Research 61 (2016) 129e135

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Cretaceous Research

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New Cretaceous caddisflies from (Insecta, Trichoptera)

* Wilfried Wichard a, Bo Wang b, c, a Institute of Biology, University of 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: Three new caddisflies species are described and illustrated from the mid-Cretaceous Burmese amber: Received 19 November 2015 Wormaldia cretacea sp. nov., W. resina sp. nov. () and Neureclipsis burmanica sp. nov. Received in revised form (). Palerasnitsynus ohlhoffi (Psychomyiidae) is re-described based on new fossils. 6 January 2016 Wormaldia are common in Burmese amber, and its diversity shows that this genus had very ancient Accepted in revised form 19 January 2016 origin and diversified at least during the mid-Cretaceous. N. burmanica sp. nov. is the oldest record of Available online 3 February 2016 Neureclipsis, revealing this genus originated at least in the mid-Cretaceous. So far, six extinct species representing the small Order Trichoptera have been found in Burmese amber. Keywords: © Trichoptera 2016 Elsevier Ltd. All rights reserved. Philopotamidae Polycentropodidae Burmese amber Cretaceous

1. Introduction Burminoptila bemeneha Botosaneanu, 1981 (), Wor- maldia myanmari Wichard & Poinar, 2005 (Philopotamidae), and Trichoptera (caddisflies) are an advanced order of aquatic in- Palerasnitsynus ohlhoffi Wichard et al., 2011 (Psychomyiidae). Here sects, and their adults are easily distinguished by the presence of we report two new species of Philopotamidae and one new species two pairs of wings covered with hair and held in a roof like manner of Polycentropodidae, and redescribe Palerasnitsynus ohlhoffi based over the abdomen. Their adults are nocturnal and typically found on new specimens from Burmese amber. alongside lakes and streams, and larvae are found in a variety of aquatic habitats, mainly cool swift-flowing streams. The earliest 2. Material and methods definitive caddisfly is from the Lower of Germany (Ansorge, 2002a,b). Most of Mesozoic caddisflies are compression fossils; Burmese amber has been recorded from the Shwebo, Thayet- their Mesozoic amber fossils are rare but provide more important myo, Pakokku, Magway, and Pegu districts in Myanmar (Ross et al., information about the body characters. 2010; Sun et al., 2015). However, the only commercial source is the Burmese amber (amber from northern Myanmar) contains the Hukawng Valley in Tanaing Township, Myitkyina District of Kachin most diverse biota in amber from the Cretaceous and more than State. The amber under study is from an amber mine located near 250 families of have been reported from this deposit Noije Bum Village, Tanaing Town (fig. 1 in Kania et al., 2015). These (Ross et al., 2010; Grimaldi and Ross, in press). Only three species of deposits have been investigated and dated in detail by Cruickshank caddisflies have been described from Burmese amber so far: and Ko (2003) and Shi et al. (2012). We tentatively follow the age determination as early Cenomanian (98.8 ± 0.6 million years) given by UePb dating of zircons from the volcanoclastic matrix of the * Corresponding author. State Key Laboratory of Palaeobiology and Stratigraphy, amber (Shi et al., 2012). However, the Zircon date only gives the age Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, of the bed and a minimum age for the amber. Much of the resin is Nanjing 210008, China. E-mail addresses: [email protected] (W. Wichard), [email protected] rolled and bored by pholadid bivalves which demonstrate the resin (B. Wang). was hard before it was buried and therefore older (Ross et al., 2010). http://dx.doi.org/10.1016/j.cretres.2016.01.004 0195-6671/© 2016 Elsevier Ltd. All rights reserved. 130 W. Wichard, B. Wang / Cretaceous Research 61 (2016) 129e135

Thus, the geologic age of Burmese amber should be slightly older or mostly 2/4/4. Forewing with Rl unbranched; five apical forks than the Zircon date (Ross, 2015). usually present, but fork IV, or forks I and IV, or forks I, III, and IV Photographs were taken using a Leica stereomicroscope M 420, sometimes absent; intercostal crossvein usually in mid costal re- apozoom and Zerene Stacker software, and in most instances, gion; discoidal and median cells present. Hindwing with four apical incident and transmitted light were used simultaneously. The fig- forks usually present, but fork I or forks I and III sometimes absent; ures were prepared with CorelDraw X4 and Adobe Photoshop CS3. discoidal cell present, median cell absent. Abbreviations: I, apical fork I; II, apical fork II; III, apical fork III; IV, Wormaldia McLachlan, 1865 apical fork IV; V, apical fork V; S-IX, sternum segment IX; T-IX, Type species: Hydropsyche occipitalis Pictet, 1834 (subsequent tergum segment IX; T-X, tergum segment X; Dc, discoidal cell; Mc, designation by Ross, 1949) median cell; Tc, thyridium cell; IA, inferior appendage; IA-C, infe- rior appendage (coxopodite); IA-H, inferior appendage (harpago); Diagnosis (modified from Munoz-Quesada~ and Holzenthal, 2015). SA, superior and preanal appendage (cercus); PA, phallic apparatus. Adult forewings 3e8 mm; legs with tibial spur formula 2/4/4. Forewings covered with small, fine setae; apical forks I usually and 3. Systematic palaeontology II always present, R2 and R3 veins sometimes fused as R2þ3;M3- branched with apical fork IV absent or 4-branched with forks III Order Trichoptera Kirby, 1815 and IV present. Hindwings shorter than forewings; all radial veins Suborder Martynov, 1924 (R1,R2,R3,R4, and R5) usually present in forewings (e.g. in W. myanmari from Burmese amber), except R4 and R5 veins sometimes fused as R4þ5 (e.g. in W. cratacea sp. nov. and W. resina sp. nov. from 3.1. Family Philopotamidae Stephens, 1829 Burmese amber). Median vein 3-branched (M1,M2, and M3þ4, apical fork IV absent). In male genitalia sterna VII and VIII with or Diagnosis. Ocelli present. Maxillary palps 5-segmented; segment 2 without processes posteromesally. with an apical brush of setae, segment 5 longest and annulate. Wormaldia cretacea sp. nov. (Fig. 1) Labial palps shorter than maxillary palps, 3-segmented, segment 3 longest and annulate. Mesoscutum never with a pair of warts, Etymology. The species cretacea is named after the Cretaceous sometimes with scattered setae. Tibial spurs 1/4/4 (e.g. Chimarra) period of Burmese amber.

Fig. 1. Wormaldia cretacea sp. nov., holotype NIGP156999. A, Photograph in ventral view. B, Photograph of male genitalia in ventral view. C, Drawings of fore- and hindwings. D, Drawing of male genitalia in ventral view. Scale bars ¼ 1 mm in A and C; ¼ 0.7 mm in B and D. W. Wichard, B. Wang / Cretaceous Research 61 (2016) 129e135 131

Holotype. Male, NIGP156999, deposited in the Nanjing Institute of Male genitalia: Sterna VII and VIII without processes poster- Geology and Palaeontology (NIGP), Chinese Academy of Sciences. omesally. Segment IX, viewed ventrally, slightly sinuous posteri- Diagnosis. Wormaldia cretacea is distinguishable from the other orly. Superior appendages (cerci) light brown, elongate and Wormaldia species and from two species in Burmese amber by the digitate, half as long as the inferior appendages. Inferior append- following attributes: forewing fork IV absent, hindwing forks I and ages 2-segmented, consisting of basal (coxopodite) and apical IV absent, sterna VII and VIII without processes posteromesally, (harpago) segments. In ventral view, basal segments separated by male genitalia with conspicuous, long inferior appendages. deep V-shaped emargination. Each basal segment stout, widest Material examined. The adult is embedded in a clear amber piece. In anteromedially, clearly incurved, with outer margin convexly dorsal view, forewings are clearly visible and hindwings are partly curved. Apical segment slender, tubular and elongate, inner margin covered by forewings. In ventral view, hindwings are basally very weakly concave, slightly widened and rounded posteriorly. In covered by the body. Antennae, maxillary palps, labial palps and ventral view, apical part of lobate segment X arranged deep be- legs are clearly visible. Male genitalia are dorsally covered by the tween inferior appendages; apical margin of lobate segment X with wings. medial tip (Fig. 1B and D). Description. Head, thorax, legs and abdomen brown, the antennae Wormaldia resina and palps light brown, covered all over with fine small and light sp. nov. (Fig. 2) brown setae. Head with prominent eyes laterally, three ocelli Etymology. The species is named resina, for resin, the origin of clearly developed. Antennae about 2/3rds as long as forewings, Burmese amber. fl consisting of 28 agellomeres plus stronger scapus and pedicellus. Holotype. Male, NIGP157000, deposited in the Nanjing Institute of Maxillary palps 5-segmented, each terminal segment almost as Geology and Palaeontology (NIGP), Chinese Academy of Sciences. long as 1st to 4th segments. Forewing 3 mm long. Forewing Diagnosis. Wormaldia resina is different from the other Wormaldia translucent with fine, small, light brown setae, with forks I, II, III, species and from the two species in Burmese amber by the and V present. Forks I and II sessile and fork III petiolate. M 3- following attributes: forewing fork IV absent; hindwing fork I branched. In left forewing, fork III extraordinarily 3-branched present and fork IV absent; sterna VII and VIII without processes (with an additional inner branch, Fig. 1C); in right forewing, fork posteromesally; male genitalia with long tapered superior ap- III normally 2-branched. Discoidal, median and thyridial cells pendages, median with strong setae; harpagoes each bearing an closed, indicated by a line of fine setae. Intercostal crossvein usually apical setal patch. in mid costal region not visible. Hindwing translucent with fine, small, light brown setae. Fork I absent; forks II, III, and V present Material examined. The adult is embedded in Burmese amber, and (Fig. 1A and C). surrounded by scores of small-sized air bubbles which hinder the

Fig. 2. Wormaldia resina sp. nov., holotype NIGP157000, male. A, Photograph in ventral view. B, Photograph of male genitalia in lateroventral view. C, Drawings of fore- and hindwings. D, Drawing of male genitalia in lateroventral view. Scale bars ¼ 1 mm in A and C; ¼ 0.7 mm in B and D. 132 W. Wichard, B. Wang / Cretaceous Research 61 (2016) 129e135 analysis of the species. In dorsal view, the forewings are tolerably terminal segment annulated and longer than segments 3 þ 4. Labial visible and hindwings are partly covered by forewings; in ventral palps 3-segmented, terminal segment annulated, nearly as long as view, the hindwings are basally covered by the body. Antennae, segments 1 þ 2. Tibial spurs 2/4/4. Forewing length 2 mm. Fore- maxillary palps, labial palps and legs are visible. Male genitalia are wings apically rounded. Apical fork I absent, apical forks II, IV and V dorsally covered by the wings. Body, wings and legs are light brown present; fork II sessile. Discoidal cell closed. Crossveins r and r-m and translucent; all covered with fine, small and light brown setae. indistinct, crossvein m-cu apparently present defining closed The head is dorsally destroyed; therefore the three ocelli cannot be thyridial cell. verified, except very probably the front ocelli. Wing venation and the male genital structures indicate that the species belongs to the Male genitalia (Fig. 3D): In ventral view, all inferior appendages genus Wormaldia within Philopotamidae. (gonopods) 2-segmented, consisting of a coxopodite (basal Description. Head dorsally destroyed, front ocelli probably present. segment) and a harpago (terminal segment); coxopodite stocky, Right antennae about 2/3rd as long as forewings, consisting of square with rounded edges; harpag about 4 times as long as cox- probably 26 flagellomeres plus larger scapus and pedicellus; left opodite, moderately curved mesad with concave inner margin, antennae truncated. Maxillary palps 5-segmented, segment 2 bearing fine setae and at the ampullate apex, stout black teeth. rounded, with an apical brush of setae; terminal segment about as Superior appendages, digitate, about half as long as gonopods, very long as 3rd þ 4th segments. Length of male forewing 3.5 mm. slightly curved mediad, with rounded apex. Intermediate append- Forewing with forks I and II sessile and fork III petiolate. Discoidal ages each slender, small black spines, almost as long as gonopods, and median cells closed, indicated by very weak crossveins. Inter- distally curved outwards. Phallus elongate, terminating in bifid costal crossvein not visible. Hindwing with forks I and II sessile, fork aedeagus with two expanded lobes. III petiolate, fork V probably present (Fig. 2A and C). Remarks. The male genitalia of the new specimens consist of more Male genitalia (Fig. 2B and D): Sterna VII and VIII without processes detailed structures than the described holotype genitalia in posteromesally. In lateral view, segment IX broad, nearly sub- Wichard et al., 2011. Thus we revise the male genitalia. triangular in appearance. Superior appendages (cerci) light brown, elongate, distally tapered, about as long as the inferior appendages; 3.3. Family Polycentropodidae Ulmer, 1906 along the inside armed with strong spines and two long spines basally. Inferior appendages 2-segmented, consisting of basal Diagnosis. Adults without ocelli. Maxillary palps 5-segmented, first (coxopodite) and apical (harpago) segments. In ventral view, basal two segments short, terminal segment long, annulated, flexible. segments separated by deep V-shaped emargination. Each basal Antennae almost as long as forewings. Mesoscutum with a pair of segment stout, almost rectangular and little shorter than the har- setose warts. In forewing, fork I sessile (Neureclipsis and Poly- pago. Terminal segment rectangular and elongate, with an apical centropus clusters) or absent (Cyrnus and Cyrnodes clusters, sensu setal patch of short, black, peg-shaped setae, looking like a brush Olah and Johanson, 2010); forks II, III, IV, V or II, IV, V present. In border. Lobate segment X in middle of genitalia arranged between hindwing, forks II and V always present; forks I and III present or pairs of superior and inferior appendages; viewed laterally, lobe absent. Tibial spurs 3/4/4. seems triangular shaped. Remarks. The phylogeny of the family Polycentropodidae was ana- lysed by Chamorro and Holzenthal (2011) and Johanson et al. 3.2. Family Psychomyiidae Walker, 1852 (2012), a generic review by Olah and Johanson (2010). Neureclipsis McLachlan, 1864 Diagnosis. Adults without ocelli. Maxillary palps usually 5- fi segmented, segment 2 usually longer than segment 1 and almost Diagnosis (modi ed from Olah and Johanson, 2010). In forewing, as long as segment 3, terminal segment annulated and flexible. complete wing venation with apical forks I, II, III, IV and V present. Mesoscutum with two rounded setal warts; mesoscutellum with a In hindwing, apical forks I, II, III, and V present. In both wings separate pair of warts or fused setal warts. Tibial spurs 2/4/4. In discoidal and median cells closed. Distinguished from all other polycentropodid genera by the presence of fork III in hindwing. forewings, veins R2 and R3 fused, fork I absent. Hindwing with reduced venation. Neureclipsis burmanica sp. nov. (Fig. 4) Palerasnitsynus Wichard, Ross & Ross, 2011 Etymology. The new species bears the name burmanica, named after Type species: Palerasnitsynus ohlhoffi Wichard, Ross & Ross, 2011 Burma where the species is found. Diagnosis (modified from Wichard et al., 2011). Ocelli absent. Holotype. Male embedded in Cretaceous Burmese amber, SMNS Bu- Maxillary palps 5-segmented, terminal segment annulated. Tibial 173, deposited in Staatliches Museum für Naturkunde Stuttgart, spurs 2/4/4. In forewing, apical forks II, IV and V present. Distin- Germany (ex coll. Wichard/Wunderlich). guished from all other psychomyiid genera by the absence of fork III Diagnosis. Distinguished from the extant and extinct species in in forewing. Baltic amber by the male genital structure, in particular by the inferior appendages (gonopods). Redescription Material examined. The adult is embedded in clear Burmese amber Palerasnitsynus ohlhoffi Wichard, Ross & Ross, 2011 (Fig. 3) and the inclusion is completely visible. However, each base of the Material examined. NIGP157001 and NIGP157002, deposited in the fore- and hindwings overlies another and is covered by the body Nanjing Institute of Geology and Palaeontology (NIGP), Chinese ventrally. Antennae, maxillary palps, labial palps and legs are Academy of Sciences. Two adults are embedded in clear Burmese clearly visible. Male genitalia are dorsally and laterally covered by amber. Fore- and hindwing, as well as antennae, maxillary palps, wings. labial palps and legs are clearly visible. Description. Head, thorax, abdomen and antennae, palps and legs Description. Ocelli absent. Antennae almost as long as forewings, light brown, covered all over with fine, light brown setae; head consisting of 20 flagellomeres plus scapes and pedicelli. Maxillary warts, thorax (median pronotal setal warts) and tegulae with palps 5-segmented, segment 1 short, segment 2 slightly longer, brushes of long setae. Mesoscutum bearing a pair of ovoid setose W. Wichard, B. Wang / Cretaceous Research 61 (2016) 129e135 133

Fig. 3. Palerasnitsynus ohlhoffi Wichard, Ross & Ross, 2011. A, NIGP157001, photograph in ventral view. B, NIGP157002, photograph in dorsal view. C, Photograph of forewing (NIGP157001). D, Drawing of male genitalia in dorsal view (NIGP157001). Scale bars ¼ 1 mm in A, B, and C; ¼ 0.4 mm in D.

warts, scutellum with a single, rounded, mesal wart. Antennae well as the extant genera Wormaldia and Neureclipsis, whereas the slightly longer than forewings and consisting of 40 flagellomeres hydroptilid Burmenoptila and the psychomyid Palerasnitsynus are plus stronger scapus and pedicellus. Maxillary palps 5-segmented, extinct. first two segments short, 2nd segment apically with few long setae, Philoptamidae predominates with the genus Wormaldia among terminal segment annulated, flexible and slightly longer than all caddisflies found in Burmese amber. Wormaldia is the largest 3rd þ 4th segments. genus in the family Philopotamidae and is diverse and widely distributed, with about 175 extant species in all biogeographic re- Forewing length 3.5 mm. Forewing translucent with fine, small, gions except the Australasian (Munoz-Quesada~ and Holzenthal, light brown setae, with forks I, II, III, IV and V present, fork I peti- 2015). Wormaldia larvae prefer running waters and feed on fine olate and fork II sessile; nygma in fork II. Discoidal and median cells particulate organic matter from the meshes with its specialized closed; crossveins r, s, r-m, and m present. Hindwing translucent mouthparts (Munoz-Quesada~ and Holzenthal, 2015). Up to now, with fine, small, light brown setae, with forks I, II, III, V; fork I three species of Wormaldia have been reported from Burmese petiolate and fork II sessile; nygma in fork II. Discoidal cell closed; amber, and this high diversity shows that this genus had very crossveins r, s, r-m present, m not visible (Fig. 4C). ancient origin, and diversified at least during the mid-Cretaceous. Male genitalia (Fig. 4D): In dorsolateral view, tergum IX and tergum The extant family Polycentropodidae, with over 600 species, is X forming apparently a prominent functional group (probably: worldwide distributed. They are the most common caddisflies tergum IX elongated, broad at base and in the middle slightly (more than 80% of all caddisfly specimens) in Eocene Baltic amber concave bilaterally and at apex divided into two lobes, overlying (Wichard, 2013). Neureclipsis is a small genus of Polycentropodidae. tergum X; tergum X divided apically into two semi-membranous Extant Neureclipsis species are distributed in the northern hemi- lobes). Preanal appendage 2/3rds as long as tergum IX, slightly sphere, and their larval mode of life is quite similar to that of curved mediad. Preanal appendage (cercus) subtriangular in lateral Wormaldia larvae. Six fossil species have been reported, and all view, forming ventrad a semi-membranous shield, apically side by from Baltic amber. Ulmer (1912) listed four extinct species of side with rounded lobes. Inferior appendage (gonopod) 1- Neureclipsis and two species of Archaeoneureclipsis synonymized segmented, elongate, slender; its base slightly broad, then tapered, with Neureclipsis (Olah & Johanson, 2010). Neureclipsis burmanica almost rectangular near apex, with a small ampullate apex. sp. nov. is the first Mesozoic and accordingly the oldest fossil record of Neureclipsis, providing direct evidence that Neureclipsis and its 4. Discussions sister group have minimum ages of about 99 million years. Caddisflies are thought to be very rare in Burmese amber The caddisflies in mid-Cretaceous Burmese amber are repre- (Wichard et al., 2011). Our recent investigation, however, reveals a sented so far by 4 extant families: Hydroptilidae (see Botosaneanu, potential high diversity of this lineage from Burmese amber. 1981), Philopotamidae, Polycentropodidae and Psychomyiidae, as Mesozoic caddisflies are mainly described based on compression 134 W. Wichard, B. Wang / Cretaceous Research 61 (2016) 129e135

Fig. 4. Neureclipsis burmanica sp. nov., SMNS Bu-173, male in dorsal view. A, Photograph in dorsal view. B, Photograph in ventral view. C, Photograph of fore- and hindwings. D, Drawings of inferior appendages in lateroventral view (above) and male genitalia in laterodorsal view (below). Scale bars ¼ 1 mm in A, B, and C; ¼ 0.35 mm in D.

fossils which commonly preserve only wings characters. The new anonymous reviewer for their helpful input. This research was caddisflies of Burmese amber show exquisite details of body struc- supported by the National Basic Research Program of China ture, especially genitalia, thus providing insights into the evolution (2012CB821900), National Natural Science Foundation of China of this lineage. In addition, these aquatic provide robust (41572010, J1210006), and Youth Innovation Promotion Association evidence for the humid environment of Burmese amber forest. of CAS (No. 2011224).

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