Research 78 (2017) 78e83

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Cretaceous Research

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The first moth lacewing (Insecta: : Ithonidae) from the mid-Cretaceous amber of Myanmar

* ** Xiumei Lu a, c, Weiwei Zhang b, Michael Ohl c, , Xingyue Liu a, a Department of Entomology, Agricultural University, Beijing 100193, China b Three Gorges Entomological Museum, P.O. Box 4680, Chongqing 400015, China c Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitaetsforschung Invalidenstraße 43, Berlin 10115, Germany article info abstract

Article history: The lacewing family Ithonidae is reported from the mid-Cretaceous amber of Myanmar for the first time. Received 12 February 2017 A new and species, Burmithone pennyi gen. et sp. nov., is herein described based on an almost Received in revised form completely preserved female specimen. The new genus exhibits a number of remarkable forewing 10 May 2017 characters, such as the proximal branches of RP vein fused with the MA vein and the peculiar configu- Accepted in revised form 25 May 2017 ration of MP and CuA. The systematic position of Burmithone gen. nov. is briefly discussed in comparison Available online 26 May 2017 with other genera of Ithonidae. © 2017 Elsevier Ltd. All rights reserved. Keywords: Neuropterida Ithonidae

1. Introduction veinlet, and the presence of nygmata. The larval stage of Ithonidae is unique because of its peculiar scarabaeiform (grub-like C-sha- Ithonidae sensu lato presents as an archaic small family of ped) older instar larvae (Grebennikov, 2004), which are subterra- Neuroptera, comprising the moth lacewings (Ithonidae sensu nean and suggested to be phytophagous (Gallard, 1932; Faulkner, stricto), montane lacewings (former Rapismatidae) and giant 1990). Both recent morphology-based and molecule-based phylo- lacewings (former Polystoechotidae) according to the recent genetic studies suggest a sister group relationship between Itho- taxonomic treatment, which placed the preceding three family- nidae and (Aspock€ and Aspock,€ 2008; group taxa into a single Ithonidae in the broad sense based on Winterton et al., 2010; Wang et al., 2017). total-evidence phylogenetic analysis (Winterton and Makarkin, Currently, Ithonidae include 10 extant genera and ca. 40 species 2010). Nonetheless, Zheng et al. (2016a) provided morphological worldwide. The earliest definite ithonid fossil is known from the evidence to divide Ithonidae sensu lato into three genus-groups, Middle of China (Zheng et al., 2016a), and 11 genera and 28 i.e., the Ithonid genus-group, the Polystoechotid genus-group, fossil ithonids are recorded from the interval between the Middle and the Rapismatid genus-group, although the group members Jurassic to the late (Makarkin et al., 2014; Zheng et al., of the preceding three genus-groups are not equivalent to that 2016a,b). Among them, only one species was found in amber, i.e., placed in the former three families and include many fossil the late Eocene Baltic amber (Makarkin et al., 2014). genera. Here we report a new genus and species of Ithonidae from the Most adults of Ithonidae sensu lato are characterized by the mid-Cretaceous amber of Myanmar based on a nearly completed robust body, the head more or less retracted under the pronotum, female specimen. It represents the first record of Ithonidae from the broad forewing costal space with branched recurrent humeral this deposit, where a remarkably diverse paleofauna of Neuro- ptera was present (see Makarkin, 2016). Some noteworthy morphological characters in the new ithonid provide new insights * Corresponding author. for understanding the taxonomy and evolutionary history of ** Corresponding author. Ithonidae. E-mail addresses: [email protected] (M. Ohl), [email protected] (X. Liu). http://dx.doi.org/10.1016/j.cretres.2017.05.027 0195-6671/© 2017 Elsevier Ltd. All rights reserved. X. Lu et al. / Cretaceous Research 78 (2017) 78e83 79

2. Material and methods

The amber sample under present study is from the Hukawng Valley in Tanai Township, Myitkyina District of Kachin State, Myanmar (see Kania et al., 2015: fig. 1). The age of this deposit has been investigated and dated to be ~99 MA (earliest ) by UePb dating of zircons from the volcaniclastic matrix of the amber (Shi et al., 2012b). The type specimen is currently housed in the Entomological Museum, China Agricultural University (CAU), Beijing, and it will eventually be deposited in the Three Gorges Entomological Museum (EMTG), Chongqing (specimen available for study by contacting WZ). Photographs and drawings were taken and made using a Zeiss SteREO Discovery V12 stereo microscope system. The figures were prepared with Adobe Photoshop CS4. Terminology of wing venation generally follows H. Aspock€ et al. (1980) and Kukalova-Peck and Lawrence (2004). Terminology of genitalia fol- lows Aspock€ and Aspock€ (2008). Abbreviations used for wing veins are: A, anal vein; C, costa; Cu, cubitus; CuA, cubitus anterior; CuP, cubitus posterior; M, media; MA, media anterior; MP, media posterior; R, radius; RA, radius anterior; RP, radius posterior; ScA, subcosta anterior; ScP, subcosta posterior.

3. Systematic palaeontology

Order Neuroptera Linnaeus, 1758 Family Ithonidae Newman, 1853 sensu Winterton and Makarkin, 2010 Genus Burmithone gen. nov. Figs. 1e4 urn:lsid:zoobank.org:act:6F8A3661-B2AA-46C0-8C0D- 29D61C898C42 Type species: Burmithone pennyi sp. nov. Diagnosis. The new genus can be distinguished from the other species of Ithonidae by a combination of the following characters: (1) head completely retracted under a nearly hexagonal pronotum [partly protruding from pronotum in the Polystoechotid genus- group]; (2) antenna almost half of forewing length [very short in the Polystoechotid genus-group and most species of the Rap- ismatid and Ithonid genus-groups, being less than half of the forewing length]; (3) nygmata absent [present in most genera of Ithonidae]; (4) forewing recurrent humeral veinlet angulately curved [arcuately curved in most genera of Ithonidae except Prin- cipiala Makarkin & Menon and Allorapisma Makarkin & Archibald]; Fig. 1. Burmithone pennyi gen. et sp. nov., holotype female. A. Habitus photograph, (5) interlink veinlets among costal crossveins largely reduced in dorsal review; B. Habitus photograph, ventral review; C. Photograph of tarsi. Scale bar: forewing costal space [forewing costal space with several or many 2.5 mm (A, B); 1.0 mm (C). interlink veinlets among costal crossveins in many genera of the Rapismatid and Ithonid genus-groups]; (6) ScP stout, ending general irregularly arranged [regularly arranged into at least one sharply curved and fused with RA into ScP þ RA that reaches gradate series in the Polystoechotid genus-group]. anterior margin straightly before wing apex [ScP and RA distinctly Etymology. From Burma (¼ Myanmar) and Ithone (the type genus- separated in most genera of the Rapismatid and Ithonid genus- group name of Ithonidae), in reference to the occurrence of the groups]; (7) forewing RP with proximal two branches fused with new genus from the mid-Cretaceous amber of Myanmar. Gender: MA, forming two long cells [such fusion absent in most genera of Feminine. Ithonidae except Principiala and Allorapisma]; (8) forewing MP Remarks. Placement of the new genus in Ithonidae is undoubted lacking primary forking [deeply branched in most genera of Itho- based on the following features: robust body; head completely nidae except Principiala and Allorapisma ]; (9) forewing CuA pro- retracted under pronotum; proximal part of forewing costal space fusely branched anteriad from its midpoint, forming a very broad with branched recurrent humeral veinlet; presence of subtriangular area [less branched in most genera of Ithonidae numerous and irregularly arranged crossveins. except Principiala and Allorapisma, or densely branched but with branches directed posteriad in e.g. Megalithone Riek and Platys- The new genus could be placed in a suprageneric taxon, namely toechotes Carpenter (see Fig. 4)]; (10) forewing with crossveins in the Principiala group, proposed by Makarkin and Archibald (2009) 80 X. Lu et al. / Cretaceous Research 78 (2017) 78e83

Fig. 2. Burmithone pennyi gen. et sp. nov., holotype female. A. Drawing of right forewing; B. Drawing of left forewing. Scale bar: 1.0 mm. by the proximal branches of RP fused with MA and the peculiar Diagnosis. As for the genus. configuration of MP and CuA, which is however interpreted Description. Female. Body length 7.91 mm; head 0.80 mm long and differently in this paper (see Discussion). The Principiala group 1.93 mm wide; antenna length 5.36 mm; pronotum 1.50 mm long previously comprised two fossil genera, i.e. Principiala from the and 2.40 mm wide; forewing 8.89 mm long and 4.20 mm wide; Lower Cretaceous of Brazil and England and Allorapisma from the abdomen length 4.42 mm. Foreleg tarsus length: 0.81 mm; claw early Eocene of U.S.A. Obviously, the new genus, Burmithone gen. length 0.10 mm; midleg claw length 0.15 mm; hind leg claw length nov. is closely related to Principiala and Allorapisma based on the 0.08 mm. unique feature of forewing venation within Ithonidae mentioned above. However, Burmithone gen. nov. can be distinguished from Head wide, but slightly thinner than pronotum; compound eyes Principiala by the relatively long antenna (5.36 mm long) [rather globular; antenna thickly filiform, nearly half of forewing length. short, ca. 2.0e2.5 mm in Principiala], the absence of series of Pronotum nearly hexagonal, stout; meso- and metanotum slightly interlink veinlets on the forewing costal space [a long series of narrower than pronotum. interlink veinlets (¼ costal gradate series of crossveins in Makarkin Forewing covered with dense short hairs; trichosors present and Menon, 2007)inPrincipiala], the elongate cell formed by fusion along costal and distal margin, with those on costal margin more of proximal-most branch of RP and MA in the forewing [short in distinct. Costal space very broad proximally and distinctly nar- Principiala], the relatively sharp angle between primary branches of rowed near wing apex; humeral veinlet strongly recurrent, almost forewing CuA (¼ MP in Makarkin and Menon, 2007) [primary forming a right angle, with 7e8 branches, most of which have branches of forewing CuA forming much larger angle], and the additional fork; interlink veinlets absent except for a single short short hind wing humeral plate without frenulum [long with fren- one present between the first branch of recurrent humeral veinlet ulum in Principiala]. Besides the above three forewing characters, and proximal-most costal crossvein; ca. 26 costal crossvein present, the new genus differs from Allorapisma also by the narrow forewing all of them distinctly inclined toward wing apex, most costal RA space [RA space much broader than subcostal space in Allor- crossveins with additional fork near costal margin. ScA straight and apisma] and the forewing CuP with only marginal fork [CuP pec- nearly perpendicular to ScP. ScP stout, ending sharply curved and tinately branched at least from its midpoint in Allorapisma]. fused with R before reaching anterior margin straightly before wing apex; six simple veinlets of ScP þ RA present. Subcostal space Burmithone pennyi sp. nov. present 14 subcostal crossveins. R rather stout proximally, seem- Figs. 1e4 ingly fused with ScP; RP þ MA originated from R nearly at proximal urn:lsid:zoobank.org:act:653AAD4D-74DC-4AA8-AE5F- 1/5, with proximal two branches of RP fused with MA, forming two EA5384096087 long cells; remaining 5e6 branches of RP distally forked near wing X. Lu et al. / Cretaceous Research 78 (2017) 78e83 81

Type material. Holotype: EMTG BU-002284: Amber piece preser- ving an almost complete female of Burmithone pennyi sp. nov. It is polished in the form of a flattened rectangle cabochon, clear and transparent, with length width about 23.0 11.5 mm, height about 7.0 mm. Etymology. The new species is dedicated to the late Dr. Norman D. Penny, Academy of Sciences, San Francisco, who made outstanding contributions particularly on the New World Neuro- ptera, including description of the ithonid genus Adamisiana from the Neotropics.

4. Discussion

4.1. Remarkable configuration of forewing MP and CuA in Burmithone gen. nov.

Burmithone gen. nov., together with Principiala and Allorapisma, as mentioned above, belongs to the suprageneric Principiala group within Ithonidae. The remarkable characters of this group are the peculiar configuration of forewing MP and CuA. Makarkin and Menon (2007) considered the configuration of “MP” in Principiala to be unique in Neuroptera, and it stands as a complex character that includes at least three individual character states: (1) both primary branches of MP form an equal angle with the common stem of MP; (2) the anterior branch has posteriorly directed sub- branches; and (3) the posterior branch has anteriorly directed subbranches. However, based on the clearly preserved forewing venation in Burmithone gen. nov., the “MP” (as interpreted in Makarkin and Menon, 2007) is proximally fused with “CuA” (as interpreted in Makarkin and Menon, 2007), forming the proximal part to be a typical configuration of Cu in Ithonidae (see Winterton and Makarkin, 2010: fig. 2) as well as many other neuropterans. A Fig. 3. Burmithone pennyi gen. et sp. nov., holotype female. A. Photograph of genitalia, similar feature can be also seen in Allorapisma (see Makarkin and ventral view; B. Drawing of genitalia, ventral view. S: sternum; gx: gonocoxite. Scale Archibald, 2009: fig. 2), but was interpreted to be the same as bar: 0.5 mm. that in Principiala in Makarkin and Archibald (2009). Therefore the “MP” and “CuA” in Makarkin and Menon (2007) are in fact CuA and CuP. The true MP [¼ “MA” in Makarkin and Menon (2007) and margin. MP with rather weak stem that is approximating but not Makarkin and Archibald (2009)] is actually the single long vein fused to R, almost straightly directed, without primary forking, but anteriad CuA and proximally without primary forking. It should be with only 2e3 short branches present near wing margin. Cu stem noted that the “MA” and “MP” were illustrated to be proximally very stout; CuA stout on proximal half, profusely branched anteriad fused with each other in one piece of forewing in Principiala incerta from its midpoint, all main branches of CuA bearing additional Makarkin & Menon (see Makarkin and Menon, 2007: fig. 3A). As branches, thus forming a broad, nearly triangular region; CuP with such a feature was observed only in one forewing of one species but a few short branches near wing margin. A1 long, dichotomously not found in other forewings of the other species in the Principiala branched nearly from its midpoint; A2 long, pectinately branched, group, the configuration of the forewing MP and CuA should be re- with distal two branches bearing additional deep forking; A3 examined carefully in P. incerta. distally fused with proximal-most branch of A2; A4 present, distally It should be mentioned that the assumption considering the MA bifurcated. Numerous crossveins present, but irregularly arranged. proximally fused with R and represented as the posterior-most vein Hind wings longitudinally twisted and overlapped by forewings, of the RP field (sensu Kukalova-Peck & Lawrence, 2004) is recently with venation mostly unrecognizable. Only a short and thick hu- facing challenge from some authors who consider the posterior- meral plate discernible on right hind wing. most vein of the RP field is not MA (e.g. Winterton & Makarkin, Legs stout; foreleg much slender than mid- and hind leg; midleg 2010; Winterton et al., 2017). If following the latter interpreta- moderate in length, ca. 5.0 mm; two tibial spurs present; tarsi 5- tion, the single long vein anteriad CuA could be M and consist of MA segmented, tarsomere 1 much longer than each of remaining tar- and MP. Nevertheless, there is no doubt that the “MP” in Makarkin someres; tarsomere 2 and 5 nearly equal in length, and slightly and Menon (2007) is CuA irrespective different interpretation of longer than tarsomere 3 and 4, which are also equal in length; MA. simple pretarsal claws paired, strongly curved; short arolium Following our revised interpretation on the forewing MP and present. CuA in these ithonids, this feature is actually not unique but Abdomen robust, slightly narrower than thorax. Female geni- commonly found in many groups of Myrmeleontiformia, e.g. talia: Sternum 7 nearly rectangular, length about half of width, Myrmeleontidae and , in which the branches of posterolateral corners rounded in ventral view, with dense short forewing CuA occupy a broadly subtriangular area (see Shi et al., setae; putative gonocoxites 8 paired and widely separated, 2012a: figs. 8-9). Remarkably, the two primary branches of fore- distinctly inflated posteriad, densely setose; gonocoxite 9 much wing CuA also bear posteriorly and anteriorly directed sub- longer than gonocoxite 8, widely separated with each other, branches in Myrmeleontidae and Ascalaphidae, respectively, densely setose; a tiny sclerite present between gonocoxites 9. appearing nearly identical to that in Principiala and Allorapisma. 82 X. Lu et al. / Cretaceous Research 78 (2017) 78e83

Fig. 4. Forewing venations of Ithonidae. AeD. species of the Principiala genus-group: A. Burmithone pennyi gen. et sp. nov.; B. Principiala incerta Makarkin & Menon (modified from Makarkin and Menon, 2007); C. Principiala rudgwickensis Jepson, Makarkin & Jarzembowski (modified from Jepson et al., 2009); D. Allorapisma chuorum Makarkin & Archibald (modified from the drawing of the paratype in Makarkin and Archibald, 2009); EeF. species of the Rapismatid genus-group: E. Rapisma viridipenne Barnard (modified from Barnard, 1981); F. Adamsiana curoei Penny (modified from Penny, 1996); G. Megalithone tillyardi Riek (the Ithonid genus-group; modified from Riek, 1974); H. Platystoechotes lineatus Car- penter (the Polystoechotid genus-group; modified from Carpenter, 1940). Colored veins include MA (pink), MP (orange), CuA (green), and CuP (blue). Scale bar: 2.0 mm.

Nevertheless, in Myrmeleontidae and Ascalaphidae, some distal 4.2. Systematic position of Burmithone gen. nov. subbranches of the anterior primary branch of forewing CuA are considered to originate from MP, indicated by a short oblique The intergeneric phylogeny within Ithonidae including all fossil veinlet (usually called MP2) in Shi et al. (2012a). In the three and extant groups is poorly known. Obviously, Burmithone gen. nov. genera of the Principiala group, we could find a slightly oblique is closely related to Principiala and Allorapisma, and they probably crossvein-like veinlet between MP and CuA (see Fig. 2B), but it form a monophyletic group based on the unique forewing config- seems very indistinct and may be just a crossvein. Possibly, some uration within Ithonidae (i.e., the proximal branches of RP fused anterior branches of CuA might be originated from MP based on with MA and the peculiar configuration of MP and CuA). Zheng et al. the interpretation of Shi et al. (2012a), although whether this (2016a) divided Ithonidae into three genus groups, namely the interpretation is justified is still an open question. In addition, Ithonid genus-group, the Polystoechotid genus-group, and the considering the sister group relationship between Ithonidae and Rapismatid genus-group. Important characters to distinguish these Myrmeleontiformia, such similarly specialized venation character, groups include the distal parts of ScP and RA, the arrangement of probably being apomorphic, might be indicative for their close crossveins in radial sector, the position of MP first fork, and the phylogenetic relationships, although this feature is not present in branching pattern of MP2. Zheng et al. (2016a) placed Principiala all species of these groups and may be simply convergent and Allorapisma in the Rapismatid genus-group, which is defined derivation. by the distally separated and posteriorly curved ScP and RA, the X. Lu et al. / Cretaceous Research 78 (2017) 78e83 83 numerous and irregularly arranged crossveins in the radial sector, Acknowledgements the first fork of MP proximad MA divergence, and the dichoto- mously branched MP2. As a closely related genus of Principiala and This research was supported by the National Natural Science Allorapisma, Burmithone gen. nov. should be also a member of the Foundation of China (Nos. 31322501 and 31672322). Rapismatid genus-group. However, the apparently fused forewing ScP and RA that straightly reach the anterior margin before wing References apex and the sparsely branched MP in Burmithone gen. nov. contradict the placement of this genus in the Rapismatid genus- Aspock,€ U., Aspock,€ H., 2008. Phylogenetic relevance of the genital sclerites of group. Furthermore, considering the diagnostic features of RP, Neuropterida (Insecta: Holometabola). Systematic Entomology 33, 97e127. Aspock,€ H., Aspock,€ U., Holzel,€ H., 1980. Die Neuropteren Europas: eine zusam- MA, MP and CuA in Principiala and Allorapisma, these two genera € menfassende Darstellung der Systematik, Okologie und Chorologie der Neu- are also distantly related to the other genera of the Rapismatid ropteroidea (Megaloptera, Raphidioptera, Planipennia) Europas, 2 vols. Goecke genus-group. Preferably, Burmithone gen. nov., Principiala and & Evers, Krefeld, 495; 355 pp. Allorapisma should be placed in an independent genus-group, i.e. Barnard, P.C., 1981. The Rapismatidae (Neuroptera): montane lacewings of the oriental region. Systematic Entomology 6, 121e136. the Principiala genus-group, which was originally proposed by Carpenter, F.M., 1940. A revision of the Nearctic , Berothidae, Sisy- Makarkin and Archibald (2009). Besides the venational characters, ridae, Polystoechotidae and Dilaridae (Neuroptera). Proceedings of the Amer- the absence of nygmata could be another feature to distinguish the ican Academy of Arts and Sciences 74, 193e280. Faulkner, D.K., 1990. Current knowledge of the biology of the moth-lacewing genera of the Principiala genus-group from the other genus-groups. Oliarces clara Banks (Insecta: Neuroptera: Ithonidae). In: Mansell, M.W., The female genitalia of Burmithone gen. nov. has widely separated Aspock,€ H. (Eds.), Advances in Neuropterology. Proceedings of the Third Inter- gonocoxites 8, while all other species of Ithonidae with known national Symposium on Neuropterology, South African Department of Agri- cultural Development, Pretoria, pp. 197e203. female genitalia possess unpaired gonocoxites 8. Thus, the Gallard, L., 1932. Notes on the feeding habits of the brown moth-lacewing, Ithone distinctly differed female gonocoxites 8 from that in the three fusca. Australian Naturalist 8, 168e170. genus-groups also suggest that Burmithone gen. nov. cannot be Grebennikov, V.V., 2004. Grub-like larvae of Neuroptera (Insecta): a morphological placed in any genus-group proposed by Zheng et al. (2016a) but review of the families Ithonidae and Polystoechotidae and a description of Oliarces clara. European Journal of Entomology 101, 409e417. likely represents an independent genus-group together with Prin- Jepson, J.E., Makarkin, V.N., Jarzembowski, E.A., 2009. New lacewings (Insecta: cipiala and Allorapisma. Neuroptera) from the Lower Cretaceous Wealden Supergroup of southern En- e Notably, among the genera of the Principiala genus-group, the gland. Cretaceous Research 30, 1325 1338. Kania, I., Wang, B., Szwedo, J., 2015. Dicranoptycha Osten Sacken, 1860 (Diptera, forewing ScP and RA are either fused with each other [e.g., in Limoniidae) from the earliest Cenomanian Burmese amber. Cretaceous Burmithone gen. nov. and Principiala (fusion of these two veins was Research 52, 522e530. illustrated but not confirmed in Makarkin & Menon, 2007)], or Kukalova-Peck, J., Lawrence, J.F., 2004. 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