Costs and consequences of variation in the size of ruff leks

Summary We studied 13 ruff leks in a small region on the island of Gotland (Sweden) to investigate the effect of lek size on the costs and benefits of lekking for individual males. Male ruffs occur in two behaviourally and morphologically distinct forms, independents (residents plus marginals) and satellites, whose costs and benefits we have assessed separately. These ruff leks had from 1–10 resident (territory-holding) males and were visited daily by satellites, marginals and females from 5–25 May, when most copulations occurred. We used the average number of independent males, counted during censuses taken every 5 min during 2-h observation periods at each lek, as an index of mean lek size. Per independent male, the numbers of both satellites and females increased significantly with mean lek size. Female arrival rate and attendance (total female-minutes) also increased significantly with mean lek size as did the average per capita rate of mating success for resident males (that of satellites was not quite significant). Thus, the dispersion of both of these male categories did not appear to fit an ideal free distribution with respect to mating success. In addition, the number of independent-independent fights per independent and the rate of satellite-resident dyad formation per resident increased significantly with mean lek size. These results suggest that ruffs on larger leks enjoy higher mating success than those on smaller leks but also that costs increase with lek size. We suggest that independent males distribute themselves so as to maximize their own net benefits and that this factor can account for both the occurrence of ruff leks and the variation in their size. Correspondence to: J. Höglund     

Passing the buck: resource defence,lek breeding and mate choice in fallow deer

Summary Lek breeding systems, where males defend small, clustered mating territories, are thought to occur where the distribution of females is heavily clumped but males are unable to defend resources used by females. In this paper, we describe a breeding system in fallow deer where males are able to defend resources used by females but the most successful bucks instead defend small territories on a traditional mating ground; where the lek is sited in an area not heavily used by females at other times of year and is visited primarily by females in or close to oestrus; and where mating success on the lek is related to territory position and to male phenotype but not to the resources available on different lek territories. Comparisons with other ungulates suggest that lek breeding species fall into two groups: those where leks are regularly visited by herds of females many of which are not in oestrus and those, like fallow deer, where leks are visited primarily by oestrous females. In the latter species, it is unlikely that females visit the lek for ecological reasons.     

Correlates of male mating success in the ruff Philomachus pugnax,a lekking shorebird

Summary Male ruffs (Philomachus pugnax), a lekking shorebird, can generally be divided into two morphs according to behavior and plumage coloration. Predominantly dark-colored, independent male ruffs defend small territories on a lek, whereas lighter colored satellite males are non-territorial and less site-faithful to a particular lek. The traits associated with the mating success of independent male ruffs were studied at two leks during two seasons on the island of Öland, Sweden in May and June of 1985 and 1987. Using multivariate analyses, three characteristics were found to be significantly related to mating success: high frequency of visits by satellites to an independent male's residence, consistency of lek attendance, and low rates of aggressive behavior. In contrast, mating success was unrelated to the degree of darkness of the independent male nuptial plumage, territory location on the lek, or rate of courtship displays. The use of multivariate analyses helped to resolve conflicting conclusions from previous studies which employed simple statistical analyses, or none at all. Experimental manipulations are suggested which will help to further determine the influence of female mate choice in this lekking system.     

Male capacity as related to sperm production,pregnancy initiation,and sperm competition in deer mice (Peromyscus maniculatus)

Summary Limitations on male capacity may be important to the evolution of mating strategies and behavior. Sperm counts in successive ejaculates of deer mice (Peromyscus maniculatus) decreased progressively through six ejaculations and remained depressed the following day. Males were as effective in inseminating females in cycling estrus with their second two ejaculates as with their first two and as effective in inseminating postpartum females with their fifth as with their first. In sperm competition situations males competed with rested males as effectively with their fourth, fifth, and sixth ejaculates as with their first. Thus, although sperm counts decrease in successive ejaculates, males suffer no apparent functional deficiency as long as they continue ejaculating. It is not clear whether or not deer mice are honest salesmen but they appear to be honest ejaculators and usually are honest intromitters.     

Growth and survival of Mytilus edulis larvae exposed to low levels of dibutyltin and tributyltin

Two studies were conducted to observe effects of dibutyltin (DBT) and tributyltin (TBT) on larvae of Mytilus edulis for an exposure period of 25 d. Endpoints for evaluation were shell growth and mortality measured at 33 d. Larvae were cultured in a new laboratory assay chamber in a recirculating static test. The control, 2, 20, and 200 g/l DBT-treated populations had mean shell lengths of 527, 523, 417, and 180 m, respectively. Survival was 1% for the 200 g/l DBT-treated population, but ranged from 73 to 83% for controls, 2, and 20 g/l treatments. The no-observed-effect concentration (NOEC) was 2 g/l for DBT, while the lowest-observed-effect concentration (LOEC) was 20 g/l. The chronic toxicity value was 6.3 g/l. In the TBT bioassay, mean shell lengths for the control, 0.006, 0.050, and 0.130 g/l-treated populations were 565, 437, 385, and 292 m, respectively. Control survival was 74%, whereas TBT-treated populations survival ranged from 52 to 58%. The NOEC for TBT was 0.006 g/l TBT and the LOEC was 0.050. A chronic toxicity value of 0.017 g/l was calculated. The results of this study indicated that the toxicity of DBT was less than that of TBT. It was concluded that shell length was inversely related to exposure level in both DBT and TBT bioassays. In this study, we have observed TBT effects at lower exposure levels in the laboratory than previously reported, and also report the first data for DBT effects on mussel larvae.     

Sexual conflict in the house sparrow: interference between polygynously mated females versus asymmetric male investment

Summary The reproductive success of female house sparrows mated with polygynous males depended to some extent on the aid received from their mates. Polygynous males fed nestlings at the same rate as monogamous males although polygynous males gave aid almost exclusively to one of their mates (the preferred). As a consequence, the number and quality of young raised by preferred females were similar to those of monogamous females, whereas nonaided females experienced a reduction of clutch size, hatching success, and fledgling quality. Males aided the harem female that hatched her clutch earliest (in 11 out of 12 cases). However, harem females that laid eggs earliest (i.e., the potential preferred) lost eggs and/or nestlings more frequently than harem females that delayed laying. This resulted from competing females' infanticide suited for redirecting males' aid. All polygynously mated females delayed the start of a new clutch and reproduced fewer times per season than monogamous females. Consequently, monogamy seems to be the optimum mating situation for females; whereas polygynous males raised more young per year than monogamous males, interference between harem females reduced their maximum expected success.     

Carotenoids versus retinoids (Vitamin A) as essential growth factors in penaeid prawns (Penaeus semisulcatus)

Penaeus semisulcatus de Haan adult gravid females, eggs and non-feeding early larvae, including Protozoea I, were used to test the hypothesis that retinoid (Vitamin A) is not required in early decapod crustacean development. In the adult gravid females, retinoids were detected only in the eyes (1.56±0.23 g g^-1 wet mass), whereas there were up to 97 g g^-1 wet mass total carotenoid in digestive gland and epidermis. This was mostly esterified, except in the ovaries, where free astaxanthin predominated (30 g g^-1). No retinoids could be detected in the eggs, the naupliar stages or Protozoea I, but free astaxanthin was metabolised exponentially, falling from 19 g g^-1 in the eggs to 4 g g^-1 in Protozoea I. This suggests that retinoid is not essential in early development and that carotenoid could be taking its place. Also, including retinoids in artificial diets appears to be unnecessary, provided adequate carotenoid is supplied.     

Arsenic tissue concentration of immature mice one hour after oral exposure to gold mine tailings

A potentially high bioavailability of arsenic in gold mine tailings from a site in northern California has been suggested by solubility studies. To help address this issue, an in vivo dosing study was conducted using 12dayold Swiss Webster mouse pups (n=8/group). A sample of sizefractionated mine tailings from the site (<20m particle size, 691g g^–1 arsenic) was prepared as an aqueous suspension and administered by gavage in a volume that provided 4mg As/kg body weight. The control group received the same volume of a commercial soil (1g g^–1 As) of similar particle size (<60m). No mortality or toxic signs were noted in either group. Tissue samples were collected 1h after gavage, freezedried, microwavedigested and analysed for arsenic by ICP/MS (detection limit 2ng As g^–1 dry weight). Arsenic concentrations (ng As g^–1 dry weight) in tissues from the pups who received mine tailings were significantly higher than in control tissues. The mean elevation in arsenic concentration was highest in the liver (3364% of control, p<0.0001), followed by blood (818 of control, p<0.0001), skin (207% of control, p=0.07), and brain (143% of control, p<0.0001). The carcass arsenic concentration (excluding the GI tract, liver, brain and skin) was 138 of control (p=0.02). The data indicate uptake of arsenic from weathered mine tailings by the immature mouse pups after oral exposure.     

Grazing and ingestion rates of nauplii,copepodids and adults of the marine planktonic copepod Calanus helgolandicus

The average grazing and ingestion rates of all stages of the marine planktonic copepod Calanus helgolandicus (Calanoida) from nauplius stage IV to adults were measured experimentally at 15°C in agitated cultures. The chain-forming diatom Lauderia borealis and the unarmoured dinoflagellate Gymnodinium splendens were offered as food. The food concentrations were close to natural conditions and ranged from 36 to 101 g of organic carbon per liter. The medium body weights expressed in g of organic carbon of almost all larval stages raised at 49 g C/1 were identical with the weight of the same stages caught in the Pacific Ocean off La Jolla, California, USA. In a log-log system, grazing and ingestion rates increased almost linearly with increasing body weight. Grazing rates ranged from 4 to 21 ml/day/nauplius stage IV to 286 ml to 773 ml/day/female. Ingestion rates increased from 0.2 g to 0.8 g C/day/nauplius stage IV to 18 g to 69 g C/day/female. Grazing and ingestion rates per unit body weight decreased gradually with increasing body weight. The daily ingested amount of food decreased from 292 to 481% of the body weight (g C) of nauplius stage V to 28–85% of the body weight of adult females. Grazing and ingestion performances of all stages increased with increasing particle size. Grazing rates decreased and ingestion rates increased with increasing food concentrations. The published data on food intake of the different age groups of C. helgolandicus show that the young stages of herbivorous planktonic copepods can play a major part in the consumption of phytoplankton in the sea due to their high grazing and ingestion rates.     

Sexual selection and cues for female choice in leks of Jackson's widowbird Euplectes jacksoni

Summary Sexual selection through female mate choice was investigated in the lekking Jackson's widowbird by applying multivariate selection analysis to observational data from four leks. Males perform a stereotyped jump display on small display courts (dance rings) constructed by the males in open grassland. Females visit the lek solely for mating and nest on their own, away from the lek area. Few cases of interference during courtship and absence of position effects on mating success indicated that female choice within the leks was not pre-empted by male-male competition. In a set of 11 male traits with mating success as the dependent fitness measure, significant selection differentials (covariances) were found for the length of the conspicuous tail and the rate of the jump display, suggesting sexual selection of these traits. They also showed the largest selection gradients (partial effects) and thereby seem to be the cues on which females base their choice. The success of males in obtaining copulations appears to depend on two components: display rate and lek attendance affect the number of female visits, whereas tail length seems to primarily influence the chance of copulating with a visiting female. Tail length was positively related to a measure of body condition, which is of interest with regard to the suggestions that sexual ornaments may serve as indicators of male viability.     

The effect of a “permanent” clock-shift on the orientation of experienced homing pigeons

Summary A group of experienced homing pigeons vas subjected to a 6 h slow shift of their internal clock and kept under these conditions for more than 2 months. During the overlap time between the natural and artificial photoperiods they were released for training flights to familiarize them with an area while living in a permanent shift.Tested outside the permanent shift training range, the experimentals always deviated about 30° clockwise from the mean of their controls, markedly less than in a regular 6 h slow shift. Inside the permanent shift training range, however, they oriented like the controls (Fig. 2). When their internal clock was returned to normal, the birds showed a larger counterclockwise deflection on their first flight, which was roughly comparable to the effect of a regular 6 h fast shift (Fig. 3). On later flights after normalization, this large shift was no longer found; instead we observed a roughly 30° counterclockwise deflection when they were released inside the permanent shift training range in the morning. This deflection did not seem to occur in the afternoon or outside the permanent shift training range (Figs. 4, 5), and it disappeared when the birds were repeatedly released from the same site (Fig. 6).The occurrence or non-occurrence of the deflection was independent of the duration of the shift or the time passed after normalization; it seemed to depend solely on whether the birds had become familiar with a given site in the situation of the permanent shift. This argues against an effect based on the sun compass. We tend to assume that the still unknown navigational map is involved. In this case, however, as the deflection is independent of the home direction and the type of release site bias, the factors in question would act very differently from the gradients on which the traditional concepts of the navigational map are based. The processes establishing and updating the map and their possible differences are discussed.Died on August 17, 1980     

Reproductive biology of commercially exploited deep-water royal red prawns (Haliporoides sibogae,Solenoceridae) in south-east Australia

Deep-water royal red prawns (Haliporoides sibogae de Man, 1907; Solenoceridae: Penaeidea) were sampled from commercial catches off south-east Australia between November 1988 and May 1990. Four stages of sexual development were distinguished macroscopically for each sex. For the females, histology and analysis of oocyte size were used to validate the macroscopic staging of ovaries into undeveloped, developing, early ripe and ripe. Females were larger than males, reaching a maximum size of 46 mm carapace length (CL) compared to 32 mm CL for males. Average size at sexual maturity was 30.8 mm CL for females and 25.8 mm CL for males. Females apparently breed several times in their life, whereas males probably breed only once. Individual fecundity (ranging from about 58 000 to 140 000 oocytes) was relatively low compared with other penaeids, but oocytes were larger (mean diameter=0.41 mm). Two short breeding seasons occurred per year (February to April and July to August). There was a clear size-related latitudinal distribution of the prawns, with largr prawns (spawners) being more abundant in the northern region of the fishing area, and juveniles (CL20 mm) being more abundant in the south. Most of the spawning stock probably lies outside the main fishing ground, while the fishery targets mainly prawns below the size at maturity.     

Primary production in Dumbell Bay in the Arctic Ocean

Photosynthesis, chlorophyll, inorganic nutrients, and related factors were measured throughout a productive season in a small coastal inlet of the Arctic Ocean. Significant production was confined to 5 m for a very limited time period beacause of both light and nutrient limitations. Maximum chlorophyll was 8.2 mg Chl a m^-3 and maximum gross and net photosynthesis rates were 830 and 550 mg C m^-2 d^-1. Annual gross and net photosynthesis is estimated at about 12 and 9 g C m^-2 respectively. The effect of light stimulation on assimilation is described, sources of coastal zone nutrient enrichment are considered, and discrepancies between gross and net photosynthesis are discussed with reference to the growth characteristics of the phytoplankton population.Bigelow Laboratory for Ocean Sciences. Contribution No. 76008     

The structure of plankton communities of the Indian Ocean

Species diversity and trophic structure of plankton communities have been considered as most important biological features. The Fisher () and Simpson () indices and the information index H=- pi log₂ pi (Margalef) were used for evaluation of species diversity. The indices have been calculated within the group of filter feeders by numbers ( and H) and biomass ( and H) of the species; the best results were obtained in the latter case. The index H=- pi log₂ pi is proposed for the quantitative evaluation of trophic structure, where a portion of an individual trophic group is taken as pi. The plankton material was collected by the R.V. Vityaz in the northern Indian Ocean in January through February, 1960. Analysis of characteristics of zooplankton distribution, species and trophic diversity in connection with characteristics of the water regime resulted in the definition of 3 types of the plankton communities: (a) those in intensive divergence zones; (b) those in poor divergence zones; (c) those in regions with stable water stratification or with feebly marked convergence, distiguished by different values of biomass, species diversity, and trophic group ratios.     

Evaluation of Public Housing Lead Risk Assessment Data

A unique data set from lead risk assessments performed on 67 public housing developments from across the United States was made available for analyzes. The data set includes results of lead analysis from 5906 dust wipes and from 1222 soil samples. A total of 487 dwelling units in these developments, as well as associated common areas, were sampled, all by the same team of inspectors. The number of dwelling units within a development that were sampled reflected the guidelines then in force, the 1990 Interim HUD Guidelines, rather than those specified in the 1995 Guidelines. Median dust lead loadings for floors, 151gm^–2 (14gft^–2), and window sills, 936gm^–2 (87gft^–2), were much less than former HUD limits of 1076gm^–2 (100gft^–2) and 5380gm^–2 (500gft^–2), respectively and are only about one-third of the recently established limits of 431gm^–2 (40gft^–2) and 2690gm^–2 (250gft^–2). In contrast, the median lead loading for window troughs, 8560gm^–2 (795gft^–2), was almost identical to the HUD clearance limit of 8610gm^–2 (800gft^–2). There was a strong positive correlation between floor and window trough lead loading values for samples from the same dwelling units and those from common areas of the housing developments. Door threshold samples, which may reflect conditions exterior to the dwelling unit, were collected from 53 dwelling units. Median lead loading levels of these samples were more than ten times higher than those in floor samples from the same dwelling units, were about the same as window sill samples and about one-half of levels in window trough samples. Composite sample results, simulated by averaging results from four samples within a dwelling unit, revealed that in order to have the same rate of excedence of standards, the composite standards would have to be reduced, for example, from the single sample value of 1076gm^–2 (100gft^–2) to 527gm^–2 (49gft^–2) for floor samples and from the single sample value of 8610gm^–2 (800gft^–2) to 5160gm^–2 (479gft^–2) for window troughs. For this public housing data set, the portion of the units in developments containing more than 225 units which exceeded the established limit for window samples was the same when using either the full data set or a random one-half of the data set. This suggests that, for this data set, the number of dwelling units sampled was excessive . Thus, the required increase in the number of dwelling units to be sampled specified in the 1995 Guidelines for developments with more than 225 dwelling units, may not have been necessary if this data set is representative of public housing developments in the United States.     

Lead and cadmium in urban allotment and garden soils and vegetables in the United Kingdom

In order to assess the intake of lead and cadmium by consumers of home grown vegetables in urban areas, replicated experimental plots of uniform size, comprising summer and winter crops, were established in 94 gardens and allotments in nine towns and cities in England.The geometric mean lead and cadmium concentrations for the soils (n = 94) were 217 g g^–1 (ranging from 27 to 1,676 g g^–1) and 0.53 g g^–1 (<0.2–5.9 g g^–1), respectively. Compared with agricultural soils, the garden and allotment soils contained elevated levels of lead but not cadmium.Lead concentrations in the vegetables ranged from <0.25 g g^–1 to 16.7 g g^–1 dry weight and cadmium concentrations ranged from <0.025 g g^–1 to 10.4 g g^–1 dry weight. Lead concentrations were higher than reported background levels, although <1% exceeded the statutory limit for saleable food in the UK (1 g g^–1 fresh weight). Cadmium concentrations were generally similar to background levels.     

Effect of daylight variations on the energy budgets of shallow-water corals

Energy budgets were determined for small pieces (nubbins) of the coralsPocillopora damicornis, Montipora verrucosa andPorites lobata living at a water depth of 3 m on the fringing reef of Coconut Island, Kaneohe, Hawaii. The budgets were determined for three different types of day: an ideal day with no cloud and an in situ daily integrated irradiance at 3 m of 14.385 E m^–2 d^–1; a normal day with sporadic cloud cover and daily irradiance of 11.915 E m^–2 d^–1; and an overcast day with daily irradiance of 6.128 E m^–2 d^–1. On the ideal day, the energy fixed in photosynthesis was more than that required for respiration and growth of both zooxanthellae and animal components of the association, and there was a predicted loss of between 19.3 and 32.4% of the energy fixed. On a normal day, the total photosynthetic energy fixation was lower and the excess was between 12.1 and 27.9% of the energy fixed. On the overcast day, however, in bothPocillopora damicornis andPorites lobata energy expenditure exceeded photosynthetic energy fixation and the budget was in deficit. Estimates of rate of mucus secretion on an overcast day were derived and, when incorporated into the energy budget, it was predicted that all three species would have a deficit budget, necessitating the catabolism of lipid reserves. From published values for lipid storage in these species it was calculated that the reserves would last from 28 d inPocillopora damicornis to 114 d inM. verrucosa. A model is suggested in which corals draw upon their extensive lipid stores on days of sub-optimal light, replenishing the reserves again when daily light levels are high, and finally excreting the excess energy fixed, as mucus-lipid when the lipid stores are replete.     

Aqua regia extractable trace elements in surface soils of venezuela

Surface soils (0–15 cm) were sampled at 10–20 km intervals along two transects in Venezuela. One (1162 km, 70 samples) ran west to east parallel with the Caribbean coastline, the other (920 km, 92 samples) ran south to north from the frontier with Brazil to the Caribbean shore. Sampling took place in both a wet and a dry season. Trace metals were extracted from dried, sieved (<2 mm) soil with boiling aqua regia followed by analysis by ICP or flame AAS. Metal values did not differ significantly between the two seasons and dates were averaged. Geometric mean values for the west–east transect were: Cr=41.5, Cu 17.9, Cs 3.6, Li=13.9, Mn=294, Ni=21.3, Pb=17.4, Sr=39.4, V=60.4 and Zn = 83.7g g^–1, respectively. Similarly, for the south–north transect Cr=21.3, Cu=4.3, Cs=1.1, Li=2.0, Mn=55.7, Ni=4.4, Pb=6.1, Sr=13.3, V=28.2 and Zn=16.7g g^–1, respectively. A classification of samples by lithology showed surface soil composition to be related to rock composition. Metal values were low in the soils in the south of the country, in the Guyana highlands (Gran Sabana). Low Zn contents were prevalent. Lead contents were affected by roadside fallout from vehicles using leaded petrol except that high Pb contents of soils in the Gran Sabana were of more complex origin.     

Siblicide,parent-offspring conflict,and unequal parental investment by egrets and herons

Summary O'Connor (1978) generated behavioral predictions from Trivers's (1974) theory of parent-offspring conflict (POC) in relation to avian brood reduction, especially for species using escalated sibling aggression. In general, when daily mortality rates fall between certain values (the mortality difference thresholds for parents and surviving offspring: O'Connor 1978), POC is expected. At such times parents and the early-hatching senior offspring are expected to disagree on the necessity, or at least timing, of the victim chick's death. Because that death is promoted directly by the senior chicks and because parents have realistic means of nullifying those efforts, behavioral conflict is expected. I examined the activities of all family members in one siblicidal (great egrets, Casmerodius albus) and one nonsiblicidal brood-reducing species (great blue herons, Ardea herodias) of ardeidae in Texas nesting colonies. Daily mortality rates over two breeding seasons approximated the predicted POC zone for four-chick (B/4) heron broods and three-chick (B/3) egret broods. As predicted, egret sibling aggression was significantly lower in two-chick broods. Herons seldom fought, regardless of brood size. Special attention was paid to the egret B/3 data for behavioral manifestations of POC. Parents did not interfere overtly with sibling fights (99.2% of 2,829), but parents seemed to have a subtle mollifying effect on the aggression (fewer sever fights occurred when parents were present). Parents showed no favoritism in food distribution to dying victim chicks. The mixed results are consistent with two rather different POC interpretations: (1) that significant conflict exists and the offspring are winning or (2) that there is no significant evolutionary conflict. In the latter case, parents are regarded as creating asymmetries among offspring (e.g., via hatch asynchrony), then leaving competition among siblings to run its course without further intervention (laissezfaire policy). This view explicitly recognizes that siblicide and other investment-skewing selfish chick behaviors may serve parental interests well-i.e., that equal allocation of resources is not likely to be the parental optimum. Using the relative reproductive values of each nestling as an index of the parent's unequal parental investment optimum produces a closer fit to the observed food distribution skew, but does not resolve the question of whether conflict remains.     

Carapace and branchial water circulation,and water-related behaviours in the semaphore crabHeloecius cordiformis (Decapoda: Brachyura: Ocypodidae)

Water uptake, branchial water circulation and various water-related behaviours and anatomical specialisations, are described for the Australian semaphore crabHeloecius cordiformis (H. Milne-Edwards 1837). Male and female semaphore crabs were observed and collected from mangrove swamps around the south side of Botany Bay, Australia between 1983 and 1987. Semaphore crabs are active at low tide in air. Their branchial chambers are modified for both air breathing and water circulation. Water is held within the branchial chambers and is used for feeding and, to a certain extent, respiration. Crabs carry between 0.35 and 0.5 ml of water (7 g live body wt). Branchial water is continuously pumped out by the scaphognathites over specialised setae-bearing regions of the carapace and back through the branchial chambers over the gills, in a manner similar to that found in many grapsid crabs. Setae associated with water circulation are described. Water falling ventrally is collected by the abdominosternal cavity (formed between the abdomen and sternum) from where it returns to the branchial chambers via branchiosternal canals. Several novel water-related behaviours are described: Water shoogling — branchial water is drawn out between the walking legs where it is worked by scissor-like up and down movements of the legs. Abdomen flap — branchial water volume is adjusted by the extension and flexion of the abdomen against the cephalothorax. Water dumping — by tilting forwards and kissing the substrate, branchial water is voided to allow an entirely fresh load to be taken up. Statuesque posing — crabs remain erect and motionless for long periods. Branchial water is absent and external body surfaces dry out. Posing behaviour may function in thermoregulation, osmoregulation or as an anti-fouling mechanism against settling organisms. The significance of these various behaviours is discussed.