Hybrid Dysgenesis in DROSOPHILA MELANOGASTER: The Biology of Female and Male Sterility

High levels of female and male sterility were observed among the hybrids from one of the two reciprocal crosses between a wild strain of D. melanogaster known as pi2 and laboratory strains. The sterility, which is part of a common syndrome called hybrid dysgenesis, was found to be associated with the rudimentary condition of one or both of the ovaries or testes. All other tissues, including those of the reproductive system were normal, as were longevity and mating behavior. The morphological details of the sterility closely mimic the agametic condition occurring when germ cells are destroyed by irradiation or by the maternal-effect mutation, grandchildless. We suggest that sterility in hybrid dysgenesis is also caused by failure in the early development of germ cells. There is a thermo-sensitive period beginning at approximately the time of initiation of mitosis among primordial germ cells a few hours before the egg hatches and ending during the early larval stages. Our results suggest that hybrid dysgenesis, which also includes male recombination, mutation and other traits, may be limited to the germ line, and that each of the primordial germ cells develops, or fails to develop, independently of the others. This hypothesis is consistent with the observed frequencies of unilateral and bilateral sterility, with the shape of the thermosensitivity curves and with the fact that males are less often sterile than females. The features of this intraspecific hybrid sterility are found to resemble those seen in some interspecific Drosophila hybrids, especially those from the cross D. melanogaster X D. simulans.     

Nonreciprocal gonadal dysgenesis in hybrids of the chironomid midgeChironomus thummi. IV. Behavior of a female-specific protein,probably a vitellogenin

Using denaturing SDS-polyacrylamide gel electrophoresis, a protein with a subunit MW of about 148,000 daltons could be detected in the fat body of females of the reciprocal hybrids of Chironomus thummi thummi and Chironomus thummi piger, which is not present in males. This protein is presumably a vitellogenin and can be found in both hybrids during the late fourth-instar larval stage until eclosion of the adults, i.e., in early vitellogenesis. After eclosion, the reciprocal hybrids behave differently concerning the 148-kd protein. In females of the piger female x thummi male cross, which are fertile and produce yolky eggs, the 148-kd protein disappears from the fat body immediately after eclosion. In females of the reciprocal cross (thummi female x piger male) which are affected by gonadal dysgenesis and in which the oocytes only rarely contain yolk, the 148-kd protein is still present in the fat body of the adult up to 50 hr after eclosion until the fat body degrades. It is concluded that the inability of the sterile thummi female x piger male females to produce yolky eggs is caused by an impaired uptake of the presumed 148-kd vitellogenin into oocytes and not by a defective vitellogenesis. The impaired vitellogenin deposition into oocytes is taken as another aberrant trait of gonadal dysgenesis of the thummi female x piger male hybrids.     

Hybrid dysgenesis in Drosophila melanogaster: influence of temperature on cytotype determination in the P-M system

Summary In Drosophila melanogaster, the P-M system of hybrid dysgenesis is a syndrome of germ line abnormalities, including temperature dependent gonadal dysgenesis (GD sterility), high rates of mutation and male recombination, which occurs in some interstrain hybrids but only from one of the two crosses. In the P-M system, hybrid dysgenesis results from interaction between chromosomally transposable elements of the P element family and a particular extrachromosomal state referred to as the M cytotype. Cytotype (M or P) is known to be determined by the absence or presence of chromosomal factors, but principally with limited cytoplasmic transmission.In a series of experiments in which F1 hybrid females from various P and M strains were submitted to different preadult and ageing temperature treatments, it was found that the cytotype switch is strongly temperature-dependent in the F1 females from M x P but not in the reciprocal cross. In the F1 females from the former cross, a strong M cytotype occurs at a low developmental temperature (18° C) and a weak M cytotype occurs at a high developmental temperature (26.5° C). On the other hand, a high ageing temperature applied after a low developmental temperature switches the cytotype from M to P and reciprocally, a low ageing temperature applied after a high developmental temperature switches the cytotype from P to M.This thermo-reversibility of the extrachromosomal state exists only in the F1 females from M mothers but not in the F1 females from P mothers; this dissymmetrical behavior is discussed in relation to the mechanism proposed by O'Hare and Rubin (1983) which explains cytotype determination by a positive feedback of the regulator of the P transposase on its own level of activity.     

A high level of hybrid dysgenesis in Drosophila: High thermosensitivity, dependence on DNA repair, and incomplete cytotype regulation

Summary An unusually high level of P-M hybrid dysgenesis in Drosophila melanogaster is characteristic of hybrid offspring originating from both, A (M × ) and B (P × M) crosses of a subline of the Harwich P strain, termed H ^s . The novel properties induced by mobility of P elements carried by H ^s paternal chromosomes include: very high (over 95%) gonadal dysgenesis (GD) in both sexes at the low restrictive temperature of 21°C, and highly premature sterility when males are reared at 18°C and aged at 21°C. Although all three major chromosomes of the H ^s subline contributed to this atypical pattern of gonadal dysgenesis, chromosome 3 had the largest effect. Gonadal dysgenesis showed a temperature- and sex-dependent repression pattern by the defective P elements of Muller-5 Birmingham chromosomes; at 21°C there was virtually no repression of male sterility, but most effective repression of GD in females. At 29°C repression was effective in males, but declined in females. The high thermosensitive sterility, low fecundity, and premature aging of the male germ line were greatly exacerbated when males derived from either A or B crosses were deficient either in excision repair (mei-9 mutation) or in post-replication repair (mei-41 mutation). These findings demonstrate that both DNA repair pathways are essential for the repair of lesions induced by P element transposition and support the hypothesis that P element-induced chromosome breaks are responsible for the virtual abolition of the germ line. The relatively high premature sterility of cross B DNA repair-deficient males, reared at 18°C and aged at 21°C, indicates that there is incomplete cytotype regulation in H ^s subline hybrids.     

Hybrid Dysgenesis in DROSOPHILA MELANOGASTER: Morphological and Cytological Studies of Ovarian Dysgenesis

A type of intraspecific hybrid sterility, between two strains of Drosophila melanogaster, referred to as GD (gonadal dysgenesis) sterility, is observed when females from a type of strain called M are crossed with males from a second type called P. Absence of egg-laying is characteristic of female GD sterility and its manifestation is conditional on high developmental temperatures. Morphological and cytological studies of GD sterile females are described. These individuals were normal in body size and external appearance. No defects in sperm storage were observed. Both adult and larval ovaries were drastically reduced in size in comparison with control ovaries. This ovarian dysgenesis was sometimes unilateral, but more frequently it was bilateral, particularly in females developing at the highest test temperature. The ovarioles of dysgenic ovaries contained no vitellaria; the germaria lacked any cells resembling the cystocyte clusters of normal ovaries. It is concluded that sterility results from an early blockage in ovarian development, rather than from atrophy of previously developed structures. Possible mechanisms for this developmental arrest are discussed.     

Hybrid Dysgenesis in DROSOPHILA MELANOGASTER: Sterility Resulting from Gonadal Dysgenesis in the P-M System

Crosses between two types of strains, called P and M, characteristically give high frequencies of F(1) sterility and other aberrant traits. Previous studies indicated that, in addition to the direction of the parental cross, many factors influence the manifestation of this phenomenon known as "hybrid dysgenesis."-The present study is concerned with the characteristics of GD (gonadal dysgenesis) sterility associated with the P-M system and its temperature dependence. Female sterility is accompanied by a complete absence of egg-laying, and this is not attributable to an inability to mate. Thus, it seems likely that sterility results from a defect in gametogenesis of hybrid individuals. This conclusion is supported by the morphological and cytological observations presented in an accompanying paper (Schaefer, Kidwell and Fausto-Sterling 1979).-A narrow, critical, developmental temperature range was found in which both female and male sterility rose sharply from a low level to a high maximum. The critical range was 27 to 29 degrees for males, slightly higher than the range of 24 to 26 degrees for females. Two other dysgenic traits, male recombination and transmission ratio distortion, were affected by developmental temperature, but temperature response curves were quite different from those for sterility. The temperature-sensitive stage for female sterility occurs during embryonic and early larval development.-GD sterility is compared and contrasted with SF sterility, another specific type of non-Mendelian sterility resulting from a different interstrain dysgenic interaction.     

Gonadal Dysgenesis Reveals Sexual Dimorphism in the Embryonic Germline of Drosophila

In hybrid dysgenesis, sterility can occur in both males and females. At 27.5 degrees, however, we found that P element-induced germline death was restricted to females. This sex-specific gonadal dysgenesis (GD) is complete by the first larval instar stage. As such, GD at 27.5 degrees reveals the sexually dimorphic character of the embryonic germline. The only other known dimorphic trait of the embryonic germline is the requirement for ovo. ovo is required for germline development in females only and has been implicated in germline sex determination. Dominant mutations of ovo partially suppressed female GD. Although embryonic germ cells are undifferentiated and morphologically indistinguishable between males and females, the functional dimorphism seen in ovo requirement and GD at 27.5 degrees indicates that sexual identity in Drosophila germ cells is established in embryogenesis.     

The HLE hybrid dysgenesis syndrome in the midgeChironomus thummi: Differences in the dysgenic potential between strains

After crossing experiments between twoChironomus subspecies the HLE hybrid dysgenesis syndrome occurs non-reciprocally in the F1 hybrids if the males of each of the fourCh. th. thummi strains (HL, So, G, Kr) are crossed to females ofCh. th. piger, strain E. In hybrid egg masses, differences in the strength of the abnormal trait reduced egg hatch were found between the differentpiger E ×thummi interstrain crosses. From the results it is concluded that eachthummi strain has a specific hybrid dysgenesis potential which determines the severity of the HLE syndrome in the hybrids. The HLE syndrome occurs also in the hybrids of crosses of differentCh. th. thummi strains. In these hybrids the syndrome is only obtained if the male parent descends from a strain which has a higher HLE hybrid dysgenesis potential than that strain from which the female parent is derived. These results demonstrate that the HLE syndrome is not restricted to the hybrids of thethummi × piger subspecies cross. In theCh. th. thummi interstrain crosses the hybrids of that cross direction are normal, which is reciprocal to the cross direction affected by the HLE syndrome. This is in contrast to thethummi × piger subspecies crosses, where the hybrids of one cross direction always show the HLE syndrome and the hybrids of the reciprocal cross direction the Rud syndrome. Thus, the present study also demonstrates that the HLE and Rud hybrid dysgenesis syndrome inChironomus are independent from one another.     

Genetics of Hybrid Inviability and Sterility in Drosophila: Dissection of Introgression of D. simulans Genes in D. melanogaster Genome

Interspecific crosses between Drosophila melanogaster and Drosophila simulans usually produce sterile unisexual hybrids. The barrier preventing genetic analysis of hybrid inviability and sterility has been taken away by the discovery of a D. simulans strain which produces fertile female hybrids. D. simulans genes in the cytological locations of 21A1 to 22C1-23B1 and 30F3-31C5 to 36A2-7 have been introgressed into the D. melanogaster genetic background by consecutive backcrosses. Flies heterozygous for the introgression are fertile, while homozygotes are sterile both in females and males. The genes responsible for the sterility have been mapped in the introgression. The male sterility is caused by the synergistic effect of multiple genes, while the female sterility genes have been localized to a 170 kb region (32D2 to 32E4) containing 20 open reading frames. Thus, the female sterility might be attributed to a single gene with a large effect. We have also found that the Lethal hybrid rescue mutation which prevents the inviability of male hybrids from the cross of D. melanogaster females and D. simulans males cannot rescue those carrying the introgression, suggesting that D. simulans genes maybe non-functional in this hybrid genotype. The genes responsible for the inviability have not been separated from the female sterility genes by recombination.     

Developmental and reproductive biology of Scirtothrips perseae (Thysanoptera: Thripidae): a new avocado pest in California

The developmental and reproductive biology of a new avocado pest, Scirtothrips perseae Nakahara, was determined in the laboratory at five constant temperatures, 15, 20, 25, 27.5 and 30 degrees C. At 20 degrees C, S. perseae exhibited greatest larval to adult survivorship (41%), and mated females produced a greater proportion of female offspring at this temperature when compared to 15, 25, 27.5 and 30 degrees C. Average lifetime fecundity and preoviposition period was greatest at 15 degrees C at 39.6 eggs per female and 17.6 days, respectively. Jackknifed estimates of net reproduction (Ro), capacity for increase (rc), intrinsic rate of increase (rm), and finite rate of increase (lambda) were all significantly greater at 20 degrees C than corresponding values at 15, 25 and 27.5 degrees C. Population doubling time (Td) was significantly lower at 20 degrees C, indicating S. perseae populations can double 33-71% faster at this temperature in comparison to 15, 25 and 27.5 degrees C. Mean adult longevity decreased with increasing temperature, from a maximum of 52.4 days at 15 degrees C to a minimum of 2.4 days at 30 degrees C. Developmental rates increased linearly with increasing temperatures for eggs and rates were non-linear for development of first and second instar larvae, propupae, pupae, and for egg to adult development. Linear regression and fitting of the modified Logan model to developmental rate data for egg to adult development estimated that 344.8 day degrees were required above a minimum threshold of 6.9 degrees C to complete development. An upper developmental threshold was estimated at 37.6 degrees C with an optimal temperature of 30.5 degrees C for egg to adult development. Unmated females produced only male offspring confirming arrhenotoky in S. perseae.     

A novel system of fertility rescue in Drosophila hybrids reveals a link between hybrid lethality and female sterility

Hybrid daughters of crosses between Drosophila melanogaster females and males from the D. simulans species clade are fully viable at low temperature but have agametic ovaries and are thus sterile. We report here that mutations in the D. melanogaster gene Hybrid male rescue (Hmr), along with unidentified polymorphic factors, rescue this agametic phenotype in both D. melanogaster/D. simulans and D. melanogaster/D. mauritiana F(1) female hybrids. These hybrids produced small numbers of progeny in backcrosses, their low fecundity being caused by incomplete rescue of oogenesis as well as by zygotic lethality. F(1) hybrid males from these crosses remained fully sterile. Hmr(+) is the first Drosophila gene shown to cause hybrid female sterility. These results also suggest that, while there is some common genetic basis to hybrid lethality and female sterility in D. melanogaster, hybrid females are more sensitive to fertility defects than to lethality.     

Characterization of defects in adult germline development and oogenesis of sterile and rescued female hybrids in crosses between Drosophila simulans and Drosophila melanogaster

Crosses between Drosophila melanogaster and D. simulans normally result in progeny that are either inviable or sterile. Recent discovery of strains that rescue these inviability and sterility phenotypes has made it possible to study the developmental basis of reproductive isolation between these two species in greater detail. By producing both rescued and unrescued hybrids and examining the protein product staining patterns of genes known to be involved in early germline development and gametogenesis, we have found that in crosses between D. simulans and D. melanogaster, hybrid female sterility results from the improper control of primordial germline proliferation, germline stem cell maintenance, and cystoblast formation and differentiation during early oogenesis. Rescued hybrid females are fertile, yet they generally have lower amounts of adult germline from the outset and show a premature degeneration of adult germline cells with age. In addition, older rescued hybrid females also exhibit mutant egg phenotypes associated with defects in dorso-ventral patterning which may result from the improper partitioning of cytoplasmic factors during early oogenesis that could stem from the early defect. Although a variety of germline and oogenic defects are described for the hybrid females, all of them can potentially result from the same underlying primary defect. Hybrid males from these same crosses, on the other hand, have no detectable germline in adult reproductive tissues, even when hybrid sterility rescue strains are used, indicating that male sterility and female sterility stem from distinctly different developmental defects.     

The relationship between radiation-induced and transposon-induced genetic damage during Drosophila oogenesis

The combined effect of X-irradiation and transposon mobility on the frequencies of X-linked recessive lethals and dominant lethals was investigated in female hybrids in the P-M system of hybrid dysgenesis. X-linked lethals were measured in G2 hybrid dysgenic females whose X chromosome was derived from the M X P cross. To test for additivity or synergism, the mutation rate in irradiated dysgenic females was compared to that of unirradiated females as well as to irradiated nondysgenic hybrid females derived from M X M crosses. Eggs collected for 2 days after irradiation, were represented by the more radiation-sensitive A and B oocytes (about 75%) and the least sensitive C oocytes (about 25%). The production of X-linked lethal events in X-irradiated dysgenic females was 8.1%, as compared to 4.5% in dysgenic controls and 3.4% in irradiated, nondysgenic controls, demonstrating an additive effect of radiation and dysgenesis-induced genetic damage. The effect of irradiation on sterility of dysgenic hybrid females was a negative one, resulting in 20% less sterility than expected from an additive effect. The combined effect of radiation and dysgenesis on dominant lethality tested in A, B and C oocytes of the same hybrid females was synergistic. Egg broods collected for 3.5 days after irradiation showed that significantly more damage was induced in the presence of ionizing radiation in dysgenic females than in their nondysgenic counterparts. This effect was most obvious in B and C oocytes. The synergism observed may be related to the inability of cells to repair the increased number of chromosome breaks induced both by radiation and transposon mobility.     

Ovarian defects in hybrids of the species pair Drosophila virilis and Drosophila texana

In interspecific matings between Drosophila virilis and Drosophila texana female sterility is observed in F₂ hybrid females. A previous study has shown that no vitellogenin synthesis occurs in the fat body of sterile hybrid females. The results presented in this paper show that hybrid ovaries of sterile females transplanted into the abdomens of females of the parental species are not able to develop upon maturity. With few exceptions, the hybrid ovaries remained alive in the host environment, but their oocytes failed to develop to vitellogenic stages. Thus, in hybrid females between Drosophila virilis and Drosophila texana sterility is the result of defects in both the two main developmental processes of egg maturation, the synthesis of vitellogenins in the fat body and the uptake of vitellogenins by the ovary. Dev Genet 20:47–52, 1997. © 1997 Wiley-Liss, Inc.     

Essential role for gamma-tubulin in the acentriolar female meiotic spindle of Drosophila.

Microtubule nucleation in vivo requires gamma-tubulin, a highly conserved component of microtubule-organizing centers. In Drosophila melanogaster there are two gamma-tubulin genes, gammaTUB23C and gammaTUB37C. Here we report the cytological and molecular characterization of the 37C isoform. By Western blotting, this protein can only be detected in ovaries and embryos. Antibodies against this isoform predominantly label the centrosomes in embryos from early cleavage divisions until cycle 15, but fail to reveal any particular localization of gamma-tubulin in the developing egg chambers. The loss of function of this gene results in female sterility and has no effect on viability or male fertility. Early stages of oogenesis are unaffected by mutations in this gene, as judged both by morphological criteria and by localization of reporter genes, but the female meiotic spindle is extremely disrupted. Nuclear proliferation within the eggs laid by mutant females is also impaired. We conclude that the expression of the 37C gamma-tubulin isoform of D. melanogaster is under strict developmental regulation and that the organization of the female meiotic spindle requires gamma-tubulin.     

The biology of Diadromus collaris (Hymenoptera: Ichneumonidae), a pupal parasitoid of Plutella xylostella (Lepidoptera: Plutellidae), and its interactions with Oomyzus sokolowskii (Hymenoptera: Eulophidae)

The ichneumonid Diadromus collaris (Gravenhorst) has been recorded in many parts of the world as an important parasitoid of the diamondback moth, Plutella xylostella (Linnaeus), a serious pest of brassica vegetable crops worldwide. Some aspects of its biology and its interactions with Oomyzus sokolowskii (Kurdjumov), another major parasitoid of the same pest, were studied in the laboratory. At 25 degrees C, female wasps did not have mature eggs in their ovaries until about 12 h after emergence. Both males and females mated successfully 24-48 h after emergence, and females started to oviposit one to two days after emergence. Unmated females produced male progeny only; mated females produced progeny of both sexes. The development rate of the parasitoid increased linearly with temperature from 15 to 30 degrees C, with an estimated low temperature threshold of 7.4 degrees C and a thermal constant of 225.1 day-degrees for development from egg to adulthood. Rates of survival from larva to adulthood were about 90% between 20 and 28 degrees C and decreased as temperature decreased or increased. No immatures survived to adulthood at 35 degrees C. When provided with honey solution, the females lived on average 8.3, 11.5 and 7.0 days, and parasitized 26, 44 and 46 host pupae at 20, 25 and 30 degrees C, respectively. Female wasps could be stored at 15 degrees C for up to four weeks without detrimental effects on reproduction. Females of D. collaris attacked host pupae already parasitized by O. sokolowskii, inserting their ovipositor into the hosts at a similar frequency as into unparasitized host pupae, but they did not lay eggs inside the hosts.     

A Hybrid Dysgenesis Syndrome in Drosophila Virilis

A new example of ``hybrid dysgenesis' has been demonstrated in the F(1) progeny of crosses between two different strains of Drosophila virilis. The dysgenic traits were observed only in hybrids obtained when wild-type females (of the Batumi strain 9 from Georgia, USSR) were crossed to males from a marker strain (the long-established laboratory strain, strain 160, carrying recessive markers on all its autosomes). The phenomena observed include high frequencies of male and female sterility, male recombination, chromosomal nondisjunction, transmission ratio distortion and the appearance of numerous visible mutations at different loci in the progeny of dysgenic crosses. The sterility demonstrated in the present study is similar to that of P-M dysgenesis in Drosophila melanogaster and apparently results from underdevelopment of the gonads in both sexes, this phenomenon being sensitive to developmental temperature. However, in contrast to the P-M and I-R dysgenic systems in D. melanogaster, in D. virilis the highest level of sterility (95-98%) occurs at 23-25°. Several of the mutations isolated from the progeny of dysgenic crosses (e.g., singed) proved to be unstable and reverted to wild type. We hypothesize that a mobile element (``Ulysses') which we have recently isolated from a dysgenically induced white eye mutation may be responsible for the phenomena observed.     

Interaction of Nosema pyrausta and temperature on Ostrinia nubilalis egg production and hatch

Nosema pyrausta is an obligate pathogen causing reduced fecundity and longevity of Ostrinia nubilalis. This study was conducted to determine the combined effects of N. pyrausta infection and temperature on O. nubilalis egg production and hatch. N. pyrausta-infected and noninfected O. nubilalis were maintained in two different temperature regimes. The first regime allowed females to oviposit under optimum conditions (27 degrees C, 65% RH, 16:8 (L:D)), while females in the second regime were held initially under the same humidity and light conditions, but a constant temperature of 16 degrees C for 1 week after which they were transferred to optimum ovipositional conditions. Studies were performed initially with O. nubilalis populations and later with individual mating pairs. In studies with O. nubilalis populations, the mean number of eggs laid per female under optimum conditions was 660, while N. pyrausta-infected females held initially at 16 degrees C laid 116 eggs per female. In studies with individual mating pairs, N. pyrausta infection reduced egg production per female 53 and 11% in the 16 and 27 degrees C temperature regimes, respectively, compared to noninfected females under optimum conditions. Exposure to 16 degrees C temperatures early in the ovipositional period had a more profound impact on reducing egg production in N. pyrausta-infected than noninfected O. nubilalis.     

Haploid regeneration from tetraploid wheat using maize pollen.

Crosses were made between 21 tetraploid wheat genotypes (6 parents, 15 F1 hybrids) and a single F1 hybrid of maize that was used as the male parent. Plants were grown under controlled greenhouse conditions (daylength, 16 h; temperature, 25 degrees C days and 15 degrees C nights). To enhance embryo survival, 2,4-D (10 mg/L) was applied to spikes 24 h after pollination with maize. Embryos were recovered from the tetraploid wheat genotypes at a rate of 2.34-14.14/100 developed ovaries. Sixty-nine haploid plants were obtained from 3 parents and 12 F1, hybrids. Fifty-six of these were successfully doubled. General combining ability was significant for the two traits studied, indicating that additive genetic control is important for the number of developed ovaries and haploid embryo production in tetraploid wheat x maize crosses. In this report, we demonstrate the potential of using maize pollen to produce haploid plants from tetraploid wheat genotypes.     

Temperature tolerance of the diamondback moth, Plutella xylostella (Lepidoptera: Yponomeutidae) in tropical and temperate regions of Asia

Temperature tolerance was investigated in nine populations of Plutella xylostella Linnaeus from tropical and temperate regions of Asia. At all rearing temperatures between 15 and 35 degrees C, no clear differences were observed in female egg production or larval development between tropical and temperate populations. Thus, tropical populations did not show a high-temperature tolerance superior to that of the temperate populations. In all populations, the net reproductive rate (number of new females born per female) largely depended on the number of eggs laid per female, and egg production significantly decreased with increasing temperature (P < 0.001). Larval developmental rate also showed a significant positive correlation with temperature (P < 0.001). Per cent hatch of eggs and larval survival did not show a significant correlation with temperature: hatching was constant between 15 and 32.5 degrees C, but considerably lower at 35 degrees C. Larval survival was similar between 15 and 30 degrees C, appreciably lower at 32.5 degrees C and declined to 0% at 35 degrees C. Based on these results, environmental conditions under which P. xylostella can maintain a high population density throughout the year in tropical and subtropical regions are discussed.