Mechanical thinning to restore oak savanna promoted predator assemblages of gypsy moth pupae in Michigan

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Geographical variation in the periodicity of gypsy moth outbreaks

The existence of periodic oscillations in populations of forest Lepidoptera is well known. While information exists on how the periods of oscillations vary among different species, there is little prior evidence of variation in periodicity within the range of a single Lepidopteran species. The exotic gypsy moth is an introduced foliage-feeding insect in North America. Its populations are characterized by high amplitude oscillations between innocuously low densities and outbreak levels during which large regions of forests are defoliated. These outbreaks are recognized to arise periodically with considerable synchrony across much of the gypsy moth's North American range. Our analysis indicates that gypsy moth outbreaks in North America are periodic but they exhibit two dominant periodicities: a primary period of 8–10 yr (as previously reported) and a secondary period of 4–5 yr (a new finding in this study). The outbreak periodicity varied geographically and this variation was associated with forest type. We found that in the most susceptible forest types, those on xeric sites where oak is often mixed with pines, outbreak periodicity had a more dominant 5-yr period while in forest types characteristic of more mesic sites where oak was mixed with maples and other species, cycles were clearly operating on a 10-yr period.     

What causes outbreaks of the gypsy moth in North America?

The gypsy moth has been present in North America for more than 100 years, and in many of the areas where it has become established outbreaks occur with varying degrees of periodicity. There also exists extensive spatial synchrony in the onset of outbreaks over large geographic regions. Density-dependent mortality clearly limits high-density populations, but there is little evidence for strong regulation of low-density populations. Predation by small mammals appears to be the major source of mortality affecting low-density populations, but because these are generalist predators and gypsy moths are a less preferred food item, mammals do not appear to regulate populations in a density-dependent fashion. Instead, predation levels appear to be primarily determined by small mammal abundance, which is in turn closely linked to the production of acorns that are a major source of food for overwintering predator populations. Mast production by host oak trees is typically variable among years, but considerable spatial synchrony in masting exists over large geographic areas. Thus, it appears that the temporal and spatial patterns of mast production may be responsible for the episodic and spatially synchronous behavior of gypsy moth outbreaks in North America. This multitrophic relationship among mast, predators, and gypsy moths represents a very different explanation of forest insect outbreak dynamics than the more widely applied theories based upon predator–prey cycles or feedbacks with host foliage quality. Received: September 8, 1999 / Accepted: September 20, 2000     

Moth outbreaks in relation to oak masting and population levels of small mammals: an alternative explanation to the mammal-predation hypothesis

The relationship between population outbreaks of the gypsy moth (Lymantria dispar) and oak masting in North America has been interpreted as an effect of reduced predation on moth pupae from small mammals after years of acorn failure. However, moth defoliation could be a consequence of high acorn production rather than of acorn failure, as all moth outbreaks in two time series presented by Liebhold et al. [Popul Ecol (2000) 42:257–266] were reported shortly after mast years. A similar pattern has been found for the green oak leaf roller moth (Tortrix viridana) in southern Norway. Because predation from small mammals should be less important for the latter species, I argue that the acorn-moth relationships are most likely caused by mast-induced changes in the chemical composition of oak leaves. Given the high number of eggs laid by each moth female, there is a huge potential for population growth in or shortly after a mast year if larval survival is no longer limited by low food quality.     

Effects of alternative prey on predation by small mammals on gypsy moth pupae

Previous work shows that predation by small mammals is a dominant cause of mortality of low-density gypsy moths in North America and that declines in small mammal density result in increases in gypsy moth density. Here we examined whether predation by small mammals is density dependent by way of a type III functional response, and how predation is influenced by alternative prey. First we showed that the preference of predators for gypsy moth pupae was low compared to other experimental prey items, such as mealworm pupae and sunflower seeds. Predation on gypsy moth pupae was characterized by a type II functional response with percent predation highest at the lowest prey densities, whereas the functional response to sunflower seeds was characterized by a type III functional response in which predation increased with increasing prey density. These results suggest that predation by small mammals is unlikely to stabilize low-density gypsy moth populations.     

Predators ofLymantria dispar (Lep. lymantriidae) egg masses: Spatio-temporal variation of their impact during the 1988–89 pest generation in the mamora cork oak forest (Morocco)

In the Mamora cork oak forest (Morocco), oophagous predators of the gypsy mothLymantria dispar (L.) (Lepidoptera: Lymantriidae) are one of the main mortality factors influencing pest population dynamics. The predators destroy egg masses more by disrupting their cohesiveness than by predation. From 1987 to 1990, the impact of oophagous enemies significantly contributed to the collapse of a localized gypsy moth outbreak. Variation of egg mass dislocation intensity was greater between trees than among the different strata of a tree. The number of gypsy moth egg masses and egg predator attacks increased when oaks were large and unhealthy. Forest degradation probably explains why egg mass destruction rates were so high (60 to 90% of the eggs) in the infested forest. Oophagous predators find food and shelter under the dehiscent bark of unhealthy cork oaks.     

Host‐associated divergence in the activity of digestive enzymes in two populations of the gypsy moth Lymantria dispar (Lepidoptera: Erebidae)

The gypsy moth is a generalist insect pest with an extremely wide host range. Adaptive responses of digestive enzymes are important for the successful utilization of plant hosts that differ in the contents and ratios of constituent nutrients and allelochemicals. In the present study, we examined the responses of α‐amylase, trypsin, and leucine aminopeptidase to two tree hosts (suitable oak, Quercus cerris, and unsuitable locust tree, Robinia pseudoacacia) in the fourth, fifth, and sixth instars of gypsy moth larvae originating from oak and locust tree forest populations (hereafter assigned as Quercus and Robinia populations, respectively). Gypsy moths from the Robinia forest had been adapting to this unsuitable host for more than 40 generations. To test for population‐level host plant specialization, we applied a two‐population × two‐host experimental design. We compared the levels, developmental patterns, and plasticities of the activities of enzymes. The locust tree diet increased enzyme activity in the fourth instar and reduced activity in advanced instars of the Quercus larvae in comparison to the oak diet. These larvae also exhibited opposite developmental trajectories on the two hosts, i.e. activity increased on the oak diet and decreased on the locust tree diet with the progress of instar. Larvae of the Robinia population were characterized by reduced plasticity of enzyme activity and its developmental trajectories. In addition, elevated trypsin activity in response to an unsuitable host was observed in all instar larvae of the Robinia population, which demonstrated that Robinia larvae had an improved digestive performance than did Quercus larvae.     

Phenotypic plasticity and population differentiation in seeds and seedlings of the grass Anthoxanthum odoratum

Summary Seeds of Anthoxanthum odoratum were transplanted reciprocally between a xeric and a mesic field population that were genetically differentiated in adult traits. In one experiment seeds were reciprocally buried in bags in the soil, in a second experiment seeds were reciprocally sown in small plots. For most traits, site effects were much larger than seed-source effects. Germination, emergence, mortality of buried seed and recruitment were significantly higher at the xeric site than at the mesic site, irrespective of population of origin. Seed dormancy, was significantly higher for seed originating from the mesic than from the xeric population. Seedling recruits originating from the xeric population tended to be larger at both sites. Fecundity of seedling recruits depended on the environment; fecundities of plants growing in the xeric site had more than double the fecundity of plants growing in the mesic site. Phenotypic plasticity rather than population differences determined variation in performance in the seed and seedling stages.     

Population age structure of Pinus banksiana at the southern edge of the Canadian boreal forest

Abstract. To assess the effects of site type, forest initiation periods and fire regimes on the dynamics of Pinus banksiana (Jack pine), the age structure of 69 populations of the species was analyzed. Two landscapes with different fire regimes were selected in the southern part of the Canadian boreal forest in Québec: the ‘mainland landscape’ is characterized by a fire regime of large lethal fires, the ‘island landscape’ is affected by a complex fire regime including lethal and non-lethal fires. Age structure was compared between forest initiation periods and site types (mesic mainland, xeric mainland and xeric island) using the Shannon regularity index. An even-aged population structure was found within the first 100 yr following a lethal fire, while after that period the population structure becomes more uneven-aged. Under mesic conditions, populations tend to have an even-aged structure, under xeric conditions an uneven-aged structure. Natural openings present in xeric sites allow for recruitment in the absence of fire. This permits the self-maintenance of Pinus banksiana. Xeric island populations show more uneven-aged structures than xeric mainland populations. The occurrence of non-lethal fires on the islands creates uneven-aged structures. Further, the results suggest that the selection pressure of the island fire regime, favouring non-serotinous and mixed P. banksiana individuals, is one of the factors responsible for a higher recruitment in the absence of fire on islands than on the mainland.     

The potential for Entomophaga maimaiga to regulate gypsy moth Lymantria dispar (L.) (Lepidoptera: Erebidae) in Europe

Gypsy moth, Lymantria dispar L., is one of the most important pests of deciduous trees in Europe. In regular cycles, it causes large‐scale defoliation mostly of oak, Quercus spp., forests. Government authorities in the most infested countries in Europe conduct large‐scale applications of pesticides against gypsy moth. In 1999, a new natural enemy, the entomopathogenic fungus Entomophaga maimaiga, was successfully introduced into a gypsy moth population in Bulgaria. Recent investigations suggest that now E. maimaiga is quickly spreading in Europe. Herein, past studies are reviewed regarding this fungus with special emphasis on its potential for becoming an important factor regulating gypsy moth populations in Europe, focusing on the host's population dynamics in relation to the fungus, the influence of environmental conditions on fungal activity, the influence of E. maimaiga on the native entomofauna, including other gypsy moth natural enemies, and spread of the fungus. Based on this analysis, the potential of E. maimaiga for providing control in European gypsy moth populations is estimated.     

Inference of adult female dispersal from the distribution of gypsy moth egg masses in a Japanese city

1 The native range of the gypsy moth Lymantria dispar (L.) spans the temperate forests of Eurasia. Across this region, a clinal female flight polymorphism exists; gypsy moth females in eastern Asia are mostly capable of directed flight, those in western and southern Europe are largely incapable of flight and populations distributed across the centre of the distribution exhibit a range of intermediate flight behaviours. 2 Although information exists about the timing and duration of female flight from laboratory and wind tunnel studies, little or no quantitative data are available on average distances flown by Asian gypsy moth females prior to oviposition in the field. This information is critical for estimating risk of contamination at specific ports and transit terminals, as well as for predicting the spread of populations that might someday invade currently uninfested regions of the world. 3 In the present study, an extensive visual survey of gypsy moth egg masses was conducted during a walk through streets and paths in a 3.92 × 5.76 km area in Kanazawa, Japan. This area consisted of a matrix of urban, agricultural and forest land uses. The distribution of egg masses relative to distances from host forests was used to infer the magnitude of pre‐ovipositional female flight. 4 A total of 3172 egg masses was recorded from surveys conducted during the search of a path totalling 384 km. Within urban areas, egg masses were most abundant in the area <1 km from the edge of forest land. 5 These results suggest that most female gypsy moth flight is limited to the area within 1 km of host forests. They also suggest that shipping containers and other parcels located >1 km from forests are at a much lower risk of contamination with Asian gypsy moth egg masses.     

Genetic structure,admixture and invasion success in a Holarctic defoliator,the gypsy moth (Lymantria dispar,Lepidoptera: Erebidae)

Characterizing the current population structure of potentially invasive species provides a critical context for identifying source populations and for understanding why invasions are successful. Non‐native populations inevitably lose genetic diversity during initial colonization events, but subsequent admixture among independently introduced lineages may increase both genetic variation and adaptive potential. Here we characterize the population structure of the gypsy moth (Lymantria dispar Linnaeus), one of the world's most destructive forest pests. Native to Eurasia and recently introduced to North America, the current distribution of gypsy moth includes forests throughout the temperate region of the northern hemisphere. Analyses of microsatellite loci and mitochondrial DNA sequences for 1738 individuals identified four genetic clusters within L. dispar. Three of these clusters correspond to the three named subspecies; North American populations represent a distinct fourth cluster, presumably a consequence of the population bottleneck and allele frequency change that accompanied introduction. We find no evidence that admixture has been an important catalyst of the successful invasion and range expansion in North America. However, we do find evidence of ongoing hybridization between subspecies and increased genetic variation in gypsy moth populations from Eastern Asia, populations that now pose a threat of further human‐mediated introductions. Finally, we show that current patterns of variation can be explained in terms of climate and habitat changes during the Pleistocene, a time when temperate forests expanded and contracted. Deeply diverged matrilines in Europe imply that gypsy moths have been there for a long time and are not recent arrivals from Asia.     

Benefits and harm caused by the introduced generalist tachinid, <Emphasis Type="Italic">Compsilura concinnata</Emphasis>, in North America

We review North American research on Compsilura concinnata (Meigen), a highly generalist tachinid parasitoid that was introduced in 1906 to control two invasive forest insects: gypsy moth and browntail moth. The impact on gypsy moth in natural populations in North America has long thought to be minor, a view confirmed by recent work in Canada and Wisconsin, USA. However, research with experimentally created populations of gypsy moth suggests that it may be more important than previously realized. Studies on browntail moth in both naturally occurring and experimental populations suggest that C. concinnata is probably the main reason browntail moth disappeared from most of its former range in North America. Research on giant silk moths suggests that C. concinnata has become the major source of mortality among several species and may be responsible for the notable decline in their densities that has occurred over the last century. C. concinnata has been touted as a premier example of the generalist natural enemy that should be avoided in classical biological control introductions, yet in the case of browntail moth its effect has been extremely beneficial.     

Earlywood vessel size of oak as a potential proxy for spring precipitation in mesic sites

Aim In this study, we evaluate the importance of the mean earlywood vessel size of oaks as a potential proxy for climate in mesic areas. Location The study was conducted in Switzerland at three forest sites dominated by oak (Quercus petraea and Q. pubescens). The three sites were in different climatic zones, varying mainly in terms of precipitation regime. Methods Three 50‐year‐long site chronologies of mean earlywood vessel size and tree‐ring widths were obtained at each site and related to monthly meteorological records in order to identify the main variables controlling growth. The responses of mean vessel size to climate were compared with those of the width variables to evaluate the potential climatic information recorded by the earlywood vessels. Results The results show that the mean vessel size has a different and stronger response to climate than ring‐width variables, although its common signal and year‐to‐year variability are lower. This response is better in particular at mesic sites, where it is linked to precipitation during spring, i.e. at the time of vessel formation, and is probably related to the occurrence of only a few processes controlling vessel growth, whereas radial increment is controlled by multiple and varying factors. Main conclusions The mean earlywood vessel size of oak appears to be a promising proxy for future climate reconstructions of mesic sites, where radial growth is not controlled by a single limiting factor.     

Lepidoptera‐specific insecticide used to suppress gypsy moth outbreaks may benefit non‐target forest Lepidoptera

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SPECIES RESPONSE TO A MOISTURE GRADIENT IN CENTRAL ILLINOIS FORESTS

Polar and Gaussian ordination applied to data collected from 37 forest sites in central Illinois resulted in a continuous and gradual change in species composition along a moisture gradient. A series of overlapping species success curves formed by plotting Importance Values over stands ordered along the gradient varied continuously in modal location and habitat width. Blackjack oak and black oak dominated upland sandy sites. Black oak, white oak, and shagbark hickory were the most important species on exposed, upper slope positions or ridge tops with silt-loam soils. Red oak, sugar maple, American elm, and bur oak dominated sheltered locations on lower slope positions and stream terraces. Sycamore, silver maple, and cottonwood were leading tree species in floodplain forests. Conversion of black, white, and red oak forests on silt-loam sites to sugar maple, white ash, and red elm dominance is evident by high densities of these shade tolerant species in the understory. Composition of forests at the extreme ends of the moisture gradient is more stable than the mesic sites. Maximum tree diversity occurred on mesic sites and decreased toward the extreme ends of the moisture gradient. However, competitive exclusion of shade intolerant species by sugar maple and other species has caused a decrease in understory diversity on mesic sites. Diversity decreased from canopy to understory strata in lowland forests and increased on xeric sites.     

Impact of predators on artificially augmented populations of Lymantria dispar L. pupae (Lep., Lymantriidae)

The impact of predators on artificially augmented populations of Lymantria dispar pupae (Lep., Lymantriidae) and the species composition of the predator community were investigated in two oak stands in eastern Austria. The population density of L. dispar has been at innocuous levels for several years at both sites. From mid-June to mid-July, we created artificial prey populations by mounting gypsy moth pupae with beeswax on burlap bands placed at the base, 25, 50 and 100 cm height on 25 trees at each site. A total of 6600 pupae were exposed at each site. A total of 92% of exposed pupae were destroyed by predators, at site I (with dense understorey vegetation) whereas 67% were destroyed on site II (with sparse understorey vegetation). Initially, pupal mortality was highest at the base of trees, but differences in mortality among the four locations where pupae were exposed decreased during the course of the study period. Spatial differences in predation were ascertained, suggesting that the aggregation of small mammals, for example, is related to patches of dense shrub vegetation. Pupal mortality was primarily caused by mice (46.3% at site I and 36.4% at site II). Many exposed pupae simply disappeared (40.4% at site I and 22.8% at site II). Calosoma spp. and other invertebrate predators caused very little mortality among L. dispar pupae (5.5% at site I and 7.8% at site II). Trap catches on separate study plots at both sites revealed that Apodemus flavicollis (Rodentia, Muridae) was the most important predator species present, whereas Apodemus sylvaticus (Rodentia, Muridae) and Clethrionomys glareolus (Rodentia, Arvicolidae) were captured in low numbers.     

Effects of position, understorey vegetation and coarse woody debris on tree regeneration in two environmentally contrasting forests of north-western Patagonia: a manipulative approach

Aim To investigate the differential effects of position within gaps, coarse woody debris and understorey cover on tree seedling survival in canopy gaps in two old‐growth Nothofagus pumilio (Poepp. & Endl.) Krasser forests and the response of this species to gaps in two forests located at opposite extremes of a steep rainfall gradient. Location Nahuel Huapi National Park, at 41° S in north‐western Patagonia, Argentina. Methods In both study sites, seedlings were transplanted to experimental plots in gaps in three different positions, with two types of substrate (coarse woody debris or forest floor), and with and without removal of understorey vegetation. Survival of seedlings was monitored during two growing seasons. Soil moisture and direct solar radiation were measured once in mid‐summer. Seedling aerial biomass was estimated at the end of the experiment. Results Mid‐summer soil water potential was lowest in the centre of gaps, in plots where the understorey had been removed, and highest at the northern edges of gaps. Direct incoming radiation was highest in gap centres and southern edges, and lowest at northern edges. Seedling mortality was highest in gap centres, in both sites. Coarse woody debris had a positive effect on seedling survival during summer in the mesic forest and during winter in the xeric forest. The removal of understorey cover had negative effects in gap centres during summer. Seedling final aerial biomass was positively affected by understorey removal and by soil substrate in both sites. In the dry forest gaps, seedling growth was highest in northern edges, whereas it was highest in gap centres in the mesic forest. Overall growth was positively related to survival in the xeric forest, and negatively related in the mesic forest. Main conclusions Survival and growth were facilitated by the shade of gap‐surrounding trees only in the xeric forest. Understorey vegetation of both forests facilitated seedling survival in exposed microsites but competed with seedling growth. Nurse logs were an important substrate for seedling establishment in both forests; however, causes of this pattern differed between forests. Water availability positively controls seedling survival and growth in the xeric forest while in the mesic forest, survival and growth are differentially controlled by water and light availability, respectively. These two contrasting old‐growth forests, separated by a relatively short distance along a steep rainfall gradient, had different yet unexpected microenvironmental controls on N. pumilio seedling survival and growth. These results underscore the importance of defining microscale limiting factors of tree recruitment in the context of large‐scale spatial variation in resources.     

Seed survival on experimental dishes in a central European old‐growth mixed‐species forest – effects of predator guilds,tree masting and small mammal population dynamics

Predation of tree seeds can be a major factor structuring plant communities. We present a three year study on tree seed survival on experimental dishes in an old‐growth forest in central Europe in Austria. We addressed species specific, spatial and temporal aspects of post‐dispersal seed predation. Seeds of Norway spruce Picea abies, European beech Fagus sylvatica, and silver fir Abies alba were exposed on dishes in different types of exclosures which allowed access only to specific guilds of seed predators. Removal experiments were carried out in two old‐growth forests and a managed forest (macro‐sites), including micro‐sites with and without cover of ground vegetation. We conducted the experiment in three consecutive years with a mast year of beech and spruce before the first year of the study. The seed removal experiments were combined with live trapping of small mammals being potential seed predators. Our experiments showed a distinctly different impact of different predator guilds on seed survival on the dishes with highest removal rates of seeds from dishes accessible for small mammals. We observed differing preferences of small mammals for the different tree species. Seed survival in different macro‐ and micro‐habitats were highly variable with lower seed survival in old growth forests. In contrast to our assumption, and in contrast to the satiation hypothesis which assumes higher seed survival in and directly after mast years, seed survival was lower in the year following the mast year of beech when a population peak of small mammals occurred and higher in intermast periods when subsequently small mammal population crashed. This suggests a higher importance of sporadic masting shortly after mast years in intermast periods for establishment of forest trees provided that pollination efficiency is high enough in such years. Combined with the high seed mortality observed after the mast year, this corroborates the important role of seed predation for forest dynamics. An altered synchrony or asynchrony of masting of different tree species and changed masting frequencies through climate change may thus lead to strong and non‐linear effects on forest dynamics.     

The Allee effect, stochastic dynamics and the eradication of alien species

Previous treatments of the population biology of eradication have assumed that eradication can only be achieved via 100% removal of the alien population. However, this assumption appears to be incorrect because stochastic dynamics and the Allee effect typically contribute to the extinction of very low‐density populations. We explore a model that incorporates Allee dynamics and stochasticity to observe how these two processes contribute to the extinction of isolated populations following eradication treatments of varying strength (percentage mortality). As a case study, we used historical data on the dynamics of isolated gypsy moth, Lymantria dispar, populations to fit parameters to this model. The parameterized model was then used in simulations that evaluated the efficacy of various eradication strategies. The results indicated that eradication of isolated gypsy moth populations could be easily achieved following a treatment of >80% mortality as long as populations were relatively low (indicated by <100 males captured in pheromone traps).