Resource tracking in aphids: programmed reproductive strategies anticipate seasonal trends in habitat quality

Most aphids show cyclical parthenogenesis, have short generation times and complete several generations each season. Two hypotheses have been proposed to account for the adaptive seasonal trends observed in the reproductive strategies of aphids. Firstly, individuals of each generation modify their reproductive strategy in direct response to the conditions they experience during their development. Secondly, the reproductive strategies of the different generations are to a large extent programmed and anticipate seasonal trends in habitat quality. These hypotheses were tested by rearing individuals of three generations of the host-alternating willow-carrot aphid, Cavariella aegopodii, on both willow and carrot. This revealed that the way this aphid allocates resources to gonads and lipid reserves is independent of an aphid's weight and the host plant on which it is reared. In addition each generation shows a specific relationship between offspring size and adult size, which tends to keep the absolute investment in individual offspring relatively constant from generation to generation, inspite of the big differences in adult size between generations. That is, through programmed allometric engineering aphids anticipate the predictable seasonal trends in habitat quality and so more closely track their resources, investing relatively more in gonads when food quality is high and relatively more in lipoidal reserves when food quality is poor. Received: 25 August 1997 / Accepted: 19 October 1997     

Transgenic Bt maize and Rhopalosiphum padi (Hom., Aphididae) performance

Abstract.  1. The population abundance and age structure of Rhopalosiphum padi , one of the most common maize aphid species, on transgenic Bt (expressing the Cry1Ab protein) and non-Bt isogenic maize was studied in commercial plots during three crop seasons.
2. A higher density of aphids, particularly alates and young nymphs, occurred in Bt plots at very young maize development stages, corresponding to the settlement period, in the 3 years studied. Possible causes are discussed. After this period, there were no differences between Bt and non-Bt maize.
3. Mortality, development, and reproduction of the offspring of alate forms of R. padi and the offspring of different generations of apterous forms fed with Bt maize were evaluated in the laboratory under controlled conditions.
4. The developmental and pre-reproductive times of the offspring of the first generation of alatae were shorter and the intrinsic rate of natural increase ( r _m ) higher when aphids fed on Bt maize. The opposite occurs with the offspring of the first generation of apterous mothers, which have lower nymphal and adult mortality, shorter developmental and pre-reproductive times, a higher effective fecundity rate, and greater r _m , when fed on non-Bt maize. The differences in aphid development on the two cultivars may be linked to changes in host-plant quality due to pleiotropic effects of the genetic modification.
5. No differences on aphid mortality, developmental and pre-reproductive times, fecundity, and r _m were found between the offspring of apterous aphids maintained on Bt or non-Bt maize for several generations.     

Compatibility of aphid resistance in soybean and biological control by the parasitoid Aphidius colemani (Hymenoptera: Braconidae)

Biological control and soybean cultivars bred for increased resistance to the soybean aphid (Aphis glycines) are two approaches used to manage this serious pest of soybeans in North America. However, as with many other pest systems, the compatibility of these two pest management approaches has not been studied in detail. The aphidiine wasp Aphidius colemani is one of several candidate species for biological control of the soybean aphid in soybean in North America. Resistance to the soybean aphid in the USDA soybean cultivar Dowling is largely controlled by a single dominant gene Rag1, which is the focus of plant breeding programs directed against the soybean aphid. In this study, we measured developmental and behavioral differences in the parasitic wasp A. colemani when it attacked soybean aphids feeding on either the aphid-resistant Dowling or aphid-susceptible Glenwood cultivars of soybean. We used a combination of choice and no-choice experiments to examine the effects of host plant cultivar on the number of parasitized aphids formed and the sex ratio and body weights of adult offspring produced. Significantly more aphids were parasitized when they fed on Glenwood compared to Dowling and these offspring were larger when they developed in aphids that fed on Glenwood soybeans. To distinguish between effects on foraging decisions and offspring survivorship, we conducted an additional experiment that followed the oviposition decisions and fate of each parasitized aphid. Foraging female A. colemani spent less time handling individual aphids and encountered and attacked aphids at a higher rate when they fed on aphids feeding on Glenwood soybeans than aphids feeding on Dowling soybeans. Furthermore, wasp survivorship in aphids was greater on Glenwood than Dowling. Taken together, aphid-resistance in soybeans has negative effects on foraging behavior and offspring fitness of A. colemani raising concerns about the compatibility of these two pest management approaches.     

The value of three cereal aphid species as food for a generalist predator

The value of the cereal aphid species Metopolophium dirhodum (Wlk.), Sitobion avenae (F.) and Rhopalosiphum padi (L.) as prey for the linyphiid spider Erigone atra (Bl.) was assessed. Fecundity of females was determined for spiders fed on eight experimental diets: three single‐species aphid diets, a mixed diet of all three aphid species, three mixed diets with each aphid species in combination with fruit flies Drosophila melanogaster (Meig.), and pure D. melanogaster as a high quality comparison diet. The development and survival of first‐instar juveniles fed on three diets of single aphid species, and on a diet of Collembola were compared with those subjected to starvation. Prey value for adult females was assessed by egg production, hatching success and offspring size. In pure diets all three aphid species were of low value to the spiders, causing a rapid decline in egg production and supporting no growth of significance of first‐instar juveniles. No difference in value of aphid species of single‐species aphid diets was found in the fecundity experiment, while a ranking of aphid species of M. dirhodum > R. padi > S. avenae was revealed in the survivorship experiment. A mixed‐aphid diet was not found to be advantageous compared with single‐species aphid diets, and no advantage of including aphids in mixed diets with fruit flies was found. Metopolophium dirhodum and R. padi were neutral in mixed diets, while a diet of S. avenae and fruit flies caused reduced egg production compared with the pure diet of fruit flies, revealing a toxic effect of S. avenae on the spider. The value‐ranking of aphid species in mixed diets was similar to that of single‐species diets. A similar ranking of aphid species was found for different fitness parameters (fecundity of adult females and development of juveniles). A ranking of aphids by offspring size of mothers on aphid‐only diets was S. avenae > M. dirhodum > R. padi. All aphid‐fruit fly diets resulted in larger offspring than a diet of only D. melanogaster, with the overall largest offspring being produced on the diet of M. dirhodum and fruit flies.     

Multiple Cues for Winged Morph Production in an Aphid Metacommunity

Environmental factors can lead individuals down different developmental pathways giving rise to distinct phenotypes (phenotypic plasticity). The production of winged or unwinged morphs in aphids is an example of two alternative developmental pathways. Dispersal is paramount in aphids that often have a metapopulation structure, where local subpopulations frequently go extinct, such as the specialized aphids on tansy (Tanacetum vulgare). We conducted various experiments to further understand the cues involved in the production of winged dispersal morphs by the two dominant species of the tansy aphid metacommunity, Metopeurum fuscoviride and Macrosiphoniella tanacetaria. We found that the ant-tended M. fuscoviride produced winged individuals predominantly at the beginning of the season while the untended M. tanacetaria produced winged individuals throughout the season. Winged mothers of both species produced winged offspring, although in both species winged offspring were mainly produced by unwinged females. Crowding and the presence of predators, effects already known to influence wing production in other aphid species, increased the percentage of winged offspring in M. tanacetaria, but not in M. fuscoviride. We find there are also other factors (i.e. temporal effects) inducing the production of winged offspring for natural aphid populations. Our results show that the responses of each aphid species are due to multiple wing induction cues.     

Reproductive schedule and factors affecting soldier production in the eusocial bamboo aphid Pseudoregma bambucicola (Homoptera, Aphididae)

Summary: The reproductive characteristics of the soldier-producing aphid Pseudoregma bambucicola were studied in Kagoshima, Southern Japan, to know the factors affecting soldier production of eusocial aphids. The soldier proportion in aphid colonies was highest from October to November. In some large colonies, soldiers were observed in all seasons except in July when colony size was relatively small. Multiple regression analysis showed that the colony size was a principal factor affecting soldier proportion throughout a year. Other social or environmental factors such as aphid composition, host plant conditions and predator abundance were not always significant. Rearing experiments revealed that large colonies (̿,000 individuals) produced soldiers in almost all seasons while small colonies (<1,000) never produced any soldiers. The caste-production schedule of adult females was examined in the field. When solitary females produced both castes, they usually produced normal nymphs first and then soldiers. Females from large colonies tended to produce more soldiers in the earlier period of their lifetime, whereas females from newly established small colonies produced no or only a few soldiers at later times. The average number of soldiers and normal nymphs produced consecutively by a single female was >10 and >20, respectively. Because they have a small number of ovarioles (<15 on average), females should alter caste production within the same ovarioles according to changes in environmental conditions. Artificial removal or introduction of predators and reduction of colony size did not affect soldier production over two successive generations, revealing maternal effects on soldier production. Females cannot shift caste production quickly in response to changes in predator abundance and colony size. This is probably due to early developmental determination of castes within the mother's body.     

Are fecundity and longevity of female Aphelinus abdominalis affected by development in GNA‐dosed Macrosiphum euphorbiae?

Abstract. Snowdrop lectin ( Galanthus nivalis agglutinin, GNA) confers partial resistance to several aphid species when incorporated into an artificial diet and/or expressed in transgenic potato. First-tier laboratory-scale experiments were conducted to assess the potential effect of GNA on the longevity and fecundity of female parasitoid Aphelinus abdominalis (Dalman) that had developed in Macrosiphum euphorbiae (Thomas) fed artificial diet containing 0.1% GNA (w/v). In a previous study it was shown that GNA ingested by A. abdominalis larvae is not acutely toxic. It was also shown that GNA has a host-size mediated effect on parasitoid sex ratio and larval development, but no apparent direct effect.         In this study, we report that A. abdominalis larvae that developed in GNA-dosed aphids that were smaller than control aphids of the same age, produced smaller adults with a reduced longevity and fecundity. Aphelinus abdominalis larvae that developed in GNA-dosed aphids older than the control but of the same size, produced adults of similar size that lived as long as the control but had a reduced fecundity.         Our results suggest that GNA fed to aphids in artificial diet has both a host-mediated effect (via aphid-size) and a direct effect on adult parasitoid fecundity. It is not known how GNA affects parasitoid larval development and subsequently adult fecundity, but it is hypothesized that GNA acted as an antifeedant to parasitoid larvae, thus disturbing nutrient assimilation and conversion necessary for egg maturation.     

A comparison of maternal effects and current environment on vital rates of Aphis nerii, the milkweed–oleander aphid

Abstract.  1. Non-Mendelian maternal effects, the effects of maternal phenotype or environment on offspring phenotype, have been documented in numerous taxa. By affecting offspring vital rates (birth, death, and movement), maternal effects have the potential to influence population dynamics. However, relatively few studies have directly linked maternal phenotype or environment to offspring vital rates. Additionally, even fewer studies have compared the magnitude of across-generation effects (i.e. maternal effects) to within-generation effects.
2. Because of their telescoping generations, aphids can be strongly influenced by maternal effects. The effects of maternal density and maternal host-plant species on offspring survival, fecundity, and alate formation were investigated experimentally in Aphis nerii , the milkweed–oleander aphid.
3. Additionally, the relative strength of maternal effects were compared with those operating within a generation. Therefore, in another set of experiments, the effects of current density and host-plant species (within-generation effects) on aphid vital rates were examined.
4. While maternal effects were present, within-generation effects were much stronger and more strongly influenced aphid vital rates. Within a generation, aphids exhibited density-dependent survival, fecundity, and alate formation and these effects varied among host-plant species.
5. These results indicate that while maternal effects have the potential to affect population dynamics, this potential is not always met. Additionally, the current environment, not the environment of previous generations, more strongly impacts population dynamics.     

A transgenerational interval timer inhibits unseasonal sexual morph production in damson-hop aphid, Phorodon humuli

Abstract.  The induction of sexual and parthenogenetic morphs of the damson-hop aphid, Phorodon humuli , on hops is controlled by daylength. The ability of P. humuli , to produce winged pre-sexual females (gynoparae) in the short-day conditions of spring is inhibited by an interval timer present in generations immediately after hatching of the overwintering egg. The inhibition expires after three generations when nymphs are born and reared in short days (LD 12 : 12 h), irrespective of whether their parents are reared in short or long days (LD 18 : 6 h). No gynoparae are produced by aphids maintained for 13 generations in long days. Two wingless aphids from 35 survive transfer from Prunus spinosa to hops. No winged females are produced during nine generations among their progeny maintained in long days on hops, but gynoparae, followed by males, are produced one generation after these aphids are transferred to short days.     

Effects of beauvericin on Schizaphis graminum (Aphididae)

The effects of beauvericin, a toxic fungal metabolite common contaminant of maize and wheat, on aphid fitness were studied in three consecutive generations of females. Aphids were reared on wheat leaves inserted into a sandy substratum wetted with a solution of beauvericin. Ingestion of this solution through leaves did not significantly decrease the lifespan of females of all generations as compared to controls. However, the mean number of offspring from the third generation of treated females was significantly smaller than those in controls. Furthermore, treated second and third generation females produced a greater number of abortive embryos. Histological analysis revealed abundant DAPI and Feulgen positive material in the cytoplasm of some bacteriocytes of treated third generation females. This material was attributed to the endosymbionts of bacteriocytes. Tests by contact were also carried out and revealed a significantly lower survival of treated first instar aphids as compared to controls 18h after the start of the trial.     

一种室内饲养麦蚜的方法

介绍利用水培麦苗大量饲养麦蚜的技术。在不补充营养液的条件下麦苗可以维持10~15d,一般在饲养条件下可以1周换1次麦苗。小麦种子在经过24h浸种后,平铺在底部垫有滤纸的直径9cm的培养皿中,在25℃,4d后就可以接种麦蚜。适宜的试样条件为温度在18~25℃,相对湿度50%~70%,光周期为17hr∶7hr,饲养1代大约需要5~9d时间,每皿可养麦蚜500头以上。经过40代以上的连续饲养,蚜虫体重和每蚜产幼蚜数量没有明显降低趋势。     

Sexual forms of the pea aphid, Acyrthosiphon pisum, produced on an artificial diet

In recent years a number of aphid species have been successfully reared on chemically defined diets (Dadd & Mittler, 1966; Auclair, 1965; Dadd & Krieger, 1967; Ehrhardt, 1968). The development of such diets has offered an opportunity to study in detail several aspects of the physiology of these insects. In particular, the composition of the artificial diet has been shown to influence the production of winged forms by Myzus persicae (Sulzer) (Mittler & Dadd, 1966; Dadd, 1968). In all studies thus far, the aphids have been maintained in the parthenogenetic condition in order to accumulate data concerning the growth and fecundity of females of successive generations. However, it was clearly of interest to determine whether sexual forms could be produced by diet-reared aphids. If so, further information regarding the sexual reproduction of these insects would be forthcoming. Attempts to achieve this using local strains of Myzus persicae, Aphis fabae Scopoli and Acyrthosiphon pisum (Harris) were unsuccessful, presumably because these strains were anholocyclie (Kunkel, unpubl.). A clone of a green strain of A. pisum, previously found to be holocyclic (Sutherland, unpubl.), was then obtained from England and was cultured at 20° C in a photoperiodic regime of 16 hr light: 8 hr dark per day on young bean plants (Vicia faba). The artificial diet on which the experimental aphids were maintained was formulated according to Dadd (1967) and enclosed in sachets of stretched parafilm (Dadd, Krieger & Mittler, 1967). The aphids were provided with fresh sachets every 3–4 days. Adult apterous females taken from the plant culture were caged in groups of five, and exposed to a photoperiodic regime of 10 hr light: 14 hr dark per day at 15°. Successive batches of larvae deposited by these aphids were maintained in this environment until they were adult and had produced offspring for several days. These developed into oviparae and males. Compared to the sexual forms produced on host plants, these diet-reared individuals grew relatively poorly and mortality was high. Nevertheless, the surviving oviparae deposited numerous eggs, but these remained pale green whereas viable, fertilized eggs deposited by plant-reared oviparae rapidly become black. In the case of aphids reared on host plants in a short-day environment the possibility exists that the photoperiod may act indirectly via the plants and not directly on the insects themselves — a doubt which has been raised concerning Megoura viciae Buckton (von Dehn, 1967). However, our results with A. pisum reared on artificial diet demonstrate, as Lees did for M. viciae (Lees, 1960, 1967), that it is unnecessary to implicate the host plant in the determination of males and oviparae. This does not, of course, exclude the possibility that photoperiod-induced changes in a host plant may play a role as well. Nor have we ruled out the possibility that dietary composition may influence the aphids directly or their response to other environmental stimuli leading to the production of sexual forms.     

Clonal mixing in the soldier-producing aphid Pemphigus spyrothecae (Hemiptera: Aphididae)

Illuminating the genetic relationships within soldier-producing aphid colonies is an essential element of any attempt to explain the evolution of the altruistic soldier caste. Pemphigus spyrothecae is a soldier-producing aphid that induces galls on the leaf petioles of its host (trees of the genus Populus). At least a quarter of the aphids within the clonally produced gall population are morphologically and behaviourally distinct first-instar soldiers that defend the gall population from predation. Using field trapping and microsatellites, we investigated the degree of clonal mixing within natural gall populations. Field trapping in the UK showed that all the migrants of P. spyrothecae and of two other Pemphigus species were wingless first-instar soldiers. The average degree of mixing estimated from trapping P. spyrothecae migrants was 0.68% (range = 0-15%). Microsatellite genotyping of 277 aphids from 13 galls collected in Italy revealed an average mixing level of 10.4% (range = 0-59%). Six galls contained more than one clone (range = 2-5 clones). Non-kin aphids were not restricted to the soldier caste but were evenly distributed across instars. An additional gall, from which 527 occupants were genotyped, contained 12 non-kin aphids distributed among nine clones, showing that clonal diversity can be high even when mixing is very low. These observations suggest that although soldiers migrate regularly and can moult and reproduce within foreign galls, clonal mixing in this species is generally low and is unlikely to provide a barrier to the evolution of investment by the aphid clones in an altruistic soldier caste.     

Effects of plant protease inhibitors, oryzacystatin I and soybean Bowman-Birk inhibitor, on the aphid Macrosiphum euphorbiae (Homoptera, Aphididae) and its parasitoid Aphelinus abdominalis (Hymenoptera, Aphelinidae)

Transgenic plants expressing protease inhibitors (PIs) have emerged in recent years as an alternative strategy for pest control. Beneficial insects such as parasitoids may therefore be exposed to these entomotoxins either via the host or by direct exposure to the plant itself. With the objective of assessing the effects of PIs towards aphid parasitoids, bioassays using soybean Bowman-Birk inhibitor (SbBBI) or oryzacystatin I (OCI) on artificial diet were performed on Macrosiphum euphorbiae-Aphelinus abdominalis system. OCI significantly reduced nymphal survival of the potato aphid M. euphorbiae and prevented aphids from reproducing. This negative effect was much more pronounced than with other aphid species. On the contrary, SbBBI did not affect nymphal viability but significantly altered adult demographic parameters. Enzymatic inhibition assays showed that digestive proteolytic activity of larvae and adults of Aphelinus abdominalis predominantly relies on serine proteases and especially on chymotrypsin-like activity. Immunoassays suggested that OCI bound to aphid proteins and accumulated in aphid tissues, whereas SbBBI remained unbound in the gut. Bioassays using M. euphorbiae reared on artificial diets supplemented with both OCI and SbBBI showed a fitness impairment of Aphelinus abdominalis that developed on intoxicated aphids. However, only SbBBI was detected in parasitoid larvae, while no PI could be detected in adult parasitoids that emerged from PI-intoxicated aphids. The potential impact of PI-expressing plants on aphid parasitoids and their combined efficiency for aphid control are discussed.     

Evaluation of artificial diets for rearing Aphis glycines (Hemiptera: Aphididae)

Artificial aphid diets have been previously developed for the pea aphid, Acyrthosiphon pisum (Harris), and the green peach aphid, Myzus persicae (Sulzer). The ability to rear aphids on an artificial diet allows for selectively adding or subtracting compounds from an aphid's food source to determine the effect on fecundity and longevity. Five diets previously developed for the green peach aphid and the pea aphid were tested for their suitability for rearing soybean aphid, Aphis glycines Matsumura. The best diet, originally developed for the green peach aphid and based on the amino acid profile of young potato plants, allowed 12 generations of soybean aphids to develop. For all diets tested, aphid fecundity, and longevity were greatly reduced in comparison with aphids reared on soybean, Glycine max (L.) Merr., plants or on detached soybean leaves. In addition, mean developmental time was significantly longer for aphids reared on artificial diets.     

Density-dependent induction and suppression of soldier differentiation in an aphid social system

We investigated the mechanism underlying the control of soldier production in colonies of a social aphid, Tuberaphis styraci, which has a sterile soldier caste in the second instar. High aphid density was shown to induce soldier production in T. styraci. Analysis of natural colonies revealed that the soldier proportion tended to increase with aphid density but reached a plateau. Artificial diet experiments identified a similar plateau of soldier proportion under high-density conditions. In order to gain insights into the controlling mechanism of soldier production, the effect of soldiers on reproducing adult aphids was examined using the artificial diet system. It was experimentally demonstrated that soldier production was suppressed by coexisting soldiers, whereas coexisting non-soldiers facilitated soldier production. These results suggested that caste ratio in the colony of T. styraci is controlled by positive and negative feedbacks consisting of density-dependent induction and suppression of soldier differentiation.     

Sources of variation in dietary requirements in an obligate nutritional symbiosis

The nutritional symbiosis between aphids and their obligate symbiont, Buchnera aphidicola, is often characterized as a highly functional partnership in which the symbiont provides the host with essential nutrients. Despite this, some aphid lineages exhibit dietary requirements for nutrients typically synthesized by Buchnera, suggesting that some aspect of the symbiosis is disrupted. To examine this phenomenon in the pea aphid, Acyrthosiphon pisum, populations were assayed using defined artificial diet to determine dietary requirements for essential amino acids (EAAs). Six clones exhibiting dependence on EAAs in their diet were investigated further. In one aphid clone, a mutation in a Buchnera amino acid biosynthesis gene could account for the clone''s requirement for dietary arginine. Analysis of aphid F₁ hybrids allowed separation of effects of the host and symbiont genomes, and revealed that both affect the requirement for dietary EAAs in the clones tested. Amino acid requirements were minimally affected by secondary symbiont infection. Our results indicate that variation among pea aphids in dependence on dietary amino acids can result from Buchnera mutation as well as variation in the host genotype.     

Quality of two aphid species (Rhopalosiphum padi and Sitobion avenae) as food for the generalist predator Tachyporus hypnorum (Col., Staphylinidae)

Abstract:  The quality of two aphid species ( Rhopalosiphum padi and Sitobion avenae ) as food for the staphylinid generalist predator Tachyporus hypnorum was investigated. Fruit flies ( Drosophila melanogaster ) were used as prey for comparison. T. hypnorum has been reported to prey on aphids, to consume large quantities and to show a high preference for aphids. This study showed that compared with fruit flies aphids are not high-quality prey for neither adult females nor larvae of T. hypnorum . Larvae were less able to utilize aphids than adult females. Reproduction was affected by both aphid species; a diet of S. avenae reduced fecundity, and a diet of R. padi reduced egg hatching success and prolonged the time spent in the egg stage compared with a fruit fly diet. For the T. hypnorum larvae pure diets of both S. avenae and R. padi resulted in high larval mortality. Overall ranking of the three prey types based on several fitness parameters differed between larvae and adults. The adult fitness parameters gave rather conflicting rankings of all three prey types though the two aphid species were of approximately the same overall value, whereas the larval fitness parameters gave identical rankings: D. melanogaster  >  S. avenae  >  R. padi . In the larval stage survival was greatly reduced by both aphid diets, while this was not the case for the adult females.     

Introduced ants reduce interaction diversity in a multi‐species,ant–aphid mutualism

Mutualisms contribute in fundamental ways to the origin, maintenance and organization of biological diversity. Introduced species commonly participate in mutualisms, but how this phenomenon affects patterns of interactions among native mutualists remains incompletely understood. Here we examine how networks of interactions among aphid‐tending ants, ant‐tended aphids, and aphid‐attacking parasitoid wasps differ between 12 spatially paired riparian study sites with and without the introduced Argentine ant Linepithema humile in southern California. To resolve challenges in species identification, we used DNA barcoding to identify aphids and screen for parasitoid wasps (developing inside their aphid hosts) from 170 aphid aggregations sampled on arroyo willow Salix lasiolepis. Compared to uninvaded sites, invaded sites supported significantly fewer species of aphid‐tending ants and ant‐tended aphids. At invaded sites, for example, we found only two species of ant‐tended aphids, which were exclusively tended by L. humile, whereas at uninvaded sites we found 20 unique ant–aphid interactions involving eight species of ant‐tended aphids and nine species of aphid‐tending ants. Ant–aphid linkage density was thus significantly lower at invaded sites compared to uninvaded sites. We detected aphid parasitoids in 14% (28/198) of all aphid aggregations. Although the level of parasitism did not differ between invaded and uninvaded sites, more species of wasps were detected within uninvaded sites compared to invaded sites. These results provide a striking example of how the assimilation of introduced species into multi‐species mutualisms can reduce interaction diversity with potential consequences for species persistence.     

Migratory tendency in aging populations of the pea aphid,Acyrthosiphon pisum

Summary Sweep samples of the aphid, Acyrthosiphon pisum, were collected from six natural populations ranging in age from one to five years. Clones were established in the laboratory from the field-collected adults and tested for their migratory tendency in two subsequent generations by measuring the percentage of winged offspring produced in response to a standard stimulus. The number of aphids in sweep samples and the percentage of winged and wingless aphids were also determined. Tests on the first laboratory generation revealed a decline in migratory tendency with the age of the source population, but no such relationship was detected in tests on the second generation. These results are consistent with an explanation based on maternal age effects and differences in adult age structure among populations of different age. They are not consistent with one based on genetic differences among populations. Older populations also had higher densities and a lower percentage of alate adults. The percentage of larvae with wing buds was positively correlated with population density, but not population age.