Experimental Determination of Salinity, Temperature, Growth, and Metabolic Effects on Shell Isotope Chemistry of Mytilus edulis Collected from Maine and Greenland

To study the effects of temperature, salinity, and life processes (growth rates, size, metabolic effects, and physiological/ genetic effects) on newly precipitated bivalve carbonate, we quantified shell isotopic chemistry of adult and juvenile animals of the intertidal bivalve Mytilus edulis (Blue mussel) collected alive from western Greenland and the central Gulf of Maine and cultured them under controlled conditions. Data for juvenile and adult M. edulis bivalves cultured in this study, and previously by Wanamaker et al. (2006), yielded statistically identical paleotemperature relationships. On the basis of these experiments we have developed a species-specific paleotemperature equation for the bivalve M. edulis [T degrees C = 16.28 (+/- 0.10) -4.57 (+/- 0.15) {delta(18)O(c) VPBD - delta(18)O(w) VSMOW} + 0.06 (+/- 0.06) {delta(18)O(c) VPBD - delta(18)O(w) VSMOW}(2); r(2) = 0.99; N = 323; p < 0.0001]. Compared to the Kim and O'Neil (1997) inorganic calcite equation, M. edulis deposits its shell in isotope equilibrium (delta(18)O(calcite)) with ambient water. Carbon isotopes (delta(13)C(calcite)) from sampled shells were substantially more negative than predicted values, indicating an uptake of metabolic carbon into shell carbonate, and delta(13)C(calcite) disequilibrium increased with increasing salinity. Sampled shells of M. edulis showed no significant trends in delta(18)O(calcite) based on size, cultured growth rates, or geographic collection location, suggesting that vital effects do not affect delta(18)O(calcite) in M. edulis. The broad modern and paleogeographic distribution of this bivalve, its abundance during the Holocene, and the lack of an intraspecies physiologic isotope effect demonstrated here make it an ideal nearshore paleoceanographic proxy throughout much of the North Atlantic Ocean.

Experimental Determination of Salinity, Temperature, Growth, and Metabolic Effects on Shell Isotope Chemistry of Mytilus Edulis Collected from Maine and Greenland

To study the effects of temperature, salinity, and life processes (growth rates, size, metabolic effects, and physiological/ genetic effects) on newly precipitated bivalve carbonate, we quantified shell isotopic chemistry of adult and juvenile animals of the intertidal bivalve Mytilus edulis (Blue mussel) collected alive from western Greenland and the central Gulf of Maine and cultured them under controlled conditions. Data for juvenile and adult M. edulis bivalves cultured in this study, and previously by Wanamaker et al. (2006), yielded statistically identical paleotemperature relationships. On the basis of these experiments we have developed a species-specific paleotemperature equation for the bivalve M. edulis [T degrees C = 16.28 (+/- 0.10) -4.57 (+/- 0.15) {delta(18)O(c) VPBD - delta(18)O(w) VSMOW} + 0.06 (+/- 0.06) {delta(18)O(c) VPBD - delta(18)O(w) VSMOW}(2); r(2) = 0.99; N = 323; p < 0.0001]. Compared to the Kim and O'Neil (1997) inorganic calcite equation, M. edulis deposits its shell in isotope equilibrium (delta(18)O(calcite)) with ambient water. Carbon isotopes (delta(13)C(calcite)) from sampled shells were substantially more negative than predicted values, indicating an uptake of metabolic carbon into shell carbonate, and delta(13)C(calcite) disequilibrium increased with increasing salinity. Sampled shells of M. edulis showed no significant trends in delta(18)O(calcite) based on size, cultured growth rates, or geographic collection location, suggesting that vital effects do not affect delta(18)O(calcite) in M. edulis. The broad modern and paleogeographic distribution of this bivalve, its abundance during the Holocene, and the lack of an intraspecies physiologic isotope effect demonstrated here make it an ideal nearshore paleoceanographic proxy throughout much of the North Atlantic Ocean.

Sexually Different Growth Histories of the American Eel in Four Rivers in Maine

Growth histories of yellow-phase American eels Anguilla rostrata collected in four rivers in Maine, were back-calculated from sagittal otolith increments. Our objectives were to first determine whether sexually dimorphic growth rates exist and then compare the growth histories of American eels from four rivers within a geographic region. For female eels, the maximum growth rate was 31.9 +/- 1.7 mm/year at age 8, decreasing to 25.1 +/- 2.9 mm/year at age 14. Males attained a maximum of 29.8 +/- 1.6 min/year at age 3, decreasing to a minimum of 17.9 +/- 1.3 mm/year at age 11. Females grew faster than males after age 4 and had a slower reduction in growth rate with age. These faster growth rates among females were similar in all four rivers. The observed growth rates are not consistent with current life history hypotheses and may indicate an alternative life history strategy. Because female eels benefit from a larger size (i.e., size refuge, increased fecundity, and greater niche breadth), they would benefit from a higher-risk growth strategy that increases growth rate during their earlier years and reduces the amount of time spent in an unfavorable size-class. The tradeoffs (i.e., mortality, developmental rate, pathogen resistance, and longevity) associated with this faster growth rate may not favor the males' life history requirements. Male eels do not achieve the size of females and therefore are not subject to the advantages associated with being larger. Therefore, they may use a risk-averse strategy that maintains submaximum growth rates to obtain the minimum size necessary to mature and complete the spawning migration while reducing the adverse affects of faster growth rates. We postulate that, in eels, intrinsic growth rates should be considered a life history trait that has evolved to meet the life history requirements of each sex.

Growth and Persistence of Diverse Intertidal Crusts: Survival of the Slow in a Fast-Paced World

Encrusting algae are conspicuous components of hard-substratum benthic communities in the photic zone despite being poor competitors and slow growers. Little is known about their growth rates or about mechanisms controlling key processes such as wound healing and surviving overgrowth. We manipulated 12 crustose species (including red and brown algae and a lichen) from the intertidal zone of Washington, USA, studying their varying responses to identical experimental conditions. Three of 8 crust species tested showed increased growth rates with size. Species healed from standardized wounds at different rates and using different mechanisms (e.g. lateral vs vertical regeneration) as seen in cross-sections. Three species showed altered growth rates at unwounded margins of wounded crusts, suggesting intrathallus communication. Year-long experiments involving simulated overgrowth showed that some species can maintain healthy tissue in a covered area, and one (the coralline Lithothamnion phymatodeum) even grew new tissue there. Other species gradually lost color, thickness, and area in covered areas. Hildenbrandia occidentalis survived remarkably well when covered, possibly due to its very slow growth and low metabolic demand. One suggested mechanism underlying the high variation in responses among crusts is the degree to which their thalli may be anatomically integrated by features such as cell fusions; physiological work testing translocation via these features is needed. Other mechanisms allowing persistence include rapid wound healing and frequent recruitment.

Age, Growth and Population Structure of Jumbo Flying Squid, Dosidicus Gigas, Based on Statolith Microstructure Off the Exclusive Economic Zone of Chilean Waters

The jumbo flying squid, Dosidicus gigas, support an important squid fishery off the Exclusive Economic Zone of Chilean waters. However, we only have limited information about their biology. In this study, age, growth and population structure of D. gigas were studied using statoliths from 333 specimens (386 females and 147 males) randomly sampled in the Chinese squid jigging surveys from 2007 to 2008 off the Exclusive Economic Zone of Chile. Mantle lengths (MLs) of the sample ranged from 206 to 702 mm, and their ages were estimated from 150 to 307 days for females and from 127 to 302 days for males. At least two spawning groups were identified, the main spawning peak tended to occur between August and November (austral spring group), and the secondary peak appeared during March to June (austral autumn group). The ML-age relationship was best modelled by a linear function for the austral spring group and a power function for the austral autumn group, and the body weight (BW)-age relationship was best described by an exponential function for both the groups. Instantaneous relative growth rates and absolute growth rates for ML and BW did not differ significantly between the two groups. The growth rate of D. gigas tended to be high at young stages, and then decreased after the sub-adult stage (>180 days old). This study suggests large spatial and temporal variability in key life history parameters of D. gigas, calling for the collection of more data with fine spatial and temporal scales to further improve our understanding of the fishery biology of D. gigas.

Implications of Longterm Diameter-Limit Harvesting: Effects on Radial Growth of Red Spruce (Picea rubens) and Genetic Diversity of White Pine (Pinus strobus)

For over 3 centuries, diameter-limit harvesting has been a predominant logging method in the northeastern United States. Silvicultural theory asserts that such intensively selective harvesting can lead to genetic degradation. A decrease in softwood productivity has recently been reported in Maine - has a long history of dysgenic selection degraded the genetic resources of Maine softwoods, contributing to a decrease in growth and productivity? This study examines two aspects of potential implications of diameter-limit harvesting: effects on residual phenotypes of red spruce and impacts on genetic diversity of white pine. Radial growth of residual red spruce trees in stands experiencing 50 years of fixed diameter-limit harvesting was measured using annual increment rings and compared with residual red spruce trees in positive selection stands. Trees remaiaing after several rounds of diameter-limit harvesting exhibited sigdicantl y smaller radial sizes throughout their lives, and displayed significantly slower growth rates for the first 80 years of measured growth. These results strongly suggest that the largest and fastest-growing genotypes and their respective gene complexes determining good radial growth have been removed from the diameter-limit stand. Dysgenic selection can be observed in fixed diarneter-limit stands, resulting in a diminished genetic resource and decreased residual stand value. To examine more direct genetic implications of long-term diameter-limit harvesting, microsatellite DNA markers were implemented to study genetic diversity of eastern white pine in Maine. Three age groups of trees were studied: mature trees older than 200 years, juvenile trees 5-30 years old, and embryos. Trees were genotyped at 10 microsatellite loci. Overall genetic diversity levels of eastern white pine in Maine were extremely high, with an average observed heterozygosity of 0.762. Genetic differentiation was minimal among and between all three age groups, although an excess of heterozygotes was shown in the mature and juvenile groups that was not reflected in the embryo group, which actually had a slight heterozygote deficiency. Allele frequencies did not differ significantly between age groups, but did reveal more rare and low frequency alleles in the embryo groups than in the mature group. Overall, low frequency alleles comprise the largest portion of alleles in the sample population, with no common alleles evident overall. These results suggest that significant genetic degradation has either not occurred for white pine, or that the results of dysgenic selection have not yet emerged. It is clear, however, that selective harvesting could result in a loss of low frequency alleles, which are a primary reserve of evolutionary potential in a species. Implications of these studies affect industrial forestry, regional economics, and ecological concerns for the northeast. Long-term diameter-limit harvesting can lead to a degradation of residual phenotypes, and an overall decrease in stand quality. Potentially, a loss of low frequency, locally adapted alleles could result in a decrease of allelic richness and degradation of the regidnal genetic resource. Decreased genetic variation can lead to seriously limited evolutionary potential of species and ecosystems, particularly in rapidly changing environments. Based on these findings, I recommend a reassessment of any harvesting prescription that includes fixed diameter-limit removals, particularly for species that have low natural genetic diversity levels or a limited natural range, such as red spruce. Maintenance of a healthy genetic reserve can avoid effects of dysgenic harvesting.

Model-Based Estimates of Calanus Finmarchicus Abundance in the Gulf of Maine

Ocean observing systems and satellites routinely collect a wealth of information on physical conditions in the ocean. With few exceptions, such as chlorophyll concentrations, information on biological properties is harder to measure autonomously. Here, we present a system to produce estimates of the distribution and abundance of the copepod Calanus finmarchicus in the Gulf of Maine. Our system uses satellite-based measurements of sea surface temperature and chlorophyll concentration to determine the developmental and reproductive rates of C. finmarchicus. The rate information then drives a population dynamics model of C. finmarchicus that is embedded in a 2-dimensional circulation field. The first generation of this system produces realistic information on interannual variability in C. finmarchicus distribution and abundance during the winter and spring. The model can also be used to identify key drivers of interannual variability in C. finmarchicus. Experiments with the model suggest that changes in initial conditions are overwhelmed by variability in growth rates after approximately 50 d. Temperature has the largest effect on growth rate. Elevated chlorophyll during the late winter can lead to increased C. finmarchicus abundance during the spring, but the effect of variations in chlorophyll concentrations is secondary to the other inputs. Our system could be used to provide real-time estimates or even forecasts of C. finmarchicus distribution. These estimates could then be used to support management of copepod predators such as herring and right whales.

Carbon-Based Ocean Productivity and Phytoplankton Physiology from Space

Ocean biogeochemical and ecosystem processes are linked by net primary production (NPP) in the ocean's surface layer, where inorganic carbon is fixed by photosynthetic processes. Determinations of NPP are necessarily a function of phytoplankton biomass and its physiological status, but the estimation of these two terms from space has remained an elusive target. Here we present new satellite ocean color observations of phytoplankton carbon (C) and chlorophyll (Chl) biomass and show that derived Chl:C ratios closely follow anticipated physiological dependencies on light, nutrients, and temperature. With this new information, global estimates of phytoplankton growth rates (mu) and carbon-based NPP are made for the first time. Compared to an earlier chlorophyll-based approach, our carbon-based values are considerably higher in tropical oceans, show greater seasonality at middle and high latitudes, and illustrate important differences in the formation and demise of regional algal blooms. This fusion of emerging concepts from the phycological and remote sensing disciplines has the potential to fundamentally change how we model and observe carbon cycling in the global oceans.

Trace Elements in the Statoliths of Jumbo Flying Squid Off the Exclusive Economic Zones of Chile and Peru

Ontogenetic variation in 4 trace element ((88)Sr, (137)Ba, (24)Mg, (23)Na) concentrations and their ratios to Ca were measured in statoliths of the jumbo flying squid Dosidicus gigas off the Exclusive Economic Zone of Chilean and Peruvian waters using laser ablation inductively coupled plasma mass spectrometry (LA-ICP-MS). The element compositions of statoliths showed no significant differences between females and males. All of the elements in different growth zones showed significant variations, except for Mg. Sr:Ca and Mg:Ca were good indicators for distinguishing squid from autumn and winter spawning seasons. Sr:Ca and Ba:Ca distribution patterns in statoliths confirmed that paralarvae and juvenile squid inhabit surface waters, while subadult squid migrate into deeper waters. An increasing Sr: Ca ratio of subadult squid could be explained by declining temperature gradients from northern to southern sampling locations, although no significant Sr: Ca differences were observed (p > 0.05). Mg:Ca ratios decreased progressively from the nucleus to the peripheral zone, which might be correlated with statolith growth rates. Na:Ca ratios slightly declined from paralarvae to the subadult phase. Quantitative relationships between statolith trace elements and environmental conditions under different growth stages are needed to improve our understanding of life history of D. gigas.