We studied variations in genetic, physiological, and ecological traits, and the phylogenetic relationship among sexual and parthenogenetic populations of Asobara japonica, a larval parasitoid of drosophilid flies, in order to understand how they adapt to local environments and have differentiated. The strain from Iriomote-jima (IR) differed from other Japanese strains in the nucleotide sequences of its cytochrome oxidase subunit I (COI) and in not undergoing diapause and having a shorter preimaginal period and a higher adult tolerance of cold. The strains other than IR showed a low level of nucleotide variation in COI but varied in their mode of reproduction; the strains from the Ryukyu Islands were sexual, whereas those from the main islands of Japan and Ogasawara were parthenogenetic. In addition, strains from higher latitudes generally showed a high incidence of diapause, although there were some exceptions. On the other hand, preimaginal period and adult cold tolerance varied little among the strains excluding IR, and pupal cold tolerance, oviposition preference and incidence of parasitism varied little among the strains including IR. Evolution and environmental adaptations in this species are discussed, particularly focusing on parthenogenetic populations.
FMRFamide-, vasoactive intestinal polypeptide (VIP)-, somatostatin-, substance P(SP)-, cholecystokinin octapeptide (CCK-8)- and pancreatic polypeptide (PP)-like materials were immunohistochemically detected in the endocrine cells and the gut innervation of diapausing pharate and newly-hatched first-instar larvae of Antheraea yamamai. SP- CCK-8-, PP- and FMRFamide-like immunoreactive cells were distributed unequally in different midgut regions; no cells reacting with antisera against VIP and somatostatin were found. Innervation of the anterior region of midgut, close to the foregut-midgut boundary, included 90-100 FMRFamide-like bipolar neurons that project along the midgut longitudinal muscles. No immunoreactive cell bodies were found in the foregut and hindgut, but nerve fibers with FMRFamide-like material were detected in hindgut muscles and extended over the posterior midgut. No changes in the distribution and intensity of immunostaining of both the endocrine cells and the gut innervation were found in four developmental stages between pre-diapause and hatching. SP-, CCK-8-, PP- and FMRFamide-like immunoreactive cells were also found in the CNS, indicating that the corresponding antigens belong to brain-gut regulatory peptides.
Laboratory and field tests support the "voltinism-suitability hypothesis" of host selection at various latitudes as well as in local "cold pockets": The best hosts for rapid development will be selected by herbivorous insects under severe thermal constraints for completion of the generation before winter. Papilio canadensis and P. glaucus females do select the best hosts for rapid larval growth in Alaska and in southern Michigan, but not in northern Michigan and southern Ohio. In addition to latitudinal patterns, local host preferences of P. canadensis are described in relation to "phenological twisting" of leaf suitability for larval growth in cold pockets with "thermally constrained" growing season lengths. White ash leaves (Fraxinus americana) have the highest nutritional quality (relative to cherry, aspen, birch, and other local trees) throughout June and July for P. canadensis populations inside the cold pocket, but not outside. In all areas outside the cold pockets, even with bud-break occurring much later than other tree species, ash leaves rapidly decline in quality after mid-June and become one of the worse tree host species for larvae. This temperature-driven phenology difference creates a geographic mosaic in host plant suitability for herbivores. However, we also report here that the cold pocket of northern Michigan has basically disappeared since 1998. Implications of these changes are discussed in relation to voltinism and host plant selection.
Various physiological and ecological adaptations to thermal unit constraints are evident with both species of naturally hybridizing butterflies (P. canadensis to the North and P. glaucus to the South). The rapid regional climate warming since 1998 in the Great Lakes region has allowed us to document some critical aspects of gene flow via analysis of independent "species-diagnostic" trait clines (including morphological, biochemical, and physiological characters). Narrower black bands on hind-wings, larger fore-wings, larval detoxification abilities for tulip tree leaves, behavioral preference of tulip tree leaves, and species-diagnostic allozymes such as PDG (x-linked) and HK (autosomal) provide evidence of these genodynamics. Laboratory hybridization studies (providing known reference groups) and field observations along the hybrid zone show that gene flow between P. glaucus and P. canadensis has recently been extensive across the historical hybrid zone.
Seasonal cycles constitute a major challenge for organisms since they may influence the genetic composition of a population, the species structure of a community and the interactions between organisms. Diapause is frequently used by insects to synchronize their life cycle with seasonal changes and is regarded as a key factor in the coexistence of competing species. Here the occurrence, abundance and emergence patterns of three poorly-known species of carnid flies (Carnus hemapterus Nitzsch, 1818, Hemeromyia anthracina Collin, 1949 and Hemeromyia longirostris Carles-Tolrá, 1992), which overwintered in the nests of several bird species at two localities, are reported and evidence of possible interspecific competition sought. Larvae of all three species were found in association with carrion and detritus. Both Hemeromyia species co-occurred in around 50% of the nests and Carnus with each of the Hemeromyia species at a lower rate (30-40% of the nests). Coexistence of all three carnid species was rare. We did not find any evidence of interspecific competition in the larval stage. Coexistence did not reduce the number of flies that emerged of any of the three species and the abundance of some species was even positively related. Species-specific emergence patterns and different habitat selection criteria (Carnus hemapterus seemed to avoid nests lined with vegetable material), which diminished the overlap between species were found. Such spatial and temporal segregation could facilitate the coexistence of these closely related species, which have similar ecological requirements, and might influence the seasonal dynamics of this poorly-known assemblage of insects inhabiting the nests of birds.
A laboratory study was carried out on photoperiodic control of prepupal diapause in the egg parasitoid Trichogramma embryophagum (Hartig). All experiments were conducted with an isofemale parthenogenetic strain. The maternal generation was reared at 20°C and photoperiods of L:D = 3:21, 6:18, 9:15, 12:12, 15:9, 18:6, 21:3 or 24:0. The tendency to diapause in the progeny was estimated by rearing the daughter generation at 15°C in the dark. Experiments revealed a long-day type response based on maternal influence on the progeny prepupal diapause. However, significant endogenous fluctuations in the pattern of the photoperiodic curve were revealed in successive laboratory generations reared under constant conditions. The left threshold day-length was very variable, while the right threshold kept relative constancy. Experiments with individual females sequentially offered new host eggs demonstrated that the probability of the progeny entering diapause depends significantly on maternal age. At 20°C and 18L : 6D, the percentage of diapause was maximal (ca 15%) in the progeny eclosed from the eggs laid during 1st - 2nd days of maternal life. Then the proportion of diapausing progeny decreased to 0-5% at days 9-11 of female life and later slightly increased in 15-17 days old females. Thus, endogenous factors play an important role in maternal influence on progeny diapause, particularly in environments close to threshold temperature and photoperiod.
The number of larval moults, larval head capsule width and pupal weight were investigated in both direct-developing and diapausing individuals of a South-West European population of Coenonympha pamphilus. The frequency distributions of head widths of successive larval instars overlapped, partly due to variation in the number of larval moults. The larvae that entered diapause went through five instars, instead of the four reported from this species. The evidence indicates that the five instar developmental pathway represents a plastic response rather than an example of compensatory growth. This alternative growth pattern was expressed in response to short photoperiods in parallel with, or as a consequence of, larval diapause. On average, the larvae with five instars had larger heads than their normal siblings. This resulted in comparatively heavier male pupae, while the opposite trend occurred in females. It is concluded that the variation in the number of larval instars is a plastic response to diapause when temperatures remain mild and that it might have an adaptive value in areas with mild winter climates. The sexually dimorphic expression in the larval growth patterns, in terms of pupal weight, may well imply different patterns of allocation of larval resources to adult structures, although sex-dependent differences in investment into purely larval structures cannot be discounted.
We surveyed ladybirds (Coleoptera: Coccinellidae) in 10 stands of Scots pine (Pinus sylvestris), all monoculture stands 5–100 years old, in western Slovakia, Central Europe, over two successive periods, October 2013 – March 2014 and October 2014 – March 2015. The winter in each period was exceptionally mild. Ladybirds were collected from the lower branches of pine trees using beating trays and were present in 61% of the 1040 samples (one sample containing ladybirds from 20 branches, 1 m long each). In total 3965 individuals of 20 species were recorded. Non-conifer dwelling species associated with broadleaved trees or herbaceous plants prevailed (45% of species), followed by conifer specialists (40%) and generalists (15%). Although 13 species were found at least in one winter month, December, January or February, only four of them, Exochomus quadripustulatus, Coccinella septempunctata, Harmonia axyridis and Hippodamia variegata, were recorded continually during both winters. The number of species, the abundance of all ladybirds and the abundance of dominant species (E. quadripustulatus, C. septempunctata and H. axyridis) decreased from late autumn towards winter and remained lowest during this most adverse time of the year for ladybirds. Overwintering species assemblages of ladybirds changed over time and varied with age of pine stand. Our results suggest that Scots pine in Central Europe supports species rich assemblages of ladybirds from late autumn to early spring and, being widely distributed, it could be suited to winter surveying of ladybirds at large spatial scales to reveal behavioural and ecological responses of species to changing weather or different climates.
Wild females of Pyrrhocoris apterus exhibit seasonal changes in neuroendocrine activity and, consequently, reproduction. Long days (18 h light/6 h dark) (LD) stimulate reproduction, whereas short days (12 h light/12 h dark) (SD) induce reproductive arrest (diapause). This study reveals how photoperiod influences the expression of the circadian clock gene, period (per) in the insect's head. There is only a weak diurnal rhythm in per mRNA expression under LD and SD. However, levels of per mRNA are consistently higher (up to 10-fold) under SD than under LD. The influence of photoperiod on per gene expression is linked to a developmental output (diapause vs. reproduction); mutant females, reproducing under both LD and SD, show low per mRNA levels under both photoperiodic conditions. Thus, the magnitude of per gene expression may be important to the translation of photoperiodic signals into a hormonal message. Levels of per mRNA are related to properties of locomotor activity rhythms. Low per mRNA levels (displayed by wild females in LD and mutant females in both LD and SD) are associated with long free-running periods (τ~26-27 h) and late peaks of activity (ψR,L~10-12 h), whereas high per mRNA levels coincide with short free-running periods (τ~24 h) and early peaks of activity (ψR,L~4-6 h). Overall, the data provide a background for a molecular approach to the long-standing question about the role of the circadian system in insect photoperiodism.
Great progress has recently been made in cryobiology. One field, however, has been neglected: the temporal sequence of the effects of photoperiod and temperature, and their relative importance in cold hardening. This is relevant to the question of importance of diapause in cold-hardiness. Denlinger (1991) outlined the categories of such relations and stressed a great need for further detailed research. A survey of studies done over the past decade revealed many gaps in the evidence and the ambiguous nature of the data on the photoperiodic regulation of cold-hardiness. We hope that this review will stimulate further research in this field. Among several directions where research is most needed we have stressed (1) simultaneous recording of changes in survival and dynamics of suspected cryoprotectants (stressed also by Danks, 1996), (2) checking the regulation of different phases of cold hardening, and (3) discrimination between direct and indirect (mediated via neuroendocrine system) effects of environmental cues on cold hardening.
The effects of photoperiod on pre-imaginal development and reproductive maturation of adult females of the multicolored Asian lady beetle, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), were investigated at 20°C and at photoperiods from 10L : 14D to 18L : 6D. Experiments were conducted on a laboratory strain that originated from the Russian Far East. Larvae and adults were fed on the green peach aphid Myzus persicae. Under short day conditions the pre-imaginal development was faster than under long day conditions. The acceleration of pre-imaginal development occurred when day length was shorter than 16 h and the threshold day length was ca 14 h. The rate of reproductive maturation of adult females, in contrast, was twice as high under long day conditions, with the threshold between 14L : 10D and 12L : 12D, although even under short day conditions ca 10% of the females show a tendency to mature more quickly. This difference between the thresholds of these two photoperiodic reactions indicates they are relatively independent of one another. and Sergey Ya. Reznik, Nina P. Vaghina.