A practical method for predicting the short-time trend of bivoltine populations of Ips typographus (L.) (Col., Scolytidae)

Abstract: Ips typographus is the main spruce pest of European forests. In most areas of the Italian Alps there are two generations per year; overwintering adults fly in May looking for trees suitable for breeding, their offspring emerge in summer, 7–8 weeks after tree colonization, and the adults of the second generation emerge in spring of the following year after overwintering under the bark or in the litter. A long‐term population monitoring was carried out in north‐east Italy with the aim at developing a prediction model able to estimate the population density of the following year. Between 1996 and 2004, pheromone traps monitored populations of I. typographus annually. Monitoring lasted 4 months (May–August), with replacement of pheromone dispensers after 8 weeks. Insects trapped before dispenser change were called ‘spring captures’ (May–June), and included both overwintering and re‐emerging adults. Beetles caught after dispenser change were called ‘summer captures’ (July–August), and included the adults of the first generation. The results show a high positive correlation between the ratio of summer and spring captures of one year (Summer_x/Spring_x), and the ratio of total captures of the following year (Y_x+1) and those of the current year (Y_x) (Y_x+1/Y_x). Summer_x/Spring_x lower than 0.62 indicate decreasing populations in the following year (Y_x+1/Y_x <1), whereas Summer_x/Spring_x higher than 0.62 indicate increasing populations (Y_x+1/Y_x >1). The applicability of the model in the study of I. typographus risk of outbreak and in the forest management is discussed. The prediction of the short‐time trend of the population allows assessing its density in the following year, and therefore the risk of outbreak.     

Influence of trap type, trap colour, and trapping location on the capture of the pine moth, Thaumetopoea pityocampa

Studies were conducted to evaluate the effect of type, location, and colour of traps baited with 1 mg of the sex pheromone (Z)‐13‐hexadecen‐11‐ynyl acetate in polyethylene vials on the capture of the pine processionary moth, Thaumetopoea pityocampa (Denis & Schiffermüller) (Lepidoptera: Thaumetopoeidae), males. The experiments were carried out during two flight seasons in 2002 and 2003 at a low elevation Mediterranean pine forest on the hill of Goritsa (Magnesia, Thessaly, Central Greece). The hill is covered by approximately 120 ha of pines, among which Pinus brutia Ten. (Pinaceae) is predominant with Pinus halepensis Mill. (Pinaceae) as second most common species. Among the commercially available trap types used, the Delta and Pherocon II captured significantly more males than the Funnel trap. Furthermore, pine density had a significant effect on trap type and trap colour catches. Delta traps caught significantly more adults than the other two trap types in the low density pine stand, while in the medium and high‐density pine stands no significant differences were noted between Delta and Pherocon II traps. In addition, trap colour performance varied according to pine density; white‐ and yellow‐coloured traps caught significantly more males than the other two trap colours in the high‐density stand. The first adults were captured in traps during May, and traps continued to capture adults in low numbers (< two individuals per trap) until mid‐June. A period of 3–4 weeks of no trap captures followed until mid‐July, when captures restarted, at low numbers. The peak of T. pityocampa flight was observed during late August–September.     

An ordinal day model of spruce beetle trap capture phenology in northern Colorado

Spruce beetle populations (Dendroctonus rufipennis) (Coleoptera: Scolytinae) in Engelmann spruce (Picea engelmannii) stands were monitored at twelve sites for 6 years in northern Colorado using pheromone‐baited Lindgren funnel traps. During the trapping period, over 30,000 beetles were captured, and beetle abundance data were used to construct empirical models of trap capture phenology based on day of year, accumulation of thermal units and thermal day thresholds, with the goal of informing future regional trapping efforts and producing a simple model for use by forest managers. Ordinal day models outperformed thermal accumulation and thermal threshold models in both predictive power and parsimony. Mean date of earliest capture was Day 153 (June 2) and ranged from Day 126 to Day 161 (May 6–June 10), and mean date of final capture was Day 243 (August 31) and ranged from Day 220 to Day 286 (August 8–October 13). A two‐parameter logistic function was the most parsimonious of several ordinal day models, accounting for 81% of the variance in cumulative trap capture across all sites and dates. The model predicts 50% of trap captures to occur by Day 181 (June 30). This model has application as a decision support tool for forest ecosystem managers concerned with the timing of trap deployment or D. rufipennis mitigation treatments.     

Incidental capture of American Crows in coyote‐sized box traps

ABSTRACT American Crows (Corvus brachyrhynchos) are difficult to capture. I used a simple technique (box traps) to capture 151 crows incidental to my eastern coyote (Canis latrans) research at two study sites in eastern Massachusetts. Crows were captured at a higher rate during the spring (winter = 33 captures, spring = 70, summer = 22, and fall = 26). Among months, capture efficiency ranged from 0.7% in August to 8.5% in May, with an overall average of 4.1 crows captured per 100 trap days. Although other capture methods (e.g., net launchers and Australian crow traps) have been used to capture crows in greater numbers, box traps can be an important complement to the other capture techniques because they are transportable, easy to use, relatively cheap, and easy to obtain.     

Different seasonal feeding activity of diverse herbivores in two temperate forests

The aim of the present study was to understand the effects of abiotic conditions on seasonal feeding activity of diverse herbivores on the same oak tree species in two different forests. We tracked changes in herbivore feeding activities on an oak tree species (Quercus serrata) in two localities: a low elevation small hillock forest patch (Muan, MN) and a middle elevation mountain forest patch (Mt. Jirisan, JR). A total of five sites were selected in each of two forest localities. Data for leaf expansion, leaf chemical qualities, leaf damage ratio, and numbers of lepidopteran caterpillars were collected during spring (May) and summer (July to August), 2012. Leaf expansion rate was higher at the low hillock forest than the mid‐mountain forest from spring to summer. Nitrogen and carbon content decreased seasonally at both localities. Lepidopteran larval diversity was high in the mid‐mountain forest, and two‐way ANOVA showed that species richness of lepidopteran larvae was significantly affected by the interaction between season and locality. Leaf damage by all herbivores was higher in the low hillock forest than the mid‐mountain forest in spring, but was higher in the mid‐mountain forest in summer. Relative proportion of general herbivores increased from spring to summer in the mid‐mountain forest, but not in the low hillock forest. Canonical Correspondence Analysis (CCA) ordination showed that altitude‐ and season‐related variables were significant species and environment interaction factors. Our data indicate that locality and temperature disproportionally affected the feeding activities of diverse herbivores in two different temperate forests.     

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

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Seasonal activity patterns of Ixodes pacificus nymphs in relation to climatic conditions

Abstract In western North America, the tick Ixodes pacificus Cooley & Kohls (Acari: Ixodidae) is the primary vector to humans and domestic animals of the disease agents causing Lyme disease and granulocytic ehrlichiosis. We examined the seasonal activity patterns of I. pacificus nymphs over a 4‐year period, including the wet and cold El Niño winter/spring of 1998, in a dry oak/madrone woodland, and for one year in a cooler and moister redwood/tanoak woodland in Mendocino County, California. Linear regressions were used to estimate when nymphal densities first exceeded and then fell below 25, 50 and 75% of the recorded yearly peak densities. In oak/madrone woodland, nymphs typically were active by mid‐March, reached 50% of their yearly peak densities in early to mid‐April, peaked by early May, fell below 50% of their peak densities by early to mid‐June, and were absent by late July to mid‐August. The lengths of the periods with nymphal densities exceeding 50 and 75% of the recorded yearly peaks in oak/madrone woodland were associated positively with rainfall and negatively with maximum air temperatures during April–May. Moreover, nymphal numbers typically reached 50% of their peak 10–15 days later, remained at levels above 50% of the peak 1.3–1.5 times longer, and started declining 4–6 weeks later under cooler, moister climatic conditions (oak/madrone woodland in 1998 and redwood/tanoak woodland in 2000) relative to warmer, drier conditions (oak/madrone woodland in 2000–2001). In oak/madrone woodland, nymphal densities typically started to decline when mean maximum daily air temperatures exceeded 23°C. Nymphal densities were higher in dry oak/madrone relative to moist redwood/tanoak woodland from mid‐March to late May 2000, similar in both habitat types in early June, but higher in redwood/tanoak woodland from late June onwards. We conclude that large‐scale studies of the density of I. pacificus nymphs in California need to consider spatial variation in the length of nymphal activity periods and select temporal sampling regimens that yield representative data for all included habitat types.     

Characterizing spring emergence of adult Halyomorpha halys using experimental overwintering shelters and commercial pheromone traps

To improve our understanding of adult Halyomorpha halys (Stål) (Hemiptera: Pentatomidae) overwintering biology and to better inform models of its population dynamics, its temporal pattern of spring emergence was investigated using experimental overwintering shelters in screened cages within protective structures. In 2012, plastic shelters containing 100 adults were deployed in unheated, unlighted buildings, and adjacent woodlots in Virginia, USA. In 2013 and 2014, wooden shelters containing 300 paint‐marked adults were deployed in pairs in six woodlots across Virginia, West Virginia, and Maryland, USA; one in a closed cage and one in a cage with the top removed, enabling emerged adults to be counted or to disperse, respectively. In 2013 and 2014, pheromone‐baited and non‐baited pyramid traps encircled the shelters at each site. Regular counts of adults that emerged into the closed cage and of marked and ‘wild’ (unmarked) adults captured in traps were conducted from February or March through early July. In 2012, emergence patterns from shelters in buildings and woodlots were very similar and matched those recorded from woodlots in 2013 and 2014. In all years, a small peak of emergence occurred in about mid‐April, a larger and more prolonged peak was observed between mid‐May and early June, and emergence ended by early July. Of the 449 H. halys adults captured in traps between 2013 and 2014, only three were marked individuals from shelters in the open cage, suggesting that adults emerging from overwintering sites may require a dispersal flight before responding to pheromone‐baited traps. In 2013 and 2014, respectively, 98 and 93% of captures were in pheromone‐baited traps, but there was no correlation between the weekly number of adults that emerged from shelters in the closed cages and captures in traps.     

Factors contributing to coexistence of Chaoborus flavicans and C. punctipennis (Diptera,Chaoboridae) in a small meromictic lake

In Crawford Lake, a small meromictic water body in southern Ontario, Canada, the life cycles of planktonic Chaoborus flavicans and C. punctipennis were out of phase by about two months, the former pupating from mid May through mid June and the latter from July through September. C. flavicans possibly produced a second annual generation. Fourth instars of both species were strong diel migrators and occupied similar strata at most times. C. punctipennis fourth instar diet consisted almost entirely of rotifers. C. flavicans ate rotifers but fed heavily also on daphnids in May and August. C. punctipennis fourth instars showed little growth until early spring, possibly owing to a sparseness of rotifers, then grew rapidly until pupation. C. flavicans had a slow, but relatively constant growth rate at all times during the open water season, presumably because its greater mouth gape allowed it a wider range of food items.     

REPRODUCTIVE PHENOLOGY OF LAMINARIA SACCHARINA (L.) LAMOUR. (PHAEOPHYTA) AT THE SOUTHERN LIMIT OF ITS DISTRIBUTION IN THE NOTHWESTERN ATLANTIC OCEAN1,2

Laminaria saccharina (L) Lamour. Sporophytes were monitored monthly from October 1982 to September 1983 to investigate reproduction phenology and relationshiops to growth paatterns aaat its southern limit of distribution in the northwest Atlantic Ocean (Long Island Soundd). Plants exhibited an annual growth pattern. Growth raate, bladelehgth, maximum width, area, stiipe lehgth an wet weight swhoed the same seasonal pattern and reached maximum values between May and June. Only blade thickness continued to increase tthrooughout the ovservation period. Blade dissintegration occurred dduring August and September. Reproductive sporophytes occurred throughout the observation period; the greatest frequency of appearance occurred in October (43.8%) and June (37.8%). The blade area covered by sori ranged from 2.4% (Janaury) to 6.1% (August). Meiospore release under laboratory conditions was maximum in May and minimum in July. No meio-spores were released in August Sporulation was not correlated with meristematic growth of nitrogen content How ever, reproductive plants were generally larger and thicker throughout the sporulation period, and had a greater carbort content is spring than nonreproductive plants. Fecundity and reproductive success of female gametophytes were maximum in spring and minimum in winter. The growth of early sporophyte stages in the laboratory was greatest in early spring; however, juvenile macroscopic stages were hardly observed in the field during summer months due to warm water temperatures. “Over-summering” of gamelophytes and / or microscopic sporophytes may account for the annual cycle of Laminaria at its southern limit of distribution.     

Seasonal prevalence of the Culex pipiens group (Diptera: Culicidae) larvae occurring in a puddle in the basement of an apartment building

The seasonal prevalence of the Culex pipiens group larvae occurring in a puddle in the basement of an apartment building was investigated from June 1993 to May 1994 (Period I), and from June 1994 to May 1995 (Period II). Totals of 25 237 individuals in Period I and 4989 individuals in Period II were collected: they were all C. pipiens group larvae, most of which were C. pipiens molestus. In Period I, the larvae occurred at a relatively low density in the hot months July and August; density began to increase from the cooler late September, peaked in late November, and was maintained at a high level until the third week of December. In Period II, in which mean air temperature was 5°C higher as of July and August than in Period I, larvae sharply decreased in number by one‐fifth of the total volume in Period I, and showed a peak density in the third week of July, and the peak continued until mid‐December. In Period I, the low temperature was favorable for oviposition activity of C. pipiens molestus, and accounts for the higher density of larvae in Period I than in Period II. C. pipiens molestus mostly stopped laying eggs between early December and late January. Larvae were not collected in March and April 1994 and in April 1995. However, considering that adult mosquitoes had been collected throughout the two periods except in March 1994, adults of C. pipiens molestus are thought to appear throughout the year.     

Population dynamics in changing environments: the case of an eruptive forest pest species

In recent decades we have seen rapid and co‐occurring changes in landscape structure, species distributions and even climate as consequences of human activity. Such changes affect the dynamics of the interaction between major forest pest species, such as bark beetles (Coleoptera: Curculionidae, Scolytinae), and their host trees. Normally breeding mostly in broken or severely stressed spruce; at high population densities some bark beetle species can colonise and kill healthy trees on scales ranging from single trees in a stand to multi‐annual landscape‐wide outbreaks. In Eurasia, the largest outbreaks are caused by the spruce bark beetle, Ips typographus (Linnaeus), which is common and shares a wide distribution with its main host, Norway spruce (Picea abies Karst.). A large literature is now available, from which this review aims to synthesize research relevant for the population dynamics of I. typographus and co‐occurring species under changing conditions. We find that spruce bark beetle population dynamics tend to be metastable, but that mixed‐species and age‐heterogeneous forests with good site‐matching tend to be less susceptible to large‐scale outbreaks. While large accumulations of logs should be removed and/or debarked before the next swarming period, intensive removal of all coarse dead wood may be counterproductive, as it reduces the diversity of predators that in some areas may play a role in keeping I. typographus populations below the outbreak threshold, and sanitary logging frequently causes edge effects and root damage, reducing the resistance of remaining trees. It is very hard to predict the outcome of interspecific interactions due to invading beetle species or I. typographus establishing outside its current range, as they can be of varying sign and strength and may fluctuate depending on environmental factors and population phase. Most research indicates that beetle outbreaks will increase in frequency and magnitude as temperature, wind speed and precipitation variability increases, and that mitigating forestry practices should be adopted as soon as possible considering the time lags involved.     

Development and biological characteristics of <Emphasis Type="Italic">Haemaphysalis longicornis</Emphasis> (Acari: Ixodidae) under field conditions

The development and biological characteristics of Haemaphysalis longicornis were investigated under field conditions in Xiaowutai National Natural Reserve Area, North China. Unfed larvae, nymphs and adults were fed on rabbits and exposed to daylight. Three free-living stages were allowed to develop in field plot selected in a tick natural habitat. The host seeking behavior and seasonal occurrence were observed. Haemaphysalis longicornis were active from mid March to mid October. The premoult period of nymphs and preoviposition of females were regulating phases of their life cycle. The developmental durations of eggs, larvae and adults were constant under field conditions regardless when the development started. The oviposition periods in May and June were statistically shorter than those in July and August. The daily oviposition patterns of females engorged in May and June demonstrated unobvious peak, which differed from those engorged in July and August. The daily oviposition peak of the latter occurred on the 4th day of oviposition. Moreover, a positive correlation was found between the mass of the laid egg and the body weight of engorged females (r = 0.62, P < 0.001). The female reproductive efficiency indices were 2.9, 6.1, 10.5 and 9.0 in May, June, July and August, respectively. The mean weight (3.33 mg) of engorged nymphs molting to females was significantly higher than that (2.35 mg) of those molting to males (P < 0.001), but the body weights of both sexes were overlapping.     

Seasonal dynamics in community structure,abundance, body size and sex ratio in two species of Neotropical annual fishes

Seven ephemeral pools on the coastal plain of southern Brazil were found to be inhabited by three annual and 22 non‐annual fish species. Two common annual species (Austrolebias minuano and Cynopoecilus fulgens) exhibited clear seasonal dynamics, with the appearance of young fishes in the austral autumn (May to June) and a decline in abundance over the seasonal cycle. The third annual species, Austrolebias wolterstorffii, was rare. No seasonal dynamics were observed in non‐annual fishes. The relative abundance of non‐annual fishes compared with annual fishes increased over the seasonal cycle, but they coexisted widely. The size structure of annual fishes suggested the presence of a single age cohort in most pools though a second age cohort was registered in one pool in August, coinciding with a large flooding. Strong sexual dimorphism in body size was found in C. fulgens throughout the seasonal cycle, while no sexual dimorphism in body size was found in A. minuano. Female‐biased sex ratios were recorded in both common annual fish species in the last three sampling dates (in spring), but not during the first two sampling dates (in winter). The natural lifespan of annual fishes was <8 months. Annual fishes disappeared before habitat desiccation in half of the pools, while non‐annual fishes were still present.     

Monitoring of brown stink bug (Hemiptera: Pentatomidae) population dynamics in corn to predict its abundance using weather data

The brown stink bug (BSB), Euschistus servus (Say)(Hemiptera: Pentatomidae), is a serious economic pest of corn production in the southeastern United States. The BSB population dynamics was mon itored for 17 weeks from tasseling to preharvest of corn plants (i.e., late May to mid-September) using pheromone traps in three corn fields from 2005 to 2009. The trap data showed two peaks in early June and mid-August, respectively. The relationship between trap catch and pregrowing season weather data was examined using correlation and stepwise multiple factor regression analyses. Weather indices used for the analyses were accumulated growing degree day (AGDD), number of days with minimum temperature below 0℃(Subz), accumulated daily maximum (AMaxT) and minimum temperatures (AMinT) and rainfall (ARain). The weather indices were calculated with lower (10℃) and upper (35℃) as biological thresholds. The parameters used in regression analysis were seasonal abundance (or overall mean of BSB adult catch)(BSBm), number of BSB adults caught at a peak (PeakBSB), and peak week (Peakwk). The BSBm was negatively related to high temperature (AmaxT or AGDD) consistently, whereas IstPeakBSB was positively correlated to both ARain and Subz, irrespective of weather data durations (the first 4, 4.5, and 5 months). In contrast, the 7-month weather data (AGDD7) were negatively correlated to the BSBm only, but not correlated to the second PeakBSB. The 5-year monitoring study demonstrated that weather data can be used to predict the BSB abundance at its first peak in tasseling corn fields in the southeastern U.S. states.     

Prolonged development time of the bark beetle predator Thanasimus formicarius (Col.: Cleridae) in relation to its prey species Tomicus piniperda (L.) and Ips typographus (L.) (Col.: Scolytidae)

1 The generation time of the bark beetle predator Thanasimus formicarius (L.) (Col.: Cleridae) was found to be predominantly two years both in the field and in rearing experiments conducted with two of its main prey species, the pine shoot beetle Tomicus piniperda (L.) and the spruce bark beetle Ips typographus (L.) (Col.: Scolytidae). 2 Emergence of T. formicarius adults in the first summer was only observed in one of the two rearing experiments, and these individuals represented only 6% of that generation. 3 All individuals not emerging as adults in the first summer remained as larvae in their pupal chambers until the second summer. Pupae were found starting around mid-June, and adults (in pupal chambers) were found from late July through to the end of August. 4 Newly emerged adults had to feed in order to survive hibernation. 5 The existence of T. formicarius races, specialized on certain bark beetle species and with phenologies matching their hosts, could not be demonstrated. After hibernation there was no difference in feeding activity, timing of egg-laying or proportion of egg-laying females between the T. formicarius adults reared as larvae on T. piniperda (flight period in April) and those reared as larvae on I. typographus (main flight period generally starting in late May or early June) when exposed to a temperature and day-length typical of the early spring conditions prevailing during the flight period of T. piniperda. 6 T. formicarius was parasitized by Enclisis vindex (Tschek) (Hym.: Ichneumonidae) in the pupal chamber. 7 The importance of these findings for the population dynamics of bark beetles is discussed.     

The bimodal adult activity of Monochamus alternatus (Coleoptera: Cerambycidae) caught in pheromone traps in Jeju can be explained by the competitive attractiveness of dying pine trees

The Japanese pine sawyer, Monochamus alternatus Hope (Coleoptera: Cerambycidae), is known to be the primary vector of pinewood nematode Bursaphelenchus xylophilus (Steiner & Buhrer) Nickle that causes pine wilt disease. Adult activity of M. alternatus caught in a pheromone trap on Jeju Island of Korea showed a bimodal form with the first peak in mid to late June and the second peak in mid to late September. The two peaks were separated between mid and late August, showing a valley. Accumulated degree‐days predicted that the emergence of the second generation adults could be possible just before the second peak. But actually no adults of the second generation occurred in the field development experiments of M. alternatus in 2016 and 2017. Pine trees without oleoresin flow (namely dying trees by the infection of pinewood nematode) were abundant during early July to early August. The bimodal adult activity pattern of M. alternatus could be partially explained by the competitive attractiveness of dying trees against pheromone traps, when we accepted the assumption that dying pine trees attract strongly M. alternatus.     

Interruption of aggregation pheromone in Ips typographus (L.) (Col. Scolytidae) by non-host bark volatiles

Abstract 1 The antennally active nonhost bark volatiles (NHVs): trans‐conophthorin (tC), C₆‐alcohols (green leaf volatiles; GLVs) and C₈‐alcohols, were tested for their ability to reduce attraction of the spruce bark beetle Ips typographus (L) (Col. Scolytidae) to its pheromone sources in both laboratory walking bioassy and field trapping experiments. 2 In the walking bioassay with I. typographus females, individual NHVs such as tC, 3‐octanol and 1‐octen‐3‐ol, and the unsuitable host signal, verbenone (Vn), were inactive at the doses tested. However, the blend of C₆‐alcohols (3GLVs) and all the binary, ternary, or quarternary blends significantly reduced the female attraction to the pheromone sources. 3 In the field trapping experiments, individual NHV signals (tC, C6‐alcohols and C₈‐alcohols) all reduced catch of I. typographus in pheromone‐baited traps, with their inhibitory effects similar to that of the known inhibitor, Vn. The binary, ternary or quarternary combinations of these NHV signals or Vn, all caused significantly stronger reductions in trap catches than the individual signals. The blends showed similar levels of interruption, except the binary blend of C₈‐alcohols (2C8OH) and Vn. 4 Difference in trapping mechanism between pipe traps (attraction and landing) and Lindgren funnel traps (attraction) did not affect the pattern of inhibition of these active NHV signals and Vn. 5 These behaviourally active nonhost volatiles and Vn might be used effectively to protect spruce trees or stands against attacks by I. typographus.     

Effects of decadal climate change on zooplankton over the last 50 years in the western subarctic North Pacific

Decadal‐ to multi‐decadal variations have been reported in many regional ecosystems in the North Pacific, resulting in an increasing demand to elucidate the link between long‐term climatic forcing and marine ecosystems. We detected phenological and quantitative changes in the copepod community in response to the decadal climatic variation in the western subarctic North Pacific by analyzing the extensive zooplankton collection taken since the 1950s, the Odate Collection. Copepod species were classified into five seasonal groups depending on the timing of the annual peak in abundance. The abundance of the spring community gradually increased for the period 1960–2002. The spring–summer community also showed an increasing trend in May, but a decadal oscillation pattern of quasi‐30‐year cycles in July. Phenological changes coincided with the climate regime shift in the mid‐1970s, indicated by the Pacific decadal oscillation index (PDO). After the regime shift, the timing of the peak abundance was delayed one month, from March–April to April–May, in the spring community, whereas it peaked earlier, from June–July to May–June, in the spring–summer community, resulting in an overlap of the high productivity period for the two communities in May. Wintertime cooling, followed by rapid summertime warming, was considered to be responsible for delayed initiation and early termination of the productive season after the mid‐1970s. Another phenological shift, quite different from the previous decade, was observed in the mid‐1990s, when warm winters followed by cool summers lengthened the productive season. The results suggest that climatic forcing with different decadal cycles may operate independently during winter–spring and spring–summer to create seasonal and interannual variations in hydrographic conditions; thus, combinations of these seasonal processes may determine the annual biological productivity.     

Escape in space from enemies: a comparison between stands with and without enhanced densities of the spruce bark beetle

1 Populations of the spruce bark beetle, Ips typographus (L.), are known to grow rapidly in storm‐disturbed stands as a result of relaxation from intraspecific competition. In the present study, it was tested whether a second mechanism, escape in space from natural enemies, also contributes to the rapid population increase. 2 The experiment was conducted during the initiation phase of five local outbreaks of I. typographus triggered by a storm‐disturbance in November 1995 in southern Sweden. 3 The impact of natural enemies on the ratio of increase (number of daughters per mother) of I. typographus was compared pairwise between disturbed stands with high numbers of storm‐felled trees and undisturbed stands without wind‐felled trees. 4 Enemy impact was assessed by comparing the ratio of increase in uncaged (exposed to enemies) and caged (protected from enemies) bolts colonized by I. typographus prior to being placed in the stands. The experiment was conducted in the second and third summers after the storm‐felling. 5 Enemy impact was about twice as high in stands without wind‐felled trees compared with in stands with wind‐felled trees in the second summer whereas there was no significant difference between the stand types in the third summer. 6 The result demonstrates that spatial escape from enemies contributes to the rapid population growth of I. typographus after storm‐disturbances.