PROCEEDINGS of the ENTOMOLOGICAL SOCIETY of WASHINGTON

A NEW SPECIES OF RAZOWSKI (: : ) FROM GUATEMALA ATTACKING AVOCADOS (PERSEA AMERICANA) (LAURACEAE)

JOHN W. BROWN AND MARK S. HODDLE PROC. ENTOMOL. SOC. WASH. 112(1), 2010, pp. 10–21

A NEW SPECIES OF HISTURA RAZOWSKI (LEPIDOPTERA: TORTRICIDAE: POLYORTHINI) FROM GUATEMALA ATTACKING AVOCADOS (PERSEA AMERICANA) (LAURACEAE)

JOHN W. BROWN AND MARK S. HODDLE

(JWB) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, c/o National Museum of Natural History, Smithsonian Institution, Washington, DC 20013 U.S.A. (e-mail: john.brown@ars. usda.gov); (MSH) Department of Entomology, University of California, Riverside, CA 92521 U.S.A., and Center for Invasive Species Research, University of California, Riverside, CA 92521 U.S.A. (e-mail: [email protected])

Abstract.—Histura perseavora Brown, new species, from Guatemala, is described and illustrated. It is compared with Histura curvata (Meyrick) from Brazil and Histurodes costaricana Razowski from Costa Rica. All specimens of H. perseavora were reared from either fruit, fruit pedicels, or young green branches of avocados (Persea americana Mill.; Lauraceae) during efforts to identify lepidop- teran pests of this commodity in Guatemala. Coincidentally, we discovered museum specimens of H. costaricana reared from avocados in Costa Rica. We present a brief review of the fragmentary knowledge of the larval stages of Polyorthini, the tribe to which H. perseavora is assigned. Key Words: Histurodes, Central America, , pest species DOI: 10.4289.0013-8797.112.1.253.10

Avocado (Persea americana Mill.; 2007). In the U.S., avocados were Lauraceae) is a semi-deciduous tree introduced into Florida before 1850 and reaching a maximum height of about 20 into California before 1856. Commercial m. It is native to Mexico and Central varieties of avocados were introduced to America, ranging from the eastern and Chile from California in 1928. Other central highlands of Mexico south countries in South America with com- through Guatemala to the Pacific Coast mercial avocado industries include Bra- of Honduras, Nicaragua, and Costa Rica. zil, Peru, and Venezuela. Currently, U.S. There is evidence that avocado fruit has avocado fruit production is concentrated been used by indigenous peoples in primarily in California and to a much Mexico for nearly 10,000 years. The tree lesser extent in Florida and Hawaii. In was introduced into Indonesia around California the Hass cultivar is a high- 1750, Israel in 1908, and South Africa, value agricultural commodity and an New Zealand, and Australia in the late iconic crop readily identifiable with this 19th century (Index Fresh Avocado U.S. state (Hoddle et al. 2003). Fresh avocado fruit imports from * Accepted by Michael W. Gates Mexico and Central America into the VOLUME 112, NUMBER 1 11

U.S. became illegal under a federal ban and damaging California-grown avoca- imposed in 1918 to protect U.S. avoca- dos is perhaps more worrisome. Three do growers from the unwanted intro- of the last five folivorous pests to duction of specialist fruit-feeding in- become established on avocados in Cal- sects (e.g., tephritids, curculionids, and ifornia were all species new to science lepidopterans) that had evolved with at the time of first detection in the U.S. avocados in the home range of this plant This is remarkable given the long (Hoddle 2007a). However, after the history of avocado domestication and North American Free Trade Agreement commercialization in Mexico and Cen- (NAFTA) became law in 1994, barriers tral America (Hoddle 2004). to fresh avocado fruit exports were One of the most important pests lowered, allowing Mexico to legally attacking avocado fruit in Central and export avocados (mainly Hass variety) South America is the avocado seed to the U.S. Because of concerns about , catenifer Walsingham the potential introduction and establish- (Lepidoptera: Elachistidae), a species ment of unwanted invasive species, that is a specialist on members of the primarily fruit flies (Diptera: Tephriti- plant family Lauraceae. Larvae feeding dae), Mexican avocados initially were internally can cause premature fruit imported only into the northeastern U.S. drop and disfigurement of harvestable during winter. The rationale for this fruit, and production losses of 45–95% time-limited and highly geographic im- can result (e.g., Nava et al. 2005). Con- portation window was that semi-tropi- sequently within the home range of this cal fruit-feeding pests accidentally im- pest, intensive management with broad ported with fruit that escaped into the spectrum insecticides is practiced to environment would be unable to with- minimize economic losses despite sub- stand extreme winter cold, thereby stantial natural enemy activity against preventing the establishment of peren- eggs by trichogrammatid parasitoids nial populations. However, by the end and larvae by braconids and ichneumo- of 2005, Mexican avocados were grant- nids (Wysoki et al. 2002; Nava et al. ed entry into every U.S. state except 2005; Hoddle 2007a, b; Hoddle and California, Florida, and Hawaii which Hoddle 2008a, b). have commercial avocado industries. During survey work for S. catenifer On 1 February 2007, fresh avocados and other lepidopteran pests of avoca- were allowed to be exported from dos in Guatemala, a variety of micro- Mexico to California, the largest avo- lepidoptera were reared from field- cado producing and consuming state collected fruit. Among these was a in the U.S. High volumes of fruit large number of specimens of a tortricid (.130,000 tons) now are shipped an- moth, which inflicted considerable nually to all 50 U.S. states (Herald damage on avocado fruit but, only in Tribune 2007). orchards above about 1,450 m eleva- Given the enormous flow of avocado tion. After studying the material, it be- fruit into the U.S. from Mexico, there is came clear that these specimens repre- now the expectation that unwanted sented a new species in the tribe fruit-feeding pests eventually will es- Polyorthini. The generic assignment tablish in California. While the invasion was not immediately clear, and a review risk posed by known fruit pests is of morphological features of Histura identifiable and possibly can be miti- Razowski and Histurodes Razowski gated to a certain extent, the threat of suggested that the two genera may previously unknown species entering not be clearly defined and/or differ- 12 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON entiated. A comparison of larval fea- under organic management in Santa tures of the new species with those Lucia Miplas Altas, Sacatepe´quez, Gua- previously described for other Poly- temala (14835.6470N; 90840.0270W, orthini revealed a high degree of 2275 m elevation). This orchard was variation in the chaetotaxy, suggesting ;3 Ha consisting of ;300 10–12-year- that no features of the larvae alone can old trees that were ;6–8 m in height be used to unambiguously assign them with interlacing canopies. On 25 and 30 to this tribe. November 2007, 11 and 27 fruit, The purpose of this paper is to respectively, showing lepidopteran describe and illustrate this new species feeding damage were harvested at this that feeds on avocado, which may have site and returned to the laboratory for the potential to become a significant rearing. avocado pest in areas outside of its The new species of Histura also was natural range. In addition, information reared from unmanaged non-Hass fruit. is presented on its biology, and the On 11 December 2006, 354 fruit were fragmentary information on the larval purchased from a roadside vendor in stages of Polyorthini is reviewed. Santiago Atitlan, Solola´, Guatemala (14837.5810N; 91813.5570W, 1,592 m MATERIALS AND METHODS elevation). These fruit had been har- Study sites.—Avocado fruit used in vested from backyard trees by the this study was collected from commer- vendor in the immediate vicinity of cial Hass avocado orchards and from the purchase site. The second site with non-Hass trees that were growing along non-Hass avocados that yielded the roadsides, in backyards, or as shade new species was in Sumpango, Saca- trees in coffee plantations. Fruit collec- tepe´quez, Guatemala (14840.2920N; tions were part of a much larger survey 90843.1950W, 1,825 m elevation). Over being conducted for fruit-feeding Lep- the period 14–23 November 2007, 81 idoptera in Guatemala (Hoddle and fruit were harvested from one tree at Hoddle 2008a). The new species de- this site. scribed herein was reared from fruit Rearing methods.—Hass and non- collected from two commercial Hass Hass avocados were returned to the orchards. The first orchard was located laboratory and kept for 10–14 days in in San Miguel Duen˜as, Sacatepe´quez, collapsible ventilated rearing Guatemala (14831.4610N; 90846.5790W, cages (Bug-Dorm-2120 [60 cm 3 60 1,494 m elevation). This orchard was cm 3 60 cm], MegaView, Taiwan) that ;5 Ha consisting of ;800 trees; trees were labeled by collecting date and were ;5 years old and ;4–5 m in location. Rearing of larvae was con- height, and canopies were separated by ducted in a well-lit room under natural ;5–7 m of clear ground allowing full daylight and lengths (;12 h) at 22.148C sun exposure. Trees were subjected to 6 0.13 (temperature measured with a monthly rotating applications of mala- Hobo data logger [Onsett Corp., MA] at thion and endosulfan. Over the study 30 min intervals). Fruit were inspected period (24 January–23 February 2007), daily, and mature Lepidoptera larvae 2080 fruit showing damage (e.g., frass searching for pupation sites were iso- accumulations, perseitol exudates, and lated and kept in clear labeled plastic tunnel openings) from Lepidoptera cups with ventilated lids that were lined feeding were picked from trees and with paper towel to provide a pupation returned to the laboratory for rearing of substrate. were kept along with pests. The second Hass orchard was pupal cases from which they emerged; VOLUME 112, NUMBER 1 13 in most cases the exuvium is mounted vided) valva in which the abdominal on a small card beneath the pinned hairpencils from segment seven reside, specimen. Some last instars were pre- raised scales on the forewing of both served in 95% ethanol for subsequent sexes, and a finely spined anellus in the morphological examination and possi- male genitalia. Details of the muscula- ble DNA study. ture of the male genitalia are presented Systematic methods.—Dissection by Razowski (1979b), and what is methods for examining genitalia follow known of the larval stages, which are those presented in Brown and Powell poorly studied, are summarized by (1991, 2000). Dissected genitalia were Horak and Brown (1991). illustrated with the aid of a micropro- Seven genera of Polyorthini occur in jector. Terminology for genitalia struc- the New World tropics. Among these, tures follows Horak (1984); terminolo- Histura Razowski (eight species) and gy for forewing pattern elements Histurodes Razowski (two species) are follows Razowski (2003); and terminol- extremely similar in facies and genita- ogy for larval features follows Brown lia. In overall appearance (e.g., fore- (1987). wing shape, forewing pattern, length of Abbreviations of depositories are as labial palpi), they are more similar to follows: BMNH, The Natural History the Old World Lopharcha Diakonoff Museum, London, U.K.; INBio, Insti- and Lowerthantothe tuto Nacional de Biodiversidad, Santo Neotropical Polyortha Dognin, Ardeu- Domingo de Heredia, Costa Rica; UCR, tica Meyrick, Lypothora Razowski, or University of California, Riverside, Pseudatteria Walsingham. Adults of California, U.S.A.; and USNM, Nation- the aforementioned Neotropical gen- al Museum of Natural History, Smith- era have a distinctive forewing shape, sonian Institution, Washington, DC, marked polymorphism, and extremely U.S.A. long, porrect labial palpi, three or more times the diameter of the compound RESULTS eye. Histura and Histurodes possess Polyorthini, comprised of about 150 none of these features. species in 20 genera, was proposed by In his description of Histurodes, Obraztsov (1966); its classification was Razowski (1984) indicated that the developed by Diakonoff (1974) and transtilla is absent. However, we inter- revised by Razowski (1979a, b, 1981). pret the transtilla as present, represented The monophyly of the group is well by a pair of blunt appendages weakly supported (Razowski 1979a, 1981), and united mesally by the spiny anellus. A it is generally accepted that Polyorthini similar character state also is present in is the sister-group to þ Histura bicornigera Razowski. Hence, Chlidanotini (Tuck 1981, Horak and the distinction between Histura and Brown 1991, Poinar and Brown 1993, Histurodes may not be particularly well Horak 1999). These three tribes com- defined. However, the synonymy of prise the tortricid subfamily Chlidano- these two genera can be determined tinae. Polyorthini are mostly pan-trop- most convincingly through a phyloge- ical in distribution, but two genera, netic anaylsis that includes all described Guene´e and Meyrick, species, which is well beyond the scope are restricted to temperate Europe of the current work. The new species (Razowski 1979b). described below is assigned provision- Males of all Polyorthini have a ally to Histura on the basis of the unique ‘‘split’’ (i.e., longitudinally di- complete transtilla. 14 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1–6. Histura perseavora. 1, Adult moth. 2, Male genitalia—uncus, socii, and gnathos (lateral view). 3, Male genitalia (valvae spread, aedeagus removed). 4, Aedeagus. 5, Female genitalia (unmated female). 6, Ductus bursae and corpus bursae of mated female.

Histura perseavora Brown, socii, and the patch of spines of the new species aedeagus. The last character appears to Figs. 1–13, 17–19 represent an autapomorphy for the new species. Diagnosis.—Histura perseavora is Description.—Adult (Figs. 1, 19). Histurodes superficially most similar to Head: Vertex and frons dark gray costaricana Razowski and Histura cu- brown, paler on lower frons; labial prata (Meyrick), with short labial palpi palpus with length approximately 1.23 and a well-defined, two-toned forewing diameter of compound eye, mostly dark pattern with a dark proximal one-third gray brown dorsally, lighter on inner and a paler distal two-thirds (Fig. 1). The surface and ventrally, third segment ex- male genitalia of H. perseavora (Figs. 2– tending beyond scaling of second; ocel- 4) differ from those of H. cuprata by the lus large; antenna extending to ca. mid- medially narrowed transtilla and the dle of forewing costa, with two rows of presence of a linear patch of long spines scales per flagellomere. Thorax: Dor- in the distal half of the aedeagus. They sum dark gray brown, tegula concolo- differ from those of H. costaricana by rous with dorsum; scales at posterior their shorter, rounder valva, slenderer end of dorsum conspicuously flattened. VOLUME 112, NUMBER 1 15

Legs unmodified. Forewing length 5.0– sclerotized and irregular angled apical- 5.5 mm (n ¼ 15); pattern essentially ly; with linear patch of spines from ca. two-toned with darker proximal one- 0.60–0.75 distance from base, spines third distinguished from lighter distal increasing in size apically. Female two-thirds by nearly straight oblique genitalia (Figs. 5–6) with ovipositor line from costa ca. 0.3 distance from narrow, somewhat telescopic; papillae base, to dorsum ca. 0.45 distance from anales narrow; sterigma weakly sclero- base (Fig. 1); basal area nearly uniform tized, poorly developed; ostium anteriad dark gray brown; distal area pale copper and dorsad of posterior edge of segment with irregular darker scales and ill- seven (i.e., situated in slightly invagi- defined pale red-brown blotch in sub- nated intersegmental membrane); duc- terminal area; a dark spot at end of tus bursae uniform in width (in unmated discal cell; three oblique linear patches females) (Fig. 5) or gradually expanded of raised scales, one immediately basad anteriorly (Fig. 6); corpus bursae not of oblique line forming outer edge of differentiated from ductus busae, with- basal patch (interrupted near middle), out signa; ductus seminalis from ductus one at two-thirds distance from base to bursae near ostium. termen, and one about midway between Larva.—Based on three presumed two aforementioned; fringe pale copper last instars. General (Figs. 7–11, 17): to gray brown. Hind wing dingy grayish Length 6–7 mm; body color cream pink white; fringe concolorous with hind with moderately large, conspicuous, wing. Abdomen: Male with long paired dark brown pinacula; most of integu- hairpencils from invaginated pouches of ment covered with short dark spinules, segment seven. Male genitalia (Figs. 2– absent only from pinacula. Head: Am- 4) with tegumen rectangular; vinculum ber with small dark stemmatal patch. large, with rounded dilation at junction Thorax (Fig. 8): Prothoracic shield dark with tegumen (i.e., near point of brown with slender, pale dividing line at attachment of costa of valva); saccus well developed, short, rounded; uncus mid-dorsum; L-group on T1 trisetose, moderately long, strongly sclerotized, setae arranged nearly parallel to longi- slightly curved ventro-posteriorly, trun- tudinal axis of larva; spiracle round, cate-rounded apically; socii slender, relatively small, with strongly sclero- digitate, ca. as long as uncus, with tized perimeter; SV-group on T1, T2, short, fine, hairlike scales; gnathos arms and T3 2:1:1; T2 and T3 with extra unmodified, united distally into narrow, setae: MD1, MSD1, MSD2. Abdomen terminal process ca. as long as basal (Figs. 9–11): Spiracles small, round, portion of gnathos arms, with upturned slightly larger on A8; L1 and L2 on tip; transtilla moderately broad with shared pinaculum below spiracle; tiny deep rounded excavation at middle SD2 without pinaculum, separate from (transtilla frequently broken at middle large SD1 pinaculum on A1–8; SD1 on in slide mounted preparations owing to A8 dorsoanterad of spiracle; SV-group extremely narrow junction of basal on A1,2,7,8,9 2(3):3:3:2:2; D2s on halves); juxta large, U-shaped; valva shared dorsal pinaculum on A9; D1 broad, rounded, split longitudinally, and SD1 on shared pinaculum on A9; sparsely hairy throughout, with hairs L-group trisetose on A9; crochets bi- most dense in subbasal area; sacculus ordinal in complete circle; Vs about broad, well defined basally, ill defined same distance apart on A7, A8, and A9; distally, without free projection. Aedea- anal shield rounded; abdominal prolegs gus (Fig. 4) slightly curved near middle, with 29–32 crochets, A10 proleg with 16 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 7–11. Early stages of Histura perseavora. 7, Last instar. 8, Larval chaetotaxy, head, and thoracic segments T1, T2, T3. 9, Larval chaetotaxy, abdominal segments A1 and A2. 10, Larval chaetotaxy, abdominal segments A3 and A4. 11, Larval chaetotaxy, abdominal segments A7–A10.

20–22 crochets; anal fork broad, fan- of pupa with three pairs of long, shaped, with 7–8 slender, pointed tines. distally-hooked setae; cremaster not Pupa.—Based on 10 exuviae and 2 developed. preserved pupae. Typically tortricoid, Biology.—The larvae of H. persea- fusiform, 5–6 mm in length, 1.0–1.5 vora attack fruit and fruit pedicels of mm in width (Fig. 12–13, 18). Head avocados, and when fruit is scarce, and thorax typical for family as de- larvae mine succulent young branches scribed elsewhere (e.g., Horak 1999), with flush growth. Typical damage is lacking cephalic projection and con- illustrated in Figs. 14–16. The larval spicuous sculpturing. Abdomen with stages are superficially similar to those A1 lacking dorsal spines; A2–A7 with of in both appearance two rows of strong spines; A8 with and damage. These two pests are sym- single row of spines in most females patric at elevations between ;1,500 and and two rows in most males; A9 with 1,600 m, but only H. perseavora has single row in both sexes; posterior end been recorded above 1,800 m where S. VOLUME 112, NUMBER 1 17

the reared larvae were H. perseavora, the remainder were almost all S. catenifer. A single species of parasitoid was reared from larvae of H. perseavora,a Chelonus sp. (Hymenoptera: Braconi- dae: Cheloninae) (UCRC ENT 169534). We examined four specimens of Histurodes costaricana, including the type series (n ¼ 2), and all were reared from avocados in Costa Rica (USNM, INBio). Native hosts for species of Polyorthini are exceedingly poorly doc- umented and totally unknown for all other species of Histura and Histurodes. However, based on these limited data, there appears to be a tendency for species of these two genera to move Figs. 12–13. Pupa of Histura perseavora. onto cultivated Lauraceae when the 12, Ventral view. 13, Lateral view. opportunity presents itself. Given the intensity of sampling for Lepidoptera in catenifera is uncommon or absent. All Costa Rica, it is remarkable that larvae (;16) reared from avocados additional examples of H. costaricana collected at ;1,800 were H. perseavora. are not present in the collection of At elevations below 1,600 m, ,5% of INBio, suggesting that adults may not

Figs. 14–19. Histura perseavora. 14, Damage to avocado fruit. 15, Damage to avocado fruit. 16, Entrance hole of larva into avocado fruit. 17, Larva. 18. Pupa. 19, Adult. 18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON be attracted to light traps, the primary DISCUSSION method of sampling. Information on the early stages of Other tortricids reared from avocado Polyorthini is either lacking or at best during the Guatemala surveys include fragmentary and dispersed in the liter- Cryptaspasma sp. (Olethreutinae: Mi- ature. Fletcher (1921) provided superfi- crocorsini), which likely represents the cial descriptions and illustrations of the same species reported by Brown and Asian Polylopha epidesma Lower. Brown (2004) from avocados in Mi- Swatschek (1958) provided a descrip- choacan, Mexico; Amorbia santamaria tion of the larva of the European Phillips and Powell (Tortricinae: Spar- (Fabricius). ganothini), a polyphagous leafroller Dugdale (1966) presented a description recently described from Guatemala and illustration of the larva and a few (Phillips-Rodriguez and Powell 2007); details of the pupa of Lopharcha Netechma pyrodelta (Meyrick) (Tortri- insolita (Dugdale) from New Zealand. cinae: Euliini), representing the first Becker (1970) provided descriptions host record for this Neotropical and illustrations of the larva and pupa (Hoddle and Hoddle 2008a); and Apo- of Polylopha viridescens Meyrick from tomops n. sp. (Tortricidae: Euliini), a genus which has been recorded previ- Brazil. We have examined preserved ously only from Pinaceae, i.e., A. well- larvae of P. viridescens and H. persea- ingtoniana (Kearfott) on Abies and vora, preserved pupae of P. viridescens, Tsuga (Prentice 1966, Brown and and pupae exuviae of H. perseavora and Passoa 1998). P. epidesma. Our observations, in com- Distribution.—Histura perseavora is bination with descriptions and illustra- known only from Hass avocado or- tions from the literature, present a chards and non-Hass avocados growing bewildering array of character states at elevations of ;1,500 to 1,800 m in that allow few generalizations about the Guatemala. immature stages of Polyorthini. Holotype ?. Guatemala, Solola´ Prov- The SV-group on the thorax of the ince, San Lucas Toliman, near San- larva of nearly all tortricids is 2:1:1 (T1, tiago Atitlan, 1,585 m, 1483703100N, T2, T3, respectively). However, Becker 918804300W, ex non-Hass avocado fruit, (1970) reported the SV-group as 2:2:2 XII.2006-III.2007, M. & C. Hoddle in P. viridescens, possibly representing (USNM). the ancestral condition for the family Paratypes (8?,8/). Guatemala, Solo- based on the hypothesis that derived la´ Province, San Lucas Toliman, near conditions in larval chaetotaxy are Santiago Atitlan, 1,585 m, 1483703100N, manifested as losses (Horak and Brown 918804300W, ex avocado fruit, XII.2006- 1991). An examination of larvae in the III.2007 (5?,6/), M. & C. Hoddle. USNM confirms this character state for Sacatepe´quez: Finca Miguel Urias, San P. viridescens. All other Polyorthini Miguel Duen˜as, 1,495 m, 1483102700N, either examined by us or reported in 9084603400W, ex avocado fruit, XII.2006- the literature have a thoracic SV-group III.2007 (3?,2/), M. & C. Hoddle. of 2:1:1. Paratypes distributed among USNM, The SV-group on abdominal segments UCR, and BMNH. 1,2,7,8,9 is 3:3:3:2:2 in most species of Etymology.—The specific epithet is Archipini and Sparganothini and re- derived from the host genus, Persea, duced in species of most other groups: and ‘‘voro,’’ which is Latin for ‘‘eat usually 3:3:2:2:2 in Cnephasiini and greedily.’’ It is considered an adjective. Tortricini (Swatschek 1958, MacKay VOLUME 112, NUMBER 1 19

1962), variably reduced in nearly all orthini larvae we investigated are a Olethreutinae (Swatschek 1958, Mac- trisetose L-group on A9 (common to Kay 1962), and unusually multisetose in nearly all tortricids), a conspicuously Chlidanotini (Diakonoff and Arita spiny integument (common to many 1981). The SV-group is 3:3:3:3:2 in P. tortricids), and the presence of an anal viridescens (Becker 1970, confirmed by comb (ancestral for tortricids, but examination of material), which is aber- secondarily lost in many internal feed- rant, as is the thoracic SV-group de- ers). However, the anal comb is highly scribed above. It is 2(3):3:3:2:2 in H. modified in Lopharcha (Dugdale 1966), perseavora (reported here) and O. schu- and its presence/absence was not re- macherana (Swastchek 1958); 2:3:3:2:2 ported in P. cassiicola (Liu and Kawabe in L. insolita (Dugdale 1966); and 1993). In summary, none of the larval 2:3:3:2:1 in Polylopha cassiicola Liu features we examined provide evidence and Kawabe (Liu and Kawabe 1993). for the relationship of Polyorthini to any In general, the tiny SD2 seta is on the other major tortricid group, and most same pinaculum as the much larger SD1 are highly variable within the tribe. in all species of Archipini and Sparga- nothini, but the two setae are on ACKNOWLEDGMENTS separate pinacula (or the SD2 pinacu- lum greatly reduced and often incon- Illustrations of genitalia were ren- spicuous) in all species of Cnephasiini dered by David Adamski, USDA, ARS, and Tortricini, and most species of Systematic Entomology Laboratory, Olethreutinae (Swatschek 1958, Mac- and those of the larva and pupa by Kay 1962). In all species of Polyorthini Diana Marquez; illustrations were ar- but O. schumacherana, the setae are on ranged into plates by Mike Lewis, separate pinacula (similar to that found Department of Entomology, University in species of Cnephasiini, Tortricini, of California, Riverside; and the para- and Olethreutinae). Liu and Kawabe sitoid was identified by Roger Burks, (1993) reported that the SD2 seta is Department of Entomology, University absent in P. cassiicola, but we suspect of California, Riverside. We thank the that the seta may be present but in- following for providing helpful reviews conspicuous owing to the absence of a of the manuscript: Robert Kula and distinct pinaculum and the tiny size of Thomas Henry, USDA, Systematic En- the seta. tomology Laboratory, National Muse- In general, most tortricines have setae um of Natural History, Washington, D1 and SD1 on separate pinacula on A9, DC, U.S.A.; Joaquin Baixeras, Institut whereas most olethreutines (and species Cavanilles de Biodiversitat i Biologia of Hilarographini) have them on a Evolutiva, Universitat de Valencia, shared pinaculum. However, there is Spain; and Richard Brown, Mississippi substantial variation in this character in State University, Starkville, Mississippi, both subfamilies. In L. insolita (Dugdale U.S.A. Funding for field work in 1966), P. cassiicola (Liu and Kawabe Guatemala was provided in part by the 1993), and H. perseavora, D1 and SD1 California Avocado Commission. are on a shared pinaculum (as in most olethreutines), whereas in P. viridescens LITERATURE CITED and Olindia the setae are on separate Becker, V. 1970. Sobre duas especies Brazileiras pinacula on A9 (as in most tortricines). do genero Polyortha (Lepidoptera, Tortrici- Three larval character states that dae). Bulletin Universidad Federal de Parana´, appear to be consistent among Poly- Conselho de Pesquisa, Zoologia 4: 1–11. 20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Brown, J. W. and R. L. Brown. 2004. A new ———. 2007b. The avocado seed moth, Stenoma species of Cryptaspasma (Lepidoptera: Tor- catenifer Walsingham (Lepidoptera: Elachis- tricidae: Olethreutinae) from Central Amer- tidae). Website: http://www.biolcontrol.ucr. ica, the Caribbean, and southeastern United edu/Stenoma/Stenoma.html (accessed 1 States, with a catalog of the world fauna of April 2008). Microcorsini. Proceedings of the Entomo- Hoddle, M. S. and C. D. Hoddle. 2008a. Survey logical Society of Washington 106: results for Lepidoptera and associated para- 288–297. sitoids attacking Hass and non-Hass avoca- Brown, J. W. and S. Passoa. 1998. Larval dos in Guatemala. Journal of Economic foodplants of Euliini (Lepidoptera: Tortrici- Entomology 101: 1310–1316. dae): From Abies to Vitis. Pan-Pacific ———. 2008b. Bioecology of Stenoma catenifer Entomologist 74: 1–11. Walsingham (Lepidoptera: Elachistidae) and Brown, J. W. and J. A. Powell. 1991. Systematics associated larval parasitoids reared from of the Chrysoxena group of genera (Lepi- Haas avocados in Guatemala. Journal of doptera: Tortricidae: Euliini). University of Economic Entomology 101: 692–698. California Publications in Entomology 111. Hoddle, M. S., K. M. Jetter, and J. G. Morse. 87 pp. þ figs. 2003. The economic impact of Scirtothrips ———. 2000. Systematics of Anopina Obraztsov perseae Nakahara (Thysanoptera: Thripidae) (Lepidoptera: Tortricidae: Euliini). Universi- on California avocado production. Crop ty of California Publications in Entomology Protection 22: 485–493. 120. 128 pp. þ figs. Horak, M. 1984. Assessment of taxonomically Brown, R. L. 1987. Tortricidae (Tortricoidea), pp. significant structures in Tortricinae (Lep., 419–433. In F. W. Stehr, ed. Immature Tortricidae). Mitteilungen der Schweizeri- , volume 1. Kendall/Hunt, Dubuque, schen Entomologischen Gesellschaft 57: Iowa. 3–64. Diakonoff, A. 1974. The South Asiatic Poly- ———. 1999. Tortricoidea, pp. 199–215. In N. orthini with notes on species of Polyortha Kristensen, ed. Lepidoptera, moths and Dognin (Lepidoptera: Tortricidae). Zoologi- butterflies. Volume 1: Evolution, systematics, sche Verhangelingen 131: 1–86. and biogeography. Handbook of Zoology 4 Diakonoff, A. and Y. Arita. 1981. The early (35), Arthropoda: Insecta. Walter de Gruyter, stages of Thaumatographa eremnotorna Dia- Berlin & New York. konoff & Arita, with remarks on the status of Horak, M. and R. L. Brown. 1991. 1.2 the Hilarographini (Lepidoptera: Tortricoi- and phylogeny, pp. 23–48. In L. P. S. van der dea). Entomologische Berichten 41: 56–60. Geest and H. H. Evenhuis, eds. Tortricid Dugdale, J. D. 1966. A revision of the New pests, their biology, natural enemies and Zealand Schoenotenini and Cnephasiini control. Elsevier Science Publishers B. V., (Lepidoptera: Tortricidae: Tortricinae). New Amsterdam. Zealand Journal of Science 9: 731–775. Index Fresh Avocado. 2007. Website: http:// Fletcher, T. B. 1921. Life histories of Indian www.indexfresh.com/avocado_history.htm. insects, Microlepidoptera. Memoirs of the (accessed 19 March 2008). Department of Agriculture of India, Ento- Liu, Y.-Q. and A. Kawabe. 1993. A new species mological Series, volume 6(1920): 1–217. of the genus Polylopha injurious to cinnamon Herald Tribune. 2007. Mexico praises lifting of in China (Tortricidae, Chlidanotinae, Poly- last U.S. avocado import barriers. Website: orthini). Japan Heterocerists’ Journal 173: http://www.iht.com/articles/ap/2007/02/03/ 404–406. business/ MacKay, M. R. 1962. Larvae of the North LA-FIN-ECO-Mexico-US-Avocados.php. American Tortricinae (Lepidoptera: Tortrici- Published 2 February 2007. nae). Canadian Entomologist Supplement 28. Hoddle, M. S. 2004. Invasions of leaf feeding 182 pp. : why are so many new pests Nava, D. E., J. R. P. Parra, V. A. Costa, T. M. attacking California-grown avocados? Cali- Guerra, and F. L. Consoli. 2005. Population fornia Avocado Society Yearbook 87: 65–81. dynamics of Stenoma catenifer Walsingham ———. 2007a. Lurkers on the threshold: poten- (Lepidoptera: Elachistidae) and related larval tial new avocado fruit pests for California parasitoids in Minas Gerais, Brazil. Florida avocados. California Avocado Society Year- Entomologist 88: 441–446. book 89: 69–92. Obraztsov, N. S. 1966. Neotropical microlepi- VOLUME 112, NUMBER 1 21

doptera, IX. Revision of genus Pseudatteria ———. 1981. Notes on the system of Polyorthini (Lepidoptera: Tortricidae). Proceedings of (Lepiodoptera, Tortricidae). Acta Zoologica the U.S. National Museum 118: 577–622. Cracoviensia 25: 309–318. Phillips-Rodriguez, E. and J. A. Powell. 2007. ———. 2003. Reassessment of forewing pattern Phylogenetic relationships, systematics, and elements in Tortricidae (Lepidoptera). Acta biology of the species of Amorbia Clemens Zoologica Cracoviensia 46: 269–275. (Lepidoptera: Tortricidae: Sparganothini). Swatschek, B. 1958. Die larval systematik der Zootaxa 1670. 109 pp. wickler (Tortricidae and Carposinidae) aus Poinar, G. O. and J. W. Brown. 1993. A new dem zoologischen Institut der Universitat fossil tortricid (Lepidoptera: Tortricidae) Erlangen. Akademie-Verlag, Berlin. 269 pp. from Dominican amber. Entomologica Scan- [Abhandlungen zur larvalsystematik Insekt- dinavica 23: 25–29. en 3.] Prentice, R. M. 1966. Vol. 4. Microlepidoptera. Tuck, K. R. 1981. A new genus of Chlidanotini In: Forest Lepidoptera of Canada recorded (Lepidoptera: Tortricidae) from New Cale- by the Forest Insect Survey. Department of donia, with a key to the genera and checklist Forestry Canada, Publication 1142(1965): of species. Systematic Entomology 6: 543–840. 337–346. Razowski, J. 1979a. On the morphology and Wysoki, M., M. A. Van den Berg, G. Ish-Am, S. system of Polyorthini (Lepiodoptera, Tortri- Gazit, J. E. Pen˜a, and G. K. Waite. 2002. cidae). Bulletin de l’Academie Polonaise des Pests and pollinators of avocado, pp. 223– Sciences, Series des Sciences Biologiques, 293. In J. Pen˜a, J. L. Sharp, and M. Wysoki, 26(1978): 857–862. eds. Tropical Fruit Pests and Pollinators, ———. 1979b. The systematic position of Olindia Biology, Economic Importance, Natural En- and Isotrias (Lepiodoptera, Tortricidae). Zo- emies and Control. CABI Publishing, Wall- ologisches Mededelingen 54: 241–243. ingford, Oxfordshire, U.K.