Evaluation and host specificity of two seed flies Mesoclanis polana and M. magnipalpis (Diptera: Tephritidae): biological control agents for Chrysanthemoides monilifera (Asteraceae) in Australia

Larvae of the South African tephritid flies Mesoclanis polana Munro and M. magnipalpis Bezzi feed in the developing seeds of Chrysanthemoides monilifera. Host specificity evaluation using 109 plant species from 25 families indicated that complete development was restricted to their natural host C. monilifera. Minor feeding and limited development was detected on 18 species, but was of no ecological or economic significance. Mesoclanis polana and M. magnipalpis have been released in Australia and M. polana has established and dispersed widely. Mesoclanis magnipalpis has not yet become naturalized. Parasitism of M. polana in Australia by several species of Hymenoptera has been detected, but is not expected to limit the establishment and impact of these flies.     

Development of a Pesticide Exclusion Technique for Assessing the Impact of Biological Control Agents for Chrysanthemoides monilifera

Seven biological control agents have been released on the environmental weed Chrysanthemoides monilifera in Australia and two are widely established on infestations of C. monilifera subsp. rotundata in New South Wales. Five pesticides were screened for their impact on seed production of C. monilifera and two on the survival of the agent Comostolopsis germana, a shoot tip-feeding lepidopteran. The insecticides carbaryl, carbofuran, dimethoate, fluvalinate and the fungicide benomyl had no significant effect on seed production of C. monilifera when applied over a period of four months. Numbers of C. germana larvae were readily suppressed after three months by applications of fluvalinate or a mixture of carbaryl, carbofuran and dimethoate. Shoot growth was not affected by applications of carbaryl, dimethoate and benomyl. It is concluded that exclusion by pesticides of biological control agents is a valid method of measuring the impact of agents on C. monilifera.     

Establishment and rapid spread of the bitou seed fly, Mesoclanis polana Munro (Diptera: Tephritidae), in eastern Australia

The establishment and spread of the seed fly Mesoclanis polana Munro, an introduced biological control agent for Chrysanthemoides monilifera (L.) Norlindh, is described. From an initial release of 124 adult flies in August 1996, the species occupied virtually the entire range of bitou bush by October 1998, from Rainbow Beach in the north to Tathra in the south, a distance of over 1200 km. Early data on abundance are provided.     

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.     

Selection of Cassida spp. from southern Africa for the biological control of Chrysanthemoides monilifera in Australia

Four species of tortoise beetle (Chrysomelidae, Cassidinae) (Cassida spatiosa Spaeth and three undescribed Cassida species, labelled 1. 2 and 3) from South Africa, were assessed as potential biological control agents against Chrysanthemoides monilifera monilifera (L.) T. Norl. and C. m. rotundata (DC.) T. Norl. These southern African plants are important weeds of conservation areas in Australia. In South Africa, Cassida spatiosa was found feeding on C monilifera subcanescens (DC.) T. Norl., Cassida sp. 1 on Chrysanthemoides monilifera pisifera (L.) T. Norl. and C m. monilifera, Cassida sp. 2 on C m. pisifera and Chrysanthemoides incana (Burm.f.) T. Norl. and Cassida sp. 3 on C. m. rotundata. The life cycle of each species was completed in about three months on the leaves of the host. Cassida species 1, 2 and 3 were tested against eight species of plant and only fed and completed development on Chrysanthemoides species and the related species. Calendula officinalis L. Cassida species 1 and 3 showed no preference between Calendula officinalis and Chrysanthemoides species for oviposition. Cassida species 3 was selected for further assessment as a biological control agent based on match to the host subspecies and target climate in Australia.     

Seed characteristics and regeneration of some species in invaded coastal communities

The seed production, germination and regeneration characteristics of two native species (Acacia longifolia var. sophorae and Banksia integrifolia) and two invasive subspecies (Chrysanthemoides monilifera ssp. monilifera and ssp. rotundata) were compared. C. monilifera ssp. rotundata was similar in seed characteristics and regeneration to B. integrifolia in that: (1) seed production was over an extended period each year; (2) seed longevity was low (0–2% viable after burial for 3 years), but increased with depth of burial; and (3) regeneration from 26 to 28% of mature plants occurred after fire. By contrast, C. monilifera ssp. monilifera was similar to A. Iongifolia var. sophorae in that; (1) seed production was over a short period each year; (2) of the total buried, 6–13% of seeds were viable after 3 years, and again numbers increased with depth of burial (percentages of missing seeds of A. longifolia were high but all remaining seeds were viable after 3 years); and (3) regeneration after fire occurred only from seed, hence their seed longevity is an important factor for long-term survival. However, growth habit and niche occupation by ssp. rotundata is similar to that of A. longifolia and the above differences may contribute to the observed displacement of the native by the invasive species. The distribution of the two invasive subspecies is at present largely in separate areas in Australia and it is predicted this situation will continue because of their differing climatic preferences, morphology and regeneration from fire.     

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.     

“Barcoding” and ISSR data illuminate a problematic infraspecific taxonomy for Chrysanthemoides monilifera (Calenduleae; Asteraceae): Lessons for biocontrol of a noxious weed

Chrysanthemoides monilifera Tourn. ex Medik is a noxious weed in Australia and New Zealand. It is a widespread species in southern Africa, where it shows considerable morphological variation that has resulted in a confusing infraspecific taxonomy. We use DNA sequence data from the nuclear Internal Transcribed Spacer (ITS) region from 78 samples of this species from its indigenous distribution range and 10 samples from Australia and New Zealand to determine the regions of origin of the invasive plants. These data are augmented by a smaller study using ISSR markers. Bayesian Inference analysis was somewhat resolved, with many weakly supported nodes. Clades with support tended to correspond to infraspecific taxonomic entities, and were geographically coherent. In contrast, a neighbour-net analysis was not as well resolved and indicated considerable reticulation. All analyses of ITS data retrieved two major clades corresponding to Western and Eastern distributions, with some overlap. Samples from New Zealand and Australia correspond to the taxon C. monilifera subsp. monilifera, and are resolved as most closely related to samples from the greater Cape Town area. Biological control agent populations for C. monilifera subsp. monilifera should be sourced from this region in order to avoid host plant incompatibility problems.     

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.     

Distribution of Aphids (Hemiptera: Aphididae) in Potato Growing Areas of Southwestern Australia

Sticky traps, water traps and leaf samples were used to monitor aphid populations at 32 sites in southwestern Australia between October and May for four seasons between 1988–92. Twenty-seven sites were located in potato, Solanum tuberosum L., crops, with five others located in mixed vegetable crops or pastures. Sites were located in an area of approximately 90,000 km², and encompassed most of the potato growing areas of the region. Thirty-four species of aphids were detected. Potato-colonising aphids found were Myzus persicae, Aulacorthum solani, Macrosiphum euphorbiae and Aphis craccivora. M. persicae was the most widespread, and its seasonal distribution in southwestern Australia is presented. Fewer M. persicae were recorded and it arrived later in the season for 2 of 3 years at southern sites than northern sites. At sites where both sticky traps and leaf sampling were used, leaf sampling often detected M. persicae before they were found on traps. Results suggest that potato crops grown on the southern coast will be least vulnerable to infection by aphid-borne viruses.     

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.     

Alfalfa leafcutting bee population dynamics, flower availability, and pollination rates in two Oregon alfalfa fields

Since the 1970s, it has become increasingly difficult for U.S. alfalfa seed producers to maintain Megachile rotundata (F.) populations used for alfalfa, Medicago sativa L., pollination. In 1998, we monitored M. rotundata population dynamics and foraging behavior, as well as alfalfa bloom and pollination rates in two fields in eastern Oregon. Despite marked differences in bee management, establishment was very similar in the two fields (approximately 0.5 females per nesting cavity) and lagged peak bloom by approximately 2 wk. Pollination rates increased from 0-10% in the first 3 wk to 80-90% in week 4-5. By then, M. rotundata females had difficulty finding untripped (nonpollinated) flowers and visited large numbers of already tripped or not fully matured flowers. M. rotundata progeny mortality was very high (54-78%). Estimated seed yields were similar in both fields. We contend similar seed yields, and improved bee production, could be accomplished with smaller bee populations, better timed with alfalfa bloom.     

A contribution to the biology ofBangasternus fausti [Col.: Curculionidae], a potential biological control agent of diffuse knapweed,Centaurea diffusa,and its effect on the host plant

The biology of the weevilBangasternus fausti (Reitter), the larvae of which live in the flowerheads ofCentaurea diffusa (De Lamarck) and some relatedCentaurea spp., was studied in the field and in the laboratory in northern Greece and in the laboratory in Rome, Italy. The species is univoltine and overwinters as adults. Under laboratory conditions adults lived up to 90 days and up to 359 eggs were deposited by a single female (n=18). The maximum number of eggs laid by a female per day was 12 and the maximum number of days a female laid eggs was 73. Generally, there is only one larva per seedhead. One-hundred percent seed destruction was recorded in 18.7% of 502 field collected seedheads, each with oneB. fausti egg. The average mortality of the species, from egg to adult, was 94.4% in the field during 1988 (n=200) and 95.8% during 1987 (n=502).      

Production and survival of eggs of the cattle tick Boophilus microplus (Canestrini) (Acarina: Ixodidae) in the wet and dry tropics of north Queensland

Monthly observations were made on egg production and egg survival of the cattle tick Boophilus microplus , at five sites in northern Queensland from August 1978 to July 1980. Egg production was close to the potential except in the hot, dry season in the drier sites and a model was fitted to explain the data. The main factors that reduced egg production were maximum soil temperature greater than 33.6°C, minimum soil temperature less than 17.6°C, and soil moisture less than 0.26 water-holding capacity when the soil maximum temperature was greater than 37.2°C. The model explains 61% of the variation in egg production from an independent set of data from southern Queensland. Analysis of egg viability at the time of retrieval from pasture indicated that the most important variables reducing viability were maximum soil temperature greater than 25.4°C, minimum soil temperature less than 16.9°C, soil moisture below 0.21 water-holding capacity, the sum of degree-days less than 3.95 per day and stress carried over from the egg production phase.     

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.     

The Effect of Water Stress on Floral Characters, Pollination and Seed Set in White Clover (Trifolium repens L.)

White clover plants were subjected to either short-term developingwater stress or long-term stable levels of water deficit. Thehort-term stress reduced plant water status to about –2·0MPa over 16 d. The long-term stress was less severe, but wassustained for several weeks. Long-term water stress promotedthe production of inflorescences. However, water stress alsoincreased floret abortion and the premature death of whole flowerheads. The number of ovules per floret was decreased by waterstress. The most striking effect of both long- and short-term waterdeficit was to reduce pollen viability measured with the fluorochromaticassay. This was not an artefact of assay conditions. The pollenfrom water-stressed flower heads was not reversibly dehydrated;it did not score at similar viability to controls after incubationin conditions which hydrate pollen. In addition, the pollenfrom water-stressed plants lost viability more rapidly thanpollen from well-watered plants after removal from the flowerhead. The consequences of reduced pollen viability on seed set wereinvestigated by hand-crossing within and between groups of plantsmaintained for several weeks at three levels of water supply.Flower heads pollinated with pollen from water-stressed plantsset fewer seeds per floret than those pollinated with controlpollen. Key words: Trifolium repens, white clover, water stress, floral characters, seed set     

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.     

Climatic control of intra- and inter-annual wood-formation dynamics of Scots pine in northern Finland

Boreal forests are highly sensitive to climate and human impacts and therefore suitable as biological indicator for environmental changes. In this context, our study was aimed at getting deeper insight into the climate-dependence of the onset, intensity and end of wood formation of Scots pine during the growing season.We monitored the intra-annual growth dynamics of, on average, 42-year-old Scots pine trees over five consecutive years, 2000-2004, at two sites located 80 and 300 km south of the tree line in northern Finland. For that purpose, the cambium of the trees was weekly wounded with a pin and the resulting wound tissue, microscopically detectable in transverse thin-sections through the newly built wood, was taken as a time marker. During this 5-year study period, the intra-annual wood formation at the southern site was mainly positively associated with summer temperature. However, at the northern site such an association was either entirely missing or negative. At both sites, two thirds of the radial growth was produced within only 4 weeks from mid-June to mid-July, independent of whether the growing season started earlier or later.Moreover, we measured the widths of all tree rings from bark to pith (inter-annual growth) of the same study trees and assembled them to 51-year long tree-ring site chronologies. Since 1999, these two site chronologies - after having run fairly parallel over the preceding decades - were running in divergent directions thus corroborating our results derived from the intra-annual climate/growth analysis. Whereas the chronology of the southern site follows the average temperature of May and July very closely from 1961 up to 2004, the chronology of the northern site follows the July temperature, but only up to 1998, and from 1999 to 2004 is running just opposite to the distinctly rising July temperature. During the same period, there was - unlike in the years before - nearly no snow cover in May at the northern site, whereas at the southern site there was no change of the normally existing slight snow cover in May. This deviating weather situation may have led to a temperature-induced, temporary drought stress for the Scots pines at the northern site.     

Below-ground Herbivory in Carduus nutans (Asteraceae) and the Potential for Biological Control

The impact of root-feeding insects attacking the 'biennial' Carduus nutans L. in the southern part of its native European distribution was studied as part of a biological control programme against this weed for Australia. Surveys were conducted from Spain to Greece to parallel previous studies in central Europe and three populations of the weed were monitored in detail over a 2-4-year period in southern France. The damage levels and attack patterns of each insect species were recorded. The most common species damaging the roots and meristems were the weevils Hadroplontus (=Ceutorhynchus) trimaculatus F. and Trichosirocalus horridus Panzer and the syrphid fly Cheilosia corydon Harris (=grossa Fallen). In manipulative experiments, made to assess their impact on the host plant, C. corydon reduced seed production by up to 45%, while the two weevils mainly altered plant architecture. The results are discussed in relation to previous experience of using root feeders for the biological control of C. nutans.     

松墨天牛的全球潜在分布区分析

松墨天牛MonochamusalternatusHope分布在亚洲东部,是松材线虫Bursaphelenchusxylophilus(SteinerandBuhrer)在亚洲最有效的昆虫媒介,同时也是重要的蛀干害虫。利用CLIMEX模型分析松墨天牛分布区的气候限制因子,并在全球范围预测它的潜在分布区。模型分析结果表明,温度和降水是松墨天牛分布区的主要气候限制因子。温度在30°N以北地区和30°S以南地区主要表现为冷胁迫,在非洲中部、南亚和澳大利亚北部表现为热胁迫。有效积温不足可能是限制松墨天牛向北扩散的主要原因。降水在中国西北地区、非洲中北部、澳大利亚中部和西部与美国西部主要表现为干胁迫。降水量对分布区范围影响不大。预测结果表明,松墨天牛的全球潜在分布区远远大于实际分布范围。松墨天牛在东半球的潜在分布区包括亚洲东部和南部地区、地中海沿岸、非洲的中部和南部以及澳大利亚的东部和南部,在亚洲热带的潜在分布区1年3代,地中海地区1年1代,非洲1年2～3代,在澳大利亚主要1年1代。松墨天牛在西半球的潜在分布区主要集中在美国南部和东部沿海地区,中美洲以及南美洲的广大地区,美国主要1年1代,中美洲1年2～3代,南美洲主要1年2代。