Evaluation of the role of the national parks for Persian leopard (<Emphasis Type="Italic">Panther pardus saxicolor</Emphasis>, Pocock 1927) habitat conservation (case study: Tandooreh National Park,Iran)

Having knowledge of the habitat requirements of top predators is important for their conservation and also for the stability of wildlife communities since they are keystone species of ecosystems. This study aims to predict suitable habitats for the Persian leopard (Panther pardus saxicolor, Pocock 1927) in Tandooreh National Park, Iran. This species is the largest leopard subspecies in the Middle East and is at high risk of extinction. Presence points for the species have been obtained from field data, literature review, atlas data, and IUCN records. Variables related to human disturbance, terrain, land use/land cover, and the leopards’ prey were used to predict the habitat selection of the Persian leopard by employing binary logistic regression. Our model predicted that the presence of Persian leopards is restricted to the southern, north-western, and western areas of the park which are under severe fragmentation threat. Moreover, leopards avoided areas near human development such as roads. We have also found that the availability of prey such as Capra aegagrus, Ovis vignei, and Ovis orientalis is one of the important parameters to predict habitat suitability for leopards. Our model provides a suitable tool for national park managers to identify biodiversity hotspots since leopards are often associated with high biodiversity.     

Water balance in Collembola and its relation to habitat selection; cuticular water loss and water uptake

In the order Collembola a clear relationship was found between overall cuticular water loss and water conditions of the habitat. The different transpiration rates were negatively correlated with the haemolymph osmotic pressure, but there was no clear causal relationship. In two species, Orchesella cincta and Tomocerus minor, which live sympatric but have a different micro-distribution (partly due to small scale heterogeneity in water conditions), important differences exist both in rate of water loss and in speed of water uptake: Orchesella cincta had a significantly lower transpiration and a higher speed of water uptake than Tomocerus minor. The transpiration rates of both species were linearly related to the vapour pressure deficit of the ambient air. Contrary to Orchesella cincta, Tomocerus minor lost water in saturated conditions. Freshly-killed Orchesella cincta had a higher transpiration rate than living individuals, but in Tomocerus minor there was no such difference. It is suggested that the main integumentary resistance against water loss in Orchesella cincta is the epidermal cell and in Tomocerus minor the epicuticle. The important rôle of the ventral vesicles in the water relations of Collembola was confirmed.     

Le genre Postepithyris makridin (térébratulidés,brachiopodes) dans l'oxfordien et le kimméridgien Français

The questions relating to the type-species of genus PostepithyrisMakridin and to its systematic position within Terebratulids are discussed.The French species are described: besides the type-species P. cincta (Cotteau) from Upper Oxfordian, two species from Inferior Kimmeridgian can be included in this genus: P. minor (Douvillé) and P. dubisensis nov. sp.A new subspecies P. cincta mosensis nov., recently discovered in Oxfordian from East of the Paris Basin, can also be assigned to the genus.Postepithyris did not evolve through a single special facies: the species could supported different environmental conditions.     

The structure of biogenic habitat and epibiotic assemblages associated with the global invasive kelp <Emphasis Type="Italic">Undaria pinnatifida</Emphasis> in comparison to native macroalgae

Kelp forests dominate temperate and polar rocky coastlines and represent critical marine habitats because they support elevated rates of primary and secondary production and high biodiversity. A major threat to the stability of these ecosystems is the proliferation of non-native species, such as the Japanese kelp Undaria pinnatifida (‘Wakame’), which has recently colonised natural habitats in the UK. We quantified the abundance and biomass of U. pinnatifida on a natural rocky reef habitat over 10 months to make comparisons with three native canopy-forming brown algae (Laminaria ochroleuca, Saccharina latissima, and Saccorhiza polyschides). We also examined the biogenic habitat structure provided by, and epibiotic assemblages associated with, U. pinnatifida in comparison to native macroalgae. Surveys conducted within the Plymouth Sound Special Area of Conservation indicated that U. pinnatifida is now a dominant and conspicuous member of kelp-dominated communities on natural substrata. Crucially, U. pinnatifida supported a structurally dissimilar and less diverse epibiotic assemblage than the native perennial kelp species. However, U. pinnatifida-associated assemblages were similar to those associated with Saccorhiza polyschides, which has a similar life history and growth strategy. Our results suggest that a shift towards U. pinnatifida dominated reefs could result in impoverished epibiotic assemblages and lower local biodiversity, although this could be offset, to some extent, by the climate-driven proliferation of L. ochroleuca at the poleward range edge, which provides complex biogenic habitat and harbours relatively high biodiversity. Clearly, greater understanding of the long-term dynamics and competitive interactions between these habitat-forming species is needed to accurately predict future biodiversity patterns.     

European hare (Lepus europaeus) invasion ecology: implication for the conservation of the endemic Irish hare (Lepus timidus hibernicus)

European hare Lepus europaeus populations have undergone recent declines but the species has successfully naturalised in many countries outside its native range. It was introduced to Ireland during the mid-late nineteenth century for field sport and is now well established in Northern Ireland. The native Irish hare Lepus timidus hibernicus is an endemic subspecies of mountain hare L. timidus and has attracted major conservation concern following a long-term population decline during the twentieth century and is one of the highest priority species for conservation action in Ireland. Little is known about the European hare in Ireland or whether it poses a significant threat to the native mountain hare subspecies by compromising its ecological security or genetic integrity. We review the invasion ecology of the European hare and examine evidence for interspecific competition with the mountain hare for habitat space and food resources, interspecific hybridisation, disease and parasite transmission and possible impacts of climate change. We also examine the impact that introduced hares can have on native non-lagomorph species. We conclude that the European hare is an emerging and significant threat to the conservation status of the native Irish hare. Invasive mammal species have been successfully eradicated from Ireland before and immediate action is often the only opportunity for cost-effective eradication. An urgent call is issued for further research whilst the need for a European hare invasive Species Action Plan (iSAP) and Eradication strategy are discussed.     

The Value of Learning about Natural History in Biodiversity Markets

Markets for biodiversity have generated much controversy because of the often unstated and untested assumptions included in transactions rules. Simple trading rules are favored to reduce transaction costs, but others have argued that this leads to markets that favor development and erode biodiversity. Here, I describe how embracing complexity and uncertainty within a tradable credit system for the Red-cockaded Woodpecker (Picoides borealis) creates opportunities to achieve financial and conservation goals simultaneously. Reversing the effects of habitat fragmentation is one of the main reasons for developing markets. I include uncertainty in habitat fragmentation effects by evaluating market transactions using five alternative dispersal models that were able to approximate observed patterns of occupancy and movement. Further, because dispersal habitat is often not included in market transactions, I contrast how changes in breeding versus dispersal habitat affect credit values. I use an individually-based, spatially-explicit population model for the Red-cockaded Woodpecker (Picoides borealis) to predict spatial- and temporal- influences of landscape change on species occurrence and genetic diversity. Results indicated that the probability of no net loss of abundance and genetic diversity responded differently to the transient dynamics in breeding and dispersal habitat. Trades that do not violate the abundance cap may simultaneously violate the cap for the erosion of genetic diversity. To highlight how economic incentives may help reduce uncertainty, I demonstrate tradeoffs between the value of tradable credits and the value of information needed to predict the influence of habitat trades on population viability. For the trade with the greatest uncertainty regarding the change in habitat fragmentation, I estimate that the value of using 13-years of data to reduce uncertainty in dispersal behaviors is $6.2 million. Future guidance for biodiversity markets should at least encourage the use of spatially- and temporally-explicit techniques that include population genetic estimates and the influence of uncertainty.     

Restoration rocks: integrating abiotic and biotic habitat restoration to conserve threatened species and reduce fire fuel load

With rapid urban expansion, biodiversity conservation and human asset protection often require different regimes for managing wildfire risk. We conducted a controlled, replicated experiment to optimise habitat restoration for the threatened Australian pink-tailed worm-lizard, Aprasia parapulchella while reducing fire fuel load in a rapidly developing urban area. We used dense addition of natural rock (30 % cover) and native grass revegetation (Themeda triandra and Poa sieberiana) to restore critical habitat elements. Combinations of fire and herbicide (Glyphosate) were used to reduce fuel load and invasive exotic species. Rock restoration combined with herbicide application met the widest range of restoration goals: it reduced fire fuel load, increased ant occurrence (the primary prey of A. parapulchella) in the short-term and increased the growth and survival of native grasses. Lizards colonised the restored habitat within a year of treatment. Our study documents an innovative way by which conflicts between biodiversity conservation and human asset protection can be overcome.     

Biodiversity potential of <Emphasis Type="Italic">Nothofagus</Emphasis> forests in Tierra del Fuego (Argentina): tool proposal for regional conservation planning

It is difficult to map and quantify biodiversity at landscape level in areas with low data availability, despite demand from decision-makers. We propose a methodology to determine potential biodiversity pattern using habitat suitability maps of the understory plant species with highest cover and occurrence frequency in the three different forests types of Tierra del Fuego (Argentina). We used a database of vascular plants from 535 surveys from which we identified 35 indicative species. We explored more than 50 potential explanatory variables to develop habitat suitability maps of the indicative species, which were combined to develop a map of the potential biodiversity. Correlation among environmental, topographic and forest landscape variables were discussed, as well as the marginality and the specialization of the indicative species. We detected differences in the niches of the species prevailing in the three forest types. The developed map of potential biodiversity uncovered hotspots of biodiversity in the ecotone of Nothofagus pumilio and N. antarctica as well as in the wettest part of the mixed N. pumilio–N. betuloides forests. It allowed thus to identify forest areas with different conservation potential and can be readily used as a decision support system for conservation and management strategies at different scales including the identification of land-use conflicts (e.g. of biodiversity with timber production and livestock) and the development of a network of protected areas, which currently does not cover the forests of highest conservation value.     

Impacts of onshore wind energy production on birds and bats: recommendations for future life cycle impact assessment developments

Purpose

Models for quantifying impacts on biodiversity from renewable energy technologies are lacking within life cycle impact assessment (LCIA). We aim to provide an overview of the effects of wind energy on birds and bats, with a focus on quantitative methods. Furthermore, we investigate and provide the necessary background for how these can be integrated into new developments of LCIA models in future.

Methods

We reviewed available literature summarizing the effects of wind energy developments on birds and bats. We provide an overview of available quantitative assessment methods that have been employed outside of the LCIA framework to model the different impacts of wind energy developments on wildlife. Combining the acquired knowledge on impact pathways and associated quantitative methods, we propose possibilities for future approaches for a wind energy impact assessment methodology for LCIA.

Results and discussion

Wind energy production has impacts on terrestrial biodiversity through three main pathways: collision, disturbance, and habitat alterations. Birds and bats are consistently considered the most affected taxonomic groups, with different responses to the before-mentioned impact pathways. Outside of the LCIA framework, current quantitative impact assessment prediction models include collision risk models, species distribution models, individual-based models, and population modeling approaches. Developed indices allow scaling of species-specific vulnerability to mortality, disturbance, and/or habitat alterations.

Conclusions

Although insight into the causes behind collision risk, disturbance, and habitat alterations for bats and birds is still limited, the current knowledge base enables the development of a robust assessment tool. Modeling the impacts of habitat alterations, disturbance, and collisions within an LCIA framework is most appropriate using species distribution models as those enable the estimation of species’ occurrences across a region. Although local-scale developments may be more readily feasible, further up-scaling to global coverage is recommended to allow comparison across regions and technologies, and to assess cumulative impacts.

     

Urbanization Impacts on Mammals across Urban-Forest Edges and a Predictive Model of Edge Effects

With accelerating rates of urbanization worldwide, a better understanding of ecological processes at the wildland-urban interface is critical to conserve biodiversity. We explored the effects of high and low-density housing developments on forest-dwelling mammals. Based on habitat characteristics, we expected a gradual decline in species abundance across forest-urban edges and an increased decline rate in higher contrast edges. We surveyed arboreal mammals in sites of high and low housing density along 600 m transects that spanned urban areas and areas turn on adjacent native forest. We also surveyed forest controls to test whether edge effects extended beyond our edge transects. We fitted models describing richness, total abundance and individual species abundance. Low-density housing developments provided suitable habitat for most arboreal mammals. In contrast, high-density housing developments had lower species richness, total abundance and individual species abundance, but supported the highest abundances of an urban adapter (Trichosurus vulpecula). We did not find the predicted gradual decline in species abundance. Of four species analysed, three exhibited no response to the proximity of urban boundaries, but spilled over into adjacent urban habitat to differing extents. One species (Petaurus australis) had an extended negative response to urban boundaries, suggesting that urban development has impacts beyond 300 m into adjacent forest. Our empirical work demonstrates that high-density housing developments have negative effects on both community and species level responses, except for one urban adapter. We developed a new predictive model of edge effects based on our results and the literature. To predict animal responses across edges, our framework integrates for first time: (1) habitat quality/preference, (2) species response with the proximity to the adjacent habitat, and (3) spillover extent/sensitivity to adjacent habitat boundaries. This framework will allow scientists, managers and planners better understand and predict both species responses across edges and impacts of development in mosaic landscapes.     

Assessment of Habitat Fragmentation and Corridors for an Isolated Subspecies of the Sichuan Golden Snub-Nosed Monkey, <Emphasis Type="Italic">Rhinopithecus roxellana hubeiensis</Emphasis>

Understanding habitat quality and landscape connectivity and exploring corridors connecting habitat patches are crucial for conservation, particularly for species distributed among isolated populations. The Sichuan golden snub-nosed monkey, Rhinopithecus roxellana, is an Endangered primate species endemic to mountainous forests in China. Its easternmost distribution lies in the Shennongjia area, which harbors an isolated subspecies, R. roxellana hubeiensis. Unfortunately, it has experienced significant habitat loss, fragmentation, and dramatic population decline in recent decades, primarily due to increased human disturbance. To quantify habitat quality, identify suitable habitat patches, and detect possible linkages among these patches for R. roxellana hubeiensis, we conducted habitat suitability assessments and landscape connectivity analyses in the Shennongjia area based on a set of environmental factors. We created a habitat quality model and a movement cost surface for the Shennongjia area based on a habitat suitability index, graph theory, expert knowledge, field experience, and information from the literature. Our results show that suitable habitat for R. roxellana hubeiensis in Shennongjia is fragmented and limited, and that this is particularly true for highly suitable habitats. We detected six core habitat patches and six least-cost paths and corridors. Our study does not provide accurate distributions of the monkeys and their habitat use. However, it identifies the most feasible and traversable habitats and corridors, which should be conservation priorities for this subspecies, and provides valuable guidance for reevaluating habitat conservation plans.     

Rhinanthus minor population genetic structure and subspecies: Potential seed sources of a keystone species in grassland restoration projects

In the last few decades unimproved semi-natural grasslands have been affected by intensification of land use and habitat fragmentation. Because of their biodiversity these species-rich grasslands are of high conservation importance and efforts are under way to restore such habitats. Detailed knowledge of within species diversity will aid deciding on the optimal seed source for such restoration projects, e.g. local genotypes or ecotypes. Rhinanthus minor is a species that is typically found in semi-natural grasslands and is commonly used in grassland restoration projects. This is because R. minor is a hemiparasitic plant that takes minerals and nutrients from its host, which in turn decreases the host's biomass and leads to opportunities for less competitive species in the vegetation. Here, we investigate genetic diversity within and between R. minor populations. This allowed us to test whether the six different subspecies of R. minor that have been described in the UK, based on their morphology, flowering time, and habitat, can be differentiated using molecular markers. We identified moderate levels of genetic differentiation between R. minor populations within the UK. In addition, R. minor individuals from the UK appear to be distinct from R. minor and Rhinanthus angustifolius individuals from other European countries based on microsatellite genotyping and DNA sequencing of cpDNA and rDNA ITS. The molecular markers used in the current study did not separate populations of R. minor based on either their subspecies or habitat. The implication for the use of R. minor in grassland restoration projects seems to be that it is not necessary to use local seeds or seeds from the same subspecies.     

NEGATIVE MATERNAL EFFECT REVISITED: A TEST ON TWO POPULATIONS OF ORCHESELLA CINCTA L. (COLLEMBOLA: ENTOMOBRYIDAE)

A Dutch population of Orchesella cincta had been demonstrated to exhibit a negative maternal effect on age at first reproduction, which caused alternation of short and long generations. The adaptive significance of such a mechanism was assumed to be associated with the bivoltine life cycle of Dutch O. cincta. We expected that it would be absent in a non bivoltine population sampled in Siena, Italy. To test this hypothesis we performed a parent-offspring regression experiment with both populations simultaneously. The experiment showed that there was no negative maternal effect in both populations. We leave open the question of the cause of the discrepancy between the previous result with the Dutch population and the present result. The results of our experiment were also used to determine heritabilities of the traits age, mass and number of molts at first reproduction, and size of the first clutch. The estimates of heritabilities were often not significantly different from zero, especially in the Italian population which had only one significant heritability.     

Chimpanzee population structure in Cameroon and Nigeria is associated with habitat variation that may be lost under climate change

Background

The Nigeria-Cameroon chimpanzee (Pan troglodytes ellioti) is found in the Gulf of Guinea biodiversity hotspot located in western equatorial Africa. This subspecies is threatened by habitat fragmentation due to logging and agricultural development, hunting for the bushmeat trade, and possibly climate change. Although P. t. ellioti appears to be geographically separated from the neighboring central chimpanzee (P. t. troglodytes) by the Sanaga River, recent population genetics studies of chimpanzees from across this region suggest that additional factors may also be important in their separation. The main aims of this study were: 1) to model the distribution of suitable habitat for P. t. ellioti across Cameroon and Nigeria, and P. t. troglodytes in southern Cameroon, 2) to determine which environmental factors best predict their optimal habitats, and 3) to compare modeled niches and test for their levels of divergence from one another. A final aim of this study was to examine the ways that climate change might impact suitable chimpanzee habitat across the region under various scenarios.

Results

Ecological niche models (ENMs) were created using the software package Maxent for the three populations of chimpanzees that have been inferred to exist in Cameroon and eastern Nigeria: (i) P. t. troglodytes in southern Cameroon, (ii) P. t. ellioti in northwestern Cameroon, and (iii) P. t. ellioti in central Cameroon. ENMs for each population were compared using the niche comparison test in ENMtools, which revealed complete niche divergence with very little geographic overlap of suitable habitat between populations.

Conclusions

These findings suggest that a positive relationship may exist between environmental variation and the partitioning of genetic variation found in chimpanzees across this region. ENMs for each population were also projected under three different climate change scenarios for years 2020, 2050, and 2080. Suitable habitat of P. t. ellioti in northwest Cameroon / eastern Nigeria is expected to remain largely unchanged through 2080 in all considered scenarios. In contrast, P. t. ellioti in central Cameroon, which represents half of the population of this subspecies, is expected to experience drastic reductions in its ecotone habitat over the coming century.

Electronic supplementary material

The online version of this article (doi:10.1186/s12862-014-0275-z) contains supplementary material, which is available to authorized users.     

Developing characterisation factors for land fragmentation impacts on biodiversity in LCA: key learnings from a sugarcane case study

Purpose

Habitat change was identified by the Millennium Ecosystem Assessment as the main direct driver of biodiversity loss. However, while habitat loss is already implemented in Life Cycle Impact Assessment (LCIA) methods, the additional impact on biodiversity due to habitat fragmentation is not assessed yet. Thus, the goal of this study was to include fragmentation effects from land occupation and transformation at both midpoint and endpoint levels in LCIA.

Methods

One promising metric, combining the landscape spatial configuration with species characteristics, is the metapopulation capacity λ, which can be used to rank landscapes in terms of their capacity to support viable populations spatially structured. A methodology to derive worldwide regionalised fragmentation indexes based on λ was used and combined with the Species Fragmented-Area Relationship (SFAR), which relies on λ to assess a species loss due to fragmentation. We adapted both developments to assess fragmentation impacts due to land occupation and transformation at both midpoint and endpoint levels in LCIA. An application to sugarcane production occurring in different geographical areas, more or less sensitive to land fragmentation, was performed.

Results and discussion

The comparison to other existing LCIA indicators highlighted its great potential for complementing current assessments through fragmentation effect inclusion. Last, both models were discussed through the evaluation grid used by the UNEP-SETAC land use LCIA working group for biodiversity impact assessment models.

Conclusions

Midpoint and endpoint characterisation factors were successfully developed to include the impacts of habitat fragmentation on species in LCIA. For now, they are provided for bird species in all forest ecoregions belonging to the biodiversity hotspots. Further work is required to develop characterisation factors for all taxa and all terrestrial ecoregions.

     

Dispersal capacity and diet breadth modify the response of wild bees to habitat loss

Habitat loss poses a major threat to biodiversity, and species-specific extinction risks are inextricably linked to life-history characteristics. This relationship is still poorly documented for many functionally important taxa, and at larger continental scales. With data from five replicated field studies from three countries, we examined how species richness of wild bees varies with habitat patch size. We hypothesized that the form of this relationship is affected by body size, degree of host plant specialization and sociality. Across all species, we found a positive species–area slope (z = 0.19), and species traits modified this relationship. Large-bodied generalists had a lower z value than small generalists. Contrary to predictions, small specialists had similar or slightly lower z value compared with large specialists, and small generalists also tended to be more strongly affected by habitat loss as compared with small specialists. Social bees were negatively affected by habitat loss (z = 0.11) irrespective of body size. We conclude that habitat loss leads to clear shifts in the species composition of wild bee communities.     

The role of protected area zoning in invasive plant management

As anthropogenic pressure on the landscape increases, invasive alien species (IAS) pose a growing threat to areas designed to protect high biodiversity habitats. In order to assess the present danger of IAS spread, we examined 23 Czech sites of community importance (SCI) within Natura 2000 protected areas (PA) over 2015 and mapped the occurrence of four IAS: Solidago spp. (goldenrod), Impatiens glandulifera (Himalayan balsam), Heracleum mantegazzianum (giant hogweed) and Fallopia spp. (Japanese knotweed). The model areas were divided into five monitoring zones, graded by conservation importance and habitat disturbance level (core area [A], broader core area [B], semi-natural habitat [C], anthropogenically affected habitat [D], anthropogenically degraded habitat [E]). Despite a high number of IAS occurrences (3222 localities), habitats of European importance (zone A) showed a relatively low level of invasion (< 0.3% total area). Highest IAS occurrence number was in SCI border areas and disturbed habitats (zones C and E). There was a significant positive correlation between level of invasion inside and outside SCIs, related to human activities such as logging and urbanisation. A strong effect for watercourse vicinity was noted for the occurrence of I. glandulifera and Fallopia spp.; but not for H. mantegazzianum and Solidago spp. A stratified management approach, employing zones delimitation to assess what threat pose IAS to the PA objects of conservation, can be useful to prioritize control measures in IAS local action plans.     

Rural avenues as a refuge for feral honey bee population

Several honey bee (Apis mellifera) subspecies are in danger of local extinction because their feral population have almost completely disappeared. An important threat to the feral populations of bees is loss of habitat and loss of woodlands. In many places the only habitat suitable for honey bee nesting are rows of trees along roadsides. We studied a feral population of honey bees inhabiting avenues in northern Poland. We inspected 142 km of avenues and found 45 feral colonies. The estimated density of feral population inhabiting the avenues was 0.10 nest km^?2. Honey bees preferred to build their nests in trees with a thick trunk and a somewhat weak state of health. There was no strong preference of bees to any species of trees. We stress the importance of protection of existing avenues and creating new ones. This can provide suitable habitat not only for honey bees but also for other endangered species.     

Isozyme diversity in some Canarian woody endemisms of the genus Echium L. (Boraginaceae)

Oceanic archipelagos are considered hot-spots of biodiversity because they harbor unique biota with a high level of endemicity. However, the endemic biodiversity of oceanic islands is very vulnerable to extinction. In recent decades, intensive exploitation of these territories and human-mediated introduction of alien species have posed unprecedented threats to the long-term survival of the endemic contingent. The very limited population genetic information available until now for the 28 Canarian endemic Echium taxa has hindered the development of conservation strategies for the ca. 25% of them that are under threat. In this paper, we analyze the levels and distribution of genetic isozyme diversity in 23 natural populations of three endangered endemics of restricted distribution (E. acanthocarpum, CR; E. onosmifolium ssp. spectabile, EN; and E. callithyrsum, VU), and two endemics of wide distribution and in principle free of threat (E. decaisnei and E. onosmifolium). Our results reveal high levels of genetic variability in all these taxa that have plausibly been reached despite a predominance of selfing. They also point out a high incidence of inbreeding in the reproductive dynamics of populations and suggest the potential value of hybridisation processes in shaping the genetic makeup of these species. Among-population differentiation, as estimated by Gst, and genetic distances within taxa are low overall, but they do support the current taxonomic separation between the two subspecies of E. onosmifolium and do not furnish evidence that the current status of the endangered species may be attributed to genetic factors. We use the genetic parameters to suggest some guidelines to help implement a conservation strategy of these taxa.