Continuous population estimates forDendroctonus frontalis Zimm. (Coleoptera: Scolytidae) occurring in infestations

Infestations of Dendroctonus frontalisZimm. are often observed to enlarge continuously by the colonization of new hosts in a pattern similar to a forest fire. This pattern of infestation growth presents unique problems in quantitatively estimating populations of D. frontalis. Beetle populations on each infested tree in an infestation go through five processes: attack, oviposition, reemergence, survivorship, and emergence. These processes, which have been described mathematically in the literature, each take several days for completion. In order to follow the distribution and abundance of D. frontalis throughout the course of development of a spot, we need a daily estimate of the number of beetles involved in each process on every tree. Since it is not practical to sample each tree daily, we developed a procedure whereby quantitative estimation procedures for within-tree populations were used in combination with the mathematical models for the life processes to produce a daily record of the number of adults successfully attacking trees, the number of eggs oviposited, the number of beetles reemerging, number of beetles surviving within the trees, and the number of beetles emerging. These daily estimates were then summarized for all trees in the spot for the duration of the infestation. The daily record of populations of D. frontalis, used with information on infestation geometry, were suggested to be of value in describing and elucidating several important facets of population dynamics including dispersal patterns within infestations, between tree beetle loss (mortality), and time lags among the various population processes. The information reported can be used to develop simulation models of population dynamics or to validate existing models.     

Delayed benefits of paternal care in the burying beetle Nicrophorus vespilloides

Burying beetles, Nicrophorus spp., inter the carcasses of small vertebrates as a food source for their offspring. Females can bury a carcass and rear a brood on it alone, but are frequently assisted by a male whose presence reduces the risk of the carcass being taken over by other beetles. However, the male often stays for longer than the carcass is vulnerable to take-over, and he cares for the brood without conferring any further benefits on it. In a laboratory experiment using N. vespilloides, we found that, in the absence of competitors, male assistance conferred no advantages on the brood for which he was caring, but significantly increased the subsequent reproductive success of his mate, in terms of the mass and rate of development of a second brood, reared alone. We suggest that this is due to a reduced parental effort of assisted females, who spent less time feeding offspring and more time resting than unassisted females whilst rearing their first broods. In the field, a female is unlikely to pair with the same male for consecutive broods, so we discuss the possible benefits a male may accrue from increasing his mate's reproductive success. We also discuss the relevance of these results to our understanding of the evolution of biparental care in birds. Copyright 2000 The Association for the Study of Animal Behaviour.     

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

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Parental care trade‐offs in the inter‐brood phase in Barn Swallows Hirundo rustica

In seasonal environments with limited time and energy resources, double‐brooded birds face trade‐offs in the timing of their two reproductive attempts and in the effort allocated to the first and the second broods. In the Barn Swallow Hirundo rustica a long care period for the first brood enhances the survival of first‐brood chicks, but also delays the start of the second brood, which in turn reduces the survival prospects of second‐brood chicks. Probably as a response to this trade‐off, double‐brooded Barn Swallows reduce the period of post‐fledging care for first‐brood fledglings. By radiotracking whole families, we investigated the determinants of this behaviour and its consequences for the survival of the first‐brood fledglings. The end of the females’ investment in post‐fledging care of the first brood was related to the beginning of egg synthesis for the second clutch. With the start of egg synthesis, females significantly reduced provisioning rates to the first‐brood fledglings to less than one‐fifth of the previous rates, while the proportion of time they spent foraging remained high. Assuming that the females’ foraging success was constant, we conclude that their energy income was allocated to egg production rather than fledgling provision. Males did not compensate for the females’ reduced feeding rates. Thus the start of egg production for the second clutch had a marked effect on the quantity of food received by first‐brood fledglings. In parallel with the changes in parental behaviour and provisioning rates, we observed a marked drop in the daily survival rate of first‐brood chicks. These results support the hypothesis that females face a strong trade‐off in the allocation of energy to subsequent broods. Energy allocation to a second clutch involves a cost in terms of reduced provisioning, and as a result the survival of first‐brood chicks is compromised. This is probably outweighed by the improved success of an early second brood.     

Survival of parent and brood adult bark beetles, Ips typographus, in relation to size, lipid content and re-emergence or emergence day

ABSTRACT. This study investigated a possible trade-off between bark beetle reproductive effort and future survival. Parent adult Ips typographus (L.) (Coleoptera: Scolytidae) were collected when they re-emerged from a laboratory culture after completing their first brood and held at constant temperatures of 4 or 25^oC. As a comparison, emerging brood adults were collected and held at 25^oC. The longevity of the beetles was related to when they re-emerged or emerged (measured in days since parent introduction) and to their size (pronotal width), fresh weight and lipid content at collection, by multiple regression analysis. The expression (fresh weight x pronotal width ^-2.6), used as an estimate of lipid content of the living beetles, was significantly correlated (r=0.67, P<0.001) with the actual (extracted) lipid content. At 25^oC, fat content was the variable most strongly correlated with survival time in all beetle groups except parent females. In both parent and brood females, re-emergence or emergence day, respectively, contributed significantly to the explanation of survival time, whereas pronotal width and fresh weight never contributed significantly to the regression equation. At 4^oC, fat content was not estimated but, re-emergence day was negatively correlated with survival time. In conclusion, beetles with high fat content and re-emerging or emerging early have longer expected survival than beetles leaving the log later or containing less fat. This means that females laying a large first brood might suffer a somewhat higher mortality than females laying a smaller brood.     

Emergence patterns of univoltine and bivoltine Ips typographus (L.) populations and associated natural enemies

Control measures aiming at reducing bark beetle populations and preserving their natural enemies require a sound knowledge on their overwintering and emergence behaviour. These behavioural traits were investigated in univoltine and bivoltine populations of the European spruce bark beetle (Ips typographus [L.], Coleoptera: Scolytinae) and its predators and parasitoids over several consecutive years. In univoltine populations, roughly 50% of the bark beetles left their brood trees in fall together with most parasitoids and some significant predatory flies and beetles. In bivoltine populations, <10% of the second bark beetle generation emerged in fall and the remainder overwintered under the bark of their brood trees. Likewise, most predatory beetles and flies spent wintertime with their prey under the bark, while most parasitic wasps emerged in fall. The spring emergence of bivoltine predatory beetles was found to occur up to 3 weeks earlier than that of I. typographus, while that of the predatory flies and the parasitoids was delayed by up to 1 month. In univoltine populations, the bark beetles emerged several weeks prior to most antagonistic taxa. In the heat year 2003, three I. typographus generations were produced at the lower location, 36% of the third generation emerged in fall, while the proportions of overwintering predators remained largely the same as in previous years. Similar to their host, more parasitoids left their brood trees in fall after warm years. The results show that sanitation felling during winter probably kills most bark beetles in bivoltine populations, but also eliminates many natural enemies. In univoltine populations, sanitation felling might be less detrimental to both I. typographus and natural enemies because a fair fraction of their populations will already have left the trees before cutting. Warmer climates may affect the interactions of bark beetles and natural enemies and thus the impact of control measures.     

Evaluation of funnel traps for characterizing the bark beetle (Coleoptera: Scolytidae) communities in ponderosa pine forests of north-central Arizona

Lindgren funnel traps baited with aggregation pheromones are widely used to monitor and manage populations of economically important bark beetles (Coleoptera: Scolytidae). This study was designed to advance our understanding of how funnel trap catches assess bark beetle communities and relative abundance of individual species. In the second year (2005) of a 3-yr study of the bark beetle community structure in north-central Arizona pine (Pinus spp.) forests, we collected data on stand structure, site conditions, and local bark beetle-induced tree mortality at each trap site. We also collected samples of bark from infested (brood) trees near trap sites to identify and determine the population density of bark beetles that were attacking ponderosa pine, Pinus ponderosa Douglas ex Lawson, in the area surrounding the traps. Multiple regression models indicated that the number of Dendroctonus and Ips beetles captured in 2005 was inversely related to elevation of the trap site, and positively associated with the amount of ponderosa pine in the stand surrounding the site. Traps located closer to brood trees also captured more beetles. The relationship between trap catches and host tree mortality was weak and inconsistent in forest stands surrounding the funnel traps, suggesting that trap catches do not provide a good estimate of local beetle-induced tree mortality. However, pheromone-baited funnel trap data and data from gallery identification in bark samples produced statistically similar relative abundance profiles for the five species of bark beetles that we examined, indicating that funnel trap data provided a good assessment of species presence and relative abundance.     

The dispersal behaviour of the phoretic mite Poecilochirus carabi (Mesostigmata,Parasitidae): adaptation to the breeding biology of its carrier Necrophorus vespilloides (Coleoptera,Silphidae)

Summary When the phoretic mite Poecilochirus carabi reproduces in the brood chamber of its carrier Necrophorus vespilloides, a beetle with biparental brood care, the first deuteronymphs of the new mite generation aggregate on the male beetle. They do not use sex-specific traits to discriminate between male and female beetles in the brood chamber, but traits that are related to the beetles' behaviour and may be displayed by both parent beetles. When the male beetle departs, it carries virtually all deuteronymphs then present in the brood chamber. Deuteronymphs that develop later congregate on the female, which leaves the crypt some days after the male. Only those deuteronymphs that miss the female's departure disperse on the beetle larvae, meaning they have to wait in their pupal chambers until the beetles have completed their development. On average, 86% of the deuteronymphs leave the brood chamber on the parent beetles, thereby gaining the advantage of an early departure. As soon as their carrier arrives at one of the beetles' meeting places, the deuteronymphs can transfer between the beetles present. Choice experiments revealed that the deuteronymphs tend to even out density differences between congregating carriers, and prefer sexually mature to immature beetles. Therefore, transferring between beetles results in a dispersion of deuteronymphs on the sexually mature beetles of the population.     

Population density ofMonochamus alternatus adults (Coleoptera: Cerambycidae) and incidence of pine wilt disease caused byBursaphelenchus xylophilus (Nematoda: Aphelenchoididae)

Summary The seasonal occurrence ofMonochamus alternatus and newly weakened trees were investigated in aPinus thunbergii stand for 4 years. Adult beetles were present between June and September with a peak in their population occurring in early July followed by a decline then a period of about one month being in a steady number. The average number ofBursaphelenchus xylophilus (Nematoda), which is the causal agent of pine wilt disease, within beetles decreased as the season advanced. Pine trees newly weakened byB. xylophilus appeared between June and October, especially from August to October. The proportion of weakened or killed trees was directly proportional to the average beetle density per tree from June to August.     

Density-dependent decline of breeding success in an introduced, increasing mute swan Cygus olor population

An increasing population of the mute swan Cygnus olor , was studied from its very establishment in 1976 until 1998. As the number of pairs increased, there was a decline in all production parameters, including the average number of clutches per pair, the average number of broods per clutch, and the average number of fledged young per brood. In a multiple regression analysis covering the whole breeding season, the number of pairs and the average number of clutches per pair explained 71% of the variation in the average number of fledglings per pair. During the brood season, the average number of fledglings per brood was an additionally important parameter in explaining fledgling production. Habitat quality seemed to affect breeding: in places occupied earlier more cygnets fledged from a clutch than in habitats inhabited later. The decreased production of young very likely reflects density-dependent effects on reproduction. This density dependence seems to operate on the breeding grounds, since winter harshness did not affect breeding success. Density-dependent processes started acting already when pairs were beginning to nest, and continued during the brood period. Density dependence has apparently not been detected at the pair stage in earlier studies of bird populations.     

Habitat selection and offspring survival rate in three paracoprid dung beetles: the influence of soil type and soil moisture

Paracoprid dung beetles build brood chambers in the soil beneath a dung pat and provide them with dung Onthophagus species lay one egg into each chamber This paper deals with the influence of soil type and soil moisture on micro-habitat selection and survival of offspring m three middle-European Onthophagus-species ( O coenobita, O fracticonis and O vacca) Discrimination between sandy soils with three different loam contents (0%, 20%, 40%) and four different water contents (4%, 8%, 12%, 16%) was tested in the laboratory During the first 24 h of each replicate beetles which colonized one of the patches did not distinguish between different soil conditions Emigration rates, measured as time when 50% of all individuals had left the patch, and numbers of brood chambers proved to be species specific and depended on soil moisture and soil type Survival rates of the larvae in the brood chambers were influenced nearly exclusively by soil moisture The results are discussed in relation to the ecology of the three species and in context with optimal foraging theories     

Adverse effects of ivermectin on the dung beetles, Caccobius jessoensis Harold, and rare species, Copris ochus Motschulsky and Copris acutidens Motschulsky (Coleoptera: Scarabaeidae), in Japan

Effects of the antiparasitic drug, ivermectin, on the dung beetles, Caccobius jessoensis Harold, 1867 and the rare species, Copris ochus Motschulsky, 1860 and Copris acutidens Motschulsky, 1860 were studied in laboratory and field experiments in Hokkaido, Japan. Ivermectin was detected in dung from 1 to 21 or 28 days following treatment, with a peak on the first day after treatment in two pour-on administrations (500 microg kg(-1)), although there were considerable differences between the two peaks. In C. jessoensis, brood balls constructed by the female were not reduced in the dung of treated cattle except for seven days after treatment in experiment 2. Also, there was no significant difference in the mean weight of brood balls between dung from treated and control cattle. However, the emergence rates were significantly reduced in dung 1-3 days after treatment. In the field study, brood balls constructed by C. jessoensis were more abundant in dung from treated cattle in experiment 1, but adult emergence was significantly reduced at one and seven days after treatments. Adult mortality of C. ochus Motschulsky at 90 days after the beginning of rearing was 11.1% in dung from control cattle with 22 brood balls constructed, whereas it was 84% in dung from treated cattle with no brood balls and/or ovipositioning. Also, in C. acutidens Motschulsky, adult mortality at 90 days after the beginning of rearing was 3.6% in dung from control cattle with 13 brood balls constructed, whereas it was 94.1% in dung from treated cattle with no brood balls or ovipositioning. The environmental risk in the use of ivermectin during breeding period of dung beetles in pasture is discussed.     

Seasonal variation in parental care, offspring development, and reproductive success in the burying beetle, Nicrophorus vespillo

1. Beetles of the genus Nicrophorus reproduce on small vertebrate carcasses that they bury in the soil to provide the larvae with food. Usually, both parents cooperate in brood care by feeding and guarding their progeny. 2. In pairs of the common European species N. vespillo, the duration of care depended on the time of year when the beetles reproduced. Both in 1990 and in 1991, male and female parents stayed longer with their broods when reproduction started in spring than when reproduction started in early or late summer. This was probably due to the longer development time of the larvae caused by lower temperatures in spring, because laboratory experiments suggested a strong influence of temperature on both the duration of brood care and offspring development. 3. The number of adult offspring produced by a beetle pair did not vary among different times of the year. 4. The median time required for offspring development, measured as time from burial of the carcass to emergence of young adults, was between 62 and 84 days. When the beetles reproduced in late summer, only about three-quarters of the offspring left the soil and hibernated as adults. The remaining offspring stayed underground and adults appeared on the soil surface the following spring. They still showed the flexible cuticle typical of newly-hatched beetles, suggesting that they may have overwintered in a pre-adult stage.     

Efficacy of systemic insecticides for protection of loblolly pine against southern pine engraver beetles (Coleoptera: Curculionidae: Scolytinae) and wood borers (Coleoptera: Cerambycidae)

We evaluated the efficacy of the systemic insecticides dinotefuran, emamectin benzoate, fipronil, and imidacloprid for preventing attacks and brood production of southern pine engraver beetles (Coleoptera: Curculionidae: Scolytinae) and wood borers (Coleoptera: Cerambycidae) on standing, stressed trees and bolt sections of loblolly pine, Pinus taeda L., in eastern Texas. Emamectin benzoate significantly reduced the colonization success of engraver beetles and associated wood borers in both stressed trees and pine bolt sections. Fipronil was nearly as effective as emamectin benzoate in reducing insect colonization of bolts 3 and 5 mo after injection but only moderately effective 1 mo after injection. Fipronil also significantly reduced bark beetle-caused mortality of stressed trees. Imidacloprid and dinotefuran were ineffective in preventing bark beetle and wood borer colonization of bolts or standing, stressed trees. The injected formulation of emamectin benzoate was found to cause long vertical lesions in the sapwood-phloem interface at each injection point.     

Development of Copris tripartitus (Coleoptera: Scarabaeidae) in Two Different Rearing Media

ABSTRACT Like many other sub-social insects, Copris tripartitus Waterhouse requires parental care during the development period in the brood balls. Mass-rearing is achieved by using indoor rearing systems and adding more dung after taking away the brood balls containing eggs. Without parental care, larvae are often affected by desiccation, as well as by fungal or sciarid fly intrusion of the brood balls. This study developed a good substitute for brood ball rearing medium. A significantly greater number of progeny survived to adulthood from broods produced by beetles reared on vermiculite medium (90.69±10.98%) than on peat moss medium (44.82±13.92%). Results of the study suggest that to protect the brood balls from desiccation and to produce healthy adults, 15-20% moisture content vermiculite must be used as brood ball rearing medium for most species reared in the laboratory.     

Effect of host tree seasonal phenology on substrate suitability for the pine engraver (Coleoptera: Scolytidae): implications for population dynamics and enemy free space

This study evaluated the effects of seasonal phenology on the substrate quality of susceptible hosts to the pine engraver, Ips pini (Say). We also determined the effects of the duration and method of storage on host quality for purposes of laboratory rearing. Live red pine trees were felled at various times during the season, and I. pini adults from a laboratory colony were established on the logs. Subsamples of logs were stored for various intervals, and then provided to beetles. Subsamples of stored logs were waxed at both ends to prevent water loss before being submitted to the same assays. Suitability of red pine phloem tissue in susceptible hosts declined for I. pini throughout the growing season. As the season progressed, the number of beetle progeny that emerged from colonized hosts dropped substantially. This decline was associated with simultaneous reductions in phloem moisture content. Reduction in host suitability may partially offset any advantage I. pini may gain from colonizing trees after the major predators have become less abundant. Bark beetle brood production decreased significantly with length of storage, regardless of the month of tree felling or the method of storing. Implications for bark beetle population dynamics and laboratory rearing systems are discussed.     

Temporal variation in mycophagy and prevalence of fungi associated with developmental stages of Dendroctonus ponderosae (Coleoptera: Curculionidae)

Mycophagy by bark beetles is widespread. However, little is known regarding which developmental stages of bark beetles actually feed on fungi. To study this question, we sampled fungi associated with Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae) throughout development in naturally attacked trees. Isolations of fungi were made from phloem adjacent to brood and from brood exoskeletons and guts. Overall, the incidence of fungi with individual brood increased as brood development progressed. Grosmannia clavigera (Robinson-Jeffrey and Davidson) Zipfel, de Beer and Wingf. and Ophiostoma montium (Rumbold) von Arx exhibited generally opposing trends in prevalence. G. clavigera was most likely to be found in phloem adjacent to prewintering third- and postwintering fourth-instar larvae. O. montium was most likely to be found in phloem adjacent to eggs, first-instar larvae, pupae, and teneral adults. In contrast to isolations made from phloem, fungi isolated from brood guts and exoskeletons were not observed to shift in prevalence. First- and third-instar larvae were often observed migrating to older portions of their galleries, indicating that they do not spend all of their time feeding at, and extending, the apex of the gallery. Our results suggest that not only are D. ponderosae brood in contact with and feeding on fungi throughout development, but also, that during development, contact of brood with a particular fungus is likely to change. Such temporal shifts in fungal symbionts may be environmentally driven and have important implications in how these fungi interact with their hosts within and across generations.     

Egg laying, maternal care and development of young in the scolopendromorph centipede, Cormocephalus anceps anceps Porat

The methods used to observe the construction of the brood chamber and rearing of the young by the female Cormocephalus anceps are described. The period of egg laying and the maternal care, essential for the survival of both eggs and brood, were observed and the behaviour of the female during this time is recorded. Two broods were successfully reared to first year adolescent stages in the laboratory. The two embryonic and three adolescent stages are briefly described and their developmental times recorded. Subsequent stages of development are noted by increase in size until sexual maturity is attained in the second year. The length of life and number of broods per female remains to be investigated.     

A field experiment on dispersal of newly emerged adults ofMonochamus alternatus (Coleoptera: Cerambycidae)

Newly-emerged adults of Monochamus alternatus aged 1 to 5 days were code-numbered with lacquer paint and released by placing them on the trunks of one or two trees in a Pinus thunbergii stand at weekly intervals during the beetle emergence period from 1980 to 1983. Beetles were captured at weekly intervals from one week after the first day of release. Determinations were made on the distance and direction of beetle dispersal during a week after release and analysed by a method of Inoue (1978). When the stand canopy was closed, the rate of beetle's stay on trees was 0.56 per week. The beetles dispersed at random by walk and flight. When the pine stand was sparse, the rate of beetle's stay on trees was 0.02–0.30 per week. They dispersed at random by flight. The average distances traversed were estimated to be 7.1–37.8 m for the first week after emergence. Using other method, the average distance traversed was estimated to be 10–20 m for each week through the first 3 weeks after release. The results of stepwise multiple regression analysis and a simple field experiment suggested that the dispersal of newly-emerged beetles was affected by stand density, number of beetles emerging from individual dead trees, maximum air temperature, and precipitation.     

Reemerging vivax malaria: changing patterns of annual incidence and control programs in the Republic of Korea

Changing patterns of the reemerging Plasmodium vivax malaria in the Republic of Korea (South Korea) during the period 1993 to 2005 are briefly analyzed with emphasis on the control measures used and the effects of meteorological and entomological factors. Data were obtained from the Communicable Diseases Monthly Reports published by the Korea Center for Disease Control and Prevention, and webpages of World Health Organization and United Nations. Meteorological data of Kangwon-do (Province) were obtained from local weather stations. After its first reemergence in 1993, the prevalence of malaria increased exponentially, peaking in 2000, and then decreased. In total, 21,419 cases were reported between 1993 and 2005 in South Korea. In North Korea, a total of 916,225 cases were reported between 1999 and 2004. The occurrence of malaria in high risk areas of South Korea was significantly (P > 0.05) correlated with the mosquito population but not with temperature and rainfall. Control programs, including early case detection and treatment, mass chemoprophylaxis of soldiers, and international financial aids to North Korea for malaria control have been instituted. The situation of the reemerging vivax malaria in the Republic of Korea is remarkably improving during the recent years, at least in part, due to the control activities undertaken in South and North Korea.