Sensibilité des larves deSpodoptera littoralis [Lep.: Noctuidae] aux hyphomycètes entomopathogènesNomuraea rileyi etPaecilomyces fumoso-roseus

Résumé L'étude de la sensibilité des chenilles deSpodoptera littoralis Boisd. à des doses croissantes de spores deNomuraea rileyi (F.)Samson, montre que le 6^e et dernier stade est plus résistant que le 1^er et surtout les 4^e et 5^e stades larvaires. La virulence du pathotypeN. rileyi n^o 5 à l'égard des larves de cette noctuelle est élevée puisque le temps léthal 50 % (TL 50) est de 6 j en moyenne. Pendant l'incubation de la maladie les chenilles continuent de s'alimenter mais l'infection peut réduire jusqu'à 60 % la prise alimentaire par rapport à la consommation des larves des lots témoins. Toutes conditions égales par ailleurs, la mortalité provoquée parN. rileyi n^o 5 après traitement des larves nouvelles-nées est supérieure lorsque les insectes sont maintenus à 25°C par rapport à celle constatée à 20°C. Cependant pour une dose d'inoculum élevée, les conditions thermiques (20°, 25° ou 28°C), ne modifient pas sensiblement la réponse à l'infection parN. rileyi n^o 5 alors qu'elles limitent l'efficacité d'un pathotype moins virulent:Paecilomyces fumoso-roseus Wize no^o 39.

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Summary Laboratory studies were conducted to determine the susceptibility of various larval instars of the cotton leafworm,Spodoptera littoralis Boisd. to different spore doses ofNomuraea rileyi (F.)Samson to investigate the influence of temperature on the infection by this fungus and byPaecilomyces fumosoroseus Wize. Contaminations were obtained by direct spraying on larvae in tower apparatus or by feeding of larvae on treated pieces of leaf during 48 h or 72 h. The influence of cryptogamic infection on food consumption was studied by measuring surfaces of standard cabbage leaf disces submitted to treated and control larvae. Angular values mortality rates were submitted to the 2- or 3-way analysis of variance and comparisons of means were made by theDuncan's test. In some cases we have also considered the time-mortality and the dose-mortality curves. The 6th instar was more resistant than all other tested instars. TL50 were found to be 6 days in most cases. During incubation of the disease, larvae continued to feed, but food consumption could be reduced at 40 % of controls. Larval mortality due toN. rileyi N^o 5, recorded after 8 days of incubation, was higher at 25°C than at 20°C. Nevertheless, at high dosage, efficiency ofN. rileyi N^o 5 was not affected by temperature at 20°, 25° and 28°C. The other pathotype,P. fumoso-roseus N^o 39 was more effective at 20°C than at 25° and 28°. At 32°C, the temperature, unfavourable to fungal growth, limited mortality at non significant rates.

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Avec la collaboration technique deH. Vermeil de Conchard.     

Interactions between a braconid,Microplitis croceipes,and a fungus,Nomuraea rileyi,in laboratory-reared bollworm larvae

The parasite Microplitis croceipes required 1.1 days longer at 26°C to complete development in Heliothis zea larvae than was required for the fungus Nomuraea rileyi to kill the host larvae and sporulate. Host larvae parasitized by M. croceipes or infected with N. rileyi failed to complete a fifth larval molt or pupate. Of the remaining healthy larvae, one-half completed six larval stadia before popation. Larvae parasitized by M. croceipes were predisposed to infection by N. rileyi, but the fungus inhibited development of M. croceipes if host larvae were infected with N. rileyi within 1 day after parasitization.     

Susceptibility of Heliothis zea larvae to Nomuraea rileyi at various temperatures

Laboratory studies were conducted to determine the susceptibility of various larval instars of Heliothis zea to different spore doses of Nomuraea rileyi at constant and variable temperatures. The fungus was most effective at 20° and 25°C, with a mortality of 80% and 71%, respectively. At 15°C the disease progressed very slowly with larval mortality occurring in 12–28 days post-treatment. Conversely, at temperature ranges above 15°C, the mortality of the larvae occurred in 6–12 days. Three different combinations of variable temperatures included 20–30°, 25–30°, and 20–35°C, but mortality did not exceed 46%. Larvae in the third to fifth instars were more susceptible to infection than were those in the first and second instars.     

Early developmental stages of the scorpaenid fish,Scorpaena miostoma,reared in the laboratory

Embryonic development and morphology of the eggs, egg masses and larvae of the scorpaenid fish,Scorpaena miostoma are described on the basis of the laboratory-reared specimens. The gelatinous egg mass is not bilobed but simple, oval and balloon-like in shape. It usually floats at or near the surface. The individual egg is colorless and ovoid in shape, measuring 0.86 × 0.75 mm. Hatching occurs between ca. 34 and 40 hours after spawning at the water temperature of 22–24°C. A newly hatched larva measures 1.55 mm in notochord length and has ca. 25 myomeres. The finfold is balloon-like in shape and almost completely covers the larval body. Minute “cell-like” granules scatter on the findfold.     

Some effects of time,temperature and feeding on infection rates with Babesia bovis and Babesia bigemina in Boophilus microplus larvae

Dalgliesh R. J. and Stewart N. P. 1982. Some effects of time, temperature and feeding on infection rates with Babesia bovis and Babesia bigemina in Boophilus microplus larvae. International Journal for Parasitology12: 323–326. Percentages of larval ticks in which Babesia bovis and B. bigemina parasites could be detected (infection rates) were determined after the larvae had been exposed to temperatures between 9°C and 27°C for periods of 1–35 days and then either fed on calves or heated at 37°C to stimulate babesial development. Infection rates with both species increased during 2–4 weeks after the larvae hatched, regardless of the temperature of exposure. Infection rates with B. bovis were higher after exposure of larvae to 14°C than to 27°C. This effect was less pronounced with B. bigemina. Infection rates were higher in fed larvae than in unfed, ‘heat stimulated’ larvae. The findings indicate that infected larval ticks become more efficient vectors of Babesia during the first 2–4 weeks after hatching and that repeated sampling of a tick population is necessary to determine valid infection rates.     

Effects of temperature experienced during embryonic development on biomass and C and N composition at hatching in Palaemon serratus (Pennant, 1777)

In decapod crustaceans, the conditions experienced during embryonic development trigger phenotypic plasticity of the larvae at hatching. The objective of this study was to test the effects of temperature during embryonic development of Palaemon serratus on the phenotypic plasticity of hatching larvae. We incubated egg-bearing females from eggs laying to hatching at four temperatures (10, 15, 18 and 20°C). Weight, carbon and nitrogen contents were measured on newly laid eggs and on freshly hatched larvae. The duration of embryonic development was negatively correlated with incubation temperature. At 20°C, all females abandoned their eggs during development. Incubation temperature had no effect on the weight and the percentage of N of the larvae at hatching, while it did affect their percentage of C and their C/N ratio. Embryos incubated at 10°C seemed to produce larvae with fewer lipid reserves than those incubated at 15 and 18°C. They probably overconsumed their lipid reserves to compensate for the metabolic losses due to the low temperature. These results provide information on the link between maternal investment per egg and larval development in P. serratus.     

Spécificité des hyphomycètes entomopathogènes pour les larves de lepidoptèresNoctuidae

Résumé L'étude de la sensibilité des 4 espèces de noctuelles,Mamestra brassicae L.,Spodoptera littoralis Boisduval,Spodoptera frugiperda Smith etScotia ipsilon Hübner, aux hyphomycètes a été conduite avec 10 souches dePaecilomyces fumoro-roseus (Wize) Brown & Smith, 6 deNomuraea rileyi (Farlow) Samson, 1 deBeauveria bassiana (Bals.) Vuill. et 1 deMetarhizium anisopliae (Metsch.) Sor. Les chenilles nouvelles-nées agées de moins de 16 h ont été exposées pendant 24 h sur des rondelles de feuilles contaminées par aspersion des inoculums, puis élevées individuellement à 25±1°C. A la dose de 3.10⁵ conidiospores/cm², l'activité pathogène des germes sur les chenilles varie dans de larges proportions suivant l'espèce-h?te et suivant l'isolat considérés. L'espèce la plus sensible à la plupart des souches estM. brassicae et la plus résistanteS. littoralis. Les germes les plus performants présentent des temps létaux 50% compris entre 2 et 3,5 j (à la dose de 3.10⁵ spores/cm²) et des doses létales 50% de 10³ à 10⁴ conidiospores/cm² (mortalités cumulées le 6e j après traitement); ce sont les isolats:P. fumoso-roseus n^o 13, 32, 39, 40 et 45 pourM. brassicae, N. rileyi n^o 5 pourS. littoralis; P. fumoso-roseus n^o 32 pourS. frugiperda etM. anisopliae n^o 139 pourS. ipsilon. La virulence élevée du pathotypeN. rileyi n^o 5 pourS. littoralis et son agressivité à l'égard des 3 autres noctuelles, confirment l'intérêt de cet hyphomycète en dehors de son aire d'origine (Sud-Est des Etats-Unis). Par ailleurs, les aptitudes des souches européennes deP. fumoso-roseus montrent que cette espèce présente des potentialités pour la lutte biologique contre divers noctuides.

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Summary Bioassays were carried out on 1st instar larvae to determine the relative susceptibility of 4 noctuid species,Mamestra brassicae L.,Spodoptera littoralis Boisduval,Spodoptera exigua Smith andScotia ipsilon Hübner to 10Paecilomyces fumoso-roseus, 6Nomuraea rileyi, 1Metarhizium anisopliae and 1Beauveria bassiana isolates. Neonate larvae, 2 to 16 h old, were exposed for 24 h to leaf pieces treated with conidial suspensions of each fungal strain and then transferred to untreated foliage at 25±1°C. When tested at 3×10⁵ conidia/cm², pathogenic activity of hyphomycetes towards noctuids showed high variability depending on both host-species and fungal isolate.M. brassicae was more susceptible to tested isolates than the other 3 noctuids. In contrast,S. littoralis was the most resistant species. The most active pathotypes had LT 50 values ranging from 2 to 3,5 days at the concentration of 3×10⁵ conidia/cm² and LD 50 values ranging from 10³ to 10⁴ conidia/cm² (mortality recorded at 6 days post exposure). According to these data, there were 5P. fumoso-roseus isolates (nos 13, 32, 39, 40 and 45) highly active againstM. brassicae, oneN. rileyi isolate (n^o 5) againstS. littoralis, oneP. fumoso-roseus isolate (n^o 32) againstS. frugiperda and oneM. anisopliae isolate (n^o 139) againstS. ipsilon. Bioassays establishing the high virulence of the FloridaN. rileyi isolate (n^o 5) to the Egyptian cottonworm and its pathogenic activity against the other tested noctuids, confirmed the interest for this fungal entomopathogen which has been demonstrated on noctuid populations in the southeastern United States. Moreover results of the host range assays of European isolates ofP. fumoso-roseus showed that this species appears to be an other potentially valuable microbial control agent against noctuid pests.

     

In vitro storage of unfertilized eggs of the Eurasian perch and its effect on egg viability rates and the occurrence of larval malformations

Ova ageing is the most important factor affecting fish egg quality after ovulation. Long-term storage of fish ova, using cryopreservation and vitrification techniques, has been unsuccessful to date. Instead, short-term in vitro ova storage has been used successfully and optimized in some cultured fish species. In vitro ova storage can drastically improve mass production of larvae and juveniles in the hatcheries by providing the possibility of the synchronous artificial fertilization for different females. To study how long unfertilized eggs of Eurasian perch (Perca fluviatilis L.) can retain their fertilizing ability after stripping, eggs were stored at temperatures of 4°C, 8°C and 12°C for 72 h post-stripping (HPS). The stored eggs of four female perch were separately fertilized at 0 h (i.e. control eggs fertilized before storage) and at 6-hour intervals during the experimental period of 72 h. The embryos reaching the eyed-egg and hatched-larvae stages, eyed-egg mortality and larval malformation rates were recorded as indices of egg quality. The results indicated that the maximum eyed eggs and hatched larvae (86% and 63%, respectively) were observed for eggs fertilized immediately after stripping, whereas the storage of the eggs at 4°C for 48 HPS decreased the eyed-egg and hatched-larvae rates to 46% and 17%, respectively. The use of a higher storage temperature resulted in a more rapid decrease in egg viability: eyed-egg and hatched-larvae rates of 23% and 9%, respectively, were obtained after 48 HPS storage at 8°C and 2% and 1% for eggs stored at 12°C. Eyed-egg mortality and larval malformation rates were not significantly affected by post-stripping ova ageing for at least up to 36 h. Thereafter, both values increased significantly and were measured to be the highest in the most aged ova. The present study demonstrated that stripped Eurasian perch eggs can be stored for at least 12 h at 4°C to 12°C without a significant reduction in their quality.     

Susceptibility of Oryctes rhinoceros adults to Metarrhizium anisopliae

The susceptibility of Oryctes rhinoceros adults to infections due to entomopathogenic fungi (Beauveria bassiana, Beauveria tenella, Metarrhizium anisopliae, Paecilomyces fumoso-roseus, and Spicaria rileyi) was studied by spraying titrated spores suspensions on the insect integument. The results show a definite susceptibility of the adults to Metarrhizium anisopliae strains of the major type only.     

Development,metamorphosis, and natural history of the nudibranch Doridella obscura Verrill (Corambidae: Opisthobranchia)

The doridacean nudibranch Doridella obscura Verrill was raised through one complete generation in laboratory culture, and spawning behavior monitored for a year at monthly intervals in Barnegat Bay, New Jersey.The nudibranch deposited egg masses throughout the year in Barnegat Bay, and the larvae remained viable at temperatures ranging from 1.5 to 28 °C. At 25 °C the eggs hatch 4 days after oviposition, and the planktotrophic veliger larvae swim and feed for 9 days before they metamorphose. Settlement occurs specifically on the bryozoan Electro crustulenta (Pallas). The spirally coiled larval shell grows rapidly until the dorsal mantle fold is retracted from the aperture 5–6 days after hatching. Although starved larvae grow only slightly and do not metamorphose, they resume normal development on introduction of suitable food. Newly metamorphosed juveniles consume algae and debris on the surface of the bryozoan until they grow large enough to attack the living zooids of E. crustulenta.The life cycle of Doridella obscura is short (26 days at 25 °C), allowing the nudibranchs to take advantage of short-lived Electra crustulenta colonies in unstable habitats in bays and estuaries.     

Survival and behavioral characteristics of amphidromous goby larvae of Sicyopterus japonicus (Tanaka, 1909) during their downstream migration

To understand the ecology and environmental tolerances of newly hatched larvae of the amphidromous fish Sicyopterus japonicus during their downstream migration, the salinity tolerance of eggs, 0-15 day old larvae, and adults, and the temperature tolerance, specific gravity and phototaxis of hatched larvae were examined. Tolerances of adults were measured as survival after a 24 h challenge in freshwater (FW), brackish water (1/3 SW) and seawater (SW). The survival rate of adult S. japonicus was 100% in FW and 1/3 SW, while none survived in SW. Hatching success of eggs (30 eggs each) was significantly higher in FW (mean: 73%) and 1/3 SW (73%) than in SW (19%). Tolerance of newly hatched larvae to salinity and temperature was investigated in different combinations of salinities (FW, 1/3 SW and SW) and temperatures (18, 23 and 28 °C). Larval survival was significantly different in each salinity and temperature. Survival rate was significantly higher in 1/3 SW than in FW and higher in SW than in FW at 23 °C and 28 °C. At the latter part of the experiment, there was no survival in FW and at 28 °C. Survival was higher in lower temperatures, but larval development did not occur in FW. Specific gravity of newly hatched larvae was 1.036 at 28 °C and 1.034 at 23 °C. When exposed to a light source on one side of an aquarium, larval distribution was not affected. Our results indicated larval S. japonicus are more adapted to brackish water and seawater than freshwater, while the adults and eggs are more adapted to freshwater and brackish water than seawater. This is consistent with their amphidromous life history with growth and spawning occurring in freshwater and the larval stage utilizing marine habitats.     

Temperature and laboratory feeding rates in Carcinusmaenas L. (Decapoda: Portunidae) larvae from hatching through metamorphosis

Larvae of Carcinus maenas L. were reared in the laboratory and temperature-dependent stage durations of successive instars were examined at 12, 15, 18, and 25°C. Feeding rates (FR), in numbers of newly hatched Artemia nauplii, biomass, and energy consumed by a single crab larva during 24 h, were measured daily through the moulting cycles of all larval stages at the four temperatures. Dry weight (DW) and elemental content of carbon (C), nitrogen (N) and hydrogen (H) were analysed in newly hatched (0 h) and 1-day-old (24 h) Artemia nauplii at six temperatures (6, 9, 12, 15, 18, 25°C). Due to a 24 h feeding regime, the temperature dependent “mean nutritive value” of newly hatched brine shrimp nauplii is defined, individual biomass (DW, C, N, H) and energy (Joule) 12 h after hatching. General figures in changing individual daily FR, and temperature-dependent peculiarities are discussed. The total amount of food ingested by successive larval instars increases exponentially, while the increasing cumulative quantity consumed by individual crab larvae on successive days of development is described by power functions. At higher temperatures significantly less absolute biomass and energy is consumed during the entire larval development. C. maenas megalops are the main energy consumers in larval life, contributing 41 to 67% (12 to 25°C) to the total larval energy intake between hatching and metamorphosis. Larval ability to adapt to increasing metabolic costs for maintenance in higher water temperatures is discussed with average daily feeding rates (AFR). Improved efficiencies are presented for the cumulative larval energy budget, 31% in assimilation, and 4.4% in gross growth (K₁).     

Effects of grapevine bunch exposure to sunlight on berry surface temperature and Lobesia botrana (Lepidoptera: Tortricidae) egg laying,hatching and larval settlement

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Significant mortality of eggs and young larvae of two pine processionary moth species due to the entomopathogenic fungus Metarhizium brunneum

Bioassays were conducted to determine the susceptibility of egg masses and young larvae of two pine processionary moth species, Thaumetopoea pityocampa and Thaumetopoea wilkinsoni, to two strains (ARSEF4556, V275) of the entomopathogenic fungus Metarhizium brunneum. Mortality of treated eggs by both strains ranged from 96% to 99% but not all of this was caused by M. brunneum since control groups also experienced egg mortality due to saprophytic fungi. Still, larvae hatched in the laboratory from eggs treated with M. brunneum were all killed by this fungus, acquiring M. brunneum conidia, whereas larval mortality was 0% in the control groups. Young larvae of both pine processionary moth species were also highly susceptible to ARSEF4556 and V275 with larval mortality ranging between 94% and 100%, 8 days post-inoculation, with the vast majority of larvae being killed within the first 2–4 days. Larval mortality was dose dependent. Results were consistent across the two pine processionary moth species, showing that the pathogenicity of M. brunneum to both eggs and young larvae might be promising for biological control of these insect pests. The study also showed that non-target parasitoids of pine processionary moth eggs were also susceptible to M. brunneum. Further work is required to understand and reduce the M. brunneum effect on non-target insects.     

Contribution to the biology of the citrus bud mite, Aceria sheldoni (Ewing) (Acarina: Eriophyidae)*

In laboratory studies mites of Aceria sheldoni were reared on citrus fruit peels, beneath coloured cellophane hoods, to cater for their thigmotaxis and sensitivity to light of particular wave lengths. Hatching was most successful at 25°C and 98% r.h. but was reduced by low humidities (35–40% r.h.), when abnormal dwarf larvae emerged. The eggs hatched in 3–14 days; the length of a generation (egg to egg) was 12–33 days. The threshold of embryonic development was 9 °C and that for completion of the life cycle, egg to egg, was 12.5 °C. The average number of eggs laid per female was six (4–8). It increased to twelve (5–19) if the mite, during its larval stages, had been fed on buds. The vitality of both the eggs and the mature mites was tested by exposure to extreme low and high temperatures (below 0 °C, 39 °C): 50% of mites died after 30 min at 30 °C; 50% died after 30 min at –15 °C or lower.     

Soybean leaf consumption by Nomuraea rileyi (Fungi: Deuteromycotina)-infected Plathypena scabra (Lepidoptera: Noctuidae) larvae

To determine the impact of Nomuraea rileyi on consumption by the green cloverworm, Plathypena scabra, larvae were reared from eggs obtained from field-collected moths, inoculated with conidia, and placed individually in separate plastic vials with a piece of surface-sterilized soybean leaflet. No significant differences in consumption rates were found between N. rileyi-inoculated and control larvae until after 144 hr post-treatment. After this period, consumption by larvae inoculated as first, second, third, or fourth instars was significantly less than that of control larvae because of mortality. First or second instars inoculated with N. rileyi conidia had significantly longer subsequent stadia than did control larvae. The average LT₅₀ of all inoculated instar groups was 6.5 days, and there was little evidence of significantly different N. rileyi susceptibility among instar groups. In general during the N. rileyi infection period leading to their deaths, inoculated P. scabra larvae had consumption patterns that were very similar to those of healthy larvae.     

The effect of temperature on the growth,development and survival of Macropetasma africanus (Balss) (Penaeoidea:Penaeidae) larvae reared in the laboratory

Macropetasma africanus (Balss) has been successfully spawned and its larvae reared under controlled laboratory conditions. The relationship between egg number (E) and female total length (L) was E = 18.59 L^2.11. An experiment was designed to test the effect of temperature on larval development, survival and growth. Temperature effected larval development time, from 13–15 days at 25°C, to 25 days at 15°C (nauplius 1 to post-larva). Mortality was low for the naupliar stages at 25, 22 and 18°C, while at 15°C only 52% of the larvae reached nauplius 6. Mortality was highest from nauplius 6 to protozoea 1 (17, 21, and 18% at 25, 22, and 18°C, respectively), but decreased considerably for all temperatures once the mysis stage was reached. Overall survival rates from nauplius 1 to post-larva decreased with decreasing temperature (65, 54, 48, and 39% at 25, 22, 18, and 15°C respectively). Temperature also significantly affected larval growth. At 25°C mean total length was significantly (P < 0.05) larger than at 15°C (protozoea 2 to post-larva), while from protozoea 3 to post-larva total length differences were significantly different (P < 0.05) between 18 and 25°C. M. africanus has a major spawning peak in summer, suggesting that there may be a selective advantage to reproducing during the warmer months.     

Aseptic rearing of Epiphyas postvittana (Lepidoptera: Tortricidae) on a meridic diet

The light brown apple moth, Epiphyas postvittana (Walker), was reared aseptically from newly hatched larvae to adults for three generations on a meridic diet. Growth and development on this diet were comparable with that on oligidic diets and on host‐plants. At 25 ± 1°C, 50–60% R.H., and 12‐h photophase, the average larval period was approx. 28 days, pre‐pupal and pupal 11–12 days, and pre‐oviposition approx. 2 days; the incuva‐tion period of eggs was approx. 7 days. The average developmental period from egg to adult was thus about 49 days; 89 % of larvae reached the adult stage. The average fecundity was 598 eggs per female, and the sex ratio was nearly 1:1. There were no differences in the rate of development or in fecundity of insects reared on media with and without a mould inhibitor.     

Induction of larval diapause in the blowfly, Calliphora vicina R.-D. (Diptera, Calliphoridae) under field and laboratory conditions

Field experiments conducted in the environs of St. Petersburg (Russia) with a local population of Calliphora vicina showed that induction of larval diapause under natural conditions was significantly dependent on day lengths and temperature. The maternal photoperiodic response had a distinct threshold: the first diapausing larvae hatched from the eggs laid in the middle of August when the day length was 16 h; at shorter photoperiods, the fraction of diapausing larvae depended only on temperature. At the mean temperature of 16°C, larval diapause was rarely recorded; at 12–13°C, about 50% of the larvae entered diapause; at 7–9°C, nearly all the larvae entered diapause. These results of the field experiments agree well with the parameters of photoperiodic and thermal responses studied in the laboratory at constant temperatures and photoperiods.     

Effects of temperature and salinity on the eggs and early larvae of Caranx mate (Cuv. & Valenc.) (pisces: carangidae) in Hawaii

Eggs and larvae of the carangid fish, Caranx mate (Cuv. & Valenc.), were incubated at various temperature (17.2 to 33.1 °C) and salinity (10 to 42 ‰) combinations in five experiments. The following rates were directly proportional to temperature: embryonic development, yolk absorption, eye and jaw development, and increase in length. Unfed C. mate larvae attained a maximum size at 25 °C and 20 ‰ Eyes and jaws of larvae were functional by the end of the yolk sac stage at all temperature and salinity levels tested.Hatching success and larval survival at the end of the yolk sac stage were generally greater than 50 % between 22° and 32°C. Hatching success and larval survival at the end of the yolk sac stage were reduced at salinity extremes, especially in low temperature-low salinity and high temperature-high salinity combinations. The frequency of morphological abnormalities was also high at extreme temperatures and salinities.The incipient upper thermal TL_m for unfed C. mate larvae acclimated to 23.8°C increased from 31.5°C for newly hatched larvae, to 34.2°C for 72 h larvae, but decreased to 32.0°C for starving larvae after the exhaustion of the yolk supply.