Long‐term compositional responses of a South African mesic grassland to burning and mowing

Question: What is the long‐term compositional response of grass and forb species to various combinations of burning and mowing? Can these responses be predicted from simple plant traits? Location: Ukulinga research and training farm (24°24′E, 30°24′S), Pietermaritzburg, KwaZulu‐Natal, South Africa. Methods: Grass species relative abundance in 1996 in various burning and mowing treatments of a long‐term (> 50 a) experiment was calculated from data obtained using a point sampling method, whereas forb species abundance in 1999 was determined using the importance score method. The experiment consisted of different frequencies (annual, biennial and triennial) of burning and mowing in winter or spring in combination with different frequencies of summer mowing (none, early, late or both). Results: Grasses responded to the type of disturbance (burn or mow) and frequency of burning, whereas forbs responded primarily to the presence or absence of any form of disturbance and secondarily to the timing of burning (spring versus winter). Summer mowing and annual or biennial dormant‐period burning maintained communities dominated by short grasses, whereas tall grasses dominated under annual dormant‐period mowing, triennial burning or protection from disturbance. Grass tillering strategy (below‐ or above‐ground) influenced response to burning frequency. Many erect herbaceous dicot species with aerial meristems were reduced in abundance by summer mowing whereas most small creeping herbaceous dicot species appeared to be dependent upon summer mowing. Conclusions: This long‐term experiment demonstrated that: (1) grasses and forbs responded differently to burning and mowing; (2) simple plant traits, such as height, position of tiller initiation, and position of meristems have potential for predicting the response of species to the timing and frequency of burning and mowing.     

The effects of pruning,fertilizers, and organic amendments on lowbush blueberry production

A four-year field study was conducted on a Hebert gravelly sandy loam (pH 4.5) in Nova Scotia to assess the effects of pruning management and seven fertility amendments on lowbush blueberry (Vaccinium augustifolium Ait) production (yield, above ground and root tissue composition) and soil fertility. Pruning by oil burning produced higher fruit yields than flail mowing but burning had the opposite effect on the plant N content (with a lesser influence on above ground Mn and Zn). None of the fertility treatments (chicken manure, dairy manure, swine manure, urea, sawdust, NPK, NPK+S+Lime+Micronutrients) produced fruit yields significantly greater than the control. Treatments provided the equivalent of 50kg total N/ha/2-yr cycle. Treatments influenced tissue N, P, K, Ca, Mg, B, Mn, Cu, Zn and Mo levels. In general, the three manure treatments produced the highest levels of plant macronutrients; the urea treatment produced the lowest levels of plant nutrients. In most cases, extractable levels of soil P, K, Ca and Mg were highly correlated with the plant tissue content of these elements. Overall, the dairy manure treated soils were the highest in soil fertility.     

Impact of whole-canopy and systemic insecticidal treatments on Callirhytis cornigera (Hymenoptera: Cynipidae) and associated parasitoids on pin oak

The gall wasp Callirhytis cornigera (Osten Sacken) is a cynipid with alternating generations that produce large, woody stem galls and tiny blister-like leaf galls on pin oak, Quercus palustris Muenchhausen, in the United States. We tested 3 approaches to control the leaf-galling generation, and determined their impact on associated parasitoids and effectiveness in reducing numbers of new stem galls. First, trees were sprayed with bifenthrin or chlorpyrifos in late March to kill females emerging from stem galls before they oviposited into buds. Second, concentrated solutions of abamectin, imidacloprid, or bidrin were injected from pressurized containers into tree sapwood to control larvae developing in young leaf galls. Finally, systemic insecticides (acephate, abamectin, dimethoate, or imidacloprid) were sprayed at early leaf expansion (2 May) or to young, expanded leaves (17 May) to target larvae in leaf galls. Parasitoids, mostly eulophids, accounted for approximately 70% mortality of leaf-galling C. cornigera larvae on untreated trees. Whole-canopy sprays during C. cornigera emergence from stem galls reduced overall numbers of galled leaves and leaf galls. Trunk injections of bidrin or abamectin resulted in significant mortality of gall inhabitants, including parasitoids. However, neither of the aforementioned approaches significantly reduced numbers of new stem galls. Sprays of abamectin, dimethoate, or imidacloprid applied on 2 May caused high mortality of all gall inhabitants. There was no net benefit, however, because parasitism caused a similar reduction in C. cornigera survival on unsprayed shoots. Sprays applied later in leaf expansion had little impact on gall inhabitants. Of the treatments tested, bifenthrin sprays at bud break provided the greatest reduction in new leaf galls, whereas bidrin injections provided the greatest reduction in gall wasps emerging from galled leaves. This study suggests that gall wasp outbreaks are unlikely to be controlled by a single treatment, regardless of application method.     

Annual burning of semi‐natural grasslands for conservation: winners and losers among plant species

Species‐rich semi‐natural grasslands in Europe are a main target for conservation efforts, and alternative methods to the traditional management of mowing or grazing would be welcome due to the difficulties in maintaining traditional management practices. One such method proposed is burning of grassland vegetation during late winter or spring. To evaluate the effects of annual spring burning vs annual mowing on semi‐natural grassland vegetation, we compared the frequency of species in eleven field experiments in southern Sweden after ca 14 years. Out of the 88 species analyzed, five were more frequent in burnt plots compared with mowed plots (Vicia cracca, Cirsium arvense, Urtica dioica, Galium verum, Convallaria majalis). In contrast, 37 species were significantly less frequent in burnt plots compared with mowed ones, those with the largest differences being Ranunculus acris, Briza media, Veronica chamaedrys, Festuca ovina, Plantago lanceolata and Anthoxanthum odoratum. Tall‐grown species and those with preferences for N‐rich soils increased in frequency under an annual spring‐burn regime, compared with annual mowing, as did species producing larger amounts of nectar. Hence, although vegetation composition becomes more trivial with annual spring burns, there might be long‐term benefits for nectar‐feeding insects.     

Ant mutualists alter the composition and attack rate of the parasitoid community for the gall wasp Disholcaspis eldoradensis (Cynipidae)

Abstract.  1. The strength or density dependence of pairwise species interactions can depend on the presence or absence of other species, especially potential mutualists.
2. The gall wasp Disholcaspis eldoradensis induces plant galls that secrete a sweet honeydew from their top surfaces while the wasp larvae are active. These galls are actively tended by Argentine ants, which collect the honeydew and drive off parasitoids attempting to attack the gall wasp.
3. When ants were excluded, the total rate of parasitism by seven species of parasitoids increased by 36%, and the rate of gall-wasp emergence decreased by 54%.
4. The total percentage parasitism was affected by gall density when ants were excluded but not when ants were unmanipulated, suggesting a change in parasitoid functional responses due to ant tending.
5. In addition, excluding ants significantly altered the proportions of different parasitoid species that emerged from galls; one parasitoid species increased from 1% to 34%, and another decreased from 46% to 19%.
6. The invasive Argentine ants studied are capable of maintaining the mutualism with the gall wasps that evolved in the presence of different ant species and also act as a selective filter for the local community of generalist parasitoids trying to attack this gall species.     

Mowing Reduces Exotic Annual Grasses but Increases Exotic Forbs in a Semiarid Grassland

Cheatgrass (Bromus tectorum) and other exotic winter‐active plants can be persistent invaders in native grasslands, growing earlier in the spring than native plants and pre‐empting soil resources. Effective management strategies are needed to reduce their abundance while encouraging the reestablishment of desirable native plants. In this 4‐year study, we investigated whether mowing and seeding with native perennial grasses could limit growth of exotic winter‐actives, and benefit growth of native plants in an invaded grassland in Colorado, United States. We established a split‐plot experiment in October 2008 with 3 mowing treatments: control, spring‐mowed, and spring/summer‐mowed (late spring, mid‐summer, and late summer), and 3 within‐plot seeding treatments: control, added B. tectorum seeds, and added native grass seeds. Cover of plant species and aboveground biomass were measured for 3 years. In March and June of 2010, 2011, and March of 2012, B. tectorum and other winter‐annual grasses were half as abundant in both mowing treatments as in control plots; however, cover of non‐native winter‐active forbs increased 2‐fold in spring‐mowed plots and almost 3‐fold in spring/summer‐mowed plots relative to controls. These patterns remained consistent 1 year after termination of treatments. Native cool‐season grasses were most abundant in spring‐mowed plots, and least abundant in control plots. There was higher cover of native warm‐season grasses in spring/summer‐mowed plots than in control plots in July 2011 and 2012. The timing of management can have strong effects on plant community dynamics in grasslands, and this experiment indicates that adaptive management can target the temporal niche of undesirable invasive species.     

Biomass Yield and Composition of Switchgrass Bales on Marginal Land as Influenced by Harvest Management Scheme

Switchgrass (Panicum virgatum L.) is well suited to marginal croplands, but is difficult to manage sustainably both for maximum yield and optimal biomass composition. Quality can be improved by overwintering switchgrass in the field, but more information is needed on amount and consistency of yield recovery in spring. Two cultivars of switchgrass were sown on separate fields in Freeville, NY, and mowed and baled in late fall (FALL), mowed in fall and baled in spring (WINTER), or mowed and baled in spring (SPRING), using conventional field harvesting equipment. Samples were collected for analysis of plant morphological components and for determining the influence of harvest stubble height on yield and composition. Recovery of FALL biomass yields the following spring ranged from 52 to 82% and was related to both total winter snowfall and to the spring date when soil was dry enough to allow equipment traffic. Approximately 1% of dry matter yield was left in the field for each centimeter of stubble height following mowing. Bale moisture content was very low in spring, averaging 7.3%, but was much more variable and higher in the fall, averaging 22% for “Cave-in-Rock”. Inflorescence and leaf blade were the primary morphological components lost in standing switchgrass over winter. The SPRING treatment can be mowed and baled on the same day without other field operations and has higher quality than WINTER, with no consistent yield advantage for either spring baling treatment. The large and variable yield loss due to overwintering switchgrass in the field makes the practice questionable.     

Effects of origin and experience on patterns of host acceptance by the opiine parasitoid Diachasmimorpha tryoni

1. Patterns of host acceptance by Diachasmimorpha tryoni (Ashmead), a parasitoid of tephritid flies, were evaluated in relation to host–substrate complex, wasp origin, and wasp experience. 2. Naive female D. tryoni originating both from the Mediterranean fruit fly (medfly) Ceratitis capitata (Wiedemann) and the lantana gall fly Eutreta xanthochaeta Aldrich probed medfly-infested coffee fruit two to six times more often than E. xanthochaeta-inhabited lantana galls. No significant differences were detected between the two groups of parasitoids in patterns of probing response to medfly-infested coffee fruit or to E. xanthochaeta galls. 3. An 18-h pretest exposure to medfly-infested coffee fruit or E. xanthochaeta-inhabited galls affected the probing response of D. tryoni to E. xanthochaeta galls significantly, but did not affect the probing response to medfly-infested coffee fruit. Diachasmimorpha tryoni exposed to E. xanthochaeta galls probed E. xanthochaeta galls two to three times more often than naive wasps, and seven to 11 times more than wasps exposed to medfly-infested coffee fruit. Regardless of the prior exposure treatments, a high proportion (75–100%) of the test parasitoids probed medfly-infested coffee fruit. 4. Parasitoid acceptance of less-preferred hosts or host–substrate complexes may be more amenable to conditioning through prior experience (i.e. learning) than preferred host–substrate complexes. The relevance of these findings to host range expansion of parasitoids used in fruit fly biological control is discussed.     

Dogwood borer (Lepidoptera: Sesiidae) infestation of horned oak galls

Pin oak, Quercus palustris Muenchhausen, is the primary host for the gall wasp Callirhytis cornigera (Osten Sacken). Woody stem galls formed by C. cornigera may be infested by the dogwood borer, Synanthedon scitula (Harris), an important pest of flowering dogwood, Cornus florida L. Previous research has shown that S. scitula has a bimodal seasonal flight pattern, with peaks in late spring and midsummer. We tested the hypothesis that moths emerging from dogwoods largely account for the first flight pulse, whereas emergence from stem galls contributes disproportionately to the second pulse. Seasonal flight activity of S. scitula was monitored with pheromone traps baited with Z,Z-3,13-octadecadien-1-ol acetate. Traps were hung near plantings of dogwoods in suburban landscapes or near heavily galled pin oaks. Borer emergence from dogwood was monitored by sampling infested trees for pupal exuviae, and from galls that were collected and held in outdoor rearing cages. The impact of S. scitula on C. cornigera larvae was assessed by weighing, measuring, and dissecting galls. Flight activity of S. scitula began on 5 May and ended on 13 October 1999, with peaks in late May and in late July to early August. The flight pattern was similar for the two types of trapping sites, and moths emerged from both hosts during both flight periods. Proportionately more moths emerged from dogwoods during the first flight pulse than during the second, but emergence from galls was nearly evenly divided between the two flight peaks. We therefore reject the hypothesis that emergence of borers from galls contributes disproportionately to the second flight period. Approximately 12-15% of stem galls (2-3 yr old) contained S. scitula larvae. Feeding and tunneling by borers contributed to gall desiccation and reduced horn development, but rarely killed C. cornigera larvae. This study has implications for management of S. scitula because borers emerging from horned oak galls may represent a threat to dogwood.     

Evaluation of habitat management strategies for the reduction of malaria vectors in northern Belize.

Mowing and burning of emergent vegetation were evaluated as potential management strategies for the control of the malaria vector, Anopheles vestitipennis, in northern Belize, Central America. The primary aim was reduction of tall dense macrophytes (dominated by Typha domingensis) as preferred larval habitat for An. vestitipennis. Nine experimental plots were established in a Typha marsh in Orange Walk District, Belize. Three plots were burned, three were treated by subaquatic mowing, and three were unaltered controls. After treatment, Typha height was most dramatically affected by the mow treatment. Plant heights at 21 and 95 days post-treatment reflected an 89% and 48% decrease, respectively, compared to pretreatment conditions. The Typha height in the burn plots was not as severely affected. Heights at 21 days post-treatment were 39% lower than those of pre-treatment vegetation, with a return to near pre-test heights by 95 days post-treatment. Both treatments resulted in a significant reduction in the number of An. vestitipennis larvae collected as compared to control plots. Conversely, the treatments resulted in increased larval densities of several other vector and pest mosquito species. Larval population densities ofAn. albimanus, Ochlerotatus taeniorhynchus, and Culex coronator were significantly higher in burn plots. In mow plots, there were significant increases in An. albimanus and Oc. taeniorhynchus larval populations. Non-target invertebrate species affected by the treatments were adult Tropisternus collaris, larval Corythrella, and adult Parapleapuella.     

Partitioning of herbivore hosts across time and food plants promotes diversification in the Megastigmus dorsalis oak gall parasitoid complex

Communities of insect herbivores and their natural enemies are rich and ecologically crucial components of terrestrial biodiversity. Understanding the processes that promote their origin and maintenance is thus of considerable interest. One major proposed mechanism is ecological speciation through host‐associated differentiation (HAD), the divergence of a polyphagous species first into ecological host races and eventually into more specialized daughter species. The rich chalcid parasitoid communities attacking cynipid oak gall wasp hosts are structured by multiple host traits, including food plant taxon, host gall phenology, and gall structure. Here, we ask whether the same traits structure genetic diversity within supposedly generalist parasitoid morphospecies. We use mitochondrial DNA sequences and microsatellite genotypes to quantify HAD for Megastigmus (Bootanomyia) dorsalis, a complex of two apparently generalist cryptic parasitoid species attacking oak galls. Ancient Balkan refugial populations showed phenological separation between the cryptic species, one primarily attacking spring galls, and the other mainly attacking autumn galls. The spring species also contained host races specializing on galls developing on different host‐plant lineages (sections Cerris vs. Quercus) within the oak genus Quercus. These results indicate more significant host‐associated structuring within oak gall parasitoid communities than previously thought and support ecological theory predicting the evolution of specialist lineages within generalist parasitoids. In contrast, UK populations of the autumn cryptic species associated with both native and recently invading oak gall wasps showed no evidence of population differentiation, implying rapid recruitment of native parasitoid populations onto invading hosts, and hence potential for natural biological control. This is of significance given recent rapid range expansion of the economically damaging chestnut gall wasp, Dryocosmus kuriphilus, in Europe.     

A nectar-secreting gall wasp and ant mutualism: selection and counter-selection shaping gall wasp phenology, fecundity and persistence

Abstract.

• 1 The natural history of a gall wasp including interactions with inquilines, parasites, and a mutualistic ant are examined. The stability of the system is described from the perspective of influences on gall wasp life history characteristics.
• 2 An exclusion experiment demonstrated that the nectar-secreting gall of Disholcaspis perniciosa mediates a mutualism with the tending ant, Formica obscuripes. Survivorship increased from 0% in the absence of ants to 25.3% in their presence, largely due to the exclusion of inquilines.
• 3 Specialized parasites, Eudecatoma spp., attacked before the ant-gall interaction began, when the developing gall was still beneath the host plant (Quercus gambellii) epidermis and ants were not in attendance. They may select for later developing gall wasps, which benefit by having fewer individuals parasitized. However, counter-selection for earlier development may result from decreased gall wasp size, decreased fecundity, and an increase in gall failures resulting from late development.
• 4 Local persistence of the gall wasp population despite increased pressure from inquilines and parasites was attributed to gall wasp escape in time due to polymorphic emergence resulting from diapause. Most individuals emerge at the end of the summer, but approximately 15% remain in the galls as prepupae for 1–5 years.

     

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.     

Factors affecting components of fitness in a gall-making wasp (Cynips divisa Hartig)

Summary The galls of the agamic generation of the cynipid gall-wasp Cynips divisa were studied; these galls are found only on the main veins of the leaf. Galls are concentrated on the middle veins of leaves, and generally occur closer to the midrib than to the edge of the leaf. Survival of the gall-maker and fecundity of the energing wasp depend on wasp size and position of the gall. There is a striking pattern in gall size according to the number of galls on the vein. The number of galls on a vein appears therefore to have a strong effect on potential fecundity.     

Seasonal acquisition of mineral nutrients by a chalcid gall on lowbush blueberry

Seasonal accumulation of various mineral nutrients by galls of the chalcidHemadas nubilipennis on lowbush blueberry were examined in various phases of gall development. Levels of copper, nickel iron and zinc were higher in tissues of galls in initiation phase compared to tips of ungalled shoots and in general were higher than galls in growth and maturation phases. Levels of manganese and magnesium in gall tissues decrease throughout the season compared to ungalled shoots. Levels of calcium are less in gall tissue in initiation phase compared to ungalled shoots, and elevated in maturation phase. Patterns of mineral concentration within galls at various phases of development are related to known physiological roles of minerals in plant nutrition; however we conclude that patterns are variable and their significance in gall physiology requires further study.     

The effect of hosptlant and parasitoids on the reproductive success of the parthenogenetic gall wasp Diplolepis rosae (Hymenoptera,Cynipidae)

Summary The univoltine cynipid gall wasp Diplolepis rosae reproduces by an obligate homozygosity promoting system known as gamete duplication. The wasp is confined to roses (Rosa spp) on which it induces large, complex and multichambered galls. In southern Sweden, D. rosae was found to parasitize Rosa canina, R. dumalis, R. rubiginosa, R. villosa, R. sherardi and R. rubrifolia, but not R. majalis and R. rugosa. The distribution of galls shows that there are differences in the relation between wasp and hosplant with respect both to species and individual plants. There is a positive correlation between wasp size and gall (clutch) size. Parasitoid pressure was found to be high, causing D. rosae an estimated average larval loss of approximately 75%, mainly due to the attack of the ichneumonid wasp Orthopelma mediator. The very common cynipid inquiline Periclistus brandtii does not seem to have any negative effects. Overall parasitism and probability of no hatched offspring per gall decrease with increasing gall (clutch) size. The probability of loosing all of a given number of offspring decreases with the number of galls produced. It is suggested that D. rosae, in order to escape parasitoids, needs high ability to establish new colonies. Hence the production of many comparatively small galls, which increases the chance of leaving any offspring, rather than the production of few large galls, maximizing the number of offspring, should be favoured by selection.     

Fungal diversity in galls of baldcypress trees

The baldcypress midge (Taxodiomyia cupressi and Taxodiomyia cupressiananassa) forms a gall that originates from leaf tissue. Female insects may inoculate galls with fungi during oviposition, or endophytes from the leaf tissue may grow into the gall interior. We investigated fungal diversity inside of baldcypress galls, comparing the gall communities to leaves and comparing fungal communities in galls that had successful emergence versus no emergence of midges or parasitoids. Galls of midges that successfully emerged were associated with diverse gall fungal communities, some of which were the same as the fungi found in surrounding leaves. Galls with no insect emergence were characterized by relatively low fungal diversity.     

Effect of cattail (<Emphasis Type="Italic">Typha</Emphasis> spp<Emphasis Type="Italic">.</Emphasis>) mowing on water beetle assemblages: changes of environmental factors and the aerial colonization of aquatic habitats

Some studies found no, or weak evidence that dense monotypic cattail (Typha spp.) stands exclude water beetle species from aquatic habitats, or modify aquatic beetle assemblages. Other studies suggest that cattail may reduce the chance of aerial water beetle colonization, and decreases water temperature; negatively affecting these insects. We examined the response of aquatic beetle assemblages to the mowing of cattail (Typha angustifolia L., T. latifolia L.) in a freshwater marsh. Following removal of cattail at the water level in experimental plots, aquatic beetles were sampled both in mowed and intact (control) plots weekly, through a month in the spring of 2008. Aquatic beetles were more abundant in mowed plots. Species richness was the same, but it showed different patterns in mowed and intact plots. Shannon’s diversity was similar between treatments, while evenness was lower in mowed plots. 29% of the aquatic beetles showed a strong preference for mowed plots, and 15% preferred the control plots. Water temperature was an important factor, with mowed plots having higher water temperatures because of increased solar radiation. Polarization visibility of the water surface was also a factor, since aerially colonizing (flying) aquatic beetles use horizontally polarized light reflected from the water surface to seek potential locations. Using imaging polarimetry, we showed that mowing strongly enhanced the water-reflected polarized light signal, because it reduced the screening effect of cattail leaves, which made the visual detection of water easier. Our results suggest that cattail mowing is a useful method in aquatic beetle conservation: it increases the chance of aerial colonization due to the enhanced polarization visibility of the water surface, and creates a habitat for more abundant assemblages otherwise excluded by the monodominant dense cattail stands. Thus, sustaining hemi-marsh conditions with vegetated and mowed areas is advisable to maximize overall aquatic beetle diversity.     

Life-history traits in insect inclusions associated with bamboo galls

We examined the life-history traits of insect inclusions in bamboo galls induced by the gall maker, Aiolomorphus rhopaloides Walker (Hymenoptera: Eurytomidae) in a bamboo stand. Eight hymenopteran and one dipteran species were detected using soft X-ray photography of the galls and insect emergence from the galls. Aiolomorphus rhopaloides was the gall maker and Diomorus aiolomorphi Kamijo (Torymidae) was its inquiline.Homoporusjaponicus Ashmead (Pteromalidae) and Eupelmus sp. (Eupelmidae) are likely to be primary parasitoids of the larva ofA. rhopaloides. Eurytoma sp. (Eurytomidae),Sycophila sp. (Eurytomidae) and Norbanus sp. (Pteromalidae) are thought to be facultative hyperparasitoids. Cecidomyiidae sp. is thought to be the inquiline ofA. rhopaloides galls.Leptacis sp. (Platygastridae) probably parasitizes the larvae of Cecidomyiidae sp. Larvae of A. rhopaloides appeared in galls in July with the percentage of larvae decreasing in September, before overwintering as pupae. The growth ofD. aiolomorphi larvae within galls may be faster than that ofA. rhopaloides. The percentage of parasitoids in galls was low in July, but increased until winter. Aiolomorphus rhopaloides and D. aiolomorphi emerged from mid-April to early May, and from late April to early June, respectively. From overwintering galls, six other hymenopteran species emerged between late May and late June; one dipteran Cecidomyiidae sp. emerged between mid-April and early May.     

Finding hidden females in a crowd: Mate recognition in fig wasps

Multi-species mating aggregations are crowded environments within which mate recognition must occur. Mating aggregations of fig wasps can consist of thousands of individuals of many species that attain sexual maturity simultaneously and mate in the same microenvironment, i.e, in syntopy, within the close confines of an enclosed globular inflorescence called a syconium – a system that has many signalling constraints such as darkness and crowding. All wasps develop within individual galled flowers. Since mating mostly occurs when females are still confined within their galls, male wasps have the additional burden of detecting conspecific females that are “hidden” behind barriers consisting of gall walls. In Ficus racemosa, we investigated signals used by pollinating fig wasp males to differentiate conspecific females from females of other syntopic fig wasp species. Male Ceratosolen fusciceps could detect conspecific females using cues from galls containing females, empty galls, as well as cues from gall volatiles and gall surface hydrocarbons.In many figs, syconia are pollinated by single foundress wasps, leading to high levels of wasp inbreeding due to sibmating. In F. racemosa, as most syconia contain many foundresses, we expected male pollinators to prefer non-sib females to female siblings to reduce inbreeding. We used galls containing females from non-natal figs as a proxy for non-sibs and those from natal figs as a proxy for sibling females. We found that males preferred galls of female pollinators from natal figs. However, males were undecided when given a choice between galls containing non-pollinator females from natal syconia and pollinator females from non-natal syconia, suggesting olfactory imprinting by the natal syconial environment.