A N N A L E S Z O O L O G I C I (Warszawa), 2013, 63(1): 143-148

A NEW SPECIES OF DICRANOMYIA STEPHENS, 1829 FROM BALTIC AMBER (DIPTERA: LIMONIIDAE)

IWONA KANIA1, *, AGNIESZKA PENAR2, and WIESŁAW KRZEMIŃSKI3

1, *Department of Environmental Biology, University of Rzeszów, Zelwerowicza 4, 35-601 Rzeszów, Poland; e-mail: [email protected] 2Wróblik Szlachecki, Sadowa 1, 38-483 Wróblik Szlachecki, Poland; e-mail: [email protected] 3Institute of Biology, Pedagogical University of Kraków, Podbrzezie 3, 31-054 Kraków, Poland; e-mail: [email protected] *Corresponding author

Abstract.— A new species of the genus Dicranomyia Stephens, 1829 (Diptera: Limoniidae) from Baltic amber (Eocene) is described. The characteristic feature distinguishing a new species, D. gorskii sp. nov. is the presence of the surprisingly strong and elongate spine on rostral prolongation of ventral gonostylus. The morphological features of the new species are briefly discussed. New replacement name Dicranomyia alexandri nom. nov. for Dicranomyia flagellata (Alexander, 1931) is proposed to avoid with Dicranomyia flagellata Edwards, 1928.

Key words.— Dicranomyia gorskii sp. nov., Limoniidae, Diptera, Baltic amber, Eocene, taxonomy, new species, Dicranomyia alexandri nom. nov., new name, homonymy.

INTRODUCTION and Johnson 1996) could also belong to Dicranomyia, but these should be revised. The genus Dicranomyia was described by First revision of these dipterans from Baltic amber Stephens in 1829, and it belongs to the most numerous was done by Alexander (1931). In this monograph dipteran family Limoniidae. This genus comprises over Alexander proposed to change a systematic position of 800 extant species, distributed worldwide with ex- all species described previously in the genus Dicra- ception of Antarctica (Krzemiński 2000b). Till now nomyia and put them to the genus Limonia Meigen, about thirty extinct species of Dicranomyia are 1803. The descriptions and information about the fossil known. The fossil representatives of the genus were representatives of the genus Dicranomyia from Baltic described based on imprints from North America amber are to be found in the papers of Giebel (1856), (Scudder 1877, 1894), Europe (Alexander 1931, Cock- Meunier (1899, 1906, 1916), Savchenko (1967) and erell 1908, 1922, Cockerell and Haines 1921, Henriksen sKrzemiński (1985). 1922, Théobald 1937, Krzemiński 1985, 2001, Krze- Eight species could be placed without doubts in the miński and Gentlini 1992), and Asia (Krzemiński genus (and subgenus) Dicranomyia, i.e. D. (D.) ale- 2000a), of various age. xandri nom. nov., D. (D.) graciosa Meunier, 1916, D. The representatives of this genus are also frequent- (D.) grandis (Meunier, 1899), D. (D.) kalandyki Krze- ly preserved as amber inclusions. Some fossil species miński, 2000b, D. (D.) lobata Meunier, 1906, D. (D.) allotted to the other Limoniidae genera (Heer 1849, meunieri (Alexander, 1931), D. (D.) perpendicularis 1856, Giebel 1856, Alexander 1931, Piton 1940, Gelhaus (Savchenko, 1967), D. (D.) sinuata Meunier, 1916.

PL ISSN 0003-4541 © Fundacja Natura optima dux doi: 10.3161/000345413X666183 144 I. KANIA, A. PENAR, and W. KRZEMIŃSKI

MATERIAL AND METHODS Material examined. Holotype, male. No. MP/3313 (6550), Coll. Andrzej Górski, Bielsko-Biała, Poland. The The study were based on material from the collec- holotype will be deposited in Institute of Systematic tion of Mr. Andrzej Górski; Bielsko-Biała, Poland. The and Evolution of Animals, Polish Academy of Sciences, specimens were studied using a Nikon SMZ 1500 Kraków, Poland. stereomicroscope. The photographs were taken with a Nikon DS-Fi1 camera equipped with a microscope. The drawings were made on the basis of specimens New replacement name for Dicranomyia flagellata and photographs. (Alexander, 1931)

The specific name of fossil taxon from the Baltic SYSTEMATIC PALEONTOLOGY amber Dicranomyia flagellata (Alexander, 1931), proposed as new combination for Limonia flagellata Order: Diptera Linnaeus, 1758 Alexander, 1931 by Krzemiński (2000b: 347) is preoc- cupied by the Dicranomyia flagellata Edwards, Family: Limoniidae Speiser, 1909 1928: 68, recent species. To avoid the homonymy and in Genus: Dicranomyia Stephens, 1829 concordance with the Article 53 of ICZN (2000), we pro- Subgenus Dicranomyia Stephens, 1829 posed here the replacement name. The history of use of the Alexander’s name is given according to the rules of Type species. Limnobia modesta Meigen, 1818; open nomenclature as proposed by Matthews (1973) by subsequent designation of Coquillett, 1910: 533. and Bengtson (1988) for names of fossil taxa.

Dicranomyia alexandri nom. nov., Kania pro Dicranomyia flagel- Dicranomyia gorskii sp. nov. lata (Alexander, 1931: 38) non Dicranomyia flagellata Ed- wards, 1928: 68. (Figs 1A–D, 2A–C) Limonia flagellata Alexander, 1931: 38. Limonia flagellata Alexander, 1931; Keilbach 1982: 318. Diagnosis. The new species can be distinguished Limonia flagellata Alexander, 1931; Evenhuis1994: 76. by the following combination of characters: vein Sc Dicranomyia flagellata (Alexander, 1931); Krzemiński 2000b: 347. very elongate, ending opposite 4/5 of Rs; m-cu in fork of Mb; the strong and elongate single spine on rostral pro- longation of ventral gonostylus, the spine reaching half DISCUSSION the length of gonostylus. Etymology. The specific name is derived from the The newly established species distinguishes from owner of private collection of inclusions in Baltic am- the others by characteristic structure of outer gonosty- ber – Andrzej Górski. lus with enlarged rostrum provided with very long and Description. Head: not well preserved; antennae strong spine. Majority of recent and known fossil (Figs 1B, 2B) 14-segmented, about 0.61 mm; scape tu- species of Dicranomyia are characteristic of pres- bular, pedicel wide and oval; flagellomeres oval, ence of 1 to 3 spines, usually small to medium sized. tapered at apex; on all segment of flagellum tiny, wispy The huge spines are to be found exceptionally, and in setae, each flagellomere with two elongate setae; seg- these cases usually they are paired. This morphologi- ments sixth and seventh invisible. cal feature is very rarely found among extant represen- Palpi not visible. tatives of the Dicranomyia. It is without doubts the Wings (Figs 1C–D, 2C): 2.18 mm long, 0.56 mm wide; apomorphic feature. The only other species known pre- vein Sc ending at 4/5 the length of vein Rs; vein R3+4 senting the enlarged spines is Dicranomyia (Dicra- about 2 and 1/7 times longer than vein Rs; R1 ending at nomyia) lobata from the Eocene Baltic amber, howev- 2/5 of the length of R2+3+4 and R3+4 measured from er the spine is shorter and more delicate than in Dicra- Rs forking; M1+2 almost twice longer than d-cell; m-cu nomyia (Dicranomyia) gorskii sp. nov. This newly in fork of vein Mb. above described species presents all the features of the Hypopygium (Figs 1A, 2A): 0.27 mm long, gonocox- venational pattern, structures of antennae and hypopy- ite half times shorter than outer gonostylus; outer gium enabling placement of it in the subgenus Dicra- gonostylus balloon-like, rostrum on outer gonostylus nomyia. provided with a single, elongate and strong spine re- The genus Dicranomyia is recently subdivided aching half the length of gonostylus; inner gonostylus into 25 subgenera (Oosterbroek 2012). All species from narrow and hooked. the Baltic amber, as well as fossils from the other local- Age and occurrence. Eocene, Baltic amber (for the ities and strata are placed in the nominative subgenus. age discussion see Weitschat and Wichard 2010). Therefore, surprisingly the remaining 24 subgenera DICRANOMYIA GORSKII SP. NOV. FROM EOCENE BALTIC AMBER 145 are lacking fossil record. On the other hand the fossil Most of the recent subgenera recognized within the record of the species attributed to the subgenus Dicra- genus Dicranomyia are distributed in the tropical nomyia ranges back from Palaeocene (Menat, France; and subtropical zones of both Old and New World, with Piton 1940) to Miocene (Dominican amber; Arillo and highest species diversity in the Oriental, Australian/ Ortuńo 2005), with fossil species known from deposits Oceanian and Neotropical regions (Oosterbroek 2012). in Europe, North America, Asia and the Caribbean. Only a few subgenera present distribution limited to

Figure 1A–D. Dicranomyia gorskii sp. nov., holotype No. MP/3313 (6550): (A) hypopygium; (B) antenna; (C) the body of specimen; (D) wing. 146 I. KANIA, A. PENAR, and W. KRZEMIŃSKI

Figure 2A–C. Dicranomyia gorskii sp. nov., holotype No. MP/3313 (6550). (A) hypopygium; (B) antenna; (C) wing venation. Abbreviations of male hypopygium: r – rostrum with spine; ing – inner gonostylus; oug – outer gonostylus. one region, numerous are present e.g. in the Orient- morphological differences resulting in taxonomic sub- al area and islands of Australian/Oceanian region. divisions and observed distributional patterns of the It is interesting that cranefies are not generally con- species ascribed to particular subgenera of Dicra- sidered strong fiers, but the mitochondrial phylog- nomyia. It seems that this group could be a very eny of the endemic Hawaiian cranefies presented by good model for the future research on evolution- Nitta and O’Grady (2008) suggested that can present ary traits, paleoecological and paleobiogeograph- higher migrational potential than expected. This could cal analyses, molecular evolution, ecology and biogeog- be also one of the explanations for the observed raphy. DICRANOMYIA GORSKII SP. NOV. FROM EOCENE BALTIC AMBER 147

ACKNOWLEDGEMENTS from Bolshaya Svetlovodnaya (Eastern Asia, Russia). Pol- skie Pismo Entomologiczne, Gdańsk, 69: 239–245. Krzemiński, W. 2000b. A new species and other representa- We are deeply indebted to the owner of private tives of the genus Dicranomyia (Diptera: Limoniidae) in collection Mr. Andrzej Górski for lending the material the collection of the Museum of Amber Inclusions, Univer- of Limoniidae amber inclusions for our disposal. sity of Gdańsk. Polskie Pismo Entomologiczne, Gdańsk, 69: 347–353. Krzemiński, W. 2001. New fossil Tipuloidea (Diptera) from the REFERENCES Fur Formation of Denmark in the collection of the Natural History Museum in London. Polskie Pismo Entomolog- Alexander, C. P. 1931. Crane flies of the Baltic Amber (Dip- iczne, 70: 333–339. tera). Bernstein-Forschungen, 2: 1–135. Krzemiński, W. and G. Gentlini. 1992. New information on Arillo, A. and V. M. Ortuńo. 2005. Catalogue of fossil insect Limoniidae from Monte Castellaro, Italy (Upper Miocene). species described from Dominican amber (Miocene). Acta Zoologica Cracoviensia, 35: 87–95. Stuttgarter Beiträge zur Naturkunde, Serie B, 352: 1–68. Linnaeus, C. 1758. Systema nature per regna tria naturae, Bengtson, P. 1988. Open nomenclature. Palaeontology, 31(1): secundum classes, ordines, genera, species, cum carac- 223–227. teribus, differentiis, synonymi, locis. Tomus I. Editio deci- Cockerell, T. D. A. 1908. Descriptions of Tertiary insects. Part ma, reformata. L. Salvii, Holmiae [= Stockholm], pp. 824. II. American Journal of Science, (4)25: 227–232. Matthews, S. C. 1973. Notes on open nomenclature and on syn- Cockerell, T. D. A. 1922. New name for a fossil tipulid fly. Ento- onymy lists. Palaeontology, 16(4): 713–719. mologist, 55: 17. Meigen, J. W. 1803. Versuch einer neuen Gattungs Eintheilung Cockerell, T. D. A. and F. H. Haines. 1921. Fossil Tipulidae der europäischen zweiflügligen Insekten. Magazin der from the Oligocene of the Isle of Wight (part). Entomolo- Insektenkunde, 2: 259–81. gist, 54: 81–84. Meigen, J. W. 1818. Systematische Beschreibung der bekann- Coquillett, D. W. 1910. The type species of the North American ten europäischen zweiflügeligen Insecten. Erster Theil. genera of Diptera. Proceedings of the United States F.W. Forstmann, Aachen., xxxvi + 332 + [1] p. National Museum, 37: 499–647. Meunier, F. 1899. Révision des Diptères fossiles types de Loew Edwards, F. W. 1928a. Diptera Nematocera from the Federat- conserves au Musée provincial de Koenigsberg. Entomolo- ed Malay States museums. Journal of the Federated Malay gy Miscellaneous, 7: 161–165, 169–182. States Museums, 14: 1–139. Meunier, F. 1906. Monographie des Tipulidae et Dixidae de Evenhuis, N. L. 1994. Catalogue of the fossil flies of the world l’ambre de la Baltique. Annales des Sciences Naturelles (Insecta: Diptera). Backhuys, Leiden, 1–600. Zoologie, (9)4: 349–401. Gelhaus, J. K. and R. Johnson. 1996. First record of crane flies Meunier, F. 1916. Beitrag zur Monographie des Tipuliden des (Tipulidae: Limoniidae) in Upper Cretaceous Amber from Bernsteins. Zeitschrift der Deutschen Geologischen New Jersey, U.S.A. Transactions of the American Entomo- Gesellschaft, 68: 477–493. logical Society, 122: 55–65. Nitta, J. H. and P. M. O’Grady. 2008. Mitochondrial phylogeny Giebel, C. G. A. 1856. Fauna der Vorwelt mit steter Berück- of the endemic Hawaiian craneflies (Diptera, Limoniidae, sichtigung der lebenden Thiere. Monographisch darger- Dicranomyia): Implications for biogeography and spe- stellt. Zweiter Band. Gliederthiere. Erste Abtheilung. In- cies formation. Molecular Phylogenetics and Evolution, 46 secten und Spinnen. F. A. Brockhaus, Leipzig, 411 pp. (3): 1182–1190. Henriksen, K. L. 1922. Eocene insects from Denmark. Oosterbroek, P. 2012. Catalogue of the Craneflies of the World. Danmarks Geologiske Undersøgelse København (2), 37: (Diptera, Tipuloidea: Pediciidae, Limoniidae, Cylindrotomi- 1–36. dae, Tipulidae). http://nlbif.eti.uva.nl/ccw/index.php. Last Heer, O. 1849. Die Insektenfauna der Tertiärgebilde von Oe- updated 28 December 2012. Accessed 11 January 2013. ningen und von Radoboj in Croatien. Zweite Theil: Heus- Piton, L. 1940. Paléontologie du gisement éocène de Menat chrecken, Florfliegen, Aderflügler, Schmetterlinge und (Puy-de-Dôme) (Flore et faune). Mémoire de la Société Fliegen. W. Engelmann, Leipzig, pp. 264. d'Histoire Naturelle d’Auvergne, 1: 1–303. Heer, O. 1856. Über die fossilien Insecten von Aix in der Pro- Savchenko, E. N. 1967. New fossil limoniid fly (Diptera, Limo- vance. Vierteljahrsschr. Naturforschenden Gessellschaft niidae) from the Baltic amber. Dopovidi Akademii Nauk in Zürich, 1: 1–40. URSR(B), 5: 469–473. ICZN 2000. International Code of Zoological Nomenclature Scudder, S. H. 1877. The first discovered traces of fossil (4th edition). 305 pp. insects in the American tertiaries. Bulletin of the United Keilbach, R. 1982. Bibliographie und Liste der Arten tierisch- States Geological and Geographical Survey of the Territo- er Einschlüsse in fossilen Harzen sowie ihrer Aufbe- ries. 3: 741–742. wahrungsorte. Deutsche Entomologische Zeitschrift, Scudder, S. H. 1894. Tertiary Tipulidae, with special reference 29(4–5): 301–491. to those of Florissant, Colorado. Proceedings of the Amer- Krzemiński, W. 1985. Limoniidae (Diptera Nematocera) from ican Philosophical Society, 32: 163–245. Baltic amber (in the collection of the Museum of the Earth Speiser, P. 1909. 4 Orthoptera. Orthoptera Nematocera. Wis- in Warsaw). Part I. Subfamily Limoniinae. Prace Muzeum senschaftliche Ergebnisse der Schwedischen Zoologische Ziemi, Warszawa, 37: 113–117. Expededition nach Kilimandjaro, Meru 10 (Diptera), Krzemiński, W. 2000a. The Oligocene Tipulomorpha (Diptera) 31–65. 148 I. KANIA, A. PENAR, and W. KRZEMIŃSKI

Stephens, J. F. 1829. The nomenclature of British insects; Théobald, N. 1937. Les insects fossils des terrains oligocènes being a compendious list of such species as are contained de France, Nancy, 473 pp. in the Systematic Catalogue of British Insects, and form- Weitschat, W. and W. Wichard. 2010. Baltic amber, pp. 80–115. ing a guide to their classificati on, &c. &c. Baldwin In: Penney, D. (Ed.), Biodiversity of fossils in amber from & Cradock, London, [2] + 68 p. major world deposits. Siri Scientific Press, Manchester, UK.

Received: January 17, 2013 Accepted: February 26, 2013