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.     

Habitat use by sponge-dwelling brittlestars

Cryptic organisms often associate with sessile invertebrates for refuge in space-limited environments. To examine interspecific habitat associations on coral reefs, tube- and vase-shaped sponges were surveyed for associated brittlestars at six sites on the coral reefs off Key Largo, Florida. Of 179 sponges encountered, Callyspongia vaginalis was the most abundant (43.0%), followed by Niphates digitalis (39.7%), and Callyspongia plicifera (4.5%). Three of eight sponge species surveyed did not differ from C. vaginalis in two physical refuge characteristics: oscular diameter and inner tube surface area. Brittlestars (416 total), all of the genus Ophiothrix, were only found in C. vaginalis, N. digitalis, and C. plicifera. The most abundant brittlestar, O. lineata (326), occurred on C. vaginalis (99.0%) and N. digitalis (1.0%), while O. suensonii (67) occurred on C. vaginalis (79.1%), N. digitalis (19.4%), and C. plicifera (1.5%). There was no pattern of co-occurrence of O. lineata and O. suensonii on C. vaginalis. The abundance of O. lineata increased with surface area of C. vaginalis. Differential habitat use was observed in O. lineata, with small individuals (<5 mm disk diameter) located inside and on the surface of sponge tubes and large individuals (5 mm) solely inside tubes. The number of large O. lineata in C. vaginalis never exceeded the number of tubes per sponge, and tagged O. lineata remained in the same sponge for at least 3 weeks. In density manipulations, no pattern of intraspecific competition among large O. lineata was observed; however, there was evidence for interaction between size-classes. Brittlestars selected live sponge habitat over a non-living refuge, suggesting a mechanism for sponge habitat recognition. Sponge-dwelling brittle stars prefer some tube- and vase-shaped sponge species despite similar oscular diameters and surface areas. Surprisingly, these preferred sponge species are known from previous studies to be chemically undefended against generalist fish predators; therefore, brittlestars that inhabit these sponges do not gain an associational chemical defense. Sponge habitat use by O. lineata may be governed by intraspecific interactions to maintain habitat and access to food. While past studies have suggested that O. lineata is an obligate sponge commensal, the present study suggests that O. lineata has a species-specific association with the tube-sponge C. vaginalis.Communicated by O. Kinne, Oldendorf/Luhe     

Highly repeated DNA sequences in the scleractinian coral genusAcropora: Evaluation of cloned repeats as taxonomic probes

A plasmid clone containing highly repeated DNA sequence fromAcropora formosa was used to probe slot blots of genomic DNA and Southern blots of Taq I digests from other Acroporidae. Homologous repeated DNA sequences were detected in 12 otherAcropora species, but not in three other species of Acroporidae (Astreopora sp.,Montipora digitata andM. aequituberculata). Slot-blot experiments indicated that the repeated sequences inAcropora latistella, A. tenuis, A. longicyathus andA. nobilis were distantly homologous with the clonedA. formosa repeat, whereas those inA. pulchra, A. millepora, A. valida, A. hyacinthus andA. microphthalma were highly homologous with this probe and those inA. digitifera were intermediate. Relatedness of the highhomology group toA. formosa was assessed by comparison of Taq I-digestion patterns. The predominant repeats inA. pulchra andA. hyacinthus had two Taq I sites per repeat unit (as didA. formosa), whereas repeats inA. digitifera, A. valida andA. microphthalma had one Taq I site per repeat; the pattern given byA. millepora implied that its highly homologous repeat units contained either one or two Taq I sites. Within the high-homology group, decreasing number of Taq I sites implies increasing taxonomic distance fromA. formosa. The relatedness series implied by these data differs from that based on morphological criteria.     

Energy balance of <Emphasis Type="Italic">Octopus maya</Emphasis> fed crab or an artificial diet

The present study was designed to evaluate the effect of a natural prey (the crab Callinectes sp.) and an artificial diet (pellet with squid paste and offered as a paste) on the survival and assimilation efficiency of subadult octopuses with 486 g of initial live weight. In order to reach this goal, the effects of the type of diet on energetic balance were assessed by recording ingestion rate (C), respiratory rate (R = R routine, R _rout + R apparent heat increment, R _AHI), ammonia production rate (U = U routine, U _rout + U post-prandial, U _PP) and biomass production (P) of Octopus maya during its growing process. Energy lost from faeces (H) was calculated as H=C−(U+R+P) and assimilated energy (As) as R + P. Octopuses fed an artificial diet had almost five times higher ingestion rate compared to that observed in octopuses fed crab. However, growth rate and production (P) were high in octopuses fed crab in comparison to octopuses fed artificial diet. An inverse relation between faeces (H) and type of food was observed, indicating that animals lost 77% of the ingested energy when fed artificial diet and only 5% when fed crab. A higher assimilation and production efficiency were obtained in octopuses fed crab (P/As: 61%) than in animals fed the artificial diet (P/As: −5%). The routine O : N ratio for animals in fasting was 9.1 and 2.3 for octopuses being fed crabs and the artificial diet, respectively. The post-alimentary O : N ratio was 3.6 and 2.2 for animals fed crabs and the artificial diet, respectively. This indicates that animals fed on both diets rely almost exclusively on protein. Based on energy balance data, a value of 472 kJ week⁻¹ kg⁻¹ of live octopus was estimated as the energy needed to obtain a growth rate near 9 g day⁻¹ (2.8% BW day⁻¹) for O. maya subadults. The total crab biomass needed to obtain 1 kg of fed O. maya biomass was calculated. A comparison with other different energy balance measurements made in other octopus species indicates that O. maya and Enteroctopus megalocyathus (Pérez et al. 2006) tend to be more efficient by channelling more ingested energy to biomass production (P = 69.5% of C) than O. vulgaris (P = 23% of C; Petza et al. 2006) or Paraledone charcoti (P = 4% of C; Daly and Peck 2000).     

Comparisons between the diets of four abundant species of elasmobranchs in a subtropical embayment: implications for resource partitioning

The diets of one ray species (Rhinobatus typus) and three shark species (Carcharhinus cautus, Negaprion acutidens, Rhizoprionodon acutus) undergo size-related changes and differ among these species in the nearshore waters of a large subtropical embayment (Shark Bay) in which these elasmobranchs are abundant, thereby reducing the potential for competition for food within and among these four species. R. typus fed almost exclusively on penaeid prawns and portunid crabs, which is reflected in its narrow dietary breadth, whereas different species of teleosts constituted a major component of the diets of each size class of the three shark species. The prey consumed by the three shark species was diverse, with representatives of 15 teleost families being consumed by C. cautus and substantial volumes of cephalopods being ingested by that species and R. acutus. The pronounced differences in the diet of the single ray species and three shark species reflect differences between a bottom-dwelling and more pelagic life, and between modes of feeding and relative mouth sizes. The relative contributions of the different species of teleost to the diets of the three shark species varied. Thus, although each of these species fed on atherinids, labrids and sillaginids, C. cautus also consumed substantial amounts of platycephalids and terapontids and R. acutus and N. acutidens also ingested considerable amounts of clupeids. Furthermore, R. acutus, which is the only one of the four species that typically occurs over seagrass, was the only species that fed on the centropomid Psammoperca waigensis, which is very abundant in seagrass meadows. However, the sparid Rhabdosargus sarba, which lives in unvegetated areas, was never ingested by R. acutus, but was consumed by C. cautus and N. acutidens. As the individuals of R. typus increased in size, they progressively consumed proportionately smaller volumes of the penaeid prawns Penaeus merguiensis and Melicertus latisulcatus and relatively greater volumes of the portunid crab Portunus pelagicus, which is slightly larger and has a harder exoskeleton. In addition to teleosts, large C. cautus ingested substantial volumes of portunid crabs and ophidian reptiles, presumably sea snakes, while large N. acutidens also fed on the ray R. typus.Communicated by G.F. Humphrey, Sydney     

Elemental composition (C,N, H) and energy in the development of Pagurus bernhardus (Decapoda: Pagurida) megalopa

No differences in development time and mortality were detected between starved and fed laboratory raised megalopa of Pagurus bernhardus. The average time of development in 138 megalopa was determined as 7.3±0.1 (95% CI) days. During megalopa development P. bernhardus loses about 7% in dry weight (DW), 17% in carbon (C), 6% in nitrogen (N) and 17% in hydrogen (H). During development C/N ratio and individual energy content descend about 14 and 22% respectively. Weight specific energy content decreases by 17% in the first 3 d and remains constant at 12.3±0.3 (95% CI) J·(mg DW)^-1 thereafter. About another 25% in individual energy content was lost by molting to crabs. The measured compounds do not follow a steady decrease. The possibility is discussed that a period of low energy cost (about the first half of development) alternates with times of higher energy expenditure mainly based on lipids. A fixed physiological program different from starvation capability is indicated for P. bernhardus megalopae. By comparing megalopae hatched in two different seasons and years reference is given to the variability in growth pattern.     

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     

Comparative electrophysiological analysis of plant odor perception in females of threePapilio species

Summary Antennae of femalePapilio butterflies perceive many volatile plant constituents with widely differing, constituent-specific sensitivities. We compared the responses of threePapilio species to volatiles from host and non-host plants to assess species-specificity and the degree of evolutionary conservatism in olfactory responses.Since previous studies had demonstrated that the polar constituents in odor fromDaucus carota stimulate oviposition behavior inPapilio polyxenes, we collected headspace volatiles fromD. carota, Pastinaca sativa (both Apiaceae) andArtemisia dracunculus (Asteraceae) and separated the polar fraction of these volatiles by gas chromatography. GC-coupled electroantennograms (GC-EAG) were recorded from the speciesPapilio polyxenes, P. machaon hippocrates andP. troilus. In addition, the responses of the three species to five compounds known as generally occurring constituents of plant odor were recorded. The relative sensitivities for these compounds were nearly identical in all threePapilio species. The response spectra to the separated plant volatiles also showed considerable similarities among the species.From the limited set of GC peaks evoking a response in one of the species, 64% (D. carota), 44% (P. sativa) and 29% (A. dracunculus) also evoked a response in both of the other species. The responses of the two closely related Apiaceae feeders (P. polyxenes, P. m. hippocrates) to volatiles fromD. carota were more similar to each other than was either to the response ofP. troilus, which feeds on Lauraceae. However, this was not true for the responses to volatiles fromP. sativa. The least congruence among the three species was found in the responses to volatiles fromA. dracunculus, a non-host for all of them. The differences and similarities found in the response profiles of the threePapilio species are discussed with respect to evolutionary adaptation to host odor versus evolutionary conservatism in adaptation of olfactory receptors.     

Larval development of Ocypode quadrata (Brachyura: Crustacea) under laboratory conditions

The larvae of Ocypode quadrata (Fabricius) have been reared in the laboratory, from hatching to megalopa stage, at 35 S, 25°C. The five zoeal stages and the megalopa are described, including functional appendages of each stage. On the basis of morphological characteristics, the first zoeal and megalopa stages of O. quadrata can be distinguished from similar stages of closely related Ocypodinae. At 25°C, the megalopa appeared in a minimum of 34 days following hatching.     

Ecological energetics of Nerita (Archaeogastropoda,Neritacea) populations on Barbados,West Indies

Population energy budgets estimated on the assumption of steady state conditions for Nerita tessellata Gmelin, N. versicolor Gmelin, and N. peloronta L. on Barbados, W. Indies, are presented. Large differences in population structure, and hence energetics, occurred at different localities along the beach. Relatively high proportions (81 to 88%) of the assimilated energy were lost via metabolism. Assimilation efficiencies ranged from 39 to 43%, net growth efficiencies from 5 to 13%, and ecological efficiencies from 3 to 7%. For each species, production (P), energy flow (A) and total energy consumption (C) were expressed as functions of animal size, in order to facilitate gross estimations of the energy components for other populations for which data on size-frequency and density are available. Respiration studies of all three species in the laboratory failed to detect differences between respiration rates in air or under seawater.     

Respiration rates of<Emphasis Type="Italic"> Beroe ovata</Emphasis> in the Black Sea

Metabolic rates of the ctenophore Beroe ovata within the length range from 0.4 mm (newly hatched larvae) to 60 mm were investigated. At 20° the respiration rates (Q, µg O₂ ind.^–1 h^–1) of individuals with wet weights (W, mg) less than or greater than 100 mg changed according to the equations Q=0.093W^0.62 and Q=0.016W^0.99, respectively. The weight-specific respiration rate of the juvenile ctenophores with wet body weights of 0.021–100 mg diminished approximately 20-fold (from 0.35 to 0.017 µg O₂ mg^–1 h^–1, respectively), but did not change within the range from 100 to 30,000 mg. The difference in the slope of the regression lines seems to be attributable to the ontogenetic changes in B. ovata metabolism. For the tested temperature range of 10–28°, the mean Q₁₀ coefficient was equal to 2.17±0.5. The basal metabolism of B. ovata narcotised by chloral hydrate was 4.5±0.9 times lower than total metabolism. Such a metabolic range is considered to be characteristic of aquatic invertebrates with high levels of locomotory activity.Communicated by O. Kinne, Oldendorf/Luhe     

Energy allocation in a reef coral under varying resource availability

An organism’s pattern of resource allocation to reproduction and growth over time critically impacts on its lifetime reproductive success. During times of low resource availability, there are two fundamental, mutually exclusive strategies of energy investment: maintenance of somatic tissues to support survival and later reproduction or investment into an immediate reproductive event at the risk of subsequent death. Here, we examine energy allocation patterns in the coral Montipora digitata to determine whether energy investment during periods of resource shortage favours growth or reproduction in a sessile, modular marine species. We manipulated light regimes (two levels of shading) on plots within a shallow reef flat habitat (Orpheus Island, Great Barrier Reef, Australia) and quantified energy uptake (rates of net photosynthesis), energy investment into reproduction (E _R ), tissue growth per unit surface area (E _T ) and energy channelled into calcification (E _C ). With declining resource availability (i.e. reduced photosynthesis), relative energy investment shifted from high (~80%) allocation to tissue growth (E _R :E _T :E _C  = 11:81:8%) to an increasing proportion channelled into reproduction and skeletal growth (20:31:49%). At the lowest light regime, calcification was maintained but reproduction was halted and thus energy content per unit surface area of tissue declined, although no mortality was observed. The changing hierarchy in energy allocation among life functions with increasing resource limitation found here for an autotrophic coral, culminating in cessation of reproduction when limitations are severe, stands in contrast to observations from annual plants. However, the strategy may be optimal for maximising fitness components (growth, reproduction and survival) through time in marine modular animals.     

Energetics of Euphausia pacifica. I. Effects of body carbon and nitrogen and temperature on measured and predicted production

Laboratory production during the life span of Euphausia pacifica was measured directly (as the sum of growth, molting and reproduction) and indirectly (as assimilation minus metabolism and leakage) to test the hypothesis that weight-specific production is a constant for all sizes. Euphausiids were collected in Puget Sound, Washington State, USA, from September 1973 to March 1978. Equations were determined (in terms of carbon and nitrogen at 8° and 12° C) expressing the relationships between body weight and the daily rates of growth, molting, reproduction, ingestion and metabolism. The allometric equation (R=aW ^b ) best related body weight (W) to the rate (R) for growth, molting, ingestion, respiration and excretion for life stages from late larvae through adults. As predicted by the original above hypothesis, the weight-specific coefficient (b) was close to 1.0 for ingestion and excretion; in contrast, b was 0.62 for growth, and 0.77 to 0.85 for molting and respiration. The Q₁₀ s also varied: 3.5 for growth, 2.4 for molting, about 3.0 for ingestion, and 2.0 for respiration and excretion. Assimilation efficiencies, for all weights and at both temperatures, were 81.3% of carbon and 85.9% of nitrogen ingested. The relationships between rate and body weight of early larvae for growth and molting were linear, as was the relationship for reproduction in adults. Weight-specific production was higher by I to 2% at 12° than 8° C for all life stages, and was 2 to 4% for carbon and 2 to 6% for nitrogen in adults, but 13 to 17% for carbon and 14 to 15% for nitrogen in early furcilia larvae. The null hypothesis was rejected for production measured directly, but would have been accepted if only an indirect measurement of nitrogen production had been considered. Clearly, indirect measurement incorporates all errors of measurement and assumption and makes interpretation difficult.     

Species-specific differences in lipid composition and omnivory indices in Arctic copepods collected in deep water during autumn (North Water Polynya)

To discern species-specific patterns in omnivory indices in copepods from the North Water Polynya, lipid classes and fatty acids were determined in Calanus hyperboreus Krøyer copepodite stage 5 (CV), C. glacialis Jaschnov (CV) and Metridia longa Lubbock (females) sampled below the mixed layer during autumn 1999. Generally, M. longa contained higher proportions of triacylglycerols, polar lipids and 18:1(n-9) than the other species. M. longa also had lower relative amounts of wax esters (WE), polyunsaturated fatty acids (PUFA), 20:1(n-9), 22:1(n-11), and lower absolute (g lipid copepod^-1) and relative (% lipid) lipid levels. Unsaturation coefficients (UC; ratio of polyunsaturated to total WE) were usually lowest in M. longa. These differences probably relate to a lesser dependence on phytoplankton in M. longa, and hence more ingestion of PUFA-poor prey. Moreover, levels of the carnivory index 18:1(n-9)/18:1(n-7) were highest in M. longa. Our data support the widespread contention that M. longa is more omnivorous than C. hyperboreus and C. glacialis. Proportions of bacterial fatty acids [odd-numbered and/or branched (OBFA); 18:1(n-7)] and PUFA/SFA (SFA: saturated fatty acids) ratios were largely unrelated to feeding strategy. Relationships between relative and absolute amounts of 16:1(n-7) and 18:1(n-7) in copepods suggest that the latter fatty acid is formed in vivo by chain elongation of the former. However, elevated levels of 18:1(n-7) and OBFA in M. longa at stations dominated by the microbial loop imply that these indices can be used to track ingestion of PUFA-poor protozoans when diatoms are scarce.Communicated by J.P. Grassle, New Brunswick     

Occurrence of tetrodotoxin in the gastropodsRapana rapiformis andR. venosa venosa

Paralytic toxicity was detected in four of 17 specimens and six of 28 specimens of the gastropodsRapana rapiformis andR. venosa venosa, respectively, collected from Chiching (Kaohsiung City) and Nanfangao (Ilan County), Taiwan, in 1988 and 1989. The highest toxicities, calculated as tetrodotoxin (TTX) content, were 140 and 13 mouse units (MU) g^–1 digestive gland inR. rapiformis andR. venosa venosa, respectively. The toxins obtained from each species were purified by ultrafiltration and Bio-Gel P-2 column chromatography. The toxin's specific toxicities (as TTX) were 56 MU mg^–1 (R. rapiformis) and 52 MU mg^–1 (R. venosa venosa). Results of analyses by thin-layer chromatography, cellulose acetate membrane electrophoresis and high-performance liquid chromatography showed that TTX and anhydrotetrodotoxin were responsible for the toxicity. The alkali hydrolysate of each toxin showed maximum absorption at ca. 274 nm, which is the unique absorption for the C₉-base of TTX and its related substances (such as anhydrotetrodotoxin, 4-epitetrodotoxin, etc.). Here, we report for the first time the occurrence of TTX in the gastropodsR. rapiformis andR. venosa venosa.     

Facultative lecithotrophy during larval development of the burrowing shrimp Callianassa tyrrhena (Decapoda: Callianassidae)

Survival, developmental and consumption rate (Artemia nauplii ingested per day) as well as predation efficiency (ingested per available Artemia nauplii) were studied during the larval development of the shallow-water burrowing thalassinid Callianassa tyrrhena (Petagna, 1792), which exhibits an abbreviated type of development with only two zoeal stages and a megalopa. The larvae, hatched from berried females from S. Euboikos Bay (Aegean Sea, Greece), were reared at 10 temperature–food density combinations (19 and 24 °C; 0, 2, 4, 8 and 16 Artemia nauplii d⁻¹). Enhanced starvation resistance was evident: 92 and 58% of starved zoeas I molted to zoea II, while metamorphosis to megalopa was achieved by 76 and 42% of the hatched zoeas at 19 and 24 °C, respectively. The duration of both zoeal stages was affected by temperature, food density and their interaction. Nevertheless, starvation showed different effects at the two temperatures: compared to the fed shrimp, the starved zoeae exhibited accelerated development at 19 °C (8.4 d) but delayed metamorphosis at 24 °C (5.9 d). On the other hand, both zoeal stages were able to consume food at an increased rate as food density and temperature increased. Predation efficiency also increased with temperature, but never exceeded 0.6. Facultative lecithotrophy, more pronounced during the first zoeal stage of C.tyrrhena, can be regarded as an adaptation of a species whose larvae can respond physiologically to the different temperature–food density combinations encountered in the wide geographical range of their natural habitat. Received: 28 February 1998 / Accepted: 21 October 1998     

Respiratory metabolism of Pagurus bernhardus (Decapoda: Paguridae) megalopa

Individual oxygen consumption was determined during the megalopa development of Pagurus bernhardus at 18°C in 1983. The maximum observed range was from 0.101 to 0.343 l O₂ h^-1 ind^-1. Individual respiration rate dropped by about 33% during the first two days and continued to remain at that level throughout the moult cycle. The results are discussed together with data on biomass and the fact that P. bernhardus megalopa do not feed.     

Respiration and nitrogen excretion of zooplankton. IV. The influence of starvation on the metabolism and the biochemical composition of some species

Changes in the respiration, ammonia excretion and biochemical composition were studied for three species of starving zooplankton (Calanus finmarchicus, Sagitta elegans, and Acartia clausi). Over the period of starvation, the respiration rate of all three species followed the same pattern of an initial decrease followed by a more or less constant level. A similar pattern was observed for the ammonia excretion rate of S. elegans and A. clausi, whereas C. finmarchicus excretion appeared to oscillate between high and low levels of protein catabolism. Study of the biochemical changes showed that C. finmarchicus consumed primarily lipids, and at times proteins, to meet its energy requirement whereas S. elegans and A. clausi primarily used protein. Variations in the elemental composition as well as the O:N ratio confirmed that C. finmarchicus alternated between periods of protein-dominant catabolism and lipid-dominant catabolism during starvation. No similar change in catabolism was observed in the two other species. The results are discussed in terms of physiological mechanisms of resistance to starvation and were used to calculate the energy budget of S. elegans and C. finmarchicus during the period of total starvation. The significance of such budgets is discussed and some of the sources of error examined.Bedford Institute of Oceanography Contribution.     

Gastric evacuation of different foods byNephrops norvegicus (Crustacea: Decapoda) and estimation of soft tissue ingested,maximum food intake and cannibalism in captivity

Gastric evacuation, maximum food intake and mortality during starvation were studied inNephrops norvegicus (L.) by means of simple laboratory experiments. Lobsters were collected by trawling off Barcelona (northwest Spain) at 400 m depth in winter 1986, and were held individually or in groups and fed different types of food:Nereis spp. (soft tissue, muscle), some species of Crangonidae (mostly hard tissue, carapace) andEngraulis encrassicolus (soft and hard tissues, combined muscles and bones). The digested matter was subsequently examined and analyzed from the stomach contents. Laboratory results were compared with the stomach contents of individuals collected from the field. Gastric evaluation was calculated as weight of stomach contents at Timet = stomach-content weight × 100:weight of food ration — residual weight of food, and the characteristic appearance of identifiable food items at different digestion times (0.5, 1, 2, 3, and 5 h) was recorded and illustrated, providing a basis for establishing the time of ingestion of the last meal and the underestimation of soft, compared to hard tissues. Underestimation of soft tissues may yield errors which constitute up to 10% of the soft food actually ingested. Maximum food intake was estimated at 0.25 g g^–1 body wt d^–1, and feeding did not recommence until 24 h after a previous meal. Finally, we present data on mortality resulting from cannibalism which occurred in conjunction with starvation: this ranged from 0% in isolated individuals to 36.6% in individuals held in groups.