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The First Pipizine Hoverfly from the Oligocene of Céreste, France

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The First Pipizine Hoverfly from the Oligocene of Céreste, France The first pipizine hoverfly from the Oligocene of Céreste, France VALENTIN NIDERGAS, JIŘÍ HADRAVA, ROMAIN GARROUSTE, JAKUB PROKOP, THOMAS SCHUBNEL, and ANDRÉ NEL Nidergas, V., Hadrava, J., Garrouste, R., Prokop, J., Schubnel, T., and Nel, A. 2018. The first pipizine hoverfly from the Oligocene of Céreste, France. Acta Palaeontologica Polonica 63 (3): 539–548. The Oligocene Oligopipiza quadriguttata Nidergras, Hadrava, and Nel gen. et sp. nov. is the first fossil Pipizinae found in the lacustrine outcrop of Céreste (South-East of France). It differs from the other Pipizinae in the male genitalia, with a surstylus without tooth and shorter than epandrium, and a long epandrium with a very deep and narrow median theca. It is compared to other extant and fossil Pipizinae. Its position in this clade is supported by its inclusion in previous mor- phological phylogenetic analysis of the Syrphidae. Palaeoecological inferences for the paleobiota of Céreste are made based on this taxon and point to the presence of a mixed forest. The taphonomy of these flies is discussed. They were probably embedded in surface microbial mats. The pollinator role of Oligopipiza quadriguttata is also discussed on the basis of the presence of pollen surrounding the fossil flies. Key words: Insecta, Syrphidae, Pipizinae, palaeoecology, Paleogene, France. Valentin Nidergas [[email protected]], Romain Garrouste [[email protected]], Thomas Schubnel [thom- [email protected]], and André Nel [[email protected]] (corresponding author), Institut Systématique Evolution Biodiversité (ISYEB), Muséum national d’Histoire naturelle, CNRS, Sorbonne Université, EPHE, 57 rue Cuvier, CP 50, F-75005 Paris, France. Jiří Hadrava [[email protected]] and Jakub Prokop [[email protected]], Department of Zoology, Faculty of Science, Charles University, Viničná 7, CZ-128 43, Praha 2, Czech Republic; and Institute of Entomology, Biological Centre, Czech Academy of Science, Branišovská 31, CZ–370 05 České Budějovice, Czech Republic. Received 2 May 2018, accepted 15 June 2018, available online 20 July 2018. Copyright © 2018 V. Nidergas et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License (for details please see http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. accurate fossil members of the Syrphidae and Pipunculidae Introduction are dated from the Eocene (Dirickxs 2009; Archibald et al. With more than 6000 described extant species living in all 2014). Kovalev (1979) recorded what may be the oldest ex- climatic regions (except Antarctica, Spitsbergen, and some ample of Syrphidae from the Late Cretaceous resins of the remote oceanic islands), the Syrphidae are one of largest Taimyr in Siberia, but this fossil has never been described families of Diptera in number of species. They have import- or re-examined (Evenhuis 1994). Fossil Syrphidae are ant roles in ecosystems as pollinators (Ssymank et al. 2008; poorly known compared to extant hoverflies, with ca. 106 Glaum 2017; Lefèbvre et al. 2018) and as aphid predators described species all over the world (Hull 1945; Evenhuis (Rojo et al. 2003; Bičík and Láska 2011). Grimaldi (1999) con- 1994; Kotthoff and Schmid 2005; Dirickx 2009). The last sidered that the Syrphidae radiated during the Cretaceous. description of a fossil syrphid dates back thirteen years ago Wiegmann et al. (2003) dated the clade Schizophora be- (Kotthoff and Schmid 2005). tween 107 and 84 Myrs and considered the Syrphidae as The extant Pipizinae are widely spread in all world eco- their sister group, suggesting the same minimum age for this zones except in the Polar ecozones and the Afrotropical re- family. Wiegmann et al. (2011) dated the Syrphidae from gion (Thompson 1972; Vujić et al. 2013; Mengual et. al. ca. 100 Myrs. The “mid” Cretaceous corresponds to the 2015). It contains approximately 180 species distribu ted in period of radiation and diversification of the angiosperms eight genera: Claussenia Vujić and Ståhls, 2013, Cry pto- (Barba-Montoya et al. 2018). Young et al. (2016) and Pauli pipiza Mutin, 1998, Heringia Rondani, 1856, Neo cnemodon et al. (2018) recently proposed the Syrphidae as sister group Goffe, 1944, Pipiza Fallén, 1856, Pipizella Ron dani, 1856, of the Pipunculidae + Schizophora. Nevertheless, the oldest Trichopsomyia Williston, 1888, and Triglyphus Loew, 1840. Acta Palaeontol. Pol. 63 (3): 539–548, 2018 https://doi.org/10.4202/app.00500.2018 540 ACTA PALAEONTOLOGICA POLONICA 63 (3), 2018 Pipizines have long been considered as a tribe of the sub- compressed “mummies”. These animals were probably em- families Syrphinae or Eristalinae (Ståhls et al. 2003; Hippa bedded in microbial mats that were originally floating at the and Ståhls 2005). They have strong morphological simi- water surface (Nel 1991). larity with the Eristalinae (e.g., pilose postpronotum), but Drawings were made with a drawing tube attached to they share ecological and biological characteristics with the a Nikon SMZ1500 stereomicroscope and digitized using Syrphinae (Mengual et al. 2015). Larvae are aphid predators Adobe ® Photoshop-Elements 12. Then, drawings were on several species of plants, like Rumex Linnaeus, 1753, vectorized and reworked with Inkscape v0.92. Photographs Prunus Linnaeus, 1753 or Ulmus Linnaeus, 1753 according were made with a reflex camera Nikon D800 mounted on a to the “Syrph The Net” database (Speight and Castella 2016). Nikon SMZ25 stereomicroscope, edited with DxO PhotoLab Only few fossils are currently attributed to the Pipizinae, v1.1.2 and then focus-stacked with the “D-map” algorithm viz., Pipiza melanderi Hull, 1945 (lowermost Oligocene of on Zerene Stacker v1.04. We used environmental scanning Florissant, USA), Pipiza venilia Heyden, 1870 (Oligocene of electron microscopy (ESEM) to obtain microstructural Rott, Germany), Pseudopipiza antiqua Hull, 1945, Pseudo- details of these fossils, with the back-scattered electrons pipiza europa Hull, 1945, and Palaeopipiza xenos Meunier, (BSE) mode. We used Energy-dispersive X-ray spectros- 1902, (Eocene, Baltic amber) (Heyden 1870; Meunier 1902; copy (EDS) in order to analyze the element composition of Hull 1945; Evenhuis 1994). Théobald (1937) described and the fossils and its matrix. Then the spectra were plotted in listed all the known fossil Syrphidae from France, but none R and reworked on Inkscape v.0.92. Electron microscopy in was described from the middle Oligocene of Lubéron. In BSE mode (SEM BSE) makes it possible to highlight dif- this paper, we describe one new genus and species of the ferences in chemical composition (atomic number contrast) Pipizinae from the middle Oligocene of Céreste, Lubéron, between the mineral matrix and the fossilized residues, in in Southern France. We also discuss the systematic position addition to the microtopographic differences revealed by the of the newly described pipizine fossil based on a morphol- conventional mode. EDS spectrometry makes it possible to ogy phylogenetic analysis. know the chemical nature of these differences and to iden- Institutional abbreviations.—RNGL, Public Museum of the tify the elements involved in the fossilization process, which Réserve Naturelle Géologique du Lubéron, Apt, France. can provide information on the sedimentary conditions, the taphonomic process and the initial nature of the fossilized Other abbreviations.—BSE, back-scattered electrons; dm-cu, organic elements. These two methods are mainly allowed by distal median cubital crossvein; EDS, energy-disper sive X-ray the use of the environmental mode to overcome the need to spectroscopy; M1, first branch of median vein; R1, R2+3, metallize the samples. R4+5, branches of radius; r-m, radial median crossvein; Sc, We followed Thompson (1999) for morphology nomen- subcostal vein; sv, supplementary pseudovein. clature. The systematics of hoverflies follows Mengual et al. (2015). A new phylogenetic analysis to assess the systematic position of this new fossil was performed using 111 charac- Material and methods ters and 96 species, including the new fossil taxon, based on the morphological matrix used by Mengual et al. (2015), In total, 15 fossil specimens of the new genus and new spe- based on 111 characters and 96 species including the new cies were collected, by digging and splitting the laminites fossil taxon (see SOM; Mengual et al. 2015; Hippa and Ståhls with a hammer and a spatula, during the last 25 years in the 2005). The tree research was performed using Wagner’s par- National Geological Reserve of Lubéron, Cereste, France, simony in PAUP* 4.0 b10 (Swofford 2002), with the follow- from middle Oligocene clay-limestone laminite (base of late ing parameters: heuristic search 100 replicates, non-ordered Rupelian) (Cautru and Gigot 1982; Ducreux et al. 1985). characters, and TBR recombination. The 15 specimens of the type series, described herein, are deposited at the public collection of the Réserve Naturelle Géologique du Lubéron at Apt, France. All the specimens were chosen to form the type material. Systematic palaeontology The Oligocene paleolatitude of the outcrop of Céreste was very close to the modern one. The paleoclimate in the Order Diptera Linnaeus, 1758 area was cooler than during the Eocene but still relatively Family Syrphidae Latreille, 1802 warm (Pound and Salzmann 2017). The layers that contain Subfamily Pipizinae Williston, 1885 insects at Céreste are very
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