The Genus Leptoconops Skuse (Diptera: Ceratopogonidae) in Early Cretaceous Charentese Amber

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The oldest psyllipsocid booklice, in Lower Cretaceous amber from Lebanon (Psocodea,Trogiomorpha, Psocathropetae, Psyllipsocidae)

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Joanna Choufani, Dany Azar, Vincent Perrichot, Carmen Soriano, Paul Tafforeau & André Nel

Palaeobiodiversity and Palaeoenvironments

ISSN 1867-1594

Palaeobio Palaeoenv DOI 10.1007/s12549-011-0057-1

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Palaeobio Palaeoenv DOI 10.1007/s12549-011-0057-1

ORIGINAL PAPER

The genus Leptoconops Skuse (Diptera: Ceratopogonidae) in Early Cretaceous Charentese amber

Joanna Choufani & Dany Azar & Vincent Perrichot & Carmen Soriano & Paul Tafforeau & André Nel

Received: 23 May 2011 /Revised: 14 July 2011 /Accepted: 21 July 2011 # Senckenberg Gesellschaft für Naturforschung and Springer 2011

Abstract A new species of biting midge is described and group of subgenera [Holoconops Kieffer (Ann Hist-Nat figured based on five females from the uppermost Albian Mus Natl Hung 16:31–136, 1918)+(Megaconops Wirth and amber of France. One specimen preserved in opaque amber Atchley+Leptoconops s. str. + Proleptoconops Clastrier was reconstructed by propagation phase contrast X-ray (Parassitologia 16:231–238, 1974))], making inference on its synchrotron microtomography, allowing for detailed obser- palaeoecology possible, with larvae of this clade living in vation of minute external features. Leptoconops daugeroni moist and usually saline sandy soil on coastal and inland Choufani, Azar and Nel, sp. nov. can be attributed to the beaches, which is congruent with the current reconstruction of the palaeoenvironment of this amber deposit. J. Choufani : A. Nel CNRS UMR 7205, CP 50, Entomologie, Muséum National Keywords Insecta . Leptoconops . Synchrotron imaging . ’ d Histoire Naturelle, French amber. Cretaceous . Palaeoecology 5 rue Buffon, 75005, Paris, France J. Choufani e-mail: [email protected] Introduction A. Nel e-mail: [email protected] Ceratopogonidae, or biting midges, have their earliest record in the Early Cretaceous (Neocomian, 130–120 Ma) D. Azar amber of Lebanon, but the high diversity already reported Faculty of Sciences II, Department of Natural Sciences, Lebanese from this material, viz. 24 species in eight genera as listed University, FanarP.O. Box 26110217Fanar - Matn, Lebanon in the world catalogue of Borkent and Wirth (1997), with its updated online version by Borkent (2011), suggests an D. Azar e-mail: [email protected] earlier origin of the family in the earliest Cretaceous or even the Late Jurassic. Simulidium priscum Westwood V. Perrichot (*) 1854, from the Late Jurassic Purbeck Formation CNRS UMR 6118 Géosciences, Université de Rennes 1, Campus (−150 Ma) of England, was suggested by Grogan and de Beaulieu bât. 15, 263 avenue du Général Leclerc, Szadziewski (1988) to be a species of Leptoconops Skuse 35042, Rennes cedex, France 1889 but is actually a brachyceran, not a ceratopogonid e-mail: [email protected] (Borkent 1995). Based on the antiquity of Lebanese amber, one could expect to find only extinct genera; however, C. Soriano : P. Tafforeau European Synchrotron Radiation Facility, Leptoconops (subgenus Palaeoconops Borkent 2001) and 6 rue Jules Horowitz, BP 220, 38043, Grenoble cedex, France Austroconops Wirth and Lee 1958 are also present and C. Soriano diversified, making Leptoconops+Austroconops the earliest e-mail: [email protected] extant lineage of biting midges. The biology, distribution, P. Tafforeau and fossil record of Leptoconops were discussed mostly by e-mail: [email protected] Wirth and Atchley (1973) and Borkent (1995, 2000). The Author's personal copy

Palaeobio Palaeoenv genus now comprises 150 extant species mostly distributed to generate all the 3D renderings. All the synchrotron in intertropical or subtropical regions throughout the world, microtomographic data (original stacks of slices, segmen- and 15 extinct species ranging from the Early Cretaceous to tation, animations, and pictures) are available at the ESRF the Late Eocene (from Borkent's 2011 online catalogue). paleontological online database (http://paleo.esrf.eu), and Here, we report the discovery of a new species of 3D models in ABS plastic are deposited with the paratype Leptoconops from the mid-Cretaceous Charentese amber, and other specimens in University of Rennes 1, in the in the Aquitaine Basin. Although Ceratopogonidae are MNHN amber collection (Division of Paleontology), and at abundant in French Cretaceous amber deposits (Perrichot et the ESRF. al. 2007), only three species have yet been formally described (Atriculoides cenomanensis Szadziewski and Schlüter 1992; A. incompletus Szadziewski and Schlüter Systematic palaeontology 1992; Austroconops borkenti Szadziewski and Schlüter 1992) and a poorly preserved, unnamed Leptoconops Order Diptera Linnaeus, 1758 reported (Schlüter 1978: Ceratopogonidae species D; Family Ceratopogonidae Newman, 1834 Szadziewski and Schlüter 1992: Leptoconops sp. indet.) Subfamily Leptoconopinae Noè, 1907 from Cenomanian amber of the Paris Basin. Genus Leptoconops Skuse, 1889

Leptoconops daugeroni Choufani, Azar and Nel, sp. nov. Materials and methods Ty p e s p e c i e s : Holotype IGR.ARC-182.3 (a female in The new species is based on five specimens preserved in hyaline amber), paratypes IGR.ARC-182.2, IGR.ARC- amber from the Font-de-Benon quarry between the villages 237.10, IGR.ARC-333.1 (three females in hyaline amber), of Archingeay and Les Nouillers, in Charente-Maritime, and IGR.ARC-392.1 (scan ESRF A-020_d; a female in south-western France. Two amber-bearing strata are found opaque amber); in amber collection of the Department of in this quarry, the older of which (stratigraphic level A1sl- Geosciences of the University Rennes 1, France. The A) provided the pieces considered here and is dated to latest plastic cast of the paratype IGR.ARC-392.1 which is stored Albian (Néraudeau et al. 2002). Details on the age, in the collection of the Laboratoire de Paléontologie, geological settings, and palaeobiota of this deposit are Muséum National d’Histoire Naturelle, Paris, was assigned reviewed in Perrichot et al. (2010). the reference MNHN.F.A39479. The holotype and three paratypes are preserved in Type locality: Font-de-Benon quarry, 1 km East of relatively hyaline amber and were studied using a stereo- Archingeay, Charente-Maritime, France. microscope under transmitted and reflected light. A fourth Stratigraphic horizon: Cretaceous, uppermost Albian, paratype was first detected during a large survey of opaque amber level A1sl-A sensu Perrichot et al. (2010). Charentese amber using propagation phase contrast X-ray Etymology: Named after our friend and colleague Dr. synchrotron microradiography, and then three- Christophe Daugeron, dipterist at the MNHN, Paris. dimensionally imaged using microtomography (PPC- Diagnosis: Female with 12 flagellomeres; flagellomeres SRμCT), following the protocol detailed by Lak et al. 2–11 spherical; terminal flagellomere moderately elongate, (2008). These two experiments were performed on the 3.44.5× as long as flagellomere 11; lamellar cerci elongate, Beamline ID19 of the European Synchrotron Radiation with two long apical setae; tarsal claws strongly curved Facility (ESRF) at Grenoble, France. The specimen was basally, with thick, well-developed inner tooth; ratio length of scanned in two parts using an isotropic voxel size of body/length of wing: 2.35. 1.4 μm. The beam was monochromatized using a multi- Note: The specimen IGR.ARC-392.1 reconstructed by layer (Ru/B4C) monochromator set at 30 keV. The microtomography is best for detailed examination of all propagation distance used to obtain well-adapted phase structures except for pilosity which could not be recon- contrast was set at 300 mm. The size of the amber block structed. It is thus treated below as the paratype 1 (p1) and being far larger than the tomographic field of view, we used the specimen with the best visible pilosity, i.e. IGR.ARC- local microtomography with 1,500 projections of 0.3 s each 182.3, is treated as the holotype. Other specimens IGR. on 180 degrees in continuous acquisition mode (Lak et al. ARC-237.10, IGR.ARC-182.2, and IGR.ARC-333.1 are 2008). After tomographic reconstruction, ring artefact treated as paratypes 2, 3, and 4 (p2, p3, p4), respectively. correction and 16 bits conversion, the volume was The description and measurements below are based on the segmented in 3D using the VGStudioMax 2.1 software holotype (Figs. 1a and 3c) unless otherwise noted. (Volume Graphics, Heidelberg, Germany), in order to Description: Total body length between 0.86 and virtually extract the specimen from the opaque amber and 1.41 mm. Head wider than high and long; eyes large, Author's personal copy

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Fig. 1 Assorted females of Leptoconops daugeroni sp. nov., in Albian c paratype IGR.ARC-237.10, habitus in right lateral view; d paratype Charentese amber. a Holotype IGR.ARC-182.3, habitus in left lateral IGR.ARC-182.2, habitus in left lateral view. Scale bars 0.25 mm view; b paratype IGR.ARC-333.1; habitus in right lateral view; separated by the equivalent of two ommatidia, in lateral Thorax scutum shining; sparse short pilosity on thorax; view covering most of the head except the posterior part prescutal pits each with two separated round areas. of vertex, four long setae visible on vertex; antenna 13- Wing less than twice as long as wide (length/width segmented; flagellum relatively long, 330 μminp1 values in mm: 0.59/0.35 in holotype, 0.61/0.35 in p1, 0.61/ (292 μm in p2, 265 μminp3,315μminp4);pedicel 0.32 in p2, 0.62/0.24 in p3, 0.59/0.34 in p4); radial cells 1.5× broader than long; flagellomeres 2–11 more or less very short, R1, R3 joining costa in thickened pterostigma; spherical, diameter 21.7 μm (ca. 17 μmforp2andp3); crossvein r–m absent; vein R4+5 reaching wing apex; terminal flagellomere 75 μm long (65 μm in p1 and p4, median veins short; intercal vein not running close to M 77 μm in p3, ), 3.4× as long as penultimate (4.5× in p2 (Fig. 3a). and p3); proboscis relatively short, about half length of Legs slender, all with femur and respective tarsus head (as long as head in p2); palpus four-segmented; subequal in length, barely longer than respective tibia; third palpal segment long, broad (length/width values in midleg tibia with a row of ca. 20 strong setae; hind leg μm: 66.7/25 μm in holotype, 53.8/20.6 μm in p1, 34/11 tibia with a distinct but straight apical spur; all in p3), with a long apical seta; fourth segment slender tarsomeres with thick spines, a few on basitarsomere, (length in μm:41.7inholotype,38.5inp1,23.2inp3); twoontarsomeresII–IV, and three on tarsomere V; tarsal labrum with six short thick apical spines; details of clypeus, claws strongly curved basally, with thick, well-developed mandibles and lacinia not visible. inner tooth (Fig. 3b). Author's personal copy

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Abdomen generally narrow (Figs. 1a–d), at least 3× as Discussion long as wide (excluding cerci), except for p1 dilated either by putrefaction or maybe by presence of blood inside, 1.6× These five female specimens are nearly identical in shape as long as wide (Fig. 2); spermathecae not visible; details of and proportion, especially of the long cerci with two long sternite 8, sternite 9, segment 10 not visible; cercus lamellar, apical setae, and general proportions of flagellomeres, 0.21 mm long in holotype (0.17 mm in p1, 0.14 mm in p2, except for terminal flagellomere which is 4.5 times as long 0.15 mm in p4), covered with very short setae throughout as the preceding one in IGR.ARC-237.10 and IGR.ARC- and with a series of long setae along dorsal and ventral 182.2 instead of 3.4 times in the others. This difference is margins and two longer apical setae (Fig. 3c). likely an intraspecific variation, however, and since they all

Fig. 2 Leptoconops daugeroni sp. nov., paratype female IGR.ARC-392.1 in Albian Charentese amber. Synchrotron microtromographic 3D rendering, 1.4 μm voxel size. a dorsal view; b ventral view; c left lateral view; d frontal view; e posterior view. Scale bar 500 μm Author's personal copy

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be decided on the scanned paratype because the method of reconstruction using PPC-SRμCT has led, in this particular case, to the ‘virtual elimination’ of the pilosity. This technique does allow for a precise imaging of pilosity (see Solórzano-Kraemer et al. 2011), but it would have required a second scan at higher resolution that was not considered important with regard to the additional material available in transparent amber. Fossileptoconops has similar very long setae but nearly as long as the cerci (autapomorphy after Borkent and Craig 2004). Leptoconops daugeroni also differs from Fossileptoconops in several characters, viz. the apomorphies of Leptoconops listed above, plus 12 flagellomeres instead of 13 in the female antenna, and long female cerci. Within the genus Leptoconops, the presence of 12 Fig. 3 Detailed structures of Leptoconops daugeroni sp. nov., in flagellomeres excludes affinities with the Lower Cretaceous Albian Charentese amber. a Wing, paratype female IGR.ARC-333.1; b subgenus Palaeoconops (L. amplificatus Borkent 2001 and midleg claws, paratype female IGR.ARC-333.1; c cercus in right L. antiquus Borkent 2001 from Lebanese amber). Borkent lateral view, holotype female IGR.ARC-182.3 (1995) proposed the only available phylogenetic analysis of the six subgenera known at this time. This author come from the same outcrop and IGR.ARC-182.2 was even considered that the female cerci elongate and laterally preserved in the same piece of amber as the holotype, we compressed is the synapomorphy of the group of subgenera consider it likely that they belong to the same species. [Holoconops+(Megaconops+Leptoconops s. str. + Prolep- In the phylogenetic analysis of the Ceratopogonidae toconops)], excluding affinities of L. daugeroni with the proposed by Borkent and Craig (2004), the Leptoconopinae two other subgenera Styloconops and Brachyconops. The resulted in six genera: Archiaustroconops Szadziewski lack of information on the sensillae and sensory pits of the 1996, Austroconops, Fossileptoconops Szadziewski 1996, palpomeres prevents a precise placement of L. daugeroni as Jordanoconops Szadziewski 2000, Leptoconops, and Min- compared to Borkent’s work. Nevertheless, affinities with yohelea Borkent 1995. The subfamily is supported by two Holoconops can be excluded by the presence of 12 synapomorphies, one of which applies only to males (and flagellomeres instead of 10–11 (Szadziewski and Arillo could not be tested here) and the other which is the 2003; Wirth and Atchley 1973). This character and the claw presence of four palpal segments in male and female adults with a basal tooth merely suggest it belongs to the subgenus with the character reversed in some fossil species of Leptoconops but the main features of this subgenus are Austroconops and Archiaustroconops. Leptoconops dauger- plesiomorphic. oni has the two synapomorphies supporting the clade Comparison with other Cretaceous Leptoconops species (Leptoconops+Minyohelea), i.e. ‘radial cells very short’ indicates the presence of long apical setae on the female and ‘female wing with R1 and R3 joining costa in a cerci as sufficient to characterise L. daugeroni, but this thickened pterostigma’. It also has the following synapo- character is undetermined for some other fossil species. We morphies of Leptoconops: based our comparison on the key to Mesozoic species of 1) ‘wing without r–m’. Borkent and Craig (2004) indicated Borkent (2001),amendedbySzadziewskiandArillo that this character is ‘unique within at least the Culicomorpha’; (2003). It would fall near L. curvachelus Borkent 1996, L. 2) ‘female cerci elongate and laterally compressed’. sibiricus Szadziewski 1996, and L. myanmaricus Szad- Borkent and Craig (2004) concluded that this character is ziewski 2004 for the antenna with 12 flagellomeres and the reversed in the two recent subgenera Brachyconops Wirth tarsal claws strongly curved basally, with thick, well- and Atchley 1973 and Styloconops Kieffer 1921, based on developed inner tooth. The spherical distal flagellomeres, the presence of elongate cerci in the oldest known representa- plus the rather short, 65–75 μm long, terminal flagellomere tives of the genus Leptoconops in Lebanese amber that separate L. daugeroni from L. sibiricus (in which the Borkent (2001) considered in a separate subgenus Palae- flagellomeres are increasingly elongate and the terminal oconops as sister group to all other subgenera. one 108 μm long). In L. curvachelus, the terminal Leptoconops daugeroni has two long setae apically on flagellomere is 3 times as long as the preceding one the female cerci, although theses are shorter than the cerci (Borkent 1996: fig. 2a). themselves. This character is most clearly visible on the Furthermore, the presence of spherical flagellomeres 2– holotype and on specimen IGR.ARC-237.10, but it cannot 11 of L. daugeroni excludes affinities with L. copiosus Author's personal copy

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Borkent 1996 (Borkent 1996:fig.1k;Borkent2001; littoral to lacustrine environments and intermediate Szadziewski and Arillo 2003). In L. primaevus Borkent mangrove-like areas, under a warm and wet subtropical 1995, only the flagellomeres 7–11 are spherical while the climate but with probable arid seasons”, with several flagellomere 2 is longer than wide, unlike in L. daugeroni arthropods typical of sandy sea shore (Perrichot and Girard (Borkent 1995:30–31, fig. 1f). The wing length (ca. 2009; Perrichot et al. 2010) and obvious marine influence 0.60 mm) and the ratio length of body/length of wing (Girard et al. 2008, 2009). Vertebrates present in this amber (2.35) of L. daugeroni separates it from L. boreus Kalugina forest ecosystem, that may have served as hosts for the 1991 (wing length 0.82 mm, ratio 1.95) (Szadziewski female Leptoconops, include squamates (Perrichot and 1996). Leptoconops zherikhini Szadziewski and Arillo 2003 Néraudeau 2005), non-avian theropod dinosaurs or birds also differs from L. daugeroni in the flagellomeres 2 and 3 (Perrichot et al. 2008; Vullo et al. 2007; Vullo and longer than broad, as well as the structure of the tarsal Néraudeau 2010), and mammals (Vullo et al. 2009, 2010). claws. Leptoconops nosopheris Poinar 2008 has small The substrate and its food web, as reconstructed for this simple tarsal claws, its terminal flagellomere 2.9 times as ecosystem with a variety of microalgae, fungi, and bacteria long as flagellomere 11, instead of 3.4–4.5 times in L. (Adl et al. 2011), were likely suitable as breeding sites for daugeroni, and very long cerci, 0.62 mm instead of 0.14– the immature stages. 0.21 mm in L. daugeroni (Poinar 2008). L. burmiticus Szadziewski 2004 and L. rossi Szadziewski 2004 also have Acknowledgements We are grateful to Ryszard Szadziewski, Art small simple tarsal claws and short cerci. Leptoconops Borkent and Vincent Girard for critical comments on the manuscript; to the Marchand family who generously permitted us to collect the clava Borkent 1997 (amber of Hungary, Upper Cretaceous, amber on their property and provided field assistance; and to Didier 80–90 Mya) and L. subrossicus Szadziewski and Poinar Néraudeau for his active participation in collecting and studying the 2005 (Burmese amber, Upper Albian, 100 Mya) are known amber. Partial support for this work was provided by the French only by males and are therefore hardly comparable to the National Research Agency grant BLAN07-1-184190 (project AMBRACE to D. Néraudeau, Université Rennes 1), and by the female specimen of L. daugeroni. The apex of vein R3 is in ESRF through attribution of inhouse beamtime on the beamline ID19. a more distal position in L. clava than in L. daugeroni This is a contribution to the project ‘Novambre’ (to D. Néraudeau) in (Borkent 1997: fig. 2g, h). The hind tibia of L. subrossicus the program INTERRVIE from the French National Institute for ‘ has a strongly curved apical spur, instead of being shorter Universe Sciences, and to the project Biodiversity: Origin, Structure, Evolution, and Geology’ allotted to D.A. by the Lebanese University. and nearly straight in L. daugeroni. The Baltic amber L. succineus Szadziewski 1988 differs from L. daugeroni in the flagellomeres 7–11 slightly elongate and the wing References longer (0.67–0.73 mm instead of 0.60 mm in L. daugeroni) (Szadziewski 1988). Unfortunately, it was not possible to Adl S, Girard V, Breton G, Lak M, Maharning A, Mills A, Perrichot V, locate the unique specimen of Leptoconops sp. reported Trionnaire M, Vullo R, Néraudeau D (2011) Reconstructing the from French Cenomanian amber of Bezonnais in the Paris soil food web of a 100 million-year-old forest: The case of the Basin (Szadziewski and Schlüter 1992), and no comparison mid-Cretaceous fossils in the amber of Charentes (SW France). Soil Biol Biochem 43:726–735 is possible based on the brief description and illustrations Aussel J-P (1991) Isolement et description de la larve de Leptoconops provided by Schlüter (1978: 100, fig. 83, table 12.4). (Styloconops) albiventris De Meijere, 1915 (Diptera, Ceratopo- gonidae). Rev Fr Entomol (NS) 13:109–112 Borkent A (1995) Biting Midges in the Cretaceous Amber of North America (Diptera: Ceratopogonidae). Backhuys, Leiden Palaeoecological implications Borkent A (1996) Biting midges from Upper Cretaceous of New Jersey amber (Ceratopogonidae, Diptera). Am Mus Novit 3159:1–29 As the new species belongs in the subgenus Leptoconops,it Borkent A (1997) Upper and Lower Cretaceous biting midges is possible to make a rather accurate phylogenetic inference (Ceratopogonidae) from Hungarian and Austrian amber and the Koonwarra fossil bed of Australia. Stuttg Beitr Naturk (Ser B) for its palaeobiology after those of the recent representa- 249:1–10 tives of this genus (Nel 1997). Adult female Leptoconops Borkent A (2000) Biting midges (Ceratopogonidae: Diptera) from are bloodsuckers on vertebrates, and the larvae, which feed Lower Cretaceous Lebanese amber with a discussion of the on algae, fungi, and bacteria, live in moist and usually diversity and patterns found in other ambers. In: Grimaldi D (ed) Studies on fossils in amber, with particular reference to the saline sandy to muddy substrate or in the sand of coastal Cretaceous of New Jersey. Backhuys, Leiden, pp 335–345 and inland beaches (Aussel 1991; Raspi et al. 2007; Rohani Borkent A (2001) Leptoconops (Diptera: Ceratopogonidae), the et al. 2006; Tokunaga 1963), suggesting a similar palae- earliest extant lineage of biting midge, discovered in 120–122 oenvironment for L. daugeroni. This is consistent with the millions-year-old Lebanese amber. Am Mus Novit 3328:1–11 Borkent A (2011) World Species of Biting Midges (Diptera: Charentese amber forest ecosystem as reconstructed from Ceratopogonidae). Online catalogue updated February 10, avail- previous studies: Perrichot et al. (2007) proposed “a mosaic able at www.inhs.illinois.edu/research/FLYTREE/Ceratopogoni- of coastal habitats with peculiar plant associations, from daeCatalog.pdf Author's personal copy

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