Fecundity in relation to wing-morph of three closely related species of the melanocephalus group of the genus Calathus (Coleoptera: Carabidae)

Summary In two successive years the fecundity of the carabid beetles Calathus (Neocalathus) cinctus, C. (N.) melanocephalus and C. (N.) mollis was studied in relation to wing-morph and temperature. Differences were found between the three species in both egg production and timing and length of the oviposition period. In all species the fecundity of laboratory bred beetles was significantly higher than that of females collected in the field. Long-winged females of both cinctus and melanocephalus had significantly higher egg production than short-winged females, and they also tended to produce eggs over a longer period. In mollis only the fecundity of the long-winged morph was established. The observed lower relative fitness of the short-winged morph in both cinctus and melanocephalus contradicts the supposed increase of the frequency of this morph in ageing, more or less isolated, populations of these species. The loss of long-winged genotypes, resulting from flight activities, is considered the most plausible cause of the increase of short-winged beetles in ageing populations. The higher fecundity of macropterous females makes them especially suited for (re)establishing populations.Communication No. 429 of the Biological Station WijsterPresent address and address for offprint requests: Kortenburg 31, NL-6871 ND Rentum     

Seasonal Development and Population Fluctuations of Arrowhead Scale,Unaspis yanonensis (Homoptera: Diaspididae), in Citrus orchards in Jeju,Korea

Arrowhead scale, Unaspis yanonensis (Kuwana), feeds on the foliage, stems, and fruits of citrus trees, and causes tree dieback when heavy infestations occur. The objective of this study was to collect basic data to establish seasonal management strategies for U. yanonensis. The abundances of male nymphs and female adults were monitored in unsprayed citrus orchards. The overwintered females started to produce their progenies (1st gen.) from mid-May. The populations of the 1st nymphs in the 1st generation showed a bimodal occurrence pattern, with a 1st larger peak between late May and early June, and a 2nd smaller peak in June. The nymphs in the 2nd generation, which were laid from newly developed females, were observed from late July, and the 1st peak occurred in mid-August, followed by an obscure later peak (the 2nd peak) around late September. The numbers of live females for the overwintered populations began to decrease gradually from early June by the impact of the predacious Chilocorus kuwanae, as well as by natural mortality factors, and dropped to a low level in mid-July. When there were abundant predator attacks, the U. yanonensis female populations sharply decreased from early June and were almost nonexistent in July. New female adults (1st generation adults) appeared from late June and peaked in mid-July. The 2nd generation female adults, consequently, the overwintering generation, occurred from mid- September, and gradually increased to late October. Overall, these results will be useful in terms of management strategies for U. yanonensis in citrus orchards.     

Survival, longevity and fecundity of overwintering Mesoplatys ochroptera Stål (Coleoptera: Chrysomelidae) defoliating Sesbania sesban (Leguminosae) and implications for its management in southern Africa

1 The defoliator beetle, Mesoplatys ochroptera Stål, is a serious pest of the legume tree sesbania (Sesbania sesban (L.) Merrill) in agroforestry systems in southern Africa. The survival of the overwintering stage of M. ochroptera and post‐ emergence longevity and fecundity of adults in sesbania fallows in eastern Zambia were quantified. 2 Only adult M. ochroptera survived during the winter (May–August) and the dry season (September–October) hidden under weeds, plant litter, in soil cracks and under rocks. During these periods, some beetles were occasionally found feeding on S. sesban, particularly in natural stands in humid areas. The adults were parasitized by the braconid Perilitus larvicida van Achterberg at a rate of 8.5–16.4%. The beetles stayed in the winter refuges for up to 210 days and emerged with the first heavy rains in October–November. The effective survival of beetles overwintering in an insectary was 18.1% in 1998 and 37.7% in 1999, and that of beetles overwintering in the field was estimated at 0.4 and 2.8% in 1998 and 1999, respectively. 3 The post‐emergence longevity and oviposition period of females in the insectary was 14–31 days and 5–29 days, respectively. The post‐emergence fecundity varied from 87 to 783. 4 It is concluded that the serious M. ochroptera infestation observed every year following the rains is due to synchronous emergence of overwintered resident adult populations and their high capacity for reproduction on many species of Sesbania. Recommendations for pest management in sesbania planted‐fallows are given.     

Seasonal aspects of life cycles in Calathus melanocephalus and C. micropterus (Coleoptera, Carabidae) in the northern taiga

The seasonal dynamics of the activity and the demographic structure of Calathus melanocephalus and C. micropterus populations were studied in the northern taiga of Arkhangelsk Province. The period of adult activity lasts from early June to mid-or late September with the maximum surface activity observed in the middle of summer. In C. melanocephalus, mature individuals are recorded from early June to early September, and in C. micropterus, during the entire season. The data on seasonal changes of the demographic structure of the population demonstrate that in the northern taiga, these species probably possess a biennial life cycle with summer reproduction. The geographical variability of the demographic structure of populations and of life cycles of two species of the genus Calathus was demonstrated. Northwards from the south, the period of activity decreases and the period of reproduction increases and shifts from autumn to the middle of summer. A hypothesis was formulated, according to which in the species studied the annual life cycle in the central part of the range is replaced by the biennial cycle in the north.     

A Monte Carlo simulation model for studying evolution in age-structured populations

This model provides for any number of genotypes defined by age-specific survival and fecundity rates in a population with completely overlapping generations and growing under the control of density-governing functions affecting survival or fecundity. It is tested in situations involving two alleles at one locus. Nonselection populations at Hardy–Weinberg equilibrium obey the ecogenetic law; i.e., each genotype follows Lotka's law regarding rate of increase and stable age distribution as if it were an independent true-breeding population. Populations experiencing age- and density-independent selection approximate this situation, and the changes in gene frequency are predicted by relative fitnesses bases on λ, the finite rate of increase of the genotypes. Polymorphic gene equilibria occurring at steady-state population sizes are determined by fitnesses based on R, the net reproductive rate. In examples involving differences in generation time produced by age-dependent differences in fecundity, the allele associated with longer generation time may be favored or opposed by selection, depending on whether the density-governing factor controlling population size affects survival or fecundity. If such genotypes have similar R's, a genetic equilibrium may be established if the population is governed by a density function acting upon fecundity. Received: August 23, 1999 / Accepted: July 13, 2000     

Notes on the biology and ecology of the parasitoids of the poplar clearwing moth, Paranthrene tabaniformis (Rott.) (Lep., Sesiidae) in Bulgaria. II. Eriborus terebrans (Gravenhorst, 1826) (Hym., Ichneumonidae)

The biology and ecology of Eriborus terebrans (Grav.), a parasitoid of the poplar clearwing moth, Paranthrene tabaniformis (Rott.), were studied during the period 1987–98. One-year-old poplar ( Populus spp.) shoots infested with P. tabaniformis larvae were collected from poplar seedlings at 11 localities in Bulgaria and examined in both field and laboratory conditions. Eriborus terebrans was recorded in seven localities as a solitary internal larval parasitoid of P. tabaniformis which developed two generations in early and mid-stage host larvae. Eriborus terebrans overwintered as a larva in P. tabaniformis overwintering larvae. In the field adult parasitoids of the overwintering generation appeared between late April/early May, and June or July. The peak activity of E. terebrans adults only coincided with the beginning of host emergence, which resulted in low levels of parasitism, being no more than 6.2%. Parasitoid adults of the summer generation appeared in late June–mid August. In this period enough larvae of the host were suitable for attacking and parasitism reached 24.4–39% in some cases. The average mortality of P. tabaniformis overwintering larvae caused by this parasitoid in Bulgaria during the period of the study was 4.7%. A significant part of the parasitized P. tabaniformis larvae constructed tunnel structures of frass and silk threads over the external openings of the galleries. It is possible that these structures protect the parasitoid cocoons from natural enemies – hyperparasitoids and predators.     

The life histories and population dynamics of two carabid species on a Dutch heathland

Summary We deal with the causes of the synchronously fluctuating numbers of subpopulations of the carabid species Calathus melanocephalus as compared with the asynchronously fluctuating numbers of subpopulations of the carabid Pterostichus versicolor. Both species continuously occupy a large heath area, Dwingelder Veld (1600 ha), in The Netherlands, and are studied there in the same localities with the same methods. Of the adults of C. melanocephalus, 90% do not cover more than 2 ha during the entire reproductive season, while 90% of adults of P. versicolor cover no more than 12 ha. In C. melanocephalus egg production in the field is usually similar to that under optimal feeding conditions in the laboratory, but in P. versicolor egg production seems to be much lower in the field. In the field 70–80% of the eggs most probably are killed by eelworms, followed by more than 90% mortality among the remaining larvae. Comparing mortality of developmental stages in laboratory experiments with that in field experiments in enclosures, it appears that mortality of larvae is not density-dependent, even when density in the experiments is much higher than it ever is in the field. Larval mortality mainly results from the poor ability of the larvae to find prey, even when in field experiments prey density is increased far above natural densities. We discuss why these poor prey-finding abilities are not improved by natural selection. In the spring breeder P. versicolor differences between localities both in abiotic factors, soil moisture and surface temperature, and biotic factors, reactions of prey species to abiotic factors, in spring and summer when the larvae are maturing contribute to the asynchronous fluctuations of numbers between subpopulations. In the autumn breeder C. melanocephalus possible differences in biotic factors between sites are outnumbered by the effects of winters with a higher or lower than normal amount of precipitation respectively. During a wet winter mortality among the larvae is much higher than during a dry winter. As these winter conditions are similar over large areas (many km²) the fluctuations of numbers between subpopulations are synchronous.Communication No. 443 of The Biological Station, Wijster     

Winter survival and oviposition before and after overwintering of a parasitoid wasp, Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae)

Winter survival and oviposition before and after overwintering in Ooencyrtus nezarae, an egg parasitoid of phytophagous heteropterans, were examined in Osaka, Japan. Eggs of Riptortus clavatus parasitized by O. nezarae were kept under natural photoperiod and temperature. When honey was supplied, some female adults emerging from early September to late November overwintered. The percentage of overwintering individuals increased as the date of adult emergence advanced. Most female adults supplied with honey and hosts oviposited soon after emergence, then stopped laying eggs. Female adults emerging in mid‐October and early November laid eggs and then overwintered. The induction of diapause in the field seems to vary greatly depending on host availability. Without honey, the survival time of female adults was very short, whether host eggs were supplied or not. After overwintering, most females began to lay eggs in early May if host eggs were supplied, and they produced both male and female progeny. In the study area, a legume field in Osaka, parasitization by O. nezarae was observed from early July to November.     

Breeding of the bush fly,Musca vetustissima Walker,in an over-wintering area during winter and spring

Field experiments in an area of south-western Australia where bush fly (Musca vetustissima) adults occur permanently, showed that it overwintered by continuous breeding; but only two to three generations occurred between June and September. Low survival of eggs and larvae in the food (cattle dung) from May to August was associated with rainfall rather than low temperatures. High egg-adult survival occurred in late August; lower egg and larval survival in September and early October was attributed partly to dung fauna. Egg and larval survival was high in late October until December but predation or parasitization caused low survival of puparia outside the dung. Experimental and wild adult flies were largest and therefore most fecund in early spring; smaller wild flies in late spring appeared to result from larval competition for food. Simultaneous high egg-adult survival, high fecundity and rapidly decreasing generation times in late winter and early spring provide a basis for explaining the major increase in adult bush fly abundance observed in some areas in mid-spring.     

The evolutionary significance of wing dimorphism in carabid beetles (Coleoptera: Carabidae)

A review of data on the background of wing dimorphism in carabid beetles (Coleoptera: Carabidae) and especially of the closely relatedCalathus cinctus andC. melanocephalus is given. In bothCalathus species wing dimorphism is inherited in a simple Mendelian fashion with the brachypterous condition dominant, but inC. melanocephalus the expression of the long winged genotype is under environmental control as well. The development of long winged phenotypes in the latter species is favoured by relatively favourable environmental conditions, such as high temperatures and a high food-supply. The higher fecundity of the larger and heavier long winged females of both species may compensate for losses of long winged phenotypes by flight activities. The evolutionary significance of both types of inheritance is discussed in relation to dispersal. The ‘fixed type’ as found inC. cinctus is considered an opportunistic short term ‘between sites strategy’, whereas the ‘dynamic type’ ofC. melanocephalus represents a flexible long term ‘within sites strategy’.     

Life History of Ilybius fenestratus (Fabricius) (Coleoptera,Dytiscidae) in a Central Norwegian Lake

The life history of Ilybius fenestratus was studied in Målsj?en, a lake in S?r-Tr?ndelag, Central Norway (63°14'N, 10°26'E), during 1971–1972. Adults and larvae were sampled in activity traps every week during March–October and every 2–3 weeks during November–February. Newly-emerged adults were generally found from mid-July to September, and after overwintering in torpidity, probably in the water, they again appeared in the second half of June to August/September. Laying of eggs took place from early/mid July to early August. Larvae occurred in the traps from late July to early May. I. fenestratus was thus found to be a semivoltine summer breeder, with overwintering larvae the first winter and overwintering adults the next. Both adults and larvae lived in large areas of the lake's littoral zone.     

Gyne development in the Argentine ant Iridomyrmex humilis: role of overwintering and queen control

Abstract A possible stimulatory effect of overwintering on gyne development in Iridomyrmex humilis (Mayr) was investigated. Although gyne-potent larvae are present in the nest throughout the year, small queenless colony fragments composed of freshly overwintered ants (collected in late winter or early spring) produced 3–8 times more gynes than comparable fragments made up of non-freshly overwintered ants (collected at other times of the year). Apparently, this stimulatory effect of overwintering acts on both the developmental potential of larvae and the tendency of workers to rear sexually competent larvae as gynes; queenless colony fragments in which either the brood or workers were freshly overwintered produced more gynes than fragments composed of non-freshly overwintered workers or brood but fewer than fragments containing both brood and workers freshly overwintered. This increased sexualization potential of larvae due to overwintering is enough to overcome weak pheromonal inhibition of queens associated with low queen number; a single queen in a large freshly overwintered colony fragment is insufficient to inhibit gyne development, whereas ten queens are totally inhibitory. In non-freshly overwintered colony fragments one queen is completely inhibitory. Variability in egg developmental potential according to queen age does not appear to play a major role in the seasonal production of gynes, because at least some eggs of very young queens (less than 3 weeks old) are capable of gyne development. In the field this stimulatory effect of overwintering is superimposed on a seasonal fluctuation in the combined strength of pheromonal queen control. In southern France, gynes are produced only in spring where they arise primarily from overwintered larvae just after a sharp drop in queen number, and presumably the total level of inhibitory queen pheromone, due to the massive execution of queens by workers.     

中华稻蝗长沙种群的生活史及其卵滞育的进化意义

中华稻蝗Oxya chinensis为重要的水稻害虫,在我国除青海、西藏、新疆、内蒙古等未见报道外,南起海南北至东北均有分布,在许多的分布区域1年发生1代。为探索中华稻蝗长沙种群的生活史及其季节适应特征,通过野外和实验室的研究,调查了其发生代数、孵化率和卵滞育率的季节变化及越冬卵的存活率。结果显示,中华稻蝗长沙种群为1年2代和1年1代混合发生:第1代卵产卵后大部分孵化为若虫而1年完成2代,但亦有19.4% -4.1%的卵不孵化而1年只能完成1世代。第1代成虫于6月上旬至8月上旬羽化,6月下旬至8月中旬产卵;第2代若虫于7月初开始孵化,9-10月羽化为成虫,10月上旬至11月下旬产卵。在室外自然条件下,中华稻蝗长沙种群6-8月(第1代)和10-11月(第2代)所产卵块均为部分滞育,滞育率为30%左右,皆无显著差异。然而,其卵滞育率在12月以后显著降低,仅为6.6%或以下,卵滞育快速地得以解除。因此,包括非滞育卵和滞育解除卵,中华稻蝗长沙种群的越冬卵皆以非滞育状态度过其后的寒冷季节。即使是遭遇长江流域2007年末至 2008年初异常寒冷的冬季,在长沙地区越冬后其卵的存活率亦在98%以上。非滞育状态的中华稻蝗长沙种群越冬卵完全能安全地越冬,其滞育的发生并非是为了提高其耐寒性而安全度过不适环境。并探讨了中华稻蝗长沙种群卵滞育的进化意义。     

Factors influencing the composition of mixed populations of a hemiclonal hybrid and its sexual host

Hemiclonal/hybridogenetic hybrids combine demographic superiority of asexuals and genetic diversity of sexuals, but their need for backcrossing with a parental species tightly couples them with this sexual host. How can systems like this persist in ecological and evolutionary time? Two discrete‐time mathematical models describing the complex life cycle and mating system of hybridogenetic waterfrogs (Rana esculenta) identified four factors and their interactions as important. Although female mating preferences, in combination with differences in fecundity, determine species coexistence, differences in larval competitiveness seem to be more important for the hybrid‘s actual frequency. However, coexistence is possible even when host and hybrid are equally fecund and competitive. Dispersal and competition interact in their influence on species composition, but ecological and reproductive dispersal has opposing effects. In ecological terms our results explain the remarkable stability of observed species ratios over time within natural hybridogenetic populations, and indicate why the species composition can vary so widely between localities. In evolutionary terms they explain the old age of these and other hybridogenetic systems. They also suggest interesting consequences for other tightly coupled systems.     

Herbivory below ground and biological weed control: life history of a root-boring weevil on purple loosestrife

The life history of the root-boring weevil Hylobius transversovittatus was studied in north and central Europe. The weevil develops on Lythrum salicaria, a perennial marshland plant that has become a problem weed in North America. It was found in all habitats of its host plant with the exception of permanently flooded sites. It also attacked L. salicaria in an early successional stage, devastating large parts of the storage tissue. The beetle was found in two-thirds of the field populations examined with a mean attack rate of 76.3%. Larvae developed according to a 1- or 2-year generation cycle depending on the time of oviposition. Adult beetles developing within 1 year emerged between July and October, whereas beetles with a 2-year larval period emerged within 3 weeks in July and August. Adults were long-lived and could hibernate several times. Beetles of the new generation mated right on the spot and some oviposited prior to overwintering. Hibernated females had an oviposition period from May to early September and produced 3–4 eggs/day during the peak oviposition period. Specific mortality factors were scarce. Dispersal flights ensure the regular occurrence of H. transversovittatus in scattered L. salicaria populations. The severe impact of the weevil is expected to reduce the competitive ability of its host plant after introduction into North America.     

Life-cycle variation inPanonychus akitanus Ehara (Acarina:Tetranychidae)

The life history ofPanonychus akitanus Ehara was studied in two Hokkaido populations on dwarf bamboos. The Sapporo population overwintered both as egg and female adult onSasa senanensis, and the Tomakomai population overwintered as egg onSasa apoiensis. Mites of the Sapporo population produced four or five generations from late April to late November or early December. The eggs that had overwintered began to hatch in mid-May, and this hatching period overlapped with that of eggs laid in late April by females that had overwintered. Therefore, mites with different overwintering stages would interbreed. Most eggs that had overwintered in the Tomakomai population hatched in mid-May, and about four generations were produced before early December, when only eggs were found. The density of mites per leaf of the Sapporo population reached a maximum in early May on old leaves and in late June on new leaves, and thereafter gradually decreased. The Tomakomai population initially decreased in density after hatching in the spring, but rapidly dispersed to new leaves, reached a peak in early September, and then gradually decreased. The maximal density was about 10 times higher and the distribution was less clumped (lower values of the aggregation index,m/m) than that of the Sapporo population. This was probably because the Sapporo mites could utilize only the underside of sporadically distributed leaves which were curled by spiders, whereas the Tomakomai mites inhabited any part of the leaf undersurface of the hairy host plant. Predators observed were phytoseiid mites and larvae of gall midges. The predatory effect on the Sapporo population was not clear. In the Tomakomai population, the number of gall midges correlated with the number of spider mites better than that of phytoseiid mites.     

Seasonal variations in body melanism and size of the wolf spider Pardosa astrigera (Araneae: Lycosidae)

Variations in species morphology and life‐history traits strongly correlate with geographic and climatic characteristics. Most studies on morphological variations in animals focus on ectotherms distributed on a large geographic scale across latitudinal and/or altitudinal gradient. However, the morphological variations of spiders living in the same habitats across different seasons have not been reported. In this study, we used the wolf spider, Pardosa astrigera, as a model to determine seasonal differences in adult body size, melanism, fecundity, and egg diameter both in the overwintering and the first generation for 2010 and 2016. The results showed that in 2010, both females and males of the overwintering generation were significantly darker than the first generation. Moreover, the overwintering females were markedly larger and produced more and bigger eggs than the first generation in both 2010 and 2016. Considering the overwintering P. astrigera experiencing low temperature and/or desiccation stress, these results suggest that substantially darker and larger body of the overwintering generation is adaptive to adverse conditions.     

The phenology of Lygus rugulipennis, the European tarnished plant bug, on late-season strawberries, and control with insecticides

The phenology of Lygus rugulipennis, the European tarnished plant bug, was studied in fields of strawberries producing late-season (July to September) crops. Adults invaded fields in small numbers, mainly in July. Large numbers of nymphs were found in late July and early August; these matured into adults from late August. Adults formed the overwintering population; some remained in strawberry fields but left in spring to feed on other host plants. All the insecticides tested, i.e. malathion, chlorpyrifos, heptenophos, cypermethrin, bifenthrin and trichlorfon, gave good short-term control of nymphs.     

Intraspecific variation of wing pointedness index in juvenile Acrocephalus warblers in the southeastern Baltic

Variation of wing pointedness index between groups of juveniles captured in different months (July, August, and September) and at different stages of juvenile moult was studied in three Acrocephalus warbler species captured on the Courish Spit on the Baltic Sea. Sedge warblers (Acrocephalus schoenobaenus) captured in July had less pointed wings than sedge warblers captured in August or September. Marsh warblers (A. palustris) showed no significant difference between birds in early and in late moult. No differences in wing pointedness were found between different cohorts of reed warblers (A. scirpaceus), including known locally hatched birds and late migrants captured in September. It is hypothesised that reed warbler populations in the northeastern Baltic are too evolutionarily young to have evolved a different wing shape as compared with the local Courish population.Communicated by F. Bairlein     

A comparison of the larval overwintering biology of the striped stem borer, Chilo suppressalis (Lepidoptera: Crambidae), in rice and water-oat fields

It has been suggested that the striped stem borer, Chilo suppressalis (Walker), has developed distinct populations inhabiting rice Oryza sativa (Linnaeus) and water-oats Zizania latifolia (Turcz). In this study, we compared several biological traits between overwintering larvae from rice and water-oat fields during the winter of 2010. The parasitism rate was significantly higher in the larvae that overwintered in the rice field than in those that overwintered in the water-oat field. The average body weight of overwintering larvae and the weight of 3-day-old pupae, as well as the number of eggs carried by adult females, were all significantly greater in the samples from the water-oat field. The diapause intensity, defined as the number of days required until pupation by overwintering larvae transferred to 25 °C and L:D 16:8 h, decreased progressively from January to March but decreased more rapidly in the water-oat field. Our results demonstrate differences in the overwintering biology of C. suppressalis larvae from rice and water-oat fields. These differences may contribute to the differentiation and reproductive isolation between host-plant populations in this species.