Female-extended control over their reproductive investment: the role of early mating interactions on oocyte maturation in the terrestrial crustacean Armadillidium vulgare (Latreille, 1804)

Abstract

In the terrestrial crustacean Armadillidium vulgare, the prolonged presence of males along with females is known to boost female reproductive physiology, and the so-called ‘male-effect’ was best characterized by a significant shortening of the pre-parturial intermoult (PPI) during which oocyte maturation spontaneously takes place. Continuous presence of a male over that period can speed up vitellogenesis and the moulting cycle, so to reduce female PPI by 15–20 days (shortening of 30–40%, in comparison to females reared with other females or in isolation, respectively). In the last investigation on the subject, we revealed that sexual interactions may start much sooner than previously thought in this species, and suggested that the observed ‘male-effect’ may likely result from early mating stimulations. Here we tested the specific effect of controlled mating interactions (one or two mating events at different times) on female PPI. We revealed that male presence for the time (about 2 h) that allowed a single mating to occur was enough to reduce the female intermoult by about 10 days (shortening of 19%, in comparison to females reared in similar conditions but in the absence of a mating interaction). Moreover, results indicate that the earlier and longer the copulations, the stronger the ‘male-effect’. Altogether, the data support the conclusion that A. vulgare females are adjust their reproductive physiology according to the presence/absence and the intensity of male mating stimuli. We place the findings in a broader ecological context, revise the so far prevailing view on the ‘male-effect’, and stress its possible significance in relation to the occurrence of feminising bacteria in this and other terrestrial isopod species.     

Life cycle and population dynamic of Armadillidium pelagicum Arcangeli, 1955 (Isopoda, Oniscidea) at Aouina

Armadillidium pelagicum Arcangeli, 1955 is a terrestrial isopod endemic to the circum-Sicilian islands and the North of Tunisia. The life cycle and the population structure of this species were studied on a natural population at Aouina, in the surroundings of Tunis, over 16 months from, January 2000 to April 2001, by monthly or semi-monthly samplings. The ovigerous females were present from March/April to the end of August and absent from September to February/March. These observations indicate that A. pelagicum at Aouina has a seasonal reproduction, followed by a sexual rest. The recruitment period is spread from April/May to mid-September. The fecundity, estimated by the number of eggs in the marsupium of ovigerous females, exhibited a great variability, which is related to the weight of these females. The sex ratio underwent fluctuations throughout the sampling period. It was female-biased in most samplings. Mass frequency distribution was analysed and nine cohorts were identified during the sampling period. The field growth rates are high in the first life phase, decrease during winter and increase during spring. The characteristics of the life cycle of A. pelagicum at Aouina may be summarized as follows: (i) Semi-annual species, since females appear to produce up to five broods per year, (ii) iteroparous females, since females seem to reproduce twice or more in life; (iii) bivoltine life cycle, since the population produces two generations per year; (iv) variability of cohorts' life span.     

Post-parturial reproduction in terrestrial isopods: a partial review

This review deals with some aspects of terrestrial isopod reproduction including breeding season, breeding periods, patterns and strategies, parturition, number and size of broods and mancae. Reproductive period is described by the exact dates (i.e. season) marsupial females were collected in the field, and the duration of the breeding period. The information for both aspects was generally obtained through field work by observing marsupial females. Two reproductive patterns are used by terrestrial isopods, either a discrete (i.e. seasonal) or a continuous (i.e. non-seasonal) pattern. The discrete pattern can be either annual when females breed once a year, or they can breed biannually or multi-annually (two or more times during the year). This conclusion is based generally on observing marsupial females in the field. Therefore, this information does not apply to the reproductive pattern of the individual female since a female may use either or both patterns, i.e. seasonal and continuous. Only by raising individual females singly (with a male) can this point be clarified. This way, the breeding strategy of the same individual female can be studied. The subject is discussed and reviewed based on my research data.     

Biology and population dynamics of Porcellionides sexfasciatus (Crustacea,Isopoda, Oniscidea)

The biology and the population dynamics of Porcellionides sexfasciatus Budde-Lund (1879) were studied on a field and carried out at Garat Naam (Kasserine, Tunisia) from July 1996 to June 1998. The reproduction exhibited a seasonal pattern extending from February/March to October/November. The juveniles appeared in the population from April to November. Size frequency distributions were analysed and 14 cohorts were recognised during the sampling period. Six cohorts were identified at the first sampling and eight new ones on the other samplings. Among these latter cohorts, three were tracked till they disappeared. Minimum average length of new cohorts ranged from 3.07 +/- 0.35 mm to 3.47 +/- 0.2 mm. Maximum average length of cohorts was 10.42 mm. P. sexfasciatus is a semi-annual species (females producing two or three broods per year), with iteroparous and amphogenes females (females reproducing twice or more in their life and producing both males and females), and bivoltine life cycle (two generations per year). The females are able to produce two broods, in the laboratory, without new mating. Fecundity and fertility, corresponding to the number of eggs or embryos per brood, appeared positively correlated with females' size. Although oscillating throughout the year, the sex ratio was often in equilibrium.     

栎旋木柄天牛生殖行为研究

为了阐明栎旋木柄天牛Aphrodisium sauteri Matsushita生殖行为的方式和特点,寻找有效的防治手段,本试验观察了栎旋木柄天牛的生殖行为,并在林间尝试了成虫诱集。结果表明:雌雄成虫之间表现出较强的吸引作用,近距离时雌雄虫可相互吸引,而远距离时雌虫引诱力更强。室内试验中求偶时间会随着成虫的衰弱而增加。成虫交配主要发生于白天。羽化出孔后1～14日龄雌虫和2～13日龄雄虫均可多次交配。成虫一次完整交配平均需时70.12min,交配后保护平均仅用时3.86min。一次完整交配过程中,不同日龄成虫的交配历时、交配间隔历时及交配次数都存在差异,各日龄成虫的交配间隔历时均大于交配历时。交配或产卵经历对雄虫交配行为的影响明显大于雌虫。雌虫产卵前期平均为30.71h。雌虫每产1粒卵平均需时99.58s,并随日龄增长显著加长。每头雌虫平均每日产卵量和总产卵量分别为7.89和26.20粒。雌虫多在白天产卵,用产卵器触探树皮表面寻找合适的产卵部位。卵主要产于树皮裂缝和枝条疤痕内,一般每次产卵1粒,卵表面无覆盖物。     

Mate guarding behaviour in the supralittoral isopod,Ligia dentipes (Oniscidea) from the Andaman and Nicobar Islands

Precopulatory mate guarding is a characteristic feature in the mating behaviour of many Malacostraca, and a necessary prerequisite for those species in which female receptivity for males is restricted to a short period of time after the pubertal/reproductive moult. This study deals with the pre-mate guarding behaviour of the semi-terrestrial isopod Ligia dentipes living in the crevices of coral boulders and rocks in the supralittoral region of the Andaman Islands. As in other isopods, moulting in L. dentipes is biphasic, in which the posterior body part invariably moults first. The guarding male aids the female partner in the removal of the moulted exoskeleton. Mating occurs immediately after the posterior body exuviates. The male leaves the female after copulation and goes in search of another receptive female, demonstrating a polygamous mating system in these isopods. The mated females also re-mate with several other males without mate guarding. Females that had mated several times produced more young, compared to females mated only once in the laboratory. Female receptivity ceases following moulting of the anterior half. Intrasexual encounters among males lead to the large males acquiring receptive females. This study reveals interesting deviations from the general pattern of mate guarding already reported in other isopods and decapods. The evolutionary and ecological significances of mate guarding, intrasexual and intersexual conflicts, found in these semi-terrestrial isopods, are discussed.     

Mineral in skeletal elements of the terrestrial crustacean Porcellio scaber: SRμCT of function related distribution and changes during the moult cycle

Terrestrial isopods moult first the posterior and then the anterior half of the body, allowing for storage and recycling of CaCO₃. We used synchrotron-radiation microtomography to estimate mineral content within skeletal segments in sequential moulting stages of Porcellio scaber. The results suggest that all examined cuticular segments contribute to storage and recycling, however, to varying extents. The mineral within the hepatopancreas after moult suggests an uptake of mineral from the ingested exuviae. The total maximum loss of mineral was 46% for the anterior and 43% for the posterior cuticle. The time course of resorption of mineral and mineralisation of the new cuticle suggests storage and recycling of mineral in the posterior and anterior cuticle. The mineral in the anterior pereiopods decreases by 25% only. P. scaber has long legs and can run fast; therefore, a less mineralised and thus lightweight cuticle in pereiopods likely serves to lower energy consumption during escape behaviour. Differential demineralisation occurs in the head cuticle, in which the cornea of the complex eyes remains completely mineralised. The partes incisivae of the mandibles are mineralised before the old cuticle is demineralised and shed. Probably, this enables the animal to ingest the old exuviae after each half moult.     

Sexual differences in moult–breeding overlap and female reproductive costs in pied flycatchers, Ficedula hypoleuca

1. In this study I show that a sexual difference in timing of the post-nuptial moult frequently occurs in a sub-arctic population of the pied flycatcher.
2. Most pairs started to moult after fledging of their young, but an overlap between moult and nestling feeding was more common among males than females. This sexual difference in moult–breeding overlap increased as the season progressed.
3. Females with moult–breeding overlap laid smaller clutches than non-moulting females. In addition to many other factors explaining the seasonal decline in clutch size that has been found for many bird species, it is possible that females adjust their clutch size according to their own risk of having to start moulting while still feeding the nestlings.
4. Nearly 24% of the females were deserted by their mate before the young fledged. Desertion imposed no fitness costs to males in terms of fledgling number or quality, suggesting that their females managed to adjust their care for the loss of male care.
5. Deserted females started moulting later than aided females, which may be a result of their increased reproductive investment.
6. Deserted females and females aided by moulting males had lower survival rate than females aided by non-moulting males.
7. These findings suggests that delayed moult may be one mechanism causing inter-annual reproductive costs in birds, and the relationship between a sexual difference in timing of moult and its fitness consequences may be widespread among passerine birds.     

Female genitalia,reproductive cycle and sperm storage in Armadillidium vulgare (Crustacea,Isopoda, Oniscidea)

Light and electron microscopic studies that we have published in the past have reported many aspects of the reproductive process in Armadillidium vulgare with particular emphasis on the reconstruction of female genitalia. Together this body of work provides an almost complete, albeit fragmented picture of these processes and include many data on sperm storage and sperm translocation. Females of A. vulgare have a pair of cuticular genitalia in the lumen of the oviduct. For insemination, these genitalia can receive the copulatory organs of males formed by the elongated tips of the first two pleon endopods. During transitions between reproductive (parturial) and non-reproductive (normal, non-parturial) moult cycles the genitalia undergo intriguing structural changes resulting in two types of genitalia. Throughout their reproductive lifetime, either type is reconstructed after each moult depending on the reproductive phase of the female. In this review, we integrate the events that occur during a reproductive cycle with particular emphasis on the genitalia reconstruction and sperm storage, and discuss functional aspects of the genitalia. Thereby, we provide a case model that can be useful for further studies on genitalia diversity and female reproductive strategies in terrestrial isopods.     

Reproductive costs of mating with a sibling male: sperm depletion and multiple mating in Cephalonomia hyalinipennis

Polygynous parasitoid males may be limited by the amount of sperm they can transmit to females, which in turn may become sperm limited. In this study, I tested the effect of male mating history on copula duration, female fecundity, and offspring sex ratio, and the likelihood that females will have multiple mates, in the gregarious parasitoid Cephalonomia hyalinipennis Ashmead (Hymenoptera: Bethylidae: Epyrinae), a likely candidate for sperm depletion due to its local mate competition system. Males were eager to mate with the seven females presented in rapid succession. Copula duration did not differ with male mating history, but latency before a first mating was significantly longer than before consecutive matings. Male mating history had no bearing on female fecundity (number of offspring), but significantly influenced offspring sex ratio. The last female to mate with a given male produced significantly more male offspring than the first one, and eventually became sperm depleted. In contrast, the offspring sex ratio of first‐mated females was female biased, denoting a high degree of sex allocation control. Once‐mated females, whether sperm‐depleted or not, accepted a second mating after a period of oviposition. Sperm‐depleted females resumed production of fertilized eggs after a second mating. Young, recently mated females also accepted a second mating, but extended in‐copula courtship was observed. Carrying out multiple matings in this species thus seems to reduce the cost of being constrained to produce only haploid males after accepting copulation with a sperm‐depleted male. I discuss the reproductive fitness costs that females experience when mating solely with their sibling males and the reproductive fitness gain of males that persist in mating, even when almost sperm‐depleted. Behavioural observations support the hypothesis that females monitor their sperm stock. It is concluded that C. hyalinipennis is a species with a partial local mating system.     

Adaptive strategies of overwintering adults: Reproductive diapause and mating behavior in a grasshopper,Stenocatantops splendens (Orthoptera: Catantopidae)

Abstract To understand the adaptive strategies of the overwintering adults of Stenocatantops splendens, the mechanism of maintenance and termination of the reproductive diapause, the variation in mortality between overwintering females and males, and the mating strategy of the males were investigated. The results indicated that the adult reproductive diapause in natural conditions was mainly regulated by photoperiod in the fall – long photoperiods promoted reproductive development and short photoperiods maintained reproductive diapause, and the sensitivity of the overwintering adults to photoperiod was over before the end of the winter. When transferred from natural conditions to controlled laboratory conditions on dates from September through February, pre‐oviposition became increasingly shorter with increasingly deferred transfer dates regardless of photoperiod conditions. The adults treated with low temperature for 30 days in September through November had significantly shorter pre‐oviposition, suggesting that low temperatures in winter had an important role in the termination of reproductive diapause. The female had a significantly lower supercooling point than the male, which was related to their lower mortality after winter. In addition, observations of wild populations of the species indicated that mating behavior prior to winter and the duration of pre‐mating period were not affected by photoperiod; mating and sperm transfer were mostly completed by November. Compared with females only mating before winter, females mating in the spring had shorter life span, longer pre‐oviposition, lower hatching rate and laid fewer egg pods while showing no significant difference with regard to ovipositional interval, per pod number of eggs and nymph dry weight.     

Morphology and evolution of the respiratory apparatus in the family Eubelidae (Crustacea,Isopoda, Oniscidea)

The morphology of the respiratory apparatus in the pleopodal lungs of the family Eubelidae was investigated. The family is a monophyletic group including more than 240 species in 53 genera (three of which are nomina dubia), mostly distributed in the Afrotropical Region (tropical Africa and Arabian Peninsula). In all the Eubelidae, except for the monospecific genus Parelumoides and two species of the genus Elumoides, the exopods of pleopods have lungs. All the pulmonary morphologies present in the entire suborder Oniscidea are found: 1) uncovered lungs, composed of a pleated respiratory surface, directly exposed to the air (Atracheodillo-type) or partially enclosed within the appendage (Synarmadilloides-type); 2) covered lungs with several spiracles and respiratory trees, housed within the appendages, with spiracles surrounded by a specialized, nonrespiratory, structure (perispiracular area) (Eubelum- and Somaloniscus-types); 3) covered lungs with only one spiracle, with or without perispiracular area, and one respiratory tree (Aethiopopactes- and Periscyphis-types), which in taxa with Periscyphis-type lung crosses the insertion of the appendage and penetrates into the pleon with bundles of respiratory tubules. The evolution of the various types of lungs is discussed. It is concluded that the two main evolutionary lines, i.e., uncovered lungs and covered lungs, originated independently from an ancestral respiratory structure-the semilunar area. A first mechanism of development of the semilunar area by folding of its surface produced the Atracheodillo-type (all folds coplanar with the surface of the exopod) and Synarmadilloides-type (folds partly coplanar and partly intraflexed inside the exopod) uncovered lungs. A second mechanism of development by tubular invagination of the cuticle of the semilunar area produced the polyspiracular Eubelum-type lungs (numerous arborescent invaginations) and the monospiracular Aethiopopactes-type lungs (only one arborescent invagination), probably passing through a common intermediate pattern. From the common pattern, both the poly- and monospiracular types would have inherited the characteristic concave cell arrangement of the perispiracular area. The Somaloniscus-type and Periscyphis-type lungs are forms specialized for arid environments, directly derived from the Eubelum-type and Aethiopopactes-type, respectively.     

Effect of age on reproductive attributes of an aphidophagous ladybird, Cheilomenes sexmaculata

The effect of both male and female age was investigated on certain reproductive attributes, viz. mating incidence, mating duration, fecundity, percent egg viability, ratio of reproductive and non‐reproductive periods and reproductive rate, of an aphidophagous ladybird, Cheilomenes sexmaculata (Fabricius). Females started mating at the age of 8 hours post‐emergence (PE) and males at the age of 2 days PE. Mating in the laboratory was a male‐dominated phenomenon. The mating duration and reproductive rate of 10‐day‐old females when mated with males of varying ages increased up to the male age of 60 days, and thereafter decreased, whereas, fecundity, egg viability and ratio of reproductive and non‐reproductive periods increased up to the male age of 50 days, and thereafter declined. However, when females of varying ages were mated with 10‐day‐old males, fecundity and reproductive rate increased up to 40 days of female age, respectively, then decreased. The ratio of reproductive and non‐reproductive periods increased with increasing age of females. Mating age for optimal reproductive output was 10J50‐day‐old males and NE to 40‐day‐old females. Reproductive cessation in males was recorded after 50 days PE, whereas in females at the age of 40 days PE. Higher mating durations lead to elevated reproductive rates. Delay in the reproductive phase was positively correlated with longevity. The results of this study may aid mass multiplication of this ladybird by identifying and promoting usage of adults of optimal age. Our results also enhance our understanding of the effect of age on reproductive attributes in ladybirds.     

Seasonal reproduction of Porcellio variabilis Lucas, 1846

Porcellio variabilis is common and very widespread in Tunisia. Its breeding period lasts from February to October with a cessation during the months of November, December and January. The recruitment of young takes place principally between April and July. Virgin and mated females reared with natural photoperiod in the laboratory show that the initiation of reproduction does not depend on mating; however, it is accelerated when the male is present, the preparturial moult being shorter in females reared in the presence of a male.     

Mate choice, sexual imprinting, and speciation: a test of a one-allele isolating mechanism in sympatric sticklebacks

One-allele isolating mechanisms should make the evolution of reproductive isolation between potentially hybridizing taxa easier than two-allele mechanisms, but the generality of one-allele mechanisms in nature has yet to be established. A potentially important one-allele mechanism is sexual imprinting, where the mate preferences of individuals are based on the phenotype of their parents. Here I test the possibility that sexual imprinting promotes reproductive isolation using sympatric species of threespine sticklebacks (Gasterosteus aculeatus). Sympatric species of sticklebacks consist of large benthic species and small limnetic species that are reproductively isolated and adapted to feeding in different environments. I fostered families of F1 hybrids between the species to males of both species. Preferences of these fostered females for males of either type revealed little or no effect of sexual imprinting on assortative mating. However, F1 females showed preferences for males that were similar to themselves in length, suggesting that size-assortative mating may be more important than sexual imprinting for promoting reproductive isolation between species pairs of threespine sticklebacks.     

Age-specific mating and reproductive senescence in the seven-spotted ladybird, Coccinella septempunctata

Abstract:  Adults of the seven-spotted ladybird, Coccinella septempunctata Linnaeus (Col., Coccinellidae) were paired for mating from very young to old age (1–50 days) to record the willingness to mate, attainment of sexual maturity and onset of reproductive senescence in both the sexes. Mating commenced after 4 and 2 days of emergence of male and female, respectively, and 100% mating was achieved at the young age in both cases (10 days). Willingness to mate decreased with increase in the age from 40 to 50 days of both the sexes. Ladybird exhibited protandry. Mating duration, fecundity and per cent viability of eggs of middle-aged males (20–30 days old) and females (20 days old) were the highest. Mating duration and per cent viability of eggs were male age dependent, whereas pre-oviposition and oviposition periods were mating stimulus dependent. Oviposition period and fecundity were female age-dependent responses. Fecundity was highest when 20-day-old female and 30-day-old male were paired. Onset of reproductive senescence started at the age of 30 days in males and 20 days in females. The present study confirms the effect of ageing on male and female C. septempunctata and supports the Hansen and Price [J. Evol. Biol. (1995) vol. 8, pp. 759–778] model that females mating with young and middle-aged males yield optimal quality progeny.     

Signaling of Reproductive Status in Captive Female Golden-Headed Lion Tamarins (Leontopithecus chrysomelas)

In species in which paternal care has an important impact on the offspring's fitness, concealment of reproductive status has been proposed as a strategy employed by females to prevent males from practicing desertion and polygamy, which would then lead to monogamous or polyandrous mating systems or both. We investigated whether the female's reproductive status is being concealed in golden-headed lion tamarins, which exhibit extensive paternal care and a mainly monogamous/polyandrous mating system. We used a combination of behavioral observations and endocrine data to determine female reproductive status and to examine changes in sociosexual behaviors over the ovarian cycle and between conceptive and nonconceptive cycles. Females clearly signaled their reproductive status by way of proceptive sexual presenting. Males showed increased frequencies of anogenital sniffing and mounting during the fertile period, indicating that they detected changes in olfactory and behavioral cues emitted by females, and they adjusted their mounting behavior accordingly. Males and females also remained in closer proximity before and during the fertile period, which suggests the existence of mate guarding. We discuss a possible function of behavioral advertisement of reproductive status in shaping the mating system in Leontopithecus chrysomelas.     

The evolutionary dynamics of adaptive virginity,sex‐allocation,and altruistic helping in haplodiploid animals

In haplodiploids, females can produce sons from unfertilized eggs without mating. However, virgin reproduction is usually considered to be a result of a failure to mate, rather than an adaptation. Here, we build an analytical model for evolution of virgin reproduction, sex‐allocation, and altruistic female helping in haplodiploid taxa. We show that when mating is costly (e.g., when mating increases predation risk), virginity can evolve as an adaptive female reproductive strategy. Furthermore, adaptive virginity results in strongly divergent sex‐ratios in mated and virgin queen nests (“split sex ratios”), which promotes the evolution of altruistic helping by daughters in mated queen nests. However, when helpers evolve to be efficient and increase nest production significantly, virgin reproduction is selected against. Our results suggest that adaptive virginity could have been an important stepping stone on the pathway to eusociality in haplodiploids. We further show that virginity can be an adaptive reproductive strategy also in primitively social haplodiploids if workers bias the sex ratio toward females. By remaining virgin, queens are free to produce sons, the more valuable sex in a female‐biased population. Our work brings a new dimension to the studies linking reproductive strategies with social evolution.     

The timing of gonadal development and moult in three raptors with different breeding seasons: effects of gender, age and body condition

Differences between species in breeding seasons are thought to be mediated through differences in their reproductive physiology. Little is known about how the timing and duration of gonadal maturation varies between raptor species, how the timing of moult relates to the gonadal cycle, whether the timing and degree of sexual maturation varies between juveniles and adults or whether body condition has a significant effect. To address these questions, data on gonadal size and moult for adults and juveniles of both sexes of three raptor species were extracted from the Predatory Bird Monitoring Scheme (based on birds found dead by members of the public). The three species, Sparrowhawk Accipiter nisus, Kestrel Falco tinnunculus and Barn Owl Tyto alba, have different ecologies – diurnal bird predator, diurnal mammal predator and nocturnal mammal predator, respectively. All are single‐brooded but have different breeding seasons. The duration of gonadal maturation was markedly different between the species. Barn Owls showed the earliest maturation and the latest gonad regression, and Sparrowhawks the latest maturation and earliest gonad regression. Kestrels were intermediate. In males of all species, the testes remained fully mature throughout their respective breeding seasons. In females, the ovaries remained partially mature throughout the breeding season. Moult started slightly earlier in Sparrowhawks than in Kestrels and coincided with gonadal regression in the two species. Although females of the two species started to moult earlier than males, moult duration was similar between the sexes. Barn Owls showed no distinct annual pattern of moult. In juveniles of all three species, the gonads were smaller than in adults throughout spring and started to mature later. Gonad size in birds that had starved tended to be smaller than in birds dying from other causes, but did not influence the difference in gonad mass between adults and juveniles and between seasons. Body condition had no effect on moult. Whilst ecology has led to the evolution of different breeding seasons, differences between species, and between adults and juveniles, are mediated through adaptive differences in their reproductive physiology.     

Breeding patterns in the oniscid isopod,Porcellio ficulneus Verh., at high temperature and under different photophases

Summary

Porcellio ficulneus Verhoeff inhabits xeric Mediterranean grassland habitats. During winter vitellogenesis takes place and mancas are released in April. Under natural conditions some oocytes (about 20%) were lost. Both high temperature (25°C) at long or short photophases affected the reproductive pattern. Thus at 25°C and constant light vitellogenesis was markedly shorter than under field conditions. Oocytes matured sooner, but many were resorbed. The marsupium was formed earlier and contained fewer eggs. In some cases mancas that had developed could not emerge from the marsupium and died inside. In conclusion, P. ficulneus responds to warmer conditions by breeding earlier and reducing the number of offspring.