Reproductive characteristics of the flower breeding Drosophila hibisci Bock (Drosophilidae) along a latitudinal gradient in eastern Australia: relation to flower and habitat features

For many insect species, egg and larval substrate characteristics are significantly correlated with interspecific differences in female reproductive allocation and egg size-number tradeoffs. We tested the hypothesis that a similar pattern occurred within the Australian drosophilid, Drosophila hibisci, that is restricted throughout its life cycle to flowers of species in the genus Hibiscus. These plants occur as small, isolated, normally monospecific stands that should facilitate differentiation of the fly populations in relation to specific oviposition and larval substrates. Data from 38 sites ranging from 20.8̀ to 34.4̀ S latitude in eastern Australia indicated no relationship between female body size, egg size, or ovariole numbers and floral size or mass among four species of Hibiscus. However, the flies did show a latitudinal cline in ovariole number that was independent of floral variation. Females averaged 15–20 ovarioles per female in the south (32–34̀ S latitude) and 10–12 ovarioles in the north (21–22̀ S latitude). The increase in ovariole number with latitude was due to divergence in the ovariole number of the largest females. In contrast, small females in the north and south had the same number of ovarioles. Reproductive allocation of female flies in the northern region was less than females in the southern region. The latitudinal divergence in ovariole number was not associated with habitat differences (density of trees, density of flies and beetles), nor with differences in floral characteristics (flower weight, petal length, yeast species present). Short term weather patterns in daily temperature and rainfall preceding collections pardy explain the variation in ovariole number. These observations in conjunction with preliminary genetic results suggest the cline is associated with genetic differences that interact with environmental determinants such as the temperature during larval development.     

Habitat quality and reproductive investment in aphids

ABSTRACT.

• 1 Genetically identical vetch aphids, Megoura viciae (Buckton), showed a weak but significant positive relationship between reproductive investment (number of ovarioles) and weight, here used as an indicator of habitat quality.
• 2 Apterae reared on poor quality plants showed the same range in numbers (twelve to twenty-two) but on average fewer ovarioles than those reared on high quality plants.
• 3 The variability in ovariole number was determined prior to birth.
• 4 Under constant conditions the age of a mother does not affect the range of ovariole numbers or the proportion of her offspring in each ovariole class.
• 5 Apterae reared on poor quality host plants gave birth to proportionately fewer offspring with a high ovariole number than apterae reared on high quality plants.
• 6 Winged individuals gave birth to proportionately more offspring with a high number of ovarioles than apterae.
• 7 The control and ecological significance of the intramorph variation in reproductive tactics shown by many species of aphids is discussed in the light of these results.

     

Age‐related changes in the development of oocytes and testicular follicles after emergence in Stavsolus japonicus (Plecoptera)

Abstract Female stoneflies oviposit several times during the adult stage of their life cycle. The number of eggs within the deposited egg masses decreases at successive ovipositions. To clarify the reason for this decrease and to determine the conditions of testicular follicles, the patterns of development of oocytes and testicular follicles on different days after emergence are investigated in the systellognathan species Stavsolus japonicus (Okamoto) (Perlodidae). The size of the mature oocytes in the ovariole peaks a few weeks after emergence but decreases to the lowest level by 35 days after emergence. Several maturing oocytes can be observed in the ovarioles of individuals a few weeks after emergence but only one mature oocyte is observed at 35 days after emergence. The decreased number of eggs laid per mass by older individuals may therefore be due to the lower maturation of all the ovarioles.     

Effect of biotic and abiotic factors on reproductive parameters of the synovigenic endoparasitoid Venturia canescens

Abstract.  The main effects and interactions of adult age, access to food and host deprivation, on the egg load of Venturia canescens (Gravenhorst) (Hymenoptera: Ichneumonidae), a larval parasitoid of Ephestia kuehniella (Lepidoptera: Pyralidae) and other pyralids, were studied in the laboratory. Intraspecific variation in the number of ovarioles was also studied. There was a positive and significant correlation between wasp size and both egg load and ovariole number, with the reproductive system of large wasps containing significantly more ovarioles and mature eggs than small wasps. Newly emerged adult wasps contained 27 ± 2.4 ovulated (mature) eggs in their lateral oviducts. Access to food and host deprivation were the only conditions under which egg load increased with parasitoid age. Wasps that had access to hosts immediately after emergence showed a significant decline in their egg complement, irrespective of food presence. Under conditions of both host and food deprivation, there was practically no alteration of egg load with parasitoid age. A three-way analysis of variance revealed that egg load varies significantly with food or host access but not with parasitoid age. All interactions among the three factors were significant. It is confirmed that there is no egg resorption in V. canescens and that egg production stops in the absence of food.     

Egg envelopes of Baetis rhodani and Cloeon dipterum (Ephemeroptera, Baetidae): a comparative analysis between an oviparous and an ovoviviparous species

A comparative ultrastructural investigation on the eggshell (vitelline and chorionic envelopes) has been carried out in the nymphs of two mayfly species encompassed into the Baetidae, namely Baetis rhodani and Cloeon dipterum. During oocyte differentiation in the meroistic telotrophic ovarioles, gametes are connected to discrete nurse cells by trophic cords. In B. rhodani, each ovariole contains several eggs arranged in sequence, whereas in C. dipterum each contains a single egg. Follicle cells are competent for vitelline and chorionic envelope synthesis. Baetis rhodani is an oviparous species and the chorion is fairly thick, formed by an alveolate endochorion and a fibrillar exochorion delimited by a honey‐comb roof. Cloeon dipterum stands out among Ephemeroptera for being ovoviviparous. In B. rhodani, ovulation starts in the older nymphs with dark wing‐pads, whereas in C. dipterum choriogenesis ends in the imaginal stage. Here the chorion is very thin and laid eggs hatch almost immediately, allowing the larvulae to move out. The maturation of a single egg per ovariole is synchronized with the achievement of the adult stage. The acquisition of ovoviviparity has led to remarkable changes in the ovariole organization along with a simplification of the eggshell structure.     

Effects of female body size on lifetime fecundity of <Emphasis Type="Italic">Monochamus urussovii</Emphasis> (Coleoptera: Cerambycidae)

To analyze the relationship between female body size and lifetime fecundity (LF) in Monochamus urussovii (Fischer), 17 pairs of adults were reared on Picea jezoensis (Sieb. et Zucc.) Carrière twig sections and were allowed to oviposit on Abies sachalinensis (Fr. Schmidt) Masters bolts at 25 °C and 16:8 h light:dark. The means of life span and LF were 42.5 days and 28.2 eggs for fertile and sterile females combined, respectively. The mean lengths of preoviposition and oviposition periods (OPs) of fertile females were 12.1 and 28.1 days, respectively. The potential fecundity was estimated to be 72.1. Path analysis showed that female body size, i.e., mass and elytral length, had positive effects on LF through the oviposition rate but not through the OP. Ovariole number was 24 and did not vary according to body size, indicating a higher egg productivity per ovariole in large females than in small ones. Higher egg productivity per ovariole in large females is true for each of three other synovigenic lamiine beetles with an almost fixed number of ovarioles. The body size-dependent egg productivity per ovariole is considered to be one of the reasons for the body size–LF correlation in synovigenic species with a fixed ovariole number.     

Diet, fecundity and egg size in some polyphagous predatory carabid beetles

Egg size was measured at different rates of egg laying in three polyphagous carabid species, known to be useful predators of cereal aphids; the small Bembidion lampros Herbst and the medium-sized Pterostichus cupreus L. and P. melanarius Illiger. Variations in fecundity, as well as the ability of the medium-sized species to also build up fat reserves, were obtained when beetles were subjected to different dietary regimes consisting of aphids, or foods with a lower or higher protein content. Egg size was found to be dependent on the rate of egg laying within a species. A diet of cereal aphids appeared to be adequate for egg production in these polyphagous carabids, but female P. cupreus were unable to build up fat reserves when they ingested aphids contaminated with the aphicide pirimicarb. Beetles were able to devote resources to more and larger eggs (B. lampros), or to larger eggs and/or fat reserves (P. melanarius/P. cupreus) when given access to a carbohydrate-rich food with low protein content. The highest rate of egg laying was obtained when female P. cupreus and P. melanarius were given a more varied diet at frequent intervals; including regular shifts between unsprayed aphids, carbohydrate-rich food and protein-rich maggots. Within the varied diet treatment a negative relationship was obtained between egg size and egg number among similar-sized individuals of P. cupreus and P. melanarius; females producing the largest number also laid the smallest eggs. Egg size affected larval survival, since first instars hatching from large eggs were found to survive longer than those hatching from small eggs. The influence of differences in food intake on reproduction, maintenance metabolism, and survival of fieldinhabiting carabids is discussed.     

Genetics of egg production in Drosophila sechellia

Drosophila sechellia, an island endemic that specializes on a single host plant, has a lower rate of egg production than its generalist sister species D. melanogaster, D. simulans, and D. mauritiana. Earlier work showed that part of this difference in egg production was due to a reduction in the number of ovarioles in D. sechellia relative to its sister species. Here, I extend this earlier work by genetically analyzing the difference in egg production between D. sechellia and D. simulans. In all, 10 genetic markers were used in several interspecific backcrosses to identify chromosome regions that affected the rate of egg production. While previously mapped factors affecting ovariole number appear to impact the rate of egg production, new, non-ovariole factors were also identified. Overall, the difference in egg production between D. sechellia and D. simulans appears to be a polygenic trait. The relationship between these factors and genes involved the adaptation of D. sechellia to its host plant is not yet clear. The data are consistent with the hypothesis that decline in egg production is, in part, a negative pleiotropic effect of genetic changes required for host specialization in D. sechellia, although finer-scale genetic analysis of both traits is needed to truly test this hypothesis.     

A test of three hypotheses for ovariole number determination in the grasshopper Romalea microptera

Ovariole number in insects determines potential fecundity and can be influenced by genes, environmental conditions during development and parental effects. In the present study, three hypotheses are tested for ovariole number determination in the grasshopper Romalea microptera (Beauvois), which exhibits both intra‐ and interpopulation variation in ovariole number. In hypothesis 1, variation in ovariole number is a result of genetic variation. In hypothesis 2, ovariole number is influenced by nutrition during development. In hypothesis 3, ovariole number is influenced by maternal nutritional status. Females from four treatments are compared: low‐food, high‐food, daughters of low‐food, and daughters of high‐food. There is a relationship between parent and offspring ovariole number despite different environments, supporting hypothesis 1. Also, ovariole numbers are slightly, but significantly lower in individuals fed a low‐food diet compared with a high‐food diet, supporting hypothesis 2. Hypothesis 3 is not supported: starved and well‐fed females produce eggs of similar mass, as well as offspring with similar numbers of ovarioles, suggesting that the nutritional status of mothers does not influence offspring mass or offspring ovariole number. The results imply that genetic variation and developmental conditions determine ovariole number in this species but maternal environment does not. These results conflict with previous studies of ovariole determination in grasshoppers and locusts.     

Ovariole development in Staphylinidae (Coleoptera)

Summary

Twenty percent of the species from 88% of the genera, which comprise the British Staphylinidae fauna, were dissected and their ovaries attributed to one often defined types. Three-quarters of the species studied possessed six ovarioles in each ovary although the eight species of Bledius examined were found to have only a single ovary, a condition previously thought to be unique to the family Scarabaeidae within the Coleoptera. The type of ovariole development within most staphylinid subfamilies is remarkably constant. Any inconsistencies are described. Species of Aleochara, known to have parasitic larvae, were found to have a larger number of ovarioles per ovary than any other member of the Staphylinidae examined. They also exhibited a greater variation in ovariole number within each species which is correlated with the size of the adult beetle.     

Structure of the ovaries in larvae and mature females of euholognathan stoneflies (Plecoptera)

The morphology of ovaries, oviducts and egg capsules in four species of euholognathan stoneflies was investigated. The characteristic features found were as follows: (i) numerous, long ovarioles, that open individually to the extensively folded, lateral oviducts; (ii) a thin, morphologically undifferentiated chorion; (iii) a thick gelatinous layer (extrachorion) which acts as an adhesive layer fixing the eggs to the substrate. Additionally, in the larval ovariole of Leuctra sp. the terminal filament anlage and clusters of germ cells have been found. These observations are in agreement with the classification of stonefly ovaries as primary (true) panoistic.     

The influence of a shortage of hosts on the specificity of oviposition behaviour in species of Dacus (Diptera, Tephritidae)

ABSTRACT. When deprived of their preferred hosts for up to 16 days prior to behaviour assays, three relatively specialized tephritid flies, D. cacuminatus (Hering), D. cucumis French and D. jarvisi (Tryon), retain strong preferences for their particular hosts. Moreover, in the absence of the preferred hosts, D. cacuminatus , and to a lesser extent D. cucumis , will not accept fruits that are outside their normal host range. D. jarvisi readily accepts other fruits in the absence of its preferred host, Planchonia careya , but in common with the other two shows no increase in the number of eggs laid with increasing periods of deprivation. None of these species accumulate oocytes when deprived of fruits.
In contrast, the highly polyphagous D. tryoni Frogg. lays more eggs and oviposits readily into a previously unacceptable fruit after 4 days without access to hosts. This change in behaviour is associated with a marked increase in the number of mature eggs carried by females.
These results suggest an important difference between the generalist and more specialized species in the physiological control of oocyte maturation. In D. tryoni there appears to be no inhibition of oocyte development once the primary follicle has matured, whereas in the other species each ovariole contains no more than one mature egg.
In D. tryoni , the change in behaviour which accompanies this increase in egg load, may influence its propensity to accept unusual fruits, and may help to explain its rapid acceptance of most cultivated fruits introduced to Australia.     

Egg size manipulation in the migrant skipper, Parnara guttata guttata (Lepidoptera: Hesperiidae), in response to different host plants

Female adults of the migrant skipper, Parnara guttata guttata, lay different-sized eggs on different host plants in different generations. Moreover, P. g. guttata maintains large egg size variation either in the population or in the individual. Why such phenotypic plasticity in egg size is maintained has not been clarified. In the present study we performed a series of experiments to verify whether or not females of P. g. guttata discriminate between the different host plants, i.e., rice plant Oriza sativa in the first and second generations and cogon grass Imperata cylindrica in the third (overwintering) generation and manipulate egg size accordingly on these host plants. When a caged female was allowed to lay eggs alternatively on soft-leafed rice plant and tough-leafed cogon grass, the size of the eggs laid on cogon grass was significantly larger than that on rice plant. When a caged female was allowed to lay eggs on the two host plants that were supplied on alternate days, the size of eggs laid on cogon grass was also significantly larger than that on rice plant. A preliminary experiment using crude extract from the plants suggested that females did not manipulate egg size in response to chemical stimulants alone. The skipper female was able to lay different-sized eggs instantaneously after assessing the host plant. However, the reaction norm to different host plants differed among females. Eggs of various sizes were matured in each ovariole, and egg size variation at the lowest part of the ovariole ranged from the size of the larger eggs laid on cogon grass to that of the smaller eggs laid on rice plant, although how the female chose the appropriate-sized egg at each occasion is unknown.     

Oogenesis and ovarian morphology in pollinating and non-pollinating fig wasps: evidence from adult and immature stages

Hymenopteran insects have meroistic polytrophic ovaries characterised by trophocytes associated with oocytes inside the follicles. In pro-ovigenic species, all oocytes mature before emergence and no trace of oogenesis is visible in adult females. Pro-ovigeny is a rare condition among Hymenoptera, but common in pollinating fig wasps. In the present investigation, we studied adult and pupa females of three fig wasp species with different trophic strategies. We demonstrated that females of Pegoscapus aerumnosus and Idarnes spp. have an unusual ovarian organisation (i.e. each ovariole has only one mature egg and no oocyte) that has led to misleading interpretation of fig wasp reproductive anatomy. The ovaries of these studied species have several ovarioles, recognisable by the presence of nuclei of tunica propria cells surrounding them. Each adult wasp ovariole had one mature egg. None of the pupae had mature eggs, but all of them had follicles with oocytes in different developmental stages. The studied fig wasps are pro-ovigenic, irrespective of their trophic strategy, since there were no signs of ovigeny in adult females. We discuss ecological and phylogenetic factors that could play a role in fig wasps reproductive strategies.     

Ovary structure in a presocial insect,Elasmucha grisea (Heteroptera,Acanthosomatidae)

First generation egg clusters of Elasmucha grisea are more closely guarded than second generation clusters. The ovaries of this species are structured to enhance this behavior. The population of E. grisea from S-W Poland breeds in the spring (May–June) and late summer (July–August). The second generation clutches contain fewer eggs and are destroyed 3–4 days after oviposition by predators and parasitoids.The ovary structure in the studied species differs from that found in other Heteroptera. The average number of ovarioles per ovary is 24 while in the other investigated species the number of ovarioles per ovary is 6–7. Lateral oviducts are elongated and the ovarioles are arranged in a pennate pattern. Each ovariole contains only one growing ovarian follicle. Differentiation of the ovarioles and ovarian follicles is synchronised thus enabling simultaneous oviposition. A comparative analysis of the ovary structure during the life cycle, particularly the presence of atresive ovarian follicles in the ovarioles of egg- and nymph guarding females, as well as the shape and structure of the apical part of the tropharium all support the hypothesis of cooperation between females in E. grisea. A similar ovary structure has been observed in the Coccoidea (Hemiptera, Homoptera) which indicates presocial behavior.     

Reproductive characteristics of Drosophila hibisci in the Northern Territory, Australia

Reproductive traits of Drosophila hibisci collected at 18 sites in the Northern Territory (NT) of Australia in May, 1998, as well as at two sites in north Queensland, in June, 1998, were compared to those from earlier work on a cline in ovariole number in D. hibisci along the east coast of Australia. The flies in the NT were considerably smaller, but had more ovarioles than comparably-sized flies on the east coast. Although the flies on the east coast showed an increasing number of ovarioles in populations at increasing distances from the equator, these new populations, both on the east coast and in the NT, reversed this trend, producing a generally U-shaped pattern of ovariole number with latitude among all populations. The northernmost and southernmost populations allocate more to ovariole numbers than populations in intermediate latitudes. Ovariole number is closely related to body size of females in all populations, but the regression coefficient is small at intermediate latitudes and increases at the northern and southern ends of the distribution. Egg volumes primarily varied with body size of the female (positive) and number of ovarioles per female (negatively), producing a generally inverted U-shaped pattern of egg volumes with latitude. Reproductive allocation patterns, but not thorax size or ovariole number, varied significantly in two samples taken 10 days apart at one NT site. This variation probably results from environmental differences across generations of developing larvae and is consistent with our earlier suggestion of substantial effects of the environment, primarily rainfall and temperature, on reproductive allocation in D. hibisci.     

两种寄生蜂的卵巢结构与卵成熟特点初步观察

对斜纹夜蛾的内寄生蜂双斑侧沟茧蜂MicroplitisbicoloratusChen和南美斑潜蝇的外寄生蜂潜蝇姬小蜂Diglyhusisaea(Walker)的卵巢结构进行了研究。双斑侧沟茧蜂的2个卵巢管底部膨大有大量的卵,通过凹陷的颈部与萼区相连接,在萼区里大量成熟待产的卵浸泡在萼液中。产卵量随产卵时间延长而降低,在日产卵实验中,日产卵量202(±114SE),从1000至1800每2h的产卵量分别占日产卵量的38,32,17和13%;在一生产卵量实验中,9d产卵期,每头雌峰一生平均产卵量为502(±78SE)粒,第1d产卵量占一生总产卵量的42%。潜蝇姬小蜂的每个卵巢由3条卵巢管组成,每条输卵管中仅有1粒成熟卵,整个卵巢中只有6粒成熟卵,每次产卵数不超过6粒。结果表明,M.bicoloratus是早熟卵(proovigenic)类昆虫;D.isaea是同步卵(synovigenic)类昆虫。     

Host discrimination and interspecific competition of Trissolcus nigripedius and Telenomus gifuensis (Hymenoptera: Scelionidae), sympatric parasitoids of Dolycoris baccarum (Heteroptera: Pentatomidae)

Trissolcus nigripedius Nakagawa and Telenomus gifuensis Ashmead (Hymenoptera: Scelionidae) are solitary egg parasitoids of Dolycoris baccarum L. (Heteroptera: Pentatomidae), a polyphagous seed-sucking bug of agricultural crops. Field examinations revealed that the two parasitoids are the most common species, but only a single species emerge from a host egg mass. To explain this observation we tested two hypotheses of interspecific host discrimination and asymmetry in competitive interaction between Tr. nigripedius and Te. gifuensis. Trissolcus nigripedius and Te. gifuensis could discriminate host egg pre-parasitized by either self or conspecific like other scelionid parasitoids that use external mark on host eggs. When provided with host eggs pre-parasitized by each other, both Tr. nigripedius and Te. gifuensis multiparasitized 94% and 100% of the host eggs without interspecific host discrimination, respectively. Interestingly, from the multiparasitized eggs irrespective of oviposition sequence, progeny of Te. gifuensis always survive better than Tr. nigripedius. Telenomus gifuensis is superior in immature competition probably due to shorter egg incubation period after oviposition, hence progeny of Te. gifuensis become first instar faster than that of Tr. nigripedius. However, adult Tr. nigripedius is always superior competitor in possessing and guarding the host eggs even after oviposition against Te. gifuensis. Therefore, the asymmetry in competitive interaction between Tr. nigripedius and Te. gifuensis may explain the emergence of a single species from a host egg mass in the field in spite of no interspecific host discrimination.     

The role of female search behaviour in determining host plant range in plant feeding insects: a test of the information processing hypothesis

Recent theoretical studies have suggested that host range in herbivorous insects may be more restricted by constraints on information processing on the ovipositing females than by trade-offs in larval feeding efficiency. We have investigated if females from polyphagous species have to pay for their ability to localize and evaluate plants from different species with a lower ability to discriminate between conspecific host plants with differences in quality. Females of the monophagous butterflies Polygonia satyrus, Vanessa indica and Inachis io and the polyphagous P. c-album and Cynthia cardui (all in Lepidoptera, Nymphalidae) were given a simultaneous choice of stinging nettles (Urtica dioica) of different quality. In addition, the same choice trial was given to females from two populations of P. c-album with different degrees of specificity. As predicted from the information processing hypothesis, all specialists discriminated significantly against the bad quality nettle, whereas the generalists laid an equal amount of eggs on both types of nettle. There were no corresponding differences between specialist and generalist larvae in their ability to utilize poor quality leaves. Our study therefore suggests that female host-searching behaviour plays an important role in determining host plant range.