The effect of spruce cone insects on seed production in Switzerland

Abstract: In 1989 and 1990, spruce cones were harvested at 29 sites distributed over the five main geographic regions of Switzerland, i.e. Jura, Central Plateau, North, Central and South Alps. The presence of insects and the species-specific consumption of seeds was determined for each cone by examination of the median longitudinal section. Seven seed-feeding species ( Assara terebrella , Cydia strobilella , Dioryctria abietella , Eupithecia abietaria , Megastigmus strobilobius , Plemeliella abietina , Strobilomyia anthracina ) and the spruce cone gall midge Kaltenbachiola strobi were found in the samples. The proportions of infested cones ranged from 36 to 100%. The regions did not show significant differences in terms of infestation rates, whereas differences between sites within the regions were highly significant. Seed loss ranged from 2 to 100%, its geographical distribution exhibiting a similar pattern to the infestation rates. Considering the number of infested cones, C. strobilella was the most abundant species, followed by K. strobi and P. abietina . Cones infested by conospermatophagous species ( C. strobilella , D. abietella , E. abietaria ) showed higher seed losses than those infested by spermatophages ( M. strobilobius , P. abietina ). Due to the ubiquitous occurrence of C. strobilella , its high infestation rates and the resulting seed losses caused by its offspring, this species has to be considered as the most important insect seed predator in Switzerland. A multiple linear regression analysis was carried out for each region to simultaneously explore the impact of site and tree variables as well as the effect of insect attack on the yield of viable seeds. The results indicate that site and tree conditions contributed more to the variability of seed yield than insect presence in a cone. The model predicted significant effects of insect attack mainly for cones with total seed numbers that were higher than the average.     

A survey of insects attacking seed cones of Pinus cembra in the Alps, the Pyrénées and Massif Central

The composition and impact of the entomofauna exploiting seed cones of Swiss stone pine ( Pinus cembra L.) was surveyed in native stands and plantations of France, Italy, Switzerland and Austria during 1992–1996. A total of 2785 cones were collected from 25 sites. The cone entomofauna consisted of five species, of which three ( Dioryctria abietella , Eupithecia abietaria , Cecidomyia pini ) were regularly observed. The two other insects, Polydrusus atomarius and Zeiraphera diniana were observed on first-year cones, and only in two stands. No spermatophagous insect was detected. None of these insect species could be considered specific to cones of P. cembra. The overall damage never exceeded 40% of the cone crop. The importance of cone damage varied with year, cone abundance, and the position of the tree in the stand. The cone entomofauna of Swiss stone pine largely deviated from that recorded in cones of other high altitude conifers but resembled that of other stone pines growing in Eurasia. The question: why cones of P. cembra were less damaged by insects than those of other alpine conifers is discussed.     

Potential for mating disruption to reduce cone damage by the spruce seed moth, Cydia strobilella, in spruce seed orchards

Abstract:  The spruce seed moth, Cydia strobilella (L.), is a serious and widely distributed pest of spruce seed orchards in North America and Europe. Current pest management activities in seed orchards rely mainly on chemical pesticides for insect control. Mating disruption with sex pheromone is a potential alternative pest management tool for C. strobilella . In 2002, field tests confirmed that sticky traps baited with 3  μ g of (E) -8-dodencenyl acetate ( E 8-12:Ac), the sex pheromone of C. strobilella , could capture males in Quebec, a region of Canada not previously monitored for this insect. In the following years (2003–2005), grey rubber septa loaded with 0.75, 1.5, or 2.0 mg of E 8-12:Ac were deployed separately in two white spruce seed orchards at a density of 40 and 60 dispensers/ha to test the potential for mating disruption. The results showed that the captures of male C. strobilella in the pheromone-treated plots were reduced by up to 98%. Furthermore, at the end of the experiment in 2005, 17.3% of cones were damaged by C. strobilella in the treated plot, compared with a significantly (P < 0.0001) higher 56.4% in the control plot. The results suggest that mating disruption has potential for controlling C. strobilella to protect seed cones in white spruce seed orchards.     

Bacillus thuringiensis variety kurstaki x aizawai applied to spruce flowers reduced Dioryctria abietella (Lepidoptera: Pyralidae) infestation without affecting seed quality

We investigated the effects of Bacillus thuringiensis variety kurstaki x aizawai (Bt) on infestation levels of two lepidopteran insects as well as on seed quality in Norway spruce, Picea abies L. (Karst.) in central Sweden. Spruce flowers (female strobili) were sprayed with a 0.2% suspension (wt:wt) of the Bt preparation Turex 50 WP, 25,000 IU/mg in water. To expose even those lepidopteran larvae that feed exclusively embedded within the cone tissue, the Bt treatment was applied to open flowers, before they closed and developed into cones. The experimental design included three main factors: treatment (untreated control, water, or Bt), spruce genotype (three clones), and spraying time (spraying before, during, and after the phase of highest pollen receptivity). The Bt treatment reduced the proportion of cones infested by the cone worm Dioryctria abietella Den. et Schiff. (Lepidoptera: Pyralidae) from approximately 30 to 15%. There was no statistically significant treatment effect on the infestation rate of Cydia strobilella (L.) (Lepidoptera: Tortricidae). The Bt variety kurstaki x aizawai treatment caused no reduction in seed quality as measured by seed weight or percentage of nonfilled seeds. There was no difference in number of seeds per cone between the Bt-treated and untreated control cones. There was a significant effect of genotype on insect infestation rates, as well as on number of seeds per cone and seed weight. Neither level of insect damage nor any seed quality parameters were affected by time of application of the treatments.     

在韩国危害红松球果的三种蛾类幼虫形态记述（鳞翅目）

详细地记述了韩国红松球果害虫冷杉梢斑螟Dioryctria abietella、赤松梢斑螟D. sylvestrella (螟蛾科)和小花尺蛾Eupithecia abietaria debrunneata (尺蛾科)幼虫的形态特征,并提供了形态特征图。     

Effects of different insect species on seed quantity and quality in Norway spruce

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Measuring the impact of Leptoglossus occidentalis (Heteroptera: Coreidae) on seed production in lodgepole pine using an antibody-based assay

We measured the impact of Leptoglossus occidentalis on seed production in lodgepole pine, Pinus contorta variety latifolia Engelmann, using an antibody marker developed to detect residual saliva in fed-on seeds. Nymphs, adult females, and adult males were caged on cones during early, mid- and late season cone development. Individual analysis of 12,887 seeds extracted from 365 cones revealed that 37.3% seeds tested positive for seed bug saliva. The antibody assay was 38 times more effective than radiography at detecting seed bug damage. Radiography can detect partially emptied seed but cannot discriminate between aborted seeds and those emptied by seed bugs. The antibody marker was least sensitive in detecting early season damage compared with mid- and late season damage. We hypothesize that residual saliva in seeds fed on early in the season was either absorbed by the damaged seed or degraded over time. Early season feeding resulted in the greatest number of seeds fused to cone scales and the extraction efficiency for cones exposed to feeding during this time was reduced by 64% compared with control cones. Adding fused seeds to antibody-positive seeds raised the proportion of damaged seeds to 48.3%. At all stages of cone development, adult females were the most destructive life stage, damaging up to two seeds per day late in the season. When seed losses were adjusted to damage per degree-day, female damage was greatest early in the season, while males caused the same amount of damage regardless of cone development period. The results of the antibody assay provide baseline data for developing damage prediction formulae, and establish L. occidentalis as a potentially serious pest in lodgepole pine seed orchards.     

韩国波尺蛾亚科三种幼虫记述（鳞翅目：尺蛾科）

本文详细地记述了波尺蛾亚科Eupithecia abietaria debrunneata Staudinger, E. spadix Inoue 及Telenomeuta punctimarginaria (Leech) 幼虫的形态特征.     

Insect pests of Norway spruce cones: incidences and altitude preferences with emphasis on Cydia strobilella (L.)

Pest species infesting spruce cones were identified from 109 locations approved for the collection of seed material in the Czech Republic. Four (occasionally five) cones were studied from each location, and 448 cones were examined in total. Each cone was assessed for external damage and was then cut open along the spine. The predominant pest species, Cydia strobilella, was detected in 65% of the cones, while the next most common species, Dioryctria abietella, was detected in 13% of the cones. Ernobius abietis, Eupithecia sp., and Thekopsora areolata were less numerous. Kaltenbachiola strobi and Megastigmus sp. were recorded sporadically. Most cones (68%) were infested with only one species. The maximum number of C. strobilella larvae detected in one cone was 13. Cone damage resulting from C. strobilella declined markedly with increasing elevation. Thus, cone infestation by C. strobilella was greater in those forest vegetation zones located in lowlands and uplands rather than in highlands and at lower mountain elevations. The number of cones without damage tended to increase with elevation, but infestation by D. abietella was unrelated to elevation. In some cases, C. strobilella damage to cones was severe. External markings on the cone (resin and deformation) cannot by themselves serve as reliable criteria for predicting C. strobilella infestation, although cones with such signs should be preferentially assessed when seed material is inspected.     

Performance of the fir coneworm Dioryctria abietivorella (Grote) as affected by host species and presence or absence of seed cones

1 Larval performance of Dioryctria abietivorella (Grote) (Lepidoptera: Pyralidae) was compared when insects were reared in white spruce, black spruce and Jack pine seed orchards. For each species, half of the insects developed in the presence of cones, while the other half was maintained on branches without cones. 2 Significantly faster development rates were observed on spruce species when compared to Jack pine. Significant differences in survival were also recorded between the different feeding treatments. 3 The presence of seed cones on spruce species significantly increased insect mean weight compared with branches without cones, but no significant differences were noticed between the different tree species with cones. 4 White spruce is the most suitable host tree for fir coneworm feeding and the availability of seed cones plays an important role in determining D. abietivorella larval performance.     

Cono- and seminiphagous insects of Norway spruce Picea abies (L.) Karst. and their parasitoids in lower and upper montane zone of the Tatra National Park in Poland

Abstract: A study of the insects inhabiting of cones of Norway spruce Picea abies (L.) Karst. in the Tatra National Park was conducted during 1987–92. As a result of breeding and analyses of the cones collected from trees at various altitudes above sea level, 53 746 specimens of cono- and seminiphagous insects were collected. Interior damage to cones and the losses of spruce seeds caused by pests were measured, with regard both to the variable cone crop abundance of spruce and to different altitudes above sea level. The role of parasitoids in limiting the numbers of some cono- and seminiphagous insects was also estimated.     

Impact of Leptoglossus occidentalis (Hemiptera: Coreidae) on Douglas-fir seed production

We investigated the effect of feeding by the western conifer-seed bug, Leptoglossus occidentalis Heidemann, on seed production in developing cones of coastal Douglas-fir, Pseudostuga menziesii (Mirb.) Franco, with respect to seed bug life stage and sex (nymphs, adult females, and adult males) and timing of feeding (early, mid-, and late season cone development). Feeding by females on caged cones for a 2-wk period during late season cone development reduced the proportion of full seeds in cones by approximately 70% compared with caged control cones. There was no significant difference among nymphs, adult females, and adult males with respect to the proportion of empty or partially fed-upon seeds produced during the same feeding period. Feeding by nymphs for 2 wk early in the season resulted in a threefold increase in the number of unextractable seeds fused to cones compared with the control. Weight measurements of harvested seeds indicated that radiography is an accurate tool to distinguish among Douglas-fir seeds that have sustained light, moderate, or severe damage. Determining the full impact of L. occidentalis on conifer seed production will require the development of a reliable method to distinguish between naturally aborted seeds and seeds emptied through feeding by seed bugs.     

Modelling irruptions and population dynamics of the great spotted woodpecker – joint effects of density and cone crops

Irruptive migrants are partially migrating species showing pronounced temporal variation in the number of migrants. Occasional irruptions are often explained by increased population density (per area) or lack of food. Similarly to population dynamics, these explanations are not mutually exclusive, but probably act in concert. Here we do a parallel analysis on the migration intensity and population dynamics of an irruptive migratory bird species, the great spotted woodpecker Dendrocopos major, in southern Finland. Both analyses include simultaneous effects of breeding density and cone crops of Scots pine Pinus sylvestris and Norway spruce Picea abies. A novel model for statistical analyses of irruptive migration intensity is developed. Population dynamics are investigated using log‐linear state‐space models. Woodpecker migration intensity is well explained by lack of spruce cones and increases proportionally to population density. Population dynamics is most successfully described with a density dependent model, where the natural logarithm of spruce cone availability previous autumn has a strong positive effect on population growth. This is likely to be due to lowered winter mortality and lowered emigration rates resulting from plentiful food availability. Although the species uses pine cones as a food resource in winter, the estimated impacts of pine cone crops on migration intensity and population dynamics are low, probably due to low annual variation in crop comparing to spruce cones. Large irruptions also tend to occur earlier in season, often before cones are a crucial food resource for the woodpeckers, suggesting that the woodpeckers use the amount of maturing cones as a cue for upcoming conditions. Our results show that the ecological role of different resources can differ considerably from the expected pattern based on the proportion of the resources in a species’ diet. Advantages of examining patterns of bird migration, population density and dynamics in unison are apparent.     

Evolution of seed dispersal in North American <Emphasis Type="Italic">Ephedra</Emphasis>

Related plants often produce seeds that are dispersed in very different ways, raising questions of how and why plants undergo adaptive shifts in key aspects of their reproductive ecology. Here we analyze the evolution of seed dispersal syndromes in an ancient group of plants. Ephedra (Gymnospermae; Gnetales; Ephedraceae) is a genus containing ≈50 species in semiarid ecosystems worldwide and with three distinct types of cones. We collected mature cones and seeds of ten species of Ephedra in southwestern United States and measured nine morphological traits for each species. Principal component analysis and other data characterized three types of Ephedra cones and seeds. Species with dry, winged cone bracts are dispersed by wind (i.e., E. torreyana and E. trifurca), those with succulent, brightly-colored cone bracts are dispersed by frugivorous birds (i.e., E. antisyphilitica), and those with small, dry cone bracts and large seeds are dispersed by seed-caching rodents (e.g., E. viridis and E. californica). Two species (E. funerea and E. nevadensis) have cone and seed morphologies intermediate between two seed dispersal syndromes. Seed and cones traits were mapped onto two recent phylogenies to help reveal the evolutionary history of seed dispersal syndromes. Bird dispersal is thought to be the ancestral form of seed dispersal in ephedras as it is common in the Old World where Ephedra originated, but the three North American species dispersed by birds are not monophyletic. The two wind dispersed species in North America also do not cluster together, suggesting separate origins. Seed dispersal by seed-caching rodents is common in North America and appears to have evolved several times, but this syndrome is absent form other continents. The evolutionary history of Ephedra in North America suggests that the means of seed dispersal has been malleable. Evolutionary shifts were likely linked to changes in ecological conditions.     

Sex pheromone of the cloaked pug moth,Eupithecia abietaria (Lepidoptera: Geometridae), a pest of spruce cones

The sex pheromone of the cloaked pug moth, Eupithecia abietaria Götze, an important cone‐feeding pest in spruce seed orchards in Europe, was investigated. Chemical and electrophysiological analyses of pheromone gland extracts of female moths and analogous analyses of synthetic hydrocarbons and epoxides of chain length C₁₉ and C₂₁ revealed (3Z,6Z,9Z)‐3,6,9‐nonadecatriene (3Z,6Z,9Z‐19:H) and 3Z,6Z‐cis‐9,10‐epoxynonadecadiene (3Z,6Z‐cis‐9,10‐epoxy‐19:H) as candidate pheromone components, which were found in a gland extract in a ratio of 95 : 5. In field trapping experiments, conspecific males were only attracted to a combination of 3Z,6Z,9Z‐19:H and the (9S,10R)‐enantiomer of 3Z,6Z‐cis‐9,10‐epoxy‐19:H. The (9R,10S)‐enantiomer was not attractive, which is in agreement with studies on other Eupithecia species, for which males have only been attracted by the (9S,10R)‐enantiomer of epoxides. Subsequent experiments showed that E. abietaria males were attracted to a wide range of ratios of the two active compounds and that trap catches increased with increasing dose of the binary blend. A two‐component bait containing 300 μg 3Z,6Z,9Z‐19:H and 33 μg of the (9S,10R)‐enantiomer of 3Z,6Z‐cis‐9,10‐epoxy‐19:H was efficient for monitoring E. abietaria in spruce seed orchards in southern Sweden, where this species has probably been overlooked as an important pest in the past. With sex pheromones recently identified for two other moths that are major pests on spruce cones, the spruce seed moth, Cydia strobilella L., and the spruce coneworm, Dioryctria abietella Denis & Schiffermüller, pheromone‐based monitoring can now be achieved for the whole guild of cone‐feeding moths in European spruce seed orchards.     

Variation in seed packaging of a fleshy‐fruited conifer provides insights into the ecology and evolution of multi‐seeded fruits

• The study of intraspecific seed packaging (i.e. seed size/number strategy) variation across different populations may allow better understanding of the ecological forces that drive seed evolution in plants. Juniperus thurifera (Cupressaceae) provides a good model to study this due to the existence of two subspecies differentiated by phenotypic traits, such as seed size and cone seediness (number of seeds inside a cone), across its range.
• The aim of this study was to analyse seed packaging (seed mass and cone seediness) variation at different scales (subspecies, populations and individuals) and the relationship between cone and seed traits in European and African J. thurifera populations.
• After opening more than 5300 cones and measuring 3600 seeds, we found that seed packaging traits followed different patterns of variation. Large‐scale effects (region and population) significantly contributed to cone seediness variance, while most of the seed mass variance occurred within individuals. Seed packaging differed between the two sides of the Mediterranean Sea, with African cones bearing fewer but larger seeds than the European ones. However, no differences in seed mass were found between populations when taking into account cone seediness. Larger cones contained more pulp and seeds and displayed a larger variation in individual seed mass.
• We validated previous reports on the intraspecific differences in J. thurifera seed packaging, although both subspecies followed the same seed size/number trade‐off. The higher seediness and variation in seed mass found in larger cones reveals that the positive relationship between seed and cone sizes may not be straightforward.We hypothesise that the large variation of seed size found within cones and individuals in J. thurifera, but also in other fleshy‐fruited species, could represent a bet‐hedging strategy for dispersal.

     

Asymmetrical development of biovulate cones resulting in uniovulate cones in Ephedra rhytidosperma (Ephedraceae)

Female cone morphology in Ephedra, including the number of initiated ovules and mature seeds per cone, provides important taxonomic characters used in sectional or species delimitation within Ephedra. Recent molecular phylogenies have indicated, however, that seed number per cone has changed repeatedly during the evolution of the genus. This study reports on the development of the female cone of E. rhytidosperma, based on a large sample of dissected cones studied under SEM. All cones were initially biovulate, and in the majority of cases, both female reproductive units (FRUs) developed a micropylar tube and formed mature seeds. In a few cases, the FRU pair developed asymmetrically in a cone, with one of them eventually aborting. There was no evidence of fusion of the FRU pair. Phylogenetically, E. rhytidosperma is in a clade with E. equisetina, which has uniovulate cones, and E. gerardiana and E. minuta, which have biovulate cones that also become unispermous via abortion. The biovulate condition may thus be ancestral in this clade.     

远东地区影响针叶树天然更新的球果种实害虫———球果花蝇 (双翅目 :花蝇科)( 英文)

球果花蝇是远东地区的重要球果种实害虫。尤其是在我国东北地区 ,由于球果花蝇的危害已严重影响了落叶松种子产量 ,进而影响到落叶松的造林与更新。本文系统地总结了近些年危害落叶松、云杉、冷杉种实的球果花蝇的研究进展及存在的问题。文中还涵括了作者的一些原始研究数据和尚未发表的新结果。     

Patterns of Cross-Continental Variation in Tree Seed Mass in the Canadian Boreal Forest

Seed mass is an adaptive trait affecting species distribution, population dynamics and community structure. In widely distributed species, variation in seed mass may reflect both genetic adaptation to local environments and adaptive phenotypic plasticity. Acknowledging the difficulty in separating these two aspects, we examined the causal relationships determining seed mass variation to better understand adaptability and/or plasticity of selected tree species to spatial/climatic variation. A total of 504, 481 and 454 seed collections of black spruce (Picea mariana (Mill.) B.S.P.), white spruce (Picea glauca (Moench) Voss) and jack pine (Pinus banksiana Lamb) across the Canadian Boreal Forest, respectively, were selected. Correlation analyses were used to determine how seed mass vary with latitude, longitude, and altitude. Structural Equation Modeling was used to examine how geographic and climatic variables influence seed mass. Climatic factors explained a large portion of the variation in seed mass (34, 14 and 29%, for black spruce, white spruce and jack pine, respectively), indicating species-specific adaptation to long term climate conditions. Higher annual mean temperature and winter precipitation caused greater seed mass in black spruce, but annual precipitation was the controlling factor for white spruce. The combination of factors such as growing season temperature and evapotranspiration, temperature seasonality and annual precipitation together determined seed mass of jack pine. Overall, sites with higher winter temperatures were correlated with larger seeds. Thus, long-term climatic conditions, at least in part, determined spatial variation in seed mass. Black spruce and Jack pine, species with relatively more specific habitat requirements and less plasticity, had more variation in seed mass explained by climate than did the more plastic species white spruce. As traits such as seed mass are related to seedling growth and survival, they potentially influence forest species composition in a changing climate and should be included in future modeling of vegetation shifts.     

Within-season synchrony of a masting conifer enhances seed escape

Predator satiation resulting from interannual reproductive synchrony has been widely documented in masting plants, but how reproductive synchrony within a year influences seed escape is poorly understood. We evaluated whether the intra-annual reproductive synchrony of individual white spruce trees (Picea glauca) increased seed escape from their primary predispersal seed predator, North American red squirrels (Tamiasciurus hudsonicus). Trees with cones that matured synchronously relative to those of other trees within red squirrel territories were significantly more likely to escape squirrel predation in years with both low and superabundant levels of cone production, generating a significantly positive linear selection differential for increasing intra-annual reproductive synchrony. Thus, this masting plant escapes seed predation in numbers through interannual synchrony in seed production and in time through intra-annual synchrony of seed availability.