Factors affecting size, longevity and fecundity in the reindeer oestrid flies Hypoderma tarandi (L.) and Cephenemyia trompe (Modeer)

1. Laboratory reared reindeer oestrid flies Hypoderma tarandi and Cephenemyia trompe (Diptera: Oestridae) were weighed to determine progressive weight loss and death weights at treatments with various temperature and humidity conditions.
2. Four individual measurements of size were taken: larval weight, wet weight of newly eclosed flies, wing length, and weight of flies after dehydration and fat extraction. In H. tarandi, males were bigger than females (except for wing length), whereas the reverse was true for C. trompe .
3. Size variation was not significantly related to conditions (temperature, humidity, duration) during the pupal stage, but individual reindeer produced flies (both species) of different mean sizes. These size differences were not correlated with larval burden (= number of larvae per individual host), but are hypothesized to be connected to unknown host quality factors.
4. Longevity of flies kept in vials and subjected to various temperature and humidity conditions revealed that C. trompe lived significantly longer than H. tarandi (range: 4–44 and 1.2–27 days, respectively) at 5–33 °C. Male H. tarandi survived longer than females; female C. trompe survived longer than males. Longevity was not significantly correlated to any of the size measures.
5. Most flies had a large portion of their fat reserves left at death.
6. In H. tarandi , mean number of eggs was 609 ± SD 73 (range 354–772, n = 119). Egg number was slightly dependent on larval size, but not on wet weight of newly eclosed flies or wing length. In C. trompe , mean number of eggs was 960 ± SD 208 (range 493–1349, n = 31).
7. The possible adaptive value of large size in oestrids is questioned. Benefits of flexibility in size in oestrids are hypothesized.     

Oviposition of resistant and susceptible strains of Drosophila melanogaster in the presence of deltamethrin

Oviposition of three strains of Drosophila melanogaster in the presence of deltamethrin was observed. These strains had different levels of physiological susceptibility to deltamethrin. Two-choice tests were conducted with couples of flies in petri-dish arenas containing two oviposition dishes. On the first day of the experiment, females were given a choice between a treated oviposition dish and an untreated control dish. On the second day of the experiment, two control oviposition dishes were given to females. Although individual females showed a tendency to aggregate their eggs in one of the dishes, control experiments demonstrated an overall equal distribution of eggs between the dishes. When one of the two oviposition dishes in the arena was treated with deltamethrin, the percentage of females ovipositing and the mean number of eggs laid by females were reduced, compared with control arenas. Females avoided the treated oviposition dish and laid significantly more eggs on the control dish. Furthermore, when the deltamethrin concentration was increased on the first day, female flies postponed their oviposition and laid significantly more eggs on the second day. The resistant strain, SR, demonstrated the same capacity to select the untreated site for oviposition as the susceptible strain, but it showed a smaller oviposition reduction and egg retention. The relationship between physiological and behavioural susceptibility to deltamethrin is discussed.     

Lifecycle of Rhipicephalus haemaphysaloides under laboratory conditions

Rhipicephalus haemaphysaloides Supino, 1897 (Ixodida: Ixodidae) is a three‐host hard tick widely distributed in Sri Lanka. The lifecycle of the R. haemaphysaloides population in Sri Lanka was studied under laboratory conditions using fully engorged females collected from dogs. Larvae, nymphs and adults were reared on New Zealand rabbits. Engorged females weighed a mean ± standard deviation (SD) of 470.1 ± 106.5 mg and laid a mean ± SD of 4067.4 ± 3136.2 eggs within 16.1 ± 3.7 days. The mean ± SD preoviposition period was 6.5 ± 1.1 days. The mean ± SD reproductive efficiency index was 8.5 ± 4.1, and was higher in heavier females (Pearson correlation, r = 0.8; P = 0.001). The incubation period of eggs was shorter at higher temperatures. The majority (86.9%) of larvae successfully moulted after a mean ± SD premoulting period of 7.9 ± 0.7 days following a parasitic period of 3.2 ± 0.5 days. Unfed larvae survived a mean ± SD of 103.4 ± 19.8 days. Nymphs completed the bloodmeal after 3.7 ± 0.9 days, weighed 5.7 ± 1.7 mg and demonstrated a premoulting period of 15.1 ± 1.9 days. Unfed nymphs survived a mean ± SD of 45.8 ± 3.8 days. Heavier nymphs moulted into females, whereas lighter nymphs moulted into males (two‐sample t‐test, P < 0.001). The bodyweight of adult females increased more than 100‐fold after feeding. The lifecycle of R. haemaphysaloides was successfully completed on rabbits within a mean of 91.3 days (range: 69–117 days) under laboratory conditions.     

Embryonic development and reproductive seasonality of Buccinanops globulosus (Nassariidae) (Kiener, 1834) in Patagonia,Argentina

Buccinanops globulosus mated all year round, with higher frequency from May to September, prior to spawning months. Gravid females were found between October and March. Oviposition peaked during rising temperatures and longest daylength while hatching peaked with high water temperature and declining daylength. Gravid females measured between 20 and 41?mm in shell length. The spawn consisted on average of 31 egg capsules, each containing 1266 eggs. Embryos usually completed development within each egg capsule by ingesting small fragments of the uncleaved nurse eggs, which were not a limiting resource. Egg capsules with more than one embryo were not common; in those cases, the embryos had different sizes probably related to intracapsular competition for nutrients, and were on average smaller than solitary embryos in the other capsules. Embryos hatched as crawling juveniles with a mean hatchling shell length of ~3.4?mm. In a few cases, malformed embryos were found, but it was not a common phenomenon. The information recorded in this study, as the minimum reproductive size and spawning season, is valuable for fisheries management.     

Effects of larval diet on development rates and reproductive maturation of male and female Mediterranean fruit flies

Abstract Mediterranean fruit fly Ceratitis capitata (Diptera: Tephritidae) larvae are capable of developing in one of many hosts that may vary greatly in quality. We hypothesized that they will respond to the larval environment in a manner beneficial to their subsequent reproductive performance. Accordingly, we investigated the effects of various larval diets (varying in the amount of protein and sugar they contain) on the size, development time, nutritional status and reproductive maturation (ovarian development and onset of sexual behaviour) of females and males. We found that flies which undergo larval development in artificial host fruit that contain sugar and protein ('protein-fed') were larger, developed faster and emerged with more nutritional reserves than flies that were protein-deprived as larvae. Protein-fed males, regardless of their size, became sexually active before males that developed in hosts with no protein. Protein-fed females produced more mature eggs than protein-deprived ones. Moreover, protein-fed females tended to copulate sooner than females that developed in hosts with no protein. In addition, regardless of female larval diet, females with more mature eggs tended to copulate sooner than females with less mature eggs. In light of these results, the importance of the larval environment for adult reproductive success is discussed.     

Physiological status of Drosophila suzukii (Diptera: Drosophilidae) affects their response to attractive odours

An improved understanding of the biology of the invasive pest, Drosophila suzukii (Matsumura) (Diptera: Drosophilidae), is critical for the development of effective management strategies. Trapping is one technique used for both detection and control; however, the efficacy of trapping can vary depending on the target insect's physiological state, its behavioural priorities and the type of attractant used in the trap. We conducted a series of caged trapping experiments and a greenhouse trapping experiment to investigate the effects of D. suzukii feeding status, age, mating status, ovipositional status and seasonal morph type on the capture rate of traps baited with fermentation odours. Starved flies were trapped at greater rates compared to fed flies; more virgin flies were trapped than mated flies; flies deprived of an oviposition substrate were trapped more frequently than flies given an oviposition substrate. It is still unclear whether age or seasonal morphology affect bait response. Lastly, a caged choice experiment investigated the relationship between female reproductive status and attraction to fermentation or fruit odours. Fermentation‐based traps captured female flies regardless of their reproductive status but, ripe fruit‐based traps were more attractive to flies with more than seven eggs. In summary, studies that use fermentation‐based traps should recognize that capture rates of D. suzukii will depend on the feeding, mating and oviposition experiences of the population; also, fruit‐based traps may better target gravid females.     

Effect of mating period and time‐of‐day for bloodmeal on rearing of Asian tiger mosquito (Aedes albopictus) in laboratory conditions

Colonization and maintenance of mosquitoes in the laboratory is required to study physiology, ecology, and behavior of mosquitoes and interactions between mosquito and pathogens. Artificial blood feeding systems have been widely used to maintain the laboratory colony of Aedes albopictus. In this study, we investigated the effects of mating period (1, 3, 6, and 10 days) and time‐of‐day for bloodmeal (08:00, 13:00, and 18:00) in the use of an artificial feeding system on blood‐feeding rate, female fecundity, egg hatching rate, and developmental time of the Asian tiger mosquito, A. albopictus. Younger females mated for three or fewer days reproduced more eggs compared to those of oldest females mated for ten days. Similar to the result for eggs laid, the mean egg‐hatching rate was significantly higher from the offspring of younger females than from those of older females. However, mating period and time‐of‐day for bloodmeal had no effect on the blood feeding rate and developmental time. Taken together, we suggest that three‐day mating with bloodmeal at 18:00 is optimal for maintaining colonies of A. albopictus in laboratory conditions.     

Comparative fecundity and associated factors for two sibling species of the Anopheles gambiaecomplex occurring sympatrically in The Gambia

Abstract. For two sibling species of mosquitoes belonging to the Anopheles gambiae complex of malaria vectors, the effects of body size (wing length) and bloodmeal size (haematin excretion) on fecundity of wild females were investigated in The Gambia, West Africa. Freshly blood-fed individuals from sympatric populations of An.arabiensis and An.gambiae sensu stricto were sampled by collection at 07.00–09.00 hours from within bednets during July/August 1993, at the beginning of the rainy season. The possible confounding effect of infection with Plasmodium parasites was removed by eliminating infected mosquitoes from the study samples. An.arabiensis females comprised 75% of the An.gambiae sensu law population and were significantly larger (greater mean wing length) than those of An.gambiae s.s. mosquitoes. Mean egg production per female (for the subsequent gonotrophic cycle, excluding pre-gravids) for the two species was not significantly different, though the relationship between wing length and egg production showed An.gambiae s.s. to be more fecund than the An.arabiensis of the same size. Pre-gravid An.gambiae s.s. had consumed significandy smaller bloodmeals than gravid females but the mean wing length of these two gonotrophic categories was not significantly different. In contrast, An.arabiensis pre-gravids were smaller and had consumed smaller bloodmeals than the gravids.     

Geographic variation in reproductive traits and trade-offs between size and number of eggs of the Chinese cobra (Naja atra)

We collected gravid Chinese cobras (Naja atra) from one island (Dinghai) and three mainland (Yiwu, Lishui and Quanzhou) populations in south‐eastern China to study geographical variation in female reproductive traits and the trade‐off between the size and number of eggs. We then conducted an common experiment on cobras from two of the four populations to further identify factors contributing to the observed trade‐offs. The mean size (snout–vent length) of the smallest five reproductive females increased with increasing latitude. Oviposition occurred between late June and early August, with females from the warmer localities laying eggs earlier than those from the colder localities. Maternal size was a major determinant of the reproductive investment in all populations, with larger females producing not only more but also larger eggs. Clutch size was more variable than egg size within and among populations. The observed geographical variation in clutch size, egg size, clutch mass and post‐oviposition body condition was not a simple consequence of variation in maternal size among populations, because interpopulation differences in these traits were still evident when the influence of maternal size was removed. The upper limit to reproductive investment was more likely to be set by the space availability in the island population, but by the resource availability in the three mainland populations. Trade‐offs between size and number of eggs were detected in all populations, with females that had larger clutches for their size having smaller eggs. Egg size at any given level of relative fecundity differed among populations, primarily because of interpopulation differences in the resource availability rather than the space availability. Except for the timing date of oviposition and the mean size of the smallest five reproductive females, all other examined traits did not vary in a geographically continuous trend. The common garden experiment, which standardized environmental factors, synchronized the timing date of oviposition, but it did not modify the conclusion drawn from the gravid females collected from the field. The observed geographical variation in the female reproductive traits could be attributed to the consequence of the effects of either proximate or ultimate factors. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 85 , 27–40.     

Influence of host age and host deprivation on longevity, egg load dynamics and oviposition rate in Microplitis rufiventris

Adult size, longevity, egg load dynamics and oviposition ofMicroplitis rufiventris Kok. which began their development in the first, second, third （preferred hosts） or fourth （non-preferred hosts） instar larvae of Spodoptera littoralis （Boisd.） were studied. The parasitoid size was largely determined by the initial host size at parasitism. Non-ovipositing females derived from older hosts lived for longer periods than those derived from younger ones. However, the ovipositing females, irrespective of their size, lived for almost the same periods. At emergence, the oviducts of adult females contain a significant amount of mature eggs available for oviposition for a few hours on eclosion day. Egg load increases during the early phase of adult life. The amount of additional mature eggs and rate of egg maturation per hour was greater for wasps derived from preferred hosts compared with those in females derived from non-preferred hosts. The pattern of egg production in M. rufiventris females depended on the availability of hosts for parasitization. Host-deprived females depleted the egg complement with aging; the longer the host deprivation, the lower the oviduct egg load. Marked reduction in both realized or potential fecundity of host-deprived females was observed following host availability. Host privation for more than 3 days induced a marked deficit fecundity pattern through the female＇ s life. The realized fecundity was determined by the interaction among host availability, the number of eggs that are matured over the female＇ s life span, oviposition rate and host size from which the female was derived. These results suggest that： （i） M. rufiventris wasp is a weak synovigenic species; （ii） the maturation of additional eggs is inhibited once the maximum oviduct egg load is reached; （iii） the egg load of the newly emerged female is significantly less than the realized fecundity; and （iv） because M. rufiventris females oviposit fewer eggs when they begin depleting their egg supply at 3 days, augmentative releases will require release immediately following emergence to ensure the highest parasitization rate in the field.     

Sexual dimorphism inAnastrepha suspensa (Loew) and other tephritid fruit flies (Diptera: Tephritidae): Possible roles of developmental rate,fecundity, and dispersal

Larger male Caribbean fruit flies are more likely to be chosen as mates and defeat rivals in territorial contests. Yet males are smaller than females. Adaptive explanations for relatively small male size include (1) acceleration of male development to maximize female encounter rates, (2) selection for greater female size to increase fecundity, and (3) selection for body sizes most suitable for sexually dimorphic degrees of mobility, speed, and distance flight. None of these unambiguously accounts for the degree of sexual dimorphism. Male development is not accelerated relative to that of females. On average, males remain inside fruit longer than females and those males with extended development periods are smaller than more rapidly developing individuals. There is no evidence that female enlargement alone, presumably for greater fecundity, has generated the degree of dimorphism in the Caribbean fruit fly or other fruit flies. The relationship between dimorphism and mean female body size in 27 species of Tephritidae is the opposite of what would be predicted if differences in dimorphism were due to differences in unilateral female enlargement. Larger size in a species or in one sex of a species may be an adaptation for extensive flight. In general, among 32 species of fruit flies, as body size increases, wing shape becomes progressively more suited for distance flight. However, there are important exceptions to this correlation. Both sexual selection and nonadaptive allometries may contribute to the range of dimorphisms within the family.     

Manipulation of laying effort reveals habitat-specific variation in egg production constraints in Great Tits (Parus major)

It has been suggested that a bird's clutch size is not limited by the amount of resources available at the time of laying but that differences in the availability of food for nestlings is the ultimate underlying factor determining spatio-temporal variations in clutch size. However, habitat-related variations in egg production ability has yet to be investigated explicitly. We studied the breeding of Great Tits Parus major in deciduous and coniferous forests in the same area. The sizes of both the clutches and the eggs were, on average, larger in the former habitat than in the latter. A number of females were induced to lay more eggs than usual by removing four eggs from designated experimental clutches early in the laying period. These manipulated females laid approximately one egg more than control females, with the number of additional eggs laid not differing between the habitats. However, in both study years the relative size of the extra eggs – relative to the mean size of earlier laid eggs of the same clutch – was smaller in the coniferous habitat than in the deciduous habitat, while there was no habitat-related difference in the relative size of the last-laid eggs of control clutches. This result indicates that some form of proximate limitation during egg-laying period can contribute to the relatively small clutches and eggs in the coniferous habitat. Our results emphasize the need to take egg production costs into account when attempting to account for spatial variation in the reproductive behaviour of birds.     

Effect of host deprivation on egg quality,egg load,and oviposition in a solitary parasitoid,Chetogena edwardsii (Diptera: Tachinidae)

Solitary parasitoids are limited to laying one egg per host because larvae compete within hosts. If host encounter rate is low, females should not increase the number of eggs/host in response. The tachinid fly, Chetogena edwardsii,was used to evaluate the effect of host deprivation on egg accumulation, oviposition behavior, and egg quality in a solitary parasitoid. Females deprived of hosts for 2– 7 days accumulate about 1 day's supply of eggs. Egg output of deprived females once hosts are restored does not differ from that of control females. Deprived females retain one egg in the uterus where it undergoes embryogenesis. Maggots emerging from retained eggs are more likely to survive in hosts molting in 40 h or less after receipt of an egg than are maggots emerging from eggs fertilized shortly before oviposition. Egg retention is a consequence of host deprivation that permits females to broaden the range of hosts they can exploit to include soon-to-molt hosts and possibly multiply parasitized hosts.     

Fecundity and oviposition ofEucelatoria bryani, a gregarious parasitoid ofHelicoverpa zea andHeliothis virescens

We examined longevity, fecundity, and oviposition strategies ofEucelatoria bryani Sabrosky (Diptera: Tachinidae), a gregarious endoparasitoid ofHelicoverpa zea (Boddie) andHeliothis virescens (F.) (Lepidoptera: Noctuidae). Longevity of adult femaleE. bryani was not related to body size. In contrast to longevity, largerE. bryani females had greater potential fecundity than smaller females, as determined by the number of embryonated eggs present in the common oviduct. However, female parasitoid size did not affect primary clutch size (number of eggs deposited in a host). Because embryos in eggs located in the ovisac were larger than those located elsewhere in the common oviduct, maximum primary clutch size may be physiologically limited by the number of fully mature eggs a female has available at one time.E. bryani females adjusted primary clutch size in response to host size, for bothH. zea andH. virescens. This adjustment appears to be adaptive because females did not overexploit hosts by depositing more larvae than a host could support. Adult emergence was not related to host size. Although host weight positively influencedE. bryani progeny weight, increases in progeny size with host size were counterbalanced by increases in primary clutch size with host size.     

Factors affecting the landing and feeding responses of the tsetse fly Glossina pallidipes to a stationary ox

In Zimbabwe, studies were made of the landing and feeding responses of Glossina pallidipes on an ox. Of the tsetse approaching an ox, ≈ 70% fed. Increasing densities of tsetse increased the grooming responses of the ox but had no significant effect on the percentage of tsetse that engorged. The landing site of tsetse on the ox varied with density, with ≈ 50% landing on the legs at low densities (< 20 flies per ox), compared to ≈ 80% at densities > 40 flies per ox. For male G. pallidipes , the mean bloodmeal size was 37 mg. The probability of feeding was negatively correlated with fat content, declining from 91% for flies with < 1 mg fat to < 50% for flies with > 4 mg fat. Bloodmeal size was also negatively correlated with fat content; the regression equation relating bloodmeal size and fat content indicated that the mean wet weight declined from 42 mg for flies with 1 mg of fat to 31 mg for flies with 5 mg of fat. For females, the probability of feeding was not significantly affected by age as determined by ovarian category but there was a paucity of young (ovarian category 0) flies attracted to the ox. Pregnancy status had no significant effect on the probability of feeding, but samples of flies attracted to the ox showed a relative dearth of females approaching larviposition and a preponderance just after.     

Intruder Pressure Affects Territory Size and Foraging Success in Asymmetric Contests in the Water Strider Gerris lacustris

The theory of evolutionarily stable strategies (ESS) in asymmetric contests predicts that the propensity of an individual to expose itself to risk during contests depends on the individual's resource-holding-potential (RHP) and on the value of the disputed resource (V) for the individual compared with that for an opponent. If encounters of a territory owner with individuals of high RHP and high food demands (V) increase in frequency, one should expect a decrease in total aggressiveness of the territory owner, and in consequence a decrease of its territory size. Such a decrease should result in a lower amount of food consumed by the territory owner. Using natural variability in RHP and V in Gerris lacustris, I experimentally tested these predictions. The average prey item has higher value (V) for reproductive female water striders (which probably transform most of their food into eggs), than for nonreproductive females and for males. Because males are smaller, they have lower RHP than females, as RHP depends on size. Thus the reproductive females are the class of individuals of high RHP and high food demands (high V). Most nonreproductive females defend food-based territories. I observed two groups of water striders in a seminatural laboratory setting. As predicted, there was a negative correlation between the rate of encounter with reproductive females and size of the territory, and a positive correlation between territory size and number of Drosophila flies consumed by the owner. Territories were smaller in the group with high rates of encounter between territory owners and reproductive females. Territory owners caught the same number of Drosophila flies as non-territorial individuals in this group. In contrast, in the group with fewer encounters between territory owners and reproductive females, territories were larger, and territory owners gained more food than non territorial water striders.     

Effect of food deprivation on attractiveness of food sources, containing natural and artificial sugar and protein, to three African fruit flies: Ceratitis cosyra, Ceratitis fasciventris, and Ceratitis capitata

The effects of food deprivation, age, and mating status on the responses of three fruit fly species, Ceratitis cosyra (Walker), Ceratitits fasciventris (Bezzi), and Ceratitits capitata (Wiedemann) (Diptera: Tephritidae) to natural and artificial sugar and protein food sources were investigated. Natural food sources included guava [Psidium guajava L. (Myrtaceae)] juice (a common host fruit for all three fruit fly species) and bird faeces (farm chicken). Artificial food sources included molasses (obtained from a local sugar factory) and a locally produced protein bait (the International Centre of Insect Physiology and Ecology yeast). In all species studied, sugar deprivation of immature (1–2‐day‐old) male and female flies increased their response to food odours, although it did not change their preference for the type of odour (protein or sugar). Protein deprivation of mature (14–17‐day‐old) male and female flies also increased their response to food odours compared to protein‐fed flies. Protein‐deprived females were highly attracted to odours from protein sources in particular. Odours from natural food sources, guava juice, and chicken faeces, were more attractive to food‐deprived flies than were odours from artificial sugar and protein sources. Attraction to food odours increased significantly with increasing age for protein‐deprived females of all species. For males and females of all species, nutritional state was a more important factor than mating status in influencing responses of flies to food odours. Practical implications of these findings are discussed in terms of strategies for fruit fly control using food baits.     

The effects of male body size on territorial and mating success in the landmark-defending fly Hermetia comstocki (Stratiomyidae)

Abstract.

• 1 Males of Hermetia comstocki Williston compete for territorial control of certain agaves and yuccas. Winners copulate with females that visit these plants solely to acquire a mate.
• 2 Males vary in body weight by more than an order of magnitude and larger flies almost always defeat smaller ones in aerial contests for control of landmark territories.
• 3 The mean body size (as measured by wing-length) was significantly greater for males retaining residency at a site for at least one hour compared to males unable to do so. Likewise, males able to return to a perch site in the study area on more than one day were larger on average than males unable to do so.
• 4 Male preferences for landmark territories remained similar across years. Large males dominated the perch landmarks most likely to be occupied by males and most likely to be visited by females.
• 5 Despite the fighting and territorial advantages enjoyed by large males, the mean size of males found mating with females was not significantly larger than that of the general population.
• 6 The apparent failure of large males to secure a statistically significant mating advantage may be a statistical consequence of the small sample size of males observed mating. On the other hand, any mating advantage of large males may be reduced because (a) receptive females visit many different landmarks, (b) females mate with the first male they encounter at a landmark, regardless of his size, (c) there are usually many vacant landmarks available for smaller males, and (d) even popular territories are often open to small males, thanks to the low site-tenacity of territory owners.

     

Strategies of a successful new invader in European fresh waters: fecundity and reproductive potential of the Ponto-Caspian amphipod Dikerogammarus villosus in the Austrian Danube, compared with the indigenous Gammarus fossarum and G. roeseli

1. Fecundity of a Dikerogammarus villosus population at Spitz was studied in the Austrian Danube during the 3‐year period 2002–2004. Ovigerous females were absent in October and November, and extremely scarce in December when the reproductive season started again slowly. From January to September pre‐copulatory pairs and egg‐carrying females were present. The reproductive cycle lasted for 9–10 months. 2. Various pigmentation phenotypes of D. villosus have been described in the literature. However, no significant differences were found between the reproductive variables studied here and several colour morphs. Mating was size‐assortative; mean body length of males was about 1.3 times greater than that of their potential mates, and the wet weight was approximately twice as heavy. 3. The relationship between the number of embryos per clutch and the wet weight of females was described by a 3‐parameter power equation. The population mean was 43 eggs with a range of five to 194 eggs. Eighty‐two specimens from 1359 D. villosus females had more than 100 eggs: the smallest of these females was 12 mm long (30 mg) wet weight, and the largest, which was 18 mm long (91 mg), had 194 eggs in embryonic development stage 4. 4. Numbers of embryos in developmental stages 2 (early egg stage) and 7 (newly hatched neonates) differed significantly with body wet weight of ovigerous females (P < 0.05). For an average female in the range 10–12 mm (20–30 mg) the number of juveniles in the brood pouch was 74% of the number of stage 2 eggs. This value can be interpreted as the survival rate of eggs. 5. The overall mean egg volume (EV, ±95% CL) of stage 2 eggs of D. villosus was 0.05 ± 0.001 mm³, and EV increased significantly at each stage of development. At stage 6, egg volume had increased by a factor of 2.6, and averaged 0.13 ± 0.001 mm³. In comparison, G. fossarum and G. roeseli had significantly larger eggs in all developmental stages. 6. Mean egg size of D. villosus (0.063 mm³) was maximal in January. For D. villosus (and G. roeseli) the minimum mean egg size occurred in September. In contrast to G. fossarum and G. roeseli, a second peak in egg size was not observed for D. villosus, and egg size fell more or less successively from January to September. 7. A simple index of fecundity was calculated from the number of stage 2 eggs divided by the female's wet weight. The highest values were observed in April and May, when females from the overwintering generation grew to their maximum body size. Thus the release of a large number of neonates corresponds with the availability of plentiful food and rising water temperatures for juvenile growth in the spring. The lowest value occurred in December. In June the small females of a summer generation appeared, with a naturally low fecundity. 8. The relationship between brood development time and water temperature was studied in the laboratory at a series of constant temperatures. At 16 °C, mean brood development time was 14 days for D. villosus, compared with about 3 weeks for the indigenous species. At 10 °C, mean brood development time was 24 days in D. villosus, compared with 40 days in G. fossarum and 44 days in G. roeseli. At 4 °C it was 1.8 and 3.5 times longer in G. fossarum and G. roeseli. 9. The number of offspring produced by a single clutch from a large female D. villosus is considerably higher than the total numbers produced by the indigenous freshwater gammarids, such as G. fossarum, G. roeseli and G. pulex, during their life‐spans of 1.5–2 years in seven to nine successive broods. Only one or two large ovigerous D. villosus would probably be enough to start a new population. A potentially high reproductive capacity, comparatively small eggs, optimal timing to release the maximum number of neonates per female in April/May, and a long reproductive cycle, together with rapid development of eggs, rapid growth to sexual maturation, short life span, tolerance to a wide range of environmental conditions, and exceptional predatory capabilities, all give the invasive Ponto‐Caspian gammarid an opportunity to become globally distributed in freshwater ecosystems of the temperate climate zone.     

Divergence in reproductive ecology of the three-spined stickleback,Gasterosteus aculeatus

An anadromous population (trachurus form) and three freshwater populations (leiurus form) of the three-spined stickleback,Gasterosteus aculeatus, in central Japan were compared with one another in the reproductive traits: body length at maturity, egg size, clutch size, the swell of abdomen and the number of eggs per nest. The anadromous fish which become larger in body size at maturity spawn eggs smaller in size and greater in number than the freshwater fish. The abdomen swell of gravid females expressed by the proportion of abdomen width to body length was greater in the freshwater fish. The anadromous male fish collected a mean of 2,638 eggs with a range of 1,119 to 4,052 eggs from about 6–7 females. In the three freshwater (the Yamayoke, the Tsuya and the Jizo) populations, males must have mated with about 9–22, 7–18 and 4–7 females respectively. It seems that theleiurus form increases its reproductive success by its much more mating opportunities and the parental efforts of nesting males as well as by spawning large eggs. Furthermore, among the freshwater populations, the Jizo one inhabiting the upper stream was clearly larger in body size, in egg size and in clutch size than the Yamayoke and the Tsuya ones which inhabit stable waterbodies with springs. It is possible that the Jizo population adopted the strategy of spawning a few large eggs as an adaptation to its habitat. The causal and functional explanations in reproductive characteristics among the four populations are discussed in regard to differences in the environmental conditions.