide data for comparisons withthe estimates commonly derived from palaeoclimaticreconstructions and transfer functions.

Plots with target species present (95 plots for snails;234 for plants) were classified into 11 habitat typesaccording to vegetation structure, plant species composi-tion and abiotic environmental factors (Table 2, Fig. 2).The number of sampled sites approximately reflected thefrequency of a particular habitat type in the landscape.

The variation of individual climatic values for sitescontaining each target species was graphically pre-sented using beanplots (Kampstra 2008). Apart fromshowing each measurement, the beanplots provide aneasy visual comparison of the studied groups by show-ing the density of distribution. The overall median val-ues were calculated separately from all records ofglacial relict snail or plant species (Table 1). We usedindividual species records instead of plots to give aproportionally higher importance to plots with morespecies; e.g. climatic values of a plot with five specieswere used as five independent values. Beanplots werecreated in R (version 2.15.1, R Core Team 2012) withthe use of the ‘beanplot’ package (Kampstra 2008).

Results and discussion

In the following text and graphs (Figs 3, 4), species areordered by their temperature requirements, rangingfrom the most cold-adapted to the most warm-demanding.

Land snails

From 315 plots sampled for snails in the Russian AltaiMts (Fig. 1), we obtained 109 records of six target

land-snail species (i.e. Columella columella, Pupillaalpicola, P. loessica, Vallonia tenuilabris, Vertigo parce-dentata and V. pseudosubstriata; Table 1) that, basedon loess fossil evidence from central Europe (Lo�zek1964, 2000), characterize full-glacial environments.Thirty-six records of Pupilla turcmenica and twenty-five records of Vertigo aff. genesii were also included.Both of these taxa are morphologically and ecologi-cally similar central Asian or Beringian siblings of theEuropean Pupilla sterrii (Nekola et al. 2015) and Ver-tigo genesii (Nekola unpubl. data), respectively, whichare common and abundant in central European loessassemblages (e.g. Lo�zek 2000). Pupilla sterrii is com-mon in contemporaneous limestone steppes of centraland eastern Europe (Kerney et al. 1983), whereasVertigo genesii is an arcto-alpine European endemicknown from the UK and Scandinavia and extending towestern Russia and south into the Alps (Cameronet al. 2003).

Pupilla loessica. – Pupilla loessica is a steppe-tundraspecies that is currently restricted to cold areas of thecentral Asian mountain systems (Meng & Hoffmann2009; Hors�ak et al. 2010), with related sibling speciesoccurring in southwestern (Hokkaido) and eastern(Alaska) Beringia (Nekola et al. 2015). However, it isalso a characteristic full-glacial fossil in loess depositsacross almost the whole of temperate Europe, except inareas west of the Rhine valley (e.g. Lo�zek 1964, 2001;Rousseau 2001). It was first described based on fossilmaterial from Moravia (Czech Republic) and untilrecently was considered to be extinct (cf. Meng & Hoff-mann 2009). Lately it has been found to occur com-monly in the Tien Shan and Altai Mountains,northern Mongolia and near Lake Baikal (Meng &Hoffmann 2009; Hors�ak et al. 2010).

Its modern distribution in the Altai is limited toareas with a dry and cold climate (Figs 3, 4). In someplaces it represents the only snail species capable ofsurviving in extremely harsh environments, e.g. in theChuya basin, Ukok plateau, and adjacent mountainpasses in the southeastern part of the Russian Altai(Fig. 1). In these areas it is confined to Betula rotundi-folia tundra or to sparse Larix sibirica woodlands witha well-developed moss layer. Although it commonlyinhabits both grassland and shrub tundra, it can alsobe found in a variety of other habitats, including wet-lands (Table 2) where shells may be almost twice aslarge (Nekola et al. 2015).

Given its modern ecological preferences, it is likelythat it may have lived in various glacial habitats acrossthe European steppe-tundra zone (Lo�zek 2001; Meng& Hoffmann 2009; Hoffmann et al. 2011). In someEuropean loess deposits it represents the only snailspecies, yet occurs in high numbers. This is exactly thepattern observed in its driest habitats in the Altai. It isimportant to remember, however, that in contempora-


A B

C D

E F

G H I

J K


neous environments it also occurs in damper habitatswhere it may co-occur with species like Pupilla alpicolaand Vertigo parcedentata (Table 2); however, such wetto mesic assemblages were much less likely to be pre-served through loess deposition and are thus under-represented in the fossil record.

Vertigo parcedentata. – Vertigo parcedentata is a sub-alpine wetland species originally described on the basisof fossils from loess near Wiesbaden, Germany. It waslater discovered living at four sites near Kongsvoll inthe Dovrefjell National Park in central Norway (Pok-ryszko 1993). Very recently it was also recorded in the

Fig. 3. Mean January temperatures of sites with records of relict snail and plant species. The longest grey line refers to the median value;lengths of the shorter black lines correspond to the number of sites with the same climatic value (ranges from one to six).

Fig. 4. Total annual precipitation of sites with records of relict snail and plant species. See Fig. 3 for details.

Fig. 2. Habitats in which the studied land snail and vascular plant species occur in the Russian Altai Mountains (from left to right and top tobottom): A. Dry steppe. B. Meadow steppe. C. Steppe scrub. D. Tundra grassland. E. Tundra scrub. F. Herb-rich forest. G. Taiga forest. H.Wet grassland. I. Saline grassland. J. Open fen. K. Wet woodland. The habitat classification and the order of habitat types are as in Table 1.Photographs by P. H�ajkov�a (A–C, F, G, I, J), M. Hors�ak (D, E, H) and M. Chytr�y (K).


Altai (Meng 2008; Hors�ak et al. 2010) and Tien Shan(Meng 2008) mountains of central Asia. Based on bothmitochondrial and nuclear markers, European andcentral Asian populations have been confirmed to bepart of the same species-level clade (J.C. Nekola,unpublished data).

In the Russian Altai it is predominantly associatedwith willow scrub in riparian corridors and fens. Itsassociation with such wet and patchily distributed habi-tats may explain its relatively common occurrence, butlow numbers, in loess deposits (e.g. Lo�zek 2001): it ispossible that in Pleistocene Europe this species occurredin wet habitats adjacent to the loess zone (wet depres-sions, toe-slope seepages etc.), and only accidentlystrayed into the dry loess steppe matrix. This hypothesisis supported by its record from Lateglacial calcareousfen sediments in the intermountain basins of the Wes-tern Carpathians in Slovakia (H�ajkov�a et al. 2015a).

Columella columella. – Columella columella is a cir-cumboreal arcto-alpine species typical of willow scrub,wet tundra and taiga seeps. It is common in coldregions of Norway (Pokryszko 1993), inhabits alpinetundra grasslands in the calcareous ranges of the Alpsand Carpathians (Kerney et al. 1983; Hors�ak et al.2013a), is known from several mountain systems ofcentral Asia (e.g. Sysoev & Schileyko 2009) and occursin similar habitats throughout boreal and arctic NorthAmerica (Nekola & Coles 2010).

In the Altai refugium, most populations are associ-ated with alluvial fen woodlands and treeless fens(Table 2). Despite this clear affinity for wetland sites(see also Hoffmann et al. 2011), climate data indicatethat C. columella occurs in the driest regions, whereannual precipitation is <300 mm. Perhaps this restric-tion is related to the fact that groundwater in such

regions will typically contain higher levels of dissolvedcations.

These observations may help explain the presumeddiversity of habitat preferences for this species at thePleistocene/Holocene transition, when C. columellaappeared not only as a xerophilous species able to per-sist in loess-mantled uplands (Lo�zek 1964), but wasalso capable of establishing large populations in calcar-eous wetlands (Lo�zek 2007; H�ajkov�a et al. 2015a).With Lateglacial climate amelioration and the increasein precipitation, calcareous wetlands may have becomemore frequent and provide the principal habitat forthis species. It is also possible that shells found in loessdeposits represent temporary populations that flour-ished only during wetter periods, as also suggested bythe data from the Nussloch loess series (Moine et al.2008). This is also the only characteristic glacial speciescommonly found in the Weichselian upper pleniglacialloess deposits west of the Rhine valley, especially closeto the Channel coast (Moine 2014; O. Moine, pers.comm. 2014), i.e. in areas of high humidity. In Britainit persisted into the early Holocene (e.g. Preece &Bridgland 1999).

Vallonia tenuilabris. – Vallonia tenuilabris is typical ofsteppe-tundra, but also inhabits various habitats rang-ing from dry steppes to wet forests across central andsouthern Siberia (Gerber 1996). Its occurrence is con-centrated in the mountain systems of central Asia(Meng 2009; Hors�ak et al. 2010), but it has also beenfound on the northern Siberian plains, such as incentral Yakutia (Hors�ak et al. 2013b). It is a commonand characteristic species of central European loessassemblages (Lo�zek 1964; Meng 2009), which becameextinct in Europe at the onset of the Holocene (e.g.Hor�a�cek & Lo�zek 1988; Lo�zek & C�ılek 1995).

Table 1. Climatic descriptors of the sites with records of at least one target species of snails and plants. Values are also provided for all snailspecies first and then for all snail species except Pupilla turcmenica (see text for explanation).

Median Mean SD Min. Max.

All snailsMean January temperature (°C) �18.8 �18.4 1.56 �21.8 �14.0Mean July temperature (°C) 13.5 14.2 2.58 8.3 20.5Mean annual temperature (°C) �2.4 �1.9 2.00 �6.7 3.4Total annual precipitation (mm) 283 304 93.5 146 592

Snails except Pupilla turcmenicaMean January temperature (°C) �19.0 �18.8 1.25 �21.8 �15.8Mean July temperature (°C) 13.1 13.5 2.19 8.3 19.5Mean annual temperature (°C) �2.9 �2.4 1.70 �6.7 1.6Total annual precipitation (mm) 280 295 90.7 146 592

PlantsMean January temperature (°C) �19.1 �18.4 1.83 �21.8 �14.0Mean July temperature (°C) 13.4 14.2 2.77 8.3 20.5Mean annual temperature (°C) �2.5 �1.8 2.26 �6.7 3.4Total annual precipitation (mm) 285 301 94 137 593


Table

2.Hab

itat

preferencesof

thestud

iedfull-glaciallan

d-snail(shad

ed)an

dvascular

plan

tspecies(non

shad

ed).The

relative

percentage

ofthespeciesrecordsam

ongst11

habitattypesis

show

nforeach

specieswiththehigh

estpercentage

valuegivenin

bold.T

hespeciesareclassified

aspreferring

steppe

(i.e.m

ostlyrecorded

indryor

meado

wsteppes),tun

dra(m

ostlyrecorded

intund

ragrasslan

dor

scrub)

orwet

habitats(m

ostlyrecorded

insalin

egrasslan

d,op

enor

woo

dlan

dfens).The

lastsnailspecies

was

notclassified

asitwas

rare

andexhibitedno

distinct

habitat

preferences.Dash(–)indicatesspeciesab

sencein

theha

bitat.

Hab

itat

types

Steppe

Tun

dra

Wet

habitats

Kochiaprostrata

Agropyroncristatum

Krascheninnikoviaceratoides

Pupillaturcmenica

Alliumstrictum

Carexobtusata

Artemisialaciniata

Kobresiamyosuroides

Saxifragahirculus

Artemisiarupestris

Ligulariaglauca

Pupillaloessica

Pentaphylloidesfruticosa

Astragalusfrigidus

Trichophorumpumilum

Blysmusrufus

Carexmicroglochin

Pupillaalpicola

Valloniatenuilabris

Vertigoaff.genesii

Vertigoparcedentata

Carexcapillaris

Columellacolumella

Vertigopseudosubstriata

Dry/typ

ical

steppe

9793

8856

3524

85

–11

148

6–

––

––

9–

––

––

Meado

wsteppe

3–

–11

2222

44–

–28

218

5–

––

–5

––

––

––

Steppe

scrub

–6

1217

19–

8–

–6

75

3–

––

––

4–

––

––

Tun

dragrasslan

d–

––

319

226

3731

3329

2522

–10

–10

–13

––

25–

–Tun

drascrub

––

––

–7

326

316

720

2229

––

–11

916

2513

–33

Herb-rich

forest

–2

–11

522

173

–11

1413

521

––

–5

13–

––

––

Taiga

forest

––

––

–2

––

––

–5

229

––

–5

44

––

––

Wet

grasslan

d–

––

––

–3

815

67

39

14–

8–

169

88

1311

33Sa

linegrasslan

d–

––

3–

–6

5–

––

––

–60

5440

–4

4–

––

–Openfen

––

––

––

–16

23–

–10

11–

3031

5032

1332

25–

33–

Wet

woo

dlan

d–

––

––

–6

––

––

516

7–

8–

2622

3642

5056

33

No.

ofrecords

3854

1736

3741

3638

1318

1440

6414

1013

1019

2325

128

96


Although it was found in almost all habitat types athigh elevations in the Altai (Table 2) the largest popu-lations were recorded from wet habitats such as openand wooded fens. However, it was also observed in twodry steppes and at one steppe scrub site. The ability ofthis species to occupy environmentally diverse condi-tions may explain its common occurrence in Europeanloess (Meng 2009).

Vertigo aff. genesii. – Vertigo aff. genesii is the Berin-gian sibling to the European arctic/alpine Vertigo gene-sii that ranges from the Altai through Yakutia,Mongolia, Hokkaido and Alaska to central Manitobaand the Gulf of St Lawrence in Quebec (Nekola &Coles 2010; J.C. Nekola, unpublished data). Altaianpopulations of this species have previously been identi-fied based on conchological characters as V. genesii(Meng 2008; Hors�ak et al. 2010; Hoffmann et al.2011), which is otherwise known from calcareous partsof the Scandinavian mountains with isolated popula-tions extending to the British Isles and the Alps(Schenkov�a & Hors�ak 2013). Prior contact betweenthese two species is suggested by the introgression oftypical V. genesii mtDNA into some Altaian popula-tions of V. aff. genesii which possess typical nDNAsequence for that entity (J.C. Nekola, unpublisheddata).

Throughout its extensive range Vertigo aff. genesii islargely limited to graminoid-dominated fens or otherwetlands (Nekola & Coles 2010). Its habitat thus basi-cally mimics V. genesii, although the latter speciesappears more constrained to base-rich habitats. In theAltai, V. aff. genesii is the second most common full-glacial relict snail, being recorded from 25 sites, oftenin large populations, especially in graminoid-dominatedfens and seepages. Several populations were also foundin wet tundra habitats.

In the central European Quaternary, V. genesii wascommon during the warm phases of the Lateglacialand early Holocene (Lo�zek 1964; H�ajkov�a et al.2015a). Although Lo�zek (2001) did not considerV. genesii characteristic of central European loessenvironments (fossil records from loess deposits arevery rare; see Moine et al. 2008), the observed cli-matic preferences of V. aff. genesii in the Altai refu-gium are very similar to those of the characteristicloess species Vallonia tenuilabris and Pupilla loessica(Figs 3, 4). The general absence of V. genesii fromloess assemblages is probably due to its restriction towetland habitats. It is therefore likely that it onlystrayed accidentally into upland sites where loess wasbeing deposited.

Vertigo pseudosubstriata. – Vertigo pseudosubstriatahas a similar modern and Pleistocene/Holocene distri-bution to Pupilla loessica. Originally described fromCzech Republic Pleistocene loess deposits (Doln�ı

V�estonice) in 1954 and it was discovered to be extantabout 30 years later in the Tien Shan Mts and subse-quently from several other central Asian mountainranges (e.g. Meng 2009; Sysoev & Schileyko 2009;Hors�ak et al. 2010).

It is the least frequent glacial relict snail speciesfound in the Altai, having a scattered distribution inisolated habitats such as fens, seepage slopes andhumid shrublands. It was absent from a number ofseemingly suitable sites across the region. Its climaticrange in the Altai matches that of relict species thatsurvived in temperate Europe such as Pupilla alpicola.

This species is also rare in loess deposits of centralEurope (Lo�zek 2001). Perhaps its patchy contempora-neous distribution centred on wet-mesic sites helps toexplain its rarity in loess deposits.

Pupilla alpicola. – Pupilla alpicola inhabits treelessalkaline fens and fen meadows, and until recently wasonly known from the Alps and Carpathians (Kerneyet al. 1983; Hors�ak et al. 2011). Recent molecularstudies (Nekola et al. 2015) have demonstrated thatthis species also includes the more highly calcifiedshells previously termed Pupilla pratensis. As a result,the modern distribution of this species has been con-siderably expanded both to the west (British Isles,Scandinavia) and east (Altai) (Hors�ak et al. 2010). It islikely that at least some (e.g. appendix S1 in Hors�aket al. 2010) or all records of P. muscorum reportedfrom interior Asia actually represent P. alpicola. Care-ful examination of shell microsculpture and architec-ture (rather than apertural characters) is required foraccurate species assignment in this genus (Nekola et al.2015).

In the Altai this species was found in various high-humidity habitats such as open fen and fen woodland,with some populations also occurring in wet shrubtundra (Table 2).

Pupilla alpicola, namely its form densegyrata, ischaracteristic of the younger loess sediments in centralEurope (Lo�zek 2001), and should be considered a localLateglacial relict (Lo�zek 1964; Hors�ak et al. 2011).Lo�zek (1964, 2001) reported various ecomorphologicalforms of P. alpicola from central European full-glacialdeposits that may correspond to the ecophenotypicvariation observed in modern populations.

Pupilla turcmenica. – Pupilla turcmenica is a centralAsian species that inhabits steppe and other dry, tree-less habitats (Sysoev & Schileyko 2009). Based on shellshape, whorl expansion and periostracal microsculp-ture, it resembles the characteristic central Europeanloess species Pupilla sterrii, which was widespread andone of the most abundant species of loess assemblages(Lo�zek 2001). This species remains common acrosscentral and southeastern Europe (Kerney et al. 1983)where it is mainly confined to rocky or very stony


steppe or other open-land habitats (Hors�ak et al.2013a). DNA sequence analysis has shown P. sterriiand P. turcmenica to be sibling taxa (Nekola et al.2015), with the latter perhaps being a subspecies of theformer.

Pupilla turcmenica, as an ecomorphological parallelto European P. sterrii, is the only characteristic glacialspecies found almost exclusively in the dry steppe habi-tats in the Altai, which were occupied by the species inclimatically diverse areas including those with the high-est temperature and precipitation (Figs 3, 4). Mostpopulations were found in rocky or very stony dry hab-itats and the species expressed remarkable ecopheno-typic variation in the development of aperturalcalcification, including the populations correspondingto putative Pupilla altaica, which was found to be con-specific with P. turcmenica on the basis of geneticanalyses (Nekola et al. 2015). This variability appearsto be linked to soil calcium content and aridity.Hoffmann et al. (2011) also reported P. turcmenicaonly from very dry habitats, but as they restricted it toonly that part of the species’ morphological variationcorresponding to the putative species P. altaica, theybiased the climatic affinities of the species to warmconditions. Based on our findings we suggest that loesspopulations of P. sterrii also expressed a strong affinityto very dry habitats in different macroclimatic con-texts.

It is important to note that the characteristic andwidespread snails of loess assemblages (sensu Lo�zek2001), which managed to survive across the lowlandsof temperate Europe, are species of either dry steppehabitats, e.g. Pupilla sterrii and Helicopsis striata orare generalists, e.g. Succinella oblonga and Trochulushispidus.

Vascular plants

From 570 quadrats sampled for vascular plants in theRussian Altai (Fig. 1), we identified 16 species (i.e.Agropyron cristatum, Allium strictum, Artemisia lacini-ata, A. rupestris, Astragalus frigidus, Blysmus rufus,Carex capillaris, C. microglochin, C. obtusata, Kobresiamyosuroides, Kochia prostrata, Krascheninnikovia cer-atoides, Ligularia glauca, Pentaphylloides fruticosa,Saxifraga hirculus and Trichophorum pumilum) thatoccurred in at least seven quadrats while also beingcharacteristic of full-glacial environments in centralEurope. In total, these species were represented by 425occurrences.

Kobresia myosuroides. – Kobresia myosuroides is a cir-cumpolar species distributed in the arctic zone of Asiaand North America, and in the mountains of the bor-eal and temperate zones of Eurasia and North Amer-ica (Meusel et al. 1965). In Europe, it is found in

western Scandinavia, and the Pyrenees, Alps, Carpathi-ans, Apennines and Dinarids.

In these mountain ranges it is generally confined towind-exposed summits where it grows with short hemi-cryptophytes and chamaephytes (including cushionplants) and numerous species of bryophytes andlichens (Petr�ık et al. 2005, 2006). Many common fea-tures shared by Kobresia grassland across the whole ofnorthern Eurasia (Ohba 1974) support the assumedrelict status of Kobresia myosuroides in the mountainranges of the European temperate zone.

In the Altai K. myosuroides is a typical species ofhigh-mountain tundra in the dry continental parts ofthis mountain range (Figs 3, 4). In places it is adominant or co-dominant species of mesic tundragrasslands (Table 2), but it also grows in patches ofwind-swept tundra with Dryas oxyodonta or in tundrascrub dominated by Betula rotundifolia. It occasionallyalso occurs in valley bottoms or intermountain basinswhere it is usually confined to wetter sites such as fens,and wet and saline grasslands.

Saxifraga hirculus. – Saxifraga hirculus is a circumpo-lar arctic-boreal species with disjunct occurrences inthe mountain systems of Europe and interior Asiaincluding the Caucasus (Meusel et al. 1965). InEurope, scattered populations exist, or once existed,in Romania, Switzerland, France, the British Isles,northern Poland, the Baltic Countries and Scandina-via, but the species has been declining and is becom-ing endangered across Europe (Vitozz et al. 2006). Amore continuous distribution starts in northernPoland and continues eastwards. Magyari et al.(2014a,b) found a continual pollen record of S. hircu-lus in a lake sediment dated back to the Last GlacialMaximum in the Harghita Mts in Romania, a regionwhere the species has survived in fen refugia up tothe present (Kerekes 2008). Refugial European popu-lations are limited to brown-moss fens with a stablewater table (either acidic or alkaline) situated closeto the soil surface (Vitozz et al. 2006), where the spe-cies apparently experiences low competition pressureas it probably did in the cold glacial tundra. Thesesites tend to occur in lowlands or mountain basinsrather than in once-glaciated sub-alpine or alpinezones.

In the Altai it grows in areas with a stronglycontinental climate where it prefers wet habitatssuch as tundra grasslands with Dryas oxyodontaand tundra scrub dominated by Betula rotundifolia,Pentaphylloides fruticosa and various Salix species.When the moss layer is well developed, S. hirculusreaches only low abundances. It also lives in openfens and wet grasslands (e.g. in the Dzhazator val-ley and on the Ukok plateau), where it can locallyreach high abundance.


The genetic structure of European and North Amer-ican populations reveals higher haplotype diversityoutside the limits of the last glaciations, suggesting thatit survived the cold stages in the periglacial zone(Oliver et al. 2006). Molecular sequencing of organicsediments preserved in permafrost documents its full-glacial presence in nonglaciated northeastern Russia(Sønstebø et al. 2010).

Artemisia rupestris. – Artemisia rupestris has a distri-bution range similar to that of A. laciniata (see below),including the southern half of Siberia and adjacentparts of central Asia (J€ager 1987; Meusel & J€ager1992; Yourun et al. 2011) and was also reported froma single site in the southwestern Yukon Territory(Shultz 2006). However, this population was recentlyre-evaluated as a distinct species not closely related toA. rupestris (Riggins & Seigler 2012). In Europe, iso-lated populations are situated in northern Russia andon the Baltic islands of Saaremaa, €Oland and Gotland.Occurrences in central Germany disappeared by theearly 20th century (J€ager 1987). All of these regionsare characterized by a rather continental climate.

At its German sites, A. rupestris was confined to sal-ine pastures and meadows. The sites in the Balticregion are found on shallow limestone soils of alvarsand seashores. In both habitats, the species is protectedfrom competition either by salinity or by other soilproperties (J€ager 1987).

In the Altai A. rupestris occurs in dry high-moun-tain landscapes, most typically at altitudes above2000 m. It is also occasionally found on valley bottomsat lower altitudes. It occurs in various types of grass-lands, most often in mesic high-mountain grasslands(steppe-tundra), but also in meadow steppes and drysteppes on south-facing slopes at high altitudes(Table 2). It can be rarely found in low scrub (Betularotundifolia and Caragana arborescens) or in openlarch forests.

Pentaphylloides fruticosa. – Pentaphylloides fruticosa(syn. Potentilla fruticosa) is a species with a continuousNorth American and central and east Asian distribu-tion, and several disjunct occurrences in central andwestern Europe (Meusel et al. 1965), and being morefrequent east of the Urals and Caucasus. One popula-tion is found in Bulgaria (Rhodopes Mts) and severalin the Pyrenees, British Isles, Baltic Countries andsouthern Sweden. Although this species prefers cold,but usually neither high-mountain nor arctic regions inEurasia, it occurs in North America in a wide varietyof mostly calcareous habitats (including fens and xericrock outcrops; Nekola 1990) ranging from near sealevel to alpine meadows. A phylogeographical studybased on chloroplast DNA, carried out on the Qing-hai-Tibetan plateau, suggested that P. fruticosaexpanded its range on the plateau during the periods

of climatic cooling and contracted to the interior high-mountain region during warmer periods (Shimonoet al. 2010).

In the Altai this is a common species that occupiesvarious habitats but favours tundra shrub and wetwoodlands. Considering the climatic affinities of themodern Altai populations, it may be assumed thatEuropean and central Asian populations were inter-connected during the last glacial period when the land-scape was open and temperatures at low altitudes incentral Europe were similar to those that currentlyoccur in the south Siberian mountains (e.g. Isarin &Renssen 1999). Germination experiments (Meshinev1975) showed that this species needs a sufficientamount of light to germinate and establish. This indi-cates that the bottleneck for P. fruticosa survival wasprobably the Holocene climatic optimum, when open,unproductive or disturbed habitats suitable for regen-eration of this species were rather rare.

Carex obtusata. – Carex obtusata is a circumpolar spe-cies (Meusel & J€ager 1992) with a more or less continu-ous distribution range in southern Siberia (Malyshev1990), occurring further in adjacent parts of centralAsia in Kazakhstan, Mongolia and northern China(Tang et al. 2010). It is further present in the Caucasusand in North America (Ball & Reznicek 2002). Yurtsev(1982) considered C. obtusata to be a typical species ofthe Beringian steppe-tundra (see also Kucherov &Dani€els 2005). In Europe, it is scattered throughoutEuropean Russia and Ukraine, and is found on theSwedish island of Gotland, at four sites in central Ger-many (Schultze-Motel 1967–1980), and at two locali-ties in the eastern part of the Czech Republic (�Repka2007).

The German populations, of which only one is stillextant, occur in dry grasslands or open pine forests onsandy soils (www.floraweb.de). Both Czech popula-tions are found in dry steppe grasslands on shallowacidic soil.

In the Altai C. obtusata is a widespread species ofvarious steppe and tundra grasslands and open larchforests especially in dry continental areas (Table 2). Itoften co-occurs with Artemisia rupestris, but it is morecommon than this species and shows a stronger prefer-ence for drier steppe and dry larch forests. These affini-ties suggest that it may have been common in theEuropean glacial landscape and that the current iso-lated localities may be remnants of its past, largerrange.

Astragalus frigidus. – Astragalus frigidus is a remark-ably variable Eurasian taxon with a distributionextending from northernmost Europe and the low-lands of northern Siberia to the Far East, Japan,southern Siberian mountain systems and the adjacentparts of central Asia (Meusel & J€ager 1992; Xu &


Podlech 2010). It is now divided into three subspecies(e.g. Vydrina 1994; Elven 2014), of which the type sub-species is present both in Europe and in the Altai. TheEuropean distribution of A. frigidus, now ratherfragmentary, was probably more continuous in thePleistocene, as indicated by Lateglacial fossil recordsfrom the Baltic Sea area (Hult�en & Fries 1986).

In the Alps it occurs most frequently in alpine grass-lands on steep slopes or convex ridges on slightly cal-careous soils, usually at elevations of 1700–2500 ma.s.l., whereas its northernmost localities in Norwayare situated almost at sea level (Gams 1923–1924).

In the Altai A. frigidus is a species of continentalhigh-mountain landscapes, in which it is confined tovarious types of forests and open woodlands domi-nated by Larix sibirica, Picea obovata, Pinus sibirica,scrub with Betula fruticosa (B. humilis s. lat.) and Pen-taphylloides fruticosa, and sub-alpine tall-forb vegeta-tion. It avoids dry or mesic short grasslands (Table 2).

Ligularia glauca s. lat. – Ligularia glauca is a Euro-pean-Siberian species with a strongly disjunct range inEurope and a continuous distribution in the southernparts of western and central Siberia (Ermakov et al.1999). The European populations are morphologicallyslightly different from those in Siberia, sometimesbeing considered as different subspecies or as thespecies L. carpathica (e.g. Pojarkova 1961). In allEuropean countries where the species occurs (Slovakia,Romania, Bulgaria and Ukraine) it is critically endan-gered and considered as a glacial relict (Tomescu &Chifu 2009; Kricsfalusy & Budnikov 2007). In Europe,L. glauca grows on limestone outcrops with semi-openvegetation (Slovakia; Hajd�uk 1957), semi-dry grass-lands (Romania, Ukraine), calcareous fens (Romania;Tomescu & Chifu 2009) and sub-alpine tall-herb vege-tation (Bulgaria; Roussakova 2002). Although thesehabitats differ in many attributes, they all share anopen or semi-open structure with certain continuityduring the Holocene (Fajmonov�a et al. 2013). Presum-ably this species may have been widely distributed incentral and eastern Europe under the strongly conti-nental climate of the glacial period.

In the Altai L. glauca has two ecological optima. Inthe more humid climate of the northern Altai, it pre-fers meadow steppes with Brachypodium pinnatum,Calamagrostis arundinacea, Crepis sibirica and Phlomistuberosa, but it also occurs in drier steppes (with Asteralpinus and Iris ruthenica) and in herb-rich birch for-ests (Table 2). By contrast, it occupies tundra grass-lands and tundra scrub in the continental southernpart of the Altai (e.g. Ukok plateau, Kurai and Chuyaranges).

The fact that most European localities are concen-trated at low or middle elevations in summer-warmsoutheastern Europe suggests that this species doesnot require cold climate per se. It rather benefits

from high light availability and soil conditions asso-ciated with continental climate (Fig. 2).

Blysmus rufus. – Blysmus rufus is currently distributedin three distinct regions: (i) the Atlantic and Baltic Seacoast in northern and northwestern Europe (the Neth-erlands, Scotland, Denmark, Scandinavia, BalticCountries, NW Russia); (ii) central Asian, stronglycontinental mountains and (iii) North American seacoasts (Meusel et al. 1965). Along the sea coast, itoccupies dune slacks (Hofstra 1996) and salt-marshes,whereas in central Asia (Altai and Trans-Altai region)it occupies saline meadows on clayey soils (von Wehr-den et al. 2006; e.g. Trichophorum pumilum halophyticfen grasslands, D�ıt�e et al. 2013).

In the Altai B. rufus mostly grows in saline grass-lands dominated by Halerpestes salsuginosa and Glauxmaritima, and open fens with a well-developed bryo-phyte layer. It was recorded in river floodplains of theChuya basin and Ukok plateau, and in the Menka andChuya river valleys, where it frequently co-occurredwith Trichophorum pumilum and Carex microglochin.An important environmental factor for B. rufus is highpH and high mineral or salt content. This environmen-tal feature is common for European sea-coasts andalso for central Asian alluvial meadows and fens whereit is supported by salinization because of low summerprecipitation.

In Eurasia the two modern centres of distributionmay represent refugia from a once more continuousfull-glacial range, as supported by fossil seeds found inLateglacial sediments far from modern occurrences insouthwestern Slovakia (Borsk�a lowland; H�ajkov�aet al. 2015b). Here, numerous Blysmus sp. seeds werefound along with the seeds of Triglochin maritima. Wehypothesize that during the glacials, Blysmus rufus mayhave occupied saline alluvial meadows and fens sup-plied by calcium- or sulphate-rich groundwater, i.e. thesame habitats as it occupies currently in central Asia(Table 2).

Krascheninnikovia ceratoides. – Krascheninnikovia ce-ratoides is a circumpolar species with a large distribu-tion range including mainly continental parts ofEurasia and North America (Holmgren 2003). ItsNorth American subspecies (subsp. lanata) occurs inthe intermountain region of the USA, whereas the typesubspecies has a large Eurasian distribution rangeincluding southern Siberia, central and southwesternAsia, eastern, central and western Europe, and reach-ing also to the Sinai Peninsula and the Atlas Mts ofnorthern Africa (Uotila 2011). In Europe, it is scat-tered throughout southern Russia and southernUkraine, and also occurs in Romanian Transylvania(B�ad�ar�au et al. 2002; Kov�acs 2011), Macedonia, Hun-gary, Lower Austria (Jalas & Suominen 1980) and thesoutheastern Czech Republic (now vanished; Tom�sovic


1990). The westernmost populations are found in thenortheastern Iberian Peninsula (P�erez-Collazos & Ca-tal�an 2007).

In Europe it has a similar ecology to Agropyroncristatum s. lat. and Kochia prostrata (Niklfeld 1964;B�ad�ar�au et al. 2002; Kov�acs 2011), being confined tovery dry open grasslands. In the Altai this low shruboccurs especially in desert steppes of the Chuya basin,but is also found elsewhere in dry steppes where itshares very similar habitat preferences to, but is muchrarer than, Agropyron cristatum and Kochia prostrata(Table 2).

Carex microglochin. – Carex microglochin is one of thefew Carex species with a bipolar distribution, occur-ring in the northern part of the Northern Hemisphereand in the Andean and Patagonian regions of SouthAmerica (Wheeler & Guaglianone 2003). In Eurasia, itis almost continuously distributed in the southern partof central and eastern Siberia, from the Altai east-wards. It also occurs as disjunct populations in theCaucasus and eastern Himalayas. In Europe, it is atypical arcto-alpine species and is found in the alpinezone of the Alps, northern Scandinavia, Iceland andthe northern part of the British Isles (Meusel et al.1965), where it occupies springs, fens, stream and lakemargins, and in humid regions, also rock crevices.

In the Altai C. microglochin grows predominantly inopen calcareous fens and wet saline grasslands withhigh soil pH (above 7.0) and high conductivity (300–2000 lS cm�1), mostly together with Blysmus rufusand Trichophorum pumilum. It was also recorded rarelyin tundra grasslands (Ukok plateau). Climatic condi-tions are similar for all three of these species, i.e.strongly continental climate with mean January tem-perature about �19 °C and an annual precipitation of<250 mm (Figs 3, 4). These factors, together with dis-turbances associated with dry, open continental land-scapes, may further weaken competition from otherwetland plants including bryophytes that may competewith vascular plant seedlings.

These climatic parameters suggest that in glacialEurope C. microglochin may have been able to live notonly in the mountains, but also at lower altitudes inregions with a cold and dry continental climate.Although suitable habitats, such as calcareous fens andsaline grasslands (Table 2, Fig. 2), probably occurredin the region, direct fossil evidence for the presence ofthis species is missing. In modern Europe it is limitedto cold and wet areas, with the mean July temperatureoptimum of about 11 °C documented from Norway(Saetersdal & Birks 1997) being even lower than thatfound in the Altai (about 14 °C).

Trichophorum pumilum. – Trichophorum pumilum (syn.Baeothryon pumilum) in Europe is considered an arcto-alpine species that also inhabits lower latitude/

elevation travertine sites (i.e. with tufa formation). Itoccurs in the alpine belt of the Western and SouthernAlps, in submontane travertines in the Western Carpa-thians (Slovakia) and in two regions of Norway, onenorth of the Arctic Circle and one in the mountainslocated more southwards (D�ıt�e et al. 2013). The morecontinuous distribution starts in Iran (Naqinezhadet al. 2009) and continues through the central Asianmountains (Tien-Shan, Himalaya, Altai) and south-western Siberia to Mongolia (Minayeva et al. 2005).The species ranges across the Rocky Mountains in wes-tern North America, with a disjunct eastern occur-rence along the Gulf of St. Lawrence in Quebec.

In the Altai T. pumilum occupies mostly wet salinegrasslands and, less frequently, open fens or tundragrasslands. We recorded this species in the Chuyabasin, Menka river valley and the northern Kurairange. In central Europe, T. pumilum is confined to cal-cium-rich fens of the Caricion davallianae alliance,characterized by low competition from other vascularplants, whereas in Norway it occurs in low-productivetundra (D�ıt�e et al. 2013). The floristic composition ofthe Western Carpathian travertine fens exhibits consid-erable similarity to that of Altaian saline grasslandsowing to a high representation of halophytic and sub-halophytic species such as Glaux maritima and Triglo-chin maritima. By contrast, populations in the Alpsand in Norway contain fewer (sub)halophytic andmany more arcto-alpine species (D�ıt�e et al. 2013).

Records of fossil T. pumilum seeds from the full-gla-cial period in Belgium (~700 km from the nearestextant populations in the Alps; Haest et al. 1986) andfrom the Lateglacial/early Holocene transition in Slo-vakia (outside the recent distribution range; D�ıt�e et al.2013) suggest that the species was more widespread inEurope during the last glacial period, occupying notonly travertine swards, fens and tundra, but also salinegrasslands or river banks. In the Western Carpathiansit tends to be concentrated in ancient calcareous fens,originating in the Lateglacial or Early Holocene, but isabsent from younger fens (H�ajek et al. 2011).

Carex capillaris. – Carex capillaris exhibits a boreal-circumpolar distribution and is considered as an arcto-alpine species in Europe. It occurs in the alpine andsub-alpine belts of the Pyrenees, Alps, Carpathiansand Caucasus, and rarely in the Dinaric Mountains.Generally, its distribution is rather continuous acrossthe boreal and sub-arctic zones of the Northern Hemi-sphere, but its individual populations are sometimesseparated by large distances (Hult�en & Fries 1986;Stevanovi�c et al. 2009). There is a fairly continuousdistribution range in Scandinavia and northeasternEurope.

In Europe the species occurs in wet calcareoushabitats (springs, fens or tundra) under cold climate atboth high altitudes and high latitudes (Dierssen &


Dierssen 1985; Moen et al. 2012). At lower altitudes oftemperate Europe it is confined to relict habitats, suchas Western Carpathian calcareous fens (D�ıt�e & Pu-kajov�a 2004) or extremely species-rich wooded mead-ows in Estonia (Kull & Zobel 1991). In North Americait is largely limited to cool, humid calcareous habitatswhere it is characteristic of sites supporting glacial re-licts along the Great Lakes (Voss 1972).

In the Altai the ecological affinities of C. capillarisare rather similar: it was recorded in open wet wood-lands dominated by Larix sibirica, Picea obovata andtree willows (Kurai basin, Dzhazator river valley), tun-dra grasslands with Dryas oxyodonta, tundra scrubwith Betula rotundifolia and wet grasslands with Fili-pendula ulmaria.

Fossil records of C. capillaris are very rare in Eur-ope and they come only from the sites of its currentoccurrence (e.g. middle Holocene in the Alps; Jan-kovsk�a & Rybn�ı�cek 1988). Nevertheless, based on itsrecent distribution pattern and analogy with centralAsia, we hypothesize that this species may also haveoccurred at lower altitudes of the European Pleisto-cene landscape.

Agropyron cristatum s. lat. – Agropyron cristatum is amorphologically extremely variable species with a largeEurasian distribution range, including the southernhalf of Europe, southern Siberia, large parts of centralAsia and northeastern China, the Caucasus andextending to Morocco (Meusel & J€ager 1992; Tzvelev1976). Across its entire distribution range, this speciesis usually divided into numerous subspecies (e.g.Tzvelev 1976), which are sometimes given species rank(e.g. Baikov 2005).

Especially in the forest-steppe regions of eastern cen-tral Europe, A. cristatum is traditionally considered tobe a relict species of the Pleistocene cold loess steppe,as are Kochia prostrata and Krascheninnikovia cerato-ides with which it often co-occurs (Wendelberger 1954;Niklfeld 1964), usually in open dry grasslands on loessslopes (Z�olyomi 1958; Niklfeld 1964; Dihoru & Donita1970; Chytr�y 2009).

The habitat affinities of A. cristatum in the Altai(Table 2) support the hypothesis of its glacial relictstatus in temperate Europe. It is confined to thedriest habitats of this area (short-grass steppes), where itoften co-occurs with Kochia prostrata and occasionallyKrascheninnikovia ceratoides. These interspecific associa-tions can also be found over a broader region of centralAsia (Hilbig 1995; Korolyuk 2002). In the Altai A. crist-atum is most common in the dry intermountain Kuraiand Chuya basins in the southeast, but also occurs else-where on dry, usually south-facing steppe slopes. Occa-sionally it can be also found in short steppic scrub withCaragana pygmaea and other species of Caragana andSpiraea, or in open, herb-rich larch forests, but usuallyin landscapes dominated by dry to typical steppe.

Allium strictum. – Allium strictum is a widely distrib-uted species known from the southern part of Siberia,the adjacent parts of central Asia and the Russian FarEast, including the southern part of the Kamchatkapeninsula. Its continuous distribution range reachesEurope in the middle Volga basin. Further west itexhibits a disjunct distribution occurring in the valleysof the western and eastern Alps, in central Germany,southwestern Poland (now extirpated; Ka�zmiercza-kowa & Zarzycki 2001), on the Baltic island of €Olandand in the Russian part of Karelia (Meusel et al.1965). It is further known from about 30 sites in cen-tral and northern Bohemia in the Czech Republic(Martinovsk�y 1969), from a single locality in the Wes-tern Carpathians in Slovakia as well as from a handfulof sites in western Ukraine (Krahulec et al. 2006). Allof these European occurrences have been considered asrelicts of cold glacial steppes (Martinovsk�y 1969;Krahulec et al. 2006).

The European sites are situated from the lowlandsup to elevations of about 2500 m a.s.l. in the Swisscanton of Valais. The Alpine populations are foundmainly in the continental valleys of the western andcentral Alps, where it grows with dry-grassland speciessuch as Koeleria macrantha (syn. K. cristata) andPhleum phleoides (Vierhapper 1919). Many of thesespecies have large Eurasian or even circumpolar distri-bution ranges. In Bohemia, it occurs in short-grasssteppes with Festuca valesiaca and Carex humilis andon dry rock outcrops with Festuca pallens (Toman1981), usually on base-rich volcanic or metamorphicbedrock and in areas with dry continental climate.

In the Altai A. strictum occurs in various types ofmoderately dry to mesic grasslands, ranging from typi-cal steppe through meadow steppe to tundra grassland(Table 2). These grasslands are usually dominated bysedges (Carex humilis and C. pediformis), grasses (He-lictotrichon altaicum) or Iris ruthenica. It is also foundin steppic scrub with various species of Caragana,Cotoneaster, Rosa and Spiraea, and in herb-rich larchforests that contain grassland species.

Kochia prostrata. – Kochia prostrata is a small procum-bent shrub with a Eurasian distribution range (Schu-bert et al. 1981). It is almost continuously distributedin southern Siberia and southwestern and central Asia.In Europe it occurs as scattered populations betweenthe central Iberian Peninsula in the west and the lowerVolga River in the east (Jalas & Suominen 1980). It isknown also from Morocco and Algeria in northwest-ern Africa (Uotila 2011).

In Europe it often co-occurs with Agropyron crista-tum s. lat. (Z�olyomi 1958; Chytr�y 2009) in communitiesconsidered to be Pleistocene cold-steppe relicts (Wen-delberger 1954; Niklfeld 1964). In the Altai the habitatrange occupied by K. prostrata is very similar to thatof A. cristatum s. lat., although it is hardly ever found


in meadow steppe, steppe scrub or in open woodland.These species often grow together in dry steppes. Theconstancy of the interspecific association of these twospecies between the core area of their distribution incentral Asia and scattered occurrences in Europe sup-ports the hypothesis of the relict status of the Euro-pean populations.

Artemisia laciniata. – Artemisia laciniata has a largedistribution range covering the southern half of Sibe-ria, parts of the Russian Far East, Mongolia andnortheastern provinces of China (J€ager 1987; Meusel& J€ager 1992; Krasnoborov 1997). It is also reportedfrom the Yukon Territory and Alaska (Shultz 2006)but the taxonomic identity of the North Americanpopulations is not reliably resolved. In Europe A. la-ciniata occurs in the Don basin and in eastern Austriaon the eastern shores of Lake Neusiedl. Several popu-lations also occurred near Staßfurt and Artern in cen-tral Germany but all went extinct in the late 19thcentury (J€ager 1987). In that area populations wereconfined to saline soils, thereby avoiding competition.At these sites it was less frequent than the co-occurringA. rupestris.

In the Altai A. laciniata is a species of forest-steppelandscapes, where its main habitat is species-rich mea-dow (tall-herb, closed) steppe with Carex humilis, Fra-garia viridis, Galium boreale, Geranium pratense andPhlomis tuberosa. It also occurs in open herb-rich for-ests of Betula pendula, Larix sibirica and Pinus sylves-tris, which are adjacent to steppes and contain manysteppe species in their herb layer (Table 2). Artemisialaciniata also grows in other types of grasslands, scruband forests.

Relict species in the Pleistocene environments

The distributions of the vascular plant and land-snailspecies considered in this paper were studied in an areawith mean annual temperatures ranging from �6.7 to3.4 °C (median �2.5 °C) and total annual precipita-tions of 137 to 593 mm (median 283 mm; Table 1).The analysis of the macroclimatic requirements of theglacial relict species sampled in the southern Siberiancontinental refugium yielded consistent predictions ofthe full-glacial central Europe climate derived bothfrom plant and land-snail populations. The results didnot substantially change if only characteristic glacialland snails were considered or if the central Asian Pu-pilla turcmenica and Beringian Vertigo aff. genesii werealso included (Table 1). We also observed similar pat-terns of individual species affinities of the studied vas-cular plants and snails (Figs 3, 4). In both groupsthere were species limited to areas with a colder anddrier climate (e.g. snails Columella columella and Pupil-la loessica, and plants Kobresia myosuroides and Kra-scheninnikovia ceratoides), whereas other species

required a warmer or damper climate (e.g. snails Pupil-la turcmenica and P. alpicola, and plants Artemisia la-ciniata and Carex capillaris). For snails, most of theseresults are in good concordance with those alreadyreported by Hors�ak et al. (2010) and Hoffmann et al.(2011). In comparison with these two earlier studies,however, the results presented here are based on amuch larger data set and, most importantly, they arebased on a corrected and improved taxonomic conceptof some problematic species developed using extensivemolecular examination (Nekola et al. 2015; J.C. Nek-ola unpublished data). The results presented here thusprovide a robust overview of the macroclimatic andhabitat affinities of the characteristic European full-glacial species, including those for which very fewrecords were available in Hors�ak et al. (2010) or wereentirely missing in Hoffmann et al. (2011) (Pupilla al-picola, Vertigo parcedentata and V. pseudosubstriata).

The analysis of climatic conditions also indicatesthat distributional shifts of the studied species duringthe Pleistocene/Holocene transition were closelyrelated to their climatic tolerances. Snail species thatwent extinct in Europe are currently confined in theAltai to the areas of the coldest and driest climate(i.e. Pupilla loessica and Vallonia tenuilabris). By con-trast, species that have survived at lower altitudes incentral Europe have climatic preferences that areshifted towards warmer and/or wetter conditions (Pu-pilla alpicola and P. turcmenica/sterrii). The onlyexception is Vertigo pseudosubstriata, which has van-ished from Europe yet occupies sites climatically sim-ilar to P. alpicola in the Altai. As it was found atonly six sites, with most exhibiting strong groundwa-ter influence, the actual climate experienced by thesepopulations may be significantly different from themacroclimate values encountered in this study. Like-wise, plant species with the lowest temperaturerequirements in the Altai tend to be confined to refu-gial habitats in European mountains (e.g. Kobresiamyosuroides), whereas those currently confined towarmer areas in Siberia also survive in refugial popu-lations in the European lowlands. However, theselowland populations have been strongly restricted,perhaps mainly because of the spread of competi-tively stronger species, and many survive in specifichabitats with limited competition, such as inland sal-ine grasslands (e.g. Artemisia laciniata) or dry loessslopes often affected by erosion (e.g. Kochia prostrat-a).

A basic assumption of our study is that the extantrefugial populations in the Altai region occur underclimatic conditions that are similar to those character-istic of a considerable part of Eurasia in certain peri-ods of the Pleistocene. We can conclude that theobserved climatic requirements of both plants andsnails in southern Siberia are in good agreement withthe climatic values reconstructed for full-glacial tem-


perate Europe based on both palaeoclimatic modellingand fossil pollen data. Based on a positive degree-dayglacier mass-balance model with reduced precipitationof 25–75%, Heyman et al. (2013) suggested temperaturedepressions during the Last Glacial Maximum (LGM;about 18–21 ka BP) of between 8.0 °C and 15 °C forfour central European upland regions. However, LGMclimate models have generally reconstructed a lesssevere climate than that inferred from pollen data. Forexample, Strandberg et al. (2011) predicted meanannual temperatures to have been 5–10 °C cooler thantoday for the ice-free areas of central Europe duringthe LGM. Jost et al. (2005) predicted the differencesfrom the present temperature of the coldest month tobe around 10 °C and ~350 mm for the total annualprecipitation using various climatic models. By con-trast, based on pollen data Peyron et al. (1998) deriveda mean annual temperature for 18 ka BP of 12�3 °Ccooler than today for the northern–western zone abovethe Pyrenees–Alps line, and total annual precipitationof 60�20% lower than today. This was confirmed byJost et al. (2005) based on an updated modern pollencalibration data set. These new data suggested aslightly less cold climate than reconstructed before, butstill considerably colder than today for the meantemperature of the coldest month.

Likewise, the habitat requirements of modern popu-lations of the studied species in southern Siberia sup-port the current concept of the Pleistocene loess-steppeenvironments developed on the basis of the fossilrecords (e.g. Lo�zek 2000; Hors�ak et al. 2010; Magyariet al. 2014a,b). Both snail and plant assemblages char-acteristic of the full-glacial environment include speciespreferring both dry steppe and tundra conditions(Table 2). Additionally, both in Europe and centralAsia, many of these full-glacial snails and plants arecurrently confined to wet habitats, such as alkaline fensand wet grasslands. For some snail species this findingcan explain their rare occurrence in loess deposits.Some (e.g. Columella columella and Vertigo parcedenta-ta) currently establish abundant populations in fensand wet woodlands. In the European glacial landscapesthey may have partly survived in small fens, wetlandsor woodland patches, although steppe-tundra environ-ments probably formed the landscape matrix over largeareas especially in the coldest phases (e.g. Willis & vanAndel 2004; Kune�s et al. 2008; Magyari et al. 2014a,b). Snails, in particular, may have been partly depen-dent upon shelters provided by tree stands (Hors�aket al. 2010) as they do not have any specific cryopro-tective chemicals (Riddle 1983). We can also consider aprotective role of snow cover preventing topsoil fromfreezing; however, this was probably less importantunder the dry full-glacial climate than it is today. Bycontrast, most herbaceous plant species considered asglacial relicts are typical of treeless habitats, which sug-gests that herbaceous plants may be more limited by

summer than winter temperatures and that they arenot so dependent upon the availability of shelters tosurvive cold periods. However, as the three tempera-ture characteristics (Table 1) assessed for the Altaipopulations were strongly intercorrelated and weobtained almost identical patterns for all studied snailand plant species, we present the results for individualspecies only in relation to mean January temperature.

The data derived from the Altai glacial snail popula-tions might be very useful for improving calibration ofmodern data sets by developing transfer functions forthe reconstruction of glacial climate (Rousseau 1991;Moine et al. 2002). The reconstructed glacial tempera-tures derived from modern snail species distributionsacross western and central Europe were considered tobe milder than what would be expected based on othermethods (Moine et al. 2002). Likewise, in a compari-son with our data, the existing transfer functions pro-duced temperatures about 10–15 °C warmer for thecoldest month, but were almost identical for the warm-est month (see Rousseau 1991; Moine et al. 2002). Thiscan be explained by the lack of modern data in the cali-bration data sets for many characteristic species ofloess faunas, especially those that have become extinctin western and central Europe. Their modern climaticaffinities in the Altai clearly show that they are limitedto the areas of colder climate as opposed to the speciesthat are still common in Europe.

Conclusions

Our results show that climatic affinities of modern ref-ugial populations of both land snails characteristic ofcentral European loess sediments and presumably gla-cial-relict plant species are in good concordance withclimatic scenarios derived from fossil pollen data andclimate models. The plausibility of these conclusions issupported by the observed pattern of differences in cli-matic affinities of individual species, which closelymatches the variation in their distributional shifts atthe Pleistocene/Holocene transition. In both taxa therewere species limited to areas with colder and drier mac-roclimates (e.g. snails Columella columella and Pupillaloessica, and plants Kobresia myosuroides and Krasche-ninnikovia ceratoides), whereas other species preferredareas of warmer or moister macroclimate (e.g. snailsPupilla turcmenica and P. alpicola, and plants Artemi-sia laciniata and Carex capillaris). However, in suchareas individual species may be confined to eithermoist or dry habitats regardless of the macroclimaticconditions. For example, Pupilla turcmenica prefers dryhabitats in areas of warm and moist macroclimatewhereas some wetland plant species prefer mires andriver shores in areas of cold and dry macroclimate.

This study also highlights the importance of wethabitats as refugia in the generally dry full-glacial low-land landscape. All of these findings suggest that the


habitat requirements of southern Siberian populationsof land snails and vascular plants can provide impor-tant and realistic insights for reconstructions ofEurasian, especially central European, glacialenvironments. Such observations from modern analo-gues may be very useful for explaining some question-able palaeoecological interpretations of several loessspecies as discussed in detail above.

Acknowledgements. – Many thanks go to our Russian colleaguesfor their logistic support and help in the field and Ond�rej H�ajekfor preparing the climate data and the map. We are very gratefulto Olivier Moine, John Birks and Richard Preece for many valu-able comments and improvements of the manuscript. This studywas supported by the Czech Science Foundation (P504/11/0454),the institutional support of Masaryk University, and the long-term research development project no. RVO 67985939.

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