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New fossil spongilla-flies from the lowermost Eocene amber of France (Insecta, Neuroptera, Sisyridae) André NEL Jean-Jacques MENIER Alain WALLER Gilbert HODEBERT Gaël DE PLOËG Département Histoire de la Terre, Muséum national d’Histoire naturelle, et CNRS UMR 8569, Bâtiment d’Entomologie, 45 rue Buffon, F-75231 Paris cedex 05 (France) [email protected] [email protected] Nel A., Menier J.-J., Waller A., Hodebert G. & De Ploëg G. 2003. — New fossil spongilla- flies from the lowermost Eocene amber of France (Insecta, Neuroptera, Sisyridae). Geodiversitas 25 (1) : 109-117. ABSTRACT KEY WORDS A critical list of the fossil Sisyridae is proposed. The new sisyrid genus and Insecta, species Paleosisyra eocenica n. gen., n. sp. is described from the lowermost Neuroptera, Sisyridae, Eocene amber of the Paris basin. Its relationships within the family are dis- Paleosisyra eocenica n. gen., n. sp., cussed in a first phylogenetic analysis of the Sisyridae. It falls as sister genus of amber, lowermost Eocene, the recent genus Sisyra Burmeister, 1839 and had probably a similar chorolo- Paris basin, gy. Some specimens of P. eocenica n. gen., n. sp. are covered by pollen, suggest- France, ing that it lived on flowers, as for the recent Sisyra. Also, the male specimens of phylogenetic analysis, new genus, P. eocenica n. gen., n. sp. are clearly less frequent than the female, as for the new species. recent Sisyra. This taxon is the oldest accurate record of the Sisyridae. RÉSUMÉ Nouveaux Sisyridae fossiles de l’ambre éocène inférieur de France (Insecta, Neuroptera). Une liste critique des Sisyridae fossiles connus est proposée. Les nouveaux MOTS CLÉS genre et espèce de Sisyridae Paleosisyra eocenica n. gen., n. sp. sont décrits de Insecta, l’ambre éocène basal du Bassin de Paris. Ses affinités dans cette famille sont Neuroptera, Sisyridae, discutées dans une première analyse phylogénétique des Sisyridae. Paleosisyra Paleosisyra eocenica n. gen., n. sp., n. gen. est le genre frère de Sisyra Burmeister, 1839 et avait probablement une ambre, Éocène basal, chorologie similaire. Des spécimens de P. eocenica n. gen., n. sp. sont couverts Bassin de Paris, de pollen, suggérant une alimentation au dépend de fleurs, comme pour les France, Sisyra modernes. Les mâles de P. eocenica n. gen., n. sp. semblent nettement analyse phylogénétique, nouveau genre, plus rares que les femelles, comme chez les Sisyra actuels. Il s’agit du plus nouvelle espèce. ancien Sisyridae décrit. GEODIVERSITAS • 2003 • 25 (1) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 109 Nel A. et al. INTRODUCTION Sisyra”. The preserved parts of the wing venation strongly resemble to that of Paleosisyra n. gen., The Sisyridae Handlirsch, 1906 is a small family, but also to Nevrorthus. with only five extant genera, i.e. Climacia Wichard & Weitschat (1996: fig. 9) figured two McLachlan, 1869 (Nearctic and Neotropical), adult specimens from the Baltic amber that they Sisyra Burmeister, 1839 (Cosmopolitan), Sisyrella attributed to “Sisyra (Rophalis) div. spec. – Banks, 1913 (Japan), Sisyrina Banks, 1939 Sisyridae”. The specimen figured in their “Tafel (India), and Sisyborina Montserrat, 1981 9” (lower photograph) is clearly a Rophalis sp. (Afrotropical region) (Carpenter 1940; Tjeder because it has its forewing veins Sc and R not dis- 1976; Monserrat 1977, 1981). Sisyridae strongly tally joined, but with a small cross-vein between resemble the Nevrorthidae Nakahara, 1915 in them. It is very similar to the specimen of their wing venation. After Monserrat (1977), the Rophalis relicta Hagen, 1856 described in the family Nevrorthidae comprises three extant Nevrorthidae by Nel & Jarzembowski (1997). genera, i.e. Austroneurorthus Nakahara, 1958 Wichard & Weitschat (1996: fig. 10) figured a (Australia), Nevrorthus Costa, 1863 (Palaearctic), larva they attributed to a “Neurorthus spec. – and Nipponeurorthus Nakahara, 1958 (Eastern Neurorthidae”. This larva could be that of Asia), and the Baltic amber genus Rophalis Rophalis and would confirm the presence of the Hagen, 1856. The fossil record of both families is family Nevrorthidae in the Baltic amber. The very scarce. specimen figured in the upper photograph of Schlüter (1986), in his list of fossil Neuroptera, their “Tafel 9” is not a Rophalis sp. Interestingly, indicated that the Sisyridae are known from the it is very similar to Paleosisyra eocenica n. gen., Cretaceous, without any indication of literature. n. sp. (especially in its forewing veins Sc and RA The only Mesozoic described taxon, Cratosisyrops which are distally joined, without any cross-vein gongazai Martins-Neto, 1997 (revised in between them, it has numerous cross-veins in the Martins-Neto 2000) (Lower Cretaceous, Santana costal area, etc.). It seems to be a female specimen Formation, north-eastern Brazil) was first figured because of the presence of genital appendages and described by Martins-Neto (1992) as a very similar to those of Paleosisyra eocenica “Sisyridae? gen. et sp. indet.”. Martins-Neto n. gen., n. sp. (1997) gave the following diagnostic characters: Weitschat & Wichard (1998: fig. 54a) also fig- small size; great eyes; oval wings; forewing Sc and ured a female adult specimen from Baltic amber R very close to each other, connected distally by a they attribute to “?Rophalis sp.”. The genital small cross-vein; MA forks near the base of the appendages of this specimen correspond to those wing; RP with five branches; CuA pectinated; of a Sisyridae and its wing venation to that of CuP bifurcated. Martins-Neto (1997) attributed Paleosisyra n. gen., especially in the presence of a this fossil taxon to the Sisyridae with some doubt very short fusion of MA with RP in hindwing and compared it to the genus Sibithone (see below). All these specimens probably belong Ponomarenko, 1984 (in the fossil family to this genus, but a direct revision of these speci- Mesithonidae Panfilov, 1980). After the descrip- mens will be necessary before any systematic tion and original figure of the type specimen, as attribution. proposed by Martins-Neto, it is not possible to Schumann & Wendt (1989) mentioned a fossil attribute this taxon to a precise family. It could Sisyridae from the Saxonian amber (Miocene, be related to Sisyridae, Nevrorthidae, or even Germany). Dilaridae Handlirsch, 1906. Nel & Jarzembowski (1997) revised the Baltic Jarzembowski (1980) described an impression of amber genus Rophalis Hagen, 1856 and attrib- the fore- and hindwing of a fossil Sisyridae from uted it to the Nevrorthidae. They also described a the upper Eocene of the Isle of Wight (UK). Its Sisyra sp. indet. (upper Miocene of the Centre of wing venation is “closest to the extant genus France). 110 GEODIVERSITAS • 2003 • 25 (1) New Sisyridae (Insecta, Neuroptera) in amber The present new species is the oldest accurate Paleosisyra eocenica n. sp. Sisyridae. We follow Aspöck et al. (1980) in their (Figs 1-8) interpretation of genitalia characters and wing venation of Sisyridae, and Parfin & Gurney TYPE MATERIAL. — Holotype male specimen PA 2789 (1/8), with eight adult Diptera: Chironomidae and a (1956) in their nomenclature of the body struc- Ephemeroptera in the same piece of amber. Female tures. allotype specimen PA 156 (1/2), female paratype spe- cimens PA 11, PA 52 (1/5), PA 157 (1/5), PA 2329, ABBREVIATIONS PA 2478 (1/2) and (2/2), PA 2493 (1/2), PA 2812, PA 1st m first cross-vein between MA and MP; 3921, PA 4144, PA 4864, PA 5368, PA 5374, PA 1st, 2nd, 3rd r cross-veins between R and RP; 5603. All specimens in collection De Ploëg and A1, 2, 3 anal veins; Indivision Langlois-Meurine, deposited in Muséum national d’Histoire naturelle, Paris. Specimens collect- cp coxopodites; ed in Le Quesnoy all bear the letter PA for Paris CuA cubitus anterior; (meaning Paris basin); the following number is the CuP cubitus posterior; ordinal number in the collection. epr ectoprocte; gl “gonapophyses latérales”; ETYMOLOGY. — After the age of the amber locality. M median vein; MA media anterior; TYPE LOCALITY. — Le Quesnoy, Chevrière, region of MP media posterior; Creil, Oise department, France. R radial vein; GEOLOGICAL AGE. — Lowermost Eocene, Sparnacian, RP radius posterior; level MP7 of the mammal fauna of Dormaal. We RA radius anterior; could demonstrate that the amber is autochthonous Sc subcosta. and very different from the Baltic amber in age, che- mical composition and origin (Nel et al. 1999). STATE OF PRESERVATION OF THE MATERIAL.— SYSTEMATICS Specimen PA 2789 (1/8): nearly complete in a very clear piece of amber. The upper surfaces of the tho- rax and abdomen are partly destroyed. Specimen Order NEUROPTERA Handlirsch, 1908 PA 156 (1/2): complete in a very clear piece of Family SISYRIDAE Handlirsch, 1906 amber, together with a small insect. Specimens PA 11, PA 2478 (1/2 and 2/2), PA 2493 (1/2), Genus Paleosisyra n. gen. PA 2812, PA 5368, and PA 5603: complete in very clear pieces of amber. Specimen PA 2329: complete TYPE SPECIES.— Paleosisyra eocenica n. gen., n. sp. covered with pollen. Specimens PA 157 (1/5), PA 4144, PA 4864, and PA 5374: head and thorax ETYMOLOGY. — After Paleo and the genus Sisyra. partly destroyed. Specimen PA 3921: apical parts of DIAGNOSIS.— Paleosisyra n. gen. has three characters four wings. absent in all extant Sisyridae, i.e. very short fusion of MA with RP in hindwing; prothoracic epimeron and episternum well defined and broad, with a furrow bet- DESCRIPTION ween them; two strong spurs on prothoracic legs (un- It is based on all specimens. The