ASD Oil Palm Papers, N°36, 14-33. 2011

Opsiphanes cassina Felder (: ), defoliator of the oil palm ( Jacquin) in Central America

Ramón G. Mexzón1, C.Ml. Chinchilla2

Abstract

The populations of cassina Felder associated with oil palms in Central America are normally under control by its numerous natural enemies, but occasional outbreaks occurred in the past in localized areas. This review describes some details of past out- breaks, important aspects of the (, anatomy, life cycle, behavior, sampling, control and natural enemies), its potential to cause economic damage and the possibilities to manage the pest through an integrated approach.

Key words: Life cycle, , Nymphalidae, oil palm, insect pest.

Introduction

embers of the subfamily Brassolinae are robust hairy and colorful . They Mare easy to recognize because of singu- Other important defoliations also occu- lar spots (ocellus) below the wings. The group rred in other adult plantations located along has 12 genera, nine of which are distributed the Atlantic coast, particularly in the valley of from Mexico to South tropical America. Lar- the Aguan River. vae feed on monocots, particularly Araceae, In Costa Rica, the pest caused important Musaceae, Heliconiaceae, Poaceae and Brome- damage in adult plantations located in the Cen- liaceae. Larvae of all genera, except Brassolis, tral (Quepos region, 1982) and South Pacific have cephalic horn-like appendages. (Coto, 1984). In Coto, aproximately 260 ha The genus Opsiphanes has 13 species, six of were affected when 150-600 larvae/leaf were which are in Costa Rica, and O. cassina has been found. Other outbreaks of medium to high the most frequent pest found in oil palm plan- magnitude also occurred in Panamá (Chiriquí), tations in all Central America. Reliable records Nicaragua and Guatemala. Outbreaks norma- of most defoliations are not available. The first lly occurred during the rainy season. documented damage occurred in approxima- Considering the historic importance of this tely 150 ha in 1972 in an adult plantation loca- pest in many of the oil palm plantations in Cen- ted near the city of Tela on the Atlantic coast of tral America it was considered appropriate to Honduras. Later in 1977 a new outbreak was summarize important aspects of the biology of observed in the same area, but the damage ex- the insect and several control alternatives now tended to near 250 ha and up to 380 larvae/leaf available. were counted. This focus grew to more than 1,000 ha in 1979 which was associated with 30- 60 % bunch yield decrease (Richardson 1981).

1 Museo de Insectos, Escuela de Agronomía, Universidad de Costa Rica, San José. Costa Rica. [email protected] 2 Consultor para ASD Costa Rica P.O. Box 30-1000, San José, Costa Rica. [email protected] 15

Biology and Behavior

Biology spots with a clear golden tone. Later they turn Adult. A brown moth with a corporal length of clear brown. The whole stage takes 10-15 days. about 28 mm in the male and wing span of 60- Behavior 65 mm in the male and 70-75 mm in the female. They both have characteristics transversal Larvae feed on leaves. Adults are attracted orange bands formingaYontheanterior to over ripe fruits where they obtain car- wings. On the ventral side of the wings they bohydrates and proteins from yeasts develo- present circular spots (ocellus): one on the ping on these substrates. These nutrients are anterior wing (black with a yellow halo) and the necessary fuel for flying and reaching se- two on the posterior wings (one brown with a xual maturity. black halo and other white with two rings; one black and other clear brown). Sexual dimor- Adults show two activity peaks: one during phism is well marked with males presenting a the first hours in the morning (6.00-8.00 tuft of white hairs on the posterior wings and hours) and the main one late in the afternoon yellow glandular openings on both sides of the (15.30 - 17.30 hours). Females can mate the se- abdomen. Adults may live for 7-10 days (Table cond day after emergence from papa and are 1). born with a full load of eggs which are norma- lly lay late afternoon Eggs. Eggs are globose, reticulated, and initia- lly creamy and later show reddish bands Loría et al. (2000) studied the population corresponding to the developing embryo. dynamic of the insect in the South Pacific Females lay the eggs on the abaxial (below) coast of Costa Rica and found that adults part of the leaflets, particularly near the base, emerge during a period of 2-3 months. Several individually or in small groups where they consecutive periods of emergence may occur hatch in 5-10 days of about one month each: first (10-15 days) Larva. Upon emergence they measure about 7 where males predominate, and later appear the mm and grow to ca. 80 mm in length when they females to finally reach a near 1:1 sex ratio. enter the pre-pupa stage. Initially they are rose A population increase seems to start from in color, with five yellow longitudinal bands. isolated individuals that migrate to oil palm The cephalic capsule is hirsute and black. With plantations from the surrounding natural ve- age the larvae take a green color with yellow bands (a pair on the pleuras and one Table 1. Life cycle of Opsiphanes cassina on the back that prolongs Duration (days) Length (mm)* Stage on the cephalic capsule). Mean Range Mean Range The head and caudal part of Egg 8.0 5 - 10 2.0 2 -3 the body have horn-like Larval stage 41.2 33.5 - 61.8 appendages. There are five I 7.5 6.0 - 11.0 7.5 ± 0.5 7 - 9 developmental stages that II 6.0 4.5 - 10.5 28.9 ± 2.9 23 - 30 are completed in 33-62 III 6.0 5.5 - 10.5 39.4 ± 3.3 33 - 42 days. IV 8.2 5.5 - 11.5 49.0 ± 5.3 40 - 60 Pupa. Larvae tend to V 13.5 12.0 - 18.3 69.3 ± 8.1 53 - 80 pupate on the numerous Prepupa 1.0 0.5 - 1.6 weeds that normally grow Pupa 12.0 10.0 - 15.0 on palm stems where they Adult 7.0 7.0 - 10.0 28.0 ± 1.0 26 - 31 can blend with the surroun- Total 69.2 57.1 - 98.4 dings. Initially the pupa is * n= 60, 29.0± 2°C; 87±5% R.H. light green with two small

ASD Oil Palm Papers, N° 36, 2011 16

getation. Initially, the population starts to in- of the central vein of the leaflet and then the crease in open areas (i.e. along roads and ca- larvae changes position to start a similar cut on nals) where tunnel winds are formed which are the other side of the vein, which finally leaves a supposed to favor the dissemination of phero- characteristic arrow-like cut. mones that aggregate the population even fur- ther. These areas must be closely observed to Larvae feed during most of the day except prevent any further outbreaks. when temperatures are too high. A high popu- lation can be detected by observing the nume- Larvae are gregarious up to the fourth deve- rous excreta on the ground and the noise pro- lopmental stage and from there they are soli- duced when the droppings hit the vegetation tary. Initially they do not eat much and feed which sounds like a light rain. Larvae stop fe- from the borders of the leaflets. Upon rea- eding for about a day before and soon after ching the fourth stage, the larva chooses a lea- molting. When they are ready to pupate, most flet and feeds moving its head back and forth. of them move to the epiphytes growing on the This behavior makes a slanting cut on one side stem. Damage

A single larva may eat up to three leaflets Doing this exercise for leaf 17 in the phyllo- and most of this forage is consumed by the last taxy permits to know if the potential damage is stage: up to 76% (432 cm²) of the total con- below or above a given permitted defoliation. sumption to complete the whole larval stage Assuming that an adult oil palm produces (Table 2). about 6.250 cm² of new foliage every day, this could be considered a permitted daily defolia- According to Wood et al. (1973) the oil tion, at least for some time. The results of this palm can tolerate a defoliation of 6.25 % (or 17 type of exercise appear in Tables 2,3. %) of the youngest (or oldest) leaves. This lea- ves a margin to react when an increase in the The data do not consider the fact that larvae population of a defoliator is observed. This do not feed for some time before and after time will allow for an evaluation of the popula- molting. tion of the pest and the efficacy of natural con- trol, before active control measures are taken. It is normally accepted that active control measures have to be taken when damage exce- In order to have an idea of the potential da- eds the capacity of the palm to replace the leaf mage caused by a given number of larvae per area consumed by the pest in one day, and such leaf, this number is multiplied by the leaf area damage level should not exceed 20% (about that could be consumed by each larval stage. 1.160 cm²). However, other factors have to be

Table 2. Estimated of permited daily defoliation per leaf (theoretical) Daily defoliation cm²/leaf/day of Necessary number of first stage larvae to per leaf (%) defoliation cause a given percentage of defoliation 6.²/ 250cm day 156.²// 25cm leaf day None =156. 25 =679 40leaves 023.²/cm day 6.²/ 250cm day´ 5 312.²// 5cm leaf day 5% =312. 5 =1359. 100 023.²/cm day 6.²/ 250cm day´ 20 1250.²//cm leaf day 20% =1250. =5. 435 100 023.²/cm day It is assumed that an adult palm has 40 leaves. The exercise can be done for each larval stage and the corresponding daily defoliation that could be caused 17

considered such as palm age, the occurrence of natural enemies of the pest, weather condi- previous defoliations, and the abundance of tions, plant nutrition and other.

Table 3. Estimated number of larvae of O. Cassina per leaf that could be allowed without causing economical damage Estimated number of larvae necesary to Leaf area consumed (cm²) cause a theoretical defoliation of: Larva Total for the Daily larval stage % 0% 5% 20% 50% I 0.23 1.72 0.40 679 1.359 5.435 13.587 II 0.96 5.76 1.33 163 325 1.302 3.255 III 2.18 13.08 3.02 72 143 573 1.433 IV 10.10 82.82 19.20 15 31 124 309 V 24.35 328.72 76.05 6 13 51 128 Total 432.1 100.00 The data do not consider the fact that larvae do not feed for some time before and after molting

Sampling

An efficient sampling method done with entomo-pathogens). All stages of the life cycle the appropriate frequency will allow the early should be sampled (eggs, larvae, pupa and identification of a developing pest outbreak. adults). The information obtained will permit to act accordingly, reducing control costs and mini- In some commercial oil palm plantations in mizing damage to the plantation and the popu- Colombia, the pytosanitary people check the lation of natural enemies of the pest. It is advi- plantation every other week: first week every sable that all field personnel get involved in the other section and the rest the week after phytosanitary monitoring of the plantation, (chessboard faction) Genty (1985). This may particularly harvesting personnel (quality con- be necessary when outbreaks are imminent, trol and harvester themselves). These person- but this is not the situation in most of the plan- nel can give the first voice of alarm in case they tations in tropical America and visits are nor- observe some unusual behavior of a given pest, mally done less frequently. The risk of out- particularly on the cut leaves that are left breaks is reduced if the plantation is managed behind on the ground during bunch harves- from an agro-ecological perspective, which ting. The information is picked up by the means, among other things, given an adequate phytosanitary squat that then goes to the spot nutrition to the palm (opportune, in the right and takes the necessary information. amounts and balanced), keeping the soil well aerated and maintaining an adequate weed Sampling should not be done only under an control that favors the development of the be- outbreak situation in order to determine the neficial flora that harbors and feeds a healthy number of larvae per leaf that justify the use of population of predators and parasitoids of po- an insecticide, but should be a normal part of tential pests. the phytosanitary management of the planta- tion that permits to keep a eye on the evolution During increments of the population, a re- of all potential pests and their natural enemies. liable and fast sampling of O. cassina can be When an abnormal increase in the population done by counting larvae on the 80 distal lea- of a pest is observed, further observations are flets of leaf 17 (40 leaflets on each side of the done to follow the evolution of the pest and rachis) Rhainds et al. (1993). Most larvae (7.85 their natural enemies (predators, parasitoids, ± 0.71) were found on the abaxial portion of

ASD Oil Palm Papers, N° 36, 2011 18

the leaves compared with 1.60 ± 0.42 on the palms/ha represented a critical level of popu- upper portion, which may indicate a strategy to lation. However, the population on leaf 17 reduce depredation or desiccation. It was con- could give a better idea of the potential damage sidered that 7-10 large larvae on leaf 25 in 2-4 (Rhainds et al. 1993).

Natural Enemies

Some notable increments in the popula- Some Hymenoptera are important parasi- tions of some Pentatomidae (Alcaeorrhynchus toids of O. cassina. Telenomus (Scelionidae) and grandis Dallas, Mormidea sp., Podisus spp. and to a lesser extend Ooencyrtus (Encyrtidae) para- Proxys punctulatus (Duvois de Palisoth)) accom- site eggs. Cotesia (Braconidae) and Horismenus panied the population increments of O. cassina (Eulophidae) attack larvae; Conura (Chalcidi- in the central Pacific of Costa Rica in 1990. dae) and several tachinid flies parasite larvae These insects were common in the vegetation and pupae, and Brachymeria (Chalcididae) is growing on the ground (including the cover common on pupae. Horismenus and Brachymeria crop of Pueraria phaseoloides Bentham) where may also be hiper-parasitoids of Cotesia sp. and they preyed on larvae of several Lepidoptera, Conura spp., respectively (Table 4). including Anticarsia gemmatalis Hubner (Noc- tuidae) and Estigmene acrea Drury (Arctiidae). The population of all these and other para- sitoids depends on the availability of plants Under normal conditions, the individual with nectaries and flowers where the adult bugs are dispersed and are hard to spot, but forms feed and take the necessary energy to when there is a defoliator outbreak, they aggre- move around the plantation. This is particular- gate and several thousand of individuals may ly true for Cotesia sp. and Conura spp. that only be present within one single hectare. Pentato- fly for short distances compared with tachinic mid bugs are generalist predators and during a flies that can fly longer distances. defoliator (Lepidoptera) outbreak they, and some birds (Quiscalus mexicanus and Psaracolices Some entomopathogens may also reduce monctezuma, Icteriidae), can drastically reduce the pest population. In the case of O. cassina, the pest population since they eat large Paecilomyces sp. and a nuclear polyhedrosis vi- amounts of prey in a short time. rus may infect larvae.

Table 4. Biological regulation of the populations of Opsiphanes cassina in Central America Stage Regulation factor for the different stages Eggs Ooencyrtus sp.: 1-4 micro-wasps emerge/egg. For Telenomus sp.: 6-11 wasps/egg Conura sp. (L-P): 12-16 wasps/pupa Cotesia sp. (L-L): several dozens of wasps/larva Horismenus sp. (L-L): as above. Can be an hyper-parasitoid on Cotesia sp. Tachinidae n. i.(L-P): 4-8 flies/pupa Larvae Hemiptera bugs: Alcaeorrhynchus grandis, Mormidea sp., Podisus sp. Fungus: Paecilomyces sp. Virus Birds: Quiscalus mexicanus, Psaracolices Monctezuma Brachymeria sp: a wasp/pupa. Hyper-parasitoid of Conura spp. Pupa Predator bugs Adult Birds From Mexzón and Chinchilla (1996). L-L = parasitoid completes development in the larva; L- P = parasitoid on larva and emerges as an adult from pupa 19

Viruses are very specific and effective applied with sub-lethal doses of a insecticide (Genty and Mariau 1975; Luchini et al. 1984; such as a Deltametrin (i.e. DECIS 2.5.% E.C., Orellana 1986; Sipayung et al. 1989), and when 30 cc/ha) a synergistic effect is achieved.

Nectariferous Plantas

There is a large number of plants on which leuca Schlechtandlal and Cham (Cyperaceae) many adult forms of parasitoids of O. cassina and V. sycioides; and Conura sp. feed on A. spino- feed (Table 5). For example, Cotesia sp. visits sus, B. aculeata, C. tora, M. aspera, S. melaleuca, So-

Table 5. Relative abundance of families of insects (parasitoids on Opsiphanes cassina) on several plant species in Costa Rica* Plant species Tachinidae Braconidae Chalcididae Eulophidae Encyrtidae Scelionidae Ageratum conyzoides VF A A VF VF - Amarantus spinosus A C A C VF VF Baltimora recta VF C A C VF VF Byttneria aculeata -C AFFF Cassia guatemalensis -F C--- Cassia reticulata -C CCVFF Cassia tora VF C A VF VF VF Chamaesyce hirta AF AF- - Melanthera aspera * * -C CCCF Priva aspera -C FVF-- Scleria melaleuca * * VF C A C VF VF Senna stenocarpoides ** -C ACFF Solanum jamaicense FC AFFVF Spermacoce laevis VF F C F VF VF Triunfetta semitriloba -C AF-F Urena lobata ** AF AF- - Vitis sycioides AC AFCC Tachinidae: unidentified species (parasitoid on larvae); Braconidae: Cotesia sp. (parasitoid on larvae); Chalcididae: Conura spp. (parasitoid on larvae) Eulophidae: Horismenus sp. (parasitoid on larvae); Encyrtidae: Ooencyrtus sp. (parasitoid on eggs; Scelionidae: Telenomus sp. (parasitoid on eggs). Relative abundance: VF = Very few (1-9 individuals in 10 plants of the same species); F = few (1-4 individuals per plant); C = Common (5-15 individuals per plant); A = Abundant (more than 15 individuals per plant). * From Mexzón (1997). ** These species could be planted along roads and other open areas within the oil palm plantation.

Ageratum conyzoides L., Baltimora recta L (Astera- lanum jamaicense Miller (Solanaceae), Urena lo- ceae), Amaranthus spinosus L. (Amarantaceae), bata L. (Malvaceae) and V. Sycioides. Byttneria aculeata Jacquin (Sterculiaceae), Cassia tora L. (Leguminosae), and Vitis sycioides L. (Vi- taceae); Horismenus sp. feeds on B. aculeata, Cas- sia reticulata Willdenow (Leguminosae), C. tora, Melanthera aspera L. (Asteraceae), Scleria mela-

ASD Oil Palm Papers, N° 36, 2011 20

Chemical Control

Some 'biological' insecticides such as Dipel inhibitors of chitin synthesis (triflumuron, di- (Bacillus thuringiensis) have been successfully flubenzuron, etc.) have also been used, taken used to control Opsiphanes outbreaks, and at the necessary precautions to reduce the impact the same time, causing less damage to its natu- on the natural regulators of the pest. However, ral enemies and the oil palm pollinators. Biolo- contact insectides were used only on spot gical toxins (EVISECT, PADAN and others), applications (Table 6).

Table 6. Some insecticide products used during outbreaks of Opsiphanes cassina population* Group Genetic name Commercial name Doses/ha Biological toxin Cartap Padan *** 0.5 - 1.0 kg Biological insecticide Bacillus thurigiensis Dipel 0.8 - 2.5l Chitin synthesis inhibitors Triflumuron Alsystin**** 0.075 - 0.1 kg Teflubenzuron Dart*** Diflubenzuron Dimilin *** Pyrethoids Esfenvalerate Asana **** Cypermethrin Capture **** Deltamethrin Decis **** From Monge (1985). * This list cannot be taken as recommendations by the authors ** Only on spot applications (normally less than 20 ha), when the population of natural enemies is considered too low. Its use should be considered highly restricted *** Can be used with very low doses (sub-lethal) of pyrethroids **** Used with sub-lethal doses (0.025-0.03 l/ha).

Integrated Management

The program should be based on a systema- in order to have an idea about its ability to con- tic monitoring of the different life stages of the trol the pest by themselves without additional insect across the plantation. Such monitoring intervention. An integrated management pro- should be the responsibility of well trained gram for a defoliating pest like O. cassina personnel (pest scouts), but it is also a good should consider the following: idea that other field personnel get involved, in particular those working in quality harvesting 1. Sampling of the different stages of O. cas- control, since they walk large areas of the plan- sina and its natural enemies tation and can take a look to pruned leaves on a. All stages should be considered. Eggs are exami- the ground and observe if something unusual ned to determine the percentage of parasitism. is occurring with respect to a pest population. Normal eggs are white and transparent with two If this is the case, they can give the voice of rose bands indicating the presence of a healthy developing embryo. Parasitized eggs are black alarm to the phytosanitary personnel who will or orange indicating the presence of Telenomus take the matter from there. sp. or Ooencyrtus sp. b. Counting the number of each larval stage per Once an unusual increment in the popula- leaf. Larvae are counted on the last 80 terminal tion of a potential pest is detected, the person- leaflets in leaf number 17 nel evaluate the condition of natural enemies 21

c. Counting the number of predators like hemipte- amples are Urena lobata, Scleria melaleuca, Senna rous bugs (adults and immature individuals) on stenocarpoides and Melanthera aspera. During an leaf 17. The data are complemented counting all outbreak, weed control has to be done in such a individuals on the ground vegetation growing manner that no such plants are eliminated. For within the area of four contiguous palms. The this purpose, weed control personnel must be procedure is repeated at four sites per hectare trained recognizing such plants d. Gathering parasitized pupae and larvae to 4. Biological control obtain an estimate of parasitism. Parasitized lar- vae may show a package of small white cases a. Reproduction and release of predatory bugs below the body corresponding to Cotesia sp. or and parasitoid wasps Horismenus sp. pupae. Parasitized pupae of O. cas- 5. Lures and pheromones sina are dark and have a faulty smell. Horismenus sp may act as a primary parasitoid or a Cotesia sp. a. Placement of traps containing an attractant and hyperparasitoid. food bait for adults (Loría et al. 2000). These 2. Inmediate actions traps can be maintained at a low density all year around in the plantation, and upon an increase a. Delimitation of the area affected in adult captures, the amount of traps per area is b. Use of an appropriate fill-out form to write increased. A good trap is a plastic bag which is down the information obtained collapsed to reduce the opening to a narrow en- trance, so the alate adults can get in, but once in- c. Data processing and development of a strategy side, cannot leave. The bags contain pieces of to follow sugar cane on the bottom and a plastic bottle 3. Cultural aspects with holes to release the volatiles from a mix- ture of molasses and yeast. a. An area suffering recurrent attacks of any defo- b. Planting plant species that can feed other Lepi- liator must be subjected to a serious study to doptera in order to maintain a healthy popula- determine if agronomic conditions are appro- tion of depredatory bugs. A combination of M. priate for maintaining a healthy plant. Special aspera, Lantana camara L. and Stachytarpheta ja- attention should be given to soil aeration condi- maicensis Miller (Verbenaceae) can achieve this tions (any drainage or compaction problems, result. for example), nutritional unbalances (particu- larly high nitrogen contents in the leaves) 6. Chemical control b. Collect and eliminate pupae when possible a. Spot application of insecticides c. Management of the associated vegetation by al- b. Alternate applications of B.t., pyrethroids, nere- lowing the development of a varied flora capa- istoxin, chitin inhibitors, etc. ble of sustaining a healthy population of parasi- toids and predators. There is a large variety of c. Repeated applications within a restricted are to plants that can be planted in open areas. Some ex- break the pest life cycle

Literature

Chinchilla C.M. 1993. Fauna perjudicial en palma and observations on the behavior of the pest aceitera. ASD de Costa Rica, Costa Rica. pp. in oil palm. ASD Oil Palm Papers Nº 21: 1- 8. 13-20. Luchini F., Morin J.P., Rocha de Souza R.L., da De Vries P.J. 1987. The of Costa Rica Silva J.C. 1984. Perspectivas del uso de ento- and their natural history. Princeton University movirus para el combate de Sibine sp., defolia- Press, U.K. 327 p. dor de la palma aceitera en Pará. Pesquisa en Adamento Nº 23. 5 p. Genty P, Mariau D. 1975. Utilización de un ger- men entomopatógeno en la lucha contra Sibine Mexzón R.G. 1997. Malezas atractivas de la ento- fusca. Oléagineux 30 (8/9): 349-354. mofauna en los cultivos de palma aceitera y pejibaye. IV Congreso Costarricense de Ento- Loría R., Chinchilla C., Domínguez J., Mexzón R. mología. San José, Costa Rica (17-21 nov. 2000. An effective trap to capture adults of 1997). Opsiphanes cassina (Lepidoptera: Nymphalidae)

ASD Oil Palm Papers, N° 36, 2011 22

Mexzón R., Chinchilla C.M. 1996. Enemigos natu- Rhainds M., Chinchilla C.M., Castrillo G. 1993. rales de los artrópodos perjudiciales a la palma Desarrollo de un método de muestreo de las aceitera (Elaeis guineensis Jacq.) en América larvas de Opsiphanes cassina en palma aceitera. Central. ASD Oil Palm Papers Nº 13: 9-33. Manejo Integrado de Plagas Nº 30: 15-18. Monge V., L.A. 1985. Manejo racional de insectici- Sipayung A., Desmier de Chenon R., Sudharto P.S. das: resistencia y rotación. 1 edición. Editorial 1989. Recent work with viruses in the biologi- Tecnológica de Costa Rica, Cartago, Costa cal control of leaf-eating caterpillars in North Rica. Sumatra, Indonesia. pp. 285-293. In. Proc. PORIM Int. Palm Oil Development Conf. Su- Orellana F. 1986. Control biológico del defoliador kaini, J. et al., eds. PORIM, Malaysia. de la palma aceitera, Sibine fusca Stoll (Lep., Li- macodidae). INIAP, Bol. Divulgativo Nº 170. Est. Exp. Santo Domingo, Ecuador. 10 p.

Fig. 1. Above: adults of Opsiphanes cassina. Below left: characteristic leaf cut left by a fully grown larva while feeding. Right, below, pupa Arriba: adulto de Opsiphanes cassina. Izquierda abajo. Corte en bisel característico de las larvas en las hojas de las palmeras. Derecha abajo. Pupa 23

Fig. 2. Opsiphanes cassina larvae being attacked by a group of Pentatomid nymphs (left) and a parasitoid wasp (right) Larvas de Opsiphanes cassina atacadas por un grupo de chinches pentatómidos (izquierda) y una avispa parasitoide (derecha)

Fig. 3. Trap for attracting and capturing Opsiphanes cassina adults. The plastic bag has sugar cane pieces on the bottom and a plastic bottle with a mix of molasses and yeast Trampa para la captura de adultos de Opsiphanes cassina. La bolsa lleva caña de azúcar en el fondo y una botella plástica (que se amarra con un cordel a una base peciolar) con una mezcla de melaza y levadura

ASD Oil Palm Papers, N° 36, 2011