Population dynamics of Euterpina acutifrons (Copepoda: Harpacticoida) from North Inlet,South Carolina,with reference to Dimorphic Males

The population dynamics of Euterpina acutifrons (Dana), a pelagic, harpacticoid copepod, are summarized in a life table based on field data. Highest mortality occurred in the last naupliar stage (NVI) and the first copepodite stage (CI). Overall survival in the field was 0.06% from the first naupliar stage (NI) to adult (CVI). The net reproductive rate (R _o=55.590) and intrinsic rate of increase (I _m=0.28) were sufficiently high to maintain a population with such a low survival rate in nature. E. acutifrons was present and breeding in the field from April through December. Low temperatures limited breeding, which began when the temperature reached 16.5°C and ceased when it fell to 11°C. Optimum temperature for North Inlet E. acutifrons was 25°C, with a maximum laboratory survival of 15.3% and a generation time of 10.3 days. Generation time in the field (20°C) was 14 days. Temperature also affected the abundance of dimorphic males. Small males were always most abundant, but peaked during the coldest month; large males became equal in abundance only during the varmest months.Contribution No. 298 from the Belle W. Baruch Institute for Marine Biology and Coastal Research.     

Life cycles in marine meiobenthos. Experiments at various temperatures with Monhystera disjuncta and Theristus pertenuis (Nematoda)

Monhystera disjuncta Bastian 1865 and Theristus pertenuis Bresslau and Schuubmans Stekhoven 1935 were cultured on sea-water agar in Boveri dishes at various temperatures. Generation time was measured as the period elapsing in two consecutive generations between the first egg deposit, the first hatching, or the first appearance of sexual characters. M. disjuncta has a generation time of 13 days at 17° to 22°C, 15 days at 13° to 15°C, 17 days at 9° to 12°C, 22 days at 7°C, 77 days at 0° to 2°C, and 135 days at -1° to 1°C. Low temperatures result sometimes in vivipary. T. pertenuis has a generation time of 23 days at 17° to 22°C, 41 days at 13° to 15°C, 47 days at 9° to 12°C, and 71 days at 7°C. M. disjuncta females live for up to 61 days at 17° to 22°C, T. pertenuis females up to 208 days at 7°C. Under North Sea temperature conditions, 17 generations of M. disjuncta and 7 generations of T. pertenuis could occur during the course of 1 year (calculation based upon experiments giving the shortest possible generation time). Females deposit eggs over a couple of days, therefore, the medium generation time is longer, and there will be fewer generations per year in the sea.     

Reproduction and growth of Euterpina acutifrons (Copepoda: Harpacticoida) under experimental conditions

Individual specimens of Euterpina acutifrons (Copepoda: Harpacticoida) taken from the mass cultures of the C.N.E.N.-EURATOM Laboratory at Fiascherino, Italy, were reared in new culture media prepared with suspensions of several species of algae in filtered and sterilized sea water. All the experiments were carried out at a temperature of 18°C±1 C°. The influence of food concentrations on adult life-span and reproductive activity of E. acutifrons was analyzed. A good correlation was found between concentration of algal suspension and egg production. Maximum life-span was observed at intermediate values of food concentration. Other experiments were carried out to determine egg fertility and duration of the various embryonic and postembryonic development stages. Embryonic development time was calculated as approximately 2 days; the adult females appeared 10 to 12 days after hatching of Nauplius I. Production of nauplii by 6 females reared under conditions of excess food supply was also studied. These conditions were achieved by supplying high concentrations of a mixture of 4 different algal species and by completely renewing the culture medium at frequent intervals. Under these conditions, each female laid an average of 12.5 sacs and produced an average of 294.3 nauplii. An average production of 355.5 eggs per female was estimated. An analysis was made of growth in size and weight of the females: the average daily egg production in terms of dry weight corresponded to about 32% of the biomass of the adult female. Birch's (1948) method was used to calculate net reproduction rate (R _o=70.89), intrinsic rate of increase (r _m=0.161) and mean generation time (T=26.5 days).This study was performed at: Laboratorio per lo Studio della Contaminazione Radioattiva del Mare, C.N.E.N.-EURATOM, Fiascherino, I-19030 La Spezia, Italy.     

Effects of dietary fatty acids on the reproductive success of the calanoid copepod<Emphasis Type="Italic"> Temora longicornis</Emphasis>

Egg production and hatching success of the copepod Temora longicornis were measured in laboratory experiments and in the field (North Sea). In the laboratory, ingestion of four algal species (Thalassiosira weissflogii, Phaeocystis globosa, Isochrysis sp. and Dunaliella tertiolecta) was followed and the content of fatty acids in the algae was determined. The two food types (T. weissflogii and Isochrysis sp.) that provided the highest ingestion of carbon and long chain fatty acids also resulted in the highest egg production rate (E_r) and hatching success (H%). In contrast, D. tertiolecta led to both low ingestion of carbon and long chain fatty acids, resulting in low reproductive success. There was a positive relationship between the amount of eicosapentaenoic fatty acid [20:5(n-3), EPA] ingested and E_r and H%, and of the ratio between docosahexaenoic and eicosapentaenoic fatty acid [22:6(n-3)/20:5(n-3), DHA/EPA] in the ingested food and H%. In the field, chlorophyll a and specific fatty acids were measured and protists were enumerated, in order to investigate the link between these factors and the reproductive success of T. longicornis. Hatching time was found to be related to temperature and exceeded 120 h at 6°C. No relationship was found between chlorophyll a and reproductive success (E_r or H%). E_r correlated with the concentration of diatoms and ciliates, which were the dominating protists in early spring, indicating that food quantity was the limiting factor for E_r. As in the laboratory experiments, H% was dependent on the fatty acid DHA and the ratio of DHA/EPA, which indicates that the quality of eggs (H%) is linked to the quality of food.Communicated by M. Kühl, Helsingør     

The cultivation of an appendicularian through numerous generations

Oikopleura dioica Fol has been cultivated continuously for 7 months, through 19 filial generations, in the laboratory. At 13°C, mean generation time was 9.5 days, at food concentrations close to natural phytoplankton concentrations in the North Sea. The reproductive performance did not deteriorate with increasing filial generations.     

Effect of temperature on the egg and yolk-sac larval development of common pandora,<Emphasis Type="Italic"> Pagellus erythrinus</Emphasis>

Temperature is one of the most critical environmental factors for fish ontogeny, affecting the developmental rate, survival and phenotypic plasticity in both a species- and stage-specific way. In the present paper we studied the egg and yolk-sac larval development of Pagellus erythrinus under different water temperature conditions, 15°C, 18°C and 21°C for the egg stage and 16°C, 18°C and 21°C for the yolk-sac larval stage. The temperature-independent thermal sum of development was estimated as 555.6 degree-hours above the threshold temperature (the temperature below which development is arrested), i.e. 7°C for the egg and 12.1°C for the yolk-sac larval stage. Higher hatching and survival rates occurred at 18–21°C. At the end of the yolk-sac larval stage, body morphometry differed significantly (p<0.05) between the temperatures tested. The growth rate of the total length increased as temperature rose from 16°C to 18°C, while in the range of 18–21°C it stabilized and was independent of water temperature. The estimated Gompertz growth curve for the yolk-sac larvae of P. erythrinus was (r²=0.992) for the 16°C, (r²=0.991) for the 18°C and (r²=0.981) for the 21°C treatment. The efficiency of vitelline utilization during the yolk-sac larval stage was higher at 18°C.Communicated by O. Kinne, Oldendorf/Luhe     

Respiratory adaptations of Pleuroncodes planipes to its environment off Baja California

The reproductive potential of two harpacticoid copepods, Tachidius discipes and Paronychocamptus nanus, as measured by the intrinsic rate of natural increase, r _m, has been studied. The influence of temperature on all factors used in the calculation of r _mhas been taken into account. Generation time, which has also been determined in a third species, Nitocra typica, is the single factor which is most affected by temperature and which has the most pronounced effect on r _m. Values of the reproductive potential differ considerably according to temperature, and thus to the time of the year; during the spring bloom more than half the reproductive potential is realized.     

Tisbe pori n. sp. (Copepoda: Harpacticoida) from the Mediterranean coast of Israel and its cultivation in the laboratory

Tisbe pori n. sp., from the Mediterranean coast of Mikhmoret, Israel, was cultivated in the laboratory and is described. The relative food values of 9 species of marine algae were evaluated according to the copepods' development time (length of period from hatching to reproductive maturity). Individuals fed on Dunaliella tertiolecta or D. primolecta developed the fastest. The rate of feeding of the adults on Dunaliella primolacta was determined. In the temperature range 17° to 31.5°C, the shortest development time (7 days) was at 26° to 28°C. The range of salinity tolerated by adults was 26 to 48.     

Seasonal and interannual variations in size,biomass and chemical composition of the eggs of North Sea shrimp, <Emphasis Type="Italic">Crangon crangon</Emphasis> (Decapoda: Caridea)

In the shrimp Crangon crangon, an important fishery resource and key species in the southern North Sea, we studied temporal variations in size, biomass (dry weight, W) and chemical composition (C, N, protein and lipid) of eggs in an initial embryonic stage. Data from 2 years, 1996 and 2009, consistently revealed that egg size and biomass varied seasonally, with maxima at the beginning of the reproductive season (January), decreasing values throughout spring, minima in June–July, and a slight increase thereafter. This cyclic pattern explains why “Winter eggs” are on average larger and heavier than “summer eggs”. Using a modelling approach, we estimated the duration of oogenesis in relation to seasonally changing seawater temperatures. According to an additive model of multiple explanatory variables, the C content per newly laid egg showed in both years a highly significant negative relationship with day length (r2 = 0.38 and 0.40, respectively; P < 0.0001), a weak positive relationship with temperature (r2 = 0.08 and 0.09; P < 0.05), and a weak negative relationship with phytoplankton biomass (r2 = 0.11 and 0.12; P < 0.05) at the estimated time of beginning oogenesis. Phenotypic plasticity in initial egg size and biomass is interpreted as an adaptive reproductive trait that has evolved in regions with strong seasonality in plankton production and periods of larval food limitation. In contrast to biomass per egg, the percentage chemical composition remained similar throughout the reproductive period. Both the absolute and percentage values also showed significant interannual variations, which caution against generalizations based on short-term studies of reproductive traits of C. crangon and other species of shrimp.     

Anaerobic heat production of bivalves (Polymesoda caroliniana andModiolus demissus) in relation to temperature,body size,and duration of anoxia

Anaerobic heat-production rates of two co-occurring species of estuarine bivalves (a clam and a mussel) were measured with double-twin heat-flow calorimeters, one at 20°C, the other at 30°C. There is no significant difference between the two species in metabolic rates. There is evidence of initial aerobic metabolism in some individuals, as shown by high initial rates exponentially decreasing with time, while others had fluctuating but stable average metabolic activity from the beginning. During aerobic as well as anaerobic metabolism, the bivalves showed rhythmic periods of activity and quiescence. The two species differed in their rhythmic pattern of active and resting metabolism. In the case ofPolymesoda caroliniana, periods of resting metabolism tend to be longer and periods of active metabolism shorter at 30°C than at 20°C. There is a similarity between thermograms ofModiolus demissus at 20° and 30°C. Following acute temperature changes from 5° to 20° and 30°C, the bivalves showed stable metabolic rates in a matter of hours. The stabilized average rates [pooled averages for both species of 1.34×10^-4 (standard error of the mean=0.17×10^-4) W g^-1 dry weight of tissue at 20°C and 2.10×10^-4 (SE=0.20×10^-4) W g^-1 at 30°C] signify a temperature coefficient (Q₁₀) of 1.56 between 20° and 30°C, or partial temperature acclimation. Subtracting heat production as a result of physical activity, i.e., considering only resting metabolism, the corresponding means and standard errors of the means are 1.24×10^-4 and 0.14×10^-4 W g^-1 at 20°C and 1.91×10^-4 and 0.077×10^-4 W g^-1 at 30°C. Anaerobic heat production rate at 20°C is proportional to body size (r=0.84, 9 degrees of freedom, DF). ForM. demissus, measured anaerobic heat production is on the order of 7.5% of the level of aerobic respiration reported in the literature.     

The effects of temperature,body size and growth rate on energy losses due to metabolism in early life stages of haddock (<Emphasis Type="Italic">Melanogrammus aeglefinus</Emphasis>)

Rates of routine respiration (R _R, μl O₂ fish⁻¹ h⁻¹) and total ammonia nitrogen excretion (E _R, μg NH₄–N + NH₃–N fish⁻¹ h⁻¹) were measured on larval and juvenile haddock (Melanogrammus aeglefinus) to ascertain how energy losses due to metabolism were influenced by temperature (T), dry body mass (M _D, mg) and specific growth rate (SGR, % per day). R _R and E _R increased with M _D according to y = a  · M _D ^b with b-values of 0.96, 0.98, 1.14, and 0.89, 0.78, 0.74, respectively, at 10, 7, and 4°C, respectively. Multiple regressions explained 98% of the variability in the combined effects of M _D and T on R _R and E _R in larval haddock: R _R = 0.97 · M _D ^0.98  · e^0.092 · T ; E _R = 0.06 · M _D ^0.79  · e^0.092 · T . In young juvenile (24–30 mm standard length) haddock, R _R tended to decline (P = 0.06) and E _R significantly declined (P = 0.02) with increasing SGR. O:N ratios significantly increased with increasing SGR suggesting that N was spared in relatively fast-growing individuals. Our results for young larval and juvenile haddock suggest: (1) nearly isometric scaling of R _R with increasing body size, (2) allometric scaling of E _R with increasing body size, (3) Q ₁₀ values of 2.5 for both R _R and E _R, (4) metabolic differences in substrate utilization between relatively fast- and slow-growing individuals, and (5) that rates of routine energy loss and growth were not positively related. The measurements in this study will provide robust parameter estimates for individual-based models that are currently being utilized to investigate how variability in climatic forcing influences the vital rates of early life stages of haddock. Our results also stress that inter-individual differences in rates of energy loss should not be overlooked as a factor influencing growth variability among individuals.     

Icy heritage: ecological evolution of the postglacial Baltic Sea reflected in the allozymes of a living fossil,the priapulid Halicryptus spinulosus

Genetic variation of 16 allozyme loci in 397 Halicryptus spinulosus (Priapulida) revealed overall polymorphism of P=0.438 and Hardy-Weinberg expectations for heterozygosity of H _e=0.060 for Baltic Sea stocks, H _e=0.143 for the White Sea and H _e=0.121 for Iceland. Maximal unbiased standard distances of D=0.0693 separated Baltic and White Sea populations. Nordic and Baltic populations could be distinguished by allozymes, but Baltic subsamples proved cohesive. Gene flow amounted to effective exchange values per generation of N _m=2.94 over 650 km of continuous habitat, N _m=10.65 over 175 km, and N _m=13.85 over 20 km. Gene flow started to decrease with geographic distance beyond a dispersal threshold of 20 km, but hierarchical G _ST-statistics indicated light isolation by distance beyond a minimum of 8 km. Gene flow is high for a benthic worm assumed to lack dispersal by pelagic larva, a paradox which cannot be resolved now. Baltic populations are characterized by lower heterozygosity than Nordic stocks. In the Baltic Basin, temporally continuous brackish-water conditions have only existed for the past 7000 years. The two possible colonization routes of H. spinulosus to the geologically young Baltic Sea imply genetic drift, whether by founder effect (sweepstake colonization from Iceland) or by refugial bottlenecking during the Ancylus phase of the Baltic Basin after a direct connection to the White Sea had been sequestered. Continued genetic drift is emphasized by lower heterozygosity in the ecologically unstable Belt Sea compared to the central Baltic. Allozymes falsify the reduced-mutability hypothesis to explain bradytelic evolution of Priapulida. Regional genetic homogeneity, ample polymorphism, and preference for anoxic black mud qualify H. spinulosus populations as indicators of microevolutionary responses to water circulation regimes or pollution in the Baltic Sea.     

Otolith microstructure and daily increment validation of marbled sole (<Emphasis Type="Italic">Pseudopleuronectes yokohamae</Emphasis>)

We examined the daily deposition of otolith increments of marbled sole (Pseudopleuronectes yokohamae) larvae and juveniles by rearing experiments, and estimated the growth pattern of wild larvae and juveniles in Hakodate Bay (Hokkaido Island, Japan). At 16°C, prominent checks (inner checks; ca. 19.8 µm in diameter) were observed on the centers of sagittae and lapilli extracted from 5-day-old larvae. On both otoliths, distinctive and regular increments were observed outside of the inner checks, and the slopes of regression lines between age and the number of increments (n_i) (for sagittae: n_i=0.98×Day–5.90; for lapillus: n_i=0.96×Day–5.70) did not significantly differ from 1. Inner check formations were delayed at lower temperature, and the inner checks formed 13 days after hatching at 8°C. Over 80% of larvae, just after their yolk-sac has been absorbed completely (stage C), had inner checks on both their otoliths. On the lapilli, other checks (outer check) formed at the beginning of eye migration (stage G). To validate the daily deposition of increments during the juvenile stage, wild captured P. yokohamae juveniles were immersed in alizarin complexone (ALC)-seawater solutions and reared in cages set in their natural habitat. After 6 days, the mean number of rings deposited after the ALC mark was 5.7. The age–body length relationship of wild P. yokohamae larvae and juveniles caught in Hakodate Bay was divided into three phases. In the larval period, the relationship was represented by a quadratic equation (notochord length=–0.010×Age²+0.682×Age–2.480, r²=0.82, P<0.001), and the estimated instantaneous growth was 0.38 mm day^–1 at 15 days, 0 mm day^–1 at 34 days and –0.12 mm day^–1 at 40 days. The age–body length relationship in the early juvenile stage (<50 days) and the late juvenile stage (>50 days) were represented by linear equations (standard length=0.055×Age+5.722 and standard length=0.345×Age–9.908, respectively). These results showed that the growth rates in the late larval periods and the early juvenile stage were lower than those in the early larval stage and late juvenile stage; during the slow growth period, energy appears to be directed towards metamorphosis rather than body growth. This study provided the information needed to use otolith microstructure analysis for wild marbled sole larvae and juveniles.Communicated by T. Ikeda, Hakodate     

A simulation model for the management of vicuña populations

The population dynamics of the vicuña was modelled based on field data from the Central Galeras Sector, Perú. Rains were simulated in order to obtain net primary productivity and grass availability which act upon density-dependent fecundity and mortality. The model produces a population growth curve that tends to stabilize at densities around 100 vicuñas per km². Harvest and shearing processes were simulated as part of the vicuña population management model. Harvest was based upon a fixed threshold density D_e below which no harvest occurs; for densities above D_e harvesting is applied at a rate proportional to the difference between D_e and the current population density (fixed escapement or ‘bang-bang’ harvest rule). Management optimization was analyzed by determining the optimal escapement density D_e, which maximized either net profits or the number of animals harvested. Analyzing the results as cumulative totals over 20 years of simulation, an optimum harvest production was obtained for a D_e of about 40 vicuñas per km², and maximum profit was obtained for a D_e around 70 vicuñas per km². Analyzing the results of 20 years of simulation as annual averages with an original population density of 40 vicuñas per km², the harvest production and profits were maximized for a D_e of 40 and 60 vicuñas per km², respectively.The model was validated using data from the Reserva Nacional San Guillermo, Argentina, where vicuña and guanaco populations coexist (although their interaction was not modelled). The sensitivity analysis was performed with three different techniques: (a) stepwise multiple linear regression, (b) visual graphic analysis based on a polar coordinates system, and (c) direct evaluation of the effect on management decisions. The curve shape parameters of the fecundity and mortality functions proved to be the most important ones in determining the outcome of the model.     

A controlled-temperature plankton wheel

A controlled-temperature plankton wheel is described that is suitable for use on board a ship. The IMER plankton wheel system allows the use of various sizes of experimental bottles, up to 2.2 litres, the simulation of ambient light regimes and variable speed control for the rotation of the experimental bottles. The flexibility of the system was demonstrated by investigating the relationship between temperature and ingestion rate of an herbivorous copepod. Using four of the IMER plankton wheels simultaneously at four different temperatures (5°, 10°, 15° and 20° C), the ingestion rate of Calanus helgolandicus, feeding on Thalassiosira weissflogii, was shown to increase with increasing temperature; from a transformation of log_e (ingestion rate), this relationship was calculated as a Q₁₀ (10° to 20°C) for Copepodite Stage V (Q₁₀ 4.5) and adult female (Q₁₀ 2.7) C. helgolandicus. The possibility of damaging cells, by rotation at 2 rpm, was investigated using the spinose form of the diatom T. weissflogii. Such rotation did not cause any damage to the spines of T. weissflogii, but mixing this diatom with a magnetic stirrer bar did damage the spines to varying degrees, depending on the volume being mixed.     

Genetic evidence for two sibling species within Polydora cf. ciliata (Polychaeta: Spionidae) from the Sea of Japan

Two morphologically indistinguishable but genetically very distinct polychaete species of the family Spionidae are found in the Peter the Great Bay of the Sea of Japan. Both species bore into molluscan shells and occur sympatrically. They are close in morphology to Polydora ciliata (Johnston, 1838) and some other members of the redefined P. ciliata species complex. The systematics of the two species is still under consideration by one of us (V.I.R.). A total of nine enzyme systems comprising 12 isozyme loci were examined in both the species by means of starch gel electrophoresis. Nei's genetic distance between them proved surprisingly high (D= 1.942) being the highest among estimates so far obtained for pairs of polydorid species. The levels of intraspecific genetic variation in both species (Polydora sp. 1, H _e= 0.287 ± 0.064; Polydora sp. 2, H _e= 0.111 ± 0.045) are comparable to those of other electrophoretically studied polydorids. The problem of taxonomic integrity of cosmopolitan marine invertebrate species is briefly discussed as well as the usefulness of isozymes as species-specific characters suitable for unequivocal discrimination between sibling polydorid species. A list of enzymes characterized by interspecifically variable numbers of expressed isozymes within polydorids is given and recommended for such a discrimination. Received: 22 July 1997 / Accepted: 19 February 1998     

Daily energy requirements and trophic positioning of the sand shrimp<Emphasis Type="Italic"> Crangon septemspinosa</Emphasis>

Sand shrimp, Crangon septemspinosa Say, are important to the trophic dynamics of coastal systems in the northwestern Atlantic. To evaluate predatory impacts of sand shrimp, daily energy requirements (J ind.^–1 day^–1) were calculated for this species from laboratory estimates of energy losses due to routine (R_R), active (R_A), and feeding (R_SDA) oxygen consumption rates (J ind.^–1 h^–1), coupled with measurements of diel motile activity. Shrimp used in this study were collected biweekly from the Niantic River, Connecticut (41°33N; 72°19W) during late spring and summer of 2000 and 2001. The rates of shrimp energy loss due to R_R and R_A increased exponentially with increasing temperature, with the magnitude of increase greater between 6°C and 10°C (Q₁₀=3.01) than between 10°C and 14°C (Q₁₀=2.85). Rates of R_R doubled with a twofold increase in shrimp mass, and R_SDA was 0.130 J h^–1+R_R, irrespective of shrimp body size. Shrimp motile activity was significantly greater during dark periods relative to light periods, indicating nocturnal behavior. Nocturnal activity also increased significantly at higher temperatures, and at 20°C shifted from a unimodal to a bimodal pattern. Laboratory estimates of daily metabolic expenditures (1.7–307.4 J ind.^–1 day^–1 for 0.05 and 1.5 g wet weight shrimp, respectively, between 0°C and 20°C) were combined with results from previous investigations to construct a bioenergetic model and make inferences regarding the trophic positioning of C. septemspinosa. Bioenergetic model estimates indicated that juvenile and adult shrimp could meet daily energy demands via opportunistic omnivory, selectively preying upon items of high energy content (e.g. invertebrate and fish tissue) and compensating for limited prey availability by ingesting readily accessible lower energy food (e.g. detritus and plant material).Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by J.P. Grassle, New Brunswick     

Genetic evidence of cryptic speciation within hammerhead sharks (Genus Sphyrna)

Surveys of genetic variation within cosmopolitan marine species often uncover deep divergences, indicating historical separation and potentially cryptic speciation. Based on broad geographic (coastal eastern North America, Gulf of Mexico, western Africa, Australia, and Hawaii) and temporal sampling (1991–2003), mitochondrial (control region [CR] and cytochrome oxidase I [COI]) and nuclear gene (lactate dehydrogenase A intron 6 [LDHA6]) variation among 76 individuals was used to test for cryptic speciation in the scalloped hammerhead, Sphyrna lewini (Griffith and Smith). CR and COI gene trees confirmed previous evidence of divergence between Atlantic and Indo-Pacific scalloped hammerhead populations; populations were reciprocally monophyletic. However, the between-basin divergence recorded in the mtDNA genome was not reflected in nuclear gene phylogenies; alleles for LDHA6 were shared between ocean basins, and Atlantic and Indo-Pacific populations were not reciprocally monophyletic. Unexpectedly, CR, COI, and LDHA6 gene trees recovered a deep phylogenetic partition within the Atlantic samples. For mtDNA haplotypes, which segregated by basin, average genetic distances were higher among Atlantic haplotypes (CR: D _HKY=0.036, COI: D _GTR=0.016) than among Indo-Pacific haplotypes (CR: D _HKY=0.010, COI: D _GTR=0.006) and approximated divergences between basins for CR (D _HKY=0.036 within Atlantic; D _HKY=0.042 between basins). Vertebral counts for eight specimens representing divergent lineages from the western north Atlantic were consistent with the genetic data. Coexistence of discrete lineages in the Atlantic, complete disequilibrium between nuclear and mitochondrial alleles within lineages and concordant partitions in genetic and morphological characters indicates reproductive isolation and thus the occurrence of a cryptic species of scalloped hammerhead in the western north Atlantic. Effective management of large coastal shark species should incorporate this important discovery and the inference from sampling that the cryptic scalloped hammerhead is less abundant than S. lewini, making it potentially more susceptible to fishery pressure.     

Effects of delayed feeding and temperature on the age of irreversible starvation and on the rates of growth and mortality of Pacific herring larvae

The time periods from exhausion of the yolk to the age of irreversible starvation for Pacific herring Clupea harengus pallasi larvae were 8.5, 7.0 and 6.0 d at 6°, 8° and 10°C, respectively. These periods are within the range perviously measured for Atlantic herring larvae and other temperature zone fish species; they are long compared to the periods for tropical species. The variation in the length of this period is due almost entirely to temperature; the natural logarithm of the time period from fertilization to irreversible starvation is highly correlated (r=0.91) with the mean rearing temperature for 25 species of pelagic marine fish larvae. The rates of growth and mortality, measured for 26 experimental populations of Pacific herring larvae reared at 6°, 8° and 10°C and ten ages of delayed first feeding, decreased and increased, respectively with increasing age of first feeding and increasing temperature. These rates, adjusted for the effects of rearing conditions, were compared with the rates for natural populations of herring larvae. Growth is generally faster in the sea than in experimental enclosures. Two of the eleven estimates of natural mortality rate were high enough to indicate possible catastrophic mass starvation. This is consistent with Hjort's critical period concept of year class formation and it suggests that mass starvation occurs in 18 to 36% of the natural populations of first feeding herring larvae.     

Genetic study of the extent and consequences of sexual and asexual reproduction in the deep-sea epizoic anemones Amphianthus inornata and Kadosactis commensalis (Cnidaria: Anthozoa)

The anemone Amphianthus inornata is found at bathyal depths living on colonies of the gorgonian Acanella arbuscula. Previous studies of the morphology and reproductive stage of this anemone, during different times of the year, have indicated that it reproduces sexually on a seasonal basis. A small proportion of the study population were also reported to be undergoing asexual reproduction by fission. The anemone Kadosactis commensalis is also bathyal, but lives mainly on the holothurian Paroriza prouhoi. Previous morphological studies have indicated that K. commensalis is a protandrous hermaphrodite that exhibits non-seasonal sexual reproduction only. In the present study, allozyme electrophoresis was used to examine the prevalence and genetic consequences of asexual reproduction in a population of Amphianthus inornata from 2 200 m in the Rockall Trough, North Atlantic Ocean. Genetic evidence, from five randomly selected polymorphic enzyme loci, for asexual reproduction in this species was weak. Exact tests indicated that genotype frequencies did not differ significantly from those expected under Hardy–Weinberg equilibrium. F _IS (correlation of homologous alleles with reference to local population, assuming random mating) values also did not differ significantly from zero, and observed heterozygosity (H _o =0.446) and genotypic diversity (G _o =17.0387) were very similar to Hardy–Weinberg expected frequencies (H _e =0.446; G ^* _e =17.0010). Evidence suggests that the contribution of asexual reproduction to recruitment in the study population of A. inornata is low. For a single population of K. commensalis from 4 850 m on the Porcupine Abyssal Plain, North Atlantic Ocean, the hypothesis that inbreeding due to reduced occurrence of outcrossing between anemones on a single holothurian host was examined by electrophoresis of ten randomly selected enzyme loci. Single-locus genotypic frequencies were significantly different from expected frequencies for one locus P≤ 0.05, hexokinase-1 (Hex-1)]. F _IS values were significantly different from zero for two enzyme loci (Hex-1 and Hex-2, P≤ 0.01 and P≤ 0.05, respectively), and the overall observed heterozygosity was lower than the expected heterozygosity (H _o =0.125, H _e =0.140). The hypothesis of inbreeding could not be rejected by the present study, although sample size was small (N=55), leading to possible bias in tests of significance. Genetic variation in A. inornata was higher than that recorded for most eukaryotes, although interlocus sampling error for only five loci is high. High genetic variability has been found in other sea anemones, and has been related to high longevity and mixed reproductive modes. Genetic variation in K. commensalis was in the higher range of that found in other eukaryotes, and is not unusual for anemones. Received: 5 August 1996 / Accepted: 11 December 1996