Lepidoptera: Crambidae): a New Genus and Species of Musotimine with Leaf-Mining Biology from Costa Rica
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Life history and systematics of Albusambia elaphoglossumae (Lepidoptera: Crambidae): A new genus and species of musotimine with leaf-mining biology from Costa Rica M. Alma Solis1, Donald R. Davis2 & Kenji Nishida3 1 Systematic Entomology Laboratory, PSI, Agricultural Research Service, U. S. Department of Agriculture, c/o National Museum of Natural History, Washington, D.C., 20560-0168, U.S.A., [email protected] 2 Department of Entomology, Smithsonian Institution, Washington, D.C., 20560-0127, U.S.A., [email protected] 3 Sistema de Estudios de Posgrado en Biología, Escuela de Biología, Universidad de Costa Rica, 2060 San José, Costa Rica. [email protected] Received 15-V-2004. Corrected 09-II-2005. Accepted 10-II-2005. Abstract: Albusambia elaphoglossumae Solis & Davis, a new genus and species, is described. It was discov- ered mining the fronds of the fern Elaphoglossum conspersum in Costa Rica (San José and Cartago Provinces, at elevations of 2300-3100 m). The type series was obtained by rearing of the immature stages in laboratory. The adult is defined by unique genital characters, and the pupa with a medial depression on the vertex and with two anterolateral horn-like structures on the prothorax. The larva is a gregarious leaf miner with its body flat- tened dorsoventrally and head prognathous; morphological adaptations to its leaf-mining habit are new to the Musotiminae. Fern-feeding musotimines are important to the discovery of new biological control agents for invasive ferns. Rev. Biol. Trop. 53(3-4): 487-501. Epub 2005 Oct 3. Key words: Elaphoglossum conspersum, E. biolleyi, gregarious larvae, fern, leaf miner, Neotropical, Neurophyseta, Proacrias, Undulambia. Recent collecting in the Cordillera de about 200 species worldwide; 50 or more Talamanca of Costa Rica resulted in the dis- species and five to seven genera from Asia covery of a new genus and species of a and the South Pacific are still undescribed musotimine crambid mining fronds of the (S.-H. Yen, pers. comm.). There are nine gen- fern, Elaphoglossum conspersum H. Christ era and 82 species of Musotiminae in the (Pteridophyta: Lomariopsidaceae) (Figs. 1-3). Western Hemisphere, including Albusambia. The mining habit in the Pyraloidea is not Musotimines are defined by their laterally common. Other New World musotimines compressed antennae with flattened segments, have been reared feeding on ferns. These R2 stalked with R3+R4 (but not in Albusambia), are Undulambia polystichalis (Capps), reared a reduced coecum in the aedeagus of the from leather leaf fern, Rumohra adiantiformis male genitalia, and enlarged tympanal cases [=Polystichum adiantiforme (G. Forst) Small] (Minet 1985, Yoshiyasu 1985, Yen 1996). The Ching (Dryopteridaceae) (Kuitert and Dekle relationship of Albusambia to other genera 1962). In 2002, Neurophyseta camptogram- is unknown, although it is externally most malis (Hampson) was reared from Alsophila similar to Neurophyseta Hampson. The genus firma (Baker) D.S. Conant (Cyatheaceae) Undulambia Lange contains three species from (Janzen and Hallwachs database homepage). Florida and is the only genus of Musotiminae Musotiminae is comprised of 18 genera and found in the U.S. Lange (1956) placed it in Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 53 (3-4): 487-501, September-December 2005 487 Nymphulinae, a subfamily known for its true room (16-18°C) in the Museo de Insectos, aquatic caterpillars. Munroe (1972) later placed Universidad de Costa Rica, UCR (San Pedro Undulambia in a new, separate tribe, Ambiini, de Montes de Oca, 1150 m), or were placed within Nymphulinae, because U. polystich- in a refrigerator (7.0-8.0°C) for approximately alis was terrestrial and fed on ferns. Munroe 16 hours and then at room temperature (20°C) (1972) also noted that Musotima Meyrick (in for eight hours daily. A subset of specimens of Musotimini) larvae were terrestrial and fed immature stages were preserved in alcohol and on ferns. Speidel (1984) separated and raised deposited in the USNM. to subfamily rank the Musotiminae, including The host plant, E. conspersum, is common Ambiini from Nymphulinae, based on charac- in dense understory growth at several moist, ters of the genitalia and immatures. well-shaded sites along roads (Fig. 2). Vegetation In this paper, we describe the life history associated with E. conspersum consists of a of Albusambia elaphoglossumae as well as all tropical montane rain forest densely populat- life-stages in detail. The species is compared to ed by temperate elements such as black oaks U. polystichalis and species of Neurophyseta. (Quercus costaricensis Liebm.) (Fagaceae), with more scattered white oak (Q. copeyensis C. H. Muller) (Hartshorn 1983). Other abundant, MATERIALS AND METHODS principally understory species in this association include Schefflera rodriguesiana Frodin ex M. The eggs, larvae, and pupae were collected Cannon and Cannon (Araliaceae), Weinmannia in Costa Rica in an oak forest near the Pan pinnata L. (Cunoniaceae), Cavendishia brac- American Highway at Villa Mills, Cerro de la teata (Ruiz and Pav. ex J. St.-Hil.) Hoerold, Muerte (San José province) (Fig. 1), and CATIE Vaccinium consanguineum Klotzsch (Ericaceae); (Centro Agronómico Tropical de Investigación Miconia biperulifera Cogn., M. schnellii y Enseñanza) Biological Station in Cuericí Wurdack (Melastomataceae); and several species (Cartago province), at elevations between 2800 of Chusquea (Poaceae) (KN, pers. observ.). and 3100 m. Larvae from E. conspersum were first collected in January 1998 at the end of the wet season and reared by Donald R. Davis RESULTS (DRD). The wet season in this area extends from April through December or January. Albusambia Solis and Davis, n. gen. Heavy rains are common during these months, with a yearly annual rainfall of 2 812 mm Diagnosis: Base color of body shiny (Kappelle 1996). During the dry season in the white; wing margins not incised; forewing year 2000, Kenji Nishida (KN) visited the same costal swelling absent; male genitalia with area and began collecting additional material, valva curved in above junction of tegumen by inspecting fronds and conducting extensive and vinculum and continuing less broadly to observations in the field. In this region, the dry anterior end. Larva dorsoventrally flattened; season starts in December or January and ends head prognathous, mandibles with two rows of in mid- to late April. Although the rain is infre- serrations. Pupa with vertex of head round and quent, the humidity remains high and dense fog with prominent medial dorsoventral depres- is common in the afternoons. The annual aver- sion; prothorax smooth dorsally, not square, age temperature is 10.9°C, and the temperature with two anterolateral horn-like structures that sometimes decreases to -3°C during the dry protrude only slightly; anus posteriorly with season (Kappelle 1996). lateral conical depressions. Immature stages with the fronds collect- Comparison to other taxa: Albusambia ed by KN were placed in transparent plas- is compared to Undulambia in Capps (1965) tic bags and taken into an air-conditioned and Neurophyseta in Phillips and Solis (1996). 488 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 53 (3-4): 487-501, September-December 2005 The basal color of Undulambia adults is brown, Thorax: Dorsally white; patagium and not white, as in Albusambia and Neurophyseta. tegula white, a few specimens with tegula Wings of Undulambia are incised, and grey and metathorax dorsally grey. Forelegs those of Albusambia and Neurophyseta are white, mediolaterally black; midleg and hind- entire. Forewing costal swelling is absent leg white, midleg with one pair of tibial spurs, in Albusambia and Neurophyseta, but pres- hindleg with two pairs of tibial spurs. ent in Undulambia. In the male genitalia of Forewing (Fig. 21) [Refer to Phillips and Undulambia the valva is long and equal in Solis, 1996, Fig. 7 for venation terminology]: width throughout, not widened posteriorly as length = 6.91 mm (range: 6.0-8.1 mm) (n=36); in Albusambia and some Neurophyseta spe- margins not incised. Costal swelling absent. R1 cies. The body of Undulambia larva is round, and R2 from discal cell; R5 short stalked with not dorsoventrally flattened, and not inter- R3+4, stalked about two-thirds its length. Basal segmentally constricted as in Albusambia. In band black, basal area white with scattered Undulambia the mandible consists of one line black scale, antemedial band black, antemedial of teeth, not two as in that of Albusambia. area white, reniform spot black, postmedial line Other than the host plant record, the bio- black, black spots between R4 and R5 and on logical and morphological data of the larvae M3, fringe white with black patches on M3 and of Neurophyseta are not available. Unlike CuA2; a few specimens mostly black posterior to the vertex of the pupal head of Albusambia, and slightly above M-CuA line to posterior mar- Undulambia does not have a prominent medial gin, antemedial and postmedial lines broader, dorsoventral depression. Undulambia has a black lines to outer margin from postmedial line somewhat rugose, broadly flattened prothorax on M3 and CuA2, a black line parallel to postme- with two anterolateral horn-like structures that dial line from M3 to posterior margin. Underside protrude twice as much as in the smooth pro- white with postmedial line slightly visible with thorax of Albusambia. Albusambia has lateral black scales, some black at basal area near costa; conical depressions posterior to the anus, but slightly darker specimens with similar color and Undulambia lacks these depressions. Pupal pattern on underside as on upperside. data on Neurophyseta is incomplete. Hindwing (Fig. 21): Antemedial and post- Etymology: The prefix of the generic epi- medial line black, black discal spot, c-shaped thet is derived from the Latin “albus” referring mark from Rs to costal margin at intersection of to the white base color of the adult moths, and Sc+R1 and Rs, fringe white with black patches “ambia” meaning “around” in Latin and refer- on R5, M3, and CuA2; a few specimens black ring to its relatedness to other genera in the sub- basally to postmedial line, with broad black family.