Climate signals in the ring widths and stable carbon, hydrogen and oxygen isotopic composition of Larix decidua growing at the forest limit in the southeastern European Alps

The southeastern border of the European Alps is not well resourced with high-resolution climate proxies and experiences a distinct climatic regime from the northern and western Alpine zones. Here, we present new high-resolution climatic proxies (AD 1907–2006) from ring widths and stable carbon (δ¹³C), non-exchangeable hydrogen (δ²H) and oxygen (δ¹⁸O) isotope ratios of cellulose extracted from Larix decidua tree rings, growing at the forest limit in the southeastern European Alps (Slovenia). δ¹³C, δ²H and δ¹⁸O are strongly (p < 0.001) and positively correlated with each other. June temperature has the strongest control on tree ring width (TRW), while later summer conditions (July–August) influence the stable isotope composition. All four proxies are strongly correlated (r > 0.4; p < 0.001) with summer temperature and also sunshine hours, while precipitation has less impact. A combination of TRW and δ¹³C provides the greatest potential for reconstructing past temperatures (June–August) with significant (p < 0.001) correlations with gridded temperatures extending across a very large part of southern and western Europe west of the Carpathian Mountains. The water isotopes (oxygen and hydrogen) record conditions in the Adriatic and Mediterranean, which are the source area for the air masses that bring precipitation to this region giving strong correlations with temperatures in southern Italy and the western part of the Balkan Peninsula. Combining proxies with different spatial and temporal signals allows the strength and spatial footprint of climate signals to be enhanced. These findings open new perspectives for climate reconstruction in the southeastern European Alps and Western Balkans.     

An ectomycorrhizal symbiosis differently affects host susceptibility to two congeneric fungal pathogens

The role of ectomycorrhizal fungi in modulating host susceptibility to fungal pathogens is poorly known. Pinus sylvestris susceptibility to two congeneric fungal pathogens was compared between mycorrhizal and non-mycorrhizal plants. The ectomycorrhizal fungus was Suillus luteus, while the pathogens were Heterobasidion irregulare and H. annosum. H. irregulare is native to North-America and invasive to Europe, whereas S. luteus and H. annosum are native to Eurasia. Non-mycorrhizal plants were equally susceptible to both pathogens. Mycorrhizal plants were significantly less susceptible to H. annosum than non-mycorrhizal plants, whereas there were no differences when considering H. irregulare. The abundance of ectomycorrhizas was negatively and significantly correlated to the level of host susceptibility only to H. annosum. This study shows that the protective role of ectomycorrhizal fungi may differ when the host is challenged by congeneric fungal pathogens. Results add a further clue to explain and predict the invasiveness of H. irregulare in Europe.     

Genetic structure of expanding wolf (Canis lupus) populations in Italy and Croatia,and the early steps of the recolonization of the Eastern Alps

After centuries of range contraction and demographic declines wolves are now expanding in Europe, colonizing regions from where they have been absent for centuries. Wolf colonizing the western Alps originate by the expansion of the Italian population. Vagrant wolves of Italian and Dinaric-Balkan origins have been recently observed in the Eastern Alps. In this study we compared the genetic structure of wolf populations in Italy and Croatia, aiming to identify the sources of the ongoing recolonization of the Eastern Alps. DNA samples, extracted from 282 Italian and 152 Croatian wolves, were genotyped at 12 autosomal microsatellites (STR), four Y-linked STR and at the hypervariable part of the mitochondrial DNA control-region (mtDNA CR1). Wolves in Croatia and Italy underwent recent demographic bottlenecks, but they differ in genetic diversity and population structure. Wolves in Croatia were more variable at STR loci (N_A = 7.4, H_O = 0.66, H_E = 0.72; n = 152) than wolves in Italy (N_A = 5.3, H_O = 0.57, H_E = 0.58; n = 282). We found four mitochondrial DNA (mtDNA CR1) and 11 Y-STR haplotypes in Croatian wolves, but only one mtDNA CR1 and three Y-STR haplotypes in Italy. Wolves in Croatia were subdivided into three genetically distinct subpopulations (in Dalmatia, Gorski kotar and Lika regions), while Italian wolves were not sub-structured. Assignment testing shows that the eastern and central Alps are recolonized by wolves dispersing from both the Italian and Dinaric populations. The recolonization of the Alps will predictably continue in the future and the new population will be genetically admixed and very variable with greater opportunities for local adaptations and survival.     

Stayin' alive: survival of mycorrhizal fungal propagules from 6-yr-old forest soil

Spores and sclerotia are the main propagules that allow fungi to persist through unfavorable conditions and disperse into new environments. Despite their importance, very little is known about their longevity and dormancy, especially in ectomycorrhizal fungi. To assess the viability of ectomycorrhizal fungal spores in forest soil, we collected and buried non-sterile forest soil, in pots, in the field distant from an inoculum source. After 6 yr, a subset of this soil was assayed for viable spores by baiting the fungi with Bishop pine (Pinus muricata) seedlings. Our results show that the three most frequent colonizers in year 1 continued to colonize significant percentages of seedlings in year 6: Wilcoxina mikolae (77 %), Rhizopogon vulgaris (13 %) and Suillus brevipes (9 %). While three species that colonized low percentages of seedlings in year 1, Suillus pungens (1 %), Rhizopogon salebrosus (2 %), and Thelephora terrestris (5 %) were not recovered in year 6. Laccaria proxima, a species not seen in year 1, was recovered on a single seedling in year 6. This is the first report of long-term survival of S. brevipes and W. mikolae. Our results reveal a more complete picture of ectomycorrhizal fungal spore longevity in soil spore banks.     

Common Buzzards wintering strategies as an effect of weather conditions and geographic barriers

AimMigration is a constantly changing adaptation due to the climate condition evolution. The struggle for surviving during harsh winter season is different across Europe, being more complex toward the inner parts of the continent. The current approach explores the Common Buzzard number variation during the cold season and the climatic predictors of birds of prey wintering movements in relation to the possible influences of the Carpathian Mountains, which may act as a geographical barrier providing shelter from cold air outbreak from north and northeast of the continent.LocationRomania (45°N25°E).TaxonBirds of Prey.MethodsWe applied a GLMM to investigate the relation between continental and local climatic factors with the number of Common Buzzard observations in two regions. The first region is located inside the Carpathian Arch and the other one outside, east of this large mountains chain.ResultsThe Common Buzzard numbers wintering Eastern from the Carpathian Mountains are highly influenced by AO (Z = 2.87, p < .05%), while those wintering western are influenced by NAO (Z = 2.17, p < .05%). This is the first proof of separating influences for biodiversity of AO and NAO at continental scale, outlining the influence limit placed over the Eastern Carpathian Mountains.Main conclusionsThe Carpathian Mountains act like a geographic barrier, separating the wintering Common Buzzard populations from both sides of the mountain range. While the high number of individuals in Moldova is related to their eastern and northeastern Europe origins, in Transylvania the large number of individuals observed is related to the more sheltered characteristics of the region attracting individuals from central Europe. Also, since Transylvania region is well sheltered during cold air outbreak, it represents a more favorable region for wintering. From this point of view, we can consider that the Carpathian Mountains are a geographic barrier for wintering birds of prey.     

Mitochondrial phylogeography,contact zones and taxonomy of grass snakes (Natrix natrix,N. megalocephala)

Grass snakes (Natrix natrix) represent one of the most widely distributed snake species of the Palaearctic region, ranging from the North African Maghreb region and the Iberian Peninsula through most of Europe and western Asia eastward to the region of Lake Baikal in Central Asia. Within N. natrix, up to 14 distinct subspecies are regarded as valid. In addition, some authors recognize big‐headed grass snakes from western Transcaucasia as a distinct species, N. megalocephala. Based on phylogenetic analyses of a 1984‐bp‐long alignment of mtDNA sequences (ND4+tRNAs, cyt b) of 410 grass snakes, a nearly range‐wide phylogeography is presented for both species. Within N. natrix, 16 terminal mitochondrial clades were identified, most of which conflict with morphologically defined subspecies. These 16 clades correspond to three more inclusive clades from (i) the Iberian Peninsula plus North Africa, (ii) East Europe and Asia and (iii) West Europe including Corso‐Sardinia, the Apennine Peninsula and Sicily. Hypotheses regarding glacial refugia and postglacial range expansions are presented. Refugia were most likely located in each of the southern European peninsulas, Corso‐Sardinia, North Africa, Anatolia and the neighbouring Near and Middle East, where the greatest extant genetic diversity occurs. Multiple distinct microrefugia are inferred for continental Italy plus Sicily, the Balkan Peninsula, Anatolia and the Near and Middle East. Holocene range expansions led to the colonization of more northerly regions and the formation of secondary contact zones. Western Europe was invaded from a refuge within southern France, while Central Europe was reached by two distinct range expansions from the Balkan Peninsula. In Central Europe, there are two contact zones of three distinct mitochondrial clades, and one of these contact zones was theretofore completely unknown. Another contact zone is hypothesized for Eastern Europe, which was colonized, like north‐western Asia, from the Caucasus region. Further contact zones were identified for southern Italy, the Balkans and Transcaucasia. In agreement with previous studies using morphological characters and allozymes, there is no evidence for the distinctiveness of N. megalocephala. Therefore, N. megalocephala is synonymized with N. natrix.     

Morphogenetic diversity of the ectomycorrhizal genus <Emphasis Type="Italic">Cortinarius</Emphasis> section <Emphasis Type="Italic">Calochroi</Emphasis> in the Iberian Peninsula

The taxonomic and genetic diversity of Cortinarius section Calochroi in one of the most biodiversity-rich regions in Europe, the Iberian Peninsula, was investigated through morphological and phylogenetic methods. This combined methodological approach allowed the identification of 15 known species and one new species, Cortinarius ortegae, which is described here. A dichotomous key is provided for field recognition of Calochroi species inhabiting different Iberian and Balearic Island woodlands. Polymorphism analyses within some studied species showed that the distribution of intraspecific lineages is dependent on geography and, in some cases, tree host. Furthermore, the dating analysis suggested that diversification within Calochroi started in the Pliocene and most of the current genetic diversity originated in the Pleistocene. This temporal scenario supports hypotheses in which climatic oscillations in the Quaternary may have been driving the evolution of this group of ectomycorrhizal fungi.     

European wildcat populations are subdivided into five main biogeographic groups: consequences of Pleistocene climate changes or recent anthropogenic fragmentation?

Extant populations of the European wildcat are fragmented across the continent, the likely consequence of recent extirpations due to habitat loss and over‐hunting. However, their underlying phylogeographic history has never been reconstructed. For testing the hypothesis that the European wildcat survived the Ice Age fragmented in Mediterranean refuges, we assayed the genetic variation at 31 microsatellites in 668 presumptive European wildcats sampled in 15 European countries. Moreover, to evaluate the extent of subspecies/population divergence and identify eventual wild × domestic cat hybrids, we genotyped 26 African wildcats from Sardinia and North Africa and 294 random‐bred domestic cats. Results of multivariate analyses and Bayesian clustering confirmed that the European wild and the domestic cats (plus the African wildcats) belong to two well‐differentiated clusters (average Ф_ST = 0.159, R_st  = 0.392, P > 0.001; Analysis of molecular variance [AMOVA]). We identified from c. 5% to 10% cryptic hybrids in southern and central European populations. In contrast, wild‐living cats in Hungary and Scotland showed deep signatures of genetic admixture and introgression with domestic cats. The European wildcats are subdivided into five main genetic clusters (average Ф_ST = 0.103, R_st  = 0.143, P > 0.001; AMOVA) corresponding to five biogeographic groups, respectively, distributed in the Iberian Peninsula, central Europe, central Germany, Italian Peninsula and the island of Sicily, and in north‐eastern Italy and northern Balkan regions (Dinaric Alps). Approximate Bayesian Computation simulations supported late Pleistocene–early Holocene population splittings (from c. 60 k to 10 k years ago), contemporary to the last Ice Age climatic changes. These results provide evidences for wildcat Mediterranean refuges in southwestern Europe, but the evolution history of eastern wildcat populations remains to be clarified. Historical genetic subdivisions suggest conservation strategies aimed at enhancing gene flow through the restoration of ecological corridors within each biogeographic units. Concomitantly, the risk of hybridization with free‐ranging domestic cats along corridor edges should be carefully monitored.     

Late-glacial chironomid-based temperature reconstructions for Lago Piccolo di Avigliana in the southwestern Alps (Italy)

Chironomid headcapsules were used to reconstruct late glacial and early-Holocene summer temperatures at Lago Piccolo di Avigliana (LPA). Two training sets (northern Sweden, North America) were used to infer temperatures. The reconstructed patterns of temperature change agreed well with the GRIP and NGRIP δ¹⁸O records. Inferred temperatures were high during the Bølling (ca 19 °C), slowly decreased to ca 17.5 °C during the Allerød, reached lowest temperatures (ca 16 °C) during the Younger Dryas, and increased to ca. 18.5 °C during the Preboreal. The amplitudes of change at climate transitions (i.e. Oldest Dryas/Bølling: 3 °C, Allerød/Younger Dryas: 1.5 °C, and Younger Dryas/Preboreal: 2.5 °C) were smaller than in the northern Alps but similar to those recorded at another site in northeastern Italy. Our results suggest that (1) Allerød temperatures were higher in the southern Alps and (2) higher during the Preboreal (1 °C) than during the Allerød. These differences might provide an explanation for the different responses of terrestrial-vegetation to late glacial and early-Holocene climatic changes in the two regions. Other sites on both sides of the Alps should be studied to confirm these two hypotheses.     

Higher genetic diversity in recolonized areas than in refugia of Alnus glutinosa triggered by continent‐wide lineage admixture

Genetic admixture is supposed to be an important trigger of species expansions because it can create the potential for selection of genotypes suitable for new climatic conditions. Up until now, however, no continent‐wide population genetic study has performed a detailed reconstruction of admixture events during natural species expansions. To fill this gap, we analysed the postglacial history of Alnus glutinosa, a keystone species of European swamp habitats, across its entire distribution range using two molecular markers, cpDNA and nuclear microsatellites. CpDNA revealed multiple southern refugia located in the Iberian, Apennine, Balkan and Anatolian Peninsulas, Corsica and North Africa. Analysis of microsatellites variation revealed three main directions of postglacial expansion: (i) from the northern part of the Iberian Peninsula to Western and Central Europe and subsequently to the British Isles, (ii) from the Apennine Peninsula to the Alps and (iii) from the eastern part of the Balkan Peninsula to the Carpathians followed by expansion towards the Northern European plains. This challenges the classical paradigm that most European populations originated from refugial areas in the Carpathians. It has been shown that colonizing lineages have met several times and formed secondary contact zones with unexpectedly high population genetic diversity in Central Europe and Scandinavia. On the contrary, limited genetic admixture in southern refugial areas of A. glutinosa renders rear‐edge populations in the Mediterranean region more vulnerable to extinction due to climate change.     

Les industries lithiques moustériennes de la Baume Moula-Guercy (Soyons,Ardèche). Fouilles 1993–1999

The Moula-Guercy cave is located in southeastern France in the middle part and on the west bank of the Rhone valley, 80 meters above the Pleistocene Rhone River between the Ardeche Mountains and the western part of the Alps. Excavations spanning from 1993–1999 have yielded 2595 lithic artefacts out of which over 92% are coming from the four main layers (lV, VIII, XIV and XV) corresponding to the seasonal hunting site. The stratigraphic sequence is exclusively Middle Paleolithic. Paleontological and geological analysis led to the division of the sequence into 19 layers grouped into three climatic phases corresponding to isotope stages 4 to 6. The layer XV unearthed the remains of 6 cannibalized Neanderthals: children, adolescents, and adults. The study of the lithic assemblages highlights two distinct sets corresponding to layers IV–VIII and XIV–XV. This probably indicates that the same territory was travelled by two human groups with different technical traditions ten thousands years apart in very different climatic and ecological contexts. This study is also questioning whether the discoid debitage (sensu Bordes) should be part of the Levallois philosophy.     

Croatian mayflies (Insecta,Ephemeroptera): species diversity and distribution patterns

Knowledge of the mayfly biodiversity in the Balkan Peninsula is still far from complete. Compared to the neighbouring countries, the mayfly fauna in Croatia is very poorly known. Situated at the crossroads of central and Mediterranean Europe and the Balkan Peninsula, Croatia is divided into two ecoregions: Dinaric western Balkan and Pannonian lowland. Mayflies were sampled between 2003 and 2013 at 171 sites, and a total of 66 species was recorded. Combined with the literature data, the Croatian mayfly fauna reached a total of 79 taxa. Of these, 29 species were recorded for the first time in Croatia while 15 species were not previously recorded in Dinaric western Balkan ecoregion. Based on the mayfly assemblage, sampling sites were first structured by ecoregion and then by habitat type. In comparison with the surrounding countries, the Croatian mayfly fauna is the most similar to the Hungarian and Bosnian fauna. Some morphologically interesting taxa such as Baetis cf. nubecularis Eaton, 1898 and Rhithrogena from the diaphana group were recorded. Ephemera cf. parnassiana Demoulin, 1958, the species previously recorded only from Greece, was also recorded.     

Immunoglobulin genes in Andalusia (Spain). Genetic diversity in the Mediterranean space

Andalusia is the most densely populated region of Spain since ancient times, and has a rich history of contacts across the Mediterranean. Earlier studies have underlined the relatively high frequency of the Sub-Saharan GM 1,17 5* haplotype in western Andalusia (Huelva province, n = 252) and neighbouring Atlantic regions. Here, we provide novel data on GM/KM markers in eastern Andalusians (n = 195) from Granada province, where African GM*1,17 5* frequency is relatively high (0.044). The most frequent GM haplotypes in Andalusia parallel the most common in Europe. Altogether, these data allow us to gain insight into the genetic diversity of southern Iberia. Additionally, we assess population structure by comparing our Iberian samples with 41 Mediterranean populations. GM haplotype variation across the Mediterranean reflects intense and complex interactions between North Africans and South Europeans along human history, highlighting that African influence over the Iberian Peninsula does not follow an isotropic pattern.     

Temporal dynamics of ectomycorrhizal fungal communities in Shorea siamensis forest fragments

Investigating the dynamics of ectomycorrhizal fungal (EMF) communities in seasonally dry tropical forests is essential for sustainable management and for understanding the resilience of this forest type in future climate change scenarios. EMF communities in secondary forest fragments with Shorea siamensis as a single host tree species in central Thailand were sampled seasonally for 2.5 y. Ten EMF taxa were identified from ectomycorrhizal morphotypes, with/tomentella-thelephora and/russula-lactarius as the dominant taxa. Seasonal differences in EMF diversity were not detected; the dominant morphotypes were present in both seasons and their abundance varied. Most EMF taxa exhibited wide environmental ranges and only a few taxa were correlated with soil moisture. Seasonal dynamics of ectomycorrhizal colonization was likely influenced by climatic factors and the phenology of host species. Together, these results suggested that climatic variation may have a long-term and subtle influence on the composition of ectomycorrhizal communities.     

Reynoutria niche modelling and protected area prioritization for restoration and protection from invasion: A Southeastern Europe case study

An important step in managing invasive species is determining the factors responsible for their current and potential distribution, especially when species distribution is climatically determined like in Reynoutria taxa case. The main aim of this paper was to integrate all available distribution data of the Reynoutria taxa in SE Europe and to predict in which habitats and along which corridors its future spread can be expected. Distribution data of the Reynoutria taxa were obtained from extensive field studies in the period from 2006 to 2016, as well as from the literature and herbarium sources. A total of 4081 localities in Serbia and the Kosovo region, Montenegro, Slovenia, Croatia, Bosnia and Herzegovina, Bulgaria and Romania have been recorded, most frequently in riparian and human-made habitats. Ecologically suitable ranges for the Reynoutria taxa were predicted using the MaxEnt program and 19 bioclimatic variables which were derived from the WorldClim data set. Results of this study showed that the most frequent Reynoutria taxa in SE Europe are R. japonica var. japonica and hybrid R. × bohemica. The most suitable geographical areas for R. japonica and R. sachalinensis are mostly located in the north, while for R. × bohemica the central part of SE Europe is suitable. The precipitation of the warmest quarter had a high influence on the potential distribution modelling of these closely related species. Potential distribution modelling revealed R. × bohemica’s high temperature tolerance in different seasons with high durability of drought (up to 60 mm per year less precipitation than parental species). Moreover, this research predicts which areas are most likely to be invaded, and makes suggestions where to focus management efforts for survey and removal of these invaders, especially in protected areas. According to the future climate analysis R. japonica and R. × bohemica could expand their range in riparian habitats up to 30–40%. Consequently, invasion in Slovenia’s and Croatia’s rivers requires urgent preventative measures. This study provides empirical evidence that these plants could become widespread throughout SE Europe within 25 years under the current trend and emphasizes that regionally specific management plans should be implemented to prevent further spread of these taxa.     

The bat fauna hibernating in the caves of the Polish Tatra Mountains, and its long-term changes

We monitored bats hibernating in the Tatra Mountains during winters between 1997 and 2012. The Tatras are Central Europe’s second-highest massif after the Alps. Our winter censuses identified 14 species of bats hibernating in caves of the Polish Tatras. The most characteristic features of these winter bat assemblages were the dominance of Myotis mystacinus and high numbers of Eptesicus nilssonii. During the monitoring period, we noted qualitative and quantitative changes in the hibernating bat fauna. Two thermophilous species not recorded earlier and absent during the entire Holocene appeared: Rhinolophus hipposideros and Myotis emarginatus. The abundance of M. mystacinus, M. daubentonii, E. nilssonii and Plecotus auritus increased. We found no such changes in the abundance of M. myotis or M. nattereri. The Tatra Mountains are a key region for the occurrence of bats of the mystacinus group (particularly M. mystacinus sensu stricto) in Europe, and for E. nilssonii an important region in Central Europe.     

The growth-ring variations of alpine shrub Rhododendron przewalskii reflect regional climate signals in the alpine environment of Miyaluo Town in Western Sichuan Province,China

Rhododendron przewalskii is an important dwarf shrub species in alpine environments of western Sichuan that offers a unique opportunity to expand current dendrochronological networks into extreme environments beyond the survival limit of trees. Our objectives in this study are to evaluate the dendroclimatological potentials of R. przewalskii and determine the major limiting climate factor for the species’ growth rings. We sampled 25 cross-sections of R. przewalskii at an elevation of 4050 m, about 150 m above treeline, on Zhegu Mountain of Miyaluo in western Sichuan of southeastern Tibetan Plateau. R. przewalskii has well-defined growth rings such that most stem sections could be cross-dated. The resulting 61-year long standard chronology (A.D. 1949–2009) was derived from 38 series from 19 cross-sections. Response analysis revealed that radial growth of the Zhegu Mountain R. przewalskii is significantly and negatively correlated to late winter temperature (January–February), mainly driven by maximum temperatures. This correlation indicates that colder daytime temperatures during late-winter lead to improved growth the following growing season. Minimum winter temperatures do not appear important for radial growth of this population. R. przewalskii ring widths are also strongly correlated to late-winter maximum temperatures over the southeastern Tibetan Plateau region. April maximum and July minimum temperatures are positively and significantly correlated to radial growth suggesting that warm April days and warm July nights promote current-season radial growth. In contrast, radial growth is only negatively correlated to April precipitation, indicating that wet soil conditions inhibit total radial increment. The main differences of R. przewalskii compared to tree species living at high-altitude nearby regions is that R. przewalskii has a weaker positive growth response to summer temperature and winter minimum temperature, but a much stronger negative response to winter temperature. Due to the strong climatic signal recorded in the growth curves of R. przewalskii, this dwarf shrub should be useful for climate–growth studies in alpine regions where no forests are present.     

Effects of host abundance on larch budmoth outbreaks in the European Alps

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Production dynamics of Cenococcum geophilum ectomycorrhizas in response to long-term elevated CO2 and N fertilization

Ectomycorrhizal fungi are important in many forest ecosystems, yet their production dynamics and responses to environmental changes are poorly understood. Cenococcum geophilum is a common ectomycorrhizal fungus important to plant and forest soil biogeochemical cycles. The seasonal and inter-annual patterns of production and persistence of mycorrhizas formed by C. geophilum in a pine forest exposed to elevated atmospheric CO₂ and nitrogen fertilization were monitored using a 12 y minirhizotron dataset. Production of C. geophilum mycorrhizas was distinctly seasonal and peaked in late summer/autumn. Elevated CO₂ generally increased production while nitrogen fertilization strongly decreased production. Persistence times of C. geophilum mycorrhizas was ca. 2.7 y and was unaffected by CO₂ and nitrogen addition. Total production was greater in shallow soil (0–16 cm) but persistence was longer in deeper soil (17–32 cm). These observations provide insights into the autecology of C. geophilum and suggest that its tissues may be slow to decompose compared to other ectomycorrhizal species.