The timing of oviposition and larval growth by two tephritid fly species in relation to host-plant development

Abstract. 1. The relationships between the timing of oviposition, larval establishment and host development were studied in the tephritids Tephritis bardanae (Schrank) and Cerajocera tussilaginis (Fab.) which attack flowerheads of Arctium minus (Hill) Bernh. (Compositae).
2. T. bardanae laid eggs in small sized flowerheads that were at an early stage of development, and appeared to use the external dimensions of the head as an oviposition cue. In contrast, C. tussilaginis females laid eggs into heads that were near to flowering and the timing of their attack was most closely related to particular lengths of florets and achenes inside the head. Both tephritids apparently recognized suitable heads by comparing aspects of their own body size with the size of particular flowerhead structures.
3. Individual heads were suitable for oviposition for a period of 10–11 days for both species, but the periods of susceptibility to each species did not overlap.
4. Attack by T. bardanae was timed to exploit the flowering phase of flowerhead development. Larval establishment and early development were synchronized with increasing nutrient availability occurring during the rapid growth of the florets and achenes before and during anthesis.
5. In contrast, the later attack by C. tussilaginis was timed to exploit the seed maturation phase of head development. Individual C. tussilaginis larvae operated as highly specialized 'achene parasites', utilizing the influx of nutrients into single achenes after fertilization.
6. Larvae of both tephritids induced abnormal host-tissue growth, but oviposition in both species was timed primarily to synchronize larval development with natural periods of high nutrient availability inside flowerheads, rather than with the presence of tissues suitable for gall induction.     

Resource limitation of tephritid flies on lesser burdock,Arctium minus (Hill) Bernh. (Compositae)

Summary The intensity of resource exploitation by phytophagous insects is usually considered to reflect population size. For populations of two flowerhead-attacking tephritid flies, however, the resources utilised were not related to the numbers of searching adults. Tephritis bardanae Schrank attacked 11–13% of the total flowerheads each year, and Cerajocera tussilaginis (Fab) 17–65%, despite much wider and uncorrelated variation in adult numbers. Analysis of field data showed that the proportion of flowerheads used was not limited by poor flowerhead quality, but was restricted by (1) the synchronisation of adult activity with the appearance of flowerheads at the correct age for oviposition, and (2) by factors which influenced the ability of female flies to locate available heads. These restrictions were more severe in T. bardanae and explained its relatively low rate of infestation. Both tephritids tended to avoid flowerheads in open areas. The processes governing resource exploitation in each tephritid operated independently of the other, and a partial separation of the two species between flowerhead types and habitats arose simply because of their different timing of attack.     

Molecular phylogeny of the subfamily Tephritinae (Diptera: Tephritidae) based on mitochondrial 16S rDNA sequences

The phylogeny of the subfamily Tephritinae (Diptera: Tephritidae) was reconstructed from mitochondrial 16S ribosomal RNA gene sequences using 53 species representing 11 currently recognized tribes of the Tephritinae and 10 outgroup species. The minimum evolution and Bayesian trees suggested the following phylogenetic relationships: (1) monophyly of the Tephritinae was strongly supported; (2) a sister group relationship between the Tephritinae and Plioreocepta was supported by the Bayesian tree; (3) the tribes Tephrellini, Myopitini, and Terelliini (excluding Neaspilota) were supported as monophyletic groups; (4) the non-monophyletic nature of the tribes Dithrycini, Eutretini, Noeetini, Tephritini, Cecidocharini, and Xyphosiini; and (5) recognition of 10 putative tribal groups, most of which were supported strongly by the statistical tests of the interior branches. Our results, therefore, convincingly suggest that an extensive rearrangement of the tribal classification of the Tephritinae is necessary. Since our sampling of taxa heavily relied on the current accepted classification, some lineages identified by the present study were severely under-sampled and other possible major lineages of the Tephritinae were probably not even represented in our dataset. We believe that our results provide baseline information for a more rigorous sampling of additional taxa representing all possible major lineages of the subfamily, which is essential for a comprehensive revision of the tephritine tribal classification.     

Egg–larval opiine parasitoids (Hym., Braconidae) of tephritid fruit fly pests do not attack the flowerhead-feeder Trupanea dubautiae (Dipt., Tephritidae)

Abstract:  We investigated the potential impact of three opiine tephritid fruit fly parasitoids: Fopius arisanus (Sonan), Fopius caudatus (Szépligeti) and Fopius ceratitivorus Wharton, on the non-target native Hawaiian tephritid, Trupanea dubautiae (Bryan), infesting flowerheads of the endemic Asteraceae shrub Dubautia raillardioides Hillebrand. The three species are the only known opiine fruit fly parasitoids that attack host eggs (but occasionally attack first instars). F. arisanus , which originated in southeast Asia, is now widely established in Hawaii and elsewhere in the world, while the other two are African species currently in quarantine in Hawaii. In the laboratory, field-collected flowerheads of D. raillardioides containing T. dubautiae eggs and first instar larvae were exposed to naïve female wasps of each of the three Fopius species in the absence (no-choice test) or presence (choice test) of papaya fruit infested with eggs of the Mediterranean fruit fly Ceratitis capitata (Wiedemann), the parasitoids' normal host. All three Fopius species visited the papaya fruit much more often than the flowerheads, and displayed normal ovipositional responses (probing and stinging) on the fruit in the choice test. None of the three parasitoid species showed ovipositional responses to flowerheads in either choice or no-choice tests. As a result, not a single T. dubautiae egg or larva was attacked by any of the three parasitoids, indicating that these parasitoids of pest tephritids are unable to recognize the microhabitats of flowerhead-feeding tephritids. The results suggest minimal risk of non-target impact in these biological control agents.     

Distribution of Insect Attacks in Biological Control of Weeds: Infestation of Centaurea virgata Flowerheads by a Gall Fly

The success of biological control efforts to reduce weed density through release of insects may depend as much on the distribution of insect attacks among individual plants or plant parts as on the mean level of infestation. We used an index of dispersion to describe the distribution of Urophora quadrifasciata (Diptera: Tephritidae) galls among squarrose knapweed (Centaurea virgata) flowerheads at 18 west central Utah sites in the first 5 years following introduction of the biological control agent. Two thirds of the samples showed a significantly aggregated distribution of galls among flowerheads. Statistical analysis showed that site and year accounted for relatively small proportions of the variance in the index of dispersion. The degree of gall aggregation among flowerheads was positively correlated with the mean flowerhead quality (mean number of seeds per flowerhead; P = 0.013) and tended to be negatively correlated with the mean fly density per flowerhead at a site in a given year (P = 0.097). Our data suggest that higher quality flowerheads, and possibly higher quality plants, are preferentially attacked by U. quadrifasciata and therefore are more heavily subject to reduced reproductive potential through biological control. However, an aggregated distribution of fly attacks may undercut the potential of the fly to reduce seed production by the weed population as a whole. Understanding both the distribution of insect attacks among individual plants and the behavioral mechanisms producing such distribution patterns is important to the biological control of weeds.     

Carbon dioxide assimilation in the flowerhead of Arctium

Summary The gas exchange of flowerheads was determined in Arctium tomentosum and A. lappa during their development. The light, temperature and CO₂ responses were used to estimate flowerhead photosynthesis and the in situ contribution of carbon assimilation to the carbon requirement of the plant for supporting a flowerhead. Changes in vapour pressure deficit had no effect on flowerhead photosynthesis rates and were not included in the model.In both species assimilatory capacity correlated with total bract chlorophyll content. Light, temperature and CO₂ response curves were very similar in form between species, differing only in absolute rates. During all stages of development, flowerheads always exhibited a net carbon loss, which was mainly determined by temperature. The respiration rate decreased in the light, the difference of CO₂ exchange in the dark and in the light was interpreted as photosynthesis. This rate was larger in A. lappa than in A. tomentosum. 30% of the total C requirement of A. lappa flowerheads was photosynthesized by its bracts, the total contribution offlowerhead photosynthesis in A. tomentosum was only 15%. The potential competitive advantages of variation in flowerhead photosynthesis are discussed.     

Impact of the biological control agent Mesoclanis polana (Tephritidae) on bitou bush (Chrysanthemoides monilifera subsp. rotundata) in eastern Australia

The seed fly Mesoclanis polana (Diptera: Tephritidae) was released in Australia in 1996. Its impact on seed production of bitou bush (Chrysanthemoides monilifera subsp. rotundata) was monitored at eight sites along the New South Wales coast from 1996 to 2004. Peak flowerhead production occurred in autumn (March to May); therefore, samples collected in May of each year were used to compare abundance and impact of M. polana across sites and between years. Latitude had a significant effect on abundance and impact of M. polana. By May 2004, 99.6% of flowerheads at the five most northern sites contained at least one egg, while 64% of flowerheads from the two most southern sites contained at least one egg. In May 2004, mean numbers of M. polana eggs per flowerhead were between 13 and 17 at four of the six northern sites and below two at the two southern sites. In May 2004, average seed destruction by M. polana was 58% at the four most northern sites and 11% at the two most southern sites. The highest level of seed destruction recorded was 86% at Kingscliff in May 2003. Percentage seed destruction in May provided a reasonable estimate of seed destruction for the whole year. Parasitoids were found throughout the range of M. polana in Australia. Their attack rate on M. polana was less than 10% at all sites, except at the two most northern sites where parasitism of up to 27% was recorded. Parasitism results are compared with predictions made following an earlier study. The release of Mesoclanis magnipalpis, a species suited to cooler environments, is recommended for the southern range of C. monilifera subsp. rotundata.     

The directional control of sucrose and asparagine transport in lupin by abscisic acid

Injections of exogenous abscisic acid into the primary flowerhead of Lupinus luteus cv. Weiko III reduced the movement of ¹⁴C-sucrose into the flowerhead from the uppermost leaves. Sucrose transported from below the lateral branches subtending the flowerhead, was diverted into the lateral branches by injection of the exogenous abscisic acid into the flowerhead. ¹⁴C-sucrose was also diverted from a lateral branch injected with exogenous abscisic acid to all other parts of the plant, particularly the main stem and leaves, and the roots. Transport of ¹⁴C-asparagine administered at the cotyledonary node was directed from the flowerhead into the subtending lateral branches by injection of abscisic acid into the flowerhead. Transport of both ¹⁴C-sucrose and ¹⁴C-asparagine into the flowerhead was reduced at least three fold at physiological levels of abscisic acid. No significant correlation was found between the amount of ¹⁴C-asparagine entering a sink and the dry weight of the tissues of that sink. It is concluded that distribution of ¹⁴C-sucrose and ¹⁴C-asparagine between the flowerheads and lateral branches of L. luteus is actively and dynamically controlled and that abscisic acid levels play a significant part in that control. It is suggested that the relative levels of endogenous abscisic acid in plant organs could serve as an important factor in the directional control of assimilate transport in plants.     

Nutrient addition extends flowering display, which gets tracked by seed predators, but not by their parasitoids

Although phenological matching between two and three trophic interactions has received some attention, it has largely been disregarded in explaining the lack of strong cascade dynamics in terrestrial systems. We studied the response of the specialist seed predator, Paroxyna plantaginis (Tephritidae) and associated generalist parasitoids (Chalcidoidea) to controlled fertilisation of individuals of naturally growing Tripolium vulgare (Asteraceae) on four island populations (Skeppsvik Archipelago, Sweden). We consistently found evidence of nutrient limitation: fertilised plants increased their biomass, produced more capitula (the oviposition units for tephritid flies), were more at risk of attack by the tephritids, and puparia were heavier in fertilised plants. During some parts of the season tephritids became more heavily parasitized, supporting the presence of cascade dynamics, however net parasitism over season decreased in response to nutrient addition. We found no evidence that capitulum size complicated parasitoid access to the tephritids, however the extended bud production prolonged the flowering season. Thus, tephritids utilized the surplus production of capitula throughout the entire season, while parasitoids did not expand their oviposition time window accordingly. Implications for top down regulation and cascade dynamics in the system are discussed.     

Chemical composition,antibacterial and antifungal activities of flowerhead and root essential oils of Santolina chamaecyparissus L., growing wild in Tunisia

The antimicrobial properties of essential oil from various Santolina species have not been investigated enough in the previous studies dealing with the biological activities of medicinal plants. In Tunisia, Santolina chamaecyparissus L. (Asteraceae) is the only Santolina species recorded and is used as vermifuge and emmenagogue. The chemical composition, antibacterial and antifungal properties of essential oils from the flowerheads and roots of spontaneous S. chamaecyparissus growing in Tunisia and the chemical composition which leads to the Tunisian chemotype are investigated here for the first time. Essential oils isolated by hydro distillation from flowerheads and roots of S. chamaecyparissus were analyzed by GC and GC/MS. Two methods served for antimicrobial assays of the essential oils: diffusion in a solid medium and micro-well dilution assay. Antifungal tests were carried out by the agar incorporation method. Sixty-seven constituents were identified from the essential oil of the flowerhead. The major constituents were: 1,8-cineole and β-eudesmol. Two non identified compounds were present at the highest concentration in root oil. Flowerhead oil was characterized by high contents in monoterpenes and sesquiterpenes oxygenated compounds. The flowerhead essential oil demonstrated potent of antibacterial properties against Pseudomonas aeruginosa ATCC and Enterococcus faecalis ATCC, with MIC of 0.625 μg/ml. These findings demonstrate that the flowerhead essential oils of S. chamaecyparissus have excellent antibacterial properties and for this reason they could contribute to decrease the problem of microbial resistance to antibiotics.     

Effects of prey-size and predator-instar on the predation of Daphnia by Notonecta

Abstract. 1. Attack rates and handling times are measured by a series of separate functional response experiments for each instar of Notonecta glauca attacking four size classes of Daphnia magna as prey. The resulting attack rate and handling time surfaces are complex, with maximum attack rates for small predators attacking small prey, and large predators attacking large prey. Adult Notonecta have lower attack rates than the two previous juvenile instars (4 and 5).
2. The literature on attack rates and handling times in other predator—prey interactions that involve a series of different predator and prey size or age classes is reviewed in the context of the Notonecta-Daphnia results. The data suggest that small predator instars will usually compete with large instars for food, unless there is spatial or temporal separation between them.
3. Complex attack rate and handling time surfaces are to be expected wherever a wide range of prey and predator sizes is involved.
4. Size related changes in attack rates and handling times can introduce very complex dynamics into predator-prey interactions.     

The influence of host size on sex ratios in the parasitoid Diglyphus begini (Hymenoptera: Eulophidae)

Abstract. 1. Sex ratio in the ectoparasitoid, Diglyphus begini (Ashmead), attacking larvae of the dipterous leafminer Liriomyza trifolii (Burgess) in glasshouse marigolds, is best depicted by a model where sex ratio varies as a function of host size.
2. Male D. begini progeny are produced in hosts significantly smaller in size than those producing female progeny.
3. Female wasps attack and oviposit on the largest leafminer larvae available and whether a host is large or small depends upon the size of the other hosts attacked.
4. Diglyphus begini females adjust the thre:shold size for the change-over in sex allocation relative to the size of hosts attacked; however, the sex ratio is maintained at between 60% and 70% males.
5. The patterns observed in these glasshouse studies are not due to sex-specific differential mortality or superparasitism.     

Life-history strategy in an oligophagous tephritid: the tomato fruit fly, Neoceratitis cyanescens

Abstract.  1. In phytophagous insects, life-history traits mainly depend on host plant range. Substantial longevity, high fecundity and larval competition are the major traits of polyphagous Tephritidae while species with a restricted host range generally exhibit a lower longevity and fecundity as well as mechanisms to avoid larval competition. Our aim in this study was to investigate the life history of an oligophagous species, the tomato fruit fly, Neoceratitis cyanescens (Bezzi).
2. We determined life tables under laboratory conditions in order to calculate the main demographic parameters of N. cyanescens and studied the influence of larval and adult diet on life-history traits.
3. The mean longevity of N. cyanescens females was 40 days. There was a strong synchronisation of female maturity. Oviposition showed an early peak at 9–13 days after a short pre-oviposition period (6 days). The absence of proteins in the adult diet both delayed ovarian maturation and decreased female fecundity. In addition, females originating from tomato fruits produced significantly more eggs than females originating from bugweed or black nightshade, showing that even the larval host plant may strongly affect the subsequent fecundity of adult females.
4. The traits of N. cyanescens are then discussed in the light of those documented for polyphagous and monophagous tephritids. Neoceratitis cyanescens displayed attributes intermediate between those of polyphagous and monophagous tephritids. Its smaller clutch size compared with polyphagous species and its specialisation on the Solanaceae family whose fruits contain toxic compounds may help in reducing intra- and inter-specific competition, respectively.     

Body size and host range co-determine the altitudinal distribution of Neotropical tephritid flies

Aim We addressed the following questions: (1) Does tephritid body size tend to increase in species found at higher elevations, as predicted by Bergmann's rule? (2) Do tephritids conform to Rapoport's rule, so that species found at higher elevations tend to have broader altitudinal ranges? (3) More generally, how do body size and host range jointly affect the patterns of altitudinal distribution among Neotropical tephritid flies? Location The Mantiqueira mountain range, south‐eastern Brazil, at sites ranging from c. 700 to 2500 m a.s.l. Methods At each site we collected flower heads of all Asteraceae species to rear out endophagous immatures (from January to June in 1998 and 1999). We used structural equation models (SEM) to evaluate jointly the relationships between body size, host range and altitudinal distribution (range and mid‐point). Results Neotropical tephritid body size showed a negative relationship with altitudinal distribution. SE modelling showed no significant direct effect of body size on altitudinal range; however, it had significant indirect negative effects through host range and altitudinal mid‐point. The SE model was a good predictor of observed correlations and accounted for 84% of the variation in tephritid altitudinal range. Main conclusions The altitudinal range of flower‐head‐feeding tephritids is related to host range and is indirectly affected by body size via host range and altitudinal mid‐point. As predicted by Rapoport's rule, tephritids that occur at higher elevations also present wider altitudinal ranges. Bergmann's rule does not apply to Neotropical tephritids along a tropical elevational gradient, but rather its converse was found. Body size may determine host range by imposing a restriction upon large individuals using small flower heads. Host species turnover along the altitudinal gradient may be the main factor explaining the strong relationship between host range and insect elevational distribution.     

Phenotypic plasticity in <Emphasis Type="Italic">Carlina vulgaris</Emphasis>: effects of geographical origin,population size,and population isolation

If phenotypic plasticity is under genetic control, it may vary in amount and pattern on a geographical scale, e.g. among different regions of a species distribution. It may also differ between large and small or between less and more isolated populations, due to differences in genetic diversity. In a 2-year common garden study, the responses of several traits to drought and fertilizer treatments were studied in the grassland herb Carlina vulgaris. Individuals originating from populations of different size and degree of isolation in six European countries, representing central and marginal regions, were compared. Fertilizing had a negative effect on early plant survival, as well as on flowering probability in surviving plants. However, in those plants that flowered, fertilizing strongly increased mean number of flowerheads, flowerhead area (a correlate of seed number), and seed mass. Drought had generally weaker effects but enhanced survivorship, indicating that this treatment was closer to optimal conditions than were non-drought conditions. For some traits there were significant interactions of region × fertilizer, but the geographical pattern of reaction norms was inconsistent and lent no support to the hypothesis that central and marginal populations differ in overall plasticity. Population size and isolation had hardly any influence on treatment responses, but populations within regions differed in their mean response to fertilizing with regard to survival and flowering probabilities, as well as in their response to drought with regard to survival and total flowerhead area. It is concluded that response to raised nutrient levels is highly variable within populations, ranging from death to strongly increased reproductive output, but also among populations irrespective of size or isolation. This also goes for the response to water supply, though this variation shows a more unclear pattern. There is no evidence that small or isolated/marginal populations are less plastic than large or non-isolated/central populations, and the explanation for differences in treatment responses among plant populations should be sought in other population characteristics.     

Species segregation during predator evasion in cyprinid fish shoals

SUMMARY. 1. When a predator attacks a mixed-species group, individual prey arc faced with a conflict between retaining the advantages of large group size and the disadvantages of remaining among aliens. Experiments were designed to show how this conflict is resolved in fish shoals.
2. Simulated attacks with a model predator were made upon mixed and single species shoals of three cyprinid species under semi-natural conditions in a fluvarium.
3. Measurements of elective group size, neighbour identity and distance, and encounters within and between species revealed behaviour which sorted fish into species at the onset of predator attack.
4. A control experiment demonstrated the same effect for fish of matched size.
5. The value to the individual of segregative behaviour in response to predator threat is discussed.     

Habitat characteristics,stage structure and reproduction of colline and montane populations of the threatened species <Emphasis Type="Italic">Arnica montana</Emphasis>

Arnica montana, a characteristic plant of nutrient-poor grasslands, has strongly declined during the last decades, particularly at lower altitudes. To gain insights into the underlying reasons for this decline, we recorded differences in size, stage structure and reproductive traits between 21 colline populations of A. montana in the Ardennes-Eifel region (280–600 m a.s.l.) and 10 montane populations in the Vosges Mountains (1,200 m a.s.l.). Community composition of the Violion caninae vegetation at colline sites and Nardion strictae vegetation at montane sites indicated that temperature and moisture were the main abiotic factors differentiating between low and high altitudinal relevées. The proportion of flowering rosettes decreased with altitude, indicating a shift from sexual reproduction to clonal growth. In contrast to expectation, Ellenberg values for nutrient availability as well as the proportion of young rosettes and population size did not differ between colline and montane populations. However, population size decreased with nutrient availability, indicating eutrophication as the reason for population decline. In small populations, plant density, the proportion of flowering rosettes, the number of flowerheads per rosette and the number of seeds per flowerhead were lower, indicating less suitable conditions. Thus, preventing further eutrophication in both colline and montane populations will be crucial for the conservation of the species. Reduced reproduction may not be important for the population dynamics of this clonal plant in the short term, but could affect genetic diversity and survival in the long term.     

Pollen limitation and pollinator preferences in Scorzonera hispanica

The plant life cycle is often affected by animal–plant interactions. In insect‐pollinated plants, interaction with pollinators is very important. When pollen transfer due to a lower abundance of pollinators limits seed production, selection pressures on plant traits related to plant attraction to pollinators might occur, e.g. on flowering phenology, height or number of flowerheads. Landscape changes (e.g. habitat fragmentation or changed habitat conditions) may cause plant–pollinator systems to lose balance and consequently affect population dynamics of many plant species. We studied the relationship between measured plant traits, environmental variables and pollinator preferences in Scorzonera hispanica (Asteraceae), a rare perennial, allogamous herb of open grasslands. We estimated the pollen limitation by comparing seed set of supplemental‐pollinated plants with that of open‐pollinated ones. Pollinators selected plants based on position within the locality (isolated plants close to trees) rather than on their traits. In spite of a high proportion of undeveloped seeds on the plants, we demonstrated that they are not pollen limited. Instead, seed set and weight of seeds was correlated with plant size traits (height and flowerhead number), with larger plants producing more and larger seeds. This suggests that the studied plants are likely resource limited. Overall, the results suggest that pollinators are not a selection factor in this system, in contrast to studies on various plant species, including self‐compatible species of the Asteraceae. The lack of any effect of pollinators in the system may be caused by a strong negative effect of ungulate herbivores, which could play a decisive role in functioning of the system.     

The influence of preformed defences on the dynamic wound response in Spruce bark

1. Two of the principal defences in conifer bark against attack by bark beetles and associated fungi, the flow of preformed resin and the dynamic wound response (DWR), are commonly regarded as separate, sequential responses to attack. In this paper the concentration of three preformed defences (resin, total polyphenols and lignified stone cell masses) was measured at different heights on the stem of Norway and Sitka Spruce to determine their effect on the size of lesions formed during the DWR to inoculation with three facultatively pathogenic fungi ( Phacidium coniferarum, Ophiostoma piceae and Cryptosporiopsis sp.) inoculated during the dormant season.
2. There was considerable within-tree variation in concentration of lignin and there was a dose-dependent negative effect of lignin on lesion size. The form of the relationship, however, was influenced by fungal and tree species and for some fungi, also by an individual tree effect.
3. Results suggest that resistance of conifer bark to pests and pathogens depends on an interaction between preformed and induced defences.     

The causes of over wintering mortality of Phyllonorycter on Quercus robur

Abstract. 1. An experiment was performed to examine the relationship between overwintering mortality rates and physical conditions in the leaf litter for Phyllonorycter spp. on Quercus robur .
2. Analyses also were performed to determine if survival and mortality rates vary with leaf size and the position of a leaf mine within a leaf.
3. The results indicate that survival rates are independent of variation in physical conditions in the leaf litter, as well as leaf size and the position of a mine on a leaf.
4. Mortality from leaf decomposition was higher at the bottom of the litter pack, and it may compensate for reduced mortality from predation, crushing, desiccation, and fungal attack at the level.
5. Rates of parasitism were significantly higher on small than on large leaves.