True Flies (Insecta: Diptera) from The

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Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 1–60, 2019

True flies (Insecta: Diptera) from the late Eocene insect limestone (Bembridge Marls) of the Isle of Wight, England, UK Wiesław KRZEMIN´ SKI1*, Vladimir BLAGODEROV2, Dany AZAR3, Elena LUKASHEVICH4, Ryszard SZADZIEWSKI5, Sonja WEDMANN6, Andre´ NEL7, François-Marie COLLOMB7, Alain WALLER7 and David B. NICHOLSON8 1 Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Ul. Sławkowska 17, 31-016 Krako´w, Poland. Email: [email protected] 2 Department of Natural Sciences, National Museums Scotland, Chambers St., Edinburgh EH1 1JF, UK. 3 Department of Biology, Faculty of Sciences II, Lebanese University, Fanar – Matn, P. O. box 26110217, Lebanon. 4 Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya Str. 123, 117868 Moscow, Russia. 5 Department of Invertebrate Zoology, University of Gdan´sk, Pilsudskiego 46, 81-378 Gdynia, Poland. 6 Senckenberg Research Institute and Museum, Research station Grube Messel, Markstraße 35, D-64409 Messel, Germany. 7 CNRS UMR 5202, Museúm National d’Histoire Naturelle, CP 50, Entomologie, 45 rue Buffon, F-75005, Paris, France. 8 School of Earth and Environmental Sciences, University of Portsmouth, Burnaby Road, Portsmouth PO1 3QL, UK. *Corresponding author.

ABSTRACT: The Diptera fauna from the late Eocene of the Isle of Wight (Bembridge Marls) is studied including redescriptions of formerly described material. The fauna includes the following taxa: Anisopodidae – one species; Bibionidae – 11 species; Ceratopogonidae – one described and two unidentified species; Chironomidae – undetermined species of three subfamilies; Culicidae – four species; Cylindrotomidae – one species; Dixidae – one species; Keroplatidae – one described and four unidentified species; Limoniidae – 31 species; Mycetophilidae – 14 species; Psychodidae – two species; Scatopsidae – two species; Sciaridae – three species; Simuliidae – an unnamed species; Tipulidae – nine species. Diptera Brachycera: Agromyzidae – unidentified species; Anthomyzidae – two species; Asilidae – two species; Bombyliidae – one species; Chloropidae – one species; Dolichopodidae – unidentified species; Empididae – unidentified species; Ephydridae – two species; Heleomyzidae – unidentified species; Lauxaniidae – unidentified species; Otitidae – one species; Phoridae – unidentified species; Rhagionidae – two species; Stratiomyidae – two species; Syrphidae – unidentified species; Tabanidae – one species. Following new taxa are described: Leptotarsus (Longurio) fragmentatus Krzemin´ski, n. sp., Leptotarsus (Longurio) wegiereki Krzemin´ski, n. sp., Tipula (s. lato) anglicana Krzemin´ski, n. sp. (Tipulidae); Cyttaromyia rossi Krzemin´ski, n. sp. (Cylindrotomidae); Gurnardia corami Krzemin´ski, n. gen, n. sp., Pilaria hooleyi Krzemin´ski, n. sp., Pilaria volodii Krzemin´ski, n. sp., Cheilotrichia (Cheilotrichia) duplicata Krzemin´ski, n. sp., Cheilotrichia (Empeda) szwedoi Krzemin´ski, n. sp., Symplecta (Psiloconopa) gurnetensis Krzemin´ski, n. sp., Orimarga (Orimarga) lenae Krzemin´ski, n. sp., Dicranomyia (Dicranomyia) azari Krzemin´ski, n. sp., Dicranomyia (Dicranomyia) fasciata Krzemin´ski, n. sp., Helius (Helius) edmundi Krzemin´ski, n. sp., Helius (Helius) popovi Krzemin´ski, n. sp., Dicranoptycha staryi Krzemin´ski, n. sp. (Limoniidae); Wightipsychoda Azar, n. gen. (Psychodidae); Dilophus andrewrossi Nel, Colomb & Waller, n. sp., Bibiodes massiliensis Nel, Colomb & Waller, n. sp. (Bibionidae); Azana cockerelli Blagoderov, n. sp., Aglaomyia vectis Blagoderov, n. sp., Dziedzickia oligocenica Blagoderov, n. sp., Palaeoempalia saxea Blagoderov, n. sp., Mycomya hoolei Blagoderov, n. sp., Leia gurnardensis Blagoderov, n. sp. (Mycetophilidae); Regmoclemina haennii Krzemin´ski, n. sp. (Scatopsidae); Sylvicola problematica Krzemin´ski, n. sp. (Anisopodidae); Palaeoberidops barkeri Nicholson, n. gen., n. sp. (Stratiomyidae). The Diptera fauna and palaeohabitat of the Insect Bed is discussed.

KEY WORDS: fossil insects, new taxa.

The Diptera from the late Eocene of the Isle of Wight are regions. They are a very common and diversiﬁed group in the similar to the Recent fauna in generic composition, particu- Insect Bed. Most important information about fossil Diptera larly in containing genera occurring today in subtropical from Isle of Wight were published by Cockerell (1915,

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Table 1 Current taxonomic position of the previously described species fossil Diptera form the Insect Limestone.

Name and taxonomic placement as described Valid name and current taxonomic placement Acnemia simplex Cockerell, 1921; Mycetophilidae Mycetophilidae: Sciophilinae: Acnemia simplex Cockerell, 1921 Adelphomyia disjunctula Cockerell & Haines, 1921, p. 83; Limoniidae: Limnophilidae: Pseudolimnophila (Pseudolimnophila) disjunctula Limoniidae (Cockerell & Haines, 1921) Asilus gurnetensis Cockerell, 1921; Asilidae Not revised Atarba vectensis Cockerell, 1915; Limoniidae Limoniidae: Limoniinae: Helius (Helius) vectensis (Cockerell, 1915) Bibio extremus Cockerell, 1921; Bibionidae Bibionidae: ?Bibio extremus Cockerell, 1921 Bibio gurnetensis Cockerell, 1917a; Bibionidae Bibionidae: ?Bibio gurnetensis Cockerell, 1917a Bibio oblitus Cockerell, 1921; Bibionidae Bibionidae: ?Bibio gurnetensis Cockerell, 1917a Bibio oligocenus Cockerell, 1917a; Bibionidae Bibionidae: ?Bibio oligocenus Cockerell, 1917a Bibiodites conﬂuens Cockerell, 1915; Bibionidae Bibionidae: Bibiodes conﬂuens (Cockerell, 1915) Chrysopilus anglicus Cockerell, 1921; Rhagionidae Not revised Chrysopilus stigmaticus Cockerell, 1921; Rhagionidae Not revised Chrysops vectensis Cockerell, 1921; Tabanidae Not revised Culex petrifactellus Cockerell, 1915; Culicidae Culicidae: Ochlerotatus protolepis (Cockerell, 1915) Culex protolepis Cockerell, 1915; Culicidae Culicidae: Ochlerotatus protolepis (Cockerell, 1915) Culex protorhinus Cockerell, 1915; Culicidae Culicidae: Culex protorhinus Cockerell, 1915 Culex vectensis Edwards, 1923; Culicidae Culicidae: Culex vectensis Edwards, 1923 Dicranomyia excavata Cockerell & Haines, 1921; Limoniidae Limoniidae: Limoniinae: Dicranomyia (Dicranomyia) excavata Cockerell & Haines, 1921 Dicranomyia exhumata Cockerell, 1922; Limoniidae Limoniidae: Limoniinae: Dicranomyia (Dicranomyia) excavata Cockerell & Haines, 1921 Dicranomyia undulata Cockerell & Haines, 1921; Limoniidae Limoniidae: Limoniinae: Thaumastoptera undulata (Cockerell & Haines, 1921) Dixa priscula Cockerell, 1921; Dixidae Dixidae: Dixella priscula (Cockerell, 1921) Empeda ferruginea Cockerell, 1921; Limoniidae Limoniidae: Chioneinae: Cheilotrichia (Empeda) ferruginea (Cockerell, 1921) Empeda hyalina Cockerell, 1921; Limoniidae Limoniidae: Chioneinae: Cheilotrichia (Empeda) hyalina (Cockerell, 1921) Ephydra oligocena Cockerell, 1915; Ephydridae Anthomyzidae indet. Ephydra sepulta Cockerell, 1915; Ephydridae Anthomyzidae indet. Epiphragma spilopterum Cockerell, 1921; Limoniidae Limoniidae: Dactyolabinae: Dactylolabis (Dactylolabis) spiloptera (Cockerell, 1921) Gonomyia ferrea Cockerell, 1921; Limoniidae Limoniidae: Chioneinae: Gonomyia (Gonomyia) indecisa Cockerell & Haines, 1921 Gonomyia grisea Cockerell, 1921; Limoniidae Limoniidae: Chioneinae: Cheilotrichia (Empeda) hyalina (Cockerell, 1921) Gonomyia indecisa Cockerell & Haines, 1921; Limoniidae Limoniidae: Chioneinae: Gonomyia (Gonomyia) indecisa Cockerell & Haines, 1921 Gonomyia lutescens Cockerell, 1921; Limoniidae Limoniidae: Chioneinae: Gonomyia (Gonomyia) indecisa Cockerell & Haines, 1921 Gymnastes fasciatipennis Cockerell, 1921; Limoniidae Limoniidae: Chioneinae: Gymnastes (Paragymnastes) fasciatipennis Cockerell, 1921 Hippelates brodiei Cockerell, 1915; Chloropidae Chloropidae: Hippelates brodiei Cockerell, 1915 Holorusia vasifera Cockerell & Haines, 1921, Tipulidae Tipulidae: Ischnotoma (Ischnotoma) vasifera (Cockerell & Haines, 1921) Limnobia spilota Cockerell, 1921; Limoniidae Limoniidae: Limoniinae: Limonia spilota (Cockerell, 1921) Limnophila aliena Cockerell, 1921; Limoniidae Limoniidae: Limoniinae: Dicranomyia (Dicranomyia) aliena (Cockerell, 1921) Limnophila cyclospila Cockerell & Haines, 1921; Limoniidae Limoniidae: Limnophilinae: Limnophila (Limnophila) cyclospila Cockerell & Haines, 1921 Limnophila deleta Cockerell, 1921; Limoniidae Limoniidae: Limnophilinae: Limnophila (Limnophila) deleta Cockerell, 1921 Limnophila incognita Cockerell, 1921; Limoniidae Limoniidae: Chioneinae: Ilisia (Lunaria) incognita (Cockerell, 1921) Macromastix cladoptera Cockerell & Haines, 1921; Tipulidae Tipulidae: Leptotarsus (Macromastix) cladoptera (Cockerell & Haines, 1921) Megistocera gurnetensis Cockerell, 1921; Tipulidae Tipulidae: Brachypremna gurnetensis (Cockerell, 1921) Mesomyites concinna Cockerell, 1917a; Limoniidae Limoniidae: Chioneinae: Styringomyia concinna (Cockerell, 1917a) Mongoma cruciferella Cockerell, 1917a; Limoniidae Limoniidae: Limoniinae: Trentepohlia (Mongoma) cruciferella (Cockerell, 1917a) Mongoma pallescens Cockerell, 1921; Limoniidae Limoniidae: Limoniinae: Trentepohlia (Mongoma) pallescens (Cockerell, 1921) Mycetophila vectensis Cockerell, 1915; Mycetophilidae Mycetophilidae: Mycetophilinae indet. Mycomya oblita Cockerell, 1921; Mycetophilidae Mycetophilidae: Mycomya oblita Cockerell, 1921 Palpomyia edwardsi Cockerell, 1921; Ceratopogonidae Ceratopogonidae: Palpomyia edwardsi Cockerell, 1921 Paltostomopsis ciliata Cockerell, 1915; Blephariceridae Tipulomorpha indet. Phronia virgata Cockerell, 1921; Mycetophilidae Mycetophilidae: Gnoristinae: ?Coelosia virgata (Cockerell, 1921)

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Table 1 (Continued)

Name and taxonomic placement as described Valid name and current taxonomic placement Platyura obliqua Cockerell, 1921; Mycetophilidae Keroplatidae: Macrocerinae: Macrocera obliqua (Cockerell, 1921) Plecia acourti Cockerell, 1921; Bibionidae Bibionidae: Plecia acourti Cockerell, 1921 Proctacanthus fractus Cockerell, 1921; Asilidae Not revised Protoberis obliterata Cockerell, 1915; Stratiomyidae Bibionidae: Bibio obliteratus (Cockerell, 1915) Protoscinis perparvus Cockerell, 1917a; Chloropidae Ephydridae: Ephydrinae: Scatellini: Protoscinis perparvus Cockerell, 1917a Psychoda leucospila Cockerell, 1921; Psychodidae Psychodidae: Wightipsychoda leucospila (Cockerell, 1921) Psychoda primaeva Cockerell, 1915; Psychodidae Psychodidae: Pericoma primaeva (Cockerell, 1915) Rhipidia brodiei Cockerell, 1917a; Limoniidae Limoniidae: Limoniinae: Dicranomyia (Dicranomyia) brodiei (Cockerell, 1917a) Rhyphys hooleyi Cockerell, 1921; Anisopodidae. Anisopodidae: Sylvicola hooleyi (Cockerell, 1921) ‘Rymosia’ edwardsi Cockerell, 1921; Mycetophilidae Mycetophilidae: Mycetophilinae: Mycetophilini indet. ‘Rymosia’ ferruginea Cockerell, 1921; Mycetophilidae Mycetophilidae: Mycetophilinae Mycetophilini indet. Rymosia grisea Cockerell, 1921; Mycetophilidae Mycetophilidae: Mycetophilinae indet. ‘Rymosia’ rufescens Cockerell, 1921; Mycetophilidae Mycetophilidae: Mycetophilinae: Mycetophilini indet. Sciara gurnetensis Cockerell, 1915; Sciaridae Not revised Sciara lacoei Cockerell, 1915; Sciaridae Not revised Sciara protoberidis Cockerell, 1915; Sciaridae Not revised Sphaerocera sepultula Cockerell, 1915; Borboridae Ephydridae indet. Stenomyites fuscipennis Cockerell, 1915; Otitidae Otitidae Stenomyites fuscipennis Cockerell, 1915 Stratiomys brodiei Cockerell, 1915; Stratiomyidae Stratiomyidae: Odontomyia brodiei (Cockerell, 1915) Styringomyia extensa Cockerell & Haines, 1921; Limoniidae Limoniidae: Chioneinae: Styringomyia concinna (Cockerell, 1917a) Systropus acourti Cockerell, 1921; Bombyliidae Not revised Taeniorhynchus cockerelli Edwards, 1923; Culicidae Culicidae: Coquillettidia cockerelli (Edwards, 1923) Tipula acourti Cockerell, 1921; Tipulidae Tipula (s. lato) acourti Cockerell, 1921 Tipula callarche Cockerell, 1921; Tipulidae Leptotarsus (Macromastix) cladoptera (Cockerell & Haines, 1921) Tipula gardneri Cockerell, 1917a; Tipulidae Tipula (s. lato) gardneri Cockerell, 1917a Tipula hooleyi Cockerell & Haines, 1921; Tipulidae Tipula (s. lato) limiformis Cockerell, 1915 Tipula limiformis Cockerell, 1915; Tipulidae Tipula (s. lato) limiformis Cockerell, 1915

1917a, b, 1921); Cockerell & Haines (1921); Edwards (1923); Insect Bed, which lies towards the base of the Bembridge Jarzembowski (1976); Krzemin´ski (1996); Vane-Wright (1967). Marls Member (Solent Group: Bouldnor Formation) (Gale In years 2005–2007 thanks to financial support of et al. 2006; Ross & Self 2014). INTAS (Project no. 03-51-4367) an international group of The largest collection is housed in Natural History Museum dipterologists, including Wiesław Krzemin´ski (Poland) – (NHM), London. Most of the specimens at this museum team leader, Dany Azar (Lebanon), Vladimir Blagoderow belong to the A’Court Smith (purchased 1877, 1883), P.B. (Russia), Elena Lukashevich (Russia), Andre Nel (France), Brodie (purchased 1898) and R.W. Hooley (purchased 1924) Ryszard Szadziewski (Poland), Sonja Wedmann (Germany), collections. They are labelled ‘Gurnard Bay’ (which is an old Vladimir Zlobin (Russia), had undertaken the task of deter- name for Gurnet Bay); however, Smith collected specimens mination and revision of dipteran materials from the Isle of all the way from West Cowes to Newtown River on the NW Wight. Later Francis-Marie Collomb (France), David B. side of the Isle of Wight (Jarzembowski 1980). Most of the Nicholson (UK) and Alain Waller (France) also joined the specimens probably came from Thorness Bay (Jarzembowski project. 1976). Brodie and Hooley acquired parts of Smith’s collection, Our study on old and new collections housed in UK and so that parts and counterparts of individual insects have USA museums and materials collected during field trips re- turned up in the three collections. The parts and counterparts vealed over 1,700 fossil specimens, belonging to 32 families of often have different numbers because they were registered at the Diptera; however, only around 800 specimens were studied different times. An additional collection was discovered at the for this paper. Taxonomic results of study on higher flies Sedgwick Museum, Cambridge by A.J. Ross. This collection from the Insect Bed are published elsewhere (Zlobin 2007). A has also yielded counterparts of specimens at the NHM, which summary of current taxonomic position of previously described indicates that this is another part of the Smith collection. A species is presented in Table 1. label with ‘1883’ on it suggests that the Sedgwick Museum acquired this collection in 1883, the same year that the NHM purchased specimens from Smith. Prefixes I. and In. preceding the numerals of specimens indi- 1. Material and methods cate that they come from different series. The fossil insects of the Isle of Wight originate from the late Eocene concretions or tabular bands of very fine-grained 1.1. Number of specimens studied micrite, identified as Insect Limestone of the Isle of Wight. Diptera Nematocera: Anisopodidae – five specimens; Bibioni- The unit where these concretions/bands occur is known as the dae – 19 specimens; Ceratopogonidae – nine specimens;

Holotype. NHMUK In.17090/In.17492 (part and counterpart), Smith Collection; Insect Bed; late Eocene; NW Isle of 2. Systematic palaeontology Wight, UK. Order Diptera Linnaeus, 1758 Remarks. Revision and redescription of this species was given by Krzemin´ski (1996). Suborder Nematocera Dumeril, 1805 Genus: Leptotarsus Guerin-Meneville, 1831 Superfamily Tipuloidea Latreille, 1802 by Wieslaw Krzemin´ski Type species. Leptotarsus macquartii Guerin-Meneville, 1831: Atlas: pl. 20, ﬁg. 1 (by monotypy) Family Tipulidae Latreille, 1802 Subgenus: Macromastix Osten Sacken, 1887 TIPULIDAE (Diptera Nematocera) are rather common and well preserved in Bembridge Marls of the Isle of Wight when Type species. Macrothorax ornatus Jaennicke, 1867, p. 320 compared to other contemporary faunas. These large Diptera Remarks. The subgenus Macromastix is represented in fossil are represented mostly by isolated wings or some wing frag- state by two fossil species, from the Oligocene of France and ments. Bodies are rare and mostly poorly preserved. In the from late Eocene Isle of Wight, England. Recently the sub- collections from the Bembridge Marls eight species were genus comprises 94 species living in tropical and subtropical described by Cockerell (1915, 1917a, 1921) and Cockerell & regions.

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Subgenus Longurio Loew, 1869

Type species. Longurio testaceus Loew, 1869 (by monotypy) Remarks. Recently subgenus Longurio comprises 112 species living in tropical and subtropical regions and two fossil species from the Lower Cretaceous from Russia and China (Shih et al. 2015).

Leptotarsus (Longurio) fragmentatus Krzemin´ski, n. sp. (Fig. 3, Pl. 1c)

Holotype. NHMUK In.25379, Hooley Collection; Insect Bed; late Eocene; NW Isle of Wight, UK. Diagnosis. Sc ending in Costa, opposite proximal third portion of R2 þ 3 þ 4; R3 strongly curved to costal wing margin; cross-vein m-cu about 1.25 times longer than d cell. Description. Only a portion of the wing is preserved, but characteristic venation of radial sector allows to separate it from other Tipulidae from the Isle of Wight. Wing clear, 18– 19 mm long. Venation: Sc long, ending in the costal vein, opposite 1/3 of R2 þ 3 þ 4; cross-vein sc-r at its three lengths before tip of Sc; R3 short, and strongly curved to costal wing Figures 1–4 Tipulidae, wing venation. (1) Brachypremna gurnetensis margin; Rs 1.5 times longer than R2 þ 3 þ 4; d cell long and (Cockerell, 1921), holotype NHMUK In.17090. (2) Leptotarsus narrow; petiole (distal section of M1 þ 2) very short; upper Macromastix cladoptera ( ) (Cockerell & Haines, 1921), holotype margin of d cell four times longer than petiole; M1 about ﬁve NHMUK I.10379. (3) Leptotarsus (Longurio) fragmentatus Krzemin´ski, n. sp., holotype NHMUK In.25379. (4) Leptotarsus (Longurio) times longer than petiole; cross-vein m-cu very long and about wegiereki Krzemin´ski, n. sp., holotype NHMUK In.17493. 1.25 times longer than d cell.

Leptotarsus (Longurio) wegiereki Krzemin´ski, n. sp. (Fig. 4, Pl. 7a) Leptotarsus (Macromastix) cladoptera (Cockerell & Haines, 1921) Etymology. A new species is dedicated to Piotr We˛gierek (Fig. 2, Pl. 1b) (Silesian University, Katowice, Poland) a paleoentomologist, specialist of Homoptera. 1921 Macromastix cladoptera Cockerell & Haines, p. 111– Holotype. NHMUK In.17493, Smith Collection; Insect 112, ﬁg. I10397 (published May 1921) Bed; late Eocene; NW Isle of Wight, UK. 1921 Tipula callarche Cockerell p. 455, ﬁg. 1 (published June Other material. CAMSM X.50140 (TN 76), CAMSM 1921), syn. nov. X.50140.89a (TN 141), Smith Collection; NHMUK In.25207, 1994 Leptotarsus (Macromastix) cladoptera Evenhuis, p. 56 Hooley Collection. 1994 Tipula callarche Evenhuis, p. 58 Diagnosis. Sc ending in Costa, opposite Rs fork; d cell very small; cross-vein m-cu long, about three times longer than Holotype. NHMUK I.10397(part), Brodie Collection, de- width of d cell. scribed as Macromastix cladoptera Cockerell & Haines, 1921 Description. Wing length 18–19 mm, without pattern, but and NHMUK In.17381(counterpart), Smith Collection (de- anal part is destroyed in all specimens. Venation: Sc ending scribed as Tipula callarche Cockerell, 1921); Insect Bed; late in Costal vein, opposite to fork of Rs; cross-vein sc-r at its Eocene; NW Isle of Wight, UK. three lengths before tip of Sc; R2 þ 3 þ 4 1.5 times shorter Diagnosis. Wing wide, narrowing to petiolate basally, anal than Rs; R4 twice longer than R2 þ 3 þ 4; d cell very small; lobe very narrow; A2 very close and parallel to anal wing petiole of M1 þ 2 twice longer than preceding section between margin; colour pattern on wing membrane. cross-veins r-m and m-m; M1 about twice longer than M1 þ 2; Redescription. Two wings (one incomplete) and small cross-vein m-cu 2.5 times longer than width of d cell. portion of thorax are preserved. Wing about 9.5 mm long, Remarks. Venation pattern in this new species is unique by with a well visible colour pattern. Venation: tip of Sc lost, Sc being complete (tip not atrophied) and ending in the Costa, and this vein together with sc-r terminates in Rl, opposite small d cell, and long cross-vein m-cu. All these characters are mid of Rs; R3 0.5 times length of R3 þ 4; Rs little shorter only exceptionally met in the Tipulidae, such as some Recent than R3 þ 4; petiole (most distal section) of Ml þ 2 as long as species of the genus Leptotarsus. M2 and equal preceding section between cross-veins r-m and m-m; cross-vein m-cu in fork of M3 þ 4; A2 close and parallel Genus: Tipula Linnaeus, 1758 to anal wing margin; basal part of wing is narrow and almost without anal lobe. Type species. Tipula oleracea Linnaeus, 1758, p. 585 Remarks. Macromastix cladoptera was described by Cockerell (designated by Latreille 1810) & Haines in May 1921 from one part of the specimen. One Remarks. The genus Tipula (Diptera Nematocera) is very month later, in the same year, Cockerell described another diverse; currently about 1,200 Recent species in 34 subgenera

Plate 1 Tipulidae and Cylindrotomidae. (a) Brachypremna gurnetensis (Cockerell, 1921) holotype part NHMUK In.17090. (b) Leptotarsus (Macromastix) cladoptera (Cockerell & Haines, 1921), holotype part NHMUK I.10397. (c) Leptotarsus (Longurio) fragmentatus Krzemin´ski, n. sp., holotype NHMUK In.25379. (d) Tipula (s. lato) acourti Cockerell, 1921, holotype part NHMUK I.8901. (e) Tipula (s. lato) anglicana Krzemin´ski, n. sp., holotype NHMUK In.64117. (f ) Tipula (s. lato) gardneri Cockerell, 1917a holotype NHMUK I.8663. (g, h) Tipula (s. lato) limiformis Cockerell, 1915: (g) (type of Tipula hooleyi Cockerell & Haines, 1921 syn. nov.), NHMUK In.24353; (h) holotype USNM 61438. (i) Ischnotoma (Ischnotoma) vasifera (Cockerell & Haines, 1921), holotype NHMUK In.24352. (j) Cyttaromyia rossi Krzemin´ski, n. sp., holotype CAMSM X.50140.14.

Figures 5–13 Tipulidae and Cylindrotomidae, wing venation. (5) Tipula (s. lato) acourti Cockerell, 1921, holotype NHMUK I.8901. (6) Tipula (s. lato) anglicana Krzemin´ski, n. sp. holotype NHMUK In.64117. (7) Tipula (s. lato) gardneri Cockerell, 1917a, holotype NHMUK I.8663. (8), (9) Tipula (s. lato) limiformis Cockerell, 1915: (8) – (type of Tipula hooleyi Cockerell & Haines, 1921 syn. nov.), NHMUK In.24353; (9) Holotype of Tipula limiformis USNM 61438. (10) Ischnotoma (Ischnotoma) vasifera (Cockerell & Haines, 1921), holotype NHMUK In.24352. (11)–(13) Cyttaromyia rossi Krzemin´ski, n. sp. – wing venation; holotype CAMSM X.50140.14 (TN 63): (11) left wing; (12) right wing; (13) reconstruction based on both wings.

Holotype. NHMUK In.24352/I.8852 (part and counterpart), known only from nine fossil species from the Palaeogene of Hooley/Brodie Collections; Insect Bed; late Eocene; NW Isle of North America (Scudder 1877, 1894; Cockerell 1924, 1925), Wight, UK. Denmark (Freiwald 1991), France (Seguy 1934) and Russia Other material. Besides holotype, specimens NHMUK (Freiwald & Krzemin´ski 1991) and one species from Baltic In.25210, In.25260/In.25493 (part and counterpart). amber (Krzemin´ski 1998). Diagnosis. Vein R4 strongly curved to upper wing margin; Recent Cylindrotomidae are a small family comprising 72 m-cu placed between fork of Mb and fork M3 þ 4. species inhabiting mainly Holartic and Oriental Regions. In Redescription. Wing large, about 23 mm long. Venation: Sc the Afrotropical Region the family is not represented. ending in C; sc-r at its two lengths before tip of Sc; R2 þ 3 þ 4 In the fossil state only 14 fossil species of the family have as long as R3; Rs 1.5 times longer than R2 þ 3 þ 4; R4 in been described so far (Cockerell 1920; Freiwald & Krzemin´ski distal part strongly curved to upper wing margin; d cell rather 1991; Evenhuis 1994; Krzemin´ski 1998). small; distal M1 þ 2 very short; mid section of M1 þ 2is2.5 times longer than distal; cross-vein m-cu placed between fork Genus Cyttaromyia Scudder, 1877 of Mb and fork M3 þ 4. Remarks. Holorusia vasifera described by Cockerell & Type species. Cyttaromyia fenestrata Scudder, 1877, by Haines (1921) was transferred by Vane-Wright (1967) to the monotypy genus and subgenus Ischnotoma Skuse, 1890 on the basis of similarity in wing venation. The species is particularly very similar to the Australian Ischnotoma (Ischnotoma) par (Walker Cyttaromyia rossi Krzemin´ski, n. sp. 1856). (Figs 11–13, Pl. 1j) Conclusion. Generally Tipulidae from the Isle of Wight are indicative of typical warm climate. Recent species from genera Derivation of name. The name of species is dedicated to Brachypremna, Macromastix, Leptotarsus and Ischnotoma live Andrew Ross, with many thanks for his efﬁcient help during in tropical and subtropical regions. all stages of our project. Holotype. CAMSM X.50140.14 (TN 63) Smith Collection; Insect Bed; late Eocene; NW Isle of Wight, UK. Family: Cylindrotomidae Schiner, 1863 Diagnosis. R1 terminating in R3 þ 4 (a generic character); R1 fused with r-r (R2) and terminating in R3 þ 4 in 1/3 of its Cylindrotomidae were until now unknown in the fossil material length; M3 as long as d cell; cross-vein m-cu distinctly before from the late Eocene of the Isle of Wight. There is one specimen fork of Mb. belonging to the fossil genus Cyttaromyia Scudder. This genus Description. Only two wings of the holotype are preserved; has a characteristic wing venation and until now has been body absent. Wing length about 9 mm, with distinct pterostigma.

Figures 14–25 Limoniidae, wing venation. (14) Gurnardia corami Krzemin´ski, n. sp., holotype NHMUK In.64104. (15) Limnophila (Limnophila) cyclospila Cockerell & Haines, 1921, holotype NHMUK I.9213. (16) Limnophila (Limnophila) deleta Cockerell, 1921, holotype NHMUK I.9053. (17) Pilaria hooleyi Krzemin´ski, n. sp., holotype, NHMUK In.25254. (18) Pilaria volodii Krzemin´ski, n. sp., holotype NHMUK I.9167. (19) Pseudolimnophila (Pseudolimnophila) disjunctula (Cockerell & Haines, 1921), holotype NHMUK. In.24355. (20) Dactylolabis (Dactylolabis) spiloptera (Cockerell, 1921) n. comb., holotype NHMUK I.9015. (21) Cheilotrichia (Cheilotrichia) duplicata Krzemin´ski, n. sp., holotype NHMUK II 2763. (22) Cheilotrichia (Empeda) ferruginea (Cockerell, 1921), holotype NHMUK I.8963. (23) Cheilotrichia (Empeda) hyalina (Cockerell, 1921), holotype NHMUK I.24342. (24) Cheilotrichia (Empeda) szwedoi Krzemin´ski, n. sp., holotype NHMUK In.17082. (25) Gonomyia (Gonomyia) indecisa Cockerell & Haines, 1921, holotype NHMUK. I.9087.

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Plate 2 Limoniidae. (a) Gurnardia corami Krzemin´ski, n. sp., holotype NHMUK In.64104. (b) Limnophila (Limnophila) cyclospila Cockerell, 1921, holotype NHMUK I.9213. (c) Limnophila (Limnophila) deleta Cockerell, 1921, holotype NHMUK I.9053. (d) Pilaria hooleyi Krzemin´ski, n. sp., holotype NHMUK In.25254. (e) Pilaria volodii Krzemin´ski, n. sp., holotype NHMUK I.9167. (f ) Pseudolimnophila (Pseudolimnophila) disjunctula (Cockerell & Haines, 1921) n. comb., holotype NHMUK In.24355. (g) Dactylolabis (Dactylolabis) spiloptera (Cockerell, 1921) n. comb., holotype NHMUK I.9015. (h) Cheilotrichia (Cheilotrichia) duplicata Krzemin´ski, n. sp., holotype NHMUK II 2763. (i) Cheilotrichia (Empeda) ferruginea (Cockerell, 1921), holotype part NHMUK I.8963. (j) Cheilotrichia (Empeda) hyalina (Cockerell, 1921), holotype NHMUK In.24342.

Downloaded from https://www.cambridge.org/core. Chalmers Tekniska Högskola, on 06 Oct 2019 at 16:48:14, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S1755691018000464 12 WIESŁAW KRZEMIN´ SKI ET AL. Holotype. NHMUK I.9213 (female), Hooley Collection), Genus Pilaria Sintensis, 1889 Insect Bed; late Eocene; NW Isle of Wight, UK. Additional material. NHMUK I.9059, I.9868, I.9943, Brodie Type species. Limnobia pilicornis Zetterstedt, 1851, p. 3885 Collection; In.17491 (female), Smith Collection; In.24775, (designated by Sharp 1890). In.25209, In.25269, In.25270, Hooley Collection. Remarks. Forty-six recent species are known from Palaearctic Diagnosis. Wing with conspicuous colour pattern; Sc ending (15 sp.), Nearctic (14 sp.), Afrotropic (8), Oriental (7 sp.), a little before fork of Rs into R2 þ 3 þ 4andR5;R2þ 3 þ 4 Australian (1 sp.) and Neotropic (1 sp.) Regions. Five fossil very short; R4 16 times longer than R2 þ 3 þ 4; M1 long, 1.5 species are known from Eocene and Miocene of Europe (three times longer than d cell; cross-vein m-cu in half of d cell. from the Baltic amber). Redescription. Wing length 7.0–7.8 mm, with very distinct colour pattern (Fig. 15). Body is preserved partially only in two specimens (holotype I.9213 and In.17491). Antenna: only Pilaria hooleyi Krzemin´ski, n. sp. cylindrical scapus and broad, oval pedicel is preserved in one (Fig. 17, Pl. 2d) specimen, In.17491. Venation. Sc ending before fork of Rs into R2 þ 3 þ 4and Holotype. NHMUK In.25254, Hooley Collection, Insect R5; cross-vein sc-r at its length before end of Sc; R1 rather Bed; late Eocene; NW Isle of Wight, UK. short; cross-vein r-r (R2) at its length before end of R1; Additional material. NHMUK I.9032/In.64099 (part and R2 þ 3 þ 4 very short; R4 16 times longer than R2 þ 3 þ 4; counterpart), Brodie/Hooley Collections. Rs about 10 times longer than R2 þ 3 þ 4; d cell short and Derivation of name. A new species is dedicated to R.W. broad; petiole of m1 cell a little shorter than d cell; M1 about Hooley, the famous collector of fossil insects from the Isle of twice longer than petiole of m1 cell; cross-vein m-cu in half of Wight. d cell; A2 long and somewhat wavy. Diagnosis. Sc ending opposite fork of Rs into R2 þ 3 þ 4 Remarks. The original description of (Cockerell 1921) of and R5; d cell long and narrow; petiole of m1 cell very Limnophila cyclospila has many mistakes, the most important long, about twice longer than M1 and 1.5 times longer than d being the absence of vein Sc. Cross-vein m-cu is wrongly posi- cell. tioned in distal part of d cell position. Description. Wing length 2.9–5.2 mm, without colour pattern. The colour pattern may indicate genus Austrolimnophila Venation. Sc rather short, ending opposite fork of Rs; cross- Alexander, but without a complete specimen preserved in vein sc-r twice its length before end of Sc; R1 rather long; good condition it is not possible to make this taxonomical cross-vein r-r (R2) at its length before end of R1; Rs five times decision. longer than R2 þ 3 þ 4; R3 about 1.5 times longer than Rs; d cell long and narrow; petiole of m1 cell about twice longer Limnophila (Limnophila) deleta Cockerell, 1921 than M1 and 1.5 times longer than d cell; cross-vein in third (Fig. 16, Pl. 2c) portion of d cell; A2 little curved. Remarks. Basing only on wing venation it is very difficult to 1921 Limnophila deleta Cockerell, pp. 459, 462, fig. 9 decide about generic position of this species, but in my opinion 1994 Limnophila deleta Evenhuis, 1994, p. 75 the general pattern of wing venation is more similar to the genus Pilaria than to Limnophila or Pseudolimnophila Alexander. Holotype. NHMUK I.9053 (female), Brodie Collection, Insect Bed; late Eocene; NW Isle of Wight, UK. Pilaria volodii Krzemin´ski, n. sp. Additional material. NHMUK, I.9019, I.9060, I.9067, (Fig. 18, Pl. 2e) I.9684, Brodie Collection; In.17115, In.17171, In.24414, Smith Collection; In.17484/In.64127 (part and counterpart), Smith/ Hooley Collections; In.25217, In.25474, In.64127, Hooley Holotype. NHMUK I.9167/I.9201 (part and counterpart), Collection; PI. II. 2764, Palaeontological Institute, Moscow Brodie Collection, Insect Bed; late Eocene; NW Isle of Wight, team, north end Thorness Bay, Isle of Wight. UK. Diagnosis. Sc rather long, ending opposite fork of Rs into Derivation of name. A new species is dedicated to Vladimir R2 þ 3 þ 4 and R5; Rs twice longer than R2 þ 3 þ 4; d cell (Volodia) Blagoderov (NHM, London), a paleoentomologist short; cross-vein m-cu little behind fork of Mb into M1 þ 2 and specialist on Diptera. and M3 þ 4 in 1/4 of d cell; A2 long and somewhat wavy. Diagnosis. Sc very short, little shorter than half wing length, Redescription. Wing length 5.4–8 mm, without colour pattern. ending opposite 2/3 of Rs; R3 and M1 very short; petiole of Few specimens are preserved with small portion of the body. m1 cell more than twice longer than d cell and about five times Only holotype is preserved with body, but antennae and palps longer than M1. are missing. Description. Wing length 4.7–5.0 mm, without colour pattern; Venation. Sc rather long, ending opposite fork of Rs on anal part of the wing destroyed. R2 þ 3 þ 4 and R5; cross-vein sc-r at its length before end of Venation. Sc very short, little shorter than half wing length; Sc; R1 rather long; cross-vein r-r (R2) at its length before end cross-vein sc-r at three times its length before end of Sc; R1 of R1; Rs about twice longer than R2 þ 3 þ 4; R3 about 1/4 long; cross-vein r-r (R2) at end of R1; R2 þ 3 þ 4 about 1/6 longer than Rs; d cell small; petiole of m1 cell little longer shorter than Rs and only little longer than R4; R3 very short, than d cell; M1 about 1.4 times longer than petiole of m1 about 1/3 of R4; d cell long and narrow; M1 very short, cell; cross-vein m-cu in quarter of d cell; A2 long and little petiole of m1 cell more than twice longer than d cell and about wavy. five times longer than M1; cross-vein m-cu in half length of d Remarks. Limnophila deleta is one of most common species cell. of Limoniidae in the Isle of Wight. The specimens allotted Remarks. Venation pattern of the wing venation with very herein are very variable and they may possibly represent short Sc, R3 and M1 is rather rare among Limoniidae. This more than one species, but based only on wing venation it is new species is different from all Pilaria species from Baltic not possible separate them further. amber.

Genus Gonomyia Meigen, 1818 Cheilotrichia (Empeda) hyalina (Cockerell, 1921) (Fig. 23, Pl. 2j) Type species. Limnobia tenella Meigen, 1818, p. 146 (by monotypy). 1921 Empeda hyalina Cockerell, p. 457, ﬁg. 5 1921 ‘Gonomyia’ grisea Cockerell, p. 461, ﬁg. 14, syn. nov. Subgenus Gonomyia Meigen, 1818 1994 Cheilotrichia hyalina Evenhuis, p. 65 Type species. As for genus. 1994 Gonomyia? grisea Evenhuis, p. 89 Remarks. Two hundred and sixteen recent species worldwide distributed from cold to tropical zone. Five fossil species Holotype. NHMUK I.24342, Hooley Collection; Insect of the genus Gonomyia are known, but probably only two Bed; late Eocene; NW Isle of Wight, UK. belong to the nominative subgenus: one from Italy (Miocene) Additional material. NHMUK In.24344, Hooley Collection, and one from Germany (Oligocene). Only one fossil species described as the holotype of the species Gonomyia grisea from the subgenus Electrogonomyia Alexander is described Cockerell, 1921. from Baltic amber. Diagnosis. Sc ending behind mid of Rs; Rs three times From the Isle of Wight Cockerell & Haines (1921) described longer than R3; R2 þ 3 þ 4andR3þ 4togetheraslongas Gonomyia indecisa; in the same year Cockerell (1921) described Rs; and 1.5 times longer than R4; d cell long and narrow. G. ferrea, G. grisea and G. lutescens. Revision of the holotypes

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Plate 3 Limoniidae. (a) Cheilotrichia (Empeda) szwedoi Krzemin´ski, n. sp., holotype NHMUK In.17082. (b) Gonomyia (Gonomyia) indecisa Cockerell & Haines, 1921, holotype NHMUK In.9087. (c) Gymnastes (Paragymnastes) fasciatipennis Cockerell, 1921, holotype NHMUK I.10242. (d) Symplecta (Psiloconopa) gurnetensis Krzemin´ski, n. sp., holotype NHMUK In.25266. (e) Styringomyia concinna (Cockerell, 1917a) n. comb., holotype NHMUK I.8666. (f ) Thaumastoptera undulata (Cockerell & Haines, 1921) n. comb., holotype NHMUK I.9146. (g) Orimarga (Orimarga) lenae Krzemin´ski, n. sp., holotype NHMUK In.25487. (h) Dicranomyia (Dicranomyia) aliena (Cockerell, 1921) n. comb., holotype NHMUK I.9018. (i) Dicranomyia (Dicranomyia) brodiei (Cockerell, 1917a) n. comb., holotype NHMUK I.8603. (j) Dicranomyia (Dicranomyia) excavata Cockerell & Haines, 1921, holotype NHMUK I.9946.

Subgenus Paragymnastes Alexander, 1922 Holotype. NHMUK In.25266, Hooley Collection); Insect Bed; late Eocene; NW Isle of Wight, UK. Type species. Limnobia fascipennis Thomson, 1869, p. 447 Derivation of name. The name species origin from the old (original designation). name of Gurnard Bay (Gurnet Bay). Remarks. Thirty-one recent species are known from Oriental, Diagnosis. Sc rather long, ending behind cross-vein r-r (R2); Australian and Eastpalaearctic Regions cross-vein sc-r opposite proximal 1/4 Rs; cross-vein very close to fork of Mb into M1 þ 2andM3þ 4. Gymnastes (Paragymnastes) fasciatipennis Cockerell, 1921 Description. Wing length 4.1 mm without colour pattern. (Fig. 26, Pl. 3c) Wing folded.

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Figures 26–36 Limoniidae, wing venation. (26) Gymnastes (Paragymnastes) fasciatipennis Cockerell, 1921, holotype, NHMUK I.10242. (27) Symplecta (Psiloconopa) gurnetensis Krzemin´ski, n. sp., holotype NHMUK In.25266. (28) Ilisia (Lunaria) incognita (Cockerell, 1921), holotype NHMUK I.9186. (29) Styringomyia concinna (Cockerell, 1917a), holotype NHMUK I.8666. (30) Thaumastoptera undulata (Cockerell & Haines, 1921), holotype NHMUK I.9146. (31) Orimarga (Orimarga) lenae Krzemin´ski, n. sp., holotype NHMUK In.25487. (32) Dicranomyia (Dicranomyia) aliena (Cockerell, 1921), holotype NHMUK I.9018. (33) Dicranomyia (Dicranomyia) brodiei (Cockerell, 1917a), holotype NHMUK I.8603. (34) Dicranomyia (Dicranomyia) excavata Cockerell & Haines, 1921, holotype NHMUK I.9946. (35) Dicranomyia (Dicranomyia) fasciata Krzemin´ski, n. sp., holotype NHMUK I.9058. (36) Limonia spilota (Cockerell, 1921), holotype NHMUK In.17114.

Type species. Erioptera maculata Meigen, 1804, p. 51 Holotype. NHMUK I.8666, Brodie Collection; Insect Bed; (original designation). late Eocene; NW Isle of Wight, UK. Additional material. NHMUK In.24354, Hooley Collection Subgenus Lunaria Savchenko, 1973 (H 261), described as Styringomyia extensa; I.8998 (1), Brodie Collection; II 2,785 (2) collected by the Zoological Institute, Type species. Ilisia (Lunaria) idiophallus Savchenko, 1973, St Petersburg team, Burnt Wood, Isle of Wight. p. 38 (original designation). Diagnosis. Sc very short, about 1/3 wing length, ending just Remarks. Only two recent species are known from the behind fork of Rb into R1 and Rs; R1 ending opposite proximal Palaearctic Region. 1/3 of Rs; Rs twice times longer than R2 þ 3 þ 4. Redescription. Wing length 3.5–4.8 mm. Ilisia (Lunaria) incognita (Cockerell, 1921) n. comb. Venation. Sc very short, about 1/3 wing length, ending just (Fig. 28, Pl. 7b) behind fork of Rb into R1 and Rs; cross-vein sc-r absent; only three long radial veins present; R1 very short, ending opposite 1921 Limnophila incognita Cockerell, p. 462, Fig. 16 1/3 of Rs; cross- vein r-r (R2) and vein R3 probably lost by 1994 Limnophila incognita Evenhuis, p. 75 atrophy; Rs twice longer than R2 þ 3 þ 4; R5 1.75 times longer than Rs; d cell long and narrow, about twice longer Holotype. NHMUK I.9186, Brodie Collection; Insect Bed; than M3; cross-vein m-cu in 1/5 of d cell; A2 long and little late Eocene; NW Isle of Wight, UK. wavy. Additional material. NHMUK In.64111, Hooley Collection. Remarks. Cockerell (1917a) described Mesomyites concinnus Diagnosis. Sc short, ending before fork of Rs into and mistakenly included this species in the family Empididae R2 þ 3 þ 4 and R5; cross-vein opposite proximal 1/3 of Rs; (Diptera Brachycera) and together with Haines (Cockerell & M3 1.5 times longer than d cell; cross-vein m-cu behind fork Haines 1921) described new species Styringomyia extensa. of Mb into M1 þ 2andM3þ 4. Study of the holotypes of these species shoved that it is the Redescription. Most parts of body preserved, without palps, same species from the genus Styringomyia and S. extensa is legs and genitalia. Wing length 6.3 mm. the junior synonym. Antenna. scapus rather long; pedicel short and rounded; only three flagellomeres are preserved: first two short, broad Subfamily Limoniinae Speiser, 1909 and next narrow and elongate, little widened basally; all with bristles shorter than flagellomeres. Genus Thaumastoptera Mik, 1866 Venation. Sc short, ending before fork of Rs into R2 þ 3 þ 4 and R5; cross-vein opposite proximal 1/3 of Rs, about nine Type species. Thaumastoptera calceata Mik, 1866, p. 303 times its length before end of Sc; R1 rather long; cross-vein (original designation). r-r (R2) eight times its length before end of R1; Rs long, only about 1/3 shorter than R5; R2 þ 3 þ 4andR2þ 3veryshort, Subgenus: Thaumastoptera Mik, 1866 R2 þ 3 þ 4 only twice longer than R2 þ 3; d cell narrow; M3 1.5 times longer than d cell; cross-vein little behind fork of Mb Type species. As for genus. into M1 þ 2andM3þ 4; A2 slightly wavy. Remarks. Eight recent species are known from Palaearctic Remarks. Ilisia (Lunaria) incognita is the first fossil repre- (three sp.), Oriental (two sp.), Afrotropic (two sp.) and Nearctic sentative of the genus Ilisia. Recently, Ilisia is a small genus (one sp.) Regions. Only two fossil species are known from the whose species live mostly in Palaearctic and Oriental Regions, Baltic amber (Eocene). (some are also in Afrotropical and Australian Regions). Classification of the species from the Isle of Wight in the sub- Thaumastoptera undulata (Cockerell & Haines, 1921) genus Lunaria is based only on wing venation. n. comb. (Fig. 30, Pl. 3f ) Genus Styringomyia Loew, 1845 1921 Dicranomyia undulata Cockerell and Haines, p. 110, fig. I9146 Type species. Styringomyia venusta Loew, 1845, p. 6 (by 1931 Dicranomyia undulata Alexander, pp. 8, 41 monotypy) (fossil). 1994 Dicranomyia undulata Evenhuis, p. 68

Downloaded from https://www.cambridge.org/core. Chalmers Tekniska Högskola, on 06 Oct 2019 at 16:48:14, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S1755691018000464 TRUE FLIES FROM THE LATE EOCENE INSECT LIMESTONE OF THE ISLE OF WIGHT 19 Holotype. NHMUK I.9146, Brodie Collection; Insect Bed; Type species. As for genus. late Eocene, NW Isle of Wight, UK. Remarks. Eight hundred and fifty-seven recent species Diagnosis. Sc short, ending opposite fork of Rb into R1 and worldwide distributed from cold to tropical zones. About 30 Rs: Rb forked behind mid wing; cross-vein m-cu at its length fossil species are known from the Upper Cretaceous to Miocene. before fork of Mb into M1 þ 2andM2þ 3. Unfortunately, representatives of the nominal subgenus Redescription. Wing length 6.0 mm. cannot be identified without knowledge of morphology of Venation. Sc very short, only a little longer than half wing male genitalia, and only wings are presented. Four species are length, ending opposite fork of Rb into R1 and Rs; cross-vein known from Baltic amber. sc-r twice its length before end of Sc; Rb forked behind mid wing; R1 rather short; cross-vein r-r(R2) twice its length Dicranomyia (Dicranomyia) aliena (Cockerell, 1921) n. comb. before end of R1; Rs very short , only twice longer than (Fig. 32, Pl. 3h) R2 þ 3 þ 4; R3 þ 4 three times longer than R2 þ 3 þ 4and twice longer than Rs; R5 three times longer than Rs; d cell 1921 Limnophila aliena Cockerell, pp. 461–62, fig. 15 open by atrophy of cross-vein between M3 and M4; M3 1.5 1931 Limnophila aliena Alexander, p. 8 times longer than M1 þ 2; cross-vein m-cu at its length 1994 Limnophila aliena Evenhuis, p. 75 before fork of Mb into M1 þ 2andM3þ 4; A2 strongly wavy terminally. Remarks. Cockerell & Haines (1921) erroneously included Holotype. NHMUK I.9018/I.9420 (part and counterpart), T. undulata in the genus Dicranomyia. Brodie Collection; Insect Bed; late Eocene; NW Isle of Wight, Recently genus Thaumastoptera is not represented in UK. Australian and Neotropical Regions; most species live in the Additional material. NHMUK, I.9758, Brodie Collection. Holarctic Region. Until now only two species from Baltic Diagnosis. Sc rather long, ending opposite mid Rs; d cell amber have been described (Krzemin´ski, 1985). long and broad; cross-vein m-cu in proximal 1/3 of d cell. Description. Wing length 5.4–7.0 mm, basal part and small parts of medial and cubital field destroyed. Genus Orimarga Osten Sacken, 1869 Venation. Sc rather long, ending opposite mid of Rs; cross- vein sc-r at its length before end of Sc; R1 rather long, ending Type species. Limnobia alpina Zetterstedt, 1851, p. 3894 opposite fork; of R2 þ 3 þ 4intoR3andR3þ 4; Rs as long (designated by Coquillett 1910). as R2 þ 3 þ 4 and 1.5 times longer than R3 þ 4; R5 three times longer than R5; d cell long and broad; M3 1.3 longer Subgenus Orimarga Osten Sacken, 1869 than d cell; cross-vein m-cu in proximal 1/3 of d cell; A1 straight; A2 rather short and straight. Remarks. Cockerell (1921) erroneously included this species Type species. As for genus. in the genus Limnophila. Remarks. One hundred and thirty-seven recent species are known from Neotropic (42 sp.), Oriental (41 sp.), Palaearctic (21 sp.), Australian (15 sp.), Afrotropic (14 sp.) and Nearctic Dicranomyia (Dicranomyia) brodiei (Cockerell, 1917a) n. comb. (four sp.) Regions. (Fig. 33, Pl. 3i)

Orimarga (Orimarga) lenae Krzemin´ski, n. sp. 1917a Riphidia brodiei Cockerell, pp. 373, 384, pl. 31, ﬁg. 7 (Fig. 31, Pl. 3g) 1917b Rhipidia brodiei Cockerell, p. 141 1994 Rhipidia brodiei Evenhuis, p. 84 Holotype. In.25487 Hooley Collection; Insect Bed; late Holotype. NHMUK I.8603, Brodie Collection; Insect Bed; Eocene; NW Isle of Wight, UK. late Eocene; NW Isle of Wight, UK. Derivation of name. The new species is dedicated to Lena Additional material. In.17486 (part and counterpart), Smith Lukashevich (Russia, Moscow), specialist of fossil Diptera. Collection; In.25460, In.25477/In.26051 (part and counter- Diagnosis. Sc ending opposite fork of Rs into R2 þ 3 þ 4 part), Hooley Collection. and R5; cross-vein m-cu little before fork of Mb into M1 þ 2 Diagnosis. Sc short, ending opposite fork of Rb on R1 and and M3 þ 4. Rs; d cell short and broad; cross-vein m-cu in or little before Description. Sc rather short, ending opposite fork of Rs into fork of Mb on M1 þ 2andM3þ 4. R2 þ 3 þ 4 and R5; cross-vein sc-r three time its length before Redescription. Wing 6.1–7.0 mm, with big, dark stigma. end of Sc; R1 short, ending opposite cross-vein of m-r; cross- Venation. Sc short ending opposite fork of Rb into R1 and vein r-r (R2) at its length before end of R1; R3 þ 4 three times Rs; cross-vein sc-r twice its length before end of Sc; R1 rather longer than R2 þ 3 þ 4 and little more than twice Rs; d cell long; R3 þ 4 2.3 times longer than Rs and 2.5 times longer open by atrophy of cross-vein m-m between M1 þ 2andM3; than R2 þ 3 þ 4; R5 2.5 times longer than Rs; d cell short cross-vein m-cu somewhat before fork of Mb into M1 þ 2and and broad; M3 as long as d cell; cross-vein m-cu in fork of M3 þ 4; A2 short and rather straight. Mb into M1 þ 2andM3þ 4, but in some specimens this cross Remarks. Orimarga(Orimarga) lenae is the ﬁrst fossil repre- vein is positioned little before this fork; A1 straight or little sentative of the genus Orimarga. wavy; A2 little wavy. Remarks. Cockerell (1917a, b) erroneously included this Genus Dicranomyia Stephens, 1829 species in the genus Rhipidia. More detailed study of the holotype and additional materials has shown that species described Type species. Limnobia modesta Meigen, 1818, p. 134 as Rhipidia brodei belongs to the genus Dicranomyia. (designated by Coquillett 1910). Dicranomyia (Dicranomyia) excavata Cockerell & Haines, 1921 Subgenus Dicranomyia Stephens, 1829 (Fig. 34, Pl. 3j)

Holotype. NHMUK I.9058, Brodie Collection; Insect Bed; Genus Helius Lepeletier & Seville, 1828 late Eocene; NW Isle of Wright, UK. Diagnosis. Sc long, ending opposite 3/4 of Rs; R3 þ 4 very Type species. Limnobia longirostris Meigen, 1818, p. 146 long and strongly curved. (autom.). Description. Wing length 8.0 mm, with big, dark stigma. Wing in medial, cubital and annal part destroyed; most parts of the body are preserved. Subgenus Helius Lepeletier & Seville, 1828 Head. antenna with big tubular scapus and small flat pedicel; flagellomeres short and rounded, with short bristles, not longer Type species. As for genus. than flagellomere. Remarks. One hundred and eighty recent species are known Thorax partially preserved, with only one strongly destroyed from Oriental (70 sp.), Neotropic (51 sp.), Afrotropic (23 sp.), wing. Palaearctic (20 sp.), Australian (14 sp.) and Nearctic (two sp.) Venation. Sc long, ending opposite 3/4 of Rs; cross-vein sc-r Regions. Seven fossil species are described from Upper Creta- at its length before end of Sc; R3 þ 4 very long and strongly ceous through Oligocene. curved, 2.3 times longer than Rs and about twice longer than R2 þ 3 þ 4; R5 very long, strongly curved, and more than Helius (Helius) edmundi Krzemin´ski, n. sp. twice longer than Rs. Remaining veins in most parts are de- (Fig. 37, Pl. 4c) stroyed. Abdomen. Male genitalia with outer dististyle large, balloon Holotype. NHMUK I.9810, Brodie Collection; Insect Bed; like; inner dististyle invisible; penis short and broad basally. late Eocene; NW Isle of Wight, UK. Remarks. Despite poor state of wing preservation, the Derivation of name. A new species is dedicate to Edmund venation retained (especially shape of radial veins) and male Jarzembowski (UK) specialist of fossil insects.

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Plate 4 Limoniidae, Scatopsidae and Anisopodidae. (a) Dicranomyia (Dicranomyia) fasciata Krzemin´ski, n. sp., holotype NHMUK I.9058. (b) Limonia spilota (Cockerell, 1921), holotype NHMUK In.17114. (c) Helius (Helius) edmundi Krzemin´ski, n. sp., holotype NHMUK I.9810. (d) Helius (Helius) popovi Krzemin´ski, n. sp., holotype NHMUK I.9392. (e) Dicranoptycha staryi Krzemin´ski, n. sp., holotype NHMUK In.24787. (f) Helius (Helius) vectensis (Cockerell, 1915) n. comb., holotype USNM 61439. (g) Trentepohlia (Mongoma) pallescens (Cockerell, 1921), NHMUK I.8898. (h) Trentepohlia (Mongoma) cruciferella (Cockerell, 1917a), holotype NHMUK I.8635. (i) Regmoclemina haennii Krzemin´ski, n. sp., holotype NHMUK In.9512. (j) Sylvicola hooleyi (Cockerell, 1921), holotype NHMUK In.24346. (k) Sylvicola problematica Krzemin´ski, n. sp., holotype NHMUK In.25675.

Figures 37–42 Limoniidae, wing venation. (37) Helius (Helius) edmundi Krzemin´ski, n. sp., holotype NHMUK I.9810. (38) Helius (Helius) popovi Krzemin´ski, n. sp., holotype NHMUK I.9392. (39) Helius (Helius) vectensis (Cockerell, 1915), holotype USNM 61439. (40) Dicranoptycha staryi Krzemin´ski, n. sp., holotype NHMUK In.24787. (41) Trentepohlia (Mongoma) cruciferella (Cockerell, 1917a), holotype NHMUK I.8635. (42) Trentepohlia (Mongoma) pallescens (Cockerell, 1921), holotype NHMUK I.8898.

Diagnosis. Sc ending opposite fork of Rs into R2 þ 3 þ 4 M3 1.3 times longer than d cell; A1 and A2 little wavy in and R5; R5 twice longer than Rs, in mid part strongly wavy distal part. and d cell widened in proximal broaden in 1/3. Remarks. H. popovi differs from two other Helius species Description. Wing length 5.0 mm, with small black stigma. from the Isle of Wight by the regular rectangular shape of d Most parts of thorax, abdomen and two wings are preserved; cell. head is destroyed. Venation. Sc rather long, ending opposite fork of Sc into Helius (Helius) vectensis (Cockerell, 1915) n. comb. R2 þ 3 þ 4 and R5; cross-vein four times its length before (Fig. 39, Pl. 4f ) end of Sc; R1 long; R2 þ 3 þ 41.25timesofRs;R5twice longer than Rs, strongly wavy in mid part; d cell roughly 1915 Atarba vectensis Cockerell, pp. 492–93, 499, pl. 61, rhomboid, with rounded upper margin and widened in 1/3 fig. 10 length; M1 þ 2 strongly wavy; M3 1.25 times longer than d 1994 Atarba vectensis Evenhuis, p. 64 cell; A1 little wavy in distal part; A2 only slightly curved to anal margin of the wing. Holotype. USNM 61439 (Lacoe Collection 7513); Insect Remarks. Strong and wavy veins R5 and M1 þ 2 separate Bed; late Eocene; NW Isle of Wight, UK. this single species from all other representatives of Helius. Additional material. NHMUK I.10169 (female), Brodie Collection; In.25252/In.25512 (female, part and counterpart), Helius (Helius) popovi Krzemin´ski, n. sp. In.25464/In.25383 (part and counterpart), In.64097, Hooley (Fig. 38, Pl. 4d) Collection. Diagnosis. Sc ending opposite 2/3 of Rs; R5 2,5 times longer Holotype. NHMUK I.9392, Brodie Collection; Insect Bed; than Rs; d cell broaden in 1/3 of its length. late Eocene; NW Isle of Wight, UK. Redescription. Wing length 4.8–7.8 mm, with small, dark Derivation of name. The new species is dedicated to the late stigma. Some specimens preserved with body. Yuri Popov (Russia), specialist of fossil Heteroptera. Head with long rostrum, about twice longer than head; of Diagnosis. Sc ending opposite fork of Rs into R2 þ 3 þ 4 the antenna only scapus, pedicel and three flagellomeres are and R5; R5 twice longer than Rs; d cell rectangular. preserved. Scapus long and tubular; pedicle half as long as Description. Wing length 6.0 mm, with dark stigma. Only scapus and more broad; flagellomeres short and oval. small parts of thorax and abdomen are preserved. Venation. Sc ending opposite 2/3 of Rs; cross-vein sc-r twice Venation. Sc rather long, ending opposite fork of Rs into its length before end of Sc; R1 long; R2 þ 3 þ 4 1.5 times R2 þ 3 þ 4andR5;R1long;R2þ 3 þ 4 1.3 times longer longer than Rs; R5 2,5 times longer than Rs; d cell rhom- than Rs; R5 twice longer than Rs; d cell rectangular, broad; boid, widened in 1/3 length; M3 1.3 times longer than d cell;

Downloaded from https://www.cambridge.org/core. Chalmers Tekniska Högskola, on 06 Oct 2019 at 16:48:14, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S1755691018000464 TRUE FLIES FROM THE LATE EOCENE INSECT LIMESTONE OF THE ISLE OF WIGHT 23 cross-vein m-cu just behind fork of Mb into M1 þ 2and Head rather small, preserved without antennas and palps. M3 þ 4; terminal section of A1 wavy; A2 rather straight. Thorax in good condition, but without legs and only one Abdomen. Some specimens preserved with ovipositor. wing is visible. Remarks. Helius vectensis is the most common species of Venation. Sc rather short, ending opposite 1/3 of the genus. Cockerell (1915) used old genus name Atarba,a R2 þ 3 þ 4 þ 5onR2þ 3 þ 4andR5;R1rathershort;r-r synonym of Helius. Recently the genus is rich in species and (R2) long, twice longer than last section of R1 and escaping distributed in all regions on the Earth. in fork of R2 þ 3 þ 4 þ 5; Rs 1.25 times longer than R2 þ 3 þ 4 þ 5, a little more than twice R3 þ 4 and a little Genus Dicranoptycha Osten Sacken, 1860 shorter than R5; d cell long and narrow, and about 1.3 longer than M3; cross-vein m-cu in fork of Mb into M1 þ 2and Type species. Dicranoptycha germana Osten Sacken, 1860, M3 þ 4 and strongly curved basally; Cu strongly curved in p. 217 (designated by Coquillett 1910). last section, connected with tip A1; A2 very short and wavy. Remarks. Eighty-nine recent species are known from Afro- Abdomen of specimen In.17264/I.9200 with well preserved trocic (31 sp.), Palaearctic (22 sp.), Nearctic (22 sp.), Oriental ovipositor. (six sp.) and Neotropic (three sp.) Regions. Five fossil species Trentepohlia (Mongoma) pallescens (Cockerell, 1921) are known from Upper Cretaceous through Miocene. (Fig. 42, Pl. 4g) Dicranoptycha staryi Krzemin´ski, n. sp. 1921 Mongoma pallescens Cockerell, p. 460, ﬁg. 11 (Fig. 40, Pl. 4e) 1994 Trentepohlia (Mongoma) pallescens Evenhuis, p. 87 Holotype. NHMUK I.8898, Brodie Collection; Insect Bed; Holotype. NHMUK In.24787, Hooley Collection; Insect late Eocene; NW Isle of Wight, UK. Bed; late Eocene; NW Isle of Wight, UK. Additional material. In.17117/I.9051 (part and counterpart) Derivation of name. A new species is dedicated to Jaroslav Smith/Brodie Collections; In.17123, Smith Collection; Stary (Czech Republic), specialist of recent Limoniidae In.25431, In.25439, In.64113, Hooley Collection. (Diptera). Diagnosis. Cross-vein r-r (R2) about three times longer than Diagnosis. R5 2.3 length of Rs; cross-vein m-cu in fork of last section of R1 escaping in R2 þ 3 þ 4 þ 5, before fork of Mb into M1 þ 2andM3þ 4. this vein; d cell short and narrow, about half of M3. Description. Wing length 7.0 mm. Redescription. Sc rather short, ending opposite 1/4 of Venation. Sc ending opposite 1/4 of R2 þ 3 þ 4; cross-vein R2 þ 3 þ 4 þ 5; cross-vein sc-r at its length before end of sc-r at its length before end of Sc; R1 long; cross-vein r-r (R2) Sc; R1 rather short; cross-vein r-r (R2) about three times long, twice longer than last section of Sc; R3 þ 41.25times longer than last section of R1 and escaping directly in longer than R2 þ 3 þ 4 and only a little shorter than Rs; R5 R2 þ 3 þ 4 þ 5; Rs as long as R2 þ 3 þ 4 þ 5andR3þ 4 þ 5 2.5 times longer than Rs; d cell big and broad with rounded together, three times longer than R3 þ 4 and about half as upper margin; M3 as long as d cell; cross-vein m-cu in fork of long as R5; d cell short and narrow, about half as long as Mb into M1 þ 2andM3þ 4; A1 a little curved. M3; cross-vein in fork of Mb into M1 þ 2andM3þ 4, strong curved basally; Cu strongly curved in distal section, connected Genus Trentepohlia Bigot, 1854 with tip of A1; A2 very short and wavy. Remarks. These two species of the subgenus Mongoma Type species. Limnobia limnobioides Bigot, 1854, p. 456 are common at the site and indicate tropical or subtropical (original designation). climate. All recent representatives live in warm climate zones.

Subgenus Mongoma Westwood, 1881 Superfamily Psychodoidea Newman, 1834 by Dany Azar Type species. Mongoma fragillima Westwood, 1881, p. 364 (by monotypy). Family Psychodidae Newman, 1834 Remarks. One hundred and forty-one recent species are known from tropical and subtropical regions. Two fossil species Psychodidae are a cosmopolitan family habitually associated are known. with moist or sylvan habits. Most are nocturnal and can be found resting in shaded places in the daylight hours. Trentepohlia (Mongoma) cruciferella (Cockerell, 1917a) Fossil Psychodidae fossilised in the amber are sufficiently (Fig. 41, Pl. 4h) well preserved in pristine detail for accurate comparisons with the extant representatives of the family. Unfortunately, the study of insects’ fossil impressions in marl or limestone is 1917a Mongoma cruciferella Cockerell, p. 460, fig. 11 mainly based on the wing venation. 1994 Trentepohlia (Mongoma) cruciferella Evenhuis, p. 87 Cockerell (1915, 1921) described very succinctly two psychodid flies (Psychoda primaeva Cockerell, 1915, and Psychoda Holotype. NHMUK I.8635, Brodie Collection; Insect Bed; leucospila Cockerell, 1921). late Eocene; NW Isle of Wight, UK. The recent revision of the Isle of Wight fossil insect material Additional material. In.17116, In.17130, In.17259, Smith kept in the NHMUK and the Maidstone Museum and Art Collection; In.17264/I.9200 (female, part and counterpart) Gallery (Maidstone) revealed a new species of fossil psychodid Smith/Brodie Collections; In.25505, In.64123, Hooley Collection. flies and led to improvement of our knowledge of P. primaeva Diagnosis. Cross-vein r-r (R2) twice longer than last section Cockerell, 1915, and P. leucospila Cockerell, 1921. of R1 and escaping in to fork of R2 þ 3 þ 4intoR3þ 4and Herein I redescribe P. leucospila Cockerell, 1921, and P. R5; d cell long and narrow, about 1.3 times longer than M3. primaeva Cockerell, 1915, as Wightipsychoda leucospila Redescription. Wing length 5.0–6.0 mm. Female In.17264/ (Cockerell 1921) n. comb.; Pericoma primaeva (Cockerell I.9200 preserved with body.

Derivation of name. After the Isle of Wight and Psychoda. 1915 Psychoda primaeva Cockerell, pp. 493, 494, plate 62, Type species. Psychoda leucospila Cockerell, 1921. fig. 5 Diagnosis. Presence of incomplete eye bridge. Wings shape 1994 Psychoda leucospila Evenhuis, p. 194 and venation very similar to those of the genus Psychoda with pointed wings; radial fork beyond medial, but with the differ- Holotype. USNM 61441 (Fig. 46, Pl. 5g), Lacoe Collection; ence of R4 ending at tip instead of R5. Males with white oval Insect Bed; late Eocene; NW Isle of Wight, UK. prominent tuft of scales shortly beyond costal node and be- Additional material. NHMUK I.8987 (Fig. 48), I.9011/ tween C and Sc. CuA2 well developed, strongly curved I.9272 (part and counterpart), I.9011, I.9149 (Pl. 5e), I.9165 basally and reaching costal margin. (female?, Fig. 49), I.9177 (male, Pl. 5d), I.9290 (male, Pl. 5f ), I.9319, I.9388, I.9600, I.9746, I.10403, Brodie Collection; Wightipsychoda leucospila (Cockerell, 1921) n. comb. In.25341, In.25342, In.25344, In.25345 (male, Fig. 47), (Figs 43–45, Pl. 5a–c) In.25346, In.26057, Hooley Collection; In.64535, Jarzembowski Collection (Thorness Bay); In.25456, Hooley collection; 1921 Psychoda leucospila Cockerell, p. 466, fig. 25 I.10069, Brodie collection; In.25343, Hooley collection; 1994 Psychoda leucospila Evenhuis, p. 194 I.9254, Brodie collection; II.2719, Azar Collection (Thorness Bay, May 2005); MNEMG 2018.6.0517; 2018.6.1522; Holotype. NHMUK I.9463, Brodie Collection, male (Fig. 43, 2018.6.0545/2018.6.0549 (part and counterpart); 2018.6.0159; Pl. 5a); Insect Bed; late Eocene; NW Isle of Wight, UK. 2018.6.0411; 2018.6.0684 and 2018.6.0761. Additional material. NHMUK I.10010, male (Fig. 44, Pl. 5b), Diagnosis. Presence of incomplete eye bridge. Antenna with I.9384, male (Fig. 45, Pl. 5c), I.9265 (female), I.9974, Brodie barrel-shaped flagellomeres. Wings obtusely pointed; radial Collection; In.17204 (probably male), Smith Collection; fork level very slightly distal to radial one. R5 ending in wing In.25340, Hooley Collection. margin beyond the apex. Diagnosis. As for the genus. Description. Eyes presenting a well-developed but incom- Description. Eyes with a developed eye bridge. Eyes separated plete eye bridge. Eyes separated with a distance of 0.025 mm by a distance of 0.017 mm. Wing 2.2 mm long, 1.02 mm wide. (after specimens I.8987 and I.9319). Antenna with barrel- Chalky-white large oval prominent tuft of scales shortly beyond shaped flagellomeres. The first one nearly twice longer than costal node and between C and Sc. Subcostal vein Sc distally the other flagellomeres (after specimen I.9319). Wings ob- reaching the costal margin. R1 reaching the costal margin tusely pointed 2.3–3 mm long (the holotype is 2.8 mm long), 1.4 mm from wing base. Branching of R2 þ 3 0.65 mm distal 0.9–1.2 mm wide (the holotype is 1.08 mm wide). Subcostal of wing base. R2 and R3 separated 1.1 mm distally. R2 and vein Sc distally reaching the costal margin. R1 reaching the R3 reaching costal margin. R4 reaching wing apex. Fork of costal margin 1.5–1.62 mm from wing base. Branching of M into M1 þ 2 and M3 0.8 mm distal of wing base. M3 reach- R2 þ 3 0.5–0.56 mm distal of wing base. R2 and R3 separated ing wing margin at 1.45 mm from wing base. CuA1 reaching 0.95 to 1 mm distally. R2 and R3 reaching costal margin. wing margin 1.1 mm distal of wing base. CuA2 rather developed, Fork of M into M1 þ 2 and M3 0.94 mm distal of wing strongly curved basally. Thorax (after specimen I.10010) 0.65 mm base. M3 reaching wing margin at 1.65–1.68 mm from long, 0.5 mm high. Abdomen 1 mm long, and 0.45 mm wide. wing base. CuA1 reaching wing margin 1.42–1.45 mm distal Remarks. Wightipsychoda leucospila (Cockerell 1921), shares of the wing base. CuA2 reaching wing margin at 0.83–1 mm with the Psychoda species the general known features of the from wing base. Anal vein well developed reaching wing genus such as (1) the presence of eye bridge; (2) pointed wing; margin at 0.375–0.43 mm apically. Thorax 0.6–0.85 mm long, (3) radial fork beyond medial; and (4) males with prominent 0.55–0.75 mm wide. Abdomen 0.9–1.4 mm long, and 0.5 mm tuft of scales shortly beyond costal node and between C and wide. Sc. It differs from Psychoda with R4 ending at wing apex Remarks. Cockerell (1915, pp. 493–94) described Pericoma instead of R5. The latest character allows the attribution of primaeva (Cockerell 1915) n. comb. and attributed it to the this species to a new genus Wightipsychoda. Cockerell (1921, genus Psychoda Latreille, 1796 because the ‘rather pointed p. 466) stated that no species could be found among living wingtip and general appearance indicate the reference to forms of psychodid flies with the presence of ‘chalky-white Psychoda’. But for all the species attributed to Psychoda,R5 costal patch’. A minute examination of this ovoid patch indi- ends at wingtip, and the antennal flagelomeres are not barrel- cates that it is formed of prominent tuft of scales. This charac- shaped but rather nodiform, thus Pericoma primaeva cannot ter is known in males of several taxa belonging to Psychoda belong to the genus Psychoda. According to the keys to genera genus as could be seen in Duckhouse (1966, p. 194), like Psy- given by Quate & Vockeroth (1981), Pericoma primaeva choda pennicillata Satchell, 1950; Psychoda mundula Duckhouse, (Cockerell 1915) n. comb. falls into the genera Pericoma 1966; Psychoda cristata Duckhouse, 1966; Psychoda gracilipenis Haliday, 1856 for the following reasons: (1) eyes with eye Duckhouse, 1966; Psychoda disaccus Duckhouse, 1966; Psy- bridge; (2) flagellomeres barrel-shaped; (3) R5 ending beyond choda salicorna Quate, 1954, etc. In females this patch is present wing apex; (4) radial fork on the same level as medial fork or but smaller and without scales. Excluding the general features slightly distal to it. of the family, Wightipsychoda leucospila does not share The relatively abundance of the fossils belonging to Peri- special similarities with any of the known Tertiary fossils coma primaeva allows a better understanding of this species; moth flies. it indicates that there is a large range of morphological polymorphism (not due to sexual dimorphism) in wing sizes and

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Figures 43–50 Psychodidae. (43)–(45) Wightipsychoda leucospila (Cockerell, 1921) n. comb.: (43) holotype, NHMUK I.9463, male; (44) NHMUK I.10010; (45) NHMUK I.9384. (46)–(49) Pericoma primaeva (Cockerell, 1915) n. comb.: (46) holotype, USNM 61441; (47) NHMUK In.25345, male; (48) NHMUK I.8987; (49) NHMUK I.9165, female? (50) Psychodinae? NHMUK In.26069(2). Scale bars ¼ 1 mm.

measurements. At a first examination I decided to split this sented by the unique specimen NHMUK In.26069(2), Hooley species into two different ones on the basis of the difference in Collection (Fig. 50, Pl. 5h). This species is very small (wing not wing length based on a 30 % of difference in size: a group with more than 1.35 mm long) and quite different from all the large wings and another one with small wings. But the minute known fossil psychodid flies from the Isle of Wight. The radial study of all the specimens shows clearly a gradualism in size fork apically well beyond the medial. Unfortunately the badly from small (wings of 2.3 mm) to large individuals (wings of preserved state of the specimen does not allow any further 3 mm) with a clear dominance of specimens of middle size consideration beyond the subfamily level. (six specimens with wings of 2.6 mm). Unfortunately, except Key to the fossil Eocene moth fly species from the Isle of the wing venation measurements, no preserved features could Wight be utilised to verify if the observed difference is only due to polymorphism. Except the common features of the genus, (1) Wings smaller in length than 1.5 mm...... the herein Pericoma primaeva doesn’t share particular similarities with any unnamed species of the known Tertiary fossils moth flies, but is much larger. Wings greater than 1.5 mm...... 2 (2) Wing with R4 ending in wing apex, and radial fork distal Subfamily ?Psychodinae Newman, 1834 to medial fork...... Wightipsychoda leucospila R4 and R5 beyond wing apex, and level radial fork The examination of all the Isle of Wight psychodid material slightly distal to medial one...... Pericoma primaeva allows the finding of one more fossil moth fly species repre-

Plate 5 Psychodidae and Dixidae. (a–c) Wightipsychoda leucospila (Cockerell, 1921) n. comb.: (a) holotype NHMUK I.9463, male; (b) NHMUK I.10010; (c) NHMUK I.9384. (d–g) Pericoma primaeva (Cockerell, 1915) n. comb.: (d) NHMUK I.9177, male; (e) NHMUK I.9149; (f ) NHMUK I.9290, male; (g) holotype, USNM 61441. (h) Unnamed new ?Psychodinae, NHMUK In.26069 (2). Scale bars ¼ 1 mm. (i–k) Dixella priscula (Cockerell, 1921) n. comb.: (i) NHMUK In 24345, holotype; (j) NHMUK In 17402, male; (k) NHMUK In 26055.

Holotype. NHMUK In.24345 (Fig. 54, Pl. 5i), Hooley Family Dixidae? Schiner, 1868 Collection; poorly preserved wing; Insect Bed; late Eocene; Specimens NHMUK In.25478 (well preserved impression of NW Isle of Wight, UK. isolated wing 2.6 mm long – Fig. 51) and I.8896 (partly pre- Material. NHMUK In.17402 (well preserved male without served body and wing) are not conspecific with D. priscula legs, Fig. 52, Pl. 5j), In.25470 (partly preserved wing), In.26055 due to much smaller size and can be identified only as Dixidae (well preserved wing without anal lobe, Fig. 53, Pl. 5k), Hooley inc. sedis. Collection. Description. First antennal segments indistinctly separated Family Culicidae Meigen, 1818 (probably the basal flagellomere elongate, cylindrical and no segments of flagellum fusiform). Scutum glabrous. Wing by Ryszard Szadziewski

1915 Culex protolepis Cockerell, p. 488, pl. 62, fig. 1 (Gurnet Bay, wing, sex not indicated) 1915 Culex petrifactellus Cockerell, 489, pl. 61, fig. 12 (Gurnet Bay, wing; sex not indicated) 1923 Aedes protolepis (Cockerell); Edwards, p. 142, figs 1, 2; pl. 7: figs 2–4 (Gurnet Bay, combination, syn. C. petrifactellus, male, female) 1944b Aedes protolepis (Cockerell); Statz: 114 (comments) 1994 Aedes protolepis (Cockerell); Evenhuis, p. 232 (in catalogue) 1998 Aedes protolepis (Cockerell); Szadziewski, p. 234 (comments)

Holotype. USNM. 61429, Lacoe Collection; Insect Bed; late Eocene; NW Isle of Wight, UK. Description (from Edwards 1923). Wing length 3.2–4.0 mm. Wing scales slender; subcostal vein short, ending at level of base of vein R2 þ 3; vein R2 þ 3 relatively long, only 1.4 times shorter then R2 (10:7). Male palpi are longer than proboscis with fourth and fifth segments of almost equal length. Male genitalia barely preserved; gonostyli rather long, slender, curved, with single apical tooth. Discussion. Cockerell (1915) briefly described Culex protolepis and Culex petrifactellus from single incomplete wings housed in the Smithsonian Institution, Washington. Sex was not indicated. Edwards (1923) suggested that C. protolepis is a female Figures 51–54 Dixidae. (51) Dixidae incertae sedis NHMUK In.25478, wing. (52)–(54) Dixella priscula (Cockerell, 1921) n. comb.: while C. petrifactellus a male of the same species. In the collec- (52) NHMUK In.17402, male; (53) NHMUK In.26055, wing; (54) tion of the NHM he determined a further 24 males and 14 NHMUK In.24345, holotype, wing. Scale bar ¼ 1 mm. females and completed a description of the species which was assigned to the genus Aedes. Edwards (l.c.) suggested that another 40 incomplete specimens probably belong to the The Culicidae is a well known family of nematocerous flies. same species. In the extant world fauna over 3200 species are known. They Reinert (2000) proposed to divide the composite genus are frequent parasites of mammals and birds as females feed Aedes Meigen into two genera Aedes and Ochlerotatus and on their blood. Mosquito larvae and pupae inhabit exclusively this proposition is widely accepted. The great majority of the shallow stagnant waters. extant Holarctic species belongs to the genus Ochlerotatus. Despite of the fact that the phylogenetic history of mosqui- The oldest fossil species in the genus is Ochlerotatus serafini toes goes back to the Mesozoic era, they are rarely preserved (Szadziewski 1998) from Eocene Baltic amber (Szadziewski & as fossils. The oldest true mosquitoes are reported from the Giłka 2011). Upper Cretaceous in Canadian amber (Poinar et al. 2000) In Ochlerotatus the gonostyli are long, narrow, somewhat and Burmese amber (Borkent & Grimaldi 2004). curved, with a single gonostylar tooth while in Aedes the Fossil mosquitoes from the Tertiary are reviewed by Edwards gonostyli have the distal portion expanded, bifurcated and (1923), Statz (1944a), Evenhuis (1994) and Szadziewski (1998). usually with more than one apical claw. Drawings of male Some named fossils are barely preserved and the visible genitalia of Aedes protolepis determined by Edwards (1923, characters show nothing more than that they are mosquitoes. fig. 1c–e) show that gonostyli are rather long, slender, curved, Mosquitoes are very rare in Eocene Baltic amber (Szadziewski with single apical tooth as in the genus Ochlerotatus. This 1998; Podenas 1999), while they are more common in younger indicates that the species should be placed in this genus. Miocene Dominican amber (Szadziewski & Grogan 1994; Poinar 2005), Miocene deposits of Rott (Statz 1944a) or the Genus Coquillettidia Dyar, 1905 Oligocene Insect Bed of the Isle of Wight described here. Dipterans of the family Culicidae are common fossils Type species. Culex perturbans Walker, 1856. among insects from the Bembridge Marls (Cockerell 1915; Edwards 1923). Over 140 specimens of adult mosquitoes and Coquillettidia cockerelli (Edwards, 1923) n. comb. one pupa have been found in the collection of the NHM (Figs 57, 58) (Edwards 1923; A. Ross, pers. comm.). This paper presents results of an examination of the collection of mosquitoes housed at the NHM, London. 1923 Taeniorhynchus (?) cockerelli Edwards, p. 151, fig. 4b (Gurnet Bay, female, fig. wing) Subfamily Culicinae Latreille, 1817 1944b Mansonia cockerelli Edwards; Statz, p. 114 (combination) Genus Ochlerotatus Lynch Arriba´lzaga, 1891 1994 Ochlerotatus cockerelli Edwards; Evenhuis, p. 234 (combination, in catalogue) Type species. Ochlerotatus confirmatus Lynch Arriba´lzaga, 1998 Mansonia cockerelli Edwards; Szadziewski, pp. 234, 1891 (¼Aedes scapularis Rondani, 1848). 243 (comments, in key)

Figures 55–58 Culicidae. (55), (56) Culex vectensis Edwards, 1923, holotype female, NHMUK I.9324: (55) habitus; (56) wing. (57), (58) Coquillettidia cockerelli (Edwards, 1923), holotype female, part and counterpart: (57) NHMUK I.10106; (58) NHMUK I.17190.

Chironominae (14). The other 16 barely preserved specimens length 3.4 mm; I.10060, one female, lateral view, no wings, remain undetermined. body 2.2 mm long; I.10217, one female, lateral position, thorax and abdomen 4.0 mm, all Brodie Collection; In.17470, Family Chironomidae Newman, 1834 one female, barely visible, dorsal position, Smith Collection; (Figs 59–64) In.25394, one male, lateral position, wings not preserved; In.25759, one male, barely preserved, body length 5.0 mm, Hooley Collection; all from the Insect Bed; late Eocene; NW Indeterminate material. NHMUK I.8475, one female, Isle of Wight, UK. lateral position; I.8530, one female, lateral position, no wings, length of preserved body 1.4 mm; I.8542, one female, lateral position, no wings; I.8991, one male, lateral position, no Subfamily Tanypodinae Frey, 1927 wings; I.9660, barely preserved remnants of three specimens; I.9782, one male, lateral position, no wings; I.9793, one Material. NHMUK I.9308, one female, wing 1.9 mm long; female, dorsal view, no wings; I.9880, one male, partly pre- I.9589, one female, lateral position, ﬂagellum short, with served; I.9964, one male, lateral position, no wings, body numerous ﬂagellomeres, body length 2.7 mm; I.9750, one

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Figures 59–64 Chironomidae. (59) Tanypodinae indet., female, NHMUK I.9921. (60) Orthocladiinae indet., male, NHMUK I.17390. (61) Chironominae indet., male, NHMUK I.9665. (62) Chironomini indet., male and female, NHMUK I.8912. (63) Chironomini indet., male, NHMUK I.9611. (64) Chironomini indet., male, NHMUK In.17388.

male, dorsal position, wings partly preserved, body 4.3 mm Material. NHMUK I.8912, one male, body 3.6 mm long, long; I.9921 (Fig. 59), one female, lateral position, flagellum one female, body 2.7 mm long, without wings (Fig. 62); composed of numerous flagellomeres, wings absent, all Brodie I.9611, one male, lateral position, wings not preserved, body Collection. length 4.0 mm (Fig. 63); I.9623, one female, lateral position, wings absent, body 4.2 mm long, Brodie Collection; In.17109, Subfamily Orthocladiinae Kiefer, 1911b one male, body 7.1 mm long, wings and genitalia not preserved; In.17388, three males, two females, one male very large, abdomen plus thorax 9.4 mm long (Fig. 64), Smith Material. I.17390(1), one male, body length without flagellum Collection; In.26040, part and counterpart, one female, lateral 1.9 mm (Fig. 60), together with Stilobezzia indet. (Ceratopogo- position, no wings, body 2.8 mm long, Hooley Collection. nidae), Smith Collection. Comments. Chironomids are rare fossils preserved in the Insect Bed of the Bembridge Marls. They are poorly preserved Subfamily Chironominae Newman, 1834 and have little taxonomic value. Among better preserved compression fossils from the early Miocene deposits of Rott in Material. NHMUK I.9665, one male with preserved genitalia, Germany the chironomids were represented by adults and body 4.5 mm long, in lateral position (Fig. 61); I.10071, two pupae assigned to the same subfamilies (Statz 1944b). females, lateral position, wing in one specimen visible, Brodie Collection. Family Ceratopogonidae Newman, 1834 Tribe Chironomini Newman, 1834 by Ryszard Szadziewski

Figures 65–68 Ceratopogonidae. (65) Stilobezzia sp. indet., NHMUK I.9809. (66), (67) Palpomyia edwardsi Cockerell, 1921, NHMUK I.9757. (68) Dasyhelea sp. indet., NHMUK I.9453.

Biting midges (Diptera: Ceratopogonidae) form a small n. comb. and genera Amberaspinus Evenhuis, 1994 and family of nematocerous ﬂies which is relatively well known in Aspinus Hong, 1981 are recognised as new junior synonyms the recent fauna and has well studied fossil record from the of Forcipomyia. However, two other species described by Lower Cretaceous to the Miocene. The family includes 5,471 Hong (1981) from females in the genus Aspinus (A. amblopteres extant species and 253 fossil species (Borkent & Wirth 1997; Hong, 1981 and A. stenopteres Hong, 1981) actually are Borkent, personal comm.). Fossil species from the Lower and members of the Chironomidae. I propose to leave these species Upper Cretaceous as well as from many Tertiary deposits in the original combination within the Chironomidae as species were described within 19 fossil and 25 extant genera. The best incertae sedis. known is the fossil fauna of biting midges in Eocene Baltic Dipterans of the family Ceratopogonidae are rare fossils amber (Szadziewski 1988). among insects from the Bembridge Marls. Until now only Hong (1981) described within the family Chironomidae a one biting midge Palpomyia edwardsi has been described new fossil genus Aspinus Hong, 1981, p. 52, with type-species from the collection of the NHM by Cockerell (1921). Aspinus orientalus Hong, 1981, p. 53 from Eocene Fushun Nine fossils of Ceratopogonidae from the NHMUK, were amber. Evenhuis (1994) discovered that the Aspinus was pre- examined. Nine of them were determined to genera, one barely occupied and replaced it with the name Amberaspinus Even- preserved specimen (NHMUK In.25394) remains undeter- huis, 1994, p. 260. The single known male of Amberaspinus mined. orientalus (Hong), based on the description and illustration by Hong (1981), is clearly a member of the genus Forcipomyia Tribe Ceratopogonini Newman, 1834 Meigen, 1818 of the family Ceratopogonidae. The overall habitus, presence of 13 ﬂagellomeres (10–13 elongated), elon- Genus Stilobezzia Kieffer, 1911a gated proboscis with relatively short palpi and the shape of gonocoxites and gonostyli indicate this generic placement. Type species. Stilobezzia festiva Kieffer, 1911a. The wing venation and the shape of tergite IX in genitalia are probably incorrect in the drawings. The type species Stilobezzia sp. indet. of Aspinus is here therefore named Forcipomyia orientalus, (Figs 65, 69)

Downloaded from https://www.cambridge.org/core. Chalmers Tekniska Högskola, on 06 Oct 2019 at 16:48:14, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S1755691018000464 TRUE FLIES FROM THE LATE EOCENE INSECT LIMESTONE OF THE ISLE OF WIGHT 33 Discussion. Hong (1981) described and illustrated a female of Palpomyia unca from Eocene amber of Fushun. Subsequently Hong et al. (2000) proposed a new genus Eopalpomyitis for this species. Actually the female of the type species has all femora armed with ventral spines, enlarged fore femora, two first radial cells well developed, sessile median veins, tarsal claws moderately long, equal to subequal, with basal inner tooth and other characters typical of the genus Palpomyia (Szadziewski 1988; Grogan, pers. comm.). Other characters used by Hong et al. (2000) for diagnosing a new genus like 14-segmented flagellum, first flagellomere with sensillum coeloconicum, costal vein prolonged to M1, sensory pit on third palpal segment are most probably a result of misinterpretations of structures not clearly visible in amber. In the biting midges antenna in both sexes never includes more than 13 flagellomeres, and within the tribe Palpomyiini the costal vein is never prolonged beyond R2 þ 3, sensilla coeloconica are absent on first flagellomere as well as the sensory pit is not developed Figures 69–70 Ceratopogonidae. (69) Stilobezzia sp. indet., on the third palpal segment. I suggest placing the generic NHMUK I.9809. (70) Palpomyia edwardsi Cockerell, 1921, NHMUK name Eopalpomyitis proposed by Hong et al. (2000) among I.9757. synonyms of the large, worldwide distributed extant genus Palpomyia.

Material. NHMUK I.9210, female, preserved one wing, part Palpomyia edwardsi Cockerell, 1921 of abdomen; I.9798, female, preserved one wing, abdomen in (Figs 66, 67, 70) dorsal view, scutum; I.9809, probably female, one wing 1921 Palpomyia edwardsi Cockerell, p. 469, fig. 31 (Gurnet and part of thorax preserved, Brodie Collection; In.17390(2), Bay, fig. wing, thorax) female?, wing incomplete, part of thorax preserved; In.17471, 1988 Palpomyia edwardsi Cockerell; Szadziewski, p. 240 female, one wing, thorax plus abdomen in lateral view pre- 1994 Palpomyia edwardsi Cockerell; Evenhuis, p. 251 served, Smith Collection; all from the Insect Bed; late Eocene; 1997 Palpomyia edwardsi Cockerell; Borket and Wirth, NW Isle of Wight, UK. p. 131 Description. Female. Wing length 2.40–2.51 mm, costal ratio CR 0.73–0.81. Venation typical of the genus Stilobezzia Holotype. NHMUK I.9757. One wing, thorax and partly Kieffe with median veins M1 and M2 petiolate and both first abdomen in lateral view preserved, Brodie Collection; Insect radial cells well developed (Figs 65, 69). Macrotrichia on wing Bed; late Eocene; NW Isle of Wight, UK. membrane not visible. Description. Female. Body length about 4.4 mm. Wing Male unknown. length 2.74 mm, CR 0.92. Second radial cell 2.47 times longer Discussion. Preserved wings of the fossils allow us to than first one. Median vein M2 forked proximal of cross-vein determine them as members of the extant genus Stilobezzia. r-m (Figs 66, 67, 70). Male unknown. However, they are poorly preserved females devoid of sub- Remarks. The generic position of this fossil determined generic and specific features which indicate the subgeneric solely by wing characters is not certain. Similar wing venation position or allow distinguishing them from other fossil species can be found also in some other genera of predatory biting known from Tertiary ambers and rocks. It seems that all midges within the tribes Sphaeromiini or Heteromyiini. How- specimens examined from the Bembridge Marls belong to one ever, legs which are necessary for final determination of the species. genus are not preserved. The relatively broad wing and stout Fossil Stilobezzia were reported from Paleocene amber of preserved part of abdomen suggest that it is a female. Sakhalin (Szadziewski 1990, one species indet.), Eocene Baltic Palpomyia is an extant genus of biting midges which amber (Szadziewski 1988, 1993, four sp.), Miocene Dominican evolved during Tertiary (Szadziewski 1996). It includes 11 fossil amber (Szadziewski & Grogan 1998a, two sp.), and Miocene species reported from Eocene Baltic amber (Szadziewski 1988, compression fossils from Rott (Statz 1944c; Szadziewski 1993, four sp.), Eocene Chinese amber (Hong 1981, one sp.), 1993, one sp.). Oligocene Bembridge Marls (one sp., see above), Miocene Stilobezzia is a worldwide distributed genus with 310 extant Dominican amber (Szadziewski & Grogan 1997, one sp.) and species (Borkent & Wirth 1997). Females are predators on Miocene nodules from California (Pierce 1966, four sp.). other small insects. Predatory larvae are good swimmers Palpomyia is a genus distributed worldwide that includes which live in small water bodies and semiaquatic habitats. 234 recent species (Borkent & Wirth 1997). Females are predators on small insects. In most cases the prey consists of males Tribe Palpomyiini Enderlein, 1936 of Chironomidae, or small Ephemeroptera. Swimming larvae of Palpomyia are common inhabitants of different types of Genus Palpomyia Meigen, 1818 aquatic and semiaquatic habitats where they feed on other small animals. 1818 Palpomyia Meigen, p. 82. Type species Ceratopogon fla- vipes Meigen, 1804 Subfamily Dasyheleinae Lenz, 1934 2000 Eopalpomyitis Hong, Guo & Ren, p. 226. Type species Genus Dasyhelea Kieffer, 1911c Palpomyia unca Hong, 1981, by original designation. Eopalpo- myitis: Hong, 2002, p. 165. Syn. nov. Type species. Dasyhelea halophila Kieffer, 1911c.

2002 Palpoimyitis Hong, p. 165. Nomen nudum Dasyhelea sp. indet. 2002 Palpomyitis Hong, p. 165. Nomen nudum (Fig. 68)

Downloaded from https://www.cambridge.org/core. Chalmers Tekniska Högskola, on 06 Oct 2019 at 16:48:14, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S1755691018000464 34 WIESŁAW KRZEMIN´ SKI ET AL. Material. NHMUK I.9453, female, wings absent, thorax Material: The single simuliid pupa was collected from late and abdomen in lateral view and flagellum preserved, doubt- Eocene deposits of the Isle of Wight, southern England. It is ful determination, Brodie Collection; In.25374/In.25405 (part deposited in the palaeontological collection of the NHMUK and counterpart), female, one wing, thorax and abdomen under collection no. In.36778. Both part and counterpart are partly preserved; Insect Bed; late Eocene; NW Isle of Wight, present. On the same pieces of limestone is a specimen of a UK. female Culex protorhinus Cockerell 1915 (Culicidae) is located Description. Female. Body length 1.1 mm, wing length which was mentioned by Edwards (1923) under its old collec- 0.8 mm, costal ratio CR about 0.4. Radial cells barely visible. tion no. H. 540 (former Hooley Collection, now included in Remarks. The shape of partly preserved flagellum, wing the collection of the NHMUK). venation and total habitus show that most probably it is a Methods: The fossil was examined with a Leica MZ 9.5 species of the genus Dasyhelea. The oldest records of Dasyhelea Stereomicroscope. Drawings were produced directly from the are from Eocene Baltic amber (Szadziewski 1988, 1993, four specimen with a camera lucida attachment on a Leica MZ 9.5 sp.). The genus was also reported from Miocene Dominican Stereomicroscope. Photographs were made with a Leica MZ amber (Szadziewski & Grogan 1998b, four sp.), Mexican 9.5 Stereomicroscope, and a Nikon Coolpix 4500 digital amber (Szadziewski & Grogan 1996, one species indet.) and camera. Miocene nodules from California (Palmer 1957; Pierce 1966, Terminology and systematics: Morphological terminology six sp.). follows Adler et al. (2004). Current systematic placement of The phyto- and/or saprophagous larvae of Dasyhelea are Simuliidae is in Culicomorpha (Yeates & Wiegmann 2005). A aquatic and usually inhabit small and shallow bodies of water. recent molecular analysis by Beckenbach & Borkent (2003) The genus is distributed worldwide and includes 466 recent suggests that Simuliidae is the sister group of (Ceratogogoni- species (Borkent & Wirth 1997). Adults of both sexes feed on dae þ Chironomidae). For Simuliidae the phylogeny given in honeydew or visit flowers for nectar. Adler et al. (2004) is followed. Discussion. Biting midges are rare fossils preserved in the Insect Bed of Bembridge Marls. They are poorly preserved Family Simuliidae Newman, 1834 and have a little taxonomic value. The examined Ceratopogo- Subfamily Simuliinae Newman, 1834 nidae represent large extant genera of worldwide distribution: Gen. et sp. indet. Palpomyia, Stilobezzia and Dasyhelea. Their larvae live in (Figs 71–74) aquatic and semiaquatic habitats. Larvae and female adults of Palpomyia and Stilobezzia are predaceous while those of Description. The fossil pupa is preserved in the special three- Dasyhelea phyto- and/or saprophagous. Females of the latter dimensional mode of preservation typically found in insects genus feed on nectar of flowers and honeydew like males of from the Isle of Wight. The counterparts of the fossil show biting midges. completely different aspects of the fossil. The former body left Among better preserved compression fossils from the early behind a cavity which reveals morphological features rich in Miocene deposits of Rott in Germany, the biting midges were detail. In this fossil probably also remains of the former cuticle represented by five genera: Atrichopogon Kieffer, Bezzia are preserved which have a dark colour. Kieffer, Culicoides Latreille, Stilobezzia Kieffer and Serromyia Length of the visible part of the body (without gills) is Meigen (Statz 1944c). As in Bembridge Marls they are about 2.5 mm, both from the ventral side (In.36778a, H. 540) exclusively extant genera of worldwide distribution which are and from the dorsal side (In.36778b, H. 229). Present are common in recent semiaquatic and aquatic habitats. head, thorax and two basal segments of the abdomen. The apical abdominal segments are not visible, probably they are Family Simuliidae Newman, 1834 still embedded in the sediment. The estimated length of the complete pupa probably would have been 5 mm. Maximum by Sonja Wedmann width of the preserved thorax is 1.75 mm. In.36778a shows the pupa from its ventrolateral view (Figs Fossil black flies are very rare and fossils of immature simu- 71, 73). The pupa is of the obtect type, its appendages are not liids are rarer still. Compared to the more than 2100 described free but soldered to the body. extant species (Adler & Crosskey 2011), there are only very The head lies anteroventral to the thorax. An imprint of the few fossil specimens recorded. Evenhuis (1994) gave an over- basal part of one antennal sheath is visible at the uppermost view of the fossil record of Simuliidae. Updated information part of the head. Both eyes, parts of the mouthparts and other on fossil simuliids (excluding subfossil specimens) is discussed structures are discernable in the head and in the ventral by Currie & Grimaldi (2000). Up to now, there are only 12 thoracic region. Imprints of the sheaths of the rather long fossil simuliid species described, which are listed in Adler & maxillary palps can be seen clearly. No special surface sculp- Crosskey (2011). But in the private collection of Christel and ture on head and thorax is discernible, and no cephalic or Hans Werner Hoffeins there are several yet undescribed inclu- thoracic trichomes can be identified. sions of adult black flies in Baltic and Bitterfeld amber (C. & As in In.36778b the prothoracic gills are partly visible. They H. W. Hoffeins 2007, pers. comm.). The oldest undisputed are arborescent, with many filaments preserved only frac- simuliid fossil records are from the Jurassic/Cretaceous of tionally. The imprints of the leg sheaths of the first and second Siberia (Currie & Grimaldi 2000). pair of legs are conspicuous. The imprints of the wing pads are Additionally, there are a few undetermined immature simu- preserved completely and show an uneven surface. In the tips liid fossils, e.g., from the Lower Cretaceous of Australia (Jell of the wing sheaths there seem to be some dark patches of & Duncan 1986; Currie & Grimaldi 2000). Because of the pigmented cuticle preserved. The rest of the body is missing. scarcity of records of immature simuliids, the fossil record of No trace of a cocoon is preserved. the pupa from the late Eocene Insect Bed of the Isle of Wight In.36778b shows the pupa in dorsal view with most of the is interesting even if its exact systematic placement cannot be abdomen not visible (Figs 72, 74). The thorax is enlarged determined. and most probably kind of hump-backed or arched, but no

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Figures 71–74 Simuliid pupa gen. et sp. indet. (71), (72) Photographs. (73), (74) Drawings. In the drawings, black colour indicates areas where dark pigment is present in the fossil, dotting illustrates three-dimensional preservation. (71), (73) NHMUK In.36778a, showing the ventral side of the fossil. (72), (74) NHMUK In.36778b, showing the dorsal side of the fossil. Scale bar ¼ 1 mm. Abbreviations: a1 ¼ abdominal segment 1; a2 ¼ abdominal segment 2; ant ¼ antennal sheath; e ¼ eye; g ¼ gill; l1 ¼ sheath of leg 1; l2 ¼ sheath of leg 2; mxp ¼ sheath of maxillary palp; psb ¼ postscutellar bridge; w ¼ wing sheath.

trichomes are discernible. The dark former integument pre- smooth. Anterior to the first abdominal segment the served in the thoracic region laterally shows a structure con- postscutellar bridge is visible. Only the first and second basal sisting of small fissures, but this could also be secondary due abdominal segments are preserved. Most of the abdomen is to diagenetic processes during fossilisation. not visible. At each anterolateral corner of the prothorax one gill is Owing to the very well preserved three-dimensional struc- present which consists of many long, slender branches. In the ture of this fossil imprint, it seems most probable that the fossil there is no basal stalk visible. The number of the main fossil was a pupa from which the adult fly had not emerged trunks of the gill cannot be seen, only two or three thickened yet. A moulted pupal skin could not generate such a good basal parts are preserved. The number of secondary filaments three-dimensional imprint. With the preserved structures it is is high, approaching 40 to 50 fine secondary filaments per gill. not possible to determine the sex of the pupa. The surface sculpture of the gill is not discernible in detail, but Discussion: Systematic placement. The fossil pupa can be with 60-fold magnification its surface seems to be quite placed systematically in Simuliidae: Simuliinae because

as Parasimulium, the prosimuliines, and basal simuliines the ends 0.15 mm distal of apex of R4 þ 5; Sc ends just basal of cocoon is quite shapeless and saclike (Adler et al. 2004, pterostigma; R1 reaches the costal margin in the pterostigma; p. 50). A possible reason for the lack of a cocoon in this fossil basal part of R4 þ 5 is short, 0.15 mm long in mean, distinctly might be that the pupa was ripped out of its cocoon and got shorter than r-m; r-m is 0.305 mm long in mean; distance carried away by strong water currents. There are also several between base of r-m and fork of M into M1 and M2 0.72 mm simuliid taxa whose pupae are fixed to the substrate by only a in mean; cross-vein bm-cu slightly basal of fork of M or few threads of cocoon material (Crosskey 1990, p. 296), and ending on it; bm-cu 0.30 mm long; A1 well defined, reaching this might be another explanation for the lack of a cocoon in posterior wing margin. the fossil specimen. Remarks. These fossils are attributable to the same species The length of pupal life varies with the water temperature of because of their very similar wing venation and proportions, the habitat. Under tropical conditions the adult fly can emerge and to Dilophus rather than to Bibio or Bibiodes because of within two days of pupation, in cold climates this period the presence of only two small apical spines on fore tibia rarely exceeds 14 days (Crosskey 1990, p. 320). (Hardy 1981). The very long r-m cross-vein two times as long The adult flies feed on the nectar of a wide variety of plants as basal part of Rs is a very secondary character that could be (Crosskey 1990, p. 388). Additionally, the females of many present in Dilophus, but that also occurs in the genus Bibio. species feed on the blood of warm-blooded vertebrates This structure can be very variable in the same species of a (Crosskey 1990, p. 411). recent Bibio,viz.B. hortulanus. Among the fossil bibionids from the Isle of Wight, Bibio gurnetensis Cockerell, 1917a has Superfamily Bibionoidea Fleming, 1821 also a very long r-m, about three times as long as basal part of Rs, but this species is distinctly larger than D. andrewrossi, by Andre´ Nel, François-Marie Collomb and Alain Waller with a wing 4.8 mm long. Comparison of D. andrewrossi with the recent species is impossible to achieve because of the lack Family Bibionidae Fleming, 1821 of the genital structures in our material. The fossil record of The past diversity of Bibionidae is difficult to estimate. Dilophus is extremely scarce compared to that of Bibio, with Their systematic is far from being easy as these insects have eight species, Dilophus campbelli Harris, 1983 (larva, New an important intraspecific variability, in body size and wing Zealand, Eocene), D. krantzii Heyden, 1870 (Rott am venation (see above). Also, several characters that can be Siebengebirge, Germany, Oligocene), D. magnus Du¨rrenfeldt, used for recent species (male anal appendages, body and wing 1968 (Germany, Pliocene), D. priscus Loew, 1850 (Baltic amber, Eocene/Oligocene), D. matilei Waller et al., 2000

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Figures 75–95 Bibionidae. (75–78) Dilophus andrewrossi Nel, Colomb & Waller, sp. n.: (75) holotype counterpart NHMUK I.9235; (76) paratype NHMUK In.26048; (77) holotype part NHMUK I.9073; (78) paratype NHMUK I.9148. (79–81) ?Bibio gurnetensis Cockerell, 1917a: (79) holotype part NHMUK I.8641; (80) NHMUK I.9604 (type of Bibio oblitus Cockerell, 1921 syn. nov.); (81) holotype counterpart NHMUK I.8900. (82–84) ?Bibio oligocenus Cockerell, 1917a: (82) holotype NHMUK I.8650; (83) NHMUK In.32283; (84) NHMUK In.25243. (85) ?Bibio extremus Cockerell, 1921, holotype NHMUK I.8860. (86) ?Bibio sp., NHMUK I.9363. (87) Bibiodes conﬂuens (Cockerell, 1915) n. comb., NHMUK In.43454 (scale bar ¼ 4 mm). (88) Plecia acourti Cockerell, 1921, holotype NHMUK In.17098/In.17500. (89–92) Plecia sp.: (89) NHMUK In.26049; (90) NHMUK In.17230/In.17474; (91) NHMUK In.25249; (92) NHMUK In.20558/In.20557. (93) Plecia undans Zeuner, 1941, holotype NHMUK In.37274. (94, 95) Penthetria sp.: (94) NHMUK In.24800; (95) NHMUK In.25246. (Scale bars ¼ 1 mm if not stated otherwise.)

Plate 6 Bibionidae. (a) Dilophus andrewrossi Nel, Colomb & Waller, n. sp. holotype counterpart NHMUK I.9235. (b–d) ?Bibio gurnetensis Cockerell, 1917a: (b) holotype part NHMUK I.8641. (c) NHMUK I.9604 (type of Bibio oblitus Cockerell, 1921 syn nov.). (d) holotype counterpart NHMUK I.8900. (e) ?Bibio oligocenus Cockerell, 1917a, holotype NHMUK I.8650. (f ) Bibio obliteratus (Cockerell, 1915) n. comb., holotype USNM 61442. (g) ?Bibio extremus Cockerell, 1921, holotype NHMUK I.8860. (h) ?Bibio sp., NHMUK I.9363. (i) Bibiodes conﬂuens (Cockerell, 1915) n. comb., NHMUK In.43454. (j) Plecia acourti Cockerell, 1921, holotype counterpart NHMUK In.17500. (k) ?Plecia sp., NHMUK In.26049.

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Table 2 Dilophus andrewrossi sp. n. Main wing dimensions of the type material. Abbreviations: L wing ¼ wing length; W wing ¼ wing width; L r-m ¼ length of r-m; L Rs base ¼ length of basal part of Rs; L bm-cu ¼ length of bm-cu.

Specimen/dimensions L wing W wing L r-m L Rs base L bm-cu I.9235 3.9 mm 1.5 mm 0.29 mm 0.15 mm 0.30 mm I.26048 3.8 mm 1.4 mm 0.30 mm 0.18 mm 0.30 mm I.9073 4.0 mm 1.51 mm 0.30 mm 0.13 mm 0.28 mm I.9148 3.7 mm ? 0.33 mm 0.15 mm ?

(Dominican Republic amber, Miocene), D. palaeofebrilis Sc ends just basal of pterostigma; R1 reaches the costal margin Skartveit, 2009, D. crassicornis Skartveit, 2009 and D. succineus in the pterostigma; basal part of R4+5 is short, 0.20–0.26 mm Skartveit 2009 (Evenhuis 1994; Waller et al. 2000; Skartveit long, three times shorter than r-m; r-m 0.60–0.64 mm long; 2009). Statz (1943, p. 56) added an undescribed record from distance between base of r-m and fork of M into M1 and M2 the Late Oligocene of Aix-en-Provence, France. This record is 0.9–1.0 mm; cross-vein bm-cu slightly basal of fork of M or very unlikely: in 26 years we have never collected any represen- ending on it; bm-cu 0.44–0.49 mm long; A1 well defined, tatives of this genus among the thousands of bibionids from reaching posterior wing margin. this outcrop. Remarks. Cockerell (1917a, pp. 375, 384, pl. 31, fig. 4; 1921, The original description and figures of D. krantzii are not p. 468, fig. 29) described B. gurnetensis and B. oblitus on the very convincing and this species should be redescribed. Never- basis of rather poorly preserved wings. There is no correct theless, its wing is distinctly longer than that of D. andrewrossi, character to separate to the two species. All these fossils are i.e., about 7.9 mm long, instead of 3.85 mm in D. andrewrossi attributable to the same species because of their very similar (Heyden 1870, p. 255, pl. 45, fig. 24). The wings of D. wing venation and proportions, especially in the large wing matilei are shorter: 2 mm long (Waller et al. 2000). The and the very long r-m, about three times as long as basal part wings of D. magnus are distinctly longer: 8.0 mm long (Du¨rren- of Rs. Nevertheless, its attribution to the genus Bibio rather feldt 1968, p. 50). D. palaeofebrilis, D. crassicornis and D. than Dilophus is impossible to establish as there is no wing succineus have wing length similar to that of D. andrewrossi, character that would be sufficient to separate the two genera. between 3.15 and 3.80 mm. The r-m cross-vein is distinctly more than two times as long as basal part of Rs in both D. ?Bibio oligocenus Cockerell, 1917a palaeofebrilis and D. crassicornis. D. andrewrossi has only two (Figs 82–84, Pl. 6e) small apical spines on fore tibia, instead of four or five in D. palaeofebrilis, D. crassicornis, and D. succineus. D. priscus 1917a Bibio oligocenus Cockerell, pp. 375, 384, pl. 62, fig. 3a–d has apparently never been revised and is very poorly known, 1994 Bibio oligocenus Evenhuis, p. 125 by the following information concerning the strong thoracic setae: ‘Von der Gattung Dilophus ist mir nur ein einziges Holotype. NHMUK I.8650 (Fig. 82); Insect Bed; late Weibchen vorgekommen, welches in der Stellung der dicken Eocene; NW Isle of Wight, UK. Dornen auf dem Thorax von allen bei uns einheimischen Arten Description. Wing hyaline, about 9.1 mm long, 3.7 mm abweicht. Ich nenne die Art Dilophus priscus.’ (Loew 1850, wide; pterostigma elliptic, clearly sclerified, 1.3 mm long, p. 38). Thus it is impossible to compare to our fossils. 1.42 mm wide; C ends just distal of apex of R4+5;ScandR1 reach the costal margin in the pterostigma; basal part of R4+5 Tribe Bibionini Frey, 1927 is long, 0.82 mm long, three times longer than r-m; r-m 0.27 mm long; distance between base of r-m and fork of M Subtribe Bibionina Frey, 1927 into M1 and M2 unknown; position of cross-vein bm-cu Genus Bibio Geoffroy, 1762 unknown; A1 well defined. Remarks. Cockerell (1917a, pp. 375, 384, pl. 31, fig. 5) described and figured this species on the basis of an incom- Type species. Tipula hortulana Linnaeus, 1758, by subsequent plete and rather poorly preserved wing. Its main character is designation of Latreille (1810, p. 442) (as ‘Hirtea hortulana its very large wing, with a long basal part of Rs. It is probably Fabr.’). (Validated by I.C.Z.N. (1957a, p. 86).) a Bibio but the possible attribution to Dilophus cannot be excluded. Two other specimens In.32283 (Fig. 83, Colenutt ?Bibio gurnetensis Cockerell, 1917a Collection) and In.25243 (Fig. 84, Hooley Collection) with (Figs 79–81, Pl. 6b–d) very large wings could belong to the same species, but their 1917a ?Bibio gurnetensis Cockerell, p. 375, 384, pl. 31, fig. 4 wings are slightly shorter and narrower than that of the type 1921 Bibio oblitus Cockerell, p. 468, fig. 29 syn. nov. of B. oligocenus and their r-m are very short or completely 1994 Bibio gurnetensis Evenhuis, p. 124 reduced. 1994 Bibio oblitus Evenhuis, p. 125 Bibio obliteratus (Cockerell, 1915) n. comb. Pl. 6f Holotype. NHMUK I.8641/I.8900 (part and counterpart 1915 Protoberis obliterata Cockerell, pp. 494, 499, pl. 63, Figs 79, 81, Pl. 6b, d), Brodie Collection; Insect Bed; late figs A,B Eocene; NW Isle of Wight, UK. 1992 Protoberis obliterata Carpenter, p. 422 Additional material. I.9604 (type of Bibio oblitus, Fig. 80, 1994 Protoberis obliterata Evenhuis, p. 309 Pl. 6c). Description. Wing hyaline, 4.8–4.9 mm long, 1.8–2.0 mm Remarks. Cockerell (1915, pp. 494, 499, pl. 63, fig. 1a, b) wide; pterostigma elliptic, clearly sclerified, 0.7–0.8 mm long, described and figured a very poor specimen (holotype USNM 0.2–0.3 mm wide; C ends 0.1–0.2 mm distal of apex of R4+5; 61442) he attributed to the Stratiomyidae under the name

Downloaded from https://www.cambridge.org/core. Chalmers Tekniska Högskola, on 06 Oct 2019 at 16:48:14, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S1755691018000464 40 WIESŁAW KRZEMIN´ SKI ET AL. Protoberis obliterata. A simple examination of the photo of 1915 Bibiodites confluens Cockerell, pp. 493, 499, pl. 62, the type specimen shows that it is a Bibioninae, and probably fig. 3a–d a Bibio. Its large size (wing length about 8 mm) suggests that it 1992 Bibiodites confluens Carpenter, p. 414 could correspond to the same species as B. oligocenus,butwe 1994 Bibiodites confluens Evenhuis, p. 126 have no definite proof of this hypothesis. We propose to name it Bibio obliteratus n. comb. Holotype. USNM 61440, Lacoe Collection; Insect Bed; late Eocene; NW Isle of Wight, UK. ?Bibio extremus Cockerell, 1921 Material. In addition to holotype specimen NHMUK (Fig. 85, Pl. 6g) In.43454 (Fig. 87, Pl. 6i), Hooley Collection. Description. (Based on specimen In.43454) An isolated 1921 Bibio extremus Cockerell, p. 486, fig. 30 wing, hyaline, about 4.32 mm long (about 5 mm in the holo- 1994 Bibio extremus Evenhuis, p. 124 type), 1.66 mm wide; pterostigma elliptic, clearly sclerified, 0.82 mm long, 0.25 mm wide; C ends just distal of apex of Holotype. NHMUK I.8860, Brodie Collection; Insect Bed; R4+5; Sc reaches the costal margin just basal of the pteros- late Eocene; NW Isle of Wight, UK.. tigma and R1 in the pterostigma; basal part of R4+5 is long, Description. Wing hyaline, about 5.5 mm long, 2.0 mm 0.27 mm long; Rs fused with M for 0.25 mm; cross-vein wide; pterostigma elliptic, clearly sclerified, 0.82 mm long, bm-cu deformed; A1 present. 0.27 mm wide; apex of C not preserved; Sc and R1 reach the Remarks. The attribution of In.43454 to Bibiodes confluens costal margin in the pterostigma; basal part of R4+5 is long, is based on its very close wing dimensions. Its fusion Rs-M 0.42 mm long, r-m completely reduced and Rs and M touch- slightly longer than in the type of this species supports its attri- ing in one point; distance between this point and fork of M bution to Bibiodes. into M1 and M2 1.0 mm; cross-vein bm-cu 0.42 mm long; A1 well defined. Subfamily Plecinae Duda, 1930 Remarks. The figure by Cockerell (1921, p. 468, fig. 30) is partly erroneous as he figured a short r-m cross-vein. It is Genus Plecia Wiedmann, 1828 smaller than B. oligocenus and longer than Bibiodes confluens (see below), but the complete reduction of r-m suggests possible Type species. Hirtea fulvicollis Fabricius, 1805, by subse- affinities with Bibiodes, which cannot be definitely demon- quent designation of Blanchard (1840, p. 576). strated. Another specimen (I.9363, see Fig. 86, Pl. 6h) has also its r-m completely reduced and Rs and M touching in one Plecia acourti Cockerell, 1921 point, but its wing is shorter than that of B. extremus (4.5 mm (Fig. 88, Pl. 6j) long instead of 5.5 mm in the latter). I.9363 could correspond to a different species but this difference in size is not sufficient 1921 Plecia acourti Cockerell, pp. 467–68, fig. 28 to clearly separate it from B. extremus. We prefer to consider it 1994 Plecia acourti Evenhuis, p. 129 as a Bibio? species. Holotype. NHMUK In.17098/In.17500 (part and counter- Subtribe Bibiodina Frey, 1927 part), Smith Collection; Insect Bed; late Eocene; NW Isle of Wight, UK. Genus Bibiodes Coquillett, 1904 Description. This species is based on a rather poorly pre- 1904 Bibiodes Coquillett, p. 171 served female specimen (abdomen, part of thorax and one 1915 Bibiodites Cockerell, p. 493; syn. nov. wing) (Cockerell 1921, pp. 467–68). Wing infuscate, 6.4 mm long, 2.5 mm wide; pterostigma not visible; basal part of Rs long, 1.0 mm long, r-m 0.26 mm long; R2+3 separates from Type species. Bibiodes halteralis Coquillett, 1904, by original R4+5 1.5 mm distal of r-m; R2+3 nearly straight, 0.78 mm designation. long, strongly diverging from R4+5; distance between r-m and The fossil record of this recent Nearctic genus comprises fork of M into M1 and M2 0.58 mm; cross-vein bm-cu 0.6 mm Bibiodes intermedia James, 1937, B. nanus Skartveit, 2009 and basal of r-m, 0.3 mm long; A1 well defined. Abdomen broad B. balticus Skartveit, 2009. Cockerell (1915, p. 493) separated (female), but deformed. his fossil genus Bibiodites from Bibiodes on the basis of its Remark. The generic attribution of this species is confirmed, shorter confluence between Rs and M. The ratio (length but it is nearly impossible to compare it to the other Cenozoic of confluent Rs-M/wing length) is 0.09 in the recent Bibiodes Plecia. aestivus Melander, 1912, 0.05 in the type specimen of Bibio- dites confluens, 0.06 in specimen In.43454, 0.0325 in the type ?Plecia spp. specimen of Bibiodes intermedia (Cockerell 1915; James 1937) and 0.03 in Bibiodes massiliensis sp. n. The differences are too Among the Plecia from the Isle of Wight, three other poten- weak to justify a generic separation. Thus we support the syn- tial ‘species’ can be defined: onymisation of the two genera and the transfer of Bibiodites confluens Cockerell, 1915 into the genus Bibiodes, as proposed e A smaller one (specimen In.26049, wing length 4.3 mm), by Skartveit (2009). The presence of the Nearctic genus Bibiodes with R2+3 strongly curved (Fig. 89, Pl. 6k) in the Eocene and Oligocene of Europe is of great interest for e A larger one (specimen In.17230/In.17474, wing length its age and past distribution. Its relatively recent discovery in 10.4 mm), with R2+3 elongate and sigmoidal (Fig. 90, China (Bibiodes sinensis Yang & Luo 1987) supports its ancient Pl. 9c) presence in the Palaearctic region (Yang & Luo 1987). e A species of nearly the same size as Plecia acourti (specimens In.25249 (Fig. 91, Pl. 9a), wing length 8.25 mm and Bibiodes confluens (Cockerell, 1915) n. comb. In.20558/In.20557 (Fig. 92), wing length 9.1 mm), with (Fig. 87, Pl. 6i) R2+3 very long and nearly parallel to R4+5, but less than in

Downloaded from https://www.cambridge.org/core. Chalmers Tekniska Högskola, on 06 Oct 2019 at 16:48:14, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S1755691018000464 TRUE FLIES FROM THE LATE EOCENE INSECT LIMESTONE OF THE ISLE OF WIGHT 41 the Penthetria from the same outcrop (see below). The wing widely used in the taxonomy of the subfamily. Until further of this species is very similar to that of Plecia undans advance in classification of fungus gnats it is possible to Zeuner, 1941 from the early Eocene of the Isle of Mull say that Mycetophila vectensis Cockerell, 1915 (Pl. 7g), (UK). ‘Rymosia’ edwardsi Cockerell, 1921 (Pl. 7d), ‘Rymosia’ ferruginea Cockerell, 1921 (Pl. 7e) and ‘Rymosia’ rufescens We prefer to maintain these fossils as ‘Plecia species un- Cockerell, 1921 (Pl. 7f) most probably belong to the tribe determined’, in order to avoid increasing confusion in the Mycetophilini; Rymosia grisea Cockerell, 1921 (Pl. 7h) is a Cenozoic bibionid taxonomy. representative of the subfamily Mycetophilinae. These species will not be redescribed in the present paper. Plecia undans Zeuner, 1941 Fifteen specimens of the subfamily Sciophilinae belong to (Fig. 93) 10 species of eight genera: Acnemia, Azana, Aglaomyia, Coelosia, Dziedzickia, Palaeoempalia, Mycomya and Leia,mostofthem, except Palaeoempalia, are found in recent fauna. All the genera, Holotype. NHMUK In.37274. except Aglaomya, are known from Baltic amber. Aglaomya can Additional material. Paratype NHMUK In.37266. be easily mistaken for Boletina, known from the Baltic amber Age and outcrop. Early Ypresian, early Eocene, Ardtun and Oligocene and Miocene of Europe. A species of Mycomya Head, Isle of Mull, UK. Evenhuis (1994, p. 133) indicated is also known from Palaeocene amber of the Sakhalin Island that this outcrop was Late Cretaceous, but it is now con- (Blagoderov 2007). sidered as Early Eocene (Royer et al. 2001). The detailed comparison of the fauna can be made only Description. Zeuner (1941, pp. 94–95, figs 13, 14) proposed after revision of the Baltic amber material. However, an exact figures of the type material. We had the opportunity to apparent observation can be made. From 155 species of revise the paratype specimen. Wing about 11.0 mm long, Mycetophilidae described from Baltic amber, 126 belong to about 3.52 mm wide; pterostigma not visible; basal part of Rs the subfamily Sciophilinae, while more than a half of species long, 1.12 mm long, r-m 0.4 mm long; R2+3 separates from (12) and more than two-third of specimens of the Isle of Wight R4+5 1.2 mm distal of r-m; R2+3 very long, sigmoidal, and fungus gnats are from the subfamily Mycetophilinae. Larvae nearly parallel to R4+5, 2.3 mm long; distance between r-m of Mycetophilinae are almost obligatory mycetophagous, and fork of M into M1 and M2 0.56 mm; cross-vein bm-cu developing on fruiting bodies of mushrooms. Sciophilinae, on 1.2 mm basal of r-m, 0.2 mm long. the other hand demonstrate a number of various biologies, including sapromycetophagy, zoomycetophagy, xylomycetophagy Subfamily Penthetriinae Enderlein, 1936 and occasionally eumycetophagy. The great diversity and abundance of mycetophiline fungus gnats in the Isle of Wight Genus Penthetria Meigen, 1803 oryctocenosis allows us to reconstruct warm temperate and (Figs 94, 95, Pl. 9b) rather humid forest environment.

Type species. Penthetria funebris Meigen, 1804, by subse- Family Mycetophilidae Newman, 1834 quent monotypy in Meigen (1804, p. 104). Subfamily Sciophilinae Rondani, 1840 Penthetria sp. Remarks. We discovered in the NHM collection two Genus Acnemia Winnertz, 1863 specimens, In.24800 (Fig. 94) and In.25246 (Fig. 95, Pl. 9b) (both Hooley Collection), attributable to this genus that was 1863 Acnemia Winnertz, p. 798. Type species Leia nitidicollis previously unrecorded from the Isle of Wight. The genus Meigen, 1818, by subsequent designation by Johannsen is rather frequent in the Oligocene and Miocene of Europe. 1909, p. 63 Its oldest records date from the Late Eocene of Monteils 1865 Agaricobia Philippi, p. 626. Type species Agaricobia (undescribed species, France, see above), the North American fulvicollis Philippi, 1865, by monotypy Eocene, namely Penthetria whipsawensis Rice, 1959 and P. creedensis (James 1938) and two Baltic amber species Remarks. The genus includes about 40 species distributed (Fitzgerald 1999; Skartveit 2009). It is not necessary to create worldwide, mainly from temperate to warm temperate zones. new species for the material of the Isle of Wight because it would hardly be comparable with the already described species. Acnemia simplex Cockerell, 1921 (Pl. 8a) Superfamily Sciaroidea Billberg, 1820 1921 Acnemia simplex Cockerell, pp. 463, 464, ﬁg. 19 by Vladimir Blagoderov 1994 Acnemia simplex Evenhuis, p. 146

The fungus gnats of the late Eocene of the Isle of Wight are Holotype. NHMUK I.9597, Brodie Collection; Insect Bed; surprisingly diverse and rather abundant. Keroplatidae are late Eocene; NW Isle of Wight, UK. represented by seven species (17 specimens) and Mycetophilidae Diagnosis. The species differs from A. bolsiusi Meunier, by 22 species (55 specimens studied). Cockerell (1915, 1921) 1904 in having very short section M2 (M fork is 2.5 times the described eight species of Mycetophilidae and one species of section M3 in the latter species). It differs from A. cyclosoma Keroplatidae. Although all described taxa constitute good Cockerell, 1925 in smaller size and having RS base more species, their taxonomic position should be reconsidered. proximal (at 0.4 times the wing length in the latter species). Seventy-two per cent of the studied specimens of mycetophilids Description. Wing, 2.5 mm. Wing membrane with macrotri- belong to the subfamily Mycetophilinae. Among 40 specimens chia decumbent. Costa with two rows of setae, all veins except of Mycetophilinae from the collection of the NHM one could RS and section M3 with macrotrichia. Sc2 at the level of base distinguish up to 12 different species, but their generic assign- of r-m. Terminal part of Sc (distad of Sc2) three time the r-m. ment remains unclear. Characters of wing venation are not Costa continues beyond the tip of R5 at 1/5 of distance

Plate 7 Tipulidae, Limoniidae, Mycetophilidae and Sciaridae. (a) Leptotarsus (Longurio) wegiereki Krzeminski, n. sp., NHMUK In.17493. (b) Ilisia (Lunaria) incognita (Cockerell, 1921) n. comb., NHMUK I.9186. (c) Paltostomopsis ciliata Cockerell, 1915, holotype, USNM 61433 (Tipulomorpha indet.). (d) ‘Rymosia’ edwardsi Cockerell, 1921 NHMUK I.9336. (e) ‘Rymosia’ ferruginea Cockerell, 1921, NHMUK I.9802. (f ) ‘Rymosia’ rufescens Cockerell, 1921, NHMUK I.8865. (g) Mycetophila vectensis Cockerell, 1915, USNM 61434. (h) Rymosia grisea Cockerell, 1921, NHMUK I.8956. (i) Sciara gurnetensis Cockerell, 1915, USNM 61435. (j) Sciara protoberidis Cockerell, 1915, USNM 61437. (k) Sciara lacoei Cockerell, 1915, USNM 61436.

Downloaded from https://www.cambridge.org/core. Chalmers Tekniska Högskola, on 06 Oct 2019 at 16:48:14, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S1755691018000464 TRUE FLIES FROM THE LATE EOCENE INSECT LIMESTONE OF THE ISLE OF WIGHT 43 between tips of R5 and M1. Length of r-m about the length of of r-m, and length of RS base is 0.5 times the length of r-m. section M3 and 1.2 times the length of RS1. Distance from the Section M2 is 1.87 the section M3. Base of fork M3 þ 4and wing base to RS is 0.28 times the wing length. Length of Sc is CuA proximad of the base of section M2. 0.37 times the length of wing; length of R1 is 0.65 times the Remarks. The species is assigned to genus Aglaomyia as length of wing. M fork is 10 times the length of section M3, having a combination of following characters: R4 absent; Sc section M2 is three times the length of section M3. long, reaching the level of r-m base, ending in C; Sc2 in terminal 1/3 of Sc. The genus is close to Boletina Staeger. A number of Genus Azana Walker, 1856 species of Boletina is described from the Eocene–Miocene of Europe, North America and China, but they should be revised. 1856 Azana Walker, p. 26. Type species Azana scatopsoides Some of them do not belong to Mycetophilidae at all (B. Walker, 1956, p. 234, by monotypy ¼ Boletina anomala acropteris Hong, 1981 – probably Chironomidae, B. melaina Staeger, 1840, p. 238 Hong, 1981, B. pulvinata Hong, 1981, B. uda Hong, 1981 – Sciaridae). Remarks. Fifteen species in Holarctic and Oriental Regions, temperate to tropic. One fossil species in Eocene of Baltic Genus Coelosia Winnertz, 1863 amber. 1863 Coelosia Winnertz, p. 796. Type species Boletina ﬂava Azana cockerelli Blagoderov, n. sp. Staeger, subsequent designation by Johannsen, 1909, (Fig. 96, Pl. 8b) p. 86

Holotype. NHMUK I.9264, partly preserved thorax, abdo- Twenty-four species mainly in Holarctic. Larvae in fungi men and a wing, Brodie Collection; Insect Bed; late Eocene; (Soli 1997). NW Isle of Wight, UK. Diagnosis. The species differs from A. rarissima Meunier, ?Coelosia virgata (Cockerell, 1921) n. comb. 1904 from the Baltic amber in having RS base more proximal (Fig. 98, Pl. 8e) (R base is 2.71 times the R1 in the latter species) and CuA longer (0.63 times the wing length in latter species). 1921 Phronia (?) virgata Cockerell, pp. 465–66, ﬁg. 20 Description. Measurements: body 2.8 mm, wing 2.1 mm. 1994 Phronia virgata Evenhuis, p. 161 Costa slightly extends beyond the tip of R5. Sc not apparent. Length of R þ R1 is 0.44 times the wing length. R base Holotype NHMUK I.9789, Brodie Collection; Insect Bed; (measured between humeris and RS base) is 1.89 times the late Eocene; NW Isle of Wight, UK. R1. Length of r-m is 1.56 times the length of terminal section Additional material. NHMUK In.25315, In.25290, Hooley of Sc (distad of Sc2). Length of R5 is 0.83 times the wing Collection. length, length of CuA – 0.75 times the wing length. Basal Diagnosis. The species differs from other species of Coelosia half of M vein is obsolete. M curved posteriorly at the very in absence of macrotrichia on Sc. end. CuA is evenly curved. Description. Measurements: body 3 mm; wing 2.5–3 mm (holotype 2.7). Wing: wing membrane with microtrichia not Subfamily Gnoristinae Edwards, 1925 organised in rows. Costa extends beyond the tip of R5 to 1/2 Genus Aglaomyia Vockeroth, 1980 of the distance between the tips of R5 and M1. Costa, R base, R1, R5 and r-m setose. Sc bare, ending at C at the distad of 1980 Aglaomyia Vockeroth, p. 537, type species Aglaomyia the level of r-m base, its length 0.24–0.29 times the length of gatineau Vockeroth, by original designation the wing (holotype 0.29). Sc2 absent. Length of R1 is 0.71– 0.78 times the length of the wing (holotype 0.78). Length of Two species in Nearctic and Palaearctic. Larvae in decaying R5 (measured from the base of RS) is 0.59–0.63 times the wood of deciduous trees. length of the wing (holotype 0.63). Length of section M3 is 1.21–1.44? times the length of r-m (holotype 1.44) and 0.77– Aglaomyia vectis Blagoderov, n. sp. 0.87 times the section M2 (holotype 0.87). M-ratios (Soli (Fig. 97, Pl. 8c) 1997) are 0.26 and 0.31, Cu ratios are 1.9–2.5 (holotype 1.9) and 4.47–4.8 (holotype 4.47). Thorax. Mediotergite and Derivation of name. From Vecta insula, Latin name of the laterotergites without apparent hairs. Isle of Wight. Holotype. NHMUK In.24819, partial wing, Hooley Collec- Genus Dziedzickia Johannsen, 1909 tion; Insect Bed; late Eocene; NW Isle of Wight, UK. Other material. In.25321, In.25318, In.25194, Hooley 1885 Hertwigia Dziedzicki, p. 165. Preoccupied by Schmidt. Collection. Type species Hertwigia marginara Dziedzicki, 1885, by Diagnosis. It differs from A. ingrica and A. gatineau in monotypy having section M3 longer (in recent species length of M3 is 1909 Dziedzickia Johannsen, p. 44. Replacement name for about the length of r-m). Hertwigia Dziedzicki Description. Measurements: wing: 3.7–4 mm (holotype – 4 mm). Wing membrane with microtrichia only. Costa, R1 Approximately 60 species worldwide, distributed from cold and R5 with macrotrichia. Costa extends beyond the tip of temperate to equatorial zone. Biology unknown. Five fossil R5 to 1/4 of the distance between the tips of R5 and M1. Sc species are known from the Baltic amber and the Oligocene ending in C at the level of the base of section M3. Sc2 in of Rott. terminal 1/3 of Sc, slightly proximad of the level of M3 þ 4 and CuA fork base. Veins R1 and R5 almost straight, not Dziedzickia oligocenica Blagoderov, n. sp. curved posteriorly. Length of section M3 is 2.3 the length (Fig. 99, Pl. 8g)

Figures 96–104 Mycetophilidae and Keroplatidae, wing venation. (96) Azana cockerelli Blagoderov, n. sp., holotype NHMUK I.9264. (97) Aglaomyia vectis Blagoderov, n. sp., holotype NHMUK In.24819. (98) ?Coelosia virgata (Cockerell, 1921), n. comb., holotype NHMUK I.9789. (99) Dziedzickia oligocenica Blagoderov, n. sp., holotype NHMUK In.25301. (100) Palaeoempalia saxea Blagoderov, n. sp., holotype NHMUK I.9360. (101) Mycomya oblita Cockerell, 1921, holotype NHMUK I.8973. (102) Mycomya hoolei Blagoderov, n. sp., holotype NHMUK In.25330. (103) Leia gurnardensis, n. sp., holotype NHMUK In.25295. (104) Macrocera obliqua (Cockerell, 1921), n. comb., holotype NHMUK I.9657.

Derivation of name. From Oligocene. and CuA very proximal; D. sepulta Meunier, 1917 – in Holotype. NHMUK In.25301, a wing, Hooley Collection; absence of R4 vein. D. nashi Blagoderov & Grimaldi, 2004 Insect Bed; late Eocene; NW Isle of Wight, UK. from the Upper Cretaceous New Jersey amber differs from Diagnosis. D. hadroneuroides Meunier, 1923 differs from the the new species in having small cell bigger, Sc longer and new species by larger size, P6mm; D. johannseni Meunier, base on M3 þ 4 and CuA fork more proximal. 1917 – in having section M3 shorter than r-m and base of Description. Measurements. Wing 3.1 mm (preserved part M3 þ 4 and CuA fork proximad of r-m base; D. lepida Statz, 2.9 mm). Wing membrane with microtrichia only. Costa 1944a in having RS base more proximal, small sell long and extends beyond the tip of R5 to 1/5 of the distance between rectangular, and base of M3 þ 4 and CuA fork proximally; the tips of R5 and M1. R base with 12–13 setae, R1 with D. sedula Meunier, 1923 – in having base on fork M3 þ 4 6 þ 23 setae, and R5 with 14 setae. Sc ending at R at the level

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Plate 8 Mycetophilidae and Keroplatidae. (a) Acnemia simplex Cockerell, 1921, holotype NHMUK I.9597. (b) Azana cockerelli Blagoderov, n. sp., holotype NHMUK I.9264. (c) Aglaomyia vectis Blagoderov, n. sp., holotype NHMUK In.24819. (d) Palaeoempalia saxea Blagoderov, n. sp., holotype NHMUK I.9360. (e) ?Coelosia virgata (Cockerell, 1921), n. comb., holotype NHMUK I.9789. (f ) Leia gurnardensis Blagoderov, n. sp., holotype counterpart NHMUK In.25314. (g) Dziedzickia oligocenica Blagoderov, n. sp., holotype NHMUK In.25301. (h) Mycomya hoolei Blagoderov, n. sp., holotype NHMUK In.25330. (i) Mycomya oblita Cockerell, 1921, holotype NHMUK I.8973. (j) Macrocera obliqua (Cockerell, 1921), n. comb., holotype NHMUK I.9657.

Palaeoempalia saxea Blagoderov, n. sp. Derivation of name. In honour of R. W. Hooley, collector of (Fig. 100, Pl. 8d) the type specimen. Holotype. NHMUK In.25330, wing, Hooley Collection; Derivation of name. From Latin saxeus meaning made of Insect Bed; late Eocene; NW Isle of Wight, UK. stone. Diagnosis. The new species is the closest to M. helmi Holotype. NHMUK I.9360, partial body and wing, Brodie (Cockerell, 1904) from which it differs in having M3 þ 4and Collection; Insect Bed; late Eocene; NW Isle of Wight, UK. CuA fork base proximad of the level of base of M2. Diagnosis. The new species differs from P. brongniarti in Description. Measurements: wing 5.5 mm. Sc ending at having longer section M3, and position of Sc2 and base of Costa, its length 0.46 times the wing length. Sc2 at the fork M3 þ 4 and CuA more distally. Edwards (1940) noted proximal 1/4 of the middle of small cell. Section RS1 is equal that most of the species referred by Meunier to Palaeoempalia the crossvein r-m and 0.53 times the section RS2. Section M2 may belong to Palaeoboletina. The detailed comparison can is two times the section RS2 and 0.60 times the section M2. be possible only after revision of the Baltic amber species. Length of fork M1 and M2 is 2.17 times the section M3. Description. Measurements: body 4.2 mm, wing 3 mm. Base of fork M3 þ 4 and CuA proximad of the level of Sc2, Wing: R base, R1 and R5 setose. Sc ending at C, just before M3 þ 4 and CuA stalk is 0.72 times the section M2. the level of R4, its length 0.39 times the wing length. Sc2 present, at the level of RS. Section RS2 is 1.2 times the section Genus Leia Meigen, 1818 RS1. Length of crossvein r-m is 1.44 times the section RS1. Section M3 is 3.05 times the length of crossvein r-m. Section 1818 Leia Meigen, p. 258. Type species Leia fascipennis M2 is 2.23 times the section M2. M fork is 3.27 times the sec- Meigen, by subsequent designation by Curtis, 1837, tion M3; tips of veins M1 and M2 diverging. Base of M3 þ 4 p. 645 and CuA fork at the level of base of r-m. 1856 Lejosoma Rondani, p. 195, unjustiﬁed replacement name for Leia Meigen. Type species Leia bimaculata Genus Mycomya Rondani, 1856 Meigen, by original designation 1856 Lejomya Rondani, p. 195, incorrect original spelling of 1856 Mycomya Rondani, p. 194. Type species Sciophila Lejosoma marginata Meigen, 1818, by monotypy 1863 Glaphyroptera Winnertz, p. 771. Preoccupied by Heer. 1865 Cnephaeophila Philippi, p. 618. Type species Cnephaeo- Type species Leia fascipennis Meigen, by subsequent phila fenestralis Philippi, 1865, by monotypy designation by Coquillett 1910, p. 547 1878 Neoglaphyroptera Osten Sacken, 1878, p. 10. Replace- The genus has world wide distribution and includes more ment name for Glaphyroptera Winnertz than 300 extant species. About 15 fossil species of Mycomya Leja auct. Misspelling. are described from Cainozoic localities of Europe, North America and Far East. The genus has worldwide distribution and includes about 130 species. Larvae were reported from mushroom fruiting Mycomya oblita Cockerell, 1921 bodies, on surface of fungi or decaying wood, in birds’ nests. (Fig. 101, Pl. 8i) 1921 Mycomya oblita Cockerell, p. 463, ﬁg. 18 Leia gurnardensis Blagoderov, n. sp. 1994 Mycomya oblita Evenhuis, p. 157 (Fig. 103, Pl. 8f )

Holotype. NHMUK I.8973, partial insect, Brodie Collec- Derivation of name. From Gurnard Bay of the Isle of tion; Insect Bed; late Eocene; NW Isle of Wight, UK. Wight. Diagnosis. M. unicolor, M. fossilis, M. kuhni, M. umbonata, Holotype. In.25295/In.25314 (part and counterpart), wing M. fulvescens Statz, 1944a differ from the new species in hav- without apex and partial thorax, Hooley Collection; Insect ingScendingatRandM3þ 4 and CuA fork base proximad Bed; late Eocene; NW Isle of Wight, UK.

Figures 105–106 Scatopsidae. Regmoclemina haennii Krzemin´ski, n. sp., holotype NHMUK I.9512. (105) Antenna. (106) Wing venation.

Derivation of name. This species is named in honour of Figures 107–108 Anisopodidae – wing venation. (107) Sylvicola Jean-Paul Haenni, an outstanding specialist of Scatopsidae hooleyi (Cockerell, 1921), holotype NHMUK In.24346. (108) Sylvicola problematica Krzemin´ski, n. sp., holotype NHMUK In.25675. (Museé national d’histoire naturelle in Neuchaˆtel, Switzerland), with thanks for the critical reading of the manuscript. Holotype. NHMUK I.9512, Brodie Collection; Insect Bed; late Eocene; NW Isle of Wight, UK. Anisopodidae are direct descendants of an extinct family Other material. NHMUK In.25255, Hooley Collection. Protorhyphidae, known from numerous fossils of Jurassic and Diagnosis. Antennae 13 segmented; vein A2 very strong and Cretaceous of Europe (Krzemin´ski & Krzemin´ska 2003). characteristically double bent. Eight fossil species of Sylvicola have been described so far Description. Very small specimens preserved with wings and from the Eocene and Miocene of Europe and North America. body. Wing length 1.3–1.8 mm; body length 1.8–2.8 mm. Recent Anisopodidae are a small family of ca.100 species Head small, only partially visible; antenna (Fig. 105) twice of two subfamilies, distributed on all continents apart from longer than width of head, composed of 13 segments; last Antarctida. The most numerous is the genus Sylvicola. flagellomere twice longer than penultimate; palps invisible. Species live usually at borders of forest, among higher vege- Thorax and legs only partially visible. Wings badly preserved, tation, loose shrubs and garden. The larvae feed on decaying only one wing of holotype is in state allowing its partial recon- detritus. struction (Fig. 106). Vein Sc very short; R1 as long as R4 þ 5; R4 þ 5 long, ending in Costa beyond fork of Mb; cross-vein r-m invisible, having been fused with Mb and R4 þ 5 into one Genus Sylvicola Harris, 1780 horizontal vein; fork of M1 þ 2 very long. M1 poorly visible Sylvicola hooleyi (Cockerell, 1921) and its shape in Fig. 106 may be inaccurately reconstructed, (Fig. 107, Pl. 4j) but in any case it is 1.5 times longer than Mb; false vein (f vn) present between Mb-M2 and Cu; A1 has complicate shape, 1921 Rhyphus hooleyi Cockerell, p. 467, fig. 27 characteristically double bent; membrane of anal part with 1994 Sylvicola hooleyi Evenhuis, p. 206 few macrosetae. Abdomen very broad; genitalia preserved only partially. Holotype. NHMUK In.24346, Hooley collection (H. 456); Shape of abdomen and genitalia suggests that both specimens Insect Bed; late Eocene; NW Isle of Wight, UK. are females, but in neither the spermatheca is visible. Additional material. NHMUK I.9063/In.17136 (part and Remarks. Both specimens are poorly preserved but have counterpart) Brodie/Smith Collections; II 2811 (part and well visible 13-segmented antennae. It is a unique feature in counterpart), collected by the Polish team, north end Thorness Scatopsidae family; most recent and fossil species have no Bay; I.8851, Brodie collection. more than 12 segments. I included this species from the late Diagnosis. Wing regularly oval, with several distinct spots; Eocene of the Isle of Wight in the genus Regmoclemina, R4 þ 5 ca3.5 times longer than Rs. because all visible characters (wing venation and macrosetae Redescription. Holotype specimen is a single, almost com- on anal lobe) are in concordance with reminding species of plete wing 5.2 mm long, with distinct dark spots, which are this genus. also preserved in remaining specimens allotted to this species. Superfamily Anisopodoidea Knab, 1912 Venation: Sc ending distinctly before fork of M1 þ 2 into M1 by Wiesław Krzemin´ski and M2; R2 þ 3 2 times longer than Rs; R4 þ 5almostfour times longer than Rs; M1 two times longer than upper margin Family Anisopodidae Knab, 1912 of d cell; Cu conspicuously wavy. Anisopodidae are rare in the Isle of Wight fossil material. Remarks. All specimens show almost no variation in wing The only taxon represented is Rhyphus hooleyi (¼Sylvicola venation, which is also characteristic of recent representatives hooleyi) described by Cockerell in 1921 on the basis of a of Sylvicola. single, very well-preserved wing originally labelled No. H. 456 (now NHMUK In.24346), and the specimen preserved on Sylvicola problematica Krzemin´ski, n. sp. two counterparts: In.17136 and I.9063. (Fig. 108, Pl. 4k)

Figures 109–115 Stratiomyidae. (109)–(113) Odontomyia brodiei (Cockerell, 1915): (109) NHMUK I.10114 antenna; (110) NHMUK In.17253 wing venation; (111) USNM 61443 holotype; (112) NHMUK I.8779 wing venation showing acute angled discal cell; (113) NHMUK I.8892 wing venation lacking m–cu. (114), (115) Palaeoberidops barkeri Nicholson, n. gen., n. sp.: (114) NHMUK I.9866(3) holotype part; (115) NHMUK I.9718(3) holotype counterpart.

O. brodiei must be the ‘Oligocene’ ancestor of modern Odonto- Derivation of name. Greek, palaeo, ancient, and beridops, myia. Fossil and molecular data indicates the divergence the genus to which it bears a close resemblance. between Odontomyia and Stratiomys occurred 25–35 Ma Type species. Palaeoberidops barkeri n. gen., n. sp. by (Brammer & Dohlen 2007). While the uppermost Eocene age monotypy. of the Bembridge Marls leaves open the possibility that Diagnosis. As for the type and only known species. O. brodiei could be the most recent common ancestor to Stratiomys and Odontomyia, the lack of scutellar spines Palaeoberidops barkeri Nicholson, n. gen., n. sp. (which are nearly ubiquitous in both genera today) makes it Figs 114, 115, Pl. 9h, i unparsimonious to assume this to be the case, or that it is a direct ancestor of all modern Odontomyia. Sixteen stratiomyid larvae are held in the NHM collections, Derivation of name. In honour of Dr Michael Barker in his provisionally assigned to the subfamily Stratiomyinae. Larvae retirement from the School of Earth and Environmental in this subfamily are aquatic and some are known to be able Sciences, University of Portsmouth, UK. to withstand highly saline water (James 1981), so it is possible Holotype. NHMUK I.9718/I.9866(3) (part and counter- that these were living in the lagoon, though more likely to be part), Brodie Collection, incomplete single wing Insect Bed; that they were washed in by streams. late Eocene; NW Isle of Wight, UK. Diagnosis. Absence of M3 and m-cu, with CuA1 originating at the point where CuA meets the discal cell; R strongly Subfamily Beridinae Westwood, 1838 2+3 curved anteriorly; width of discal cell two thirds the length; Genus Palaeoberidops, n. gen. length of discal cell less than half that of cell r1; wing apex

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Figures 116–135 Stratiomyidae: Odontomyia brodiei. Representations of the shape of the discal cell in fossil specimens, where preserved. For ease of comparison all cells have been made roughly the same size and oriented with the proximal wall of the cell vertical and some images reversed so all appear as on a right wing. (116) NHMUK I.8775. (117) NHMUK I.8779. (118) NHMUK I.8796. (119) NHMUK I.8802. (120) NHMUK I.8892. (121) NHMUK I.8975. (122) NHMUK I.9044. (123) NHMUK I.9172. (124) NHMUK I.17221. (125) NHMUK In.17253. (126) NHMUK In.17356. (127) NHMUK In.17478. (128) NHMUK In.25276. (129) NHMUK In.25277. (130) NHMUK In.25390. (131) NHMUK In.25414. (132) NHMUK In.25508. (133) NHMUK In.25744. (134) CAMSM X.50140.8. (135) USNM 61443 (Lacoe 7507).

not visible but trajectories suggest R4 less than half the length curved at base of fork, otherwise straight; pterostigma black; of R5;R4+5 substantially thicker than all other veins; pteros- branches of medial vein (M1 and M2) fade distally; anterior tigma present in cell r1 and possibly extends into subcostal wing margin not completely visible. Preserved section of the cell (sc). wing 3.8 mm long. Description. Single partial wing with colouration preserved Remarks. Initial comparison to Beris miocenica James, in veins. Preservation fades in from proximal end of basal cells 1937, from the Eocene Florissant Beds, USA, shows P. barkeri through nearly to the apex of the wing; costal vein from has a much more strongly curved R2+3, resulting in less crowded anterior end of R1 out towards apex; CuA2 and relatively radial cells. Comparison with extant species of Beris (e.g., B. short A1 join more anteriorly than is commonly seen in the fuscipes, B. chalybata, B. geniculata, B. morisii and B. vallata) Subfamily; CuA2 short and slightly recurved; R4 only slightly shows P. barkeri has a markedly less elongate discal cell, with

Plate 9 Bibionidae and Stratiomyidae. (a) Plecia sp., NHMUK In.25249. (b) Penthetria sp., NHMUK In.25246. (c) Plecia sp., NHMUK In.17230. (d–g) Odontomyia brodiei (Cockerell, 1915): (d) NHMUK In.64530 (¼ In.64647), dorsal view showing abdominal colour patterning; (e) holotype USNM 61443 (Lacoe Collection 7507); (f) NHMUK In.17253, showing exceptional wing preservation; (g) NHMUK I.10114, lateral view showing full antenna. (h, i) Palaeoberidops barkeri Nicholson, n. gen., n. sp., holotype: (h) NHMUK I.9866(3), part; (i) I.9718(3), counterpart.

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Figures 136–137 Hybotidae and Phoridae. (136) Hybotidae, NHMUK I.8981. (137) Phoridae, NHMUK I.9488.

a shape more like that seen in the tribe Stratiomyini. With the Holotype.In.24349 exception of Beridops spp., all other genera in the Beridinae have the CuA1 originating on the discal cell and the discal Chrysopilus stigmaticus Cockerell, 1921 cell markedly elongate. Beridops differs from Palaeoberidops (Pl. 10e) barkeri in that the discal cell shows some elongation and is at 1921 Chrysopilus stigmaticus Cockerell, p. 471, fig. 36 (Leptidae) least two-thirds the length of cell r1. 1994 Chrysopilus stigmaticus Evenhuis, p. 292 The following taxa have been previously described or newly recognised from the Insect Limestone, but have not been Holotype. NHMUK In.24350. studied for this project. Family Tabanidae Latreille, 1802 Family Bombyliidae Latreille, 1802 Tabanus vectensis (Cockerell, 1921) (Pl. 10g) Systropus acourti Cockerell, 1921 (Pl. 10a) 1921 Chrysops vectensis Cockerell, p. 472, fig. 37 1994 Tabanus vectensis Evenhuis, p. 289 1921 Systropus acourti Cockerell, p. 469, fig. 32 1973 Systropus acourti Hull, p. 59 Holotype. NHMUK In.24351. 1994 Systropus acourti Evenhuis, p. 336 Family Empididae Latreille, 1804 Holotype. NHMUK In.17208/In.17293. Specimens NHMUK I.8981, I.9289, I.9371, I.9401, I.9465, Family Asilidae Latreille, 1802 I.9730, I.10275, I.10372, I.10378 (Pl. 10h), In.17199, In.25288, In.25264, In.43435. Asilus gurnetensis Cockerell, 1921 (Pl. 10b) Family Dolichopodidae Latreille, 1809 1921 Asilus gurnetensis Cockerell, p. 470, fig. 33 1962 Asilus gurnetensis Hull, p. 24 Specimens NHMUK I.8443, I.10260, I.8666, I.8850, I.8930, 1994 Asilus gurnetensis Evenhuis, p. 324 I.8974, I.9080, I.9094, I.9328, I.9391, I.9765, I.9826, In.17110, In.17488, In.20534, In.24635, In.25173, In.25224, In.25233 Holotype. NHMUK In.24348. (Pl. 10i), In.25262, In.25273, In.25348, In.25377, In.25397, In.25438, In.25506, In.25732. Proctacanthus fractus Cockerell, 1921 (Pl. 10c) Family Hybotidae Meigen, 1820 1921 Proctacanthus fractus Cockerell, p. 470, fig. 34 1962 Proctacanthus fractus Hull, p. 24 Specimen NHMUK I.8981 (Fig. 136) 1994 Proctacanthus fractus Evenhuis, p. 327 Family Syrphidae Latreille, 1802 Holotype. NHMUK In.24347. Specimens NHMUK In.17355, In.20551, In.24809 (Pl. 10j), Family Rhagionidae Latreille, 1802 In.25261, In.25264. Chrysopilus anglicus Cockerell, 1921 Family Phoridae Curtis, 1833 (Pl. 10d)

1921 Chrysopilus anglicus Cockerell, p. 471, ﬁg. 35 (Leptidae) Specimens NHMUK I.9488, I.9512, In.20534b (Fig. 137). 1994 Chrysopilus anglicus Evenhuis, p. 292

Plate 10 Bombyliidae, Asilidae, Rhagionidae, Stratiomyidae, Tabanidae, Empididae, Dolichopodidae, and Syrphidae. (a) Systropus acourti Cockerell, 1921 (Bombyliidae), holotype NHMUK In.17208. (b) Asilus gurnetensis Cockerell, 1921 (Asilidae), holotype NHMUK In.24348. (c) Proctacanthus fractus Cockerell, 1921, holotype NHMUK In.24347. (d) Chrysopilus anglicus Cockerell, 1921 (Rhagionidae), holotype NHMUK In.24349. (e) Chrysopilus stigmaticus Cockerell, 1921, holotype NHMUK In.24350. (f ) Odontomyia brodiei? (Cockerell, 1915) (Stratiomyidae) larva, NHMUK II.2978, A. Yule Coll. (g) Tabanus vectensis (Cockerell 1921), holotype NHMUK In.24351. (h) Empididae, NHMUK I.10378. (i) Dolichopodidae, NHMUK In.25233. (j) Syrphidae, NHMUK In.24809.

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Table 3 Diptera known from Bembridge Marls.

Tipulidae ?Bibio gurnetensis Cockerell, 1917a Brachypremna gurnetensis (Cockerell, 1921) ?Bibio oligocenus Cockerell, 1917a Ischnotoma (Ischnotoma) vasifera (Cockerell & Haines, 1921) Bibiodes conﬂuens (Cockerell, 1915), n. comb. Leptotarsus (Longurio) fragmentatus Krzeminski, n. sp. Dilophus andrewrossi Nel, Colomb & Waller, n. sp. Leptotarsus (Longurio) wegiereki Krzeminski, n. sp. Penthetria indet. Leptotarsus (Macromastix) cladoptera (Cockerell & Haines, 1921) Plecia acourti Cockerell, 1921 Tipula (s. lato) acourti Cockerell, 1921 Keroplatidae Tipula (s. lato) gardneri Cockerell, 1917a Macrocerinae: Macrocera obliqua (Cockerell, 1921), n. comb. Tipula (s. lato) limiformis Cockerell, 1915 Keroplatinae: Orfelini Tipula (s. lato) anglicana Krzeminski, n. sp. Mycetophilidae Tipulomorpha indet. Acnemia simplex Cockerell, 1921 Paltostomopsis ciliata Cockerell, 1915 Aglaomyia vectis Blagoderov, n. sp. Cylindrotomidae Azana cockerelli Blagoderov, n. sp. Cyttaromyia rossi Krzeminski, n. sp. ?Coelosia virgata (Cockerell, 1921), n. comb. Limoniidae Dziedzickia oligocenica Blagoderov, n. sp. Cheilotrichia (Cheilotrichia) duplicata Krzemin´ski, n. sp. Leia gurnardensis Blagoderov, n. sp. Cheilotrichia (Empeda) ferruginea (Cockerell, 1921) Mycetophila vectensis Cockerell, 1915 Cheilotrichia (Empeda) hyalina (Cockerell, 1921) Mycomya hoolei Blagoderov, n. sp. Cheilotrichia (Empeda) szwedoi Krzemin´ski, n. sp. Mycomya oblita Cockerell, 1921 Dactylolabis (Dactylolabis) spiloptera (Cockerell, 1921), n. comb. Palaeoempalia saxea Blagoderov, n. sp. Dicranomyia (Dicranomyia) aliena (Cockerell, 1921), n. comb. ‘Rymosia’ edwardsi Cockerell, 1921 Dicranomyia (Dicranomyia) brodiei (Cockerell, 1917a) ‘Rymosia’ ferruginea Cockerell, 1921 Dicranomyia (Dicranomyia) excavata Cockerell & Haines, 1921 Rymosia grisea Cockerell, 1921 Dicranomyia (Dicranomyia) fasciata Krzemin´ski, n. sp. ‘Rymosia’ rufescens Cockerell, 1921 Dicranoptycha staryi Krzemin´ski, n. sp. Sciaridae Gonomyia (Gonomyia) indecisa Cockerell & Haines, 1921 Sciara protoberidis Cockerell, 1915 Gurnardia corami Krzemin´ski, n. gen., n. sp. Sciara gurnetensis Cockerell, 1915 Gymnastes (Paragymnastes) fasciatipennis Cockerell, 1921 Sciara lacoei Cockerell, 1915 Helius (Helius) edmundi Krzemin´ski, n. sp. Scatopsidae Helius (Helius) popovi Krzemin´ski, n. sp. Regmoclemina haennii Krzemin´ski, n. sp. Helius (Helius) vectensis (Cockerell, 1915), n. comb. Anisopodidae Ilisia (Lunaria) incognita (Cockerell, 1921), n. comb. Sylvicola hooleyi (Cockerell, 1921) Limnophila (Limnophila) cyclospila Cockerell, 1921 Sylvicola problematica Krzemin´ski, n. sp. Limnophila (Limnophila) deleta Cockerell, 1921 Rhagionidae Limonia spilota (Cockerell, 1921) Chrysopilus anglicus Cockerell, 1921 Orimarga (Orimarga) lenae Krzemin´ski, n. sp. Chrysopilus stigmaticus Cockerell, 1921 Pilaria hooleyi Krzemin´ski, n. sp. Tabanidae Pilaria volodii Krzemin´ski, n. sp. Chrysops vectensis Cockerell, 1921 Pseudolimnophila (Pseudolimnophila) disjunctula (Cockerell & Haines, Stratiomyidae 1921), n. comb. Odontomyia brodiei (Cockerell, 1915) Styringomyia concinna (Cockerell, 1917a) Palaeoberidops barkeri Nicholson, n. gen., n. sp. Symplecta (Psiloconopa) gurnetensis Krzemin´ski, n. sp. Bombyliidae Thaumastoptera undulata (Cockerell & Haines, 1921), n. comb. Systropus acourti Cockerell, 1921 Trentepohlia (Mongoma) cruciferella (Cockerell, 1917a) Asilidae Trentepohlia (Mongoma) pallescens (Cockerell, 1921) Asilus gurnetensis Cockerell, 1921 Psychodidae Proctacanthus fractus Cockerell, 1921 Pericoma primaeva (Cockerell, 1915), n. comb. Empididae (undescribed, NHM collection) Wightipsychoda leucospila (Cockerell, 1921), n. comb. Dolichopodidae (undescribed, NHM collection) Dixidae Hybotidae (undescribed, NHM collection) Dixella priscula (Cockerell, 1921), n. comb. Syrphidae (undescribed, NHM collection) Culicidae Phoridae (undescribed, NHM collection) Coquillettidia cockerelli (Edwards, 1923), n. comb. Otitidae Culex protorhinus Cockerell, 1915 Stenomyites fuscipennis Cockerell, 1915 Culex vectensis Edwards, 1923 Lauxaniidae indet. (Zlobin, 2007) Ochlerotatus protolepis (Cockerell, 1915), n. comb. Agromyzidae indet. (Zlobin, 2007) Chironomidae Anthomyzidae Chironominae: Chironomini indet. ‘Ephydra’ sepulta Cockerell, 1915 Orthocladiinae indet. ‘Ephydra’ oligocena Cockerell, 1915 Tanypodinae indet. Chloropidae Ceratopogonidae Hippelates brodiei Cockerell, 1915 Dasyhelea indet. Heleomyzidae indet. (Zlobin, 2007) Palpomyia edwardsi Cockerell, 1921 Ephydridae Stilobezzia indet. Protoscinis perparvus Cockerell, 1917a Simuliidae ‘Sphaerocera’ sepultula Cockerell, 1915 Simuliidae gen. et sp. indet. Bibionidae Bibio obliteratus (Cockerell, 1915), n. comb. ?Bibio extremus Cockerell, 1921

Downloaded from https://www.cambridge.org/core. Chalmers Tekniska Högskola, on 06 Oct 2019 at 16:48:14, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S1755691018000464 56 WIESŁAW KRZEMIN´ SKI ET AL. 3. Diptera fauna and habitat of the insect bed 4. Acknowledgements SW wants to thank Dr Andrew Ross (London) and Prof. Dr A list of Diptera known up to date from Insect Limestone is Rainer Willmann (Go¨ttingen) for their support of this re- provided in Table 3. In total, 102 species and 32 families are search, Theresa Howard (London) for access to extant simu- recorded from the locality. Among the Diptera from the Isle liid pupae and INTAS (Project-No. 03-51-4367) for financial of Wight the most abundant are those that depend directly on support. D.N. extends special thanks to Andrew Ross, Wieslaw the water habitat. Thus most abundant are the Limoniidae Krzemin´ski, Vladimir Blagoderov, David Martill, Erica (135 specimens), Chironomidae (170 specimens), Culicidae McAlister and Michael Barker. RS is much indebted to Dr A. (140 specimens), Psychodidae (39) and Stratiomyidae (41); rare Ross, for arranging the loan of materials and his kind hospital- are the Dixidae (6 specimens) and Simuliidae (one specimen). ity during his visit to England; to Dr A. Borkent of Salmon Surprisingly numerous are the Culicidae, which are scarcely Arm, Canada, Dr N. Evenhuis of Bernice P. Bishop Museum, represented in older sediments and in Baltic amber (Eocene) Honolulu, Hawaii and Dr W. L. Grogan of Salisbury State (Szadziewski & Szadziewska 1985; Szadziewski 1998; Podenas University for their advice and discussions on synonymy and 1999; Wichard et al. 2009). The Isle of Wight is the oldest nomenclatural problems. locality with so rich representation of culicid adults. Culicidae from the Isle of Wight belong to three extant genera of worldwide distribution (Coquillettidia, Ochlerotatus and Culex), whose extant species are common inhabitants of shallow reservoirs of stagnant water. Larvae and pupae of the genus 5. References Coquillettidia live among water plants and take air from plant Adler, P. H. and Currie, D. C. & Wood, D. M. 2004. The black flies tissues. (Simuliidae) of North America. Ithaca: Cornell University Press. Some of the Tipulidae (represented by 32 specimens in the Adler, P. H. & Crosskey, R. W. 2011. World blackflies (Diptera: Simuliidae): a comprehensive revision of the taxonomic and fauna) also prefer semiaquatic habitats or live directly in geographical inventory. http://entweb.clemson.edu/biomia/pdfs/ water, as several species of Tipula and Ischnotoma (Alexander blackflyinventory.pdf (accessed 21 December 2011). 1944, 1971). Alexander, C. P. 1919. The crane-flies of New York. Part I. Distribu- Generally, the Diptera of the aquatic and semiaquatic con- tion and taxonomy of the adult flies. Memoirs, Cornell University Agricultural Experiment Station 25, 767–993. ditions indicate the environment of a spacious shallow bay Alexander, C. P. 1920. New or little-known crane-flies in the Queens- with rich vegetation. Some lived in small slowly running land museum (Tipulidae, Diptera). Memoirs of the Queensland streams (Dixidae, Limoniidae of the genera Gonomyia, Limno- Museum 7, 52–63. phila, Pseudolimnophila, Cheilotrichia, Ilisia, Thaumastoptera) Alexander, C. P. 1922. New or little-known Tipulidae (Diptera). I. or wet soil on the borders of lakes or streams (Savchenko Proceedings of the Linnaean Society of New South Wales 47, 581–90. 1982, 1985, 1986). Alexander, C. P. 1931. Crane-flies of the Baltic amber (Diptera). The second, much less numerous group of Diptera are the Bernstein-Forschung 2,1–135. species living in the detritus, fungi, rotten wood, or feeding Alexander, C. P. 1944. New or little-known species of Australian on plant juices or are herbivorous. These are Anisopodidae Tipulidae (Diptera). II. Proceedings of the Linnaean Society of New South Wales 69,1–15. (five specimens), Bibionidae (19), Ceratopogonidae (nine), Alexander, C. P. 1956. Tipulidae. Ruwenzori Expedition 1934–35 1, Scatopsidae (two), Cylindrotomidae (one specimen), Keroplatidae 129–380. (17), some Tipulidae and Limoniidae, and the majority of the Alexander, C. P. 1971. New or little-known Tipulidae from Chile and Mycetophilidae (55 specimens), Agromyzidae, Anthomyidae Peru (Diptera: Tipulidae). Part III. Anales del Museo de Historia and Lauxanidae. Natural 4, 157–81. Ansorge, J. 1992. Dixa cimbrica n. sp., ein Vertreter der Dixidae Slowly running rivulets and streams transported dead (Diptera nematocera) aus der oberpaleoza¨nen/untereoza¨nen insects of the forest habitats to the lake, mainly those that Fur-Formation (Moler) Ju¨tlands (Da¨nemark). Neues Jahrbuch lived on the banks of the water body. The Diptera which prefer fur Geologie und Palaöntologie, Stuttgart 9, 513–18. fast-running, well-oxygenated water are either completely absent Armbruster, L. 1938. Versteinerte Honigbiennen aus dem Obermioca¨nen Randecker Maar. 3. Archiv fu¨r Bienenkunde 19, 97–133. from the collection (Thaumaleidae and Blephariceridae; speci- Beckenbach, A. T. & Borkent, A. 2003. Molecular analysis of the men previously described as Paltostomopsis ciliata Cockerell biting midges (Diptera: Ceratopogonidae), based on mitochon- 1915 (Pl. 7c) is an indeterminate member of Tipulomorpha). drial cytochrome oxidase subunit 2. 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MS received 28 March 2014. Accepted for publication 31 July 2017