PROC. ENTOMOL. SOC. WASH. 109(3), 2007, pp. 649–655

LONGIOCULUS BURMENSIS, N. GEN., N. SP. (: ELCANIDAE) IN

GEORGE POINAR,JR., ANDREJ V. GOROCHOV, AND RON BUCKLEY

(GP) Department of Zoology, Oregon State University, Corvallis, OR 97331, U.S.A. (e-mail: [email protected]); (AG) Zoological Institute, Russian Academy of Sciences, S. Petersburg 199034, Russia (e-mail: [email protected]); (RB) 9635 Sumpter Road, Florence, KY 41042-8355, U.S.A. (e-mail: [email protected])

Abstract.—An adult orthopteran in Early Burmese amber is described as a new genus and species, Longioculus burmensis Poinar, Gorochov, and Buckley.On the basis of its tegminal venation, three-segmented tarsi, and large spines on the hind tibia, it is placed in the extinct family Elcanidae. This fossil differs from previously described members of the family by its relatively small and slender body, protruding eyes, enlarged scapes and antennal cavities, short pronotum, and unique venational and leg armament characters. Key Words: Orthoptera, Elcanidae, Longioculus, Longioculus burmensis, Burmese amber, Early Cretaceous

Adult orthopterans in amber are rare, unpublished research 2002). Palyno- since these are usually strong morphs from the amber beds where the enough to free themselves from the resin. fossil originated have been assigned to the A well-preserved Burmese amber orthop- Upper of the Lower Cretaceous teroid previously depicted in Poinar et al. (,100 mya) (Cruickshank and Ko 2003); (2005) is the topic of this paper. The however, since the amber is secondarily specimen has three-segmented tarsi, deposited, the age could be older. a short pronotum, enlarged hind femora, The piece of amber containing the and venational characters that align it specimen is rectangular in shape, mea- with members of the family Elcanidae suring 13 mm in length, 10 mm in width (Sharov 1968; Carpenter 1992; Gorochov and 6 mm in depth. The fossil is well 1995; Gorochov and Rasnitsyn 2002). preserved and complete except for the tips of the antennae and wings, one side MATERIALS AND METHODS of the apex of the abdomen (including Amber from Burma occurs in lignitic one cercus and the genital apparatus), seams in sandstone-limestone deposits in the left metatibia and metatarsus and the the Hukawng Valley, southwest of Main- apical portion of the right metatibia. gkhwan in the state of Kachin (26u209N, Other fossils in the same amber piece are 96u369E). Nuclear magnetic resonance three small adult beetles, one mite, one (NMR) spectra of amber samples taken adult nematoceran and several branched from the same locality as the fossils plant hairs. Observations and photo- indicated an araucarian (possibly Agathis) graphs were made with a Nikon SMZ- source of the amber (Lambert and Wu, 10 stereoscopic microscope and Nikon

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Optiphot optical microscope (with mag- Cretaceous Cratoelcana Martins- Neto nifications up to 6503). (Martins-Neto 1991), by the apices of the branches of tegminal ‘‘C’’ positioned Family Elcanidae Handlirsch, 1906 within a short distance of each other and Longioculis Poinar, Gorochov, and the most proximal crossvein in the area Buckley, new genus between R+ RS (possibly the basal part of RS) situated near the bases of the third and Description.—Body relatively small fourth branches of RA. From Cratoelcana, and slender; scapes and antennal cavities with species over 20 mm in length, the new greatly enlarged, occupying entire area genus differs in being much smaller (under between compound eyes; compound eyes 8mminlength). large, protruding from head; ocelli lack- Type species.—Longioculis burmensis, ing; pronotum short, hind tibia with n. sp. dorsal row of denticles and row of much larger spines in middle part (and possibly Longioculis burmensis Poinar, Gorochov, in distal part); ventral surface of hind and Buckley, new species basitarsus with row of rather long spines; tegmina with both branches of false (Figs. 1–12) costa (‘‘C’’) meeting within short dis- Description.—Characters as listed un- tance of each other on anterior wing der generic description. Alate male; body margin; tegminal area between ‘‘C’’ and brown with transparent wing; length Sc narrow; tegminal crossveins in visible body, 7.14 mm; length pronotum, 2.4 area between R+ RA reduced (most mm; length hind femur, 6.4 mm. proximal crossvein in the area, possibly Head: Narrow; without rostral tuber- basal part of RS, situated between bases cles between antennal cavities; ocelli not of third and fourth branches of RA). observed; antennal cavities in contact with Etymology.—‘‘Longioculis’’ is from each other, with edges prominent; anten- the Latin ‘‘oculus’’ for eye and ‘‘longus’’ nal scapes enlarged; antennal flagellum for long, in reference to the protruding longer than body (both antennae cut off at eyes. The gender is masculine. edge of amber piece); filiform; antenno- Diagnosis.—On the basis of the nar- meres short, equal or subequal in size; row, large spines on the hind tibiae, the each antennomere bears one to several new genus may belong to the Late spines; genae narrow; clypeus and mouth- -Early Cretaceous subfamily Elca- parts more or less normal for ninae (5 Baisselcaninae). Longioculis is (only lower edge of labrum exposed). similarinsizetotheEarlyCretaceous Thorax: Pronotum saddle-shaped, with genus Minelcana Gorochov, Jarzembow- anterior margin convex; pronotal dorsum sky, and Coram (Gorochov et al. 2006) with seven circular light blotches with and the smallest species of the Early dark borders located in depressions sym- Cretaceous genus Panorpidium Westwood metrically positioned on each side; hind (Westwood 1854), but can be distinguished part of pronotum convex, not prolonged from both these genera as well as from over abdomen; legs covered with short other genera of Elcanidae, except the Early hairs; fore- and midfemora with pair of

r

Figs. 1–4. Longioculis burmensis in Burmese amber. 1, Lateral view of entire specimen. Bar 5 1.6 mm. 2, Metatarsus showing long spine (arrow) and two denticles (arrowheads). Bar 5 88 mm. 3, Dorsal view of anterior portion showing protruding eyes (arrows) and round colored areas on pronotum. Bar 5 542 mm. 4, Portion of antenna showing scattered setae. Bar 5 63 mm.

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Figs. 10–12. Longioculis burmensis in Burmese amber. 10, Venation of tegmina. Dotted portions represent reconstructed areas. Bar 5 1 mm. 11, Frontal view showing protruding eyes, prominent antennal cavities and enlarged antennal scapes. Bar 5 1 mm. 12, Hind leg showing denticles and spines on dorsal surface of metatibia and spines on ventral surface of basitarsus. Dotted portions represent reconstructed areas. Bar 5 1 mm. rounded apical lobules, lacking denticles; dorsal surface bearing single row of small fore tibia with single apical outer spine; denticles from 25–31 mminheightand mid- and hind tibiae with pair of apical 25–31 mm in width at base; distance spines; basal portion of hind tibia on between denticles 201–246 mm; denticles

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Figs. 5–9. Longioculis burmensis in Burmese amber. 5, Frontal view. Bar 5 375 mm. 6, Mouthparts. Bar 5 700 mm. 7, Metatarsus (arrow points to small spines on first tarsal segment of metatarsus) and mesotarsus (arrowhead). Note rows of large spines opposite small spines on ventral surface of basimetatarsus. Bar 5 800 mm. 8, Denticles on proximal portion of metatibia. Bar 5 575 mm. 9. Portion of abdomen showing remaining cercus. Bar 5 800 mm.

Proceedings of the Entomological Society of Washington went-109-03-13.3d 10/4/07 11:02:20 653 Cust # 06- 654 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON replaced on middle part (and possibly on suggests that the entire distal portion distal part) of dorsal surface of hind tibia contained a row of large spines, which is by large spines (only one complete spine, typical for elcanids. The depressed col- 265 mm long and 63 mmwide,andone ored areas on the dorsum of the prono- partial second spine visible); distance tum probably served as camouflage. between first and second large tibial spines, 315 mm; all tarsi with first segment longer DISCUSSION than 2nd and 3rd combined; apex of all Fossils of the superfamily Elcanoidea basitarsi with 2 pairs of apical spines; first appeared in the Early and midtarsus very short, less than J as long extended into the mid-Cretaceous. Mem- as basitarsus; third tarsal segments with bers of the grasshopper-appearing family well- developed, simple, slightly curved Elcanidae first appeared in the Early claws; arolium absent; fore- and midbasi- Jurassic and representatives of the sub- tarsi with 2 rows of short spines on dorsal family Elcaninae, which ranged from and ventral surfaces; hind basitarsus with Siberia to England and Brazil, were rather two rows of spines, dorsal row composed common in Lower Cretaceous localities of 27 short spines and ventral row with 7 (Martins-Neto 1991; Gorochov 1995; large spines; midtarsus without spines; Gorochov and Rasnitsyn 2002). The tegmina membranous (wings folded); ve- small size of Longioculis may indicate nation as shown in Fig. 10; tegmina how it became entrapped in amber. coloration light (transparent) with dark veins and clear area between first and second branches of RA. ACKNOWLEDGMENTS Abdomen: Short, robust; cercus slen- We thank Jim Davis for supplying the der, pointed, approximately 1/3 length of amber, J. B. Lambert and Y. Wu, De- abdomen; base of cercus with single, partment of Chemistry, Northwestern small, outer sensilla; anal plate, subgeni- University, for their analysis of the tal plate, and genital apparatus missing. amber, and Roberta Poinar for comments Material examined.—Holotype male on an earlier draft of this manuscript. in Burmese amber from the Hukawng Valley, southwest of Maingkhwan in the LITERATURE CITED state of Kachin (26u209N, 96u369E), Carpenter, F. M. 1992. Superclass Hexapoda. In northern Myanmar (Burma), deposited Kaesler, R. L. ed. Treatise on Invertebrate in the amber collection of Ron Buckley, Paleontology, Part R, Arthropoda 4, Vol- Florence, Kentucky (accession # ab ume 3. The Geological Society of America 307). The specimen is available for study and The University of Kansas, Boulder and Lawrence. 277 pp. by contacting R. Buckley. Cruickshank, R. D. and K. Ko. 2002. Geology of Etymology.—The specific name ‘‘bur- an amber locality in the Hukawng Valley, mensis’’ indicates the place of origin of northern Myanmar. Journal of Asian Earth the fossil. Sciences 21: 441–455. Comments.—Unfortunately, the left Gorochov, A. V. 1995. System and evolution of the suborder Ensifera (Orthoptera). Part 1. Pro- hind tibia is missing and only the ceedings of the Zoological Institute, Russian proximal half of the right hind tibia is Academy of Sciences [Trudy Zoologicheskogo present (the remainder is obliterated by Instituta RAN], v. 260: 1–224 (in Russian). a cavity in the amber) but the right Gorochov, A. V. and A. P. Rasnitsyn. 2002. midtarsus is present on the other side of Superorder Gryllidea. Laicharting, 1781 (5 Orthopteroidea Handlirsch, 1903). pp. 293–303. the cavity. One complete and a second In Rasnitsyn, A. P. and D. L. J. Quicke, eds. partial spine positioned at the distal end History of Insects. Kluwer Academic Publishers, of the remaining portion of the hind tibia Dordrecht.

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Gorochov, A. V., E. A. Jarzembowski, and R. A. Poinar, Jr., G. O., R. Buckley, and A. Brown. 2005. Coram. 2006. Grasshoppers and crickets (In- The secrets of Burmese amber. Mid Atlantic sects: Orthoptera) from the Lower Cretaceous Paleontology Society 29: 20–29. of southern England. Cretaceous Research 27: Sharov, A. G. 1968. Phylogeny of orthopteroid 641–662. insects. Proceedings of the Paleontological Martins-Neto, R. G. 1991. Sistema´tica dos En- Institute, Academy of Sciences of the USSR sifera (Insects, Orthopteroida) da Formac¸aˆo 118: 1–216 (in Russian). Santana, Creta´ceo Inferior do nordeste do Westwood, J. D. 1854. Contributions to fossil Brasil. Estudio Technolo´gicos, Acta Geologica entomology. Quarterly Journal of the Geo- Leopoldensia 32: 3–162. logical Society of London 10: 378–396.

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