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9-2003 PARASITOIDS AND PARASITES OF SPODOPTERA FRUGIPERDA (LEPIDOPTERA: NOCTUIDAE) IN THE AMERICAS AND CARIBBEAN BASIN: AN INVENTORY Jaime Molina-Ochoa Universidad de Colima, [email protected]

James E. Carpenter USDA-ARS

Elvis A. Heinrichs University of Nebraska-Lincoln, [email protected]

John E. Foster University of Nebraska-Lincoln, [email protected]

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Molina-Ochoa, Jaime; Carpenter, James E.; Heinrichs, Elvis A.; and Foster, John E., "PARASITOIDS AND PARASITES OF SPODOPTERA FRUGIPERDA (LEPIDOPTERA: NOCTUIDAE) IN THE AMERICAS AND CARIBBEAN BASIN: AN INVENTORY" (2003). Faculty Publications: Department of Entomology. 546. http://digitalcommons.unl.edu/entomologyfacpub/546

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254 Florida Entomologist 86(3) September 2003

PARASITOIDS AND PARASITES OF SPODOPTERA FRUGIPERDA (LEPIDOPTERA: NOCTUIDAE) IN THE AMERICAS AND CARIBBEAN BASIN: AN INVENTORY

1 2 3 3 JAIME MOLINA-OCHOA , JAMES E. CARPENTER , E. A. HEINRICHS AND JOHN E. FOSTER 1Facultad de Ciencias Biológicas y Agropecuarias, Universidad de Colima Apartado postal no. 36, Tecomán, Colima 28100, México

2United States Department of Agriculture, Agricultural Research Service Crop Protection & Management Research Laboratory, P.O. Box 748, Tifton, GA 31793-0748, USA

3University of Nebraska Lincoln, Department of Entomology 312F Plant Industry Building, Lincoln, NE 68583-0816, USA

ABSTRACT

An inventory of parasitoids and parasites of fall armyworm (FAW), Spodoptera frugiperda (J. E. Smith), was conducted using references describing parasitized FAW eggs, larvae, pu- pae and adults collected from different crops or habitats throughout the Americas and the Caribbean Basin. The crops and countries where these parasites were reported occurring in the Americas is also inventoried. Maize was the crop where the FAW was more frequently collected followed by rice. Overall, Chelonus insularis (Cresson) had the broadest natural distribution in the Americas. For the North American region C. insulares, Chelonus sp., and Euplectrus platyhypenae (Howard) were the most relevant parasitoids. In Central America, C. insularis was the most prevalent parasitoid, and in the South American region the most prevalent parasites were Archytas incertus (Macq.), A. marmoratus (Tns.), C. insularis, and Meteorus laphygmae (Viereck). Diapetimorpha introita (Cresson) is the most important pu- pal parasitoid of FAW occurring mainly in North America. An acugutturid, Noctuidonema guyanense (Remillet & Silvain), is the most important ectoparasitic nematode attacking adults of FAW and other noctuid moths in South and Southeastern US, and Mexico in North America, Caribbean Basin, Central America, and Northern South America.

Key Words: parasitoids, fall armyworm, Chelonus insularis, Diapetimorpha introita, Noctu- idonema guyanense, maize, natural distribution, biological control

RESUMEN

Un inventario de los parasitoides y parásitos del gusano cogollero, Spodoptera frugiperda (J. E. Smith) se llevó a cabo usando referencias relacionadas con parásitos de huevos, larvas, pupas y adultos del insecto plaga colectados de diferentes cultivos en su ámbito de distribu- ción. Además, un inventario se realizó de los cultivos y países donde estos parásitos atacaron al gusano cogollero en América. La plaga fue colectada principalmente en el maíz, seguido por el arroz. Chelonus insularis (Cresson) fué el parasitoide distribuido más ampliamente en todo el ámbito de distribución del. C. insulares, Chelonus sp. y Euplectrus platyhypenae (Howard) fueron los parasitoides más prevalecientes en Norteamérica. En Centroamérica, C. insularis fue el parasitoide más prevalente, y en la región Sudamericana lo fueron Archy- tas incertus (Macq.), A. marmoratus (Tns.), C. insularis y Meteorus laphygmae (Viereck). Di- apetimorpha introita (Cresson) fue el parasitoide de pupas más importante y éste habitó en Norteamérica, principalmente. Un acugguturido, Noctuidonema guyanense (Remillet & Sil- vain) fue el nematodo ectoparásito más importante atacando adultos de gusano cogollero y otros noctuidos en el Sur y Sureste de los Estados Unidos de América y México en Nortea- mérica, Cuenca del Caribe, Centroamérica y Norte de Sudamérica.

Descriptores: parasitoides, gusano cogollero, Chelonus insularis, Diapetimorpha introita, Noctuidonema guyanense, maíz, ámbito de distribución, control biológico

Biodiversity in agro-ecosystems can be as var- biodiversity can be used for improved pest man- ied as the crops, weeds, arthropods, and microor- agement (Altieri 1991). A major problem in all ar- ganisms themselves, and may differ according to eas of agriculture is the lack of basic research on geographical location, climate, soil, and human taxonomy of insect pests and their natural ene- factors. Experimental evidence suggests that mies. This problem is greatest in tropical coun-

Armyworm Symposium 2002: Molina-Ochoa et al. 255

tries where the needs are also greater (Claridge MATERIALS AND METHODS 1991). Waage (1991) stated that agricultural sys- tems are generally simpler than the natural hab- Sources of Information itats from which they are developed. These systems have fewer plant species, fewer primary Research was conducted to obtain information, consumers, and generally fewer natural enemy papers, and bibliographic references reporting species. In tropical systems, despite their great the collection of parasitized FAW from the field. biodiversity, natural enemies could be particu- In the US, we used the Agricola and CAB Ab- larly susceptible to local extinction as a result of stracts database at the University of Nebraska- habitat destruction and unfavorable cropping Lincoln in Lincoln, Nebraska, and the University practices such as indiscriminate use of pesticides of Georgia, Coastal Plain Experiment Station in (Claridge 1991; Waage 1991). Tifton, Georgia. We collected references cited in More than 200 years ago, the fall armyworm reviews and catalogs of FAW parasites (Guima- (FAW), Spodoptera frugiperda (J. E. Smith), was rães 1971; Marsh 1978; Ashley 1979; Ashley recognized as a destructive pest of many agricul- 1986; Andrews 1988) to verify information con- tural crops (Luginbill 1928). In the continental cerning host plant of parasitized FAW, location of United States the costs for chemical control and the collected FAW, and stage of FAW attacked. We losses due to this pest exceeded $300,000,000 dur- also used various internet search engines to iden- ing 1977 (Gross & Pair 1986). The biological con- tify published reports of parasitized FAW col- trol of FAW in areas of overwintering and lected from the field. In México, we collected throughout its annual geographical distribution papers from Latin American journals and pro- is a highly desirable alternative to conventional ceedings from meetings of International and Mex- control methods (Gross & Pair 1986). Luginbill ican Entomological and Biological Control (1928) and Vickery (1929) recognized the value of Societies. Information was also obtained from the parasitoids in reducing larval populations of libraries in the Facultad de Ciencias Biológicas y FAW. Parasitoids and parasites can be highly ef- Agropecuarias (School of Biological, Agricultural fective at little or no cost, serve as biotic insecti- & Animal Sciences) of the Universidad de Colima, cides in place of chemicals, provide long-term and in the Centro Nacional de Referencia de Con- control without the target pest developing signif- trol Biológico (CNRCB)-Comisión Nacional de icant resistance to them, and impose minimal or Sanidad Agropecuaria-SAGARPA (National Cen- no harm to humans or the environment (Wilson & ter for Reference on Biological Control) in Huffaker 1976; Stary & Pike 1999). Tecomán, Colima, México, during 2001 and 2002. Sivasubramaniam et al. (1997), and Dent (2000) advocated that the first step in any inves- Organization of the Information tigation of the role of natural enemies in pest con- trol should involve a field survey to determine The classification (order, family, genus and which species are present and how their numbers species) of each parasite and parasitoid collected vary in relation to those of the pest insects. Sur- from FAW is presented in Table 1. The crop from veys on the FAW parasitoids and other natural which parasitized FAW were collected, the FAW enemies in different parts of its range have been stage attacked, the country from which parasit- conducted because of increasing economic and en- ized FAW were collected, and the bibliographic vironmental concerns (Carrillo 1980; Ashley references for each record were included in Table 1986; Castro et al. 1989; Gross & Pair 1991; Cave 1 whenever the information was available. We 1993; Lezama-Gutiérrez et al. 2001; Molina- preferred to list only original references that re- Ochoa et al. 2001). However, information about port the collection of FAW from the field. There- distributions and host plants of the FAW and the fore, review articles that contain lists of insects accompanying parasites and parasitoids are scat- collected by other authors (Guimarães 1971; tered throughout the published literature, and Marsh 1978; Ashley 1979; Ashley 1986; Andrews most reviews of FAW parasitoids have empha- 1988) usually are omitted for the entries in Table sized those attacking the egg and larval stages 1. Because of the quantity involved, all references (Ashley 1986). The aim of this paper is to summa- listed in Table 1 were not cited in the text. Data rize the information and provide an inventory of on FAW parasitoids and parasites from Table 1 the known FAW parasites and parasitoids occur- have been summarized as the number of species ring in the Americas and the Caribbean, indicat- in each taxon reported from different geographi- ing the host stage attacked, the crops from which cal regions (Table 2). Number of species and FAW parasitized fall armyworm were collected, and stage attacked reported from different countries the country of collection. In addition, because par- (Table 3), and number of species in each taxon re- asites and parasitoids of the pupal and adult ported from FAW collected from different host stages of FAW have received little attention in plants (Table 4). Omission of pertinent literature most previous reviews on this subject, we provide from this paper is the authors’ responsibility and an expanded discussion of these natural enemies. was unintentional.

256 Florida Entomologist 86(3) September 2003 WITH

COLLEC-

BASIN OF

ARIBBEAN COUNTRIES

C AND AND

Reference MERICAS A COLLECTED

THE

IN WERE

irla et al. 1999 irla et al. irla et al. 1999 irla et al. erán 1974 erán 1974 erán 1974 erán 1974 Cave 1993 Cave Maes 1989 Enkerlin 1975 T V Guimarães 1977 1957 Etcheverry T Guimarães 1977 1993 Cave Notz 1972 T V Alam 1979 1993 Cave & Stehr 1984 Ravlin Maes 1989 Luginbill 1928 T Cave 1993 Cave Maes 1989 Lacayo 1977 Lacayo Ashley 1986 Ryder & Pulgar 1969; ARMYWORM

PARASITOIDS

2 AND FALL

(N/G) M (N/G) (N/G) M (N/G) M (N/G) (N/G) (N/G) M (N/G) M M M,R,O,T M M (N/G) (N/G) (N/G) M M,S M (N/G) Luginbill 1928 WHICH

PARASITES ), FROM

CROPS , OCTUIDAE collection Crop Country of : N S enezuela enezuela enezuela enezuela Honduras Nicaragua V Argentina Brazil Chile V Brazil Honduras V Barbados Honduras Mexico Nicaragua US V Nicaragua Nicaragua ATTACKED

1 EPIDOPTERA STAGE

L L L LU ) (L attacked Host stage HOST , MITH FAMILY (J. E. S

INDICATING

(Fabricius) L1921 US BG Allen Aldrich (Walker) (Reinhard) Spodoptera frugiperda (Townsend) L Honduras M 1993 Cave Maes 1989; , REFERENCES (Weideman) L US M Dew 1913 (Coquillett) L US (N/G) Luginbill 1928

(Weideman) L Lesser Antilles M 1947 Fennah (Walker) (Aldrich) Townsend (Reinhard) (Walker) L US (N/G) & Stehr 1984 Ravlin (Walker) Fabricius L Argentina sp. L US ARMYWORM .

sp. L Mexico sp. L Honduras ALL ACCOMPANYING TION Sarcophaga morionella Sarcophaga assidua Spallanzania vetula Archytas divisus Archytas 1. F

hinidae c Syn: Syn: Diptera: Phoridae Diptera: Poecilanthrax (Anthrax) lucifer Megaselia Helicobia morionella Syn: Rivinia assidua Sarcophaga georgiana Sarcophaga lambens Sarcophaga Sarcodexia sternodontis Ta Acroglossa vetula Syn: Admontia degeerioides analis Archytas apicifer Archytas incasana Archytas Syn: Syn: ABLE Diptera: Bombyliidae Diptera: Classification of parasite and parasitoid Diptera: Tachinidae Diptera: Diptera: Sarcophagidae Diptera: T

Armyworm Symposium 2002: Molina-Ochoa et al. 257 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

kery 1929 c irla et al. 1999 irla et al. valos 1988 WHICH an Dine 1913 alicente & Barreto 1999; Virla et al. 1999 et al. Virla alicente & Barreto 1999; alicente 1989 1973 Valdivia alencia & an Dither 1960 arker et al. 1953 arker et al. et al. Milward Guimarães 1977; 1953; arker et al. Silveira & Ruffinelli 1956 1953; arker et al. air et al. 1986 air et al. & Fuxa 1992 Pantoja 1985; antoja et al. ennah 1947 aseen 1979 ones 1913

V 1997 Avalos Molinari & P Alam 1979 1984, & Habib 1982, Patel Almeida 1980; & Lucchini Silva et al. 1989; Valicente 1987; Silveira et al. 1986; 1997 P 1991a,b,c,d,e 1957 Etcheverry Curran 1927 V J Segeren & Sharma 1979 Hynes 1942 Vi Luginbill 1928 P V 1997 Avalos Molinari & A Alam 1979 V Guimarães 1977 V 1975 Bruner et al. 2000 Benzing et al. 1983 Malausa 1981, Canas & O’Neil 1998 1993 Cave F P 1977 Lacayo Maes 1989; Huis 1981; Sarmiento & Razuri 1978 P V Y 1984 Soteres et al. PARASITES FROM

), 2 CROPS , M SY (N/G) M M (N/G) M SC SC SC,M M,PN M M,PN (N/G) (N/G) M SY (N/G) M M (N/G) (N/G) (N/G) Ve M M P,S,M M M,S M M R N/G) M A OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST , eru Barbados Brazil Chile Mexico Puerto Rico Suriname Trinidad US Uruguay Barbados Brazil Chile Cuba Ecuador Guadeloupe Honduras Lesser Antilles Mexico Nicaragua P Puerto Rico Suriname Trinidad US MITH 1 FAMILY

(J. E. S

attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , . ARMYWORM

ACCOMPANYING

ALL (Townsend) L Argentina ) F WITH

(Macquart) L Argentina COLLECTION

OF BASIN

ONTINUED TRIES BEAN 1. (C

Archytas incertus Archytas marmoratus Archytas ABLE Classification of parasite and parasitoid T 258 Florida Entomologist 86(3) September 2003 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

all & Berberet 1975; Nickle 1976 Nickle all & Berberet 1975; irla et al. 1999 irla et al. WHICH era et al. 1995 era et al. erán 1974 erán 1974

Campos 1965; Enkerlin 1975; Hogg et al. 1982; Gross 1982; Hogg et al. Enkerlin 1975; Campos 1965; 1993 1992, Riggin et al. 1991; 1986, & Pair 1994 Tingle et al. McCutcheon 1991 1986; et al. Pair Rohlfs & Mack 1985; Butler 1958a Reed 1980 & Stehr 1984 Ravlin & Clavijo 1984 Fernández Notz 1972; T V Guimarães 1977 King & Saunders 1984 1993 Cave Estrada 1960 Hynes 1942 V 1997 Avalos Molinari & Guimarães 1977 Enkerlin 1975 1989 Wheeler et al. Carrillo 1980 Estrada 1960 1976 Nickle 1975 Bruner et al. W T 1993 Cave Maes 1989 Sabrosky 1981 1980 Ashley et al. 1994 Tingle et al. Sabrosky 1981 PARASITES FROM

), 2 CROPS , ,A,M,S ,PN ,PN (M M,C M,S C M,BG,Mi (N/G) M (N/G) M (N/G) M (N/G) M M M SY (N/G) PN M M M PN (N/G) PN (N/G) S M S M,C S OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST , enezuela enezuela V Brazil Honduras Nicaragua Trinidad Brazil Chile Honduras Mexico Nicaragua US US V Nicaragua US Brazil S,PN Sabrosky 1981 MITH 1 FAMILY

(J. E. S

L attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , . ARMYWORM

ACCOMPANYING

ALL (Townsend) L (Coquillett) L Cuba (Townsend) L Peru (N/G) Enkerlin 1975 (Coquillet) L US M ) F WITH

Sabrosky L Honduras Curran L Argentina COLLECTION

OF BASIN

sp. L Argentina (N/G) 1953 et al. Parker sp. L Honduras M1993 Cave ONTINUED sp. L Argentina TRIES BEAN 1. (C

Archytas marmoratus Archytas plangens Archytas Archytas Chetogena Cuphocerini Eucelatoria armigera Eucelatoria australis Eucelatoria bryani Eucelatoria guimaraesi Eucelatoria rubentis ABLE Classification of parasite and parasitoid T Armyworm Symposium 2002: Molina-Ochoa et al. 259 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

all & Berberet 1975 WHICH an Dine 1913 arker et al. 1953; Silveira & Ruffinelli 1956 1953; arker et al. ennah 1947 erán 1974 ernández & Clavijo 1984 erán 1974 ones 1913

Blanchard 1963 Guimarães 1977 1957 Etcheverry Enkerlin 1975 P Enkerlin 1975 Maes 1989 Goncalves & Goncalves 1973; Silva et al. 1997 Silva et al. Goncalves & 1973; Guimarães 1977 1976 Nickle Guimarães 1977 Goncalves & 1973; 1993 Cave F V J Luginbill 1928 T Guimarães 1977 Enkerlin 1975 Alam 1979 1989 Valicente 1987; Silveira et al. Guimaraes 1977 Enkerlin 1975 Gladstone 1991 Maes 1989; F T PARASITES FROM

), 2 CROPS , NW (N/G) (N/G) M (N/G) (N/G) (N/G) (N/G) (N/G) Luginbill 1928 M (N/G) (N/G) (N/G) S M SC SC,M (N/G) (N/G) (N/G) (N/G) M M (N/G) PN M M (N/G) OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST S SP , enezuela enezuela eru CubaBrazil Chile Uruguay (N/G) 1975 Bruner et al. Nicaragua US Honduras Lesser Antilles Puerto Rico US V P Brazil Chile Nicaragua V MITH 1 FAMILY

(J. E. S

L LU LU attacked Host stage INDICATING

REFERENCES Zenillia blanda

Spodoptera frugiperda , (Townsend) . Townsend L Brazil (Townsend) ARMYWORM

ACCOMPANYING (Coquillet) (Reinhard) Syn:

Townsend L Honduras (N/G) 1993 Cave

ALL ) F WITH

(Aldrich) L Argentina (Fabricius) L Brazil COLLECTION

(Townsend) L Brazil M 1997 Silva et al. Guimarães 1983; DeGeer L US (N/G) Luginbill 1928 (Walker) L US PN & Berberet 1975 Wall Reinhard L Cuba (N/G) 1975 Bruner et al. OF BASIN

sp. L Brazil

sp. L Barbados ONTINUED TRIES BEAN sp. L Chile Exorista ceratomiae Chetogena tachinomoides 1. (C

Incamyia chilensis Lespesia affinis Syn: Syn: Hyphantrophaga hyphantriae Hyphantrophaga collina Gonia Euphorocera Exorista mella Gonia capitata Gonia crassicornis Gonia (Reaumuria) pacifica Gonia texensis Eucelatoria Euphorocera floridensis Euphorocera tachinomoides Syn: ABLE Classification of parasite and parasitoid T 260 Florida Entomologist 86(3) September 2003 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

kery 1929 c all & Berberet 1975; Nickle 1976 Nickle all & Berberet 1975; all & Berberet 1975 irla et al. 1999 irla et al. 1999 irla et al. WHICH an Dine 1913 atel & Habib 1984, 1986; Guimarães 1983; Valicente Valicente Guimarães 1983; 1986; atel & Habib 1984, Bruner et al. Ryder & Piedra 1968; 1953; arker et al. ainter 1955 1986 air et al. air et al. 1986 air et al. ennah 1947 1977 erán 1974, ones 1913

V P 1989 Guimarães 1977 1957 Etcheverry P 1975 1983 Malausa 1981, P Canas & O’Neil 1998 1993 Cave F Carrillo 1980 Gladstone 1991; Maes 1989; Huis 1981; Estrada 1960; 1977 Lacayo V J 1984 Soteres et al. 1994 Tingle et al. 1993 1992, Riggin et al. 1986; Gross & Pair W Vi P Butler 1958a Reed 1980 Luginbill 1928 Silveira & Ruffinelli 1956 & Clavijo 1984 Fernández Notz 1972; T V Blanchard 1963 Guimarães 1977 Cave 1993 Cave W P Reed 1980 PARASITES FROM

), 2 CROPS , ,A,M,S M M (N/G) M (N/G) M M,Te M M,S M M M SC SC,M A C M PN M PN, M,S C M,BG,Mi,S (N/G) M (N/G) M (N/G) (N/G) (N/G) PN M,S M,BG,Mi OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST , enezuela Argentina Brazil Chile Cuba Guadeloupe Guatemala Honduras Lesser Antilles Mexico Nicaragua Puerto Rico US Uruguay V Brazil US MITH 1 FAMILY

(J. E. S

L attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , (Wile) . (Scudder) ARMYWORM

ACCOMPANYING

ALL (Riley) ) F WITH

COLLECTION (Williston) L US (N/G) Luginbill 1928

(Riley) L Honduras (Blanchard) L Argentina OF BASIN

ONTINUED TRIES BEAN rontina archippivora F Achaetoneura archippivora Achaetoneura 1. (C

Lespesia frenchi Lespesia grioti Syn: Syn: Lespesia aletiae Lespesia archippivora Syn: ABLE Classification of parasite and parasitoid T Armyworm Symposium 2002: Molina-Ochoa et al. 261 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

ilson 1923 irla et al. 1999 irla et al. irla et al. 1999 irla et al. WHICH alicente 1989 argas & Sanchez 1983 arker et al. 1953; Silveira & Ruffinelli 1956 1953; arker et al. 1986 atel & Habib 1984, 1953 arker et al. Silveira & Ruffinelli 1956 1953; arker et al. arker et al. 1953 arker et al. & Fuxa 1992 Pantoja 1985; antoja et al. 1953 arker et al. erán 1974 erán 1974 erán 1974

V 1993 Cave Maes 1989 Enkerlin 1975 Luginbill 1928 T Soteres et al. 1984 Soteres et al. W Luginbill 1928 Guimarães 1977 P P Guimarães 1977 V P P 1984 Soteres et al. Luginbill 1928 T V P Almeida 1980 & Lucchini Guimarães 1977 Goncalves & 1973; V Ryder & Pulgar 1969 1989 Wheeler et al. Maes 1989 P 1980 Ashley et al. McCutcheon 1991 P Notz 1972 T PARASITES FROM

), 2 CROPS , M (N/G) (N/G) PN (N/G) (N/G) C (N/G) (N/G) (N/G) (N/G) M (N/G) (N/G) A (N/G) (N/G) M (N/G) M (N/G) R (N/G) M M R M M,S (N/G) M (N/G) OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST , enezuela enezuela enezuela Brazil Honduras Nicaragua US V Uruguay Uruguay V Brazil Colombia Cuba Honduras Nicaragua Puerto Rico US Uruguay V MITH 1 FAMILY

(J. E. S

L LUS A LUS attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , . (Macquart) ARMYWORM (Townsend)

ACCOMPANYING

ALL (Macquart) (Townsend) ) F (Townsend) L Venezuela M 1977 Terán WITH (Walker)

(Fallen) L Honduras (N/G) 1993 Cave Wiedeman L Chile (N/G) 1957 Etcheverry COLLECTION

(Walker) (Townsend) L Brazil M sp. L Brazil OF BASIN

sp. L Ecuador M 2000 Benzing et al. ONTINUED sp. L Argentina sp. L Argentina TRIES BEAN Chetogena claripennis Chetogena floridensis Exorista pyste 1. (C

arasetigena atelloa similis atelloa eleteria robusta P P P P Syn: Phorocera floridensis Syn: Phorocera claripennis Lespesia Linnaemya annalis Linnaemya comta Linnaemya Nemorilla pyste Syn: ABLE Classification of parasite and parasitoid T 262 Florida Entomologist 86(3) September 2003 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

alton 1913; Luginbill 1928 alton 1913; all & Berberet 1975 irla et al. 1999 irla et al. irla et al. 1999 irla et al. irla et al. 1999 irla et al. WHICH era et al. 1995; Virla et al. 1999 et al. Virla 1995; era et al. alicente 1989 arker et al. 1953 arker et al. erán 1974 erán 1974

Notz 1972 T Costa-Lima 1949 1957 Etcheverry V 1997 Avalos Molinari & P Guimarães 1971 Guimarães 1977 Butler 1958a Silveira & Ruffinelli 1956 Dew 1913 V Guimarães 1977 Etcheverry 1957 Etcheverry W T Soteres et al. 1984 Soteres et al. 1994 Tingle et al. Riggin 1980; Ashley et al. Hofmaster & Greenwood 1949; 1993 1992, et al. W 1986 et al. Pair Rohlfs & Mack 1985; Enkerlin 1975 Danks 1975 Reed 1980 V V Guimarães 1977 V Silva et al. 1986; & Habib 1984, Patel Escalante 1974; 1997 Guimaraes 1977 PARASITES ,C FROM

), 2 ,SY ,BG CROPS , ,A,M,S ,A,M,S (N/G) (N/G) M M SY (N/G) M (N/G) C M M,S M (N/G) M (N/G) (N/G) C M PN M,S PN,BG C M,BG,Mi,S M M (N/G) M M (N/G) OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST S , enezuela Chile Brazil US Uruguay Brazil US V ChileChile (N/G) (N/G) 1973 Valdivia & Valencia 1973 Valdivia & Valencia Brazil Brazil MITH 1 FAMILY

(J. E. S

LU L L attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , . Fabricius L Chile ARMYWORM

ACCOMPANYING

(Kirkpatrick) ALL (Bigot) L US A (Thompson) L Argentina (Kirkpatrick) Townsend L Venezuela M ) F WITH (Reinhard) L US PN & Berberet 1975 Wall

(Reinhard) L Argentina COLLECTION

OF BASIN (Fallen) L Argentina

sp. L Argentina sp. L Brazil ONTINUED TRIES BEAN Nemorea leucanae 1. (C

inthemia leucanae inthemia mima inthemia quadripustulata inthemia reliqua inthemia roblesi inthemia rufopicta inthemia sinuata inthemia trinitatis inthemia oria ruralis Pronemorilla mima Pseudokea V W Syn: Syn: W W W W W W W W ABLE Classification of parasite and parasitoid T Armyworm Symposium 2002: Molina-Ochoa et al. 263 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

kery 1929; Bianchi 1944; Ashley et al. 1982; Mitch- 1982; Ashley et al. Bianchi 1944; kery 1929; c all & Berberet 1975 WHICH argas & Sánchez 1983 arker et al. 1953 arker et al. 1986 air et al. antoja & Fuxa 1992 ennah 1974 erán 1974 aseen 1979; Hynes 1942 aseen 1979;

Etcheverry 1957; Campos 1965 1957; Etcheverry Enkerlin 1975 V 1993 Cave F 1979 Guevara et al. Sarmiento & Razuri 1978 Y W McCutcheon 1991 T et al. Virla De Santis & Esquivel 1966; De Santis 1967; 1999 P 1997b Cruz et al. 1957 Etcheverry Canas & O’Neil 1998 1989; Wheeler et al. 1993 Cave 2001 Molina-Ochoa et al. P Gladstone 1991 Maes 1989; Huis 1981; Estrada 1960; P Vi 1992, Riggin et al. 1986; Gross & Pair 1984; ell et al. 1993 Rohlfs & Mack 1985 McCutch- Isenhour 1988; 1986; et al. Pair Reed 1980; eon 1991 1983 Ashley et al. 1951 Muesebeck et al. Luginbill 1928; 1951 Muesebeck et al. 1951 Muesebeck et al. 2001 Marsh & Shaw Luginbill 1928; PARASITES FROM

), 2 CROPS , M M,PN R M,S (N/G) M M (N/G) PN M,S (N/G) (N/G) M A,M,C M M,S M M,S M R M S M,S M,BG,PG (N/G) A,M,C A,M,C (N/G) OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST , enezuela eru Colombia Honduras Lesser Antilles Mexico P Trinidad US V NicaraguaBrazil Chile Honduras (N/G)Mexico Nicaragua Andrews 1988 Puerto Rico US Canada US MITH 1 FAMILY

(J. E. S

L L L attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , . ARMYWORM

ACCOMPANYING

(Viereck) ALL (Cresson) (Viereck) ) F WITH (Cresson) (Cresson) E Argentina M

COLLECTION

OF BASIN

sp. L Chile sp. L US M Bianchi 1944 ONTINUED TRIES BEAN Rogas caphimal Rogas laphygmae Rogas terminalis 1. (C

: inthemia erisierola W P Aleiodes caphimal Syn: Alieodes laphygmae Syn: Aleiodes terminalis Syn Agathis stigmatera ABLE Hymenoptera: Bethylidae Hymenoptera: Hymenoptera: Braconidae Hymenoptera: Classification of parasite and parasitoid T 264 Florida Entomologist 86(3) September 2003 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

irla et al. 1999 irla et al. irla et al. 1999 irla et al. WHICH argas & Sánchez 1983 arker et al. 1953 arker et al. 1989; Valicente 1986; 1984, atel & Habib 1982, assoa 1983 arker et al 1953 ennah 1947 aseen 1979

V P Alam 1979 P Silva et al. 1997a,b; Cruz et al. 1995a,b; Rezende et al. 1997 1994 Rezende et al. 1957 Etcheverry 1988 Medina et al. V Myers 1932 Ryder & Pulgar 1969 1975; Bruner et al. Ryder & Piedra 1968; Myers 1932 Canas & O’Neil 1998 1989; Wheeler et al. 1993 Cave 1989; Castro et al. F Alam 1979 Y Cave 1993; Passoa 1983; Wheeler et al. 1989; Canas & 1989; Wheeler et al. 1983; Passoa 1993; Cave O’Neil 1998 Maes 1989 Huis 1981; V Ryder & Piedra 1968 Ryder & Pulgar 1969; P 1977 Lacayo Maes 1989; Estrada 1960; 1984 Soteres et al. P 1993 Cave Alam 1979 Ryder & Pulgar 1969 Canas & O’Neil 1998 1993; Cave 2001 Molina-Ochoa et al. Huis 1981 PARASITES FROM

), 2 CROPS , M (N/G) (N/G) M M,C A,M,C M R BG M (N/G) BG M M,S M (N/G) M M M M M M M A (N/G) (N/G) M M M M,S M OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST , Argentina Barbados Brazil Chile Colombia Cuba Haiti Honduras Lesser Antilles Trinidad Honduras Nicaragua Argentina Cuba Honduras Nicaragua US Honduras Nicaragua Honduras Mexico Nicaragua MITH 1 FAMILY

(J. E. S

E L L E attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , . (Cresson) ARMYWORM

ACCOMPANYING

(Cresson) ALL (Viereck) L US C 1994 Tingle et a. (Sonan) E Barbados Muesebeck (Marshall) E Barbados Wilkinson L Mexico (N/G) 1980 Pena Moya 1980; ) F WITH (Cresson)

(Muesebeck) COLLECTION Cresson

OF BASIN

sp. ONTINUED sp. sp. L Argentina TRIES BEAN Chelonus texanus Rogas vaughani Rogas cautus Microchelonus 1. (C

Chelonus insularis Syn: Aleiodes vaughani Syn: Aleiodes Syn: Bassus Bracon kirkpatricki nigriceps Cardiochiles Chelonus antillarum Chelonus cautus Syn: Chelonus formosanus ABLE Classification of parasite and parasitoid T Armyworm Symposium 2002: Molina-Ochoa et al. 265 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

all & Berberet 1975 Pair Rohlfs & Mack 1985; Whitcomb 1982; addill & WHICH alicente 1989 air et al. 1986 air et al. antoja & Fuxa 1992 Silveira & Ruffinelli 1956 1953; arker et al. ennah 1947 aseen 1979 1981 aseen et al.

Bahena & García 1991; Nava & Castro 1991; Arce & & Castro 1991; Nava Bahena & García 1991; 2001 Molina-Ochoa et al. Garcia 1995; P Maes 1989; Ashley 1986; Huis 1981; Estrada 1960; Gladstone 1991 1977 Lacayo P Y Y 1984 Soteres et al. 1994 Tingle et al. Bianchi 1944; 1929; Vickery 1912; Pierce & Holloway Ash- 1982; 1980, Ashley et al. 1951; Muesebeck et al. 1986; Gross & Pair 1984; et al. Mitchell 1986; ley 1983, 1992 Riggin et al. W 1983 Ashley et al. Enkerlin 1975 W McCutcheon 1991 1986; et al. Butler 1958b Luginbill 1928 P & Clavijo 1984 Fernández Notz 1972; V Cortez & Coronado & Ruíz 1991; Loya-Ramírez 1978; Carrillo 1980 1998; et al. Sánchez-García Trujillo 1994; 2001 Molina-Ochoa et al. Herrera-Aranguena 1998 Cave 1993 Cave Maes 1989 1999 et al. Virla 1989; Wheeler et al. 1984, & Habib 1982, Patel Almeida 1980; & Lucchini 1997b Cruz et al. 1986; 1957 Etcheverry 1993 Cave F PARASITES FROM

), 2 CROPS , ,A,M,S M S M, M M,S R M (N/G) A C M PN M,BG M,PN M,S C (N/G) (N/G) M M M M,S C (N/G) M M M A,M,C M M,S OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST , enezuela eru Mexico Nicaragua Puerto Rico Trinidad US Uruguay V Mexico P Honduras Nicaragua BarbadosArgentina Brazil Chile Honduras MLesser Antilles Alam 1979 MITH 1 FAMILY

3 3 3 (J. E. S

E,L E attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , (Cresson) (Linnaeus) E,L (Say) E,L . (Viereck) (Ashmead) ARMYWORM (Ashmead)

ACCOMPANYING

(Cresson) ALL ) F WITH (Cresson)

COLLECTION

OF BASIN

ONTINUED sp. E Brazil TRIES BEAN Protapanteles harnedi Chelonus texanus Apanteles laphygmae Apanteles grenadensis 1. (C

Syn: Syn: Syn: Chelonus Cotesia (Apanteles) congregata Cotesia (Apanteles) glomeratus Cotesia (Apanteles) marginiventris Syn: Chelonus insularis Syn: ABLE Classification of parasite and parasitoid T 266 Florida Entomologist 86(3) September 2003 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

all & Berberet 1975; Nickle 1976 Nickle all & Berberet 1975; WHICH an Dither 1960 argas & Sánchez 1983 air et al. 1986; Molina-Ochoa et al. 2001 Molina-Ochoa et al. 1986; air et al. antoja & Fuxa 1992 Silveira & Ruffinelli 1956 1953; arker et al. 1997 Silva et al. 1986; atel & Habib 1984, erán 1980 aseen 1979* aseen 1979

Marsh 1978 P Gladstone 1991 Maes 1989; Huis 1981; Estrada 1960; P Segeren & Sharma 1979 V 1984 Soteres et al. 1994 Tingle et al. Bianchi 1944;Hofmas- 1929; Vickery Muesebeck 1921; Marsh 1951; Muesebeck et al. ter & Greenwood 1949; Ash- 1982; Hogg et al. 1982; 1980, Ashley et al. 1978; 1986; Gross & Pair 1984; et al. Mitchell 1986; ley 1983, Ru- 1994; Hamm et al. 1994; 1993, 1992, Riggin et al. Whitfield 1996 berson & McCutcheon 1991 1986; et al. Pair Rohlfs & Mack 1985; W 1983 Ashley et al. Reed 1980 Luginbill 1928 P & Clavijo 1984 Fernández Notz 1972; T Y 1985*** Rajapakse et al. 1983**; McCutcheon et al. Alam 1979 P V 1988 Medina et al. Sinha 1982 Maes 1989 1977 Lacayo Herrera-Aranguena 1998 Y Hofmaster & Greenwood 1949 1993 Cave Rohlfs & Mack 1985 1951 Muesebeck et al. PARASITES FROM

), 2 CROPS , M M,S M R M,PN (N/G) A C M M,S PN M,BG M,S,C,Mi (N/G) (N/G) M SY,To M (N/G) M M R M M M M,S C M M M S (N/G) OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST , enezuela eru Mexico Nicaragua Puerto Rico Suriname US Uruguay V Tobago Trinidad & US Barbados Brazil Colombia Guyana Nicaragua P Trinidad US US MITH 1 FAMILY

3 3 3 (J. E. S

E,L E,L attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , (Cresson) . (Haliday) (Viereck) (Ashmead) ARMYWORM (Ashmead)

ACCOMPANYING

(Ashmead) (Walsh) L Honduras ALL sp. E,L ) F WITH

COLLECTION

OF BASIN

sp. L Honduras M 1993 Cave ONTINUED TRIES BEAN Protapanteles harnedi Microplitis manilee Apanteles laphygmae Apanteles grenadensis 1. (C

Syn: Syn: Syn: Syn: Syn: Cotesia (Apanteles) ruficrus Cotesia (Apanteles) Distatrix Glyptapanteles militaris Glyptapanteles Cotesia (Apanteles) marginiventris Syn: ABLE Classification of parasite and parasitoid T Armyworm Symposium 2002: Molina-Ochoa et al. 267 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

kery 1929; Wall & Berberet 1975; Riggin et al. Riggin et al. & Berberet 1975; Wall kery 1929; Bianchi 1944 kery 1929; c c WHICH argas & Sánchez 1983 an Dither 1960 air et al. 1986; McCutcheon 1991 1986; air et al. 1986 air et al. 1993 1992, Riggin et al. air 1986; McCutcheon 1991 1986; air et al.

Cave 1993 Cave Maes 1989 Huis 1981; 1984 Soteres et al. Vi 1993 1992, P Reed 1980 Luginbill 1928 Alam 1979 1997 Silva et al. 1993 Cave Maes 1989 P 1984 Soteres et al. 1994 Tingle et al. Gross & 1984; et al. Mitchell 1982; 1980, Ashley et al. P 1976 Nickle Rohlfs & Mack 1985 P Reed 1980 1983 Ashley et al. 1951 Muesebeck et al. Luginbill 1928; 1957 Etcheverry 1988 Medina et al. V 1988 Ortegón et al. 1993 Cave 2001 Molina-Ochoa et al. 1995 Ciceros et al. Gladstone 1991 Segeren & Sharma 1979 V Vi Enkerlin 1975 Luginbill 1928 PARASITES FROM

), 2 CROPS , M,S M A M M,S M,S,C (N/G) (N/G) M (N/G) M M,S A C M PN S M,S M,BG,Mi M,BG,PG (N/G) M M R C,S M,S M S (N/G) M,PN (N/G) M M,BG (N/G) OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST , Honduras Nicaragua US Brazil Nicaragua US Colombia Honduras Mexico Nicaragua Suriname US MITH 1 FAMILY

(J. E. S

L attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , . ARMYWORM

ACCOMPANYING

ALL (Muesebeck) L Mexico (Say) (Muesebeck) L Honduras (Viereck) L Chile ) F WITH

COLLECTION (Cresson)

sp. L Honduras (N/G) 1993 Cave sp. L Barbados OF BASIN

ONTINUED TRIES BEAN Zele mellea 1. (C

Gnathopleura Homolobus truncator Syn: Macrocentrus Meteorus arizonensis Meteorus autographae Meteorus laphygmae ABLE Classification of parasite and parasitoid T 268 Florida Entomologist 86(3) September 2003 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

ilson 1923 ilson 1923 ilson 1923 irla et al. 1999 irla et al. WHICH antoja & Fuxa 1992 eraza-Lizarraga 1982; Coronado & Ruíz 1991 eraza-Lizarraga 1982; 1953 arker et al. ernández & Terán 1990a,b Terán ernández &

W 1951 Muesebeck et al. Luginbill 1928; Maes 1989 Luginbill 1928 De Santis & Fidalgo De Santis 1989; De Santis 1979; 1999 et al. Virla 1994; P 1989 Wheeler et al. Huis 1981 F P 2001 Molina-Ochoa et al. Herrera-Aranguena 1998 1993 Cave Maes 1989 V P Ashley 1979 W Luginbill 1928 1993 Cave Maes 1989 W Luginbill 1928 1993 1992, Riggin et al. 1993 Cave Maes 1989 PARASITES FROM

), 2 CROPS , (N/G) M (N/G) M R M (N/G) M S M, C (N/G) M M (N/G) M (N/G) (N/G) (N/G) C (N/G) (N/G) M OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST SM , eru US Puerto Rico Nicaragua P Nicaragua US Nicaragua US Honduras Nicaragua MITH 1 FAMILY

(J. E. S

LUS C LU L attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , (Cresson) (Ashmead) (Fabricius) L Honduras (Burks) L US M Hofmaster & Greenwood 1949 (Ashmead) . (Cameron) ARMYWORM

(Ashmead) ACCOMPANYING

Howard L Nicaragua ALL (Cresson) P US C Howard L Honduras (Viereck) L Venezuela M & Clavijo 1984; Fernández 1980; Terán Notz 1972; Walker L Argentina (Say) L Panama (N/G) Andrews 1988 ) F WITH (Say) P Argentina

(Cresson) L US (N/G) 1951 Muesebeck et al. Luginbill 1928; (Kirby) COLLECTION

OF BASIN

sp. L Uruguay (N/G) Silveira & Ruffinelli 1956 1953; et al. Parker sp. L Honduras ONTINUED sp. L Mexico sp. L Trinidad M 1979 Yaseen TRIES BEAN Splilochalcis hirfitemora Splilochalcis Conura fulvomaculata vittata Splilochalcis 1. (C

alinzele Euplectrus comstockii Euplectrus comstockii Euplectrus furnicus Euplectrus hircinus Euplectrus insularis Meteorus laphygmae Meteorus vulgaris Meteorus Microplitis ovata Brachymeria robusta Brachymeria femorata Conura (Spilochalcis) Conura (Ceratosmicra) hirtifemora Syn: Conura (Ceratosmicra) immaculata Syn: Conura (Ceratosmicra) meteori Conura igneoides Syn: P Stantonia ABLE Hymenoptera: Eulophidae Hymenoptera: Hymenoptera: Chalcididae Hymenoptera: Classification of parasite and parasitoid T Armyworm Symposium 2002: Molina-Ochoa et al. 269 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

kery 1929; Muesebeck et al. 1951; Ashley et al. Ashley et al. 1951; Muesebeck et al. kery 1929; c all & Berberet 1974, 1975 all & Berberet 1974, eller 1980 irla et al. 1999 irla et al. WHICH argas & Sánchez 1983 antoja & Fuxa 1992 1986 air et al. 1986 air et al. Blanchard 1963 1953; arker et al. ennah 1947 aseen 1979

Alam 1979 Costa-Lima 1962 1957 Etcheverry V Myers 1932 1975 Bruner et al. Ryder & Piedra 1968; Sinha 1982 F Gutiérrez- Montoya-Burgos 1980; 1979; Guevara et al. 2001 Molina-Ochoa et al. Rodriguez 1982; Ryder & Pulgar 1969 P Y Vi 1993 1992, Riggin et al. 1982; Hogg et al. 1982; 1980, W Luginbill 1928 Marín-Acosta 1966 Guagliami 1962 Almeida 1980 & Lucchini Myers 1932 1993 Cave Trujillo 1994 Cortez & 1980; Montoya-Burgos 1979, P Huis 1981 1977 Lacayo Reed 1980 K P Enkerlin 1975 V P Cortes 1980 PARASITES FROM

), 2 CROPS , M (N/G) M M Pm&Pb (N/G) M M M M R (N/G) M PN (N/G) M S,C M BG M M M,S M M,S BG M M,S (N/G) M (N/G) (N/G) OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST , enezuela Brazil Chile Colombia Cuba Guyana Lesser Antilles Mexico Nicaragua Puerto Rico Trinidad US V Cuba Honduras Mexico Nicaragua US BrazilBrazil (N/G) Guimaraes 1977 MITH 1 FAMILY

(J. E. S

L attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , . ARMYWORM

ACCOMPANYING

Ashmead L Nicaragua M Maes 1989 Howard L Barbados ALL (Ferriere) P Barbados M Alam 1979 (Bigot) L Argentina ) F WITH

(Brethes) L Brazil M De Santis 1980 COLLECTION

OF BASIN

sp. L Argentina (N/G) 1953 et al. Parker sp. L Brazil ONTINUED TRIES BEAN 1. (C

richodischia caerulea richodischia soror richodischia pupivora richospilus richospilus Euplectrus ronnai Euplectrus T T T T Euplectrus marginatus Euplectrus plathypenae ABLE Classification of parasite and parasitoid T 270 Florida Entomologist 86(3) September 2003 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

all & Berberet 1975 irla et al. 1999 irla et al. WHICH era et al. 1995 era et al. air et al 1986 Isenhour 1988 1986; air et al. atel & Habib 1982, 1984, 1986; Cruz et al. 1997a,b Cruz et al. 1986; 1984, atel & Habib 1982, 1981 et al. Yaseen 1953; arker et al. 1986 air et al.

Cruz et al. 1997b Cruz et al. 1989 et al. Machuca Canas & O’Neil 1998 1993 Cave P 1984 Soteres et al. 1993 1992, Riggin et al. 1986; Gross & Pair P V & Bar- Valicente 1987; Silveira et al. 1997; Silva et al. reto 1999 1999 Cruz et al. Maes 1989 McCutcheon 1991 Costa-Lima 1949 1957 Etcheverry P 1993 Cave 2001 Molina-Ochoa et al. Maes 1989 Huis 1981; 1984 Soteres et al. 1982 Hogg et al. W P V Cruz et 1987; Silveira et al. Almeida 1980; & Lucchini 1997b al. Ashley 1983 Morey 1971 1951 Muesebeck et al. P Luginbill 1928 PARASITES FROM

), 2 CROPS , M M M M,S M,S A M M,S M M (N/G) M M,S (N/G) M M (N/G) M,S M A M PN (N/G) M M M M M,S (N/G) OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST , Chile Honduras Mexico US Brazil Nicaragua US Chile Honduras Mexico Nicaragua US Uruguay Brazil US Uruguay MITH 1 FAMILY

(J. E. S

LUS M attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , . ARMYWORM

ACCOMPANYING

(Blanchard) L Peru M 1993 Ayqui-Vilca ALL (Ashmead) L Brazil (Cresson) L Brazil (Say) L US M,S 1986 et al. Pair (Cresson) (Vierech) L Barbados M Alam 1979 ) F WITH

(Cresson) (Blanchard) L Argentina COLLECTION

OF sp. L Argentina (N/G) 1953 et al. Parker BASIN

sp. L Brazil sp. L Argentina ONTINUED TRIES BEAN Sagaritis oxylus 1. (C

Campoletis Amblyteles Amblyteles Ancyloneura Campoletis chloridae Campoletis curvicauda Campoletis flavicincta Campoletis grioti Campoletis oxylus Syn: Campoletis sonorensis Anomalon ejuncidum ABLE Hymenoptera: Ichneumonidae Hymenoptera: Classification of parasite and parasitoid T Armyworm Symposium 2002: Molina-Ochoa et al. 271 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

ilson 1923 yala-Sifontes et al. 1978 yala-Sifontes et al. WHICH arker et al. 1953; Silveira & Ruffinelli 1956 1953; arker et al. orter 1998; Virla et al. 1999 et al. Virla orter 1998; 1989 air & Gross 1984, 1986 air et al. Sil- 1989; Valicente 1986; 1984, atel & Habib 1982, 1986 air et al. ernández & Clavijo 1984 aseen et al. 1981 aseen et al. aseen et al. 1981 aseen et al.

Cruz et al. 1997a; Silva et al. 1997 Silva et al. 1997a; Cruz et al. Trujillo 1994 Cortez & Hynes 1942 Y F W Luginbill 1928 1951 Muesebeck et al. 1975 Bruner et al. 1993 Cave Maes 1989 Dew 1913 Luginbill 1928 1993 Cave P P 1997 Silva et al. A 1991 Flores-Dávila et al. 1993 Cave P P Ashley 1979 Y P 1997b Cruz et al. 1987; veira et al. 1988 Medina et al. Canas & O’Neil 1998 1989; Wheeler et al. 1993 Cave P Huis 1981 Ashley 1983 1995 1994a,b, 1992, & Garcia-R. Giraldo-Vanegas PARASITES FROM

), 2 CROPS , M M M (N/G) M (N/G) M M (N/G) M C,M (N/G) M (N/G) M M M M M M M,S M (N/G) M M M M,S M,S M M (N/G) OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST , enezuela enezuela Mexico Trinidad V Argentina Cuba Honduras Nicaragua US Uruguay Brazil Cuba Mexico US Brazil Colombia Honduras Mexico Nicaragua US V MITH 1 FAMILY

(J. E. S

LUS C L attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , . ARMYWORM (Say)

(Cresson) ACCOMPANYING

(Cresson) P Honduras (Cresson) L Venezuela M Notz 1972 (Gravenhorst) ALL (Cresson) L Bolivia ) F WITH

(Fabricius) (Cresson) P US M & Gross 1989 Pair COLLECTION

OF BASIN

sp. L Brazil M 1997 Silva et al. sp. L Brazil sp. L Argentina sp. L Brazil M 1997 Silva et al. sp. L Honduras ONTINUED TRIES BEAN Enicospilus concolor Enicospilus purgatus 1. (C

Enicospilus flavus Syn: Enicospilus merdarius Syn: Goryphina Hyposoter Campoplex Cryptus albitarsis Diadegma Diapetimorpha introita Eiphosoma annulatum Eiphosoma vitticole Eiphosoma ABLE Classification of parasite and parasitoid T 272 Florida Entomologist 86(3) September 2003 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

kery 1929; Bianchi 1944 kery 1929; kery 1929 c c all & Berberet 1975 irla et al. 1999 irla et al. WHICH atel & Habib 1984, 1986 atel & Habib 1984, 2001 Molina-Ochoa et al. 1986; air et al. 1986 air et al. 1953 arker et al. assoa 1983

1993 1989 Wheeler et al. 1993 Cave Escalante 1974 1984 Soteres et al. P Canas & O’Neil 1998 1989; Wheeler et al. 1993 Cave Carrillo 1980 P Maes 1989; Gladstone 1991; Huis 1981; Estrada 1960; 1977 Lacayo 1984 Soteres et al. 1994 Tingle et al. Vi W P Luginbill 1928 1999 et al. Virla De Santis 1967; P Silveira & Ruffinelli 1956 1957 Etcheverry Vi Enkerlin 1975 Luginbill 1928 V Sil- 1986; 1984, & Habib 1982, Patel Goncalves 1973; 1987 veira et al. Canas & O’Neil 1998 1989; Wheeler et al. 1993 Cave P PARASITES FROM

), 2 CROPS , S M A M M M,S M M,S M A C M PN M,S (N/G) M (N/G) (N/G) M M PN (N/G) M M M M,S (N/G) OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST , US Brazil Honduras Mexico Nicaragua US Uruguay US Brazil Honduras MITH 1 FAMILY

(J. E. S

L attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , (Viereck) . (Morley) L Uruguay (N/G) Silveira & Ruffinelli 1956 1953; et al. Parker ARMYWORM

(Gupta) L Honduras M Fabricius P US M New record (JEC) ACCOMPANYING (Cresson) P US M New record (JEC)

(Say) L US M 1992, Riggin et al. Hofmaster & Greenwood 1949; ALL (Fabricius) (Wichsee) L Argentina ) F WITH

(Howard) L US M 1993 1992, Riggin et al. (Say) L Chile COLLECTION (Brulle) L Argentina

OF BASIN

(Cushman) L US C 1994 Tingle et al. ONTINUED TRIES BEAN sp. L Peru Neopristomerus appalachianus 1. (C

arania (Atrometus) tricolor Ichneumon ambulatorius Ichneumon promissorius Ichneumon anticarsiae Microcharops Netelia sayi Netelia P Pristomerus spinator Syn: Ophion ancyloneura Ophion bilineatus Ophion flavidus Isdromas lycaenae disceitergus Mesochorus ABLE Classification of parasite and parasitoid T Armyworm Symposium 2002: Molina-Ochoa et al. 273 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

kery 1929 c WHICH era et al. 1995 era et al. air et al. 1986 air et al. air et al. 1986 air et al. 1953 arker et al.

Canas & O’Neil 1998 1993 Cave Gladstone 1991 1989 Wheeler et al. Maes 1989 1983 1982, 1980, Ashley et al. 1993 Cave Notz 1972 Mitchell et al. 1984; Gross & Pair 1986 Gross & Pair 1984; et al. Mitchell P Molina-Ochoa et al. 2001 Molina-Ochoa et al. P Gladstone 1991 Maes 1989; Riggin et al. 1991; 1986, Gross & Pair 1982; Hogg et al. 1993 1992, 1986 et al. Pair Rohlfs & Mack 1985; Reed 1980 Maes 1989 Luginbill 1928 V 1997 Silva et al. Ashley 1986 Maes 1989 Huis 1981; Escalante 1974 1994 Tingle et al. 1984 et al. Mitchell 1976 Nickle 1983 Ashley et al. P Vi Luginbill 1928 PARASITES FROM

), 2 CROPS , M M,S M M M M M M M,S M M,S M M M,S M,BG,Mi,S M (N/G) M M M M M C M PN M,PG (N/G) (N/G) OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST , enezuela eru Nicaragua Nicaragua US Honduras V Nicargua US Nicaragua US Brazil Mexico Nicaragua P US Uruguay MITH 1 FAMILY

(J. E. S

L L attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , . (Say) P US M & Gross 1989 Pair ARMYWORM (Gravenhorst)

ACCOMPANYING

(Cushman) L Honduras ALL (Viereck) L Peru M Escalante 1974 (Say) (Dasch.) L US M (Cresson) L US M ) F WITH

(Gravenhorst) COLLECTION (Brulle) L Mexico

OF BASIN

sp. L Honduras ONTINUED sp. L Argentina TRIES BEAN Enicospilus purgatus 1. (C

rachysphyrus cleonis rachysphyrus ulgichneumon brevicintor ulgichneumon erilampus hyalinus emelucha emelucha emelucha grapholithae emelucha emelucha difficilis emelucha (Reported as hiperparasitoid) P T V T T Ophion flavidus Ophion merdarius Syn: Ophion Sagaritis dubitatus T ABLE Hymenoptera: Perilampidae Hymenoptera: Classification of parasite and parasitoid T 274 Florida Entomologist 86(3) September 2003 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

ojcik et al. 1976 ojcik et al. 1976 ojcik et al. addill 1977; Waddill & Whitcomb 1982 & Waddill addill 1977; WHICH argas & Sánchez 1983 aseen et al. 1981 aseen et al. 1981 aseen et al. aseen 1979

Irving 1978 Alam 1979 1999 et al. Correa-Figueiredo Alvarez & Roa 1995 Y Y Sinha 1982 Acosta 1999 & Cave Maes 1989 W Segeren & Sharma 1979 Y W W 1996a,b; Hernandez & Díaz 1995, 1989; Hernández et al. 1998a,b Ferrer Gonzalez-Narvaez & Zocco 1996; 1999 Cruz et al. Alvarez & Roa 1995 V Ayala-Sifontes 1987; Ayala-Sifontes Armas-García & 1992 et al. Malausa 1983 Canseco- Morales-Pérez 1982; Montoya-Burgos 1979; 1989 Barilla-Vera García-Lagunas 1988; Román 1988; Alam 1979 Maes 1989 Maes 1989 1951 Muesebeck et al. Luginbill 1928 Hofmaster & Greenwood 1949; Riggin et al. 1992, 1993 1992, Riggin et al. Hofmaster & Greenwood 1949; PARASITES FROM

), 2 CROPS , M M M M (N/G) (N/G) M,S M M (N/G) M M M,S (N/G) M M M R M M M M M M M (N/G) OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST SM , enezuela Barbados Brazil Colombia Dominican Republic Guadeloupe Guyana Honduras Nicaragua Puerto Rico Suriname Trinidad US V Colombia Cuba Guadeloupe Mexico Nicaragua US MITH 1 FAMILY

(J. E. S

LU attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , . (Walsh) ARMYWORM

(Zucchi) E Chile M 1999 Pratissoli et al. Perkins E Barbados ACCOMPANYING Riley E Nicaragua

(Girault) L US M Hofmaster & Greenwood 1949 ALL ) F WITH

(Nixon) E Antigua COLLECTION

OF BASIN

sp. E Brazil ONTINUED TRIES BEAN Eupteromalus viridescens 1. (C

richogramma fasciatum richogramma richogramma demoraesi richogramma minutum richogramma richomalopsis viridescens richomalopsis elenomus elenomus remus T T T T T Syn: Catalaccus aeneoviridis T ABLE Hymenoptera: Trichogrammatidae Hymenoptera: Hymenoptera: Pteromalidae Hymenoptera: Scelionidae Hymenoptera: Classification of parasite and parasitoid T Armyworm Symposium 2002: Molina-Ochoa et al. 275 ARIB- COUN-

C AND

AND

MERICAS COLLECTED A

THE

WERE

IN

Reference ARMYWORM

PARASITOIDS

FALL

AND

addill & Whitcomb 1982 addill & irla et al. 1999 irla et al. WHICH argas & Sánchez 1983 aseen et al. 1981 aseen et al.

V 1994 De Sa & Parra 1999 Cruz et al. V Ayala-Sifontes 1987; Ayala-Sifontes Armas-García & 1988 et al. Malausa 1983 Y 1980; Montoya-Burgos 1979, Loya-Ramírez 1978; Wong- Rodríguez-Luna 1982; 1979; Guevara et al. & Carrillo-Sánchez Johannes-Toonders Arevalo 1982; 1987 Bahena & García 1991 1990 Mulock et al. Maes 1989; Huis 1981; W Simmons & Rogers 1990b Simmons & Rogers 1990b 1991 Simmons et al. Simmons & Rogers 1990b; Simmons & Rogers 1990b Simmons & Rogers 1990b Simmons & Rogers 1990b Simmons & Rogers 1990b Simmons & Rogers 1990b Rogers et 1990; Marti et al. Remillet & Silvain 1988; Marti Silvain & Remillet 1993; 1993; 1991, 1990a, al. 2000 et al. 1991 Simmons et al. Simmons & Rogers 1990b; 2000 Marti et al. Simmons & Rogers 1990b 1990 Marti et al. Simmons & Rogers 1990b; Simmons & Rogers 1990b 1991 Simmons et al. Simmons & Rogers 1990b; 1991 Simmons et al. Simmons & Rogers 1990b; De Sa & Parra 1994; Zucchi et al. 1991 et al. Zucchi 1994; De Sa & Parra Huis 1981 PARASITES FROM

), 2 CROPS , M M (N/G) R M M (N/G) M (N/G) M M,S (N/G) (N/G) (N/G) (N/G) (N/G) (N/G) (N/G) (N/G) (N/G) (N/G) (N/G) (N/G) (N/G) (N/G) (N/G) (N/G) M M OCTUIDAE : N ATTACKED

STAGE

EPIDOPTERA collection Crop Country of ) (L HOST , rench Guiana rench anama Brazil Colombia Cuba Guadeloupe Mexico Nicaragua US Belize Bermuda Islands Cayman Colombia Costa Rica Dominica El Salvador F Grenada Guadeloupe Honduras Martinique Mexico P Puerto Rico Nicaragua MITH 1 FAMILY

(J. E. S

attacked Host stage INDICATING

REFERENCES

Spodoptera frugiperda , . ARMYWORM Remillet & Silvain A Bahamas

(Riley) E Brazil ACCOMPANYING

ALL ) F WITH

COLLECTION

sp. E Argentina OF BASIN

ONTINUED TRIES BEAN 1. (C

richogramma richogramma pretiosum richogramma Noctuidonema guyanense T T ABLE Nematoda: Acugutturidae Nematoda: Classification of parasite and parasitoid T 276 Florida Entomologist 86(3) September 2003 L.),

ARIB- COUN-

C AND

AND

L.), O = Onion

Arachis hypogaea MERICAS Lycopersicum esculentum COLLECTED A

THE

WERE

IN

Panicum miliaceum Reference L.), PN = Peanut (

L.), T = Tomato ( ARMYWORM

PARASITOIDS

FALL L.), Mi = Millet (

AND

P. maxiumum Glycine max WHICH

Simmons & Rogers 1990b Simmons & Rogers 1990b Rogers et Simmons & Rogers 1990b; 1990; Marti et al. Sim- 1991; Simmons et al. 1996; 1993, 1991, 1990b, al. 2000; Marti et al. 1994; 1991, mons & Rogers,1990a,b, 1996 Rogers & Marti 1994, Simmons & Marti 1992 Marti & Rogers 2000 1993a; Rogers & Marti 1992a,b, Zea mays L.), Pm = (

PARASITES FROM

), 2 CROPS L.), SY = Soybean ( , (N/G) (N/G) (N/G) M S M, OCTUIDAE spp.), M = Maize (

: N Brachiarie mutica ATTACKED

Gossypium STAGE

EPIDOPTERA collection Crop Saccharum officinarum Country of ) (L HOST irgin Island (N/G) Simmons & Rogers 1990b , Trinidad US V L.), PG = Paragrass (

L.), Ve = Vegetable Crops, (N/G) Not given. MITH L.), C = Cotton ( 1

FAMILY

(J. E. S

attacked Host stage INDICATING Nicotiana tabacum

Panicum barbinode Cynodon dactylon L. (Moench.)), SC = Sugarcane ( L.), Pb = (

REFERENCES

Spodoptera frugiperda , Sorghum bicolor Schard), To = Tobacco ( . L.), BG = Bermudagrass ( Capsicum annuum

ARMYWORM Remillet & Silvain A Suriname

ACCOMPANYING

ALL ) F WITH

L.), S = Sorghum ( COLLECTION

Euchlaena mexicana OF BASIN Medicago sativa

L.), P = Hot pepper ( ONTINUED

TRIES BEAN Oryza sativa 1. (C

Reported attacking eggs and larva (no confirmation). 3 Host stage attacked: E (Egg), L (Larva), P (Pupa), A (Adult). A = Alfalfa ( *Imported from Pakistan, ** Imported from Australia, *** Imported from Thailand & Imported from India. *** Imported from Australia, ** Imported from *Imported from Pakistan, 1 2 Noctuidonema guyanense ABLE Allium cepa Classification of parasite and parasitoid ( R = Rice ( L.), Te = Teosinte ( T Armyworm Symposium 2002: Molina-Ochoa et al. 277

TABLE 2. NUMBER OF SPECIES IN EACH TAXON REPORTED FROM DIFFERENT REGIONS.

Region

North America Central America Order Family (Mexico-US) & Caribbean Basin South America

Diptera Bombyliidae 1 Phoridae 1 Sarcophagidae 3 3 1 Tachinidae 29 24 38 Hymenoptera Bethylidae 1 Braconidae 18 22 11 Chalcididae 6 2 1 Eulophidae 3 8 7 Ichneumonidae 27 18 21 Perilampidae 1 1 Pteromalidae 2 Scelionidae 2 2 2 Trichogrammatidae 2 4 3 Nematoda Aphelenchoididae 1 1 1

RESULTS AND DISCUSSION Parasitoid species that attack FAW eggs were more prevalent in Nicaragua (11) Brazil (8), Bar- A great diversity of parasitoids and parasites bados (7) and Mexico (6). Parasitoid species that of FAW has been reported occurring in the Amer- attack FAW pupae were reported from only 4 icas and the Caribbean basin. In this inventory, countries, with the greatest number of these spe- approximately 150 species of parasitoids and par- cies collected from the US (6). asites have been recorded from 14 families, nine Parasitized FAW were collected from 19 differ- in Hymenoptera, four in Diptera and one in Nem- ent host plants (Table 4). FAW collected from corn atoda (Table 1). Ichneumonids and braconids were parasitized by species from all families ex- were the most diverse families in Hymenoptera, cept Bombyliidae. The number of parasite and represented by 36 and 28 species, respectively. parasitoid species attacking FAW was greatest The most diverse family in Diptera, as well as for FAW collected from corn (134), sorghum (40), overall, was Tachinidae with 55 species. cotton (28), peanut (24), alfalfa (17), rice (13), and Distribution among the taxa was highly vari- Bermuda grass (11). Six of the 14 parasitoids and able. Three of the 14 families were only reported parasite families were recovered from FAW that from one country, and five of the 14 families were had been collected from a single host plant spe- reported from two or fewer countries. Bethylid and cies. Ten of the 14 parasitoids and parasite fami- bombyliid parasitoids were only reported in the lies were recovered from FAW that had been United States (US) during the 1920s and 1940s collected from three or fewer host plant species. (Table 1). Of the ≈150 species, 74 were reported The most diverse parasitoid families also were re- from only one country, and 102 were reported from covered from more FAW host plant species. Ta- only one geographical region. In contrast, nine of chinids, braconids, eulophids, and ichneumonids the 14 families and 18 of the ≈150 species were re- were recovered from FAW collected from 15, 11, 8 ported from all geographical regions (North Amer- and 8 FAW host plant species, respectively. ica, Central America and the Caribbean, and A considerable number of parasitoid species South America), and eight species were reported (≈33%) were reported only from one geographical from 10 or more countries (Tables 1 and 2). region (Table 1). These findings emphasize the The number of parasitoid and parasite species need for more surveys and taxonomical studies of reported from each FAW stage attacked in each the natural enemies in the different habitats of country is summarized in Table 3. The highest the natural distribution of the pest (Molina-Ochoa number of parasitoid species attacking FAW lar- et al. 2001). It is important to consider the differ- vae was reported from the US (75), followed by ent developmental stages of the pest and the Brazil (45), Honduras (43) and Nicaragua (42). growing season of the crop to determine if differ- 278 Florida Entomologist 86(3) September 2003

TABLE 3. NUMBER OF SPECIES OF PARASITOIDS AND STAGE HOST ATTACKED REPORTED FROM DIFFERENT COUNTRIES.

Number of parsitoid species

Country Egg Larval Pupae Adult

Antigua 1 Argentina 4 28 1 Bahamas 1 Barbados 7 9 1 Belize 1 Bermuda 1 Bolivia 1 Brazil 8 45 Cayman Is. 1 Canada 1 Chile 3 19 Colombia 5 6 1 Costa Rica Cuba 3 11 Dominica 1 Dominican Republic 1 Ecuador 2 El Salvador 1 French Guinea 1 Grenada 1 Guadeloupe 3 2 1 Guatemala 1 Guyana 2 2 Haiti 1 Honduras 5 43 1 1 Lesser Antilles 2 8 Martinique 1 Mexico 6 22 Nicaragua 11 42 Panama 1 Peru 2 10 Puerto Rico 3 9 1 Suriname 2 4 1 Tobago 1 1 Trinidad 4 9 1 United States 3 75 7 1 Uruguay 1 16 Venezuela 2 29 Virgin Is. 1

ences in natural distribution of the parasitoids ex- ity on egg and larval populations. An understand- ist (Ashley 1979; Molina-Ochoa et al. 2001) or if ing of induced parasitization of the complex of the records related to unique occurrence are due parasitoids is needed to determine the species to inadequate surveys. It also is important to de- with higher capability to attack each stage of the termine if the occurrence of FAW parasitoids is as- FAW. It is necessary to unite institutional efforts sociated with the developmental stage of the pest to establish programs of release of FAW parasi- and the host plant. Studies related to tritrophic toids in overwintering areas and to reduce the mi- interactions are needed to elucidate the role of the gration of the pest northward (Gross & Pair 1986). plant metabolites on the susceptibility of the pest Because much of the published work does not in- to parasitoids and pathogens, as well as their suit- clude environmental data with the collections, we ability (Molina-Ochoa et al. 1999). A high diver- encourage authors to include this information in sity of FAW parasitoids has been reported in its the future. These data would help select candi- natural distribution exerting significant mortal- date parasitoids for a specific or broad region. Armyworm Symposium 2002: Molina-Ochoa et al. 279 Te To Ve PN = Peanut, R = Rice, PN = Peanut, , P. maximum . PLANTS

PG = Para grass, Pm = grass, PG = Para , HOST

NR SSCSYT DIFFERENT

host plants Panicum barbinode FROM

FAW COLLECTED FAW FROM

REPORTED

TAXON

EACH

IN

BCMOP PbPGPmP ABGCMMiO SPECIES

PARASITOID

OF

hinidae 6 4 7 34 4 1 1 15 4 13 3 5 1 1 1 c erilampidae 1 UMBER BombyliidaePhoridaeSarcophagidaeTa BethylidaeBraconidae 1 ChalcididaeEulophidaeIchneumonidaeP Pteromalidae 7Scelionidae 4Trichogrammatidae 4 4 1 1 12 1Acuguttuidae 24 5 3 1 1 36 2 4 9 1 5 2 2 1 2 1 1 4 1 4 5 1 11 2 11 2 1 1 1 1 1 1 1 4. N A = Alfalfa, BG = Bermuda grass, C = Cotton, M = Maize, Mi = Millet, O = Onion, P = Hot Pepper, Pb = P = Hot Pepper, O = Onion, Mi = Millet, M = Maize, C = Cotton, BG = Bermuda grass, Alfalfa, A = ABLE S = Sorghum, SC = Sugarcane, SY = Soybean, T = Tomato, Te = Teosinte, To = Tobacco, Ve = Vegetable crops Vegetable = Ve Tobacco, = To Teosinte, = Te Tomato, T = SY = Soybean, SC = Sugarcane, S = Sorghum, Diptera Hymenoptera Nematoda Order Family T 280 Florida Entomologist 86(3) September 2003

Parasites and Parasitoids of Spodoptera frugiperda reared in the laboratory on S. frugiperda pupae. Pupae and Adults Carpenter & Greany (1998) compared the devel- opmental time, weight, fecundity, longevity, and Spodoptera frugiperda pupae and adults are ability to parasitize hosts for D. introita wasps de- attacked by several parasite and parasitoid spe- veloping on artificial diet and wasps reared on cies. Five species of Ichneumonidae: Diapetimor- S. frugiperda pupae. They conclude that the abil- pha introita (Cresson), Cryptus albitarsis ity to rear D. introita on an inexpensive, artificial (Cresson), Ichneumon promissorius (Erichson), diet significantly enhances the potential of mass Ichneumon ambulatorius and Vulgichneumon rearing this parasitoid for inundative releases brevicinctor (Say) have been reported attacking against species in the genus Spodoptera. pupae of S. frugiperda and other noctuids (Bechinski & Pedigo 1983; Pair & Gross 1984; Studies on Ichneumon promissorius Wilson 1983; Pair & Gross 1989; Fitt & Daly and Collection of Ichneumon ambulatorius 1990; Pavuk & Stinner 1991). Two generalist pu- pal parasitoids of Chalcididae, Brachymeria I. promissorius was collected in Australia from ovata (Say) and B. robusta (Cresson), have been Helicoverpa armigera (Hübner), and H. punctig- collected from S. frugiperda (Wilson 1923; Lugin- era (Wallengren) pupae (Chadwick & Nikitin bill 1928; Parker et al. 1953; Ashley 1979; Virla et 1976; Wilson 1983; Fitt & Daly 1990). The parasi- al. 1999). Only one eulophid species, Trichospilus toid was imported into the US and released in Ar- pupivora (Ferriere), has been reported as a gener- kansas, Georgia, Oklahoma, and Texas from 1992 alist parasitoid of Lepidopteran pupae, including to 1997 (J. E. C. et al., unpublished data). Follow- S. frugiperda (Alam 1979, Anantanarayanan ing releases of I. promissorius in ear-stage corn in 1934). Also, only one species (the ectoparasitic the lower Rio Grande valley in Texas during 1993 nematode, Noctuidonema guyanense Remillet & and 1994, feral noctuid pupae, including FAW, Silvain (Remillet & Silvain 1988)), has been ob- were removed from the soil. During 1993, 575 served parasitizing S. frugiperda adults. FAW pupae were collected, 8 of which were para- sitized by I. promissorius. Of the 13 FAW pupae Studies on Diapetimorpha introita collected in 1994, none were parasitized by I. promissorius. A similar study was conducted in a Diapetimorpha introita, a parasitoid which at- cornfield in Rabun County, Georgia. During 1993, tacks FAW in the pupal stage, was reported for 141 of the 300 FAW pupae collected were parasit- first time by Pair & Gross (1984) in Tifton, Geor- ized by I. promissorius and 24 were parasitized by gia. The levels of parasitization in six trials ranged I. ambulatorius, a new host record. Only 5 FAW from 0.0 to 23.7%, and the percent of intact pupae pupae collected in 1994. Two of these FAW pupae that were parasitized during that study averaged were parasitized by I. promissorius and one was 13.5%. Later, Gross & Pair (1986) emphasized the parasitized by I. ambulatorius. need for more efforts to explore the role of species Carpenter et al. (1994) compared several in- of parasitoids that employ similar strategies. Pair digenous lepidopteran species in Tifton, GA, for I. & Gross (1989) reported the seasonal incidence of promissorius acceptance and development. Pupae D. introita with rates of parasitism that averaged of H. zea, H. virescens (F.), H. subflexa (Guenée), 5.2% (range 0-23.7%) and 8.4% (range 0-50.0%), S. frugiperda, S. exigua (Hübner), Trichoplusia ni respectively, during 1983 and 1984, with the high- (Hübner), Agrotis ipsilon (Hufnagel), and Antic- est rate occurring during September to November arsia gemmatalis (Hübner) were used as hosts. of each year. Because male D. introita are at- Ninety pupae/species were tested, resulting in 74, tracted to chemicals emitted by the female wasps 72, 68, 66, and 62 wasps emerging from T. ni, A. (Jewett & Carpenter 1998) sticky traps baited ipsilon, H. suflexa, S. exigua, and S. frugiperda, with live female wasps were used to study the sea- respectively. Lowest emergence of wasps was ob- sonal abundance of D. introita (Jewett & Carpen- tained on A. gemmatalis (6). The most acceptable ter 2001). The highest number of adult males was hosts of I. promissorius were H. zea and H. vire- caught during early autumn which corresponded scens, as expected because I. promissorius was to the rates of parasitism reported by Pair and collected in Australia from heliothid species (Fitt Gross (1989). As the investigation of their impor- & Daly 1990). Carpenter et al. (1994) considered tance to biological control of Spodoptera spp. pro- that H. zea should be the primary host species of ceeds, more convenient methods of monitoring D. this pupal parasitoid in the US. introita in the field are needed. Carpenter (1995) examined the influence of Pair (1995) studied the biology and rearing of host species, host availability, and mating status D. introita on host and non-host noctuid pupae, on I. promissorius fecundity and oviposition. The Spodoptera spp., Helicoverpa zea (Boddie), and results from this study suggest that the females Heliothis virescens (F.) to identify factors that in- budget their energy expenditures and regulate fluence the reproduction and developmental rate oogenesis to maximize their reproductive poten- of D. introita. This parasitoid was successfully tial. Host size, as well as the host species, may Armyworm Symposium 2002: Molina-Ochoa et al. 281 have contributed to differences in weight of the Pyralidae, and Sphingidae were naturally infested wasps, and contributed to differences in fecun- with this nematode species. Twenty-five species of dity, longevity, and oviposition between wasps Noctuidae were infected by N. guyanense, the reared on S. exigua and H. zea pupae. Virgin fe- hosts most commonly infected were Lesmone for- males reared on S. exigua pupae laid fewer eggs mularis (Hübner), S. dolichos (F.), S. frugiperda, than virgin females reared on H. zea pupae. and Xanthopastis timais (Cramer) (Rogers et al. 1990a). Using the same methodologies, Rogers et Studies on Vulgicheumon brevicinctor al. (1990b) determined that multiple species of and Cryptus albitarsis Mocis and Spodoptera were parasitized by the nematode in Florida and Georgia, and it was the Two V. brevicinctor were recovered from FAW first record of this parasite in North America. Sim- pupae collected in 1982 from a cornfield near Tif- mons & Rogers (1990a) determined the distribu- ton, GA (Pair & Gross 1989). Five V. brevicinctor tion and prevalence of the nematode in tropical were recovered from a sample of 300 FAW pupae Americas, occurring in northern South America, collected from a corn field in Rabun County, GA, Mexico, Texas, Florida, Bermuda, most of the Car- in 1993, and one V. brevicinctor was recovered ibbean basin countries, and Central America. from a sample of 3 FAW pupae collected from the Since 1990, numerous studies have been con- same field in 1994 (J. E. C., unpublished data). V. ducted on this nematode including biology (Sim- brevicinctor also has been reported from other mons & Rogers 1994; Marti & Rogers 2000), noctuids and species of other lepidopteran fami- pathological effects on the host (Marti et al. 1990; lies (Carlson 1979). For example, Bechinski & Rogers et al. 1993), infestation dynamics (Rogers Pedigo (1983) studied the population dynamics of & Marti 1992a; Silvain & Remillet 1993; Rogers the green cloverworm (Plathypena scabra F.) in & Marti 1993b), geographical distribution (Rog- soybeans in Iowa during 1979 and 1980. They and ers et al. 1993; Rogers et al. 1991; Simmons et al. found that V. brevicinctor acted in a delayed den- 1991; Rogers et al. 1997; Marti et al. 2000), ecol- sity-dependent manner on green cloverworm pu- ogy (Silvain & Remillet 1993), population profiles pal mortality. Pavuk & Stinner (1991) reported (Rogers & Marti 1992a, 1994), mating behavior that V. brevicinctor was reared from pupae of an (Simmons & Marti 1992), prevalence (Simmons & arctiid, Cisseps fulvicollis (Hübner), and pupae Rogers 1990b), maintenance of colonies (Rogers & from a pyralid, Ostrinia nubilalis (Hübner). Marti 1993a, b), host range (Simmons & Rogers Pair & Gross (1989) collected C. albitarsis in 1996; Rogers & Marti 1996; Marti et al. 2000), Tifton, GA, from a single sample taken November and the bionomics of host insects of N. guyanense 2, 1984. Four C. albitarsis also were recovered (Rogers et al. 1996; Marti et al. 2000). Subse- from a sample of 300 FAW pupae collected from a quently however, a study on the speciation in corn field in Rabun County, GA, in 1993 (J. E. C., Acugutturidae shows that N. guyanense is limited unpublished data). Pair & Gross (1989) reported to the lepidopteran genera Spodoptera and Pseu- that C. albitarsis had been established in a labo- daletia (Marti et al. 2002). ratory colony in Tifton, GA. C. albitarsis cohorts Simmons et al. (1991) studied the seasonal from this laboratory colony were successfully chronology of the nematode in the tropical and reared on an artificial diet devoid of any host com- subtropical America, and determined that host ponents (Greany and Carpenter 1998). parasitism and nematode population density var- ied among locations and over time. Examples in- Studies on Ectopasitic Nematodes Attacking Adults clude higher parasitism on male moths in Grenada of Spodoptera frugiperda (77%) than in Texas (1%), and higher nematode populations at lower latitudes than higher lati- Remillet & Silvain in 1982 discovered and re- tudes. Simmons & Rogers (1991) also studied the ported an ectoparasitic nematode, Noctuidonema dispersal and seasonal occurrence of N. guyanense guyanense Remillet & Silvain infecting Spodoptera on FAW adults in the US. They found that nema- androgea (Cramer) in French Guiana. It was sub- tode populations and parasitism of FAW males sequently described as a new genus and species were higher in Eastern States than in the Plains, (Remillet & Silvain 1988) most commonly found on Midwestern, and Central states, and that the per- moths of FAW, Spodoptera latifascia (Walker), S. cent parasitism and the number of nematodes per marima (Schaus), Anicla infecta (Ochsenheimer), infested FAW changed over time at each location. and Leucania spp. (Remillet & Silvain 1988). Rogers et al. (1990a) determined the life cycle and host range for N. guyanense in French Gui- ACKNOWLEDGMENTS ana. They collected moths using a white sheet illu- The authors thank J. J. Hamm and O. G. Marti minated by UV light, by pheromone traps, and (USDA-ARS-Crop Protection & Management Research sweeping vegetation from a variety of habitats in Laboratory, Tifton, GA), for their critical review of ear- Northeastern French Guiana. Moths in five fami- lier versions of this manuscript, and thank O. G. Marti lies, Lasiocampidae, Noctuidae, Notodontidae, for providing information on ectoparasitic nematodes. 282 Florida Entomologist 86(3) September 2003

The authors appreciate the assistance of Duncan Mc- ASHLEY, T. R. 1983. Growth pattern alterations in fall Clusky (University of Georgia Libraries, Tifton, GA), armyworm, Spodoptera frugiperda larvae, after par- and Marco Antonio Mellín-Rosas (CNRCB, Tecomán, asitization by Apanteles marginiventris, Campoletis Colima, México) in obtaining scientific literature. The grioti, Chelonus insularis, and Eiphosoma vitticole. authors gratefully acknowledge and thank Susan Florida Entomol. 66: 260-266. Drawdy (USDA-ARS-Crop Protection & Management ASHLEY, T. R., C. S. BARFIELD, V. H. WADDILL, AND E. R. Research Laboratory, Tifton, GA) for her exceptional MITCHELL. 1983. Parasitization of fall armyworm technical assistance and dedication to this project. In- larvae on volunteer corn, bermuda grass, and para- sect identifications and verification of insect classifica- grass. Florida Entomol. 66: 267-271. tions were provided by several scientists associated ASHLEY, T. R. 1986. Geographical distributions and par- with the USDA/ARS Systematic Entomology Labora- asitization levels for parasitoids of the fall army- tory, including B. Carlson, E. E. Grissell, P. Marsh, N. E. worm, Spodoptera frugiperda. Florida Entomol. Woodley, and N. W. Gates. This paper is a contribution of 69:516-524. the University of Nebraska Agricultural Research Divi- AVALOS, D. S. 1988. Moscas tachinidae de la provincia de sion, Lincoln, NE 68583. Journal Series No. 13681, De- Córdoba (Argentina). Rev. Peruana Entomol. 31: 48-50. partment of Entomology, University of Nebraska- AYALA-SIFONTES, J. L., M. GONZÁLEZ-ALVAREZ, H. Lincoln. The authors thank Universidad de Colima, and GRILLO-RAVELO, Y S. CABALLERO-FIGUEROA. 1978. CONACYT-Mexico for supporting the senior author. Observaciones preliminares sobre el ciclo biológico de Diadegma sp., parásito de Heliothis virescens (Fabricius). Centro Agrícola 5: 1-13. REFERENCES CITED AYALA-SIFONTES, J. L., J. RAMÍREZ-OBREGÓN, S. 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