Divorce in common murres (Uria aalge): relationship to parental quality

Behavioral precursors of 12 divorces were examined in 30 color-banded pairs of common murres (Uria aalge) over six breeding seasons. Common murres are long-lived seabirds that typically return each year to the same mate and nest site in dense colonies. At least one parent is present continuously from egg lay to chick fledging. Murres, therefore, have considerable opportunities to compare their mates parental behavior with that of several familiar neighbors. Previous reproductive success was lower for divorcing birds than for reuniting pairs. As predicted by the better option hypothesis, there were clear choosers (seven females and five males) that initiated divorce by moving to a new birds site or by courting a new partner at their current site. Choosers improved their reproductive success after the divorce, whereas their previous partners, the victims did not. Yearly divorce rates (average 8.2% per year) were significantly correlated with yearly mortality rates. Divorces appear to be opportunistic: pairs divorced after varying numbers of reproductive failures with the immediate precursor usually being the disappearance (death) of a murre from a successful neighboring site. In contrast to the delays experienced by victims, choosers formed new pairs quickly and laid their eggs no later than reuniting pairs. Prior to the divorce, victims fed their chicks less often than choosers, and some engaged in other behaviors that compromised egg or chick survival. These observations suggest that deficiencies in parental behavior were precursors to the divorce. This report is one of the first cases where reproductive failure of divorcing pairs has been linked to deficits in the parental behavior of the subsequent divorce victim.Communicated by C. Brown     

Expansive covers of turf-forming algae on human-dominated coast: the relative effects of increasing nutrient and sediment loads

Turf-forming algae form more extensive habitat on subtidal rock adjacent to urban than non-urban coast of South Australia. This pattern is frequently observed on the worlds temperate coasts and is variously considered to be a result of enhanced concentration of nutrients or rates of sediment deposition on urban coasts. We experimentally tested which of three components of environmental change (increased nutrients in water, increased nutrients in sediments and increased sediment deposition) best explain the expansive covers of turf-forming algae on urban coasts. All three treatments had independent and positive effects on the percentage cover of turf-forming algae. The addition of nutrients from the water column had the largest influence (²=0.55), which was more than six times greater than the effect of nutrients added to sediments (²=0.08). An increase in rate of deposition of sediments had substantial effects (²=0.35), which were about one third less than those of water-borne nutrients. Importantly, the combined effect of all three treatments caused a 77% increase in percentage cover of turf-forming algae, which is comparable to the observed difference in covers between urban and non-urban coast in South Australia (93%). These results suggest that human activities that reduce water quality in both nutrient and sediment loads account for major change observed on human-dominated coasts. Despite this knowledge, we still lack complete information on the mechanisms that switch the primary subtidal habitat from canopy-forming algae to turf-forming algae on human-dominated coasts.Communicated by M.S. Johnson, Crawley     

Stable isotopic investigation of physiological and environmental changes recorded in shell carbonate from the giant clam Tridacna maxima

The aragonitic shell of the photosymbiont-bearing bivalve Tridacna maxima contains a record of the physiological and environmental changes the organism has experienced during its lifetime. This record is preserved as chemical and microstructural variations throughout the shell. Stable isotopic analyses of oxygen (¹⁸O/¹⁶O) and carbon (¹³C/¹²C) in shell carbonate were combined with growth increment studies to interpret the shell record of specimens collected from the Rose Atoll (Lat. 14°31S; Long. 168°10W) in April 1982. The seasonal water temperature cycle is recorded in the oxygen isotopic signature of the clams, permitting the recognition of annual cycles in the ¹⁸O profile. The total number of these cycles corresponds to the age of a specimen, while the cycle length is a measure of the yearly growth rate. Large-amplitude cycles, reflecting year-round calcification, characterize the early portion of the growth record. With the onset of sexual maturity and slower growth at an age of approximately ten years, the cycles decrease in amplitude and become more erratic. During this later growth phase calcification is limited to the cooler months of the year, perhaps in response to a re-ordering of energy priorities between growth and gametogenesis. A growth curve developed from the ¹⁸O profile indicates rapid juvenile shell growth followed by slower growth thereafter producing a lifespan of several decades. Carbon isotopic analyses of T. maxima were compared to analyses of the symbiont-barren gastropod Terebra areolata collected from the same locality in April 1984. A 2 depletion in the ¹³C composition of T. maxima shell carbonate is attributed to a symbiontenhanced metabolic rate and an increased flow of isotopically light, respired CO₂ into the carbon pool used in calcification. Such a depletion may prove useful in identifying the presence of photosymbionts in extinct species of fossil mollusks.     

Spatial mesoscale patterns of West Pacific picophytoplankton as analyzed by flow cytometry: their contribution to subsurface chlorophyll maxima

Vertical distributions of picophytoplankton (ppp) (<2 m) were studied by ship-board flow cytometry during two cruises in Western Pacific waters to Palau and to Australia in 1990. Weak red-fluorescing small ppp, supposed to be free-living prochlorophytes (Chisholm et al. 1988), were abundant in the area surveyed. These ppp, designated the prochlorophytes, were abundant in the surface waters (>10⁴ cells ml^-1) at the northern region (27°03N; 7°11N) in November, whereas in December at the southern tropical stations (0°23.54S; 9°20.30S; 13°50.6S), they formed subsurface maximum layers (>10⁵ cells ml^-1) on a nitracline at a depth of 3.5 to 5.4% surface irradiation. Their fluorescence intensity increased with depth below 10% surface irradiation. The prochlorophytes at a depth of 1% surface irradiation had ten times higher fluorescence than those at the surface layer. The total fluorescence intensity of the prochlorophytes accounted for 32 to 63% of the sum of the total fluorescence intensity of all fluorescing phytoplankton detected at subsurface chlorophyll maxima in the tropical area. These results suggest that distribution of the prochlorophytes is greatly affected by nitracline and by light intensity and that their chlorophyll is a major contributor to the subsurface chlorophyll maximum in the pelagic West Pacific Ocean.     

Hyperiid amphipods in the eastern part of the South Pacific gyre

Studies were made of Amphipoda Hyperiidea collected in 1987 with a pelagic trawl in the South Pacific anticyclonic gyre in the vicinity of the Nazca and Sala y Gómez ridges (17°35 to 26°01S, 80°10 to 100°50W), at depths ranging from 50 to 300 or 600 m. Forty-three hauls contained 7 198 specimens, comprised of 119 species within 43 genera. Two of the species appeared to be new for science; two others were found in the Pacific Ocean for the first time. A comparison with hyperiid collections from other tropical Pacific regions revealed great stability of oceanic faunal composion. However, the collections differed considerably in their dominant species. Furthermore, comparison of Hyperiidea from various parts within the investigated area of the South Pacific gyre also reflected differences in dominant species. The degree of these mesoscale spatial differences was comparable to that among the central regions of the northern and southern anticyclonic gyres.     

Habitat and age of the giant squid (<Emphasis Type="Italic">Architeuthis sanctipauli</Emphasis>) inferred from isotopic analyses

The age and habitat of the giant squid, Architeuthis sanctipauli Velain, 1877, were determined based on isotopic analyses of the statoliths of three female specimens captured off Tasmania, Australia, between January and March 1996. Assuming that the aragonite of the statoliths formed in equilibrium with seawater, ¹⁸O analyses indicated that the squid lived at temperatures of 10.5–12.9°C, corresponding to average depths of 125–250 m and maximum depths of 500 m. The capture records indicated that these squid may have occasionally ranged still deeper, to as much as 1000 m. All the statoliths were labeled with bomb ¹⁴C (¹⁴C=+22.9 to +44.6), consistent with the depths inferred from ¹⁸O. A thin section through one of the statoliths revealed 351 growth increments grouped into check-ring structures every 10–16 increments. A model for statolith growth and the pattern of temporal change in ¹⁴C in the water column was used to estimate the ages of the three specimens. These estimates were very sensitive to the choice of depth range over which ¹⁴C values were integrated. Assuming that the capture depths represented the maximum habitat depths of these individuals, the calculations suggested an age of 14 years or less. More refined age estimates require a better understanding of the variation of ¹⁴C and temperature with depth in the areas in which the squids live.Communicated by J.P. Grassle, New Brunswick     

Hydrogen, carbon and sulphur isotope ratios in peat: the role of diagenessis and water regimes in reconstruction of past climates

We demonstrate that carbon, hydrogen and sulphur isotope ratios both, in the total peat (¹³C_tp=–25.52 to –28.27, D_tp=–78.67 and –109.24, ³⁴S=4.35 to 19.87), and in cellulose from the peat (¹³C_nc=–25.06 to –27.33 and D_nc=–92.43 to –118.02) are not affected by postdepositional changes. Therefore, the original isotope composition of plants are in general preserved in the peat and represent an archive of the past environmental variations. These can be supported by (i) good correlations between ¹³C_tp and ¹³C_nc, and between D_tp and D_nc, (ii) high horizontal homogeneity of ¹³C_tp and ¹³C_nc in the scale of one peat-bog – the same major factor(s) control(s) C isotopic ratios, (iii) no correlation between organic sulphur concentrations and ³⁴S value –³⁴S results from variations in the water level.This revised version was published online January 2005 with corrections to the title.     

The role of male vs male interactions in maintaining population dialect structure

Summary Nonmigratory populations of Whitecrowned Sparrows in coastal California exhibit dialects in territorial male songs that are stable in space and time. By field playback experiments, we tested a prediction from the hypothesis that male aggressive interactions prohibit mixing of song dialects. Playback of the home dialect Clear song to territorial males singing the Clear dialect resulted in less response than that given to playback of the neighboring Buzzy dialect. Response to both Buzzy and Clear dialect songs by target Clear males, however, was greater than that given to the Bodega dialect recorded 55 km away (Figs. 2 and 3). We conclude that these results are consistent with the hypothesis that male-male aggressive interactions play an important role in reducing dialect mixing.     

The evolution of cheating and selfish behavior

Summary We examine the necessary conditions for the spread of genes that determine selfish and cheating behaviors and the rate of spread of these genes through structured populations, in order to address the question of the invadability of altruistic systems by anti-social mutations. We find that, although cheaters always have a higher relative fitness than altruists within groups, population structures which permit the evolution of altruism also preclude invasion by anti-social mutations. These results are related to a discussion by Hamilton (1971) concerning the limits to the evolution of altruistic and selfish behaviors.     

On the evolution of male workers in the Hymenoptera

Summary The effects of brood sex-ratio (investment ratio) and the presence of laying workers on relatedness in the Hymenoptera are analyzed. It is shown that the conditions of brood composition that generate degrees of relatedness favorable to the evolution of eusocial type helping behavior among females will select against such helping behavior among males, and vice versa (see Fig. 1). Thus, societies in the Hymenoptera can be expected to have male workers or female workers, but not both. It is argued that the conditions leading to degrees of relatedness favorable to male helping are quite restrictive and unlikely to be met in haplodiploid species.The presence of laying workers is shown to lead to biasses in relatedness such that females may be selected to be workers even when the sex-ratio is male-biassed. This result sheds new light on a possible pathway to eusociality in the Hymenoptera. It is argued that offspring parasitism of the natal nest may have been important in the evolution of eusociality.     

Territory and group sizes in Eurasian beavers (Castor fiber): echoes of settlement and reproduction?

According to current theories of territoriality, an animal is expected to defend the smallest area that can provide resources for maximisation of reproduction, known as the economically defendable area. In group territorial species however, the strategies behind resource defence are likely to be more complex with corporate territoriality, cooperative breeding, delayed dispersal and intra-group competition all potentially playing a role. Here we examined group territoriality in a social herbivorous rodent, the Eurasian beaver Castor fiber. Beavers in our study do not inhabit economically defendable territories. Instead the sequence of arrival of pairs into unoccupied areas seems to play a more important role in determining the size of the territory, whereas group size is determined by past reproductive success. We argue that the settlement pattern and reproductive history have a lasting impact in the territorial system of beavers due to a combination of the low adult mortality, high dispersal costs, and avoidance of resource depletion.R.D. Campbell and F. Rosell contributed equally to this work     

Resource partitioning by reef corals as determined from stable isotope composition

The pattern of resource partitioning vs depth by corals collected in February 1983 from Jamaica and the Red Sea was determined from their stable carbon isotope composition. Observations were made on isolated zooxanthellae and corresponding algae-free animal tissue from eight species at four depths over a 50 m bathymetric range. Zooxanthellae ¹³C was high in shallow water and became lower as depth increased. This trend correlated significantly with the anual integrated photosynthetic rate. The trend is interpreted according to a depletion-diffusion hypothesis; in shallow water, at high rates of photosynthesis, metabolic CO₂ is nearly depleted and the supply of CO₂ from seawater bicarbonate is limited by diffusion. Since most of the available CO₂ is fixed, isotope fractionation is minimal. In deeper water, at lower rates of photosynthesis, metabolic CO₂ is ample, and isotope fractionation is greater. Animal tissue ¹³C was slightly lower than corresponding zooxanthellae values in shallow water. As depth increased the difference between zooxanthellae and animal tissue ¹³C increased and the latter approached the ¹³C of oceanic particulate organic carbon. These data suggest that carbon is translocated at all depths and that deep-water corals draw significantly on allocthonous sources of carbon.Contribution No. 436 of the Discovery Bay Marine Laboratory of the University of the West Indies     

Problems in the assessment of the package effect in five small phytoplankters

The magnitude of the package effect in five small phytoplankters [Thalassiosira sp., Clone 2601 (an unidentified eucaryote), Nannochloris atomus, Synechococcus Syn and Synechococcus WH 7803] was assessed by comparison of the absorption spectra of intact and disrupted cells. The package effect was considerably reduced with reductions in cell size and this was broadly in agreement with theoretical predictions based on Mie theory. However, the quantitative assessment of the package effect is confounded by an inability to assign attenuation (apparent absorption) measurements at =750 nm to either scattering or absorption. The magnitude of the apparent absorption at =750 nm was greatest with the smallest picoplankton species examined, and was reduced, but not eliminated, after cell disruption. Whilst the apparent absorption at =750 nm is commonly thought to be due to residual scattering losses, the available evidence does not exclude the possibility that this may be due in part to absorption by cells or cell constituents and this requires further examination. Although these difficulties are particularly evident with the small picoplankton species, there is no reason to expect that they will not complicate the assessment of the package effect in larger phytoplankton cells.     

Nutrition studies in the nauplius larva of Calanus pacificus (Copepoda: Calanoida)

Nauplii of Calanus pacificus were raised on a mixture of algae. Details of the mouth-parts, such as denticles, labial palps and lobes, setations and structure of the masticatory teeth were examined by scanning electron microscopy (SEM). Under the experimental conditions (15°C and 300 gC l^-1), exponential growth coefficients for the period Nauplii II–VI were 0.179 for carbon and 0.228 for nitrogen. C:N ratios dropping from 5.1 to 4.7. Growth was isochronous, each stage lasting 1.5 days. Respiratory losses were 15 to 19.6% of body carbon daily. Nauplii raised on a given alga showed higher rates of ingestion in the presence of this food, compared to nauplii switched to other algae. Minimal threshold concentrations for feeding were found, depending on the size of the food offered and ranging from 5.8 gC I^-1 for Lauderia borealis (28.7 m spherical diameter) to 47.1 gC 1^-1 for Chlamydomonas sp. (11.0 m). Unlike the Copepodite I stage, Nauplii II–VI larvae were not able to ingest small cells such as Isochrysis galbana (4.3 m), or very large ones such as Ditylum brightwellii (47.5 m) at more than maintenance rations. Below the critical concentration for maximal feeding, ingestion was clearly dependent on size of the cells offered, but the size-dependent relationship was different for diatoms and non-diatoms. Filtering rates increased from a threshold concentration to a maximal rate at about 50 gC 1^-1, and decreased at higher concentrations. Critical concentrations ranged from 125 gC 1^-1 for L. borealis to 1000 gC 1^-1 for Chlamydomonas sp. Maximal daily rations ranged between 100 and 150% of body carbon.     

Applying the discrete vortex method in environmental fluid mechanics: A study of the time-averaged near wake behind a circular cylinder

This work illustrates the discrete vortex method (DVM) as a tool for simulating environmental fluid mechanics problems involving transport in the wake of a bluff body. The DVM was used to model both the long-time-averaged and instantaneous features of flow past a circular cylinder. Simulations were performed for Re = 140, 000. Verification testing was accomplished by refining time-step and vortex element circulation. The DVM was validated through comparison with experimental data from Cantwell and Coles. Verification testing demonstrated that, while global convergence is not possible for an unsteady flow simulation, it is possible to have convergence to physical results. This verification entails identifying a range of parameters in which a flow containing coherent structures and matching physical conditions is attainable. Validation tests demonstrated excellent agreement between experimental and simulated results for time-averaged velocity and shear stress profiles, as well as Strouhal number.     

Reproductive effort in relation to maternal social rank in reindeer (<Emphasis Type="Italic">Rangifer tarandus</Emphasis>)

In polygynous mammals, high-quality females may increase their fitness by providing superior care to their offspring. Based on the agonistic interactions of female reindeer in an experimental herd during two consecutive years (1997 and 1998), we tested whether maternal social rank influenced: (1) winter body-mass change of females, (2) preparturition reproductive effort (measured as fecundity, the birth mass and the birth date of their calves), (3) preweaning maternal effort (measured as calves preweaning mortality, early preweaning and late preweaning growth rate and September body mass of calves), and (4) postweaning maternal effort (measured as calves body-mass change during their first winter). In the models, we included September females body mass as a covariate to separate the effects of maternal rank and body mass. We also tested whether the effect of social rank on maternal efforts was dependent on offspring sex. High-ranked females gained body mass whereas low-ranked females lost weight during the winter. Fecundity was higher and date of birth was earlier in high-ranked females than in subordinates, whereas no effect of females rank on birth mass of calves was found. Early preweaning growth rate and September body mass of calves increased with increasing females social rank, whereas late preweaning daily growth rate of calves was not influenced by females rank. Calves preweaning mortality was only influenced by year, which also explained most of the variance in the winter body-mass change of calves. The effects of females rank on the reproductive-efforts parameters studied were not specific to offspring sex. These findings suggest that females rank influences reproductive effort during the preparturition, as well as the preweaning, period, the effect being sex independent.Communicated by R. Gibson     

Aggregation and foraging behavior of whirligig beetles (Gyrinidae)

Summary Whirligig beetles aggregate in the daytime into dense single-and multispecies groups (rafts) of hundreds or thousands of individuals. On the 22km shoreline of Lake Itasca in northern Minnesota, these aggregations were on the average 0.8 km apart, and they were usually found day after day in the same ocations.Most beetles apparently do not home to the aggregation of their origin after dispersing at night because (a) the species composition of some aggregations changed greatly, and (b) paint-marked beetles (Dineutus horni) moved overnight from one aggregation as far as 4km, joining 11 of the 14 large (>300 beetles) D. horni groups on the lake.Throughout the night, the largest concentrations of beetles remained within 100m of the diurnal aggregation sites. Beetles reconvened into the compact rafts before daybreak, in part by following each other in sometimes long single files or trains. Their forward motion stopped after they joined large number of other beetles. We infer that following behavior enables those individuals that have dispersed from their original aggregations (during their nocturnal foraging) to find and join other aggregations before daylight.Naive fish ate the beetles despite their noxious secretions. However, fish living near rafting sites and feeding on insects on the water surface in daylight should soon learn to avoid the beetles. The rafting sites would then become safe places. We observed fish attacking only those beetles that had been either dispersed from their rafts or released into open water away from raft sites in the daytime. We speculate that the evolutionary significance of the aggregation behavior is related to predator (fish) avoidance.     

Mercury budget estimates for the State of New York

The development of two comprehensive modelling systems and a test application to estimate the mercury budget for the State of New York is discussed in the present study. The modelling tools developed are based on two well known meteorological modelling systems namely, RAMS and SKIRON/Eta. Both models have been modified and the developed modules describing the physico-chemical processes of mercury have been incorporated in each modelling system. Model calculations and measurements have been compared during the 14 to 26 August 1997 simulation period. Two different simulations were performed, scenario1 where all available sources of mercury were used and scenario2 where the New York State mercury sources were excluded. An attempt was made to identify and quantify critical gaps in the current understanding of regional scale transport deposition and fate of mercury in New York State, improving modelling capabilities and understanding of mercury as an air pollutant, and provide a tool for policy makers in mitigating the impacts of mercury pollution.     

Acute toxicity and sublethal behavioral effects of copper on barnacle nauplii (Balanus improvisus)

This study documents the effects of short-term (24h) sublethal copper exposures on undirected swimming activity and photobehavior of Balanus improvisus stage II nauplii. All Cu treatments were static, with temperature and salinity conditions at 20°C and 15 or 30. The 24h LC 50 estimate for Cu is 88 ppb at 15 and >200 ppb at 30. Sub-lethal Cu concentrations cause reductions in swimming speed, which decrease progressively with increasing Cu dose. At 50 ppb Cu, this was significant primarily at light intensities below the phototactic threshold. At higher Cu concentrations, significant reductions in mean linear velocity occurred at most light intensities tested. At 30, 50 and 100 ppb Cu also reduce the positive phototactic response and 150 ppb Cu causes reversal of phototaxis at optimal intensities. Photokinesis is reduced at 100 ppb Cu and disappears at 150 ppb Cu. At 15, the behavioral effects of 50 ppb Cu resemble those occurring with 150 ppb Cu at 30. Swimming speed and photobehavior show promise as sensitive behavioral indicators of copper toxicity. Additional research is required to determine if these responses apply to a broad range of pollutants and to other planktonic organisms. There is also a need to further evaluate the significance of these behavioral effects ecologically.Contribution No. 181 from the EPA Environmental Research Laboratory, Narragansett, RI 02882, USA     

Discrimination between aluminium held within vegetation and that contributed by soil contamination using a combination of Electron Probe Micro Analysis (EPMA) and Inductively Coupled Plasma-Atomic Emission Spectrometry (ICP-AES)

Aluminium concentrations in NIST (formally NBS) Standard Reference Material Tomato Leaves and Citrus Leaves were determined by ICP-AES after acid digestion. The normal total acid attack (nitric and perchloric acids) gave very low recovery (40%) of Al in NIST tomato leaves, but not for citrus leaves. The contribution of Al from Al-rich soil particles in both tomato leaves and citrus leaves was estimated semi-quantitatively by computer-controlled EPMA. The Al held in these particles corresponded in approximate concentration to the shortfall between the acid soluble component determined in this study and the certified value. Analysis by EPMA can provide, therefore, a method of estimating Al contributed by soil contamination to plant materials. The limitations of the NIST tomato and citrus leaf reference material for the validation of methods for the determination of Al in vegetation are demonstrated.