Mythimna unipuncta (rice armyworm)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Natural enemies
- Notes on Natural Enemies
- Plant Trade
- Wood Packaging
- Impact Summary
- Detection and Inspection
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Mythimna unipuncta Haworth
Preferred Common Name
- rice armyworm
Other Scientific Names
- Cirphis unipuncta Haworth
- Heliophila Butl.
- Heliophila unipuncta Haworth
- Leucania antica Walker
- Leucania extranea Guenée
- Leucania saccharivora Butl.
- Leucania unipuncta Haworth
- Leucania unipuncta tseki Koutsaftikis
- Noctua unipuncta Haworth
- Pseudaletia adultera
- Pseudaletia unipuncta (Haworth)
- Pseudaletia unipuncta quecha Franclemont
- Sideridis unipuncta Haworth
International Common Names
- English: American army worm; American wainscot; armyworm; armyworm, American; armyworm, true; rice cutworm; true armyworm; white-speck
- Spanish: cuncunilla de las chagras; gusano soldado; gusano soldado del maiz (Mexico); isoca militar verdadera (Arg); lagarta de los cereales; oruga desfoliadora de los pastos
- French: chenille legionaire; legionaire uniponctuee; légionnaire uniponctuée; légionnaire unipunctée; noctuelle des graminees
- Portuguese: lagarta das pastagens
Local Common Names
- Brazil: lagarta do trigo; lagarta do trigo
- Germany: Eule, Amerikanische Reis-; Heerwurm
- Japan: Awa-yoto
- USA/Hawaii: peelua; poko
- PSEDUN (Mythimna unipuncta)
Summary of InvasivenessTop of page It is commonly known that M. unipuncta migrates and it is only likely to establish when the climatic conditions change. Following migration it can then remain a resident instead of being a visitor.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Lepidoptera
- Family: Noctuidae
- Genus: Mythimna
- Species: Mythimna unipuncta
Notes on Taxonomy and NomenclatureTop of page It should be noted that this species is still referred to as Pseudaletia unipuncta in North America (see Franclemont, 1951). Most taxonomists regard Pseudaletia as a subgenus of Mythimna, so the species can be known as Mythimna (Pseudaletia) unipuncta.
Past records of Mythimna (Cirphis, Leucania or Pseudaletia) unipuncta in South-East Asia, Australia and certain Pacific Islands are now known to refer to other species (CIE, 1967) including Mythimna separata (South-East Asia) or Mythimna convecta (Australia).
DescriptionTop of page Eggs
At the time of oviposition, eggs are white to pale-yellow spheres, 6-7 mm in diameter. They darken in colour as embryonic development progresses and just prior to hatching are a dark, metallic grey.
Young larvae are pale-green, whereas the more mature larvae vary from yellow-brown to grey-green, depending on diet and climatic conditions. Segmental patterning is such that three dorsal and two lateral longitudinal lines are visible on the body. Breeland (1958) provided a detailed description of the sixth-instar larva and a key to separate M. unipuncta larvae from other noctuids found in the same habitat in Tennessee, USA. See also Carter (1984).
A typical, 12-19 mm long, noctuid pupa, changing in colour from pale-amber to dark-brown during pupal development. Females are larger than males, and the sexes may be separated using the structures on the ventral surface of the terminal abdominal segments (see Figure 12 in Breeland, 1958). Carter (1984) gives details of the pupal cremaster.
Moths vary in colour from pale-beige to a dark reddish-brown, with a distinctive single white spot on each forewing. Breeland (1958) reproduced Riley's 1883 detailed description of the M. unipuncta adult. Complete descriptions of adult genitalia may be found in Franclemont (1951).
Suspected presence in South-East Asia (e.g. through introduction) should be confirmed by examination of genitalia (see also M. separata entry).
DistributionTop of page Past records of Mythimna (Cirphis, Leucania or Pseudaletia) unipuncta in South-East Asia, Australia and certain Pacific Islands are now known to refer to other species (CIE, 1967) including Mythimna separata (South-East Asia) or Mythimna convecta (Australia). A record from China (Fang et al., 1993) has not been confirmed as M. unipuncta.
M. unipuncta has been reported from all states of the USA except Alaska, Colorado, Idaho, Nevada and Wyoming. In some cases this may just reflect low densities and/or little trapping. Similarly, it has been reported from all the Canadian Provinces, but not from the Yukon Territory or the North-West Territories (JN McNeil, Laval University, Quebec, Canada, personal communication, 1995).
In northern Europe, M. unipuncta is often recorded as a migrant, but population densities have never reached pest levels. In the distribution list, where the species is occasionally present ('present, few occurrences'), it occurs as a migrant and it is only likely to establish if the conditions change (e.g. rising temperatures).
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|China||Present||Introduced||Fang et al., 1993|
|-Gujarat||Present||Patel et al., 1987|
|-Haryana||Present||Rishi et al., 2004; Rishi et al., 2004|
|-Haryana||Present||Rishi et al., 2004; Rishi et al., 2004|
|-Indian Punjab||Present||Garg and Chaudhary, 1979|
|-Jharkhand||Present||Rabindra and Devendra, 2006|
|-Karnataka||Present||Vijaykumar, and Patil, 2003; Vijaykumar, and Patil, 2003|
|-Karnataka||Present||Vijaykumar, and Patil, 2003; Vijaykumar, and Patil, 2003|
|-Maharashtra||Present||Gawande et al., 1979|
|-Rajasthan||Present||Khan and Sharma, 1971, publ. 1972|
|-Tamil Nadu||Present||Balasubramaniam, 1973|
|Turkey||Present||Introduced||Karsholt and Razowski, 1996|
|-Canary Islands||Present||Introduced||CIE, 1967|
|-Manitoba||Present||Ayre and Lamb, 1990|
|-Nova Scotia||Present||Specht, 1972|
|-Ontario||Present||McClanahan and Elliott, 1976, publ. 1977|
|-Quebec||Present, few occurrences||Fields and McNeil, 1984; McNeil et al., 2005|
|-Quebec||Present||Fields and McNeil, 1984; McNeil et al., 2005|
|-Arkansas||Present||Steinkraus and Mueller, 2003|
|-Connecticut||Present||Magnarelli and Andreadis, 2004|
|-Florida||Present||Koehler et al., 1977|
|-Illinois||Present||Roberts et al., 1977|
|-Iowa||Present||Hendrix and Showers, 1992|
|-Maryland||Present||Taylor and Shields, 1990|
|-Minnesota||Present||Wynn et al., 1988|
|-Missouri||Present||Hendrix and Showers, 1992|
|-New York||Present||Taylor and Shields, 1990|
|-Ohio||Present||Willson and Stinner, 1994|
|-Oklahoma||Present||Soteres et al., 1984|
|-Tennessee||Present||Fields and McNeil, 1984|
|-Texas||Present||Hendrix and Showers, 1992|
|-Washington||Present||Landolt and Higbee, 2002; Landolt and Higbee, 2002|
|-Washington||Present||Landolt and Higbee, 2002; Landolt and Higbee, 2002|
Central America and Caribbean
|Costa Rica||Present||Native||CIE, 1967|
|Puerto Rico||Present||Native||CIE, 1967|
|-Sao Paulo||Present||Native||CIE, 1967|
|Bulgaria||Present||Introduced||Karsholt and Razowski, 1996|
|Czech Republic||Present||Introduced||Karsholt and Razowski, 1996|
|Denmark||Present, few occurrences||Introduced||Karsholt and Razowski, 1996|
|-Corsica||Present||Introduced||Karsholt and Razowski, 1996|
|Germany||Present, few occurrences||Introduced||Karsholt and Razowski, 1996|
|Hungary||Present||Introduced||Karsholt and Razowski, 1996|
|Iceland||Present, few occurrences||Introduced||Karsholt and Razowski, 1996|
|Ireland||Present, few occurrences||Introduced||Karsholt and Razowski, 1996|
|Luxembourg||Present, few occurrences||Introduced||Karsholt and Razowski, 1996|
|Malta||Present||Introduced||Karsholt and Razowski, 1996|
|Netherlands||Present, few occurrences||Introduced||Karsholt and Razowski, 1996|
|Poland||Present, few occurrences||Introduced||Karsholt and Razowski, 1996|
|Romania||Present||Introduced||Karsholt and Razowski, 1996|
|Russian Federation||Present, few occurrences||Introduced||CIE, 1967|
|-Central Russia||Present, few occurrences||Introduced||CIE, 1967|
|-Russia (Europe)||Present, few occurrences||Introduced||CIE, 1967|
|Slovakia||Present||Introduced||Karsholt and Razowski, 1996|
|Sweden||Present, few occurrences||Introduced||Karsholt and Razowski, 1996|
|Switzerland||Present||Introduced||Karsholt and Razowski, 1996|
|UK||Present, few occurrences||Introduced||CIE, 1967|
History of Introduction and SpreadTop of page M. unipuncta was originally a neotropical species and has been known in Europe since the nineteenth century. As it is strongly migratory, it is always ready to spread but will only do so if the conditions are favourable, for example, it has not normally reached pest numbers in the Swiss Alps. However, it has begun to reach pests numbers in the Magadino plain, Tessin, Switzerland, due to climatic changes (Hächler and Brunetti, 2002).
Hosts/Species AffectedTop of page The larvae of M. unipuncta cause the most important economic losses in small grains (barley, millet, oats, rice, rye and wheat), maize and grasses (e.g. Phleum pratense). However, at high larval densities, they will feed on less preferred hosts, such as lucerne and clover (Trifolium sp.), that occur in or close to infested sites.
Forbes (1905) reported larvae feeding on amaranthus, apples, beans, cucumbers, honeysuckle, parsley, peppers, ragweed (Ambrosia sp.), strawberries, sweet potatoes and watermelons, with Beirne (1971) adding cabbages, marigolds, potatoes, sugarbeets, and turnips to the list. In addition, Forbes (1905) successfully reared M. unipuncta in the laboratory on beet, carrots, lettuces, onions, parsnips, peas, poppies, radishes and raspberries. Carter (1984) provides references to European sources.
Host Plants and Other Plants AffectedTop of page
|Avena sativa (oats)||Poaceae||Main|
|Brassica oleracea var. botrytis (cauliflower)||Brassicaceae||Other|
|Brassica oleracea var. gemmifera (Brussels sprouts)||Brassicaceae||Other|
|Brassica oleracea var. italica (broccoli)||Brassicaceae||Main|
|Chenopodium quinoa (quinoa)||Chenopodiaceae||Other|
|Hordeum vulgare (barley)||Poaceae||Main|
|Medicago sativa (lucerne)||Fabaceae||Other|
|Oryza sativa (rice)||Poaceae||Main|
|Panicum miliaceum (millet)||Poaceae||Main|
|Saccharum officinarum (sugarcane)||Poaceae||Main|
|Secale cereale (rye)||Poaceae||Main|
|Solanum lycopersicum (tomato)||Solanaceae||Other|
|Solanum tuberosum (potato)||Solanaceae||Other|
|Zea mays (maize)||Poaceae||Main|
|Zea mays subsp. mays (sweetcorn)||Poaceae||Main|
Growth StagesTop of page Fruiting stage, Seedling stage, Vegetative growing stage
SymptomsTop of page Irregular defoliation of leaves, with the midrib often left intact, is typical on small grains. On maize, larvae may be found in the youngest furled leaves. When M. unipuncta infestations coincide with heading in small grains, larvae may cut off the seed heads (barley) or feed on panicles (wheat, rice). Evident quantities of frass pellets on the ground near plants or in leaf axis may also be used to detect infestations.
List of Symptoms/SignsTop of page
|Leaves / external feeding|
|Seeds / external feeding|
|Stems / external feeding|
Biology and EcologyTop of page Life Cycle
A detailed description of the life cycle may be found in Breeland (1958). Carter (1984) reviews this species in a European context.
Females lay their eggs in tight places, such as between the leaf sheath and stem or in cracks on dry vegetation, making eggs difficult to locate under field conditions. (This is reflected by the fact that nobody claimed the $10,000 that CV Riley, the Missouri state entomologist, offered for a field-collected egg mass in the late 1800s (cited by Breeland, 1958).)
Eggs hatch after 6-12 days (at temperatures between 15-25°C). During the first two larval instars, larvae are somewhat gregarious, feeding during the day on the youngest leaves. During this period, leaves are skeletonized. By the third instar, larvae become solitary and are nocturnal feeders, chewing holes in the leaves. During the day the larvae are found in leaf axils or under dense vegetation or debris on the ground. There are six or seven larval instars, the number depending on both the type and quality of host plants. In the absence of a proper scouting system, larvae are only detected when they reach the last two instars, the time at which defoliation is the greatest and when, under epidemic conditions, mass movement of larvae in search of food occurs.
At the end of larval development (varying between 20 and 65 days from 15 to 25°C), the prepupa burrows 2-10 cm into the soil, forms a pupal cell lined with a thin silk webbing, and pupates. Moths emerge several weeks later and there is a pre-reproductive period of at least two days. Adults are nocturnal, feeding early in the scotophase on nectar or decomposing fruit. Behaviour associated with reproduction is generally expressed in the later half of the night. Both males and females may mate several times, with females producing 400-2,000 eggs over a 2 to 3 week period.
The geographic range of M. unipuncta may be divided into two zones. In the first, at sites such as Tennessee, USA (Breeland, 1958), southern France (Bues et al., 1986) and the Azores (Tavares, 1989), populations are present all year round. In such locations, the number of generations will be directly related to prevailing temperature conditions, for this species does not enter diapause (Fields and McNeil, 1984; Ayres, 1985; Bues et al., 1987) and development is continuous. Growth rates of the different developmental stages of M. unipuncta over a range of temperatures have been studied (see Guppy, 1969; Bues et al., 1987; Taylor and Shields, 1990).
However, in other locations where climatic conditions do not permit permanent occupation, such as the northern USA and Canada as well as western France, temporary populations are established annually through the arrival of immigrants. It has been hypothesized that these are not dead-end populations, but represent one part of a seasonal migration pattern in response to predictable habitat deterioration (McNeil, 1987; McNeil et al., 1994). In North America, for example, it is proposed that Mythimna unipuncta may overwinter in the southern USA but leave in the spring, before temperatures become too high, to establish summer populations in sites further north where temperature conditions are suitable. In the autumn the newly formed adults undertake a southward migration to avoid lethal winter conditions. In both cases it is proposed that the onset of migration is induced by low-temperature, short-day conditions (Turgeon and McNeil, 1983; Delisle and McNeil, 1987; McNeil et al., 1994).
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Bacillus thuringiensis thuringiensis||Pathogen||Larvae|
|Damaster blaptoides||Predator||Larvae||Hawaii||cereals; pasture plants; Poaceae|
|Ichneumon deliratorius cinctitarsis||Parasite||Pupae|
Notes on Natural EnemiesTop of page Breeland (1958) compiled a list of all published records for parasitoids of the true armyworm. However, a number of these species may have been reared from other Mythimna species, and a list restricted to M. unipuncta (sensu Franclemont, 1951) was prepared by Guppy (1967). As noted by McNeil and Turgeon (1988), a few of the Diptera listed therein may be necrophagous species and not actual parasitoids of the armyworm. However, M. unipuncta is attacked by a wide range of generalist parasitoids: detailed lists have been published for Ontario (Guppy, 1967) and Quebec (McNeil and Turgeon, 1988) in Canada; Louisiana (Burrell, 1967), Oklahoma (Soteres et al., 1984), Tennessee (Breeland, 1958) and Virginia (Laub and Luna, 1992) in the USA.
As seen in these publications, the relative importance of the different species within the parasitoid guild varies from year to year, although Marcovitch (1958) suggested that in North America the occurrence of true armyworm epidemics may be closely related to the influence of weather on the population dynamics of one of the most important larval parasitoids, Glyptapanteles (Apantales) militaris. This braconid is also the dominant parasitoid species found attacking the armyworm in the Azores (Tavares, 1989). Masses of yellow silken cocoons, containing prepupae or pupae of Glyptapantales militaris may be found associated with moribund larvae in the field.
The presence of dipteran parasitoids in the population may often be inferred from the presence of eggs, usually laid close to the head capsule on the dorsal surface of the caterpillar. It should be noted that many of the dipteran parasitoids attacking larvae may complete their development once the host has pupated and thus in some references may be referred to as pupal parasitoids as this is the stage from which they emerge. Examples include: Archytas apicifer, Athrycia cinerea, Exorista larvarum-mella, Periscepia helymus, Periscepia laevigata, Phryxe pecosensis, Winthemia rufopicta and Enicospilus purgatus.
The impact of predators has received less detailed attention: Breeland (1958) provided a review from available literature at the time he wrote his monograph. Subsequently, it has been shown that insect generalists, especially certain Coleoptera, may have a significant impact on larval populations (Laub and Luna, 1992; Clark et al., 1994).
Studies on the natural enemies of M. unipuncta have reported that it is attacked by a number of different pathogens (see, for example, Breeland, 1958; McNeil and Turgeon, 1988). The most widely studied have been several baculoviruses (see Harvey and Tanada, 1985; Tanada, 1985 and references therein): workers have examined their mode of action and also the aspects of mixed viral infections and viral-parasitoid interactions. Tanada and Hukuhara (1971) demonstrated the presence of a synergistic factor, in the capsules of a granulosis virus, that enhances the infection of a nuclear-polyhedrosis virus, while Kaya and Tanada (1973) reported a toxin of viral origin that negatively affected parasitoids.
Plant TradeTop of page
|Plant parts not known to carry the pest in trade/transport|
|Fruits (inc. pods)|
|Growing medium accompanying plants|
Wood PackagingTop of page
|Wood Packaging not known to carry the pest in trade/transport|
|Loose wood packing material|
|Processed or treated wood|
|Solid wood packing material with bark|
|Solid wood packing material without bark|
Impact SummaryTop of page
|Fisheries / aquaculture||None|
ImpactTop of page Although M. unipuncta populations are present annually, epidemics occur sporadically on local or wide scales. Due to the unpredictable nature of outbreaks, considerable crop losses may be incurred. This was particularly true when regular monitoring programmes were not in place. For example, losses in excess of $10,000,000 were reported in Kentucky and Minnesota in the 1950s (Pfadt, 1985).
However, if densities are not so high that the entire crop is destroyed, losses are of lesser importance. Attacked plants may support >25% defoliation without significant losses and the impact of larval feeding will vary with environmental conditions and the timing of infestation (Rice et al., 1982a, b; Mulder and Showers, 1986).
The implementation of conservation-tillage practices to reduce soil erosion and conserve energy may affect the population dynamics of the true armyworm. The true armyworm is favoured by maize produced under no tillage management (Tonhasca and Stinner, 1991; Willson and Eisley, 1992), especially if maize is planted in old hayfields or where crops such as rye had been grown previously (Willson and Eisley, 1992).
Detection and InspectionTop of page In many places there are now well-established scouting programmes, where in-field sampling for a variety of pests (insects, weeds and diseases), including M. unipuncta, is carried out on a regular basis.
Light traps have been used to monitor adult populations early in the season. For example, in the Maryland IPM scouting guide for small grains it notes that cumulative April trap catches that exceed 200 moths are generally associated with subsequent larval outbreaks. This information is used to alert scouts of potential problems in the following 4-6 weeks and allows for intervention, if necessary, before the more damaging late larval instars occur. Light trap catches are also useful to detect the arrival time and the density of immigrants in locations where there are no permanent populations (McNeil, 1987), again providing advanced warning of potential outbreaks.
A new attractant for trapping both sexes of this pest, comprising a combination of acetic acid and 3-methyl-1-butanol, was found to be effective by Landolt and Higbee (2002). Also see Landolt and Hammond (2001).
The identification of the M. unipuncta female sex pheromone (Hill and Roelofs, 1980; McDonough et al., 1980; Farine et al., 1981), the elaboration of an effective lure (Steck et al., 1983) and field trials evaluating different trap parameters (Turgeon et al., 1983; Hendrix and Showers, 1990) have provided an additional tool for these monitoring networks. Captures of 50 males per trap in a night have been reported in epidemic years, whereas under endemic conditions it is less than 10 per trap in a week (Turgeon et al., 1983). In the event of high trap catches (a threshold of 10 moths per trap in a night is used in the province of Quebec, Canada), agronomists, extension personnel and farmers are warned to intensify visual inspections for both damage and larvae in high-risk areas.
Zones where lodging has occurred provide excellent microclimatic conditions for resting larvae, so high densities at these sites are a good indicator of future damage. However, decisions to intervene must be based on whole field evaluations, as within-field distribution may be highly variable.
The larval densities required to justify treatment will vary depending on the crop. For example, in Maryland (USA) the threshold is 1 larva per linear foot between rows in barley, but 3-4 larvae per linear foot for wheat. A higher density is tolerated in wheat because larvae tend to feed on the tips of seeds rather than cutting off the whole seed head. In Quebec, where damage is more frequent on maize, the thresholds for treatment are when densities exceed 1 larva for every four plants and an average density of 54-64 larvae per square metre.
Prevention and ControlTop of page
If insecticide treatments are used they must be applied before caterpillars reach the most destructive last instar. When larval densities exceeding the tolerated limits are detected through monitoring, carbaryl is often used.
Alternative Control Methods
Given the unpredictable nature of outbreaks of M. unipuncta, the establishment of preventative control methods has not received a great deal of attention. Although preliminary tests on the efficacy of an entomogenic nematode have been conducted (Kaya, 1985) and this has been further explored by Medeiros et al. (2000) and Rosa et al. (2002). Furthermore, trials examining the efficacy of inundative releases of the egg parasitoid Trichogramma minutum found that satisfactory control could be achieved. The releases were in July and August from 1987 to 1990 in the region around Arribanas (at 200 m altitude) on the island of San Miguel (Azores) (J Tavares, Department of Biology, University of the Azores, personal communication). At present, the costs of rearing and application prohibit the deployment of this parasitoid, but if rendered cost effective it would be a viable means of control, especially in sites where populations persist all year long.
Pilcher et al. (1997) tested genetically engineered wheat which is prone to infestation by Bacillus thuringiensis.
ReferencesTop of page
Abbasipour H, 2006. An identification of Mythimna species in the west of Mazandaran rice fields and population fluctuations of the dominant species. Iranian Journal of Agricultural Sciences, 37(4):Pe687-Pe695, en14.
Ambethgar V; Kumaran K, 1998. Mycosis of rice cutworm, Pseudaletia unipunctata Haworth by the green muscardine fungus, Nomuraea rileyi (Farlow) Samson. Annals of Plant Protection Sciences, 6(1):114-115; 4 ref.
APPPC, 1987. Insect pests of economic significance affecting major crops of the countries in Asia and the Pacific region. Technical Document No. 135. Bangkok, Thailand: Regional Office for Asia and the Pacific region (RAPA).
Beirne BF, 1971. Pest insects of annual crops in Canada. I Lepidoptera. II Diptera. III Coleoptera. Memoirs of the Entomological Society of Canada, 78:41-42.
Breeland SG, 1958. Biological studies on the armyworm, Pseudaletia unipuncta (Haworth) in Tennessee (Lepidoptera: Noctuidae). Journal of the Tennessee Academy of Science, 33:263-347.
Bues R., Poitout S, Anglade P, Robin JC, 1986. Cycle evolutif et hivernation de Mythimna (Syn. Pseudaletia) unipuncta Haw, Lep. Noctuidae) dans le sud de la France Acta Ecological/Ecologia Applicata, 7:151-166.
Bues R; Poitout S; Robin JC; Anglade P, 1987. Studies in controlled conditions of the thermal limits for the development of Mythimna unipuncta Haw. (Lep. Noctuidae). Acta Oecologica, Oecologia Applicata, 8(1):79-89
Burrell RW, 1967. Parasites of the armyworm in Louisiana. Journal of Economic Entomology, 60:111-114.
Carter DJ, 1984. Pest Lepidoptera of Europe with special reference to the British Isles. Series Entomologica, 31:285.
Clark MS; Luna JM; Stone ND; Youngman RR, 1994. Generalist predator consumption of armyworm (Lepidoptera: Noctuidae) and effect of predator removal on damage in no-till corn. Environmental Entomology, 23(3):617-622
Farine JP; Frérot B; Isart J, 1981. Facteurs d'isolement chimique dans la sécrétion phéromonale de deux noctuelles hadeninae: Mamestra brassicae (L) et Pseudaletia unipuncta (Haw.) Compte rendu de l'Academie des Sciences. Paris, Serie III 292, 101-104.
Forbes SA, 1905. The armyworm. Annual Report of the State Entomologist of Illinois, 23:47-51.
Franclemont JG, 1951. The species of the Leucania unipuncta group, with a discussion of the generic names for the various segregates of Leucania in North America. Proceedings of the Entomological Society of Washington, 53:57-85.
Guppy JC, 1967. Insect parasites of the armyworm, Pseudaletia unipuncta (Lepidoptera: Noctuidae), with notes on species observed in Ontario. Canadian Entomologist, 99:94-106.
Guppy JC, 1969. Some effects of temperature upon larval mortality of the armyworm, Pseudaleia unipuncta (Lepidoptera: Noctuidae) under controlled conditions. Canadian Entomologist, 101:1320-1327.
Hendrix WH III; Showers WB, 1992. Tracing black cutworm and armyworm (Lepidoptera: Noctuidae) northward migration using Pithecellobium and Calliandra pollen. Environmental Entomology, 21(5):1092-1096.
Hächler M; Brunetti R, 2002. Appearance of a new pest, Pseudaletia (Mythimna, Cirphis) unipuncta Haw. (Lepidoptera: Noctuidae), in the Magadino plain (Ticino). Revue Suisse d'Agriculture, 34(5):211-215; 9 ref.
Karsholt O; Razowski J, 1996. The Lepidoptera of Europe: a distributional checklist. Stenstrup, Denmark: Apollo Books, 380 pp.
Kaya HK, 1985. Susceptibility of early larval stages of Pseudaletia unipuncta and Spodoptera exigua (Lepidoptera: Noctuidae) to the entomogenous nematode Steinernema feltip (Rhabditida: Steinernematidae). Journal of Invertebrate Pathology, 46(1):58-62
Khan RM; Sharma SK, 1971. Some new pest records on sugarbeet in India. Indian Journal of Entomology, 33(1):105-106.
Landolt PJ; Hammond PC, 2001. Species' composition of moths captured in traps baited with acetic acid and 3-methyl-1-butanol, in Yakima County, Washington. Journal of the Lepidopterists' Society, 55(2):53-58; 19 ref.
Landolt PJ; Higbee BS, 2002. Both sexes of the true armyworm (Lepidoptera: Noctuidae) trapped with the feeding attractant composed of acetic acid and 3-methyl-1-butanol. Florida Entomologist, 85(1):182-185; 14 ref.
Marcovitch S, 1958. Some climatic relations of armyworm outbreaks. Journal of the Tennessee Academy of Science, 33:348-350.
Marino PC; Landis DA, 1996. Edge of landscape structure on parasitoid diversity and parasitism in agroecosystems. Ecol Appl, in press.
McNeil JN; Cusson M; Delisle J; Tobe SS, 1994. Hormonal control of sexual behaviour in moths that migrate in response to predictable and unpredictable habitat deterioration. In: Davey KG, Peter RE, Tobe SS, eds. Perspectives in Comparative Endocrinology. Ottawa: National Research Council of Canada, 464-468.
McNeil JN; Maury M; Bernier-Cardou M; Cusson M, 2005. Manduca sexta allatotropin and the in vitro biosynthesis of juvenile hormone by moth corpora allata: a comparison of Pseudaletia unipuncta females from two natural populations and two selected lines. Journal of Insect Physiology, 51(1):55-60.
Medeiros J; Rosa JS; Tavares J; Simoes N, 2000. Susceptibility of Pseudaletia unipuncta (Lepidoptera: Noctuidae) to entomopathogenic nematodes (Rhabditida: Steinernematidae and Heterorhabditidae) isolated in the Azores: effect of nematode strain and host age. Journal of Economic Entomology, 93(5):1403-1408; 31 ref.
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