Patterns of larval and early juvenile growth in a semiterrestrial crab, Armases angustipes (Decapoda: Sesarmidae): comparison with a congener with abbreviated development

In a semiterrestrial and estuarine tropical crab, Armases angustipes Dana (Grapsoidea: Sesarmidae), changes in biomass (measured as dry mass, W; carbon, C; nitrogen, N; and hydrogen, H; per individual) and relative elemental composition (C, N, H, in percent of W; C:N mass ratio) were studied during development from an early egg stage through hatching, the complete larval phase, metamorphosis and the first juvenile crab stage (CI). In the megalopa and CI, growth was measured also within the moulting cycle, and biomass and elemental composition were determined in cast exuviae. From an early egg stage to the freshly hatched larva, A. angustipes lost about 20% of W, 29% of C, 5% of N and 32% of H. Proportionally higher losses in C than in N were reflected also in a significantly decreasing C:N mass ratio (from 5.02 to 3.74). These results indicate that lipids mobilised from yolk reserves represented the principal metabolic substrate for embryonic energy production, while proteins were catabolised at a much lower rate. The present data of growth and exuviation are compared with previously published data from a congener, A. miersii Rathbun, which has an abbreviated and facultatively lecithotrophic mode of larval development (with three instead of four zoeal stages; stages I and II in principle independent of food). When growth is measured as an increase in the final (premoult) biomass of successive developmental stages, both species show an exponential pattern. Within the moulting cycles of the megalopa and the first juvenile, both species show parabola-shaped growth curves, with a rapid biomass increase in postmoult and intermoult stages, and losses in the premoult phase. Thus, the two Armases species show, in general, similar patterns of larval and early juvenile growth. However, the initial size of eggs and larvae is about four times larger in A. miersii, and its biomass remains higher throughout the period of larval and early juvenile development. A. angustipes is able to partially make up for this difference, as it has an additional zoeal stage, and its megalopa and CI stages show higher relative biomass increments (in percent of initial values). Due to this compensatory growth pattern, A. angustipes reaches in its CI stage about half the biomass of a juvenile A. miersii. When exuvial losses of megalopae and juveniles are compared between these two species, A. miersii shows higher biomass losses per individual (corresponding with its larger size), but lower relative losses (C, N, H, in percent of late premoult body mass or in percent of previously achieved growth increments). Differences in larval and early juvenile growth and in the exuvial losses of megalopae and juveniles of these two congeners are discussed in relation to their differential ecology, life history and reproductive strategy.     

Linking the EU emissions trading scheme: economic implications of allowance allocation and global carbon constraints

We investigate the role of domestic allowance allocation and global emissions constraints for the carbon-market impacts of linking the EU Emissions Trading Scheme (ETS) internationally. Employing a quantitative simulation model of the global carbon market, we find that the economic benefits from connecting the European ETS to emerging non-EU schemes strongly depend on the regional allowance allocation of the linking participants: In a world of moderate carbon constraints, an economically efficient regional allowance allocation induces a much stronger fall in total compliance costs than a sub-optimal (i.e. too high) domestic allocation of emissions permits. However, a more efficient (i.e. stricter) allocation shifts abatement efforts and compliance costs to energy-intensive industries which are covered by the domestic ETS. We further find that committing to ambitious global emissions reduction targets (compatible with stabilizing CO₂ concentrations at 450 ppm) induces much stronger regional abatement efforts and substantially higher compliance costs for the abating regions. In such an ambitious climate policy regime, an efficient domestic allocation of allowances is even more important from an economic perspective: Here, linking emissions trading schemes diminishes the associated compliance costs on the largest scale.

Digestive enzymes in the ontogenetic stages of the southern king crab, Lithodes santolla

The early ontogenetic stages of the sub-Antarctic king crab Lithodes santolla were analyzed for the presence and activities of a set of important digestive enzymes. The eggs and non-feeding larvae (zoea I-III, megalopa) showed high activities of esterases, phosphatases, and exopeptidases indicating the enzymatic ability to utilize endogeneous yolk reserves. SDS-PAGE showed a continuous decrease of proteins or proteids in the range of 59–81 kDa during ontogenetic development from the eggs through the zoeal stages to the first juvenile crab stage, CI. This reduction reflects the degradation of storage compounds during lecithotrophic larval development. Activities of the endopeptidases, trypsin and chymotrypsin, were low in eggs and larvae but increased significantly in the first juvenile crab stage. These enzymes typically facilitate the first steps of proteolysis in the extra-cellular spaces of the midgut gland and in the stomach. Their scarcity indicates that the larvae of L. santolla are physiologically not prepared to digest external food. This ability seems to appear first in the CI stage. Extracts of juvenile midgut glands and the gastric fluids of adults showed high activities of a variety of digestive enzymes including phosphatases, carbohydrases, as well as endo- and exopeptidases. High activities of digestive enzymes in adults may compensate for scarce food supply and rate-limiting low temperatures in the predominantly sub-Antarctic habitats of L. santolla.     

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.     

Energetics,moult cycle and ecdysteroid titers in spider crab (Hyas araneus) larvae starved after the D0 threshold

Zoea I larvae of Hyas araneus L. (Decapoda: Brachyura: Majidae) were dredged in January 1986 from the German Bight and reared in the laboratory at constant 12°C, until they reached the transition of stages C/D₀ of the moult cycle (4 d after hatching). This developmental stage had previously been found to correspond with the point of reserve saturation (PRS) which allows autonomous (food-independent) development through the rest of the moult cycle and hence, was termed the D₀ threshold. One part of the larvae was continually fed (control), another group was starved from the D₀ threshold until moulting to the zoea II instar. In these two experimental groups, as well as in the two groups of zoea II larvae obtained from the different feeding conditions, the course of the moult cycle, biomass (dry weight, W; carbon, C; nitrogen, N; hydrogen, H; energy, E; the latter estimated from C), and ecdysteroid titers (measured with a radio-immuno-assay as ecdysone equivalents) were investigated. When the larvae reached the PRS, they had gained 90% in W, 72% in C, 32% in N, 53% in H, and 65% in E, since hatching, corresponding to an accumulation of 87% of final W and 62 to 69% of C, N, and H reached later, at the end of the mould cycle in the control. The period of starvation caused a 2.5-d delay of the moult cycle, mainly in late premoult, and significant losses of biomass and energy. Starved and fed larvae secreted similar amounts of moulting hormone per individual, but with a reduced rate in the starved group, thus causing developmental delay. Zoea II larvae moulting after starvation contained less than half of the control biomass and energy, and even less than a freshly hatched zoea I. Growth rate was only slightly enhanced in these zoea II larvae as compared to the fed control, but losses of biomass, mainly of lipids, were partly compensated by a 4-d prolongation of their moult cycle, chiefly (3 d) in stage C. Biomass curves were almost parallel in the two experimental groups of zoea II larvae, with significantly higher values in the control during all stages of the moult cycle. However, similar relative proportions (74 to 89%) of late premoult biomass and energy were reached at the D₀ threshold, regardless of different feeding history and initial or final values in a given group. The ecdysteroid titer curve of the zoea II which had moulted from starved zoea I was very similar to that in control larvae, but with a 3-d delay in the occurrence of premoult peak concentration (in both groups in stage D₁). Regulation and coordination of moult cycle, ecdysteroid titers, and growth in the larval development of decapod crustaceans are discussed, with special reference to the D₀ threshold.     

Physico-chemical and biological treatment of MSW landfill leachate

This paper analyses the evolution of the physico-chemical characteristics of the leachate from the Central Landfill of Asturias (Spain), which has been operating since 1986, as well as different treatment options. The organic pollutant load of the leachate, expressed as chemical oxygen demand (COD), reached maximum values during the first year of operation of the landfill (around 80,000 mg/L), gradually decreasing over subsequent years to less than 5000 mg/L. The concentration of ammonium, however, has not decreased, presenting values of up to 2000 mg/L. When feasible, recirculation can greatly decrease the organic matter content of the leachate to values of 1500–1600 mg COD/L. Applying anaerobic treatment to leachates with a COD between 11,000 and 16,000 mg/L, removal efficiencies of 80–88% were obtained for organic loading rates of 7 kg COD/m³ d. For leachates with lower COD (4000–6000 mg/L), the efficiency decreased to around 60% for organic loading rates of 1 kg COD/m³ d.Applying coagulation–flocculation with iron trichloride or with aluminium polychloride, it was possible to reduce the non-biodegradable organic matter by 73–62% when treating old landfill leachate (COD: 4800 mg/L, BOD₅: 670 mg/L), also reducing turbidity and colour by more than 97%. It is likewise possible to reduce the non-biodegradable organic matter that remains after biological treatment by adsorption with activated carbon, although adsorption capacities are usually low (from 15 to 150 mg COD/g adsorbent). As regards ammonium nitrogen, this can be reduced to final effluent values of 5 mg/L by means of nitrification/denitrification and to values of 126 mg/L by stripping at pH 12 and 48 h of stirring.     

Physiological and biochemical changes during lecithotrophic larval development and early juvenile growth in the northern stone crab,Lithodes maja (Decapoda: Anomura)

Larvae of the northern stone crab, Lithodes maja L., were reared in the laboratory from hatching to the second crab stage. complete larval development (at constant 9°C) lasted about 7 wk, invariably consisting of three pelagic zoeal stages and a semibenthic Megalopa; only two zoeal stages have been described in the literature. All larval stages are lecithotrophic. First feeding was consistently observed only after metamorphosis, in the first juvenile crab stage. In short intervals (every 1 to 5 d), developmental changes in biomass, B (expressed as: dry weight, W; carbon, C; nitrogen, N; hydrogen, H) and oxygen consumption (respiration, R) were measured in larvae and early juveniles; additionally, protein and carbohydrates were measured, but only in the zoeal stages and early Megalopa. Unusually high C contents (varying between 56 and 61% of W in eggs and freshly hatched Zoea I larvae from 12 different females) and high C:N weight ratios (8 to 11) indicate enhanced initial lipid stores, which are utilized as the major metabolic substrate during both embryonic and lecithotrophic larval development. Predominant degradation of lipids is shown indirectly; the C:N ratio decreased significantly, from 10 (at hatching) to 6 (at metamorphosis), while larval protein decreased only little, from ca. 55% of W (at hatching) to 48% (in the Megalopa). From hatching to metamorphosis, about 27% of the initially present W, 48% of C, 18% of N, and 52% of H were lost. This decrease in larval biomass can be described as an exponential function of development time. The major part of these losses were associated with metabolic energy requirements, while exuvial losses were comparably small. In each of the zoeal stages, only about 1 to 2% of late premoult (LPM) B was shed with the exuvia. The Megalopa, which produces a much thicker, calcified exoskeleton, lost 20% of LPM W, but only 5 to 8% of organic constituents (C, N, H). Much higher exuvial losses were measured in the Crab I stage (51% in W, 21% in C, 5% in N, and 7% in H). Maximum respiration was found in the actively swimming zoeal stages, a minimum in the predominantly benthic, mostly inactive Megalopa. The Crab I stage exhibits also a sluggish behaviour and low R, in spite of beginning food uptake and growth. Immediately after metamorphosis, the juvenile crab gained rapidly in W, in particular in its C fraction. A transitorily steep increase in the C:N ratio indicates a replenishment of partially depleted lipid stores, but also a rapid initial increase of inorganic C in the heavily calcified exoskeleton. Instantaneous rates of growth, assimilation, and net growth efficiency (K ₂) were high during the initial (postmoult) phase in the first juvenile crab stage (C-specific growth rate: 6% d^-1; K ₂:70%), but decreased towards zero values during laterstages of the moulting cycle; metabolism remained practically constant during the Crab I stage. Entirely lecithotrophic larval development from hatching to metamorphosis in L. maja is considered an adaptation to seasonally short and limited planktonic food production in subarctic regions of the northern Atlantic.     

Growth and elemental composition (C,N, H) in Inachus dorsettensis (Decapoda: Majidae) larvae reared in the laboratory

Larvae of the spider crab Inachus dorsettensis were reared in the laboratory at constant 12 °C. Development lasted 8 to 10 d in the Zoea I, 10 to 12 d in the Zoea II and 14 to 20 d in the megalopa stage. During this time, larval growth was measured in samples taken every 2 to 4 d as dry wt (W), carbon (C), nitrogen (N), and hydrogen (H); energy content (E) was calculated from C. Biomass and energy (per individual) increased in each larval stage as a parabola-shaped function of age, which could be fitted by a power equation. C, H, and E show a higher percentage gain (relative to the initial values at hatching) than W or N, suggesting that proportionally more lipid than protein is accumulated during larval development. There are cyclical changes in the relative (per unit of W) biomass and energy figures, corresponding to the larval moult cycles: immediately after each ecdysis all these values decrease, presumably due to rapid uptake of water and minerals, then they increase again due to tissue growth and remain high until the next moulting occurs. Cyclical changes in the C/N ratio suggest that proportionally more lipid than protein is accumulated during the initial (postmoult) phase of the moult cycle, followed by a period of balanced or protein-dominated gain during the intermoult and premoult phases. These patterns of growth and elemental composition observed during the complete larval development and in single moult cycles of I. dorsettensis are compared with those described in the literature for other decapod species. This comparison suggests a high degree of similarity in biochemical composition and growth characteristics of larval decapod crustaceans.     

China’s air pollution reduction efforts may result in an increase in surface ozone levels in highly polluted areas

China, as a fast growing fossil-fuel-based economy, experiences increasing levels of air pollution. To tackle air pollution, China has taken the first steps by setting emission–reduction targets for nitrogen oxides (NO_x) and sulphur dioxide (SO₂) in the 11th and 12th Five Year Plans. This paper uses two models—the Energy–Environment–Economy Model at the Global level (E3MG) and the global Chemistry Transport Model pTOMCAT—to test the effects of these policies. If the policy targets are met, then the maximum values of 32 % and 45 % reductions below ‘business as usual’ in the monthly mean NO_x and SO₂ concentrations, respectively, will be achieved in 2015. However, a decrease in NO_x concentrations in some highly polluted areas of East, North-East and South-East China can lead to up to a 10% increase in the monthly mean concentrations in surface ozone in 2015. Our study demonstrates an urgent need for the more detailed analysis of the impacts and designs of air pollution reduction guidelines for China.     

In-situ studies on the life cycle of Diastylis rathkei (Cumacea: Crustacea)

Aspects of the life cycle of the cumacean Diastylis rathkei Kröyer, the most important food item of demersal fishes in the Western Baltic Sea, were investigated from the Underwater Laboratory (UWL) Helgoland in Lübeck Bay. In August the population density was 10.1±3.6 individuals 100 cm^-2 in fine-medium sand (Md=292 m), and 3.6±2.4 individuals in coarser medium sand (Md=470 m). In both substrata, abundance values decreased until October, at each station to a different extent. Particular attention was paid to vertical migrations of D. rathkei. Plankton catches and numerous direct observations provided evidence that the pelagic phase, observed at night, is always combined with ecdysis. During short periods of high swarming and moulting activities, the population structure changed considerably: juveniles of the 3rd stage dominated in early August; thereafter they were replaced by prematures, the last juvenile stage. The first mature individuals were observed in October. Some data are given on growth rate.