Genetic relations between the bivalves Cardium (Cerastoderma) edule,Cardium lamarcki and Cardium glaucum,studied by means of crossed immunoelectrophoresis

Rabbit antisera were raised against Cardium edule (L.) from the Waddensea locality, Skallingen, against Cardium lamarcki (Reeve) from the Danish locality, Dybsø Fjord, and against Cardium glaucum (Bruguière) from the French Mediterranean coast at Ile des Embiez. Allopatric and sympatric populations of C. lamarcki and C. edule from Swedish and Danish localities, and allopatric populations of C. glaucum from the Mediterranean Sea, Red Sea and Black Sea were studied by means of the described method and by starch gel electrophoresis of malate dehydrogenase (MDH). By means of crossed immuno-electrophoresis, using intermediary gels containing antibodies against one of the Cardium types and receiver gels containing antibodies against one of the others, type-specific and type-non-specific reactions were recorded. The results are compared with a similar investigation of Mytilus edulis, M. galloprovincialis and M. californianus. Based on immunological properties, it is shown that C. edule is a valid species different from the C. lamarcki/C. glaucum complex. It is further suggested that C. lamarcki and C. glaucum should still be referred to by these two names as significant genetic differences between them is revealed by this immunological study.     

Seasonal changes in the physiology,reproductive condition and carbohydrate content of the cockle Cardium (=Cerastoderma) edule (Bivalvia: Cardiidae)

Pronounced seasonal cycles in the rates of oxygen consumption and feeding were found for Cardium (=Cerastoderma) edule L. measured in the field under ambient conditions. The cockles had a maximum rate of oxygen consumption (0.89 ml O₂ g^-1 h^-1) in April which declined to a minimum of 0.35 ml O₂ g^-1 h^-1 in March. Their feeding rate was variable but had a maximum value (3.91 l g^-1 h^-1) in April and a minimum value (0.73 l g^-1 h^-1) in October. There was no apparent seasonal variation in absorption efficiency, with a mean value of 67.6%. Gametogenesis was initiated in January and the population reached a peak in reproductive condition in April/May, followed by a 3 month spawning period. Carbohydrate reserves were synthesised during spawning, and were then utilised during the winter and early spring. An adaptive function for a reduction in time spent feeding is postulated, and correlations between the rates of certain physiological processes and some exogenous and endogenous variables are discussed.     

Habitat selection of two congeneric bivalves,Cardium edule and C. glaucum in sympatric and allopatric populations

Habitat selection of two bivalve species, Cardium edule (L.) and C. glaucum (Bruguière) is described for sympatric and allopatric populations of both sibling species. It is demonstrated that C. edule grows faster than C. glaucum and is absent from localities with periodic low food-availability. The ability to burrow of C. glaucum is low compared with that of C. edule; the former species does not occur in very loose, well-sorted sand near the coast. Earlier studies which suggest that tidal range determine the species distribution could not be confirmed.     

Evolution of Cardium (Cerastoderma) edule,C. lamarcki and C. glaucum: studies of DNA-variation

Genetic relations between populations of Cardium (Cerastoderma) edule (L.) and C. lamarcki (Reeve) from Danish waters and C. glaucum (Bruguiére) from the Mediterranean Sea were studied using DNA analysis. Repetitive sequences of nuclear DNA were compared by Southern blotting of enzyme digested DNA fragments, separated on agarose gel by electrophoresis. Fragments were visualized by autoradiography using ³²P labelled probes. These were nick translated, randomly sheared fragments of the whole chromosomal genomes of two species of distant taxa — Mytilus edulis and Mya arenaria, respectively. The number of detected DNA bands was 18 for C. edule, 32 for C. lamarcki, and 24 for C. glaucum. Measurements of genetic distances, based on comparisons of these repetitive sequences using a simple dissimilarity coefficients, show the greatest distance, i.e., 0.84 between C. edule and C. lamarcki, 0.69 between C. edule and C. glaucum, and the smallest, 0.52 between C. lamarcki and C. glaucum. Comparisons of the three Cardium types, using gene spliced and cloned C. edule nuclear DNA (nDNA) fragments as probes, accorded with these findings. Also, analysis of mitochondrial DNA (mtDNA) from eggs and hepatopancreas, although not in itself conclusive, accorded with the results of the analyses of nDNA. MtDNA was measured by electron micrograph to 15 000 base pairs (bp), and sizes of restriction enzyme digested fragments were compared by gel electrophoresis. The results of the comparisons based on repetitive sequences supported by the two other studies suggest that C. edule resembles the ancestral form more than the two others, and that C. lamarcki is the most different and recently evolved form. This hypothesis is discussed in relation to the geographical distribution of the species complex.     

Energetics of growth and reproduction in cockles (Cerastoderma edule): Seasonal and age-dependent variations

Seasonal patterns of growth were compared interannually between 1983 and 1985 for different ageclasses of an intertidal population of cockles,Cerastoderma edule L., in the Mundaka Estuary (Basque Country, North Spain). Growth data and simultaneous determinations of respiratory metabolism were integrated over defined periods of time in order to compute assimilation and net growth efficiency. Consistent differences between the exponents that scale both assimilation and metabolism to body size led to growth efficiency values that were a decreasing function of age. Total growth was partitioned into three components: somatic growth, reproductive output and reserve storage, and the corresponding effort indexes were calculated. Despite important differences in total growth, reproductive output was constant between years, indicating a higher reproductive effort under poorer growth conditions. Increased costs of reproduction that appeared to be associated with this higher reproductive effort did not affect somatic maintenance, since previously stored carbohydrate reserves were primarily used to fuel gametogenesis.     

Enzymatic and morphological criteria for distinguishing between Cardium edule and C. glaucum of the Portuguese coast

Cardium edule L. and C. glaucum Bruguière are two sibling species of cockles commonly found in littoral systems along the Portuguese coast, where they sometimes occur as mixed populations. The problem of morphological separation of these two cockle species has not been completely solved. Indeed, the distinctive morphological criterion proposed by Brock after identifying the specimens by means of electrophoretic patterns of hepatopancreatic malate dehydrogenase was not effective when applied to Portuguese Cardium spp. The object of the present study was to establish morphological discrimination criteria for distinguishing Portuguese C. edule and C. glaucum on the basis of initial identification of individuals through the enzymatic criterion established by Brock. Cockles were collected from 1988 to 1991 at 16 estuarine and lagoonal systems along the Portuguese coast. Hepatopancreatic extracts were submitted to horizontal starch gel electrophoresis following Brock's procedure. The malate dehydrogenase patterns obtained were identical to those found by Brock. Clear interspecific differences in the leucine aminopeptidase and esterase isozyme systems were also detected. The morphological study of the shells involved 7 morphometric and 9 qualitative and semiquantitative characters. Canonical variate analysis of the morphometric characters showed that these could not be used for identification purposes. However, a clear separation between the two species was obtained by a principal-component analysis of qualitative and semiquantitative characters. The characters mainly useful for distinguishing between the two species were valve profile, shape of ventral valve junction, type of calcareous scales and shell colour.     

Assessment of intertidal growth and capacity adaptations in suspension-feeding bivalves

Curves relating instantaneous growth rate to aerial exposure were determined for six species of bivalves in laboratory and shore experiments. A dimensionless index equatable with the relative intertidal growth performance of a species was calculated by integration of the growth curve after converting both growth rate and aerial exposure to decimal fractions. Intertidal growth performance of the bivalves tested corresponded well with their natural levels of occurrence on the shore, and improved in the following order: Modiolus modiolus < Argopecten irradians < Ostrea edulis < Mytilus edulis < Crassostrea virginica. Geukensia demissa, for which an index value could not be determined, grew faster intertidally than subtidally. The relative contributions made to intertidal growth performance by energy-conserving and energy-supplementing capacity adaptations were assessed by resolving the growth curves into energy-input and energy-loss components. The rate of energy loss due to intertidal exposure was lower in the high-shore species, and also less affected by harsher subaerial conditions, than in the low-shore species. Moreover, M. edulis and C. virginica showed abilities to supplement energy input such that growth per unit immersion time was better at certain intertidal levels than subtidally. Energy conservation and supplementation in these forms made roughly equal contributions to their improved intertidal growth relative to species occurring lower on the shore.     

Vertical distribution, population structure and life cycle of Neocalanus cristatus (Crustacea: Copepoda) in the Oyashio region, with notes on its regional variations

Vertical distribution and population structure of Neocalanus cristatus were investigated at Site H in the Oyashio region from September 1996 through October 1997 to evaluate their life cycle mode. Additional temporary samplings were also made at several stations covering the entire subarctic Pacific, Okhotsk Sea and Japan Sea, as a basis for regional comparison of life cycles of this species. At Site H, N. cristatus spawned throughout the year below 500 m depth, with a peak from October to December. The resulting eggs and nauplii floated/migrated upward, and formed an abundance peak of Copepodite Stage 1 (C1) in the surface layer in February. In the surface layer, the C1 developed and reached C5 by early June through a phytoplankton bloom which occurred in mid-March to end of June. The C5 migrated to deeper layers in July and August, where they molted to adults. Apparently, the developmental time from C5 to adults was highly variable (>1 month), and some might overwinter. The life cycle of N. cristatus appeared to be annual for the major portion of the population. Taking into account sampling season, temporal changes in vertical distribution and population structure data collected from regions other than Site H, there was a close correlation in the timing of the life cycle over the entire subarctic Pacific, but the reproduction season (April to June) was observed to be different in the Okhotsk and Japan Sea populations. Regional comparison of prosome length of C5 individuals, including those in the Bering Sea, indicated significantly larger sizes of specimens from the Japan Sea and Okhotsk Sea, as compared with those from the entire subarctic Pacific. Possible causes for regional variability in life cycle patterns and body sizes are discussed. Received: 18 December 1998 / Accepted: 19 April 1999     

Annual variation in the growth rate of Yoldia notabilis (Bivalvia: Nuculanidae) in Otsuchi Bay,northeastern Japan,analyzed using shell microgrowth patterns

Annual variation in the growth rate of the protobranch bivalve Yoldia notabilis (collected in Otsuchi Bay, northeastern Japan between 1989 and 1991) was determined by measuring shell length at successive growth lines. Factors affecting growth rates were assessed by examining long-term environmental data. Shell cross-sections of Y. notabilis showed a clear pattern of internal growth lines which formed simultaneously with the annual external lines on the outer shell surface. Hence, they were used as an age marker. A maximum lifespan of 17 yr was determined for this species, and a sigmoidal growth curve was obtained by the internal growth line analysis. Examination of the growth lines also revealed a large annual variation in growth rate, fluctuating as much as 32-fold during the past 9 yr. The variation correlated negatively with water temperature and positively with chlorophyll a content in the water column. It is suggested that the variation in annual growth rate is dependent on food supply during the spring phytoplankton bloom which varies from year to year according to the flow of the cold Oyashio current each spring.     

川东北大气沉降中水溶态重金属季节变化:自然和人为因素的影响

大气沉降地球化学研究对解释表生环境中元素地球化学行为具有重要意义.但以往对大气沉降地球化学的研究主要集中在城市或者受工业活动强烈影响的地区.本研究选择地处偏远、无明显工业活动影响的川东北地区,于2011年8月—2014年7月采集大气沉降样品,对其中水溶态重金属元素Cr、Co、Ni、Cu、Zn、Ga、As、Cd和Pb进行了研究.结果表明,这些重金属元素的年平均沉降通量最大的为Zn,158μg·(10~2cm~2·a)~(-1),最小的为Ga,0.06μg·(10~2cm~2·a)~(-1),相差达到4个数量级.相对于我国城镇地区或受工业活动影响的偏远地区,川东北地区重金属元素沉降通量较低.总体上Co和Ni的沉降通量在冬-春季节较高,可能与这些季节的大气粉尘活动有关,Ga、Pb、Cr、Cd、Zn、As等则一般在夏季沉降通量较高,可能与降水增加和旅游旺季的人类活动加强有关.2012年12月—2013年9月的当地道路改建工程导致了重金属元素沉降通量的明显提高.     

Variability of shell growth and morphology of the wall-plate junctions of the intertidal barnacle Tesseropora rosea (Cirripedia: Tetraclitidae)

Variability in shell growth and morphology of the wallplate junctions of Tesseropora rosea (Krauss) were studied in 1983 in New South Wales, Australia, at two sites with differing wave exposure. The wall-plate junctions, a combination of dado (geometrically interlocking) and lap bints, increase in complexity with age, but do not differ between sites. The number of parietal tubes is fixed at settlement and does not differ between sites, but differs among wall plates in a manner consistent with their evolutionary history (rostrum > laterals > carina). The number of secondary septa increases with age, but does not differ between sites. In epilithic T. rosea, growth of the alae contributes to orifice enlargement. Epizoic T. rosea (on limpets) have conspicuous radii and display diametric growth. At the more exposed site, the shell wall was sleeper, and the aperture length, height and weight of the shell increased at a faster rate than at the less exposed site, but the maximum height of the shell was less. These differences appear to be due to a combination of greater feeding rates and greater erosion rates at the exposed site. The greater maximum height attained at the more sheltered site may be due to greater individual longevity. Juveniles at the sheltered site had thinner septa in the laterals than those at the exposed site. Septa in adults were rhinner at the exposed site, but this is due to age differences between sites in the adult populations.     

Behavioral ecology of the neotropical termite-hunting ant Pachycondyla (= Termitopone) marginata: colony founding,group-raiding and migratory patterns

This study provides the first detailed field account of colony founding, group-raiding and migratory habits in the neotropical termite-hunting ant rPachycondyla marginata, in a semi-deciduous forest in south-east Brazil. New colonies can originate by haplometrosis, pleometrosis, or colony fission. Incipient colonies with multiple foundresses persisted longer in the field, and most excavated nests contained more than one dealated female. A total of 202 group raids by P. marginata were registered, and in all cases the raided termite species was Neocapritermes opacus. Nearly 20% of the workers within a colony engage with raiding activity. Colonies of P. marginata hunt for termites approximately every 2–3 weeks, and group-raids may last for more than 24 h. Target termite nests are up to 38 m from the ant colony, and occasionally two nests are simultaneously raided by one ant colony. Raiding ants carry 1 or 2 paralysed prey, and nearly 1600 termites can be captured during a 9-h raid. Migration by P. marginata colonies lasted over 2 days and covered distances of 2-97 m (n = 48). Average residence time at a given location was 150 days. Three basic migratory patterns were noted: colony fission (only part of the colony moves), long-distance migrations, and short-distance migrations. Both raiding and migratory activities appeared to be strongly affected by seasonal factors. The group raiding and migratory patterns of P. marginata are compared with other ant taxa with similar habits. It is concluded that P. marginata presents a rudimentary form of the so-called army ant behavior, which is highly developed in the subfamilies Dorylinae and Ecitoninae. The extremely specialized diet of P. marginata and the associated high costs of migration are features likely to prevent it from evolving a full army ant life pattern.     

Individual growth and size-selective mortality of larval turbot (Scophthalmus maximus) reared in enclosures

Larval turbot (Scophthalmus maximus) were reared in a large marine enclosure and in plastic bags in southern Norway. Samples of larvae in the enclosure were taken during the first 12 d of life to estimate individual body growth based on back calculations from daily growth rings on their otoliths. Size selective mortality was documented for these larvae in the predator-free enclosure. Starvation in the laboratory occurred on the seventh day. In the enclosure, a mortality rate of 18.1% d^-1 prevailed. Our data indicates that the survivors beyond the starvation period are larger by 0.18 mm on average. This result is important with regard to the question of whether starvation is an important mechanism for larval mortality in the sea. A possible means of estimating the relative effects of starvation versus predation based on these results and the backcalculation technique is suggested.     

Interannual variation in the reproductive cycle and biochemical composition of the cockle Cerastoderma edule from Mundaca Estuary (Biscay,North Spain)

Seasonal determinants of body weight, biochemical composition and reproductive condition in the cockle Cerastoderma edule L. from the Mundaca Estuary, Spain, were performed from December 1983 to July 1985. Interannual differences in timing of gametogenesis and spawning between 1984 and 1985 were correlated with temporal variations in the cycles of storage and utilisation of carbohydrate reserves. Young individuals exhibited larger annual fluctuations in soft-body weight, displaying higher growth rates in spring and greater diminishment of energy reserves during autumn and winter than older individuals. Carbohydrate content displayed a similar tendency, levels being lower in young individuals throughout the year.     

Short-term variations in the pigment composition of a spring phytoplankton bloom from an enclosed experimental ecosystem

Phytoplankton samples taken during the spring bloom in the experimental enclosed ecosystem bags at Loch Ewe, Scotland, during 1983 were analysed for carotenoids and chlorophyll compounds using high-performance liquidchromatography (HPLC). Changes in the relative proportions of these pigments were related to day-to-day changes in the composition of the bloom and the physiological state of the algae. There is clear evidence for a change in the chlorophyllide a:chlorophyll a ratio, which reached a maximum as nutrient limitation occurred. No major qualitative changes in the carotenoid components were seen during the bloom; the relative proportion, however, of some carotenoids does provide useful information on the relative abundance of certain algal type in the phytoplankton.     

Differential ingestion of zooplankton by four species of bivalves (Mollusca) in the Mali Ston Bay,Croatia

This study provides information about differences in composition of ingested zooplankton amongst bivalve species coexisting in the same area in a period from May 2009 to December 2010. The study was conducted at the Mali Ston Bay (42°51′ N, 17°40′ E)—the most important bivalve aquaculture area in the eastern Adriatic Sea. Stomach content analysis was performed on cultured species—Ostrea edulis and Mytilus galloprovincialis, and commercially important bivalve species from their natural environment—Modiolus barbatus and Arca noae. Results confirmed carnivory in bivalves, both from natural and cultured populations, but cultured species had higher numbers of zooplankters than those living on the seabed. The most abundant taxa were bivalve larvae, followed by tintinnids, copepods, unidentified eggs and gastropod larvae. Recorded numbers of bivalve larvae in M. galloprovincialis stomach were the highest so far reported and show that mussels impact the availability of natural spat.     

Age structure,growth, and morphometric variations in the Atlantic surf clam,Spisula solidissima,from estuarine and inshore waters

The surf clam, Spisula solidissima (Dillwyn), population in the estuarine waters of Long Island Sound, New York, USA, was characterized in 1984 and again in 1988 by an age structure restricted to just two age-classes, and an apparent lifespan of only about 10 yr. In the inshore coastal waters off Fire Island, New York, a wide age range from 2 to 22 yr old was present. The age structure at Long Beach, New York, a third geographic region which is coastal but influenced by the Hudson River estuary, was similar to Long Island Sound. Juvenile surf clams grew at similar rates in all three geographic regions. However, adults from the Long Island Sound population grew significantly slower and reached an asymptotic maximum size which was 37% smaller than Fire Island adults. Long Beach adults had intermediate growth rates and maximum sizes. The shells of Long Island Sound clams were also 25% thinner than those from the other two regions. Density dependent effects on growth, evaluated over abundances ranging from 0.5 to 294 ind. m^–2, were present but were too small to account for observed regional differences. Results suggest that adult surf clams may be physiologically stressed by the reduced salinity and more extreme temperatures found in estuarine waters.     

Accumulation of polychlorinated biphenyls in the cockle Cerastoderma edule and the tellin Macoma balthica

Two species of marine bivalve, Cerastoderma edule (Linnaeus) and Macoma balthica (Linnaeus), were exposed to particles contaminated with three commercial polychlorinated biphenyl (PCB) mixtures, Aroclors^® 1242, 1254 and 1260. The accumulation of PCB in bivalve tissue was monitored for periods of up to 40 days. In addition to measuring total PCB concentrations, which reached 60 ppm of Aroclor 1242 in M. balthica, the fate of individual PCB homologues of different chlorine content, ranging from 2 to 8 chlorine atoms per biphenyl molecule, was also determined. Selective accumulation of PCBs with 5 chlorine atoms per biphenyl molecule occurred, whilst isomers of very low and very high molecular weights were accumulated at much slower rates.