The causes of seasonal changes in numbers of the yellow dung fly, Scathophaga stercoraria (Diptera: Scathophagidae)

ABSTRACT. 1. Counts of adult Scathophaga stercoraria (L.) on cow pats were made in Houghall, County Durham, in 1964 and 1965.
2. A spring peak of numbers was due to adults (overwintered mainly as pupae or larvae) maturing and going to dung to breed. Numbers then dropped, rising to one or more peaks in late June—early July. In 1964 there was then a summer drop in numbers until late September. In 1965 high numbers persisted in summer associated with cooler, wetter weather. Autumn peaks in both years persisted until severe frosts or snow.
3. Mature adults, developed from eggs laid during the spring peak, form the first generation when breeding in mid-late June. No clear generations can be identified after this, due to eggs being laid daily (females have successive gonotrophic cycles). Changes in adult numbers breeding reflects survival of eggs and newly-hatched larvae 5–6 weeks earlier, and lower survival rates of adults in mid-summer compared with spring and autumn.
4. Adult Scathophaga numbers in vegetation rose as numbers on dung dropped. Females dissected to count ovariole tunica dilatations showed that most flies in vegetation were immature, with some parous flies hunting insects to develop the next batch of eggs.
5. Females on dung were dissected and found to range from immature to seven-parous. Those gravid for the first time were grossly under-represented, possibly due to wider dispersal.
6. It is suggested that seasonal changes in this r-strategist cannot be explained simply in terms of generations nor by the occurrence of adult diapause.     

Observations on the biology and ecology of the chrysomelid beetle Gastrophysa polygoni in cereal fields

Abstract. 1. Observations were made on the biology of Gastrophysu polygoni (L.) (Coleoptera: Chrysomelidae) in cereal fields in southern England in 1977, 1978 and 1979. Adults of the overwintering generation emerged in late April/ May and there were usually two generations during the spring and summer months. In 1979 there was some evidence for at least a partial third generation.
2. In the field, the oviposition period was 44 days in the first generation and c . 25 days in the second. Fecundity varied from 586 to 1028 eggs per female and was higher in the first than in the second generation in both 1977 and 1979; in 1978 the reverse was true.
3. Every year there were Iarge losses in the numbers within a generation. However, only one parasite was bred from the developmental stages and a pathogen attacking the larvae was found only in 1977.
4. In some fields and in some years, harvesting and straw burning operations were carried out when eggs were present on the plants. Harvesting did not result in a significant reduction in the numbers of eggs. Burning reduced the numbers of egg batches. The effect was most severe when the straw was spread over the field prior to burning.
5. In the field, significantly more eggs were laid on plants of Polygonum aviculare than on P.convolvulus . In the laboratory, larval survival was higher and duration of development shorter on these two species than on other Polygonaceae found on the farm.     

Mating system and sex allocation in the gregarious parasitoid Cotesia glomerata

The gregarious parasitoid Cotesia glomerata (L.) is often presumed to possess the characteristic attributes of a species that manifests local mate competition (LMC), as it commonly produces female-biased broods. However, our field surveys of sex ratio and laboratory observations of adult behaviour showed that this species is subject to partial local mate competition caused by natal dispersal. On average, 30% of males left their natal patch before mating, with the proportion of dispersing males increasing with an increase in the patch's sex ratio (i.e. proportion of males). Over 50% of females left their natal patch before mating, and only 27.5% of females mated with males emerging from the same natal patch. Although females showed no preference between males that were and were not their siblings, broods from females that mated with siblings had a significantly higher mean brood sex ratio (0.56) than broods from females that mated with nonsiblings (0.39). Furthermore, brood sex ratios increased as inbreeding was intensified over four generations. A field population of this wasp had a mean brood sex ratio of 0.35 over 3 years, which conformed well to the evolutionarily stable strategy sex ratio (r=0.34) predicted by Taylor's partial sibmating model for haplodiploid species. These results suggest that the sex allocation strategy of C. glomerata is based on both partial local mate competition in males and inbreeding avoidance in females. In turn, this mating system plays a role in the evolution of natal dispersal behaviour in this species.Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Factors underlying the late seasonal appearance of the lepidopterous leaf-mining guild on oak

Abstract. 1. Most lepidopterous leaf mining species found on the oak Quercus robur in Britain develop in summer. At this time of year, externally feeding caterpillars remove little leaf area since most of these free living folivores are predominantly spring feeders.
2. I forced Phyllonorycter harrisella (L.) miners to oviposit in spring, then exposed developing larvae to a wide range of leaf damage levels.
3. Leaf miner survivorship and mean female pupal weight were significantly greater in the experimental spring generation on undamaged oak leaves, this being when oak foliage is of the highest nutritional quality.
4. Leaf miner survivorship in all generations is correlated with natural leaf damage levels. Experimental leaf damage also reduces miner survivorship.
5. Leaf damage reduces miner survivorship by increasing the probability of larval death due to wound induced responses.
6. The role of asymmetrical competitive interactions between caterpillars and leaf-miners in determining the late seasonal appearance of miners is discussed. It is suggested that wound induced responses may play an important part in the structuring of phytophagous insect communities.     

Superparasitism in Cotesia glomerata: response of hosts and consequences for parasitoids

Abstract.  1. Superparasitism occurs in Cotesia glomerata (Hymenoptera: Braconidae), a gregarious endoparasitoid of Pieris spp. (Lepidoptera: Pieridae). The response of P. brassicae larvae to superparasitism and the consequences for the parasitoid were examined in order to elucidate the ecological significance of this behaviour.
2. Field surveys of a Swiss population revealed that C. glomerata brood sizes from P. brassicae larvae ranged from three to 158, and both the female ratio and the body weight of emergent wasps correlated negatively with brood size. In the laboratory, single oviposition on P. brassicae larvae did not produce any brood size larger than 62, but brood size increased with superparasitism.
3. Laboratory experiments demonstrated that both naive and experienced female wasps were willing to attack hosts that had been newly parasitised by themselves or conspecifics. Superparasitism reduced survivorship but increased food consumption and weight growth in P. brassicae larvae. Superparasitism lengthened parasitoid development and prolonged the feeding period of host larvae.
4. Despite a trade-off between maximising brood size and optimising the fitness of individual offspring, two or three ovipositions on P. brassicae larvae resulted in a greater dry female mass than did a single oviposition on the host. Thus, superparasitism might be of adaptive significance under certain circumstances, especially when host density is low and unparasitised hosts are rare in a habitat.     

Sex ratio and maternal investment in the multivoltine large carpenter bee Xylocopa sulcatipes (Apoideh: Anthophoridae)

Abstract. 1. Females of the multivoltine carpenter bee Xylocopa sulcutipes (Maa) (Hymenoptera: Anthophoridae) usually excavate a straight tunnel in dead twigs and mass provision a linear array of up to ten brood cells with pollen and nectar. An egg is deposited upon each food mass within one cell.
2. Female offspring generally receive a higher provisioning mass (0.180 ± 0.048 g) than males, a significant difference ( P > 0.001). There are, however, male larvae that receive as much food or more as their sisters or female larvae reared in another nest.
3. There is a close positive association between the size of a mother and the weight of provisions for individual daughters, but not for sons.
4. Female offspring are positioned in the innermost brood cells (Gositions 1, 2 and 3). The sex ratio of the outer cells is either significantly male biased (positions 4–6) or skewed towards males (positions 8 and 9). Positions 7 and 10 are in equilibrium.
5. Solitary females produce a significantly female biased sex ratio ( P < 0.01). Sex ratio in social nests is skewed toward females, but not significantly so ( P < 0.2). There is no significant difference between the sex ratio of solitary and social nests ( P = 0.361). The population sex ratio (pooled sex ratio of all broods produced) is significantly female biased ( P = 0.003).
6. Females kept in the laboratory produced female biased sex ratios whilst unmated females produced all-male broods indicating that insemination and ovarian development are not causally related.
7. The expected sex ratio (ESR) under equal investment, calculated as 1/CR (CR = mean male provision weight/mean female provision weight), is 137.5:117.5 (males:females), and differs significantly from that observed, 104:151 (males:females) ( P < 0.001). The 'Local Resource Enhlancement' hypothesis best explains the female biased sex ratio found in X.sulcatipes and its maintenance in the population.     

Brood-size and sex-ratio variation in field populations of three species of solitary aphid parasitoids (Hymenoptera: Braconidae, Aphidiinae)

Solitary parasitoids of colony-forming hosts may produce quasi-gregarious broods, which favours sibmating on the natal patch and local mate competition (LMC). We examined seasonal variations in brood size and sex ratio in three species of solitary parasitoids of aphids associated with trophobiotic ants. Adialytus arvicola, a parasitoid of Sipha agropyrella on grasses, had the smallest broods (mean=4.2, maximum 19), while Lysiphlebus hirticornis, a parasitoid of Metopeurum fuscoviride on tansy, had the largest broods (mean=32.0, maximum 265). In Pauesia pini, a parasitoid of Cinara piceicola on Norway spruce, broods comprised an average of 5.8 (maximum 41) individuals; brood size increased during early summer when hosts became more available but remained the same later in the season. In all three species the sex ratio at eclosion was female-biased, with broods containing approximately two daughters for each son in both A. arvicola and L. hirticornis; the degree of female-bias was least in P. pini. The sex ratio did not vary with brood size. In A. arvicola, the variance of the number of sons declined with an increase in brood size, consistent with "precise" sex allocation. In contrast, in L. hirticornis, the overall sex-ratio variance was greater than its binomial expectation, while it did not differ from binomial in P. pini. A large proportion of broods contained only sons or only daughters, especially in A. arvicola. An excess of male-only broods is expected if constrained females (which can produce only sons) contribute offspring prior to mating. The number of male-only broods, however, did not differ from the number expected if all females are mated and allocate offspring sexes binomially, except in P. pini. In the latter species, broods with two daughters (as opposed to two sons) exceeded binomial expectations. We propose that P. pini is largely outbreeding, while the strongly female-biased sex ratio in A. arvicola and L. hirticornis is consistent with partial sibmating and LMC. Ant-parasitoid interactions could account for a different population mating structure in the three parasitoid species. Both A. arvicola and L. hirticornis mimic the epicuticular hydrocarbon pattern of their aphid hosts. Eclosing wasps are ignored by honeydew-collecting ants and hence can mate and forage on the natal patch. In contrast, P. pini generally depart the mummy area to avoid attacks by trophobiotic ants and mate off patch.     

Internest sex-ratio variation and male brood survival in the ant Pheidole pallidula

Sex allocation in social insects has become a general modelin tests of inclusive fitness theory, sex-ratio theory, andparent-offspring conflict. Several studies have shown that colonysex ratios are often bimodally distributed, with some coloniesproducing mainly females and others mainly males. Sex specializationmay result from workers assessing their relatedness to malebrood versus female brood, relative to the average worker-relatednessasymmetry in other colonies of their population. Workers thenadjust the sex ratio in their own interest This hypothesis assumesthat workers can recognize the sex of the brood in their colonyand selectively eliminate males. We compared the primary sexratio (at the egg stage) and secondary sex ratio (reproductivepupae and adults) of colonies in the ant Pheidole pallidula.There was a strong bimodal distribution of secondary sex ratios,with most colonies producing mainly reproductives of one sex.In contrast, there was no evidence of a bimodal distributionof primary sex ratios. The proportion of haploid eggs producedby queens was 0.35 in early spring and decreased to about 0.1in summer. Male eggs also were present in virtually all fieldcolonies sampled in July, although eggs laid at this time ofyear never give rise to males. All male brood is, therefore,selectively eliminated beginning in July and continue to beeliminated through the rest of the year. Finally, the populationsex-ratio investment was female-biased. Together, these resultsare consistent with the hypothesis that workers control thesecondary sex ratio by selectively eliminating male brood inabout half the colonies, perhaps those with high relatednessasymmetry.[Behav Ecol 7: 292–298 (1996)]     

Insemination Capacity and Dispersal in Relation to Sex Allocation Decisions in Goniozus legneri (Hymenoptera: Bethylidae): Why Are There More Males in Larger Broods?

Models considering sex ratio optima under single foundress strict local mate competition predict that female bias will be reduced by stochasticity in sex allocation, developmental mortality of males and limited insemination capacity of males. In all three cases the number of males per brood is expected to increase with brood size. Sex ratio optima may also be less female biased when several mothers contribute offspring to local mating groups or if non‐local mating occurs between members of different broods; again more males are expected in larger broods. In the parasitoid wasp Goniozus legneri (Hymenoptera: Bethylidae), sex allocation has only a small stochastic component, developmental mortality is low and non‐siblings are unlikely to develop in the same brood. However, the number of males per brood increases with the size of the brood (produced by a single mother). We investigated the further possibilities of limited insemination capacity and non‐local mating using a naturalistic experimental protocol. We found that limited insemination capacity is an unlikely general explanation for the increase in number of males with brood size. All males and females dispersed from both mixed and single sex broods. Although most females in mixed sex broods mated prior to dispersal, these data suggest that non‐local mating is possible, for instance via male immigration to broods containing virgin females. This may influence sex ratio optima and account for the trend in male number.     

The influence of host size on sex ratios in the parasitoid Diglyphus begini (Hymenoptera: Eulophidae)

Abstract. 1. Sex ratio in the ectoparasitoid, Diglyphus begini (Ashmead), attacking larvae of the dipterous leafminer Liriomyza trifolii (Burgess) in glasshouse marigolds, is best depicted by a model where sex ratio varies as a function of host size.
2. Male D. begini progeny are produced in hosts significantly smaller in size than those producing female progeny.
3. Female wasps attack and oviposit on the largest leafminer larvae available and whether a host is large or small depends upon the size of the other hosts attacked.
4. Diglyphus begini females adjust the thre:shold size for the change-over in sex allocation relative to the size of hosts attacked; however, the sex ratio is maintained at between 60% and 70% males.
5. The patterns observed in these glasshouse studies are not due to sex-specific differential mortality or superparasitism.     

Lake outlet blackflies - the dynamics of filter feeders at very high population densities

Larvae of the blackfly Simulium noelleri aggregated at very high population densities (up to 1.2 × 10⁶ individuals m⁻²) at a lake outlet in Kent, United Kingdom. During 1983 and 1984 their first appearance in these large numbers was in late-June and they completed three summer generations before the overwintering larval generation appeared in October. It is not known where the larvae overwinter but they recolonized the concrete steps of this outlet in May, together with larvae of the S. ornatum group which, however, were not found after completing one generation at this location.
Female flies from the overwintering generation oviposited en masse during late-June and the result was a well-synchronized growth of larvae in the first summer generation. Within this, and other generations, there was A wide range of emergence times for adults; they could emerge early and were then relatively small, or could emerge later and were then relatively large. Females were always larger than males and the emergence of flies was protandrous. A very similar pattern of growth and emergence times was found at a site in Finland.
In all generations, sex ratio was biased to males and the sex ratio in each generation was inversely correlated with population density. This ensured that there were sufficient males emerging, and surviving adult mortality, to guarantee fertilisation of the females which were more expensive to produce.     

A population model for diapausing multivoltine insects under asymmetric cannibalism

Pupa-eating cannibalism occurs naturally in several insect species. Byasa alcinous is a multivoltine species of Red-bodied Swallowtail butterfly found in East Asia, which diapauses as pupa over the winter and whose larvae cannibalize eggs and pupae. We investigate the effects on population dynamics of increasing the asymmetric cannibalistic attack rate of a general insect species in different environmental conditions. We do this by theoretically formulating a generalized system of univoltine and bivoltine larvae over two generations in the spring and summer months. We predict that a lack of resources over the summer can force the population to become entirely univoltine, unless the second-generation bivoltine larvae increase their cannibalistic attack rate, and consume the diapausing pupae from the first generation. The model shows that under extreme environmental conditions, the persistence of univoltine larvae is favoured when faced with the threat of extinction. The model also predicts the conditions for the coexistence of both univoltine and bivoltine larvae, and the degree to which they can both coexist, which decreases as the resource in the second generation increases. This work provides the grounding for future theoretical and experimental consideration of the role of cannibalism in determining insect voltinism.     

A Comparative Study of Sex Ratio and Clutch Size in Gregarious Ichneumonoid Wasps

Some convincing support for sex ratio theory comes from the cross-species relationship between sex ratio and brood size in gregarious bethylid wasps (Hymenoptera: Bethylidae), in which the proportion males declines as brood size increases as predicted under local mate competition. It is unknown how widely such relationships hold within parasitoid wasps as a whole. We assemble a dataset on sex ratio and brood size for gregarious Braconidae and Ichneumonidae. Their sex ratios deviate substantially from those of bethylids; sex ratios differ widely across species; and they are not significantly related to brood size across species. Several factors explain the heterogeneity in sex ratios including across-species differences in mating system, sex determining mechanism, and sexual asymmetries in larval competition and polyembryony leading to single-sexed broods.     

FIELD STUDIES ON THE INFLUENCE OF WEATHER ON EGG-LAYING BY THE CABBAGE ROOT FLY, ERIOISCHIA BRASSICAE BCHÉ. II

Observations on the intensity and duration of egg-laying by Erioischia brassicae Bché were continued in 1953 on the same site as in 1952. Eggs were first observed on 31 April and the spring peak period of egg-laying lasted from 3 to 30 May with a daily average of 40 eggs/plant. The mean total in spring 1953 was 1287 eggs/plant. Summer peaks occurred on 1–20 July (average 17 eggs/day/plant) and 5–11 August (average 21 eggs/day/plant).
In the field the threshold of reproduction was approximately 60° F. The rate of egg-laying was highest at 65–70° F. and it declined during prolonged periods of higher temperatures.
The annual cycle of generations, as shown by egg counts and confirmed by the examination of puparia, consisted of two generations and a partial third. The spring generation developed without a prolonged diapause, and there was complete emergence during the summer. The overwintering puparia came from the second and third generations of larvae.     

Towards a genetic theory for the evolution of the sex ratio. III. Parental and sibling control of brood investment ratio under partial sib-mating

Models of sex ratio evolution under partial sib-mating are investigated in haplodiploids and diploids. In the cases of parental and sibling control of the brood investment ratio between the sexes in diploids, we find that the “unbeatable” investment ratio obtained by W. D. Hamilton (Science156, 477–488) for his local mate competition model corresponds in our inbreeding models to a weak form ESS (evolutionary stable strategy) fixation state and also to the population investment ratio at certain internal equilibria of our models. For haplodiploids, “strong form ESS” values exist under inbreeding in models involving father and sister control. Under brother and mother control, however, the ESS derived from local mate competition models is unstable in our inbreeding models to the introduction of any other investment ratio. We stress important qualitative differences between models involving local mate competition and inbreeding.     

Evolution of sex ratios in social hymenoptera: Consequences of finite brood size

Evolutionarily stable sex ratios are determined for social hymenoptera under local mate competition (LMC) and when the brood size is finite. LMC is modelled by the parameterd. Of the reproductive progeny from a single foundress nest, a fractiond disperses (outbreeding), while (l-d) mate amongst themselves (sibmating). When the brood size is finite,d is taken to be the probability of an offspring dispersing, and similarly,r, the proportion of male offspring, the probability of a haploid egg being laid. Under the joint influence of these two stochastic processes, there is a nonzero probability that some females remain unmated in the nest. As a result, the optimal proportion of males (corresponding to the evolutionarily stable strategy, ESS) is higher than that obtained when the brood size is infinite. When the queen controls the sex ratio, the ESS becomes more female biased under increased inbreeding (lowerd). However, the ESS under worker control shows an unexpected pattern, including anincrease in the proportion ofmales withincreased inbreeding. This effect is traced to the complex interaction between inbreeding and local mate competition.     

Observations on the age, growth, reproduction and food of the dace, Leuciscus leuciscus (L.), in two rivers in southern England

Scales of 497 dace from the river Stour and 573 from the river Frome were used for age determinations. Annuli were laid down in late May to early June at the start of the growth period. Minimal growth occurred from November to April. Male dace grew minimally faster than females; Frome dace faster than those from the Stour. Growth rates in the two rivers were similar to those published for most other European waters. Variation in growth rates in different years was not great, but there were marked differences in recruitment success. Spawning occurred in the second half of March and elaboration of the gonads between September and March. Immature dace have an annual cycle in condition with a maximum in June and a minimum during the winter months. The condition of the mature females is affected by the gonad cycle. The fecundity of Stour dace is represented by the formula:

and for Frome dace by:

Most dace were mature by age IV, but there were more mature II and III group fish in the Frome population. Molluscs and Trichoptera larvae comprised the bulk of the winter diet of Stour dace, and Ephemeroptera nymphs, Simulium and chironomid larvae were the principal constituents during the summer.     

Very high secondary production at a lake outlet

SUMMARY. 1. Larvae and pupae of Simulium noelleri Fried, coated the concrete of parts of an artificial lake outlet in southern England.
2. In the first two (of three) summer generations, development was synchronous and this allowed the calculation of their secondary production by the instantaneous growth method. The production of the two summer generations was, respectively, 229.1 g C m⁻² (7.4 g C m⁻²day⁻¹) and 185.5 g C m⁻² (8.8 g C m⁻² day⁻¹) The contribution of the third summer generation, and the overwintering generation, to annual production would be less than that of the first two summer generations. Nevertheless, annual production will have exceeded 500 g C m⁻² at this site.
3. Larvae are suspension feeders and they captured the rich supply of particulate and dissolved organic material which passed over them after export from the lake. As food is brought to the larvae they only require space for attachment and can thus build up very high population densities (which exceeded 1 × 10⁶ m⁻² on some occasions during the summer). The high population densities result in a high biomass and hence in the high levels of production.     

Activity patterns, diets and interspecific interactions of coexisting spring and autumn breeding carabids: Carabusyaconinus and Leptocarabus kumagaii (Coleoptera, Carabidae)

Abstract. 1. Activity patterns, diets and interspecific interactions of two coexisting carabid beetles, Carabus yaconinus B. and Leptocarabus kumagaii K. et K., were studied in a lowland habitat in Japan.
2. Larvae of C. yaconinus appeared from May to October and fed on earthworms. Those of L. kumagaii appeared from October to the next May and fed on dipteran and lepidopteran larvae.
3. In the adult stages, feeding habits and daily activities of the two species were similar. The beetles fed mainly on earthworms and other animal materials, but various plant materials were also utilized.
4. Direct competition for food items was observed both intra- and interspecifically. Interspecific interactions occurred only in early summer, however, because of the difference in breeding seasons (i.e. spring for C. yaconinus and autumn for L. kumagaii).
5. The difference in seasonal life history patterns between these carabids is a possible mechanism of coexistence.     

Generation-dependent female choice: behavioral polyphenism in a bivoltine butterfly

Climatic and biotic circumstances vary as seasons shift, anddifferent cohorts of multivoltine species are likely subjectedto different selection regimes. The bivoltine butterfly Leptideareali (Réal's wood white; Lepidoptera: Pieridae) appearsduring May and June in central Sweden and has a partial secondgeneration in late July. We manipulated both generations toappear simultaneously and performed laboratory mating experimentsthat showed the presence of a behavioral polyphenism in matingpropensity, which is induced during the developmental stages.Females of the summer generation expressed higher mating propensitiesthan spring generation females. Spring females showed an increasein mating propensity with increasing age, whereas summer femalesaccepted most matings already when they were only 1 or 2 daysold. It is likely that larval time constraints, a lower abundanceof males and a lower risk of accepting a male of their univoltinesister species Leptidea sinapis (wood white), have relaxed selectionon mate discrimination among summer generation females. A majorchallenge for future research is to further investigate thedevelopmental pathways causing the polyphenism and the adaptiveimplications of cohort-dependent behaviors.