Entomopathogenic nematodes for the biological control of Colorado potato beetles- Nematode information

Entomopathogenic nematodes and Colorado potato beetle

• Colorado potato beetles (Leptinotarsa decemlineata) are a most damaging pest of potatoes but they can also cause a significant damage to tomatoes and eggplants.
• Generally, both adults and larvae feed voraciously on leaves causing hundreds of millions dollars in yield loss each year in the United States.
• Many chemical insecticides have been recommended to control these beetles but unfortunately beetles have an ability to develop resistance to insecticides.
• Entomopathogenic nematodes as biological control agents could provide an alternative to chemical pesticides in management of Colorado potato beetles.
• As entomopathogenic nematodes naturally found soil, they are very effective against soil dwelling stages of host insect pests.  For example, mature larvae of Colorado potato beetle that moves in the soil for pupation can be a very good target for entomopathogenic nematodes.
• Commercially available entomopathogenic nematode species including Steinernema carpocapsae, Steinernema feltiae, Heterorhabditis megidis, Heterorhabditis marelata and Heterorhabditis bacteriophora have showed a very high efficacy against adult, larval and prepupal stages of Colorado potato beetles when tested in soil under laboratory conditions.

Publications:

1. Ebrahimi, L., Niknam, G. and Lewis, E. E. 2011.   Lethal and sublethal effects of Iranian isolates of Steinernema feltiae and Heterorhabditis bacteriophora on the Colorado potato beetle, Leptinotarsa decemlineata.  Biocontrol 56: 781-788.
2. Ebrahimi, L.,Niknam, G.and Dunphy, G.B. 2011. Hemocyte responses of the Colorado potato beetle, Leptinotarsa decemlineata, and the greater wax moth, Galleria mellonella, to the entomopathogenic nematodes, Steinernema feltiae andHeterorhabditis bacteriophora . Journal of Insect Science 11, Article Number: 75.
3. Armer, C.A., Berry, R.E., Reed, G.L. and Jepsen, S.J. 2004.  Colorado potato beetle control by application of the entomopathogenic nematode Heterorhabditis marelata and potato plant alkaloid manipulation. Entomologia Experimentalis et Applicata. 111: 47-58.
4. Berry, R.E., Liu, J. and Reed, G. 1997.  Comparison of endemic and exotic entomopathogenic nematode species for control of Colorado potato beetle (Coleoptera : Chrysomelidae). Journal of Economic Entomology. 90: 1528-1533.
5. Cantelo, W.W. and Nickle, W.R. 1992. Susceptibility of prepupae of the Colorado potato beetle (coleoptera, chrysomelidae) to entomopathogenic nematodes (Rhabditida, Steinernematidae, Heterorhabditidae). Journal of Entomological Science. 27: 37-43.
6. Nickle, W.R., Connick, W.J. and Cantelo, W.W. 1994. Effects of pesta-pelletized Steinernema-carpocapsae (all) on western corn rootworms and colorado potato beetles. Journal of Nematology. 26: 249-250.
7. Trdan, S., Vidrih, M., Andjus, L. and Laznik, Z. 2009. Activity of four entomopathogenic nematode species against different developmental stages of Colorado potato beetle, Leptinotarsa decemlineata (Coleoptera, Chrysomelidae. Helminthologia. 46: 14-20.

Entomopathogenic nematode identification with a quantitative real-time PCR (qPCR)

Entomopathogenic nematodes and qPCR Quantitative real-time PCR (qPCR) technique can be used for the identification of entomopathogenic nematodes in the both Heterorhabditidae and Steinernematodae families directly from soil samples.

Species specific primers and TaqMan (R) probes from the ITS rDNA region for the EPNs were used for the identification of four species of entomopathogenic nematodes including Heterorhabditis bacteriophora, Steinernema carpocapsae, Steinernema feltiae and Steinernema scapterisci (Campos-Herrera et al., 2011).

A publication on indentification of entomopathogenic nematodes using quantitative real-time PCR (qPCR) technique.

Campos-Herrera, R., El-Borai, F.E., Stuart, R.J., Graham, J.H. and Duncan, L.W. 2011.   Entomopathogenic nematodes, phoretic Paenibacillus spp., and the use of real time quantitative PCR to explore soil food webs in Florida citrus groves. Journal of Invertebrate Pathology 108: 30-39.

First record of entomopathogenic nematodes in Labanon

A presence of entomopathogenic nematode species including Heterorhabditis bacteriophora and Steinernema feltiae has been reported for the first time in Lebanon (Noujeim et al., 2011). Read following paper for survey methods

Noujeim, E., Khater, C., Pages, S., Ogier, J.C., Tailliez, P., Hamze, M. and Thaler, O. 2011. The first record of entomopathogenic nematodes (Rhabiditiae: Steinernematidae and Heterorhabditidae) in natural ecosystems in Lebanon: A biogeographic approach in the Mediterranean region. Journal of Invertebrate Pathology 107: 82-85.

Influence of potting media on the virulence of entomopathogenic nematodes against black vine weevil, Otiorhynchus sulcatus

It has been demonstrated that five different types of commercial potting media including peat, bark, coir, and peat blended with 10% and 20% compost green waste can influence the virulence of entomopathogenic nematodes against third-instar black vine weevil, Otiorhynchus sulcatus.  For example, Heterorhabditis species including Heterorhabditis bacteriophora UWS1 strain, H. megidis, H. downesi can cause 100% mortality of black vine weevil grubs in all the five types of media but  Steinernema species including Steinernema feltiae, S. carpocapsae, and S. kraussei can cause 100% black vine weevil grub mortality only in the peat blended with 20% compost green waste.  These results suggest that when growers are selecting entomopathogenic nematodes to control black vine weevil, Otiorhynchus sulcatus in their nurseries/greenhouses, they should take into consideration the type of potting media used in growing their plants. Please read following paper for the information on the method of nematode application rates and timings.

Ansari, M. A. and Butt, T. M. 2011.  Effect of potting media on the efficacy and dispersal of entomopathogenic nematodes for the control of black vine weevil, Otiorhynchus sulcatus (Coleoptera: Curculionidae). Biological Control 58: 310-318.

Ansari, M.A., Shah, F.A. and Butt, T.M. 2010.  The entomopathogenic nematodeSteinernema kraussei and Metarhizium anisopliae work synergistically in controlling overwintering larvae of the black vine weevil, Otiorhynchus sulcatus, in strawberry growbags. Biocontrol Science and Technology. 20: 99-105.

Compatibility of entomopathogenic nematodes with chemical pesticides

Recently, Radova (2011) reported that the chemical pesticide fenpyroximate showed no adverse effect on virulence of entomopathogenic nematode Heterorhabditis bacteriophora but it reduced the virulence of Steinernema feltiae against the insect called mealworm Tenebrio molitor under laboratory conditions. For more information, read following papers on related topics

Garcia-Del-Pino, F. and Morton, A. 2010.  Synergistic effect of the herbicides glyphosate and MCPA on survival of entomopathogenic nematodes  Biocontrol Science and Technology.  20: 483-488.

Gutierrez, C., Campos-Herrera, R. and Jimenez, J. 2008.  Comparative study of the effect of selected agrochemical products on Steinernema feltiae (Rhabditida : Steinernematidae).  Biocontrol Science and Technology.  18: 101-108.

Negrisoli, A.S., Garcia, M.S., Negrisoli, C.R.C.B. 2010a.  Compatibility of entomopathogenic nematodes (Nematoda: Rhabditida) with registered insecticides for Spodoptera frugiperda (Smith, 1797) (Lepidoptera: Noctuidae) under laboratory conditions.  Crop Protection 29: 545-549.

Negrisoli, A.S., Garcia, M.S., Negrisoli, C.R.C.B., Bernardi, D. and da Silva, A. 2010b.  Efficacy of entomopathogenic nematodes (Nematoda: Rhabditida) and insecticide mixtures to control Spodoptera frugiperda (Smith, 1797) (Lepidoptera: Noctuidae) in corn. Crop Protection. 29: 677-683.

Radova, S.  2011.  Effects of selected pesticides on survival and virulence of two nematode species. Polish Journal of Environmental Studies.  20: 181-185.

Biological control of codling moth, Cydia pomonella with entomopathogenic nematodes

It has been demonstrated that the Entomopathogenic nematodes including Steinernema carpocapsae and Steinernema feltiae have a potential to use as effective biological control agent against diapausing cocooned codling moth, Cydia pomonella larvae in miniature fruit bins. Read following paper for more information on efficacy of entomopathogenic nematodes against codling moth

Lacey, L.A., Neven, L.G., Headrick, H.L., Fritts, R. 2005.  Factors affecting entomopathogenic nematodes (Steinerneniatidae) for control of overwintering codling moth (Lepidoptera : Tortricidae) in fruit bins. Journal of Economic Entomology 98: 1863-1869.

Entomopathogenic nematodes as biological control agents for sheep lice, Bovicola ovis

Biological control of sheep lice, Bovicola ovis with entomopathogenic nematodes Four entomopathogenic nematodes including Steinernema carpocapsae, Steinernema riobrave, Steinernema feltiae and Heterorhabditis bacteriophora have showed a very high efficacy against sheep lice, Bovicola ovis when tested under laboratory conditions at different incubation temperatures (James et al., 2010).  However,  the efficacy all the four species of entomopathogenic nematodes varied with the nematode species and incubation temperature.

For more information on the interaction between entomopathogenic nematodes and sheep lice read following paper.

1. James, P. J., Hook, S.E. and Pepper, P. M. 2010. In vitro infection of sheep lice (Bovicola ovis Schrank) by Steinernematid and Heterorhabditid nematodes. Veterinary Parasitology 174: 85-91.

Entomopathogenic nematodes can be used for controlling pests of stored grains

It has been demonstrated that the efficacy of entomopathogenic nematodes (Heterorhabditis bacteriophora, Steinernema carpocapsae, and Steinernema feltiae against various stored grain pests (Mediterranean flour moth, Ephestia kuehniella, lesser grain borer, Rhyzopertha dominica, rice weevil, Sitophilus oryzae and confused flour beetle, Tribolium confusum) of wheat (Triticum aestivum L.) varied with nematode dosages and temperature in the storage structures. Please read following papers for detailed information on the interaction between entomopathogenic nematodes and stored grain pests.

Athanassiou, C.G., Kavallieratos, N.C., Menti, H. and Karanastasi, E. 2010.  Mortality of four stored product pests in stored wheat when exposed to doses of three entomopathogenic nematodes.  Journal of Economic Entomology. 103: 977-984.

Athanassiou, C.G., Palyvos, N.E. and Kakoull-Duarte, T. 2008.  Insecticidal effect of Steinernema feltiae (Filipjev) (Nematoda : Steinernematidae) against Tribolium confusum du Val (Coleoptera : Tenebrionidae) and Ephestia kuehniella (Zeller) (Lepidoptera: Pyralidae) in stored wheat  Journal of Stored Products Research. 44: 52-57.

Mbata, G.N., and Shapiro-Ilan, D.I. 2005.  Laboratory evaluation of virulence of heterorhabditid nematodes to Plodia interpunctella Hübner (Lepidoptera: Pyralidae). Environmental Entomology. 34: 676 – 682.

Ramos-Rodríguez, O., Campbell, J. F. and Ramaswamy, S. 2006.  Pathogenicity of three species of entomopathogenic nematodes to some major stored- product insect pest. Journal of Stored Product Research 42: 241 – 252.

Ramos-Rodríguez,O., Campbell, J. F. and Ramaswamy, S. 2007.  Efficacy of the   entomopathogenic nematodes Steinernema riborave against the stored-product pests Tribolium castaneum and Plodia interpunctella. Biological Control 40:15 -21.

Tradan, S., Vidric, M. and Valic, N. 2006.  Activity of four entomopathogenic nematodes against young adult of Sitophilus granarious (Coleptera: Curculionidae ) and Oryzophilus surinamensis ( Coleoptera: Silvanidae ) under laboratory condition. Plant Disease and Protection. 113: 168 – 173.

Kill cereal leaf beetles, Oulema melanopus with entomopathogenic nematodes

Recently, it has been demonstrated that the entomopathogenic nematodes including Steinernema feltiae strain B30, S. carpocapsae strain C101, and Heterorhabditis bacteriophora strain D54 have a potential to use as biological control agents against cereal leaf beetles (Oulema melanopus), which is a most common pest of many cereal crops including barley, corn, oats, wheat, rye, millet and rice.

For more information on interaction between entomopathogenic nematodes and cereal leaf beetles read following research paper.

Laznik, Z., Toth, I., Lakatos, T., Vidrih, M. and Trdan, S. 2010.  Oulema melanopus (L.) (Coleoptera: Chrysomelidae) adults are susceptible to entomopathogenic nematodes (Rhabditida) attack: results from a laboratory study. Journal of Plant Diseases and Protection. 117: 30-32.

Occurrence of entomopathogenic nematode Steinernema feltiae in Slovenia

Presence of an entomopathogenic nematode, Steinernema feltiae (Rhabditida: Steinernematidae) was recorded for first time in soil samples collected from grasslands and field crops in central part of Slovenia. Nematodes were isolated using Galleria-baiting technique (Bedding and Akhurst, 1975) and identified using molecular technique. Read following literature for more information

Bedding, R.A. and R.J. Akhurst. 1975. A simple technique for detection of insect parasitic rhabditid nematodes in soil. Nematologica. 21: 109-110.

Laznik, Z., Toth, T., Lakatos, T., Vidrih, M. and Trdan, S. 2009.  First record of Steinernema feltiae (Filipjev) (Rhabditida: Steinernematidae) in Slovenia. Helminthologia. 46: 135-138.

How entomopathogenic nematodes find their insect hosts (Foraging Strategies)

Infective juveniles of entomopathogenic nematodes use three different strategies to find their insect hosts.1. Ambush foraging: Ambushers such as Steinernema carpocapsae and S. scapterisci have adapted "sit and wait" strategy to attack highly mobile insects (billbugs, sod webworms, cutworms, mole-crickets and armyworms) when they come in contact at the surface of the soil.  These nematodes do not respond to host released cues but infective juveniles of some Steinernema spp can stand on their tails (nictate) and easily infect passing insect hosts by jumping on them.  Since highly mobile insects live in the upper soil or thatch layer, ambushers are generally effective in infecting more insects on the surface than deep in the soil. 2. Cruise foraging: Cruiser nematodes such as Heterorhabditis bacteriophora, H. megidis, Steinernema glaseri and S. kraussei generally move actively in search of hosts and therefore, they are distributed throughout the soil profile and more effective against less mobile hosts such as white grubs and black vine weevils.  Cruisers never nictate but respond to carbon dioxide released by insects as cues. 3. Intermediate foraging: Some nematode species such as Steinernema feltiae and S.riobrave have adapted a strategy in between ambush and cruise strategies called an intermediate strategy to attack both the mobile and sedentary/less mobile insects at the surface or deep in the soil.  Steinernema feltiae is highly effective against fungus gnats and mushroom flies whereas S.riobrave is effective against corn earworms, citrus root weevils and mole crickets.

Kill Shore flies (Scatella stagnalis) with Entomopathogenic Nematodes

• The shore fly, Scatella stagnalis (Fallén) (Diptera: Ephydridae) is an important insect pest of greenhouse plants.

• Larvae of these flies mainly feed on blue-green algae grown on the surface of plant growing media, walls, floors, benches, and pots.

• But larvae can also cause a serious damage to tender plant tissues thus reducing quality and productivity of plants.

• The adults are not considered as plant feeders but they are nuisance to people and disseminate pathogens such as Fusarium and Pythium from plant to plant as they disperse through the greenhouse.

• Currently, most growers rely on chemicals that kill host plants such as blue-green algae to reduce the incidence of shore flies. However, this method has not been proved effective in reducing shore fly incidence.

• Biological control agents including Bacillus thuringiensis var. thuringiensis (Bt) and entomopathogenic nematodes have been considered as alternatives to chemical pesticides.

• For successful control of shore flies, entomopathogenic nematodes can be easily applied in water suspension as spray application to the surface of plant growing medium.

• Entomopathogenice nematodes including Heterorhabditis megidis, Steinernema arenarium and Steinernema feltiae when applied at the rate of 50 nematodes/cm2 can cause 94- 100% mortality of shore flies.

How Entomopathogenic Nematodes kill Shore flies

• When the infective juveniles are applied to the surface of plant growing substrate, they start searching for their hosts, in this case shore fly larvae.

• Once a larva has been located, the nematode infective juveniles penetrate into the larval body cavity via natural openings such as mouth, anus and spiracles.

• Infective juveniles of Heterorhabditis spp also enter through the intersegmental members of the larval cuticle.

• Once in the body cavity, infective juveniles release symbiotic bacteria (Xenorhabdus spp. for Steinernematidae and Photorhabdus spp. for Heterorhabditidae) from their gut in the larval blood.

• In the blood, multiplying nematode-bacterium complex causes septicemia and kills shore fly larvae usually within 48 h after infection.

• Nematodes feed on multiplying bacteria, mature into adults, reproduce and then emerge as infective juveniles from the cadaver to seek new larvae in the potting medium/soil.

For more information on the interaction between entomopathogenic nematodes and leafminers, please read following research and extension publications.

• Foote, B.A. 1977. Utilization of blue-breen algae by larvae of shore flies. Environmental Entomology 6, 812-814.

• Goldberg, N.P. and Stanghellini, M.E. 1990. Ingestion-egestion and aerial transmission of Pythium aphanidermatum by shore flies (Ephydrinae: Scatella stagnalis). Phytopathology 80, 1244-1246.

• Lindquist, R., Buxton, J. and Piatkowski, J. 1994. Biological control of sciarid flies and shore flies in glasshouses. Brighton Crop Protection Conference, Pests and Diseases, BCPC Publications 3, 1067-1072.

• Morton, A., Garcia del Pino, F., 2007. Susceptibility of shore fly Scatella stagnalis to five entomopathogenic nematode strains in bioassays. Biocontrol 52: 533-545.

• Morton, A. and Garcia del Pino, F. 2003. Potential of entomopathogenic nematodes for the control of shore flies (Scatella stagnalis). Growing Biocontrol Markets Challenge Research and Development. 9th European Meeting IOBC/WPRS Working Group "Insect Pathogens and Entomopathogenic Nematodes", Abstracts, 67.

• Vanninen, I., Koskula, H. 2000. Biological control of the shore fly (Scatella tenuicosta) with steinernematid nematodes and Bacillus thuringiensis var. thuringiensis in peat and rockwool. Biocontrol Sci. Technol.. 13: 47-63.

• Zack, R.S. and Foote, B.A. 1978. Utilization of algal monoculture by larvae of Scatella stagnalis. Environmental Entomology 7, 509-511.

Kill fungus gnats using biological control agents: Insect-parasitic nematodes

• Several fungus gnat species including Bradysia coprophila, B. impatiens and B. difformis are considered economically important indoor and greenhouse pests in Europe and the US.
• Fungus gnat flies are black or gray in color with clear wings, relatively small (3-4 mm) in size and commonly associated with compost and natural soils with high organic contents.
• You can see these hopping flies when you water your plants.
• Fungus gnat maggots (larvae) are white-bodied with black heads and can be found just under the surface of the potting medium/soil.
• These maggots primarily feed on fungi and organic matter but they can also cause a serious damage to many ornamental plants.
• Maggots often chew or strip plant roots and tunnel stems affecting water and nutrient absorption in severely injured plants resulting in lost vigor, turn off-color and eventually death.
• Maggots are also capable of transmitting fungal pathogens (Fusarium, Phoma, Pythium and Verticillium) during feeding.
• Adult flies are nuisance to people and disseminate fungal spores from plant to plant as they disperse through the greenhouse.
• Females often laying over 1000 eggs in a lifetime on the media surface and completing egg-to-egg life cycle within 20-25 days at 20-25oC.
• Continuous and overlapping generations of fungus gnats in the greenhouse have made most control strategies difficult.
• Currently, most growers rely on insecticides to manage fungus gnats in floriculture.
• However, use of these insecticides is restricted due to their environmental pollution and human health concerns, development of resistance to pesticides and removal of some of the most effective products from the market.
• Biological control agents including Bacillus thuringiensis (Bt), the predatory mite, Hypoaspis miles and entomopathogenic nematodes have been used as alternatives to chemical pesticides.
• The entomopathogenic nematodes species including Heterorhabditis bacteriophora GPS11 strain, H. indica LN2 strain and Steinernema feltiae UK strain have a potential to use as biocontrol agents against fungus gnats.
• These nematodes kill both maggots (larvae) and pupae, but the second and fourth stages are most susceptible than pupae.
• Nematodes are generally applied in water suspension as spray applications to the surface of plant growing medium to target larval and pupal stages.
• The potting medium (Ball-mix, Nursery-mix or Pro-mix) can influence the survival, persistence and efficacy of entomopathogenic nematodes in greenhouse production.
• In the Nursery-mix, H. bacteriophora can survive longer and perform better than H. indica, H. marelatus Oregon, H. zealandica X1 and Steinernema feltiae against fungus gnats.
• In the Pro-mix, only H. indica have performed better than all other nematode species that tested against fungus gnats.
• Application of S. feltiae can cause 40% reduction in fungus gnat population in Ball-mix, 50% in Metro-mix and 56% in Pro-mix, but only 27% in the Nursery-mix.
• In the greenhouse, temperature can influence efficacy of nematodes. For example, H. bacteriophora and H. indica can survive and cause very high mortality of fungus gnats at warmer (above 25oC) temperatures whereas S. feltiae is generally effective against fungus gnats at cooler (below 25oC) temperatures.
• Application of an appropriate concentration of nematodes is a crucial step in the cost effective control of fungus gnats in greenhouse production.
• Generally, application of one billion infective juveniles of H. bacteriophora, H. indica or S. feltiae per acre can kill over 50% fungus gnats in greenhouse productions.

How entomopathogenic nematodes kill fungus gnats

• When the infective juveniles are applied to the surface of plant growing medium, they start searching for hosts, in this case fungus gnat maggots (larvae) and pupae.
• Once a maggot/pupa has been located, the nematode infective juveniles penetrate into the maggot body cavity via natural openings such as mouth, anus and breathing pores called spiracles.
• Infective juveniles of Heterorhabditis spp also enter through the intersegmental members of the maggot/pupal cuticle.
• Once in the body cavity, infective juveniles release symbiotic bacteria (Xenorhabdus spp. for Steinernematidae and Photorhabdus spp. for Heterorhabditidae) from their gut in the fungus gnat blood.
• Multiplying nematode-bacterium complex causes septicemia and kills the host usually within 48 h after infection.
• Nematodes feed on multiplying bacteria, mature into adults, reproduce and then emerge as infective juveniles from the cadaver to seek new maggots in the potting medium/soil.

Nematodes are now commercially available from many suppliers distributed throughout in the USA.

For more information on biological control of fungus gnats, please read following research papers/book chapters:

• Binns, E.S., 1973.  Fungus gnats (Diptera: Mycetophilidae, Sciaridae) and the role of mycophagy in soil: a review. Rev. Ecol. Biol. Sol. 18, 77-90.
• Chambers, R.J., Wright, E.M., Lind, R.J., 1993.  Biological control of glasshouse sciarid larvae (Bradysia spp.) with the predatory mite, Hypoaspis miles on Cyclamen and Poinsettia. Biocontrol Sci. Technol. 3, 285-293.
• Ecke, P.Jr., Faust, J.E., Williams, J., Higgins, A., 2004.  The Poinsettia Manual. Ball Publishing, The Paul Ecke Ranch, Encinitas, California, USA.
• Freeman, P., 1983.  Sciarid flies, Diptera; Sciaridae. Handbooks for the identification of British insects 9, Part 6. London, Royal Entomol. Soc. pp 68.
• Gillespie, D.R., Menzies, J.G., 1993.  Fungus gnat vector Fusarium oxysporum f. sp. radicislycopersici.  Ann. Appl. Biol. 123, 539-544.
• Gouge, D.H., Hague, N.G.M., 1994.  Control of sciarids in glass and propagation houses with Steinernema feltiae. Brighton Crop Protection Conference: Pest Dis. 3, 1073-1078.
• Gouge, D.H., Hague, N.G.M., 1995.  Glasshouse control of fungus gnats, Bradysia paupera, on fuchsias by Steinernema feltiae. Fundam. Appl. Nematol. 18, 77-80.
• Grewal, P.S., Richardson, P.N., 1993.  Effects of application rates of Steinernema feltiae (Nematoda: Steinernematidae) on control of the mushroom sciarid fly, Lycoriella auripila (Diptera: Sciaridae).  Biocontrol Sci. Technol. 3, 29-40.
• Grewal, P.S., Tomalak, M., Keil, C.B.O., Gaugler, R., 1993. Evaluation of a genetically selected strain of Steinernema feltiae against the mushroom sciarid fly, Lycoriella mali. Ann. Appl. Biol. 123, 695-702.
• Harris, M.A., Oetting, R.D., Gardner, W.A., 1995.  Use of entomopathogenic nematodes and new monitoring technique for control of fungus gnats, Bradysia coprophila (Diptera: Sciaridae), in floriculture. Biol. Control 5, 412-418.
• Jagdale, G. B., Casey, M. L., Grewal, P. S. and Lindquist, R. K. 2004.  Application rate and timing, potting medium and host plant on the efficacy of Steinernema feltiae against the fungus gnat, Bradysia coprophila, in floriculture. Biol. Contrl. 29: 296-305.
• Jagdale, G. B., Casey, M. L., Grewal, P. S. and Luis Cañas. 2007.  Effect of entomopathogenic nematode species, split application and potting medium on the control of the fungus gnat, Bradysia difformis (Diptera: Sciaridae), in the greenhouse at alternating cold and warm temperatures. Biol. Control. 43: 23-30.
• Kim, H.H., Choo, H.Y., Kaya, H.K., Lee, D.W., Lee, S.M., Jeon, H.Y., 2004.  Steinernema carpocapsae (Rhabditida: Steinernematidae) as a biological control agent against the fungus gnat Bradysia agrestis (Diptera: Sciaridae) in propogation houses. Biocontrol Sci. Technol. 14, 171-183.
• Lindquist R., Piatkowski J. 1993. Evaluation of entomopathogenic nematodes for control of fungus gnat larvae. Bull. Int. Organiz. Biol. Integr. Control Noxious Animals and Plants. 16, 97-100.
• Lindquist, R.K., Faber, W.R., Casey, M.L., 1985.  Effect of various soilless root media and insecticides on fungus gnats.  HortScience. 20, 358-360.
• Menzel, F., Smith, J.E., Colauto, N.B., 2003.  Bradysia difformis Frey and Bradysia ocellaris (Comstock): two additional neotropical species of black fungus gnats (Diptera : Sciaridae) of economic importance: a redescription and review. Ann. Entomol. Soc. Am. 96, 448-457.
• Nielsen, G. R., 2003. Fungus gnats. http://www.uvm.edu/extension/publications/el/el50.htm
• Oetting, R.D., Latimer, J.G., 1991.  An entomogenous nematode Steinernema carpocapsae is compatible with potting media environments created by horticultural practices. J. Entomol. Sci. 26, 390-394.
• Olson, D.L., Oetting, R.D., van Iersel, M.W., 2002.  Effect of soilless media and water management on development of fungus gnats (Diptera: Sciaridae) and plant growth. HortScience. 37: 919-923.
• Richardson, P.N., Grewal, P.S., 1991.  Comparative assessment of biological (Nematoda: Steinernema feltiae) and chemical methods of control of mushroom fly, Lycoriella auripila (Diptera: Sciaridae).  Biocontrol Sci. Technol. 1, 217-228.
• Tomalak, M., Piggott, S. and Jagdale, G. B. 2005.  Glasshouse applications. In: Nematodes As Biocontrol Agents. Grewal, P.S. Ehlers, R.-U., Shapiro-Ilan, D. (eds.). CAB publishing, CAB International, Oxon. Pp 147-166.
• Wilkinson, J.D., Daugherty, D.M., 1970.  Comparative development of Bradysia impatiens (Diptera: Sciaridae) under constant and variable temperatures. Ann. Entomol. Soc. Am. 63, 1079-1083.