372 S. L. Winterton Et Al. Are Obligate Predators of Freshwater Sponges and Bryozoans, Whereas Nevrorthidae Are Generalist Benth

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372 S. L. Winterton et al. are obligate predators of freshwater sponges and bryozoans, Berothidae and Mantispidae whereas Nevrorthidae are generalist benthic predators in lotic habitats. Sometimes incorrectly referred to as semiaquatic, Mantispidae (mantid lacewings) are distinctive lacewings some osmylid larvae (e.g. Osmylinae, Kempyninae) are found with raptorial forelegs resembling preying mantids (Mantodea). in moist stream-bank habitats, whereas other species (e.g. The phylogenetic placement of Rhachiberothinae (thorny Stenosmylinae, Porisminae) live under bark in drier habitats. lacewings) is contentious, having been proposed as a subfam- Our data (Figs 5, 7; Figure S1) support a clade comprising ily of Berothidae (Tjeder, 1959; MacLeod & Adams, 1968), Nevrorthidae, Sisyridae and Osmylidae sister to the rest of a subfamily of Mantispidae (Willmann, 1990) and as a sep- Neuroptera after Coniopterygidae. Unfortunately, this clade has arate family entirely (Aspock¨ & Mansell, 1994; Grimaldi & weak statistical support, and in the pruned analysis Sisyridae Engel, 2005). Our analyses recovered a monophyletic clade are recovered as sister to Dilaridae. A close relationship composed of Mantispidae + Berothidae with relatively strong between these two families was supported by Sziraki´ (1996) support (PP = 1.00, PB = 91%, DI = 9) (Fig. 5). Unfortu- based on female internal genitalia. Using molecular data, nately, internal relationships between and within these families Haring & Aspock¨ (2004) also placed Nevrorthidae, Sisyridae were not recovered with strong support and varied among and Osmylidae in sequence as sister taxa to the rest of analyses (Figs 4–7; Figs 4, 5). The enigmatic Ormiscocerus Neuroptera. The placement of Nevrorthidae as sister to the was transferred recently from Hemerobiidae to Berothidae: rest of Neuroptera by these authors supported a previous Cyrenoberothinae (Penny & Winterton, 2007). The oldest fos- proposal by Aspock¨ et al. (2001) (see also Beutel et al., 2009), sils known for Berothidae and Mantispidae are all from Late based on morphological evidence, for an aquatic origin of the Jurassic- (Ren & Guo, 1996) and Cretaceous-aged amber order based on the presence of plesiomorphic characters in deposits (Panﬁlov, 1980; Whalley, 1980; Klimaszewski & the larva such as a well-formed gula plate and absence of Kevan, 1986). Estimates of divergences times presented here cryptonephric Malpighian tubules. The presence of a gula in indicate that the group arose during the Early Triassic (Fig. 7). the larval head capsule is found throughout Neuropteroidea, but is lost in Neuroptera except for Nevrorthidae (Aspock¨ Curved versus straight jaws in larval Neuroptera et al., 2001; Beutel et al., 2009), Ithonidae, Polystoechotidae and Myrmeleontiformia (MacLeod, 1964), in all of which it is All of the families discussed above have larvae with probably retained as a symplesiomorphy. straight jaws (with the exception of Nevrorthidae). The remaining Neuroptera (Chrysopidae, Hemerobiidae, Ithonidae, Polystoechotidae and members of Myrmeleontiformia) have Dilaridae larval jaws that are curved, with bases usually widely spaced (characters 28 and 36). The jaws of larval Ithonidae and Pleasing lacewings (Dilaridae) are an enigmatic group Polystoechotidae are much shorter than those of the other of lacewings found in all biogeographic regions except families, but in at least some members the jaws are notably Australasia. Males have distinctive pectinate antennae and curved in at least one or more instars (MacLeod, 1964; females have a long ovipositor. In our combined data anal- Grebennikov, 2004). Larvae in this clade typically have yses, Dilaridae were recovered as sister to the paraphyletic re-enforcing modiﬁcations of the posterior rim of the head Hemerobiiformia + Myrmeleontiformia. In the pruned taxa set capsule. In Chrysopidae, Ithonidae and Polystoechotidae (but analysis it was recovered as sister to Sisyridae, although not Hemerobiidae) this involves a bracing of the posteroventral with relatively weak statistical support. Authors have sug- rim by a close approximation of the posterior tentorial arms. gested a close relationship between Dilaridae, Mantispidae and In Ithonidae and Polystoechotidae this also involves some Berothidae (MacLeod, 1964; Aspock¨ et al., 2001; Aspock¨ & sclerotization of the gula region. Larvae of Myrmeleontiformia Aspock,¨ 2008; Beutel et al., 2009), although support of this re-enforce the posterior rim through sclerotization of the group of families is equivocal when all the evidence is consid- entire ventral surface of the head capsule and restriction of ered. The molecular study of Haring & Aspock¨ (2004) placed the maxillary and gula sclerites to a small region anteriorly Dilaridae as sister to Coniopterygidae. Morphological support (MacLeod, 1964). This latter strategy allows larvae to take for Dilaridae as an early branching lineage along with Osmyl- larger or more active prey items and may be correlated with idae, Nevrorthidae and Sisyridae has been provided by Sziraki´ larger overall body size; some of the largest lacewings are (1996) based on the structural arrangement of the female inter- found in families such as Ascalaphidae and Myrmeleontidae nal genitalia, whereas Handlirsch (1906–1908) grouped Sisyri- (MacLeod, 1964). In larvae with straight jaws and a head dae with Dilaridae based largely on fossil data. These families capsule without ventral sclerotization, the jaws move largely in share with Megaloptera and Raphidioptera characters such as the longitudinal plane and thus larvae are unable to manipulate paired spermathecae and a bursa copulatrix forming a com- their prey items except through movements of the entire head plex with the ductus seminalis. It appears, based on molecular capsule. Larvae with straight jaws feed on small or relatively data (presented herein and by Haring & Aspock¨ (2004)) and immobile prey items (Withycombe, 1925) and are more likely various morphological characteristics (e.g. Sziraki,´ 1996), that to be specialized feeders on a single prey type (e.g. Sisyridae, this anomalous family is not closely related to Mantispidae and Mantispidae, Berothidae). By contrast, with the exception of Berothidae as was proposed previously. highly specialized Ithonidae and Polystoechotidae, lacewing