Sound and vibration are not obligatory host finding stimuli for the bark beetle parasite,Coeloides brunneri [Hymenoptera: Braconidae]

Previous researchers, have speculated or concluded thatCoeloides brunneri Viereck females detect their bark beetle hosts by perception of the vibrations or sound made by boring larvae. However, when placed on logs containing various actively mining stages ofDendroctonus pseudotsugae Hopkins,C. brunneri females actively searched for the host only on logs infested with young or maturing brood larvae, and oviposited only in logs with maturing brood larvae. Moreover, when offered larvae in logs that had been frozen at approximately ? 50°C, and then allowed to thaw at room temperature for 2 days, they found the motionless, dead larvae, and oviposited through the bark on to them. Therefore,C. brunneri is able to find ist host by perception of some stimulus (or stimuli) other than sound or vibration.     

Effect of temperature on germination,growth, and infectivity of the mosquito pathogen Tolypocladium cylindrosporum (deuteromycotina; hyphomycetes)

The mosquito pathogen Tolypocladium cylindrosporum was examined with regard to its response to temperature. Similar temperature ranges were found for growth, germination, and infectivity of blastospores and conidia. Germination occurred at 8° and 33°C but not at 6° and 35°C. Optimal germination and growth was noted between 24° and 27°C for both spore types. Infectivity of blastospores and conidia at different temperatures was examined by exposing L₂Aedes sierrensis larvae to concentrations of 5 × 10⁵ blastospores/ml or 5 × 10⁶ conidia/ml. Larvae were incubated at 12°, 15°, 25°, and 30°C. Infection occurred at all temperatures tested with LT₅₀ values ranging from 22.7 days (12°C) to 5.6 (25°C) days for conidia and 4.7 days (12°C) to 0.6 day (25°C) for blastospores. These results confirmed earlier findings that blastospores infected and killed host larvae more rapidly than conidia and suggested that this difference is largely due to the more rapid germination rate of blastospores. These experiments demonstrated that T. cylindrosporum can be active against mosquito larvae over a broad range of temperatures encompassing both the cold-water habitat of certain temperate mosquito species as well as the habitat of tropical vector species.     

Egg laying and host stage preference at constant temperatures inEncarsia tricolor [Hym.: Aphelinidae]

Encarsia tricolor Foërster is a facultative autoparasitoid that develops on the important pestTrialeurodes vaporariorum (Westwood) in outdoor crop conditions, which makes this aphelinid species promising for biological control programs in regions where field and protected crops coexist. In this paper we report the results obtained in the study of daily and totalE. tricolor egg laying and of adult female preference for different host stages in which to lay eggs at constant temperatures in the range 10 to 32 °C. Only whitefly nymphs were present in the searching arena (tomato leaflets). The mean number of eggs laid per female in one day ranged from 4.0 (10 °C and 32 °C) to 15.2 (24 °C). The mean total number of eggs increased with temperature from 10 to 28 °C, reaching a maximum of 123 eggs per female at 28 °C, and decreased sharply from 28 to 32 °C. The relation between the intrinsic rate of increase (r_m) and temperature in the range 10 to 28 °C followed a straight line whose equation was r_m=?0.076+0.011*T (R²=0.99). The r_m ofE. tricolor was greater than the r_m ofT. vaporariorum when temperature was higher than 9.2 °C. The preference for any particular host instar in which to lay eggs was not always significant. However, N4 was the host instar preferred whenever preference was statistically significant.     

The susceptibility ofTribolium castaneum [Col.: Tenchrionidae] toFarinocystis tribolii [Protozoa: Schizogregarinida]

The LT₅₀ ofFarinocystis tribolii Weiser to larvae ofTribolium castaneum (Herbst) increased with the age of the insect indicating that older larvae were relatively more tolerant to the infection though there was 100 % mortality ultimately. The adults were less susceptible than larvae and between sexes, females were more susceptible than males. The number of spores produced increased with the stage of the larvae, but there was no variation in the size of spores in the different instars. The LC₅₀ on 20th and 40th day of inoculation were 1.4×10⁷ and 2.1×10⁶ respectively. Mortality-time due toF. tribolii was shorter at 35 °C than at 25 °C. Sporulation occurred earlier at 35 °C than at 25 °C.     

Metarhizium spp. isolates from madagascar: Morphology and effect of high temperature on growth and infectivity to the migratory locust,Locusta migratoria

Five strains ofMetarhizium anisopliae (Metsch.) Sorokin and one strain ofMetarhizium flavoviride Gams &; Rozsypal originally isolated in Madagascar were studied. Measurements of conidia and, for the first time, also of blastospores produced in a liquid medium were used for species and variety determination. Blastospores ofM. flavoviride were more homogenous in their size than those ofM. anisopliae. Growth at high temperatures between 25° and 40°C showed that 4 isolates ofM. anisopliae grew at 36°C andM. flavoviride grew at 38°C. Using alternating day/night temperatures (8/16 h) the three strains tested could also tolerate 40°/25°C. In bioassays, fiveMetarhizium spp. isolates were tested against third and fourth instar larvae ofLocusta migratoria (L.) at two alternating day/night temperatures of 30°/25°C and 36°/25°C. In the cooler regime, all strains caused a mortality of 50% within 5.9 to 8.5 days (median lethal time), while in the 36°/25°C treatment only the thermophilicM. flavoviride and oneM. anisopliae strain isolated from a soil sample gave comparable results with median lethal times of 6.8 and 7.3 days, respectively.     

Bioecology ofLydella thompsoni Herting, [Dip. Tachinidae] within the Rhone Delta in Southern France

Lydella thompsoni Hertin,Tachinidae, is an endoparasitoid frequently associated in southern France with larvae ofOstrinia nubilalis, Pyralidae, Sesamia nonagrioides, Noctuidae, Archanara geminipuncta andA. dissoluta, Noctuidae. The tachinid was reared successfully at 21°C in the laboratory by providing the adult flies with a high humidity, a light intensity of 8,000–10,000 lux for mating and a mixture of casein proteolysate and honey as food. Under such conditions, the flies lived for about 30 days and about half of them mated successfully. Hosts were infested by dissecting mature tachinid females and placing 1–2 of the extracted planidia onto each moth larva. Half of the planidia successfully entered their larval hosts. The biology of the tachinid larvae was studied on the 2 main hosts,O. nubilalis andS. nonagrioides. At 21°C, 25°C and 28°C, larval development took less time onO. nubilalis than onS. nonagrioides. At 25°C, female larvae onO. nubilalis required 9.0±0.5 days and onS. nonagrioides 10.5±0.3 days, male larvae onO. nubilalis required 8.3±0.5 days and onS. nonagrioides 10.6±0.3 days. Pupal duration was also influenced by the larval host. In winter, 2nd instar larvae ofL. thompsoni enter a resting or quiescent condition. This condition is terminated sooner (December–January) in larvae developing onS. nonagrioides than in those developing onO. nubilalis (February–March). The life cycle of the tachinid in the field was studied by trapping flies in water dishes and by collecting parasitized host larvae from various plants. Flies were caught from April to October, mainly in September. Larvae of the spring generation of the parasitoid developed on larvae of species ofArchanara that fed on the reed,Phragmites communis. From 1976 to 1982, parasitism averaged about 16%. Parasitism by summer generation onS. nonagrioides was highest (4–5%) in June. Parasitism ofO. nubilalis did not change by more than 2 fold in either of the 2 summers studied (10–17 % in 1981, 6–10 % in 1982). By September the numbers of host larvae had increased to 10–20 times the number available earlier in the season. Larval populations ofL. thompsoni similarly increased from 3–400 larvae per ha in June to 4–5,000 per ha in September. The stability of the relationship between this parasitoid and its hosts in southeastern France is discussed and compared to relationship described elsewhere.     

Superparasitism ofEpilachna varivestis [Col.: Coccinellidae] byPediobus foveolatus [Hym.: Eulophidae]: Influence of temperature and parasitoid-host ratio

Superparasitism ofEpilachna varivestis Mulsant larvae by the hymenopterous parasitoid,Pediobius foveolatus (Crawford), occurred under laboratory conditions. However,P. foveolatus avoided previously parasitized larvae in a manner which was directly related to the number of times host larvae were initially parasitized. Increasing the parasitoid-host ratio also increased percent host mortality and highest overall host mortality occurred at 15.6°C when the parasitoid-host ratio was 10∶10. Higher temperatures (22° and 28°C) and higher parasitoid-host ratios yielded higher numbers of parasitized larvae although a significant number of parasitoids failed to emerge at the highest parasitoid-host ratio. Higher temperatures along with increasing parasitoid-host ratios favored production of more male parasitoids.     

Longevity ofBracon brevicornis [Hym: Braconidae] adults as influenced by nourishment on artificial and natural foods

Laboratory trials were undertaken to investigate the nutrition ofBracon brevicornis Wesmael adults on: (1) artificial diets (sucrose, treacle, honey bee and tap water); (2)Sesamia larvae (exposed under 3 manipulations); (3) 5 host species (Galleria, Ephestia, Sesamia, Spodoptera andOstrinia); in addition to starvation at 12, 15, 20, 25, 30, 35 and 40°C. Among artificial diets, honey bee solution was the most suitable that prolonged significantly each sex to be more than 1 month at 25°C.A Bracon female supplied daily with a freshSesamia-larva was significantly longer-lived than when kept with the “unchanged”Sesamia-larva. Larvae of borers (Sesamia andOstrinia), gave significantly the highest longevity for females among other host species. Adult-longevity was significantly decreased with an increase in temperature. Females were significantly longer-lived than males when kept only with living host insects. However, such sex difference was generally uncertain when either was supplied with an artificial diet or starved under temperatures ranging from 12–40°C.     

Energy budget for the larval development ofElminius modestus (Crustacea: Cirripedia)

Biomass (CHN), respiration rate and food uptake were estimated for the larval development ofElminius modestus at three temperatures (12, 18, 24°C). Mean values of dry weight, elemental composition and energy equivalents increased exponentially with the development from nauplius II to VI. Dry weight, elemental composition and energy content exhibited the highest values at 18°C. Respiration rates increased with the larval stages expressed by a power function, but increased logarithmically with the dry weight of the larvae. The cypris larvae showed a reduced respiration rate compared with nauplius VI. The ingestion rate was measured at a concentration of 100 cells ofSkeletonema costatum μl⁻¹. At 12 and 18°C ingestion rates increased exponentially and at 24°C by a logarithmic function. The fittings were used to estimate the energy budget ofE. modestus during larval development. The energy content of the larvae increased during the development from nauplius II to VI by a factor of 21 at 12°C, 25 at 24°C and 31 at 18°C. The estimated energy content of the freshly metamorphosed barnacle is 100 mJ (12°C), 130 mJ (24°C) and 150 mJ (18°C). The assimilation- (A/I) and gross growth efficiencies (K₁) increased strongly during the development from nauplius II to VI (A/I: 6–14% in nauplius II to 50–90% in nauplius VI; K₁: 4% in nauplius II to 75% in nauplius VI). The net growth efficiency (K₂) showed a relatively constant level ranging between 57 and 83%.     

Detection of food source by PCR analysis of the gut contents of Aldrichina grahami （Aldrich） （Diptera： Calliphoridae） during post-feeding period

The last meal of sarcophagous maggots may be useful in identifying the species on whose flesh they have fed （the＂host＂species）. The DNA profile of the host species may indeed be detectable in the＂last meal＂. In this paper, mitochondrial DNA analysis of gut contents was used to identify the prior host of post-feeding larvae of Aldrichina grahami （Aldrich） （Diptera： Calliphoridae）. A modified logistic equation was fitted to estimate the probability of identifying the host under five different constant temperatures （16, 20, 24, 28 and 32 ℃）. Our results shows that the detected time ranged from a maximum of 24 h at 32℃ to 42 h at 16℃ and a minimum of 12 h at 32~C to 30 h at 16℃. Furthermore, the host detection time was also calculated to give the maximal time after larval hatching from the egg. These results indicate that, in criminal cases where the maggots stray from the corpse, the last meal of the larvae should not be overlooked as potentially critical evidence.     

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.     

Temperature,salinity tolerance,and buoyancy during early development and starvation of Clyde and North Sea herring,cod, and flounder larvae

Tolerance limits, at which 50% of larvae could survive high temperature and low salinity for 24 h, were determined for the yolk-sac larvae of Clyde and North Sea herring (Clupea harengus L.), cod (Gadus morhua L.) and flounder (Platichthys flesus L.) during early development and starvation. Clyde and North Sea herring, cod and flounder from hatching to the end of the yolk-sac stage, could withstand 21–23.5 °C, 20.5–23 °C, 15.5–18 °C and 21.5–24°C, respectively. The temperature tolerance was reduced by about 3.5–4 °C for Clyde herring and cod, 4–4.5 °C for North Sea herring and 8–8.5 °C for flounder when the larvae reached the point-of-no-return (PNR, when 50% of larvae, although still alive, are no longer strong enough to feed). The lowest salinity tolerance between hatching and the end of yolk-sac stage was 1–1.5‰ for Clyde and North Sea herring, 2–3‰ for cod and 0–1‰ for flounder. In no instance was there a loss of tolerance to low salinity during starvation. In fact, tolerance improved somewhat until the larvae became moribund. At hatching Clyde and North Sea herring larvae were negatively buoyant with a sinking rate of 0.35–0.4cm · s⁻¹ which steadily decreased until the larvae became moribund. Cod and flounder larvae, however, were positively buoyant at hatching but became progressively less buoyant and, by the end of the yolk-sac stage they were negatively buoyant with a sinking rate of 0.06–0.07 cm · s⁻¹. This sinking rate then decreased slightly until the PNR stage. The low salinity tolerance of all three species varied in a similar fashion to buoyancy.     

Effect of temperature on development rate and intrinsic rate of increase ofAphis gossypii reared on greenhouse cucumbers

Thermal requirements for development and life table statistics ofAphis gossypii Glover (Homoptera, Aphididae) were determined over a range of constant temperatures from 10 to 30°C. The lower development threshold and the sum of effective temperatures were 6.9°C and 90.1°C, respectively, for preimaginal development, and 5.8°C and 113.6°C from birth to the onset of reproduction. Mean total fecundity ranged from 36 larvae per female at 10°C to 76 larvae at 30°C. On a time scale of days, net reproductive rate (R _o ) increased with increasing temperature while generation time (T) decreased causing the intrinsic rate of increase (r _m ) to increase linearly from 0.115 to 0.465. On a day-degree scaler _m only varied from 0.019 to 0.028 because the growth ofR _o was compensated by an increase inT with increase in temperature. The nearly constantr _m in terms of day-degrees, over a wide range of temperatures, greatly simplifies the prediction of future population numbers ofA. gossypii.     

Observations on the morphology and infectivity for cattle of Babesia bovis parasites in unfed Boophilus microplus larvae after incubation at various temperatures

Dalgliesh R. J. and Stewart N. P. 1979. Observations on the morphology and infectivity for cattle of Babesia bovis parasites in unfed Boophilus microplus larvae after incubation at various temperatures. International Journal for Parasitology9: 115–120. The temperature of incubation of unfed Boophilus microplus larvae infected with Babesia bovis influenced the morphology and infectivity of the Babesia within the tick. Incubation at 37°C for 1–3 days stimulated the development of parasites morphologically similar to those usually observed in fed larvae harvested from cattle; similar forms appeared more slowly in larvae incubated at 31°C or 25°C. Extracts prepared from larvae after incubation at 37°C for 3–5 days or 30°C for 8 days were consistently infective for cattle. Prior storage of larvae at 14°C for up to 28 days enhanced the development of infectivity at 37°C; infectivity could still be produced after 65 days storage at 14°C but not after 76 days. Larvae released on a host transmitted B. bovis sooner if they had been incubated at 37°C for 4 days. It was concluded that the development of B. bovis to an infective stage in B. microplus is temperature dependent and does not require the stimulus of feeding by the host.     

Some pathogenic effects of a species ofPleistophora [Protozoa,Microsporida] forAgrotis exclamationis and other noctuids

Larvae of noctuids were inoculatedper os with spores of a species ofPleistophora isolated fromAgrotis exclamationis (L.). The mean median lethal dose for mainly 3rd instar larvae ofA. exclamationis was 1.38×10⁵ spores per larva accumulating 34 or 35 days after inoculation and the mean slope for the regression of mortality on dose was 0.82. Third instar larvae ofA. exclamationis andSpodoptera frugiperda (J.E. Smith) inoculated with 2.5×10⁴ spores gained weight quicker than uninfected ones until between 8 days (A. exclamationis) and 13 days (S. frugiperda) post-inoculation. Thereafter they grew slower than uninfected individuals. Correspondingly, the feeding rate of inoculated larvae ofA. exclamationis was greater than that of untreated ones until 14 days post-inoculation but thereafter was less. Larvae ofNoctua pronuba (L.)Phlogophora meticulosa L. andSpodoptera littoralis (Boisduval) but notAgrotis segetum (Denis & Schiffermüller) were also susceptible to infection. The species ofPleistophora considered here corresponds toP. schubergi noctuidae (Veremtchuk & Issi) in spore morphology, tissue specificity and host range, except that is was non-infective for the typehostA. segetum. It is probably insufficiently pathogenic for use in the biological control of noctuids.     

Biology ofAnastatus biproruli (Hym.: Eupelmidae) a parasitoid ofBiprorulus bibax (Hem.: Pentatomidae)

The effect of temperature on the rate of development ofAnastatus biproruli (Girault) was determined by rearing individuals on eggs ofBiprorulus bibax Breddin under a range of constant temperatures (17.5–40.0°C). Rate of development changed in a linear fashion from 17.5–35.0°C and the lower developmental threshold was estimated to be 12.8°C. An estimated 331.8 degree days were required for development. Survival of developing parasitoids was optimal (72–90%) between 25–35°C. Longevity of virgin adults ranged from 12–26 days at 22.5–35°C but was only 3.6 days at 37.5°C. Mated females produced a mean of 54.6 progeny during an average lifespan of 36.4 days at 30°C. Progeny were produced only during the first 21 days of adulthood but post-reproductive females still attacked and killed hosts. FemaleA. biproruli overwintered successfully, and were able to parasitise host eggs if direct sunlight was available.A. biproruli also overwintered as immature stages in host eggs. The biology ofA. biproruli is discussed with regard to its importance as a natural enemy ofB. bibax.     

Studies on Prolonged Storage of Beauveria bassiana Conidia: Effects of Temperature and Relative Humidity on Conidial Viability and Virulence against Chickpea Borer,Helicoverpa armigera

Unformulated conidia of Beauveria bassiana were stored at five different temperatures (0°, 10°, 20°, 30° and 40°C) at six different relative humidities (RH) (0, 33, 53, 75, 85 and 98%). Conidial viabilities and virulence against third instar larvae of Helicoverpa armigera were determined over a 24‐month period. Conidia survived longest at lower temperatures (0–20°C) and lower RH levels (0–53% RH). At higher temperatures (30–40°C) conidia did not survive. When the temperature was decreased from 30°C to 0°C, at nearly all RH levels the longevity of conidia increased. Conidia remained virulent for third instar larvae of H. armigera under favourable storage conditions for 24 months.     

Development of the bitterling,Paracheilognathus himantegus (Cyprinidae), with a note on minute tubercles on the skin surface

The development of the eggs and larvae and minute tubercles on the skin surface ofParacheilognathus himantegus larvae were observed. The egg began to hatch approximately 68 hours after insemination and the larvae reached the free-swimming stage 23 days after hatching at water temperature of 22±1°C. The larval development and minute tubercles on the skin surface of this species were similar to those ofAcheilognathus lanceolata, A. limbata, A. signifer andTanakia tanago. However, the shape of the ripe eggs ofP. himantegus differed from those of the four species. As regards the shape of eggs, there was a common characteristic amongP. himantegus, Rhodeus uyekii andA. limbata from Korea. As regards larval development,P. himantegus had two characters also found inRhodeus. These facts seem to suggest thatP. himantegus is closely related toA. lanceolata, A. limbata, A. signifer andT. tanago but is more specialized than these four species, except forA. limbata from Korea.     

Characteristic of parasitism of diamondback moth by two larval parasites

Laboratory and greenhouse studies were conducted to investigate the suitability of 2 hymenopterous parasites,Diadegma eucerophaga Horstmann andApanteles plutellae Kurdjumov for introduction to control diamondback moth (DBM),Plutella xylostella (L.), a destructive pest of crucifers in tropical to subtropical Southeast Asia. Parasitism byD. eucerophaga was high at temperature range of 15°C to 25°C and that ofA. plutellae, at 20°C to 35°C. Both parasites were active in searching for host and oviposited only during photophase. No parasitism was observed during darkness. WhereasA. plutellae could parasitize all instars of DBM larvae,D. eucerophaga parasitized only the first 3 instars and failed to parasitize the 4^th. Parasitism byD. eucerophaga was greater when DBM larvae were feeding on common cabbage (Brassica oleracea var.capitata L.), than on cauliflower (B. oleracea var.italica L.), broccoli (B. oleracea var.botrytis L.) or Chinese cabbage [Brassica campestris L. ssp.pekinensis (Lour) Olsson].A. plutellae parasitism was greater when DBM larvac were feeding on Chinese cabbage than on common cabbage, cauliflower or broccoli. Storage of pupae at 0°C and 4°C to 6°C for up to 2 weeks reduced emergence ofD. eucerophaga adults more than that ofA. plutellae. A non-selective insecticide, deltamethrin, was toxic to adults of both parasites but selective ones such asBacillus thuringiensis, teflubenzuron, and pirimicarb were not. Pupae were more tolerant than adults to insecticides. The insecticide-resistant Luchu strain and susceptible laboratory strain of DBM suffered an equal level of parasitism by both parasites.      

Development of the bitterling,Tanakia tanago (cyprinidae), with a note on minute tubercles on the skin surface

The development of eggs and larvae and minute tubercles on the skin surface in larvae ofTanakia tanago were observed. The eggs began to hatch approximately 52 hours after insemination and the larvae reached free-swimming stage 19 days after hatching at water temperature of 22±1°C. The egg and larval development and minute tubercles on the skin surface in larvae of this species were similar to those ofAcheilognathus lanceolata andA. limbata. However,T. tanago was distinguishable in egg and larval development fromA. lanceolata andA. limbata by the following characters: the perivitelline space was narrower, embryonic and larval development was faster, and minute tubercles on the skin surface of the anteriormost parts of the yolk sac, and of the body and head were hemispheric in shape. From these characters,T. tanago is considered to be more specialized thanA. lanceolata andA. limbata.