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.     

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     

Assessment of acute toxicity of ��-cyhalothrin to a freshwater catfish, Clarias batrachus

Acute static bioassays were conducted for 96 h period with λ-cyhalothrin to determine its acute toxicity to a freshwater catfish, Clarias batrachus. The 96 h LC₅₀ value was estimated to be 5.00 μg l⁻¹ (95% confidence limit: 4.114–5.712). The alterations in behavioral pattern, such as change in the color of skin, hyperactivity, loss of balance, rapid swimming, increased surfacing activity, enhanced rate of opercular activity, as well as prominent rates of convulsions in treated fish were observed with the increasing concentrations of insecticide as compared to the control fish. The results indicate that λ-cyhalothrin is highly effective even at very low concentrations.     

Effect of body mass,temperature and food deprivation on oxygen consumption rate of common cuttlefish <Emphasis Type="Italic">Sepia officinalis</Emphasis>

Predictions of short and long term changes in Sepia officinalis metabolism are useful, since this species is both economically important for aquaculture and also is an ideal experimental laboratory organism. In this study standard and routine oxygen consumption rates of newly hatched and juvenile laboratory raised cuttlefish S. officinalis ranging between 0.04 and 18.48 g dry body mass (Dm), were measured over a range of temperatures (10, 15, 20 and 25°C). The mass exponent (b) ranged between 0.706 and 0.992 for standard oxygen consumption and between 0.694 and 0.990 for routine oxygen consumption. Oxygen consumption scaled allometrically (b = 0.7) with body mass for cuttlefish <2 g Dm and isometrically (b = 1) thereafter. No significant differences were apparent amongst the slopes of oxygen consumption and body mass at different temperatures for standard and routine oxygen consumption. However, the intercepts differed significantly amongst the regression lines, indicating a significant effect of temperature on the magnitude of oxygen consumption. The combined effect of temperature (T) and dry body mass (Dm) are best described by the following equations: cuttlefish <2 g, MO₂ = 0.116Dm^0.7111.086 ^T and >2 g, MO₂ = 0.076Dm^0.9831.091 ^T for standard oxygen consumption; cuttlefish <2 g, MO₂ = 0.538Dm^0.7291.057 ^T and >2 g, MO₂ = 0.225Dm^0.9621.081 ^T for routine oxygen consumption. Using these equations it was estimated that a cuttlefish of 1 g Dm held at 20°C, eating 5% Dm day⁻¹ and undergoing standard and routine metabolism consumes 21.3 and 35.4%, respectively of its total daily energy intake. Juvenile cuttlefish (3.32–5.08 g Dm) held at 15°C and deprived of food for 27 days maintained a stable standard oxygen consumption rate for the first 6 days following starvation. By the 18th day without food, oxygen consumption rate had declined by 53% and further declined to 65% below the standard oxygen consumption rate on the 27th day. Upon resumption of feeding, the respiration rate returned immediately to the initial level prior to food deprivation. The present study defines the basic energy requirements and general physiological state of young cuttlefish at temperatures of 10–25°C with and without food.     

Tail beat frequency as a predictor of swimming speed and oxygen consumption of saithe (<Emphasis Type="Italic">Pollachius virens</Emphasis>) and whiting (<Emphasis Type="Italic">Merlangius merlangus</Emphasis>) during forced swimming

Oxygen consumption and tail beat frequency were measured on saithe (Pollachius virens) and whiting (Merlangius merlangus) during steady swimming. Oxygen consumption increased exponentially with swimming speed, and the relationship was described by a power function. The extrapolated standard metabolic rates (SMR) were similar for saithe and whiting, whereas the active metabolic rate (AMR) was twice as high for saithe. The higher AMR resulted in a higher scope for activity in accordance with the higher critical swimming speed (U _crit) achieved by saithe. The optimum swimming speed (U _opt) was 1.4 BL s⁻¹ for saithe and 1.0 BL s⁻¹ for whiting with a corresponding cost of transport (COT) of 0.14 and 0.15 J N⁻¹ m⁻¹. Tail beat frequency correlated strongly with swimming speed as well as with oxygen consumption. In contrast to swimming speed and oxygen consumption, measurement of tail beat frequency on individual free-ranging fish is relatively uncomplicated. Tail beat frequency may therefore serve as a predictor of swimming speed and oxygen consumption of saithe and whiting in the field.     

The effect of acute hypoxia on swimming stamina at optimal swimming speed in flathead grey mullet <Emphasis Type="Italic">Mugil cephalus</Emphasis>

Flathead grey mullets Mugil cephalus are commonly found in Mediterranean lagoons, which are regularly subject to high environmental variations. Oxygen is one of the factors that shows extremely high variation. The objective of this study was to test the effects of acute hypoxia exposure at two experimental temperatures (i.e. 20 and 30°C) on the stamina (time to fatigue) in M. cephalus swimming at the minimal cost of transport (i.e. optimal swimming speed; U _opt). At each temperature, a relationship was established between swimming speed and oxygen consumption (MO₂). This allowed estimation of U _opt at 45 cm s⁻¹ (~1.12 Body Length s⁻¹). Independent of temperature, stamina at U _opt was significantly reduced in severe hypoxia, i.e. at 15% of air saturation (AS). In these conditions, oxygen supply appears therefore to be insufficient to maintain swimming, even at the low speed tested here. After the stamina test, MO₂ measured in fish tested at 15% AS was significantly higher than that measured after the test in normoxia. Therefore, we suggest that in hypoxia, fish used anaerobic metabolism to supplement swimming at U _opt, leading to an oxygen debt. Since flathead grey mullet is a hypoxia-tolerant species, it is possible that hypoxic conditions less severe than those tested here may reduce stamina at low speed in less tolerant species. In addition, we suggest that testing stamina at these speeds may be relevant in order to understand the effect of hypoxia on behavioural activities carried out at low speed, such as food searching.     

Determination of the swimming trajectory and speed of chain-forming dinoflagellate <Emphasis Type="Italic">Cochlodinium polykrikoides</Emphasis> with digital holographic particle tracking velocimetry

The marine dinoflagellate Cochlodinium polykrikoides is a harmful and highly motile algal species. To distinguish between the motility characteristics of solitary and chain-forming cells, the swimming trajectories and speeds of solitary cells and 2- to 8-cell chains of C. polykrikoides were measured using a digital holographic particle tracking velocimetry (PTV) technique. C. polykrikoides cells exhibited helical swimming trajectories similar to other dinoflagellate species. The swimming speed increased as the number of cells in the chain increased, from an average of 391 μm s⁻¹ (solitary cells) to 856 μm s⁻¹ (8-cell chain). The helix radius R and pitch P also increased as the number of cells in the chain increased. R increased from 9.24 μm (solitary cell) to 20.3 μm (8-cell chain) and P increased from 107 μm (solitary cell) to 164 μm (8-cell chain). The free thrust-generating motion of the transverse flagella and large drag reduction in the chain-forming cells seemed to increase the swimming speed compared to solitary cells. The measured swimming speeds agreed with those from field observations. The superior motility of chain-forming C. polykrikoides cells may be an important factor for its bloom, in addition to the factors reported previously.     

Swimming behavior of juvenile anchovies (<Emphasis Type="Italic">Anchoa</Emphasis> spp.) in an episodically hypoxic estuary: implications for individual energetics and trophic dynamics

Diel swimming behaviors of juvenile anchovies (Anchoa spp.) were observed using stationary hydroacoustics and synoptic physicochemical and zooplankton profiles during four unique water quality scenarios in the Neuse River Estuary, NC, USA. Vertical distribution of fish was restricted to waters with DO greater than 2.5 mg O₂ l⁻¹, except when greater than 70% of the water column was hypoxic and a subset of fish were occupying water with 1 mg O₂ l⁻¹. We made the prediction that an individual fish would select a swim speed that would maximize net energy gain given the abundance and availability of prey in the normoxic waters. During the day, fish adopted swim speeds between 7 and 8.8 bl s⁻¹ that were near the theoretical optimum speeds between 7.0 and 8.0 bl s⁻¹. An exception was found during severe hypoxia, when fish were swimming at 60% above the optimum speed (observed speed = 10.6 bl s⁻¹, expected = 6.4 bl s⁻¹). The anchovy is a visual planktivore; therefore, we expected a diel activity pattern characteristic of a diurnal species, with quiescence at night to minimize energetic costs. Under stratified and hypoxic conditions with high fish density coupled with limited prey availability, anchovies sustained high swimming speeds at night. The sustained nighttime activity resulted in estimated daily energy expenditure over 20% greater than fish that adopted a diurnal activity pattern. We provide evidence that the sustained nighttime activity patterns are a result of foraging at night due to a lower ration achieved during the day. During severe hypoxic events, we also observed individual fish making brief forays into the hypoxic hypolimnion. These bottom waters generally contained higher prey (copepod) concentrations than the surface waters. The bay anchovy, a facultative particle forager, adopts a range of behaviors to compensate for the effects of increased conspecific density and reduced prey availability in the presence of stratification-induced hypoxia.     

Swimming ability of eels (<Emphasis Type="Italic">Anguilla rostrata,Conger oceanicus</Emphasis>) at estuarine ingress: contrasting patterns of cross-shelf transport?

The transport of eel early life stages may be critical to their population dynamics. This transport from ocean spawning to freshwater, estuarine and coastal nursery areas is a combination of physical and biological processes (including swimming behavior). In New Jersey, USA, the American eel (Anguilla rostrata) enters estuaries as glass eels (48.7–68.1 mm TL) in contrast to the Conger eel (Conger oceanicus) that enters as larger (metamorphosing) leptocephali (68.3–117.8 mm TL). To begin to understand the mechanisms of cross-shelf transport for these species, we measured the potential swimming capability (critical swimming speed, U _crit) under ambient conditions throughout the ingress season. A. rostrata glass eels were collected over many months (January–June) at a range of temperatures (4–21°C), with relative condition declining over the course of the ingress period as temperatures warmed. C. oceanicus occurred later in the season (April–June) and at warmer temperatures (14–24.5°C). Mean U _crit values for A. rostrata (11.7–13.3 cm s⁻¹) and C. oceanicus (14.7–18.6 cm s⁻¹) were comparable, but variable, with portions of the variability explained by water temperature, relative condition, ontogenetic stage, and fish length. Travel times to Little Egg Inlet, New Jersey, estimated using 50% U _crit values, indicate it would take A. rostrata ~30 and ~60 days to swim from the shelf edge and Gulf Stream, respectively. Travel times for C. oceanicus were shorter, ~20 days from the shelf edge, and ~45 days from the Gulf Stream. Despite differences in life stage, our results indicate both species are competent swimmers, and suggest they are capable of swimming from the Gulf Stream and/or edge of the continental shelf to estuarine inlets.     

Metabolism,enzymic activities and cold adaptation in Antarctic mesopelagic fishes

Several species of Antarctic mesopelagic fishes that have different minimal depths of occurrence but the same environmental temperature were collected in November–December 1983 and in March 1986 between 0 and 1 000 m in the open water near the marginal ice zone in the vicinity of 60°S 40°W (1983) and 65°S 46°W (1986), and oxygen consumption rate (V _{O 2}) and the activity of two metabolic enzymes, lactate dehydrogenase (LDH, an indicator of the anaerobic potential of locomotory muscle) and citrate synthase (CS, an indicator of citric acid cycle activity or aerobic potential), were determined. In four dominant species, whole-individual oxygen-consumption rate (y, ml O₂ individual^–1 h^–1) varied with weight (X, g) according to the equation y=aX ^b, with b values falling between 0.889 and 1.029. The relation of weight-specific LDH activity (y, U g^–1 wet wt) with weight (x, g) was also described by the equation y=aX ^b, with b values varying between 0.229 and 1.025. Weight-specific CS activity declined with weight, with b values from-0.031 to-0.369. V _{2 O}, LDH activity and CS activity all declined markedly with increased species' minimum depth of occurrence (the depth below which 90% of a species' population lives). Comparisons with previous studies on ecologically equivalent species of the California Borderland indicate that depth-related decreases in metabolism are the result of adapted traits of deeper-living species, not declining temperature within the water column. The metabolic rate of Antarctic mesopelagic fishes is approximately twice that of California species at equivalent temperatures; similar rates were found at the normal habitat temperatures of the two groups. Thus, a well-developed compensation for temperature is present in the Antarctic fishes: cold adaptation. Differences in enzymic activity among species, and among different sized individuals of a species are related to differences in metabolic rate and locomotory capacity. Enzymic indices can be used to estimate metabolic rates and evaluate ecological parameters such as predatory strategies and niche separation.     

How size relates to oxygen consumption,ammonia excretion,and ingestion rates in cold (<Emphasis Type="Italic">Enteroctopus megalocyathus</Emphasis>) and tropical (<Emphasis Type="Italic">Octopus maya</Emphasis>) octopus species

The scaling of metabolic rates with body mass is one of the best known and most studied characteristics of aquatic animals. Herein, we studied how size is related to oxygen consumption, ammonia excretion, and ingestion rates in tropical (Octopus maya) and cold-water (Enteroctopus megalocyathus) cephalopod species in an attempt to understand how size affects their metabolism. We also looked at how cephalopod metabolisms are modulated by temperature by constructing the relationship between metabolism and temperature for some benthic octopod species. Finally, we estimated the energy balance for O. maya and E. megalocyathus in order to validate the use of this information for aquaculture or fisheries management. In both species, oxygen consumption and ammonia excretion increased allometrically with increasing body weight (BW) expressed as Y = aBW ^b . For oxygen consumption, b was 0.71 and 0.69 for E. megalocyathus and O. maya, respectively, and for ammonia excretion it was 0.37 and 0.43. Both species had low O/N ratios, indicating an apparent dependence on protein energy. The mean ingestion rates for E. megalocyathus (3.1 ± 0.2% its BW day⁻¹) and O. maya (2.9 ± 0.5% its BW day⁻¹) indicate that voracity, which is characteristic of cephalopods, could be independent of species. The scope for growth (P = I − (H + U + R) estimated for E. megalocyathus was 28% higher than that observed in O. maya (320 vs. 249 kJ day⁻¹ kg⁻¹). Thus, cold-water cephalopod species could be more efficient than tropical species. The protein and respiratory metabolisms of O. maya, E. megalocyathus, and other octopod species are directly dependent on temperature. Our results offer complementary evidence that, as Clarke (2004) stated, the metabolic response (R and U) cannot be determined mechanistically by temperature, as previously proposed (Gillooly et al. 2002).     

ATP content and adenylate energy charge of the mussel Mytilus edulis during the annual reproductive cycle in Lindåspollene,Western Norway

The oxygen consumption rates ( VO₂) of 6 specimens (6 to 13 kg) of the albacore tuna Thunnus alalunga were measured at sea, using specimens collected 300 km west of San Diego, California (USA) during July and August, 1981. Fish were tested in a closed continuous-flow respirometer, where they swam at about 1.3 body lengths s^-1 velocity in 15° to 19°C water. The albacore tuna is a temperate pelagic species experiencing water temperatures from about 10° to 20°C and attaining a maximum weight of 45 kg. The VO₂ ranged from 1 249 to 3 336 ml h^-1 (the mean VO₂ for the 6 fish was 2 228 ml h^-1); such values approach those of mammals of a similar size and are 3 to 4 times those of most active fishes (e.g. sockeye salmon). Among fishes, the only higher VO₂ values yet recorded were for the skipjack tuna Katsuwonus pelamis, a tropical species. The remarkably high metabolic rates of tunas are presumably correlated with their continuous swimming activity and the maintenance of endothermy. The exponent relating VO₂ to body weight (1.18), although large, is not statistically different from the exponents for most other active vertebrates.     

Among-habitat variation in herbivory on Sargassum spp. on a mid-shelf reef in the northern Great Barrier Reef

Herbivory is widely acknowledged as a key process determining the benthic community structure and resilience of coral reefs. Despite numerous studies that have examined herbivory across reef gradients in the Caribbean, few studies have directly quantified this process on Pacific reefs. Bioassays of two species of erect macroalgae (Sargassum swartzii and S. cristaefolium) were used to quantify variation in grazing intensity across seven habitats of varying depth and wave exposure on a mid-shelf reef in the northern Great Barrier Reef. Removal rates of Sargassum varied significantly among habitats, with both species displaying broadly similar patterns. The shallow habitats on the exposed aspect of the reef (i.e. reef crest, flat and back reef) experienced the highest reductions in mass (81.4–91.6% day⁻¹) for both S. swartzii and S. cristaefolium, while the deeper exposed habitats (reef slope and base) displayed the lowest reductions (3.8–13.4% day⁻¹) over a 24 h period. In contrast, the grazing intensity varied between the two species in the three habitats on the leeward aspect of the reef. Reductions in mass remained relatively high for S. swartzii on the patch reef and sheltered reef base and flat (62.7–76.5% day⁻¹) but were considerably lower for S. cristaefolium (37.9–63.5% day⁻¹) across the same habitats. Surprisingly, the rates of removal of Sargassum displayed no relationship with the density or biomass of roving herbivorous fishes or those species known to consume erect macroalgae, either collectively or independently. These results suggest that the relationship between browsing rates and herbivorous fish biomass is complex and may be driven by species that are underestimated in visual surveys. Direct quantification of browsing intensity using assays revealed a different pattern to inferences based on herbivore densities and highlights the potential difficulties of evaluating ecosystem processes based on visual census data alone.     

Photosynthetic performance of giant clams, Tridacna maxima and T. squamosa, Red Sea

Two species of giant clams, Tridacna maxima and T. squamosa, coexist in the Red Sea, but exhibit distinctly different depth distributions: T. maxima mostly occurs in shallow waters (reef flat and edge), while T. squamosa may occur down to the lower fore-reef slope. Giant clams have been described as mixotrophic, capable of both filter-feeding and photosynthesis due to algal symbionts (zooxanthellae), therefore, observed depth preferences were investigated in relation to possible differences in autotrophy vs. heterotrophy. This study was conducted from April to June 2004, at the reef near the Marine Science Station, Aqaba, Gulf of Aqaba, Red Sea, and in May 2007, at a reef near Dahab, Sinai Peninsula, Egypt. In situ measurements using a submersible pulse amplitude modulated fluorometer (Diving PAM), revealed no significant differences in effective PSII quantum yield (ΔF/Fm′) and relative electron transport rates (ETR) between the two species; but rapid light curves (ETR vs. light, photosynthetically active irradiance, PAR) showed significant differences in maximum photosynthetic rates (ETR_max), with 20% higher values in T. maxima. Chamber incubations displayed higher net and gross oxygen production by T. maxima (88.0 and 120.3 μmol O₂ cm⁻² mantle area day⁻¹) than T. squamosa (56.7 and 84.8 μmol O₂ cm⁻² mantle area day⁻¹); even under shading conditions (simulated depth of 20 m) T. maxima still achieved 93% of the surface gross O₂ production, whereas T. squamosa reached only 44%. A correlation was found between ETR and net photosynthesis measured as oxygen production (T. maxima: R ² = 0.53; T. squamosa: R ² = 0.61). Calculated compensation depth (CD) (gross photosynthesis equals respiration) in T. maxima (16 m) matches the maximum depth of occurrence in this study (17 m). By contrast, the CD of T. squamosa (9 m) was much shallower than the maximum vertical range (42 m). Findings suggest T. maxima is a strict functional photoautotroph limited by light, whereas T. squamosa is a mixotroph whose photoautotrophic range is extended by heterotrophy. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Oxygen consumption of the semi-terrestrial crab <Emphasis Type="Italic">Pachygrapsus marmoratus</Emphasis> in relation to body mass and temperature: an information theory approach

Pachygrapsus marmoratus is a semi-terrestrial crab and the most common grapsid crab in the intertidal belt of rocky shores throughout the Mediterranean Sea, Black Sea and northeastern Atlantic. In this study, the combined effects of temperature (T), body mass (M), and sex (S) on the routine oxygen consumption rate (R) in P. marmoratus were quantified. The blotted wet body mass of the specimens ranged between 43 mg and 18.0 g, and five test temperatures were used between 13.5 and 28.0°C. Six candidate models that reflected different assumptions regarding the dependence of R on S and T were compared. Model selection was based on Kullback–Leibler’s information theory and Akaike’s information criterion (AIC). The model had the highest support by the data (E is the activation energy, B = 8.618 × 10⁻⁵ eV K⁻¹ is Boltzmann’s constant, T _a is the absolute temperature in Kelvin, and b the allometric scaling exponent); for P. marmoratus it was found that No sex dependence of R was supported by the data. Following a multi-model inference (MMI) approach, the mean (± SE) allometric exponent was 0.750 (± 0.013) having a 95% (bootstrap) confidence interval of 0.726–0.774. Thus, it was established that P. marmoratus follows Kleiber’s 3/4 law, as seems to be generally true for intertidal crabs. The allometric exponent was independent of temperature as has also been reported for many other marine invertebrates (at normal temperatures). Q ₁₀ values were relatively low, indicating wide thermal tolerance of the species. Model selection based on information theory is recommended for respiration studies, as an effective method in finding a parsimonious approximating model. MMI by model averaging, based on Akaike weights, is an effective way to make robust parameter estimations and deal with model selection uncertainty.     

Population dynamics,reproduction and growth of the Indo-Pacific horned sea star, <Emphasis Type="Italic">Protoreaster nodosus</Emphasis> (Echinodermata; Asteroidea)

The horned sea star (Protoreaster nodosus) is relatively common in the Indo-Pacific region, but there is little information about its biology. This study of the population biology of P. nodosus was carried out in Davao Gulf, The Philippines (7°5′N, 125°45′E) between September 2006 and May 2008. Protoreaster nodosus was found in sand and seagrass dominated habitats at a mean density of 29 specimens per 100 m² and a mean biomass of 7.4 kg per 100 m², whereas a significantly lower density and biomass was found in coral and rock dominated habitats. Adult specimens (mean radius R = 10.0 cm) were found at depths of 0–37 m, whereas juveniles (R < 8 cm) were only found in shallow sandy habitats with abundant seagrass (water depth ≤2 m). Increased gonad weights were found from March to May (spawning period), which coincided with an increasing water temperature and a decreasing salinity. Density and biomass did not change significantly during reproduction, but sea stars avoided intertidal habitats. All specimens with R > 8 cm had well developed gonads and their sex ratio was 1:1. Protoreaster nodosus grew relatively slowly in an enclosure as described by the exponential function G = 7.433 e^{−0.257 × R} . Maturing specimens (R = 6–8 cm) were estimated to have an age of 2–3 years. Specimens with a radius of 10 cm (population mean) were calculated to have an age of 5–6 years, while the maximum age (R = 14 cm) was estimated as 17 years. Potential effects of ornamental collection on the sea star populations are discussed. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

PIXGRO: A model for simulating the ecosystem CO2 exchange and growth of spring barley

A model, PIXGRO, developed by coupling a canopy flux sub-model (PROXEL_NEE; PROcess-based piXEL Net Ecosystem CO₂ Exchange) to a vegetation structure submodel (CGRO), for simulating both net ecosystem CO₂ exchange (NEE) and growth of spring barley is described. PIXGRO is an extension of the stand-level CO₂ and H₂O-flux model PROXEL_NEE, that simulates the NEE on a process basis, but goes further to include the dry matter production, partitioning, and crop development for spring barley. Dry matter partitioned to the leaf was converted to leaf area index (LAI) using relationships for the specific leaf area (SLA). The canopy flux component, PROXEL_NEE was calibrated using information from the literature on C3 plants and was tested using CO₂ flux data from an eddy-covariance (EC) method in Finland with long-term observations. The growth component (CGRO) was calibrated using data from the literature on spring barley as well as data from the Finland site. It was then validated against field data from two sites in Germany and partly via the use of MODIS remotely sensed LAI from the Finland site.Both the diurnal and the seasonal patterns of gross CO₂ uptake were very well simulated (R² = 0.92). A slight seasonal bias may be attributed to leaf ageing. Crop growth was also well simulated; simulated dry matter agreed with field observed data from Germany (R² = 0.90). For LAI, the agreement between the simulated and observed was good (R² = 0.80), giving an indication that functions describing the conversion of fixed CO₂ to dry matter and the subsequent partitioning leaf dry matter and LAI simulation were robust and provided reliable estimates.The MODIS LAI at a resolution of 1000 m agreed poorly (R² = 0.45) with the PIXGRO simulated LAI and the observed LAI at the Finland site in 2001. We attributed this to the coarse resolution of the image and/or the small size of the barley field (about 17 ha or 0.25 km²) at the Finland site. By deriving a regression relation between the observed LAI and NDVI from a higher resolution MODIS (500 m resolution), the MODIS-recalculated LAI agreed better with the PIXGRO-simulated LAI (R² = 0.86).PIXGRO provides a prototype model bridging the disciplines of plant physiology, crop modeling and remote sensing, for use in a spatial context in evaluating carbon balances and plant growth at stand level, landscape, regional, and with some care, continental scales. Since almost 50% of the European land surface is covered by crops, such a model is needed for the dynamic estimation of LAI and NEE of croplands.     

On the estimation of overwater buoyancy length from routine measurements

The stability parameter (z/L, where z is the height and L the buoyancy length) is an essential parameter in atmospheric boundary layer studies. From routine measurements, the bulk Richardson number (R_b) is usually computed. On the basis of appropriate field measurements at sea, it is shown that z/L = A R_b where A = 10.2 (with R ² = 0.97) for unstable and 6.3 (with R ² = 0.97) for stable conditions, respectively. It is also demonstrated that the proposed simplified equations are in excellent agreement with those more complicated formulations.     

Swimming ability and its rapid decrease at settlement in wrasse larvae (Teleostei: Labridae)

Wrasses are abundant reef fishes and the second most speciose marine fish family, yet little is known of their larval swimming abilities. In August 2010 at Moorea, Society Islands, we measured swimming ability (critical speed, Ucrit) of 80 settlement-stage larvae (11–17 mm) of 5 labrid species (Thalassoma quinquevittatum [n = 67], Novaculichthys taeniourus [n = 6], Coris aygula [n = 5], Halichoeres trimaculatus [n = 1] and H. hortulanus [n = 1]) and 33 new recruits of T. quinquevittatum. Median (mdn) larval Ucrit was 7.6–12.5 cm s⁻¹. In T. quinquevittatum (n = 67), larvae of 12.5–14.5 mm swam faster (mdn 16.9 cm s⁻¹) than smaller or larger larvae (mdn 3.9 and 3.2 cm s⁻¹, respectively). Labrid larvae Ucrit is similar to that of other similar-sized tropical larvae, so labrids and species with comparable settlement sizes should have similar abilities to influence dispersal. Ucrit of T. quinquevittatum recruits decreased to 47–56% of larval Ucrit in 2 days, implying rapid physiological changes at settlement.     

Heterotrophic nitrogen fixation (acetylene reduction) during leaf-litter decomposition of two mangrove species from South Florida, USA

Heterotrophic nitrogen-fixation (acetylene reduction) was measured during decomposition (under dark conditions) of Rhizophora mangle L. and Avicennia germinans (L.) Stearn leaf litter. Nitrogen-fixation rates in leaf litter increased following 24 d incubation, then decreased after ≃44 d for both species. Maximum rates of 66.2 and 64.6 nmol C₂H₄ g⁻¹ dry wt h⁻¹ were reached by R. mangle and A. germinans leaf litter, respectively. Higher fixation rates of leaf litter were associated with an increase in water content and sediment particles on leaf surfaces of both species. Rates of nitrogen fixation by diazotrophs attached to sediment particles were not significantly different from zero. With additions of d-glucose, ethylene production rates increased by factors of 625-, 34- and 7-fold for sediment, R. mangle and A.␣germinans leaf litter, respectively, compared to rates prior to enrichment. These organically enhanced rates of nitrogen fixation on leaves could be accounted for by increased activity associated with attached sediment particles and not the leaf material. Total phenolics [reported as tannic acid equivalent (TAE) units] decreased nitrogen-fixation rates when added to d-glucose-enriched sediment at >20 mg TAE l⁻¹. Phenolic compounds could explain the initial lag in rates of nitrogen fixation during leaf-litter decomposition of R. mangle (initial content of 110.8 mg TAE g⁻¹ dry wt), but not of A. germinans (initial content of 23.4 mg TAE g⁻¹ dry wt). The higher phenolic content and reportedly lower carbon substrate of R. mangle did not result in species-specific differences in either the magnitude or temporal pattern of nitrogen fixation compared to A. germinans leaf litter. We conclude that the availability of organic substrates leached from the leaf litter along with colonization by the heterotrophic diazotrophs (as indicated by sediment accumulation) controls nitrogen-fixation rates in a similar manner in the leaf litter of both species. Received: 8 August 1997 / Accepted: 4 December 1997