Differential increment-deposition rate in embryonic statoliths of the loliginid squid Loligo vulgaris

 Apart from one study that reported growth of less than one increment per day in statoliths of the squid Alloteuthis subulata, most studies so far have presumed that one increment was laid down per day in the statoliths of the squid species they examined. The present study provides evidence of differential daily growth rates in embryonic statoliths of the squid Loligo vulgaris Lamarck, 1798, thus confirming a previous report for A. subulata. Incremental growth rates of L. vulgaris statoliths differ as a function of temperature. Squid embryos were incubated in the laboratory at three temperatures (12.0, 15.5 and 21.1 °C), and tetracycline staining was used to follow statolith growth. This growth slowed in squid exposed to the lowest temperature, but recovered when the squid were returned to warm conditions, indicating statolith adaptation. Statolith growth rate after incubation at 12 °C was 1.3% d⁻¹ and reached 6.1% d⁻¹ for squids exposed to 21.1 °C. Statoliths from embryos incubated at 15.5 °C yielded a rate of 1 increment d⁻¹ and a mean daily growth of 2.2 μm in the dorsal dome area of the statolith. In contrast, the slow growth of statoliths incubated at 12 °C yielded a mean daily growth of only 0.9 μm in the dorsal dome and the readings resulted in a less-than-daily increment-deposition rate. Received: 9 October 1999 / Accepted: 30 March 2000     

Effect of temperature on statolith growth of the European squid Loligo vulgaris during early life

Over the past decade, statolith interpretation has resulted in a major advance in our knowledge of squid population-dynamics, but the way in which environmental conditions affect the statolith increment-deposition ratio remains virtually unknown. The object of the present study was to determine the effect of temperature on this process, using tetracycline marks to validate statolith growth in Loligo vulgaris Lamarck, 1798 under rearing conditions equivalent to severe winter (11 °C) and summer (19 °C) temperature regimes. Tetracycline marking was performed every 10 d (at 10, 20, 30, 40, 50 and 60 d of age). The newly hatched squid paralarvae were slightly smaller in summer than those hatched in winter. Survival rates were similar in both cultures, but growth rates (wet mass) of summer squids were double those in winter. At hatching, statoliths were already longer in the summer squids, and growth rates were 2% d⁻¹ as opposed to 0.9% d⁻¹ for winter statoliths. For the dorsal dome area of the statolith, where more increment counts were made, statolith growth was of 3.25 μm d⁻¹ in summer, and daily increment deposition was confirmed in 87% of the statoliths. The slow growth of statoliths at winter temperatures yielded a mean growth of 1.1 μm d⁻¹– insufficient to discern the increments using light microscopy. Subsequent SEM observation enabled only 21% of the winter statoliths to be read; these also indicated a deposition rate of one increment d⁻¹. Since the life span of L. vulgaris is ≃1 yr, squids will experience at least one winter during their life cycle, and this might be visible on the statolith. Received: 28 June 1999 / Accepted: 20 December 1999     

The influence of ration level on growth and statolith increment width of the tropical squid Sepioteuthis lessoniana (Cephalopoda: Loliginidae): an experimental approach

Juvenile squids were grown in individual 2.6-l floating enclosures and were fed either a high- or a low-ration diet of fish and the crustacean Acetes. Squids were maintained for a maximum of 44 days in two experiments. The high-ration individuals reached a significantly larger size in both experiments (27, 25.5 mm mean mantle length, ML) compared to their low-ration siblings (19 mm mean ML) in both experiments. The statolith increment widths prior to the start of the experiment were significantly wider (between 3 and 4 μm) compared to the increment widths after the start of the experiment (between 2 and 3 μm) both for the low- and the high-ration squids. High-ration squids also had significantly wider increments and larger statoliths than their low-ration siblings. Even though we detected consistent trends in daily statolith increment widths for the different feeding regimes, we could not detect variation in increment widths at a daily level of resolution (i.e. as a result of differences in day-to-day food intake at an individual level). This was probably due to the relatively consistent diet experienced by each individual. These experiments revealed that ration level influences squid growth rate, statolith size and daily statolith increment width. Received: 30 March 2000 / Accepted: 30 October 2000     

Age and growth of the mesopelagic squid Ancistrocheirus lesueurii (Oegopsida: Ancistrocheiridae) from the central-east Atlantic based on statolith microstructure

Statolith microstructure was studied in 56 Ancistrocheirus lesueurii (25 to 423 mm of mantle length, ML) caught in the central-east Atlantic. Statolith growth increments were grouped into three main growth zones, distinguished mainly by increment width. The second transition in the statolith microstructure (from Zone 2 to Zone 3) coincides with the life history shift from epipelagic and upper mesopelagic to a bathyal habitat. Second-order bands (mean 27.65 growth increments) and sub-bands (mean 13.6 growth increments) within statolith microstructure appeared to be related to the lunar cycle. Striking sexual dimorphism is reflected in the age and growth rates: males live ca. 1 yr, while females only start maturing at this age and obviously live >1.5 yr. A. lesueurii is a slow growing squid, attaining 25 to 30 mm ML at the age of 100 d. After ontogenetic migrations into bathypelagic waters at ML > 30 to 35 mm, growth rates gradually decrease to the minimum known values for squids. Based on back-calculated hatching dates, A. lesueurii hatches throughout the year with a peak between November and March. Received: 28 August 1996 / Accepted: 31 January 1997     

Strontium/Calcium ratios in statoliths of the neon flying squid, Ommastrephes bartrami (Cephalopoda), in the North Pacific Ocean

Strontium to calcium ratios were observed along longitudinal sections of statoliths of nine neon flying squid, Ommastrephes bartrami (LeSueur, 1821), including three mature females (422 to 454 mm mantle length, ML; 207 to 306 d old) obtained from the North Pacific (27–35°N; 144–150°E) during winter and six immature males and females (187 to 226 mm ML; 126 to 164 d old) collected from 39°N; 145°E and 39°N; 169°W during summer. The distances between the nucleus (core) and the edge of the dorsal dome were approximately 660 to 690 μm in mature females and 450 to 510 μm in the immature squid. Sr/Ca ratios were determined at intervals of 30 μm between the nucleus and edge of the dorsal dome. Sr/Ca ratios were higher in areas near the nuclei and peripheral portions of the dorsal dome than in the middle portions of the statoliths (270 to 420 μm from the nuclei, corresponding to ages of 60 to 90 d) in mature females; thus a U-shaped pattern was evident. Sr/Ca ratios in the six immature squid decreased from nucleus to the dorsal dome; in three squid the ratios slightly increased toward the dorsal dome edge. The observed Sr/Ca ratios in immature squid were considered to represent younger portions of the U-shaped pattern. In the present study we discuss this pattern in relation to environmental and biological conditions of O. bartrami, which undertakes seasonal migrations between spawning grounds in the Subtropical Domain and feeding grounds in the Subarctic Domain and Transitional Zone in the North Pacific Ocean. Although Sr/Ca ratios are potentially affected by ambient water temperature and ontogenetic conditions, including somatic growth and statolith growth, it was impossible to evaluate each environmental and biological effect separately, as variations in these factors are complicated and effects could be interdependent. Received: 11 April 1997 / Accepted: 27 December 1997     

Chemical and structural characteristics of statoliths from the short-finned squid Illex illecebrosus

Statoliths of the short-finned squid Illex illecebrosus were chemically analyzed to define their chemical composition and surveyed by scanning electron microscope to differentiate internal structural patterns. X-ray diffraction data demonstrated that I. Illecebrosus statoliths were composed principally of CaCO₃ in the aragonite crystal form. The crystals occurred in a protein matrix to form incremental patterns which radiated from the nucleus to the edge of the statoliths. The protein matrix comprised approximately 5% of the statolith by weight. The protein was principally composed of acidic amino acids. A high abundance of aspartic acid in the protein matrix indicated that the matrix would function as a template in the initiation and acceleration of the crystal growth of CaCO₃. The rhythmic microstructural patterns, constructed of aragonite crystals in the protein matrix, were suggested to be daily in formation and subsequent growth estimations were in agreement with known life history information. The stable isotopic composition of the carbonate of I. illecebrosus statoliths suggested that oxygen may be deposited in isotopic equilibrium with the surrounding environment while carbon appeared to be related to biological processes. The information recorded in the statoliths as incremental growth and stable isotopic composition could provide valuable insights into the ecological history of squid.     

Influence of temperature and salinity on the trace element incorporation into statoliths of the common cuttlefish (<Emphasis Type="Italic">Sepia officinalis</Emphasis>)

The use of statolith chemistry to trace migration pathways and distinguish populations of cephalopods is based on the assumption that the elemental composition of statoliths is influenced by physicochemical properties of the ambient environment. However, such influences have not been investigated experimentally up until now. This study presents the first microchemical analyses of cephalopod statoliths obtained from laboratory experiments under different controlled temperature and salinity conditions. Our results show that statolith chemical composition is strongly related to both salinity and temperature in ambient waters. The Ba/Ca ratio is negatively related to temperature and shows no relation to salinity. The I/Ca ratio is positively related to temperature and negatively to salinity. No Sr/Ca relation was found to either salinity or temperature, suggesting that the well-established proxy strontium is not as useful in cephalopod statoliths as in other biomineralized aragonites. Microanalysis of trace elements, however, shows an enormous potential for field studies on distribution, migration and stock separation of cephalopods. Furthermore, Synchrotron X-ray Fluorescence Analysis is introduced as a promising novel method for statolith analysis, providing a spatial resolution of typically 10–15 μm combined with detection limits down to 0.5 ppm.     

Geographic variation in statolith trace elements of the Humboldt squid, Dosidicus gigas, in high seas of Eastern Pacific Ocean

We applied the solution-based inductively coupled plasma mass spectrometry to quantify trace elements in statoliths of Humboldt squid, Dosidicus gigas, collected from the high seas off Chile, Peru, and Costa Rica by Chinese squid jigging vessels during 2007–2009. All squid samples were aged and their spawning dates were back-calculated based on daily increments in statoliths. The most abundant trace elements in the whole statolith were Ca and Sr followed by other elements in the order of Fe, Mg, Zn, Ba, Cu, Mn, Ni, Al, Cr, Co, and U. Significant differences in Mn and Sr were found among samples from the three regions. Sr, Ni, Mn, and Co contributed significantly to the discrimination among the regions, with Co responsible for explaining most of the variation, followed by Ni, Mn, and Sr. Squid from the high seas off Costa Rica could be separated from those off Peru and Chile mostly due to the differences in Ni, Sr, and Co, while samples off Peru and Chile could be distinguished mainly because of differences in Mn and Co. Discriminant function analysis suggested that the overall cross-validated classification rate was 85.6 % with samples off Chile having the highest correct identification rates and samples off Costa Rica having the highest false classification rates. Significant positive relationships were found between sea surface temperature (SST) and Cr/Ca, Mn/Ca, and U/Ca, and there was a negative relationship between SST and Cu/Ca, Sr/Ca. This study suggests that the spatial difference in trace elements of statolith can be used to separate geographic populations of D. gigas and that elements having significant relationships with SST can be considered as natural indicators of ambient temperature.     

Food effects on statolith composition of the common cuttlefish (Sepia officinalis)

The concentration of trace elements within cephalopod statoliths can provide a record of the environmental characteristics at the time of calcification. To reconstruct accurately the environmental characteristics at the time of calcification, it is important to understand the influence of as many factors as possible. To test the hypothesis that the elemental composition of cuttlefish statoliths could be influenced by diet, juvenile Sepia officinalis were fed either shrimp Crangon sp. or fish Clupea harengus under equal temperature and salinity regimes in laboratory experiments. Element concentrations in different regions of the statoliths (core–lateral dome–rostrum) were determined using laser ablation inductively coupled plasma mass spectrometry (LA- ICPMS). The ratios of Sr/Ca, Ba/Ca, Mn/Ca and Y/Ca in the statolith’s lateral dome of shrimp-fed cuttlefish were significantly higher than in the statolith’s lateral dome of fish-fed cuttlefish. Moreover, significant differences between statolith regions were found for all analysed elements. The fact that diet adds a considerable variation especially to Sr/Ca and Ba/Ca must be taken into account in future micro-chemical statolith studies targeting cephalopod’s life history.     

Age and growth of Electrona antarctica (Pisces: Myctophidae), the dominant mesopelagic fish of the Southern Ocean

Numerically and in biomass, the lanternfish Electrona antarctica is the dominant fish in the vast pelagic region of the Southern Ocean bounded on the north by the Antarctic Convergence and in the south by the Antarctic continental shelf. It is an important krill predator, and in turn is important in the diets of flighted and swimming seabirds. Further, it is the southernmost and coldest-dwelling representative of the globally distributed fish family Myctophidae. The present study was undertaken to estimate the species' growth rate and average life span, to incorporate the information in a basic energy budget, and to compare the growth of E. antarctica with more northerly confamilials. Fishes were aged using primary growth increments that were resolved on sagittal otoliths using three sequential techniques: thin-section grinding and polishing, etching, and scanning electron microscopy (SEM). Based on increment width (0.8 to 1.2 μm), continuity, and previous studies on confamilials, the microincrements were assumed to be deposited on a daily basis. Montages of SEM photomicrographs were constructed for each sagitta to allow the daily rings to be counted over the entire life span of 31 individuals representing the entire size range of the species. Results suggest a larval stage of 30 to 47 d and a maximum life span of 3.5 yr, with females growing faster than males in the last 1.5 yr of life and reaching a larger maximum size. Construction of a simple energy budget using the best information available suggests that a surplus of energy is available to support the observed growth rates (0.05 to 0.07 mm d⁻¹). The results of the present study contrast markedly with previous estimates of an 8 to 11 yr maximum age for E. antarctica. These results provide important data addressing the ecology and population dynamics of the pelagic Antarctic ecosystem. E. antarctica is the end-member species in the continuum of vertically migrating myctophids that extend from the equator to the polar circle. Its growth rate is consonant with that of all other myctophid species examined using primary growth increments to determine age. The present study, in conjunction with earlier studies, suggests that growth rates of mesopelagic species are far higher than previously thought. Received: 12 September 1997 / Accepted: 25 July 1998     

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     

The effects of an extraordinary El Niño / La Niña event on the size and growth of the squid<Emphasis Type="Italic"> Loligo opalescens</Emphasis> off Southern California

The population structure of the California market squid Loligo opalescens was studied for the Channel Islands region off Southern California between June 1998 and March 2000. During this time Californian waters were exposed to an extraordinary El Niño event that was possibly the most dramatic change in oceanographic conditions that occurred last century. There was then a rapid transition to record cool La Niña conditions. Statolith increments were used to determine age parameters and increment periodicity was validated for the first 54 days of life. Based on statolith increment counts, the oldest males and females were 257 and 225 days respectively and individuals matured as young as 129 and 137 days respectively. No distinct hatching period was detected. There was a general trend of increasing body size throughout the study period. Squid that hatched and grew through the El Niño were strikingly smaller and had slower growth rates compared to squid that grew through the La Niña. This was related to oceanography and associated productivity. There was a positive correlation between squid mantle length and upwelling index and a negative correlation between mantle length and sea temperature. The 'live-fast die-young' life history strategy of squid makes them ideal candidates for following the effects of the dramatic changes in oceanographic conditions off California. We propose that squid can serve as ecosystem recorders and productivity integrators over time and space and are useful organisms to tie oceanography to biology.     

The gastropod statolith: a tool for determining the age of<Emphasis Type="Italic"> Nassarius reticulatus</Emphasis>

The microstructure, shape and appearance of the growth rings in statoliths of Nassarius reticulatus (L.) were investigated. This species possesses two statocysts, each containing a single spherical statolith of calcium carbonate of up to 0.22 mm in diameter in the largest animals. The relationship between statolith diameter (SD) and total shell height (TSH) is exponential [ln(TSH)=26.3SD–0.842], although the function is site specific. Statoliths of the largest whelks (>29 mm) contained three or four clearly defined rings, corresponding to TSH values of ~1.1, 4.6–5.3, 12.0–13.5 and 18.5 mm, respectively. The first ring likely represents the metamorphic ring that was deposited at the time of larval metamorphosis when the post-larval whelk adopted a benthic lifestyle. The estimated size of the whelks at formation of the second, third and fourth statolith rings closely matched the TSH inferred from the shell rings. It is concluded that the patterns of growth rings present in statoliths can provide information about the age and growth of N. reticulatus.Communicated by O. Kinne, Oldendorf/Luhe     

Comparison of the statolith structures of <Emphasis Type="Italic">Chironex fleckeri</Emphasis> (Cnidaria,Cubozoa) and <Emphasis Type="Italic">Periphylla periphylla</Emphasis> (Cnidaria,Scyphozoa): a phylogenetic approach

The rhopalia and statocysts of Periphylla periphylla (Péron and Lesueur in Ann Mus Hist Nat Marseille 14:316–366,1809) and Chironex fleckeri Southcott (Aust J Mar Freshw Res 7(2):254–280 1956) were examined histologically and showed several homologous characteristics. Differences in sensory area distribution could be connected to a slightly different functionality of equilibrium sensing. In P. periphylla, the statoliths (crystals) grow independently of each other; whereas in C. fleckeri, one large crystal covers the smaller ones. The structures of both statoliths were examined in detail with single-crystal diffraction, microtomography and diffraction contrast tomography. The single compact statolith of C. fleckeri consisted of bassanite as was previously known only for other rhopaliophoran medusae. An origin area with several small oligocrystals was located in the centre of the cubozoan statolith. The origin areas and the accretion of statoliths are similar in both species. Our results lead to the assumption that the single bassanite statolith of C. fleckeri (Cnidaria, Cubozoa) is a progression of the scyphozoan multiplex statolith. It is therefore suggested that the Cubozoa are derived from a scyphozoan ancestor and are a highly developed taxa within the Rhopaliophora.     

Impact of temperature on juvenile growth and age at first egg-laying of the Pacific reef squid Sepioteuthis lessoniana reared in captivity

Cephalopod mollusks exhibit highly plastic life cycle traits influenced primarily by the interactive effects of food availability, light cycle and temperature, with the latter perhaps the most influential. Hatchlings of the tropical reef squid Sepioteuthis lessoniana were hatched from field-collected eggs in the laboratory and cultured at different temperatures to evaluate the effect of temperature on growth rates. All groups showed rapid, sustained growth rates from hatching to a size of 10–25 g. Beyond this size range, growth was slower and not clearly exponential in form. Growth rate was closely linked to temperature. Squids grown at approximately 27 °C attained a size of 10 g in as little as 45 days at sustained growth rates of 12.2% body weight day⁻¹ (%bw day⁻¹), while squids cultured at 20 °C required almost 100 days to attain the same size at rates of 5.7%bw day⁻¹. At an age of 55 days and approximately 1 g body weight, juvenile squids cultured at 20 °C were able to accelerate growth rates from 5.7%bw day⁻¹ to over 12%bw day⁻¹ when temperature was raised to 27 °C. They maintained this growth rate to a size of about 10 g and an age of at least 75 days post-hatching, indicating that body size and not age is the limiting factor for this rapid post-hatching growth. By comparison, conspecifics cultured near 27 °C from hatching had shifted out of the rapid post-hatching growth phase by day 50 at sizes between 10 and 50 g. The hatchlings from temperate to subtropical Japan had consistently higher growth rates at comparable temperatures than hatchlings from tropical Okinawa. When plotted as growth rate versus temperature, the Japanese group had a clearly higher slope to the relationship than the tropical populations, equivalent to a 2%bw day⁻¹ difference in growth rate at 25 °C. Age at first egg-laying was decreased at higher culture temperatures; however, overall life span was not. Received: 21 February 2000 / Accepted: 6 September 2000     

Age studies based on daily growth increments in statoliths and growth lamellae in cuttlebone of cultured Sepia officinalis

One hundred and six individuals of Sepia officinalis were cultured throughout its life cycle in two different temperature regimes: 13-15°C and 18-20°C. The number of increments in the statoliths and the number of lamellae in the cuttlebone were read at known ages in different individuals. The formation of growth increments in the statoliths was linearly related with individual age, but it was independent of temperature. By comparison between the slope of these linear relationships and the bisecting line, the hypothesis "1 increment=1 day" was validated in individuals as old as 240 days. In specimens older than 240 days, the number of increments was underestimated due to the poor resolution of the later growth increments. The maximum number of days in captivity (420) was about 80% of the life span estimated for this species in the area studied. Preliminary validation was also obtained from statoliths marked with tetracycline. Increments between tetracycline marks were not visible. The mean distance between marks in ten statoliths was 44 µm (Lj). Considering the same distance in statoliths of wild individuals, the mean number of increments counted was 37 (Lj). This result is very close to the 40 days passed between both tetracycline marks. A straight line significantly different for each culture temperature defined the relationship between number of the cuttlebone lamellae and age. The number of lamellae in the cuttlebone does not correspond to the real age. The time necessary for the formation of one lamella was 8DŽ.3 days at 13-15°C, whereas the deposition of one lamella lasted 3.1ǃ.06 days at 18-20°C. In conclusion, the periodicity for lamellar deposition can be defined only if the temperature where the animal lived is considered.     

Microstructure of settlement-marks in the otoliths of tropical reef fishes

The morphology and ultrastructure of the otolith settlement-mark was examined in 44 tropical reef-fish species spanning nine families. A classification scheme based on similar otolith characteristics is presented. Three major categories are identified based on changes in increment width and optical qualities of the settlement-mark. Of the 44 species examined, 39 possessed “abrupt” settlement-marks (Type I) characterised by a rapid decrease in increment width (up to 50% reduction) over settlement. Type I settlement-marks were found in all nine families examined. The 39 species spanned the whole range of possible larval durations (Pomacentrus moluccensis, 15 d ± 0 SE; Naso hexacanthus, 91.2 d ± 2.97 SE). Four of the 44 species possessed “zonal” settlement-marks (Type II), featuring a band of increments that are wider than pre-settlement increments. Species in this category are the labrids Corisaygula, Thalassoma bifasciatum, T. lunare and an unidentified acanthurid (Acanthurus sp. 2). One species of acanthurid (N. brevirostris) possessed a “gradual” settlement-mark (Type III), manifest as a gradual decrease in increment width during the settlement period. A possible fourth type was identified from the literature. Gnatholepis thompsoni and Coryphopterus glaucofraenum possessed a settlement-mark with increment widths that increased post-settlement. Available data suggest a poor relationship between the structure of the settlement-mark and the magnitude of metamorphosis (previously reported as internal and external morphological change). Evidence suggests that the increment profile over early development and the increment transitions associated with the settlement event are taxon-specific and may enable late-larval stage fishes to be identified to species level. Received: 21 May 1997 / Accepted: 3 February 1999     

Temporal life history plasticity of the Southern Ocean squid <Emphasis Type="Italic">Todarodes filippovae</Emphasis> from waters off Tasmania,Australia

Population dynamics and maturity parameters were analysed for seasonal samples of the oceanic ommastrephid squid Todarodes filippovae from off the coast of Tasmania Australia from 2002 to 2004. Based on assumed daily periodicity in statolith increments, T. filippovae had a life cycle of about a year with the sexually dimorphic females reaching their larger size by predominantly growing faster than males. Due to the small sample size of males, analysis was undertaken on female individuals only. Growth in all samples was best described by a power curve and varied annually, with significantly faster growth in 2001 compared to the subsequent 2 years. Seasonal growth rates also varied with autumn- and winter-hatched squid significantly faster than summer-hatched squid. Spring growth rates were intermediate but not significantly different to the other three seasons. Peak hatching periods occurred in late autumn and early winter. ANOVA revealed a season × year interaction for mantle length and total body weight. Pairwise comparisons showed that the annual differences were likely driven by smaller squid in autumn 2002 compared to autumn samples in the other 2 years. Pairwise comparisons also revealed seasonal differences with winter-caught squid smaller than those from the majority of other samples. There were no seasonal effects on mature female gonad weights but females caught in 2002 had significantly lighter gonad weights than females from the following 2 years. There were no consistent trends among seasons or years in the age structure of mature females. This study revealed the plasticity and flexibility in growth and maturity parameters in this species, with a preference for faster growth during cooler periods.     

Feeding, metabolism and growth in the Antarctic limpet, Nacella concinna (Strebel 1908)

Post-prandial increases in metabolism, the specific dynamic action of feeding (SDA), were evaluated in the Antarctic limpet Nacella concinna. O₂ consumption rose to a peak value 2.3 times higher than pre-feeding standard metabolic rates. This peak rise is low for marine ectotherms, but is typical of polar species. There were three peaks in the SDA, the first lasted only for the 1st day, was caused by handling, and was minor. The second was the major peak. It lasted from post-prandial days 4–9 inclusive, and accounted for around 70% of the SDA response. The third peak lasted from day 11 to day 15 and accounted for 30% of the total SDA. A 15-day SDA is much longer than values for temperate species, but is again typical for polar marine ectotherms. NH₃ excretion declined post-prandially from around 0.4 μmol animal⁻¹ h⁻¹ to values between 0.025 and 0.223 μmol animal⁻¹ h⁻¹ throughout the SDA. The total O₂ consumed in the SDA was 90.2 μmol O₂, which converts to 44.7 J of energy. This was 45–50% of the energy consumed in the meal (93.5 J). Pre-feeding O:N ratios, after 26 days without food, were around 1, indicating protein as the sole metabolic substrate prior to initiating the SDA. After feeding, O:N ratios rose to between 2.5 and 19, indicating significant use of lipid or carbohydrate from the food. Experiments were conducted in ambient seawater with enhanced levels of Sr (SrCl added at 800 mg kg⁻¹), and limpets were fed microalgal films also grown in enhanced Sr media. Sr incorporated in the shells during the experiment allowed the measurement of shell increments deposited during the SDA. Between five and eight microgrowth bands were present in the Sr-enhanced increments, which was similar to the number of days in the second SDA peak. The mean shell increment laid down was 17.6 μm. Estimating tissue deposition from measured growth increments and published ash-free dry mass (AFDM) to length relationships produced a value of 0.81 mg AFDM, which converted to 26.4 J of energy, or 25–30% of the energy ingested in the meal. Estimates of growth increments associated with a single SDA have not previously been possible. Overall energy used in the SDA and tissue deposition accounted for 75–80% of the energy ingested; the remainder was probably accounted for by unmeasured costs such as mucus production. Received: 6 June 2000 / Accepted: 20 September 2000     

Observations on shell deformities,ultrastructure, and increment formation in the bay scallop Argopecten irradians

Shell morphology and ultrastructure were examined in the bay scallop Argopecten irradians, cultured in recirculating seawater systems under various conditions of feeding, lighting, and handling. On a unialgal diet of Thalassiosira pseudonana, scallop growth ranged from 120 to 183 m d^-1 at 20°C in the laboratory, about two-thirds of the growth rate found in the field. However, shell deposited in the laboratory differed from that in the field in several ways. In the field, scallops formed costae as an unpigmented, corrugated marginal shell layer; shell deposited in the laboratory lacked this layer and was therefore darker. Also, microstructure of the exterior shell surface of field scallops was coarsely granular, while that of cultured scallops was relatively smooth. Excessive handling of scallops in the laboratory resulted in marginal thickening of valves, a deformity which was completely arrested by a change from daily to weekly handling. Scallops cultured in the same tank with oysters developed shell-thickening on the interior of the valves. It is postulated that shell abnormalities in bivalves result from disruption of complex behavioral processes associated with shell deposition and may be elicited by a variety of natural and experimental irritants. Under natural lighting regimes and optimal conditions for growth, scallops deposited exactly one shell increment per day, but under continuous lighting, deposition of growth increments often became aphasic. In one 28-d experiment, there was a strong correlation between number of growth increments formed and increase in shell height, suggesting that shell ridge formation occurred intermittently, rather than daily, when shell growth rates fell below approximately 150 m d^-1.