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American Museum Novitates AMERICAN MUSEUM NOVITATES Number 3770, 20 pp. February 22, 2013 Coniopterygidae (Neuroptera: Aleuropteryginae) in amber from the Eocene of India and the Miocene of Hispaniola DAVID GRIMALDI,1MICHAEL S. ENGEL,2 PAUL C. NASCIMBENE,1 AND HUKAM SINGH3 ABSTRACT The genusSpiloconis Enderlein is comprised of six Recent Asian and Pacific species, rang- ing from Sri Lanka (and possibly Madagascar) in the west to eastern Australia and Fiji in the east. Two species previously described in Miocene amber from the Dominican Republic, Spi- loconis glaesaria Meinander and S. oediloma Engel and Grimaldi, are redescribed for some details. Spiloconis eominuta Grimaldi and Engel, new species, is described in earliest Eocene Cambay amber from Gujarat, India, which has well-preserved male terminalia. Neoconis paleo- caribis Grimaldi and Engel, new species, is described from Dominican amber; this Recent genus is known from the southern United States and neotropics. The fossil Spiloconis may have sig- nificant biogeographic implications, but definitive determination of this requires a phylogenetic analysis of the 16 Recent genera in the monophyletic subfamily Aleuropteryginae. A Dominican amber–Australasian distribution pattern is known for 18 genera of insects in myriad orders (including Spiloconis), which are briefly reviewed. 1 Division of Invertebrate Zoology, American Museum of Natural History, New York. 2 Division of Invertebrate Zoology, American Museum of Natural History; and Division of Entomology (Paleoentomology), Natural History Museum, and Department of Ecology and Evolutionary Biology, 1501 Crestline Drive – Suite 140, University of Kansas, Lawrence, Kansas 66045. 3 Birbal Sahni Institute of Palaeobotany, 53 University Road, Lucknow 226 007, India. Copyright © American Museum of Natural History 2013 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3770 INTRODUCTION The Coniopterygidae is a family of approximately 560 described Recent species in 23 gen- era (Sziráki, 2011), comprised of diminutive lacewings that feed on small arthropods such as mites, aphids and scale insects. The family is definitively monophyletic, and the last major revisionary treatment was the monograph by Meinander (1972) on the world genera. Since then, there has been a world catalog of the species (Meinander, 1990; Sziráki, 2011), and vari- ous taxonomic treatments of particular faunas, the most notable being coverage of the Eur- asian, North African, Arabian, and New Guinean faunas (e.g., Sziráki, 2004; Sziráki and van Harten, 2006). Unfortunately, as of yet there is no comprehensive phylogenetic treatment of the genera. The one molecular study utilized just a region of one gene with 15 exemplar species in nine genera (Wang and Liu, 2007). The fossil record of Coniopterygidae, in contrast, is well explored since these tiny lacewings are preserved in most major amber deposits from the Cre- taceous and Tertiary periods. Excluding species in copal (which are presumably Recent), some 21 fossil species in 13 genera are preserved in amber from eight localities, as follows: Lebanon (Hammana, Early Cretaceous, Neocomian: Whalley, 1980; Azar et al., 2000; Nel et al., 2005); France (Char- ente-Maritime, late Albian, Cretaceous: Nel et al., 2005); Myanmar (Kachin, Albian-Ceno- manian boundary, Cretaceous: Engel, 2004); New Jersey (Raritan Formation, Turonian, Cretaceous: Grimaldi, 2000; Engel, 2002); Siberia (Taimyr Peninsula, Cretaceous: Santonian: Meinander, 1975); France (Oise, Les Quesnoys, Eocene: Nel et al., 2005); Baltic and the Ukraine (Eocene: Enderlein, 1910, 1930; Meinander, 1975; Dobosz and Krzemiński, 2000; Kupryjanowicz and Makarkin, 2008; Engel, 2010); and the Dominican Republic (Miocene: Meinander, 1998; Engel and Grimaldi, 2007). As yet, Coniopterygidae are unknown from amber of western Canada (Campanian: Late Cretaceous) or southern Mexico (Miocene). A summary of the fossil species in amber is provided in Engel and Grimaldi (2008), with additions by Kupryjanowicz and Makarkin (2008) and Engel (2010). There is a coniopter- ygid compression fossil from the Late Jurassic, Juraconiopteryx zherichini Meinander, but as described it is too poorly preserved for any meaningful interpretation (Meinander, 1975). It is interesting to note that most Cretaceous species, but not all, have three branches of the median vein in both wings, whereas all Tertiary and Recent species have two such branches in the forewing (Archiconiocompsa prisca Enderlein, 1910, in Baltic amber, has three M branches in the hind wing only). In lieu of a phylogenetic analysis, a three-branched vein M is probably plesiomorphic, and so many of these Cretaceous species are probably stem groups to Recent and Tertiary lineages. The present report has primarily biogeographic significance.Spiloconis Enderlein today has six species, distributed according to Meinander (1990) in the Indo-Pacific Region from Sri Lanka in the west, east to eastern Australia and the Fijian archipelago, and also including Thailand, southern China, and the Philippines. Meinander (1972) also mentioned “possibly Madagascar” for the distribution of Spiloconis, in reference to the inadequate description of Spiloconis nebulosa Fraser, 1957, the type of which may be lost. Fossils of the genus in Miocene amber from the Dominican Republic (Meinander, 1998; Engel and Grimaldi, 2007) thus rep- 2013 GRIMALDI ET AL.: AMBER CONIOPTERYGIDAE 3 resent a dramatic range extension. Here we assess assignment of most of the Dominican species to Spiloconis, based on new specimens and restudy of the original specimens. According to Meinander (1998), Spiloconis is polyphyletic due to the dramatic differences in male genitalia, although phenetic disparity per se is not an index of relationships. This study was prompted by the discovery of two coniopterygids in a piece of amber from the Eocene of India, which also appear to be in Spiloconis or a related genus, and which are described here. MATERIAL AND METHODS Dominican amber specimens, obtained from dealers, were originally excavated from mines just north of Santiago, Dominican Republic (reviewed in Grimaldi, 1995a). Age of the amber is Miocene, ca. 17–20 Ma (Iturralde-Vinent and MacPhee, 1996). Cambay amber from India was excavated directly by the authors from the Tadkeshwar lignite mines in Surat Dis- trict, Gujarat state, India. Here, the amber is found in thick lignitic outcrops of the Cambay Shale Formation, dated as earliest Eocene, ca. 52 Ma. A review of the Cambay amber deposits and stratigraphy is provided by Rust et al. (2010). The Cambay amber is a dammar-type fossil resin, which is poorly cross-linked; most pieces, in fact, have interiors that are still tacky. Thus, it requires embedding in a high-quality synthetic resin to stabilize it and facilitate working with it. Embedding and preparation techniques followed that of Nascimbene and Silverstein (2000), with some modifications such as use of EpoTek-301 resin for the embed- ding. Material is deposited in the following repositories: AMNH, Division of Invertebrate Zoology, American Museum of Natural History, New York, New York, and BSIP, Birbal Sahni Institute of Palaeobotany, Lucknow, India. Those specimens from the large Cambay amber study (Rust et al., 2010) that derive from the Tadkeshwar mine have a database/col- lection number beginning with “Tad”; there are two such specimens in this study. Morpho- logical terminology for the descriptive work follows that of Meinander (1972), and vein nomenclature is indicated in figure 5B. SYSTEMATIC PALEONTOLOGY Genus Spiloconis Enderlein Spiloconis Enderlein, 1907: 6. Type species: Spiloconis sexguttata Enderlein, 1907, by original designation. Diagnosis (modified from Meinander, 1972): Frons largely membranous or faintly sclero- tized, extending to clypeus. Antenna with 22–33 segments in Recent species, fossils with 17–24; scape large in Recent and some fossil species, length 3–4× width; pedicel large in Recent spe- cies, length 2.5–3× width. Forewing with 2 M setae; hind wing with basal fork of Rs in apical third of wing (vs. apical half), bases of M and Cu parallel and very close, separated by a distance approximately equal to diameter of vein. Male genitalia varied, stVIII (hypandrium) protruding apicad, scoop shaped in Recent species (see Meinander, 1972, for further description). Plica- turae on segments 2–7 in males, 2–6 in females. 4 AMERICAN MUSEUM NOVITATES NO. 3770 Spiloconis glaesaria Meinander Figures 1A, B; 2B; 4A Spiloconis glaesaria Meinander, 1998: 33. Engel and Grimaldi, 2007: 19. Diagnosis: Oral margin in frontal view broad (width 0.7× the distance between com- pound eyes), much of frontal portion of head membranous/lightly sclerotized, most of it cov- ered with fine setulae; dorsal margin of head barely protruding (cf. Spiloconis oediloma Engel and Grimaldi); wing with r-rs proximal to fork of R2+3–R4+5 (i.e., r-rs connected to Rs); L/W basal discal cell 3.2; antenna with 21–22 flagellomeres (23–24 antennomeres), size of basal flagellomere nearly equal to that of other flagellomeres. Type and Other Material: Known only from Dominican amber. Holotype, AMNH DR14-1094, in clear yellow amber 9 × 4 × 5 mm in size, embedded in EpoTek 301 resin, 12 × 3 × 5 mm. The amber contains many bubbles, and the holotype is best observed ventrally. Meinander (1988) stated that the body of the specimen was covered with “wax” (a fine layer of which indeed coats most modern dustywings), but the fossil is actually covered with a milky froth, a preservational artifact common among inclusions in
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