Depth regulation of crab larvae in the absence of light

Behavioral responses to gravity and hydrostatic pressure have been investigated in two species of xanthid crabs Leptodius floridanus (Gibbes) and Panopeus herbstii Milne-Edwards to determine whether such responses provide a mechanism for depth regulation in the absence of light.In laboratory experiments, the four zoea stages and one megalopa stage of each species assume a differential vertical distribution in darkness, with succeeding stages showing a deeper overall distribution. Passive sinking rates increase in succeeding zoea stages and drop to an intermediate level after the molt to the megalopa stage. All zoea stages exhibit a negative geotaxis in the absence of light; the megalopa shows a positive geotaxis. The first zoea stage of Leptodius floridanus responds to an increase in hydrostatic pressure (up to 1 atmos above ambient) with an increase in swimming rate. This pressure response is shown to be reversible and not subject to short-term acclimation. The swimming rate of the last zoea stage does not increase in response to an increase in pressure.It is concluded that the responses of these larvae to gravity and hydrostatic pressure together with their characteristic passive sinking rates provide a mechanism for depth regulation in the absence of light that varies during ontogeny.     

Differential thermal tolerance and energetic trajectories during ontogeny in porcelain crabs,genus Petrolisthes

Thermal tolerance limits of marine intertidal zone organisms are elevated compared to subtidal species, but are typically just slightly higher than maximal habitat temperatures. The small thermal safety margins maintained by intertidal zone organisms suggest that high thermal tolerance is associated with a physiological cost. If true, we hypothesize that species that transition between intertidal zone and planktonic habitats during ontogeny, will adjust their thermal tolerance accordingly to capitalize upon potential energy savings while in a thermally benign habitat. We tested this hypothesis in porcelain crabs that transition between the thermally stressful, intertidal zone as embryos, to the thermally benign pelagic zone as larvae, and back at settlement. We found the more thermally tolerant, mid-intertidal zone species, Petrolisthes cinctipes, and the less thermally tolerant, subtidal zone species, Petrolisthes manimacilis, exhibited reduced thermal tolerance (LT₅₀) in the transition from embryos to larvae. This was associated with an increased oxygen consumption rate in both species, though P. cinctipes exhibited a significantly greater increase in oxygen consumption. P. cinctipes also showed an increase in thermal tolerance in settled juveniles compared to pelagic zoea I larvae, resulting in an overall V-shaped thermal tolerance relationship during ontogeny, while in P. manimaculis thermal tolerance was significantly lower in juveniles compared to zoea I. In neither species were these changes (zoea I to juvenile) associated with a significant change in metabolism. While embryos and juveniles of P. cinctipes have thermal tolerance limits near intertidal habitat thermal maxima (∼32.5 °C), all three life-history stages in P. manimaculis (especially embryos and larvae) exhibit considerable thermal safety margins. The mechanisms underlying this “excess” thermal tolerance in P. manimacilis embryos are unknown, but suggest that patterns of thermal tolerance in early life history stages are species-specific.     

Larval development of the subantarctic squat lobster Munida subrugosa (White, 1847) (Anomura: Galatheidae), reared in the laboratory

The larval development of the squat lobster Munida subrugosa from subantarctic waters of the Beagle Channel (Tierra del Fuego, Argentina) was studied under controlled laboratory conditions of temperature, salinity, and food supply. Developmental times, survival, and growth of larvae and early juveniles were investigated. Hatching of the entire brood always occurred during one night. Larvae were kept in 100ml individual bowls with filtered seawater at 8 ± 0.5°C and fed with Artemia spp. nauplii three times a week. Larvae passed through 6 zoeal instars and one megalopa. Previously, only five zoeal instars were known from this species. Mean cumulative durations of the zoeal stages I to VI were: 20.5 ± 2.5, 33.9 ± 4.1, 43.3 ± 5.4, 52.6 ± 5, 61.2 ± 3.9, and 83days, respectively. By adding the 28days that a single megalopa took to metamorphose to crab I stage, the complete larval development lasted 111days. Highest mortality occurred prior to the moult from the zoea I to zoea II stage (79.21% ± 18.65%) and during the moult from zoea VI to megalopa (92.86%). Carapace length was 1.64 ± 0.06, 1.52 ± 0.16, 1.57 ± 0.26, 1.64 ± 0.21, 2.11 ± 0.35, and 2.58 ± 0.19mm, for zoeal stages I to VI, respectively. Carapace length of megalopae and crab I instars was similar (2.85 ± 0.28 and the 2.84 ± 0.05, respectively). Unlike other subantarctic decapods, which show a tendency towards abbreviated larval development and/or some degree of endotrophy, M. subrugosa shows an extended planktotrophic larval development synchronized with short seasonal plankton production in austral summers.     

The significance of diet in the growth and development of larvae of the blue crab,Callinectes sapidus rathbun,under laboratory conditions

The following protistan diets were tested on blue crab larvae: the algae Isochrisis galbana Parke, Monochrisis lutheri Droop, Dunaliella sp., and an unknown mixture; and the ciliated protozoans Euplotes vannus Muller and Parauronema virginianum(2/1) Thompson. None of these diets resulted in development past the first zoea stage, although some apparently were ingested and delayed mortality as compared to unfed controls.The rotifer Brachionus plicatilis Müller sustained good survival through early zoea development; however, rotifer-fed larvae did not metamorphose to the megalopa. Larvae of the polychaete Hydroides dianthus (Verrill) sustained crab larvae throughout zoea development, resulting in 17% survival to metamorphosis. The percentage mortality per stage was significantly lower in polychaete-fed larvae when compared with rotifer-fed larvae during zoea stages III, VI, and VII. Mean intermolt duration varied between diet treatments during the first three stages, but showed no differences during later zoea development. In tests on groups of late stage sibling larvae, Artemia salina L. nauplii gave development to metamorphosis, whereas rotifers did not.All the diets so far tested on blue crab larvae are classified according to their ability to sustain development. It is demonstrated that the two diets which allow completed development, Hydroides dianthus larvae and Artemia salina nauplii, contain 2–3 times as much lipid per dry weight as do rotifers. A metabolic requirement for lipid late in development may be indicated. Invertebrate larvae derived from yolky telolecithal and centrolecithal eggs may be an important dietary component for brachyuran larvae.     

Moult cycle and morphogenesis inHyas araneus larvae (decapoda,majidae), reared in the laboratory

Moult cycle and morphogenesis in larval instars (zoea I, zoea II, megalopa) of the spider crabHyas araneus (L.) were studied in the laboratory. Changes in the epidermis and cuticle were documented photographically at daily intervals to characterize the stages of the moult cycle. Stage A (early postmoult) is a very short period during which the larva takes up water. During late postmoult (B) and intermoult (C) the endocuticle is secreted, and there is conspicuous epidermal tissue condensation and growth. The onset of early premoult (D₀) is characterized by epidermal apolysis, occurring first at the bases of the setae in the telson of zoeal instars or in the rostrum of the megalopa, respectively. Intermediate premoult (D₁) is the main period of morphogenesis, in particular of setogenesis: in the setae of the zoeal telson and carapace there is invagination or (in the zoea II) degeneration of epidermal tissues. Formation of new setae in the interior of epidermal tubules was observed in zoeal maxillipeds and in the antennae of the zoea II and megalopa instars. During late premoult (Stages D_2–4) part of the new cuticle is secreted, and the results of morphogenesis become clearly visible. For technical reasons (rigid exoskeleton) only a preliminary account of the moult cycle in the megalopa can be given. A time schedule is suggested for the stages of the moult cycle. It is estimated that postmoult (A–B) takes ca 9 to 15 % of total instar duration, intermoult (C) ca 22 to 37 %, and premoult (D) ca 48 to 69 %. There is an increasing trend of relative portions of time (% of total instar duration) from instar to instar in Stages A–C (mainly in the latter) and a decreasing trend in Stage D (mainly in D₀ and D_2–4).     

Neurogenesis in larval stages of the spider crab Hyas araneus (Decapoda,Brachyura): proliferation of neuroblasts in the ventral nerve cord

Larvae of the spider crab Hyas araneus were reared in the laboratory from hatching through to metamorphosis. Neurogenesis was recorded in the ventral nerve cord during development of successive larval stages, zoea 1, zoea 2, megalopa and crab 1. Proliferating cells were detected immunocytochemically after in vivo labelling with 5-bromo-2-deoxyuridine (BrdU) which, as a thymidine analogue, is incorporated into the DNA of dividing cells. Segmental sets of mitotically highly active neuroblasts were found in newly hatched larvae. A dorsal neuroblast, a ventral-median neuroblast, 3–6 anterior-ventral neuroblasts and 1–3 lateral neuroblasts could be distinguished in each thoracic ganglion. Significantly fewer neuroblasts were labelled in the suboesophageal ganglion as compared to the thoracic ganglia. The number of active labelled neuroblasts was high throughout zoea 1 and about 30% of zoea 2 development and then dramatically decreased towards premetamorphosis. In the newly moulted megalopa, only a reduced set of neuroblasts was labelled which ceased dividing within the first few days of megalopa development. There is an indication that, although most ganglion mother cells born by unequal division of neuroblasts may go through their final division at an early stage, certain clusters of ganglion mother cells obviously delay their final mitosis. These results are discussed with regard to neuronal integration which necessarily changes during the course of metamorphosis in relation to the altered behavioural repertoire of the larvae.     

Larval and early juvenile development of Lithodes santolla (Molina, 1782) (Decapoda: Anomura: Lithodidae) reared at different temperatures in the laboratory

The southern king crab, Lithodes santolla Molina, is distributed in cold-temperate and subantarctic waters ranging from the southeastern Pacific island of Chiloé (Chile) and the deep Atlantic waters off Uruguay, south to the Beagle Channel (Tierra del Fuego, Argentina/Chile). Recent investigations have shown that its complete larval development from hatching to metamorphosis, comprising three zoeal stages and a megalopa, is fully lecithotrophic, i.e. independent of food. In the present study, larvae were individually reared in the laboratory at seven constant temperatures ranging from 1 to 18 °C, and rates of survival and development through successive larval and early juvenile stages were monitored throughout a period of 1 year. The highest temperature (18 °C) caused complete mortality within 1 week; only a single individual moulted under this condition, 2 days after hatching, to the second zoeal stage, while all other larvae died later in the zoea I stage. At the coldest condition (1 °C), 71% of the larvae reached the zoea III stage, but none of these moulted successfully to a megalopa. A temperature of 3 °C allowed for some survival to the megalopa stage (17-33% in larvae obtained from two different females), but only a single individual passed successfully, 129 days after hatching, through metamorphosis to the first juvenile crab instar. At all other experimental conditions (6, 9, 12 and 15 °C), survival through metamorphosis varied among temperatures and two hatches from 29% to 90% without showing a consistent trend. The time of nonfeeding development from hatching to metamorphosis lasted, on average, from 19 days at 15 °C to 65 days at 6 °C. The relationship between the time of development through individual larval or juvenile stages (D) and temperature (T) was described as a power function (D=aT^b, or log[D]=log[a]blog[T]). The same model was also used to describe the temperature dependence of cumulative periods of development from hatching to later larval or juvenile stages. One year after hatching, the 7th (6 °C) to 9th (15 °C) crab instar was reached. Under natural temperature conditions in the region of origin of our material (Beagle Channel, Argentina), L. santolla should reach metamorphosis in October-December, i.e. ca. 2 months after hatching (taking place in winter and early spring). Within 1 year from hatching, the crabs should grow approximately to juvenile instars VII-VIII. Our results indicate that the early life-history stages of L. santolla tolerate moderate cold stress as well as planktonic food-limitation in winter, implying that this species is well adapted to subantarctic environments with low temperatures and a short seasonal plankton production.     

The systematic position of Ergasticus (Decapoda,Brachyura) and allied genera,a molecular and morphological approach

The complete larval series (zoea I, zoea II and megalopa) of the crab Ergasticus clouei is described and illustrated based on plankton samples from Mediterranean waters. The zoea II and megalopal stages, previously unknown, are described here for the first time. Nucleotide sequence analysis of two gene regions (16S rDNA and Cox1 genes) confirmed the assignment of these larvae to Ergasticus clouei. The molecular analyses and the morphology of the larval stages revealed large differences between Ergasticus and Inachus, the type genus of the family Inachidae. In fact, E. clouei larvae presented a series of morphological characters, such as antennal shape and mouthparts setation pattern that placed them closer to the family Oregoniidae. The phylogenetic analyses also showed significant support for the monophyly of the Oregoniidae + Ergasticus group. The data argue for removal of Ergasticus and the related genera (Bothromaia, Pleisticanthoides, Parapleisticantha and Pleistacantha) from the Inachidae and their placement within the Oregoniidae as a separate subfamily, Pleistacanthinae ?tev?i?, 2005. Our results demonstrate that larval stages provide reliable morphological traits, independent from those of adults, to help resolving relationships among Majoidea genera.     

The accumulation of age pigments during larval development of the spider crab,Hyas araneus (Decapoda,Majidae)

• 1.1. Lipofuscin, body carbon and respiration rates were measured in Hyas araneus from hatching to metamorphosis. Lipofuscin was measured spectrofluorometrically from the chloroform phase of chloroform/methanol extracts.
• 2.2. Excitation/emission spectra of both the chloroform and the methanol/aqueous phase showed one distinct fluorescence peak in the chloroform (410–415 nm emission/340–350 nm excitation) and the methanol/aqueous phase (405/350 nm) of zoea I (directly after hatching) and megalopa (0 and 24 days old).
• 3.3. Individual lipofuscin concentrations increased continuously during zoea I and halfway through zoea II, but remained constant through the entire megalopa despite high metabolic activity in this stage.
• 4.4. Individual lipofuscin concentrations were positively correlated with body carbon and carbonspecific lipofuscin was negatively correlated.
• 5.5. Moulting caused considerable loss of lipofuscin. During the first two larval ecdyses 17–18% were lost, with the shed moults containing only 3.4–4.5% of the lipofuscin found in late premoult individuals.
• 6.6. The different patterns of lipofuscin accumulation in respective larval stages is discussed in regard to mitotic activity of tissues. While in the zoea, growth is more related to lipid formation and biomass accumulation, in the megalopa morphogenetic processes require substantial epidermal growth, i.e. protein accumulation. However, the question why in the megalopa no increase in lipofuscin is found, remains unanswered.

     

Structure of the stomach cuticle in adult and larvae of the spider crab Maja brachydactyla (Brachyura,Decapoda)

The stomach of decapods is a complex organ with specialized structures that are delimited by a cuticle. The morphology and ontogeny of the stomach are largely described, but few studies have focused on the morphology of its cuticle. This study examined the morphology of the stomach cuticle of cardiac sacs, gastric mill ossicles, cardio-pyloric valve and pyloric filters, and during various stages (zoea I and II, megalopa, first juvenile, and adult) of the common spider crab Maja brachydactyla using dissection, histology and transmission electron microscopy. The results show that cuticle morphology varies among structures (e.g., cardiac sacs, urocardiac ossicle, cardio-pyloric valve, pyloric filters), within a single structure (e.g., different sides of the urocardiac ossicle) and among different life stages. The cuticle during the larval stages is very thin and the different layers (epicuticle, exocuticle, and endocuticle) are infrequently distinguishable by histology. Major changes during larval development regarding cuticle morphology are observed after the molt to megalopa, including the increment in thickness in the gastric mill ossicles and cardio-pyloric valve, and the disappearance of the long thickened setae of the cardio-pyloric valve. The cuticle of all the stomach structures in the adults is thicker than in larval and juvenile stages. The cuticle varies in thickness, differential staining affinity and morphology of the cuticle layers. The structure–function relationship of the cuticle morphology is discussed.     

Publications of Anthony Calhoun Clement (1909–1984)

Summary

Two zoeae and the megalopa of the majid crab Macropodia parva were obtained from laboratory material. At 25±1°C and 35% salinity, the first crab appeared 12 days after hatching; survival to the first crab instar was 27%. The larval stages are described in detail and compared with those of the previously described species of the genus Macropodia. Zoeal characteristics of M. parva that differentiate it from other known Macropodia larvae are the naked telson furcae and the absence, in zoea II, of the exopodal setae on the basis of the maxillule.     

Seasonal variation in the occurrence of planktonic larvae of sympatric hermit crabs

Patterns of seasonal change in the abundances of the planktonic larvae of four sympatric hermit crabs were examined at five fixed stations over 18 months in Christmas Bay, Texas. Clibanarius vittatus (Bosc) was present in the plankton from April through October, Pagurus longicarpus Say from September through May and P. pollicaris Say from December through February. Pagurus annulipes (Stimpson) zoeae were the most frequently encountered zoeae and were found throughout the year, but were most abundant in late spring and summer. Zoea I of P. annulipes and P. pollicaris were found to be reliably distinguishable on the basis of their carapace lengths. Clibanarius zoea I were very abundant (3149) and 42 megalopa but no zoea II–V were found. Inadequate sampling and the short durations of these missing stages do not seem sufficient to explain their absence.     

Occurrence of an alternative pathway in the larval development of the crab Chasmagnathus granulata Dana, 1851 under laboratory conditions

Larvae of the crab Chasmagnathus granulata were collected in a salt marsh located in the Lagoa dos Patos, Brazil and reared from eclosion to metamorphosis under different dietary regimes. Larvae reared individually in beakers of 40 ml and fed Tetraselmis chuii (zoea III and zoea IV), showed a supplementary stage, here designated as zoea V, with morphological characteristics intermediary between zoea IV and megalopa. No zoeae V molted to megalopa stage. To confirm the occurrence of the supplementary stage, mass cultures of larvae of C. granulata were fed Artemia sp. at high densities, we again detected the fifth zoeal instar. However, when zoeae V were individually placed in beakers and fed Artemia nauplii, they succeeded in molting into megalopae. We observed the occurrence of two types of zoeae IV — a smaller type (from which originated the zoeae V) and a larger type (which directly developed into megalopae). We conclude that stressful nutritional/environmental conditions were responsible for the occurrence of this alternative path of development.     

Effects of various salinities on Na(+)-K(+)-ATPase, Hsp70 and Hsp90 expression profiles in juvenile mitten crabs, Eriocheir sinensis

Eriocheir sinensis is a euryhaline crab migrating from sea to freshwater habitats during the juvenile stage. We used quantitative real-time polymerase chain reaction (qRT-PCR) to investigate the gene expression profile of Na(+)-K(+)-ATPase, Hsp70 (heat shock protein 70) and Hsp90 in megalopa exposed to salinities of 0, 2, 5, 10, and 15 parts per thousand. Both low and high salinities markedly stimulated expression of Na(+)-K(+)-ATPase, Hsp70 and Hsp90 genes of Chinese mitten crab megalopa; salinity had different effects on Na(+)-K(+)-ATPase, Hsp70 and Hsp90 levels depending on the duration of salinity stress, implying that Na(+)-K(+)-ATPase, Hsp70 and Hsp90 may play an important role in salinity tolerance in this crab species.     

Foregut morphology and ontogeny of the spider crab Maja brachydactyla (Brachyura,Majoidea, Majidae)

We describe the morphology of the foregut of the spider crab Maja brachydactyla Balss, 1922, from first larval stage to adult, with detailed stage‐specific documentation using light and scanning electron microscopy. A total of 40 ossicles have been identified in the foregut of adults of M. brachydactyla using Alizarin‐Red staining. The morphological pattern of the ossicles and gastric mill is very similar to other Majoidea species with only a few variations. The foregut of the zoeae stages appeared as a small and simple cavity, with a cardio‐pyloric valve that separates the stomach into cardiac and pyloric regions. The pyloric filter is present from the first zoea, in contrast to the brachyuran species which have an extended larval development. Calcified structures have been identified in the cardio‐pyloric valve and pyloric region of the zoeal stages. The most significant changes in foregut morphology take place after the metamorphosis from ZII to megalopa, including the occurrence of the gastric mill. In the megalopa stage, the foregut ossicles are recognizable by their organization and general morphology, but are different from the adult phase in shape and number. Moreover, the gastric teeth show important differences: the cusps of the lateral teeth are sharp (no molariform); the dorsal tooth have a small, dentate cusp (not a well‐developed quadrangular cusp); and the accessory teeth are composed of one sharp peak (instead of four sharp peaks). The gastric mill ontogeny from megalopa to adult reveals intermediate morphologies during the earlier juvenile stages. The relationship between gastric mill structures with food preferences and their contribution to the brachyuran phylogeny are briefly discussed. J. Morphol. 276:1109–1122, 2015. © 2015 Wiley Periodicals, Inc.     

Morphological features of the larvae of spider crab <Emphasis Type="Italic">Pugettia quadridens</Emphasis> (Decapoda: Majidae) from the Northwestern Sea of Japan

The entire cycle of larval development of the spider crab Pugettia quadridens (de Haan, 1850) (Decapoda: Majidae), widespread in Peter the Great Bay (Sea of Japan) is studied under the laboratory conditions. The development cycle of this species comprises prezoea, zoea I, zoea II, and megalopa. At a temperature of 18–20° C larval development took from 11 to 15 days. Zoea II is described in detail for the first time. Many morphological characters are found distinguishing zoea and megalopa of P. Quadridens in Russian waters from the larvae of this species in Japanese and Korean waters. Some characters of larvae are similar in P. Quadridens and the related species of the genus Pugettia. The larvae of P. Quadridens occur in the plankton of Vostok Bay from late June to late October with a density up to 5 ind/m³ at a surface water temperature of 13–21°C. They are easily distinguished from the other brachyuran larvae of this region by the absence of lateral spines on the carapace.Original Russian Text Copyright © 2004 by Biologiya Morya, Kornienko, Korn.     

Adaptive diversity in congeneric coastal crabs: Ontogenetic patterns of osmoregulation match life-history strategies in Armases spp (Decapoda, Sesarmidae)

Ontogeny of osmoregulation and salinity tolerance were investigated throughout the larval development of two congeneric species of sesarmid crab, Armases ricordi (H. Milne Edwards) and A. roberti (H. Milne Edwards), and compared with previous observations from two further congeners, A. miersii (Rathbun) and A. angustipes (Dana). In the semiterrestrial coastal species A. ricordi, the zoeal stages were only at moderately reduced salinities (17-25.5‰) capable of hyper-osmoregulation, being osmoconformers at higher concentrations. The megalopa was the first ontogenetic stage of this species, which exhibited significant hyper-osmoregulation at further reduced salinities (≥ 5‰), as well as a moderately developed function of hypo-regulation at high concentrations (32-44‰). The riverine species A. roberti showed similar overall patterns in the ontogeny of osmoregulation, however, also some striking differences. In particular, its first zoeal stage showed already at hatching a strong capability of hyper-osmoregulation in salinities down to 5‰. Interestingly, this early expressed function became significantly weaker in the subsequent zoeal stages, where survival and capabilities of hyper-osmoregulation were observed only at salinities down to 10‰. The function of hyper-regulation in strongly dilute media re-appeared later, in the megalopa stage, which tolerated even an exposure to freshwater (0.2‰). Differential species- and stage-specific patterns of osmoregulation were compared with contrasting life styles, reproductive behaviours, and life-history strategies. In A. ricordi, the larvae are released into coastal marine waters, where salinities are high, and thus, no strong hyper-osmoregulation is needed throughout the zoeal phase. The megalopa stage of this species, by contrast, may invade brackish mangrove habitats, where osmoregulatory capabilities are required. Strong hyper-osmoregulation occurring in both the initial and final larval stages (but not in the intermediate zoeal stages) of A. roberti correspond to patterns of ontogenetic migration in this species, including hatching in freshwater, larval downstream transport, later zoeal development in estuarine waters, and final re-immigration of megalopae and juvenile crabs into limnic habitats, where the conspecific adults live. Similar developmental changes in the ecology and physiology of early life-history stages seem to occur also in A. angustipes. A. miersii differs from all other species, showing an early expression and a gradual subsequent increase of the function of hyper-osmoregulation. This ontogenetic pattern corresponds with an unusual reproductive biology of this species, which breeds in supratidal (i.e. land-locked) rock pools, where variations in salinity are high and unpredictable. Matching patterns in the ontogeny of osmoregulation and life-history strategies indicate a crucial adaptive role of osmoregulation for invasions of (by origin marine) crabs into brackish, limnic and terrestrial environments.     

Physiological and biochemical changes during the larval development of a brachyuran crab reared under constant conditions in the laboratory

Larvae of the spider crabHyas araneus were reared in the laboratory at constant conditions (12°C; 32‰S), and their feeding rate (F), oxygen consumption (R), nitrogen excretion (U), and growth were measured in regular intervals of time during development from hatching to metamorphosis. Growth was measured as dry weight (W), carbon (C), nitrogen (N), hydrogen (H) protein, and lipid. All these physiological and biochemical traits revealed significant changes both from instar to instar and during individual larval moult cycles. AverageF was low in the zoea I, reached a maximum in the zoea II, and decreased again in the megalopa. In the zoeal instars, it showed a bell-shaped pattern, with a maximum in the middle (zoea I) or during the first half of the moult cycle (zoea II). MaximumF in the megalopa was observed still earlier, during postmoult. Respiration (R) increased in the zoeal instars as a linear function of time, whereas it showed a sinusoidal pattern in the megalopa. These findings on variation inF andR during larval development confirm results obtained in previous studies onH. araneus and other decapod species. Excretion (U) was measured for the first time with a high temporal resolution in crab larvae. It showed in all three larval instars a bell-shaped variation pattern, with a maximum near the middle of the moult cycle, and significantly increasing average values from instar to instar. The atomicO/N ratio followed an inverse pattern, suggesting a maximum utilization of protein as a metabolic substrate during intermoult. Growth data from the present study and from a number of previous studies were compiled, showing consistency of growth patterns, but a considerable degree of variability between larvae from different hatches reared under identical conditions. The data show the following consistent tendencies: during the first part of each larval moult cycle (in postmoult, partly in intermoult), lipids are accumulated at a higher rate than protein, whereas an inverse growth patterns is typical of the later (premoult) stages. These two different growth phases are interpreted as periods dominated by reserve accumulation in the hepatopancreas, and epidermal growth and reconstruction (morphogenesis), respectively. Differences between individual larval instars in average biochemical composition and growth patterns may be related to different strategies: the zoeal instars and the early megalopa are pelagic feeding stages, accumulating energy reserves (principally lipids) necessary for the completion of larval development, whereas the later (premoult) megalopa is a semibenthic settling stage that converts a significant part of this energy to epidermal protein. The megalopa shifts in behaviour and energy partitioning from intense feeding activity and body growth to habitat selection and morphogenesis, preparing itself for metamorphosis, i.e. it shows an increasing degree of lecithotrophy. Data from numerous parallel elemental and biochemical analyses are compiled to show quantitative relationships betweenW, C, N, H, lipid, and protein. These regressions may be used as empirical conversion equations for estimates of single chemical components in larvalHyas araneus, and, possibly, other decapods.