Effects of large‐scale host plant addition and removal on parasitoid‐mediated associational resistance in the gall midge Asphondylia borrichiae

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Non-random distribution among a guild of parasitoids: implications for community structure and host survival

Abstract 1. Immature stages of the gall midge, Asphondylia borrichiae, are attacked by four species of parasitoids, which vary in size and relative abundance within patches of the gall midge’s primary host plant, sea oxeye daisy (Borrichia frutescens). 2. In the current study, a bagging experiment found that the smallest wasp, Galeopsomyia haemon, was most abundant in galls exposed to natural enemies early in the experiment, when gall diameter is smallest, while the wasp with the longest ovipositor, Torymus umbilicatus, dominated the parasitoid community in galls that were not exposed until the 5th and 6th weeks when gall diameter is maximal. 3. Moreover, the mean number of parasitoids captured using large artificial galls were 70% and 150% higher compared with medium and small galls respectively, while stem height of artificial galls significantly affected parasitoid distribution. Galls that were level with the top of the sea oxeye canopy captured 60% more parasitoids compared with those below the canopy and 50% more than galls higher than the plant canopy. 4. These non‐random patterns were driven primarily by the differential distribution of the largest parasitoid, T. umbilicatus, which was found significantly more often than expected on large galls and the smallest parasitoid of the guild, G. haemon, which tended to be more common on stems level with the top of the plant canopy. 5. Large Asphondylia galls, especially those located near the top of the Borrichia canopy, were more likely to be discovered by searching parasitoids. Results using artificial galls were consistent with rates of parasitism of Asphondylia galls in native patches of sea oxeye daisy. Gall diameter was 19% greater and the rate of parasitism was reduced by almost 50% on short stems; as a result, gall abundance was 24% higher on short stems compared with ones located near the top of the plant canopy. 6. These results suggest that parasitoid community composition within galls is regulated by both interspecific differences in ovipositor length and preferences for specific gall size and/or stem length classes.     

Biology of Asphondyliini (Diptera: Cecidomyiidae)

The family Cecidomyiidae (Diptera) including about 6100 described species displays diverse feeding habits. The tribe Asphondyliini is a well‐circumscribed monophyletic group of Cecidomyiidae and all species are known as gall inducers. Species belonging to this tribe exhibit fascinating ecological traits such as host alternation, polyphagy, extended diapause, induction of dimorphic galls and association with fungal symbionts. For these reasons, biogeographical and phylogenetic studies of Asphondyliini are of interest in elucidating the evolution of these traits, and particularly the processes of host‐range expansion, host‐plant shift and shifts in gall‐bearing organs. In order to facilitate further evolutionary studies of Asphondyliini, I review studies of systematics, biogeography, phylogeny, speciation, cytology, behavior, ecology, physiology, biological interaction and economic importance in this tribe.     

Bottom-up effects shift top-down community structure of the parasitoid guild that attacks Asphondylia borrichiae (Diptera: Cecidomyiidae) on Borrichia frutescens (Asteraceae)

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Adaptive radiation of gall-inducing insects within a single host-plant species

Speciation of plant-feeding insects is typically associated with host-plant shifts, with subsequent divergent selection and adaptation to the ecological conditions associated with the new plant. However, a few insect groups have apparently undergone speciation while remaining on the same host-plant species, and such radiations may provide novel insights into the causes of adaptive radiation. We used mitochondrial and nuclear DNA to infer a phylogeny for 14 species of gall-inducing Asphondylia flies (Diptera: Cecidomyiidae) found on Larrea tridentata (creosote bush), which have been considered to be monophyletic based on morphological evidence. Our phylogenetic analyses provide strong support for extensive within-host plant speciation in this group, and it demonstrates that diversification has involved numerous shifts between different plant organs (leaves, buds, flowers, and stems) of the same host-plant species. Within-plant speciation of Asphondylia is thus apparently facilitated by the opportunity to partition the plant ecologically. One clade exhibits temporal isolation among species, which may have facilitated divergence via allochronic shifts. Using a novel method based on Bayesian reconstruction, we show that the rate of change in an ecomorphological trait, ovipositor length, was significantly higher along branches with inferred shifts between host-plant organs than along branches without such shifts. This finding suggests that Larrea gall midges exhibit close morphological adaptation to specific host-plant parts, which may mediate ecological transitions via disruptive selection.     

Plant water stress and gall formation (Cecidomyiidae: Asphondylia spp.) on creosote bush

Abstract. 1. Populations of creosote bush ( Larrea tridentata (DC) Coville), were studied in Arizona to determine whether associated gallformers (Cecidomyiidae: Asphondylia spp.) were more abundant on water-stressed or nonstressed plants. Gall densities were measured along a steep elevational gradient that extended from mesic, higher elevations to lower elevations in the desert; and in the Grand Canyon where severely water-stressed and relatively unstressed plants occurred adjacently. At the Grand Canyon site, the responses of creosote bush to water stress were also studied.
2. The number and densities of Asphondylia species increased both at lower elevations and locally on water-stressed plants in the Grand Canyon, indicating that climatic and local conditions influence gallformer abundance in the same way.
3. Five of the eight Asphondylia species studied at the Grand Canyon site were more abundant on stressed plants, two species were more abundant on nonstressed plants and one species showed no preference for either plant type.
4. Densities of most species on stressed plants were positively correlated with the number of meristematic terminals per branch, which were more numerous on stressed plants, due to a bushier architecture. Flower gallformers were more abundant on nonstressed plants, which produced more flowers. Gall densities did not correlate with chemistry measurements, although these parameters also varied with level of stress.
5. These results suggest that gallforming species respond variably to plant stress, even within a closely-related lineage, and that there are effects of stress on plants, including architectural changes, that may be more important to herbivores than biochemical effects emphasized by White (1984) and others.     

The biology of Dasineura dielsi Rübsaamen (Diptera: Cecidomyiidae) in relation to the biological control of Acacia cyclops (Mimosaceae) in South Africa

Abstract  Acacia cyclops is an invasive Australian tree in South Africa and a target for biological control using seed-reducing agents. In southern Australia, two gall-forming Cecidomyiidae, Dasineura dielsi (Small Fluted Galler) and Asphondylia sp., develop on the flowers and seeds of A. cyclops , respectively. The larvae of D. dielsi form woody fluted galls on the ovaries of flowers and prevent the development of fruit. Immature Asphondylia sp. develop in the loculi of green fruit and destroy developing seeds. Dasineura dielsi was selected as a biological control candidate for A. cyclops in South Africa and was approved for official release after host specificity evaluation and consideration of potential conflicts of interest. Dasineura dielsi naturalised in South Africa in 2001 and after 3 years dispersed up to 450 km from a single population at Stellenbosch, Western Cape. At sites where D. dielsi has been present longest, high gall densities occur on A. cyclops during the peak flower season in summer. Four hymenopterans, ? Synopeas sp., Mesopolobus sp., Torymus sp. and an unidentified Platygastridae, were reared from D. dielsi galls and are suspected parasitoids of the cecidomyiid, with incidence levels less than 10%. Monitoring is required to evaluate trends in the population status of D. dielsi , its parasitoids and seed production of A. cyclops . Importantly, field monitoring should determine the extent and nature of possible competitive interactions between D. dielsi and an introduced seed-feeding weevil, Melanterius servulus .     

The interactions of plant clone and abiotic factors on a gall-making midge

Within and around Tampa Bay, Florida, monoclonal populations of the sea daisy, Borrichia frutescens, can be found on small, isolated islands growing within the intertidal zone. Stem tips of Borrichia are attacked by the gall-making cecidomyiid, Asphondyliaborrichiae. We used reciprocal transplants of Borrichia clones between islands to assess the importance of plant genotype and local environmental conditions (shade and host-plant nitrogen) on gall abundance. In another experiment, we controlled for host genotype effects by inducing differences in local environmental conditions through the addition of NH₄NO₃ fertilizer and/or shade to field plots at the only monoclonal site with a large enough population of Borrichia to facilitate the experiment. We also examined the effect of these variables on attack levels of Asphondylia by parasitoids. In the reciprocal transplant, while some Borrichia clones always supported more galls than others, regardless of environmental conditions, all four clones developed more galls when they were placed in the shade, compared to those in the sun, at all four sites. In addition, some islands always supported more galls than others and we found a significant clone × site interaction. In the single-clone experiment, Borrichia in fertilized- and shaded-only plots developed more Asphondylia galls than those from nonmanipulated control plots, and plants that received both shading and fertilizer developed the most galls. Although shade and fertilization produced an additive increase in plant nitrogen content, their effects resulted in a synergistic decrease in C:N ratio. Neither shading nor host plant nitrogen content had a significant effect on levels of parasitism between experimental and control plants. Our results suggest that genetic differences in Borrichia's susceptibility to Asphondylia attack are important in shaping the distribution of galls, but environmental factors such as soil nitrogen and degree of shading are at least as important as genetic differences between host plants. Received: 12 June 1997 / Accepted: 6 April 1998     

Indirect competitive effects of stemborers on a gall community

Interspecific competition between phytophagous insects using the same host plant occurs frequently and can strongly affect population densities of competing species. Competition between gallmakers and stemborers could be especially intense because both types of herbivore are unable to avoid competition by relocation during their immature stages. For apical meristem gallmakers the main result of competition is likely to be the interruption of resources to the gall by the stemborers' devouring of stem contents. The proximate effect of such competition could be to reduce gall size, thereby increasing the number of chambers per gall unit volume, and reducing the size and potential reproductive output of the gallformer. In addition, smaller galls may be more susceptible to attack from size‐limited parasitoids, resulting in a second indirect effect of competition. Using a community of galling and stemboring insects on the saltmarsh shrub Iva frutescens L. (Asteraceae), we measured for indirect effects of competition. We examined the primary indirect effect of competition on gall midge crowding and the secondary effects on parasitism rates and parasitoid guild composition. Results indicated that galls co‐occurring with stemborers were smaller, crowding of gall inhabitants was 22% greater, and the composition of the parasitoid guild was altered relative to galls on unbored stems. The overall parasitism rate was not different between galls on bored vs. unbored stems. These results show that competition resulting from the presence of stemborers has the potential to affect the gall midge Asphondylia borrichiae Rossi & Strong (Diptera: Cecidomyiidae) and secondarily to affect its guild of hymenopteran parasitoids.     

Key‐factor/key‐stage analysis of long‐term life table data for a fruit gall midge,Asphondylia sphaera (Diptera: Cecidomyiidae)

1. Population dynamics of Asphondylia sphaera Monzen (Diptera: Cecidomyiidae), a species that induces fruit galls on Ligustrum japonicum (Oleaceae), was studied from 1970 to 1996 in broad‐leaved evergreen forests in Kagoshima, southern Japan. The numbers of fruit galls and emerging adults fluctuated greatly from year to year along with alternate year flowering of the host plant. 2. To detect density‐dependent and independent forces operating on the A. sphaera population and to assess the relative strength of top‐down and bottom‐up effects, we used the method of Key‐factor/key‐stage analysis, which allowed us to avoid various problems of the conventional key‐factor analysis. 3. Five factors and seven stages were distinguished in the life tables. Key‐factor/key‐stage analysis indicated that the number of flower buds made the largest contribution to the annual changes in total survival rate by operating through the proportion of eggs and first instars that survived abortion of host flower buds and flowers, which contributed most to density‐dependency. 4. The population dynamics of A. sphaera has been strongly influenced by the bottom‐up effects of both the abundance of flower buds and the abortion of flowers of L. japonicum. 5. A top‐down effect of Bracon asphondyliae Maeto (Hymenoptera: Braconidae) was weakly density‐dependent and contributed less to the total survival rate. Abiotic factors, such as summer and winter temperatures and the effect of typhoons were negligible.