Population regulation in the lime aphid

The results from a long-term field study (10 years) and laboratory experiments have been used to develop a simulation model of lime aphid populations. This has revealed that the numbers of this aphid are regulated by an interaction between predation and aphid flight. These normally pre-empt regulation by aphid induced changes in plant quality causing increased flight and mortality later in the year.     

Interaction of a population of the black-kneed capsid, Blepharidopterus angulatus, and its prey, the lime aphid

Abstract.

• 1 The population dynamics of Blepharidopterus angulatus (Heteroptera: Miridae), preying on the lime aphid, were investigated from 1965 to 1974. Numbers increased or remained stable until 1969, then declined, at least intermittently. K-factor analysis showed that the key factor was k₅₊₆(loss of adult females, eggs and newly-hatched nymphs).
• 2 Detailed study of numbers and mortalities from 1968 to 1971 showed that loss of adult females was the largest part of k₅₊₆. Studies of flight activity showed most females probably emigrated from the trees before laying eggs, but k₅₊₆ was not related to aphid numbers at the time of peak migration. However, variation in k₅₊₆ depended on the length of time that aphid numbers remained favourable for egg-laying.
• 3 Because of the consistently high emigration of adult females, B.anguhtus numbers increased on average by only 10% when aphid numbers favoured egg-laying. Nett emigration was usually high, probably because the trees were isolated or in rows, with few sources of immigrants to the South-West (direction of prevailing wind). Since the population declined by 85% in years when aphids favoured egg-laying for ≤ 20 days, recovery would be slow after a bad year. Since 1969, 1971 and 1973 were all bad years, this explains the intermittent decline in B.angulatus numbers after 1969.
• 4 Simulation showed that neither increased numbers nor faster population growth of B.angulatus would enhance stability of predator and prey populations, but would lead to over-exploitation of the latter. Given its low observed rate of increase, it is unlikely that B.angulatus could persist on lime at levels much above those recorded.

     

Two‐year oscillation cycle in abundance of soybean aphid in Indiana

• 1 The present study evaluated the population dynamics of the heteroecious soybean aphid Aphis glycines Matsumura (Hemiptera: Aphididae) during an 8‐year period in Indiana, shortly after its detection in North America. Sampling conducted at multiple locations revealed that A. glycines exhibited a 2‐year oscillation cycle that repeated itself four times between 2001 and 2008: years of low aphid abundance were consistently followed by years of high aphid abundance.
• 2 Similar patterns of abundance of A. glycines and coccinellids (Coleoptera: Coccinellidae) in soybean fields, both within and between‐years, suggest that late season predation by coccinellids plays a role in the oscillatory cycle of aphids. Insidious flower bugs Orius insidiosus (Say) (Hemiptera: Anthocoridae) were numerically more abundant than coccinellids, although the lack of synchrony between aphids and predatory bugs suggests that O. insidiosus has a limited influence on between‐year variations in aphid density.
• 3 The inverse relationship between aphid densities before and after the start of the autumn migratory period changes direction in alternate years. High aphid density on soybean in the summer is associated with a reduced number of alate migrants produced in the autumn. Conversely, years with low density aphids on soybean in the summer are characterized by high numbers of alates that migrate to the primary host in the autumn.
• 4 From a pest management perspective, the 2‐year oscillation cycle of A. glycines is a desirable attribute with respect to population dynamics because it implies that aphids cause significant economic damage only in alternate years (as opposed to every year). Cultural practices enhancing the conservation biological control of Coccinellidae may help to preserve the periodicity of aphid infestation and restrict the pest status of A. glycines.

     

The population dynamics of the large pine aphid,Cinara pinea (Mordv.)

The large pine aphid, Cinara pinea lives exclusively on Pinus species, where it feeds on the foliated shoots of the current and previous year. The paper describes the development of a computer model designed to simulate the aphid's population dynamics on saplings in the controlled environment of the laboratory, i.e. in the absence of natural enemies. The model was able to account for about 80% of the variation in aphid numbers within and between trees over a three month period. Sensitivity analysis revealed that the number of pine aphids is limited primarily by nymphal emigration, the operation of which is sensitive both to density and to plant quality as reflected in aphid growth rates. Of secondary importance are changes in reproduction acting through increased reproductive delay, again a result of altered growth rates and adult size. Development, too, has an important secondary influence. Contrary to expectation and conventional belief, however, alate production proved to be of negligible importance, either in limiting or regulating population numbers. Alatae are produced in too few numbers and for too short a period to significantly alter the pattern of population change.     

The role of food quality and competition in shaping the seasonal cycle in the reproductive activity of the sycamore aphid

The hypothesis that seasonal changes in sycamore aphid,Drepanosiphum platanoidis (Schr.), recruitment are determined by changes in food quality and aphid population density was tested. There was no clear association between the reproductive activity of the sycamore aphid and the seasonal changes in specific amino acids or groups of amino acids in extracts of sycamore,Acer pseudoplatanus L., leaves. Seasonal changes in reproductive activity tracked the changes in total amino acids of the leaf tissue of the host, but with a short time delay. High numbers of adult aphids appeared to depress reproductive activity. A regression analysis of the results revealed that total amino acids the previous week and current numbers of adult aphids significantly affected sycamore aphid reproductive activity. The results of this analysis support the above hypothesis, that the marked seasonal changes in the total quantity of amino acids in sycamore leaves and intraspecific competition for this resource, through its effect on adult weight, shape the seasonal cycle in the reproductive activity of the sycamore aphid.     

Effects of plant spacing and interplanting with oilseed rape on colonisation of dwarf hops by the damson-hop aphid, Phorodon humuli

The pattern of colonisation of dwarf hops (Humulus lupulus) by damson-hop aphid (Phorodon humuli (Schrank)) migrating from Prunus spp. was investigated at six plant spacings and where some of the hops were replaced by oilseed rape (Brassica napus L.), a non-host of the aphid. The number of migrant aphids that accumulated on hop stems (bines) increased with increasing bine size and density. The numbers of aphids that colonised hops interplanted with oilseed rape reflected the density of the hop plants only and not the overall plant density. As the physical size of the wind shadow within which flying aphids can manoeuvre and land was unimportant unless provided by a host plant, the finding supports the theory that flying aphids respond to olfactory stimuli associated with their hosts. Variation in bine height (as a measure of plant size) explained 29–93% of the variance in aphid counts during the 3 years' study and bine density 1–14%. Standardising the data as the numbers of aphids per metre of bine and taking a square-root transformation of these standardised counts improved the precision of the analyses and, by stabilising variances, facilitated comparisons between years as growth became more vigorous as the plants matured. Each year, the rate of increase in numbers of aphids settling on plots of hops declined curvilinearly with increasing bine density. Maximum colonisation by P. humuli occurred at a bine density of five per metre row, a density similar to that used commercially by growers of dwarf hops.     

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.     

The population ecology of the Cinnabar Moth,Tyria jacobaeae L. (Lepidoptera,Arctiidae)

Summary This paper describes the results of a study of the factors determining the abundance and distribution of the Cinnabar Moth in Britain. The main part of the study was on a population of the moth at Weeting Heath in Norfolk. This is an area of sandy heath which is heavily overgrazed by rabbits. Here the moth fluctuates violently in number and periodically it completely defoliates its food plant (ragwort, Senecio jacobaea L.) over large areas. This happened in 1960, 1961, 1967, and 1968.Since 1966, the numbers of the moth have been studied in detail and Life Tables are presented for five years.Adult fecundity varies considerably between years. This is due mainly to fluctuations in adult size resulting from changes in larval density. By comparison adult mortality and dispersal have little effect on fecundity; although there is some evidence to suggest that dispersal is density dependent. Because the number of eggs laid in one generation depends on the size of the previous generation, fecundity tends to be acting as a delayed density dependent factor.Mortality is low during the egg stage, but is high amongst young larvae, due mainly to arthropod predation. As the caterpillars grow they become immune from this predation. They are distasteful to vertebrate predators.A larval parasite, Apanteles popularis, kills many of the fully grown larvae. The highest rates of parasitism have coincided with the lowest densities of the moth, however, so that over the five years it has acted as an inverse density dependent factor.In 1967, the population became so large that the ragwort was completely defoliated, and about 20% of the caterpillars died from starvation. In the following year the population was still large and the ragwort plants were small, owing to the effects of defoliation in the previous year. Food ran out early in the season and about 50% of the larvae starved. Because of the overriding effect of starvation, total larval mortality tends to be density dependent.Mortality is high at, or immediately after, pupation and this is thought to be due primarily to predation by moles (Talpa europaea). Pupal mortality does not appear to be density dependent.The upward growth of the population is limited by food supply. Starvation led to a population crash, so that in 1969 only 62 eggs/150 m² were laid compared with 17110 and 16493 in the previous two years. The rate of recovery of the population after this crash was dependent upon the rapid recovery of the ragwort plants. Owing to the wet summer in 1968, plant numbers actually increased after defoliation, due to regeneration from root buds. This was in spite of the fact that no seed was produced in the area in either 1967 or 1968.The only factor which appears to buffer the population against extinction in years when food runs out early in the season, is the heterogeneity within the moth and ragwort populations. The earliest individuals manage to obtain sufficient food in those patches of ragwort which survive longest.Large fluctuations in number only occur in some localities. Other populations of the moth persist at low density and never eat out their food supply. Some data are presented from such a population at Monks Wood. This site is on a heavy clay soil, rabbit grazing is less marked than at Weeting, and ragwort occurs only at a low density. The lusher vegetation supports a very large population of arthropod predators and these take a higher percentage of the young caterpillars than was found at Weeting. Pupal survival is also low due probably to waterlogging of the soil. Pupae can withstand considerable desiccation, but excessive moisture soon leads to their death.The distribution of the moth in Britain and its use for the biological control of ragwort are discussed.     

Frequency‐dependent selection acting on the widely fluctuating sex ratio of the aphid Prociphilus oriens

Frequency‐dependent selection is a fundamental principle of adaptive sex ratio evolution in all sex ratio theories but has rarely been detected in the wild. Through long‐term censuses, we confirmed large fluctuations in the population sex ratio of the aphid Prociphilus oriens and detected frequency‐dependent selection acting on these fluctuations. Fluctuations in the population sex ratio were partly attributable to climatic factors during the growing season. Climatic factors likely affected the growth conditions of host plants, which in turn led to yearly fluctuations in maternal conditions and sex ratios. In the process of frequency‐dependent selection, female proportion higher or lower than ca. 60% was associated with a reduction or increase in female proportion, respectively, the next year. The rearing of aphid clones in the laboratory indicated that mothers of each clone produced an increasing number of females as maternal size increased. However, the mean male number was not related to maternal size, but varied largely among clones. Given genetic variance in the ability to produce males among clones, selection should favour clones that can produce more numerous males in years with a high female proportion. Population‐level sex allocation to females was on average 71%–73% for three localities and more female‐biased when maternal conditions were better. This tendency was accounted for by the hypothesis of competition among foundresses rather than the hypothesis of local mate competition. We conclude that despite consistent operation of frequency‐dependent selection, the sex ratio continues to fluctuate because environmental conditions always push it away from equilibrium.     

Interactive effects of host resistance and drought stress on the performance of a gall-making aphid living on Norway spruce

Four full-sib families of Norway spruce (Picea abies), of which two were resistant to a galling aphid (Adelges abietis) and two were susceptible, were exposed to drought stress for 2 years. The primary aim was to test the hypothesis that the various life stages of the aphid differ in their response to environmentally induced changes in the host plant. The drought treatment had a significant negative effect on tree growth. This was reflected in gall size which responded in a similar way to the drought stress, thus supporting the plant vigour hypothesis. Drought affected the survival of aphid stem-mothers negatively in susceptible trees but positively in resistant trees. This result was matched by the response of an individual phenolic compound which, contrary to the total phenolic concentration, tended to increase in susceptible trees exposed to drought and decrease in similarly exposed resistant trees. Thus it is possible that this single, as yet unidentified, phenolic compound could be used as a marker of resistance. The performance of surviving stem-mothers, evaluated by measuring the diameter of the wax cover they produced (a correlate of fecundity), was not significantly affected by drought, but aphids on susceptible trees produced more wax than those on resistant trees. The change in gall density over time (analysed separately for each treatment and phenotype) correlated best with patterns of stem-mother survival. This indicates that stem-mother survival, which is closely linked to host plant quality and resistance, may play a significant role in the population dynamics of this aphid. However, drought-stress-induced changes in host plant quality affected survival, and hence gall density, less than the genetically determined level of resistance. The results also support the hypothesis that an insect may respond differently to environmentally induced changes in the host plant at different stages in its development. Received: 15 September 1999 / Accepted: 3 December 1999     

Collembola respond to aphid herbivory but not to honeydew addition

Abstract. 1. Euedaphic collembola alter their soil distribution in response to above‐ground aphid herbivory of Poa annua L. Graminae, a host grass. 2. Two mechanisms potentially underpin this effect. Carbon‐rich aphid honeydew falling onto the soil surface may affect mycophagous collembola; alternatively aphid‐induced changes in root biomass may be necessary to produce changes in collembola abundance. 3. When compared to a plant‐only control, aphid herbivory increased the number of collembola in the top 5 cm of soil, reduced both foliar and root biomass, and increased shoot/root ratio. Honeydew addition had no effect on collembola numbers or any recorded host‐plant parameter. 4. Honeydew deposition is not responsible for the increased numbers of collembola found in the upper soil after aphid herbivory; aphid‐induced reductions in root biomass may be the most important factor explaining knock‐on effects of aphid herbivory on soil fauna.     

Climate change and cereal aphids: the relative effects of increasing CO2 and temperature on aphid population dynamics

Experimental evidence regarding the responses of cereal aphids to rising atmospheric CO₂ has been ambiguous. Some studies suggest increased population sizes under future CO₂ levels, others suggest decreased population sizes, and still others suggest little or no difference. Recently, Newman et al. (2003) constructed a general mathematical model of the aphid–grass interaction to investigate whether or not we should, in fact, expect a general aphid response to rising CO₂. They concluded that aphid populations are likely to be larger under future CO₂ concentrations if soil N levels are high, the aphid species' nitrogen requirement is low and the aphid species' density‐dependent response in winged morph production is weak. In that model, and in field experiments, CO₂ concentration influences aphid population dynamics through the effect it has on plant quality. However, future CO₂ concentrations are also likely to be accompanied by higher ambient temperatures, a combination that has received little focus to date. In the present paper, the Newman et al. model is used to consider the combined effects of increased CO₂ concentrations and temperature on aphid population sizes. It is concluded that, when both factors are elevated, aphid population dynamics will be more similar to current ambient conditions than expected from the results of experiments studying either factor alone. This result has important implications for future experimentation.     

Aggregation of polyphagous predators in response to multiple prey: ladybirds (Coleoptera: Coccinellidae) foraging in alfalfa

The spatial distribution of polyphagous predators may often reflect the integration of aggregative responses to local densities of multiple species of prey, and as such may have consequences for the indirect linkages among the prey sharing these predators. In a factorial field experiment in which we manipulated local prey densities within a field of alfalfa in Utah (USA), we tested whether aphidophagous ladybirds would aggregate not only in response to their primary aphid prey, but also in response to an abundant alternative prey, the alfalfa weevil (Hypera postica [Gyllenhal]). Native North American ladybirds (primarily Hippodamia convergens Guerin and H. quinquesignata quinquesignata [Kirby]) responded only to spatial variation in aphid density. In contrast, the introduced ladybird, Coccinella septempunctata L., aggregated also at local concentrations of the weevil late in the experiment when weevil density was high and aphid density was relatively low throughout all experimental plots. The results support the hypothesis that C. septempunctata is more responsive than are native ladybirds to the availability of alternative prey in alfalfa, which may account in part for the displacement of native ladybirds from alfalfa by the introduced species as aphid numbers have declined. The differing responses of the native and introduced ladybirds to spatial patterns of the alternative prey underscore the importance of extending the study of predator aggregation to understand better how polyphagous predators distribute themselves in response to spatial patterns of multiple species of potential prey.     

Assessment of the relative impact of different natural enemies on population dynamics of the grain aphid Sitobion avenae in the field

1. A detailed population dynamics model was devised to provide a tool for integrated pest management against the cereal aphid Sitobion avenae on winter wheat. 2. This model allowed investigation of the relative impact of different natural enemies on aphid population dynamics. 3. The output of the model was compared with a set of data collected in the western part of France from 1976 to 1986. 4. Fungal diseases accounted for 75% of the reduction in peak aphid density and were the key factor acting on aphid dynamics in this region. 5. This study highlights the importance of detailed population dynamics modelling of keystone species, like aphids, for elucidation of the relations between the keystone species and other species associated in the ecosystem.     

The effects of host-plant properties on gall density,gall weight and clone size in the aphidAploneura lentisci (Pass.) (Aphididae: Fordinae) in Israel

Summary Samples of shoots ofPistacia lentiscus carrying galls of the aphid,Aploneura lentisci, were collected at three localities in Israel. Shoots growing near pruning scars carried more galls than elsewhere on the plant, but these galls weighed less and contained fewer aphids (smaller clones). The proportion of empty galls increased with gall density. Crowding of galls at such sites may be due to the early burst of buds at the time of aphid emergence from the overwintering eggs, and not to active search for preferred sites. Shoots bearing larger numbers of leaves carried heavier galls, which contained larger aphid clones. The position of the galled leaf on the shoot had no effect on gall weight nor on clone size. The physiological condition of the plant may be an important environmental (ecological) factor affecting the variation in clone-size and in aphid morphology among galls.     

Properties and isolates of barley yellow dwarf virus

Barley yellow dwarf virus is persistently transmitted by a number of aphid species of which three, Rhopalosiphum padi, Sitobion avenae and Metopolophium dirhodum, are common in most years. Other aphids may be locally important. Isolates of the virus differ in their virulence and geographical distribution and are not transmitted equally well by all aphid vectors. Isolates with similar properties are grouped into strains according to their transmission by vectors and their severity. Changes in strain and aphid occurrence from year to year alter the incidence of virus and its effect on yield. These changes emphasize the need for detailed knowledge of cereal aphid biology and epidemiology of BYDV before effective control can be used.     

Predation on Mindarus abietinus infestingbalsam fir grown as Christmas trees: the impact ofcoccinellid larval predation with emphasis on Anatis mali

The impact of natural coccinellid larvalpredation on the balsam twig aphid was evaluated bysystematically removing coccinellid egg masses in a6–8 year-old balsam fir (Abies balsamea)Christmas tree plantation in southwesternQuebec. Among coccinellid species hunting on firfoliage during development of Mindarus abietinusfundatrices in May, the indigenous Anatis mali was by far the most abundant and themain one to oviposit on trees. Comparison of trees onwhich coccinellid larval predation was excluded withcontrol trees showed that A. mali had a markedimpact both during and after the phase of rapid M. abietinus population growth that followedfundatrix maturation. On trees where coccinellidlarvae were allowed, aphid colonies became inactive(i.e. no live aphids in the colony) about two weeksearlier than on controls. A strong dampening effect onaphid density was also observed in those colonies thatremained active until the end of the aphid life cycle.Predation on aphid colonies reduced sexualsproduction, as the density of M. abietinusoverwintering eggs per shoot subsequently was reducedby 32%. Predation by coccinellid larvae occurred toolate to prevent needle damage to current year shoots,which affects the aesthetic value of Christmas trees.However, current year shoots measured in the mid-crownof trees late in the season were 19% longer on treeswhere aphid predation by coccinellid larvae wasallowed, compared with trees where they were excluded.Rearing all larval stages of A. mali on 4thinstar and adult sexuparae of M. abietinusindicated an average consumption of 269 aphids tocomplete larval development and pupate, which wasequivalent to at least seven colonies of M.abietinus at maximum aphid density at theexperimental site. Anatis mali is an importantnatural control factor of balsam twig aphid inChristmas tree plantations, hence its activity shouldbe protected and possibly stimulated by favourablepest management practices.     

Responses of summer cereal aphid populations to reduced rate aphicide applications in field plots of winter wheat

Abstract 1 Recommended and reduced rate applications of pirimicarb and alpha‐cypermethrin were applied to winter wheat crops to control summer infestations of grain aphid (Sitobion avenae) and rose‐grain aphid (Metopolophium dirhodum). 2 Aphid numbers were assessed weekly and the yield response to treatment application was compared with accumulated aphid days on the crop. 3 Responses to aphicide treatment varied between sites according to variations in the subsequent development of aphid populations under varying weather conditions and differential pressures from aphid natural enemies. 4 Alpha‐cypermethrin treatment reduced spider density at most sites, and also resulted in a resurgence of aphid populations at three sites.     

The role of refuge areas in the phenology of Rhopalosiphum padi in low rainfall cropping areas of South Australia

Extensive surveys of possible aphid habitats in South Australia indicated that irrigated perennial grass pastures in the Mount Lofty Ranges and Lower Murray Valley were summer refuges for Rhopalosiphum padi (L.) (Hemiptera: Aphididae). Large numbers of aphids build up in these pastures each year during autumn (April and May) with numbers peaking in May. The size of the May peak was related to the number of aphids surviving the summer. The proportions of alates were highest in May and August/September. Both peaks coincided with a photoperiod of between 11.2 and 11.5 h, and partial correlations suggested that aphid density, photoperiod and temperature were all significant determinants of alate production.     

The effects of nutrition and density on the production of alate Elatobium abietinum on Sitka spruce

Summary Observations over a period of 10 years showed that, in Northeastern Scotland, alatae of E. abietinum regularly appeared in mid-May, the timing being unrelated to aphid density. The peak number of alatae produced was, however, correlated with aphid density. Following an initially high level the proportion of alatae dropped to virtually nil by mid-June, whilst over the same period the aphid population density increased. Amino acid levels in spruce needles were considerably higher during the period of alate formation than they were at the termination of alate production. It is suggested that a high amino acid level was the main factor controlling the formation of alatae and that population density affected the proportions of these alatae only when nutritional levels were favourable for alate formation.