La Flavescence dorée est une maladie à phytoplasme incurable qui affecte la vigne. Cette pathologie est uniquement transmise de plante à plante par la cicadelle vectrice Scaphoideus titanus. Malgré l'importance de cette maladie, peu d'études ont été menées sur le vecteur. Ce travail de thèse vise à apporter des connaissances sur la biologie et l'écologie de S. titanus mais aussi de manière plus générale sur l'écologie des insectes phytophages. Trois grands thèmes ont été abordés dans ce mémoire : i) les relations entre l'insecte et son environnement à travers l'influence des températures hivernales sur les œufs en diapause, ii) les relations de l'insecte avec sa plante hôte en examinant l'attractivité de cette dernière et le comportement alimentaire du vecteur, iii) les interactions entre congénères par l'étude du comportement d'agrégation des larves. Les résultats obtenus indiquent que 1) des températures froides ne sont pas indispensables à la levée de diapause mais affectent la protandrie en régulant le sex-ratio opérationnel, 2) l'attractivité de la plante hôte est en grande partie liée à sa couleur et il existe des différences de comportement en fonction du stade de développement, 3) les mâles et les femelles n'ont pas le même comportement alimentaire, ce qui pourrait expliquer leur différentes capacités de vection, 4) l'agrégation des larves ne semble pas due à un recrutement actif de leur part et pourrait contribuer à un transfert horizontal de phytoplasme d'insecte à insecte via la plante. A terme, ces connaissances pourront contribuer à l'élaboration de pistes alternatives à la lutte chimique actuelle contre cette cicadelle dans le cadre d'une réduction des intrants. / The Flavescence dorée is an incurable phytoplasma disease of grapevine. This pathology is exclusively transmitted from plant to plant by the leafhopper vector Scaphoideus titanus. Despite the importance of this disease, few studies have been conducted on the vector. This thesis aims to provide knowledge on the biology and ecology of S. titanus but also more broadly on the ecology of phytophagous insects. Three major themes were addressed in this thesis: i) the relationships between the insect and its environment through the influence of winter temperatures on diapausing eggs, ii) the relationships of the insect with its host plant by examining the attractiveness of the plant and the feeding behaviour of the vector, iii) interactions between congeners in the study of the aggregative behaviour of larvae. Our results indicate that 1) cold temperatures are not essential to break the diapause but contribute to protandry through operational sex ratio regulation, 2) the attractiveness of the host plant is largely due to its colour and different instars exhibit differences in behavioural choices, 3) males and females have different feeding behaviour which may explain their different efficiencies at vection, 4) the aggregation of larvae did not seem to result from active recruitment and could contribute to horizontal insect to insect transfer of phytoplasma through the plant. Such knowledge can contribute to the development of alternative S. titanus pest management techniques.

Depuis les années 1990, on observe une accélération de la vitesse de propagation après établissementd’une grande partie des insectes introduits accidentellement en Europe. L’invasion fulgurante de lapyrale du buis, Cydalima perspectalis, est un exemple de ce phénomène. Observée pour la premièrefois en 2007 en Allemagne, elle s’est ensuite propagée jusqu’au Caucase en une décennie. Il a étérapidement supposé que le commerce de buis d’ornement entre les pays européens et la Chine auraitpermis son introduction tandis que l‘intensification des échanges commerciaux entre pays européensaurait facilité sa propagation à travers le continent. Cependant, certains traits biologiques de l’insecteont également pu favoriser son invasion. Pour mieux comprendre les facteurs impliqués dans soninvasion fulgurante, une approche pluridisciplinaire a été utilisée au cours de cette thèse. Tout d’abord,les routes d’invasion ont été appréhendées avec deux types de marqueurs génétiques. Puis, deux traitsbiologiques ayant pu favoriser son établissement et son expansion ont été étudiés : ses capacités devol et la diapause. Les résultats génétiques ont permis de cibler la partie côtière de la Chine commezone source probable, et suggèrent au moins deux introductions indépendantes depuis la Chine suivipar un phénomène tête de pont probablement lié au commerce d’ornement. Les capacités de volestimées sont élevées et expliqueraient la dispersion de l’insecte à l’échelle locale. De plus, la sortie dediapause influencée par la température a pu permettre l’adaptation du nombre de générations enfonction du climat annuel. Ces deux traits biologiques ont par conséquent dû favoriser son invasion àl’échelle locale mais il apparaît que les activités humaines, en particulier le commerce des plantesornementales, ont joué un rôle majeur dans son invasion fulgurante à large échelle en Europe et auCaucase. / Since 1990, recently established non-native insect species appear to be spreading across Europesignificantly faster than before. The box tree moth, Cydalima perspectalis, is a representative of such arapid spread. First observed in 2007, it spread across Europe to Asia Minor in a decade, damaging onBuxus plants in urban areas as well as in forests. The trade of ornamental box plants between Europeand China has been hypothesized as the invasion pathway of the insect while the trade betweenEuropean countries could explain its fast expansion. The biological traits of the moth may also havefavored its spread. To understand the factors involved in both its invasion and fast expansion, we useda multidisciplinary approach. First, the invasion routes were tentatively assessed using two geneticmarkers. Then, two biological traits, flight capacities and diapause, were studied because they may havepromote its spread. The genetic analyses clearly suggest an origin from Eastern China origin but alsocomplex invasion pathways. Two independent introduction events probably happened directly fromChina followed by bridgehead effects due to ornamental plant trade. The moth flight capacities areimportant but can only explain dispersal at local scale. Moreover, diapause termination was influencedby temperature, which may lead to an adapted number of generation depending on local climate. Thesebiological traits may have favored local invasions. However, our study suggests that human-mediateddispersal played a key role in the fast expansion of C. perspectalis across Europe and Asia Minor.

The Tardigrada are hydrophilous, segmented, molting micrometazoans that occupy a diversity of niches in freshwater, marine, and terrestrial habitats. A sister group of the arthropods, this phylum of bilaterally symmetrical lobopods, most less than 1 mm in length, have a hemocoel, a complete digestive tract, a dorsal gonad with one or two gonoducts, and a dorsal lobed brain with a ventral nerve cord and five ganglia. About 1000 species have been described based on the morphology of sclerified structures, especially the claws and buccal-pharyngeal apparatus. Reproduction occurs through fertilized or unfertilized eggs, with individuals being either gonochoric, unisexual, or hermaphroditic, and eggs are deposited either freely or within the shed exuvium. Parthenogenesis, very frequent in limnic and terrestrial tardigrades, allows them to colonize new territories by passive dispersal of a single individual. Quiescence (cryptobiosis: anhydrobiosis, anoxybiosis, cryobiosis, and osmobiosis) and diapause (encystment and resting eggs) occur during the tardigrade life history. Ecological parameters and global distribution patterns are poorly known or understood. Methods for collection, microscopy, and culturing have been developed.

In order to complete their life cycles, vertebrates require oxygen and water. However, environments are not always forgiving when it comes to constantly providing these basic needs for vertebrate life. The annual killifish Austrofundulus limnaeus is possibly the most well described extremophile vertebrate and its embryos have been shown to tolerate extremes in oxygen, salinity, and water availability. This phenotype is likely a result of the annual killifish life history, which includes periods of temporary habitat desiccation and oxygen deprivation, and requires the production of stress-tolerant embryos that depress metabolism in a state of suspended animation, known as diapause. Over the past several decades, the basic morphology and physiology of annual killifish development has become better characterized. However, there are still basic cellular processes that remain to be described in annual killifish such as A. limnaeus. Specifically, changes in DNA structure, expression, and copy number are known to have profound impacts on the phenotype and survival of an organism. Little is known as to how A. limnaeus maintains genome integrity during cell stress, nor how the A. limnaeus nuclear and mitochondrial genomes may have evolved under the unpredictable conditions in which A. limnaeus thrive. Early annual killifish embryonic development is also characterized by a complete dispersion and subsequent reaggregation of embryonic blastomeres prior to formation of the embryonic axis. This unusual period of early development may provide a functional adaptation that allows annual killifish embryos to survive these extreme conditions. The overall goals of this project were to (1) characterize the ability of A. limnaeus to tolerate and repair DNA damage through enzymatic and developmental mechanisms, (2) to determine possible consequences of mitochondrial DNA sequence and copy number on the metabolism of A. limnaeus, and (3) to establish a draft genome of A. limnaeus for future comparative genome studies. The results of this project show that embryos of A. limnaeus have an impressive ability to survive and reverse high doses of DNA damage induced by ultraviolet-C (UV-C) radiation, especially when allowed to recover under photoreactivating light. Surprisingly, embryos that survived irradiation during blastomere dispersion phases were able to develop normally. Characterization of gene expression during embryonic development for genes important for axis formation and cellular differentiation suggests that A. limnaeus embryos may delay axis formation until several days after epiboly is complete, thus allowing time for cells that become damaged to be replaced by surrounding pluripotent cells. This outcome would represent first case of a developmental buffering stage in a vertebrate. A. limnaeus embryos are also unique in their mitochondrial response to anoxia. Whereas in other species the amount of mitochondrial DNA (mtDNA) copy number fluctuates following extremes in oxygen availability, A. limnaeus embryonic mtDNA remains stable. Additionally, characterization of the fully sequenced A. limnaeus mitochondrial genome reveals possible evolutionary adaptations that may have facilitated dormancy and anoxia tolerance when compared to other species within the Order Cyprinodontiformes. The final chapter of this project characterizes the draft genome of A. limnaeus and I provide evidence suggesting that epigenetic DNA methylation that may be involved in regulating diapause.

Animals and plants in temperate regions must adapt their life cycle to pronounced seasonal variation. The research effort that has gone into studying these cyclical life history events, or phenological traits, has increased greatly in recent decades. As phenological traits are often correlated to temperature, they are relevant to study in terms of understanding the effect of short term environmental variation as well as long term climate change. Because of this, changes in phenology are the most obvious and among the most commonly reported responses to climate change. Moreover, phenological traits are important for fitness as they determine the biotic and abiotic environment an individual encounters. Fine-tuning of phenology allows for synchronisation at a local scale to mates, food resources and appropriate weather conditions. On a between-population scale, variation in phenology may reflect regional variation in climate. Such differences can not only give insights to life cycle adaptation, but also to how populations may respond to environmental change through time. This applies both on an ecological scale through phenotypic plasticity as well as an evolutionary scale through genetic adaptation. In this thesis I have used statistical and experimental methods to investigate both the larger geographical patterns as well as mechanisms of fine-tuning of phenology of several butterfly species. The main focus, however, is on the orange tip butterfly, Anthocharis cardamines, in Sweden and the United Kingdom. I show a contrasting effect of spring temperature and winter condition on spring phenology for three out of the five studied butterfly species. For A. cardamines there are population differences in traits responding to these environmental factors between and within Sweden and the UK that suggest adaptation to local environmental conditions. All populations show a strong negative plastic relationship between spring temperature and spring phenology, while the opposite is true for winter cold duration. Spring phenology is shifted earlier with increasing cold duration. The environmental variables show correlations, for example, during a warm year a short winter delays phenology while a warm spring speeds phenology up. Correlations between the environmental variables also occur through space, as the locations that have long winters also have cold springs. The combined effects of these two environmental variables cause a complex geographical pattern of phenology across the UK and Sweden. When predicting phenology with future climate change or interpreting larger geographical patterns one must therefore have a good enough understanding of how the phenology is controlled and take the relevant environmental factors in to account. In terms of the effect of phenological change, it should be discussed with regards to change in life cycle timing among interacting species. For example, the phenology of the host plants is important for A. cardamines fitness, and it is also the main determining factor for oviposition. In summary, this thesis shows that the broad geographical pattern of phenology of the butterflies is formed by counteracting environmental variables, but that there also are significant population differences that enable fine-tuning of phenology according to the seasonal progression and variation at the local scale. / <p>At the time of the doctoral defense, the following paper was unpublished and had a status as follows: Paper 4: Manuscript.</p>

L’implantation retardée ou diapause embryonnaire décrit l'arrêt ou le retardement pendant l'embryogenèse. Chez le vison, la diapause est corrélée avec une sécrétion pituitaire insuffisante de la prolactine, ayant pour résultat la différentiation incomplète du corpus luteum et réduction de la progestérone. Des études antérieures suggèrent que le blastocyste de vison en diapause demeure dans un état de quiescence ou se développe lentement. Pour élucider ceci, la réplication de l'ADN a été étudiée. Les résultats démontrent synthèse de l’ADN et prolifération cellulaire dans les embryons au stade de morula, avant la diapause et dans les blastocystes après la réactivation. La réplication de l'ADN a été également détectée dans des blastocystes en diapause et en diapause prolongée. L'implantation est considérée comme une interaction bidirectionnelle entre le blastocyste et l'utérus. Il a été montré que les prostaglandines sont importantes pour la vascularisation de l’utérus au moment de l’implantation et peuvent réactiver l'utérus des visons après la diapause. La concentration protéinique et la localisation de la phospholipase citosolique A2 (CPLA2) et de la cyclooxygenase 2 (COX2) ont été étudiées dans l'utérus de vison. L’expression de la CPLA2 et COX2 étaient sur-régulées au moment de l'implantation. Il est connu que la prolactine active les corpus luteum des visons. L'idée de un lien entre la prolactine et la voie de signalisation des prostaglandines a été testée en mesurant les récepteurs de prolactine. Les résultats montrent une augmentation de l’expression des récepteurs de prolactine à l'implantation suggérant que la prolactine pourrait activer la voie de prostaglandine à l'utérus par son propre récepteur. La conclusion, les embryons pendant la diapause ne sont pas arrêtées complètement et les protéines liées à la voie de prostaglandine sont implique dans la réactivation de l'utérus. / Delayed implantation or diapause describes arrest or retardation during embryogenesis. In mink, diapause is related to insufficient pituitary prolactin secretion, resulting in incomplete differentiation of the corpus luteum with reduced progesterone concentration. The mink blastocyst at diapause was believed to be totally quiescent or expanding at a low rate. To explore this, DNA replication was studied. Results showed synthesis of DNA, and thus cell proliferation at the morulae stage before diapause and at the blastocyst following activation. DNA replication was detected not only at diapause but also at extended diapause. Furthermore, implantation is considered as a two-way interaction between the blastocyst and the uterus. It has been shown that prostaglandins are important for vascularization of the uterus and products of the prostaglandin pathway could reactivate the mink uterus following diapause. Protein concentration and localization was studied for cytosolic phospholipase A2 (CPLA2) and cyclooxygenase 2 (COX2) in mink uterus. Expression of CPLA2 and COX2 was up regulated at implantation. It is know that prolactin is the factor that activates the mink corpus luteum. The idea of a link between prolactin and prostaglandin pathway was investigated by quantifying the prolactin receptors in the uterus. Results showed an increase of prolactin receptors at implantation suggesting that prolactin could activate the prostaglandin pathway at the uterus through its own receptor. In conclusion, embryos during diapause are not completely arrested, and proteins related to the prostaglandin pathway are implicated in reactivation of the uterus.

Os meliponíneos são abelhas que ocorrem nas regiões tropicais e subtropicais da Terra. Neste momento em que as mudanças globais atraem tanto interesse científico, consideramos oportuno o estudo detalhado da organização social de Plebeia remota, espécie de abelha que apresenta diapausa reprodutiva no outono e inverno. Embora outras pesquisas tenham sido realizadas nesta espécie, indicando principalmente os efeitos dos fatores abióticos sobre a atividade de vôo, a divisão de trabalho entre as operárias durante o forrageamento não tinha sido estudada, tanto no período de diapausa quanto no período em que ocorre oviposição pela rainha. A ausência da construção de células de cria durante alguns meses na colônia, e o reinício desta atividade em agosto, implicam em profundas alterações fisiológicas em um grupo de operárias e na rainha, aumentando a complexidade da divisão de trabalho e da organização social. Analisamos as atividades de operárias individualmente nos dois períodos, e tratamos do comportamento individual na construção do coletivo nas atividades de forrageamento. Uma das abordagens foi a ritmicidade das forrageiras nesses dois períodos. Também estudamos o efeito da temperatura e da umidade relativa no padrão de forrageamento. Além disso, outro processo de grande importância para a reprodução das colônias foi estudado, a enxameação. Tanto a biologia dos enxames quanto a ritmicidade da construção de células e do POP no início do estabelecimento das colônias recém-enxameadas foram estudadas. Também discutimos os fatores que podem estar envolvidos no crescimento inicial das colônias e seu estabelecimento. O forrageamento e a enxameação são processos muito importantes para o crescimento e reprodução das colônias, e a compreensão deles são importantes para o desenvolvimento de técnicas de manejo e de medidas para a promoção da conservação das abelhas. / The meliponine are bees that occur in the tropical and subtropical regions of the Earth. In this moment that the global changes attract so much scientific interest, we consider opportune the detailed study of the social organization of Plebeia remota, a bee species that present reproductive diapause in autumn and winter. Although other researches had been made in this species, indicating mainly the effects of abiotic factors on the flight activity, the division of labor among the workers during the foraging had not been studied either in the diapause period or in the queen oviposition period. The absence of brood cells construction during some months in the colony, and the restart of this activity in august, imply profound physiologic alterations in a group of workers and in the queen, increasing the complexity of the division of labor and social organization. We analyzed the workers activities individually in the two periods and considered the individual behavior in the construction of the collective on the foraging activities. One of the approaches was the rhythm of the foragers in these two periods. We also studied the effect of temperature and relative humidity on the foraging pattern. Furthermore, another process of great importance to the reproduction of the colonies was studied, the swarming. Either the biology of the swarms as the rhythm of the construction of brood cells and of the POP in the beginning of the establishment of the colonies was studied. We also discussed the factors that may have been involved in the initial growth of the colonies and their establishment. The foraging and the swarming are processes very important to the growth and the reproduction of the colonies, and their comprehension is important to the development of management techniques and measurements to the promotion of conservation of bees.

Os meliponíneos são abelhas que ocorrem nas regiões tropicais e subtropicais da Terra. Neste momento em que as mudanças globais atraem tanto interesse científico, consideramos oportuno o estudo detalhado da organização social de Plebeia remota, espécie de abelha que apresenta diapausa reprodutiva no outono e inverno. Embora outras pesquisas tenham sido realizadas nesta espécie, indicando principalmente os efeitos dos fatores abióticos sobre a atividade de vôo, a divisão de trabalho entre as operárias durante o forrageamento não tinha sido estudada, tanto no período de diapausa quanto no período em que ocorre oviposição pela rainha. A ausência da construção de células de cria durante alguns meses na colônia, e o reinício desta atividade em agosto, implicam em profundas alterações fisiológicas em um grupo de operárias e na rainha, aumentando a complexidade da divisão de trabalho e da organização social. Analisamos as atividades de operárias individualmente nos dois períodos, e tratamos do comportamento individual na construção do coletivo nas atividades de forrageamento. Uma das abordagens foi a ritmicidade das forrageiras nesses dois períodos. Também estudamos o efeito da temperatura e da umidade relativa no padrão de forrageamento. Além disso, outro processo de grande importância para a reprodução das colônias foi estudado, a enxameação. Tanto a biologia dos enxames quanto a ritmicidade da construção de células e do POP no início do estabelecimento das colônias recém-enxameadas foram estudadas. Também discutimos os fatores que podem estar envolvidos no crescimento inicial das colônias e seu estabelecimento. O forrageamento e a enxameação são processos muito importantes para o crescimento e reprodução das colônias, e a compreensão deles são importantes para o desenvolvimento de técnicas de manejo e de medidas para a promoção da conservação das abelhas. / The meliponine are bees that occur in the tropical and subtropical regions of the Earth. In this moment that the global changes attract so much scientific interest, we consider opportune the detailed study of the social organization of Plebeia remota, a bee species that present reproductive diapause in autumn and winter. Although other researches had been made in this species, indicating mainly the effects of abiotic factors on the flight activity, the division of labor among the workers during the foraging had not been studied either in the diapause period or in the queen oviposition period. The absence of brood cells construction during some months in the colony, and the restart of this activity in august, imply profound physiologic alterations in a group of workers and in the queen, increasing the complexity of the division of labor and social organization. We analyzed the workers activities individually in the two periods and considered the individual behavior in the construction of the collective on the foraging activities. One of the approaches was the rhythm of the foragers in these two periods. We also studied the effect of temperature and relative humidity on the foraging pattern. Furthermore, another process of great importance to the reproduction of the colonies was studied, the swarming. Either the biology of the swarms as the rhythm of the construction of brood cells and of the POP in the beginning of the establishment of the colonies was studied. We also discussed the factors that may have been involved in the initial growth of the colonies and their establishment. The foraging and the swarming are processes very important to the growth and the reproduction of the colonies, and their comprehension is important to the development of management techniques and measurements to the promotion of conservation of bees.

L’implantation retardée ou diapause embryonnaire décrit l'arrêt ou le retardement pendant l'embryogenèse. Chez le vison, la diapause est corrélée avec une sécrétion pituitaire insuffisante de la prolactine, ayant pour résultat la différentiation incomplète du corpus luteum et réduction de la progestérone. Des études antérieures suggèrent que le blastocyste de vison en diapause demeure dans un état de quiescence ou se développe lentement. Pour élucider ceci, la réplication de l'ADN a été étudiée. Les résultats démontrent synthèse de l’ADN et prolifération cellulaire dans les embryons au stade de morula, avant la diapause et dans les blastocystes après la réactivation. La réplication de l'ADN a été également détectée dans des blastocystes en diapause et en diapause prolongée. L'implantation est considérée comme une interaction bidirectionnelle entre le blastocyste et l'utérus. Il a été montré que les prostaglandines sont importantes pour la vascularisation de l’utérus au moment de l’implantation et peuvent réactiver l'utérus des visons après la diapause. La concentration protéinique et la localisation de la phospholipase citosolique A2 (CPLA2) et de la cyclooxygenase 2 (COX2) ont été étudiées dans l'utérus de vison. L’expression de la CPLA2 et COX2 étaient sur-régulées au moment de l'implantation. Il est connu que la prolactine active les corpus luteum des visons. L'idée de un lien entre la prolactine et la voie de signalisation des prostaglandines a été testée en mesurant les récepteurs de prolactine. Les résultats montrent une augmentation de l’expression des récepteurs de prolactine à l'implantation suggérant que la prolactine pourrait activer la voie de prostaglandine à l'utérus par son propre récepteur. La conclusion, les embryons pendant la diapause ne sont pas arrêtées complètement et les protéines liées à la voie de prostaglandine sont implique dans la réactivation de l'utérus. / Delayed implantation or diapause describes arrest or retardation during embryogenesis. In mink, diapause is related to insufficient pituitary prolactin secretion, resulting in incomplete differentiation of the corpus luteum with reduced progesterone concentration. The mink blastocyst at diapause was believed to be totally quiescent or expanding at a low rate. To explore this, DNA replication was studied. Results showed synthesis of DNA, and thus cell proliferation at the morulae stage before diapause and at the blastocyst following activation. DNA replication was detected not only at diapause but also at extended diapause. Furthermore, implantation is considered as a two-way interaction between the blastocyst and the uterus. It has been shown that prostaglandins are important for vascularization of the uterus and products of the prostaglandin pathway could reactivate the mink uterus following diapause. Protein concentration and localization was studied for cytosolic phospholipase A2 (CPLA2) and cyclooxygenase 2 (COX2) in mink uterus. Expression of CPLA2 and COX2 was up regulated at implantation. It is know that prolactin is the factor that activates the mink corpus luteum. The idea of a link between prolactin and prostaglandin pathway was investigated by quantifying the prolactin receptors in the uterus. Results showed an increase of prolactin receptors at implantation suggesting that prolactin could activate the prostaglandin pathway at the uterus through its own receptor. In conclusion, embryos during diapause are not completely arrested, and proteins related to the prostaglandin pathway are implicated in reactivation of the uterus.

Recently, the cold resistance becomes a common topic in scientific articles. If we focus on order Heteroptera, we can find a lot of studies dealing with the cold resistance or with the supercooling point (SCP) and also with the factors that influence them. According to the general rule, the acclimation at low temperatures increases the cold resistance. Therefore, the main goal of this thesis is to find out, whether the acclimation also increases the cold resistance of the water cricket Velia caprai Tamanini, 1947 (Veliidae), with the intention to measure SCP of the water crickets depending on the acclimation temperature and the length of exposure. Due to the high mortality (the 284 individuals remained alive of total 1300) and due to the technical difficulties in measuring of SCP (measured only 36 individuals) it was necessary to modify the original goal of the thesis. The new goal was to test the influence of acclimation (during one month vs. one week) and the influence of different temperatures (0°C, 5°C, 10°C) on SCP of V. caprai. The individuals used for this measurement were collected during the year 2010 and in March 2011. The results of this thesis did not confirm the hypothesis that the acclimation at low temperatures increases the cold resistance. The individuals who were acclimated at a higher temperature have the values of SCP demonstrably lower than the individuals who were acclimated at low temperatures. The individuals who overwinter for the first time have arguably higher cold resistance than the individuals who overwinter for the second time. Probable cause of these results is the fact that the water cricket V. caprai does not start with diapause in the fall but it overwinters in quiescence.