Relative importance of density-dependent regulation and environmental stochasticity for butterfly population dynamics

The relative contribution of density-dependent regulation and environmental stochasticity to the temporal dynamics of animal populations is one of the central issues of ecology. In insects, the primary role of the latter factor, typically represented by weather patterns, is widely accepted. We have evaluated the impact of density dependence as well as density-independent factors, including weather and mowing regime, on annual fluctuations of butterfly populations. As model species, we used Maculinea alcon and M. teleius living in sympatry and, consequently, we also analysed the effect of their potential competition. Density dependence alone explained 62 and 42% of the variation in the year-to-year trends of M. alcon and M. teleius, respectively. The cumulative Akaike weight of models with density dependence, which can be interpreted as the probability that this factor should be contained in the most appropriate population dynamics model, exceeded 0.97 for both species. In contrast, the impacts of inter-specific competition, mowing regime and weather were much weaker, with their cumulative weights being in the range of 0.08–0.21; in addition, each of these factors explained only 2–5% of additional variation in Maculinea population trends. Our results provide strong evidence for density-dependent regulation in Maculinea, while the influence of environmental stochasticity is rather minor. In the light of several recent studies on other butterflies that detected significant density-dependent effects, it would appear that density-dependent regulation may be more widespread in this group than previously thought, while the role of environmental stochasticity has probably been overestimated. We suggest that this misconception is the result of deficiencies in the design of most butterfly population studies in the past, including (1) a strong focus on adults and a neglect of the larval stage in which density-dependent effects are most likely to occur; (2) an almost exclusive reliance on transect count results that may confound the impact of environmental stochasticity on butterfly numbers with its impact on adult longevity. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Differing contributions of density dependence and climate to the population dynamics of three eruptive herbivores

1. Although both endogenous and exogenous processes regulate populations, the current understanding of the contributions from density dependence and climate to the population dynamics of eruptive herbivores remains limited. 2. Using a 17‐year time series of three cereal aphid species [Rhopalosiphum padi L., Metopolophium dirhodum (Walker), and Diuraphis noxia (Kurdumov)] compiled from a trapping network spanning the northwestern U.S.A., temporal and spatial patterns associated with population fluctuations, and modelled density dependence in aphid abundances were tested. These models were used to analyse correlations between climate and aphid abundances in the presence and absence of residual variance as a result of density‐dependent effects. 3. The temporal dynamics of aphid population fluctuations indicated periodicity, with no clear evidence for a spatial pattern underlying population fluctuations. 4. Aphid abundances oscillated in a manner consistent with delayed density dependence for all three aphid species, although the strength of these feedbacks differed among species. 5. Diuraphis noxia abundances were negatively correlated with increasing temperatures in the absence of density‐dependent effects, whereas M. dirhodum abundances were positively correlated with increasing cumulative precipitation in the presence of density‐dependent effects; yet, R. padi abundances were unrelated to climate variables irrespective of population feedbacks. 6. Our analysis suggests that endogenous feedbacks differentially regulate aphid populations in the northwestern U.S.A., and these feedbacks may operate at an expansive spatial scale. It is concluded that the contributions of density dependence and climate to aphid population dynamics are species‐specific in spite of similar ecological niches, with implications for assessing species responses to climate variability.     

Can polyphagous predators control the bird cherry-oat aphid (Rhopalosiphum padi) in spring cereals?: A simulation study

Predation by two carabid species,Bembidion lampros (Herbst.) andPterostichus cupreus L., on the cereal aphid,Rhopalsiphum padi L., was described using a discrete event simulation model. The results were incorporated into an aphid population growth model. The aphid model was run using actual temperatures,R. padi emigration rates, field densities and distributions from three years with distinctly different aphid population levels. Predation was incorporated at different time periods. Predation during the arrival of emigrants and establishment of aphid populations had approximately the same effect as predation over the entire aphid period, whereas predation during the 20 days prior to peak had very little effect on resulting peak populations. We conclude that a) yearly levels ofR. padi infestation are mainly determined by the volume and intensity of emigrants; b) peak levels are sensitive to changes in predator density; and c) in some cases an outbreak ofR. padi may be prevented by the action of spring active polyphagous predators alone.     

Survival, growth, and reproduction of two aphid species on sucrose solutions containing host or non-host honeydews

Aphids, like most phloem-feeding insects, commonly exhibit a high degree of host specificity. Plant-specific chemical compounds are likely to serve as important host selection cues for monophagous aphids and such substances could be present in aphid honeydew. Apterous virginoparae ofMyzus persicae (Sulzer) andPhorodon humuli (Schrank) were reared on a buffered sucrose solution containing various aphid honeydews or a mixture of amino acids. In two separate experiments, the host-specificP. humuli (hop aphid) could grow and reproduce only on diets containing honeydew collected from hop (Humulus lupulus L.).M. persicae (the green peach aphid, GPA) did not perform well on diets containing hop honeydew, perhaps because hop is a poor GPA host. Honeydew collected from preferred GPA host plants rape,Brassica napus L., and jimsonweed,Datura stramonium L., allowed GPA growth and reproduction. Hop aphids, however, performed poorly on rape and jimsonweed honeydew diets. Bell pepper,Capsicum annuum L., honeydew supported neither the hop aphid nor GPA. The study of aphid honeydew components may contribute towards a more complete understanding of host preference and selection phenomena in aphids.     

Estimating susceptibility of biological control agents to pesticides: influence of life history strategies and population structure

In this study we examined the influences that differing life history strategies and population structures at the time of pesticide exposure have on population susceptibility to pesticides. We used life table data and a matrix projection model to incorporate combinations of mortality (lethal effect) and reductions in fecundity (sublethal effect) into estimates of intrinsic population growth rates (r) for a predator, the seven-spot lady beetle, Coccinella septempunctata L., and its prey, the pea aphid, Acyrthosiphon pisum Harris, and an aphid parasitoid, Diaeretiella rapae (M’Intosh). All three species exhibited differences in key life history variables. The aphid had the highest r and shortest generation time, the ladybeetle had the lowest r and longest generation, while the parasitoid exhibited intermediate life history characteristics. When the model was run with populations started as neonates (aphids) or eggs (lady beetle, parasitoid) for each species, ladybeetle populations were much more susceptible than either aphid or parasitoid populations 30 days after simulated exposure to a pesticide. For example, 50% mortality and a 50% reduction in fecundity resulted in a population headed toward extinction (negative r) for the ladybeetle while the parasitoid population grew exponentially (positive r) even after sustaining 70% mortality and a 70% reduction in fecundity. The aphid species maintained exponential growth after sustaining 80% mortality and an 80% reduction in fecundity. Thus, differences in life history variables accounted for the greater susceptibility of the ladybeetle to a pesticide than its aphid prey or the parasitoid over a set time interval. These differences in susceptibility were greatly reduced when the model was run starting with a mixed age/stage population (the stable age distribution) for each species indicating that population structure at the time of pesticide exposure plays a critical role in population susceptibility. These results suggest that life history attributes as well as population structure at the time of pesticide exposure both play a major role in population susceptibility to pesticides, highlighting the need to explicitly consider differences in life history variables among species when calculating compatibility of pesticides and biological control agents as well as the population structure of beneficial species at the time of pesticide application.     

Density-dependent mortality in Pinus sylvestris caused by the snow blight pathogen Phacidium infestans

Summary Relationships between disease incidence and the density of host plant populations were investigated in the Pinus sylvestris-Phacidium infestans host-fungal pathogen association, in which the season of death of plants killed up to 3 years previously could be accurately determined. Significant (P<0.05), positive density-dependent relationships between the proportion of plants dying in the winters of 1987–1988, 1988–1989 or 1989–1990 and the original stand density were detected in 12 of 26 comparisons. Of the remaining comparisons, all but three had positive regression coefficients for the same association. Plants killed up to 2 years previously contributed to inoculum production. The use of standing dead as a predictor in the analyses showed that the proportion of plants dying in the winters of 1988–1989 or 1989–1990 was generally better correlated with standing dead in the previous summer than with the density of the original population. Significant (P<0.05), positive density-dependent associations were also found between the proportion of living plants in 1990 infected with P. infestans and the number of standing dead plants in all nine comparisons. In contrast, only four of the nine associations between these proportions of infected plants and population density were significant. The strength of the density-dependent relationships varied substantially within and between sites. Much of this variation appears to be due to differences in the stage of development of the epidemics occurring at different sites.     

A time series and geographical analysis of population dynamics of the red-backed vole in Hokkaido,Japan

Summary A time series and geographical analysis of the long term (25-years and 16-years) census data of the red-backed vole, Clethrionomys rufocanus bedfordiae, populations in Hokkaido, Japan was carried out. Eighty seven populations from all over Hokkaido were grouped into the following 4 types according to their flucturation patterns: type A: low density-constant type; type B: medium density-gradual increase type; type C: medium density-wavy change type; type D: high density-large amplitude type. The border between type B and type C was not clear. Although the distribution of the fluctuation types did not show any geographical cline, the high density type (type D) occurred in limited areas where the winter was severe and spring came later and almost all the populations in the Oshima peninsula were the low density type (type A). The most common type of population in Hokkaido, type C, greatly fluctuated from year to year with the maximum density level/the minimum one being over 10. The periodicity of the peak years was not statistically significant, although peaks often occurred at 3 or 4 year intervals. The population grew from spring to autumn in almost all years so that the pattern of the seasonal change was qualitatively constant. The population density levels in spring were not significantly different between peak years and others, while those in autumn were greatly different between them, so that the population growth rates from spring to autumn were variable from year to year. This differential pattern was also found between high and low density areas.     

Fungal endosymbionts affect aphid population size by reduction of adult life span and fecundity

The symbiosis between grasses and endophytic fungi is a common phenomenon and can affect herbivore performance through acquired, chemical plant defence by fungal alkaloids. In laboratory experiments, two species of common grass aphids, Rhopalosiphum padi and Metopolophium dirhodum were tested, in a population experiment (on four plant cultivars) and individually (on one plant cultivar) for the effects of the endophyte, Neotyphodium lolii, that forms symbiotic associations with perennial ryegrass Lolium perenne. In the population experiment that lasted for four aphid generations both aphid species showed decreased population sizes when feeding on each of the four endophyte-infected cultivars. Individuals of R. padi tested individually showed reduced adult life span and fecundity when feeding on infected plants. Individuals of M. dirhodum showed no response in any of the traits measured. This suggests that R. padi individuals are more sensitive to endophyte infection than M. dirhodum individuals. However, all infected grass cultivars reduced population sizes of both aphid species over four generations. Therefore, fungal endophytes can reduce populations of aphid herbivores independent of plant cultivars.     

Negative dietary effects of Colorado potato beetle eggs for the larvae of native and introduced ladybird beetles

Aphidophagous ladybird beetles (Coleoptera: Coccinellidae) are attracted to and feed heavily on aphids, but many species will also feed opportunistically on other prey that they encounter. In potatoes (Solanum tuberosum L.) in Washington State, USA, coccinellids feed on both green peach aphids (“GPA,” Myzus persicae Sulzer) and eggs of the Colorado potato beetle (“CPB,” Leptinotarsa decemlineata Say). The guild of aphidophagous ladybirds includes two native species, Hippodamia convergens Guérin-Méneville and Coccinella transversoguttata Brown. Recently, an introduced species, Coccinella septempunctata L., has invaded and apparently displaced its native congener. A second exotic, Harmonia axyridis Pallas, has colonized the area and is becoming more abundant. We compared larval development of each species on a monotypic diet of GPA, a monotypic diet of CPB eggs, or a mixed diet of both GPA and CPB eggs. Our goal was to answer two questions: (1) do larvae of the four ladybird species benefit from including CPB eggs in their diet and (2) do the four ladybird species differ in their ability to utilize CPB eggs as prey? No larva of any species completed development on a pure diet of CPB eggs, and survivorship was highest for all species when they fed on a pure diet of GPA. One native species, H. convergens, and one exotic species, H. axyridis, exhibited significantly lower survivorship on a mixed diet of both CPB eggs and GPA, compared to a pure GPA diet; H. axyridis also took longer to develop from egg to adult when both prey were provided. Survivorship of the two Coccinella spp. was not altered by the inclusion of CPB eggs with GPA, although CPB eggs lengthened the development time of C. transversoguttata. Adult size was not consistently affected by diet for any of the coccinellids. Overall, no ladybird species benefited from the inclusion of potato beetle eggs in its diet. The two Coccinella species responded similarly to the inclusion of CPB eggs, and so we would not expect any difference in the success of coccinellid larval development in potato fields following the replacement of C. transversoguttata by C. septempunctata. Hippodamia convergens and H. axyridis, the two species whose survivorship was depressed by combining CPB egg and aphid prey, were also the two species that consumed the greatest number of CPB eggs during successful larval development. A comparison of total egg consumption by each species cohort suggested that displacement of the other species by H. axyridis would not alter CPB biological control, because the higher per capita feeding rate by H. axyridis larvae compensated for individuals’ greater mortality risk on a diet including CPB eggs.     

Classical biological control of Aphis gossypii (Homoptera: Aphididae) with Neozygites fresenii (Entomophthorales: Neozygitaceae) in California cotton

In August 1994 and 1995 classical biological control releases were made in cotton in the San Joaquin Valley, California, with an Arkansas strain of the entomopathogenic fungus, Neozygites fresenii, a pathogen of the cotton aphid, Aphis gossypii. Pre-release samples in both years indicated that N. fresenii was not naturally present in A. gossypii populations in the San Joaquin Valley. Two release methods were compared: dried N. fresenii-infected cotton aphid “cadavers” and chamber inoculation of A. gossypii. Both methods were successful in introducing N. fresenii to cotton aphids in California; however, higher prevalence of fungal infection resulted with the cadaver treatments. N. fresenii persisted and spread in the aphid population until early October 1994 and late September 1995. The highest mean percentage infection in the cadaver treatment in 1994 reached a level (14%) considered imminent for epizootics (12–15%). The use of predator exclusion cages resulted in higher N. fresenii prevalences.     

Demography in relation to population density in two herbivorous marsupials: testing for source–sink dynamics versus independent regulation of population size

We compared demography of populations along gradients of population density in two medium-sized herbivorous marsupials, the common brushtail possum Trichosurus vulpecula and the rufous bettong Aepyprymnus rufescens, to test for net dispersal from high density populations (acting as sources) to low density populations (sinks). In both species, population density was positively related to soil fertility, and variation in soil fertility produced large differences in population density of contiguous populations. We predicted that if source–sink dynamics were operating over this density gradient, we should find higher immigration rates in low-density populations, and positive relationships of measures of individual fitness—body condition, reproductive output, juvenile growth rates and survivorship—to population density. This was predicted because under source–sink dynamics, immigration from high-density sites would hold population density above carrying capacity in low-density sites. The study included 13 populations of these two species, representing a more than 50-fold range of density for each species, but we found that individual fitness, immigration rates and population turnover were similar in all populations. We conclude that net dispersal from high to low density populations had little influence on population dynamics in these species; rather, all populations appeared to be independently regulated at carrying capacity, with a balanced exchange of dispersers among populations. These two species have suffered recent reductions in range, and they are ecologically similar to other species that have declined to extinction in inland Australia. It has been argued that part of the cause of the vulnerability of species like these is that they exhibit source–sink dynamics, and disturbance to source habitats can therefore cause large-scale population collapses. The results of our study argue against this interpretation.     

Comparative population structure of Chinese sumac aphid <Emphasis Type="Italic">Schlechtendalia chinensis</Emphasis> and its primary host-plant <Emphasis Type="Italic">Rhus chinensis</Emphasis>

Most of our current understanding of comparative population structure has been come from studies of parasite–host systems, whereas the genetic comparison of gallnut-aphids and their host-plants remain poorly documented. Here, we examined the population genetic structure of the Chinese sumac aphid Schlechtendalia chinensis and its unique primary host-plant Rhus chinensis in a mountainous province in western China using inter-simple sequence repeat (ISSR) markers. Despite being sampled from a mountainous geographic range, analysis of molecular variance (AMOVA) showed that the majority of genetic variation occurred among individuals within populations of both the aphid and its host. The aphid populations were found to be structured similarly to their primary host populations (F _ST values were 0.239 for the aphid and 0.209 for its host), suggesting that there are similar patterns of gene flow between the populations of the aphid and between populations of its host-plant. The genetic distances (F _ST/1 − F _ST) between the aphid populations and between its host-plant populations were uncorrelated, indicating that sites with genetically similar host-plant populations may not always have genetically similar aphid populations. The lack of relationships between genetic and geographical distance matrices suggested that isolation by distance (IBD) played a negligible role at this level. This may be mainly attributed to the founder effect, genetic drift and the relative small spatial scale between populations. Zhumei Ren and Bin Zhu contributed equally to this work.     

Comparison of transmission efficiency of various isolates of Potato virus Y among three aphid vectors

Potato virus Y (PVY) strains are transmitted by different aphid species in a non‐persistent, non‐circulative manner. Green peach aphid (GPA), Myzus persicae Sulzer, is the most efficient vector in laboratory studies, but potato aphid (PA), Macrosiphum euphorbiae Thomas (both Hemiptera: Aphididae, Macrosiphini), and bird cherry‐oat aphid (BCOA), Rhopalosiphum padi L. (Hemiptera: Aphididae, Aphidini), also contribute to PVY transmission. Studies were conducted with GPA, PA, and BCOA to assess PVY transmission efficiency for various isolates of the same strain. Treatments included three PVY strains (PVY^O, PVY^N:O, PVY^NTN) and two isolates of each strain (Oz and NY090031 for PVY^O; Alt and NY090004 for PVY^N:O; N4 and NY090029 for PVY^NTN), using each of three aphid species as well as a sham inoculation. Virus‐free tissue‐cultured plantlets of potato cv. Russet Burbank were used as virus source and recipient plants. Five weeks post inoculation, recipient plants were tested with quantitative DAS‐ELISA to assess infection percentage and virus titer. ELISA‐positive recipient plants were assayed with RT‐PCR to confirm presence of the expected strains. Transmission efficiency (percentage infection of plants) was highest for GPA, intermediate for BCOA, and lowest for PA. For all aphid species, transmission efficiency did not differ significantly between isolates within each strain. No correlations were found among source plant titer, infection percentage, and recipient plant titer. For both GPA and BCOA, isolates of PVY^NTN were transmitted with greatest efficiency followed by isolates of PVY^O and PVY^N:O, which might help explain the increasing prevalence of necrotic strains in potato‐growing regions. Bird cherry‐oat aphid transmitted PVY with higher efficiency than previously reported, suggesting that this species is more important to PVY epidemiology than has been considered.     

Ladybird population dynamics in potato: comparison of native species with an invasive species, Harmonia axyridis

Following the detection of the harlequin ladybird, Harmonia axyridis, in 2003 in potato crops in Belgium, a study was carried out between 2004 and 2006 on the phenology of this species compared to native species in potato. The results confirmed the success of H. axyridis, with high population levels in 2004 and 2005. In 2006, aphid populations were very low and no H. axyridis larvae were sampled in potato, but the indigenous species Coccinella septempunctata and Propylea quatuordecimpunctata were detected. A species by species comparison of the date of first larvae detection, the larvae population peak, and the difference between this peak and the aphid population peak was performed. Results showed a clear correlation between C. septempunctata and P. quatuordecimpunctata and potato aphids, with a delay of 3.5 and 6.5 days between the aphid and ladybird population peaks for the two native species. H. axyridis arrived 7–8 days after the two indigenous species and the larval peak population occurred 15.8 days after the aphid population peak. This meant that H. axyridis had to complete its larval development with very low aphid populations or even with no aphids at all. The reason for its late arrival and the possible food resources used by H. axyridis larvae are discussed.     

Population responses of a conifer-dwelling aphid to seasonal changes in its host

Abstract.  1. Current evidence suggests that seasonal changes in spruce needle sap nutrients have a decisive influence on green spruce aphid ( Elatobium abietinum ) population density, but the mechanisms of population change, the roles of development rate, fertility and mortality, and the existence of density-dependent processes, are not clearly understood.
2. Experimental studies of aphid populations were conducted in controlled environments to estimate seasonal patterns in aphid mean relative growth rate, prenatal development, fertility, and mortality. Studies were also made of the effect of aphid crowding on vital rates.
3. Independent of the degree of aphid crowding, seasonal changes in the amino acid concentration of needle sap were tracked by aphid growth rate, fertility (and adult size), but not by rates of aphid mortality. The most pronounced change in vital rates, and the one most likely to drive seasonal population change, was in fertility. Prenatal development time actually became shorter in periods when nutrients were scarce, but the resulting adult aphids were smaller and less fertile than during periods of improved nutrition.
4. Density dependence of vital rates was only observed during mid-summer when nutrients were least available. Mortality, growth rate, and prenatal development were the most strongly density-dependent processes. In contrast, there was no evidence that fertility rates were likely to respond to crowding.
5. There were no important differences between populations reared on small, potted spruce trees and those on plantation trees aged 25 years. This gives confidence that demographic data from a variety of field and laboratory sources could be used to compile data appropriate for population models.     

醉马草水浸液对豌豆蚜触杀活性及种群增长的影响

为探讨醉马草水浸液对豌豆蚜触杀活性及种群增长的影响,采用带虫浸叶法比较不同生育期醉马草带菌（E+）和不带菌（E-）水浸液对豌豆蚜触杀活性及种群生命表,测定了豌豆蚜的死亡率及触杀后对其生殖期,平均繁殖力,繁殖率及生命表参数的影响。结果表明,不同生育期醉马草带菌（E+）水浸液触杀豌豆蚜后对其各项指标均有显著影响。在苗期时,E+水浸液触杀豌豆蚜后校正死亡率最高,繁殖力最低,内禀增长率（r_m=0.145 d^-1）和净生殖率（R₀=4.802头）均为最小值。在成熟期时,E+水浸液触杀豌豆蚜24 h、48 h和72 h后校正死亡率分别为26.15%,19.01%,9.07%;繁殖期（3.87 d）,平均繁殖力（8.80头）,繁殖率（1.40%）,内禀增长率（r_m=0.208 d^-1）和净生殖率（R₀=8.820头）。在枯黄期时,E+水浸液触杀豌豆蚜后校正死亡率最低,繁殖力最强,内禀增长率（r_m=0.247 d^-1）和净生殖率（R₀=13.647头）均为最大值。不同生育期醉马草E-水浸液触杀豌豆蚜后对其种群繁殖无显著影响,与对照差异不显著（P>0.05）。综上,苗期醉马草E+水浸液对豌豆蚜有较好的触杀效果,校正死亡率高,且触杀后当代繁殖力减弱,种群扩建时间延长,不利于其种群繁殖和增长;故苗期醉马草E+水浸液具有很好的杀虫潜力,所采用水浸液方法制备简单,成本低,可为新型植物源农药研发提供重要理论依据。     

Effects of crowding on populations of Aedes albifasciatus larvae under laboratory conditions

Aedes (Ochlerotatus) albifasciatus(Macquart 1838) (Diptera: Culicidae) is a neotropical flood water mosquito, incriminated as the main vector of the western equine encephalitis virus, and which affects beef and milk production in central Argentina. The short time required to hatch and develop from egg to adult, usually in temporary pools, suggests a strategy which allows for exploitation of transient pools, thus evading predation and interspecific competition. Under these conditions intra specific competition could represent a major density-dependent source of larval mortality, but the relative importance of density-dependent regulation of mosquito populations has generated controversy. Therefore we examined the effects of larval density on basic population characteristics of Ae. albifasciatusin the laboratory. Larvae were obtained by synchronous hatching of eggs laid by field-trapped females. Emerging larvae (L1) were used to build cohorts of different initial densities, kept in plastic trays with 400 ml of distilled water, and food supplied daily during the first 10 days (0.1 g per larvae day^–1). Age-specific development time and specific and relative mortality were estimated, and their relation to initial larval density was assessed through linear and non-linear regressions and correlation analysis. First hatching was registered 3 h after flooding the eggs. Higher levels of pre-adult mortality were detected in groups with higher densities. Specific mortality and average time to enter a stage of L1 to L3 could directly be related to initial larval density, but no significant relations were found for L4 and pupae. Results suggest that crowding could be a factor capable of regulating the density of natural populations of Ae. albifasciatus.     

Temporal and spatial variation of larval parasitism in non-outbreaking populations of a folivorous moth

In order to assess the role of parasitoids in the regulation of non-outbreaking populations of Epirrita autumnata, a geometrid lepidopteran with outbreaking populations in northern Europe, we examined the temporal and spatial variation of larval parasitism in southwestern Finland during 6 successive years. The study was carried out on two spatial scales, among trees within sites of about 1 ha and among sites separated by distances of 2–10 km, using experimental and observational approaches respectively. The overall percent parasitism was independent of host density on both spatial scales, while temporally it fluctuated only little. Of the two main parasitoids, the commoner one, Protapanteles immunis, showed a variable response to host density on the larger spatial scale and negative density dependence on the smaller scale. Temporally, parasitism caused by this species was independent of host density. Another parasitoid, Phobocampe bicingulata, showed positive density dependence on the smaller spatial scale and had a variable response on the larger scale, but exhibited negative density dependence over time. The results of this study caution against drawing conclusions concerning population regulation on the grounds of spatial density dependence alone. Larval parasitoids apparently do not maintain low densities in the E. autumnata populations studied. However, they may suppress E. autumnata densities to a level low enough for density-dependent mortality factor(s) to become regulating. Among other mortality factors of E. autumnata, pupal predation has been found to be temporally positively density-dependent. Received: 19 October 1999 / Accepted: 10 January 2000     

Population dynamics of a tree-dwelling aphid: regulation and density-independent processes

1. A population of the Turkey-oak aphid ( Myzocallis boerneri Stroyan) was sampled at approximately weekly intervals on two Turkey-oak trees for 19 years.
2. On one tree (A), the aphids exhibited a distinct seasonal pattern with a spring increase, summer decrease, early autumn increase, and late autumn decline. On the other tree (B) the aphids remained at low densities after the decrease in summer.
3. On tree A, significant undercompensating density dependence occurred during all periods of the seasonal population development, and their strength varied little during the course of the season. On tree B, significant density dependence compensated exactly for increase, but appeared only after the decrease in summer when the population remained at very low densities for the rest of the season.
4. Density-independent weather variables affected the population dynamics very little. Their influence was marginally significant only at very low densities when the aphids were regulated exactly by compensating density-dependent factors.
5. The results suggest a curvilinear density dependence, with strong regulation at low densities, and weak at high densities. That is, this aphid was most regulated not at the peak but at the trough densities.