Within-plant distribution of the Melaleuca quinquenervia biological control agent Lophodiplosis trifida

The gall-producing midge Lophodiplosis trifida Gagné (Diptera: Cecidomyiidae) is an established biological control agent of the exotic tree Melaleuca quinquenervia (Cav.) Blake, which invades wetland systems of Florida (USA). Host use patterns within the tree canopy were investigated. Plant height affected within-plant distribution of galls as L. trifida attack rates were greater in lower versus higher portions of the M. quinquenervia canopy. Nonetheless, galls occurred even at tree-top levels of 13 m.     

Nonselective oviposition by a fastidious insect: the laboratory host range of the melaleuca gall midge Lophodiplosis trifida (Diptera: Cecidomyiidae)

Invasion by the Australian paperbark, Melaleuca quinquenervia, has degraded large areas of south Florida wetlands. Restoration of these wetlands requires the removal of expansive monocultures of this large tree while simultaneously curtailing its spread. Management strategies developed by federal and state agencies include biological control to halt the spread of this species and to prevent reinfestation of cleared areas. This requires biological agents able to reduce flowering, seed production, and growth while increasing mortality of seedlings and saplings. Two of the three introduced agents (Oxyops vitiosa Pascoe and Boreioglycaspis melaleucae Moore) partially meet these needs but outcomes are not spatially or temporally consistent. Thus, additional agents are needed. The bud-gall fly Fergusonina turneri Taylor, with its mutualistic nematode Fergusobia quinquenerviae Davies and Giblin-Davis, is actively being released but has not established. A fourth promising agent, the gall midge Lophodiplosis trifida Gagné, manifested an extremely narrow host range during laboratory testing. Oviposition was indiscriminant in caged environments. Small, incipient, unilocular galls were initiated on Melaleuca viminalis, but larval development ensued only on M. quinquenervia. The unilocular galls on M. viminalis did not grow and produced no adult flies. As a result, M. viminalis test plants suffered only minor cosmetic damage. Observations from both Australia and Florida attest to the ability of this midge to impede M. quinquenervia growth and kill small plants. Thus, L. trifida is safe to release and will likely contribute to management objectives for control of this pernicious wetland invader.     

Herbivory alters resource allocation and compensation in the invasive tree Melaleuca quinquenervia

Abstract. 1. Plants may compensate for the effects of herbivory, especially under favourable growing conditions, limited competition, and minimal top‐down regulation. These conditions characterise many disturbed wetlands dominated by introduced plants, implying that exotic, invasive weeds in these systems should exhibit strong compensatory responses. 2. The Australian native Melaleuca quinquenervia is highly invasive in the Florida Everglades, U.S.A., where it experiences limited competition or herbivory from native species, making it a likely candidate for compensation. The introduced biological control agent Oxyops vitiosa feeds exclusively on the seasonal flushes of developing foliage at branch apices, which represents ≈15% of the total foliar biomass. 3. The hypothesis that M. quinquenervia compensates for folivory by O. vitiosa was tested in a series of field‐based experiments. Trees experiencing folivory over four consecutive years maintained similar levels of foliar biomass after attack yet possessed twice the number of leaf‐bearing terminal stems as undamaged trees. The biomass of these stems was similar among attacked and unattacked trees, indicating that herbivore‐damaged trees produce greater quantities of smaller terminal branches. However, undamaged trees were 36 times more likely to reproduce than herbivore‐damaged trees. 4. In a separate herbivore exclusion study, a single bout of herbivory on previously undamaged M. quinquenervia trees caused an 80% reduction in reproductive structures the following season. Herbivore‐damaged trees also possessed 54% fewer fruits than undamaged trees. An increase in the herbivory frequency (two bouts per year) or magnitude (100% simulated herbivory) did not result in a further reduction in fitness. 5. It has been concluded that M. quinquenervia partially compensates for herbivory by producing new stems and replacing foliage, but this compensation results in a substantial reduction in reproduction.     

Effects of soil fertility and flooding regime on the growth of Ambrosia trifida

Understanding the effects of abiotic environmental factors on invasive plants species traits is of importance for practical prevention. To examine the effects of soil fertility and flooding regime on the growth of Ambrosia trifida L., a mesocosm experiment was conducted for 18 weeks. Two levels of soil fertility (high and low) and three types of flooding regime (non-flooded, flooded, and periodically flooded) were prepared. Shoot height and dry weight of each plant were measured. We found both individual and interactive effects of soil fertility and flooding regime on the overall growth performance of A. trifida (p?<?0.05). The highest shoot height (154.7?±?4.4 cm) and total dry weight (TDW, 13.0?±?1.4 g) were obtained under high fertility and non-flooded condition. Height and weight were relatively low under flooding conditions (flooded and periodically flooded). In particular, shoot height (102.3?±?3.2 cm) and TDW (3.2?±?0.3 g) were the lowest under low fertility and periodically flooded condition. On the other hand, the ratio of above- to below-ground dry weight was relatively high under flooded conditions, showing the adaptive phenotypic plasticity. Adventitious root formation and more biomass allocation to shoots were a flooding-adaptive mechanism of A. trifida, well developed under high fertility condition. We suggest maintaining appropriate water regime and avoiding eutrophication in wetlands would be necessary to prevent A. trifida from invading. These findings will contribute to the conservation of biodiversity in wetlands by effective management of A. trifida.

     

基于ISSR的豚草和三裂叶豚草遗传多样性研究

[目的]研究不同地理种群的豚草和三裂叶豚草的遗传多样性水平和遗传结构。[方法]应用筛选的13条ISSR引物对8个豚草居群和7个三裂叶豚草居群共240个样品进行分子标记。[结果]8个豚草居群128个样品,共扩增出304条带,多态性位点比率为98.68%;Shannon''s信息指数为0.6716,Nei''s基因多样性指数为0.4788。7个三裂叶豚草居群112个样品,共扩增出266条带,多态性位点比率为95.86%;Shannon''s信息指数为0.6593,Nei''s基因多样性指数为0.4670。豚草和三裂叶豚草各居群遗传距离较近。[结论]豚草和三裂叶豚草在居群间具有一定的遗传稳定性;居群内具有丰富的遗传多样性。豚草和三裂叶豚草遗传变异主要来源于居群内部。豚草各居群遗传距离和地理距离有显著相关性,三裂叶豚草各居群遗传距离和地理距离无显著相关性。     

Host range of Boreioglycaspis melaleucae Moore (Hemiptera: Psyllidae), a potential biocontrol agent of Melaleuca quinquenervia (Cav.) S.T. Blake (Myrtaceae), under quarantine

Research on the natural enemies of the Australian tree Melaleuca quinquenervia, a dominant, adventive weed of wetland habitats in south Florida, USA began in the late 1970s. Since then host-range tests have been conducted in Australia and Florida on three insect species: a weevil, a sawfly, and a psyllid. Herein we discuss the host-range studies of the psyllid, Boreioglycaspis melaleucae conducted in quarantine in Florida. Tests showed that the host range is very narrow, limited to one Melaleuca species, M. quinquenervia. One time, a few psyllids developed on a different species, Callistemon (=Melaleuca) citrinus, broad-leaf form, but repeated efforts to rear it again failed. In its Australian homeland, B. melaleucae has been collected on 5 closely related broad-leaved Melaleuca species including M. quinquenervia; it developed on 2 of 4 species tested, M. viridiflora (28%) and M. quinquenervia (37%). There was also 2% development on a 6th narrow-leaved Melaleuca species, M. nodosa. In the New World, there are no native Melaleuca species; in the United States, there is one introduced group, Callistemon (=Melaleuca), which is of commercial value and just one naturalized species, M. quinquenervia, which has no commercial value. Because of the near absence of Melaleuca species and B. melaleucae’s narrow host range and ability to severely damage or kill M. quinquenervia, we concluded that field release of B. melaleucae in south Florida would be safe. The first releases were made in February 2002.     

Invasion by an exotic tree alters above and belowground ecosystem components

With the widespread introduction and invasion of exotic plants there is a need for studies that quantify alterations of basic ecosystem structure and function. Ecosystem invasion by Melaleuca quinquenervia significantly altered both above- and belowground ecosystem components in this study. We measured the quantity and nutrient concentration of the litterfall, litter layer, and soil; microbial biomass pools; and rates of potentially mineralizable nitrogen and soil oxygen demand. Annual litterfall was 4.9 times higher in the non-invaded sites and contained 1.9 times more phosphorus than invaded sites. Non-invaded plots contained a larger litter layer compared to invaded plots: 2.4 ± 1.2 kg m⁻² and 0.62 ± 0.3 kg m⁻² , respectively. Lower nutrient concentration and quantity of the litter layer in the invaded plots led to changes in the aboveground storage of nutrients. In the invaded plots there was four times less carbon, seven times less nitrogen, and ten times less phosphorus stored in the organic litter layer compared to the non-invaded plots. Microbial biomass nutrient pools were consistently lower at both the 0–5 cm and 5–15 cm depth in the invaded soils compared to non-invaded soils, indicating a plant mediated change. Although M. quinquenervia altered microbial community structure, microbial activities were not different between invaded and non-invaded plots at either depth as measured by rates of soil oxygen demand and potentially mineralizable nitrogen. These changes may affect both native plant growth and water quality, and may act to promote and maintain site dominance by M. quinquenervia.     

Root development and competitive ability of the invasive species Melaleuca quinquenervia (Cav.) S.T. Blake in the South Florida flatwoods

Melaleuca quinquenervia (Cav.) S.T. Blake is an aggressive, invasive species in sub-tropical Florida that is considered a serious threat to the existing biological integrity of many subtropical ecosystems in south Florida. It prevents other species from thriving through its high rate of seed production/germination and the formation of a dense tree canopy. However, its ability to take over a site between initial seedling establishment and crown closure is not well understood. The objective of this study was to determine (i) the nature of root development with time and soil depth, and (ii) the ability of M. quinquenervia to invade and absorb nutrients from soil already occupied by native vegetation. The working hypotheses were that M. quinquenervia captures a site either by (i) tolerating competition by prolifically growing roots into soil already occupied by native plants, or (ii) avoiding competition by rooting to depths where inter-root competition is less and water supply during a drought is available. Soil trenches and in-growth trays were used to measure root distribution and growth. Root number (# m^–2), root length density (m root m^–3 soil volume), and root biomass (g root m^–3 soil) were determined. This study demonstrated that M. quinquenervia (1) is a prolific rooter with or without the presence of competing vegetation; (2) can develop root densities higher than many mature native species at an early age; (3) can develop roots in the soil surface during soil drying periods, even while competitive grasses are dying out; (4) can develop a deep root system at an early age; and (5) is an effective rooter in both moist and dry water regimes in this fluctuating water table soil. The data suggested that this species is a strong competitor through the use of both competition avoidance and tolerance mechanisms and that the rooting habit of M. quinquenervia should be an important consideration when evaluating its ability as an invasive species.     

Physiological host range of two highly specialised mutualistic symbiotes: the fly Fergusonina turneri and its obligate nematode Fergusobia quinquenerviae,potential biocontrol agents of Melaleuca quinquenervia

In Australia, galls develop on Melaleuca quinquenervia (Cav.) S.T. Blake (Myrtaceae) as a result of the mutualistic association between the fly Fergusonina turneri Taylor (Diptera: Fergusoninidae) and its obligate nematode Fergusobia quinquenerviae Davies & Giblin-Davis (Tylenchida: Sphaerulariidae). The nematode induces gall formation, whereas the fly promotes gall maturation. Together they exploit M. quinquenervia buds and may inhibit stem elongation and flower formation. We delimited the physiological host range of this pair to determine their suitability as biological control agents of invasive M. quinquenervia populations in Florida, USA. Host use was assessed for eight species of Myrtaceae native to Florida, eight phylogenetically related ornamental species and oviposition alone on five non-myrtaceous species. Although oviposition was less specific, galls developed and matured only on M. quinquenervia. After establishment, galls are predicted to prevent flower and seed production, thereby reducing the regenerative potential of M. quinquenervia. This is the first example of an insect/nematode mutualism released as biological control agents of an invasive plant.     

Electrical Penetration Graph analysis reveals population differentiation of host-plant probing behaviors within the aphid species Uroleucon ambrosiae

Uroleucon ambrosiae collected from the eastern and the southwestern United States were studied in relation to host-plant probing. In the field, eastern aphids are highly specific on Ambrosia trifida, while southwestern aphids feed on Ambrosia trifida plus many other species in the family Asteraceae. Electrical penetration graphs of insects on the principal host, A. trifida, and an additional host of southwestern populations, Heterotheca subaxillaris, revealed regional differentiation in host-associated aphid behaviors, specifically in the first phase of subcuticular probing, and in phloem finding activities. Eastern aphids used A. trifidamore efficiently than southwestern aphids, but were demonstrably less vigorous in their probing activities on H. subaxillaris.     

Biocontrol without borders: the unintended spread of introduced weed biological control agents

An underlying assumption of classical biological control implies that intentionally introduced natural enemies will remain within the boundaries that delineate the program’s area of implementation. A weed biological control program targeting Melaleuca quinquenervia in Florida, USA has resulted in the release and establishment of Oxyops vitiosa and Boreioglycaspis melaleucae. An international survey of M. quinquenervia populations in 13 other states or countries where the insects have not been intentionally introduced was initiated to monitor the long range dispersal of O. vitiosa and B. melaleucae beyond the herbivores’ intended geographic range (Florida). Surveys in 2006 resulted in the discovery of B. melaleucae within the canopies of several M. quinquenervia trees near San Juan, Puerto Rico. In 2007, O. vitiosa was observed on the island of New Providence in the Bahamas but neither herbivore was detected on nearby Grand Bahama or Andros islands. In 2009, B. melaleucae was observed attacking M. quinquenervia trees in Los Angeles, California (USA). The herbivores have not been detected on other surveyed M. quinquenervia populations in Cuba, Jamaica, Texas (USA), Costa Rica, Brazil, Hawaii (USA) or South Africa. There is no evidence to suggest that herbivore colonization of New Providence, Puerto Rico, or California was influenced by linear distance between Florida and the recipient M. quinquenervia stand. While the dispersal pathway(s) remains unknown, biological control agents were detected from 200 to >3500 km from their original release location (Florida) and at locations that have strong links via tourism and trade as indicated by the number of airline flights connecting south Florida with colonized tree populations. Implications of this unintended spread are discussed in relation to permeability of biogeographical barriers and risk assessment of biological control agents.     

豚草和三裂叶豚草不同植株部位种子萌发与入侵扩散关系

豚草(Ambrosia artemisiifoliaLinn.)和三裂叶豚草(A. trifidaLinn.)是我国恶性入侵物种,分布常呈高密度单一种群,且结实量庞大。探讨二者不同植株部位种子萌发(休眠)与扩散特点,对了解二者入侵机制具有重要意义。以新疆伊犁新源县发生的豚草和三裂叶豚草为材料,在种子成熟期,根据植株高度、枝条长度,按比例从上到下分为9个部位,对不同植株部位种子的形态特征、数量和萌发特性进行比较,分析这两种植物不同植株部位种子萌发与扩散的共性和差异性,研究二者种群密度调节和入侵的关系。结果表明:1)两个物种内不同植株部位间种子的长、宽、百粒重无显著差异,但三裂叶豚草种子的长度和宽度分别是豚草的2—3倍,百粒重高7倍。结合两个物种在伊犁地区分布差异,认为种子大小是两个物种分布区域性差异的原因之一。2)豚草和三裂叶豚草植株外部的上顶、中顶、上中部位种子数占植株总种子数量的50%,中中、下顶占比约23%,而下部的上基、中基、下中、下基的种子数占比约27%,表明当年生产的种子有近73%的比例具有远距离扩散的潜力。3)豚草和三裂叶豚草不同植株部位种子的萌发率具有上端中端下端的趋势;初始萌发时间为下端中端上端;萌发持续时间为上端中端下端。这种萌发方式避免了同一生长季大批种子同时萌发有可能导致高密度死亡的风险。基于上述研究分析,认为豚草和三裂叶豚草不同植株部位种子具有不同的适应功能。其中,上部所产生的种子具较强的扩散能力和低休眠性,有利于两物种快速占据新生境并扩大种群;而中、下部位的种子在母株周围就近扩散,翌年萌发率低,缓解了种群竞争。豚草和三裂叶豚草不同植株部位生产的种子特性和萌发差异是两个物种进行种群密度调节和扩散入侵的重要原因。     

Microsatellite segregation analysis and cytogenetic evidence for tetrasomic inheritance in the American yam Dioscorea trifida and a new basic chromosome number in the Dioscoreae

Despite the economic and cultural importance of the indigenous “Amerindian” yam Dioscorea trifida, very little is known about their origin, phylogeny, diversity and genetics. Consequently, conventional breeding efforts for the selection of D. trifida genotypes resistant to potyviruses which are directly involved in the regression of this species have been seriously limited. Our objective of this paper is to contribute to the clarification of the cytogenetic status, i.e., inheritance and chromosome number. Our results provide genetic evidence supporting tetrasomic behaviour of the genome of D. trifida based on chromosomal segregation pattern analysis using eight SSRs markers in three different crosses. This is the first reliable evidence of an autopolyploid species in the genus Dioscorea. The second major result in this study is the revealing of a new base chromosome number in the botanical section Macrogynodium to which D. trifida belongs. To date, our assumptions about the ploidy level of yams are based on the observations that the basic chromosome number is 10 or 9, and D. trifida was described as octoploid. The chromosome number of D. trifida accessions was also assessed using somatic chromosomic count techniques. Flow cytometry did not show significant variation of 2C DNA content among 80 accessions indicating homogeneity of the ploidy level of the cultivated D. trifida. This suggests that autotetraploidy is well established as well as the rule for the cultivated pool of D. trifida, even if the direct diploid ancestor remains to be identified. The data presented in this paper are significant and important for the effective breeding and conservation of the species and for elucidating the phylogeny and the origins of the yam and the evolution of the genus Dioscorea.     

MORPHOMETRIC ANALYSIS OF POLLEN FROM FOUR SPECIES OF AMBROSIA (COMPOSITAE)

A morphometric approach proved useful for characterizing pollen from the Ambrosia species that are responsible for most ragweed hayfever and for correctly identifying individual pollen grains. Pollen was harvested from 4 species (A. trifida, A. artemisiifolia, A. bidentata, A. psilostachya) grown in Champaign County, Illinois, and observed with a scanning electron microscope. Quantitative measurements of 6 different exine characters were made on micrographs of 96 pollen grains from each species. Discriminant analysis based on these characters was used to obtain a population centroid for each species and indicated that these centroids were distinctly different. This analysis also included a species classification for individual pollen grains and indicated probability of species membership. About 86% of all individuals were correctly classified when the 4 species were included, with a range from 80% (A. artemisiifolia) to 95% (A. psilostachya). When three species (A. trifida, A. artemisiifolia, A. bidentata) were analyzed, about 87% of the individuals were correctly classified, and almost all individuals (97%) were classified correctly when only A. trifida and A. artemisiifolia were included. The validity of the derived discriminant functions was also tested by treating herbarium specimens as unknowns. The species and percentages of these individuals correctly classified were A. trifida (100%), A. artemisiifolia (58%), A. bidentata (100%), and A. psilostachya (31%). The classification results using three species were A. trifida (100%), A. artemisiifolia (75%), and A. bidentata (100%); and using two species were A. trifida (100%) and A. artemisiifolia (92%). Hence, the disciminant functions do seem valid for 3 species, but further investigation of A. psilostachya is needed. This type of approach may be useful in pollen analysis, especially where these species significantly contributed to aeroallergens.     

Chemotype variation of the weed Melaleuca quinquenervia influences the biomass and fecundity of the biological control agent Oxyops vitiosa

Host plant nutritional and non-nutritional variability can have a significant effect on herbivore populations by influencing survival, larval performance, and fecundity. The effect of chemical and physical variation of the leaves of two chemotypes of the weed Melaleuca quinquenervia was determined on the biomass and fecundity of the biological control agent Oxyops vitiosa (Coleoptera: Curculionidae). M. quinquenervia chemotypes were distinguished by the principal terpenoids E-nerolidol and viridiflorol using gas chromatography and mass spectroscopy. Not only were the terpenoid profiles of the two chemotypes different but the viridiflorol leaves had greater toughness (1.2-fold) and reduced nitrogen (0.7-fold). When the larvae and adults were fed leaves of the E-nerolidol chemotype increased adult biomass (1.1-fold) and fecundity were found (2.6- to 4.5-fold) compared with those fed leaves of the viridiflorol chemotype. Regardless of the larval diet, when adults were fed the E-nerolidol chemotype leaves they had greater egg production compared with those adults fed the viridiflorol leaves. Moreover, adult pre-oviposition period was extended (1.5-fold) when individuals were fed the viridiflorol leaves compared with those fed the E-nerolidol leaves. By rearing the O. vitiosa weevil on the more nutritious chemotype plants these results assisted in the mass production and establishment of the M. quinquenervia biological control agent.     

Population abundance and host use pattern of Ophraella communa (Coleoptera: Chrysomelidae) in its native and introduced range

We found that the abundance of Ophraella communa, a specialist herbivore of the invasive weeds Ambrosia artemisiifolia and A. trifida, is higher in the introduced range than in its native range. We also found that the native O. communa does not feed on A. trifida, but introduced O. communa does so extensively.     

Can genomics clarify the origins of Boreioglycaspis melaleucae in California,USA?

The Australian psyllid Boreioglycaspis melaleucae is a biological control agent of Melaleuca quinquenervia in Florida (USA) but was observed attacking M. quinquenervia trees in southern California (USA). Genotyping revealed the California population matched three of eight Australian haplotypes and all three Florida haplotypes. It remains unclear if the California psyllid population arrived directly from Australia or via Florida.     

The influence of plant size on attack rates of Oxyops vitiosa,an introduced herbivore of Melaleuca quinquenervia

Oxyops vitiosa Pascoe (Coleoptera: Curculionidae) was released in Florida in 1997 as a biological control agent of the invasive tree Melaleuca quinquenervia (Cav.) S.T. Blake. Three years after release, the weevil’s dispersal into three replicated M. quinquenervia stands was monitored to determine if O. vitiosa preferentially selects larger trees during patch colonisation. Data indicated that tree size did not influence the likelihood of an attack by O. vitiosa and smaller individuals within host patches did not escape herbivory through enemy-free space.     

Puccinia xanthii forma specialis ambrosia-trifidae

The rustPuccinia xanthii forma specialisambrosia-trifidae attacks leaves of giant ragweed,Ambrosia trifida L., significantly reducing seed and pollen production, seed weight and seedling vigor. In tests, teliospores did not infectA. artemisiifolia L.,Xanthium strumarium L., and cultivated varieties of chrysanthemum, safflower, sunflower, cosmos, marigold, lettuce, endive, dahlia, zinnia, gloriosa daisy, aster, scabiosa, and gallardia. This fungus commonly attacksA. trifida in eastern and central North America, and it should be of value for the biological control ofA. trifida where introduced in Eurasia.

---

Puccinia xanthii forma specialis ambrosia-trifidae

     

Consequences of Melaleuca quinquenervia Invasion on Soil Nematodes in the Florida Everglades

The tree Melaleuca quinquenervia invades all types of habitats of South Florida leading to up to 80% loss of aboveground diversity. To examine impacts on the belowground ecosystem, we investigated the composition and diversity of nematodes from soils dominated by the invasive tree and compared them with soils supporting native plant communities at six locations across the Florida Everglades over three years. Despite the significant differences in soil type, hydrology, and native plant composition of the sites, there were consistent differences in nematode communities between soil environments under the native and invaded plant communities. The total abundance and diversity of nematodes in soils dominated by M. quinquenervia was 60% and 80% of adjacent soils under native plants. Fungal-feeding and plant-parasitic nematodes were twice as abundant under native plants as under M. quinquenervia. Nematode communities under M. quinquenervia were bacterivore-dominated, while under native vegetation plant-parasite dominated. The overall diversity of nematodes was 20% lower under the exotic than under native plants, with plant parasites being 36% and fungivores being 30% less diverse. Soil moisture, % of Ca, Mg, and clay particles and total soil C and N were greater in M. quinquenervia soils, but plant-available concentrations of P, K, Ca, and Mg as well as CEC were reduced. Overall, data suggests that the invasion process may modify soil biotic and abiotic conditions that in turn promote the advancement of the exotic M. quinquenervia and displacement of the native plants.