Hatching rhythms in Uca pugilator (Decapoda:Brachyura)

Gravid female Uca pugilator (Bosc) maintained in conditions of 14h L: 10h D light regime and constant temperature without simulated tides showed a significant positive correlation between time of hatch and time of the nighttime high tide. Light: Dark experiments showed no entrainment by LD cues. Larval hatching was delayed until dark when high tide occurred between 1900 and 2130 hrs. Hatch time was a positive linear function of the time of the nighttime high tide when high tide occurred after 2130 hrs. Time of hatch was delayed by the light period on days lacking a nighttime high tide. Experiments conducted under constant light (LL) and temperature, without simulated tides, indicated a persistent free-running rhythm of approximately 25 h 03 min, suggesting an endogenous timer. Rhythmic larval release on the nighttime high tide may be adaptive as a predator-avoidance mechanism for adult gravid females.Contribution No. 366 from the Belle W. Baruch Institute for Marine Biology and Coastal Research, University of South Carolina     

Multivariate analysis of the mud crab Scylla serrata (Brachyura: Portunidae) from four locations in Southeast Asia

The large, edible mud crab Scylla serrata (Forskål) exhibits different phenotypes which are recognised by the fishermen of Southeast Asia and are given local names such as “white”, “green”, “red” or “black” crabs on the basis of their colouration. A preliminary study using multivariate techniques was undertaken in order to examine the degree of dissimilarity between the different morphs of commercially fished stocks of S. serrata. Twenty-two morphometric and 20 meristic characters were measured on male crabs from four locations in three countries in Southeast Asia, including Surat Thani in Thailand where “black” and “white” morphs exist sympatrically. Canonical variate analysis (CVA) on the morphometric data discriminated three groups with no chain-linking. Surat Thani “white” crabs and those from south Vietnam were phenotypically similar and formed one distinct group; a second group contained crabs from Ranong (Thailand) and Sarawak which were phenotypically “black”; the third group contained “black” crabs from Surat Thani. CVA on meristic data confirmed two groupings, but implied that the Surat Thani “black” morph may only be a variant of the Surat Thani/Vietnam “white” form, rather than a third species. These findings are discussed in relation to the ecology, fishery management and aquaculture potenial of S. serrata.     

Effects of temperature and salinity on larval development ofNecora puber (Brachyura: Portunidae)

Temperature and salinity affected both length of larval development and mortality inNecora puber collected in the Ría de A Coruña during December 1984 and January 1985. Development time decreased considerably with increased temperature. This decrease was sharper when temperature increased from 15° to 20°C than when it increased from 20° to 25°C. At 35S, average development took 48, 32 and 28 d at 15°, 20° and 25°C, respectively. At the three salinities tested (25, 30 and 35), larval development was completed only at 15°C, at 20°C/30 and 35S, and at 25°C/35S. Development times at 15° and 20°C were highly significantly different at both 35 and 30S (P 0.01). However, there were no significant differences between development times at 20° and 25°C (P > 0.05). Within any one specific temperature series, no significant difference was observed between the salinity values tested (P > 0.05). The duration of each of the five zoeal stages was similar within each and the same temperature/salinity combination, whereas the duration of the megalop was twice as long as any of the zoeal stages. The combination of the lowest temperature (15°C) and the highest salinity (35) tested resulted in the greatest larval survival of 28%. Highest mortality occurred at 25°C, at which temperature development was completed only at 35S. A sharp drop in larval survival was observed in the transition period Zoea V — megalop in all combinations of temperature and salinity tested. Within the limits of tolerance to temperature and salinity, the former effected more pronounced differences in the duration of larval development, while salinity appeared to constitute a limiting factor for survival.     

Geostatistical analysis of densities of Liocarcinus depurator (Brachyura: Portunidae) on the western Mediterranean from 1994 to 2003

Bottom trawl estimates of densities of the portunid crab Liocarcinus depurator along the Mediterranean coast of Spain from 1994 to 2003, covering a depth range of 50–800 m, were analysed using non-linear geostatistical techniques. The variation of the spatial distribution was modelled through variogram analysis using depth as a covariate. Patch size ranged from 20 to 75 km. Crab density showed a general decrease from 1994 to 2003. L. depurator high-density areas were characterised as either occasional or permanent (always present every year). High-density patches appeared to be associated with either upwelling or strong river influence, i.e. the Western Alborán Sea and the Ebro Delta. The application of non-linear geostatistical techniques proved useful in identifying permanent high-density population patches, thereby highlighting regions of special interest for the ecology of the species.     

Feeding and mouthpart morphology in the semaphore crabHeloecius cordiformis (Decapoda: Brachyura: Ocypodidae)

The mouthparts of the Australian semaphore crabHeloecius cordiformis (H. Milne-Edwards, 1837) are described. Male and female semaphore crabs were observed and collected from mangrove swamps around the south side of Botany Bay, Australia between 1983 and 1987. Semaphore crabs inhabit mangrove mud/sand flats between the midlittoral intertidal zone and the supralittoral fringe of bordering saltflats. Scanning electron micrographs provide detailed pictures of mouthpart morphology and the setae present on the third, second, and first maxillipeds; the second and first maxillae; and the mandibles. Amongst other setal types, spoon-tipped, corn, and bottle-brush setae are described. While semaphore crabs are primarily deposit feeders (microphagous), they have well developed mandibles and are, therefore, also able to eat larger plant and animal material (macrophagous). A brief comparison is made between the mouthparts of semaphore crabs and fiddler crabs (Uca spp.), since the two genera are closely related, occupy similar niches, and resemble each other in appearance. This comparison reveals that while the second maxilliped setal morphology differs between the two genera (Heloecius possesses corn setae,Uca possesses spoontipped setae) these setae may have similar functions. The feeding behaviour ofH. cordiformis is related to substrate composition, burrow construction and various mouthpart specialisations. It is conlcuded that, within the context of the present study, certain structural e.g. mouthpart modifications allowing both scavenging and deposit feeding, and behavioural modifications e.g. mound building in wet muddy habitats, enable semaphore crabs to occupy a wide range of habitat types within the mangrove.Please address all correspondence and requests for reprints to Dr Maitland at his present address at the University of the Witwatersrand     

Population dynamics and growth of juveniles of the velvet swimming crab Necora puber (Decapoda: Portunidae)

Recruitment variability plays a critical role in determining local population densities of benthic organisms, but extreme vulnerability at the onset of juvenile life is a trait that is largely responsible for population survivorship trends. The aim of the present study was to determine the role of juvenile recruitment in the population structure of Necora puber. Juveniles of N. puber were collected from the lower intertidal of rocky shores of Plymouth Sound (southwest coast of the UK) and monthly size–frequency distribution were used to determine the dynamics and the growth of the population. The parameters of the von Bertalanffy growth function were estimated (K=0.281 year⁻¹; t ₀=0.043; C=0.103; and t _s=0.268) assuming a L _∞=105 mm. Growth was markedly seasonal and present results indicated a slower juvenile growth rate than described previously for N. puber. The recruitment period was extensive and was two times higher in 2001 than in 2000 at the start of the 1+ year, but levelled off at the end of the 1+ year class on three of the four shores studied. Instantaneous mortality as high as 5.1 year⁻¹(99.4% year⁻¹) was observed during the higher recruitment year. Early juvenile mortality appears to be density dependent and a demographic bottleneck appears to limit the number of juveniles on some shores.     

Measuring the effects of thermoclines on the vertical migration of larvae of Callinectes sapidus (Brachyura: Portunidae) in the laboratory

An experimental system is described that is capable of maintaining precise and stable thermoclines of specified magnitude for laboratory study of behavior of planktonic organisms. Behavioral responses to encounters with thermoclines of various magnitudes were measured in stage I larvae of the portunid crab Callinectes sapidus Rathbun. Larvae adapted to an initial temperature of 15°C were able to swim up through a temperature gradient of 10°C, but showed increasing inhibition of upward migration when t=12.5°C. Larvae adapted to 17.5°C were unable to penetrate t=10°C. Thermoclines of 5° or 10°C slowed passive sinking rates. Results suggest that larvae avoid an upper temperature of 27.5°C by a cessation of swimming, accompanied by passive sinking. Thermal gradients of the magnitude present in the natural habitat are unlikely to alter larval vertical migration that is regulated by other exogenous stimuli. Naturally occurring thermoclines should seldom influence dispersal characteristics of C. sapidus larvae.Contribution No. 1535 of the Center for Environmental and Estuarine Studies, University of Maryland     

Ultrastructure of the spermatozoa of Hymenosomatidae (Crustacea: Brachyura) and the relationships of the family

The spermatozoa of the genus Odiomaris Ng and Richer de Forges, 1996 (=Amarinus Lucas, 1980) have the components typical of eubrachyuran (Heterotremata + Thoracotremata) sperm, but differ significantly from all other investigated eubrachyurans in at least ten characteristics: (1) presence of an epiopercular dome; (2) separation of all but the central region of the operculum from the remainder of the acrosome by an infra-opercular rim; (3) the fact that the acrosome is smaller in volume than the nucleus; (4) the acrosome is strongly emergent from the nucleus, being surrounded only basally by nuclear material; (5) the cytoplasmic sheath, ending anteriorly with the nucleus, is also basal; (6) division of the acrosome contents into an inner and outer acrosome zone is scarcely apparent in longitudinal section as the inner zone is narrow and of doubtful homology; (7) the thin, putative inner acrosome zone is anteriorly almost septate owing to several longitudinal corrugations; (8) basally there is a unique “fringe zone”; (9) the acrosome, including the epinuclear dome, is longer than wide; (10) the unique helical and posterolateral disposition of the nuclear arms. From a purely spermatological viewpoint, Odiomaris (as exemplified by O. pilosus and O. estuarius), and provisionally the Hymenosomatidae, are thus excluded from the Thora- cotremata, in which they were formerly placed, nor are they readily placeable in the Heterotremata. Received: 30 December 1996 / Accepted: 4 February 1997     

Larval development of the spider crab Pisoides edwardsi (Decapoda,Brachyura) under laboratory conditions

Larvae of Pisoides edwardsi (Bell, 1835) have been reared in the laboratory at 2 different temperatures (13.8° and 18.5°C), from hatching to megalops stage. The two zoeal stages and the megalops, as well as the setation of the functional appendages are described and illustrated. The main characteristics useful to differentiate the larvae of P. edwardsi from those of Libidoclaea granaria, the other Chilean species belonging to the same sub-family, are discussed. Data on duration of zoeal development, length of moulting intervals, and mortality at the 2 test temperatures, are also given.This study was financially supported, in part, by the Chilean Ministry of Agriculture.     

The annual reproductive cycles of Uca annulipes,Portunus pelagicus and Metapenaeus affinis (Decapoda: Crustacea) from the South-west coast of India

Employing the gonad index method, the reproductive cycles of three decapod crustaceans, Uca annulipes (Latreille), Portunus pelagicus (Linnaeus) and Metapenaeus affinis (Milne-Edwards) have been studied. In these crustaceans breeding is not continuous all the year round, but extends over several months of the year with distinct peak periods of gonadal activity. The male and female reproductive cycles are not concurrent. The peak of the reproductive cycle of males occurs slightly earlier in the breeding season than that of females. These studies indicate the possibility of production of successive broods of eggs during the same breeding season. In these species, the low saline conditions of the monsoon period are unfavourable for breeding. The medium and high saline conditions during the post-monsoon and pre-monsoon months, respectively, with plenty of planktonic food for the larvae, seem to be the favourable periods for breeding activity.     

Podotreme affinities of Raninoides sp. and Lyreidus brevifrons: evidence from spermatozoal ultrastructure (Crustacea: Brachyura: Raninoidea)

Spermatozoal synapomorphies which singly or collectively distinguish Raninoidea are: (1) the presence of single (Ranina, Raninoides) or multiple (Lyreidus) keel-like projections of the acrosomal capsule; (2) a very large, weakly electron-opaque peripheral acrosomal zone (Ranina, Raninoides) and an homologous large outer zone in Lyreidus; (3) poor differentiation of the operculum from the capsule (autapomorphy); (4) a very well developed, perforate subopercular zone, of variable form; (5) presence of unique inward longitudinal projections (septa or corrugations) in the wall of the subacrosomal chamber (autapomorphy). Shared, presumably synapomorphic characters of Ranina and Raninoides but not of Lyreidus within the Raninidae, are: (1) branching of some of the subacrosomal septa (unbranched in Lyreidus); (2) the subspheroidal form of the acrosome in Ranina with a length: width ratio (L:W) of 0.76, and, although slightly more depressed, in Raninoides (L:W 0.73), considered apomorphic relative to the more depressed form in Lyreidus (L:W 0.52); (3) single or multiple coiled perforatorial filaments (Ranina, Raninoides) contrasting with a capitate perforatorium with “amoeboid” head in Lyreidus; (4) division of the capsule wall to give one posterior (Ranina) or multiple enclaves, plesiomorphically (?) absent in Lyreidus. Similarities of Lyreidus with other podotremes include the capitate perforatorium, questionably related to the radiate spiked-wheel structure of homolids in which acrosome proportions are similar or less closely to the bilateral capitate perforatorium of dromiids and dynomenids, and basal capsular projections as in the dromiid Stimdromia (=Petamolera) lateralis and in cyclodorippoids. No spermatozoal synapomorphies support a sister-group relationship of raninoids and heterotreme-thoracotreme crabs.     

Elemental composition (C,N,H) in larval and crab-1 stages of Pagurus bernhardus (Decapoda,Paguridae) and Carcinus maenas (Decapoda,Portunidae)

Analyses of individual content of carbon (C), nitrogen (N), and hydrogen (H) were carried out for all larval stages of Pagurus bernhardus and Carcinus maenas, and for newly metamorphosed crabs. Maximum range in total larval development is 12.8 to 165.8 g C, 3.2 to 35.1 g N, and 1.9 to 24.9 g H in P. bernhardus and 3.1 to 43.2 g C, 0.7 to 10.1 g N, and 0.4 to 6.3 g H in C. maenas. From these data energy equivalents were calculated. Maximum range in total larval life is 0.43 to 6.38 J ind. ^-1 in P. bernhardus and 0.1 to 1.49 J ind. ^-1 in C. maenas. There is a 32.4% mean loss of energy in P. bernhardus megalopa development; this seems to describe the normal developmental pattern in this stage. Biomass was determined as fresh and dry weight respectively. Individual dry weight is about 3.6 to 5.6 times higher in P. bernhardus (44 to 340 g) than in C. maenas (12 to 93 g) larvae.Contribution to research project Experimentelle marine Ökosystemanalyse sponsored by Bundesministerium für Forschung und Technologie, Bonn (Grant No. MFU-0328/1)     

Widespread occurrence of extensive epimural rod bacteria in the hindguts of marine Thalassinidae and Brachyura (Crustacea: Decapoda)

Direct observation by scanning electron microscopy indicates that the presence of bacteria (epimural and unattached) in the hindguts of Crustacea is widespread, occurring across taxa (two infraorders, i.e., Brachyura and Thalassinidae, nine genera, 16 species), feeding types (detritivores, scavengers and carnivores), habitats (mangroves, saltmarshes, sand/mudflats) and continents (North America, South Africa, Australia). Crustacean hindguts clearly represent suitable environments for colonization by micro-organisms, despite the lack of specialized structures or modifications of the gut to facilitate this. Mats of closely-packed epimural rods and scattered epimural rods were the most common types of bacteria observed in the guts of the Crustacea examined, although unattached rods and scattered epimural cocci occurred in some species. There were, however, taxon-dependent differences in colonization characteristics of hindgut bacteria, possibly related to differences in gut lining. Abundance of hindgut microflora was unrelated to the host's taxon, habitat or geographical locality, but appeared to be affected by the feeding habits of the animal. Mats of epimural rods were associated exclusively with detritivores, while cocci were only observed in the hindguts of scavengers and carnivores. Moreover, extensive colonization by epimural rod bacteria (covering >50% of the hindgut lining) was observed in detritivores only, while carnivores harboured few or no rod bacteria. The detritivore hindgut appears to provide a better environment for microbial habitation than does that of carnivorous crustaceans. In all cases the rod bacteria were monocultures of morphologically identical bacteria and were remarkably similar among crustacean species. The potential significance of prolific microbial colonization in the hindguts of crustaceans deserves consideration.Communicated by O. Kinne, Oldendorf/Luhe     

Carapace and branchial water circulation,and water-related behaviours in the semaphore crabHeloecius cordiformis (Decapoda: Brachyura: Ocypodidae)

Water uptake, branchial water circulation and various water-related behaviours and anatomical specialisations, are described for the Australian semaphore crabHeloecius cordiformis (H. Milne-Edwards 1837). Male and female semaphore crabs were observed and collected from mangrove swamps around the south side of Botany Bay, Australia between 1983 and 1987. Semaphore crabs are active at low tide in air. Their branchial chambers are modified for both air breathing and water circulation. Water is held within the branchial chambers and is used for feeding and, to a certain extent, respiration. Crabs carry between 0.35 and 0.5 ml of water (7 g live body wt). Branchial water is continuously pumped out by the scaphognathites over specialised setae-bearing regions of the carapace and back through the branchial chambers over the gills, in a manner similar to that found in many grapsid crabs. Setae associated with water circulation are described. Water falling ventrally is collected by the abdominosternal cavity (formed between the abdomen and sternum) from where it returns to the branchial chambers via branchiosternal canals. Several novel water-related behaviours are described: Water shoogling — branchial water is drawn out between the walking legs where it is worked by scissor-like up and down movements of the legs. Abdomen flap — branchial water volume is adjusted by the extension and flexion of the abdomen against the cephalothorax. Water dumping — by tilting forwards and kissing the substrate, branchial water is voided to allow an entirely fresh load to be taken up. Statuesque posing — crabs remain erect and motionless for long periods. Branchial water is absent and external body surfaces dry out. Posing behaviour may function in thermoregulation, osmoregulation or as an anti-fouling mechanism against settling organisms. The significance of these various behaviours is discussed.     

Laboratory-reared larvae of Cancer anthonyi (Decapoda: Brachyura) with a brief description of the internal anatomy of the megalopa

The larval development of Cancer anthonyi Rathbun (Decapoda, Brachyura) is described from laboratory rearing experiments. The external anatomy of the various larval stages is illustrated. A prezoeal stage, 5 zoeal stages and 1 megalopa stage were identified. At 17.5°C it took an average of 32.5 days for the first-stage zoeae to develop through the fifth zoeal stage and molt to the megalopa stage. The general internal anatomy of C. anthonyi larvae is discussed, and a drawing of a parasagittal section of a megalopa-stage larva is included.     

Rhythms in larval release of the sublittoral crab Neopanope sayi and the supralittoral crab Sesarma cinereum (Decapoda: Brachyura)

Ovigerous females of the subtidal xanthid crab Neopanope sayi (Smith) and the high intertidal grapsid crab Sesarma cinereum (Bosc) were collected during the summers of 1986 and 1987 in the Beaufort, North Carolina (USA), area and brought into the laboratory, where rhythms in larval release were monitored. When crabs with late-stage embryos were put under a 14 h light:10 h dark cycle in an otherwise constant-environment room, an apparent tidal rhythm in release of larvae was observed for both species, with N. sayi releasing near the time of day and night high tides, and S. cinereum releasing around the time of night high tides. The time of sunset relative to high tide was a complicating factor, since larval release for both species was often concentrated around sunset when evening high tides fell several hours before sunset. When a group of N. sayi and S. cinereum were brought into the laboratory and placed under constant lowlevel light for 5 d, the release rhythm of the population persisted, thus implying that the rhythm is endogenous. Larval release near the time of high tide and often at night is common among brachyurans living in tidal areas, regardless of specific adult habitat, suggesting a common functional advantage. Possibilities include transport of larvae from areas where predation and the likelihood of stranding and exposure to low-salinity waters are high, as well as a reduced probability of predation on adult females. Results of the present study suggest that the importance of release after darkness may increase with increasing tidal height of the adult.     

Population genetics of the swimming crab<Emphasis Type="Italic"> Callinectes bellicosus</Emphasis> (Brachyura: Portunidae) from the eastern Pacific Ocean

The population genetics and historical demography of the swimming crab Callinectes bellicosus from the eastern Pacific were assessed using mitochondrial DNA (mtDNA) sequences from portions of the cytochrome c oxidase subunit I (COI) and cytochrome b (Cytb) genes. Analysis of molecular variance of sequence data from crabs collected from nine localities, ranging from the upper to lower Gulf of California and the outer coast of the Baja California peninsula, revealed an absence of population structure, suggesting a high level of gene flow over a wide geographic area. Maximum-likelihood estimates of long-term effective population size obtained with the program FLUCTUATE, in addition to highly significant values obtained from neutrality tests (Tajimas D, Fu and Lis D, and Fus F_S) and mismatch analysis, are consistent with a population expansion dating to the Pleistocene epoch. Phylogenetic analysis of C. bellicosus sequences using both neighbor-joining and Bayesian methods revealed a widely distributed subclade (clade II) cryptically embedded at low frequency in the main (clade I) population. Although sequence divergence between the two clades was low (1.1% COI; 0.6% Cytb), statistical support for the split was high. The Kimura-2-parameter genetic distance between C. bellicosus and the sympatric and morphologically similar C. arcuatus was high (d=0.17) and similar to the genetic distance between C. bellicosus and the allopatric C. sapidus from the western Atlantic (d=0.18), suggesting an ancient (Miocene) divergence of C. bellicosus and C. arcuatus.Communicated by P.W. Sammarco, Chauvin     

Feeding ecology of Liocarcinus depurator (Decapoda: Portunidae) in the Ria de Arousa (Galicia,north-west Spain): effects of habitat,season and life history

The diet of the portunid crab Liocarcinus depurator (L.) in soft-bottom areas of the Ría de Arousa (Galicia, north-west Spain) was studied by analysis of stomach contents, and by comparison of habitats of contrasting abundance, composition and diversity of prey communities. Monthly samples were taken from July 1989 to June 1990 (N=3747) in mussel (Mytilus galloprovincialis) raft-culture areas and central channel zones. Ninety-two food items were identified: crustaceans, molluscs, polychaetes, ophiuroids and fishes were the dominant prey. The raft epifauna and the mussels constituted the dominant group of prey in the raft areas and inner channel station (the epifaunal crab Pisidia longicornis comprised 30 to 52% of the diet), whereas the benthic macroinfauna was the dominant food component in the mid-outer channel (polychaetes constituted 32% of the diet). In the epifaunal community, diet selection for P. longicornis was positive and for amphipods was negative, whereas within the infauna diet selection for bivalves and ophiuroids was positive and for polychaetes negative or neutral. The seasonal importance of P. longicornis and M. galloprovincialis varied in the diet of L. depurator parallel to seasonal activities connected with mussel culture (seeding, thinning, harvesting) that determine the availability of prey from the rafts. Major changes in the composition of the diet related to the life history of L. depurator occurred during ontogeny. In the raft zones, consumption of mussels and fishes increased with increasing body size, while the consumption of Natantia, non-decapod crustaceans, gastropods, polychaetes Pectinariidae and plants decreased with increasing crab size. Maximum consumption of P. longicornis was by L. depurator individuals of intermediate size (15 to 44 mm carapace width). Stomach fullness (measured as dry weight of food) increased with increasing body size, but with negative allometry. During the late pre-moult and early post-moult periods L. depurator did not consume any food; maximum stomach fullness was recorded for the late post-moult and intermoult stages. Female food consumption decreased during egg incubation. Food consumption was greater in autumn and winter than at other seasons; this appears to be related to seasonal changes in reproduction and moulting.     

Distribution,habitat preferences and breeding periods in the crustaceans Sesarma cinereum and S. reticulatum (Brachyura: Decapoda: Grapsidae)

The distribution and habitat preferences of Sesarma cinereum and S. reticulatum were studied in the region of Beaufort, North Carolina, USA. S. cinereum inhabits the supralittoral zones of marshes characterized by high salinities (mean=27.9%) and sandy substrates, while s. reticulatum were found to prefer eulittoral marshes with brackish salinities (mean=16.2%) and silty substrates. Major differences have also been found in the cycling of egg production between these two species. Populations of S. cinereum produce approximately 4 to 6 egg batches in close synchrony with the lunar phase, whereas populations of S. reticulatum produce only from 2 to 3 batches of young per breeding season. It appears that female S. cinereum invest less time in carrying their eggs (approximately 1 lunar month), while female S. reticulartum invest a greater amount of time (eggs carried for approximately 45 days). These and other specific differences in reproductive strategies suggest that populations of S. cinereum are responding to density-independent selective pressures, while populations of S. reticulatum exhibit characteristics of density-dependent selective effects.     

Influence of limited starvation periods on growth and elemental composition (C,N,H) of Carcinus maenas (Decapoda: Portunidae) larvae reared in the laboratory

Zoea-1 larvae of Carcinus maenas L. (Decapoda: Brachyura: Portunidae) were from Helgoland in March 1984 and reared in the laboratory at 18°C through ecdysis. Dry weight (DW) and elemental composition of carbon (C), nitrogen (N), and hydrogen (H) were analyzed in newly hatched zoea-1, after different initial starvation periods, and in newly moulted zoea-2. Continually starved zoea-1 lost biomass and energy steadily, and logarithmic functions show best fit of empirical and predicted data. Biomass and energy equivalents of newly moulted zoea-2 are significantly correlated with starvation periods in the zoea-1, showing lower values with longer initial starvation. After about 25 to 34% individual biomass and energy losses, larvae exceed the point-of-noreturn (PNR), and do not recover or moult to the zoea-2, even if re-fed. When starvation ceases before the PNR, larvae moult to the zoea-2, and develop with lower average growth rates (AGR) after prolonged periods of initial food deprivation. The later larvae were re-fed, the less absolute amounts of DW, C, H, and individual energy, but more DW-related energy equivalents and N accumulated during subsequent feeding towards ecdysis. It is suggested that lipid, rather than protein, is the main source of energy controlling the maintenance of larval moult cycles. After lipid reserves are depleted, zoea-1 larvae live on body protein, and lose the ability to absorb and restore sufficient lipid if re-fed later than the PNR.Contribution to research project An-145/1-1 granted by the Deutsche Forschungsgemeinschaft (DFG)