| Butterflies are important ornamental insects with great artistic,ornamental and economic value.Meanwhile,they also play an important role in scientific research and environmental instruction.In this paper,we first observed the comprehensive behaviors of Tirumala limniace which included flight,foraging,courtship,mating and oviposition.Then we explored the mechanisms of species recognition in T.limniace and intraspecific recognition in Cethosia cyane cyane.Furtherly,we studied the associative learning ability of pupae and adult of T.limniace.Aiming at deepening our understanding of butterflies communication mechanism in species and sexes,the relationship between evolutionary adaptation of butterflies and their environment were discussed in this study.Also we provide a theoretical basis for the construction of the butterfly flying landscape.The main conclusions are as follows:1.Adult behavior of Tirumala limniaceAdult activities including flight,foraging,courtship,mating,and oviposition were observed.We found that from a behavioral point of view,the adult stage of this butterfly could be divided into four stages: the precourtship adult stage(from the 1st to the 6th day after emergence),the nectar-feeding stage,which promotes egg development(from the 2nd to the 16 th day),courtship and mating(from the 5th to the 13 th day),and oviposition(from the 7th to the 15 th day).Adults emerged from pupae when temperatures were above 23.5°C and eclosion took only 1 min,typically followed by a small amount of flight on the first day.Flight activity peaked from the ninth to the thirteenth day after eclosion,and there were two daily peak flight times: 10:00~13:00 and 15:00~18:00.The peak of flower-visiting activity was from the eighth to the thirteenth day after eclosion,and there were two daily peak foraging times: 11:00~12:00 and 16:00~17:00.Courtship and mating took place on the fifth day after eclosion,while oviposition occurred the following day.Oviposition occurred over 8 d.The average life expectancy of males was 16.5 d,while that of females was 15 d.In conclusion,males spent 22.1% of its time flying,14.1% foraging,63.8% in courtship and mating.And females spent 30.8% of its time flying,10.1% foraging,57.1% in courtship and mating,and 2% ovipositing.Adult activities are mainly for the reproductive process,and the mating time which is directly related to reproduction is longer.The flight may be related to the maturation of the reproductive system,while flower-visiting provides nutrition and energy for the adult's activities.2.The characteristics of butterflies courtship behavior and the use of visual cues during courtshipWe analysisd males' and females' courtship behavior,the frequency of chasing models and the visual characteristics of T.limniace,Danaus genutia,and Papilio demoleus.Results showed that males always took the initiative and females were passive during courtship.Four kinds of courtship behavior were observed: male chasing male,male chasing female,female chasing male,and female chasing female.,It is generally believed that wing color plays an important role in the identification of the sexes during courtship.We suggest that the three butterfly species tested exhibit three different patterns of information utilization to distinguish between the sexes: only olfaction was used,D.genutia;olfaction was given priority over vision,T.limniace;vision was given priority over olfaction,P.demoleus.3.The role of visual and olfactory cues in species recognition in Tirumala limniace during courtshipIn addition to visual cues,olfactory cues(body volatiles)are also playing an important role in species recognition of butterflies during courtship.We first explored the olfactory cues used in the identification of conspecifics.A total of 15 volatiles were detected in the bodies of females and 18 volatiles in males.Among them,there were 14 common volatiles.The larger proportion of common volatiles were ?-ocimene and ?-pinene,and both of them were derived from body volatiles rather than sex pheromones.EAG and behavioral experiments showed that ?-ocimene is an important compound for species recognition in T.limniace,while ?-pinene does not participate in the recognition process.The role of visual and olfactory cues in the identification of conspecifics showed that this butterfly could use both visual and olfactory cues,but olfaction was given priority over vision.4.The role of visual and olfactory cues in sex recognition in Cethosia cyane cyane with sexually dimorphic wing coloration and pheromone characteristicswe explored the roles of visual and olfactory cues in sex recognition during courtship.The attractant experiments of winged models and print paper models showed that males could distinguish the gender of conspecifics using visual cues alone.Behavioral assays showed that males visually recognized females not by wing size,but by their sexually specific wing color.The movement pattern of the model also exerted some influence on male courtship chasing behavior.A total of 21 volatiles were detected in the bodies of adults,but only cedrol played a role in the process of male recognition of females at close range.Therefore,males rely on both visual and olfactory cues to distinguish females during courtship.Visual cues play a major role in attracting males at the beginning of the courtship chase,while olfactory cues play a role in accurately identifying partners at close range.Three kinds of models,dissected natural wings(? and ?),printed paper wings attached(? and ?),and live adults(? and ?)were used to investigate the roles of visual and olfactory cues in sex recognition in the female C.cyane cyane.Our results showed that female C.cyane cyane use both visual and olfactory cues to distinguish the gender of conspecifics during courtship.Females are more likely to prefer males with the composite,multimodal display.5.Associative learning of Tirumala limniaceThe associative learning ability in pupae and adult stages of T.limniace,and the differences in the learning ability between males and females were investigated.We first found that T.limniace had the learning and memory ability in pupae stage,and this experience could affect adult foraging choices after eclosion.In the adult stage of T.limniace,the ability of associative ?-pinene with the food resources was strong.The learning efficiency was related to the frequency of training.The more training,learning efficiency was not necessarily higher,and the learning efficiency was better for 3 to 4 days.We found that the alteration of the olfactory preference during conditioning was not induced by physiological maturation,but the result of learning and training.We also found that the learning ability for ?-pinene and ethyl acetate in males was better than females,contrary to the generally considered female learning efficiency was higher than male.This may be caused by the differences between males and females in the demand for nectar,in learning ability of the color and in antennal sensilla.6.Releasing and trapping adults and butterfly flying landscape constructionThe role of color and odor in attracting adults during foraging was explored,and we designed a set of lure equipment to attract butterflies after releasing the adults.The effective equipment included the appropriate color of artificial flowers(white,yellow and red),attractant(?-pinene)and lights(seven-colors lamp).According to the releasing environment,the utility device could be made into the curtain flower rattan and the ringed flower rattan.The feeding method was using ?-pinene for learning and training for 3 days in adults stage.Better way of releasing butterflies was divided into several times,and the number of releasing was 200 adults for each time.In the wild,the highest attraction rate of the curtain flower rattan to T.limniace and D.chrysippus was 59% and 84%,and the ringed flower rattan was 71.1% and 69.5%,respectively,indicating that the luring equipment has good effect on butterfly attraction,and achieves the purpose of constructing the natural landscape of the butterfly flying to a certain extent. |
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