Research 69 (2017) 198e203

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Cretaceous Research

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Short communication A remarkable caddisfly with bipectinate antennae in Cretaceous (Insecta, Trichoptera)

* Wilfried Wichard a, Bo Wang b, c, a Institute of Biology, University of Koeln, Gronewaldstr. 2, D 50931 Koeln, Germany b State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China c Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Science, Beijing 100101, China article info abstract

Article history: A new caddisfly (Trichoptera), Palaeopsilotreta xiai gen. et sp. nov. is described based on three well- Received 8 August 2016 preserved male specimens from mid-Cretaceous Burmese amber. It is assigned to the extant family Received in revised form . Palaeopsilotreta is similar to the extant genus but differs from the latter by 19 September 2016 partially bipectinate antennae which are unknown among living Trichoptera. Our are not only the Accepted in revised form 28 September only Mesozoic Odontoceridae, but also hitherto the earliest record of this family. 2016 © Available online 29 September 2016 2016 Elsevier Ltd. All rights reserved.

Keywords: caddisfly Palaeopsilotreta xiai Psilotreta Odontoceridae

1. Introduction insights into the evolution of this lineage. In this paper, we describe a new extinct genus and species placed in the family Odontocer- Burmese amber (from northern Myanmar) contains the most idae: Palaeopsilotreta xiai gen. et sp. nov., based on three well- diverse biota in amber from the mid-Cretaceous and more than 250 preserved male specimens. It exhibits partially bipectinate families of have been reported from this deposit (Ross antennae which are unknown among living Trichoptera. et al., 2010; Grimaldi and Ross, in press). The caddisflies in Bur- mese amber are represented to date by four extant families: 2. Material and methods , , and Psycho- myiidae (Botosaneanu, 1981; Wichard and Poinar, 2005; Wichard The specimens are from an amber mine located near Noije Bum et al., 2011; Wichard and Wang, 2016). Our recent investigation Village, Tanaing Town, Myanmar (Kania et al., 2015). The age given reveals a potentially high diversity of this lineage in Burmese by UePb dating of zircons from the volcanoclastic matrix of the amber. Mesozoic caddisflies are mainly described based on amber is early Cenomanian (98.8 ± 0.6 million years) (Shi et al., compression fossils which commonly preserve often only wing 2012), but the geological age of Burmese amber should be slightly characters. Some of compression fossils have discernible body parts older than the zircon date (Ross, 2015). and enable detailed descriptions (e.g., Gao et al., 2013; Zhang et al., The fossil specimens are embedded in small amber blocks cut 2016). The new caddisflies from Burmese amber show exquisite out from larger Burmese amber pieces. The adult are nearly details of body structure, especially genitalia, thus providing completely preserved and clearly visible in ventral and dorsal view. The male genitalia are flattened. The hindwings are partly covered by the somewhat distorted forewings. Some legs or parts of legs are * Corresponding author. State Key Laboratory of Palaeobiology and Stratigraphy, missing. The head, thorax and abdomen show signs of decompo- Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, sition and maceration. Antennae are incomplete. Nanjing 210008, China. E-mail addresses: [email protected] (W. Wichard), [email protected] Photos were taken using a Leica stereomicroscope M 420 Apo- (B. Wang). zoom in combination with a Canon EOS 600D, EOS utility software http://dx.doi.org/10.1016/j.cretres.2016.09.012 0195-6671/© 2016 Elsevier Ltd. All rights reserved. W. Wichard, B. Wang / Cretaceous Research 69 (2017) 198e203 199 and the Zerene Stacker software or were taken by the digital mi- Description. Head, thorax and abdomen brown covered partly with croscope Keyence VHX-900F. All images and figures were prepared small and light brown setae; laterally with prominent eyes. Antennae with CorelDraw X4 and Adobe Photoshop CS4. The wing venation partially bipectinate, about as long as forewings, consisting of a strong terminology in general follows Holzenthal et al. (2007):Ie apical scapus, approximately twice the length of the short pedicellus, then fork I; II e apical fork II; V e apical fork V; R e radius, M e media, follow 15e20 sub-cylindrical flagellomeres: the basal 10 flag- Cu1 e cubitus anterior, Cu2 e cubitus posterior, A e analis, DC e ellomeres each bear distally bipectinate rami, the terminal 6e10 discoidal cell. All taxonomic acts established in the present work flagellomeres are simple, rami are lacking. Maxillary palps 5- have been registered in ZooBank (see below), together with the segmented, sparsely covered with setae; terminal segment not flex- electronic publication LSID: urn:lsid:zoobank.org:pub:668567C5- ible or annulated, apically rounded. A47B-4866-ADA6-59D6C5B752AC. Head dorsal ocelli absent, with a pair of antennal (anterior) setal 3. Systematic palaeontology warts, small and ovoid and a pair of posterior setal warts, large and subtriangular. Pronotum transversally with two setal warts, small Order Trichoptera Kirby, 1815 and rounded. Mesoscutum bearing a pair of small ovoid scutal Suborder Martynov, 1924 warts, the scutellum with a single dome-shaped, large, ovoid wart. Family Odontoceridae Wallengren, 1891 Forewings light brown, 2.3e2.6 mm long, apically rounded. Vein R1 basally thickened, strong, running to wing margin. Forks I and II Palaeopsilotreta gen. nov. present and sessile; discoidal cell closed by crossveins. DC about as (urn:lsid:zoobank.org:act:211F3FC8-0A53-4FBF-BD2F- long as fork I, fork II a little longer. Media simple, running to apical 0777F996CA15) margin, without branching into M1þ2 and M3þ4. Thyridial cell Diagnosis. Male antennae partially bipectinate, about as long as absent. Cu1 basally thickened, running straight, apically bifurcated forewings. Maxillary palps 5-segmented, first segment without into fork V; Cu2 simple, unforked. Anal vein complete and basally apicomesal nodule; terminal segment not flexible or annulated. thickened. Mesoscutum with a pair of ovoid setal warts, mesoscutellum Hindwings light brown, translucent, relatively narrow, smaller covered by a single wart, large, dome-like and ovoid. In fore- and than forewings, about 2 mm long, so far as visible, venation prob- hindwings forks I, II and V present; media simple, unbranched ably similar to forewing venation. Radius R1 vein conspicuously running to apical margin; discoidal cells closed, median and thyr- strong and slightly thickened. Forks I (R2 þ R3), II (R4 þ R5) and V idial cells lacking. Tibial spurs: 1/4/4. (Cu1a þ Cu1b) visible, as well as the apical junctions of media (M) Remarks. The new extinct genus can be assigned to the family and Cu2 at apical wing margin. Odontoceridae by the following combination of features: maxillary Legs with apical tibial spurs, mid- and hindlegs with additional palps 5-segmented, terminal segment not flexible or unnulated; pre-apical tibial spurs, however, in forelegs pre-apical tibial spurs mesoscutellum covered by large single wart, dome-like and ovoid, absent: Tibial spurs: 1/4/4. sparsely covered with setae; at mesoscutum a pair of ovoid setose Male genitalia are visible only in ventral view, so the description warts present. Tibial spurs: 0e2/0e4/2e4. In the forewings dis- is preliminary and incomplete (Fig. 3). Inferior appendages (gon- coidal cell closed, median cell absent; the thyridial cell lacking in ocoxites) 2-segmented. Coxopodites cone-shaped, bearing mesad the males of the genera and Psilotreta; the M stem missing scattered long setae; almost 2/3rd in length of the gonocoxites; the in Pseudogoera and simple and unbranched in Psilotreta. In order to harpagoes originating probably from the apex of the coxopodites, distinguish odontocerids from other families, a combination of elongate, pencil-shaped, about 1/3rd in length of the gonocoxites, different morphological characters is essential (Olah and Johanson, each harpago ending in a small crown with few black spines. 2010, 2011). Preanal appendages membranous, dorsum slightly convex, inner side slightly hollowed, at apex triangular and pointed; the preanal The genus Palaeopsilotreta is closely related to the extant genus appendages longer than the gonocoxites. Segment X and the gen- Psilotreta by the simple and unbranched media in forewings, but ital apparatus unclear, therefore undefined. The positions of fore- differs from the latter in having the partially bipectinate antennae and hindwings hinder a clear description of the flattened male and only one apical spurs of the fore legs, instead of the two apical genitalia, visible ventrally. spurs in Psilotreta. In forewing the vein combination of forks I and II, discoidal cell closed, M simple and unbranched and the median and 4. Discussion thyridial cells absent, is only known from the odontocerid genera Psilotreta and the extinct new Palaeopsilotreta. The family Odontoceridae is relatively small and is known from all zoogeographical regions. It contains about 150 extant species in Etymology. The generic name refers to the extant genus Psilotreta,to 15 genera (Holzenthal et al., 2010) and 7 fossil species: Phenacop- which the new extinct genus is closely related. syche vexans Cockerell, 1909 from the of Florissant, USA; Marilia altrocki Wichard, 1986, Marilia succinea Wichard, 2013, Palaeopsilotreta xiai sp. nov. Marilia ophthalmica Ulmer, 1912, Electrocerum pedestre Ulmer, 1912 Figs. 1e4 and Electropsilotes rara Ulmer, 1912 from Eocene Baltic amber (urn:lsid:zoobank.org:act:400ACD13-B750-4FB6-9257- (Wichard, 2013). Our fossils are not only the only Mesozoic Odon- 32ADD57C67EF) toceridae, but also hitherto the earliest record of this family. Holotype. NIGP164781, deposited in the Nanjing Institute of Geol- Pectinate antennae have a single row of comb-like rami along ogy and Palaeontology, Chinese Academy of Sciences. the flagellum, one ramus for each flagellomere, forming all together Paratype. Two specimens, BA16001 and BA16002, deposited in the a comb-like shape. Bipectinate antennae have two rows of comb- Lingpoge Amber Museum (Shanghai). like rami along the flagella and a pair of rami at each flag- Etymology. The specific name is in honour of Mr. Xia Fangyuan, ellomere; they look often like flea combs. Pectinate and bipectinate Director of Lingpoge Amber Museum, for his contribution to the antennae are well known in some orders, e.g. in Coleoptera study of Burmese amber. and (Beutel et al., 2014; Gao et al., 2016). In some Diagnosis. As for the genus. and beetles, males possess such distinct antennae for 200 W. Wichard, B. Wang / Cretaceous Research 69 (2017) 198e203

Fig. 1. Palaeopsilotreta xiai sp. nov., holotype NIGP164781, male. A, Photograph in ventral view. B, Bipectinate antennae with flagellomeres and pairs of rami. W. Wichard, B. Wang / Cretaceous Research 69 (2017) 198e203 201

Fig. 2. Palaeopsilotreta xiai sp. nov., paratype BA16001, male. A, Photograph in dorsal view. B, Head in ventral view, with 5-segmented maxillary palps and bipectinate antennae. detecting the female pheromones at a distance. Since females do only from a few extant caddisfly species: the Australian Ram- not need to detect the males, they have simpler antennae. The iheithrus kocinus Neboiss, 1974 and Ramiheithrus virgatus Neboiss, Trichoptera, males and females usually have filiform antennae, 1974 (Philorheitridae) and Phraepsyche pectinata Olah & Johanson, which are often as long as the forewings and most often covered 2010 (Odontoceridae) found in Vietnam; their females are un- with small setae. Pectinate antennae are generally rare and known known. In the first half of the antennae of Ramiheithrus kocinus, the 202 W. Wichard, B. Wang / Cretaceous Research 69 (2017) 198e203

Fig. 3. Palaeopsilotreta xiai sp. nov., male genitalia and hindwing. A, paratype, BA16001. B, Holotype NIGP164781. W. Wichard, B. Wang / Cretaceous Research 69 (2017) 198e203 203

Fig. 4. Palaeopsilotreta xiai sp. nov., holotype NIGP164781, forewing drawing. basal 15e16 flagellomeres bear rami which are longer than the Gao, T.P., Shih, C.K., Labandeira, C.C., Santiago-Blay, J.A., Yao, Y.Z., Ren, D., 2016. fi flagellomeres, forming the comb-like ramified antennae; in the of rami ed antennae in insect lineages from the Early Cretaceous of Northeastern China. Proceedings of the Royal Society B, second half, the antennae are filiform. In Phraepsyche pectinata the 20161448. first 13 segments of the antennae bear long branch-like processes, Grimaldi, D.A., Ross, A.J., 2016. Extraordinary lagerstatten€ in amber, with particular reference to the Cretaceous of Burma. In: Fraser, N.C., Sues, H.D. (Eds.), giving pectinate antennae. On the scapus and pedicellus, the rami € fi Terrestrial Conservation Lagerstatten: Windows into the Evolution of Life on are always lacking. These extant structures of comb-like rami ed Land. Dunedin Academic Press Ltd, Edinburgh (in press). antennae are very similar to the bipectinate antennae of the extinct Holzenthal, R.W., Blahnik, R.J., Prather, A.L., Kjer, K.M., 2007. Order Trichoptera Palaeopsilotreta xiai, consisting of the first 10 flagellomeres bearing Kirby, 1813 (Insecta), Caddisflies. Zootaxa 1668, 639e698. e fl Holzenthal, R.W., Morse, J.C., Kjer, K.M., 2010. Order Trichoptera Kirby, 1813. Zoo- rami and of the last 6 10 agellomeres without rami. taxa 3148, 209e211. Kania, I., Wang, B., Szwedo, J., 2015. Dicranoptycha Osten Sacken, 1860 (Diptera, Limoniidae) from the earliest Upper Cretaceous Burmese amber. Cretaceous 5. Concluding remarks Research 52, 522e530. Kirby, W., 1815. , a new order of insects proposed, and the characters of the order, with those of its genera. Transactions of the Linnean Society of A new species of Trichoptera, Palaeopsilotreta xiai gen. et sp. London Zoology 11, 86e122. nov., is described from Cretaceous Burmese amber. It is the only Martynov, A.V., 1924. Rucheiniki (caddisflies). Prakticheskaya Entomologiya 5, e Mesozoic Odontoceridae, and represents the earliest record of the 1 384 (in Russian). Neboiss, A., 1974. A new caddisfly-genus from Victoria and Tasmanis (Philorhei- family. P. xiai exhibits bipectinate antennae which are unknown tridae: Trichoptera). Victorian Naturalist 91, 322e325. among living Trichoptera. So far, the caddisflies in Burmese amber Olah, J., Johanson, K.A., 2010. Description of 33 new species of , are represented by five extant families: Hydroptilidae, Phil- , Odontoceridae and Philorheithridae (Trichoptera), with detailed presentation of their cephalic setal warts and grooves. Zootaxa 2457, 1e128. opotamidae, Polycentropodidae, , and Odontocer- Olah, J., Johanson, K.A., 2011. Description of a new endemic genus of Trichoptera idae. Our find augments the diversity and morphological disparity from Madagascar (Odontoceridae). 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