Inter- and Intraspecific Effects of Volatile and Nonvolatile Sex Pheromones on Males, Mating Behavior, and Hybridization in Eretmocerus mundus and E. eremicus (Hymenoptera: Aphelinidae)

Eretmocerus species (Hym. Aphelinidae) are solitary parasitoids of Bemisia tabaci (Gennadius). Mate finding and mating behavior of two species, E. mundus and E. eremicus, were studied under laboratory conditions. We used three populations of Eretmocerus: typical arrhenotokous populations of E. eremicus (from USA) and E. mundus (from Spain), and an atypical thelytokous population of E. mundus (from Australia). We studied the intra- and interspecific responses of males to volatile and nonvolatile components of the female sex pheromones, mating behavior, and hybridization between populations and species. In both arrhenotokous populations, males reacted to volatile pheromones by walking toward conspecific virgin females. Males also reacted to nonvolatile pheromones by spending more time on and around patches on leaves of poinsettia plants that had been exposed to virgin females. Males of E. eremicus showed the same reaction to the nonvolatile sex pheromone of E. mundus females, but E. mundus males did not show any reaction to the nonvolatile sex pheromone of E. eremicus. There was no response of males of both species to thelytokous females of E. mundus. In both species three phases were distinguished in the mating behavior: premating, mating, and postmating. The duration of the phases differed between the three populations. Successful copulation between the two Eretmocerus species did not occur. In contrast, we recorded some successful copulations between Australian males and Spanish females of E. mundus, but they did not produce any hybrid females.     

Aspects of the biology of the big-handed crab,Heterozius rotundifrons (Decapoda: Brachyura), from Kaikoura,New Zealand

Abstract

From January 1975 to June 1976, samples of Heterozius rotundifrons A. Milne Edwards, 1867 were taken monthly from the intertidal zone at First Bay, Kaikoura Peninsula (42°25′S, 173°42′E). Small crabs (< 8 mm carapace width) were never common; they were either too cryptic to be collected in quantity, or have a different habitat from larger crabs. Ovigerous females were present in all months except February (in both 1975 and 1976), but generally formed less than 50% of the total sample of females each month. The number of newly deposited eggs (0.75×0.81 mm) carried by females increased with increase in carapace width according to the equation y = -1123.56+102.97x (r ² = 0.8213). Egg development lasted 3–5 months, and egg mortality during this period was almost 10%. The overall sex ratio for the 18-month sampling period was 1846 ♀ : 993 ♂, which suggests that females were approximately twice as numerous as males. However, when crabs were sorted into size classes it was evident that the sex ratio was not significantly different from 1:1 in crabs of 6–12 mm carapace width, whereas females significantly (P<0.001) outnumbered males in the 13–21 mm size range. The right and left chela were approximately equal in length in females of all sizes, but the right chela of large males was greatly enlarged compared with the left, and with the chelae of females of comparable carapace width. Increase in the growth rate of the males’ right chela commenced at a carapace width of approximately 11.0 mm. Since the smallest ovigerous female collected also had a carapace width of 11.0 mm, it is concluded that both males and females attain sexual maturity at this size.     

Polymorphism of Philaenus spumarius L. (Hemiptera: Cercopidae) in the Samsun (Mid-Black Sea Region) populations of Turkey

Variation in the frequency of colour/pattern phenotypes of Philaenus spumarius (L.) is studied in the three local populations of Samsun (Mid-Black Sea Region), Turkey. The data, obtained from the 91 samples including 10,427 adult spittlebugs, 5,340 females and 5,087 males were analysed according to phenotype distributions in the study sites. The adult spittlebugs were collected during eleven months except with April. The mean sample size was 114.6±19.9 (Mean±S.E.) for 91 samplings. Eight different phenotypes of P spumarius were detected in the Samsun populations. Of these, four were non-melanic and four were melanic phenotypes with the lack of the “O” group. In females, non-melanic frequency was over 95%, which TYP (typicus) was the predominant phenotype, but POP (populi) in males. However the phenotype frequency distributions were not different between the sites studied. Although, frequency of females reached up to 85% in some samples, the overall sex ratio did not differ from the 1:1.     

GATA repeats in the genome of Asellus aquaticus (Crustacea,Isopoda)

A 500 bp fragment of Drosophila genomic DNA containing 37 copies of the tetranucleotide GATA was used to probe, by Southern DNA blotting and in situ hybridization, two natural populations of the isopod crustacean Asellus aquaticus collected from the Sarno and Tiber rivers. This species does not have a recognizable sex chromosome pair. In a number of males from the Sarno population chromomycin A₃ staining reveals a heteromorphic chromosome pair. The heterochromosome has two blocks of heterochromatin. After digestion of genomic DNA with six restriction endonucleases and hybridization with the GATA probe, the two populations exhibit different fragment length patterns. No sex-linked pattern was observed in either population. In situ hybridization to chromosomes of males and females from the Sarno population does not reveal any sex-specific pattern of labelling and indicates a scattered distribution of GATA sequences on most chromosomes with some areas of preferential concentration. The heterochromatic arcas of the male heterochromosome are not labelled.by E.R. Schmidt     

Reproductive features of the non-native Siganus luridus (Teleostei, Siganidae) during early colonization at Linosa Island (Sicily Strait, Mediterranean Sea)

In July 2003, the finding of a newly settled population of Siganus luridus at Linosa Island (Sicily Strait, Mediterranean Sea) gave us the unusual opportunity to examine the reproductive condition of a Lessepsian migrant during early phases of colonization. Aspects of gonad morphology, fecundity, atresia and oocyte dynamics were investigated by using 43 pioneer specimens collected in concomitance with their first record in the Pelagie Islands. Ovarian development was consistent with the group‐synchronous type, and testicular organization was of the unrestricted spermatogonial testis type, with cystic spermatogenesis. Both males and females had reached final stages of gonad maturation. The rates of follicular atresia were moderate: out of 17 adult females, 10 individuals did not present atretic oocytes; six exhibited <15.1% of secondary growth phase (SGP) oocytes in α‐atresia, while one female presented 45.7% of SGP in α‐atresia. Fecundity estimates did not diverge from what was observed in a reference population along the Lebanese coast. Absolute fecundity ranged from 115 739 to 740 433 oocytes per female (16.5–24.5 cm L_T). Relative fecundity ranged from 1239 to 3162 oocytes g⁻¹, with a mean of 1885 ± 868 oocytes g⁻¹. Our observations indicated that these early settled siganids are reproductively active at Linosa and suggested the forthcoming of self‐maintaining populations across the central Mediterranean area.     

OBSERVATIONS ON THE DISTRIBUTION,EMERGENCE AND BEHAVIOUR OF CENTRAL AFRICAN CHAOBORIDAE

SUMMARY

The distribution of four species of Chaoboras over selected parts of Zimbabwe is given. The effect of temperature on the duration of the larval life cycle is discussed in relation to the generation time and the lunar periodicity of the adult emergence period. There were apparently two generations of larvae present in the habitat at any one time, although these generations were not distinct due to the variations in the time taken by the larvae to complete development at any temperature. The emergence was synchronized to the lunar cycle, but the actual moon phase at which emergence occurred was variable, as the two populations under observation both changed from new moon emergences to full moon emergences during the study. Some observations on the behaviour of adult Chaoboras edulis are given.     

Annual gametogenic cycle of the freshwater pearl mussel,Lamellidens marginalis (Lamarck, 1819) collected from a perennial lentic habitat of Bangladesh

ABSTRACT

We investigated the annual gametogenic phenology of the freshwater pearl mussel, Lamellidens marginalis (Lamarck, 1819), collected from a lentic habitat at Mymensingh, Bangladesh, using biometry and histology through monthly sampling from August 2015 to July 2016. After biometric measurements, thin slices of dorso-ventral sections were cut from the middle of the mussels for histology to determine sex and level of gonadal maturation. The condition index (CI) ranged from 0.64 (March) to 0.99 (January) over the study period. The CI peaked three times (January, April and July) indicating that the mussels were ripe during these months and subsequent decreases in CI indicated spawning, which was consistent with histology. Both males and females exhibited similar patterns in terms of gonadal development, maturation and spawning activity. It was confirmed that natural populations of L. marginalis spawn throughout the year with remarkable temporal variations, except during December when the surface water temperature reaches annual minima (16.5°C). Highest spawning activity of L. marginalis was noted during February–March, May and July–November. The data obtained in the present study could be useful for the conservation managers of this commercial species by allowing harvesting of better quality mussels to be timed without interrupting major spawning activity.     

Sex ratio and spatial pattern of males and females in the dioecious sandhill shrub, Ceratiola ericoides ericoides (Empetraceae) Michx

Sex ratio, size, age, and spatial pattern were investigated for males and females of the dioecious shrub Ceratiola ericoides ericoides (Empetraceae) Michx. within seven mapped populations in Georgia and South Carolina, USA. Among the sites studied, two are regularly burned and one site long-unburned. Age was estimated from node counts of individual shrubs. Only one (fire-suppressed) population showed a female-biased sex ratio, while all others did not differ significantly from 1:1. Mean age estimates did not differ between sexes at any site nor did mean shrub canopy diameter. Bivariate Ripley’s K analysis with a null hypothesis of random labeling was used to investigate whether any of the mapped populations exhibited spatial segregation of the sexes (SSS). No population showed strong evidence of SSS. Rather all sites but one showed males and females to be associated (though not significantly) at a scale of 1–10 m. At a scale of 10–35 m male and female shrubs were located randomly with respect to one another at all sites.     

Genetic variability, body characteristics and reproductive parameters of neighbouring rural and urban common kestrel (Falco tinnuculus) populations

We tested the genetic and ecological differences between neighbouring urban and rural populations of common kestrels (Falco tinnuculus) in southern Bohemia. The aims were to (1) assess the genetic variability of the studied kestrel populations using microsatellite markers, (2) check the genetic relatedness of individuals within the urbanization gradient, and (3) compare possible gradients of body characteristics and reproductive parameters on the urbanization gradient. The mean expected allelic polymorphism did not differ among the studied populations, which were not genetically separated (F _ST  = 0.0003, P = 0.781). Also, an individual assignment test did not show a separation of these populations. Urban kestrels that bred in the city centre were indicatively more related than others, and no relationship was found in the rural kestrel population. Kestrel females were heavier towards the city centre, but males did not show this relationship. Nest distance from the city centre had no significant effect on any of the tested reproductive parameters. Our results do not support the notion of genetic differentiation between rural and urban kestrels, but revealed trends in body characteristics and genetic relatedness along the urbanization gradient.     

GENETICS OF PRECALLING PERIOD IN THE ORIENTAL ARMYWORM,MYTHIMNA SEPARATA (WALKER) (LEPIDOPTERA: NOCTUIDAE), AND IMPLICATIONS FOR MIGRATION

Migration in insects usually occurs prereproductively so the precalling period, between emergence and the first release of pheromone, determines the number of nights over which females of migrants such as Mythimna separata can express their migratory potential. Precalling periods (PCP's) ranged from 3 to 21 nights in F₁ females from insects collected in the field in eastern China. As technical problems precluded the use of a sib-analysis design, the genetic basis of this variation was investigated by analysis of the correlation between daughters' and mother's PCP's and by selection in isofemale lines followed by reciprocal crosses. The results of these analyses indicated that PCP is strongly influenced by genes located on the X chromosome, inherited from the male parent in Lepidoptera. Genetic regulation of migratory potential, of which PCP is a crucial component, can be expected to evolve in tropical species such as Mythimna separata, whose larval habitats are associated with unpredictable tropical rainfall. This species has no preimaginal diapause but makes regular seasonal incursions every spring and early summer to high temperate latitudes at which it cannot overwinter. Where migrants are carried by winds, as occurs in these migrations, the predominantly poleward winds at this time of year must create a genetic cline across latitude, in which migratory potential of the summer populations founded by the migrants increases with increasing latitude. We suggest that X-linkage of genes influencing PCP may reduce the genetic load associated with this polymorphism by reducing the frequencies of inappropriate PCP genotypes in these summer populations. Thus, in autumn when winds are predominantly northerly, fewer individuals at any latitude will lack the capability to make the return journey southward. Some support for this model is provided by the observed distributions of PCP's of F₁ females from moths collected in the field in central and northern eastern China. If X-linkage of genes influencing PCP contributes to the stability of these migratory cycles, it can be expected in other species with similar migratory strategies.     

Grouping and Inhibition of Oogenesis in Two Strains of Acanthoscelides obtectus Say (Coleoptera Bruchidae) of Different Geographical Origin-Specificity of This Inhibitory Effect

Summary

Reproduction of A. obtectus females originating from Rubona (Rwanda) was inhibited after grouping (2 females, or 1 male—1 female). Only a few females produced mature oocytes; vitellogenin was synthesized, released into the haemolymph, but not incorporated into the oocytes. When females produced mature oocytes, their ovarian production was lower than in isolated virgin females. After pairing of a male and a female, 38% of the females mated, but mating under these conditions did not stimulate oogenesis.

Oogenesis of females originating from Tours (France) was only slightly inhibited by the presence of males or females of the same origin. “France” males and females inhibited the reproduction of “Rwanda” females under grouping conditions. However, “France” females showed only a slight sensitivity to grouping with “Rwanda” bruchids. In the presence of Phaseolus vulgaris seeds, grouping had no effect and all females, whatever their origin, produced mature oocytes. Inhibition weis relatively specific as cohabitation with bruchids of other genera (Callosobruchus maculatus-Zabrotes subfasciatus) had no effect. Cohabitation with two species of the same genus (Acanthoscelides obvelatus—A. argilaceus) inhibited oogenesis of A. obtectus “Rwanda” strain. The significance of this regulation and its importance in the maintenance of populations in nature are also analyzed.     

Population estimates,trends and habitat associations of breeding Lapwing Vanellus vanellus,Curlew Numenius arquata and Snipe Gallinago gallinago in Northern Ireland in 1999

Capsule Declines in the breeding populations of Snipe, Lapwing and Curlew were recorded between April and June 1999 and compared with previous estimates in 1987.

Aims To compare populations of non-coastal breeding waders between 1987 and 1999.

Methods In 106 2 km × 2 km square tetrads, observers recorded the number of breeding pairs of waders and habitat details on 1:10 000 scale maps.

Results A significant decline of c. 60% for Lapwing and Curlew, and a non-significant decline of 30% for Snipe was recorded over 12 years. Concentrations of these species were found in County Tyrone, but Counties Antrim, Down and Armagh supported few breeding pairs of any species. Very few pairs of any species were recorded on improved grassland despite its widespread availability.

Conclusion A successful conservation strategy for these species must address the wider countryside and not just key sites. Intensive pastoral farming in upland and lowland areas and activities such as drainage and peat extraction will further reduce the suitability of open habitats for these wader species.     

Life stages of the non-native Ommatoiulus moreleti (Lucas, 1860) (Julida,Julidae) in Australian small grain systems

1. The Portuguese millipede, Ommatoiulus moreleti (Lucas, 1860), is increasingly a pest of grains crops in Australian agricultural ecosystems. With the rapid uptake of minimum-tillage practices, habitat quality has been enhanced for several species of soil dwelling invertebrates, in particular O. moreleti.
2. To understand the population dynamics of O. moreleti in Australian grains crops, populations were sampled at multiple sites for more than 2 years using pitfall traps and cardboard roll traps. Specimens were dissected to investigate reproductive status and developmental stage.
3. Millipede trapping rates varied between sites and across the year, and tended to be lower when soil water content (m³/m³) was high. Both sexes were active year round, however females were relatively more abundant when ground temperatures were higher.
4. Males in a copulatory state and females with mature eggs were collected year round although females lacking mature eggs were more common when ground temperatures were higher and the soil was drier, and female egg load was higher under cooler conditions. Females at a stadium 9 or later carried mature eggs and copulatory males tended to be from stadium 8 or later. Different developmental stages could be recovered at all times of the year.
5. These findings indicate persistent populations of O. moreleti across fields throughout the year, with a key breeding period in autumn but likely opportunistic breeding all year. Control of millipedes will need to focus on increasing the impact of biological agents, decreasing suitable habitat for millipede breeding and intensive control at susceptible crop periods.

     

Reproductive strategies of the exotic Pharaoh's ant,Monomorium pharaonis (L.) (Hymenoptera: Formicidae) in Brazil

Summary

The production of sexual forms in laboratory colonies of Monomorium pharaonis was followed over a 19-month period. Cycles of production occurred at intervals of 4 months, with male investment inversely proportional to the number of gynes produced. There was a tendency for colonies to produce only males or females as total sexual production increased. Sperm counts of known-aged males suggest that they mate twice, in contrast to European populations. Sex-ratio was heavily female-biased, contrary to predictions for polygynous ant species.     

Characterization of age and cuticular hydrocarbon variation in mating pairs of house fly,Musca domestica,collected in the field

House flies, Musca domestica L., were collected in copula over two summers from six dairies located in three climatically distinct regions in the U.S.A. southern California, Minnesota and Georgia. Ages of males and females from a total of 511 mating pairs were estimated using pterin analysis. Cuticular hydrocarbon profiles and gonotrophic ages of females also were evaluated. Mean age of mating males ranged from 54 to 102 degree‐days (DD) (4–10 days based on field air temperatures), depending on the farm. Very young males (< 10–20 DD) and old males (> 200 DD) were rare in mating pairs. Mean female age at mating ranged from 20 to 46 DD (2.5–4 days). All mating females had eggs in the early stages of vitellogenesis and 99.2% were nulliparous. However, some older and parous females were collected, demonstrating that re‐mating can occur in the field. Head width measurements of mating pairs suggested that assortative mating by size did not occur. The cuticular hydrocarbon profiles of females were determined, with emphasis on (Z)‐9‐tricosene (muscalure). Overall, only 55% of mating females had detectable amounts (> 4 ηg per fly) of (Z)‐9‐tricosene. Of the females that had detectable (Z)‐9‐tricosene, variation in amount per female was high in all fly populations, and thus was not statistically related to the size or age of the mating female. The proportion of mating females with detectable levels of (Z)‐9‐tricosene varied by geographic region. Seventy‐one, 63, and 27% of females from southern California, Minnesota and Georgia had detectable amounts of (Z)‐9‐tricosene. Principal components analysis of the eight most abundant hydrocarbons from mating females, by state, revealed state‐level distinctiveness of hydrocarbons in house fly populations, which may reflect genetic variation associated with environmental stresses in those geographical zones.     

Tree wetas (Hemideina crassicruris) (Orthoptera: Stenopelmatidae) of Stephens Island,Cook Strait

Abstract

Tree wetas (Hemideina crassicruris) were studied in kohekohe (Dysoxylum spectabile) forest on Stephens Island, Cook Strait, between 5 and 17 January 1978. Mark and recapture observations suggested a population density of at least 5300 wetas ha^?1 and a biomass of 42 kg ha^?1. Activity of wetas increased with temperature and humidity. Tuataras are important predators on Stephens Island and probably deter wetas from moving onto the forest floor. The enlarged head and jaws of male wetas are associated with head-on fights for occupancy of galleries. Single, adult males occupy galleries with a harem of up to 9 females. Stridulation was associated with agonistic encounters and was most frequent shortly before dawn. Seventeen percent of males and 42% of females were damaged. Moulting, copulation, and oviposition were observed, and females carried up to 96 mature eggs. Stephens Island tree wetas reach larger sizes overall, and males have relatively larger heads, than in other populations of the Cook Strait region. The number and size of galleries and the presence or absence of predators may be important determinants of weta population density, sex ratio, harem formation, and activity on the ground.     

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.     

Studies on the embryogenesis of Aulocara elliotti (Thomas) (Orthoptera, Acrididae). II. Developmental variability and the effects of maternal age and environment

The stages of diapause embryos of A. elliotti reared in the laboratory were described, and the lengths of embryos of stage 19 were compared with a second group of diapause embryos of advanced ages. Collections of diapause embryos of the same species were made from wild populations during the autumn and winter of two consecutive years. The stages of embryos present in these samples were found to vary widely. The mean lengths of embryos in stage 19 from different populations were compared and found to differ with statistical significance. The numbers of retarded embryos in different populations were found to vary significantly between populations of diapause embryos. In two populations, mean lengths of embryos of stage 19 were found to differ from one year to the next. Post-diapause embryos were collected from wild populations in the spring of two consecutive years and their stages described. No significant difference was found in the incidence of retardation in post-diapause samples. Results of these studies and those of other authors indicate that there may be wide developmental variation between populations of acridid embryos as well as in one population in two different years. Rearing adult A. elliotti in varying numbers under two different environmental conditions was found to produce significant effects upon the rate of development and variability of their embryonic progeny. Embryos laid by young females were found to be more variable, in general, and to develop more slowly than those laid by older females.     

Difference in age,growth, and sexual demography of black‐spot tuskfish,Choerodon schoenleinii (Valenciennes, 1839), in two adjacent populations,Ryukyu Archipelago,southern Japan

To better understand the plasticity of life history traits in the blackspot tuskfish, Choerodon schoenleinii (Valenciennes, 1839), the characteristics of the population around the Yaeyama Islands (24°N, 124E) were examined and compared to those around Okinawajima Island (26°N, 128E) that had been investigated in a previous study. Age and growth of the Yaeyama population were examined based on 103 specimens collected at fish markets between 2006 and 2016. Specimens included 83 females (25.2–69.0 cm total length [TL]), and 20 males (43.1–71.8 cm TL). Ages determined from sectioned otoliths ranged from 1–9 for females, and 4–15 for males. Values for von Bertalanffy growth functions were L_t = 74.2 {1?exp[?0.23 (t + 0.38)]}, and the growth of the Yaeyama population was significantly faster than that of the previously studied population. Sexual demography of the two populations was compared using body length data on landings measured at the fish markets. In the Yaeyama population, females and males ranged from 24–65 cm TL and 39–75 cm TL, respectively; length at 50% individual sex change size was estimated at 54.7 ± 0.56 cm (±95% C.I.). In contrast, in the Okinawajima population, females and males ranged from 16–65 cm TL and 30–75 cm TL, respectively; meanwhile, 50% sex change size was estimated to be 50.0 ± 0.25 cm. There were thus significant differences in the size at sex change between the two populations. This difference may be related to the difference in population density between the sites.     

Interpopulation differences in pupal size and fecundity are not associated with occurrence of outbreaks in Epirrita autumnata (Lepidoptera, Geometridae)

Abstract.

• 1 Among-population differences in pupal mass were studied in a geometrid, Epirrita autumnata. Some Epirrita autumnata populations regularly reach outbreak densities while others are never known to do so. Because adults do not feed, pupal mass of females correlates strongly with fecundity.
• 2 Larvae were collected from twelve field sites. Ten of our sample populations originated within the outbreak range of the species and represented different phases of outbreaks. Two populations originated outside the outbreak range.
• 3 Pupal mass of field-collected E. autumnata varied significantly among populations. The peak phase populations had the smallest pupae and the biggest were found in low density populations outside the outbreak range.
• 4 Offspring of moths from each population were reared under identical conditions in two larval densities. Significant differences were not found in pupal mass among populations. That is, the inherent size, correlated with fecundity of moths, was not different between populations originating within and outside the outbreak range, nor among collections from different densities or phases of the outbreaks.
• 5 Rearing density did not interact in a consistent way with population.
• 6 As far as size and fecundity are concerned, the results do not support Chitty's hypothesis that differences in genetic composition of the population at low and high density phases generate cyclic fluctuations of population density.
• 7 Because no hereditary or maternal differences were found in size and fecundity between E.autumnata originating within and outside the outbreak range, variation in reproductive capacity cannot explain why outbreaks occur only in some populations.