The roles of juvenile hormone and biogenic amines on pheromone response plasticity and diapause termination in male Caloptilia fraxinella

In insects that exhibit a period of delayed reproduction, the timing of mating and reproduction is controlled by environmental conditions that regulate endogenous factors such as hormones and biogenic amines (BAs). Caloptilia fraxinella (Ely) (Lepidoptera: Gracillariidae) undergoes a 9‐month reproductive diapause from adult eclosion in the summer until diapause termination the following spring when adults mate. Male response to female sex pheromone is plastic, and is most acute when moths are reproductively active. The aim of this study is to further elucidate the mechanisms involved in the regulation of male response to pheromone in C. fraxinella, and to test whether the application of BAs with and without a juvenile hormone analogue (JHA) to males in different physiological states impacts pheromone responsiveness, as measured by electroantennogram and wind tunnel bioassays. Treatment of male C. fraxinella in reproductive diapause with one application of a JHA induces the highest subsequent pheromone response in the fall, but does not alter pheromone response earlier in reproductive diapause in the summer. The JHAs methoprene and pyriproxyfen similarly enhance pheromone response in the fall. Treatment with methoprene alone or in combination with one of the BAs octopamine, dopamine or serotonin increases male pheromone responsiveness in the fall. The increase in pheromone response can be attributed to methoprene only, as treatment with any of the BAs alone does not enhance male response to pheromone. Biogenic amine treatment lowers male responsiveness to pheromone in some experiments, indicating that there may be a role for BAs in maintaining low pheromone response during reproductive diapause in this species.     

Nutrition,sex and season contribute to variation in fat and glycerol levels in the long‐lived moth Caloptilia fraxinella

The ash leaf cone roller Caloptilia fraxinella Ely (Lepidoptera: Gracillariidae) is an invasive leaf‐mining moth pest of horticultural ash Fraxinus spp. in the Canadian Prairie Provinces. Caloptilia fraxinella overwinter as adults in reproductive diapause and mating occurs after overwintering in the spring. The effect of a carbohydrate food source on fat and glycerol reserves throughout the long adult life stage of this moth is investigated. Insects collected as pupae are given access to either water or sugar water upon adult eclosion. Moths held under the different feeding regimes are sampled before (summer and autumn) and after overwintering in the spring. Analysis of either glycerol or lipid content is conducted for male and female moths from each collection period. Both moth weight and glycerol concentration are affected by moth sex, food regime and season of collection. Although female moths are heavier than males, a higher glycerol concentration occurs in males. Moths fed sugar are heavier and have a higher glycerol concentration than water‐fed moths late in reproductive diapause and after overwintering. Moths collected in the spring after overwintering are lighter and have a lower glycerol content than moths collected before winter. There is a significant influence of feeding regime and season on moth body lipid content, with sugar‐fed moths having more fat than water‐fed moths; however, this difference is smaller in the summer than the autumn or spring. An initial understanding of the overwintering biology and diapause of this pest is provided in the present study.     

Overwintering success in adults of the Japanese common grass yellow Eurema mandarina

The evolution of reproductive diapause is controversial in males as compared to females that must overwinter to leave offspring, because late‐autumn males can obtain offspring by pre‐overwintering copulation. The Japanese common grass yellow Eurema mandarina is suitable to examine the evolution of male reproductive diapause, because direct comparisons are possible between males that do and do not exhibit reproductive diapause. Approximately one‐half of males are insensitive to diapause‐inducing conditions, and emerge as non‐diapause summer‐form. Most autumn‐form females mate with summer‐form males in late autumn. Females that have overwintered re‐mate with autumn‐form males before the onset of oviposition. Because last‐male‐precedence is general in sperm competition in Lepidoptera, it is unclear why half of males emerge as summer‐form in late autumn. A potential adaptive benefit for emerging as summer‐form is increased sperm overwintering success, if autumn‐form females have a higher overwintering success than autumn‐form males. In the present study, overwintering success was estimated for both sexes of autumn‐form adults by rearing under seminatural conditions and a mark–release–recapture technique. Both approaches estimated an overwintering success of approximately 5% for both sexes. The absence of difference in overwintering success between the sexes suggests that pre‐overwintering copulation does not increase sperm overwintering success. However, a considerably low overwintering success may explain, at least partly, the presence of summer‐form males in late autumn. The degree of overwintering success might be more important than the sexual differences of overwintering success in the evolution of male reproductive diapause.     

Influence of energy stores on activation of reproductive function in male golden-mantled ground squirrels

Summary Male golden-mantled ground squirrels were captured in the field in late summer and placed on either unlimited or restricted (80% normal) food rations in the laboratory until each animal began to hibernate. At entrance into hibernation mean body mass was 274.5 g for the unlimited group and 224.5 g for the restricted group. Only six of 21 males subsequently underwent reproductive maturation during winter. Each of these six received the unlimited ration prior to hibernation, and these six included the five heaviest animals in the experiment. In addition to activating their reproductive systems, these heavier squirrels began hibernation later and ended hibernation earlier than the lighter squirrels. Squirrels that remained reproductively quiescent averaged 25.5% body mass as fat in early spring and thus were not severely limited in energy stores during winter.In another experiment a group of 13 male and female squirrels were housed together in a common outdoor enclosure in order to examine the possible relationship between reproductive condition and social standing of males. Although there appeared to be a social hierarchy among males, each male fattened substantially in fall and became reproductively active during winter.These results suggest that the level of energy stores accumulated as fat prior to hibernation in fall affects the potential of male ground squirrels to breed in spring. This dependence of reproductive development on energy stores may reflect the high energetic costs associated with breeding in males when they emerge from hibernation in spring when food availability is low.     

The prediapause copulation and its significance in the butterflyEurema hecabe

In the pierid butterflyEurema hecabe, which shows seasonal polymorphism (summer and autumn morphs) and overwinters at adult stage, whether or not the prediapause copulation may be of usual occurrence and reproductively functional was examined. From the counts of the spermatophores carried by the females, it is clear that the prediapause copulation characteristically occurs in reproductively inactive females of the autumn corph. From behavioral observations in the field, mating partners of those females are mostly males of the summer morph rather than from the autumn morph. In autumn, males of the summer morph remained abundant and searched for females on the larval food plants. Furthermore, they mated frequently with autumn morph females. Autumn morph males seemed to be sexually less active or inactive before hibernation. Microscopic examinations of the spermathecae were made in mated autumn-morph females collected before and after hibernation. The results indicate that sperm is passed by the males at autumnal copulation. The sperm may be stored in the female reproductive tract and utilized for fertilization in spring. This supposition is strongly supported by field data; that is, once-mated autumn-morph females laid fertilized eggs in spring. Finally, physiological basis of the prediapause copulation, its adaptive signficance and the behavioral advantage inE. hecabe are discussed from the viewpoint of seasonal adaptation.     

Plasma steroid concentrations of adult male Atlantic sharpnose sharks, <Emphasis Type="Italic">Rhizoprionodon terraenovae,</Emphasis> in the northern Gulf of Mexico,with notes on potential long term shifts in reproductive timing

In previous literature it was proposed that Atlantic sharpnose sharks, Rhizoprionodon terraenovae, from the western Atlantic Ocean (WAO) population exhibited differences in their reproductive timing when compared to the northern Gulf of Mexico (GOM) population. The objective of the current study was to examine the reproductive cycle of mature male sharpnose sharks from the northern GOM population. Plasma testosterone (T) and estradiol (E₂) were measured from 76 mature male sharpnose sharks sampled from 1999 to 2001, and the gonadosomatic index (GSI) was determined for 53 additional mature males sampled from 2007-2008. Plasma T, E₂, and GSI exhibited similar cycles with peak values occurring in late spring (April/May) and lowest values occurring during the summer. These results suggest that male sharpnose sharks in the northern GOM are reproductively active one to two months earlier than previously reported, which now brings them into synchrony with the WAO population. Interestingly, this shift in reproductive timing corresponds with a 3.0°C (13.4%) increase in spring sea surface temperature (SST) from the Mississippi Sound over a 29 year period, which could translate into GOM male sharpnose sharks becoming reproductively active earlier in the year thus explaining the late spring peak in GSI.     

Reproductive ecology and movement of pallid sturgeon in the upper Missouri River,Montana

Successful recruitment of endangered pallid sturgeon has not been documented in the upper Missouri River basin for decades, and research on the reproductive ecology of pallid sturgeon has been hindered by low sample size. A conservation propagation program was initiated in the 1990s, and the oldest age class of hatchery‐origin pallid sturgeon are becoming sexually mature increasing the number of reproductively‐active fish in the system. However, it is currently unknown how the reproductive ecology of hatchery‐origin pallid sturgeon relates to the few remaining wild fish. Following spring reproductive assessments, weekly relocations were recorded for each individual from late‐May to mid‐July to facilitate comparisons of spawning season movements among reproductive classifications and between spring hydrographs (2015 and 2016) for male pallid sturgeon. Mean total movement distances (±SE) were 104.5 km (18.9) for reproductively‐active wild males, 116.0 km (18.1) for reproductively‐active 1997‐year class males, and 20.6 km (3.0) for non‐reproductively‐active fish of unconfirmed sex. Movement characteristics of reproductively‐active males did not differ between 2015 and 2016 despite a difference of eight days in the timing of peak discharge and a difference of 79 m³/s (16.7%) in magnitude. Male aggregations were observed on the descending limb of the hydrograph in 2016 during temperatures suitable for spawning, but female pallid sturgeon underwent follicular atresia, similar to the other years of the study. Hatchery‐origin pallid sturgeon from the conservation propagation program appear to have retained reproductive characteristics from the wild broodstock, a key finding for a population where local extirpation of the wild stock is imminent.     

Environmental conditions terminate reproductive diapause and influence pheromone perception in the long‐lived moth Caloptilia fraxinella

Reproductive diapause enables long‐lived insects to time mating with environmental conditions suitable for offspring development. Plasticity in the perception of pheromones used in sexual communication may enable mate‐finding at the appropriate time of year. The moth Caloptilia fraxinella (Ely) (Lepidoptera: Gracillariidae) undergoes a 9‐month reproductive diapause, during which the male response to pheromone is plastic and is highest during the period of reproductive activity. The mechanisms controlling this pheromone response plasticity are not well‐understood, and the aim of the present study is to determine the main factors involved. In the present study, the impact of temperature, photoperiod, juvenile hormone analogue (JHA) and adult nutrition on diapause termination are tested using electroantennogram (EAG) and behavioural response to pheromone in male C. fraxinella. Eclosion in a state of reproductive diapause occurs in most males; diapause is maintained under short‐day or cool conditions indoors, or under natural conditions outdoors. Exposure to long‐day, warm conditions over a period of 4 weeks causes a small number of males to become behaviourally responsive to pheromone; a larger number of males become behaviourally responsive over a period of 3 months of post‐eclosion. Treatment with a JHA impacts male EAG and the behavioural response to pheromone during the period of reproductive diapause. A carbohydrate food source is not required by reproductively active adult male C. fraxinella to respond to pheromone and express mate location behaviours. The main factors involved in controlling male pheromone response plasticity and the implications of these factors for the C. fraxinella population in its expanded range are discussed.     

Effects of nutrition and methoprene treatment upon reproductive diapause in Caloptilia fraxinella (Lepidoptera: Gracillariidae)

Abstract Female Caloptilia fraxinella exhibit a prolonged reproductive diapause immediately post adult emergence in mid‐summer until the next spring when mating, egg development and oviposition on fresh Fraxinus spp. leaflets occur. Factors that effect the termination of reproductive diapause are investigated in this species. Caloptilia fraxinella diapausing adults held in overwintering conditions (2 °C, LD 0 : 24 h) for 24 weeks terminate diapause after placement for 2 weeks in simulated summer conditions (24 °C, LD 16 : 8 h) only if they are provided with 10% sugar water. Exogenous application of the Juvenile Hormone (JH) analogue methoprene to moths in both early‐ (summer) and mid‐ (autumn) reproductive diapause demonstrates that JH affects diapause termination but a carbohydrate nutrition source also mediates mating and vitellogenesis. Mating between moth pairs early in diapause occurs only after treatment with methoprene and provision with sugar water. However, there is no impact of mating on the propensity of females to produce vitellogenic oöcytes. Moths collected in the autumn in mid‐diapause respond in a dose‐dependent fashion to methoprene treatment and the response is greater than that of moths early in diapause collected in the summer. Treatment with methoprene and access to sugar water results in vitellogenic oöcytes in 18.75% of females from mid‐diapause moth pairs treated with 0.01 μg methoprene per insect and in all females from pairs treated at the two highest doses of methoprene (0.1 and 1 μg per insect). Mating occurs only between moths in mid‐diapause treated with the two highest doses of methoprene and these doses induce 91% and 100% mating, respectively. Both control and methoprene‐treated males in mid‐diapause held under summer conditions mate successfully and pass a spermatophore to their methoprene‐treated female partner. These data demonstrate that female C. fraxinella undergo a prolonged reproductive diapause in which termination is dependent on JH and further mediated by a carbohydrate nutrition source. The production of vitellogenic oöcytes is independent of mating. These data also provide evidence that response of moths in diapause to exogenous applications of methoprene differs throughout the diapause period and between male and female C. fraxinella.     

Is lack of mating competitiveness in spring linked to mating asynchrony between wild and mass‐reared codling moths from an operational sterile insect programme?

Mating competitiveness and pheromone trap catches of mass‐reared, male codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), from the Osoyoos, British Columbia, Canada, mass‐rearing facility operated by the Okanagan‐Kootenay Sterile Insect Release Board, were compared to wild males using mark–release–recapture field experiments in spring, summer, and autumn at Summerland, British Columbia. In spring, significantly more wild diapause males mated with tethered, wild females than did non‐irradiated (0 Gy) or irradiated (100 or 250 Gy) non‐diapause, mass‐reared males. A lower dose of radiation did not improve mating competitiveness, nor catches of mass‐reared males released in spring. Median mating time for wild males was approximately 45 min earlier than mass‐reared males with most wild males (70.5%) mating before sunset and mass‐reared males mating at or shortly after sunset in spring. Superior mating competitiveness of wild males in spring was mirrored by greater recapture rates in pheromone‐baited traps. In summer, mating competitiveness of mass‐reared moths improved relative to wild males and there was a significant inverse relationship between radiation dose (0, 100, and 250 Gy) and competitiveness of mass‐reared males. In autumn, untreated, wild males were significantly more responsive to pheromone traps than non‐diapausing mass‐reared males receiving 250 Gy of radiation. Mass‐reared males, subjected to diapause‐inducing conditions as larvae and emerged from diapause before this irradiation treatment, were recaptured significantly more often than similarly irradiated, non‐diapause, mass‐reared males, but not more than untreated, diapause wild males. We hypothesize that differences between wild and mass‐reared males in daily timing or speed of responses to natural or synthetic pheromone sources under montane weather patterns typical of spring in British Columbia may partially explain poor activity of sterile males, and low sterile : wild overflooding ratios during spring when measured using pheromone traps by the sterile insect release programme in British Columbia.     

Diapause affects cuticular hydrocarbon composition and mating behavior of both sexes in Drosophila montana

Envipnmental cues,mainly photoperiod and temperature,are known to control female adult reproductive diapause in several insect species.Diapause enhances female survival during adverse conditions and postpones progeny production to the favorable season.Male diapause(a reversible inability to inseminate receptive females)has been studied much less than female diapause.However,if the males maximized their chances to fertilize females while minimizing their energy expenditure,they would be expected to be in diapause at the same time as females.We investigated Drosophila montana male mating bchavior under short-day conditions that induce diapause in females and found the males to be reproductively inactive.We also found that males reared under long-day conditions(reproducing individuals)court reproducing postdiapause fermales,but not diapausing ones.The diapausing fies of both sexes had more long-chain and less short-chain hydrocarbons on their cuticle than the reproducing ones,which presumably increase their survival under stressful conditions,but at the same time decrease their attractiveness.Our study shows that the mating behavior of females and males is well coordinated during and afier overwintering and it also gives support to the dual role of insect cuticular hydrocarbons in adaptation and mate choice.     

Seasonal variations in feeding range and flock structure of the Rook Corvus frugilegus in eastern Ireland

The feeding range and flock structure of Rooks showed temporal variations caused mainly by the dispersion of food and reproductive behaviour.
Feeding range was restricted in spring, autumn and early winter when food availability was high and Rooks were reproductively active. A large feeding range occurred in late summer and late winter, when food availability was either generally low or locally distributed and when Rooks were reproductively inactive. The occurrence of flocks common to several rookeries mirrored variations in feeding range; mixed rookery flocks were more common in late summer and late winter. Similarly, the size of the rookery (as measured by the number of nests) was related to feeding range only when Rooks were reproductively inactive and at such times larger rookeries had greater feeding ranges.
Flock structure showed similar seasonal variations; small widely spaced flocks predominated in summer, autumn, early winter and spring, whereas large dense flocks occurred in late winter. Diurnal variations in flock structure occurred within any one season. Both seasonal and diurnal variations in flock structure may be determined by the dispersion of the prey and the feeding strategy used to obtain it, reproductive behaviour and the risk of predation.     

Environmental Cues,Endocrine Factors,and Reproductive Diapause in Male Insects

Environmental cues, mostly photoperiod and temperature, mediated by effects on the neuroendocrine system, control reproductive diapause in female insects. Arrest of oocyte development characterizes female reproductive diapause, which has two major adaptive functions: It improves chances of survival during unfavorable season(s), and/or it confines oviposition to that period of the year that is optimal for survival of the eggs and progeny. Although reproductive diapause is less well studied in male insects, there may be no sex-dependent differences in regard to the first of these functions. The second one, however, is not valid for the male; instead, selection pressure directs the male's reproductive strategy toward maximum chances of fertilization of the female's eggs with minimum waste of energy. Therefore, in species with female reproductive diapause, the males may or may not exhibit diapause, but if they do, their diapause must be adapted to that existing in conspecific females. Male reproductive diapause is defined as a reversible state of inability of the male to inseminate receptive females. In relation to reproductive diapause, there are several patterns of coadaptations between male reproductive strategy and timing of female receptivity, (a) In some insects, the females are receptive in the early part of their diapause; mating occurs during this period and there is no diapause in the male. The male dies shortly after copulation and the female stores the sperms to fertilize the eggs that develop after termination of the female's diapause, (b) In some species, as in the grasshopper Anacridium aegyptium, females are receptive during diapause; though oocyte development is arrested, copulation occurs and the stored sperms fertilize the eggs when the female's diapause ends. Males were claimed to have no diapause, but recent studies have revealed the presence of a reproductive diapause in a proportion of the males. This and other cases show that female receptivity during reproductive diapause may or may not be accompanied by male reproductive diapause. If there is a reproductive diapause in the male, it is controlled by the same endocrine mechanism, the corpora allata (CA), as in the females, (c) In many species females are refractory during their diapause. In these cases, males exhibit reproductive diapause, which may be light, as in the beetle Oulema melanopus, or well established, as in certain grasshoppers, butterflies, and beetles. In the latter cases, male diapause is controlled by similar environmental cues (photoperiod, temperature) and by the same intrinsic mechanism (neuroendocrine system, especially CA) as female diapause. Nevertheless, male diapause is less intense; the environmental cues leading to its termination are less complex and/or less extreme, so male diapause terminates before that of the females. Presumably, male diapause is under two antagonistic selection pressures: A male should not waste energy by courting dia-pausing refractory females, but he should be ready to copulate as soon as the females become receptive, otherwise he may lose in the competition between males for females. Some further strategies, which do not seem to fit the above patterns, are also outlined.     

Seasonal occurrence and diapause induction of a predatory bug Andrallus spinidens (F.) (Heteroptera: Pentatomidae)

The bug Andrallus spinidens (F.) (Heteroptera: Pentatomidae) is a polyphagous predator of insect larvae, which is distributed in tropical and warm temperate zones worldwide. Seasonal occurrence and diapause induction of this bug were studied in a population in Miyazaki, southern Kyushu, Japan. The field research showed that A. spinidens produces three or four generations a year and is most abundant from mid‐summer to autumn. Reproductive activity of field‐collected adults decreased from late summer to autumn, indicating that this bug enters adult diapause in autumn. Only adults were found in early spring and these were reproductive. Laboratory experiments showed that, irrespective of photoperiod, adult diapause is induced at lower temperatures (≤22.5°C), whereas it is avoided at higher temperatures (≥25°C). Ambient temperature falls across the critical range from late summer to autumn. Thus, the bug clearly overwinters in adult diapause induced by low temperatures and this diapause is terminated during the course of winter.     

Annual variation in energy reserves in male three-spined stickleback, Gasterosteus aculeatm L. (Pisces, Gasterosteidae)

Wet and dry weights of tissue were measured and concentrations of glycogen, lipid and protein were estimated for the liver, gonad and carcass of male sticklebacks from an annual population collected each month over one complete year. In young-of-the-year there is one period of rapid weight gain, in all three body regions (liver, carcass and gonads) but particularly of the carcass, in the autumn and a second in spring and early summer. This is accompanied by an increase in the water content of all three body regions. The gonadosomatic index also increases sharply in spring and early summer. Young sticklebacks accumulate lipid and glycogen slowly during the autumn and winter of their first year of life and more rapidly from late winter to early summer. Thus, the period of most rapid accumulation of these reserves coincides with the time when body weight and gonad maturation are also increasing sharply. Lipid and glycogen levels fall during the reproductive season in those males that breed, so that by July they are reduced to 43% and 37% (respectively) of their peak values in May. Levels of protein increase throughout the year as the fish grow, but in breeding males by July the concentration of protein in the carcass falls to 70% of pre-breeding levels. Breeding males therefore reach the end of the reproductive season with their total energy reserves severely reduced, and consequently they suffer a very high mortality. In contrast, adult males that fail to reproduce survive beyond the breeding season. They continue to gain weight and to accumulate lipid and glycogen from August to September, but these energy reserves fall (to levels comparable to those of post-breeding fish) in December, when these fish probably die. These results demonstrate that in male sticklebacks, growth and gonad maturation can be sustained in parallel with the accumulation of energy reserves, which are subsequently extensively depleted as a result of reproductive activities.     

Female receptivity, embryonic diapause, and superfetation in the European badger (Meles meles): implications for the reproductive tactics of males and females

The European badger Meles meles is thought to mate throughout the year with two mating peaks occurring in late winter/spring and summer/autumn. After mating, fertilized ova enter embryonic diapause (delayed implantation) at the blastocyst stage, which lasts up to eleven months. Even if mating is successful, however, the estrous cycle may continue during embryonic diapause, which suggests that female badgers are capable of superfetation (conception during pregnancy). This may increase female fitness by facilitating polyandry, and reduce the risk of infanticide by resident males through paternity confusion. Detailed understanding of female receptivity, specifically the association of superfetation with embryonic diapause, may explain field observations of seemingly inconsistent reproductive tactics of male badgers with regard to, for instance, whether or not they guard mates or defend territories. The combination of embryonic diapause and superfetation may occur in other mustelids; if so, the sociobiology of mustelids will need reevaluating, and the Mustelidae may prove to be a good model taxon for studies of sexual conflict in the reproduction of eutherian mammals.     

Demographic study of the reproductive potential of pear psylla, Cacopsylla pyri

Longevity and fecundity of pear psylla adults, Cacopsylla pyri (L.) (Homoptera: Psyllidae), were studied in Greece under field conditions, during 1991/92 and 1992/93. Net fecundity (mean fecundity of cohort) of the overwintered females (winter form) was low (24.1 and 43.4 eggs per female for 1991/92 and 1992/93 respectively) because only a small fraction of the females survive reproductive diapause. Surviving females, however, are capable of producing a relatively high number of eggs (135.0 and 151 9 eggs per female for the two seasons respectively). Net fecundity of summer form females was very high in spring (343.2 and 406.8 eggs per female for 1992 and 1993 respectively), relatively high in early autumn (130.4 and 192.5 eggs per female) and very low in summer (21.9 and 46.5 eggs per female). Male longevity was in all cases shorter than that of female. Longevity of both sexes of the summer form was higher in spring and autumn (for females it ranged from 22.3 to 28.4 days) than it was in summer (for females it ranged from 14.3 to 17.4 days).     

Environmental Cues, Endocrine Factors, and Reproductive Diapause in Male Insects

Environmental cues, mostly photoperiod and temperature, mediated by effects on the neuroendocrine system, control reproductive diapause in female insects. Arrest of oocyte development characterizes female reproductive diapause, which has two major adaptive functions: It improves chances of survival during unfavorable season(s), and/or it confines oviposition to that period of the year that is optimal for survival of the eggs and progeny. Although reproductive diapause is less well studied in male insects, there may be no sex-dependent differences in regard to the first of these functions. The second one, however, is not valid for the male; instead, selection pressure directs the male's reproductive strategy toward maximum chances of fertilization of the female's eggs with minimum waste of energy. Therefore, in species with female reproductive diapause, the males may or may not exhibit diapause, but if they do, their diapause must be adapted to that existing in conspecific females. Male reproductive diapause is defined as a reversible state of inability of the male to inseminate receptive females. In relation to reproductive diapause, there are several patterns of coadaptations between male reproductive strategy and timing of female receptivity, (a) In some insects, the females are receptive in the early part of their diapause; mating occurs during this period and there is no diapause in the male. The male dies shortly after copulation and the female stores the sperms to fertilize the eggs that develop after termination of the female's diapause, (b) In some species, as in the grasshopper Anacridium aegyptium, females are receptive during diapause; though oocyte development is arrested, copulation occurs and the stored sperms fertilize the eggs when the female's diapause ends. Males were claimed to have no diapause, but recent studies have revealed the presence of a reproductive diapause in a proportion of the males. This and other cases show that female receptivity during reproductive diapause may or may not be accompanied by male reproductive diapause. If there is a reproductive diapause in the male, it is controlled by the same endocrine mechanism, the corpora allata (CA), as in the females, (c) In many species females are refractory during their diapause. In these cases, males exhibit reproductive diapause, which may be light, as in the beetle Oulema melanopus, or well established, as in certain grasshoppers, butterflies, and beetles. In the latter cases, male diapause is controlled by similar environmental cues (photoperiod, temperature) and by the same intrinsic mechanism (neuroendocrine system, especially CA) as female diapause. Nevertheless, male diapause is less intense; the environmental cues leading to its termination are less complex and/or less extreme, so male diapause terminates before that of the females. Presumably, male diapause is under two antagonistic selection pressures: A male should not waste energy by courting dia-pausing refractory females, but he should be ready to copulate as soon as the females become receptive, otherwise he may lose in the competition between males for females. Some further strategies, which do not seem to fit the above patterns, are also outlined.     

Population regulation in the Wood Mouse Apodemus sylvaticus (L.)

Watts 1969 described the population cycle of the Wood Mouse Apodemus sylvaticus as a decline in numbers in spring, a stable period in summer and an increase phase in autumn. Aggression by adult males was considered an important aspect of population regulation limiting male survival in spring and juvenile recruitment in summer. However, recent studies of range behaviour and experimental investigations suggest that male A. sylvaticus are not relevant in population regulation and that the principal regulating factors act on female reproductive success. Female reproductive activity is determined by spatial interactions which, in turn, are determined primarily by the density of females and the proportion of female immigrants in the population. The only impact adult males may have on population size is through infanticide and competition with the females for food.