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Home , Spodoptera, Spodoptera dolichos, Spodoptera latifascia, Spodoptera ornithogalli

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Center for Systematic Entomology, Gainesville, Insecta Mundi

September 1991

Color identification of economically important larvae in (: )

S. Passoa USDA/APHIS/PPQ

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Passoa, S., "Color identification of economically important Spodoptera larvae in Honduras (Lepidoptera: Noctuidae)" (1991). Insecta Mundi. 414. https://digitalcommons.unl.edu/insectamundi/414

This Article is brought to you for free and open access by the Center for Systematic Entomology, Gainesville, Florida at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Insecta Mundi by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Vol. 5, No. 3-4, September-December 1991 185

Color identification of economically important Spodoptera larvae in Honduras (Lepidoptera: Noctuidae)

S. Passoa USDAIAPHISIPPQ 8995 E. Main Street Bldg. 3, Room 109 Reynoldsburg, OH 43068

Abstract Spodoptera frzgiperda (J.E. Smith), S. exigua (Hiibner),S. latifascia (Walker),S. ornithogalli (Guenb), S. dolichos (Fabricius), S. sunia (Guenee), and S. eridania (Cramer) are commonly associated with crops in Honduras. A key to these species with color illustrations of rare and typical forms is presented. Potential problems in identifying Spodoptera species are discussed.

Additional key words: morphology,

Introduction mens are misidentified (Figures 9.1, 9.4, and 11.2 The noctuid Spodoptera contains many label three forms of as S. pests throughout the world (Kranz et al. 1977, Hill dolichos, S. eridania, and , 1975). At least eight species are known to occur in respectively). A guide to soybean pests in Honduras. Spodoptera frugiperda (J. E. Smith), (Gazzoni et al. 1981) incorrectly labeled a photo- Spodoptera exigua (Hiibner),Spodoptera latifascia graph of S. sunia as S. eridania. (Walker), and Spodoptera sunia (Guenee) are con- This work should facilitate identification of sidered economically important pests whereas common Spodoptera larvae in Honduras by presen- (Cramer), Spodoptera orni- tingillustrations of typical andunusual color forms. thogalli (Guenbe), and (Fabri- Careful attention to details of the color pattern will cius) are rarely abundant enough to cause serious allow accurate identification of most Spodoptera damage to Honduran crops (Passoa 1983). An adult species, although greatvariation in larval markings Spodoptera androgea (Cramer) was collected on the makes the identification of all aberrant individuals north coast of Honduras (La Lima, Department of impossible. Cortes) in October, but its larva is unknown. The host range of Spodoptera is very wide including Methods and Materials most vegetable crops and several basic grains (Passoa Most of the specimens used in this study were 1983, Andrews 1984, King and Saunders 1984). collected in Honduras, Central America, from 1979- Information on the distribution, phenology, Span- 1981. Additional specimens from , and ish common names, biology, and control of Spodoptem Florida were also studied. Field-collected larvae spp. can be found in the works cited above. were photographed, reared to adult, and then iden- In spite of a logical and thorough study by Levy tified with Todd and Poole (1980). Eggs laid by and Habeck (1976), workers in Latin America female in captivity provided another source sometimes have trouble identifying Spodoptera of larval material. Beans (Phaseolus vulgaris L.), larvae. Two examples illustrate this problem. In sweet potato (Ipomoea batatas (L.) Poir.) or wheat King and Saunders (19841, several SpaEoptera speci- germ artificial diet were all accepted by Spodoptera Insecta Mundi larvae in laboratory feeding trials. Larval speci- Dorsal pinacula usually conspicuous (Figs. 12,14) mens were killed in a 9:l mixture of 80% ethanol but sometimes pale in the green form (Fig. 13), and glacial acetic acid by volume, then transferred their diameter equal to or greater than the dia- to 80% ethanol after 24 hours for permanent stor- meter of the abdominal spiracles dorsum of ab- age. Larval terminology follows Stehr (1987). The dominal segments granulated under a mag- nification of 25 power or greater (Fig. 4) hypopharyngeal complex was mounted in glycerin ...... Spodoptera frugiperda (J. E. Smith) and examined under a compound microscope. Only Dorsal pinacula never conspicuous, their diameter the most common synonyms are mentioned under less than the diameter of the abdominal spir- each species, for a more complete list see Todd and acles dorsum of abdominal segments smooth Poole (1980). under magnification of 25 power ...... 2 Characters to separate Spodoptera from other Abdominal segments never with dorsal triangular common pest noctuids in Honduras markings, pattern consists of a series of dorsal dashes (Fig. 6) or, more commonly, an irregular The following combination of structural and series of white dots and lines (Fig. 15) lateral color characteristics will separate Spodoptera spp. spot, if present, is on the mesothorax (Fig. 1) from other common pest noctuids in Honduras. The ventral tonofibrilllary platelets between abdominal polarity of most of these characters are unknown prolegs form a "Y' with a short stem about as although the presence of a sclerotized bar around long as the arms (Fig. 9) ...... the mesothoracic SD 1seta seems to define a clade ...... Spodoptera exigua (Hiibner) which includes Spodoptera and related genera. Abdominal segments with at least one pair of dorsal triangular markings (Figs. 2, 11, 23) 1. Four abdominal prolegs present (larva not a lateral spot, if present, is on the first abdominal segment (Figs. 2,19) ventral tonofibrillary plat. looper). lets between abdominal prolegs form a straight 2. Two setae present on first abdominal seg- SV line or, more rarely, a "Y" with gvery long stem ment (Fig. 2). compared to the arms (Fig. 10)...... 3 3. Lateral spot usually present on first abdominal segment (this spot is a practical method of rec- Lateral spot on the first abdominal segment, when ognizing Spodoptera larvae in the field) (Fig. present, interrupts the subspiracular line to 2,26). form a pale, thinner line on the thorax (Fig. 19) 4. Mesothorax with a sclerotized bar connecting if lateral spot and/or subspiracular line are the SD 1setaand a tonofibrillaryplatelet (~ig. absent (Figs. 18, 20), then dorsal triangles of 3). first abdominal segment larger than dorsal tri- 5. No retinaculum on the mandible (Fig. 5). angles of fourth abdominal segment (Fig. 17) Spodoptera eridunia (Cramer) 6. Hypopharyngeal complex with coarsely spined ...... Lateral spot on the first abdominal segment, when posterior portion lacking a dense brush of present, does not interrupt the subspiracular stout bristles (Fig. 7). line and therefore the line maintains equal in- 7. Adfrontal sutures do not extend to epicranial tensity and width on the thorax compared to the notch (Fig. 8). abdomen if lateral spot and/or subspiracular 8. Skin smooth under a magnification of 25 power line are absent, then dorsal triangles of first (except in Spodoptera frugiperda) and never abdominal segment not larger than dorsal tri- with microspines. angles of fourth abdominal segment (Fig. ll) 9. Head with an inverted "Y'' because adfrontal areas are outlined in white (see discussion under Spodoptera frugiperda) (Fig. 32). Dorsal triangles with a narrow white line (either solid or composed of dashes) passing through them (Fig. 27), if lines are inconspicuous or absent in dark forms (Figs. 24-25,28), then area Key to common Spodoptera larvae below subdorsal stripe with several longitudinal in Honduras narrow white lines (Fig. 24) dorsal mesothoracic (modifiedfromLevy and Habeck 1976, Oliver and Chapin marking either triangular, elongate-oval,or semi- 1981, Crumb 1956, and Godfrey 1987) circular (Figs. 11, 32) ...... Spodoptera ornithogalli (GuenBe) Dorsal triangles never with a narrow white line passing through them, although a series of white Vol. 5, No. 3-4, September-December 1991

dots may be present (Figs. 31, 34) area below with cutworms that have granulated skin and large subdorsal stripe without several longitudinal pinacula, for example, Agrotis ipsilon (Hufnagel). narrow white lines except for the black form of However, cutworms have the adfrontal suture ex- S. latifascia (Fig. 34) dorsal mesothoracic mark- tending to the epicranial notch (top ofhead) while in ings semicircular or trapezoidal (Figs. 32, 33), S. frugiperda the adfrontal area extends only half never triangular or elongate-oval ...... 5 that distance. 5. Dorsal triangles with a white spot close to or at their apex (Fig. 22) if triangles are absent then Spodoptera (= Laphygma) exigua (Hu bne r) ; at least some of the white dots are bordered by black semicircles (Fig. 23) ...... A heavily-marked form of the beet armyworm ...... Spodoptera sunia (GuenBe) was illustrated by Levy and Habeck (19761, but this 5'. Dorsal triangles without a white spot close to their color pattern is rare in Honduras. Instead, most of apex although a white spot may be present below the triangles on the subdorsal line (Fig. the larvae are speckled with white on a green 30) ...... 6 background (Fig. 15). Head color varies from green to black and a thin white spiracular line may be 6. Mesothoracic dorsal markings usually semicir- present (Fig. 16). Spodoptera exigua can usually be cular (Figs. 32,34) and smaller than dorsal tri- recognized by a lateral spot on the mesothorax (see angles on eighth abdominal segment (Fig. ll) discussion under S. fiugiperda). However, preserved rarely mesothoracic dorsal markings are re- specimens of S. exigua usually fade in alcohol and placed by a white spot (Fig. 31) ...... lose their color pattern so most identifications must ...... Spodoptera latifascia (Walker) be confirmed with structural characters. Among 6'. Mesothoracic dorsal markings trapezoidal (Fig. common noctuids in agricultural fields in Hon- 30) and subequal to dorsal triangles on eighth duras, the presence of two subventral setae on the abdominal segment (Fig. 291, these markings 21, never replaced by a white spot ...... first abdominal segment (Fig. a sclerotized bar ...... Spodoptem dolichos (Fabricius) connecting the mesothoracic SD 1seta to a tonofib- rillary platelet (Fig. 3), the ventral tonofibrillary platelets in the form of a "Y" with a short stem (Fig. Discussion 91, and smooth skin under a magnXication of 25 power will identify S. exigua. Spodoptera (= Laphygma) frugiperda (J. E. Smith); . Spodoptera (=Xylomyges) eridania (Crame r) ; Usually Spodoptera frugiperda is readily distin- southern armyworm guished by its large pinacula (Figs. 12 and 14). Despiteits name, the southern armyworm is not However, the green form of S. frugiperda can be as common in Honduras as S. frugiperda, S. sunia, confused withSpodoptera exigua (compareFigs. 13, or S. latifascia. Spodoptera eridania has been tradi- 15 and 16) because the pale pinacula of S. frugiper- tionally distinguished by the spot on the first ab- da are easy to overlook, especially in field surveys. dominal segment which interrupts the subspiracu- Cuticular texture can also be used to distinguish lar line (Levy and Habeck, 1976).However, the term these two species. Spodoptera frugiperda has a "interrupted" is somewhat misleading since the spot granulated cuticular texture (Fig. 4) under a mag- does not physically sit in the path of the line. nification of 25 power or more whereas S. exigua has Instead, the subspiracular line loses intensity as it a smooth cuticle at the same magnification. The lat- passes below the lateral spot (Fig. 19).Most Spodop- eral spot of S. frugiperda, when present, is on the tera larvae have a lateral spot so the condition of the first abdominal segment and not the mesothorax as subspiracular line must be noted. The illustration of is the case with S. exigua. Although the literature S. latifascia (Fig. 9.4) in King and Saunders (1984) suggests S. frugiperda is distinguished by the white was probably misidentified asS. eridania because of "inverted Y" on its head (Angulo and Weigert 19751, its prominent lateral spot. pale adfrontal areas are characteristic of other Unfortunately, sometimes the subspiracular line Spodoptera spp. (for example, Fig. 22) and many of S. eridania is pale (Fig. 9) or absent (Fig. 8). representatives in scattered families throughout Under these circumstances the southern army- the Lepidoptera. worm is dEcult to recognize. One character which The fall armyworm can potentially be confused appears unique to S. eridania is the large size of the Insecta Mundi dorsal triangles on the first abdominal segment mesothoracic markings can be helpful in identifying (Fig. 17).The presence of a white spot posterodorsad Spodoptera larvae. The mesothoracic spot of S. to the spiracle wasused by Oliver and Chapin (1981) ornithogalli may be round, elongate-oval, or trian- in their key to economically important noctuids. Al- gular When a triangular or elongate-oval mesotho- though this spot is common on S. eridania, it also racic spot is present (Figs. 11,281, this separates S. occurs in other Spodoptera species, for example, S. ornithogalli from S. latifmcia and S. dolichos. In latifmcia and S. ornithogalli. Levy and Habeck ( the latter two species the mesothoracic markings 1976)mentioned a uniform brown head as a backup are semicircular (Fig. 32) and trapezoidal (Fig. 30), character for recognizing the southern armyworm. respectively. This head coloration characterizes most typical specimens ofS. eridania but sometimes the anterior Spodoptera (= Prodenia) dolichos (Fabrici us) ; portion of the epicrania may be dark brown with banded armyworm strong reticulations. Too few specimens ofS. dolichos are available to examine color variation. In a series of about 50 Spodoptera (=Xylomyges) sunia (G ue n ee) ; larvae reared from a female collected in Honduras, white-spotted armyworm all specimens showed the characteristic mesothor- Spodoptera sunia is readily identified by a white acic trapezoidal markings when mature (Fig. 30). spot at the apex of the dorsal triangles (Fig. 22). If Early instars of S. dolichos resemble S. latifascia the dorsal triangles are reduced, then at least sev- because both species have semicircular mesothor- eralof the spots will be ringed with black (Fig. 23), acic markings. representing remnants of the dorsal triangles. A new common name, the banded armyworm, is The variegated form of S. sunia (Fig. 21) may be proposed here to call attention to the distinctive hard to identify because the dorsal triangles are bands on the adult thorax which readily separate inconspicuous and the dorsum is flecked with white. this species from other members of the genus (Todd With careful observation, a subdorsal triangle with and Poole, 1980). white spot can usually be discerned. Several species of Spodoptera have white spots Spodoptera (= Prodenia) latifascia (Walker) ; at the bases of the dorsal triangles (Fig. 23) which black armyworm potentially could be confused with the spotted tri- Spodoptera latifascia is very common in Hon- angles in S. sunia. Only white spots at, or near, the duras. The body color usually is a shade of black, apex of the dorsal triangle should be used to recognize brown or gray. Dorsal triangular markings are S. sunia. either present or absent, rarely the mesothoracic dorsal markings are reduced to white dots. S. lati- Spodoptera (= Prodenia) ornithogalli fascia should not be confused with either S. dolichos, (Guenee); yellow-stri ped armyworm S. sunia, S. frugiperda, or S. exigua. Spodoptera Spodoptera ornithogalliis characterized by a ornithogalli can be confused with the black form of thin white line, either solid or composed of dashes, S. latifucia if the white markings passing through which passes through the abdominal dorsal tri- the dorsal triangles are not examined carefully. In angles. Often the line is readily visible (Fig. 27) al- S. latifascia these markings are composed of a though sometimes it may be pale and difficult to see series of dots, never a line or series of dashes asis the (Figs. 24,25 1. In the latter case, most individuals of case with S. ornithogalli. In addition, the subdorsal S. ornithogalli can be recognized by having several stripe is either yellow, white, or brown (Figs. 24-26, narrow white lines under the subdorsal stripe (Figs. 28) in S. ornithogalli while it is nearly always 24, 26). Spodoptera ornithogalli gets its common orange in the black form of S. latifascia (Fig. 34). name from the conspicuous yellowish-whitesubdor- If additional species of Spodoptera larvae are sal stripe (Figs. 24,27). Even though several other included aspart of a revision of the New World taxa, Spodoptera larvae have a prominent subdorsal stripe, several characters listed below may merit further and the stripe is not always yellow in S. orni- study. Whelen (1935) used tarsal claw shape in a thogalli, a contrasting yellow subdorsal stripe is a key to Nebraska armyworms. Differences in spira- good indication that an unknown larva probably is cle color helped Okumura (1961)distinguish among S. ornithogalli. Spodoptera larvae in cotton. Head color and man- Despite great variation, the shape of dorsal dible shape were generally too variable for use in Vol. 5, No. 3-4, September-December 1991

identification of Honduran Spodoptera larvae, al- Angulo, A. O., and G. T. Weigert. 1975a. Estados though these characters might be significant if inmaduros de lepid6pteros n6ctuidos de impor- additional species are considered. The presence and tancia econ6mica en Chile y claves para su color of the middorsal line varies and it is especially determinacibn (Lepidoptera: Noctuidae). Socie- obvious and intense in some S. sunia and S. eri- dad de Biologia de Concepci6n (Chile) Publi- dania. Ideally, Spodoptera larvae need to be studied cation Especial 2.153 pp. by rearing females in laboratory cultures for many generations, atseveral densities and temperatures, Angulo, A. 0.1975b. Noctuidae (Lepidoptera) de on awide variety ofhostplants. Only in this way can interes econdmica del valle de Ica, : Clave we begin to understand the extent of color variation para estados inmaduros. Rev. Peruana Entomol. in these . 18 (1):98-103.

Commonwealth Institute of Entomology. 1977. Acknowledgements Distribution maps of pests. Spodoptera or- I am grateful to all Peace Corps volunteers and nithogalli (Gn.) Series A (Agriculture), map no. agronomists of the Department of Natural Resour- 367. London, England. 2 pp. ces who helped me collect specimens in Honduras. Dr. D. Habeck (University of Florida) deserves Crumb, S. E. 1929. Tobacco cutworms. U.S. Dep. special thanks for all his time and effort in review of Agric. Tech. Bull. 88. 179 pp.

L this manuscript. The comments ofDrs. K. Andrews, R. Cave and Crumb, S.E. 1956. Larvae of the Phalaenidae. U.S. R. Caballero (Panamerican Agriculture School, Dep. Agric. Tech. Bull. 1135. 356 pp. Honduras), Dr. G. Godfrey (Illinois Natural History Gazzoni, D., E. B. de Oliveira, I. C. Corso, B. S. Survey) and Dr. M. Berenbaum (University of Illi- L.V. nois) greatly improved this paper. Dr. M. Spellman C. Ferreira, G. B6as, F. Moscardi, and (University of ) kindly provided a photo- A. R Panizzi. 1981. Manejo de pragas da soja. graph of S. ornithogalli (Fig. 26). Embrapa Cir. Tecn. 5.44 pp. Godfrey, G. 1987. Family Noctuidae. p. 549-579. In Bibliography F. Stehr (Ed.), Immature Insects. KendalVHunt Publishing Co. Dubuque, . 754 pp. The following is a selected bibliography of refer- ences useful in studying Spodoptera spp. because of Hill, D.S. 1975. Agricultural pests of. the their illustrations or biological notes. tropics and their control. Cambridge University Press. Cambridge, England. 516 pp. Andrews, K. 1980. The whorlworm, Spodoptera frugiperda, in Central America and neighboring Howell, Jr., H.N. 1978. Fall armyworm in a pine areas. Fla. Entomol. 63 (4): 456-467. nursery in Honduras, C.A. Ceiba 22 (1):35-37.

Andrews, K. 1984. El manejo integrado de plagas King, A.B.S., and J.L. Saunders. 1984. The in- invertebradas en cultivos agronomicos, horticolas vertebrate pests of annual food crops in Central y frutales en la Escuela Agricola Panamericana. America Overseas Development Administration. Guia para el laboratorio de campo en entomol- London, England. 166 pp. ogia. El Zamorano, Honduras. 86 p. Kranz, J., H. Schmutterer, and W. Koch. 1977. Angulo, A. 0,and C. Jana S. 1982. La pupa de Diseases, pests and weeds in tropical crops. Spodoptera Guende, 1852, en el norte de Chile Verlag Paul Parey. Berlin, Germany. 666 pp. (Lepidoptera: Noctuidae). Agric. Tecnica (Sa- ntiago) 42 (4): 347-349. Levy, R, and D.H. Habeck. 1976. Descriptions of the larvae of Spodoptera sunia and S. latifascia with a key to the mature Spodoptera larvae of the Eastern . Ann. Entomol. Soc. Am. 69 (4): 585-588. 190 Insecta Mundi

Luginbill, P. 1928. The fall armyworm. U.S. Dep. Passoa, S. 1983. Lista de 10s insectos asociados Agric. Tech. Bull. 34.91 pp. con 10s granos basicos y otros cultivos selectos en Honduras. Ceiba 25 (1): 1-98. Marti, O.G., and C.E. Rogers. 1988. Anatomy of the ventral eversible gland of fall armyworm, Peairs, F.B., and JL. Saunders. 1979. The fall Spodoptera frugiperda (Lepidoptera: Noctuidae), armyworm, Spodoptera frugiperda, a review. larvae. Ann. Entomol. Soc. Am. 81 (2): 308-317. Ceiba 23 (2): 93-113.

Ngo Dong, T.C. Carlysle, H. L. Cromroy, and D. Peterson, A. 1962. Larvae of Insects. Edwards H. Habeck. 1980. Morphological studies on the Brothers Inc. Ann Arbor, . 732 pp. beet armyworm, Spodoptera exigua (Hiibner) (Lepidoptera: Noctuidae) Florida Exp. Sta Bull. Redfern, RE. 1967. Instars of southern army- 816.38 pp. worm determined by measurement of head capsule. J. Econ. Entomol. 60 (2): 614-615. Okumura, G.T. 1961. Identification of lepidop- terous larvae attacking cotton. Dep. Rings, RW., and G.J. Musick. 1976. A pictorial Agric. Special Pub. 282. 80 p. field key to the armyworms and cutworms at- tacking corn in the North Central States. Ohio Oliver, AD., and J.B. Chapin. 1981. Biology and Res. and Dev. Center Circ. 221.36 pp. illustrated key for the identification of twenty species of economically important noctuid pests. Todd, EL., and RW. Poole. 1980. Keys and Sta. Agric. Exp. Sta. Bull. 733.26 pp. illustrations for the armyworm moths of the noctuid genus Spodoptera Guen6e from the Pashley, D.P., SJ. Johnson, and A.N. Sparks. Western Hemisphere. Ann. Entomol. Soc. Am. 1985. Genetic population structure ofmigratory 73 (6): 722-738. moths: the fall armyworm (Lepidoptera: Noc- tuidae). Ann. Entomol. Soc. Am. 78 (6): 756-762. Walkden, EH. 1950. Cutworms, armyworms and related species attacking cereal and forage crops Pashley, D.P., S.J. Johnson, and A.N. Sparks. in the Central Great Plains. U.S. Dep. Agric. 1986. Host-associated genetic differentiation in Circ. 849. 52 pp. fall armyworm (Lepidoptera: Noctuidae): a sib- ling species complex? Ann. Entomol. Soc. Am. Whelan, D.B. 1935. A key to the Nebraska cut- 79 (6): 898-904. worms and armyworms that attack corn. Ne- braska Agric. Exp. Sta. Bull. 81. 25 pp. Pashley, D.P., S.J. Johnwn, and AN. Sparks. 1988. Quantitative genetics, development, and physiological adaptation in host strains of fall armyworm. Evolution 42 (1): 93-102.

Figure 1-11. 1) Lateral view of typical Sphptera head, prothorax and mesothorax. 2) Lateral view of a typical Spodoptera first abdominal segment. 3) SD 1 pinaculum of a typical Spodoptera larva. Arrow points to a tonofibrillary platelet which is connected to the pinaculum by two scl erotized bars. 4) Granulated cuticular texture of Spodoptera frugiperda dorsum at high power. 5) Mandible of a typical Spodoptera larva showing the lack of aretinaculum. 6) Dorsal view of Spodoptera erigua abdominal markings that are not triangular. 7) Lateral view of typical Spodoptera larval hypopharyngeal complex. Arrow points to a hypopharyngeal blade which is sometimes present. 8) Anterior view of typical Spodoptera head showing that adfrontal area (bottom arrow) does not reach the epicranial notch (top arrow). 9) Tonofibrillary platelets between the prolegs of A3-6 in Spodoptera ezigua. Arrow points to a 'Y" with a short stem. 10) Tonofibrillary platelets between the prolegs of A3-6 in most Spodoptera la~aethat form a straight line or a 'Y' with a long stem (see arrow). 11) Dorsal view of Spodoptera latifascia showing that the mesothoracic markings (top arow) are smaller than the dorsal abdominal triangles of A8 (bottom arrow). A = antenna; Ad = adfrontal area; D = dorsal area; F = front; G = prothoracic ventral gland; H = hypopharyngeal complex and spinneret; L = lateral area; L3 = third lateral seta; Mn = mandible; P = maxillary palpus; S = lateral spot of first abdominal segment; S = lateral spot of the mesothorax; SA = subdorsal area; SB = subspiracular line; SD1= first subdorsal seta; SL = subdorsal line; SV = subventral area; SV3 = third subventral seta; SV1= first subventral seta; T = dorsal abdominal triangle; V = ventral area. Vol. 5, No. 3-4, September-December 1991 191

F'ignra 1Bl7. Spadapdero spp. larvae. Figure U-14.Spodaptem wpda. 12) Darh form, showing c~nepicuous dorsal pinacuk 13) Oreen f-, arrow pinto to pale pixmculum; 14) Naamal brown f&m, showingconq' ICUW. dorsal pinacula. F'igure l.5-16.Spdbptara sxigw. 16) Normal form, showing variation in the calm ofthe doreum and an bmgular pattern of white data and e,16) LsGral view, showing white epiraclee and a white mbspi- rarmlar Iine. Figure 17. Spiwbptero wkhia,dormlateral view, arrow points ta the large doreal triangle on the f'lrst abdominal E6gment. Figum 1888. Spodoptms opp, larvae. F3gure l&S.&udoptwa sridanio. 18) kkfarm, &owing the kkof a IMP; 10)N~fmm,Ihoalaptb.nd~o11din~tydwidthofth~~~ultpu~~s~-~ mpnt Orthe Mabdominal uegment. Amown point to dspbacular line before and after it underthe laW spat; a0) Dark form, showing a pale Nbspiracular stripe. Figure 11-25. Spbptum .units. 21) ztedmlm fan;22) Doraal triangle lmarlred by arrow^), ),g a white @potnear tbe apeq 23)Dodvim, ahowing reduction of the dorsal onm&abdo~segmente.~poi~~~toa~hitespot~withbWmthe~mm~.~~aS~~ twuavdh&ii. %)Blwkfol.m,ahowing several white hies belowthe mubdorsal @tripe;%)Dark form, ahowinginco~picu- ow lina paaeiag through the dorsal triangles; a6) Brown Porm, arrow point. to lateral epot. Vd. 5. No. 3-4. Sewr-December 1991 195

Figure 27-34. Spodoptera spp. larvae. Figure 27-28. Spalop&ra ornithogolli. 27) Normal form, showing lines passing through the dorsal triangles (8ee arrow) and a yellowish-white subdorsal stripe; 28) Dark form, showing inconspicuous lines passing through the dorsal triangles. Figure 29-30. Spodoptem dolich. 29) Dorsal view, showing mesothoracic dorsal marking (top arrow) as large as the dorsal triangles of the eighth abdominal segment (bottom arrow); 30) Dorsal new, showing trapezoidal mesothoracic dorsal markings. Figure 31-34. Spodaptem latifascia. 31) Black form, arrow points to a series of white dots on the dorsum; 32) Dorsal view, arrow points to the semicircular mesothoracic dorsal marking; 33) Brown form, showing the lack of dorsal abdominal triangles; 34) Black form, showing the orange subdorsal stripe and round mesothoracic dorsal markings. 196 Insecta Mundi