Effects of temperature on oxygen consumption,growth, and development of embryos and yolk-sac larvae of Siganus randalli (Pisces: Siganidae)

Eggs from laboratory spawnings of the coralreef fish Siganus randalli Woodland were incubated at two temperatures (27 and 30 °C). Eggs and larvae were sampled until larval starvation, while changes in oxygen consumption, growth, yolk utilization, and development were monitored. Oxygen consumption, which peaked at hatching, was higher for embryos incubated at 30 °C than at 27 °C. Rates of oxygen consumption (nl h^-1 individual^-1) at hatching were similar to those for other temperate and tropical species. Rates of oxygen consumption by yolk-sac larvae were highly variable, and these data suggest that larval oxygen consumption prior to yolk-sac absorption may not be significantly influenced by temperature. Rates of yolk depletion were higher for larvae at the higher temperature. After an initial rapid increase in length, length of larvae at 30 °C decreased with age. Egg size, egg weight, and maximum notochord length of larvae differed significantly between spawns. Age-specific oxygen consumption rates by the embryos varied between spawns, but regressions describing oxygen consumption as a function of age did not differ significantly. The initiation and completion of eye pigmentation were used as developmental markers to calculate the amount of yolk remaining for larvae at the different temperatures. Larvae maintained at 30 °C completed eye pigmentation approximately 3 h sooner than those maintained at 27 °C, but had less endogenous reserves. This finding indicates a trade-off between rapid development and efficient utilization of the endogenous reserves. The completion of eye pigmentation in larvae incubated at the higher temperature occurred at midnight and, depending on the amount of time that the larvae have to initiate feeding prior to the point-of-no-return, the timing of completion of eye pigmentation could influence larval survival.     

The influence of temperature on the development of Baltic Sea sprat (Sprattus sprattus) eggs and yolk sac larvae

In spring 2004 and 2005 we performed two sets of experiments with Baltic sprat (Sprattus sprattus balticus Schneider) eggs and larvae from the Bornholm Basin simulating ten different temperature scenarios. The goal of the present study was to analyse and parameterise temperature effects on the duration of developmental stages, on the timing of important ontogenetic transitions, growth during the yolk sac phase as well as on the survival success of eggs and early larval stages. Egg development and hatching showed exponential temperature dependence. No hatching was observed above 14.7°C and hatching success was significantly reduced below 3.4°C. Time to eye pigmentation, as a proxy for mouth gape opening, decreased with increasing temperatures from 17 days post hatch at 3.4°C to 7 days at 13°C whereas the larval yolk sac phase was shortened from 20 to 10 days at 3.8 and 10°C respectively. Maximum survival duration of non-fed larvae was 25 days at 6.8°C. Comparing the experimental results of Baltic sprat with existing information on sprat from the English Channel and North Sea differences were detected in egg development rate, thermal adaptation and in yolk sac depletion rate (YSDR). Sprat eggs from the English Channel showed significantly faster development and the potential to develop at temperatures higher than 14.7°C. North Sea sprat larvae were found to have a lower YSDR compared to larvae from the Baltic Sea. In light of the predictions for global warming, Baltic sprat stocks could experience improved conditions for egg development and survival.     

Heat waves,baby booms,and the destruction of kelp beds by sea urchins

Large populations of sea urchins, Strongylocentrotus droebachiensis (Müller), destroyed kelp beds along the Atlantic coast of Nova Scotia in the 1960's and 1970's. The origin of these large sea urchin populations is not understood. We have investigated the potential influence of variable growth and development of the planktonic larvae of sea urchins (in response to temperature and food abundance) on recruitment of benthic juveniles. The adult sea urchins were collected at Sandy Cove, Digby County, Nova Scotia, Canada, in December 1986. Temperature strongly affected larval size and the growth of the echinus rudiment within the range 3° to 9°C, and larvae grew most rapidly at 14°C. Food abundance had a smaller effect on larval growth, and these effects were apparent only at high temperature. Larvae fed the same concentration of two different algal food species grew and developed similarly. Correspondence between spring temperature variation and qualitative variation in sea urchin recruitment, as well as strong temperature effects on larval growth in culture, and the occurrence of a large, positive temperature anomaly in June 1960, all suggest that temperature effects on larval growth and development may have led to intense sea urchin recruitment in 1960 and the appearance of large adult populations 4 to 6 yr later. This result invites further research.     

Laboratory study of predation by Aurelia aurita on larvae of cod,flounder, plaice and herring: development and vulnerability to capture

Capture success of the medusa Aurelia aurita preying on various developmental stages of fish larvae was measured together with larval reactivity and escape speed after being stung. These experiments were conducted in the spring of 1983 with A. aurita medusae collected from Loch Etive, Scotland and laboratory-reared larvae of Gadus morhua L., Platichthys flesus L., Pleuronectes platessa L. and Clupea harengus L. Capture success of the medusae increased with medusa size, but decreased with advancing larval development. Smaller species of larvae were more vulnerable to capture. Larval reactivity to encounters with medusae increased with advancing development, and larger species of larvae were more reactive to encounters. Larval escape swimming speeds also increased with advancing larval development and size. These results indicate that earlier stages of larvae within a species and smaller species of larvae at a given stage are more vulnerable to predation by medusae since they are less reactive to encounters. Apparently they are more susceptible to the effects of neurotoxins. Predation rates on different developmental stages of herring larvae are documented and compared with rates predicted by a predation model. Predictions fell within the range of observed predation rates, but tended to overestimate rates by larger medusae feeding on larger herring larvae. This indicates the possibility of predator satiation and/or behavioural avoidance.     

Yolk utilization and growth to yolk-sac absorption in summer flounder (Paralichthys dentatus) larvae at constant and cyclic temperatures

Rates of development, growth and yolk conversion efficiency were determined in larvae of the summer flounder Paralichtys dentatus at constant temperatures of 21°, 16°, 12° and 5°C and in temperature cycles of 21°–16°, 16°–11°, and 11°–5°C. In constant incubation temperatures, development rate increased with increasing temperature. Larvae reared in the cyclic temperature regimes exhibited development rates intermediate to those at the temperature extremes of the cycle. All larvae reared at 5°C and in the 11°–5°C cycle regime died prior to total yolk-sac absorption. Although development rates were temperature dependent, no significant differences in notochord length ash-free dry weight or yolk utilization efficiency were found at the time of total yolk-sac absorption. The similarity in growth and yolk utilization efficiency for larvae reared under these various temperature regimes suggests that the physiological mechanisms involved are able to compensate for temperature changes encountered in nature.Contribution No. 195 from EPA, Environmental Research Laboratory, Narragansett, Rhode Island 02882, USA     

Effect of temperature on viability and axial muscle development in embryos and yolk sac larvae of the Northeast Arctic cod (Gadus morhua)

Cod (Gadus morhua L.) eggs may develop and hatch within temperatures of −1.5 to 12 °C, but little is known about the effects of very low temperatures on larval characteristics. Eggs of the Northeast Arctic cod (Gadus morhua) were incubated at 1, 5 or 8 °C from Day 1 after fertilisation until hatching, and transferred to 5 °C after hatching. Histological samples of the axial musculature were taken at hatching and 5 d after hatching, and the data on muscle cellularity from these samples were related to survival and hatching, size, developmental data and viability of the yolk sac larvae. All larvae hatched at the same developmental stage. Incubation of eggs at 1 °C produced shorter larvae with a larger yolk sac and more, small deep fibres at hatching than larvae from eggs incubated at 5 or 8 °C. The larval size difference was still present 5 d after hatching, a time at which the larvae from 1 °C-incubated eggs were less developed and less resistant to an acute viability stress test (65 ppt salinity). Although there were no differences between temperature groups in number and size of muscle fibres 5 d after hatching, the deep fibres of the 1 °C-group contained less myofibrils than the two other groups. The phenotype of the larvae at hatching was thus affected within these incubation temperatures. Although all groups were transferred to the same temperature after hatching, the lowest egg incubation temperature (1 °C) still had a negative effect 5 d after hatching, as these larvae were both smaller, less resistant to stress and had less functional muscles at the time of first feeding. Our conclusion is therefore that 1 °C is close to, or below, the lower thermal tolerance limit for normal functional development of Northeast Arctic cod. The results are discussed in relation to larval viability and recruitment of this species in the wild. Received: 4 February 1998 / Accepted: 10 July 1998     

Rearing<Emphasis Type="Italic"> Pandalus borealis</Emphasis> (Krøyer) larvae in the laboratory

Northern shrimp Pandalus borealis (Krøyer) larvae hatch in the northern Gulf of St. Lawrence from early May to the end of June, and larval development occurs over a range of relatively cold water temperatures. Because of the long duration of the pelagic phase and the difficulty of sampling all successive larval stages at sea, we used laboratory experiments to assess the effects of water temperature on larval development and growth. In spring 2000, P. borealis larvae were reared from hatching to the first juvenile stages (i.e., stage VI and VII) at three temperatures (3, 5, and 8°C) representing conditions similar to those in spring in the northern Gulf of St. Lawrence. Larval development and growth were dependent on temperature, with longer duration and smaller size (cephalothorax length, CL, and dry mass, DM) at 3°C relative to the 5 and 8°C treatments. There were no significant differences in the morphological characters of the different stages among treatments, indicating that regular moults occurred at each temperature. The results suggest a negative impact of cold temperatures (lower intra-moult growth rates and smaller size) and, possibly, higher cumulative mortality due to longer development time that could affect the success of cohorts at sea. However, CL and DM for stage III and later larvae were smaller than those of larvae identified at the same developmental stage in field locations. It is possible that the diet offered to larvae in this experiment (Artemia nauplii, either newly hatched nauplii or live adults, depending on the developmental stage) was not optimal for growth, even though it is known to support successful P. borealis larval development. In the field, there is the possibility that phytoplankton contributes to the larval diet during the first stages and stimulates development of the digestive glands. Furthermore, the nutritional quality of the natural plankton diet (e.g., high protein content, fatty acid composition) might be superior and favourable to higher growth rates even at lower temperatures.Communicated by R.J. Thompson, St. Johns     

Morphological and functional development of larval and juvenile Limanda yokohamae (Pisces: Pleuronectidae) reared in the laboratory

Morphological and behavioural aspects in larval development need to be studied in detail to understand the early life history better, and to gain a comprehensive knowledge on early life stages for fish species important in aquaculture and fisheries. In the present study, larvae of Limanda yokohamae (Günther) were reared to observe their behavioural development, and to obtain specimens for studying the morphological features and the intestinal development at Ohno, Hiroshima, Japan, in 1987. Swimming activity was monitored at several larval stages, and swimming speed was recorded until settlement and after-feeding behaviour was initiated. A slight increment of swimming speed was observed with larval growth. Larvae changed their swimming behaviour from surface waters to the bottom of the rearing tank when their eyes began to move. Morphological development of pigmentation patterns, fin development, squamation and the development of the digestive tract were described and illustrated in detail to characterize development stages, especially those relating to metamorphosis. During metamorphosis, growth ceased and rapid changes in allometric growth were accompanied by differentiation of the digestive tract. After metamorphosis there was steady growth, allometric growth achieved a constant value, and both the scales and digestive organs were fully formed. Metamorphosis was therefore a crucial developmental milestone, including a critical phase during which survival potential was lowered.     

Feeding behaviour of North Sea turbot (Scophthalmus maximus) and Dover sole (Solea solea) larvae elicited by chemical stimuli

The effect of chemical stimuli on the feeding behaviour of turbot (Scophthalmus maximus L.) and sole (Solea solea L.) larvae was investigated under controlled laboratory conditions. Laboratory-reared flatfish larvae exposed to chemical stimuli showed significant differences in the frequency of various behaviour patterns related to feeding (such as swimming, snapping and darting) compared to larvae exposed to blanks of filtrated seawater. The chemical substances Il-asparagine, glycine, inosine 5-monophosphate and betaine evoked the strongest behavioural responses in turbot larvae. In sole larvae the most potent substances were Il-phenylalanine, Il-lysine, Il-asparagine, inosine 5-monophosphate and betaine. These results show that feeding of turbot and sole larvae is influenced by chemosensory processes at an early larval stage, and indicate that chemoreception may be an integrated part of turbot and sole larvae feeding strategy.     

Experimental analysis of the contribution of swimming and drifting to the displacement of reef fish larvae

The extent to which behaviour affects the dispersal of pelagic larvae in reef fishes has been a topic of major discussion among marine ecologists. Here, we experimentally quantified the extent to which the displacement of late-stage larvae of Abudefduf saxatilis is due to active movement (i.e. swimming) and drifting. We consider drifting as the component of larval displacement accounted for by the current. Drifting was quantified by comparing larval displacement to the displacement of passive particles in an extended flow chamber that gave larvae the free choice of swimming (i.e. swim with or against the current or not swim at all). We also determine whether drifting results from currents exceeding larval swimming capabilities or from the behavioural choice of larvae of not to swim against adverse currents. To do this, we compare the speeds of larval swimming in the extended flow chamber to those obtained in a smaller chamber in which larvae are behaviourally forced to swim due to space constraints and a retaining fence (most available data on larval swimming is based on this sort of chamber). Within the extended chamber, larvae tended to face the current and swim slower than it. This resulted in a net displacement increasingly determined by drifting. We also found that in the extended chamber, larvae swam at speeds between one and six times slower than the speeds they achieved in the “behaviourally modifying” smaller chamber. This suggests that the net displacement in the extended chamber was in part due to the behavioural choice of the larvae of not to swim. The importance of this “behavioural drifting” is discussed in terms of energy savings required for successful completion of the larval period and post-settlement survival. The idea that larvae may modulate their swimming behaviour raises caution for the use of published data regarding swimming capabilities of reef fish larvae when assessing the extent to which these fish actively affect their dispersal.     

Feeding,conversion efficiencies,and growth of larval mummichogs,Fundulus heteroclitus

Feeding rates, conversion efficiencies and growth of larvae of the mummichog Fundulus heteroclitus, an extremely abundant estuarine fish, were measured at temperatures ranging from 18° to 30°C. The food used was Artemia salina nauplii. At the time of total yolk sac absorption (5 to 7 days after hatching), the feeding rate decreased for a short time, an indication of a shift in metabolism. Higher feeding rates and growth occurred at higher rearing temperatures. The highest conversion efficiency (gross growth efficiency) was 1.1%, at 22°C. Mummichog larvae may be energetically inefficient compared with other fish species, but efficiency might not be critical for this fish, which is an opportunistic omnivore in an energy-rich environment.Contribution No. 291 of the Belle W. Baruch Institute for Marine Biology and Coastal Research, supported by DOE contract No. EY-76-5-09-0869.     

Yolk resorption,onset of feeding and survival potential of larvae of three tropical marine fish species reared in the hatchery

This paper provides basic early life-history information on milkfish (Chanos chanos), seabass (Lates calcarifer) and rabbitfish (Siganus guttatus) which may explain in part the observed differences in their survival performance in the hatchery. Egg size, larval size, amount of yolk and oil reserves and mouth size are all greater in milkfish than in seabass, and greater in the latter than in rabbitfish. During the first 24 h after hatching, rabbitfish larvae grow much faster than milkfish and seabass larvae at similar ambient temperatures (range 26°–30°C, mean about 28°C). The eyes become fully pigmented and the mouths open earlier in seabass and rabbitfish (32–36 h from hatching) than in milkfish (54 h). Seabass larvae learn to feed the earliest. Yolk is completely resorbed at 120 h from hatching in milkfish, and yolk plus oil at 120 h in seabass and 72 h in rabbitfish at 26° to 30°C. Milkfish and seabass larvae have more time than rabbitfish to initiate external feeding before the endogenous reserves are completely resorbed. Delayed feeding experiments showed that 50% of unfed milkfish larvae die at 78 h and all die at 150 h from hatching. Milkfish larvae fed within 54 to 78 h after hatching had improved survival times: 50% mortality occurred at 96 to 120 h, and 10 to 13% survived beyond 150 h. Unfed seabass larvae all died at 144 h, while 6 to 13% of those fed within 32 to 56 h after hatching survived beyond 144 h and well into the subsequent weeks. Unfed rabbitfish larvae all died at 88 h, while 7 to 12% of those fed within 32 to 56 h after hatching survived beyond 88 h. A delay in initial feeding of more than 24 h after eye pigmentation and opening of the mouth may be fatal for all three species.Contribution No. 167 from the SEAFDEC Aquaculture Department     

Larval ecology of the great barracuda, <Emphasis Type="Italic">Sphyraena barracuda</Emphasis>, and other sphyraenids in the Straits of Florida

The great barracuda (Sphyraena barracuda) is a widespread, ecologically and socioeconomically important coastal fish, yet very little is known about its larvae. We examined spawning and larval ecology of Western Atlantic sphyraenids using monthly ichthyoplankton samples collected over 2 years along a transect spanning the east–west axis of the Straits of Florida (SOF). Samples were dominated by the great barracuda (92.8%) and sennets (Sphyraena borealis and Sphyraena picudilla; 6.6%). While larval sennets and S. barracuda displayed similar vertical distributions (majority in upper 25 m), horizontal and temporal patterns of abundance suggested a spatial and temporal species replacement between larval S. barracuda and sennets that tracks adult ecology. The diet of both taxa consisted largely of copepods, with inclusion of fish larvae at 8 mm SL, and in S. barracuda alone, a switch in the wet season to exclusive piscivory by 12 mm SL (18 days post-hatch). A lack of piscivory in S. barracuda larvae captured in the dry season corresponded to slower larval growth than in the wet season. Larval growth was also related to size-at-hatch and larval age such that larvae that were larger at hatch or larger (older) at capture grew faster at earlier ages, suggesting faster larval growth, and indirectly larger hatch size, conveys a survival advantage. Unlike larval growth, instantaneous mortality rate did not differ with season, and no lunar cyclic patterns in spawning output were identified. Our results provide insight into the pelagic phase of sphyraenids and highlight the importance of both diet and hatch size to the growth and survival of fish larvae in low latitude oceanic environments.     

Effects of temperature on morphological landmarks critical to growth and survival in larval Atlantic cod (Gadus morhua)

The morphology and function of structures important to energy acquisition were studied from spawning to the stage of transformation of larva to pelagic juvenile in Atlantic cod, Gadus morhua L., from December 1991 to July 1992. Fertilized eggs produced by adult fish from two genetically discrete populations (Newfoundland and Scotian Shelf) were raised under similar conditions in the laboratory at temperatures of 5 and 10°C. Subsamples of larvae were removed from cultures daily for 10 d, and then less frequently, and fixed for light microscopy and scanning electron microscopy. Nine functional morphological landmarks important to feeding, respiration and locomotion were chosen from observation of 280 ind. These landmarks defined 12 major developmental stages, from hatching to the pelagic juvenile stage. One of the feeding landmarks, intestinal stage, varied as a function of age and size and the variance in development was higher at 10°C than at 5°C; Newfoundland larvae developed more complex intestines than did Scotian Shelf larvae. In addition, Newfoundland larvae had significantly higher growth rates than those of Scotian Shelf larvae. Despite the higher growth rates and greater structural complexity of the intestine in Newfoundland larvae, the rate of yolk utilization was not significantly different between Newfoundland and Scotian Shelf larvae. Staging of respiratory landmarks showed that the gill arches were probably used preferentially in feeding while respiration was cutaneous. The gills, operculum and gill rakers developed late in larval life and accompanied the transition from cutaneous to branchial respiration. In the yolk-sac period, development of feeding and respiratory structures may be largely genetically controlled. During exogenous feeding, extrinsic factors also become important, as shown by the size and age-independent variation in intestinal development of larval cod raised at different temperatures.     

Relationship between the growth and survival of larval Pacific bluefin tuna, Thunnus orientalis

We tested the hypothesis that a large body size and rapid growth rate affect the survival of larval Pacific bluefin tuna, Thunnus orientalis (PBT), and analyzed larval growth in relation to environmental conditions. Seven high density larval patches of PBT were tracked with reference buoys in the northwestern Pacific Ocean for 28–171 h in May–June from 2004 to 2008. The otolith radii and daily growth rates of the survivor larvae (collected on later tracking days of each tracking session) tended to be larger and more rapid, respectively, than those of original larvae (collected on earlier tracking days). A large body size was found to positively affect the survival of larval PBT, as did a rapid growth rate, even at an early larval stage (7 days after hatching). Generalized linear modeling showed that the otolith radius was influenced positively by the sea temperature, stratification parameter and food density, while the growth rate was influenced positively by the sea temperature and food density.     

Growth and mortality in young larval herring (Clupea harengus); effects of repetitive changes in food availability

Following yolk resorption, laboratory-reared larval Baltic herring (Clupea harengus L.) were exposed to two sequences of food restriction for 5 d and re-alimentation for 10 d. Comparisons regarding larval growth (standard length and content of water-soluble protein), mortality and content of the sum of trypsin and trypsinogen were made with larvae at a continuous high ration. Larvae exposed to varying prey abundance grew less in length than the control, and during the second high-ration period (Day 22 to 32) growth in length ceased. From the first low-ration period onwards, the content of water-soluble protein in these larvae was lower than that of the control larvae, and the survival rate of the low-high ration group was 59% compared to 77% in the larvae at a continuous high ration. In contrast, the effects of varying food availability were minor on larval content of trypsin and trypsinogen. Results are compared with previous findings in larval Clyde herring, and the effects of larval stock and timing and duration of food restriction on larval growth performance are discussed.     

Combined effects of temperature and salinity on fed and starved larvae of the mediterranean mussel Mytilus galloprovincialis and the Japanese oyster Crassostrea gigas

The combined effects of temperature, salinity and nutrition on survival and growth of larvae of the Mediterranean mussel Mytilus galloprovincialis and the Japanese oyster Crassostrea gigas were studied over a period of 7 d in the laboratory. Ripe adults, collected in spring and summer 1987 from natural populations in the Bay of Arcachon, France, were induced to spawn. Larvae of both species were cultured at four temperatures (15°, 20°, 25° and 30°C), four salinities (20, 25, 30 and 35S) per temperature, and two levels of nutrition (fed and unfed) per temperature/salinity combination. The fed larvae received a mixed algal diet of 50 cells each of Isochrysis galbana and Chaetoceros calcitrans forma pumilum per microlitre. In both bivalve species, larvae survived over a wide range of temperature and salinity, with the exception of mussel larvae, which died at 30°C. Statistical analysis indicated that nutrition had the greatest effect on larval development, explaining 64 to 75% of the variance in growth of M. galloprovincialis and 54 to 70% in growth of Crassostrea gigas. Unfed mussel larvae displayed little growth. Compared with temperature, the effect of salinity was very slight. M. galloprovincialis larvae exhibited best growth at 20°C and 35S and C. gigas at 30°C and 30S.     

Egg survival,embryonic development,and larval characteristics of northern shrimp (<Emphasis Type="Italic">Pandalus borealis</Emphasis>) females subject to different temperature and feeding conditions

Laboratory experiments on ovigerous females of northern shrimp (Pandalus borealis) were used to assess the effects of temperature and food ration on female condition during incubation and examine how combined effects of temperature and female condition influenced egg survival, embryonic development, and larval characteristics. Ovigerous females were maintained at 2°C, 5°C, and 8°C and fed on a low (three times/week; 2–2.7% W/W) or high ration (five times/week at satiation). The increase in temperature accelerated the developmental time of the eggs but their survival at 8°C was reduced. Conversion efficiency of yolk reserves in developing embryos was significantly reduced at elevated temperatures and larvae hatching at 2°C and 5°C were significantly larger and heavier than those hatching at 8°C. The experimental design did not result in any effect of food ration on the energetic condition of females or on egg characteristics and their biochemical composition. However, lower energy reserves were observed for females held at 8°C.     

Effects of algal and larval densities on development and survival of asteroid larvae

Effects of larval and algal culture density and diet composition on development and survival of temperate asteroid larvae were studied in the laboratory at Santa Cruz, California, USA, during summer and fall of 1990. Larvae of Asterina miniata were reared at two densities, 0.5 or 1.0 ml^-1, and fed one or two species of cultured phytoflagellates — Dunaliella tertiolecta alone or mixed with Rhodomonas sp. — at three concentrations of 5x10², 5x10³, and 5x10⁴ total cells ml^-1. Algal concentration strongly influenced larval development; however, larval density also had a marked effect. Development progressed further with increasing algal concentration. Larval growth and differentiation were sometimes uncoupled; i.e., growth measures were directly related to food level, while differentiation indicators were less so. At the lowest food level, growth was negative and differentiation was arrested at early precompetent stages; these larvae never formed juvenile rudiments or brachiolar attachment structures. Development times of larvae given more food ranged from 26 to 50 d and depended directly on food availability. Development time to metamorphosis at the highest food concentration was similar for siblings fed D. tertiolecta alone or mixed with Rhodomonas sp. In contrast, when food level was an order of magnitude lower, larvae fed the algal mixture metamorphosed significantly earlier than larvae fed the unialgal diet. This suggests interactive effects of food quantity and food quality. Survival was little affected by larval or food density, except at the lowest ration. Feeding experiments in well-controlled laboratory conditions are useful to predict and compare the physiological or developmental scope of response of larvae to defined environmental factors; however, results from such studies should not be extrapolated to predict rates and processes of larval development in nature.