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.     

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.     

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.     

Stability of eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis> L.) depth limits: influence of habitat type

Seagrass meadows are generally considered stable although few studies have specified and tested this statement. On the basis of a large monitoring dataset from Danish coastal waters, we aimed to test whether the stability of deep eelgrass populations changes along a eutrophication gradient extending from inner bays over outer bays to open coastal waters. We defined stability in terms of the stability properties of constancy, resilience, and/or persistence. Our data allowed us to investigate the stability property constancy expressed as temporal variability in eelgrass depth limits over a time scale of 10 years. We hypothesised that colonisation depths were large and relatively constant along open coasts because of low and relatively constant levels of nutrients and turbidity. Conversely, colonisation depths were hypothesised to be low and variable in protected bays due to higher and more variable levels of nutrients and turbidity. We found that depth limits increased from inner bays towards open coastal waters, matching declines in nutrient concentration and increases in water clarity and oxygen concentration. Stability expressed as constancy of depth limits did not differ significantly between habitat types, and neither did stability of physicochemical variables. However, when data from all habitat types were analysed together, they showed that eelgrass populations at the depth limit were significantly more constant and thus, in this respect, more stable when occurring in deep waters as compared to shallow waters. Areas of good water quality may thus obtain the double benefit of deeper-growing and more stable eelgrass populations. The most likely reason why this pattern did not appear at habitat-type level is that the habitat types studied represented wide spatial variation in water quality and depth limit. We conclude that the question of whether eelgrass populations are stable depends on the stability property and the ecological situation in question. Populations may be considered unstable in terms of inter-annual variation but stable in terms of long-term persistence. Therefore, the common statement that eelgrass populations are stable is not universally true.Communicated by M. Kühl, Helsingør     

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     

Calcification in the maerl coralline alga Phymatolithon calcareum: Effects of salinity and temperature

The coralline alga Phymatolithon calcareum was dredged from 13 m in the Kattegatt, Baltic Sea, in December, 1980, and its rate of calcification was measured by ⁴⁵Ca⁺⁺-uptake methods. Light-saturated calcification rates at 5°C ranged from 15.8 g CaCO₃ g^-1 dry wt h^-1 for the basal parts of the plants to 38.7 g CaCO₃ g^-1 dry wt h^-1 for the tips. These age gradients were not apparent when calcification rates were expressed on the basis of surface area. Experiments with salinity (10, 20, 30) and temperature (0°, 5°, 10°, 20°C) indicated that optimum conditions for calcification were at 30 S and at temperatures above 10°C. Salinity had a greater influence on calcification rate than did temperature, and there was a positive relationship between salinity and calcification rate at all temperatures. In 6 mo old cultures, salinity was again the important factor, with all plants remaining healthy at 30 except those at the highest temperature (20°C). These trends, and the low calcification rates at 10S (4.6 g CaCO₃ g^-1 dry wt h^-1 at 5°C to 8.6 g CaCO₃g^-1 dry wt h^-1 at 20°C) suggest that low salinity may be the explanation for the general absence of P. calcareum from the brackish waters of the Baltic Sea. Short-term experiments in which salinity was kept constant while Ca⁺⁺ concentration was altered, and experiments in which salinity was varied and Ca⁺⁺ concentration kept constant, suggest that it is the calcium ion concentration and not salinity per se which affects calcification rates.     

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.     

Effects of nitrogen and phosphorus enrichement on in vivo symbiotic zooxanthellae of Pocillopora damicornis

The reef coral Pocillopora damicornis (Linnaeus) was grown for 8 wk in four nutrient treatments: control, consisting of ambient, unfiltered Kaneohe Bay seawater [dissolved inorganic nitrogen (DIN, 1.0 M) and dissolved inorganic phosphate (DIP, 0.3 M)]; nitrogen enrichment (15 M DIN as ammonium); phosphorus enrichment (1.2 M DIP as inorganic phosphate); and 15 M DIN+1.2 M DIP. Analyses of zooxanthellae for C, N, P and chlorophyll a after the 8 wk experiment indicated that DIN enrichment increased the cellular chlorophyll a and excess nitrogen fraction of the algae, but did not affect C cell^-1. DIP enrichment decreased both C and P cell^-1, but the decrease was proportionally less for C cell^-1. the response of cellular P to both DIN and DIP enrichment appeared to be in the same direction and could not be explained as a primary effect of external nutrient enrichment. The observed response of cellular P might be a consequence of in situ CO₂ limitation. DIN enrichment could increase the CO₂ (aq) demand by increasing the net production per unit area. DIP enrichment could slow down calcification, thus decreasing the availability of CO₂ (aq) in the coral tissue.Hawaii Institute of Marine Biology Contribution No. 920     

What is the sex ratio of harpacticoid copepods in the deep sea?

The sex ratios of deep-sea harpacticoids have been thought to be greatly skewed toward females. The representation of males is notably more equitable (approximately 1:2) at three deep-sea sites (San Diego Trough: 32°52.4N, 117°45.5W, 1 050 m, January 1987; Porcupine Seabight: 51°36.85N, 12°57.30W, 1 369 m, August 1984; northwest Atlantic, 40°27N, 62°20W, 4 820 m, July 1982, June 1983). This ratio conceals extensive variation at the species level. For some species, males are unknown. For other species, males are more numerous than females. Sampling bias against males can occur and may explain the more extreme reports of male rareness in the literature.     

Effect of heavy metals (Zn,Hg, Cu,Cd, Pb,Ni) on the length growth of Mytilus edulis

The shortterm (10–22 d) effect of Zn, Hg, Cu, Cd, Pb, and Ni on the length growth of Mytilus edulis is studied. Significant reductions of growth rate was found at 0.3 g Hgl^-1, 3 g Cul^-1, 10 g Znl^-1, and 10 g Cdl^-1 added to the local sea water, while concentrations of up to 200 gl^-1 of Pb and Ni had no effect on the growth. With exposure to Cu and Zn, there was a linear reduction in growth rate with increasing metal concentration up to about 6 g Cul^-1 and 100 g Znl^-1. Above these levels, growth stopped with Cu, while with Zn it was stabilized at about 20% of control growth. When Hg and Cd were added, a curvilinear relationship between growth and metal concentration is indicated. With Hg, growth rate is nearly zero above 3–4 g Hgl^-1, while the growth rate was 50% of control after 10 d of exposure to 100 g Cdl^-1. At 2 g Cdl^-1 there was a significant stimulation of length increase. Observed EC₅₀-values for growth were 0.3–0.4 g Hgl^-1, 3–4 g Cul^-1, 60 g Znl^-1, and 100 g Cdl^-1.     

Importance of the fatty acids 20:5ω 3 and 22:6ω 3 in the diet of plaice (Pleuronectes platessa) larvae

The requirement of plaice (Pleuronectes platessa L.) larvae for the fatty acids 20:5 3 and 22:6 3 was studied. Larvae were reared from first feeding to beyond metamorphosis on Artemia sp. nauplii (EG brand), whose nutritional content had been manipulated by enrichment. Some larvae were fed Artemia rich in 20:5 3 and 22:6 3, and others were fed nauplii with no 22:6 3 and low levels of 20:5 3 (1.9% of the total fatty acids). The differences in diet had no significant difference on growth or survival of the plaice larvae. The results indicate that the plaice larvae in this study did not require 22:6 3 in their diet, even when the levels of 20:5 3 were low.     

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.     

Environmental Geochemistry of Soils and Waters of Susaki Area, Korinthos, Greece

Soil and water samples were collected from the Susaki area of Korinthos and analysed for heavy metals in order to evaluate their environmental impact. The geology of the studied area includes ultrabasic rocks and Neogene and Quaternary deposits whereas magnesite veins are found within the ultrabasic rocks. In the north part of the studied area post volcanic emissions of H₂S, CO₂ and H₂O vapor continue to the present day. All the samples were analysed for heavy metals by the ICP method. The element ranges (in g g^–1) for soil samples are: Cu 11–63, Pb 5–256, Zn 21–604, Ni 183–2665, Co 12–124, Mn 456–1434, As 5–104, Sr 44–730, V 21–84, Cr 163–2346, Ba 48–218, Zr 3–41, Y 3–13. The metals Pb, Zn, Ni, Co, Cr, Fe, Cu, Mn, As and Sr are enriched in the Susaki soils. The element ranges for water samples are: Cu 65–103ppb, Pb<10ppb, Zn<5ppb, Ni 21–163ppb, Co 2–12ppb, As<30ppb, Cr<20ppb, Ba 36–785ppb, Sb<10ppb, W<10ppb, Bi<30ppb, Mn 0.0–0.9 g g^–1, Fe 0.01–0.22 g g^–1, Na 843–3076 g g^–1, K 98–278 g g^–1, Si 39–65 g g^–1, P 0.1–0.2 g g^–1. There is a natural pollution of soils with elevated concentrations of Ni, Co, Mn, Fe and Cr due to the presence of ultrabasic rocks. Another natural case of As pollution of soils is due to the volcanic activity and the geothermal field in the area. The geochemical data of ground waters and also the ¹⁸, D data showed a mixing in different proportions between sea water and meteoric water.     

13C/12C ratios and the trophic importance of algae in Florida Syringodium filiforme seagrass meadows

Over 380 stable carbon isotope (¹³C) analyses made during 1981–82 showed that Syringodium filiforme Kutz seagrass meadows in the Indian River lagoon of eastern Florida have food webs based on algal rather than seagrass carbon. Seagrasses averaging approximately-8 were isotopically distinct from algae epiphytic on seagrass blades (X=-19.3) and particulate organic matter in the water column X=-21.6. ¹³C values of most fauna ranged between-16 and-22, as would be expected if food web carbon were derived solely from algal sources. These results counter the idea that seagrass detritus is the dominant carbon source in seagrass ecosystems. Two factors that may contribute to the low apparent importance of seagrass in the study area are high algal productivities that equal or exceed S. filiforme productivity and the high rates of seagrass leaf export from meadows.     

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.     

A Preliminary Study on Soil–Gas Radon Geochemistry According to Different Bedrock Geology in Korea

Three study areas of Kwanak campus (Seoul National University), Boeun (Choongbuk) and Gapyung (Kyonggi) were selected and classified according to their bedrock types in order to investigate soil–gas radon concentrations. The mean values of radon concentrations decreased in the order of Gapyung (40BqL^–1) > Kwanak campus (30BqL^–1) > Boeun (22BqL^–1), and decreased in the order of granite gneiss > banded gneiss > granite > black slate–shale > mica schist > shale–limestone > phyllite schist according to bed rock types. Variation in radon emanation with water content in soils and with soil grain size was assessed by the modified Morse (1976) 3min method. Soil–gas radon concentrations increased with increasing water content in the range of less than about 6–16wt.%, but decreased above 6–16wt.%. Radon concentrations also increased with decreasing soil grain size. Radioactivity analysis of radionuclides of ²³⁸U series in some soil samples indicated their possible radioactive disequilibrium between ²²⁶Ra and ²³⁸U due to the differing geochemical behaviour of intermediate radionuclides. Thus, a radioactive isotope geochemical approach should be necessary for soil–gas radon study.     

Le système assimilation-régénération des sels nutritifs dans les eaux superficielles de l'océan Austral

Low primary-productivity levels in high-nutrient environments of the Southern Ocean are not yet well understood. An accurate knowledge of nutrient assimilation by phytoplankton, at the base of the pelagic food-web, therefore appears to be essential. A study of the mesoscale hydrological structure and chemical observations in the upper layers of the Indian sector of the Antarctic Ocean (Cruise MD-25 FIBEX of M.S. Marion Dufresne, January–February 1981) permitted us to develop a method for estimating nutrient consumption by phytoplankton. After strong vertical mixing in winter, the homogeneous Antarctic Surface Water (ASW) is, in summer, divided into two parts by a well-formed pycnocline. In the upper layer, corresponding approximately to the photic zone, we observed nutrient depletion resulting from photosynthetic activity, while the underlaying thermal minimum layer displayed unaltered winter characteristics, including nutrient concentrations typical of winter surface-water conditions. Taking into account the nutrient depletion in summer, we calculated the assimilation ratios for Antarctic phytoplankton as follows: ([NO ₃ ^- lower]-[NO ₃ ^- upper]):([PO ₄ upper])=N:P=11.2±2.4 and N:Si=0.27±0.05. These ratios are lower than the classical Redfield ratios (N:P=16 and N:Si=1), but are in good agreement with the elementary composition of phytoplankton samples collected during the same cruise. Moreover, they agree with previously published data on stoechiometric determinations of particulate matter in surface-water samples. Besides, in the transition layer between the lower layer and Warm Deep Water (WDW), the calculated mineralization ratios were slightly higher than the assimilation ratios: N:P=14.6 and N:Si=0.37. From these ratios, it would appear that, in these surface layers of the Southern Ocean (down to 250 m), assimilation-regeneration mechanisms operate in such a way that phosphate and silicate contents decrease much more than nitrate content during the northward drift of the surface waters. The determination of assimilation ratios taking into account the nutrient depletion of the upper layer allowed us to evaluate the carbon net-production which integrates time-space variations in the photic layer over the whole early summer period. We estimated a production rate (0.4 g C m^-2 d^-1) of the same order of magnitude as the mean value obtained by the ¹⁴C method (0.2 g C m^-2 d^-1). The theoretic silicium consumption by phytoplankton along a meridian transect, calculated using an estimated assimilation ratio of C:Si=1,85, indicates that the marked south-north decrease in silicate concentration in the Antarctic Surface Water would mainly result from biological activity. The silicate concentrations thus calculated were in good agreement with concentrations measured along a previous transect across the Southern Ocean.

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Le système assimilation-régénération des sels nutritifs dans les eaux superficielles de l'océan Austral

     

Carbon-isotope ratio gradients in western arctic zooplankton

Zooplankton from 87 stations in the Bering, Chukchi and Beaufort Seas sampled in 1985, 1986 and 1987 showed a geographic gradient in stable carbon-isotope ratios (¹³C). The zooplankton most depleted in ¹³C were found in the central and eastern Beaufort Sea and those most enriched were from the Bering and Chukchi Seas. Average ¹³C values ranged from-20.9 to-26.7 for copepods and from-19.4 to-25.1 for euphausiids. Euphausiids show a minimum of 1.0 enrichment relative to copepods throughout the study area. Relative biomasses of the major zooplankton taxa varied significantly across the Alaskan Beaufort Sea in October 1986, with euphausiids dominating in the west and copepods in the east. These differences in taxonomic composition affected the weighted ¹³C values for total zooplankton and may produce an even more pronounced geographic gradient in zooplankton ¹³C than that found within a single taxon. The bowhead whale Balaena mysticetus migrates between wintering areas in the Bering Sea and summering areas in the Beaufort Sea and feeds over this geographic range. The zooplankton ¹³C gradient is the probable source of ¹³C oscillations found along the baleen plates of this planktivore.     

Carbon fixation in marine phytoplankton: carboxylase activities and stable carbon-isotope ratios; physiological and paleoclimatological aspects

We measured the activity of three carboxylases: RuBP carboxylase, PEP carboxylase and PEP carboxykinase of marine phytoplankton species in culture and in natural communities. Activities of the three carboxylases were measured simultaneously with stable carbon-isotope ratios. The enzymatic activities have been used to estimate the importance of carboxylation and its impact on the ¹³C:¹²C ratio (expressed as ¹³C). The marine phytoplankton species in culture were Fragilariopsis kerguelensis, Nitzschia turgiduloides, Skeletonema costatum, Phaeodactylum tricornutum, Isochrysis galbana, Dunaliella marina, and Prorocentrum micans, and the field samples were collected from different depths off the coast of Portugal (August/September, 1981). Our results indicate that, as in terrestrial plants, the ¹³C value is a good indicator of the extent of carboxylation. RuBP carboxylase activity was always predominant, whereas the ¹³C value never reached values typical of the C₄ pathway. The carboxylases could be PEP carboxylase (in dinoflagellates) or PEP carboxykinase (in diatoms). carboxylation increased at the end of the exponential growth phase in a diatom culture and with increased biomass in natural samples. We interpret these increases as an adaptative response mechanism to poor environmental conditions, especially to low light intensity.     

Precision and accuracy of stable isotope signals in otoliths of pen-reared cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) when sampled with a high-resolution micromill

A high-resolution sampling technique for stable-isotope measurements in otoliths is now available, but little is known about the precision and accuracy of the results when using this technique. Otolith samples of pen-reared cod (Gadus morhua L.) with well-defined temperature history were obtained using this high-resolution sampling technique. Samples from different-aged cod otoliths were collected in two yearly increments corresponding to the calendar years 1994 and 1995, and the stable-oxygen (¹⁸O) and -carbon (¹³C) isotope compositions of the samples were measured. Otolith ¹⁸O values had a clear seasonal pattern in response to the seasonal water temperature. Otolith material deposited in the same calendar years of different-aged fish showed similar ¹⁸O values. High precision of the temperature estimate using otolith ¹⁸O values was obtained when comparing results from different otoliths. The accuracy of the temperature estimates, relative to measured mean monthly water temperature, was also high, but reduced sampling resolution in the otoliths significantly reduced the accuracy due to attenuation of the ¹⁸O signals. We found that the otolith ¹³C values had a cyclic pattern, roughly in phase with the ¹⁸O values. This was most likely caused by temperature-increased metabolism. Indications of age-specific otolith ¹³C values are also presented.Communicated by M. Kühl, Helsingør