Prey preferences of six species of Carabidae (Coleoptera) and one Lycosidae (Araneae) commonly found in UK arable crop fields

Abstract: Laboratory experiments were used to determine the feeding preferences of six carabid beetles and one lycosid spider on aphid and collembolan prey. The first investigation used only five species of carabid Pterostichus melanarius Illiger, Pterostichus madidus F., Harpalus rufipes DeGeer, Nebria brevicollis F. and Carabus violaceus L., which had been caught most commonly in pitfall traps in the headland region of arable crop fields. When offered Brevicoryne brassicae L., Sitobion avenae F., Metopolophium dirhodum Walker and Rhopalosiphum padi L. as prey items, the species consistently consumed in high numbers was M. dirhodum . In subsequent experiments four carabids Pterostichus cupreus L., P. melanarius, P. madidus. H. rufipes and a lycosid spider Trochosa ruricola DeGeer whose distribution was shown by pitfall trapping to extend throughout the arable crop, were the chosen predators. These predators were offered a choice between M. dirhodum and entomobryid collembolans (a recognised alternative prey item) to gauge their preference between the two prey types. Both male and female P. cupreus and P. melanarius showed a significant preference for the aphid prey, while there was no significant preference displayed by the other species. The effect of temperature on the voracity of these five predators fed on M. dirhodum was investigated. There were significant differences in the number of aphids consumed by the species and sexes at the different temperatures. Regression analysis on the mean numbers of aphids eaten by each sex of the five predators, showed that in the majority of cases there was a significant increase in predation with increasing temperature. In considering the dietary preferences illustrated by these experiments, it appeared that P. cupreus and P. melanarius offered the greatest potential in controlling aphids on arable crops.     

Monoclonal antibodies reveal changes in predator efficiency with prey spatial pattern

Spatially explicit predator–prey interactions can alter the predatory potential of natural enemies augmented through conservation biological control. To test hypotheses regarding such interactions and predatory efficiency, we used a combination of molecular techniques and mark–release–recapture to study the foraging behaviour of a generalist carabid predator, Poecilus cupreus , in response to spatial patterns of its cereal aphid prey ( Metapolophium dirhodum and Sitobion avenae ). Beetle and aphid numbers were measured across two grids of sampling locations, within which aphid spatial pattern had been manipulated to generate patchy and more homogenous distributions. Aphid consumption was measured by enzyme-linked immunosorbent assays (ELISA) of beetle gut contents, using an aphid-specific monoclonal antibody. Movement and distribution patterns suggest that P. cupreus does not aggregate at, nor instigate prey-taxis within, aphid patches. However, more than two-thirds of the 2169 P. cupreus tested by ELISA had consumed aphids and the proportion of beetles containing aphid proteins was positively related to aphid density. Against expectation, the proportion of predators feeding on aphids was greatest where prey were homogenously distributed, and this was attributed to the loss of partial refuges for prey in aphid patches. The functional value of this type of uniform foraging strategy is ideally suited to early colonization of the crop habitat, when aphid numbers are low, before populations build up and form strong spatial patterns.     

The role of large-scale spatially explicit and small-scale localized processes on the population dynamics of cereal aphids

A field-scale study of the spatially explicit interaction between the carabid Poecilus cupreus Linnaeus, and two common aphid species (Sitobion avenae (Fabricius) and Metopolophium dirhodum (Walker)) in winter wheat was conducted. All three species showed considerable spatial pattern at the field scale. Activity-density of P. cupreus was an order of magnitude higher in the central part of the field compared to its periphery. Where P. cupreus activity-density was highest, S. avenae and M. dirhodum population peaks were delayed. Additionally, in the case of M. dirhodum, lower maximum counts were evident where P. cupreus activity-density was highest. An analysis of the movement of individual P. cupreus using release-recapture indicated that those beetles within the centre of the field exhibited reduced displacement, which may have caused the generation or maintenance of spatial pattern. Crop density was also measured throughout the field. Although crop density had no large-scale spatial pattern, its variability at the small-scale was consistent with an influence on aphid population dynamics. This study demonstrates empirically that both large-scale spatially explicit and small-scale localized processes influenced aphid population dynamics simultaneously.     

Consumption by carabid beetles of three cereal aphid species relative to other prey types

The cereal aphidRhopalosiphum padi has previously been found to be a low quality prey for a range of generalist arthropod predators. The aim of this study was to reveal, using food consumption experiments whether this applies to other cereal aphids. The question of whether predator feeding capacity increased when several aphid species were offered relative to a single aphid species was also addressed by measuring food consumption on a mixed aphid diet relative to single aphid diets. Food consumption by five carabid beetles of the three cereal aphid speciesRhopalosiphum padi, Sitobion avenae andMetopolophium dirhodum was determined relative to fruit fliesDrosophila melanogaster and the collembolanIsotoma anglicana. Feeding rate was measured as food consumption over 24 hour both for previously satiated and beetles starved for 7 days. Generally the largest aphid consumption was ofM. dirhodum and the lowest ofR. padi, withS. avenae in between. The mixed aphid consumption experiments did not reveal a higher feeding rate on mixed aphid diets relative to single aphid diets. The results indicate low preference forR. padi andS. avenae.     

The reactions of two cereal aphid parasitoids, Aphidius uzbekistanicus and A. ervi to host aphids and their food-plants

ABSTRACT. A Y-tube olfactometer was used to test the reactions of the hymenopteran cereal aphid parasitoids Aphidius uzbekistanicus Luzhetski and A. ervi Haliday to odours from aphids and their host plants. Only females responded to aphids but both sexes responded to plant odours. A. uzbekistanicus responded to the cereal aphids Sitobion avenae (F.) and Metopolophium dirhodum (Walker) whilst A. ervi , which has a broad host range, responded to M. dirhodum and the pea aphid, Acyrthosiphon pisum. Female A. uzbekistanicus responded to wheat leaves only but males responded to a range of plant material. Both male and female A. ervi responded to wheat and bean leaves. The failure of A. ervi to respond to either nettle aphids, Microlophium carnosum (Bukt.), or nettle leaves, despite its frequent parasitization of this aphid in the field, suggests the existence of more than one race of the parasitoid and casts doubts on the usefulness of alternative hosts as reservoirs for A. ervi in integrated control programmes. Males of both species responded to their respective females suggesting the presence of a sex specific attractant.     

Predation on the slug Deroceras reticulatum by the carabid beetles Pterostichus madidus and Nebria brevicollis in the presence of alternative prey

1 Slugs are important pests in many agricultural crops and potential biological control agents are being studied as an alternative to molluscicides. Carabid beetles may be able to reduce slug populations, but their role as control agents may be influenced by the presence of alternative prey. 2 Attacks on the pest slug Deroceras reticulatum (Müller) by the carabid beetles Pterostichus madidus (Fabricius) and Nebria brevicollis (Fabricius) were investigated in the presence of alternative prey (earthworms and Calliphora fly larvae). Consumption of slug eggs and aphids was also investigated. 3 All five prey types were consumed to varying degrees during the experiments. Both beetle species showed a significant preference for Calliphora larvae over slugs. Pterostichus madidus showed a significant preference for earthworms over slugs. No preference was shown between earthworms or Calliphora larvae by P. madidus females or N. brevicollis. However, P. madidus males showed a significant preference for Calliphora larvae over worms. Pterostichus madidus showed no preference between slug eggs and aphids; N. brevicollis showed a significant preference for aphids over slug eggs. 4 The results from this study indicate that generalist beetles will often attack other prey in preferences to adult slugs. Slugs may not be preferred because of their mucus. Other prey items occur frequently in arable soils and generalist carabids may ignore slugs altogether and may only feed on them when slug density is high or other prey are unavailable.     

Aspects of host—parasitoid interactions of two aphid parasitoids, Aphidius urticae (Haliday) and Aphidius uzbeckistanicus (Luzhetski) (Hymenoptera, Aphidiidae)

Abstract. 1. Various aspects were studied of two host-parasitoid interactions: Aphidius urticae parasitizing the cocksfoot aphid Hyalopteroides humilis and Aphidius uzbeckistanicus parasitizing the cereal aphid Metopolophmm dirhodum. The work formed part of a study on the population dynamics of grassland and cereal aphids in Berkshire, England.
2. A.uzbeckistanicus had a higher instantaneous attack rate but a slightly longer handling time than A.urticae. In both cases the functional response was curvilinear.
3. A.urticae was specific to H.humilis and failed to develop in M.dirhodum. A.uzbeckistanicus could develop successfully in H.humilis but the functional response was sigmoid and the progeny were less fecund than those reared on the natural host. This species showed a clear preference for M.dirhodum when supplied with mixtures of M.dirhodum and H.humilis.
4. A.uzbeckistanicus had a faster rate of development and a lower developmental threshold than A.urticae , a situation paralleled by their natural hosts.
5. The results are discussed in relation to field observations. A.uzbeckistanicus may have considerable potential for reducing cereal aphid numbers. However, its preference for M.dirhodum mitigates against a reservoir being maintained in the economically unimportant grassland aphid, H.humilis.     

Prey preference and egg production of the carabid beetleAgonum dorsale

In order to elucidate whether the aphidRhopalosiphum padi is low quality food for the carabid beetleAgonum dorsale, as it has previously been found to be for cereal spiders, we performed a series of experiments using fruit fliesDrosophila melanogaster as a standard alternative prey (‘control prey’): 1) Prey preference, 2) Aphid consumption for satiated and starved beetles, 3) Egg production on different diets and 4) Preference experiments with fruit flies coated with a taste of different prey types. Other alternative prey types used were earthworms and sciarid midges, all potential prey in the field.A. dorsale has a low preference for aphids compared to fruit flies. Apparently there is an upper limit to the consumption of aphids independent of hunger and much below the beetles' total food demand. Egg production on a pure diet of aphids is lower than on a pure diet of fruit flies; it is extremely low on a pure diet of earthworms; the highest fecundity is found on a mixed diet. As fruit flies coated with a taste of aphids or sciarid midges are less prefered than flies covered with a taste of fruit flies there may be a chemical factor, acting through taste, involved in determining prey preference.     

Potential cotton aphid,Aphis gossypii,population suppression by arthropod predators in upland cotton

The cotton aphid, Aphis gossypii Glover, predation rate of convergent lady beetle, Hippodamia convergens Guerin‐Meneville, was determined by assigning a single predator randomly to each of four prey density treatments in the laboratory. Prey densities included 25, 50, 100, and 200 aphids per Petri dish arena. Predation response was recorded at 1, 4, 8, 16, 24, and 48 h after assigning predators to their prey treatments. Rate of consumption increased through time, with all 25 aphids consumed during the first 4 h of the experiment. At the highest density, adult lady beetle consumed on average 49, 99, 131, 163, 183, and 200 aphids within 1, 4, 8, 16, 24 and 48 h, respectively. Predators showed a curvilinear feeding response in relation to total available time, indicating that convergent lady beetles have the potential to suppress larger populations of aphids through continuous feeding by regulating their predation efficiency during feeding. The analysis of age‐specific mortality in absence of prey revealed that lady beetles could survive for an extended period of time (more than 2 weeks) without prey. The ability of a predator to survive without prey delays or prevents the rebound of pest populations that is a significant factor in natural biological control. A two‐year field sampling of 10 cotton arthropod predator species showed that spiders (27%) were the most dominant foliage dwelling predators in the Texas High Plains cotton followed by convergent lady beetles (23.5%), hooded beetles (13.5%), minute pirate bugs (11%), green lacewings (9.5%), bigeyed bugs (7.5%), scymnus beetles (3%), soft‐winged flower beetles (2%), damsel bugs (1.5%), and assassin bugs (1.5%). A field cage study showed that one H. convergens adult per plant released at prey density of one aphid per leaf kept the aphid population below economic threshold for the entire growing season.     

Behavioral responses of three species of ground beetle (Coleoptera: Carabidae) to olfactory cues associated with prey and habitat

The responses ofPterostichus melanarius Illiger,Harpalus rufipes DeGeer, andNebria brevicollis Fabricius (Coleoptera: Carabidae) to olfactory cues of prey and habitat were studied in a four-arm continuous-airflow olfactometer. The process was semiautomated using time-lapse image analysis by microcomputer. The primary constituent of the aphid alarm pheromone, (E)--farnesene (EBF), was synthesized and tested for a kairomonal role in prey detection by carabids. In addition, individual beetles were exposed to odors from live aphids, live collembolans, and a crude extract of wheat. All three beetle species showed evidence of olfaction.P. melanarius responded to all the odors except collembolans,H. rufipes responded to EBF and wheat, andN. brevicollis to collembolans. The use of a defence allomone as a prey-finding kairomone by certain carabid species has implications for pest management. Manipulation of predator chemical ecology by the inclusion of behavior-modifying compounds in a crop spray mix with reduced amounts of insecticide may allow for efficient aphid control with less environmental contamination.     

Factors determining the pest status of the rose-grain aphid, Metopolophium dirhodum (Walker), on winter barley in the United Kingdom

To test the hypothesis that early maturation of barley affords it some resistance to cereal aphids, the colonisation, survival, growth, reproduction and emigration of M. dirhodum on barley was measured and compared with previously published results for this aphid on wheat.
Barley plants were colonised as readily as wheat plants. Survival to maturity was similar on both hosts. Relative growth rate was higher on wheat than on barley for most of the season while reproduction was similar on both. Emigration, measured alate production, was significantly higher on mature wheat than on mature barley.
It is proposed that the increase of M. dirhodum populations on barley is normally terminated early in the year directly plant maturation and consequent drying out of the leaves, resulting in the death of aphids. This is different to the situation on wheat, which matures later, and where emigration following plant induced alate induction has been shown to be the major factor affecting the decline in population growth.
These results support the hypothesis that early maturation of barley provides some resistance to M. dirhodum.     

Habitat age increases reproduction and nutritional condition in a generalist arthropod predator

We studied the nutritional and reproductive response of Poecilus cupreus (synonymous with Pterostichus cupreus), one of the most abundant carabid beetles in arable land, to the succession in sixteen 1- to 4-year-old wildflower areas. A total of 390 male and 373 female beetles was examined. Each female was dissected and the number of ripe eggs counted. The nutritional state was expressed by a condition factor, which was calculated for each individual based on the observed weight and elytra length of male and female P. cupreus. Carabids in the 1-year-old wildflower areas contained significantly less ripe eggs than those from the 4-year-old areas. The condition factor of female and male beetles was significantly lower in 1- than 2- to 4-year-old areas. We examined the influence of habitat parameters (vegetation cover, soil water content, coarse and fine sand, pore volume, habitat size and age, surrounding landscape structure) on the reproductive success and nutritional state of P. cupreus in the 16 wildflower areas. The number of eggs was best explained by habitat age, accounting for 53.4% of the variability. The variation in the condition factor of female and male beetles was best explained by habitat age, which accounted for 73% and 71% of the variation, respectively. Moreover, the beetles' reproductive potential and nutritional condition were significantly associated with vegetation cover, and occasionally also with soil water content. The potential of wildflower areas as a reservoir for the generalist predator P. cupreus was shown to increase with successional age.     

Observation on the initial inoculum source and dissemination of Entomophthorales-caused epizootics in populations of cereal aphids

A total number of 1092 migratory alates were trapped from air in wheat grown area of Yuanyang County, Henan Province from early April through May 2002 in order to confirm the source and dissemination of entomophthoralean inocula to cause epizootics of cereal aphids. Those included 415 Sitobion avenae, 642 Rhopalosiphum padi, 22 Metopolophium dirhodum, and 13 Schizaphis graminum. The trapped alates were daily collected and individually reared for 7 days on wheat plants in laboratory. Of those 341 alates died of fungal infection, taking 31.2% in the trapped alates. These included 224 S. avenae, 106 R. padi, 8 M. dirhodum, and 3 S. graminum. Deaths of all infected alates occurred during the first 5 days and 78.9% of the deaths occurred within the first 3 days. Individual examination under microscope proved that all deaths were attributed to entomophthoralean fungi. Of those Pandora neoaphidis accounted for 84.6%, Conidiobolus obscurus for 9.9%, and Entomophthora planchoniana for 5.5%. Four alate deaths died of cross infection of P. neoaphidis and C. conidiobolus. Based on the high infection rate of the migratory alates trapped from air and the field occurrence of epizootics in populations of cereal aphids during the trapping period, Entomophthorales-caused epizootics were likely disseminated by infected alates through their flight and colonization. This makes it reasonable to interpret worldwide distribution of aphid epizootics, particularly caused by P. neoaphidis that has no resting spores discovered.     

Observation on the initial inoculum source and dissemination of Entomophthorales-caused epizootics in populations of cereal aphids

A total number of 1092 migratory alates were trapped from air in wheat grown area of Yuanyang County, Henan Province from early April through May 2002 in order to confirm the source and dissemination of entomophthoralean inocula to cause epizootics of cereal aphids. Those included 415 Sitobion avenae, 642 Rhopalosiphum padi, 22 Metopolophium dirhodum, and 13 Schizaphis graminum. The trapped alates were daily collected and individually reared for 7 days on wheat plants in laboratory. Of those 341 alates died of fungal infection, taking 31.2% in the trapped alates. These included 224 S. avenae, 106 R. padi, 8 M. dirhodum, and 3 S. graminum. Deaths of all infected alates occurred during the first 5 days and 78.9% of the deaths occurred within the first 3 days. Individual examination under microscope proved that all deaths were attributed to entomophthoralean fungi. Of those Pandora neoaphidis accounted for 84.6%, Conidiobolus obscurus for 9.9%, and Entomophthora planchoniana for 5.5%. Four alate deaths die     

Population dynamics of cereal aphids: influence of a shared predator and weather

1 Aphid populations may show strong year-to-year fluctuations, but questions remain regarding the dominance of factors that cause this variation, especially the role of natural enemies. To better understand the dynamics of aphid species that occur as pests in cereals, we investigated the relative influence of top-down control by a predator and weather (temperature and precipitation) on population fluctuations of three cereal aphid species.
2 From 1987 to 2005, populations of Metopolophium dirhodum , Sitobion avenae and Rhopalosiphum padi in insecticide-free stands of winter wheat were monitored in the Praha-Ruzyné region of the Czech Republic. Densities of an aphidophagous predator, the ladybeetle Coccinella septempunctata , were recorded from an overwintering site in the landscape. Weather was quantified using historical records.
3 A significant bottom-up effect of densities of aphids on those of C.   septempunctata was found, but evidence of direct top-down regulation of aphids by C.   septempunctata was only significant in the case of R.   padi . There was no significant periodicity in the dynamics of the aphid or C.   septempunctata , suggesting that there was no clear predator-prey cycle. Combinations of C.   septempunctata and weather variables could be used to explain M.   dirhodum and R.   padi per capita rate of change. There were also indications that weather directly affected peak density of M.   dirhodum .
4 We conclude that regional estimates of C.   septempunctata densities are not sufficient to determine whether aphid population dynamics are driven by predator–prey interactions. Feasibility of time series analysis as an investigative tool in aphid population dynamics studies is discussed.     

Foraging by the carabid Agonum dorsale in the field

Abstract. 1. Small arenas in the field were used for observing adult Agonum dorsale (Pont.) foraging in a wheat crop.
2. The carabid showed no significant change in behaviour between low and high aphid density arenas.
3. Individuals foraged nocturnally and spent most of their time on the ground searching for prey. They climbed infrequently and then not high enough to reach aphid aggregations on the ears and flag leaves of wheat.
4. Gut dissection showed that more individuals had eaten aphids in the high aphid density arena than the low. Recording of aphid distribution within the arenas showed that this increase in aphid consumption could be explained by A.dorsale capturing aphids on the ground. Prey availability per unit area was in fact higher on the ground than on the wheat.
5. The contribution of ground-zone predators such as A.dorsale to the control of cereal aphids will be determined by the frequency with which aphids arrive on the ground and the proportion of these which reclimb the wheat and reproduce in the absence of predation.     

Molecular analysis of predation on parasitized hosts

Predation on parasitized hosts can significantly affect natural enemy communities, and such intraguild predation may indirectly affect control of herbivore populations. However, the methodological challenges for studying these often complex trophic interactions are formidable. Here, we evaluate a DNA-based approach to track parasitism and predation on parasitized hosts in model herbivore-parasitoid-predator systems. Using singleplex polymerase chain reaction (SP-PCR) to target mtDNA of the parasitoid only, and multiplex PCR (MP-PCR) to additionally target host DNA as an internal amplification control, we found that detection of DNA from the parasitoid, Lysiphlebus testaceipes, in its aphid host, Aphis fabae, was possible as early as 5 min. post parasitism. Up to 24 h post parasitism SP-PCR proved to be more sensitive than MP-PCR in amplifying parasitoid DNA. In the carabid beetles Demetrias atricapillus and Erigone sp. spiders, fed with aphids containing five-day-old parasitoids, parasitoid and aphid DNA were equally detectable in both predator groups. However, when hosts containing two-day-old parasitoids were fed to the predators, detection of parasitoid prey was possible only at 0 h (immediately after consumption) and up to 8 h post consumption in carabids and spiders, respectively. Over longer periods of time, post-feeding prey detection success was significantly higher in spiders than in carabid beetles. MP-PCR, in which parasitoid and aphid DNA were simultaneously amplified, proved to be less sensitive at amplifying prey DNA than SP-PCR. In conclusion, our study demonstrates that PCR-based parasitoid and prey detection offers an exciting approach to further our understanding of host-parasitoid-predator interactions.     

Effect of plant nutrition on aphid size,prey consumption,and life history characteristics of green lacewing

Plant quality can directly and indirectly affect the third trophic level. The predation by all the instars of green lacewing, Chrysoperla carnea （S.） （Neuroptera： Chrysopidae） on the cereal aphids, Rhopalosiphum padi （L.）, and Sitobion avenae （F.） at varying nitrogen fertilizer levels was calculated under laboratory conditions. Wheat plants were grown on four nitrogen fertilizer levels and aphids were fed on these plants and subsequently offered as food to the C. carnea. Aphid densities of 10, 30, and 90 were offered to first, second, and third instar larvae of green lacewing. Increased nitrogen application improved nitrogen contents of the plants and also the body weight of cereal aphids feeding on them. Aphid consumption by green lacewings was reduced with the increase in nitrogen content in the host plants of aphids. Predation of both aphid species by first, second, and third instars larvae of C. carnea was highest on aphids reared on plants with the lowest rate of fertilization, suggesting a compensatory consumption to overcome reduced biomass （lower aphid size）. Total biomass devoured by C. carnea on all nitrogen fertilizer treatments was not statistically different. Additionally, the heavier host prey influenced by the plant nutrition had an effect on the life history characteristics of green lacewings. The larval duration, pupal weight, pupal duration, fecundity, and male and female longevity were significantly affected by the level of nitrogen fertilization to the aphid＇s host plants, except for pupal duration when fed on S. avenae. This study showed that quantity of prey supplied to the larvae affects the prey consumption and thereafter the life history characteristics of green lacewings.     

Pitfall trap catches of and aphid predation byPterostichus melanarius andPterostichus madidus in insecticide treated and untreated potatoes

Aphids were countedin situ and carabid beetle populations assessed by pitfall trapping betwen 1983–1985 in potato fields in Scotland treated with demeton-S-methyl (DSM) or untreated.Macrosiphum euphorbiae (Thomas) (Homoptera: Aphididae) was the most abundant aphid. Nineteen species of carabid beetles were trapped but the fauna was dominated byPterostichus melanarius (Ill.) andPterostichus madidus (Fabr.) (Coleoptera: Carabidae). Pitfall trap catches of these carabids were variable but generally lower in sprayed than unsprayed plots for a few days after DSM application and higher a week or two later. Of 1800P. melanarius and 910P. madidus dissected, 14.4 per cent and 30.5 per cent respectively, contained aphid remains. The proportion that had eaten aphids often was higher in sprayed than unsprayed plots immediately after treatment. In additional experiments, aphids treated with DSM fell from plants; four to five per cent of the theoretically applied dose of DSM reached the ground in a closed-canopy potato crop; and no mortality occurred whenPterostichus spp. were exposed to DSM in a field bioassay. It is proposed that observed patterns in trap catch ofPterostichus spp. around times of pesticide application may be due in part to aphids falling to the ground after treatment. Beetles that eat these fallen aphids may be less hungry, less active and less likely to be trapped. Later increases in catches may be attributed to hungry, more active beetles, a consequence of prey removal by the insecticide. Although this hypothesis requires further research, it is clear that the effects of insecticide treatment on carabid populations cannot be measured by pitfall trapping alone.     

Comparing growth patterns among field populations of cereal aphids reveals factors limiting their maximum abundance

Cereal stands in central Europe are commonly infested with three species of aphids that may become serious pests. With increasing abundance, the proportion of a particular species in the total aphid population may remain constant, suggesting a density-independent exponential growth, or the proportion can change, suggesting density-dependent constraints on growth. The constraints that affect particular species, and thus their relative abundance, were studied. The proportionality between maximum abundances of the cereal aphids was studied using a 10-year census of the numbers of aphids infesting 268 winter wheat plots. For two species their abundance on leaves and ears was compared. With increasing aphid density the maximum abundance of Rhopalosiphum padi (Linnaeus) remained proportional, but not that of Sitobion avenae (Fabricius), which was constrained by the smaller surface area of ears compared to leaves. There was no evidence of inter-specific competition. Maximum abundance of R. padi and Metopolophium dirhodum (Walker) on leaves did not change proportionally as the proportion of M. dirhodum decreased with increasing overall aphid density. This decrease was probably caused by the restricted distribution of M. dirhodum, which is confined to leaves, where space is limiting. No change in proportion between populations was detected when the average densities were below 0.54 aphids per leaf or ear. Non-proportional relationships between aphid populations appeared to be due to spatial constraints, acting upon the more abundant population. Detecting the limitation of population growth can help with the assessment of when density-independent exponential growth is limited by density-dependent factors. This information may help in the development of models of cereal aphid population dynamics.