Cryptaspasma (Cryptaspasma) perseana Gilligan and Brown

Gilligan, Todd M., Brown, John W. & Hoddle, Mark S., 2011, A new avocado pest in Central America (Lepidoptera: Tortricidae) with a key to Lepidoptera larvae threatening avocados in California, Zootaxa 3137, pp. 31-45 : 32-38

Cryptaspasma (Cryptaspasma) perseana Gilligan and Brown , new species

( Figs. 1–27 )

Systematics. Cryptaspasma perseana is assigned to the subgenus Cryptaspasma based on the following combination of characters ( Aarvik 2005): uncus with hair-pencil; valva not swollen; pulvinus without spike(s); sacculus of valva without triangular, pointed prominence; and outer edge of valva not emarginate, thus sacculus and cucullus undifferentiated.

Diagnosis. Cryptaspasma perseana is separated from other species in the subgenus Cryptaspasma by the following characters: C. bipenicilla Brown and Brown has two lengths of coremata on the male abdomen versus one length in C. perseana ; C. lugubris (Felder & Rogenhofer) and C. acrolophoides Meyrick males have an uncus that is parallel-sided versus widened above the base and tapering distally in C. perseana ; C. athymopis Diakonoff and C. microloga Diakonoff females have signa that taper to a dull point versus signa that are distally rounded in C. perseana . Other undescribed males in the subgenus Cryptaspasma from Central and South America differ from C.

perseana in having either two lengths of coremata, setae absent from the base of the valva and/or sex scaling on the male hindwing; females of undescribed species generally lack the ventrolateral triangular processes on the sterigma and/or have differently shaped signa.

Description. Adult. Male. Head ( Fig. 7 ): Vertex rough scaled, dark brown intermixed with golden-brown; upper portion of frons rough scaled, concolorous with vertex, lower portion of frons smooth scaled, dark brown to black intermixed with orange-brown; labial palpus porrect, all segments combined ca. 1.3 times diameter of compound eye, first and second segments golden brown, second segment enlarged, third segment rounded, ca. 0.3 as long as second segment, dark brown distally; antenna ca. 0.4 as long as forewing, dark brown basally, brown apically, scape dark brown to black, sensory cilia ca. 0.2 times width of flagellomere; ocellus conspicuous. Thorax ( Fig. 1 ): Dorsal and ventral surfaces brown; tegula covered with long pale-brown scales; fore leg densely covered with dark brown scales; mid- and hind femur with long broad brown scales on ventral surface; mid- and hind tibia with long hairlike brown scales on dorsal and ventral surfaces; metathorax with posterior pair of scale tufts composed of long thin pale brown scales. Forewing length 9.8–10.9 mm (mean = 10.3, n = 10); costal fold absent; costal margin weakly convex; ground color pale brown intermixed with black and grayish purple; costal strigulae present as paired or single pale brown marks, strigulae 3–9 paired or unpaired on costa, strigula 10 unpaired between R4 and R5 on apex; conspicuous orange spot between bases of M1 and M2; fasciae dark brown to black, poorly defined; basal fascia expressed as black scaling at base along dorsum; subbasal and median fasciae coalesced to form broad band of black scaling extending from costa to dorsum across discal cell; postmedian and preterminal fasciae undefined, apical half of wing variably mottled with dark brown and grayish-purple; fringe brown. Hindwing: Uniformly brown; fringe brown with lighter apices. Abdomen ( Figs. 3, 5–6, 8–10 ): Brown to pale brown; eighth abdominal tergite with posterior pair of coremata ( Figs. 6, 8–10 ) consisting of a single tuft of scales composed as follows: outer scales elongate and pale brown, middle scales rhopaloid and dark brown, inner scales larger, rhopaloid, and covered with secretions; eighth abdominal sternite ( Fig. 5 ) tapered distally with large mesal excavation forming two distal lobes. Genitalia ( Fig. 3 ) with uncus variably elongate, weakly widened above base and tapered towards apex with tuft of setae extending from distal half, setal tuft approximately same length as uncus; socii a pair of setose arched lobes laterad to uncus; gnathos a narrow band with triangular subscaphium; tegumen narrow; pedunculus a triangular lobe; valva triangular, covered in long thin setae concentrated at base and rounded apex; a row of small pointed spinelike setae running entire length of saccular margin projecting ventrally; sacculus with several rows of long blunt spinelike setae ca. 0.50–0.85 distance from base to apex of valva; median portion of valva with weakly sclerotized pulvinus densely covered in long rhopaloid scales; phallus ca. 0.50 length of valva, broad, tapering weakly to blunt apex; cornuti absent.

Female. Head: As in male. Thorax ( Fig. 2 ): As in male except: Forewing length 10.5–12.1 mm (mean = 11.3, n = 9); costal margin weakly convex; ground color grayish brown; costal strigulae subdued, remnants of strigulae 5– 9 visible in some individuals; conspicuous white and orangish-brown spot between bases of M1 and M2; fasciae undefined; basal half of wing infused with orangish-brown, apical half of wing variably mottled with dark brown and grayish-purple; black patch surrounded by orangish-brown in area of median fascia below discal cell; fringe brown. Hindwing: Uniformly brown; fringe brown. Abdomen ( Fig. 4 ): Brown to pale brown, without modified scales. Genitalia with papillae anales broad and densely setose; apophyses anteriores and posteriores slender, anteriores ca. 1.6 times as long as posteriores; sterigma composed of a pair of ventrolateral, slightly convex, triangular processes separated posteriorly by a ventromedial gap, each process strongly and finely reticulated with cellular spaces bordered by walls and covered with fine spicules, posteromedial corners of processes with several long, thin setae; ductus bursae widened asymmetrically at middle and sclerotized on ventral side from antrum to middle of ductus, forming a ring encircling ductus at middle posterior to inception of ductus seminalis; bulla seminalis absent; corpus bursae large, pear-shaped, with reticulated wall; a pair of large, hollow, paddle-shaped signa, nearly equal in size, with base of right signum arising more posterad than that of left.

Egg ( Figs. 18–19 ). Flat, oval, translucent, white when first laid, turning grey as larva develops. Eggs are laid in masses (described below).

Larva ( Figs. 11–17 , 21–22 ). Mandibles ( Fig. 13 ) with inner teeth (sensu Passoa 1985) modified into long ridge with tooth at the distal end, retinaculum present as a triangular projection; distance between P1 and AF2 on head ( Fig. 12 ) ca. 0.7 times distance between P1 and P2; a horizontal line connecting the AF2 setae on head ( Fig. 12 ) passes closer to P2 than P1; all pinacula weakly sclerotized; L pinaculum on T1 ( Fig. 11 ) anteriad to spiracle; SV groups on A1, 2, 7, 8, 9 with 3:3:3:2:2 setae; SD2 on A1–8 ( Fig. 14 ) reduced, located on same pinaculum as SD1; SD pinaculum on A8 ( Fig. 11 ) anteriad to spiracle; D2 setae on A8 ( Fig. 15 ) on same “saddle” pinaculum and closer together than D1 setae; D1, D2, and SD1 setae on A9 ( Fig. 17 ) all on same large “saddle” pinaculum; D1 and SD1 setae on A9 ( Fig. 17 ) closer together than D2 and D1 setae; L pinaculum on A9 ( Fig. 11 ) bisetose; distance between D1 setae on anal shield ( Fig. 16 ) ca. 1.7 times distance between D1 and SD1 setae; anal comb absent.

Pupa ( Figs. 23–26 ). Length 9.0– 11.6 mm (mean = 10.6, n = 13); abdominal segments 3-8 with two dorsal rows of spines: anterior row larger with ca. 18-32 spines; posterior row smaller with ca. 25-42 spines.

HOLOTYPE: 3: GUATEMALA, Sacatepéquez, Finca San Miguel Urias, San Miguel Dueñas, 1495m, 14°31’27” N, 90°46’34” W, ex. Hass avocado fruit, XII 2006 - III 2007, M. & C. Hoddle, USNM.

PARATYPES: 40 3, 60 Ƥ: GUATEMALA, same data as for holotype (4 3 CSCA; 1 3 TMG; 5 3 UCR; 8 3 USNM, genitalia slides USNM 124,660, TMG 488, TMG 489; 6 Ƥ CSCA; 1 Ƥ TMG; 5 Ƥ UCR; 8 Ƥ USNM, genitalia slides USNM 124,661, TMG 491); Sacatepéquez, Palin, Finca El Recuerdo, 1390m, 14°26’07” N, 90°40’52” W, ex. Hass avocado fruit, 8 XI 2006 - 16 XII 2006, M. Hoddle (5 3 UCR; 1 3 USNM, genitalia slide USNM 124,475; 6 Ƥ UCR); Sacatepéquez, San Pedro de las Huertas, 14°31’55” N, 90°44’4” W, ex. Non-Hass avocado fruit, lab-reared, II-III 2007, M. Hoddle (15 3 UCR; 30 Ƥ UCR); MÉXICO, Michoacán, near Morelia, Nuevo Parangaricutiro, Huerta Ladera 3, ex-larva, 7 XI 2002, Luis Cervantes (1 3 USNM, genitalia slide TMG 492; 4 Ƥ USNM, genitalia slide TMG 493).

Etymology. The specific name is derived from the larval host, Persea americana .

Distribution. Cryptaspasma perseana is recorded from Michoacán, México and central Guatemala.

Biology. Hoddle and Hoddle (2008) described the biology of C. perseana (as “ Cryptaspasma sp. nr. lugubris ”) reared under laboratory conditions; their findings are summarized here. Females laid eggs in masses ( Figs. 18–19 ) containing an average of approximately 21 individual eggs (range of 2 to 73 eggs), primarily on smooth plastic surfaces, although some eggs were laid directly on intact avocado fruit and exposed avocado seeds. Eggs hatched in approximately 11 days (at 22°C) and larvae bored into avocado seeds where they completed development. A single avocado seed supported between 5–8 individual larvae. Larvae completed development in approximately three weeks (at 22°C) and left the seeds in search of pupation sites, although 5% of larvae pupated within feeding tunnels in the seed ( Fig. 20 ). Larval damage was typically characterized by feeding tunnels in seeds, damaged pulp, and accumulation of frass at the opening of feeding tunnels. In the laboratory, larvae pupated between layers of paper towels; we hypothesize that pupation occurs primarily under bark or in leaf litter in natural settings. Larvae remained in a prepupal stage for approximately 2–4 days and in the pupal stage for 8–12 days (at 22°C). The pupal exuvium extrudes from the pupal chamber upon adult eclosion ( Fig. 26 ). Adult sex ratio was found to be 46: 54 male:female under laboratory conditions.

Adult females use pheromones to attract males. Pheromone gland extracts have been made, potential pheromone compounds isolated and identified, but field testing has not been conducted (Millar and Hoddle unpublished). Female calling was observed in the laboratory, characterized by rapid wing vibrations and a slight upturn of the abdomen with large pheromone glands protruding from the posterior. Calling was observed to commence almost immediately following sunset and lasted for approximately one hour, after which all obvious locomotor activity associated with mating by males and females ceased. At sunrise, adult moths rapidly moved to dark concealed hiding places at ground level to rest. When provided with access to a mix of 10% honey and water, adults lived for up to seven days (at 22°C).

It is unclear if females prefer to oviposit on fresh fruit still on the tree or on intact fruit that has recently fallen to the ground. Brown and Brown (2004) hypothesized that Cryptaspasma species are specialists on hard seeds of fallen fruit, suggesting that females do not oviposit on fruit that remains on the tree. Hoddle and Hoddle (2008) reviewed this issue and concluded that, based on field observations of dropped fruit, the likelihood of dropped avocado fruit being consumed by animals in a few days was very high. Thus any eggs laid by Cryptaspasma females on freshly dropped fruit would be eaten by animals as would any larvae that had not yet tunneled into the seed. Because Hoddle and Hoddle (2008) reared C. perseana from intact green fruit that was stated to be freshly picked from trees by a vendor (this was corroborated by circumstantial evidence; the fruit, at time of purchase were in a net sack on the vendor’s back and the vendor was carrying a fruit picking pole), they suggest that oviposition and subsequent larval boring to the seed could have occurred before intact fruit dropped to the ground. Fruit may drop because of larval feeding, or because of strong winds causing fruit drop independently of larval infestation.

One species of endoparasitoid, a Pseudophanerotoma sp. ( Hymenoptera : Braconidae : Cheloninae), was reared from ca. 30% of field collected C. perseana larvae. This is the first host record for a Pseudophanerotoma species; members of the Cheloninae are generally egg and larva parasitoids of tortricoids and pyraloids (J. Luhman pers. comm.).

Additional color photographs of all life stages of C. perseana and Pseudophanerotoma sp. are available ( Hoddle 2011).

Figures

FIGURES 1 – 9. C. perseana adult characters. 1, male holotype. 2, female. 3, male genitalia. 4, female genitalia. 5, male eighth abdominal sternite. 6, male coremata, slide mounted. 7, head. 8, male coremata, posterior aspect. 9, male coremata, lateral aspect.

FIGURE 10. SEM of male coremata scales.

FIGURES 11 – 17. C. perseana larval morphology. 11, complete setal map. 12, setal map of head. 13, mandible. 14, SD pinaculum on A 1 – 8. 15, D 1 and D 2 setae on A 8. 16, anal shield. 17, D 1, D 2, and SD 1 setae on A 9.

FIGURES 18 – 27. C. perseana immature stages. 18 – 19, egg masses. 20, larval damage to avocado fruit. 21 – 22, larvae. 23 – 25, pupae. 26, pupa exuvium extruding from avocado. 27, adult resting on avocado.