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Lacewings (Neuroptera) Lacewings (Neuroptera) Shaun L. Wintertona,* and Brian M. Wiegmannb been identiA ed, but they usually diB er greatly between aEntomology, Queensland Department of Primary Industries & subsequent authors with regard to membership of fam- Fisheries, 80 Meiers Road, Indooroopilly, Queensland, Australia ilies in each (1). Recent quantitative analyses of mor- b 4068; Department of Entomology, North Carolina State University, phological characters and phylogeny by Aspöck et al. (4) Raleigh, NC 27695, USA led to consolidation of this classiA cation into only three *To whom correspondence should be addressed (wintertonshaun@ gmail.com) Suborders Nevrorthiformia (containing Nevrorthidae only), Myrmeleontiformia (containing A ve families: Nemopteridae, Psychopsidae, Nymphidae, Ascalaphidae, Abstract and Myrmeleontidae), and Hemerobiiformia (contain- ing 11 families: Ithonidae, Polystoechotidae, Chrysopi- Lacewings (~5700 species) are divided into 17 families dae, Hemerobiidae, Mantispidae, Rhachiberothidae, distributed on all continents. This group of insects is well Berothidae, Coniopteryigdae, Dilaridae, Sisyridae, and defi ned by complex larval characteristics such as modifi ed Osmylidae). 7 e most signiA cant aspects of this work sucking jaws, incomplete gut, and modifi ed Malphigian were (a) placement of Ithonidae (moth lacewings) not as tubules used for spinning a silken cocoon in which the the closest relative of remaining neuropterans but rather immature pupates. Recent molecular evidence supports a in a more derived position closer to Myrmeleontiformia Permian (299–251 million years ago, Ma) origin of the order and (b) the placement of Nevrorthidae, instead, as the with Coniopterygidae as the closest relative of all other earliest-branching family. neuropterans. The only well-defi ned group of families is A monophyletic Hemerobiiformia as circumscribed Myrmeleontiformia. This group of fi ve families diverged by Aspöck et al. (4) has never been recovered as a natural from the rest of the order ~184 Ma and has become the g r oup i n a ny m o l e c ul a r a n a l y s i s 5,( 6). Myrmeleontiformia most species-rich lineage of Neuroptera. 7 e lacewings comprise a moderately sized group of insects whose adults are characterized by delicate wings, oJ en large in size, with highly reticulate or lacelike venation (Fig. 1). 7 e order is presently divided into 17, mostly well deA ned, families with ~5700 species worldwide. Lacewings, together with Megaloptera and Raphidioptera, constitute the Superorder Neuropterida, which is assumed to be the closest relative of Coleoptera. 7 ree unique larval characters distinguish Neuroptera from other insects: modiA ed jaws for sucking, incom- plete gut, and modiA ed Malphigian tubules for spinning a silken cocoon in which the immature pupates 1(). Here, we review the relationships and divergence times of the families of lacewings (Fig. 2). Elucidating the evolutionary history of lacewings has proven to be highly problematic. Neuroptera clas- siA cation has traditionally been founded largely on the intuitive phylogenies by Handlirsch (2) for fossils and by Withycombe (3) for larval anatomy and morphology. Fig. 1 A split-footed lacewing (Norfolius howensis) from From these works a number of higher-level groups have Australia. Credit: S. L. Winterton. S. L. Winterton and B. M. Wiegmann. Lacewings (Neuroptera). Pp. 290–292 in e Timetree of Life, S. B. Hedges and S. Kumar, Eds. (Oxford University Press, 2009). HHedges.indbedges.indb 229090 11/28/2009/28/2009 11:27:38:27:38 PPMM Metazoa; Arthropoda; Insecta; Neuroptera 291 Hemerobiidae 11 Chrysopidae 8 Ithonidae 10 Polystoechotidae 6 Nemopteridae 13 Myrmeleontidae 14 12 Ascalaphidae 9 Nymphidae 5 Psychopsidae Myrmeleontiformia Mantispidae 3 Berothidae 7 Rhachiberothidae Osmylidae 4 2 Nevrorthidae Sisyridae 1 Dilaridae Coniopterygidae Triassic Jurassic Cretaceous Pg Ng MESOZOIC CENOZOIC 250 200150 100 50 0 Million years ago Fig. 2 A timetree of lacewings (Neuroptera). Divergences are shown in Table 1. Abbreviations: Ng (Neogene) and Pg (Paleogene). is a strongly supported group in all published phyloge- 7 e most recent analysis of lacewing phylogeny is nies based on morphological (3, 4, 7–9) and molecular by Winterton and Wiegmann (submitted), using both data (Winterton and Wiegmann, submitted; 5, 6). A molecular and morphological data with much greater suite of adult and larval morphological characters deA ne taxon sampling than in previous studies of the group. this group. 7 ey produced the most comprehensive phylogeny of the Relative to morphological data, very few molecular order to date, by using DNA sequences for16S, 18S riboso- phylogenies of Neuroptera have been published. Haring mal genes, COI and CAD, combined with morphological and Aspöck (5) published the A rst attempt at elucidat- data for representatives of all families of Neuroptera. ing lacewing phylogeny using molecular data by sequen- Results from this study show that Coniopteryigdae cing representatives of most neuropteran families for rather than Nevrorthidae is the earliest-branching fam- amino acid sequences of cytochrome c oxidase subunit ily in the order, and that Hemerobiiformia is a para- III. 7 e results of this study recovered monophyletic phyletic group having a laddered topology leading to a Myrmeleontiformia and Nevrorthiformia, but a poly- monophyletic Myrmeleontiformia. Dilaridae is again phyletic Hemerobiiformia, with considerable discord supported as a basal group closely related to Sisyridae between molecular and morphological data regarding and Coniopteryigdae. the placement of the families Osmylidae and Dilaridae. Neuropterans have a rich fossil record with deA nitive Using 18S rDNA sequences Winterton (6) also showed neuropteridan ancestors dating back to the Permian (1, strong support for Myrmeleontiformia, but did not 10). Only Winterton and Wiegmann (submitted), using recover a monophyletic Hemerobiiformia. In this analysis the aforementioned gene sequences, have estimated Dilaridae were also placed as a basal group rather than divergence times among lacewing families based on gen- being closely related to Mantispidae, Rhachiberothidae, etic data. Coniopterygidae diverged from the rest of the and Berothidae. order during the late Permian (~257 Ma) followed by HHedges.indbedges.indb 229191 11/28/2009/28/2009 11:27:42:27:42 PPMM 292 THE TIMETREE OF LIFE Table 1. Divergence times (Ma) and confi dence/ Neuroptera is a relatively ancient group of insects with credibility intervals (CI) among lacewings (Neuroptera) a rich fossil record. 7 is, along with tremendous mor- based on Winterton and Wiegmann (submitted). phological disparity between families and highly disjunct Timetree biogeographical patterns across related taxa, indicate Node Time CI that the golden age of lacewings has long since past (4). Divergence estimates support this view with all major 1 257 268–244 divergences occurring during the Mesozoic (251–66 Ma) 2247259–235(Winterton and Wiegmann, submitted). Our under- 3242254–229standing of the evolutionary history of this group is still 4228242–212developing, with morphology providing only fragmen- 5227239–214tary insights. Further studies using molecular data (e.g., 6219231–207more molecular markers) and intermediate fossil mater- 7214230–199ial are needed to fully elucidate the evolutionary history of Neuroptera. 8213226–200 9208221–195 10 197 211–184 Acknowledgment 11 197 214–177 Support was provided by the U.S. National Science 12 187 200–176 Foundation to the authors. 13 172 183–167 14 153 162–150 References Note: Two protein-encoding genes and two ribosomal genes 1. T. R. New, Handbook of Zoology. Planipennia: Lacewings, were used to estimate divergence times in Winterton and Vol. IV. Part 30 (Walter de Gruyter, Berlin, 1989). Wiegmann (submitted). 2. A. Handlirsch, Die fossilen Insekten Und Die Phylog- enie Der Rezenten Formen: Ein Handbuch Für Paläon- all major family-level divergences occurring during the tologen Und Zoologen (Engelmann, Leipzig, Germany, Jurassic (>145 Ma). 7 is is reP ected in the fossil record, 1906–1908). with fossil taxa of most extant families present in Jurassic 3. C. L. Withycombe, Trans. Entom. Soc. Lond. 72, 303 or early Cretaceous aged deposits (1, 2, 10). 7 e estimated (1925). 4. U. Aspöck, J. D. Plant, H. L. Nemeschkal, Syst. Entom. age of the divergence of Myrmeleontiformia from its 26, 73 (2001). closest relative is ~216 Ma. 7 e most recent family-level 5. E. Haring, U. Aspöck, Syst. Entom. 29, 415 (2004). divergence is that of Ascalaphidae from Myrmeleontidae 6. S. L. Winterton, Entomol. Abhand. 61, 158 (2003). (~153 Ma). 7. E. G. MacLeod, A Comparative Morphological Study Based on these data, interesting questions may be of the Head Capsule and Cervix of Larval Neuroptera posed regarding the origins of certain families. One (Insecta), Ph.D. 7 esis (Harvard University, Cambridge, example is that of the origins of freshwater sponges and Massachusetts, 1964). spongilla P ies (Sisyridae). Freshwater sponges are clas- 8. C. S. Henry, Psyche 85, 265 (1978). siA ed in the Suborder Spongillina and likely colonized 9. M. W. Mansell, in Current Research in Neuropterology. freshwaters from marine ancestors during the early Proc. Fourth Int. Symp. on Neuropterology, M. Canard, Jurassic (210–141 Ma) (11). Considering that the origin H. Aspöck, M. W. Mansell, Eds. (Privately printed, Toulouse, France, 1992), pp. 233–241. of Sisyridae is estimated at ~247 Ma (Winterton and 10. D. Grimaldi, M. S. Engel, Evolution of e Insects Wiegmann, submitted), and that this group are specialist (Cambridge University Press, Cambridge, UK, 2005). predators on freshwater sponges and bryozoans, ances- 11. R. Manconi, R. Pronzato, in Systema Porifera: A guide tral sisyrids possibly fed on freshwater bryozoans dur- to the classiA cation of sponges, J. N. A. Hooper, R. W. M. ing the Triassic before expanding to freshwater sponges Van Soest, Eds. (Kluwer Academic/Plenum Publishers, during the Jurassic. New York, 2002), pp. 921–1021. HHedges.indbedges.indb 229292 11/28/2009/28/2009 11:27:42:27:42 PPMM.
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