SYSTEMATICS Phylogenetic Analysis of Cosmopterosis (: : ) With Discussions on Male Secondary Sexual Characters and Larval Feeding on Capparis () in the 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 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 . 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 Guene´e that Domingo de Heredia, Costa Rica; (RMNH) National occur worldwide and are serious pests on 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. 14Ð17). and 10); (1) expanded dorsolaterally and medially 11. Dorsal articulation between vinculum and valva: invaginate, signiÞcantly overlapping posterior (0) condylate (Figs. 9 and 10); (1) adjacent (Figs. margin of tegumen (Fig. 16). In all of the species 14Ð17). See Solis and Maes (2002), character 10. of Cosmopterosis, the base of the uncus is concave 12. Uncus with dorsal recurved setae: (0) absent ventrally and expanded dorsolaterally giving it a (Figs. 9 and 10); (1) present (Figs. 12 and 14Ð17). carapacelike appearance. Likewise, there is a deep 13. Pseudognathos: (0) small, simple (Figs. 9 and 10); notch medially at the anterior margin. This invag- (1) large, complex (Figs. 12 and 14Ð17). Outgroup ination seems to accommodate the phallus when taxa have an extremely short pseudognathos that it everts during copulation. is evident as an abrupt terminus of cuticle. All 10. Ninth segmental ring of male terminalia with vin- species of Cosmopterosis have a pseudognathos culum: (0) composing no Ͼ50% of circumference, that is not only extremely long, but constricted September 2009 SOLIS ET AL.: PHYLOGENETIC ANALYSIS OF Cosmopterosis 769

Figs. 7–10. 7Ð8. Venter VII in ventral view: 7. Cosmoperosis thetysalis (Walker) lacking distinctly robust setae; 8. Cosmopterosis spatha Solis, n. sp. showing the distinct set of more robust setae on the posterolateral corner. 9Ð10. Ventral view of the male genitalia of the outgroup taxa: 9. Aureopteryx argentistriata (Hampson); 10. Pseudoligostigma punctissimalis (Dyar).

subbasally before widening, and sharpened by sacculus that is located above the posteroventral carinae dorsolaterally, dorsomedially, and ventro- process in the previous character. medially. The dorsomedial carina is heavily armed 17. Juxta with deep median channel: (0) absent (Figs. with recurved spines apically. 10 and 14); (1) present (Figs. 15Ð17). The males 14. Pseudognathos with “apical cone:” (0) absent of three of the species of Cosmopterosis have a (Figs. 9 and 10); (1) present (Figs. 12 and 14Ð17). juxta that is bilaterally bisected by a deep median In addition to the uniformity in the shape of the channel running its length. pseudognathos, the species of Cosmopterosis also 18. Anellus with eversible setae: (0) absent (Figs. 9 possess a membranous process at the apex of the and 10, 14, 17); (1) present (Figs. 15 and 16). The pseudognathos. This process is conical in shape membranous area that spans the ring of the ninth with a pair of anteriorly pointing, setiform exten- segment is described by Klots (1970) as the dia- sions. phragm and the eversible cone that connects to 15. Sacculus with enlarged ventral process: (0) absent the zone of the phallus as the anellus. Two species (Figs. 9 and 10, and 14); (1) present (Figs. 15Ð17). of Cosmopterosis have a set of setae on the anellus Excluding one new species (C. hispida), the males lateral to the juxta. When the phallus is everted, of Cosmopterosis all have an extension of the pos- these setae seem to have an association with the teroventral margin of the sacculus near its apex. In denticles of the sacculus (character 16) based on two of these species, C. jasonhalli and C. thetysalis, their close, physical proximity. the process is pointed (Figs. 15 and 17), and in C. 19. Anterior apophyses with pointed process in spatha it is spatulate (Fig. 16). middle: (0) absent; (1) present (Figs. 18Ð20). 16. Sacculus with denticles above process: (0) absent; The females of all the species here analyzed (1) present (Figs. 15 and 16). Cosmopterosis jas- have anterior apophyses that are thickened at onhalli and C. spatha have a patch of thickened, the middle. All of the species analyzed except P. denticulate extensions on the inner surface of the punctissimalis have an additional pointed, tooth- 770 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 5

Figs. 11–13. 11. Partial view of the left forewing of a male specimen of Cosmopterosis jasonhalli Solis, n. sp. The radiodiscal process is elevated and moved anteriorly showing the appressed and erect setae of the discal cell. 12. Anterodorsal view of the base of the uncus of Cosmopterosis hispida Solis, n. sp. showing the expanded carapacelike dorsum and the deep, anteromedial invagination. 13. Single most parsimonious tree resulting from the analysis of the morphological data with unambiguous character state transformations indicated. See text for tree statistics. Solid dashes on nodes indicate forward changes with no homoplasy. Boxes contain values of that nodeÕs decay index.

like process extending medially from this thick- tance between signa (Fig. 19). C. spatha has the ened area. autapomorphic state of the thickened denticles 20. Signa with thickened denticles: (0) absent; (1) forming the signa to be continuous, that is the present (Figs. 18Ð20). The surface of the mem- outer edge of the left and right signa meet at the brane forming the corpus bursae has denticulate anterior end. microsculpturing in all females here analyzed. All of the females except P. punctissimalis have signa Phylogenetic Analysis. The exhaustive search in that are formed from a distinct thickening of these PAUP* produced a single shortest tree (Fig. 13) with denticles with the denticles becoming increas- a length of 22 steps (CI ϭ 0.95, RI ϭ 0.94). A mono- ingly more robust toward the center of the signa. phyletic grouping of the species related to C. thetysalis 21. Signa: (0) shorter, length less than distance be- is strongly supported (decay index ϭ 8). The charac- tween signa; (1) longer, length at least 1.5X dis- ters that strongly support this are the presence of a September 2009 SOLIS ET AL.: PHYLOGENETIC ANALYSIS OF Cosmopterosis 771

Figs. 14–17. Male genitalia of Cosmopterosis. 14. C. hispida Solis, n. sp. 15. C. jasonhalli Solis, n. sp. 16. C. spatha Solis, n. sp. 17. C. thetysalis Walker. radiodiscal process (character 1) and a dense patch of postmedial line is strongly angled toward the wing erect setae (character 2) in the discal cell of the male base at the tornus and the area anterior of this longi- forewing, an uncus with a concave base (character 9), tudinal section, distal to the discal cell, is species spe- a ninth segmental ring of the male terminalia with the ciÞc in its coloration. The area between the subter- vinculum at least 80% of circumference (character minal and terminal lines is yellow. Males have a 10), the dorsal articulation between the vinculum and radiodiscal process (character 1) as described above. the valva adjacent (character 11), the uncus with All species have distinct cataclystiform spots on the dorsal recurved setae (character 12), and the pseu- hindwing and have erect piliform scales between veins dognathos large and complex (character 13) with an CuA2 and CuP. In addition to these external features, apical cone (character 14). the males possess distinctive characters in their gen- italia described above (characters 9, 12Ð15). Species Cosmopterosis Amsel are all easily recognized by characters in the genitalia but can also be distinguished by a combination of Cosmopterosis Amsel 1956: 109. Type species Cata- external characters. clysta thetysalis Walker, 1859: 440, by original des- Adult. Head (Figs. 1Ð4). Frons creamy white later- ignation, Amsel 1956: 110. ally and dorsally, brown medially. Vertex with setae Diagnosis. Species are easily recognized by the dis- creamy white to yellow basally and yellow to reddish tinct patterns on the fore- and hindwings. The forew- brown apically, vertex looking mostly reddish brown, ings (Figs. 1Ð4) are mostly reddish and dark brown but with mixture of pale yellow and creamy white. with obscure lines except the postmedial line, which Antennae Þliform, laminate, dull gold; scape covered is a compound line of silver apically and brown basally; with mixed creamy white and gray scales; pedicel and the subterminal, which is silver; and the terminal, ßagellomeres covered with silver-gray scales dorsally, which is composed of a line of isolated black spots. The with short, golden setae ventrally, and two rows of 772 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 5

Figs. 18–20. Female genitalia of Cosmopterosis. 18. C. jasonhalli Solis, n. sp., inset provides a ventral view of the signa. 19. C. spatha Solis, n. sp., inset provides a ventral view of the signa. 20. C. thetysalis Walker. longer, golden setae ventromedially and ventrolater- margin; area between postmedial line and subterminal ally. Labial palpus three-segmented, basal segment line mostly rufous, golden yellow in narrow band ap- arcuate, apical segment with an internal channel open proximate to postmedial line; terminal line black, dis- apically, basal and middle segments creamy white ba- continuous and showing up as spots at apices of lon- sally and apically, grayish brown in middle, apical gitudinal veins; apical fringe double, dorsal fringe dark segment creamy white. Maxillary palpus three-seg- gray, half length of ventral fringe, ventral fringe whit- mented, basal and middle segments cylindrical, apical ish silver. Male with a dense patch of appressed, segment spheroidal, mixed creamy white and grayish brown-tipped scales from middle of R to furcation of brown. Proboscis much longer than labial palpus. radial sector that obscures most of discal cell and Ocelli present, disc mostly black, lens metallic gold. dense patch of erect, Þliform setae underneath, and a Chaetosemata absent. patch of dense, erect scales at apex of discal cell to

Thorax (Figs. 1Ð4). Pronotum brown medially with anterad base of M1; these absent in female. Ventral lateral margins creamy to yellowish white. Mesono- surface mostly yellowish silver to pale brown, inner tum mostly brown medially, only extreme bases of posterior margin whitish silver to 1Aϩ2A, medial line scales creamy white, creamy to yellowish white lat- and postmedial line pale brown, medial line fading erally. Pleura and sternites yellowish silver. Legs yel- toward posterior margin. Retinaculum composed of lowish silver. Forecoxa grayish brown anterodorsally. dense patch of anteriorly facing, appressed scales be- ϩ Fore- and midlegs grayish brown dorsally on femora tween bases of CuA1 and 1A 2A. and tibiae, and basally on tarsomeres. Midleg with one Hindwing (Figs. 1Ð4). Costa almost straight; termen pair of tibial spurs. Hindleg with two pairs of tibial slightly crenulate to tornus. Above basal half white to spurs. yellowish white; apical half with diffuse pattern of Forewing (Figs. 1Ð4). Costa almost straight, slightly brown-tipped scales; margin with six large, silver-me- arcuate; apex acute, but evenly rounded; termen tallic, black-margined cataclystiform spots, spots straight to CuA1; tornus obtuse and evenly rounded; rounded along crenulae of termen distally, deeply distal apex of discal cell poorly deÞned; M1 straight, invaginated proximally; apical fringe double, dorsal base closer to base of Rs4 than M2, but not approxi- fringe yellowish silver proximally, dark grayish brown mate; M2 arising separately from discal cell, but ap- distally, half length of ventral fringe, ventral fringe proximate to origin of M3. Postmedial line continuing yellowish silver. Below mostly yellowish silver, apex from costa posterolaterally in gentle arc, approximate pale brown, costa more yellow. Ventral surface lacks to termen at CuA2, then strongly angled basally for a patch of erect scales between veins CuA2 and CuP. short distance before angling toward tornus, reßec- Female frenulum with two setae. tive, metallic, silver when viewed dorsally, gray when Abdomen. Male tergite I whitish silver, tergites IIÐ viewed obliquely, margined basally by thin, rufous VII yellowish silver with pale brown basal border, band, silver component not reaching costa anteriorly; tergite VIII and dorsal terminalia mostly pale brown, subterminal line silver, not reaching costa or tornal sternites yellowish silver slightly darker basolaterally, September 2009 SOLIS ET AL.: PHYLOGENETIC ANALYSIS OF Cosmopterosis 773

Figs. 21–23. Pupa of C. spatha Solis, n. sp. 21. Lateral view. 22. Ventral view. 23. Lateral view of preserved larva of Cosmopterosis spatha (some extremely long setae digitally reconstructed). apex of valva pale brown. Tympanal organs crambi- but sharply angled, posterior width of tympanal organs form; tergo-sternal sclerite strongly sclerotized; bulla half width of anterior width; zona glabra tympani tympani ovate, longer than wide; fornix tympani extremely short; puteolus tympani absent. weakly sclerotized, posteromedial margin extended Male Genitalia (Figs. 12 and 14Ð17). Tegumen almost to pons tympani, thus ala tympani extremely short, vinculum composing all but transverse dorsal short and ligna tympani almost transverse; tympanum portion of ninth segmental ring, saccus robust, thick- greatly enlarged, conjuctivum reduced to transverse ened, lateral arms narrowing dorsally; uncus expanded strip; spinula pyriform; processes tympani enlarged, dorsolaterally and medially invaginate, signiÞcantly broadly lamellate; pons tympani strongly divergent; overlapping posterior margin of tegumen, narrowly ramus tympani weakly sclerotized, arcuate, not extended, dorsally convex and arched in both longi- strongly angled medially; venula prima almost straight, tudinal and lateral planes, densely, but Þnely setose 774 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 5 dorsally, dorsal subapex with two pairs of strong, re- plicitly state which characteristics were unique to C. cumbent setae, extreme apex in lateral view recurved, thetysalis. Amsel never saw the holotype and erected in dorsal view bifurcate, ventrolateral margin carinate, the genus based on voucher specimens “determined sclerotized medially on ventral surface, membranous by Martin” at the BMNH that we subsequently deter- laterally, median sclerotization acutely pointed pos- mined to be an undescribed species. Cosmopterosis teriorly; pseudognathos continuous with tegumen, remains a valid taxon with no species having been transverse basally, narrowing subbasally, and ex- added by description or combination. panded distally before becoming apically acute, dor- Munroe (1964) placed Cosmopterosis in Glaphyri- solateral margins carinate, medially carinate and inae where it currently resides. Forbes (1920) Þrst heavily armed with recurved spines apically, extreme deÞned Glaphyriinae based on a patch of erect scales apex with posteroventral, conical extension and pair of on the hindwing between veins CuA2 and CuP, but dorsoanteriorly directed spines, ventral surface fully Solis and Adamski (1998) demonstrated that these sclerotized with a median carina; transtilla weakly scales were not present on all species within a genus sclerotized, but continuous medially, simple, straplike, and were completely lacking among the species in and arched medially; juxta ovate, much higher than some genera such as Cosmopterosis. Munroe (1964) wide; valva spatulate subapically, apex bluntly atten- added several new taxa to the subfamily using the uate, surfaces and margins sparsely setose, inner sur- concept of Forbes (1920, 1924), but to date no cladistic face with transverse rugae in distal half, costa strongly analyses based on either morphological or molecular thickened basally, attenuate distally, almost reaching characters among the glaphyriine taxa exist. Generic- apex; coecum long, 0.75ϫ length of remaining phallus, level relationships within the subfamily are com- phallus biÞd apically, moderately sclerotized, cornu- pletely unknown so some of the currently recognized tus a dense patch of long, straight spicules. genera are likely to be either paraphyletic or polyphyl- Female Genitalia (Figs. 18Ð20). Papillae anales lon- etic. gitudinally shortened, in posterior view kidney- shaped, concave medially, setae densely covering sur- face, setae stronger at external margin and becoming Key to Species of Cosmopterosis weaker toward center; tergite VIII weakly sclerotized dorsomedially, membranous laterally; venter VIII 1. Hindwing upper surface with cataclystiform membranous; posterior apophyses slender, turning spots incomplete, middle area with brown medially near midpoint; anterior apophyses robust, 2ϫ scales concolorous as those of adjacent area of length of posterior apophyses, thickened near mid- hindwing (Fig. 4). Sacculus with posteroven- point, with a medially directed process at approxi- tral ßange Ͼ5ϫ longer than wide and apex mately one third its length; ostium bursae narrowing rounded (Fig. 17). Internal, sclerotized part anteriorly to cervix bursae; corpus bursae spherical, of phallus with spicules (Fig. 17). Apical half Ϸ0.5ϫ to subequal in length of segment VII, weakly of valva parallel-sided, apex evenly rounded spiculate; ductus bursae 2ϫ length of segment VII, (Fig. 17). Basal (posterior) one fourth of duc- slightly expanded near corpus bursae, inserting at cor- tus bursae Ͻone-third width of segment VII pus bursae posteroventrally; signa oval, 3ϫ longer than and smooth, with no rugosity (Fig. 20).... wide, medially concave, densely spiculate, spicules ...... Cosmopterosis thetysalis larger centrally, spicules grading to Þne setiform struc- 1Ј. Hindwing upper surface with cataclystiform tures toward margin; ductus seminalis inserted into spots complete, middle area with black and ventral surface of ductus bursae anterad cervix bursae; yellow scales differing in color from adjacent spermathecal duct coiled, spermathecal vesicle area of hindwing (Figs. 1Ð3). Sacculus with weakly sclerotized laterally. posteroventral ßange Ͻ4X longer than wide Species Variation. There is considerable variation or absent (Figs. 14Ð16). Internal, sclerotized among specimens within any given species. Subtle part of phallus smooth (Figs. 14Ð17). Apical variations of hue and tone exist, but are not explicitly half of valva attenuate (Figs. 14Ð17). Basal noted. (posterior) one fourth of ductus bursae Distribution. Western Mexico (Nayarit, San Luis Ͼone-half width of segment VII, usually rug- Potosi, Sinaloa) east to the Republic of Trinidad and ose (Figs. 18Ð19) ...... 2 Tobago and south to Paraguay. 2. Forewing medial area, basal to dorsal portion of Biology. Specimens from Costa Rica have been postmedial line, yellow and brown, with no reared from Capparis and Forchhammeria (Cap- white scales (Fig. 1). Male forewing with cir- paraceae). cular patch of erect setae apical to overhang-

Remarks. Amsel (1956) described Cosmopterosis for ing setae of radiodiscal process above M2 (Fig. Cataclysta thetysalis Walker, which at the time was 1). Sacculus lacking substantial posteroven- placed in Aulacodes Guene´e. He was impressed by its tral ßange or spicules near posterior margin ornate wing pattern (cosmo- ϭ ornamental, ptero- ϭ (Fig. 14). Anellus lacking patch of eversible wing) and stated that the morphology precluded it setae (Fig. 14). Juxta with sclerotized dorsal from belonging to Aulacodes, in the or extension with strong, long spines forming any of its synonyms because those type species had a large brush (Fig. 14) ...... completely different morphology. But he did not ex- ...... Cosmopterosis hispida Solis September 2009 SOLIS ET AL.: PHYLOGENETIC ANALYSIS OF Cosmopterosis 775

2Ј. Forewing medial area, basal to dorsal portion of Abdomen (Fig. 1). Male tergite I pale brown medi- postmedial line, with broad areas composed ally. of white scales (Figs. 3Ð4). Male forewing Male Genitalia (Figs. 12 and 14). Juxta with shallow lacking circular patch of erect setae apical to median channel that dissipates dorsally at anellus, dor- overhanging setae of radiodiscal process sally acuminate; anellus bordered laterally by sclero-

above M2 (Figs. 2Ð3). Sacculus with substan- tized extensions with strong, long spines forming large tial posteroventral ßange clearly extending brush; sacculus more strongly sclerotized than rest of beyond margin and with spicules near poste- valva, apex with very short, bluntly pointed extension. rior margin (Figs. 15Ð16). Anellus with patch Female. Unknown. of eversible setae (Figs. 15Ð16). Juxta with Immature Stages. Unknown. short spicules, but lacking sclerotized dorsal Material Examined. Five males. extension (Figs. 15Ð16) ...... 3 Type Material. HOLOTYPE: 1 male, BRAZIL, Rio 3. Hindwing ventral surface with or without cata- de Janeiro, 10 km SW Maraca´, “restinga,” sand dune, clystiform spots near margin almost mirroring 11Ð12-X-1985, coll. SE. Miller (USNM, slide 113,540), those on dorsal surface with metallic silver or (USNM). PARATYPES: BRAZIL: 1 male Rio de Ja- black scales present. Posteroventral ßange of neiro, Maraca´, 5 m, 11-X-1986, coll. V. O. Becker, sacculus broadly or acutely pointed (Fig. 15). (USNM, slide 112,922); 2 males (USNM), same data; Signa smaller, length less than distance between 1 male (USNM), same locality, 50 m, 29-I-1985, V. O. signa, clearly independent of each other (Fig. Becker. Paratypes at the USNM and BMNH. 18)...... Cosmopterosis jasonhalli Solis Distribution. This species is known only from Þve 3Ј. Hindwing ventral surface without cataclysti- males collected in and near Maraca´ in the state of Rio form spots. Posteroventral ßange of sacculus de Janeiro, Brazil. broadly spatulate (Fig. 16). Signa larger, Biology. Unknown. length at least 1.5X distance between signa, Etymology. hispidus (Latin, adjective) ϭ bristly; a sometimes looking conßuent anteriorly (Fig. feminized adjective describing the brushlike lateral 19)...... Cosmopterosis spatha Solis projections forming the sclerotized dorsal extension from the base of the juxta.

Cosmopterosis hispida Solis, New Species Cosmopterosis jasonhalli Solis, New Species (Figs. 1, 7, 12, 14) (Figs. 2 and 5, 11, 15, 18)

Diagnosis. Males of C. hispida are distinguished Diagnosis. Externally, this species can only be rec- from other species of Cosmopterosis by the coloration ognized by comparison and exclusion and is most basal to the transverse area of the postmedial line and easily identiÞed using the key to species. It shares with anterior to the longitudinal portion of the postmedial its sister species, C. spatha, the absence of a patch of line (Fig. 1). This area is wholly colored yellow and erect setae distal to the apex of the discal cell on the brown, with no white scales. All other species have male forewing as well as the absence of the diagnostic white scales posteriorly in this area. coloration of the forewing of C. hispida and the hind- Adult. Head (Fig. 1). Proboscis with basodorsal wing of C. thetysalis (cf. Figure 2 with Figs. 1 and 3, and scales mostly pale reddish brown with few creamy 4). Specimens can be distinguished from C. spatha by white. the presence of cataclystiform spots on the lower Thorax (Fig. 1). Patagium brown medially, with surface of the hindwing, which are absent in C. spatha. lateral margins creamy to yellowish white. Tegula Adult. Head (Fig. 2). Frons pale gray ventrally. mostly brown basally, creamy white centrally, medi- Proboscis with basodorsal scales grayish brown. ally and laterally with reddish brown-tipped scales. Thorax (Fig. 2). Patagium creamy white, brown Forewing (Fig. 1). Above, apical two thirds mostly medially. Tegula mostly brown mixed with some rufous; most of costa and basal one fourth creamy to creamy white scales. yellowish white with brown-tipped scales scattered Forewing (Fig. 2). Above, costa and apical one third throughout; basal, subbasal, antemedial, and medial golden yellow; basal one fourth ocherous to very pale lines obscured by scattered brown-tipped scales; area brown; basal, subbasal, and antemedial lines diffuse, basal to anterior portion of postmedial line yellow and with brown-tipped scales, but indicated by pale brown brown with no white scales. Male with another patch intersections with costa; medial line pale brown to of erect scales beyond appressed scales. Ventral sur- brown at costa and posteriorly to M1, fuscous to hind face with costa and apical one Þfth pale brown. margin; area between antemedial and medial lines Hindwing (Fig. 1). Above, subbasal and medial lines rufous; area basal to anterior portion of postmedial line not evident, area between with subbasal rufous spot; yellow dorsally, white toward middle and posteriorly. postmedial and subterminal lines evident as more Ventral surface with costa and termen reddish yellow, densely spaced, brown-tipped scales. Below, medial area between dorsal postmedial and medial lines yel- line not visible, postmedial line pale brown, cataclys- lowish white. tiform spots distinct, mostly silver with narrow black Hindwing (Fig. 2). Above subbasal and medial lines margin apically and broader black margin basally. composed of diffuse brown-tipped scales; area be- 776 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 5 tween rufous; postmedial and subterminal lines com- 2 males, 7 females (USNM), Rondoˆnia, Cacaulaˆndia, posed of diffuse brown-tipped scales, slightly ob- 140 m, XI-1991, coll. V. O. Becker; 1 female (USNM, scured by adjacent scales of similar color. Below slide 112,921), same data (USNM); 3 males, 1 female, medial line and postmedial line obscure, cataclysti- (USNM), same data, XI-1994; 1 female (USNM), same form spots visible. data, 28Ð31-III-1994; 2 females (USNM), same data, Abdomen (Fig. 2). Male tergite I with pair of sub- 13Ð31-XII-1997; 1 male (USNM, slide 113,543), 1 fe- medial brown spots. Female tergite I yellowish silver male (USNM), same data, 15Ð18-X-1993; 1 female with transverse stripe of brown-tipped scales, tergites (USNM), Rondoˆnia, 62 km S Ariquemes, Fazenda IIÐVIII mostly pale brown with apical margins creamy Rancho Grande, 165 m, 10Њ 32Ј S, 62Њ 48Ј W, 14Ð white, posterolateral corner of venter VII with patch 25.XI.1993, coll. R. Leuschner; 1 female (USNM), same of distinctly longer, more robust setae with enlarged data, 29-XÐ10-XI-1991. COSTA RICA: PROVINCIA DE sockets. GUANACASTE: 1 female (INBIO, 90-SRNP-1227, Male Genitalia (Fig. 15). Juxta with deep median slide 113,551), Area de Conservacion Guanacaste, Sec- channel that opens posterodorsally to ßuted, biÞd ap- tor Santa Rosa, Ojochal, 10.78506Њ N, 85.6637Њ W, 10 m, erture at anellus; anellus bordered laterally by mem- 7-VII-1990, feeding on Capparis frondosa (Cap- branous patch with bristlelike setae; sacculus robust, paraceae); 2 males (INBIO, CRI000563858, slide posterior surface deeply concave subapically, medial 563,858; CRI000563866, slide 563,866), Parque Nacio- surface of subapex with broad strip of short spicules, nal Santa Rosa, Playa Naranjo, L-N-309300-354200, apex with sharply pointed, posteroventral ßange. V-1991, coll. E. Alcazar; 1 female (INBIO, Female Genitalia (Fig. 18). Ostium bursae poste- INB0003166516, slide 113,552), Parque Nacional Palo riorly subequal in width to posterior margin of seg- Verde, Sector Palo Verde, Bagaces, L-N-260952- ment VII, rugose on all surfaces; cervix bursae 0.5ϫ 385020, 0Ð50 m, 3Ð8-IV-2000, de Luz, coll. H. Me´ndez; width of ostium bursae, with weakly sclerotized ring 1 female (INBIO, CRI0003118596, slide 113,553), anteriorly, rugose and Þnely spiculate dorsally and Parque Nacional Palo Verde, Sector Palo Verde, Bag- laterally. aces, L-N-260952-385020, 0Ð50 m, 2Ð6-VI-2000, de Larva (Fig. 5). Preserved larvae not available; larval Luz, H. Me´ndez; 1 female (INBIO, INB0003485867, description based on photographs (see Janzen and slide 113,554), Z. P. Nosara, Fila Maravilla, L-N- Hallwachs 2005). Last instar cylindrical, tapered an- 221350-381700, 800 m, 9Ð14-V-2002, Trampa de Luz, teriorly and posteriorly, head dark orange, and A8Ð coll. H. Me´ndez; 2 males (INBIO, 91-SRNP-1836.1, A10 dorsally orange. Black dorsally to just above SD1, slide 113,556; 91-SRNP-1836.2, slide 113,555), 1 female with an interrupted white subdorsal line. Primary se- (INBIO, 91-SRNP-1835), Area de Conservacion tae arising from distinct pinacula or chalazae (D and Guanacaste, Sector Santa Rosa, Las Mesas, 10.84653Њ SD groups); D2 on white chalazae, larger on A1, A3, N, 85.60915Њ W, 305 m, 16-VII-1991, reared from Cap- A5, and A7, smaller on T2, T3, A8, and A9. Yellow, paris flexuosa (Capparaceae); 1 male (INBIO, 90- lateral stigmatal line from prothorax to A7, transparent SRNP-1740, slide 113,557), Area de Conservacion ventrally looking greenish. Guanacaste, Sector Santa Rosa, Bosque San Emilio, Variation. The ventral surface of the hindwing is 10.84389Њ N, 85.61384Њ W, 300 m, 31-VII-1990, reared marked with distinct cataclystiform spots that mirror from Capparis frondosa Jacq. (Capparaceae); 4 males those on the dorsal surface, but on some specimens (INBIO, 01-SRNP-12685, 01-SRNP-12686, 01-SRNP- these are faint to almost completely absent, reduced to 12690, 01-SRNP-12698); 2 females (INBIO, 01-SRNP- only a hint of silvery scales. The apex of the posterov- 12692), Area de Conservacion Guanacaste, Sector entral ßange of the male sacculus ranges in shape from Santa Rosa, Bosque San Emilio, 10.84389Њ N, 85.61384Њ an acutely sharp point, to a longer, bluntly rounded W, 300 m, 1-VI-2001, feeding on Capparis frondosa terminus; however, despite the broad distribution of (Capparaceae); 1 female (INBIO, 01-SRNP-12691, the species, no other character or distributional dif- slide 113,559), same data; 1 male (INBIO, 01-SRNP- ferences were found to consistently covary with the 12699, slide 113,558), same data; 1 female (INBIO, shape of the ßange. 04-SRNP-13837, slide 113,570), Area de Conservacion Material Examined. Forty-Þve males, 73 females. Guanacaste, Sector Santa Rosa, Bosque San Emilio, Type Material. HOLOTYPE: 1 male, INBIO, 90- 10.84389Њ N, 85.61384Њ W, 300 m, 3-VIII-2004, feeding SRNP-1227.1, COSTA RICA, Guanacaste, Parque Na- on Capparis frondosa (Capparaceae), coll. K. Araya; 1 cional Santa Rosa, Ojochal, 10 m, 10.78506Њ N, 85.6637Њ male (INBIO, CRI001115178, slide 103,726), Santa W, 7-VII-1990, reared from Capparis frondosa Rosa National Park, 10Ð12-VII-1980, D. H. Janzen and (Capparaceae), (USNM slide 113,113), (INBIO). W. Hallwachs; 1 female (INBIO, CRI001115159, slide PARATYPES: ARGENTINA: Formosa, 2 females 113,560), Santa Rosa National Park, 300 m, 1Ð15-VIII- (USNM, slide 113610), Riacho Pilaga´, 5Ð19-II-1990. 1982, colls. D. H. Janzen and W. Hallwachs; 1 female L. A. Pereira. BRAZIL: 1 male (USNM, slide 113,544), (INBIO, 94-SRNP-328, slide 113,561), Area de Con- Bahia, Camaca, 400Ð700 m, 21Ð30-IX-1991, coll. V. O. servacion Guanacaste, Sector Santa Rosa, Bosque Hu- Becker; 1 female (USNM, slide 113,545), Bahia, Jequie´, medo, 10.85145Њ N, 85.60801Њ W, 290 m, 18-I-1994, 500 m, 15-IV-1992, coll. V. O. Becker; 1 male (USNM, feeding on Capparis frondosa (Capparaceae); 1 male slide 113,542), 1 female (USNM, slide 113,541), Mato (INBIO, 01-SRNP-12682, slide 113,563); 3 females Grosso do Sul, Corumba, 600 m, 20Ð22-IV-1985, coll. (INBIO, 01-SRNP-12674, 01-SRNP-12675, 01-SRNP- V. O. Becker (USNM); 1 female (USNM), same data; 12680), Area de Conservacion Guanacaste, Sector September 2009 SOLIS ET AL.: PHYLOGENETIC ANALYSIS OF Cosmopterosis 777

Santa Rosa, Sendero Natural, 10.83575Њ N, 85.61253Њ 25Њ 53ЈS, 57Њ 08Ј W, 13Ð14-III-1986, colls. M. Pogue and W, 290 m, 1-VI-2001, feeding on Capparis frondosa M. Solis; 1 female (USNM), Departamento de Para- (Capparaceae); 1 male (INBIO, 01-SRNP-12679, slide guari, 26.2 km SE Ybycui, Parque Nacional Ybycui, 26Њ 113,562), same data; 1 female (INBIO, 01-SRNP- 07Ј S, 56Њ 47Ј W, 15Ð18-III-1986, M. Pogue and M. Solis. 15771), same data, 18-VII-2001; 1 female (INBIO, 01- PERU: 1 male (USNM), coll. Dognin; 1 male (USNM), SRNP-16131, slide 113,564), same locality, 30-VII- Madre de Dios, Rio Tambopata Res, 30 air km. SW Pto. 2001, feeding on Capparis flexuosa (L.) L. Maldonado, 290 m, 6Ð10-XI-1979, subtropical moist (Capparaceae), coll. D. Garcia; 1 female (INBIO, forest, J. B. Heppner; 2 females, same data, 11Ð15-XI- CRI001115182, slide 113,566), W of Carmona Nicoya, 1979; 1 male, 1 female, same data, 16Ð20-XI-1979; 1 600Ð700 m, 19-VIII-1982, colls. D. H. Janzen and W. female, same data, 21Ð25-XI-1979. REPUBLIC OF Hallwachs; 1 female (INBIO, CRI001115184, slide TRINIDAD AND TOBAGO: 1 male, 1 female 113,565), same data. PROVINCIA DE PUNTARENAS: (USNM), Montserrat, coll. W. Schaus. VENEZUELA: 1 female (INBIO, CRI000345554, slide 113,568), 1 male (USNM), 1 female (USNM), Guarico, Hato Reserva Biolo´gica Carara, Estacio´n Quebrada Bonita, Masaguarai, 45 km S Calabozo, 8.57Њ N, 67.58Њ W, 50 m, VI-1991, L-N-194500, 469850, coll. R. Zuniga; 1 Gallery Forest 9, 75 m, 10-VII-1989, uv light, colls. M. female (CRI000345646, slide 113,567), same data; 1 Epstein and M. Deza; 1 male (USNM, slide 113,550), male (INBIO, CRI000127283, slide 127,283), same 1 female (USNM, slide 113,549), same data; 1 male data, X-1989; 1 male (INBIO, CRI001655645, slide (USNM), same locality, Gallery Forest 8, 8Ð9-VII- 655,645), same data, IX-1993, coll. J. Saborio; 1 female 1989, light, coll. M. Epstein; 1 male (USNM), same (INBIO, CRI002409749), Estacio´n San Miguel Send- locality, Gallery Forest 6, 20Ð21-IX-1990, uv and mv ero Mirador, 120 m, L-N-174350-411450, 3-XÐ2-XI- light, colls. M. Epstein and J. Wilterding III; 1 female 1997, coll. F. Alvarado, Trampa de Luz; 1 female (USNM), same locality, Gallery Forest 4, 29-VI-1989, (INBIO, CRI001115179, slide 113,569), Parque Nacio- uv light, colls. M. Epstein and M. Deza; 1 female nal de Corcovado, Osa Peninsula, Sirena, 10Ð12-VIII- (USNM), same locality, Gallery Forest 10, 23Ð24-IV- 1980, colls. D. H. Janzen and W. Hallwachs. GUATE- 1988, uv light, colls. M. Epstein and R. Blahnik; 1 MALA: 2 males, 3 females (USNM), Cayuga, IV, colls. female (USNM), same locality, Gallery Forest 20, 13Ð Schaus and Barnes; 1 male (USNM), same locality and 16-V-1988; 2 males, 1 female (USNM), Aroa, coll. W. collectors, IX; 1 female (USNM), same locality Schaus; 1 male (USNM) Valera, XI-1922, coll. H. Pit- and collectors, II; 1 female (USNM), same locality and tier; 2 females (USNM), same locality, coll. E. P. de collectors, V; 1 female (USNM), same locality and Bellard; 1 male (USNM), Barinas, Rio Caparo Re- collectors, VI; 1 female (USNM, slide 108,146), same search Station, 32 km E El Canton, b light, 3Ð5-II-1978, data. MEXICO: 2 males, 5 females (USNM), Sinaloa, seasonal forest, coll. J. B. Heppner; 1 female (USNM), Presidio [Presidia] River, coll. B. P. Clark; 1 male San Esteban, Carabobo, 1Ð20-XII-1939, coll. P. J. An- (USNM, slide 113,547), 1 female (USNM, slide duse; 1 female (USNM), Aragua, Playa de Cata, 9 km 113,546), same data; 2 females, Sinaloa, Venado [Ve- E El Playon, 21-I-1978, riverine forest, J. B. Heppner; nadio], coll. B. P. Clark; 2 males, 2 females, (SDNHM), 1 female (USNM), Lara, 4 km NW La Pastora, 2Ð3- Sinaloa, Micro. Ej. El Indio, N. Mazatlan, 20-VIII-1985, III-1978, riparian forest, blacklight, coll. J. B. Heppner; Faulkner, BloomÞeld; 1 female (USNM), Nayarit, Islas 1 female (USNM), Zulia, Guasare coal camp, 45 km W. Marias, Isla Maria Magdalena, 20-V-1925, coll. H. H. Carrasquero, Sierra Perija, 19Ð25-VIII-1981, 80Ð140 m, Keifer; 1 male (SDNHM), Nayrarit, Micro. Penitas, nr. J. Heppner. Tuxpan, 17-VIII-1985, Faulkner, BloomÞeld; 1 female One additional female (USNM) from the W. Schaus (USNM), San Luis Potosi, 2 miles N Tamazunchale, collection with the labels reading “52” and “Trillista 400 feet, 16Ð18-VII-1963, colls. Duckworth and Davis. thetysalis W. comp. B.” in hand-written ink has no PANAMA: 2 males (USNM), Cerro Campana, nr. locality label. Paratypes deposited at BMNH, INBIO, Chica, 2Ð5-IV-1965, colls. S. S. Duckworth and W. D. and USNM. Duckworth; 1 male (USNM), Barro Colorado Island, Distribution. Sinaloa, Mexico south to Paraguay, 10Ð17-V-1964, W. D. Duckworth and S. S. Duckworth; Peru to Venezuela, and on the islands of Trinidad and 1 female (USNM), same data, 1Ð9-V-1964; 1 female Tobago. (USNM), same data, 18Ð28-IV-1964; 1 female (USNM, Biology. Several larvae collected in Costa Rica were slide 110,135), same data; 2 males (USNM), same lo- reared from Capparis frondosa Jacquin and Capparis cality, at light, lot no. 42-4149, 20-III-1942, coll. J. flexuosa (L.) L. (Capparaceae) (Janzen and Hallw- Zetek; 1 male (USNM, slide 110,054), 1 female achs 2005). The adult ßy year-round and are (USNM, slide 110,055), same data, lot no. 41-17178, found between 50 and 900 m. 26-IX-1941; 1 female (USNM), same locality, at light, Etymology. A patronym honoring the spouse and 4-III-1979, colls. Silberglied and Aiello; 1 male good friend of the Þrst author, Dr. Jason P. W. Hall, an (USNM), Alhajuelo, 14-IV-1911, coll. A. Busck; 1 fe- expert on the Riodinidae. male (USNM), San Lorenzo, 9Њ 17Ј N, 79Њ 58Ј W, 22.X.2003, coll. R. L. Kitching. PARAGUAY: 1 female Cosmopterosis spatha Solis, New Species (USNM, slide 113,548), Departamento de Concep- (Figs. 3 and 6, 8, 16, 19, 21, 22, 23) cio´n, Arroyo Tagatiya-mi, 22Њ 39Ј S, 57Њ 32Ј W, 3Ð5- IV-1986, colls. M. Pogue and M. Solis; 1 female Diagnosis. Specimens of this species lack cataclys- (USNM), Departamento de Paraguari, Cerro Acahay, tiform spots on the lower surface of the hindwing. 778 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 5

Adult. Head (Fig. 3). Frons grayish brown ventrally. dible with one row of serrations. T1-3 and A1-10 in- Proboscis with basodorsal scales mostly grayish brown tegument smooth, all segments with prominent cha- with few creamy white. lazae of D and SD setal groups and small pinacula, Thorax (Fig. 3). Patagium creamy to yellowish setae with concolorous pinacula. Prothoracic shield white. Tegula mostly brown basally grading to creamy approximately twice as broad as long, anterior to lat- white laterally and posteriorly. eral margin ridged and forming an L-shaped sclero- Forewing (Fig. 3). Above, costa and apical one third tized pattern from lateral view, D2 2.5ϫ as long as D1, golden yellow; basal one fourth ocherous to very pale XD1, XD2, SD1, and SD2 located on prominent cha- brown; basal, subbasal, and antemedial lines diffuse, lazae, XD1 very short, SD1 3ϫ as long as SD2; L group with brown-tipped scales, but indicated by pale brown bisetose, anterior to spiracle, L1 and L2 on same elon- intersections with costa; medial line pale brown to gate pinaculum; SV group bisetose, on different small Ϸ ϫ brown at costa and posteriorly to M1, fuscous to hind pinacula. T2-T3 with developed chalaza, D2 2 as margin; area between antemedial and medial lines long as D1; SD group bisetose on similar chalaza as D rufous; area basal to anterior portion of postmedial line group; L group trisetose, L2 on larger pinaculum, Ϸ3ϫ yellow dorsally, white toward middle, often with ru- as long as L1 and L3; SV1 present, shorter than L2; V fous or brown scales through the middle. Ventral sur- seta short. A1ÐA2 with D group very short on separate face with costa and termen reddish yellow, area be- small pinacula; SD group unisetose, SD1 as long as L2 tween dorsal postmedial and medial lines yellowish of T2Ð3; L group bisetose, L2 1.5ϫ as long as L1; SV2 white. ventrad to SV 1, approximately equal in length; V1 Hindwing (Fig. 3). Above subbasal and medial lines present, approximately one third of SV in length. Setal composed of diffuse, brown-tipped scales; area be- map of A3-6 in general similar to A1-2, but D and SV tween rufous; postmedial and subterminal lines com- group located on chalazae, L3 present, SV group tri- posed of diffuse brown-tipped scales, slightly ob- setose at base of prolegs. A7 with L2 much longer than scured by adjacent scales of similar color. Below those on A3-6. D1 and D2 on A8 located on conical medial line and postmedial line pale brown, cataclys- chalazae; L group trisetose, nearly equal in length. A8 tiform spots absent. and A10 dorsally orange. A9 with D2 on large conical Abdomen (Fig. 3). Male tergite I with pair of sub- chalaza; SD group unisetose; L group unisetose. Anal medial brown spots. Female tergite I yellowish silver shield on A10 with prominent D and SD chalazae and with transverse stripe of brown-tipped scales, tergites D2, SD1 and SD2 at outer margin of anal shield. Pro- IIÐVIII mostly pale brown with apical margins creamy legs with crochets triordinal in a mesal penellipse. white, posterolateral corner of venter VII with patch Pupa (Figs. 21, 22). Segments digitally enhanced of distinctly longer, more robust setae with enlarged with black lines. Ventral view: Truncate top of head sockets. with lateral protruded clypeus; labrum narrow; pilifers Male Genitalia (Fig. 16). Juxta with deep median slightly visible; maxillary palpi absent; labial palpi channel that opens posterodorsally to ßuted, biÞd ap- barely visible; maxilla extending to A4; forefemur vis- erture at anellus; anellus bordered laterally by mem- ible, prothoracic legs (foreleg) three-quarters length branous patch with peglike setae; sacculus robust, of forewing, reaching to middle of A3; mesothoracic posterior surface deeply concave subapically, medial legs (midleg) extending beyond hind margin of A4, surface of subapex with narrow strip of short spicules, slightly longer than antennal sheath; hind tarsi not apex with broadly spatulate, posteroventral ßange. visible; A5 and A6 with vestigial prolegs; A8-10 with Female Genitalia (Fig. 19). Ostium bursae poste- genital oriÞce anteriorly; anus posteriorly with lateral riorly subequal in width to posterior margin of seg- conical depressions. Dorsal view: prothorax smooth; ment VII, rugose ventrally and laterally; cervix bursae mesothorax without seta; all spiracles on abdominal 0.5ϫ width of ostium bursae, with weakly sclerotized segments located on developed conical protuberanc- ring anteriorly, rugose and Þnely spiculate dorsally es; surface of all abdominal tergites and wing sheath and laterally. granulate; paired lateral protuberance present along Larva (Figs. 6, 23). Last instar cylindrical, tapered margin of A9; cremaster somewhat ßattened dorso- anteriorly and posteriorly. Thick and extremely long ventrally with sharp bifurcate apex and four pairs of setae arising from distinct pinacula or chalazae of D curved setae. and SD groups (Fig. 23). Black dorsally to just above Material Examined. Fifty-nine males, 56 females. SD1 except T2, A1, A3, A5, A7 with only posterior half HOLOTYPE: 1 male, COSTA RICA, Guanacaste, black. Yellow lateral stigmatal line from prothorax to (INBIO, 02-SRNP-33785), Area de Conservacion A7, transparent ventrally looking greenish. Head hy- Guanacaste, Sector Del Oro, Puente Mena, 11.04562Њ pognathous, dark orange; epicranial suture present; N, 85.45742Њ W, 280 m, 1-XII-2002, feeding on Capparis frontoclypeus and labrum dark brown, Ϸ1.25ϫ as long mollicella (Capparaceae), coll. L. Rõ´os, (USNM slide as wide; six stemmata, 1 and 2 approximate, 3 and 4 113,114) (INBIO). approximate, 5 anterio-ventrad to 6; S2 1.3ϫ as long as PARATYPES. COSTA RICA: ALAJUELA: 1 female S1, SS3 longer than SS2, MG1 short, F1 present near (USNM slide 113,571), Reserva Biologica Alberto middle of frons, AF1 and AF2 on adfrontal area rather Manuel Brenes, San Ramon, Alrededores Estacio´n short, La and Aa present, P1 longer than P2, P2 base Biologica, L-N-470400-244400, 850 m, 7Ð11-II-2005, slightly elevated; labrum with three setae on each side coll. J. B. Sullivan; GUANACASTE: 1 male (INBIO, externally and no seta on each side internally; man- 00-SNRP-11271), 1 female (INBIO, 00-SRNP-11215, September 2009 SOLIS ET AL.: PHYLOGENETIC ANALYSIS OF Cosmopterosis 779 slide 113,572), Area de Conservacion Guanacaste, Sec- 3047, 98-SRNP-3048, 98-SRNP-3054, 98-SRNP-3057, tor San Cristobal, Potrero Argentina, 520 m, 10.89021Њ 98-SRNP-3058, 98-SRNP-3066, 98-SRNP-3075), 12 fe- N, 85.38803Њ W, 26-V-2000, feeding on Capparis mol- males (INBIO, 98-SRNP-3038, 98-SRNP-3043, 98- licella (Capparaceae), coll. F. Quesada; 1 male SRNP-3049, 98-SRNP-3050, 98-SRNP-3051, 98-SRNP- (INBIO, 00-SRNP-11249, USNM slide 113,573) same 3053, 98-SRNP-3059, 98-SRNP-3061, 98-SRNP-3064, data; 2 males (INBIO, 02-SRNP-220, 02-SRNP-262), 2 98-SRNP-3065, 98-SRNP-3067, 98-SRNP-3070), same females (INBIO, 02-SRNP-285, 02-SRNP-286), Area data, coll. H. Ramirez; 1 male (INBIO, 98-SRNP-3071, de Conservacion Guanacaste, Sector San Cristobal, USNM slide 113,582), same data; 1 male (INBIO, 98- Potrero Argentina, 520 m, 10.89021Њ N, 85.38803Њ W, SRNP-3063, USNM slide 113,585), same data; 1 female 16-I-2002, feeding on Capparis mollicella (Cap- (INBIO, 98-SRNP-3042, USNM slide 113,583), same paraceae), coll. O. Espinoza; 1 male (INBIO, 03- data; 1 male (INBIO, CRI000356904), 1 female SRNP-6592), 2 females (INBIO, 03-SRNP-6593, 03- (INBIO, CRI000356824), Parque Nacional Guana- SRNP-6607), same data, 13-VI-2003; 1 female (INBIO, caste, Lado SO Volcano Cacao, Estacio´n Cacao, L-N- 03-SRNP-6604), same data, 8-VI-2003, coll. C. Cano; 1 323300-375700, 1,000Ð1,400 m, IX-1991, col. C. Chaves; female (INBIO, 04-SRNP-618), Area de Conservacion 1 male (INBIO, 02-SRNP-33771, USNM slide 113,586), Guanacaste, Sector San Cristobal, Corrales Viejos, 1 female (INBIO, 02-SRNP-33777, USNM slide 495 m, 10.89974Њ N, 85.38085Њ W, 27.I.2004, feeding on 113,590), Area de Conservacion Guanacaste, Sector Capparis mollicella (Capparaceae), coll. E. Araya; 2 Del Oro, Puente Mena, 11.04562Њ N, 85.45742Њ W, males (INBIO, 03-SRNP-5567, 03-SRNP-5572), 3 fe- 280 m, 1-XII-2002, feeding on Capparis mollicella males (INBIO, 03-SRNP-5563, 03-SRNP-5568, 03- (Capparaceae), coll. L. Rõ´os; 1 female (INBIO, 02- SRNP-5578), same data, 21-I-2003, feeding on Cappa- SRNP-33769, USNM slide 113,589), same data; 2 males ris mollicella (Capparaceae), coll. O. Espinoza; 1 male (INBIO, 02-SRNP-27630, 02-SRNP-27633), 2 females (INBIO, 03-SRNP-5566, USNM slide 113,580), same (02-SRNP-27641, 02-SRNP-27642), Area de Conser- data; 1 male (INBIO, 03-SRNP-5573, slide 113,581), vacion Guanacaste, Sector Del Oro, Camino Mangos, same data; 3 females (INBIO, 03-SRNP-11680.1, 03- 11.00766Њ N, 85.47926Њ W, 480 m, 27-VII-2002, feeding SRNP-11680.21, 03-SRNP-11680.30), Area de Conser- on Capparis mollicella (Capparaceae), coll. L. Rõ´os; 1 vacion Guanacaste, Sector San Cristobal, Vado Rio male (INBIO, 02-SRNP-27629, USNM slide 113,576) Cucaracho, 640 m, 10.8702Њ N, 85.39153Њ W, 11-VII- same data; 1 female (INBIO, CRI002135396, slide 2003, feeding on Capparis mollicella (Capparaceae), 113,597), Estacio´n Pitilla, 9 km S de Santa Cecilia, coll. J. Perez; 3 males (INBIO, 03-SRNP-11036, 03- L-N-329950-380450, 700 m, II-1995, coll. P. Rios; 1 male SRNP-11051, 03-SRNP-11085), 3 females (INBIO, 03- (INBIO, CRI002210640), same locality, VI-1995, coll. SRNP-11048, 03-SRNP-11053, 03-SRNP-11091), same C. Moraga; 1 male (INBIO, CRI001115615), 1 female data, 24-V-2003; 2 males (INBIO, 03-SRNP-11348, 03- (INBIO, CRI001115181, slide 113,594), Estacio´n SRNP-11353), 1 female (INBIO, 03-SRNP-11355, Mengo, SW side Volcan Cacao, 1,100 m, 13Ð26-VI- USNM slide 113,591), same data, 11-VI-2003; 1 male 1987, colls. D. H. Janzen and W. Hallwachs. LIMON: (INBIO, 03-SRNP-11360, USNM slide 113,579) same 1 male (INBIO, CRI001641981), Sector Cerro Cocori, data; 2 males (INBIO, 03-SRNP-11256, 03-SRNP- Finca de E. Rojas, L-N-286000-567500, 150 m, X-1993, 11259), 1 female (INBIO, 03-SRNP-11254), same data, coll. E. Rojas; 1 male (INBIO, CRI001879644), A. C. 9-V-2003, coll. M. Carmona; 1 male (INBIO, 01-SRNP- Tortuguero, Sector Cocori, 30 km N. de Cariari, Finca 3455), 3 females (INBIO, 01-SRNP-3452, 01-SRNP- E. Rojas, L-N-286000-567500, 100 m, V-1994, coll. E. 3454, 01-SRNP-3456), Area de Conservacion Guana- Rojas. PUNTARENAS: 1 male (INBIO, caste, Sector San Cristobal, Rio Blanco Abajo, CRI001919250), 1 female (INBIO, CRI001919266, 10.90037Њ N, 85.37254Њ W, 500 m, 15-IX-2001, feeding USNM slide 113,593), A. C. Arenal, Monteverde, San on Capparis mollicella, (Capparaceae), coll. O. Espi- Luis, L-N-250850-449250, 1,040 m, IX-1993, coll. Z. noza; 1 male (INBIO, 01-SRNP-3634, USNM slide Fuentes; 1 female (INBIO, CRI0003554791, USNM 113,574), 1 female (INBIO, 01-SRNP-3625, USNM slide 113,595), Parque Nacional Piedras Blancas, Al- slide 113,575), same data, 27-IX-2001; 1 male (INBIO, rededor de Estacio´n el Bonito, L-S-292700, 548350, 05-SRNP-2559, USNM slide 113,588), 2 females (IN- 100Ð200 m, XI-2002, “Red de Golpe,” coll. M. Moraga; BIO, 05-SRNP-2557, 05-SRNP-2560), same data, 6-V- 1 female (INBIO, CRI001190187, USNM slide 2005; 1 female (INBIO, 05-SRNP-2558, USNM slide 113,596), Peninsula de Osa, Rancho Quemado, L-S- 113,592), same data; 1 male (INBIO, 05-SRNP-2561, 292500, 511000, 200 m, IX-1991, coll. F. Quesada; 1 USNM slide 113,587), same data. GUANACASTE: 11 male (INBIO, CRI000483369), same locality, XII-1991, males (INBIO, 98-SRNP-3016, 98-SRNP-3017, 98- coll. F. Quesada; 1 male (INBIO, CRI001110125), Osa SRNP-3018, 98-SRNP-3020, 98-SRNP-3022, 98-SRNP- Peninsula, Corcovado National Park, Sirena, 1-V-1984, 3024, 98-SRNP-3028, 98-SRNP-3030, 98-SRNP-3033, colls. D. H. Janzen and W. Hallwachs; 1 female 98-SRNP-3034, 98-SRNP-3036), 3 females (INBIO, 98- (INBIO, CRI001115185), same data, 10Ð12-VIII-1980; SRNP-3023, 98-SRNP-3026, 98-SRNP-3029), Area de 1 male (INBIO, CRI000946789), Reserva Biologica Conservacion Guanacaste, Sector Cacao, Sendero Monteverde, Estacio´n La Casona, L-N-253250-449700, Nayo, 10.92446Њ N, 85.46953Њ W, 1,090 m, 14-VI-1998, 1,520 m, IX.1992, coll. N. Obando; 1 male (INBIO, feeding on Forchhammeria trifoliata (Capparaceae), CRI000992268), Peninsula de Osa, Fila Draque, L-N- coll. M. Pereira; 11 males (INBIO, 98-SRNP-3040, 98- 294000-508200, 5-IV-1992, coll. F. Quesada. Paratypes SRNP-3041, 98-SRNP-3044, 98-SRNP-3045, 98-SRNP- deposited at BMNH, INBIO, and USNM. 780 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 5

Distribution. Costa Rican provinces of Alajuela, Type Material. HOLOTYPE: 1 female (BMNH, ex- Guanacaste, Limon, and Puntarenas. amined): BRAZIL, Rio Amazonas, Villa Nova Biology. Several larvae were collected and reared [Parintins], coll. H. W. Bates. Additional specimens from Capparis mollicella (Capparaceae), as well as examined. FRENCH GUIANA: 1 male (USNM), Ca- Forchhammeria trifoliata (Capparaceae), (Janzen and cao, 30-III-1986, coll. C. V. Covell, Jr.; 1 male (USNM), Hallwachs 2005). The adults moths ßy year-round and Cayenne, coll. W. Schaus; 2 females (USNM), Saint have been collected between 50 m and 1,600 m. Jean, Saint Laurent du Maroni, coll. W. Schaus; 2 males Etymology. spatha (Latin, noun, feminine) ϭ broad (USNM), Saint Laurent du Maroni, coll. Dognin. blade; a noun in apposition describing the broadly SURINAME: 1 male (BMNH slide 21,198), 1 female spatulate, posteroventral ßange on the sacculus of the (BMNH slide 21,199), Maroewym Valley, Aroewarwa male. Creek, June 1905, coll. S. M. Klages. VENEZUELA: 1 female (USNM), Amazonas, Rio Orinoco. Distribution. Southern Venezuela and northwest Cosmopterosis thetysalis (Walker) Brazil north to the coast of Suriname and French (Figs. 4, 7, 17, 20) Guiana. Cataclysta thetysalis Walker, 1859: 440; Amsel 1956: Biology. Unknown. 110. 1 female (USNM slide 113,539) from BRAZIL (Ba- hia, Jequie´, 500 m, 15-IV-1992, coll. V. O. Becker) may Diagnosis. This species can be recognized by the represent an additional species, but more material is distinct pattern of the cataclystiform spots on the needed before we can be certain. upper hindwing (Fig. 4). The middle of each spot has brown scales the same color as those of the adjacent Discussion area of the hindwing rather than the silver and black invaginated pattern present in the other species. There has been signiÞcant progress in the last two Adult. Head (Fig. 4). Setiform scales on vertex apex decades on the study of Lepidoptera, but many aspects yellow. about their biology and morphology remain poorly Thorax (Fig. 4). Patagium uniformly creamy white. known. In Costa Rica progress has been made by Tegula reddish brown laterally and medially, but D.H.J. and W. Hallwachs where a massive rearing creamy white centrally. effort has resulted in data about immature stages and Forewing (Fig. 4). Above, costa and apical one third in long series of adult specimens of Lepidoptera. Such golden yellow; basal one fourth ocherous to very pale studies, and other similar studies all over the world, brown; basal, subbasal, and antemedial lines diffuse, expand our knowledge about species: we learn what it with brown-tipped scales, but indicated by pale brown looks like during its many early stages of life, what it intersections with costa; medial line pale brown to is doing in its , what it eats, and perhaps what brown at costa and posteriorly to M1, fuscous to hind eats it. In addition, the morphological and biological margin; area between antemedial and medial lines data that the immature stages provide has resulted in rufous; area basal to anterior portion of postmedial line more robust phylogenetic studies. The long series of yellow dorsally, white toward middle. Male with an- specimens collected in the same locality has opened other patch of erect scales beyond appressed scales. the doors to the study of intraspeciÞc variation in Hindwing (Fig. 4). Above subbasal spot obscure, adults and the discovery of behaviorally interesting medial line composed of diffuse rufous scales; post- characters such as secondary sexual structures. medial and subterminal lines composed of diffuse Secondary sexual structures such as scent-produc- brown-tipped scales, slightly obscured by adjacent ing structures, associated modiÞed scales, and their scales of similar color. Cataclystiform spots discontin- glands (Brown and Miller 1983, Birch et al. 1990) uous centrally, scales the same color as adjacent area provide a window into the diversity of lepidopteran of hindwing. Below medial line and postmedial line adult reproductive behavior. Lepidopteran behavior, pale brown, vestiges of cataclystiform spots, black glands, and structures such as coremata, androconia, scales visible. hairpencils, StobbeÕs gland, brushes, alar organs, and Abdomen (Fig. 4). Male tergite I with pale brown- osmophores are succinctly summarized by Hallberg tipped scales medially. and Poppy (2003). In the past male secondary sexual Male Genitalia (Fig. 17). Juxta with deep median structures on various parts of the body were thought channel that opens posterodorsally to ßuted, biÞd ap- to be too complex and too variable to provide any erture at anellus; sacculus slightly thickened, surface phylogenetic information above the species level. Re- smooth, lacking denticles, apex with narrow posterov- cent articles have shown that they can be used to entral ßange 6ϫ longer than wide. deÞne taxa at different levels, for example, at the Female Genitalia (Fig. 20). Ostium bursae poste- superfamily level in Yponomeutoidea (Hsu and Pow- riorly 0.5ϫ width of posterior margin of segment VII, ell 2005, Landry 2007), at the subfamily level in Noc- surfaces smooth; cervix bursae 0.75ϫ width of ostium tuidae (Wagner 2007), at the tribal level in Ctenuchini bursae, rugose and Þnely spiculate dorsally and later- (Weller et al. 2000) and Tortricinae (Brown 1990), at ally; left and right signa isolated anteriorly. the tribal, generic, and species level in Riodinidae Immature Stages. Unknown. (Hall and Harvey 2002), at the generic level in noc- Material Examined. Five males, 5 females. tuids (Troubridge and Lafontaine 2007), at the ge- September 2009 SOLIS ET AL.: PHYLOGENETIC ANALYSIS OF Cosmopterosis 781 neric and species level group in Heliodinidae (Hsu with complex basal structures to be an apomorphy of and Powell 2005), and at the species group level in Glyphodes Guene´e group 1 and coremata with simple conjunction with mitochondrial data in Ctenuchini basal structure with broad, long scales, with sculp- (Schneider et al. 1999) and with host data and tured scale tufts as an apomorphy for the monophy- ultrasound production in milkweed tussock moths letic group of Agrioglypta ϩ Talanga. In Musotiminae, (DaCosta et al. 2006). Phillips and Solis (1996) mention specialized thoracic In Pyraloidea, many species have secondary sexual and leg scales at the species level in Neurophyseta structures with specialized scales, singly or in patches, Hampson not found in other musotimines, and Solis et sometimes associated with membranous structures al. (2005) mention a costal forewing swelling or such as folds or hairpencils on various parts of the pocket in Undulambia Lange. body such as antennae, maxillary and labial palpi, alar Glaphyriinae had been historically deÞned by the organs such as pockets, costal wing folds, and forewing presence of spatulate scales on the male hindwing sound producing organs, hairpencils on the thorax, between CuA2 and CuP, but Solis and Adamski (1998) legs, and abdomen, and coremata associated with the illustrated that not all genera had these scales and that eighth abdominal segment. Often this information is these scales could be piliform, spatulate, or a combi- buried in published taxonomic studies and unpub- nation of these scales. In this paper we described the lished taxonomic theses (Cashatt 1968, Clavijo 1990). radiodiscal process, a new secondary sexual character In the , secondary sexual structures in- on the male forewing, but the diversity of the Glaphy- clude alar organs in Chrysauginae (Cashatt 1968) and riinae is such that we suspect many secondary sexual Epipaschiinae (Solis 1993), antennal, maxillary and structures will be discovered to be apomorphies for labial palpi modiÞcations of epipaschiine heads (Solis other genera. 1993), and in the antennal modiÞcations In addition, caterpillar collecting efforts in Costa and abdominal coremata (Heinrich 1956; Horak 1997; Rica (Janzen and Hallwachs 2005) yielded consider- Neunzig 1986, 1990, 1997, 2003; Simonsen 2008). Con- able morphological and ecological data about Cos- trary to prevalent opinion at the time, Janse (1931), mopterosis larvae reared on Capparis L. In the Area de after his outstanding study of epipaschiinae male Conservacion Guanacaste (ACG) in northwestern heads, stated: “I now consider [secondary sexual or- Costa Rica, C. jasonhalli larvae (Fig. 5) were encoun- gans] of so much importance, that I hold that they tered feeding in presumably sibling groups at very low should always be taken into consideration, not only in frequency on the both sides of mature leaves of just deÞning the species, but also in the limiting of a ge- three (C. frondosa, C. flexuosa, and C. discolor)ofthe nus...” A phylogenetic analysis on a monophyletic eight species of Capparis (Capparaceae) that occur in group of genera of the Epipaschiinae of the Western the lowland dry forest that covers the PaciÞc coastal Hemisphere by Solis (1993) corroborated JanseÕs idea plain of the ACG between 0- and 600-m elevation (see that secondary sexual characters could be informative http://janzen.sas.upenn.edu for individual records). at the generic level despite the homoplasy associated Larvae skeletonized the leaf blades and spun light silk with these structures that was often observed super- over the surface of the leaves, but moved about freely Þcially and/or casually from a higher level taxonomic on the leaf surface. There may be 1Ð100ϩ larvae in one perspective. Most recently, Simonsen and Roe (2009) group depending on how much the initial group of investigated the taxonomic signiÞcance of complex siblings has been disturbed. They fed both night and scale brushes associated with the eighth abdominal day, and were very conspicuous in having red-to- sternite in Dioryctria Zeller (Phycitinae). They found orange heads and rears, and strong white-cream dorsal the majority to be homplasious, but eight characters patches against a black background. In ACG, C. jas- were unique apomorphies. onhalli larvae have been found only in dry forest (14 Descriptions of secondary sexual structures in the different larval groups discovered in 26 yr), and the Crambidae exist in taxonomic research papers, but very similar C. spatha was restricted to ACG rain forest there is a paucity of major phylogenetic analyses that and the lower margins of cloud forest (400Ð1,200 m). cite examples of informative secondary sexual char- Like its congener, its conspicuous (Fig. 6) and social/ acters. In the Odontiinae, Leraut and Luquet (1982) gregarious larvae fed on the mature leaves of two described a tribe based on the presence of highly species of Capparis (C. frondosa and C. mollicella) and modiÞed structures associated with the male genitalia on Forchammeria trifoliata Radlk (Capparaceae), an and androconia associated with sternite VIII. In the upper elevation treelet in the lower margins of the , Clavijo (1990) discussed the generic cloud forest (22 different larval groups discovered in distribution (Diaphania Stephens, Omiodes Guene´e, 11 yr). Palpita Hu¨ bner, and Sparagmia Guene´e) of an ane- Adults of C. jasonhalli have been collected with light pisternal scale organ, two clusters of scale-like struc- traps from the semideciduous intermediate elevation tures on the mesothorax. He also proposed the term rain forests near Tamazunchale, S.L.P., Mexico, and “parategumen sclerites” for commonly found lateral the very deciduous dry forest of Isla Maria Magdalena, sclerites located near the base of the tegumen with Nayarit, Mexico, south to the dry forests of Paraguay, long hairpencils or brushes. He described the “anal with the in between ranging from strongly tuft” as specialized scales on the parategumen sclerites deciduous (e.g., Mexican and ACG PaciÞc coastal for- and the eighth abdominal segment and as an apomor- ests) to rain forest (Sirena, Osa Peninsula, Costa Rica) phy for Diaphania. Sutrisno (1999) found coremata to lower cloud forest (Cerro Campana, Panama). This 782 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 5 very broad ecosystem coverage over their entire range on Capparaceae in Costa Rica, a gallmaker and a leaf- stands in strong contrast to the apparent restriction of miner on fruit (M.A.S., unpublished). It is expected C. jasonhalli to dry forest in ACG. Light-caught adults that other species in the pyraloid genera listed above of C. spatha are known only from Costa Rican rain also feed on Capparis or other genera of Capparaceae. forest and lower elevation cloud forest sites, just as are the larvae of C. spatha in ACG. These two species undoubtedly co-occur in the area at 400Ð600-m ele- Acknowledgments vation on the ACG west slopes of the Cordillera Guanacaste where dry forest and rain forest inter- We thank various individuals worldwide who care for grade. But in general terms, C. jasonhalli is the dry Pyraloidea collections and assisted our search for material, forest Cosmopterosis and C. spatha is the rain forest but especially K. Tuck (BMNH), E. J. van Nieurkerken Cosmopterosis. Both are unambiguously specialists on (RMNH), M. Wall (SDNHM), E. Phillips (formerly with Capparaceae. The ACG caterpillar inventory has INBio), and V. O. Becker who provided extensive material reared Ͼ350,000 wild-caught larvae of Ͼ3,000 species from Brazil from his personal collection. We thank E. Araya, C. Cano, E. Cantillano, M. Carmona, O. Espinosa, R. Espi- on thousands of food plant species without encoun- nosa, R. Franco, D. Garcia, W. Hallwachs, C. Moraga, R. tering the very distinctive Cosmopterosis larvae on any Moraga, G. Pereira, M. Pereira, J. Perez, F. Quesada, H. other plant family (as well as being conspicuously Ramirez, L. Rios, P. Rios, G. Sihezar, and F. Vargas for cat- absent from at least Þve other species of Capparis). erpillar hunting and husbandry; R. Espinosa, J. Gonzales, A. Cosmopterosis larvae are rarely attacked by parasi- Guadamuz, B. Hammel, N. Zamora, and the herbarium staff toids. During a 27-yr period, rearing of 738 wild-caught at INBio for foodplant identiÞcations; and S. R. Shaw, J. B. ACG Cosmopterosis caterpillars have yielded only four WhitÞeld and D. M. Wood for wasp and ßy identiÞcations. tachinids (one species in C. jasonhalli) and 12 bra- We thank Shen-Horn Yen (National Sun Yat-Sen University, conids (a species of Hypomicrogaster and a species of Taiwan) and F. Christian Thompson, Tom Henry, John Microchelonus in C. spatha). Brown (SEL, USDA) for helpful suggestions on the manu- script. Special thanks to Thomas Simonsen who provided a Finally, another interesting observation is the pau- rigorous review that only a true morphologist could do. The city of Capparaceae larval feeders in the Lepidoptera. ACG caterpillar inventory has been variously supported by With an upper estimate of Ϸ482,000 lepidopteran spe- National Science Foundation grants BSR 90-24770, DEB 93- cies (Solis and Pogue 1999), larval feeding of Cap- 06296, DEB 94-00829, DEB 97-05072, DEB 00-72730, and DEB paraceae is relatively rare in lepidopteran families and 05-15699 and by Area de Conservacio´n Guanacaste, Consejo restricted to speciÞc genera presumably due to glu- Nacional para Investigaciones CientiÞcas y Technolo´gicas cosinolates that may serve as either detection cues to (CONICIT), INBio, and the government of Costa Rica. the adapted or as a deterrent to unadapted lepidop- terans (Futuyma 1986). The Pieridae is an exception where hundreds of species are known to feed on References Cited Capparaceae, Ϸ90 species on Capparis, and their eco- logical interactions with Capparaceae and Brassi- Amsel, H. G. 1956. Microlepidoptera Venezolana. Bol. En- caceae have been well studied (e.g., Renwick 2001). tomol. Ven. 10: 1Ð336. But there are only 11 known Capparis-feeding species Birch, M. C., G. M. Poppy, and T. C. Baker. 1990. Scents and eversible scent structures of male moths. Annu. Rev. in other lepidopteran families: Cosmopterigidae (one Entomol. 35: 25Ð58. species), Gelechiidae (one species), Gracillariidae Brown, J. W. 1990. Taxonomic distribution and phyloge- (two species), Lymantriidae (one species), Tortrici- netic signiÞcance of the male foreleg hairpencil in the dae (three species), Yponomeutidae (two species), Tortricinae (Lepidoptera: Tortricidae). Entomol. News and Thyrididae (two species) (Robinson et al. 2007). 101: 109Ð116. Capparaceae-feeding in the Pyraloidea has infre- Brown, R. L., and P. R. Miller. 1983. Studies of Lepidoptera quently evolved independently geographically and hindwings with emphasis on ultrastructure of scales in taxonomically in several subfamilies of the Crambidae; Cydia caryana (Fitch) (Tortricidae). Entomography 2: there are no published records of species reared on 261Ð295. Cashatt, E. D. 1968. Revision of the Chrysauginae of North Capparis or Capparaceae in Pyralidae. Published America. Ph.D. dissertation. Catholic University of Amer- records of Capparis-feeding includes Crocidolomia ica, Washington, DC. Zeller (11 species; ; two species reared) Clarke, J.F.G. 1941. The preparation of the slides of the in Asia and Africa, Zerny (four species; genitalia of Lepidoptera. Bull. Brooklyn Entomol. Soc. 36: ; one species reared) in the Middle East 149Ð161. and Russia, and Styphlolepis Hampson (six species; Clavijo A.J.A. 1990. 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