A sex-specific tri-trophic-level effect in a phoretic association

Hemisarcoptes coccophagus (Acari: Astigmata: Hemisarcoptidae) is an obligate parasite of armored-scale insects (Homoptera: Diaspididae). Hypopodes (non-feeding heteromorphic deutonymphs) of this mite cannot complete their development without a phoretic sojourn on the coccinellid beetle Chilocorus bipustulatus. We tested whether the feeding history of the beetles affects the probability of the hypopodes completing their development. Two diets were offered: armored-scale insects (Homoptera: Diaspididae) (suitable for both beetles and mites) and soft-scale insects (Homoptera: Coccidae) (suitable for the beetles but not for the mites). The hypopodial survival was similar for mites which had been on beetles reared on the different scales. However, female hypopodes which had stayed on beetles reared on soft scales suffered higher mortality than those from Chilocorus reared on armored scales. Male survival was not affected. A tri-trophic-level effect on sex-specific survival in a phoretic association was thus demonstrated.     

Modeling the age-specific per capita growth and reproduction ofRhizobius lophanthae (Col: Coccinellidae)

A per capita model for the growth, development and reproduction of the coccinellid predatorRhizobius lophanthae (Blaisd) feeding on the oleander scale(Aspidiotus nerii Bouche (Homoptera: Diaspididae)) was developed. A thermal threshold for development of 9.4°C was found. Under conditions of unlimited food, the relationship of mass at time t + 1 to that at t (in days at 25°C) suggests an 8.7 percent growth rate per mg larvae per day at 25°C. An adult female beetle produces approximately 20 eggs per day while consuming an average of 8.5 scales/day. This is approximately 2.16 eggs per scale consumed above the maintenance level of 1.88 scales per day. More precisely, this compensation point is 0.12 mg of prey/mg of predator/day at 25°C and the egestion rate is 1 - β = 0.63     

Intra-tree foraging behavior ofDacus dorsalis flies in relation to host fruit quantity, quality and type

Promising diets were screened and the most successful modified with additives used in artificial diets for other entomophagous insects. Two suitable diets were obtained, one for adults and one for larvae ofChilocorus nigritus (Fabricius) (Coleoptera: Coccinellidae). They were still inferior to natural prey and not adequate as the sole food source for rearing consecutive generations. They are valuable as substitute food in the insectary during shortages of natural prey. Oleander scaleAspidiotus nerii Bouché andAsterolecanium miliaris (Boisduval) were evaluated as natural prey forC. nigritus and two other potential biocontrol agents in southern Africa,C. bipustulatus (Linnaeus) andC. infernalis Mulsant.A. nerii andA. miliaris were suitable for all life stages ofC. nigritus and adults ofC. bipustulatus andC. infernalis. A. miliaris was inadequate for larvae ofC. bipustulatus andC. infernalis.     

Larval <Emphasis Type="Italic">Cryptothelea gloverii</Emphasis> (Lepidoptera: Psycidae), an arthropod predator and herbivore on Florida citrus

The orange bagworm (OBW), Cryptothelea gloverii (Packard) (Lepidoptera: Psychidae) was previously reported feeding on citrus fruit and foliage and preying upon the camphor scale Pseudaonidia duplex (Cockerell) (Homoptera: Coccidae). In this study using laboratory assays, OBW preyed upon citrus rust mite, Phyllocoptruta oleivora (Ashmead) (Acari: Eriophyidae) and consumed eggs and adults of both P. oleivora and Panonychus citri (McGregor) (Acari: Tetranychidae), two important pest mites on Florida citrus. OBW was also observed to feed on the purple scale, Lepidosaphes beckii (Newman) (Homoptera: Diaspididae) and on a fungus (Penicillium sp.). OBW fed on orange and grapefruit leaves by starting from the border and eating part of the leaf, by chewing holes, or consuming the outer epithelium of either the axial or abaxial surface of the leaf without penetrating through the leaf. OBW was observed in orange orchards in association with fruit extensively russeted by P. oleivora feeding. Laboratory assays revealed that OBW larvae preferred to feed on oranges infested with P. oleivora rather than on clean fruits that were free of mite feeding damage. Feeding damage to citrus fruit by OBW larvae results in one to several holes being eaten into the rind or albedo, without damage to the fruit sacs.     

Spatial density‐dependent parasitism and specificity in the robber fly Mallophora ruficauda (Diptera: Asilidae)

The density‐dependence in parasitism by the robber fly Mallophora ruficauda (Diptera: Asilidae) on scarab beetle larvae (Coleoptera: Scarabaeidae) populations was studied in the present research. Mallophora ruficauda is a pestiferous species common in the open grasslands of the Pampas region of South America. Adults are predators of insects and larvae are solitary parasitoids of third instar larvae of several species of scarab beetle (Coleoptera: Scarabaeidae). In contrast with most studied host‐parasitoid interactions, host searching by M. ruficauda is carried out by both larvae and adults. Typically, robber fly females lay eggs on tall grasses from where larvae drop to the ground, and attack hosts which are buried in the soil. We carried out our study at two spatial scales close to 14 apiaries located in the provinces of Buenos Aires and Entre Ríos (Argentina). We found that parasitism is density‐independent at the larger spatial scale and inversely density‐dependent at the smaller one. We also found that M. ruficauda selects Cyclocephala signaticollis among several scarab beetle species. Specificity is observed both at large and small spatial scales. We discuss the implications of both host specificity and host searching behaviour on the observed parasitism patterns.     

Divergent preferences of Rhagoletis pomonella host races for olfactory and visual fruit cues

The quality of different species of aphids as food for aphidiphagous ladybird beetles varies greatly. The optimal oviposition theory predicts that a female should lay eggs preferentially in patches of suitable prey and should be reluctant to oviposit in patches of less suitable prey. A no‐choice experiment was used to test whether aphids (Homoptera: Aphididae) of different suitability influence the oviposition behaviour of the two‐spot ladybird beetle, Adalia bipunctata (L.) (Coleoptera: Coccinellidae). The results indicate that A. bipunctata females are not more reluctant to lay eggs in the presence of moderately suitable compared to highly suitable aphids. However, females laid fewer eggs in the presence of toxic aphids.     

Differential predation by Coleomegilla maculata on Colorado potato beetle strains that vary in growth on tomato

This study tests the hypothesis that the generalist predator Coleomegilla maculata DeGeer causes differential mortality of Colorado potato beetle, Leptinotarsa decemlineata (Say), larvae differing in their degree of genetic adaptation to tomato (Lycopersicon esculentum Mill.) as a host plant. Results of a series of laboratory experiments demonstrate that adult C. maculata can cause higher mortality to nonadapted than adapted Colorado potato beetle larvae. The extent of differential mortality caused by C. maculata depended on age of potato beetle larvae; presence of potato beetle eggs; whether or not the predator had a choice among prey items; and, in choice situations, the ratio of adapted to nonadapted potato beetle larvae. Although adult C. maculata have the potential to prey differentially on tomato-adapted and nonadapted Colorado potato beetle larvae in mixed populations, the magnitude of differential predation in a natural setting could be highly variable.     

Dietary complementation across life stages in the polyphagous lady beetle Coleomegilla maculata

We investigated the life history consequences of changes in diet between larval and adult life stages in the polyphagous lady beetle Coleomegilla maculata DeGeer (Coleoptera: Coccinellidae). Beetles were reared on three larval diets: greenbug, Schizaphis graminum Rondani (Homoptera: Aphididae), eggs of the flour moth, Ephestia kuehniella Zeller (Lepidoptera: Pyralidae), and bee pollen. The reproductive performance of females was then evaluated on an adult diet of either greenbug or moth eggs. Moth eggs appeared to be the most suitable diet for larvae, yielding the largest adults, and pollen the least suitable, resulting in the smallest adults and greatly extended developmental time. Pollen‐reared beetles tended to have lower fecundity and fertility than those reared on animal protein, regardless of adult diet. Female fitness was generally increased by a change in diet upon emergence to the alternative source of animal protein, suggesting that dietary complementation occurred across life stages. Among females reared on greenbug, a change of diet to moth eggs reduced the period required for production of 12 clutches and increased egg fertility compared to continued feeding on greenbug. Among females reared on moth eggs, a change of diet to greenbug increased fecundity compared to continued feeding on moth eggs. Among females fed an adult diet of greenbug, those fed moth eggs as larvae had faster production of 12 clutches and higher fecundity. We discuss these novel results in the context of coccinellid life history and ecology and their potential implications for other insects that are predatory as both larvae and adults.     

Effects of temperature and host age upon the encapsulation of Metaphycus stanleyi and Metaphycus helvolus eggs by brown soft scale Coccus hesperidum

Encapsulation of eggs inserted by Metaphycus stanleyi (Hymenoptera: Encyrtidae) into the brown soft scale Coccus hesperidum (Homoptera: Coccidae) became more frequent as the host matured. This occurred with both laboratory reared and field-collected parasites. After parasitism for 24 hr at 27°C, encapsulation frequency did not differ in hosts reared at 20° or at 27°C, but significantly increased in hosts reared at 33°C. When parasitism and rearing were carried out at the same temperature, the percentage of eggs encapsulated increased from 48.7% at 27°C to 94.1% at 33°C. With M. helvolus, the percentage of eggs encapsulated was considerably higher than with M. stanleyi; e.g., 99.3 vs 48.7%, respectively, at 27°C. At 20° and 27°C, some M. helvolus development occurred in the larvae of brown soft scale but none at 33°C; the adult stages of the host encapsulated all the parasite eggs at these temperatures.     

Feeding range studies of Rodolia cardinalis (Mulsant), a candidate biological control agent of Icerya purchasi Maskell in the Galápagos islands

The immediate threat of the cottony cushion scale, Icerya purchasi Maskell (Homoptera: Margarodidae), to the conservation of endangered flora in the Galápagos islands prompted conservation groups to assess the risks associated with the introduction of its natural enemy, Rodolia cardinalis (Mulsant) (Coleoptera: Coccinellidae). Although R. cardinalis has been widely used for controlling this exotic pest, little information was found to confirm its presumed narrow feeding range. Consequently, studies were deemed necessary to determine whether the introduction of R. cardinalis would harm the island’s native invertebrate fauna, in particular rare or threatened species. Using no-choice trials, we tested neonate and third instar larvae of R. cardinalis against 16 and 11 potential prey species, respectively. Adults with prior feeding experience on I. purchasi were tested against eight non-target species and naïve adults (those that had not fed on I. purchasi) were tested against six. These trials included up to 35% of the Homoptera species of conservation value presumed to have the highest risk of being preyed upon by R. cardinalis. To maximize the range of species exposed to R. cardinalis, feeding trials were also carried out with some introduced species representative of groups containing potential non-target species that were not located for testing. R. cardinalis was unable to complete its life cycle on any of the test prey species and only fed on Margarodes similis Morrison (Homoptera: Margarodidae), a species closely related to the cottony cushion scale. M. similis, however, is subterranean and in its natural habitat is not at risk from foraging by R. cardinalis. Based on these trials, we believe that immature stages of R. cardinalis will have no impact on the non-target invertebrate fauna of the Galápagos islands because they specialize on Margarodidae. Although the limited nature of our testing prevents us from reaching a definitive conclusion about the prey range of R. cardinalis adults, our results indicate that it is also narrow. According to our field and laboratory studies, niche overlap with native predators of Homoptera will be minimal and intraguild predation should not occur.     

Suitability of exotic and native lady beetle eggs (Coleoptera: Coccinellidae) for development of lady beetle larvae

Predation upon lady beetle (Coleoptera: Coccinellidae) eggs in the field is most often instances of egg cannibalism by larvae or adults while the majority of the remaining predation events upon coccinellid eggs is done by other species of Coccinellidae. Thus the recent introduction and establishment of Harmonia axyridis in the US could negatively affect native species of Coccinellidae via egg predation. However, little is known regarding the suitability of interspecific coccinellid eggs as a food source for larval development. In this study, it was found that native first or third instar Coleomegilla maculata and Olla v-nigrum larvae were incapable of surviving to the adult stage when provided solely exotic H. axyridis eggs. In stark contrast, H. axyridis larvae survived equally well when cannibalizing eggs or eating eggs of either native species. When C. maculata and O. v-nigrum were grouped as ‘native’ and compared with the exotic H. axyridis, more native eggs were attacked than exotic eggs and a higher percentage of eggs was attacked by H. axyridis larvae. Native and exotic larvae attacked a similar percentage of native eggs but native larvae attacked significantly fewer exotic eggs than did exotic larvae. These data suggest that H. axyridis may prey upon the eggs of these native species, when encountered in the field, compared with the likelihood of the native species preying upon H. axyridis eggs. Therefore, eggs of the native species C. maculata and O. v-nigrum will continue to be subjected to cannibalism and also to possible predation by other native species and the exotic H. axyridis.     

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

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

Larval Cryptothelea gloverii (Lepidoptera: Psycidae), an arthropod predator and herbivore on Florida citrus

The orange bagworm (OBW), Cryptothelea gloverii (Packard) (Lepidoptera: Psychidae) was previously reported feeding on citrus fruit and foliage and preying upon the camphor scale Pseudaonidia duplex (Cockerell) (Homoptera: Coccidae). In this study using laboratory assays, OBW preyed upon citrus rust mite, Phyllocoptruta oleivora (Ashmead) (Acari: Eriophyidae) and consumed eggs and adults of both P. oleivora and Panonychus citri (McGregor) (Acari: Tetranychidae), two important pest mites on Florida citrus. OBW was also observed to feed on the purple scale, Lepidosaphes beckii (Newman) (Homoptera: Diaspididae) and on a fungus (Penicillium sp.). OBW fed on orange and grapefruit leaves by starting from the border and eating part of the leaf, by chewing holes, or consuming the outer epithelium of either the axial or abaxial surface of the leaf without penetrating through the leaf. OBW was observed in orange orchards in association with fruit extensively russeted by P. oleivora feeding. Laboratory assays revealed that OBW larvae preferred to feed on oranges infested with P. oleivora rather than on clean fruits that were free of mite feeding damage. Feeding damage to citrus fruit by OBW larvae results in one to several holes being eaten into the rind or albedo, without damage to the fruit sacs.     

Selection for pesticide resistance in Aphytis

Field-collected populations of Aphytis holoxanthus DeBach (Hymenoptera: Aphelinidae), a parasite of Chrysomphalus aonidum (L.) (Homoptera: Diaspididae), were tested for malathion and for azinphosmethyl tolerance, and selected for increased resistance. Initial tolerance for malathion was not significantly different between populations, and could not be significantly increased by 30 selection cycles. Two field populations showed relatively high levels of azinphosmethyl tolerance, which were further increased by selection to 24–48 times that of a susceptible laboratory population.The differences in response to the two organophosphorus insecticides are discussed in relation to differences in their modes of application, detoxification by insects and the genetic bottleneck experienced by an imported species.     

Effect of beetle size on predation of cabbage root fly eggs by ground beetles

Laboratory studies were done to determine the numbers of cabbage root fly (Delia radicum L.) (Diptera: Anthomyiidae) eggs eaten by sixty species of ground beetles (Coleoptera: Carabidae) collected from a range of habitats. For 35 species that were between 2.7 mm and 10 mm long, there was a linear relationship between the numbers of eggs eaten and beetle length. For each 1 mm increase in length above 2.7 mm, an additional 18 eggs were eaten/beetle/day. Eight further species within the above size range did not eat any fly eggs, as these species are known to be either phytophagous or to feed on moving prey. Large numbers of eggs were eaten by beetles in 13 of the 25 genera tested. Based on size, maximum numbers of eggs were eaten by 7 species of Agonum Bonelli, 5 species of Amara Bonelli and seven species of Bembidion Latreille. Only one of the 6 species of Pterostichus Bonelli tested, P. strenuus (Panz.), ate the expected numbers of eggs, the other species being too large to feed from such small prey items as cabbage root fly eggs. The numbers of beetles larger than 10 mm that ate eggs was highly variable and so 10 mm was considered the upper size limit of carabid predators of cabbage root fly eggs.     

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.     

Handling time, prey preference, and functional response for Chrysoperla rufilabris in the laboratory

The predaceous larvae of Chrysoperla rufilabris (Burmeister) exhibited some success-motivated searching, particularly when feeding on Heliothis virescens (F.) eggs, but handling time did not decrease with experience. Handling time for H. virescens larvae was more than twice that for eggs. H. virescens larvae were preferred to cotton aphids (Aphis gossypii (F.)) while aphids were preferred to H. virescens eggs. C. rufilabris larvae exhibited a linear functional response to the three prey types tested, over the prey densities tested.USDA, ARS, Retire. Present address: 200 Highland, College Station, TX 77840, USA     

Benthic ctenophores (Platyctenida: Coeloplanidae) in South Florida: predator–prey interactions

The primary goal of this study was to demonstrate, from field observations and laboratory experiments, some key trophic roles of benthic ctenophores as predators and prey in subtropical communities. We examined individuals of two benthic platyctenid species: Coeloplana waltoni, a minute epibiont on octocorals in exposed, open‐water settings; and Vallicula multiformis, an associate of calm‐water biofouling communities and floating Sargassum spp. Laboratory observations of individuals of both ctenophore species revealed frequent capture and ingestion of diverse zooplankton taxa, especially crustaceans. Laboratory predation trials demonstrated the capture of dolphinfish (Coryphaena hippurus) eggs and larvae by both ctenophore species. Dolphinfish eggs and larvae larger than individuals of C. waltoni were captured but not ingested during 2‐h trial periods. These prey items were sometimes purloined and ingested by polyps of the ctenophore's octocoral host. Ingestion of dolphinfish eggs and larvae by individuals of C. waltoni was observed, however, after longer periods of exposure to prey. In predation trials, dolphinfish eggs and larvae were both captured and ingested by larger individuals of the ctenophore species V. multiformis. Field and laboratory observations revealed diverse invertebrate and fish taxa that prey on both ctenophore species. In the laboratory, the mean daily per capita consumption of individuals of C. waltoni by a pomacanthid fish ranged 0.5–2.8 individuals, and ranged 2.6–3.6 individuals for predation by an ovulid mollusc. Field population densities of these predators ranged 0.1–0.7 individuals per m² for the pomacanthid, and 0.2–1.1 individuals per m² for the mollusc. Laboratory feeding observations demonstrated frequent consumption of individuals of V. multiformis by a sea anemone, and by three species of brachyuran crabs. Field observations revealed eight fishes that probably feed incidentally on individuals of V. multiformis. These findings add to the limited knowledge base of predator–prey dynamics in both C. waltoni and V. multiformis.     

Oviposition in Cryptolaemus montrouzieri stimulated by wax filaments of its prey

The oviposition responses of Cryptolaemus montrouzieri Mulsant (Coleoptera: Coccinellidae) to the soft scale Eupulvinaria hydrangeae (Steinweden) (Homoptera: Coccidae) and to the mealybug Planococcus citri (Risso) (Homoptera: Pseudococcidae) have been compared in the laboratory. The females delay oviposition and withhold mature eggs in their lateral oviducts in the absence of wax filaments produced by the prey (only present in the ovisac of E. hydrangeae, present in all stages of P. citri). Contact chemical cues perceived by females when probing the wax filaments with their mouthparts are the signals inducing the search for oviposition sites. The second step is under the control of the ovipositor by which females locate confined sites to lay eggs. This oviposition behaviour could have a considerable impact on the prey exploitation strategy of this important biocontrol agent and might help to understand its apparent ineffectiveness in situations of low prey density.     

Effects of augmentative releases of the coccinellid, Adalia bipunctata, and of insecticide treatments in autumn on the spring population of aphids of the genus Dysaphis in apple orchards

The impact of augmentative releases of indigenous predators and insecticide applications to control the autumn aphid forms of the genus Dysaphis (Homoptera: Aphididae), major pest insects on apple trees, was assessed in one-year field experiments in Switzerland. Eggs and larvae of the two-spot ladybird beetle Adalia bipunctata (L.) (Coleoptera: Coccinellidae) were released on 4-year old apple trees in various numbers at five different dates in autumn 1998 when sexuales of the aphids were present. Additionally, Pyrethrum HP was sprayed at the same five dates to compare the effectiveness of these augmentative releases to a commonly applied insecticide. Augmentative releases of larvae before mid-October significantly prevented the deposition of overwintering eggs by aphids of the genus Dysaphis and consequently reduced the number of hatched fundatrices in spring, 1999. There was a significant negative functional response among the number of released coccinellid larvae in autumn 1998 and the number of observed fundatrices on apple trees in spring 1999. Applications of Pyrethrum HP before mid-October were more effective than augmentative releases of larvae of A. bipunctata. The release of coccinellid eggs had no impact on the number of fundatrices of the genus Dysaphis in the next spring because they did not hatch due to bad weather conditions. The weather conditions in autumn seemed to have an impact on the autumn migration of the winged aphids back to their primary host. The prevention of egg deposition of aphids in autumn is a promising control strategy and deserves further exploration for practical use.