With Discussions on Male Secondary Sexual Characters and Larval Feeding on Capparis (Capparaceae) in the Pyraloidea and Lepidoptera (Insecta)
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SYSTEMATICS Phylogenetic Analysis of Cosmopterosis (Lepidoptera: Crambidae: Glaphyriinae) With Discussions on Male Secondary Sexual Characters and Larval Feeding on Capparis (Capparaceae) in the Pyraloidea and Lepidoptera (Insecta) 1,2 1 3 M. ALMA SOLIS, MARK A. METZ, AND DANIEL H. JANZEN Ann. Entomol. Soc. Am. 102(5): 766Ð784 (2009) ABSTRACT New species of Cosmopterosis Amsel were discovered feeding on Capparis L. (Cap- paraceae) during exploration for caterpillars in the Area de Conservacio´n Guanacaste (ACG), Costa Rica. Cosmopterosis is revised and now includes four species. Three new species, C. hispida, C. jasonhalli, and C. spatha, and the immatures of C. spatha and biology for two species, C. jasonhalli and C. spatha, are described; the type species, C. thetysalis (Walker), is redescribed. A key and illustrations for the identiÞcation of the species is provided. We propose a hypothesis for the relationship between species in Cosmopterosis and the placement of Cosmopterosis in the subfamily. The cladistic analysis, the Þrst such analysis in the Glaphyriinae, included 21 morphological characters one of which, the radiodiscal process, a male secondary sexual character and presumably an androconial scent pouch is described and considered a autapomorphy for the genus. Male secondary sexual characters and larval feeding on Capparis in Pyraloidea and Lepidoptera is discussed. KEY WORDS Pyraloidea, systematics, larval morphology, Costa Rica, Capparaceae Cosmopterosis Amsel has been a monotypic genus areas, strong white-cream dorsal patches and lines since 1956 when it was described. Recently, we dis- against a black background, and commonly occur ag- covered two new species that D.H.J. reared from lar- gregated in large numbers. Aggregation and coloration vae in Costa Rica on Capparaceae, and an additional may reßect antipredator evolution: the anal area looks undescribed species in material from museum collec- like the head, and the dark dorsal area with white tions. We provide a hypothesis of monophyly for Cos- patches or lines provides crypsis (see Fig. 23). This mopterosis and for relationship between the species. same pattern is present in a large mimicry complex of One of the characters that deÞnes Cosmopterosis is the similarly colored caterpillars (D.H.J., personal obser- radiodiscal process on the male forewing, a secondary vation). sexual character and presumably an adronconial scent pouch. Recent systematic studies have shown that secondary sexual characters can be phylogenetically Materials and Methods informative. We discuss the previously unknown bi- ology of the genus and the paucity of larval feeding on Material Examined. We studied Cosmopterosis Capparaceae by Lepidoptera. specimens in 12 collections worldwide and 11 collec- Cosmopterosis is a member of the Glaphyriinae. This tions in the United States. Institutions with material subfamily currently includes 33 genera and 174 species that contributed signiÞcantly to this study included: (Solis and Maes 2002). Most species occur in the (BMNH) The Natural History Museum, London, U.K.; Western Hemisphere (Munroe 1995), with the excep- (INBIO) Instituto Nacional de Biodiversidad, Santo tion of and including species of Hellula Guene´e that Domingo de Heredia, Costa Rica; (RMNH) National occur worldwide and are serious pests on Brassicaceae Natuurhistorisch Museum, Leiden, The Netherlands; (Solis and Adamski 1998; Landry and Roque-Albelo (USNM) National Museum of Natural History, Smith- 2008). Larvae of Cosmopterosis reared by D.H.J. on sonian Institution, Washington, DC, U.S.A.; and the Capparis L. (Capparaceae) are brightly colored and (SDNHM) The San Diego Natural History Museum, conspicuous. They have red-to-orange heads and anal San Diego, CA, U.S.A. We deposited type material in these institutions as designated within the text. Spec- 1 SEL, USDA, Smithsonian Institution, P.O. Box 37012, National imens are listed at the end of each species description Museum Natural History, E-517, MRC 168, Washington, DC 20013- with identiÞcation number such as slide numbers for 7012. either the USNM or BMNH and/or specimen num- 2 Corresponding author, e-mail: [email protected]. 3 Department of Biology, The University of Pennsylvania, Phila- bers, including those in the format “nn-SRNP-nnnnn” delphia, PA 19104. from the database of Janzen and Hallwachs (2005). 0013-8746/09/0766Ð0784$04.00/0 ᭧ 2009 Entomological Society of America September 2009 SOLIS ET AL.: PHYLOGENETIC ANALYSIS OF Cosmopterosis 767 Table 1. Character matrix Character/state Taxon 123456789101112131415161718192021 argentistriata 000101000 000000000110 punctissimalis 000010000 000000000000 hispida 111101001 111111111111 jasonhalli 110101010 101111111111 spatha 110101111 111111111111 thetysalis 111101001 111111010110 We examined and dissected pinned specimens after Table 1. All characters are binary. Seven characters are abdomens were soaked in 10% potassium hydroxide from the wings, one from the abdomen, 10 from the and wings soaked in bleach. Dissections were stained male genitalia, and three from the female genitalia. in chlorozal black for genitalia, and Eosin-Y for wings. Genitalia were placed in vials with glycerine and/or 1. Male forewing with radiodiscal process: (0) ab- slide mounted in Canada balsam or Euparol (Clarke sent; (1) present (Fig. 11). The males of Cos- 1941, Holloway et al. 1987). Measurements were made mopterosis possess a set of strong, fused setae with an ocular micrometer. Terminology follows Hin- whose bases originate above the discal cell at the ton (1946), Neunzig (1979), Wooton (1979), Klots conßuence of the radial sector. This Þngerlike (1970), Maes (1985, 1995, 1997), Yoshiyasu (1985), process is hinged along the radial vein and is cov- Phillips and Solis (1996), and Solis and Maes (2002). ered with long, dense scales that lay over the discal Morphological structures were photographed using cell. The discal cell is obscured by the scales on the Microptics imaging system and retouched with this process as well as similar scales more basal Adobe Illustrator (Adobe Systems, Mountain View, along the radial sector and more apical along vein CA). Rs4. The specimens with this process also possess Cladistic Methods. We studied six taxa for poten- a dense patch of appressed setae along the an- tially informative characters and disregarded only terobasal border of the discal cell that may rep- those that were not stable within a species. Autapo- resent an additional character. This radiodiscal morphic characters were included in the matrix be- process (often referred to as a costal fold) has cause they may be informative for future studies. We never been described previously and is consid- conducted an exhaustive search in PAUP* (Swofford ered to be a autapomorphy for Cosmopterosis. 1998) to Þnd all possible trees. Decay indices were 2. Distal discal cell of male forewing with a dense calculated using TreeRot (Sorenson 1999). Character patch of erect setae: (0) absent; (1) present (Fig. state changes were plotted on the hypothesized tree 11). The males of Cosmopterosis have the mem- using unambiguous character transformation using brane at the apex of the discal cell covered by an Winclada (Nixon 1999). erect patch of setae that is distinct from the ap- pressed setae along the anterobasal border. 3. Male forewing with a patch of erect setae distal to Systematics the discal cell above M2: (0) absent; (1) present Taxa. Because of the lack of knowledge of the phy- (Figs. 1 and 4). Cosmopterosis thetysalis and C. logenetic relationships among the 33 Neotropical hispida have males with an additional patch of glaphyriine genera, representatives of each genus erect setae as described above. were examined externally. Outgroup taxa were chosen 4. Forewing postmedial line color: (0) brown; (1) from the currently described genera of Glaphyriinae, silver (Figs. 1Ð4). with special consideration given to those genera that 5. Hindwing spatulate scales between veins CuA2 shared one or more diagnostic characters with the and CuP: (0) absent; (1) present. holotype of the type species C. thetysalis. These shared 6. Hindwing cataclystiform spots: (0) circular, en- features of particular importance included the pres- tire; (1) basally biÞd, medially separate (Figs. ence of cataclystiform spots (Munroe 1991) as well as 1Ð4). squamiform and/or piliform scales between CuA2 and 7. Hindwing cataclystiform spots ventrally: (0) vis- CuP on the dorsal surface of the hindwing. Species ible; (1) not visible. with these shared characteristics were then assessed 8. Venter VII with a distinct set of more robust setae for similarities in the male and female genitalia, no- on the posterolateral corner: (0) absent (Fig. 7); tably a prolonged uncus and a posteroventral exten- (1) present (Fig. 8). The abdomens of all the sion of the sacculus. Ingroup species were determined investigated taxa are covered with dense scales to possess hypothesized autapomorphies for the ge- and setae; C. jasonhalli and spatha have a patch of nus, which included specialized scales on the male setae located on the posterolateral corner of ven- forewing and male genitalic characters that were ter VII. These setae are distinct in that they are tested in the cladistic matrix. longer and stronger and the sockets Þtting to their Characters. The 21 characters and their states used bases are larger and noticeable even after the setae for the analysis are listed below and presented in are removed during dissection. 768 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 5 Figs. 1–6. Cosmopterosis male habitus. 1. C. hispida Solis, n. sp. 2. C. jasonhalli Solis, n. sp. 3. C. spatha Solis, n. sp. 4. C. thetysalis (Walker). 5. Living larva of C. jasonhalli Solis, n. sp. in its natural setting (photo by D.H.J.). 6. Living larvae of C. spatha Solis, n. sp. in its natural setting (photo by D.H.J.). 9. Base of uncus: (0) narrow, not expanded much tegumen equal in size or larger (Figs. 9 and 10); wider than rest of uncus, subtended or terminate, (1) composing at least 80% of circumference, adjacent to posterior margin of tegumen (Figs. 9 tegumen much smaller in size (Figs.