Effectiveness of habitat management in the recovery of low-density populations of wild rabbit

Understanding the relationship between spatial patterns of landscape attributes and population presence and abundance is essential for understanding population processes as well as supporting management and conservation strategies. This study evaluates the influence of three factors: environment, habitat management, and season on the presence and abundance of the wild rabbit (Oryctolagus cuniculus), an important prey species for Mediterranean endangered predator species. To address this issue, we estimated wild rabbit presence and abundance by latrine counting in transects located in 45 plots within a 250?×?250 m grid from June 2007 until June 2009 in a 1,200 ha hunting area in southern Portugal. We then analyzed how wild rabbit presence and abundance correlate with the aforementioned factors. Our results showed that the main variable influencing wild rabbit presence and abundance was the distance to the artificial warrens. North and northeast slope directions were negatively related to wild rabbit presence. Conversely, rabbit presence was positively correlated with short distances to ecotone, artificial warrens, and spring. Regarding rabbit abundance, in addition to artificial warrens, soft soils, bushes, and season also had a positive effect. We found that environmental variables, management practices, and season each affect wild rabbit presence and abundance differently at a home range scale in low-density population. Thus, our major recommendations are reducing the distance to artificial warrens and ecotone, ideally to less than 100 m, and promoting habitat quality improvement on slopes with plenty of sun exposure.     

Provision of artificial warrens as a means to enhance native wild rabbit populations: what type of warren and where should they be sited?

In Spain, wild rabbits are essential for some highly endangered species, and, therefore, many actions have been undertaken to increase their populations. In the present study, artificial warrens are provided as a means to increase shelter for native wild rabbit populations in a given area. We evaluate the use of three types of warrens by rabbits and the effect on that use of five habitat characteristics at two spatial scales (500 × 500-m grids and 25-m plots). To evaluate that use, we identified pre-established signs at the entrances to each warren, and based on this, we calculated occupancy rate and activity. Our results indicate that rabbit abundance within a grid is the only variable which simultaneously explains both the greater occupancy and the higher activity in the artificial warrens located in that grid. Some 73.2% of the grids showed signs of rabbit use at the time of the evaluation. However, the pre-existing rabbit populations within the grids were not quantified and, hence, we cannot state that the warrens contributed to an increase in the rabbit abundance. Regarding the habitat, our results reveal that warrens should be situated in grids with food coverage of less than 50%, while the use of each individual refuge is greater where food availability in the immediate surroundings is at least 20% and shelter at least 50%. The tube warrens showed significantly greater rabbit use than the other types while there was little difference between the stone and pallet warrens in terms of use.     

Potential Component Allee Effects and Their Impact on Wetland Management in the Conservation of Endangered Anurans

Effective management of wetland quantity and quality is crucial for effective conservation of declining amphibian populations. In particular, frogs and toads that employ aggregative breeding strategies may suffer negative population impacts in response to changes in availability of aquatic breeding habitat, including overabundance of suitable habitat, if density of conspecifics attending aggregations is positively correlated with reproductive success. Here we document such a positive relationship, potentially the first example of a component Allee effect in an anuran, in the critically endangered Houston toad (Bufo houstonensis). We assessed the relationship between mean yearly chorus size and reproductive success of males at the pond level using an information theoretic model selection approach and a two-sample t-test. The chosen model contained the single variable of mean yearly chorus size to predict probability of reproduction, as selected using the Akaike Information Criterion corrected for small sample size and Akaike weight. Mean chorus sizes were significantly higher among ponds exhibiting evidence of reproduction than in those that showed no evidence of reproduction. Our results suggest that chorusing alone is a poor proxy for inference of population stability and highlight a need for reassessment of widely-used amphibian monitoring protocols. Further, amphibian conservation efforts should account for potential Allee effects in order to optimize benefits and avoid underestimating critical population thresholds, particularly in species exhibiting rapid population declines.     

Nest site selection and efficacy of artificial nests for breeding success of Scarlet Macaws Ara macao macao in lowland Peru

Psittacidae (parrots) have the most threatened species of any bird family in the world. Most parrots are obligate secondary cavity nesters, and can be limited in their breeding success by the availability and quality of nest hollows. However, nesting opportunities for parrots can be increased by provision of artificial nest boxes. The Tambopata Macaw Project has been studying the breeding ecology and natural history of the Scarlet Macaw Ara macao macao in the south-eastern Peruvian Amazon for over 20 years by monitoring natural nest hollows and two types of artificial nest (wooden and PVC). We present data for breeding success in natural and artificial nests over 12 consecutive breeding seasons. The aims of this study were to: (a) determine the nesting requirements and reproductive success of breeding macaws; and, (b) compare the efficacy of the two types of artificial nests and natural nest cavities. Our data showed a high rate of reoccupation of successful nests in consecutive years and that nests in artificial and natural nests had very similar reproductive parameters. Our results indicate that artificial nest types can be used by conservation managers seeking to assist A. macao populations where nest hollows are in short supply, and that artificial nests can contribute important data to natural history studies of species where access to natural nests is limited.     

Is habitat management an effective tool for wild rabbit (<Emphasis Type="Italic">Oryctolagus cuniculus</Emphasis>) population reinforcement?

Wild rabbit scarcity in Spain is a serious problem due to its economic and ecological value; thus, management techniques to reinforce their populations are being implemented. Translocations are traditionally applied but high short-term mortality by predation makes them unsuccessful and increases their biological cost. An alternative to translocation would be to strengthen local reduced populations through habitat management (building artificial warrens and increasing food availability). Here, we test this method to determine its efficiency as an exportable protocol to recover wild rabbit populations. We compare two areas with and without artificial warrens and increased food availability and assess relative rabbit abundance by means of pellet counts. Results show that, during the 2 years following habitat management, rabbit numbers increased in the experimental plot in contrast to the not managed one, suggesting its effectiveness to reinforce low-density rabbit populations.     

Cavity-nesting birds are limited by nesting habitat in Neotropical agricultural landscapes

Most studies comparing biodiversity between natural and human-modified landscapes focus on patterns in species occurrence or abundance, but do not consider how different habitat types meet species' breeding requirements. Organisms that use or nest in tree cavities may be especially threatened by habitat conversion due to the loss of their nesting sites. Although cavity-nesting bird diversity is highest in the tropics, little is known about how tropical birds use cavities, how agriculture affects their reproductive biology, and how effective nest boxes could be as a conservation strategy in tropical agriculture. Here, we explored how habitat conversion from tropical forests to pasture affects the abundance, nesting habitat availability, and nest success of cavity-nesting birds in Northwest Ecuador. We conducted bird surveys and measured natural cavity availability and use in forest and agriculture. We also added artificial nest boxes to forest and agriculture to see whether cavity limitation in agriculture would elicit higher use of artificial nest boxes. We found evidence of cavity limitation in agriculture—there were many more natural cavities in forest than in agriculture, as well as more avian use of nest boxes placed in agriculture as compared to forest. Our results suggest that it is important to retain remnant trees in tropical agriculture to provide critical nesting habitat for birds. In addition, adding nest boxes to tropical agricultural systems could be a good conservation strategy for certain species, including insectivores that could provide pest-control services to farmers. Abstract in Spanish is available with online material.     

Improving predictions of the location and use of warrens in sensitive rabbit populations

The location and use of warrens is a key factor in the population dynamics of the European rabbit Oryctolagus cuniculus L., which can influence stability and persistence in the long term. Within the species' original distribution range, its numbers have declined sharply in recent decades, which is a serious problem for the conservation of Mediterranean ecosystems. Artificial warrens are commonly used to bolster rabbit populations and the characterization of natural warrens in those areas might improve their efficiency. In this study, we use binomial generalized linear models to identify the factors associated with the location and use of warrens in a low-density area and we evaluate the effect of including the spatial structure of the data in the model. In addition, we generate a map that predicts the most suitable areas for artificial warrens. Contrary to what was expected, habitat variables are only secondary factors, and the location and use of warrens is influenced mainly by spatial factors, such as proximity to nearby warrens. Furthermore, the aggregated spatial pattern of warrens suggests that, at the local scale, for example, the hunting estate, intra-specific interactions might be playing a primary role in these low-density populations. To identify the most suitable sites for artificial warrens and, thereby, improve the efficiency of artificial warren building for conservation purposes, information about spatial structures should be included in models predicting natural warrens of European rabbits.     

Conservation of deer: contributions from molecular biology, evolutionary ecology, and reproductive physiology

Molecular phylogenetics, interspecific comparisons, and assisted reproductive techniques are recent approaches to understanding and facilitating conservation of endangered species. This paper reviews the contribution of these approaches to a small but well-studied group of mammals, deer, many of which are endangered in the wild. Conservation efforts require a comprehensive understanding of the biology and history of these animals. The value of assisted reproductive technologies for conservation of deer has received increased awareness especially for captive populations. Such breeding programmes are designed to assist propagation of threatened species and to maximize genetic diversity within populations through the movement of genetic material across the globe, but will only be successful if we understand the genetic and reproductive potential of various lineages. Here we discuss the phylogenetic status of deer, the distinctiveness and evolution of their reproductive patterns, and current approaches for improving the success of controlled breeding programmes for the conservation of endangered lineages. Only by combining both theoretical and practical approaches to conservation efforts can we hope to salvage the remaining organismal diversity of our planet.     

Breeding like rabbits: global patterns of variability and determinants of European wild rabbit reproduction

The importance of European wild rabbits Oryctolagus cuniculus both as a pest in some areas and as a key prey species in others emphasizes the need to understand what controls its population dynamics worldwide. In this study we aim to describe the variability in rabbit breeding parameters and identify the main factors that govern it at a global scale. Despite the species' wide distribution, some reproductive traits such as short sexual maturity age, duration of gestation period, and existence of post-partum oestrus are similar in all populations conferring the species a high breeding potential. Nevertheless, other aspects vary substantially among regions resulting in highly different population productivities and also across years. These latter parameters are the length of breeding season, proportion of pregnant females, age of first reproduction, and number and size of litters. Our results show that variability in these attributes is mainly affected by a combination of environmental controls (i.e. temperature, resource availability, and photoperiod) and individual properties (age and body weight). On the other hand, the effect of other factors such as population density could not be demonstrated. Knowledge about the factors driving global reproduction patterns of European wild rabbits will improve our understanding about their population dynamics, and thus will help to optimize the management and conservation of their populations.     

Competition between honey bees and wild bees and the role of nesting resources in a nature reserve

The European honey bee exploits floral resources efficiently and may therefore compete with solitary wild bees. Hence, conservationists and bee keepers are debating about the consequences of beekeeping for the conservation of wild bees in nature reserves. We observed flower-visiting bees on flowers of Calluna vulgaris in sites differing in the distance to the next honey-bee hive and in sites with hives present and absent in the Lüneburger Heath, Germany. Additionally, we counted wild bee ground nests in sites that differ in their distance to the next hive and wild bee stem nests and stem-nesting bee species in sites with hives present and absent. We did not observe fewer honey bees or higher wild bee flower visits in sites with different distances to the next hive (up to 1,229 m). However, wild bees visited fewer flowers and honey bee visits increased in sites containing honey-bee hives and in sites containing honey-bee hives we found fewer stem-nesting bee species. The reproductive success, measured as number of nests, was not affected by distance to honey-bee hives or their presence but by availability and characteristics of nesting resources. Our results suggest that beekeeping in the Lüneburg Heath can affect the conservation of stem-nesting bee species richness but not the overall reproduction either of stem-nesting or of ground-nesting bees. Future experiments need control sites with larger distances than 500 m to hives. Until more information is available, conservation efforts should forgo to enhance honey bee stocking rates but enhance the availability of nesting resources.     

Benefits of extra food to reproduction depend on maternal condition

The amount of food resources available to upper‐level consumers can show marked variations in time and space, potentially resulting in food limitation. The availability of food resources during reproduction is a key factor modulating variation in reproductive success and life‐history tradeoffs, including patterns of resource allocation to reproduction versus self‐maintenance, ultimately impacting on population dynamics. Food provisioning experiments constitute a popular approach to assess the importance of food limitation for vertebrate reproduction. In this study of a mesopredatory avian species, the lesser kestrel Falco naumanni, we provided extra food to breeding individuals from egg laying to early nestling rearing. Extra food did not significantly affect adult body condition or oxidative status. However, it increased the allocation of resources to flight feathers moult and induced females to lay heavier eggs. Concomitantly, it alleviated the costs of laying heavier eggs for females in poor body condition, and reduced their chances of nest desertion (implying complete reproductive failure). Extra food provisioning improved early nestling growth (body mass and feather development). Moreover, extra food significantly reduced the negative effects of ectoparasites on nestling body mass, while fostering forearm (a flight apparatus trait) growth among highly parasitized nestlings. Our results indicate that lesser kestrels invested the extra food mainly to improve current reproduction, suggesting that population growth in this species can be limited by food availability during the breeding season. In addition, extra food provisioning reduced the costs of the moult–breeding overlap and affected early growth tradeoffs by mitigating detrimental ectoparasite effects on growth and enhancing development of the flight apparatus with high levels of parasitism. Importantly, our findings suggest that maternal condition is a major trait modulating the benefits of extra food to reproduction, whereby such benefits mostly accrue to low‐quality females with poor body condition.     

Reproductive biology of Ranunculus weyleri (Ranunculaceae), a narrowly endemic plant from the Balearic Islands with disjunct populations

Ranunculus weyleri is a species endemic to Mallorca Island (western Mediterranean basin) that is currently threatened due to its reduced geographical distribution and disjunct, isolated populations. It is a small, perennial herb that inhabits rocky environments, together with other endemic plants with creeping life-forms. In the present study, for the first time the reproductive biology of the species was investigated in terms of its floral phenology (floral stages and times of formation), pollen/ovule (P/O) ratios, and fruit and seed setting under natural and pollen manipulation experiments, assessing the breeding system, pollination vectors and pollen limitations. Cross-pollinations among populations were performed to evaluate the levels of compatibility between populations. In addition, the floral displays and the reproductive outputs of four wild populations were determined. The observed high P/O ratios, results of the hand-pollination experiments, and flower traits indicate that this species is predominantly allogamous. Combinations of both insects (belonging to Coleoptera, Diptera, Lepidoptera and Hymenoptera) and wind (ambophily) constitute the main pollination vectors, and pollen limitation was observed. Interpopulation crossing was effective, and exogamic depression was not present, as evidenced by the fecundity levels. The predation of the floral stems by herbivores (mainly goats) severely limits the reproductive success of this endemic species in wild populations. Finally, several ways of action are proposed for R. weyleri conservation.     

Provisioning habitat with custom‐designed nest‐boxes increases reproductive success in an endangered finch

Manipulating resource availability to assess the strength and effects of resource limitation on an animal population is relatively straightforward, and thus, common in the scientific literature. Resource quality, however, is rarely manipulated, as this requires a priori knowledge of a relationship between some measurable variation in the resource and individual preference and/or fitness. Recent research on nest‐site selection was used to inform the design of custom‐built nest‐boxes for an endangered hollow‐nester, the Gouldian finch (Erythrura gouldiae). By provisioning breeding populations with these ‘high quality’ artificial nest sites over 3 years, we experimentally investigated the strength and effects of nest‐site limitation in this species. Breeding pairs using nest‐boxes initiated nesting earlier, produced larger clutches and fledged more offspring per season than those in natural hollows. Total and mean reproductive output also increased in nest‐box provisioned sites over the duration of the study. All of these effects were predominantly driven by pairs initiating clutches earlier, potentially because of the reduced intra‐ and interspecific competition for nest sites at optimal breeding times. Our findings suggest that reproduction in wild Gouldian finch populations may be limited by the availability of high quality nest sites in the landscape, and that nest‐boxes could be used as a tool for enhancing reproduction in recovering populations. Furthermore, we conclude that resource quality is equally, if not more important than quantity when supplementing resource‐limited habitats.     

From population structure to individual behaviour: genetic analysis of social structure in the European wild rabbit (Oryctolagus cuniculus)

The European wild rabbit (Oryctolagus cuniculus) lives in stable, territorial breeding groups, with male-biased natal dispersal, female philopatry, and a polygynous mating system. It was introduced into Britain in the 11 th century and kept in captive warrens as a food and fur resource. Populations expanded in the wild in the 18th century. Microsatellite markers were employed to examine the genetic structure of wild rabbit populations on three spatial scales: macrogeographic structure of 17 populations in East Anglia, microgeographic structure of a tagged population in the grounds of the University of East Anglia over four consecutive years, and pairwise kinships and individual movement within this tagged population. Populations across East Anglia were found to be genetically distinct, and heterozygote deficits were observed at all loci indicating sub-division within sampled populations. Analysis of the tagged population confirmed that rabbit populations are genetically sub-divided among social groups. Studying this population over four consecutive years revealed that as the population size increased, the number of social groups increased. Analysis of individual pairwise relatedness of females indicated that individuals did not necessarily group with kin, and behavioural data indicate that an optimum group size may exist which maximizes reproductive success.     

Burrowing by translocated boodie (Bettongia lesueur) populations alters soils but has limited effects on vegetation

Digging and burrowing mammals modify soil resources, creating shelter for other animals and influencing vegetation and soil biota. The use of conservation translocations to reinstate the ecosystem functions of digging and burrowing mammals is becoming more common. However, in an increasingly altered world, the roles of translocated populations, and their importance for other species, may be different. Boodies (Bettongia lesueur), a commonly translocated species in Australia, construct extensive warrens, but how their warrens affect soil properties and vegetation communities is unknown. We investigated soil properties, vegetation communities, and novel ecosystem elements (specifically non‐native flora and fauna) on boodie warrens at three translocation sites widely distributed across the species’ former range. We found that soil moisture and most soil nutrients were higher, and soil compaction was lower, on warrens in all sites and habitat types. In contrast, there were few substantial changes to vegetation species richness, cover, composition, or productivity. In one habitat type, the cover of shrubs less than 1 m tall was greater on warrens than control plots. At the two sites where non‐native plants were present, their cover was greater, and they were more commonly found on boodie warrens compared to control plots. Fourteen species of native mammals and reptiles were recorded using the warrens, but, where they occurred, the scat of the non‐native rabbit (Oryctolagus cuniculus) was also more abundant on the warrens. Together, our results suggest that translocated boodie populations may be benefiting both native and non‐native flora and fauna. Translocated boodies, through the construction of their warrens, substantially alter the sites where they are released, but this does not always reflect their historic ecosystem roles.     

The effects of habitat and spatial features of wetland fragments on the abundance of two rallid species with different degrees of habitat specialization

The management of wild gamebirds for shooting involves a combination of habitat management (woodland, field boundaries and game crops), winter feeding and control of potential nest predators, any of which could benefit other birds, including nationally declining species associated with farmland habitats. Changes in numbers of passerines were monitored over six years in relation to game management on farmland in Leicestershire. Nest success was monitored over four years and, for some species, was inversely related to abundance of breeding corvids. Abundance of breeding passerines increased during the period of game management. Species whose breeding populations have declined nationally (coincident with agricultural intensification) showed the greatest increases in abundance relative both to other species, and to the same nationally declining species in nearby farmland. The precise mechanism by which the game management package contributes to increased breeding numbers is not understood and is likely to differ between species. However, these results show that further integration of wild game management into farming systems could have conservation benefits for nationally declining farmland birds.     

A Neglected Science: Applying Behavior to Aquatic Conservation

Behavioral theories, insights, and techniques are too frequently ignored by conservation biologists. Yet an animal's survival and reproductive success clearly depend on its behavior. Using examples from marine, freshwater, and terrestrial realms, I assert that behavioral information is invaluable in five conservation areas: (1) managing wild species (e.g., designing marine reserves; reducing animal–human conflicts; understanding and managing species' responses to human-induced environmental stress such as fishing, introduced species, and chemical, visual, and acoustic pollution); (2) actively reversing the decline of imperiled wild species (e.g., reducing bycatch by improving selectivity of fishing gear; re-establishing breeding populations and boosting reproduction); (3) assessing biodiversity (e.g., modeling population viability; censusing and monitoring populations and species); (4) captive breeding and reintroduction programs (e.g., minimizing loss of valuable phenotypes; teaching or maintaining valuable survival skills); and (5) changing human behavior in resource exploitation (e.g., using principles from social psychology). Both realized and potential applications to fishes are stressed. Finally, behavioral diversity, a valuable but neglected element of biodiversity, needs to be explicitly conserved to maintain diverse populations. Arguments are presented that the conservation of species diversity and genetic diversity alone does not necessarily protect important behavioral diversity. The maintenance of both individual and population variability may be essential for the preservation of a species.     

Reproduction in the endangered African wild dog: basic physiology, reproductive suppression and possible benefits of artificial insemination

The African wild dog (Lycaon pictus) is an endangered exotic canid with less than 5500 animals remaining in the wild. Despite numerous strategies to conserve this species, numbers of free-living animals are in decline. It is a highly social species with a complex pack structure: separate male and female dominant hierarchies with, typically, participation of subdominant adults in the rearing of the dominant breeding pairs' pups. Basic reproductive knowledge is largely missing in this species, with only limited information available on the profile of reproductive hormones, based on non-invasive endocrine monitoring. The dominant or alpha male and female are reproductively active and the subdominants are generally reproductively suppressed. However, the occasional production of litters by subdominant females and evidence of multiple paternity within litters suggests that fertility of subordinates is not completely inhibited. In this respect, there are still considerable gaps in our knowledge about the mechanisms governing reproduction and reproductive suppression in African wild dogs, particularly the influence of dominance and pack structure on both male and female fertility. Given concerns over the long-term survival of this species, further research in this area is essential to provide valuable information for their captive breeding and conservation. Reproductive information can also be applied to the development of Assisted Reproductive Techniques for this species; the utility of which in African wild dog conservation is also discussed.     

Factors influencing breeding success,ovarian cyclicity,and cub survival in zoo-managed tigers (Panthera tigris)

Understanding factors that influence reproduction and offspring survival in zoo populations is critical for management of threatened and endangered species. Examination of long-term data (1989–2011) compiled from the Association of Zoos and Aquarium's zoo-managed tiger breeding program provides the basis for a more thorough understanding of reproduction and scientifically based decisions for effective population management in this endangered felid. Biological and management-related factors that could influence tiger breeding success and cub survival were evaluated using logistic mixed models. Breeding success improved with female age until approximately age five, then declined thereafter. Experienced female breeders had greater breeding success than inexperienced females. Litter size was most predictive of cub survival, with average-sized litters (3–4 cubs) experiencing the highest proportional survival. Management-related factors, such as whether the breeding institution had a recent tiger litter and whether both animals were already located at the same institution, also influenced breeding success and cub survival. These results highlight the importance of institutional husbandry experience and the need to retain knowledge through staff turnovers to achieve optimal reproductive success. Using fecal estrogen data, frequency of ovarian cyclicity and mean cycle length did not differ by female age or parity; thus, lack of cyclicity and/or increased cycle duration are not likely explanations for declining breeding success with age. These results provide valuable reproductive information that should improve scientific management of zoo-based tiger populations.     

Breeding success of an endangered island endemic kestrel increases with extent of invasion by an alien plant species

The Mauritius Kestrel Falco punctatus, once the rarest kestrel worldwide, became an icon of bird conservation after it recovered from four to six individuals in 1974 to some 800 by 2005 following intense conservation management. Its population however then halved within about a decade prompting a re-evaluation of the IUCN status and up listing of the species in 2014 and an increased conservation attention. Drivers of this new decline are unclear and the influence of habitat structure and diet on breeding success may be important contributors but have received relatively little attention, particularly in the way they may interact to influence production of new fledglings. We address this knowledge gap by studying whether breeding success is influenced by habitat structure (in terms of cover of the invasive Ravenala in native habitats, an alien plant causing strong structural shift in the forests that it invades, and extent of cleared area), diet composition and food pass frequency (as a proxy for food intake) and food quality at 28 nests of a re-introduced kestrel population in south east Mauritius during the 2015–2016 breeding season. The kestrel’s diet comprised native and alien birds, reptiles, insects, and small alien mammals, with a disproportionately high proportion of Phelsuma gecko. A higher frequency of food provisioning and percentage cover of Ravenala both contributed to higher breeding success. Ravenala may increase gecko density or increase gecko detectability and predation by the kestrel, or both, while changed land use (pasture and sugar cane fields) may increase prey diversity in the form of non-forest prey known to be eaten by Kestrels (e.g. alien agamids, small mammals and birds). These prey related influences on breeding suggest that the Bambou mountain range provides a human-generated novel ecosystem altering food availability and increasing the kestrel’s breeding success. However, Ravenala is an invasive alien species harmful to the wider forest biodiversity. Progressive weeding of Ravenala and concurrent re-introduction and augmentation of native palms and Pandanus species which geckos can use at comparable densities to Ravenala, is recommended. This would likely improve the kestrel’s hunting habitat quality and maintain high gecko density or detectability and the vegetation structure required for hunting manoeuvrability and prey availability without the negative consequences of invasive Ravenala.