Reproductive behaviour of the glass frog Hyalinobatrachium cappellei (Anura: Centrolenidae) in the Southern Amazon

Frog species of the Centrolenidae family exhibit a high variation in reproductive behaviours. Herein we describe the reproductive, ecological, acoustic and behavioural features of Hyalinobatrachium cappellei in Southern Amazon. Two hypotheses were also evaluated: (1) are ecological characteristics of the reproductive site related to male size and quantity of eggs and clutches? And (2) do males with one or more existing clutches at reproductive sites accumulate more new clutches than those males initially without them? This study was conducted at two streams, with sampling at 1 km each in Southern Amazon. Species reproduction occurred during the wet season and was considered prolonged breeding. Although male vocalization occurred on both the upper and underside of leaves, clutches were most often deposited on the underside of leaves. Males emitted a courtship call upon female approach. The process of female approach until oviposition lasted approximately eight hours. After oviposition, females left the breeding site while males remained near the clutch, covering it with the belly in ventral brooding behaviour. Thus, egg attendance is reported for the first time for this species. No intraspecific or interspecific aggressive behaviour was recorded. There was no relationship between habitat characteristics of the reproductive sites and male size, number of eggs and clutches. However, after 10 days of monitoring, we found evidences that parental males had a higher number of clutches than those males initially without clutches. Similar to other glass frog species, the behavioural characteristics of H. cappellei make this species an excellent model to use in evaluating the cost-effectiveness of parental care and the importance of egg attendance for offspring survival. Therefore, these data contribute towards a better understanding of the complex phylogenetic and biogeographic relationships between centrolenid lineages.     

Oviposition site preference and reproductive ecology of Teratohyla midas (Anura: Centrolenidae) in the Colombian Amazon

ABSTRACT

The family Centrolenidae (Glass frogs) includes 157 species of anurans with a Neotropical distribution. Although glass frogs have been the subject of several studies encompassing various aspects of their biology, natural history data on their reproductive ecology is still scarce. One of the important aspects of reproductive ecology is the oviposition site since it can impact the reproductive success of a species. Herein, we provide detailed information on the reproductive ecology of a population of Teratohyla midas from the Colombian Amazon foothills and test whether this species shows a preference on substrate and height of oviposition site. We monitored a population using capture-mark-recapture of individuals identified by a photograph of their unique pattern of dorsal yellow spots. Wild-ID Software was used to corroborate if the dorsal pattern was different between individuals. Additionally, we monitored the presence of egg clutches, amplexus and oviposition behaviour. We observed 110 males and 5 females of T. midas, seen at an average height of 3.1 m ± 2.9 (0–17.8 m) from the ground. Likewise, we recorded the amplexus behaviour of two pairs for approximately 150 min. Once the oviposition happened, the male retreated, and the female covered the egg clutches for approximately 60 min. We observed 25 egg clutches with 60% being deposited in leaves of Selaginella sp and found at an average height of 2.9 ± 1.7 m (1.1–7.0 m). Thus, the selection of oviposition site was not random but mostly specific to one type of substrate. Furthermore, there was an overlap of the re-sampled median height distributions of active individuals and clutches, which suggests that oviposition does not occur at a preferred height within the vertical distribution of individuals. The information presented here is the first detailed record of reproductive ecology for this species and we hope to provide a basis to improve our knowledge on the biology of this species and other glass frogs.     

Mating behaviour,territoriality and natural history notes of Phyllomedusa ayeaye Lutz, 1966 (Hylidae: Phyllomedusinae) in south-eastern Brazil

The mating (male courtship, amplexus and oviposition) and male territorial behaviours of Phyllomedusa ayeaye are described from a high-altitude site in the state of Minas Gerais, south-eastern Brazil. Information related to male breeding success and territoriality, as well as egg clutch parameters, is provided, together with qualitative information regarding tadpole phenology and juvenile recruitment. In addition, estimations of sexual dimorphism and numbers of marked individuals are available through capture-mark-recapture sampling. Females bred with males within spatially clustered oviposition sites (broadleaf plants). In some cases, the amplectant couple actively searched for the oviposition site. Males defended territories from other males, employing both acoustic and physical interactions. Some males successfully maintained their calling sites over successive nights, and others seemed to switch among nearby sites during successive nights and tried to disrupt ongoing ovipositions. No significant relationship was found between physical attributes (snout vent length or body mass) of males with breeding success. Also, no influence of the number of nights a male was active in chorus and its breeding success was detected. Therefore, it is proposed that the mating system in P. ayeaye may be opportunistic. Additional information related to reproduction (egg clutch parameters and breeding behaviours) is also discussed for other species of the Phyllomedusa hypochondrialis group and compared with the results of the present study.     

On the natural history of the South American pepper frog,Leptodactylus labyrinthicus (Spix, 1824) (Anura: Leptodactylidae)

Some species of Leptodactylus of the L. pentadactylus group lay their eggs outside water but the tadpoles need to reach water to complete the larval phase; other species complete development in terrestrial nests. Here we present details of the reproduction of L. labyrinthicus in south‐eastern Brazil. The proportion of tadpoles and trophic eggs in aged egg clutches was determined, as well as the growth of the tadpoles while in the nest. The gut contents of tadpoles that were in egg clutches of frogs were analysed. Adult males did not differ from females in size and had hypertrophied forearms and an enlarged spine on the thumb. Reproduction was initiated with the first rains of August/September and extended to mid‐January. Calling and spawning occurred at permanent or temporary water bodies. The foam nests were built in excavated basins outside of, but close to the water. The male determined the place of the basin construction; after amplexus, the female completed the excavation. The amplexus was axillary. One female spent the day after spawning in the foam. The eggs were pale grey, the yolk averaging 2.3?mm in diameter. The mean number of eggs was 2101 per egg clutch. The number of tadpoles in individual nests varied between 0.05% and 11.40% in relation to the total laid eggs. The tadpoles entered water when rains flooded the basin. The tadpoles grew to 12 times the weight of an individual egg while in the nest; no nesting tadpole was beyond stage 25. The longest time we followed tadpoles in a nest was 25 days. Tadpoles were found preying upon eggs of three other frog species and upon conspecific eggs. Males fought by grasping each other in a belly‐to‐belly position; the powerful arms and the thumb spines represent weapons. Even though males can reach maturity in the season following birth, small size would prevent them from establishing their own territory. All the species of the L. pentadactylus group may build their foam nests within excavated basins. The basins may protect the eggs and embryos from cannibalistic tadpoles and may have an anti‐desiccation effect. In order to produce trophic eggs, the female may delay laying additional unfertilized eggs until after the male has abandoned the foam nest. Anuran eggs represent an important food item for tadpoles after they leave the nest.     

Reproductive ecology of a Tibetan frog Nanorana parkeri (Anura: Ranidae)

Endemic to the Tibetan Plateau, Nanorana parkeri figures among the highest altitude amphibians in the world. The present work, which was conducted in an alpine marsh at 4300 m altitude, is the first report of the species’ reproductive ecology. The breeding season, as indicated by the presence of amplectant pairs and fresh eggs in ponds, lasted from early Mary to early July. The ratios of adult male to female were 1.3 in hibernation ponds and 2.3 in spawning ones. Males were smaller than females. Amplexus was typically axillary (81.7%), with a few exceptional amplexus in which a male clasped another male or object rather than gravid females. Scramble competition occurred in 6.8% of the normal amplexus located. The body sizes of amplexed couples correlated positively. Deposited eggs were attached to submerged plants solitarily (60.5%) or in small clusters (39.5%), and the scattered arrangement of individual eggs could be a strategy to ensure embryonic development in oxygen-poor environments. Clutch size averaged 189 [standard deviation (SD) = 64], and correlated positively with female body size; egg size averaged 2.18 mm (SD = 0.10), independent of female body size; both traits within a clutch were negatively related with each other even controlling for female body size. Relatively few and large eggs laid by the Tibetan frogs, compared with those by lowland aquatic-spawning anuran species, should be adaptive to the harsh high-altitude conditions. Larvae hatched from eggs after 16.4 days on average (SD = 5.8). Froglets emerged between late July and early August, but some tadpoles were present in winter. These findings may help to understand the evolution of amphibians in extreme environments and to protect these remote species.     

Breeding behaviour of Bokermannohyla nanuzae (Anura: Hylidae) at an Atlantic Forest site in southeastern Brazil

We studied the breeding activities of Bokermannohyla nanuzae at an Atlantic Forest site in southeastern Brazil from August 2010 to April 2012. We described courtship behaviour, acoustic parameters of calls, egg clutch features and the egg-laying site, and characterized the reproductive mode. We recorded calling males and gravid females almost year-round, except for a few months in the dry season. We observed the interaction of five male–female pairs that exhibited stereotyped behavioural sequences including visual and tactile signals. We identified three different types of calls that we considered as the advertisement call and two types of courtship calls. Egg clutches were placed in rocky crevices with water or in streamside puddles, among rocks, that would be likely to be flooded as stream water level raises, which characterizes a reproductive mode different from the one previously recognized for this species. The breeding biology of B. nanuzae is complex, with elaborate courtship behaviour and signalling.     

Breeding behaviour and mating success of Phyllomedusa rohdei (Anura,Hylidae) in south‐eastern Brazil

Observations on the courtship behaviour, mating behaviour and mating success of a leaf‐frog, Phyllomedusa rohdei, population were conducted in a temporary flooded site of the Atlantic Forest, south‐eastern Brazil. We did not find any influence of size and body weight on male mating success across three scales: seasonal, nightly chorus and local groups (males found within 1?m of a mated pair). In addition, no evidence was found that female Phyllomedusa rohdei exhibit overt mate choice. The formation of mated pairs in P. rohdei occurred because males moved toward the females, rather than reverse. The only variable that was significantly related to male mating success was the number of nights of participation in chorus activity. Besides calling tactics, males showed active searching behaviour. There was no evidence of size‐ and mass‐assortative mating, and a high percentage of fertilization at all size and mass ratios of males and females was observed. Clutches in this species are produced in the same manner found in others members of Phyllomedusa; a purse‐like nest is made over water by folding a leaf around the egg mass (embryos and fluid‐filled capsules). Finally, we supply a list of several studies that examined the relationship between size or mass ratios of males and females in amplexus and its influence on fertilization rate, and show that size‐assortative mating in anurans is inconsistent (present in some species but not in other).     

‘On the rocks’: reproductive biology of the endemic toad Xanthophryne (Anura: Bufonidae) from the Western Ghats,India

Xanthophryne is a toad genus endemic to the northern Western Ghats of India, with two extant sister species – Xanthophryne koynayensis and Xanthophryne tigerina. Both species are local endemics and endangered. We studied reproductive biology of these toads and found that they are specialized to the lateritic rocky outcrops at mid-elevations in high rainfall areas. Xanthophryne toads have sporadic, multiple spawning bouts lasting 2–4 days during early monsoon. In this explosive breeding behaviour, we observed male toads to engage in ‘pelvic thrusts’, a unique and novel behaviour among anurans. Females oviposit in shallow pools in depressions of lateritic boulders where their tadpoles metamorphose. These ephemeral rocky pools have limited resources and they desiccate rapidly with a break in the rains. To mitigate the stochastic risk of desiccation and subsequent large-scale egg/tadpole mortality, females may disperse their reproductive investment spatially and temporally in multiple clutches, and tadpoles metamorphose rapidly. Here, we describe the amplexus, spawning and male advertisement call, and provide a comparative account of the life history traits of the two Xanthophryne species. These toads face numerous threats and are in need of urgent conservation action. These toads, seemingly well adapted to the isolated rocky outcrops, offer an excellent opportunity to understand endemism, mating systems, anuran ecology and behaviour.     

Male pheasant‐tailed jacanas commit infanticides to avoid cuckoldry when paternity of eggs is doubtful

Many hypotheses have been proposed to explain infanticide, killing the young or destruction of eggs by conspecific adults. We report cases of infanticide in the polyandrous pheasant‐tailed jacana (Hydrophasianus chirurgus) of Taiwan. In 2004, three males and one female bred at our study site. The female mated with the three males sequentially and laid clutches for all of them. Infanticidal behaviour was observed in all males and in all but one clutch, the first clutch of the breeding season. The infanticidal males threw out the first one or two eggs of a clutch, but then accepted replacement eggs and incubated normally. Because the intervals between copulation with different males were short (1–10 days) and sperm can remain viable within the female reproductive tract for at least 4–5 days, sperm competition among the males may have been intense, and the paternity of the first few eggs in each clutch was doubtful. We suggest that male pheasant‐tailed jacana may use infanticide as a strategy to avoid cuckoldry when the paternity of eggs is in doubt.     

Reproductive patterns of Trachycephalus venulosus (Laurenti, 1768) and Scinax fuscovarius (Lutz, 1925) from the Cerrado,Central Brazil

Animal mating systems are moulded by ecological factors, mainly the temporal patterns of reproduction. We report the reproductive period, use of microhabitats, clutch and egg sizes, reproductive efforts, occurrence of sexual size dimorphism, and the relation between body size and fecundity in the gravid females of Trachycephalus venulosus and Scinax fuscovarius at Serra da Bodoquena, Central Brazil. The reproduction of both species was correlated with the rainy season. Females of T. venulosus produced more eggs and their eggs were larger than those of S. fuscovarius. The females of both species were larger and heavier than males. The reproductive effort was measured as the ratio of gonad mass in relation to body mass. Females of both species presented similar reproductive efforts, but males of S. fuscovarius invested more in gonads compared to males of T. venulosus. Trachycephalus venulosus presented significant correlation between snout–vent length versus number of mature eggs and body mass versus number of mature eggs and ovarian mass. Scinax fuscovarius showed significant correlation between body mass versus ovarian mass. Trachycephalus venulosus is an explosive breeder, whereas S. fuscovarius has prolonged reproduction. These species are able to adapt to several types of habitat, due to behavioural and physiological plasticity.     

Reproductive biology and strategies of nine meloid beetles from Central Europe (Coleoptera: Meloidae)

The reproductive biology of meloid species from Central Europe is investigated by means of laboratory breeding experiments. They show that the total reproductive potential of meloids, with up to 40,000 eggs, largely exceeds prior data. Furthermore, the number of laid eggs strongly relates to the way by which the triungulins find their host. Clutch size correlates significantly with the size of the beetle, while egg size is independent of this. Both clutch size and egg size decrease with each oviposition. For some species, reproductive data are used to demonstrate how they have adapted to their habitat and to point out existing trade‐offs. Based on the obtained results and on an evaluation of the relevant literature, three types of reproductive strategy can be distinguished within the meloids: (1) very high reproductive rates of open field species with phoretic larvae; (2) average reproductive rates of forest populations of species with phoretic larvae and of species the larvae of which search the nests of their hosts actively; and (3) small to average reproductive rates of those species depositing their clutches in the immediate proximity of their hosts' nests.     

The reproductive strategies of clausiliid land snails from Northern Vietnam (Gastropoda: Stylommatophora)

Under laboratory conditions we scrutinized the reproduction and development of two clausiliid land snail species from northern Vietnam: Phaedusa paviei (Morlet, 1892) and Oospira vanbuensis (Bavay and Dautzenberg, 1899). These species, very similar in shell size and shape, feature different reproductive modes. The former is viviparous, giving birth to one or two neonates on a single occasion, with the number of developing embryos retained in the reproductive tract ranging from 1 to 11. The embryonic shells are 1.8–4.7 mm in height. The latter species is oviparous and produces eggs in clutches (four to six per clutch). Hatching occurs 18–21 days after egg-laying. Both species exhibit a similar growth pattern: the ultimate shell size is attained after 6.5–8.5 months and reproduction starts 5–7 months later. The distribution of viviparous reproduction among the Phaedusinae and its taxonomic importance are discussed.     

Breeding behaviour of the Omei tree frog Rhacophorus omeimontis (Anura: Rachophoridae) in a subtropical montane region

Breeding behaviour of the Omei tree frog, Rhacophorus omeimontis, was studied during the egg-laying period (mid-April to late June) in south-west China. The operational sex ratio of the breeding population was male-biased at 3.0?±?0.8 (mean?±?SD) on a nightly basis in Pond A and 4.3?±?1.1 in Pond B. Pairs with a male clasping a female occurred in ponds first; the amplectant pairs moved to a neighbouring plant and then released foam in leaves 14–370 cm above the pond's water surface. Immediately, other males came to join the pair, forming an amplexing group of 2 to 7 males (for all 45 breeding units observed). Among the males no direct physical competition for possession of the females was observed. Females produced foam which surrounded the subsequently-laid eggs. The whole process of oviposition lasted 45–65 minutes. After ovipositing, males left the female, the female folded and lined neighbouring leaves with more foam produced to surround the eggs further. This process took 15–45 minutes. The numbers of eggs in a clutch ranged from 457 to 837 and eggs were between 2.9 and 3.4 mm in diameter; those two measures were negatively correlated with each other. Eggs hatched after 12–17 days; the larval period lasted 63–78 days. Fertilization success was positively related to the number of males participating in matings.     

Life cycle and population structure of the terrestrial isopod Hemilepistus klugii (Brandt, 1833) (Isopoda: Oniscidea) in Iran

The life cycle and population structure of Hemilepistus klugii were studied in a population in Varamin, Iran. The population was sampled monthly (or fortnightly during the breeding season) from February 2008 to June 2009 and a total of 7015 individuals, comprising 1069 males, 1079 females and 4867 juveniles, were collected. As in other Hemilepistus species, five distinct phenophases, namely pair formation, gestation, hatching, growth and stationary, were recorded during the life cycle of H. klugii. The overall sex ratio was 1 : 1 but varied over time. Ovigerous females were observed only in April, indicating a seasonal and very short breeding period. With a short lifespan after breeding, females demonstrated true semelparity. The mean cephalothorax width for ovigerous/post-ovigerous females was higher in 2008 than in 2009. These females attained the largest size in the population throughout the year. The number of eggs per female ranged from 28 to 147 (mean ± SE, 78 ± 1.8). There was a positive correlation between female size and fecundity. Recruitment occurred in late April and resulted in the highest population density in this month, whereas the lowest densities were observed during November to January. Despite a high percentage of ovigerous females carrying undeveloped eggs (72.3%), intramarsupial mortality was low (3.5%).     

Breeding biology of the threatened Campo Miner Geositta poeciloptera (Aves: Scleruridae), a Neotropical grassland specialist

The Campo Miner is a threatened grassland passerine endemic to the South American Cerrado, whose life history is almost unknown. In this paper, we studied during three breeding seasons (2014 to 2016) the breeding biology of a colour banded population of the species found in the Upper Rio Grande Grasslands, south-eastern Brazil. We found 98 nests, 81 of which became active and were monitored. The Campo Miner breeds in frequently burnt-and-grazed natural grasslands, successfully nesting in highly disturbed sites, such as dirt banks along roads and even in mine pits. The species is socially monogamous and both parents build the nest, which is a cavity/with-tunnel/simple/platform type. The nest chamber is lined with a platform made of grass fragments, charcoal, hairs, and mammal faeces. The most common clutch size is three eggs (n = 66), with some nests containing one (n = 1), two (n = 12) or four eggs (n = 2). The egg is white and pyriform and the incubation, performed by both parents, lasts 17.5 days. Mean nestling period is 15.5 days, with both parents feeding the young. Breeding season lasted for about 125 days (August to December) and multiple breeding attempts in a single season were common, with a maximum of three attempts recorded. All species of Scleruridae built their nests inside cavities dug in the soil with an access tunnel to it, where they lay a small clutch (usually 2–3 white eggs), but no other species in the family has been studied in detail to date. Further studies are required to understand why a species apparently tolerant to anthropogenic impacts such as G. poeciloptera can be so rare, patchily distributed and threatened throughout its range.     

Natural history of the Lined Seedeater Sporophila lineola (Aves: Thraupidae) in southeastern Brazil

The Lined Seedeater (Sporophila lineola) is a migratory species that inhabits a variety of open habitats in South America. We studied the breeding biology and territorial behaviour of a colour-banded population of the species in the Universidade Federal de Viçosa, Campus Florestal (1–19,8808ºS, –44,4136ºW), during two breeding seasons (2014/2015 and 2015/2016), which spans from December to April. We monitored 74 nests of this species. The nest is a low cup supported between a fork. Nests are mainly built with grass stems and rootlets, with spider-web used to hold the material together and to bind the nest to its supporting plant. Females are solely responsible for nest building and incubation, which is synchronic, and also for feeding nestlings to a large extent. Males are responsible for defending the territory, which corresponds to a small portion of the home range restricted to the nest environs, and also feed the nestlings. Mean clutch size is two eggs (78% of nests monitored), with clutches of three (20.3%) and four (1.7%) eggs also observed. Eggs are whitish, covered with dark brown blotches and spots to a variable extent. Mean incubation period, considered as the period between the onset of incubation and hatching of the first egg, is 11 days. Nestling period, considered as the period between hatching of the first egg and fledging of the last young, is 10 days. During the first breeding season, the simple percentage of successful nests was 34.1%, while the Mayfield success was 29.8%, with slightly higher values observed during the second breeding season, with 39.4% and 35.7%, respectively. We recorded, for the first time, three cases of polygamy in the species. We also recorded breeding site fidelity for the first time in the species, with males returning to the same territory owned in a previous breeding season.     

Activity and reproduction in Megalobulimus paranaguensis (Gastropoda,Eupulmonata): implications for conservation in captivity for a South American land snail

ABSTRACT

In South America, Megalobulimus includes a number of threatened species, as well the largest land snails on the continent. The activity patterns and reproductive aspects of this group have not been documented. This work describes the daily and seasonal activity patterns and reproduction of M. paranaguensis. We maintained specimens in the laboratory for one year, and we quantified their behaviour for one hour at four different times of the day (0 h, 6 h, 12 h and 18 h) during three days in four months (August, September, April and May). The number of postures, hatching rate, time of hatching since oviposition and mortality rate among juveniles for each month were also quantified. Megalobulimus paranaguensis was more active in August, and had a egg laying peak one month after. Fifty-one eggs were laid by 12 captive individuals throughout the year, with a mean value of 4.25 eggs per individual. The hatching rate was 80.39%, and the time of hatching since oviposition was 56.7 ± 4.3 days. In two eggs, we observed the presence of twins. The mortality rate among juveniles was low (9.30%) indicating that rearing land snails in captivity has the potential to be an important and viable tool for the management and conservation of these organisms.     

Hatching phenology,life history and population dynamics of the Oriental clam shrimp Eulimnadia indocylindrova Durga Prasad and Simhachalam with notes on phenology patterns in the Spinicaudata

We studied the natural history, hatching phenology and egg bank composition of the Oriental spinicaudatan clam shrimp Eulimnadia indocylindrova Durga Prasad and Simhachalam using both field studies and ex situ sediment rehydration. Field observations revealed that hatching began very early (1–2 days) after inundation, and continued for about 5 days. Mature adults could be observed by 10 days, and they survived up to 16 days. The population showed a largely hermaphrodite-biased sex ratio (male:hermaphrodite 1:3) observed over three years, with a decrease in number of males throughout the hydroperiod. Both amphigenic and monogenic hermaphrodites were observed. The total lifetime fecundity recorded was about 300 eggs laid in multiple clutches. The egg bank composition showed a high proportion of intact eggs, indicative of predictable hydrations and low sediment adversity. Hatching began on the first day post-inundation for all successive cyclical hydration treatments, with peak hatching on days 2 and 3. Hatching rate was highest (57% of total hatching in successive cycles) for the first hydration, decreasing subsequently for the further hydrations. Hatching duration decreased with successive hydrations and was the longest (around 7 days) for the first hydration. Maximum hatching (93%) occurred in the first 10 days for the continuous hydration treatment. Overall, the total emergence of nauplii in successive hydrations was larger than that observed for the continuous hydration treatment, indicative of a risk-spreading strategy across hydroperiods. Early and concentrated naupliar emergence, along with decreased hatching durations for successive cycles, was observed for all the hydrations. A survey of literature revealed a general lack of data on hatching phenology of clam shrimps, particularly from tropical and sub-tropical regions. Based on the available data, it appears that hatching patterns, particularly high, early hatching fractions, are commonly observed in Spinicaudata species, and do not seem to differ much across biogeographical regions.     

Effective strategy of the overwintering of semiaquatic bugs: overwintering of Velia caprai (Heteroptera: Gerromorpha: Veliidae)

Overwintering of Velia caprai was studied during the winter months 2006–2007 in the area of South Bohemia (Czech Republic) by (1) field phenological observations and samples, (2) study of artificially established experimental cohorts in an unheated glasshouse, and (3) auxiliary experiments in a laboratory. Velia caprai has a unique way of overwintering within the gerromorphan bugs that is rare among insects generally; it can overwinter in both an adult and an egg stage. Most females reach sexual maturity before overwintering, mate with mature males, overwinter actively on the water surface and lay eggs during the winter. The dormant eggs can be induced to hatch by increasing temperatures. Winter oviposition can be interpreted as a sign of a risk‐spreading reproductive strategy. The overwintering of both adults and eggs provides a chance to adapt to changes in the local environmental conditions and could be an important determinant in broad variability of life‐history traits.