SYSTEMATICS Phylogeny of Moth Lacewings and Giant Lacewings (: Ithonidae, Polystoechotidae) Using DNA Sequence Data, Morphology, and Fossils

1 2 SHAUN L. WINTERTON AND VLADIMIR N. MAKARKIN

Ann. Entomol. Soc. Am. 103(4): 511Ð522 (2010); DOI: 10.1603/AN10026 ABSTRACT A phylogeny of lacewing families Ithonidae and Polystoechotidae is presented based on three gene markers (16S and 18S ribosomal DNA and CAD) and 23 morphological characters. Living and fossil genera presently placed in Polystoechotidae (Fontecilla Nava´s, Platystoechotes Carpenter and Burmeister) and Ithonidae (Adamsiana Penny, Allorapisma Makarkin & Archibald, Ithone Newman, Megalithone Riek, Oliarces Banks, Principiala Makarkin & Menon, Rapisma Walker and Walker) were included in phylogenetic analyses (parsimony and Bayes- ian) and compared with outgroups selected from various families of Neuroptera. The resulting phylogeny recovered a monophyletic clade comprising Ithonidae and Polystoechotidae as hypoth- esized previously. Rapismatidae as a separate family is not supported and Ithonidae are rendered paraphyletic with three extant genera previously placed in Ithonidae (Adamsiana, Oliarces, and Rapisma), recovered deep within Polystoechotidae. The fossil genera Allorapisma and Principiala formed a sister-group relationship with Rapisma, also within Polystoechotidae. Due to the lack of mutually exclusive synapomorphies for either Ithonidae or Polystoechotidae, a single more inclusive family Ithonidae is proposed, including all ithonid genera and all genera previously placed in Polys- toechotidae. Synapomorphies characterizing the revised concept of Ithonidae s.l. are discussed.

KEY WORDS phylogeny, Rapismatidae, Ithonidae, Polystoechotidae, lacewing

Ithonidae (moth lacewings) and Polystoechotidae known from Southeast Asia (Rapisma McLachlan) (giant lacewings) are small families of robust and often containing Ϸ20 species from montane areas in Nepal hairy lacewings. Ithonidae presently comprises seven to tropical lowlands in Thailand and Malaysia (Bar- extant genera from Australia, Southeast Asia, and the nard 1981, Barnard and New 1985, New 1985, Yang New World, whereas Polystoechotidae comprise 1993). Rapisma was previously considered as the sole three extant New World genera. Three genera of in the family Rapismatidae, although there ithonids are found in Australia; Ithone Newman (Fig. seems little evidence for maintaining this separation 1A) contains 10 species, whereas Megalithone Riek and (Penny 1996, Makarkin and Menon 2007). Varnia Walker contain two species each (Riek 1974). Polystoechotidae are represented by Platystoechotes Three genera also are described from the New World, Carpenter, Polystoechotes Burmeister (Fig. 1C), and and all are monotypic, although additional unde- Fontecilla Nava´s consisting of four species in total. scribed species are known in collections (Oswald Fontecilla graphicus Nava´s and Polystoechotes gazullai 1998b). Oliarces clara Banks (Fig. 1B) is known from Nava´s are known only from Chile, whereas Platysto- arid areas of southwestern . Adamsiana echotes lineatus Carpenter is known from the south- curoi Penny is a montane species from Central Amer- western United States () (Oswald 1998b). A ica and is unique in that females are apterous, whereas second species of Polystoechotes (P. punctatus F.) is males are macropterous (Penny 1996). Narodona recorded from Canada south to Panama (Oswald mexicana Nava´s is only known from the original de- 1998b, Archibald and Makarkin 2006; Fig. 1). scription and a hind wing Þgure of a single specimen Specimens of both Ithonidae and Polystoechotidae from Mexico (Nava´s 1929a). A single ithonid genus is are relatively rare in collections, although localized mass emergences have been recorded in Oliarces Banks and Ithone after seasonal rains (Riek 1974, Any opinions, Þndings, and conclusions or recommendations ex- pressed in this publication are those of the authors and do not nec- Faulkner 1990). Larvae of Ithonidae and Polysto- essarily reßect the views of the National Science Foundation. echotidae are known for Oliarces Banks, Megalithone, 1 Corresponding author: School of Biological Sciences, University Ithone, Polystoechotes, and Platystoechotes (MacLeod of Queensland, St. Lucia, Queensland, Australia 4072 (e-mail: 1964, Grebennikov 2004, Winterton et al. 2010). All are [email protected]). 2 Institute of Biology and Soil Sciences, Far East Branch of the fossorial and in Ithonidae the mature larvae are scar- Russian Academy of Sciences, Vladivostok, 960022, Russia. abaeiform. Larvae of Ithonidae were originally re-

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Fig. 1. Adult Ithonidae and Polystoechotidae. (A) Ithone fulva Tillyard (Australia: Queensland). (B) Oliarces clara Banks. (USA: California). (C) Polystoechotes punctatus (F.) (USA: Washington). Photograph credits: S. L. Winterton (A and B); Karl Volkman (C). (Online Þgure in color.) ported as predatory by Tillyard (1922), but Gallard combination of the two have conÞrmed this hypoth- (1932) subsequently proposed that they may be phy- esis (Aspo¨ck et al. 2001, Winterton 2003, Haring and tophagous or at least saprophagous on the bark of plant Aspo¨ck 2004, Winterton et al. 2010). The divergence roots. This is yet to be conÞrmed, but obvious simi- time for the Ithonidae and Polystoechotidae clade is larities in larval morphology between ithonids and estimated as during the Late based on molec- polystoechotids suggest a common feeding biology. ular data, with Ithonidae diverging from Polysto- Ithonid larvae were recorded by Gallard (1932) to echotidae during the (Winterton et al. 2010). congregate around the bases of eucalyptus trees and Fossil Polystoechotidae are described from the Middle observed “chaÞng,” shredding and actively feeding on Jurassic to Late (Lambkin 1988, Ren et al. the bark and roots to extract either sap or decaying 2002, Makarkin and Archibald 2003, Archibald and liquid material. Similar congregational patterns have Makarkin 2006), whereas “rapismatid-like” fossil been recorded by Faulkner (1990) around the bases of ithonids have recently been described from the Early creosote bushes [Larrea tridentata (DC) Coville] by and Early Eocene (Makarkin and Menon Oliarces and around trees in montane regions by 2007, Makarkin and Archibald 2009). Polystoechotidae. The family-level placement of some ithonids has Based on detailed studies of larval head morphol- been questioned previously. Tillyard (1926) included ogy, MacLeod (1964) proposed that Ithonidae and Oliarces in Ithonidae that was supported by Carpenter Polystoechotidae were sister-families closely related (1951), but Lameere (1936) suggested that the genus to (containing the families As- belonged to Polystoechotidae. This was supported by calaphidae, Myrmeleontidae, Nemopteridae, Nym- a phylogenetic analysis using morphology and multi- phidae, and Psychopsidae). Phylogenetic studies us- ple molecular markers by Winterton et al. (2010), ing morphology, DNA sequence data, or a which strongly indicated that Oliarces should be trans- July 2010 WINTERTON AND MAKARKIN:ITHONIDAE AND POLYSTOECHOTIDAE PHYLOGENY 513

Table 1. Exemplars used in this study for sequencing and/or morphological scoringa

Specimen Family/subfamily Exemplar 16S 18S CAD Collection data or reference voucher/code Outgroup Myrmeleontidae Stilbopteryx costalis CASENT8092178 EU734908 EU815291 EU860159 Winterton et al. (2010) Newman Nymphidae Myiodactylus osmyloides CASENT8092179 EU734873 EU815255 EU860126 Winterton et al. (2010) Brauer Osmylidae Porismus strigatus CASENT8092210 EU734894 EU815278 EU860147 Winterton et al. (2010) (Burmeister) Ingroup Ithonidae Adamsiana curoi Penny HONDURAS: Cortes: Cusuco ͓15Њ 30ЈN, 88Њ 13ЈW͔, 1900 m, 20 July 1994, Dan Curoe Allorapisma chuorum Makarkin and Archibald (2009) Makarkin & Archibald Ithone fulva Tillyard CASENT8092184 EU734865 EU815247 EU860118 Winterton et al. (2010) Megalithone tillyardi CASENT8092183 EU734866 EU815248 EU860119 Winterton et al. (2010) Riek Oliarces clara Banks CASENT8092185 EU734885 AF012527 EU860138 Winterton et al. (2010) Principiala incerta Makarkin and Menon (2007) Makarkin & Menon Rapisma cryptunum New CASENT8092186 EU734899 U815282 EU860151 THAILAND: Nakhon Nayok Khao & Barnard Yai Khao Keow spirit house, 14Њ 22.960Ј N, 101Њ 23.253Ј E, 750 m, Malaise trap, 5Ð12-IX-2006, Pong Sandeo leg. T909 Varnia implexa (Nava´s) AUSTRALIA: Queensland: 5 km E Charleville, 17.vi.1983, R. Silcock, T.6185 Polystoechotidae Fontecilla graphicus CASC201 EU734863 EU815244 EU860115 Winterton et al. (2010) Nava´s Platystoechotes lineatus CASENT8092170 EU734890 EU815274 EU860143 Winterton et al. (2010) Carpenter Polystoechotes punctatus CASENT8092171 EU734893 EU815277 EU860146 Winterton et al. (2010) (F.)

a Accession numbers indicate specimens sequenced for various genes, with gene sequences deposited in GenBank (http://www.ncbi. nih.gov/). Voucher numbers identify individual specimens deposited in the California Academy of Sciences, San Francisco, CA. ferred from Ithonidae to Polystoechotidae. Recent Platystoechotes, and Fontecilla) and Ithonidae (Adam- collections of various genera of Ithonidae and Polys- siana Penny, Ithone, Megalithone, Oliarces, Rapisma, toechotidae, suitable for DNA sequencing, have pre- and Varnia), except Narodona Nava´s as this genus is sented an opportunity to undertake a molecular phy- known only from a brief description and hind wing logenetic study of these comparatively rare to Þgure (Table 1; Fig. 2). Two relatively complete fossils try to elucidate their phylogenetic placement. A phy- of ithonids also were included (Principiala and Al- logeny for all extant genera of Polystoechotidae and lorapisma ). Outgroups were included from lacewing Ithonidae is presented based on morphology and mo- families Osmylidae (Porismus strigatus [Burmeister]), lecular DNA sequence data. Phylogenetic analyses Nymphidae (Myiodactylus osmyloides Brauer), and were undertaken using maximum parsimony and Myrmeleontidae (Stilbopteryx costalis Newman). Bayesian inference methods. Two extinct species of Twenty-three adult and larval characters were scored Ithonidae (Principiala Makarkin & Menon and Al- for all taxa, where data are available (Appendix 1). The lorapisma Makarkin & Archibald) also are included two extinct species represented by compression fossils in the analysis. The status of Ithonidae and Polysto- could not be scored for all characters as some could echotidae as separate families, or alternatively, united not be observed. as a single family, is discussed. We conclude that the Gene Sequencing. All gene sequences were down- observed phylogeny better corresponds to the latter, loaded from GenBank (Table 1) except Rapisma cryp- and a revised diagnosis and membership of the family tunum New & Barnard. DNA sequencing for this ad- Ithonidae s.l. is proposed. ditional taxon was carried out following the identical protocol outlined by Winterton et al. (2010). Gen- Bank accession and specimen voucher numbers are Materials and Methods presented in Table 1. Terminology. Wing venation terminology used Sequence Alignment and Phylogenetic Analysis. here generally follows Oswald (1993) and genitalic Alignment of all sequences was done manually, al- terminology follows Oswald (1998a). though CAD was aligned with reference to translated Exemplar Selection. Ingroup taxa included all ex- amino acid sequences (standard eukaryote) by using tant genera of Polystoechotidae (Polystoechotes, MacClade, version 4.06 (Maddison and Maddison 514 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 103, no. 4

Fig. 2. Wings of Polystoechotidae and Ithonidae. (A) Rapisma cryptunum New & Barnard. (B) Oliarces clara Banks. (C) Platystoechotes lineatus Carpenter. (D) Varnia implexa (Nava´s). (E) Ithone fulva Tillyard. Scale ϭ 2.0 mm. Abbreviations: 1A, 2A, 3A, Þrst to third anal veins; CuA, anterior cubitus; CuP, posterior cubitus; MA and MP, anterior and posterior branches of media; phs, prehumeral space; R1, Þrst branch of radius; Rs, radial sector; Sc, subcosta.

2000). Parsimony analyses were conducted using pseudoreplicates of resampled data sets, each with 30 PAUP*4.0b10 (Swofford 1999) using a branch and random additions (constant characters excluded). bound search protocol. Bootstrap support values for Bayesian analyses were preformed using MrBayes the parsimony analyses were calculated from 1,000 3.1.2 (Ronquist and Huelsenbeck 2003). The data heuristic search (tree bisection and reconnection) were partitioned by data type (DNA sequence and July 2010 WINTERTON AND MAKARKIN:ITHONIDAE AND POLYSTOECHOTIDAE PHYLOGENY 515

Table 2. Character matrix for 23 morphological characters lithone. Ithonidae were consistently divergent from for three outgroup taxa (Porismus, Myiodactylus, and Stilbopteryx) Polystoechotidae by 7.5Ð10.9% (inclusive of Rapisma and 11 ingroup taxa representing all extant, and two extinct, genera of Ithonidae and Polystoechotidaea and Oliarces). Inclusion of fossil exemplars in the analysis is likely Porismus 0021001000 0000200000 000 to weaken the Þnal estimate due to the low amount of Myiodactylus 0020110000 1000200100 001 data available (18 informative characters). Therefore Stilbopteryx 0020001001 0000100000 001 initial parsimony and Bayesian analyses of the com- Adamsiana 1100010001 1100110120 002 Allorapisma ?101010101 0001?0???? ??2 bined molecular and morphological data were under- Fontecilla 1110110010 0100200000 002 taken with Allorapisma and Principiala excluded. Ithone 1100000000 0010000011 112 The results of the Bayesian analysis are presented in Megalithone 1100000000 0010000011 112 Fig. 3; six most parsimonious trees were recovered Oliarces 1100000001 1000100021 002 ϭ Platystoechotes 1110110110 0100200000 002 from the parsimony analysis (tree length 2106, con- Polystoechotes 1110100010 0100000000 002 sistency index [CI] ϭ 0.708, retention index [RI] ϭ Principiala 1101110100 0001?0???0 ??2 0.492), although the resolved parts of the topology Rapisma 1101010101 1/00101100 002 were the same as the Bayesian tree. Two polytomies Varnia 1101000100 0010200011 112 in the parsimony tree represented unresolved rela- a Polymorphisms are indicated by a forward slash representing tionships among Adamsiana, Oliarces, and Rapisma as states 0 or 1. well as Ithone, Megalithone, and Varnia, respectively. All other branches on the tree were resolved with relatively high branch support. Adamsiana, Oliarces, morphology), locus, and by the remaining two codon and Rapisma were recovered as a clade deep within positions for each protein-coding locus. A separate Polystoechotidae, subtended by a laddered grade con- GTR ϩ ␥ nucleotide substitution model was applied to sisting of Platystoechotes, Polystoechotes, and Fontecilla. each DNA partition. The mk1 model (Lewis 2001), Ithone, Megalithone, and Varnia formed a well sup- with coding set to variable, was applied to the mor- ported clade sister to Polystoechotidae. phology partition. Each analysis consisted of four In a more inclusive analysis, all 14 taxa (including Markov chain Monte Carlo chains run simultaneously Allorapisma and Principiala ) were compared in a for 10 million generations. Trees were sampled every combined molecular and morphological data analysis, 1,000th generation. The Þrst 1 million trees were dis- consisting of 616 (18%) parsimony informative char- carded as burn-in. A majority rule consensus tree was acters and 3,886 parsimony uninformative characters computed with posterior probabilities (PP) for each (including 3,395 constant characters). The parsimony node. Morphological characters were treated as un- analysis yielded three trees (length ϭ 2,110 steps; CI ϭ ordered and equally weighted, with unknown states 0.707; RI ϭ 0.494) of similar topology to the reduced treated as Ô?Õ taxon set phylogeny, with Allorapisma and Principi- ala placed a sister to the clade consisting of Adam- siana, Oliarces, and Rapisma. A Bayesian phylogram, Results with parsimony bootstrap values and Bayesian poste- The combined sequence length for our concate- rior probability values is presented in Fig. 4. In this nated molecular data set was 4479 bp (CAD ϭ 2145 bp; analysis, the two fossil genera are placed as sister to 18S ϭ 1848 bp; 16S ϭ 486 bp) once ambiguously Rapisma; otherwise, the tree topology is the same as aligned sites (i.e., hypervariable stem regions in ribo- the parsimony analysis. Lower posterior probability somal genes) were removed. Addition of 23 morpho- values on certain branches correspond to lower sup- logical characters (Table 2) increased the combined port or unresolved topologies in the parsimony tree. data set to 4,502 characters. Average base frequencies Adamsiana, Oliarces, and Rapisma are again placed across the data set for 16S were greatly A/T biased (A, deep within Polystoechotidae, with relatively high 34.6%; C, 11.04%; G, 17.83%; T, 36.53%) (␹2 ϭ 17.419, branch support. Ithonidae are again a well supported df ϭ 39, P ϭ 0.998), whereas for 18S sequences they clade (88% bootstrap; 0.9 PP) represented by the ex- were in relatively equal proportions except for a tant Australian genera Megalithone, Ithone, and Varnia. slightly lower cytosine content (A, 25.73%; C, 22.27%; Ithonidae ϩ Polystoechotidae were recovered as a G, 26.69%; T, 25.30%) (␹2 ϭ 4.757.593, df ϭ 39, P ϭ monophyletic clade with relatively strong branch sup- 1.000). Average base frequencies for CAD (all sites) port (73% bootstrap; 0.9 PP). were A, 32.87%; C, 15.38%; G, 20.48%; and T, 31.26% Morphological characters supporting the mono- (␹2 ϭ 77.666, df ϭ 39, P ϭ 0.002) and as is common with phyly of the clade Ithonidae ϩ Polystoechotidae in- third positions, A/T bias was considerably greater clude head retracted under prothorax (char. 1), hu- (77.78%) than in either Þrst (51.04%) or second meral veinlet recurrent with multiple branches (62.56%) codon positions. Average uncorrected se- toward wing base (char. 2), and larval jaws short and quence divergences across taxa for all DNA loci varied broad for chafÞng (char. 23); characters one and 23 between the outgroup Porismus and most ingroup taxa are apparently unique within the order and are a at 13.9Ð15.5%, whereas between various ingroup gen- synapomorphy for Ithonidae ϩ Polystoechotidae. As era, pairwise divergences ranged from 1.2% between was found in Winterton et al. (2010), in all analyses closely related Ithone and Megalithone and 9.9% be- Ithonidae was rendered paraphyletic with respect to tween the more distantly related Rapisma and Mega- Polystoechotidae with relatively high levels of sup- 516 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 103, no. 4

Fig. 3. Bayesian phylogram of relationships of extant taxa of Ithonidae and Polystoechotidae based on analyses of DNA sequence data and 23 morphological characters. Branch length represents number of changes. Bayesian posterior probability and parsimony bootstrap support values are presented in order on each branch. Outgroups are Osmylidae (OSMY), Nymphidae (NYMP), and Myrmeleontidae (MYRM). port. Three genera presently placed in Ithonidae, i.e., ces) (char. 20), male sternite 9 shovel-like (char. 21), Adamsiana, Oliarces, and Rapisma, were placed as a and female sternite 9 forming a psammorotrum (char. monophyletic clade deep within Polystoechotidae, al- 22). though no discreet morphological characters could be found to support inclusion in this clade in the family. Discussion Adamsiana, Rapisma, and Oliarces are supported as a clade by wing trichosors being absent (char. 10) and The results of this analysis using a combination of pale wing venation (char. 11). When the fossil genera DNA sequence data and morphology support revised Allorapisma and Principiala were included in the deÞnitions and taxon composition for both Polysto- analysis, they formed a sister-group relationship with echotidae and Ithonidae, although treating the clade Rapisma, although branch support within the clade as a single family should also be considered. comprising Adamsiana, Rapisma, Oliarces, Al- Ithonidae, based on these results, comprises only lorapisma , and Principiala became nonexistent. The the endemic Australian genera Ithone, Megalithone, sister-group relationship between Allorapisma and and Varnia. Synapomorphies supporting Ithonidae in Principiala is supported by the divergent branching this context include forewing with two radial branches pattern of the forewing medial vein (char. 14) (Makar- originating on R, mediuncus triangular, male sternite kin and Menon, 2007, Þgs. 3 and 5; Makarkin and 9 shovel-like, and female sternite 9 forming a psam- Archibald 2009, Þgs. 2 and 3). The Australian genera morotrum. Here, genera previously placed in of Ithonidae (Ithone, Megalithone, and Varnia) were Ithonidae, i.e., Rapisma, Oliarces, and Adamsiana, recovered as a monophyletic clade in all analyses were recovered within Polystoechotidae along with (bootstrap support ϭ 88Ð99%; PP ϭ 0.89Ð1.0) with Polystoechotes, Platystoechotes, and Fontecilla. Conse- Megalithone weakly supported as sister to Varnia in the quently, Polystoechotidae are predominantly distrib- pruned analysis. Five characters are synapomorphies uted in the New World (Þve genera), with the single, for this clade: forewing with two radial branches orig- relatively species-rich genus Rapisma, distributed inating on R (char. 13), mediuncus triangular (char. throughout Southeast Asia. Multiple fossils of putative 19), male ectoprocts enlarged (homoplasious in Oliar- Polystoechotidae have been described, but deÞnitive July 2010 WINTERTON AND MAKARKIN:ITHONIDAE AND POLYSTOECHOTIDAE PHYLOGENY 517

Fig. 4. Phylogram of relationships of genera of Ithonidae and Polystoechotidae based on matching topologies from Bayesian and parsimony analyses of DNA sequence data and 23 morphological characters. Two additional extinct genera (Allorapisma Makarkin & Archibald, Principiala Makarkin & Menon) of Ithonidae also are included. Branch length represents number of changes. Bayesian Posterior Probability and bootstrap support values are presented in order on each branch. Wing drawings of Principiala and Allorapisma are modiÞed after Makarkin and Menon (2007) and Makarkin and Archibald (2009), respectively. 518 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 103, no. 4 family-level assignments have been problematic for clade. Convincing mutually exclusive synapomorphies some taxa (Lambkin 1988, Ren et al. 2002, Makarkin supporting both families are lacking, either as previ- and Archibald 2003, Archibald and Makarkin 2006). ously deÞned or based on our combined analyses. It The reason for this confusion is that synapomorphies seems that the most reasonable conclusion is to com- deÞning Polystoechotidae are mostly lacking and bine Ithonidae (including Rapismatidae) with Polys- characters used to differentiate the family are often toechotidae as a single family containing all the genera plesiomorphies (Makarkin and Archibald 2003, presently included in these two families. Archibald and Makarkin 2006). In conclusion, it is clear from the combined mor- The placement of Oliarces and Rapisma is not sur- phological and DNA sequence data presented here prising with similarities between the two genera, and that the separation of Ithonidae and Polystoechotidae with Polystoechotidae, noted by previous authors. is not warranted. Our results strongly support either Lameere (1936) suggested that Oliarces belonged to the transfer of Adamsiana, Oliarces, and Rapisma to Polystoechotidae, and Grebennikov (2004) described Polystoechotidae, or combining all the taxa into a a close morphological similarity of Oliarces Þrst instars single family. Morphological characters deÞning to a putative polystoechotid. Makarkin and Archibald Polystoechotidae relative to Ithonidae are few, but (2009) questioned the identity of this larva, suggesting there are some putative synapomorphies deÞning a that it could be an ithonid. Tillyard (1916, 1919b) single family, including adult and larval morphology suggested that Rapisma and Oliarces do not belong in and larval biology. Consequently, we have chosen the Ithonidae but require separate family status rather latter option as a more reasonable alternative to main- than placement in Polystoechotidae. Barnard (1981) taining Ithonidae and Polystoechotidae as separate, supported the separation of Rapismatidae as a distinct but poorly deÞned, family groups. Further studies are family and listed eight characters that separated this required on the group, in particular, acquisition of family from Ithonidae. In his description and discus- DNA sequence data for Narodona, Adamsiana, and sion of Adamsiana, Penny (1996) considered each of Varnia as well as biological studies of the larval mor- these characters in turn, with the new genus exhibiting phology and feeding biology of all species. character states of both Ithonidae and Rapismatidae and therefore suggested that the separation of the two families was not necessary. Neither Barnard (1981) Family Ithonidae Newman, 1853, sensu novo nor Penny (1996) considered Polystoechotidae in Ithonesidae Newman, 1853: ccii [incorrect formation their studies. Based on internal female genitalic mor- of family name]. Type genus: Ithone Newman, 1838. phology, Szira´ki (1998) indicated that Rapisma Ithonidae: Tillyard, 1916: 274 [corrected name for showed a closer similarity with Polystoechotidae than Ithonesidae]. Ithonidae. Our results are congruent with this con- Polystoechotidae Handlirsch, 1906Ð1908 [1906]: 42. clusion, with Rapisma placed within Polystoechotidae Type genus: Polystoechotes Burmeister, 1839. Syn. and that the genus does not deserve separate family nov. status. Rapismiden Kru¨ ger, 1922: 139. Type genus: Rapisma The Mexican ithonid genus Narodona is known only MacLachlan, 1866. from the original description and Þgure of the hind Rapismidae: Kru¨ ger, 1923: 72, 73. wing, with the type probably destroyed (Monserrat Rhapismidae Nava´s, 1929b: 376 (as “fam. nov.”). Type 1985). Based on the wing Þgure, this genus shows a genus: Rhapisma [unjustiÞed emendation of Rap- clear afÞnity with Adamsiana (Penny 1996) and is isma]. most likely closely related. Given that Adamsiana is Rapismatidae: Barnard, 1981: 122 (corrected name for placed within Polystoechotidae in this analysis, it Rhapismidae). is clear that Narodona is probably a polystoechotid as well. Diagnosis. Robust, medium-sized to large neurop- The incomplete yet highly autapomorphic wing ve- terans (forewing length, Ϸ15Ð50 mm) distinguished nation of Principiala and Allorapisma suggests from similar families by the presence of the following placement within Polystoechotidae. The sister-group features: Head more or less retracted under pronotum; relationship with Rapisma supports previous sugges- forewing elongate oval to subtriangular (broad-trian- tions by Makarkin and Menon (2007) of a close rela- gular in Brongnairtiellidae, Osmylopsychopidae); an- tionship among these genera based on overall simi- tehumeral space long and broad in both wings, dis- larities in wing venation and body shape. proportionately large in genera with relatively narrow Is there justiÞcation for maintaining two separate costal space; humeral veinlet well developed, recur- families, Ithonidae and Polystoechotidae? The sister rent, pectinate branched in both wings (in hindwing group relationship between Ithonidae ϩ Polysto- with at least one branch) (absent in hindwing of echotidae is supported by two morphological synapo- ; absent in both wings of Dilaridae); ϩ morphies: head retracted under prothorax in adults apical area between C and R1 (or Sc R1) markedly and larval jaws very short and broad. Although larval expanded (except perhaps Rapisma), with long, ϩ biological data are incomplete for extant taxa (and mainly forked veinlets of R1 or Sc R1 in both wings nearly impossible for extinct taxa), it is likely that (not markedly expanded in Hemerobiidae, Dilaridae); being fossorial, along with the probable phytophagous distal nygma present between two proximal branches feeding biology, represent synapomorphies for this of Rs in both wings (absent in Brongnairtiellidae, Psy- July 2010 WINTERTON AND MAKARKIN:ITHONIDAE AND POLYSTOECHOTIDAE PHYLOGENY 519 chopsidae, Hemerobiidae). Larval jaws relatively Ñ Hind wing with prominent maculae, partic- short and very broad basally; larva fossorial, associated ularly in posterior region ...... 9 with plant roots. 9. Six to nine R1-Rs cross-veins in both wings (south- Genera Included. Kasachstania PanÞlov 1980, Os- western United States)...... Platystoechotes myloides PanÞlov 1980 and Paleopterocalla Oswald, Ñ Less than three R1-Rs cross-veins in both 2007 (ϭPterocalla PanÞlov 1980; all from the Late wings (Chile) ...... Fontecilla Jurassic of Karatau, ); Principiala Makar- kin et Menon, 2007 (Early Cretaceous of Brazil and Appendix 1: Descriptions of Morphological England); Palaeopsychops Andersen, 2001 (Early Eo- Character States cene of Denmark and western North America); Al- lorapisma Makarkin et Archibald, 2009 (Early Eo- 1. Head. (0) clearly visible from above; (1) partially cene of western North America); Ithone Newman, concealed beneath anterior margin of prothorax (Fig. Megalithone Riek, Varnia Walker (Australia); Rapisma 1). The derived state of this character is a synapomor- (Oriental Region); Adamsiana Penny, Oliarces Banks, phy for Polystoechotidae ϩ Ithonidae and has been Narodona Nava´s, Polystoechotes Burmeister, Platysto- noted previously by Makarkin and Menon (2007). echotes Carpenter and Fontecilla Nava´s (North and 2. Forewing Humeral Veinlet. (0) simple, cross- South America). vein-like (Oswald 1998a: Þg. 1); (1) recurrent, Comments. Ithonidae in a broader sense (i.e., in- branched (Fig. 2BÐE). The presence of the well de- clusive of Polystoechotidae and Rapismatidae) as de- veloped recurrent and branched humeral veinlet is Þned here includes six extinct and 10 extant genera characteristic of the order Neuroptera. It occurs in restricted to Australia, the Oriental region and the psychopsoid-like families, Ithonidae, Polystoechoti- New World. The family originated during the Triassic- dae, Hemerobiidae, and some Berothidae and Man- Jurassic, more widely distributed during the Jurassic tispidae. In this analysis, the state is considered a than the present (with many undescribed taxa from synapomorphy of the genera of Ithonidae and Polys- Chinese Jurassic deposits). toechotidae; it has probably evolved independently at least three times, in Hemerobiidae, Psychopsidae (and related fossil groups), and the clade Berothidae ϩ Key to Extant Genera of Ithonidae s.l. Mantispidae. 1. Wing venation pale green or white; wings rarely 3. Proximity of Forewing Subcostal (Sc) and Radial

maculate ...... 2 (R) Veins. (0) Sc and R1 completely separate distally Ñ Wing venation dark; wings usually strongly (Fig. 2A and B, C, E); (1) closely approximated but not maculate or smoky infuscate ...... 4 fused (Fig. 2C); (2) fused (Oswald, 1998a: Þg. 1). Most 2. Wing venation brilliant white; body dark of the ingroup exhibit the plesiomorphic state (0) with

(southwestern United States) .....Oliarces the Sc and R1 veins separate along entire length, Ñ Wing venation and body mostly pale (vivid whereas Fontecilla, Polystoechotes, and Platystoechotes green in live individuals) ...... 3 have the veins closely approximated distally (1). Only

3. Wings usually spotted or banded; subcostal the outgroups have Sc and R1 fused toward the apex veinlets connected by numerous cross-veins of the wing (2). In the majority of the fossil species of so that serial ranks of cells present; female Polystoechotidae not included in this analysis, how- macropterous (Oriental region) . . . Rapisma ever, these veins are fused.

Ñ Wings without infuscate patterning, mem- 4. Number of Forewing Subcostal (Sc-R1) Cross- brane hyaline; subcostal veinlets not con- Veins. (0) one, near wing base; (1) numerous cross- nected by cross-veins, single rank of cells; veins along length of subcostal area. The plesiomor- female apterous (New World) . . Adamsiana phic state was most numerous, with the derived state 4. Forewing R vein with additional branch, basal to present in Porismus, Varnia, Rapisma, Allorapisma , pectinate Rs vein (Fig. 2E) (Australia) . . . 5 and Principiala . Ñ Forewing R vein with a single pectinate Rs 5. Number of Forewing Rs Branches. (0) 4Ð8; (1) vein (New World) ...... 7 9Ð12. Relatively few Rs branches is considered in this 5. Wings mottled; cross-veins interlinking sub- analysis as plesiomorphic and is found in Porismus, costal veinlets, creating multiple ranks of Stilbopteryx, and Ithonidae. The polarity of this char- cells ...... Varnia acter is not clear and in general, the states of character Ñ Wings uniform infuscate; single row of cells may depend largely on the wing size. in costal space ...... 6 6. Forewing Costal Space. (0) relatively narrow 7. Fore basitarsus at most no longer than apical basally and equal width along entire length (Fig. 2B); tarsomere; male ectoprocts enlarged and wid- (1) distinctly dilated basally (Fig. 2C). In Porismus, ened at apex ...... Megalithone the costal space is narrow basally and distinctly dilated Ñ Fore basitarsus much longer than apical tar- at some distance from the base; state (0) is used in the somere; male ectoprocts narrow and rounded analysis. This character is highly variable among all at apex; wing uniform infuscate; single row of ingroup and outgroup taxa. cells in costal space ...... Ithone 7. Forewing Subcostal Veinlets. (0) forked (Fig. 8. Hind wing without prominent maculae 2AÐE); (1) simple (Carpenter 1940: Þg. 73). The (pterostigma slightly marked) ....Polystoechotes forewing subcostal veinlets have extensive terminal 520 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 103, no. 4 branching in all ingroup taxa; the amount of branching hindwing sigmoid vein was considered evidence for is much less in Myiodactylus. coalescence of MA with the radial vein in the forewing 8. Forewing Subcostal Veinlets. (0) not intercon- (e.g., Carpenter 1951), but now is interpreted as a nected (Fig. 2B and E); (1) interconnected by cross- cross-vein or “a cross-vein brace” (e.g., see Kukalova´- veins, forming multiple ranks of cells between costal Peck and Lawrence 2004: Þg. 5). The sigmoid form (0) margin and Sc (Fig. 2A and C, D). The derived state, is plesiomorphic for Neuropterida and is exhibited by with the subcostal veinlets interconnected and form- Megalithone, Ithone and Polystoechotes. Stilbopteryx, ing multiple ranks of cells is found in Principiala , Principiala , Adamsiana and Oliarces have a straight Allorapisma , Varnia, Platystoechotes, and Rapisma. vein. In Porismus the vein is apparently absent or The genus Fontecilla possesses a few cross-veins, ir- greatly reduced (New 1983), but it is sigmoid in other regularity placed but not is serial ranks. Osmylidae genera (e.g., Osmylus Latreille). Rapisma 9. Forewing Gradate Series. (0) irregularly ar- and Varnia show interspeciÞc polymorphism in this ranged, more than two series present (Fig. 2A and D, character as the vein is present or absent between E); (1) two distinct series only (Fig. 2C). The derived species. state, with gradates arranged in two series is found 16. Female Wings. (0) macropterous; (1) apterous. only in Platystoechotes, Polystoechotes and Fontecilla. The apterous female is autapomorphic for Adamsiana 10. Forewing Trichosors. (0) present along entire (Penny 1996). wing margin; (1) absent entirely, or present in only 17. Tarsal Claw Shape. (0) narrow, without projec- small portion of wing margin. The presence of wing tions (Riek 1974: Þgs. 1 and 2); (1) relatively broad trichosors is a synapomorphy for Neuroptera. The basally with small projection (Barnard 1981: Þg. 7). trichosors are frequently reduced in number or com- The derived state of this character is autapomorphic pletely absent in many families, as found here in Stil- for Rapisma (Penny 1996, Makarkin and Menon 2007). bopteryx and the clade Adamsiana, Oliarces, Al- 18. Tibial Spurs. (0) present (Riek 1974: Þgs. 1 and lorapisma , and Rapisma. 2); (1) absent. Tibial spurs are likely plesiomorphic 11. Wing Venation. (0) dark (black or brown) (Fig. throughout Neuroptera with numerous instances of 2CÐE); (1) pale (white to light green) (Fig. 2A and B). secondary reduction, here exempliÞed by Myiodacty- Pale wing venation is derived and is present in Myio- lus and ingroup genera Adamsiana and Rapisma. dactylus and ingroup genera Oliarces, Rapisma, and 19. Male Genitalia, Mediuncus Shape. (0) as a lobed Adamsiana. In Adamsiana, Myiodactylus, and Rapisma, plate with ventral processes (Barnard 1981: Þgs. 12 and the body and wings are green in living specimens, 15); (1) triangular (Acker 1960: Þgs. 58 and 60; as whereas in Oliarces the body is dark and wing veins are coxopodite 9); (2) reduced or absent (Penny 1996: vivid white (Fig. 1B). Þgs. 4 and 5). Based on examinations of specimens of 12. Forewing Shape. (0) symmetrical, largely ovoid Oliarces in this study, it is clear that previous inter- (Fig. 2E); (1) asymmetrical tending toward falcate (Fig. 2C). An asymmetrical forewing is a synapomor- pretations of the gonarcus being divided medially into phy for Polystoechotidae, although it is secondarily two parts as illustrated by (Acker 1960: Þg. 52) are symmetrical in Oliarces, Allorapisma , Principiala , incorrect, with only a single plate present. An inter- and some Rapisma. pretation of the gonarcus in Ithone as triangular is 13. Forewing Radial Branches. (0) single, pectinate incorrect as well (Barnard 1981), as the darkly scle- Rs originating from R (Fig. 2AÐC); (1) two branches rotized mediuncus is triangular (1) in Ithone, Megali- originating from R, the distal branch being the pecti- thone, and Varnia, with a weakly sclerotized and nate Rs (Fig. 2D and E). The plesiomorphic condition arched gonarcus present immediately dorsal of the occurs throughout Neuroptera. The derived condition mediuncus. The mediuncus is a medial, lobed, plate- represents a synapomorphy for Ithonidae s.s. (i.e., like structure as the plesiomorphic state (0) in the Ithone, Megalithone, and Varnia) and has evolved in- outgroups, Rapisma, Fontecilla, Polystoechotes, and dependently in Hemerobiidae and some Dilaridae and Platystoechotes, whereas it is weakly formed in Oliarces Kalligrammatidae. Carpenter (1951) incorrectly Þg- and Adamsiana (2). ured the wing of Ithone fusca Newman by omitting this 20. Male Ectoprocts. (0) not enlarged (Penny 1996: basal free radial branch, instead showing only a single, Þg. 3); (1) enlarged (Riek 1974: Þgs. 4 and 8). The male pectinate Rs originating on R. ectoprocts are enlarged in Ithone, Megalithone, Varnia, 14. Fork of Forewing MP. (0) branches closely and Oliarces. proximate, diverging acutely and only composed of 21. Male Sternite 9. (0) simple, not elongated (Bar- two to three branches reaching wing margin (Fig. nard 1981: Þg. 29); (1) elongated and shovel-like (Riek 2AÐE); (1) branches widely divergent composed of 1974: Þgs. 4 and 8). The derived state is exhibited by more than Þve branches reaching wing margin extant members of Ithonidae s. str. (i.e., Megalithone, (Makarkin and Menon 2007: Þgs. 3 and 5; Makarkin Ithone, and Varnia). This character is not observable and Archibald 2009: Þgs. 2 and 3). The derived state is in the extinct species Principiala and Allorapisma . nine gonocoxites). (0) not؍) found only in the fossil genera Allorapisma and Prin- 22. Female Sternite 9 cipiala . enlarged (Barnard 1981: Þgs. 8 and 9); (1) enlarged 15. Hindwing Basal Cross-Vein r-m Shape. (0) sig- (Riek 1974: Þgs. 6 and 41). The enlarged female ster- moid (Fig. 2E); (1) straight (Fig. 2B); (2) reduced or nite 9 of Megalithone, Varnia, and Ithone forms a psam- apparently absent (New 1983: Þg. 2). The shape of the morotrum (Tillyard 1919a). July 2010 WINTERTON AND MAKARKIN:ITHONIDAE AND POLYSTOECHOTIDAE PHYLOGENY 521

23. Larval Jaw Shape. (0) elongate and straight for laridae (Neuroptera). Proc. Am. Acad. Arts Sci. 74: 193Ð piercing prey; (1) elongate, but curved for seizing 218. prey; (2) relatively short, with very broad base Carpenter, F. M. 1951. The structure and relationships of (Grebennikov 2004: Þgs. 7 and 24, 31). Elongate and Oliarces (Neuroptera). Psyche 58: 32Ð41. straight jaws modiÞed for piercing (0) is plesiomor- Faulkner, D. K. 1990. Current knowledge of the biology of phic for Neuroptera (Winterton et al. 2010) and is the moth-lacewing Oliarces clara Banks (Insecta: Neu- found in Porismus; elongate and incurved apically jaws roptera: Ithonidae), pp. 197Ð203. In M. W. Mansell and H. Aspo¨ck (eds.), Advances in Neuropterology, Third In- of Myiodactylus and Stilbopteryx are used for seizing ternational Symposium, Kruger National Park, South Af- prey (1). Larval stages known for Ithone, Megalithone, rica, 3Ð4 February 1988. Department of Agricultural De- Polystoechotes, Oliarces, and Platystoechotes and all ex- velopment, Pretoria, South Africa. hibit the derived state of short jaws with a board base Gallard, L. 1932. Notes on the feeding habits of the brown (2) (MacLeod 1964, Faulkner 1990, Grebennikov moth-lacewing, Ithone fusca. Aust. Nat. 8: 168Ð170. 2004, Winterton et al. 2010). This type of jaw is ob- Grebennikov, V. V. 2004. Grub-like larvae of Neuroptera viously not suitable for piercing or seizing prey but (Insecta): a morphological review of the families instead is used for scraping or chafÞng the surface of Ithonidae and Polystoechotidae and a description of plant roots, as observed in Ithone by Gallard (1932). All Oliarces clara. Eur. J. Entomol. 101: 409Ð417. ingroup taxa were scored for this state. This type of Handlirsch, A. 1906–1908. Die fossilen Insekten und die specialized feeding biology and head morphology is a Phylogenie der rezenten Formen. Ein Handbuch fu¨ r putative synapomorphy for the entire clade Pala¨ontologen und Zoologen. W. Engelmann, Leipzig. Germany. Ithonidae ϩ Polystoechotidae. Haring, E., and U. Aspo¨ck. 2004. Phylogeny of the Neurop- terida: a Þrst molecular approach. Syst. Entomol. 29: 415Ð 430. Acknowledgments Kru¨ ger, L. 1922. Hemerobiidae. Beitra¨ge zu einer Monog- raphie der Neuropteren-Familie der Hemerobiiden. We thank Justin Bartlett for help with preparation of wing Stett. Entomol. Zeit. 83: 138Ð172. images. Specimens of Rapisma were collected as part of the Kru¨ ger, L. 1923. Neuroptera succinica baltica. Die im bal- Thailand Inventory Group for Entomological Research tischen Bernstein eingeschlossenen Neuropteren des project supported by National Science Foundation (NSF) Westpreussischen Provinzial-Museums (heute Museum project DEB-0542864. We also thank Drs. John Oswald fu¨ r Naturkunde und Vorgeschichte) in Danzig. Stett. (Texas A&M University) and Norman Penny (California Entomol. 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