Prospects for improving biological control of olive fruit fly,Bactrocera oleae (Diptera: Tephritidae), with introduced parasitoids (Hymenoptera)

Olive fruit fly is a key pest of olive and consequently a serious threat to olive fruit and oil production throughout the Mediterranean region. With the establishment of Bactrocera oleae in California a decade ago, interest was renewed in classical (introduction) biological control of the pest. Here we discuss the prospects of identifying natural enemies of B. oleae in Africa and Asia that may help reduce B. oleae populations in California and elsewhere. Based on the current understanding of Bactrocera phylogenetics, early opinions that B. oleae originated in Africa or western Asia rather than the Mediterranean region or the Near East are taxonomically and ecologically supportable. Closely related to cultivated olive, the wild olive Olea europaea cuspidata is widely distributed in southern and eastern Africa, the Arabian Peninsula, and eastwards into Asia as far as southwestern China. Little is known regarding the biology and ecology of B. oleae in Africa and eastern Asia, especially in wild olives. While the diversity of parasitoids of B. oleae in the Mediterranean region is low and unspecialized, a diverse assemblage of parasitoids is known from B. oleae in Africa. Conversely, regions in Asia have remained largely unexplored for B. oleae and its natural enemies.     

Biology and prey range of Cryptognatha nodiceps (Coleoptera: Coccinellidae), a potential biological control agent for the coconut scale, Aspidiotus destructor (Hemiptera: Diaspididae)

Cryptognatha nodiceps Marshall is an important natural enemy used in biological control programmes against Aspidiotus destructor (Signoret), an introduced pest of coconut in many countries. In order to increase our understanding of C. nodiceps, the present study to quantify aspects of the developmental, reproductive and feeding biology under constant environmental conditions (26±2°C, 55-65% RH) was carried out. In addition, methods for culturing the scale and the beetle were developed. The average duration of development was 19.2±0.1 days for males and 19.6±0.2 days for females. Female longevity and lifetime fecundity was 35.6±5.1 days and 141.0±35.6 eggs, respectively. Life table statistics were as follows: reproductive rate, 9.99, intrinsic rate of increase, 0.09, finite rate of increase, 1.1, doubling time, 7.8 days and generation time, 41.1 days. Studies on feeding biology showed that C. nodiceps was oligophagous. The coccinellid fed and reproduced on prey species in two families, Diaspididae and Coccidae. Adults also fed on the coconut whitefly Aleurodicus cocois (Curtis) (Aleyrodidae) but no reproduction occurred on this prey. Aphididae, Psyllidae and Pseudococcidae were not fed upon.     

Energy budgets of the Chinese green lacewing （Neuroptera： Chrysopidae） and its potential for biological control of the cotton aphid （Homoptera： Aphididae）

Energy budgets of larval stages of the Chinese green lacewing, Chrysopa sinica (Tjeder) (Neuroptera: Chrysopidae) were determined under laboratory conditions at photo‐period of 14:10 L:D, 27 ± 1°C and 75%± 2% RH. The energy used as ingestion, assimilation, respiration, productivity and feces was constructed for each developmental stage. In addition, under these experimental conditions, the potential of C. sinica as a biological control agent was evaluated according to the ingestion by this predator and the energy content of cotton aphid, Aphis gossypii (Glover) (Homoptera: Aphididae). The larval stage of C. sinica was able to consume 1281.4 1‐day‐old aphids, 1018.7 2‐day‐old aphids, 626.9 3‐day‐old aphids, 393.5 4‐day‐old aphids, 312.1 5‐day‐old aphids or 203.5 9‐day‐old aphids, respectively. No significant difference was detected between the estimated number of aphids consumed by the lacewings using energetic methods and the actual number of aphids consumed by the lacewings in this experiment. Our results showed that C. sinica is an important natural enemy of the cotton aphid, and energetic methods are very useful to quantify biological control efficacy of natural enemies.     

Life history and demography of Psyllaephagus zdeneki (Hymenoptera: Encyrtidae), a potential candidate for biological control of olive psylla,Euphyllura pakistanica (Hemiptera: Psyllidae)

Life history and demographic parameters of Psyllaephagus zdeneki Noyes and Fallahzadeh (Hymenoptera: Encyrtidae) were studied on its host the olive psyllid, Euphyllura pakistanica Loginova (Hemiptera: Psyllidae). Experiments were conducted in a growth chamber at 20±1 °C, relative humidity of 60±5%, and a photoperiod of 16:8 (L: D) hours. Four different olive cultivars (Fishomi, Shenge, Oil and Yellow) were used to test possible host plant influence on parasitoid performance. The pre-imaginal developmental period of female P. zdeneki varied from 24.96 (on Fishomi) to 26.34 (on Shenge) days, and for males from 21.63 (on Fishomi) to 24.44 (on Yellow) days. Adult female longevity differed significantly among the four cultivars, ranging from 12.46 (on Fishomi) to 14.97 (on Shenge) days. For each cultivar, adult female longevity was significantly greater than male longevity. Life table parameters showed survival rates (l _x ) in newly emerged females were 84.61, 82.25, 85.71 and 78.12% on Fishomi, Yellow, Shenge and Oil, respectively. Female egg deposition was highest on Yellow (138.4 eggs per female) and lowest on Fishomi (116.3 eggs per female). The highest and lowest intrinsic rate of increase were 0.28 (on Shenge) and 0.24 (on Oil), respectively. The mean generation time ranged from 14.6 (on Shenge) to 16.7 (on Oil) days. These results are discussed with respect to the potential impact P. zdeneki as a natural enemy of E. pakistanica, the most important pest of olive in the Fars province of Iran, as well as the influence of olive cultivar on parasitoid life table parameters.     

Natural occurrence of Gibberella fujikuroi and its potential for control of the woolly aphid Ceratovacuna lanigera (Homoptera: Aphididae) in Indian sugarcane plantations

Studies were conducted to determine the natural occurrence of entomopathogenic fungi and their potential for utilisation in biological control of the woolly aphid (Ceratovacuna lanigera Zehntner) in Indian sugarcane plantations. A fungus Gibberella fujikuroi (Sawada) Wr. (Synonym: Fusarium verticillioides (Sacc.) Nirenberg, Synonym: Fusarium moniliforme (Sheld.) emend. Snyd. and Hans.) was found to be a promising biocontrol agent. It was further characterised and its field performance was studied. Reductions of aphid populations up to 60% were observed when the fungus was applied as two sprays at a week interval in the field.     

Host range of the European,rhizome-stem feeding scale Rhizaspidiotus donacis (Hemiptera: Diaspididae), a candidate biological control agent for giant reed,Arundo donax (Poales: Poaceae) in North America

The armored scale Rhizaspidiotus donacis (Leornardi) was evaluated as a potential biological control agent of the invasive reed grass Arundo donax in North America. No-choice tests, native range field surveys and non-target host exposures were used to determine the fundamental host range of the scale collected from Caloma, Spain and Perpignan, France. Thirty-five species, including two genotypes of A. donax and seven ecotypes of Phragmites australis, along with closely related grasses, economic grasses and habitat associates were tested. In quarantine no-choice testing using releases of 200 crawlers per plant, normal development of R. donacis was observed on A. donax and A. formosana, with very limited survival to the adult stage on Spartina alterniflora and Leptochloa spp. In follow-up studies using 1000 crawlers per plant, 10 live adult females were found on Leptochloa virgata, and one adult female on Spartina alterniflora, but average adult female abundance per plant was (2580%) 26-times lower on L. virgata and over (39,090%) 100-times lower on S. alterniflora than on A. donax. Field surveys were conducted at five locations in Spain and France at which A. donax infested with R. donacis, co-occurred with two non-target species of concern and R. donacis was only found on A. donax. Six-month field host exposures in Spain using potted Leptochloa plants entwined with heavily infested A. donax confirmed that R. donacis is specific to Arundo under field conditions. Based on our results, the scale R. donacis appears to be specific to the genus Arundo and is unlikely to harm native or cultivated plants in the Americas.     

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 taxonomic status of Larinus cynarae F. and Larinus latus Herbst (coleoptera: Curculionidae), and its implications for the biological control of Onopordum (asteraceae: Cardueae) in Australia

Analyses were made of the allozyme frequencies of sympatric populations of the capitulum weevils L. latus Herbst and Larinus cyarae F. in Greece. It was found that the two taxa are genetically isolated and that they can therefore be considered as separate species. This complemented ecological data showing separation of the L. latus and L. cynarae populations by their choice of Onopordum illyricum L. and Cynara cardunculus, respectively, as hosts. Onopordum spp. are serious weed problems in Australia while Cynara contains several crop plants. L. latus is a potential biological control agent of Onopordum spp., but its use had been compromised by the uncertainty of the taxonomic distinctness from L. cynarae, biotypes of which are pests of artichoke Cynara scolymus L. Clarification of its taxonomic status lifts these reservations and clears the way for the introduction of L. latus into Australia for the biological control of Onopordum thistles.     

The Oryctes virus: its detection, identification, and implementation in biological control of the coconut palm rhinoceros beetle, Oryctes rhinoceros (Coleoptera: Scarabaeidae)

In view of the increasing and devastating damage by rhinoceros beetle (Oryctes rhinoceros) to coconut palms in the middle of last century, many efforts were made to find an efficient natural control factor against this pest, which could not be controlled by pesticides. The basic procedures of these monitoring programmes are outlined together with the final detection of a virus disease in oil palm estates in Malaysia in 1963. In extensive laboratory studies, the virus was isolated and identified as the first non-occluded, rod-shaped insect virus, morphologically resembling the baculoviruses. Infection experiments clarified the pathology, histopathology, and virulence of the virus and demonstrated that the virus was extremely virulent to larvae after peroral application. These findings encouraged the first pilot release of virus in 1967 in coconut plantations of Western Samoa where breeding sites were contaminated with virus. Surprisingly, the virus became established in the Samoan rhinoceros beetle populations and spread autonomously throughout the Western Samoan islands. As a consequence, there was a drastic decline of the beetle populations followed by a conspicuous recovery of the badly damaged coconut stands. This unexpected phenomenon could only be explained after it was shown that the adult beetle itself is a very active virus vector and thus was responsible for the efficient autodissemination of the virus. The functioning of the beetle as a 'flying virus factory' is due to the unique cytopathic process developing in the midgut after peroral virus infection. Pathological details of this process are presented. Because of the long-term persistence of the virus in the populations, rhinoceros beetle control is maintained. Incorporation of virus into integrated control measures and successful virus releases in many other countries are recorded.     

Host specificity of <Emphasis Type="Italic">Tectococcus ovatus</Emphasis> (Hemiptera: Eriococcidae), a potential biological control agent of the invasive strawberry guava, <Emphasis Type="Italic">Psidium cattleianum</Emphasis> (Myrtales: Myrtaceae), in Florida

Strawberry guava, Psidium cattleianum Sabine, is a woody tree or shrub native to coastal southeastern Brazil. Strawberry guava was introduced into Florida in the late 1800s as an ornamental species. The plant escaped cultivation and is invading natural areas throughout the southern half of the state. In addition to negative effects on Florida’s native ecosystems, strawberry guava also is a preferred host of the Caribbean fruit fly, Anastrepha suspensa Loew (Diptera: Tephritidae). In total, 57 plant species representing 21 families were included in the host range tests. First instar nymphs of Tectococcus ovatus Hempel fed on two closely related guava species, Brazilian guava (Psidium friedrichsthalianum O. Berg), and Costa Rican guava (Psidium guineense Sw.). However, none of the nymphs completed their development on these two non-target species. The results of the host specificity tests suggest that T. ovatus is a suitable candidate for classical biological control of strawberry guava in Florida.     

Importation,releases, and establishment ofNeochetina spp. (Col.: Curculionidae) for the biological control of water hyacinth,Ecchhornia crassipes (Lil.: Pontederiaceae), in Benin,West Africa

Water hyacinth,Eichhornia crassipes (Martius) Solms-Laubach (Pontederiaceae) was first reported in Bénin in 1977 and about 10 years later became the major floating water weed in the south east, obstructing boat traffic and fisheries. Water hyacinth multiplies in permanently fresh water in the swampy upper reaches of the Sô River and in tributaries of the Ouémé River. From there it is moved by wind and water flow to the coastal lagoons. The coastal lagoons are brackish during the dry season and water hyacinth eventually dies. In 1991,Neochetina eichhorniae (Warner) (Col.: Curculionidae) of South American origin was imported from Australia via quarantine in Britain to Bénin. A small infestation of the fungusBeauveria bassiana (Bals.) Vuill. (Hyphomycetes) was eliminated from the colony before release by sterilizing eggs and rearing a fungus-free generation. Between late 1991 and mid 1993, about 23,900N. eichhorniae were released at 11 localities along the Ouémé River and in the head waters of the Sô River. Regular monitoring revealed feeding scars by adults on leaves and tunnelling by larvae in petioles at all release sites. By October 1993,N. eichhorniae had spread up to 20 km from some release sites.Neochetina bruchi Hustache was imported in 1992. A total of about 5,700 weevils has been released in six localities since mid 1992. Recoveries of offspring were made in all but one locality. Despite the negative impact of water flow, wind, penetration of salt water, and removal of infested water hyacinth by fishermen,N. eichhorniae andN. bruchi are established in Bénin in a situation typical for coastal West Africa.