Outbreaks of the winter moth on Sitka Spruce in Scotland are not influenced by nutrient deficiencies of trees,tree budburst,or pupal predation

Summary Since the early 1980s, the winter moth, Operophtera brumata L. (Lepidoptera: Geometridae) has emerged as a serious pest of Sitka Spruce, Picea sitchensis Bong. plantations in southern Scotland. Outbreaks are characterised by susceptible sites within plantations which can occur immediately adjacent to resistant sites. We investigated the level of some nutrients in the trees, the date of budburst of the trees, and the numbers of some potential predators of winter moth pupae. None could satisfactorily explain outbreak patterns. Although foliage analysis demonstrated that many trees were marginal or deficient in phosphorus, nitrogen and potassium, these deficiencies were not related to the susceptibility of a site. Within sites, the numbers and weights of O. brumata were positively related to phosphorus content and negatively related to calcium content of foliage. Other evidence suggests, however, that these correlations may not represent direct effects of phosphorus and calcium on larval growth and survival. Date of budburst, which commonly determines susceptibility of deciduous hosts to O. brumata, was unrelated to density, and pupal predators were more, not less, abundant in susceptible sites. Although it is difficult to distinguish between factors that initiate outbreaks and those that maintain them, these data suggest that nutrient deficiencies of trees, budburst date, and the distribution of pupal predators of the winter moth cannot explain patterns of outbreak of the winter moth on spruce.     

Delayed density-dependent parasitism of eggs and pupae as a contributor to the cyclic population dynamics of the autumnal moth

Many populations of forest Lepidoptera exhibit 10-year cycles in densities, with impressive outbreaks across large regions. Delayed density-dependent interactions with natural enemies are recognized as key factors driving these cyclic population dynamics, but emphasis has typically been on the larval stages. Eggs, pupae and adults also suffer mortality from predators, parasitoids and pathogens, but little is known about possible density relationships between mortality factors and these non-feeding life stages. In a long-term field study, we experimentally deployed autumnal moth (Epirrita autumnata) eggs and pupae to their natural enemies yearly throughout the 10-year population cycle in northern Norway. The abundance of another geometrid, the winter moth (Operophtera brumata), increased in the study area, permitting comparisons between the two moth species in predation and parasitism. Survival of autumnal moth eggs and pupae was related to the moth abundance in an inverse and delayed manner. Egg and pupal parasitoids dominated as density-dependent mortality factors and predicted the subsequent growth rate of the host population size. In contrast, effects of egg and pupal predators were weakly density dependent, and generally predation remained low. Parasitism rates did not differ between the autumnal and winter moth pupae, whereas predators preferred winter moth pupae over those of the autumnal moth. We conclude that parasitism of the autumnal moth by egg and pupal parasitoids can be related to the changes of the moth density in a delayed density-dependent manner. Furthermore, egg and pupal parasitoids cannot be overlooked as causal factors for the population cycles of forest Lepidoptera in general.     

Responses of generalist invertebrate predators to pupal densities of autumnal and winter moths under field conditions

• 1 Generalist natural enemies are usually not considered as being capable of causing population cycles in forest insects, but they may influence the population dynamics of their prey in the low density cycle phase when specialist enemies are largely absent.
• 2 In the present field study, the total response of the generalist invertebrate predator community to experimentally established pupal densities of the closely related autumnal (Epirrita autumnata) and winter moths (Operophtera brumata) was analysed.
• 3 Due to the high amount of variation in the dataset, the exact shape of the response curve could not be convincingly estimated. Nevertheless, two important conclusions can be drawn from the analyses.
• 4 Firstly, the natural invertebrate predator community seems to become saturated at rather low densities of both autumnal and winter moth pupae. Secondly, the predator community seems to become saturated at much lower densities of autumnal than of winter moth pupae.
• 5 Furthermore, pupal mass was significantly negatively correlated with invertebrate predation probability in autumnal moth pupae.
• 6 These results indicate that differences in the predator assemblage being able to consume pupae of the two moth species, as well as different handling times, could be responsible for the substantially higher predation rates in winter than in autumnal moth pupae.
• 7 As a consequence, the population dynamics of autumnal moths might be less affected by generalist invertebrate predators than those of winter moths, as autumnal moths seem able to escape from the regulating influence of generalist predators at much lower population densities than winter moths.

     

Local outbreaks of Operophtera brumata and Operophtera fagata cannot be explained by low vulnerability to pupal predation

• 1 One of the unresolved questions in studies on population dynamics of forest Lepidoptera is why some populations at times reach outbreak densities, whereas others never do. Resolving this question is especially challenging if populations of the same species in different areas or of closely‐related species in the same area are considered.
• 2 The present study focused on three closely‐related geometrid moth species, autumnal Epirrita autumnata, winter Operophtera brumata and northern winter moths Operophtera fagata, in southern Finland. There, winter and northern winter moth populations can reach outbreak densities, whereas autumnal moth densities stay relatively low.
• 3 We tested the hypothesis that a lower vulnerability to pupal predation may explain the observed differences in population dynamics. The results obtained do not support this hypothesis because pupal predation probabilities were not significantly different between the two genera within or without the Operophtera outbreak area or in years with or without a current Operophtera outbreak.
• 4 Overall, pupal predation was even higher in winter and northern winter moths than in autumnal moths. Differences in larval predation and parasitism, as well as in the reproductive capacities of the species, might be other candidates.

     

Parasitism and predation as agents of mortality of winter moth populations in neglected apple orchards in Nova Scotia

Abstract.

• 1 This study compared the roles of pupal mortality and parasitism in winter moth (Operophtera brumata) population dynamics in Nova Scotian apple orchards and assessed the importance of beetles as pupal predators.
• 2 The component of pupal mortality termed predation accounted for greater stage-specific mortality of winter moth than parasitism by Cyzenis albicans in four neglected orchards.
• 3 Parasitism by Cyzenis albicans was not spatially density-dependent in any orchard, whereas the predation component of pupal mortality was spatially density-dependent in the two orchards most densely populated by winter moth.
• 4 Field experiments indicated that over 60% of pupal predation may be attributed to beetles, and that about 46% of pupal predation occurred within 4 weeks after pupal drop.
• 5 Mortality of introduced populations of winter moth in Nova Scotia resembles that of native populations in England where density-dependent predation regulates the winter moth population and reduces the parasitoid population to minimal levels. The situation in Nova Scotia appears to have changed appreciably since the establishment of parasitoids into the system in the 1950s.

     

Predicted changes in the synchrony of larval emergence and budburst under climatic warming

Summary The impact of climatic warming on the synchrony of insect and plant phenologies was modelled in the case of winter moth (Operophtera brumata) and Sitka spruce (Picea sitchensis) in the Scottish uplands. The emergence of winter moth larvae was predicted with a thermal time requirement model and the budburst of Sitka spruce was predicted from a previously published model (Cannell and Smith 1983) based on winter chilling and thermal time. The date of emergence of winter moth larvae was predicted to occur earlier under climatic warming but the date of budburst of Sitka spruce was not greatly changed, resulting in decreased synchrony between larval emergence and budburst. The general question of how a change of climate might affect phenological synchrony and insect abundance is discussed.     

Testing the enemies hypothesis in forest stands: the important role of tree species composition

Numerous studies conducted in agro-ecosystems support the enemies hypothesis, which states that predators and parasites are more efficient in controlling pest densities in polycultures than in monocultures. Few similar studies, however, have been conducted in forest ecosystems, and we do not yet have evidence as to whether the enemies hypothesis holds true in forests. In a 2-year study, we investigated whether the survival of autumnal moth (Epirrita autumnata) larvae and pupae differs between silver birch monocultures and two-species mixtures of birch with black alder, Norway spruce and Scots pine. We placed young larvae on birch saplings and monitored their survival until the end of the larval period, when we checked whether they had been parasitized. After the larvae had pupated, pupal survival was tested in a field trial. In 2002, the larvae disappeared earlier and their overall survival was lower in birch–pine mixtures than in other stand types. In 2003, survival probability was lowest in birch–pine stands only during the first week and there were no differences between stands in overall survival. Larval parasitism was not affected by tree species composition. Pupal weight and pupal survival were likewise not affected by stand type. Among the predators, wood ants were more abundant on birches growing in birch–pine mixtures than in other stand types probably because colonies of myrmecophilic aphids were common on pines. In contrast, spider numbers did not differ between stand types. Ant exclusion by means of a glue ring around the birch trunk increased larval survival, indicating that ants are important predators of the autumnal moth larvae; differences in larval survival between stands are probably due to differential ant predation. Our results provide only partial support for the enemies hypothesis, and suggest that it is both tree species composition and species diversity which affect herbivore survival and predation.     

Biodiversity and ecology of the invertebrates of Atlantic Oakwoods

Summary

Woodland in general supports the richest invertebrate fauna of any habitat in Britain. The number of species depends on the structure of the wood and the age of the trees within it. The greatest biodiversity is associated with oaks although the difference between the communities found on the two oak species is not clearly understood, the Sessile Oak woods have been less well studied than their lowland equivalents, particularly so in Scotland. A number of minor habitats within the wood are important for many invertebrates and the presence of many of these may be due to past management. Every part of the oak tree, the leaves, flowers, acorns, bark and wood, supports invertebrates, herbivores, predators and parasites, and the fallen leaves and deadwood provides habitat for a complex of detritus feeders. Within the woodland the other trees, the fungi and the ground flora all have characteristic woodland invertebrates. Only a few studies, on beetles and spiders, have looked at which invertebrates are particularly associated with acidic oakwoods and there have been no specific studies of Atlantic oakwoods. It is known that there are some rare and local invertebrate species associated with western woods on ancient sites, including some recently discovered suggesting that the biodiversity of these sites and their management for invertebrates might be more important than currently thought.     

Macrofungal diversity and ecology in four Irish forest types

The macrofungal communities of Irish native tree species (ash and oak) and exotic tree species (Scots pine and Sitka spruce) forests were examined through the collection of sporocarps over 3 yr. Sampling of 27 plots revealed 186 species of macrofungi, including 10 species new to Ireland. The species richness of non-native Sitka spruce and Scots pine forests was similar to that of native oak forests. However, specific communities of macrofungi existed in each of the forest types as confirmed by non-metric multidimensional scaling and multi-response permutation procedure. Indicator species analysis was used to identify macrofungi which are indicative of the four forest types. The oak community lacked certain species/genera known to be distinctive of oak woods in Britain, possibly due to low inoculum availability as a result of historic removal of Ireland’s oak forests. Our results indicate that, while being similar to native forests in species richness, non-native forests of Sitka spruce and Scots pine in Ireland harbour many fungal species which are not typical of native forests, particularly members of the genus Cortinarius.     

瑞典南部落叶林动物对种子捕食的实验研究

在瑞典南部落叶林,于1985年和1986年秋季分别研究了不同动物对欧洲七叶树、夏栎、山楂和黑刺李等4种种子的捕食作用,即种子释放1周后被动物捕食的情况。实验分以下3种处理:排除小兽和鸟、排除鸟和对照(小兽和鸟均不排除),但昆虫可进入所有处理。在不同落叶林和小型林地共建有16个重复实验,即每个林分类型于每年秋季各设4个重复。在所有林分(包括落叶林和小型林地),不同处理的种子捕食表明:小兽是欧洲七叶树、夏栎和黑刺李等植物种子的主要捕食者,而无脊椎动物则是山楂种子的重要捕食者,但鸟类很少取食上述种子,因而捕食作用极小。在其中1年,动物对种子的总捕食率在落叶林和小型林地存在较大差异,以小型林地较高。     

Forest type affects predation on gypsy moth pupae

Abstract 1 Predation by small mammals has previously been shown to be the largest source of mortality in low‐density gypsy moth, Lymantria dispar (L.), populations in established populations in north‐eastern North America. Fluctuations in predation levels are critical in determining changes in population densities. 2 We compared small mammal communities and levels of predation on gypsy moth pupae among five different oak‐dominated forest types along this insect's western expanding population front in Wisconsin. Comparisons of predator impact can provide critical information for predicting variation in susceptibility among forest types. 3 The results indicated that small mammals caused more mortality than did invertebrates. 4 Both abundance of Peromyscus sp. predators and predation levels were lower in urban and xeric forest types than in mesic sites. 5 These results suggest that, because predation pressures will probably be greater in the mesic sites, gypsy moths may be less likely to develop outbreaks in these habitats, and that defoliation will probably be more frequent in urban and xeric oak‐dominated sites.     

Elevationally biased avian predation as a contributor to the spatial distribution of geometrid moth outbreaks in sub‐arctic mountain birch forest

1. Population dynamics and interactions that vary over a species' range are of particular importance in the context of latitudinal clines in biological diversity. Winter moth (Operophtera brumata) and autumnal moth (Epirrita autumnata) are two species of eruptive geometrids that vary widely in outbreak tendency over their range, which generally increases from south to north and with elevation. 2. The predation pressure on geometrid larvae and pupae over an elevational gradient was tested. The effects of background larval density and bird occupancy of monitoring nest boxes on predation rates were also tested. Predation on larvae was tested through exclusion treatments at 20 replicate stations over four elevations at one site, while pupae were set out to measure predation at two elevations at three sites. 3. Larval densities were reduced by bird predation at three lower elevations, but not at the highest elevation, and predation rates were 1.9 times higher at the lowest elevation than at the highest elevation. The rate of predation on larvae was not related to background larval density or nest box occupancy, although there were more eggs and chicks at the lowest elevation. There were no consistent differences in predation on pupae by elevation. 4. These results suggest that elevational variation in avian predation pressure on larvae may help to drive elevational differences in outbreak tendency, and that birds may play a more important role in geometrid population dynamics than the focus on invertebrate and soil predators of previous work would suggest.     

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     

Long-term microtine dynamics in north Fennoscandian tundra: the vole cycle and the lemming chaos

Densities of microtine rodents in two habitat complexes in the tundra of Finnmarks-vidda, Norwegian Lapland, were studied during 1977-89 by means of snap trapping (Small Quadrat Method) Predator populations were studied by mapping breeding raptors and by snow-tracking small mustelids During 1977-85, snow-trackmg was conducted only during peak and decline years, whereas during 1986-89, snow-tracking was conducted every winter (November-December) and live-trapping (in August) was used as an additional method
Lowland vole populations had regular density fluctuations with peaks in 1978-79. 1982-84 and 1987-88 Highland vole populations fluctuated less regularly and at lower over-all densities Highland lemming populations had two outbreaks, in 1978 and 1988, ending in abrupt winter crashes In the lowland, outbreak levels were reached only in 1978 All microtine declines in relatively productive lowland habitats were accompanied by intense activity of small mustelids. whereas avian predators were common only in 1983 Lowland declines also showed clear between-habitat asynchrony they started in areas with an exceptional abundance of productive habitats and then spread to more barren areas These lowland data are consistent with the hypothesis of a mustelid-microtine limit cycle, although also several other hypotheses remain unrefuted The highland lemming data suggest a simple exploiter-victim interaction between lemmings and the vegetation     

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.     

Density dependence and the stabilization of animal numbers

Summary Latto and Hassell (1987) disagree with the conclusion of Den Boer (1986), that the winter moth population at Wytham Wood, studied by Varley and Gradwell, was not regulated. They attempt to demonstrate regulation by means of a simulation model. In the present paper the validity of this model is tested step by step. The fixing of the initial and final densities, as practised by Den Boer and rejected by Latto and Hassell, did not prevent population explosions and extinctions, as was assumed by Latto and Hassell. It is shown that the deterministic formulation of the density dependence of pupal predation, as used by Latto and Hassell, deviates systematically from the field data. Replacing the values of the key-factor (k₁) by random values drawn from a normal distribution (Latto and Hassell) affects the dynamics such that the ability of pupal predation to govern density is improved in the model. Changing mortalities other than the key-factor does not significantly influence the pattern of fluctuations nor the limits of density. Models should leave intact the essentials of the reality under study, while removing distracting elements (Levins 1968). As both the timing of the key-factor, and its correlation with pupal predation are essential features of the winter moth population at Wytham Wood between 1950 and 1968, the model of Latto and Hassell does not apply to this population. By simply changing log₁₀ (eggs/female) it is shown that the power of the density dependence of pupal predation to govern possible trends in density of the winter moth population at Wytham Wood is weak. On the other hand, the model of Latto and Hassell gives insight into the conditions that might favour regulation of numbers. Although the model of Poethke and Kirchberg (1987) preserves more features of the pertinent winter moth population than that of Latto and Hassell (1987) it still deviates in one essential aspect: the succession in time of both the (coupled) mortalities and the deviations from the deterministic density dependence are taken at random. Therefore, also this model is still too far from the field population to be a sound base for the statistical speculation proposed by Poethke and Kirchberg.Communication No. 339 of the Biological Station, Wijster     

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.     

Seed predation by mammals and forest dominance by Quercus oleoides,a tropical lowland oak

Summary Quercus oleoides Cham. and Schlecht is an unusual tree in several respects: it is an oak found in neotropical lowland forests, its distribution is not continuous but ratherdivided into many patches of various sizes, and it is a dominant in all the forests in which it occurs, attaining densities far higher than most species of tropical trees. This density pattern is related to the vulnerability of Q. oleoides acorns to predation by mammals. Observations of agoutis, deer, peccaries, squirrels, pocket mice and other seed consumers in Santa Rosa National Park, Costa Rica, showed that these mammals act only as predators, not dispersers, of Q. oleoides acorns. Experiments which involved placing acorns in deciduous forest where Q. oleoides does not occur, demonstrated that, due to high predation rates, the number of acorns produced by an isolated tree is far too low for adults to replace themselves.In oak forest, on the other hand, where the combined acorn crops of many oaks satiate the seed predators, acorn survivorship until germination is high enough to maintain the population. Furthermore, acorn survivorship in oak forest areas is inversely proportional to the apparent mammal density in those areas. Thus the pattern of forest dominance and patchy distribution is related to positively density-dependent acorn survivorship: where Q. oleoides is the forest dominant, it will survive, but if its density falls to the level typical of tropical trees, it will go locally extinct.     

Habitat effects on second-order predation of the seed predator Harpalus rufipes and implications for weed seedbank management

Seed predators provide a valuable ecosystem service to farmers by reducing densities of weed seeds, and, in turn, densities of weed seedlings they must manage. The predominant invertebrate weed seed predator in Maine, USA, agroecosystems is the carabid beetle Harpalus rufipes DeGeer. Pitfall trapping has shown that H. rufipes prefers sites with vegetative cover to fallow sites, preference speculated to be driven by predator avoidance behavior. To test this hypothesis, ‘second-order predation assays’ were developed, in which live H. rufipes prey were presented to second-order predators. Field experiments were conducted to determine foremost if H. rufipes was subject to second-order predation, and secondly, whether (a) vegetative cover affords H. rufipes protection from second-order predators, and (b) high rates of second-order predation correspond with decreased invertebrate seed predation rates. Two 72-h experiments were conducted (mid August and September 2012) at crop and non-crop sites across a 28 ha diversified farm in Stillwater, ME, USA.Second-order predation was 2.8% per day. Based on images from motion-sensing cameras, H. rufipes’ predators included birds and small mammals. Neither a relationship between second-order predation and vegetative treatment, nor an empirical relationship between second-order predation and invertebrate seed predation were detected. However, a simulation model predicted that 2.8% per day second-order predation could increase the number of seeds entering the seedbank by more than 17% annually. Additionally, complex habitats supported higher rates of second-order predation than did simple habitats.     

Post-settlement effects of habitat type and predator size on cannibalism of glaucothoe and juveniles of red king crab Paralithodes camtschaticus

Postlarval (glaucothoe) and juvenile (first crab stage, C1) red king crab Paralithodes camtschaticus actively select structurally complex substrata for settlement. Such habitats may provide them with shelter from predation during critical early stages. We tested this hypothesis by placing glaucothoe and juvenile crab in aquaria with or without natural or artificial habitats, and with or without predators (1-3-year-old red king crab) of two different sizes. Predators caused increased mortality of glaucothoe, but predator size, habitat presence and habitat type had no effect on survival. Predators caused significant mortality of C1 crabs in the absence of habitat, and mortality was inversely related to predator size. Density of glaucothoe on habitats was similar with or without predators, but density of C1 crab on habitats was higher than that of glaucothoe, and increased in the presence of large predators. Active selection for complex substrata by settling glaucothoe does not reduce cannibalism, but may pre-position them for improved survival after metamorphosis. In contrast, juvenile crabs modify their behavior to achieve higher densities in refuge habitats, which tends to dampen the effect of predation. These survival strategies may have evolved to compensate for the greater risk of predation in open habitats.