M. Mackauer

Instar-specific defense of the pea aphid,Acyrthosiphon pisum: Influence on oviposition success of the parasiteAphelinus asychis (Hymenoptera: Aphelmidae)

The parasite Aphelinus asychisWalker (Hymenoptera: Aphelinidae) oviposits in all four instars of the pea aphid, Acyrthosiphon pisum(Harris) (Homoptera: Aphididae). Searching females display a highly stereotyped sequence of behaviors when encountering a host. Once recognized, an aphid is examined and probed by the wasp with the everted ovipositor prior to oviposition. Oviposition success is influenced by aphid behavior that is related to aphid size and expressed through instarspecific escape and defense reactions. Being smaller and less able to defend themselves, first and early-second instars of pea aphid are more susceptible to successful parasitism than third and fourth instars, in that order. Observed patterns of preference by Aphelinus females for particular aphid species and instars reflect the outcome of behavioral interactions between the hosts and the parasites, rather than preference in the strict sense.

Influence of aphid size, age and behaviour on host choice by the parasitoid wasp Ephedrus californicus : a test of host-size models

SummaryWhen host quality varies, parasitoid wasps are expected to oviposit selectively in high-quality hosts. We tested the assumption underlying host-size models that, for solitary species of wasps, quality is based on host size. Using Ephedrus californicus, a solitary endoparasitoid of the pea aphid, we evaluated the influence of aphid size (= mass), age and defensive behaviours on host selection. Experienced parasitoid females were given a choice among three classes of 5-day-old apterous nymphs: small aphids that had been starved daily for 4 h (S4) and 6 h (S6) respectively, and large aphids permitted to feed (F) normally. Wasps attacked more, and laid more eggs in, small than large aphids (S6>S4>F). This rank-order for attack did not change when females could choose among aphids of the same size that differed in age; however, wasps oviposited in all attacked aphids with equal probability. Host size did not influence parasitoid attack rates when aphids were anaesthetized so that they could not escape or defend themselves. As predicted by host-size models, wasp size increased with host size (F>S4; S6), but large wasps required longer to complete development than their smaller counterparts (S4<S6<F). For parasitoids of growing and feeding hosts (= koinobionts), host quality is not a linear function of current host size; it is influenced by the hosts age and development stage which determine the hosts potential to support parasitoid growth. Host choice in E. californicus reflects a trade-off between maximization of fitness gains per egg and the economics of search-time allocation. Because large aphids are more likely to escape parasitization, a wasp must balance her potential gain in fitness by ovipositinng in a high-quality (large) aphid against her potential cost in terms of lost opportunity time if the attack fails.

Host discrimination and larval competition in the aphid parasite Ephedrus californicus

Naive and experienced females of Ephedrus californicus Baker (Hymenoptera: Aphidiidae) were tested for their ability to discriminate between parasitized and unparasitized pea aphids, Acyrthosiphon pisum (Harris) (Homoptera: Aphididae). Attacks lasting ≥ 6 s generally resulted in oviposition; the average length was 11.8 s. The proportion of parasitized aphids that was rejected varied with the interval length between attacks. It is suggested that host discrimination is time‐dependent and can be induced by a pheromone‐like external marker left by a first‐attacking female (0–9 h), or by changes in host quality associated with parasite development (≥ 14 h). Experienced, but not naive, females responded to the external marker, which became less effective with time. In superparasitized aphids, the older of two E. californicus larvae usually eliminated a younger competitor; but younger larvae survived under certain conditions. Mechanisms for the elimination of supernumerary larvae varied with the relative developmental stage of the competitors and included physical combat and physiological suppression. Host instar had no effect on larval competition or the females ability to discriminate.

The scaling of body size and mass in a host-parasitoid association : influence of host species and stage

We describe the allometry of body mass and body size as measured by hind‐tibia length in males of Monoctonus paulensis (Ashmead) (Hymenoptera: Braconidae, Aphidiinae), a solitary parasitoid of aphids. To assess the influence of host quality on allometric relationships, we reared parasitoids on second and fourth nymphal instars of four different aphid species, Acyrthosiphon pisum (Harris), Macrosiphum creelii Davis, Myzus persicae (Sulzer) and Sitobion avenae (F.), under controlled conditions in the laboratory. Dry mass was positively correlated with hind‐tibia length, and could be predicted from it, in unparasitized aphids, in aphid mummies containing parasitoid pupae, and in the parasitoid. The reduced‐major‐axis scaling exponents for the regression of dry mass on hind‐tibia length were species‐specific in aphids, reflecting differences in volume and shape between species. In mummified aphids, the stage at death influenced the size/mass relationship. In males of M. paulensis, the allometric exponent varied between parasitoids developing in different kinds of host. Individuals developing in pea aphid were absolutely larger in dry mass as well as proportionately larger relative to their hind‐tibia length. We discuss the allometry of body size and body mass in relation to parasitoid fitness.

Interactions between ants attendingAphis fabae ssp.cirsiiacanthoidis on thistles and foraging parasitoid wasps

We describe the behavioral interactions between honeydew-collecting workers of the ants Lasius nigerand Myrmica ruginodisand females of three species of aphidiid wasps (Lysiphlebus cardui, Lysiphlebus testaceipes, Trioxys angelicae)foraging for their aphid host, Aphis fabaessp. cirsiiacanthoidis,on thistles. Using field and laboratory experiments, we show that the ant-parasitoid interactions are species specific. Workers of both ant species generally attacked and killed females of T. angelicae,but they ignored those of L. cardui.This pattern was not altered when we anesthetized the wasps slightly with carbon dioxide to reduce their mobility. Prior contacts between L. carduiand either conspecific L. nigerfrom a different nest or workers of a different ant species (M. ruginodis)did not influence L. nigersnonaggressive behavior. The number of aphids parasitized by L. testaceipeswas significantly reduced in aphid colonies attended by L. niger,although this parasitoid was rarely attacked by ants. In encounters between these species of ants and wasps, ant aggression is consistent with differences in wasp behavior. We suggest that, in addition, chemical cues located in the cuticula may enable L. carduito avoid detection by honeydew-collecting ants.

The use of visual cues in host evaluation by aphidiid wasps. II: Comparison between Ephedrus californicus, Monoctonus paulensis, and Praon pequodorum

We examined host evaluation behaviour in three species of aphid parasitoids, Ephedrus californicus Baker, Monoctonus paulensis (Ashmead), and Praon pequodorum Viereck (Hymenoptera: Aphidiidae). Mated females were provided with pairwise choices among three kinds of hosts in the laboratory: (green) pea aphid, Acyrthosiphon pisum (Harris), and a green and a pink colour morph of alfalfa aphid, Macrosiphum creelii Davis. Patterns of attack and host acceptance were species‐specific. Females of E. californicus did not respond to the presence of aphids prior to making antennal contact. Variations in rates of parasitization (pea aphid>green alfalfa aphid>pink alfalfa aphid) were consistent with differences in aphid defensive behaviours; no ‘preference’ for any host type was evident when aphids were anaesthetized with carbon dioxide. In M. paulensis, the order of preference (pea aphid>green alfalfa aphid>pink alfalfa aphid) did not vary when aphids were immobilized, or presented in the dark, or both. Host movement did not influence the rate of attack by M. paulensis. In contrast, the ranked order of preference in P. pequodorum varied with circumstance. In the light, females attacked pea aphid and green alfalfa aphid with equal frequency, but parasitized significantly more of the former; both kinds of aphids were attacked and parasitized at higher rates than pink alfalfa aphid. In the dark, P. pequodorum females parasitized green and pink alfalfa aphids equally and at higher rates than pea aphids. Whereas E. californicus was more successful ovipositing in immobilized hosts, P. pequodorum females attacked and laid more eggs in normal than anaesthetized aphids. Patterns of host recognition and evaluation are compared across six species representing four genera in the family Aphidiidae.

Growth and developmental interactions in some aphids and their hymenopterous parasites

Abstract The growth and development of apterous and alate morphs of viviparous pea aphids, Acyrthosiphon pisum , was compared with that of aphids parasitized at age 60 h by Aphidius smithi . The growth pattern, adult weight and time to reach the adult stage in A. smithi males were not affected by aphid morph. In unparasitized pea aphids, the dry-to-wet-weight ratio varied little during development and early adult life, while in parasitized aphids it increased in accordance with parasite development. It is proposed that, in parasites that develop in growing host stages, the development time and adult weight covary, both being influenced by the hosts size at the beginning of parasitism and its capacity to grow during parasitism. The importance of developmental interactions for successful parasitism is discussed.

The developmental strategy of an idiobiont ectoparasitoid, Dendrocerus carpenteri : influence of variations in host quality on offspring growth and fitness

Dendrocerus carpenteri (Curtis) (Hymenoptera: Megaspilidae) is a solitary hyperparasitoid, which attacks prepupal and pupal stages of hymenopteran parasitoids inside mummified aphids. The larva feeds externally on the host, which is envenomed by the female at oviposition. To evaluate the influence of variations in host quality on the growth, development and fitness of D. carpenteri, we varied the size and developmental stage of the primary parasitoid host (Aphidius ervi Haliday), which was reared on different instars of pea aphid [Acyrthosiphon pisum (Harris)] and English grain aphid [Sitobion avenae (F.)]. Within each kind of host, females eclosed from the relatively larger mummies, while males eclosed from the smaller mummies. Host size and hyperparasitoid size were correlated, and females were larger than males. In hyperparasitoids developing on prepupal and pupal hosts, development time from oviposition to adult eclosion was proportional to size; females required more time for development than males. The mean relative growth rate was the same in males and females and increased with host quality, as predicted by the growth model of Mackauer and Sequeira (1993) for idiobiont parasitoids. Larvae developing on late-pupal stages and pharate adults of A. ervi were unable to consume sclerotized host tissues; they were smaller and needed more time for development. The average number of mature eggs at eclosion was six, except in females developing on suboptimal hosts, which contained only one egg or none. Egg volume was correlated with female size, possibly reflecting differences in larval ontogeny. We provide equations describing the relationship between host quality as indexed by hind-tibia length of the mummified aphid and adult body size in terms of dry mass, development time and mean relative growth rate of D. carpenteri. We discuss the usefulness of host size as a proxy of host quality for idiobiont parasitoids, and provide examples of exceptions.

The use of visual cues in host evaluation by aphidiid wasps: I. Comparison between three Aphidius parasitoids of the pea aphid

Host evaluation behaviour was examined in three species of aphid parasitoids, Aphidius ervi Haliday, A. pisivorus Smith, and A. smithi Sharma & Subba Rao (Hymenoptera: Aphidiidae). Parasitoids were provided under laboratory conditions with three kinds of hosts representing two aphid species: (green) pea aphid, Acyrthosiphon pisum (Harris), and green and pink colour morphs of the alfalfa aphid, Macrosiphum creelii Davis. Females of all three Aphidius species distinguished between aphids on the basis of colour, movement, and host species. Patterns of host acceptance by parasitoids were species‐specific. In A. ervi, host preference was the same in light and dark conditions: pea aphid > green alfalfa aphid „ pink alfalfa aphid. In contrast, A. pisivorus attacked and accepted pea aphid and green alfalfa aphid equally in the light and preferred both of these over pink alfalfa aphid; however, it made no distinction between pea aphid and pink alfalfa aphid in the dark. Females of A. smithi attacked all three kinds of hosts (pea aphid > green alfalfa aphid „ pink alfalfa aphid) but apparently laid eggs only in pea aphid. The frequencies of attack and oviposition by all wasps were higher on ‘normal’ pea aphids than on those anaesthetized with CO2. Host recognition is confirmed by chemical cues in the aphid cuticle that are detected during antennation, and host acceptance is dependent on an assessment of host quality during ovipositor probing.

Decision to superparasitize based on larval survival: competition between aphid parasitoids Aphidius ervi and Aphidius smithi

Oviposition by a parasitoid wasp in an already parasitized host (= superparasitism) may result in larval competition and the loss of offspring. Consequently, in solitary species, the decision to superparasitize should be based on the probability of offspring survival. Females of Aphidius ervi Haliday and A. smithi Sharma & Subba Rao (Hymenoptera: Aphidiidae), two parasitoids of the pea aphid, discriminated between conspecific‐ and heterospecific‐parasitized hosts. Both species showed partial preference for aphids previously attacked by A. smithi, a result predicted from the fact that A. ervi is superior to A. smithi in larval competition under most conditions. Females of A. smithi selectively attacked conspecific‐parasitized pea aphids when given a choice between these and self‐parasitized hosts. Results indicated that females of both species responded to an external pheromone‐like marker to discriminate between self‐, conspecific‐ and heterospecific‐parasitized aphids. In addition, A. smithi may use internal cues to recognize, and avoid oviposition in, aphids previously parasitized by A. ervi.