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The Function and Phylogenetic Implications of the Tentorium in Adult Neuroptera (Insecta)

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The Function and Phylogenetic Implications of the Tentorium in Adult Neuroptera (Insecta) Arthropod Structure & Development 40 (2011) 571e582 Contents lists available at ScienceDirect Arthropod Structure & Development journal homepage: www.elsevier.com/locate/asd The function and phylogenetic implications of the tentorium in adult Neuroptera (Insecta) Dominique Zimmermann a,*, Susanne Randolf a, Brian D. Metscher b, Ulrike Aspöck a a Natural History Museum, 2nd Zoological Department, Burgring 7, 1010 Vienna, Austria b Department of Theoretical Biology, University of Vienna, Althanstrasse 14, 1090 Wien, Austria article info abstract Article history: Despite several recent analyses on the phylogeny of Neuroptera some questions still remain to be Received 11 April 2011 answered. In the present analysis we address these questions by exploring a hitherto unexplored Accepted 12 June 2011 character complex: the tentorium, the internal cuticular support structure of the insect head. We described in detail the tentoria of representatives of all extant neuropteran families and the muscles Keywords: originating on the tentorium using 3D microCT images and analyzed differences in combination with Neuroptera a large published matrix based on larval characters. We find that the tentorium and associated Tentorium musculature are a source of phylogenetically informative characters. The addition of the tentorial Musculature Phylogeny characters to the larval matrix causes a basad shift of the Sisyridae and clearly supports a clade of all Function Neuroptera except Sisyridae and Nevrorthidae. A sister group relationship of Coniopterygidae and the Laminatentorium dilarid clade is further corroborated. A general trend toward a reduction of the dorsal tentorial arms and the development of laminatentoria is observed. In addition to the phylogenetic analysis, a correlation among the feeding habits, the development of the maxillary muscles, and the laminatentoria is demonstrated. Ó 2011 Elsevier Ltd. All rights reserved. 1. Introduction In light of previous work we concentrate on the following three points: Neuroptera are a small but biologically and morphologically highly heterogeneous insect order. Together with Megaloptera and 1. The monophyly of Hemerobiiformia. The Hemerobiiformia, Raphidioptera, they constitute the superorder Neuropterida, which comprising 12 families, were recognized as “hemerobioid larval is one of the oldest holometabolan lineages (Aspöck et al., 2001; type” by MacLeod (1964) on the basis of features of the larval Grimaldi and Engel, 2005). In recent years many analyses have head anatomy. In a holomorphological cladistic analysis per- been conducted to illuminate the phylogeny of the Neuroptera, formed by Aspöck et al. (2001) these features were interpreted including molecular (Haring and Aspöck, 2004) morphological, as synapomorphies of this suborder. In a molecular analysis by (Aspöck et al., 2001; Aspöck and Aspöck, 2008; Beutel et al., 2010b, Haring and Aspöck (2004) the clade Hemerobiiformia was 2010a), and combined analyses (Winterton et al., 2010; review and fractured as the Sisyridae and the Osmylidae emerged as two documentation: Aspöck and Aspöck, 2010). In short, two contro- separate branches, a view also held in an analysis based on versial approaches have been conveyed: the triple suborder concept genital sclerites by Aspöck and Aspöck (2008). Keeping things with Neuroptera comprising three monophyletic suborders, Nev- complicated, Beutel et al. (2010b) re-established the mono- rorthiformia, Myrmeleontiformia, and Hemerobiiformia; and Hem- phylum Hemerobiiformia. erobiiformia as paraphyletic with respect to Myrmeleontiformia. 2. The position of the Sisyridae. Traditionally, the Sisyridae are Within Hemerobiiformia the Sisyridae and the Coniopterygidae part of the Hemerobiiformia. The position as sister group of the compete for the most conflicting status. Coniopterygidae in Aspöck et al. (2001) is strongly affected by the incorrect interpretation of the enlarged labial palps which are, in fact, an independent acquisition (Zimmermann et al., 2009). In the molecular analysis of Haring and Aspöck (2004) * þ þ Corresponding author. Tel.: 1 43 1 521 77 316; fax: 1 43 1 521 77 302. the Sisyridae emerge as sister group of all others except E-mail addresses: [email protected] (D. Zimmermann), [email protected] (S. Randolf), [email protected] Nevrorthidae, and the same relationships are hypothesized by (B.D. Metscher), [email protected] (U. Aspöck). Aspöck and Aspöck (2008). The molecular-morphological 1467-8039/$ e see front matter Ó 2011 Elsevier Ltd. All rights reserved. doi:10.1016/j.asd.2011.06.003 572 D. Zimmermann et al. / Arthropod Structure & Development 40 (2011) 571e582 combined analysis of Winterton et al. (2010) results in combination with a previously published matrix based on larval Coniopterygidae þ [(Osmylidae þ (Sisyridae þ Nevrorthidae)) þ characters (Beutel et al., 2010b). In addition, we discuss evolu- (all other families)]. tionary changes of the tentorium in the light of their adaptive value 3. The position of the Coniopterygidae. Due to reductions corre- and their correlation with feeding habits. lated with their small body size the Coniopterygidae are prob- ably the most difficult neuropteran family to classify. They have 2. Material and methods taken various positions: as sister group of all other Neuroptera (Withycombe, 1925; Winterton et al., 2010), as sister group of 2.1. Specimens examined the Sisyridae (Aspöck et al., 2001), as sister group of the Dilar- idae (possibly a result of a long-branch attraction; Haring and At least one species from each of the 17 neuropteran families Aspöck, 2004), and finally as sister group of the whole dilarid was examined. A detailed list of the material is given in Table 1.As clade (Aspöck and Aspöck, 2008; Beutel et al., 2010b). far as possible the same species as in the larval analysis by Beutel et al. (2010b) were used. Otherwise the availability of fresh mate- In the present study we address these questions by exploring rial determined our species choice. a hitherto unexplored character complex in Neuroptera: the ten- For morphological investigations the specimens were killed in torium, the inner skeleton of the insect head (Fig. 1). The tentorium 90% alcohol, fixed in alcoholic Bouin’s fluid for 3 h, and stained in 1% serves as a stabilizing element and muscle attachment structure for iodine-ethanol solution overnight. The specimens were microCT muscles of the antennae, pharynx, maxillae, labial and hypopha- scanned in 95% alcohol. The specimens are vouchered in the ryngeal muscles which are also considered in the course of this Natural History Museum Vienna. study. The tentorium is formed by the intersegmental invagination of the head capsule at two anterior and two posterior points which 2.2. Imaging and 3D-Reconstruction are externally visible as anterior and posterior tentorial pits. The anterior and posterior tentorial arms converge to the tentorial The specimens were imaged with an Xradia MicroXCT x-ray bridge. Dorsal tentorial arms are evaginations of the anterior ten- microtomography system (University of Vienna, Department of torial arms (Snodgrass, 1928, 1935). They are connected to the head Theoretical Biology) with a tungsten or rhodium source at 40- capsule by fibrillae in the antennal area (Staniczek, 2001). 80kVp and 4-8W and images were reconstructed using the soft- The tentorium has been studied in adults and larvae of various ware provided with the microCT system. The microCT data were insect taxa and has provided informative characters for recon- reconstructed with 2 Â 2 pixel binning to reduce noise and file size, structing phylogenies for certain taxa (e.g. Klass and Eulitz, 2007; and reconstructed volume images were exported as TIFF image Koch, 2000; Symmons, 1950). In Neuroptera the tentoria and its stacks. The software Amira 5.1 was used for 3D-visualization and muscles have thus far only been described for a few species in the analysis of the data. MicroCT image stacks and histological sections scope of morphological investigations of the adult head (Ferris, are deposited in the Natural History Museum Vienna. 1940; Korn, 1943; Morse, 1931; Beutel et al., 2010a), but were The classification of musculature follows von Kéler (1963). never analyzed further. The aim of our study is to elucidate the tentorial complex in an 2.3. Cladistic analysis evolutionary and phylogenetic context and to help clarify the classification of the Neuroptera and the Neuropterida. To this end For the phylogenetic analysis we only scored differences that we have used 3D microtomographic (microCT) images to obtain could be defined as qualitative characters, in order to minimize detailed morphological data for adult tentoria and associated subjectivity. As these characters alone are insufficient to infer musculature in all 17 Neuropteran families. In order to test the a phylogeny, we added them to a large published matrix based phylogenetic value of the tentorial characters we analyzed them in on larval characters (Beutel et al., 2010b). We chose the study of Fig. 1. Scheme of neuropteran head with tentorium. Abbreviations: af - antennal foramen, ata - anterior tentorial arm, atp - anterior tentorial pit, dta - dorsal tentorial arm, lt - laminatentorium, pta - posterior tentorial arm, tb - tentorial bridge. D. Zimmermann et al. / Arthropod Structure & Development 40 (2011) 571e582 573 Table 1 List of taxa examined. Family Species Sex Method Ascalaphidae Libelloides macaronius (Scopoli, 1763) ? microCT
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