from Baltic amber

E

N. Jb. Geol. Paläont. Abh. 277/2 (2015), 167–174Article Stuttgart, August 2015

A new species of Dicranomyia Stephens, 1829 (Diptera: Limoniidae) from Baltic amber (Eocene) Wiesław Krzemiński, Iwona Kania, and Roma Durak With 5 figures

Abstract: The description of a new species of the genus Dicranomyia Stephens, 1829 (Diptera: Limoniidae) from Baltic amber (Eocene) is presented. The morphological features of the newly described species Dicranomyia herczeki Krzemiński & Kania sp. nov. indicate that this new species probably belongs to the subgenus Sivalimnobia Alexander, 1963 and it is its first representative in Baltic amber and among fossil taxa of Dicranomyia. Key words: Diptera, Limoniidae, Baltic amber, Eocene, morphology, taxonomy, new species.

1. Introduction

While the subgenus Sivalimnobia is currently represented by 15 extant species, presently distributed mainly in Oriental Region (14 species), known also from West Palearctic (one species) (Oosterbroek, 2015) seems to be extremely rare among fossils. The present finding is the first fossil report of this subgenus.

Representatives of the genus Dicranomyia Stephens, 1829 (Diptera: Limoniidae) are cosmopolitan in distribution and are subdivided into 24 Recent subgenera. The subgenus Dicranomyia s. str. is the most numerous in species with over 800 extant species. The fossils of the genus Dicranomyia are also well represented either as impression fossils or amber inclusions and comprises over 40 extinct species known from different since Eocene (Lutetian) (Giebel 1856; Scudder 1877, 1894; Meunier 1899, 1906, 1916; Cockerell 1908, 1922; Cockerell & Haines 1921; Henriksen 1922; Alexander 1931; Théobald 1937; Savchenko 1967a, b; Krzemiński 1985, 2000a, b, 2001; Krzemiński & Gentlini 1992, Podenas & Poinar 2012; Kania et al. 2013; Kania 2014). Most of them are classified within the subgenus Dicranomyia, one of them within the subgenus Caenolimonia – Dicranomyia (Caenolimonia) alexbrowni Podenas & Poinar, 2012 and only one to the subgenus Melanolimonia – Dicranomyia (Melanolimonia) krzeminskii Kania, 2014. Members of other subgenera are not known in the fossil record (Evenhuis 2014).

2. Material and methods This study is based on material from the collection of the Institute of Systematic and Evolution of Animals, Polish Academy of Sciences (ISEA PAS), Kraków. The specimen was studied using a Nikon SMZ 1500 stereomicroscope. The microphotographs were taken with a Nikon DS-Fi1 camera equipped with a microscope. The drawings were made on the basis of specimen and photographs. The measurements of specimen were taken with NIS-Elements D 3.0 software and were given only in case when relevant structures were not distorted. Wing venation terminology follows Krzemiński & Krzemińska (2003). Vein abbreviations are as follows: C, costa; Sc, subcosta; Rs, sector radii; R1 to R5, radial veins (radius); M1+2 to M4, medial veins (media); Cu, cubital veins (cubitus); A1 to A2 anal veins; m-cu, medial-cubital cross-vein; sc-r; subcostal-radial cross-vein.

©2015 E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany

DOI: 10.1127/njgpa/2015/0512

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168 Wiesław Krzemiński et al.

Fig. 1. Dicranomyia (Sivalimnobia) herczeki sp. nov., No. MP/3428 (male), holotype; A – antenna: the last flagellomeres; B – wing venation; C – hypopygium. Abbreviations: aed – eadeagus; gx – gonocoxite; ing – inner gonostylus, tg IX – tergite IX; d – discal cell (d-cell).

Type species: Dicranomyia fortis Brunetti 1912; by original designation.

3. Systematic palaeontology Order Diptera Linnaeus 1758 Family Limoniidae Speiser 1909 Subfamily Limoniinae Speiser 1909 Genus Dicranomyia Stephens 1829

Dicranomyia (Sivalimnobia) herczeki Krzemiński & Kania sp. nov. Figs. 1-3

Type species: Limnobia modesta Meigen 1818; by subsequent designation of Coquillett 1910, p. 533.

Etymology: The specific name is dedicated to Professor Aleksander Herczek, the eminent specialist on fossil and Recent Hemiptera.

Subgenus Sivalimnobia Alexander, 1963

Type material: Holotype: No. MP/3428 (male), Baltic amber (Eocene; Lutetian), housed in the Institute of Systematic and Evolution of Animals, Polish Academy of Sciences (ISEA PAS), Kraków.

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A new species of Dicranomyia Stephens, 1829 169

Fig. 2. Dicranomyia (Sivalimnobia) herczeki sp. nov., No. MP/3428 (male), holotype; A – antenna; B – the body, lateral view; C – head with antennae and thorax.

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Fig. 3. Dicranomyia (Sivalimnobia) herczeki sp. nov., No. MP/3428 (male), holotype; A – wing venation; B-D – hypopygium. Abbreviations: gx – gonocoxite; ing – inner gonostylus. Arrows indicate two processes, one on each inner gonostylus.

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Fig. 4. The morphology of hypopygium of some Recent representatives of the subgenus Sivalimnobia and the fossil Dicranomyia (Sivalimnobia) herczeki sp. nov. Abbreviations: aed – aedeagus, gx – gonocoxite; ing – inner gonostylus, oug – outer gonostylus, p – parameres, pr – process, r – rostrum, tg IX – tergite IX.

partially covered by the leg of specimen and milky veil, male genitalia visible only in one position – in lateral view, outer gonostylus and rostral arm invisible. The visible part of the specimen show a unique morphology most similar to those characterizing the subgenus Sivalimnobia: short gonocoxite with wide process occurred in 2/3 of its length, inner balloon-like gonostylus with distinctly very long and tiny process, in new species 4 times longer than length of inner gonostylus. Large and straight aedeagus is only slightly longer than narrow and straight paramere visible in lateral view (Fig. 4).

Diagnosis: Vein Sc ending just after 1/2 of vein Rs, m-cu behind the bifurcation of Mb, in 1/4 of d-cell base, inner gonostylus balloon-like but relatively small with very elongate process, 4 times longer than the length of inner gonostylus. Description: Body (Fig. 2B): pale brown, 5.81 mm long. Head (Fig. 2C): antenna (Figs 1A, 2A-C) not very elongate, if bent backward reaching mesothorax, 1.70 mm long, 14-segmented; scape cylindrical, pedicel barrel-like, shorter than scape, flagellomeres 1-3 short and crowded, flagellomeres 4-14 elongate, each of these flagellomeres slightly narrowed in the middle; last flagellomere elongate and cylindrical, longer than penultimate one; antenna with three elongate setae on each flagellomere, on segments 4-14 longer or equal in length (last one) to segments bearing them, on segments 1-3 shorter than length of segments bearing them and much shorter in comparison to elongate setae on flagellomeres 4-14; last segment of antenna with additional two elongate setae at apex and three elongate setae at base, these elongate setae longer or same length as segment bearing them; pedicel with rosette of elongate setae at base. Palpus elongate, 4-segmented, last segment very short, distinctly shorter than penultimate one. Wing (Figs. 1B, 2B, 3A): 5.24 mm long, 1.71 mm wide; pterostigma present; vein Sc comparatively elongate, ending just behind 1/2 of vein Rs; sc-r short, its length from the tip of Sc; vein Rs slightly arcuate, about 1/2 length of R3+4; R1 ending approximately 1/3 of length of R2+3+4; r-r (R2) short, approximately its length before end of R1; M3 shorter than length of d-cell; cross-vein m-cu just behind the bifurcation of Mb, in 1/4 the length of d-cell base; A1 and A2 slightly sinuous. Abdomen: hypopygium (Figs. 1C, 3B-D.) 0.86 mm long,

Remarks: According to Alexander (1963) the representatives of the subgenus Sivalimnobia are characterized by wing venation and the structures of hypopygium with some features characteristic for Dicranomyia and typical of Limonia and he gives the features which characterize subgenus Sivalimnobia like wing venation with vein Sc ending beyond origin of Rs, usually at or before midlength of the vein and male hypopygium with the ventral gonostylus bearing a slender rostral arm that is triangularly expanded at outer end, with stout spine and a second separate spinous arm placed near basal part of gonostylus. The morphological features of the new species distinctly indicate that it belongs to the subgenus Sivalimnobia. The new described species is distinguished especially by characteristic, very elongate process on balloon-like, not atrophied inner gonostylus. A similar, very elongate process on inner gonostylus occurred between balloon-like gonostylus and rostral arm of the inner gonostylus, could be found among some recent species of the subgenus, e.g. in Dicranomyia (Sivalimnobia) kali (Alexander, 1963), especially the distinct similarities in morphology of its process can be found in Dicranomyia (Sivalimnobia) rahula (Alexander 1963) (Fig. 4).

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Fig. 5. The distribution of Recent and fossil Dicranomyia (Sivalimnobia), Dicranomyia (Melanolimonia), and Dicranomyia (Caenolimonia).

15 extant species (Oosterbroek 2015) and it seems to be extremely rare among fossils. The representatives of extant Sivalimnobia, like Dicranomyia (Sivalimnobia) aquosa, are associated with waterfalls, the flies occur near shaded wet rock faces that are kept permanently damp, such as the splash zones at the sides of waterfalls, or seepages on cliffs, usually bordering streams (Boardman 2007; Stubbs 2003); they are also found in the vicinity of small waterfalls and projecting boulders in woodlands streams (Crossley 2001); also near flushed red marl slopes and flushed slopes (Ashe et al. 2008). Larvae of these flies are assumed to be saprophagous in crevices in wet rocks (Stubbs 2003). Palaeobotanists identified over 200 species of spore-bearing, herbaceous and arborescent plants from the Baltic amber forests (Kohlman-Adamska 2001; Pielińska 2008). These include plants from diverse habitats: mountains, lowlands and swampy areas. Three main amber-bearing forest communities were distinguished (Kohlman-Adamska 2001). Coniferous forests in the higher mountains were composed of sequoias (Sequoia Endlicher), umbrella pines (Sciadopitys Siebold & Zuccarini), firs (Abies Miller),

4. Discussion Among Diptera, with generic taxa subdivided into subgenera, the common situation is that only a few of these subgenera are recorded in the fossil record. The same situation is observed within the genus Dicranomyia, with a few only of subgenera recognized present among fossils (Fig. 5). The subgenus Sivalimnobia of the genus Dicranomyia was until now not reported from fossil material. We have pointed out the importance of a new species Dicranomyia (Sivalimnobia) herczeki sp. nov. as the first representative of the subgenus Sivalimnobia among Baltic amber inclusions and fossils. This fossil put a new light on the period of diversification of the genus Dicranomyia and taxonomic diversity of the Eocene Limoniidae. With this finding three of over 20 subgenera recognized within Dicranomyia are found among Baltic amber inclusions (Melanolimonia and Sivalimnobia), and subgenus Caenolimonia is reported from Miocene Mexican amber. The new species is established on a single male specimen, and no more specimens representing this subgenus was found so far among inclusions or compression/impression fossils. Sivalimnobia is one of the not numerous in species subgenus Dicranomyia, with

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spruces (Picea A. Dietrich), larches (Larix Miller), and numerous representatives of the cypress family (Cupressaceae): the Californian incense-cedar Calocedrus Kurz, Thujopsis Siebold & Zuccarini ex Endlicher, Chamaecyparis Spach, and Thuja Linnaeus. In the lightly forested steppes, which covered lower parts of mountains, there were mainly species of pines (Pinus Linnaeus), palms, and numerous species of oaks (Quercus Linnaeus), both evergreen and with falling leaves; other trees grew there as well: beeches (Fagus Linnaeus), chestnuts (Castanea Miller), maples (Acer Linnaeus), cycads of the genus Zamia Linnaeus; shrubs, such as the magnolias, holly and some laurel family (Lauraceae) plants; grasses dominated the undergrowth. Swamp forests grew in the damp river valleys, and in them grew the exotic Chinese swamp cypresses (Glyptostrobus Endlicher), shrubs of the families Salicaceae, Myricaceae and Clethraceae, as well as herbaceous plants of the family Commelinaceae (Kohlman-Adamska 2001; Pielińska 2008; Szwedo & Sontag 2009). Conditions during exudation of resin, later transformed into Baltic amber were not uniform, comprising montane/submontane forest habitats as well as lowland forest and forest-steppe habitats, and wet and swampy habitats, with streams, creeks and rivers (Sontag & Szwedo 2009, 2013; Szwedo & K ania 2015). Presence of resin exuding trees, forming the amber traps in microhabitats suitable for extinct representatives of Sivalimnobia, influenced the probability of preservation of these flies in amber. The rrarity of the subgenus Sivalimnobia among Baltic amber inclusions most probably results from habitats and biology of these insects. We can assume that extinct representatives of this subgenus preferred similar habitats as the extant ones.

Alexander, C.P. (1963): New or little-known Tipulidae from eastern Asia (Diptera). LI. – Philippine Journal of Science, 92: 205-246. Ashe, P., O’Connor, J.P., Alexander, K.N.A., Chandler, P.J. & Oosterbroek, P. (2008): Additions, with some corrections, to the records of the craneflies of Ireland (Diptera: Cylindrotomidae, Limoniidae, Pediciidae and Tipulidae). – Bulletin of the Irish Biogeographical Society, 32: 6-20 Boardman, P. (2007): A provisional account and atlas of the craneflies of Shropshire. – 96 pp.; Oswestry (Privately published). Brunetti, E. (1912): Diptera Nematocera (excluding Chironomidae and Culicidae). – Fauna of British India, including Ceylon and Burma, 1: 1-581. Cockerell, T.D.A. (1908): Descriptions of Tertiary insects. Part II. – American Journal of Science, (4), 25: 227-232. Cockerell, T.D.A. (1922): New name for a fossil tipulid fly. – Entomologist, 55: 17. Cockerell, T.D.A. & Haines F.H. (1921): Fossil Tipulidae from the Oligocene of the Isle of Wight (part). – Entomologist, 54: 81-84. Coquillett, D.W. (1910): The type species of the North American genera of Diptera. – Proceedings of the U. S. National Museum, 37: 499-647. Crossley, R. (2001): Entomological report: Diptera (Tipuloidea and Empidoidea). – Naturalist, 126 (1039): 193-196. Evenhuis, N.L. (2014): Catalogue of the fossil flies of the world (Insecta: Diptera). Last revised 18 November 2014. http://hbs.bishopmuseum.org/fossilcat/ Giebel, C.G.A. (1856): Fauna der Vorwelt mit steter Berücksichtigung der lebenden Thiere. Monographisch dargestellt. Zweiter Band. Gliederthiere. Erste Abtheilung. Insecten und Spinnen. – 411 pp.; Leipzig (Brockhaus). Henriksen, K.L. (1922): Eocene insects from Denmark. – Danmarks Geologiske Undersøgelse, (2), 37: 1-36. Kania, I., Penar, A. & Krzemiński, W. (2013): A new species of Dicranomyia Stephens, 1829 from Baltic amber (Diptera: Limoniidae). – Annales Zoologici, 63 (1): 143-148. Kania, I. (2014): Subfamily Limoniinae Speiser, 1909 (Diptera, Limoniidae) from Baltic amber (Eocene): the genus Dicranomyia Stephens, 1829. – Zoological Journal of the Linnean Society, 170 (4): 748-778. Kohlman-Adamska, A. (2001): A graphic reconstruction of an “amber forest”. – In: Kosmowska-Ceranowicz, B. (Ed.): The amber treasure trove. – Museum Earth Documentary Studies, 18: 15-18. Krzemiński, W. (1985): Limoniidae (Diptera Nematocera) from Baltic amber (in the collection of the Museum of the Earth in Warsaw). Part I. Subfamily Limoniinae. – Prace Muzeum Ziemi, 37: 113-117. Krzemiński, W. (2000a): The Oligocene Tipulomorpha (Diptera) from Bolshaya Svetlovodnaya (Eastern Asia, Russia). – Polish Journal of Entomology, 69: 239-245. Krzemiński, W. (2000b): A new species and other representatives of the genus Dicranomyia (Diptera: Limoniidae) in the collection of the Museum of Amber Inclusions, University of Gdańsk. – Polish Journal of Entomology, 69: 347-353. Krzemiński, W. (2001): New fossil Tipuloidea (Diptera) from the Fur Formation of Denmark in the collection of the Natural History Museum in London. – Polish Journal of

Acknowledgements We would like to thank Prof. Dr. Martin Langer and Dr. Günter Schweigert and three anonymous reviewers for valuable comments and corrections. Mr. Marcin Gąsior, M.Sc (Museum and Institute of Zoology, Polish Academy of Sciences, Warsaw) for the graphical preparation of the map. We are deeply indebted to Mr. Joseph Ohimor, M.Sc. (Centre for Language Studies, University of Rzeszów) for English language correction.

References Alexander, C.P. (1931): Crane flies of the Baltic Amber (Diptera). – Bernstein-Forschungen, 2: 1-135.

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Scudder, S.H. (1894): Tertiary Tipulidae, with special reference to those of Florissant, Colorado. – Proceedings of the American Philosophical Society, 32: 163-245. Speiser, P. (1909): 4 Orthorapha. Orthorapha Nematocera. – Wissenschaftliche Ergebnisse der Schwedischen Zoologischen Expedition nach Kilimandjaro, Meru, 10 (Diptera): 31-65. Stephens, J.F. (1829): The nomenclature of British insects; being a compendious list of such species as are contained in the Systematic Catalogue of British Insects, and forming a guide to their classification. – [2] + 68 pp.; London (Baldwin & Cradock). Stubbs, A.E. (2003): Tipulidae and Allies – Craneflies. – Managing Priority Habitats for Invertebrates, 17: 1-158. Szwedo, J. & Kania I. (2015): Climatic reconstructions based on inclusions. – Amber news review 2014/2015, World Amber Council, Gdańsk, Poland, 2015, 6-21. Szwedo, J. & Sontag, E. (2009): The traps of the “amber trap”. How inclusions could trap scientists with enigmas. – Denisia, 26: 155-169. Szwedo, J. & Sontag, E. (2013): The flies (Diptera) say that amber from the Gulf of Gdańsk, Bitterfeld and Rovno is the same Baltic amber. – Polish Journal of Entomology, 82 (4): 379-388. Théobald, N. (1937): Les insects fossils des terrains oligocènes de France. – 473 pp.; Nancy.

Entomology, 70: 333-339. Krzemiński, W. & Gentlini, G. (1992): New information on Limoniidae from Monte Castellaro, Italy (Upper Miocene). – Acta Zoologica Cracoviensia, 35: 87-95. Linnaeus, C. (1758): Systema nature per regna trianaturae, secundum classes, ordines, genera, species, cum caracteribus, differentiis, synonymi, locis. Tomus I. Editiodecima, reformata. – 824 pp.; Holmiae (L. Salvii). Krzemiński, W. & Krzemińska, E. (2003): Triassic Diptera: descriptions, revisions and phylogenetic relations. – Acta Zoologica Cracoviensia, 46 (Supplement – Fossil Insects): 153-184. Meigen, J.W. (1818): Systematische Beschreibung der bekannten europäischen zweiflügeligen Insecten. Erster Theil. – xxxvi + 332 + [1] pp.; Aachen (Forstmann). Meunier, F. (1899): Révision des Diptères fossiles types de Loew conserves au Musée provincial de Koenigsberg. – Entomology Miscellaneous, 7: 161-182. Meunier, F. (1906): Monographie des Tipulidae et Dixidae de l’ambre de la Baltique. – Annales des Sciences Naturelles, Zoologie, (9), 4: 349-401. Meunier, F. (1916): Beitrag zur Monographie des Tipuliden des Bernsteins. – Zeitschrift der Deutschen Geologischen Gesellschaft, 68: 477-493. Oosterbroek, P. (2015): Catalogue of the Crane-flies of the World. (Diptera, Tipuloidea: Pediciidae, Limoniidae, Cylindrotomidae, Tipulidae). http://nlbif.eti.uva.nl/ccw/ index.php. Last updated 05 Jun 2015. Pielińska, A. (2008): Families and genera of the Eocene amberiferous forests after the list by Professor Hanna Czeczott, 1961, “The flora of the Baltic amber and its age”. – Bursztynisko 30: 20-22. Podenas, S. & Poinar, G.O. (2012): New short-palped crane flies (Diptera: Limoniidae) from Mexican amber. – Proceedings of the Entomological Society of Washington, 114 (3): 347-371. Savchenko, E.N. (1967a): New fossil limoniid fly (Diptera, Limoniidae) from the Baltic amber. – Dopovidi Akademii Nauk URSR, (B), 5: 469-473. Savchenko, E.N. (1967b): In: Savchenko, E.N. & Krivolutskaya, G.O. (Eds.): Limoniidae of the south Kuril Islands and south Sakhalin. – 160 pp.; Kiev (Akademia Nauk Ukrayinskoyi SSR) (in Russian). Scudder, S.H. (1877): The first discovered traces of fossil insects in the American tertiaries. – Bulletin of the United States Geological and Geographical Survey of the Territories, 3: 741-742.

Manuscript received: March 23rd, 2015. Revised version accepted by the Stuttgart editor: May 13th, 2015.

Addresses of the authors:

Wiesław Krzemiński, Institute of Biology, Pedagogical University of Kraków, Podbrzezie 3, 31-054 Kraków, Poland; e-mail: [email protected] Iwona Kania (corresponding author), Department of Environmental Biology, University of Rzeszów, Zelwerowicza 4, PL35-601 Rzeszów, Poland; e-mail: [email protected] Roma Durak, Department of Invertebrate Zoology, University of Rzeszów, Zelwerowicza 4, PL35-601 Rzeszów, Poland; e-mail: [email protected]

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