Abundance,breeding and growth of the crab Scylla serrata in two South African estuaries

Populations of the crab Scylla serrata (Forskal) were studied in two South African estuaries from 1971 until 1974. One estuary was open to the sea, the other was closed by a sandbar. Population density in the closed estuary was estimated by means of mark-release-recapture data from tagged crabs. The population of S. serrata was estimated as 1 crab/124m² and the production at 3.4 g/m²/annum. Catch per unit effort data from the closed estuary indicated a natural mortality of 41% in the crabs' second year and 60% in the third. Growth was studied in both estuaries by means of size-frequency analysis and from tagging returns. Growth was rapid in the first 12 to 15 months, when the crabs attained a carapace width of 80 to 160 mm. Thereafter growth slowed, and after 3 years crabs were between 140 and 180 mm in carapace width. Females mated in summer at a carapace width of 103 to 148 mm, males at 141 to 166 mm. After mating, females migrated out of both estuaries into the sea.     

Leaf consumption by <Emphasis Type="Italic">Sesarma plicata</Emphasis> in a mangrove forest at Jiulongjiang Estuary,China

The feeding ecology of Sesarma plicata (Grapsidae: Sesarminae), the most abundant crab species in a mangrove forest dominated by Kandelia candel at Jiulongjiang Estuary, China, was investigated through field and laboratory experiments. Feeding preference and consumption rates were determined on mature, senescent and decomposed leaves of Kandelia candel, Bruguiera gymnorrhiza and Aegiceras corniculatum. In the laboratory, S. plicata preferred leaves of K. candel over those of B. gymnorrhiza and A. corniculatum, and consumed significantly more decomposed leaves than mature and senescent ones, irrespective of crab size. Field experiments with limited power failed to reveal detectable species preferences despite more consumption of K. candel, but decomposed leaves of each species were again preferred. Leaf characteristics associated with preference changed with plant species and leaf state. Low tannins and high water content characterized the preference for decomposed state of leaves. Species preference was significantly and negatively related to crude fibers and C:N ratios for mature leaves, and crude fiber for senescent leaves, but significantly and positively related to water content for decomposed leaves. Leaf consumption rates averaged for all leaf categories from laboratory no-choice feeding experiments were 0.101, 0.055 and 0.017 gDW ind⁻¹ d⁻¹ for large, medium and small crabs, respectively. In this forest, mean density of S. plicata was 20.5 ind m⁻² as assessed by a manual catching method. Leaf litter removal rate during neap tides by sesarmid crabs was about 1.33 gDW m⁻² d⁻¹ in April 2006. The leaves removed by crabs were grazed on the sediment surface or taken into crab burrows, shredded and stored before being eaten.     

Influence of mussel rafts on spatial and seasonal abundance of crabs in the Ría de Arousa,North-West Spain

The crab community in the Ría de Arousa, North-West Spain, was studied for 2 yr (1974–1975; 1975–1976), firstly to detect any differences due to mussel (Mytilus edulis) raft aquaculture; secondly to study temporal changes in spatial distributions in beach, raft and non-raft areas throughout the ría. Portunid crabs dominated the fauna, the dominant species being Macropipus puber and M. depurator in the raft areas, M. depurator in non-raft subtidal areas, and M. arcuatus in beach stations. No substantial seasonal changes were observed in species composition throughout the ría. In general, the highest densities occurred in fall and winter. Raft stations showed the highest values of abundance, up to 0.47 individual per m², and 8.20 g wet wt m^-2. The food resource provided by the rich epifauna associated with the mussel rafts, and changes in the sediment due to shell deposits could account for the high density values in raft areas. Raft areas (polygons) cover about 10% of the surface of the Ría de Arousa and, thus, the raft aquaculture of mussels significantly increases crab production.     

Early growth and mortality of the Caribbean king crab Mithrax spinosissimus reared in the laboratory

The berried females of the Caribbean king crab Mithrax spinosissimus (Lamarck) used in this study were collected from canals on Big Pine Key, Sugarloaf Key and Lower Matecumbe Key (south Florida, USA) on 9 August, 8 October and 15 November 1986. Viable spawns hatched as first zoeae and molted to second zoeae within ca. 10 to 12 h. Most of the larvae reached the megalopa stage 1 d later, and molted to first crab 4 to 8 d after hatching (water temperature: 27.2° to 28.8°C). Low water temperature and/or early lack of food had a negative effect not only on stage duration, but also on the size of the early crab stages. Successful molt to first crabs occurred, however, in the absence of food. The growth rate (carapace length) between molts in early crab stages varied between ca. 20 and 30%. When provided with good water exchange, stocking density could be very high (>22 500 individuals m^-2), with no increase in mortality. The highest mortality rate was recorded when the larvae molted to first crab, and the highest rates of survival were always recorded when feeding was not initiated until after 5 to 8 d after hatching. No cannibalism was observed among larvae, and cannibalism was low in early crab stages. The study indicates that to achieve viable hatches and high larval survival in rearing M. spinosissimus, a continuous and adequate supply of high-quality seawater is a prerequisite both in larviculture and in maintaining brooding females.Contribution No. 93, Department of Oceanography and Ocean Engineering, Florida Institute of Technology     

Life history and population dynamics ofKinbergonuphis simoni (Polychaeta: Onuphidae)

The relationship between life-history characteristics and population dynamics were investigated in the onuphid polychaeteKinbergonuphis simoni (Santos, Day and Rice) between 1982 and 1987. The studied population is located in Upper Tampa Bay, Florida, USA. This worm attains sexual maturity at 5 to 10 mo of age, depending on temperature. Several consecutive broods are produced during a female's life time, with 7 to 26 young per brood. Adults die in June–July after the breeding season. Generations of breeding individuals do not overlap and the replacement of generations occurs in July–August. Only one extended breeding season is experienced per life time. Life span does not exceed 2 yr. Field population density in 1982 was high in fall and spring (2000 to 9000 individuals/m²), low in summer (1600 individuals/m²), and slightly depressed in winter (2500 to 4000 individuals/m²). Changes in population density may be explained by seasonal, temperature-controlled changes in instantaneous birth and death rates. Birth rates are high in fall and spring, while death rates are high in early summer. Death of juveniles contributes to the winter density decrease.     

Influence of predation by a crab on the distribution of the size-groups of three intertidal gastropods in South Australia

Two South Australian rocky intertidal platforms were sampled in April 1981 and December 1982 (Marino Rocks) and in December 1982 (Lady Bay), respectively. Three snail species, Nerita atramentosa Reeve, 1855, Bembicium nanum (Lamarck, 1822) and Austrocochlea concamerata (Wood, 1828), exhibited shore-level size-gradients, with smaller individuals occupying the higher intertidal levels. The reef crab (Ozius truncatus M.-Edwards, 1834), whose distribution overlaps that of the gastropods, is an important predator of these gastropods. The shore-level distribution of shell sizes can be explained by differential selection against smaller individuals by reef crabs. The decreased density of small snails at lower intertidal levels may be the result of either their consumption by reef crabs and/or their movement to higher shore levels where crab predation is less intense.     

A genetic analysis of the pea crabs (Decapoda: Pinnotheridae) of New Zealand

Seven populations of the pea crabPinnotheres atrinicola Page were sampled from around the North Island of New Zealand from February to October 1987, and individuals were scored at 23 presumptive enzymatic loci. For a brachyuran crab,P. atrinicola revealed high levels of polymorphism and heterozygosity. Of the loci scored, phosphoglucose isomerase (Gpi) and phosphoglucomutase (Pgm) were distinguished by high variability ( =0.602 and 0.526, respectively). A clinal variation in electromorph frequency was evident at several loci, and atGpi in particular. Statistical analyses revealed that, despite relatively small genetic distance separation, a high degree of structuring was present between the geographic populations. The degree of population subdivision observed in this study is atypical of brachyuran crabs. It is suggested that the genetic differentiation observed between pea crab populations is maintained by life-history attributes and current movements which restrict gene flow between populations and, to some extent, by random genetic drift.Publication No. 41 from the Evolutionary Genetics Laboratory, University of Auckland     

Life-history patterns in two forms of Macrophthalmus japonicus (Crustacea: Brachyura)

Two behaviorally distinct forms — courtship signal — (Form L and V) of Macrophthalmus japonicus (De Haan) are known. Monthly samples of each form were collected from two areas in Japan: Form L from Amakusa, western Kyushu, between April 1985 and February 1986; Form V from Fukuoka, northern Kyushu, between November 1982 and December 1983. Life-history traits were compared between two populations and discussed. Individuals from the Amakusa population grew more slowly and matured at a smaller size than those from the Fukuoka population. Maturation period and longevity did not differ significantly. In the first year of reproduction, females in the Fukuoka population produced more, and larger broods than those in the Amakusa population and crab mortality was higher in every age group. Egg size did not differ significantly. The major environmental factors — air-temperature, nitrogen content of the surface mud and period of aerial exposure were similar in both areas, and it is suggested that the differences in life-history exhibited by these crabs are genotypic, and perhaps related to their different form.     

Drastic decline of an extensive eelgrass bed in Nova Scotia due to the activity of the invasive green crab (Carcinus maenas)

In 2001–2002, there were severe declines of eelgrass (Zostera marina) in estuaries along the Nova Scotia coast of the southern Gulf of St. Lawrence. We examined the relationship between the recent invasion of European green crabs (Carcinus maenas) and the eelgrass decline in Benoit Cove in Tracadie Harbour. The abnormally abundant eelgrass wrack consisted mainly of entire shoots, not the usual blades. Three separate methods yielded similar estimates of the rate of shoot removal from the eelgrass beds (direct quadrat counts, “mark–recapture” of tagged shoots in the eelgrass bed, and rate of shoot appearance in the shore wrack). From 14 July to 8 September 2002, the shoot density in the bed fell by about 75 %. Similar rates of decline occurred in crab enclosures (4.4 crabs m^?2) placed in the eelgrass bed. Green crab foraging, involving the tearing of shoots and the digging of large pits, was the reason for the drastic decline of the eelgrass bed in Benoit Cove. It is clear that an invasion of green crabs to a region where they had not previously existed can both destroy eelgrass beds and restrict their recovery.     

Phenotype-environment matching in the shore crab (Carcinus maenas)

The shore crab (Carcinus maenas) exhibits a range of carapace pattern polymorphisms, but little is known regarding their function or maintenance. If patterns represent some form of crypsis, then associations between carapace colouration and substrate are expected; to determine whether such relationships exist, frequency of crab morphs and quantity of substrate type were measured from fifteen 10×40 m² quadrats at each of three sites along the southern shore of the Firth of Forth, Scotland. Five thousand one hundred and thirty-seven crabs and 3.6 km of line intercept transect data were collected during a 9-week period. Crab abundance, relative frequency of morphs and substrate type varied significantly among the three sites. Plain crabs were strongly associated with macro-algal substrates whereas patterned crabs were associated with mussel beds. This pronounced phenotype-environment matching, as well as various characteristics of the carapace patterns themselves, suggests that patterned crabs are cryptic on polychromatic backgrounds. The frequency of patterned crabs and the percentage of white pigment on the carapace both declined significantly with carapace width. The loss of pattern coincides with an ontogenetic shift in habitat use and we present evidence to suggest that individual crabs lose their pigment, rather than larger patterned crabs being preferentially removed from the population by predators. Throughout their life history, shore crabs encounter high variation in predation, food supply, and physical habitat; to survive they have evolved a strategy that includes elements of pattern polymorphism, crypsis, ontogenetic shifts, and plastic responses.     

Shell-size selection by intertidal sympatric hermit crabs

This study evaluated selection for shell size by three species of tropical intertidal hermit crabs, Clibanarius antillensis, C. sclopetarius, and C. vittatus, from species of shells which are frequently used in nature. Crab size and weight were strongly and significantly related to all measured parameters of the selected shells. The strength of these relationships (r² values) depended neither on the crab nor on the shell variables taken into account. The relationships between crab size and the dimensions of the selected shells showed higher r² values than the corresponding relationships with the shells that the crabs had occupied when they were collected (0.482–0.903 in comparison to 0.091–0.652, respectively), indicating that the crabs were occupying sub-optimal shells in nature. Negative allometry was frequently found in the relationships between crab and shell variables, indicating that large crabs select and use proportionally lighter shells than do small crabs. This negative allometry was stronger for the shells used in nature (except for C. antillensis), i.e. larger crabs tended to select heavier shells in the laboratory than in nature. Different allometric relationships were also recorded among the dimensions of shells used in nature and those selected by the hermit crabs in free-access experiments: as shell length increased, the selected shells were heavier and had larger apertures than the shells used in nature. The relationships between crab size and the length and weight of the selected shells did not depend on the species of crab or species of shell, but only on crab size. Therefore, analyses using these variables can be performed without taking the species of crab or shell into account, i.e. data from different crab or shell species can be pooled. The influence of crab and/or shell species was recorded only in the models fitted for aperture length and width, variables which were more related to shell architecture than did shell length or weight. In contrast, if crab weight is used as an independent variable, different crab or shell species can be analyzed together independently of the particular shell parameter. This indicates that crab weight may be less susceptible than crab shield length to shell morphological constraints. Finally, the results indicate that the preferred size of a given shell type chosen by a given hermit crab will depend more on crab size or weight, than on the crab or shell species under consideration, i.e. crab shell-size relationships are not species specific.Communicated by P.W. Sammarco, Chauvin     

Population dynamics of the commensal spider crab Inachus phalangium (Decapoda: Maiidae)

The ecology of the spider crab Inachus phalangium (Fabricius, 1775) (Decapoda: Maiidae) was studied in the field. I. phalangium inhabits the sublittoral on the sea anemone Anemonia sulcata Pennant. From July 1981 to April 1984 in the Mediterranean (Banyuls sur Mer, southern France) more than 3000 anemones were examined and ca 1800 I. phalangium were found on them. The population dynamics' generation cycles, reproductive activities and the dynamics of the sex ratio were investigated. The density of juveniles (crabs before the pubertal moult) on anemones changed in a yearly cycle from low in the first six months to very high in the second six months. The first occasional young crabs of a generation appeared in March/April (3rd and 4th decapodite stages) on the anemones. Their density increased enormously in the following months. The generation grew gradually on the anemones and moulted into puberty in September-January. Density of adults (crabs after the pubertal moult) on anemones changed in a yearly cycle from low to high from the summer to winter months. A new adult generation was recruited every autumn through the pubertal moult and disappeared in the following summer. Female reproductive activity continued throughout the year. Females carried several broods in succession, but the frequency of breeding females fluctuated on a yearly cycle. The highest percentage of egg-carrying females, i.e. the peak of the reproductive season, lay in the first half of the year. The maximal life span of a generation, from the hatching of the first larvae to the disappearance of the last adults, lasted 1.5 to 2 years. Males moulted into puberty ca one month later than females. The moulting distribution of adult males had roughly the same course as in females. An adult male generation diet out about one to two months before the female generation. Life expectancy was therefore 14 to 17 months for females and 12 to 15 months for males. The sex ratio of juveniles shortly before the pubertal moult was balanced. The sex ratio of adults shifted from 1:1 at the beginning of the reproductive period to ca 1:9 in favour of females at its end.     

Effect of predation threat on repeatability of individual crab behavior revealed by mark-recapture

The persistence of behavioral types in situ and the drivers of persistence are central to predicting the ecological effects of intraspecific behavioral variation. We surveyed individual refuge use of mud crabs (Panopeus herbstii), a behavior related to the strength of a trait-mediated trophic cascade in oyster reefs, in the absence and presence of toadfish (Opsanus tau) predation threat. We then released these crabs into the field and using mark-recapture, measured the repeatability of this behavior in the absence and presence of threat, and how behavioral change was affected by time in the field (a month on average, up to 81 days), crab size, and sex. Because crabs exhibited some evidence of a circatidal rhythm in refuge use, we also tested how tidal height during observation influenced behavioral change. Predation threat increased refuge use, and small crabs used the refuge more than large crabs, particularly under threat. In recaptured crabs, refuge use was more repeatable under threat. Neither time in the field, crab size, crab sex, nor tidal height had any effect on behavioral change. Our results support the non-mutually exclusive hypotheses that (1) prey organisms in the presence, rather than absence, of predation threat should exhibit less behavioral variability because the fear of dying (a severe fitness consequence) should take precedence over less immediately important influences on behavior (e.g., hunger) and that (2) individual behaviors tied to fixed traits (e.g., the body size dependence of refuge use under threat in this study), rather than variable traits, should be more repeatable over time.     

Epibiosis on cerith shells in a seagrass bed: correlation of shell occupant with epizoite distribution and abundance

The cerith Cerithium atratum (Born 1778) is an abundant gastropod in the seagrass beds at Cabo Frio, Brazil. In order to estimate the ecological importance of cerith shells as a rare hard substratum in the seagrass bed, the abundances of C. atratum and of cerith shells occupied by hermit crabs were quantified. The mean densities of C. atratum and hermit crabs were 1887 and 100 individuals m⁻², respectively, and these provided 0.5 m² shell area m⁻² available for epizoite colonization. The tube-forming polychaete Hydroides plateni (Kinberg 1867) and oyster Ostrea puelchana Orbigny, 1841 were the dominant visible epizoites on inhabited cerith shells. These epizoite populations were compared in order to investigate whether the temporal and spatial patterns in the epibiotic community were related to ecological and behavioral aspects of the occupant species (cerith or hermit crab). Larger cerith shells had a greater abundance of epizoites. Each epizoite showed a preference for a different occupant of the shells (the oyster for C. atratum and the polychaete for cerith shells occupied by hermit crabs). The oyster showed a seasonal pattern in abundance on C. atratum, being more common in fall (March–April). The distribution of the epizoites on the shells depended on the shell occupant species and was probably related to their different foraging activity –C. atratum ploughs half buried through the sediment, while the hermit crab crawls on the sediment surface. In both cases, the activity of the shell occupant was considered to be beneficial to the epizoites, as empty shells and shell fragments did not support a macroepifauna. Received: 1 May 2000 / Accepted: 8 August 2000     

Feeding habits of the dungeness crab Cancer magister as determined by the index of relative importance

Specimens of the Dungeness crab Cancer magister, were collected subtidally and intertidally from an estuary in Washington State, USA in June and September 1980, and January, April, May and July, 1981. Gut contens of freshly collected crabs were analyzed by the Index of Relative Importance; for each prey taxon, this method measured frequency of occurrence, percentage of total biomass, and percentage of total numbers consumed. The most important higher taxon eaten was fish; however, the most important prey genus was the shrimp Crangon spp. There was greater predation on Crangon spp. at night at the intertidal site, and during winter and spring when the shrimp were most abundant there. Feeding activity, as indicated by a weight-specific gut-fullness index, showed no consistent diel pattern. There were significant ontogenetic changes in feeding patterns: first-year crabs preyed primarily on very small bivalves or small crustaceans including their conspecifics; second-year crabs preferred Crangon spp. and fish, and third-year crabs preyed less on Crangon spp. and more on fish. Such changes in feeding habits with ege could be purely due to mechanics of food handling, but might also reduce competition among age groups of crabs, possibly partitioning resources within the estuary. Findings are discussed in terms of optimal foraging and compared to other similar studies.Contribution No. 599, School of Fisheries, University of Washington, Seattle, Washington 98195, USA     

Hydrocarbon metabolism in the intertidal fiddler crab Uca pugnax

The fiddler crab Uca pugnax was examined for its ability to metabolize foreign hydrocarbons. The microsomal mixed function oxidase system was identified in U. pugnax tissues using Aldrin epoxidation rates as the assay. Rates were slow: 96 pM Dieldrin per mg microsomal protein per hour in the hepatopancreas, 438 pM mg^-1 h^-1 in the gill, and 228 pM mg^-1 h^-1 in claw-muscle microsomes. Using standard methods, no difference in rates could be detected between crabs living in clean areas and those living in environments highly contaminated with foreign hydrocarbons. In vivo rates of naphthalene oxidation were measured and used to calculate a clearance time for U. pugnax body tissues based on the aromatic hydrocarbon content of crabs collected from an oil-polluted salt marsh. Calculated clearance time was beyond the life span of the crab. It is concluded that this minimal ability of U. pugnax to metabolize foreign hydrocarbons partially accounts for its sensitivity to oil pollution in the environment.     

Differences between native and invasive<Emphasis Type="Italic"> Caulerpa taxifolia</Emphasis>: a link between asexual fragmentation and abundance in invasive populations

The tropical and subtropical marine green alga Caulerpa taxifolia has invaded several temperate regions throughout the world, including southeastern Australia. In this study, I examined how invasive C. taxifolia from temperate southeastern Australia differed from native C. taxifolia from subtropical Moreton Bay, Australia, in the traits proposed as being important to its invasion success: thallus size and density, levels of asexual reproduction (fragmentation) and total biomass. Against the prediction of a large size for invasive C. taxifolia, native populations from Moreton Bay had larger stolons and fronds than invasive populations. However, invasive populations consistently had much higher densities of stolons, fronds and fragmented fronds; and a greater biomass compared to native populations. Average densities at invasive sites exceeded 4,700 stolons and 9,000 fronds/m² and were as high as 27,000 stolons and 95,000 fronds/m², which are the highest reported for C. taxifolia anywhere. Average densities of fragmented fronds at invasive sites were as high as 6,000/m² and up to 45% of all stolons at invasive sites could be directly linked to asexual recruitment via fragmented fronds. Importantly, at invasive locations there was a strong association between asexual reproduction and abundance demonstrated by positive correlations between the density of fragmented fronds and total biomass. These findings are the first to describe quantitative differences between native and invasive C. taxifolia and to demonstrate a link between the high levels of asexual reproduction and high abundance in invasive populations. Although the causes and consequences of high levels of asexual reproduction remain to be explored, this study suggests that changes in demographic and life-history traits during the invasion by C. taxifolia into temperate habitats may contribute to its success there.Communicated by M. S. Johnson, Crawley     

Population dynamics and reproduction of Mysella bidentata (Bivalvia: Galeommatacea) in Galway Bay,Irish west coast

Mysella bidentata (Montagu) is found in high densities in an Amphiura filiformis (O. F. Müller) community in Inner Galway Bay, west coast of Ireland. As part of an ongoing investigation at one station within this community, the population dynamics and reproductive pattern of this bivalve were investigated during the period November 1978-December 1981. Monthly sampling over the 3 yr period revealed a mean population density of 745±672 individuals m^-2. Recruitment occurred during August-October and three year-classes were identified. Growth rates were assessed by analysis of length-frequency histograms and winter growth checks. Individuals from the Y1 and Y2 year-classes functioned as hermaphrodites during the 1980 and 1981 reproductive seasons. The Y0 year-class contained both males and hermaphrodites. Results are compared with available information on the same species from other locations.Contribution No. 225 from the School of Marine Sciences, University College, Galway Ireland     

Growth and longevity of the mud snail Batillaria attramentaria

Crab pots were used to sample a population of Scylla serrata (Forskal) in an estuarine area in Queensland, Australia. Pots were laid 100 m apart at fixed positions for 4 d each month for 1 yr (April 1980–June 1981). Data from recapture of tagged crabs showed that males larger than 140 mm carapace width and females larger than 150 mm had a higher capture probability than did smaller crabs; thus size-frequency distributions based on crab-pot captures are biased. Spacing trials showed that pots positioned 50 m apart fished competitively but that there was no difference in catch between those placed 100 and 200 m apart. Catch distribution indicated that the presence of a crab in a pot reduced the probability of further captures. Temperature and incidence of recently moulted crabs accounted for 66% of variation in monthly catches. Catch-per-unit-effort (CPUE) data can be used as a measure of relative abundance of adults if allowance is made for temperature and the incidence of moulting. The capture-mark-release-recapture method for making population estimates was tested but it is concluded that, because of cost and bias in collecting techniques, it cannot be recommended as a technique for studying S. serrata.     

Kinetics of uranium uptake by the crab Pachygrapsus laevimanus and the zebra winkle Austrocochlea constricta

The crab Pachygrapsus laevimanus and the zebra winkle Austrocochlea constricta were exposed for 40 d to uranium (1.5 to 10 mg l^-1) in continuous-flow sea water in separate starved and fed treatments, and the kinetics of uranium bioaccumulation were estimated from an exponential model. Starved and fed crabs took up U at a similar rate, which suggests that sea water was the major source of U to the crab; the fed crabs excreted U more rapidly than the starved crabs and this led to a lower net uptake of U by fed crabs. Fed and starved winkles took up U at similar rates and excreted it at similar rates, so the sea water was also the major source of U to winkles. Crabs took up more U than winkles; the concentration factors were 7 to 18 and 4, respectively. Uranium turnover was quite slow for both species (11 to 36 d) as it was also for winkle shells (6 d); this suggests that the rate-limiting processes which control turnover are biological (e.g. growth or tissue replacement) or physical (e.g. diffusion into the shell) rather than chemical (e.g. precipitation, adsorption or exchange). There was no effect of increasing U concentration in water on the U kinetic parameters.