Tight insect–host phenological synchrony constrains the life‐history strategy of European pine sawfly

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Seasonal flight patterns of Ips typographus in southern Sweden and thermal sums required for emergence

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Diapause and overwintering of two spruce bark beetle species

Diapause, a strategy to endure unfavourable conditions (e.g. cold winters) is commonly found in ectothermic organisms and is characterized by an arrest of development and reproduction, a reduction of metabolic rate, and an increased resistance to adversity. Diapause, in addition to adaptations for surviving low winter temperatures, significantly influences phenology, voltinism and ultimately population growth. We review the literature on diapause and overwintering behaviour of two bark beetle species that affect spruce‐dominated forests in the northern hemisphere, and describe and compare how these strategies can influence population dynamics. The European spruce bark beetle Ips typographus (L.) (Coleoptera, Curculionidae) is the most important forest pest of Norway spruce in Europe. It enters an adult reproductive diapause that might be either facultative or obligate. Obligate diapausing beetles are considered strictly univoltine, entering this dormancy type regardless of environmental cues. Facultative diapausing individuals enter diapause induced by photoperiod, modified by temperature, thus being potentially multivoltine. The spruce beetle Dendroctonus rufipennis (Kirby) (Coleoptera: Curculionidae) infests all spruce species in its natural range in North America. A facultative prepupal diapause is averted by relatively warm temperatures, resulting in a univoltine life cycle, whereas cool temperatures induce prepupal diapause leading to a semivoltine cycle. An adult obligate diapause in D. rufipennis could limit bi‐ or multivoltinism. We discuss and compare the influence of diapause and overwinter survival on voltinism and population dynamics of these two species in a changing climate and provide an outlook on future research.     

Development, reproduction, voltinism and host synchrony ofMeteorus trachynotus with its hostsChoristoneura fumiferana andC. rosaceana

The form and variability of temperature-dependent development responses were measured for the braconid parasitoidMeteorus trachynotus (Vier.), developing inChoristoneura fumiferana (Clem.) andC. rosaceana (Harr.). It was found that parasitism reduces the duration and variability of postdiapause development inC. rosaceana larvae, and that the development of the parasitoid does not resume until its host emerges from diapause.M. trachynotus development and entry into diapause are controlled by the host. In large, nondiapausing hosts, the parasitoid can complete one generation in about 348 degree-days above 5.9°C. Male and female adults live an average of 22.0 and 23.8 days at 20°C, and the preoviposition period is 1.7 days. Total fecundity averages 194.2 eggs per female and is reduced considerably by low host density. Under our experimental conditions, femaleM. trachynotus lay a maximum of 14.3 eggs per day, with a maximum of 3.7 eggs per host, on average. Simulations based on degree-day accumulation in 13 locations in eastern Canada showed that the parasitoid was well synchronized with its two alternative hosts, thatM. trachynotus could complete three generations in most locations, and that voltinism was essentially determined by entry ofC. rosaceana larvae into diapause. Overlap between adult parasitoids and larvae of the spruce budworm is incomplete, and inadequate synchrony seems to be an important aspect of the ecological interactions between these two organisms.

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Résumé Nous avons mesuré la forme et la variabilité des réactions de cause à effet entre la température et le développement de la braconideMeteorus trachynotus (Vier.) se développant à l'intérieur de larves deChoristoneura fumiferana (Clem.) et deC. rosaceana (Harr.). Le parasitisme réduit la durée et la variabilité de la postdiapause deC. rosaceana, et le développement du parasito?de ne reprend qu'une fois l'h?te émergé de diapause. Le développement et l'entrée en diapause deM. trachynotus sont sous le contr?le de l'h?te. Dans des larves-h?tes agées et non diapausantes, le parasito?de peut engendrer une génération en moins de 348 degrés-jours au-dessus de 5,9°C. Les adultes males et femelles vivent en moyenne 22,0 et 23,8 jours à 20°C, et la période de prèoviposition est de 1,7 jour. La fécondité totale est de 194,2 oeufs par femelle, et une faible densité d'h?tes la réduit considérablement. Dans nos conditions expérimentales, les femelles deM. trachynotus pondent au plus 14,3 oeufs par jour, la moyenne maximale étant de 3,7 oeufs par h?te. Des simulations basées sur l'accumulation de degrés-jours pour 13 endroits dans l'est du Canada ont indiqué que le parasito?de est bien synchronisé avec ses deux h?tes alternants, queM. trachynotus peut engendrer jusqu'à trois générations par année dans la plupart des endroits, et que son voltinisme est déterminé essentiellement par l'entrée en diapause des larves deC. rosaceana. Le chevauchement entre la période de vol des adultes du parasito?de et celle de la disponibilité des larves de la tordeuse des bourgeons de l'épinette est incomplet, et il semble que ce synchronisme inadéquat soit un aspect important dans les interactions écologiques entre ces deux organismes.

     

Effect of predators and host phenology on the maternal and reproductive behaviors ofGargaphia lace bugs (Hemiptera: Tingidae)

Lace bugs of the genus Gargaphia specialize on taxonomically and ecologically diverse host plants. To examine the impact of predation pressure and host phenology on Gargaphia subsocial and reproductive behavior, we compared G. tiliae, a woodland species restricted to riparian trees in the genus Tilia, and G. solani, a specialist on Solanum in early successional habitats. Both lace bug species exhibit maternal care of eggs and nymphs. Predator densities associated with Solanum were higher than those associated with Tilia and were correlated with higher levels of maternal aggression exhibited by G. solani. In both habitats, artificial exclusion of predators significantly increased nymphal survivorship over maternally guarded broods. Both species reduced the costs associated with maternal care by ovipositing into the egg masses of conspecifics. Gargaphia solani and G. tiliae differed most strikingly in voltinism and thus the potential for iteroparity. Leaf age determined the reproductive future of G. tiliae nymphs independent of photoperiod or temperature. Feeding on plants with young foliage was sufficient to trigger oviposition. Constraints imposed by host plants on life-history patterns and maternal options are discussed.     

Mitochondrial DNA analysis of the introduced fall webworm, showing its shift in life cycle in Japan

The fall webworm, Hyphantria cunea (Drury) (Lepidoptera: Arctiidae), was introduced from North America into Japan in 1945. For the first three decades after its introduction, its life cycle was bivoltine. Thereafter, its life cycle shifted to trivoltine in south‐western areas of Japan. Two hypotheses have been proposed for the process that led to the shift in voltinism: one based on a single and the other on multiple independent colonizations. To test these hypotheses, mitochondrial (mt)DNA sequences were analyzed in the black‐headed type of 14 Japanese, one Korean and two North American populations of H. cunea. In addition, the same regions of mtDNA were compared with the red‐headed type of two North American populations. In the black‐headed type, mtDNA sequences were the same in all Japanese populations and in the Korean population, but sequences of the North American populations differed from each other and from those of the other populations. These results suggest that the process of the shift in voltinism occurred originally in Japan, and that the Japanese and the Korean population of H. cunea originated from a relatively small area in North America.     

Survivability and post‐diapause fitness in a scolytid beetle as a function of overwintering developmental stage and the implications for population dynamics

1. It has recently been suggested that expected increases in temperature might lead to an additional generation per season in bark beetles. Thus, populations would grow more rapidly. However, an additional but not fully developed generation could lead to high winter mortality in the pre‐imaginal stages. 2. Winter survivability and post‐diapause fitness as a function of the overwintering developmental stage of the spruce bark beetle Ips typographus L. were studied along an altitudinal cline. Fitness was tested in terms of weight, lipid content and flight capacity. 3. Mortality was significantly lower and emergence per brood system significantly higher for fully developed adults that entered overwintering than for larvae, pupae or callow adults. 4. Post‐diapause fitness in terms of dry weight and flight capacity was significantly higher in individuals that completed development before winter, and lipid contents also showed a trend for being higher in those individuals. 5. In conclusion, in a scenario where effective temperature sums are not adequate for the complete development of an additional generation, models may overestimate population growth by neglecting increased mortality and reduced post‐diapause fitness. The results highlight the importance of considering life‐history traits and indirect effects in addition to abiotic factors such as temperature when modelling population dynamics.     

Population persistence of the parasitoid fly Zaira cinerea (Fallén) (Diptera: Tachinidae) utilizing multiple host carabid beetles with different seasonality and quality

Zaira cinerea (Fallén) is a parasitoid fly (Diptera: Tachinidae) that attacks adult carabid beetles. To better understand mechanisms of population persistence in this species, we examined seasonality of host beetle abundance, the frequency of parasitism, and the timing of fly eclosion. In addition, we evaluated host quality using numbers of larvae or puparia per individual beetle as a measure of quality. The fly parasitized only large carabids (≥15 mm body length); the lengths of fly puparia reached 7.4–10.8 mm during development in beetle abdomens, and larger hosts are likely essential. Of the 18 large carabid species collected in this study, we chose two, Carabus maiyasanus Bates and Leptocarabus procerulus (Bates), because they were large and abundant (87% of total catch). The two carabids had different phonologies; C. maiyasanus was abundant from spring to summer, and its abundance dropped sharply in autumn, while L. procerulus was abundant in autumn and rare from spring to summer except July. Parasitism was observed in all the months from May to November except June, and adult flies eclosed more than once a year (in early summer, late summer, and mid‐autumn), indicating that the species is multivoltine. Host quality of L. procerulus was higher than that of C. maiyasanus. Carabus maiyasanus was mainly used as a host from spring to summer, and L. procerulus was used in autumn. Thus, adult beetles of one or both species are available over most of spring, summer, and autumn, allowing population persistence of this fly species over time.