Biological control of an alien tree, Acacia cyclops, in South Africa: impact and dispersal of a seed-feeding weevil, Melanterius servulus

Several Australian Acacia species have become highly invasive in a number of ecologically sensitive areas of South Africa. Many have useful attributes that, to an extent, have hampered biocontrol efforts by restricting potential agents to those reducing seed production whilst not affecting vegetative growth. The outcome of the introduction of a seed-feeding weevil, Melanterius servulus, against Acacia cyclops in 1994, was assessed by investigating the plant’s reproductive phenology, as well as levels of damage caused by the beetles. The study provided essential information on host/agent interactions and, on the build up and spread of populations of the agent. Seed damage is commonly as high as 90% (exceptionally it reaches 100%), and dispersal rates of the beetles on average is almost 2 km per year. The consistently high levels of seed damage achieved hold promise for the future of the program, however, its ultimate success will be further enhanced through the integration of biological control with other manual clearing operations that are currently employed on a large scale in South Africa.     

Implications of individual variation in insect behavior for host specificity testing in weed biocontrol

This study shows that individual behavioral variation is an under-recognised source of error that may affect the outcome of host range tests in a stenophagous species. Original specificity testing of the broom seed beetle, Bruchidius villosus (F.) (Coleoptera: Chrysomelidae: Bruchinae), a biocontrol agent for Scotch broom, Cytisus scoparius (L.) Link (Fabaceae: Genisteae), failed to detect its ability to oviposit in the field on a congeneric non-target plant, the exotic Cytisus proliferus L.f. (Fabaceae: Genisteae). These tests were repeated using individual beetles from the original UK collection sites and from New Zealand, 15 generations post release. In the original tests, low replication of small batches of females masked high levels of individual variation in oviposition preference. Although most beetles showed strong preference for the target weed, there was some indication that New Zealand beetles showed higher preference for the non-target than UK beetles.     

Confinement Behavior of Cape Honey Bees (Apis mellifera capensis Esch.) in Relation to Population Densities of Small Hive Beetles (Aethina tumida Murray)

We quantified the effects of increasing small hive beetle (Aethina tumida Murray) populations on guarding behavior of Cape honey bees (Apis mellifera capensis, an African subspecies). We found more confinement sites (prisons) at the higher (50 beetles per colony) rather than lower (25 beetles per colony) beetle density. The number of beetles per prison did not change with beetle density. There were more guard bees per beetle during evening than morning. Neither guard bee nor beetle behavior varied with beetle density or over time. Forty-six percent of all beetles were found among the combs at the low beetle density and this increased to 58% at the higher one. In neither instance were beetles causing depredation to host colonies. Within the limits of the experiment, guarding behavior of Cape honey bees is relatively unaffected by increasing beetle density (even if significant proportions of beetles reach the combs).     

Short-distance dispersal behavior and establishment of the parasitoid Gonatocerus ashmeadi (Hymenoptera: Mymaridae) in Tahiti: Implications for its use as a biological control agent against Homalodisca vitripennis (Hemiptera: Cicadellidae)

The egg parasitoid Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae), was introduced into French Polynesia as a biological control agent to control the invasive plant feeding pest Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae). The short-distance dispersal of G. ashmeadi was monitored as part of the biological control program. G. ashmeadi showed exponential dispersal capacity with 47 m/day being a minimum estimate of its natural rate of spread at high host densities (>150 nymphs per minute of sweep net sampling) in urbanized areas at sea level, which were characterized by a high diversity of exotic ornamental plants. This rate of spread contrasted starkly with almost nonexistent establishment and dispersal where host densities were very low (<2 nymphs per minute of sweep net sampling) at high elevation (800 m) with relatively undisturbed native vegetation. Survey results across different altitudes revealed an effect of vegetative diversity and host density on the measurable mobility and establishment of G. ashmeadi. In contrast, no significant influence of wind direction was found on G. ashmeadi dispersal rate or direction. Survey results for G. ashmeadi from French Polynesia suggest that the best release establishment strategies for classical biological control of H. vitripennis are: (1) many small releases where host density is high, or (2) larger and fewer releases where host densities are low.     

The Natural Host Range of Beetle Species Feeding on Broom, Cytisus scoparius (L.) Link (Fabaceae), in Southwest Europe

The natural host range of beetles feeding on broom ( Cytisus scoparius ) and 14 other species (including six other Cytisus species) in the tribe Genisteae was investigated at 39 sites in Spain , Portugal and France in 1989 and 1992 as part of a biological control programme for broom . Data on host - plant associations were analyzed for 36 phytophagous beetle species from 18 sites , and host records were listed for an additional 58 species . Nine species were apparently restricted to the genus Cytisus : Cryptocephalus octoguttatus, Gonioctena olivacea, Bruchidius lividimanus, B. villosus, Exapion elongatissimum, E. fuscirostre, Lepidapion sp . 1 , Polydrusus confluens and Tychius parallelus. These field records suggest a narrower host - plant range for some beetle species than laboratory host - range tests , and may assist in interpreting host - plant associations reported in the literature . Beetle species with a restricted host - plant range were rarely found on related non - host plants .     

Safe Site and Seed Limitation in Cytisus Scoparius (Scotch Broom): Invasibility, Disturbance, and the Role of Cryptogams in a Glacial Outwash Prairie

In all plant populations, establishment is controlled by two factors: the supply of propagules and their access to ‘safe sites’ for growth. An infestation of invading pest plants results in a seed-production gradient, from the edge where seeds are limiting, to the center where seeds may be in excess. Do invaded sites become ‘saturated’ with seeds? How rapidly does this occur, and how does the process depend on the availability of safe sites? Are safe sites, and consequently invasion, promoted by disturbance? I quantified the response of seedling establishment to seed input and disturbance in Cytisus scoparius (Scotch broom), an exotic shrub invading glacial outwash prairie remnants in western Washington, USA. I used disturbance treatments to investigate the role of the thick cryptogamic layer in these prairies, disturbing cryptogams by scraping or by fire. The effect of fire was partitioned into two factors: burning of the background vegetation/substrate versus breaking C. scoparius seed dormancy, by adding seeds either before or after the burn. Seed treatments ranged from 20 to 1000 seeds per m². Both seed number and surface treatment showed significant effects on seedling density, along with a significant interaction between the two factors. Disturbance did not promote C. scoparius establishment; undisturbed plots produced more seedlings than burned or scraped plots. Within the burned plots, fire scarification appeared to increase germination but this effect was not significant. For germinated seedlings, mortality through the dry season (June–August) was not significantly different among surface treatments, nor did survivorship depend on density, with the result that initial differences in germination among the treatments persisted. The message that the undisturbed cryptogam layer facilitates C. scoparius establishment suggests that ‘ecosystem management’ strategies promoting healthy, undisturbed sites will not always be effective against invasive pest species.     

Post hoc assessment of an operational biocontrol program: efficacy of the flea beetle Aphthona lacertosa Rosenhauer (Chrysomelidae: Coleoptera), an introduced biocontrol agent for leafy spurge

Mixed populations of Aphthona lacertosa and Aphthona czwalinae were released at more than 50 locations in Alberta in 1997. Two and 3 years post-release, beetle populations were primarily A. lacertosa, with A. czwalinae forming less than 0.5% of the sampled populations. Beetle densities were moderate (10–70 beetles per m²) or high (>70 beetles per m²) at 14% and more than 60% of the sampled sites in 1999 and 2000, respectively. Larger beetles had greater instantaneous egg loads (r²=0.424,P=0.003). In 2000, the largest beetles were found at moderate density sites and there was a significant negative relationship between beetle size and the time taken to accumulate a degree day threshold of 1230 (for females: r²=0.678,P=0.001). Sites with the most rapid accumulation of degree days have the greatest potential for beetle population growth based on potential fecundity. Changes in leafy spurge percent cover, stem density, and canopy height from 1997 to 2000 were assessed across sites with low (<10 beetles per m²), moderate, and high beetle densities in 2000. Sites with high beetle densities had significantly greater reductions of leafy spurge within 5 m of the release point than sites with low beetle densities (P<0.017). Damage caused by the beetles at high-density sites was often visible as a halo-shaped patch of dead leafy spurge stems. The significant overall reduction of leafy spurge within release patches makes A. lacertosa a promising biocontrol agent for leafy spurge in Alberta.     

Evidence for the influence of conspecific chemical cues on <Emphasis Type="Italic">Aphthona nigriscutis</Emphasis> (Coleoptera: Chrysomelidae) behaviour and distribution

Although the distribution of biological control agents may have a significant effect upon their impacts, the mechanisms regulating these distributions are often unknown. Such is the case with Aphthona nigriscutis, a classical biological control agent of leafy spurge in North America. These beetles assume aggregated distributions at some sites but disperse rapidly at others. The potential influence of plant and insect-factors upon aggregation and dispersal was investigated to try to explain these observations. Male beetles produce a putative aggregation pheromone. Responses of conspecifics to male-associated cues are greater when beetles are feeding on host plants. Densities of beetle groups greatly impact their attractiveness. Males are more sensitive to dispersal cues and females are more sensitive to congregation cues.     

Behavioral ecology ofMyiopharus doryphorae (riley) andM. aberrans (townsend), tachinid parasitoids of the colorado potato beetle

The behavior ofMyiopharus doryphorae andM. aberrans, North American tachinid parasitoids of the Colorado potato beetle, was recorded under field and laboratory conditions throughout three growing seasons in western Massachusetts. Eight common behaviors associated with resting, searching, feeding, and larviposition were distinguished, which together accounted for nearly all daytime activity of the females of both tachinids. Several of these behaviors, and in particular larviposition, were closely related to temperature but differed between species. A sequence of five defensive behaviors by the different larval stages of the Colorado potato beetle prevented larviposition in 49% of resisted attempts and perhaps one-fourth of total larviposition attempts byMyiopharus species, yet both parasitoids were highly successful in allocating their progeny during most of the summer. Second- and third-instar beetle larvae were least effective in resisting larviposition. Females of bothMyiopharus species actively guarded recently parasitized hosts from otherMyiopharus females for a period of several minutes after larviposition during the last month of the growing season when second- and third-instar Colorado potato beetle larvae were most scarce. Laboratory studies based on the field observation that femaleM. aberrans doggedly pursued circum-diapausing adult beetles led to the first recorded account ofM. aberrans larvipositing in adult hosts. Flies gained access to a beetle’s vulnerable abdominal dorsum at the instant it lifted its elytra to initiate flight. The late-season switch ofM. aberrans to adult Colorado potato beetles contributed to a seasonal sequence of larviposition-related behaviors concordant with prevailing host densities, which should lend complementarity toM. doryphorae andM. aberrans as biological controls of pest populations.     

Host race formation in the meadowsweet and strawberry feeding leaf beetle Galerucella tenella

We investigated host race formation in Galerucella tenella, a leaf beetle which feeds preferentially on meadowsweet (Rosaceae) in its natural habitats, but has become an important pest on strawberry (Rosaceae) in agricultural areas. Beetles from two isolated sites (Skeppsvik in Sweden and Solf in Finland) were compared with respect to preference and performance. At Skeppsvik the beetles were found feeding on meadowsweet, while at Solf large populations only develop in strawberry plantations, despite the presence of meadowsweet.In reciprocal field transplantations and laboratory bioassays, beetles from both sites discriminated against their foreign host, using their natal plant to a significantly higher degree for both egg laying and feeding, but with one interesting exception, namely that beetles from Solf increased their use of and even preferred meadowsweet for feeding in the laboratory. This increased use of meadowsweet by adult Solf beetles in the laboratory (without conspecific competitors) may be attributed to a density-dependent strategy, whereby mothers avoid pre-emptying the nutritional resources of the plants that will host their offspring in cases where no competitors are present.Larval fitness did not differ significantly between host plants, although larval survival of Solf beetles was halved when reared on their non-natal host plant (meadowsweet). Although beetles from both sites preferred to oviposit on their local host plant, our results provide little evidence that this presumed adaptation has to do with the nutritional quality of the plants involved. Instead, we suggest that other factors associated with the plants, such as enemy-free space and resistance to drought may be more important selective agents, shaping host preference in the field.     

Native-range research assists risk analysis for non-targets in weed biological control: the cautionary tale of the broom seed beetle

Abstract  This case study considers the broom seed beetle, Bruchidius villosus , a narrowly oligophagous species within the Fabaceae, subtribe Genistinae for which in-depth native-range studies have been vital to help understand the likely field host specificity following release. Bruchidius villosus has been used in three countries as a classical biological control agent against Scotch broom, Cytisus scoparius . Original host-specificity testing of a UK population, where this species had only been observed developing on C. scoparius , suggested this population was specific to the target. The beetle was released in New Zealand. Following release, however, the agent exhibited a broader host range than in the tests, but not a broader host range than that of the species as a whole. Subsequent studies in the native range using surveys and field testing have helped to show why B. villosus populations exhibit higher specificity in the native range than would be expected from the species' host range. This case is used to illustrate the contribution native-range studies can make to science-based risk analysis of biological control agents against weeds. By doing so, they also highlight the associated risks of ignoring native-range studies and adopting of a 'grab-and-run' approach to obtaining classical biological control agents.     

The interactions of tropical soda apple mosaic tobamovirus and Gratiana boliviana (Coleoptera: Chrysomelidae), an introduced biological control agent of tropical soda apple (Solanum viarum)

Tropical soda apple (Solanum viarum Dunal (Solanaceae) is a South American invasive plant of rangelands, pastures and natural areas in Florida. A chrysomelid beetle from South America, Gratiana boliviana Spaeth, has been released at >300 locations in Florida for biological control of tropical soda apple since 2003. Tropical soda apple is a host of several plant viruses, including the newly described tropical soda apple mosaic virus (TSAMV). We investigated the influence of TSAMV infection of tropical soda apple plants on developmental time, leaf tissue consumption, longevity, fecundity, and feeding preference of G. boliviana, and also tested transmission of the virus by the beetle. Developmental time was approximately 10% slower, and adults consumed only about 50% as much leaf tissue, for beetles fed on infected plants compared to uninfected plants. Longevity did not differ between females reared on infected and uninfected plants, but females fed on uninfected plants produced 71% more eggs than those fed on infected plants. Adult G. boliviana preferentially fed on uninfected plants when given a choice. There was no evidence of TSAMV transmission by G. boliviana. The potential impacts of TSAMV infection on the effectiveness of G. boliviana as a biological control agent are discussed.     

Establishment and dispersal of the biological control weevil Rhinoncomimus latipes on mile-a-minute weed, Persicaria perfoliata

Mile-a-minute weed, Persicaria perfoliata (L.) H. Gross (Polygonaceae), is an annual vine from Asia that has invaded the eastern US where it can form dense monocultures and outcompete other vegetation in a variety of habitats. The host-specific Asian weevil Rhinoncomimus latipes Korotyaev (Coleoptera: Curculionidae) was first released in the US in 2004 as part of a classical biological control program. The weevil was intensively monitored in three release arrays over 4 years, and field cages at each site were used to determine the number of generations produced. The weevil established at all three sites and produced three to four generations before entering a reproductive diapause in late summer. Weevils dispersed at an average rate of 1.5–2.9 m wk⁻¹ through the 50 m diameter arrays, which had fairly contiguous mile-a-minute cover. Weevils dispersing in the broader, more variable landscape located both large monocultures and small isolated patches of mile-a-minute 600–760 m from the release within 14 months. Weevil density ranged from fewer than 10 to nearly 200 weevils m⁻² mile-a-minute weed. Mile-a-minute cover decreased at the site with the highest weevil density. The production of P. perfoliata seed clusters decreased with increasing weevil populations at two sites, and seedling production declined over time at two sites by 75% and 87%. The ability of the weevil to establish, produce multiple generations per season, disperse to new patches, and likelihood of having an impact on plants in the field suggests that R. latipes has the potential to be a successful biological control agent.     

Open field host selection and behavior by tamarisk beetles (Diorhabda spp.) (Coleoptera: Chrysomelidae) in biological control of exotic saltcedars (Tamarix spp.) and risks to non-target athel (T. aphylla) and native Frankenia spp.

Biological control of invasive saltcedars (Tamarix spp.) in the western U.S. by exotic tamarisk leaf beetles, Diorhabda spp., first released in 2001 after 15 years of development, has been successful. In Texas, beetles from Crete, Greece were first released in 2004 and are providing control. However, adults alight, feed and oviposit on athel (Tamarix aphylla), an evergreen tree used for shade and as a windbreak in the southwestern U.S. and México, and occasionally feed on native Frankenia spp. plants. The ability of tamarisk beetles to establish on these potential field hosts was investigated in the field. In no-choice tests in bagged branches, beetle species from Crete and Sfax, Tunisia produced 30–45% as many egg masses and 40–60% as many larvae on athel as on saltcedar. In uncaged choice tests in south Texas, adult, egg mass and larval densities were 10-fold higher on saltcedar than on adjacent athel trees after 2 weeks, and damage by the beetles was 2- to 10-fold greater on saltcedar. At a site near Big Spring, in west-central Texas, adults, egg masses and 1st and 2nd instar larvae were 2- to 8-fold more abundant on saltcedar than on athel planted within a mature saltcedar stand being defoliated by Crete beetles, and beetles were 200-fold or less abundant or not found at all on Frankenia. At a site near Lovelock, Nevada, damage by beetles of a species collected from Fukang, China was 12–78% higher on saltcedar than on athel planted among mature saltcedar trees undergoing defoliation. The results demonstrate that 50–90% reduced oviposition on athel and beetle dispersal patterns within resident saltcedar limit the ability of Diorhabda spp. to establish populations and have impact on athel in the field.     

Horizontal transmission of a microsporidium from the convergent lady beetle, Hippodamia convergens Guérin-Méneville (Coleoptera: Coccinellidae), to three coccinellid species of Nova Scotia

Convergent lady beetles, Hippodamia convergens Guérin-Méneville, are a popular choice for aphid control in North America. An unidentified microsporidium was found in H. convergens adults that were purchased from a commercial insectary in 2004. This study examined egg cannibalism and egg predation as a means of horizontal transmission of the unidentified microsporidium among H. convergens larvae and three coccinellid species found in Nova Scotia: Coccinella septempunctata (seven-spotted lady beetle), C. trifasciata perplexa (three-banded lady beetle), and Harmonia axyridis (multicolored Asian lady beetle). The microsporidium was transmitted with 100% efficiency when first instars fed on microsporidia-infected eggs. Mean spore count data from smear preparations of infected beetles suggest that the infection was as heavy in C. trifasciata perplexa (a native coccinellid) (11.2 ± 0.96 spores/100 μm²) as it was in H. convergens (the natural host) (12.8 ± 1.16) but lighter in the introduced species C. septempunctata (7.5 ± 0.65) and H. axyridis (0.8 ± 0.11). For all of the beetle species examined, larval development was significantly longer for microsporidia-infected individuals than for their uninfected cohorts. The microsporidium had no effect on larval mortality. Based on the results of this study, field-collected H. convergens should be examined for microsporidia and uninfected individuals should be used to rear individuals for release in biological control programs. However, this is unlikely to happen because H. convergens are relatively easy and inexpensive to collect from their overwintering sites for redistribution.     

Mustard biofumigation disrupts biological control by Steinernema spp. nematodes in the soil

Mustard green manures or seed meal high in glucosinolates, which produce a natural biofumigant upon incorporation into the soil, form an alternative to synthetic fumigants. However, the non-target impacts of these biofumigants in the field are unclear. We examined the effectiveness of soil incorporation of Brassica carinata seed meal both in controlling the plant-parasitic Columbia root-knot nematode (Meloidogyne chitwoodi), and on the biological control exerted by the entomopathogenic nematodes Steinernema feltiae and Steinernema riobrave on root-knot nematodes and the Colorado potato beetle (Leptinotarsa decemlineata). Singly, both the seed meal and Steinernema spp. reduced root-knot nematode damage to potato tubers and increased marketable tuber yields. However, there was a negative interaction between the two bioagents such that their combination did not further improve suppression of plant-parasitic nematodes. Thus, mustard seed meal applications harmful to the target root-knot nematode also disrupted the ability of Steinernema spp. to act as biocontrol agents. Further, we observed modest disruption of the biological control of potato beetles following biofumigation. But, the potato beetles were less likely to lay eggs on potato plants grown in mustard-amended soil, suggesting a counteracting benefit of mustard application. Multiple, complementary controls must be integrated to replace the very effective pest suppression typical of synthetic soil fumigants. Our study suggests significant interference between biofumigation and biocontrol agents in the soil, presenting challenges in combining these two environmentally friendly approaches to managing plant-parasitic nematodes and other pests.     

Individual variation in two host plants of the ladybird beetle,Epilachna pustulosa (Coleoptera: Coccinellidae)

Individual variation in two species of host plants (thistle,Cirsium kamtschaticum, and blue cohosh,Caulophyllum robustum) of the herbivorous ladybird beetleEpilachna pustulosa was examined under laboratory conditions for their acceptability to adult beetles as a food resource, for adult preference and for larval performance. When clones of these plants were subjected to non-choice tests using posthibernating female beetles, there was found to be significant intraspecific variation among clones in terms of their acceptability, but interspecific variation was not detected. Significant intraspecific as well as interspecific variation were frequently detected in the two host plants when clones of these plants were subjected to choice tests using posthibernating female beetles; the magnitude of interspecific plant variation for beetle preference is not necessarily larger than that of intraspecific plant variation. Individual variation across plant species with respect to beetle larval performance was also significant. A positive correlation between adult preference and larval performance is suggested across the two taxonomically remote host plant species, thistle and blue cohosh, although this needs further investigation.     

植物园迁地保护环境下纹瓣兰的繁殖生物学研究

深入了解濒危物种的繁殖特性,能为物种制定有效保护策略提供科学依据。于2022和2023年对中国科学院西双版纳热带植物园迁地保存的纹瓣兰(Cymbidium aloifolium)的繁殖生物学特征进行了观察和研究,主要包括开花物候、繁育系统、花部形态特征、昆虫传粉特征以及花的挥发性成分等。结果表明,纹瓣兰的盛花期在4月中旬,单花花期7 d;群体花期约为36 d。花粉活力在开花第1天时最高,约为47.15%,之后呈下降趋势。柱头在整个单花期一直具可授性,开花第2天可授性最强。人工授粉实验表明纹瓣兰自交亲和,不存在无融合生殖和自动自花授粉,繁育系统属于兼性异交繁育并依赖传粉者,自然结实率较低为6.9%。唇瓣的先端和基部的表皮细胞分别为锥形和指状突起,基部的指状突起细胞外附着有油脂颗粒,可能是传粉者的报酬。药帽黄色,其表皮细胞为锥形细胞,能够反射光线,可能起到吸引昆虫的作用,中华蜜蜂(Apis cerana)是纹瓣兰唯一的有效传粉昆虫。纹瓣兰的挥发性气味中(E)-乙酸-2-癸烯-1-醇酯含量最高。因此,纹瓣兰的花部结构与传粉昆虫具有显著的适应性。     

Introduction, Rearing, and Host Range of Aerenicopsis championi Bates (Coleoptera: Cerambycidae) for the Biological Control of Lantana camara L. in Australia

The biology and host range of the cerambycid beetle Aerenicopsis championi Bates, a potential biological control agent for the weed Lantana camara L., were studied. A. championi is a univoltine species associated with L. camara, L. urticifolia, and L. hirsuta in Mexico and Central America. In Mexico, adult emergence occurred in May and June at the start of the rainy season. Larvae fed within the stems over a 9- to 12-month period and caused damage to the plant. The insect was imported into Australia, where a procedure for rearing it in the laboratory was developed. Host-range tests indicated that adults oviposited and larvae commenced development in L. camara and L. montevidensis but not in any of 57 other species tested. A CLIMEX model indicated that most areas infested with lantana in Australia would have a favorable climate for A. championi. Permission to release this insect in Australia was obtained and three small releases were made in southern Queensland in February 1995.