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Susanne Randolf, Dominique Zimmermann & Ulrike Aspöck Natural History Museum Vienna, A-1010 Vienna, e-mail: [email protected] In recent years many analyses have been conducted to illuminate the phylogeny of the Neuroptera, including molecular (Haring and Aspöck 2004), morphological (Aspöck et al. 2001; Aspöck and Aspöck 2008; Beutel et al. 2010a, 2010b; Zimmermann et al. 2011; Randolf et al. in press) and combined analyses (Winterton et al. 2010). Major conflicts are found in the first bifurcation events which have considerable implication on the interpretation of character evolution. We combined 41 head anatomical characters of adult Neuroptera with published larval (Beutel et al. 2010a) and genital (Aspöck et al. 2001; Aspöck and Aspöck 2008) characters and combined them in different ways to figure out their influence on the tree topology. aquatic lifestyle A Nevrorthidae and Sisyridae are the only neuropteran families with aquatic larvae poison gland and poison channel In Neuroptera, the larvae of all families except Sisyridae have poison glands and poison channels associated with their sucking tubes Sialis cryptonephric Malpighian tubules 1 21 27 33 50 60 92 109 Sisyra Several cryptonephric Malpighian tubules are found in terrestrial larvae of Neuroptera. In the aquatic larvae of Sisyridae only one Malpighian tubule is 4 5 6 12 15 78104107 cryptonephric, none in the aquatic larvae of Nevrorthidae (Gaumont 1976) Nevrorthus 0 17 25 26 37 38 39 41 51 53 70 83 84 85 95 4 108 1 3 11 19 26 28 68 Polystoechotes 15 78 85109 Ithone 36 52 88 89 27 80 82 84 98104106108113 Osmylus 21 35 15 50 68 92101 34 57 67 110 Hemerobius Hemerobiiformia partim 70 74 Chrysopa 60 61 65 66 75 76 86 87 113 5 57 73 108 Psychopsis 4 6 7 9 10 12 32 44 84 26 72 78 85 89 larval characters: brown Nemoptera 15 16 head anatomical characters of the adults: black 106 Nymphes characters of genital sclerites: olive green 29 55 64 72109111 Myrmeleontiformia 16 56 68 69 79 80 105 Libelloides 15 70 106 Sisyridae as sister group to the remaining Neuroptera is retrieved in: Euroleon combinations of larval + head anatomical characters + characters of the 14 18 24 40 45 46 47 49 Helicoconis genital sclerites (6 of 6 trees), larval + head anatomical characters (2 of 16 26 51 88 89 2 trees), Zimmermann et al. 2010: larval + head anatomical characters, Coniopteryx 21 27 64 72 78 Randolf et al. in press: larval + head anatomical characters. 73 Mantispa 8 15 54 57 15 66 75 113 Nallachius 13 27 43 111 64 Hemerobiiformia partim B Nevrorthus Hemerobiiformia 6 Lomamyia Sisyra Mucroberotha Myrmeleontiformia The combined analyses of larval and adult characters provoke conflicts. Cryptonephric Malpighian tubules are homologous in all Neuroptera in B and C The classical three-suborder-concept Nevrorthiformia, Myrmeleontiformia with the difference that in B cryptonephry evolved in a terrestrial environment and Hemerobiiformia (Aspöck et. al. 2001) is retrieved in: combination of and in C in an aquatic one. The function of cryptonephric Malpighian tubules larval characters + characters of the genital sclerites (4 of 29 trees), Beutel to reabsorb water might be necessary for the aquatic larvae of Sisyridae when et al. 2010 a: larval characters, Beutel et al. 2010b: larval characters + head anatomical characters. they migrate to a dry habitat for pupation. The aquatic larvae of Nevrorthidae, lacking cryptonephric Malpighian tubules, pupate in water. In A cryptonephry evolved independently in Sisyridae and Neuroptera with terrestrial larvae. C Nevrorthus In B and C a poison gland and a poison channel belong to the ground pattern Sisyra of Neuroptera and are lost in Sisyridae while in A these characters constitute a synapomorphy of Neuroptera excl. Sisyridae. Primarily aquatic larvae in Osmylus Neuroptera as proposed by Aspöck (1995) are supported in all combinations; in B Nevrorthidae are primarily aquatic and Sisyridae secondarily. paraphyletic Hemerobiiformia It has been demonstrated that the combination of independent data as e. g. Myrmeleontiformia adult and larval characters increase both, character conflict as well as tree support and resolution (Meier and Lim 2009). Of course, also nodes supported Nevrorthidae as sister group to the remaining Neuroptera and Sisyridae as sister group to all others except Nevrorthidae is retrieved in: combination of by synapomorphies of independent data can be regarded as most reliable. larval characters + characters of the genital sclerites (1 of 29 trees), Haring In our analyses a clade of Neuroptera exclusive Sisyridae and Nevrorthidae and Aspöck 2004: molecular analysis, Aspöck and Aspöck 2008: larval is corroborated by larval, adult head and genital characters as well as by characters + characters of the genital sclerites. molecular data and thus appears as the preferable result. References: Aspöck, U. 1995. Neue Hypothesen zum System der Neuropterida. Mitt. Dtsch. Ges. allg. angew. Ent. 10: 633-636. Aspöck, U., Aspöck, H. 2008. Phylogenetic relevance of the genital sclerites of Neuropterida (Insecta: Holometabola). Syst. Entomol. 33, 97-127. Aspöck, U., Plant, J. D., Nemeschkal, H. L. 2001. Cladistic analysis of Neuroptera and their systematic position within Neuropterida (Insecta: Holometabola: Neuropterida: Neuroptera). Syst. Entomol. 26, 73-86. Beutel, R. G., Friedrich, F., Aspöck, U. 2010a. The larval head of Nevrorthidae and the phylogeny of Neuroptera (Insecta). Zool. J. Linn. Soc. 158, 533-562. Beutel, R. G., Zimmermann, D., Krauß, M., Randolf, S., Wipfler, B. 2010b. Head morphology of Osmylus fulvicephalus (Osmylidae, Neuroptera) and its phylogenetic implications. Org. Divers. Evol. 10, 311- 329. Gaumont, J. 1976. L'appareil digestif des larves de Planipennes. Ann. Sci. Nat., Zool. Biol. Anim. 18:145-250. Haring, E., Aspöck, U. 2004. Phylogeny of the Neuropterida: a first molecular approach. Syst. Entomol. 29, 415-430. Meier, R., Lim, G. S. 2009. Conflict, convergent evo- lution, and the relative importance of immature and adult characters in endopterygote phylogenetics. Annu. Rev. Entomol. 54, 85–104. Randolf , S., Zimmermann, D., Aspöck, U. in press. Head anatomy of adult Sisyra terminalis (Insecta: Neuroptera: Sisyridae) – functional adaptations and phylogenetic implications. Arthropod Struct. Dev. Winterton, S. L., Hardy, N. B., Wiegmann, B. M. 2010. On wings of lace: phylogeny and Bayesian divergence time estimates of Neuropterida (Insecta) based on morphological and molecular data. Syst. Entomol. 35, 349-378. Zim- mermann, D., Randolf, S., Metscher B. D., Aspöck, U. 2011. The function and phylogenetic implications of the tentorium in adult Neuroptera (Insecta). Arthropod Struct. Devel. 40, 571-582..
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