Phylogenetic analysis of the Cenozoic family Sieblosiidae (Insecta: Odonata), with description of new taxa from Russia, Italy and France

Geobios 38 (2005) 219–233 http://france.elsevier.com/direct/GEOBIO/

Original article

Phylogenetic analysis of the Cenozoic family Sieblosiidae (Insecta: Odonata), with description of new taxa from Russia, Italy and France Analyse phylogénétique de la famille cénozoïque Sieblosiidae (Insecta : Odonata), avec la description de nouveaux taxons de Russie, Italie et France André Nel a,*, Julian F. Petrulevicˇius a,b, Guiseppe Gentilini c, Xavier Martínez-Delclòs d a

Laboratoire d’Entomologie and UMR CNRS 8569, Muséum National d’Histoire Naturelle, 45, rue de Buffon, 75005 Paris, France Laboratoire d’Entomologie, Muséum National d’Histoire Naturelle, 45, rue de Buffon, 75005 Paris, France and Conicet, Argentina c Via Adriatica 78, 47843 Misano Adriatico, Rimini, Italy d Departament d’Estratigrafia, Paleontologia i Geociències Marines, Facultat de Geologia, Universitat de Barcelona, 08071 Barcelona, Spain b

Received 1 September 2003; accepted 19 October 2003 Available online 26 February 2005

Abstract We describe the following Sieblosiidae: an unamed “gen. and sp. A” from the Miocene of Italy, Miostenolestes zherikhini nov. gen., nov. sp., Paraoligolestes stavropolensis nov. sp., Stenolestes fasciata nov. sp. (all from the Miocene of North Caucasus), Stenolestes (?) adygeianensis nov. sp. (Oligocene of North Caucasus), and Stenolestes cerestensis nov. sp. (Oligocene of France). The genus Sieblosia Handlirsch, 1906 is restored. A new phylogenetic analysis of the Sieblosiidae is proposed. The two taxa “gen. and sp. A” and Oligolestes fall in most inclusive positions in the same clade with the Sieblosiidae. Within the Sieblosiidae sensu stricto, the two clades (Paraoligolestes + (Parastenolestes + Stenolestes)) and (Parastenolestes + Stenolestes) are the best supported. The family Sieblosiidae seems to be restricted to the Oligocene–Miocene of Europe. © 2005 Elsevier SAS. All rights reserved. Résumé Nous décrivons les Sieblosiidae suivants : un « gen. et sp. A » du Miocène d’Italie, Miostenolestes zherikhini nov. gen., nov. sp., Paraoligolestes stavropolensis nov. sp., Stenolestes fasciata nov. sp. (tous du Miocène, Caucase du nord), Stenolestes (?) adygeianensis nov. sp. (Oligocène, Caucase du nord), et Stenolestes cerestensis nov. sp. (Oligocène de France). Le genre Sieblosia Handlirsch, 1906 est restauré. Une nouvelle analyse phylogénétique des Sieblosiidae est proposée. Les deux taxons « gen. et sp. A » et Oligolestes sont en positions les plus internes mais dans le même clade que les Sieblosiidae. Au sein des Sieblosiidae sensu stricto, les deux clades (Paraoligolestes + (Parastenolestes + Stenolestes)) et (Parastenolestes + Stenolestes) sont les mieux soutenus. La famille Sieblosiidae semble être restreinte à l’OligocèneMiocène d’Europe. © 2005 Elsevier SAS. All rights reserved. Keywords: Insecta; Odonata; Sieblosiidae; Phylogeny; Taxonomy; Oligocene–Miocene; France; Italy; Russia Mots clés : Insecta ; Odonata ; Sieblosiidae ; Phylogénie ; Taxonomie ; Oligocène-Miocène ; France ; Italie ; Russie

* Corresponding author. E-mail address: [email protected] (A. Nel). 0016-6995/$ - see front matter © 2005 Elsevier SAS. All rights reserved. doi:10.1016/j.geobios.2003.10.007

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1. Introduction The small damselfly family Sieblosiidae is strictly fossil, only known from the Oligocene and Miocene (30-15 M.y. ago) of Western Europe (France, Germany and Spain), with five described genera (Nel and Papazian, 1986; Nel, 1986, 1991; Nel and Escuillié, 1992, 1993; Nel and Paicheler, 1994; Riou and Nel, 1995; Nel et al., 1997). The exact affinities of this group remain uncertain. Their wing venation shows strong similarities with those of the Cenozoic and extant Lestidae but also with the Paleogene Thaumatoneuridae: Dysagrioninae and Frenguelliidae (Bechly, 1996; Petrulevicˇius and Nel, 2003). Nel and Paicheler (1994) considered that Sieblosiidae were related to Lestidae, because of the great similarities in their wing venation. Bechly (1996) transferred them to the very base of the zygopterid clade Caloptera sensu Bechly (1996) (=Sieblosiidae and Amphipterygida and Calopterygomorpha) but Fleck et al. (2004) doubted this position, relating them instead with the Epiproctophora Bechly, 1996 (formerly “Anisozygoptera”). Under the hypotheses of both Bechly (1996) and Fleck et al. (2004), the Sieblosiidae should be in a very basal position in the clade Odonata sensu Bechly (1996). This would imply a very important gap in their fossil record, till now restricted to the Oligocene and Miocene, because both sister clades Epiproctophora and Zygoptera are known since at least the Triassic. Another problem is their restricted palaeogeographic distribution in a relatively small

Fig. 1. Gen. and sp. A, specimen N 1734 (scale bar represents 10 mm). Fig. 1. Gen. and sp. A, spécimen N 1734 (l’échelle représente 10 mm).

area. Thus the new discoveries of Sieblosiidae in the Eastern part of Europe (Caucasus, Russia) and of a new Miocene genus, probably closely related to Sieblosiidae in Italy are of great interest for the diversity and palaeogeographic distribution of this family. In this work, we follow the wing venation nomenclature of Riek (1976); Riek and Kukalová-Peck (1984); amended by Kukalová-Peck (1991); Nel et al. (1993); Bechly (1996). The higher classification of fossil and extant Odonatoptera is based on the phylogenetic system of Bechly (1996).

2. Systematic palaeontology Family uncertain. Genus and species A. Figs. 1–3. Genus diagnosis: Broad wing; petiole short; base of RP2 in a very distal position; presence of an oblique crossvein “O”; bases of RP3/4 and IR2 distinctly midway between arculus and nodus; discoidal cells elongate; CuA curved; long pterostigma covering numerous cells; no pterostigmal brace; ScP not crossing through nodus; subnodus and nodal crossvein Cr of pronounced normal obliquity; part of CuA distal of nodus level shorter than proximal part; IR1 not strongly curved.

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Fig. 2. Gen. and sp. A, specimen N 1734, interpretive drawing (scale bar represents 5 mm). Fig. 2. Gen. and sp. A, spécimen N 1734, dessin interprétatif (l’échelle représente 5 mm).

Material: This specimen will be deposited in the future Museum of Pesaro, Italy; temporally deposited in the personal collection of Gabrielle Stroppa (N 1734), Pesaro, Italy. Species diagnosis: Presence of a dark zone, four cells basal and to middle half of pterostigma, and reaching level of RP3/4. Geological settings: Lower Messinian, Upper Miocene, base of the unit “Bituminous Marls”, 10 m below the level “Strato degli insetti”, in which more than 90% of all fossil dragonflies of Monte Castellaro have been discovered (Gentilini, 1989, 1992; Gentilini and Peters, 1993). Its age may be several thousand years older than that of other dragonflies from the same deposit. Monte Castellaro, Pesaro, Marches, Central Italy. Description: Thorax with four basal segments of abdomen and four wings partly overlapping. Thorax 7.0 mm long, 5.2 mm wide, meso-metathoracic suture present although weak; abdomen 1.3 mm wide; four wings nearly identical, hyaline except for a dark zone, four cells basal and to middle half of pterostigma, and reaching level of RP3/4; hind wing 31.0 mm long, 4.9 mm wide at nodus; distance from base to arculus about 5.2 mm; from arculus to nodus 6.2 mm, from nodus to pterostigma 14.2 mm, from pterostigma to apex

0.2 mm; nodus in a basal position; pterostigma long and broad, 3.5 mm long, 1.0 mm wide, covering six cells; pterostigmal brace absent; only three primary antenodal cross-veins Ax0, Ax1 and Ax2 present, Ax2 aligned with arculus, distance between Ax1 and Ax2 1.9 mm; discoidal cell unicellular, 1.8 mm long and 0.8 mm wide, rather long and narrow, not broadened in its distal part, and with its distal side MAb not parallel with basal side, basal side of discoidal cell 0.6 mm long, anterior side 1 mm long, posterior side 1.8 mm long, MAb 2.1 mm long; ScP not crossing through nodus, obliquity of nodal cross-vein Cr and subnodus Sn of normal type but well pronounced; 19 postnodal cross-veins, not aligned with the 23 postsubnodal cross-veins; bases of RP3/4 and IR2 between arculus and nodus; RP3/4 closer to arculus than to nodus; IR2 closer to nodus; base of RP2 six cells and 4.6 mm distal of subnodus; base of IR1 three cells and 2.2 mm distally from the base of RP2; oblique cross-vein “O” seven cells and 4.2 mm distal of base of RP2; vein CuP 0.4 mm distal of base of AA; cubito-anal area rather narrow, with two rows of cells between CuA and posterior wing margin; CuA reaching posterior wing margin distally, 2.1 mm distal of nodus level; part of CuA basal of nodus level 5.0 mm long,

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Fig. 3. Gen. and sp. A, specimen N 1734, wing bases (scale bar represents 2 mm). Fig. 3. Gen. and sp. A, spécimen N 1734, bases des ailes (l’échelle représente 2 mm).

part of CuA distal of nodus level 3 mm long; area between MP and posterior wing margin broad, with five rows of cells; postdiscoidal area narrow, 1 mm wide; area between MA and RP3/4 distally very broad; area between RP3/4 and IR2 narrow, 0.4 mm wide; IR2 very slightly zigzagged in its distal fourth; four secondary longitudinal veins in area between IR2 and RP2; basal part of IR1 straight and distal part slightly curved; two secondary longitudinal veins in area between RP2 and IR1; one row of cells in area between IR1 and RP1; one row of cells in area between RP1 and RA distal of pterostigma; CuP clearly curved between AA and MP + Cu. Discussion: The new species shares with the Sieblosiidae the following characters: broad wing, petiole short, base of RP2 in a very distal position, presence of an oblique crossvein “O”, bases of RP3/4 and IR2 distinctly midway between

arculus and nodus, discoidal cells quadrangular and closed; CuA curved; long pterostigma covering numerous cells, no pterostigmal brace. It strongly differs from Oligolestes and the sieblosiid genera and falls in the most basal position in the present phylogenetic analysis (see below) because of its subnodus and nodal cross-vein Cr of pronounced normal obliquity, and its ScP not crossing through nodus (plesiomorphies). Nevertheless, it shares with (Paraoligolestes + (Parastenolestes + Stenolestes)) the derived character state “part of CuA distal of nodus level shorter than proximal part”. Remarks: Gen. and sp. A has a CuP clearly curved as those of the Sieblosiidae genera Stenolestes and Parastenolestes. This character is a potential synapomorphy within Epiproctophora (Fleck et al., 2004).

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Family SIEBLOSIIDAE Handlirsch, 1906 sensu nov. Type genus: Sieblosia Handlirsch, 1906 stat. rest. Other genera: Stenolestes Scudder, 1895, Parastenolestes Nel and Paicheler, 1994, Paraoligolestes Nel and Escuillié, 1993, Miostenolestes nov. gen. The two genera Oligolestes Schmidt, 1958 and gen. and sp. A are probably related to the stem group of the Sieblosiidae. Diagnosis: Highly specialised nodus apparently traversed by ScP, as the terminal kink of CP is shifted basally together with the nodal and subnodal veinlets and the nodal membrane sclerotisation is reduced. This character is the only one to be restricted to the set [Sieblosia (with some doubt, see below), Stenolestes, Parastenolestes, Paraoligolestes, Miostenolestes], but it is plesiomorphically absent in the two taxa gen. and sp. A and Oligolestes. In order to define the Sieblosiidae as a monophyletic family, we characterise it on this last character alone, and thus exclude the two taxa gen. and sp. A and Oligolestes from it. Remarks: Bechly (1996) proposed the following other synapomorphies for the Sieblosiidae: (1) nodal furrow reduced, character shared by the Frenguelliidae Petrulevicˇius and Nel, 2003; (2) nodal and subnodal veinlets less oblique than in other Odonata, transverse or even with reversed obliquity, character shared by Frenguellidae and Dysagrioninae Cockerell, 1908, and plesiomorphically absent in gen. and sp. A; (3) pterostigmata very elongated, character shared by the Frenguellidae. The wing venations of gen. and sp. A and Oligolestes are very similar to those of the Sieblosiidae sensu n. They all share the following unique combination of characters: broad wing; petiole short; base of RP2 in a very distal position, four cells or more distal of subnodus; presence of an oblique crossvein “O”; bases of RP3/4 and IR2 distinctly midway between arculus and nodus; discoidal cells quadrangular and closed; CuA curved; long pterostigma covering six cells or more; no pterostigmal brace; a long and well defined IR1. Thus, gen. and sp. A and Oligolestes could be closely related to the Sieblosiidae sensu stricto, but there is no proof supporting this hypothesis because all these characters are individually present in other damselfly lineages. We consider gen. and sp. A and Oligolestes as Odonata of uncertain family position, probably belonging to the stem group of the Sieblosiidae. Wedmann (2000: Pl. 9, Fig. 1) figured from the Oligocene of Germany an adult specimen probably related to the genera Oligolestes or Sieblosia, because it has a subnodus subvertical, elongate discoidal cell, and long CuA. This fossil is of great interest because its colours are preserved, viz. its thorax is green metallic and its abdominal tergites are blue metallic, as in extant Zygoptera: Calopterygoidea or Lestoidea. The presence of very short apical abdominal appendages in the type specimen of Stenolestes cerestensis nov. sp. is not congruent with the hypothesis of affinities between the Sieblosiidae and the Lestida and Eucaloptera that have adult males with long to very long forceps-like cerci (Bechly, 1996). The very short appendages of this Stenolestes are more similar to those of the Coenagrionomorpha, but in this last clade,

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the lestine oblique vein “O” is absent (apomorphy after Bechly, 1996). Because of the state of preservation of this fossil, it is not possible to decide definitively if this specimen had four abdominal appendages as in Zygoptera or three as in Epiproctophora, but only three are apparently visible, i.e. a very small median one with a median furrow and two slightly longer lateral appendages, which could correspond to the “epiproctophoran” epiproct and paraprocts (see in S. cerestensis nov. sp.). Under this interpretation, the Sieblosiidae should be considered as very basal Epiproctophora, supporting Fleck et al. (2004). Genus Miostenolestes nov. Type species: M. zherikhini nov. sp. Etymology: Ater Miocene and Stenolestes. Diagnosis: ScP crossing through nodus; subcosta nearly perpendicular to RP; discoidal cell elongate; part of CuA distal of nodus level longer than proximal part; oblique crossvein “O” close to base of RP2; IR1 without any strong curve; One row of cells between RP3/4 and IR2, except for the presence of two rows close to posterior wing margin; postnodal cross-veins numerous; one row of cells in cubito-anal area. This last character can be considered as a “weak” autapomorphy of the genus because it corresponds to a reversal in our analysis. Miostenolestes zherikhini nov. sp. Figs. 4–6. Material: Holotype PIN 254/2169, Palaeoentomological Laboratory at the Academy of Science of Russia, Moscow. Etymology: In honour of our friend and colleague, the late Professor Vladimir Zherikhin, palaeontomologist at the Palaeoentomological Laboratory at the Academy of Science of Russia. Diagnosis: As for genus. Geological settings: Middle Miocene, Stavropol, Stavropol Province, North Caucasus, Russia.

Fig. 4. M. zherikhini nov. gen., nov. sp., holotype PIN 254/2169 (scale bar represents 10 mm). Fig. 4. M. zherikhini nov. gen., nov. sp., holotype PIN 254/2169 (l’échelle représente 10 mm).

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Fig. 5. M. zherikhini nov. gen., nov. sp., holotype PIN 254/2169, fore wing (scale bar represents 2 mm). Fig. 5. M. zherikhini nov. gen., nov. sp., holotype PIN 254/2169, aile antérieure (l’échelle représente 2 mm).

Fig. 6. M. zherikhini nov. gen., nov. sp., holotype PIN 254/2169, hind wing (scale bar represents 2 mm). Fig. 6. M. zherikhini nov. gen., nov. sp., holotype PIN 254/2169, aile postérieure (l’échelle représente 2 mm).

Description: Four wings and the five basal abdominal segments attached to thorax. Thorax 4.6 mm high; male secondary genital organs visible in abdominal segment 2; all wings hyaline; hind wing 37.4 mm long, 8.6 mm wide; distance from base to arculus 5.5 mm; from arculus to nodus 4.7 mm, from nodus to pterostigma 19.3 mm, from pterostigma to apex 4.9 mm; nodus in a very basal position; pterostigma long and broad, 4.4 mm long, 1.0 mm wide, covering six cells; pterostigmal brace absent; only three primary antenodal crossveins Ax0, Ax1 and Ax2 present, Ax2 aligned with arculus, distance between Ax1 and Ax2 2.0 mm; discoidal cell unicellular, 1.6 mm long and 0.6 mm wide, rather long and narrow, not broadened in its distal part, and with its distal side MAb not parallel with basal side, basal side of discoidal cell 0.4 mm long, anterior side 1.0 mm long, posterior side 1.6 mm long, MAb 0.9 mm long; ScP crossing through nodus, nodal crossvein Cr nearly perpendicular to RA and subnodus Sn only slightly oblique; 23 postnodal cross-veins, not aligned with the 20 postsubnodal cross-veins; bases of RP3/4 and IR2 between arculus and nodus, that of RP3/4 closer to arculus than to nodus, 1.5 mm distal of arculus, that of IR2 closer to nodus, 2.8 mm distal of arculus; base of RP2 seven cells and 6.5 mm distal of subnodus; base of IR1 three cells and 2.6 mm distally; oblique cross-vein “O” two cells and 1.4 mm distal of base of RP2; vein CuP 0.3 mm distal of base of AA; cubitoanal area narrow, with only one row of cells between CuA and posterior wing margin; CuA reaching posterior wing margin very distally, 3.7 mm distal of nodus level; part of CuA basal of nodus level 3.5 mm long, part of CuA distal of nodus

level 4.3 mm long; area between MP and posterior wing margin broad, with four rows of cells; postdiscoidal area narrow, 0.7 mm wide; area between MA and RP3/4 distally very broad; area between RP3/4 and IR2 narrow, 0.7 mm wide, but with two rows of cells near posterior wing margin; IR2 slightly zigzagged in its distal fourth; four secondary longitudinal veins in area between IR2 and RP2; basal part of IR1 weakly zigzagged and distal part slightly curved; two secondary longitudinal veins in area between RP2 and IR1; one row of cells in area between IR1 and RP1, except for two rows for a short distance just distal of pterostigma; one row of cells in area between RP1 and RA distal of pterostigma; preserved part of fore wing very similar to hind wing, except in the following points: wing 38.2 mm long; discoidal cell shorter and broader, 1.5 mm long and 0.7 mm wide; oblique cross-vein “O” three cells distal of base of RP2; 24 postnodal cross-veins and 22 postsubnodal cross-veins; curvature of IR2 just distal of pterostigma slightly more pronounced. Discussion: Its shortly petiolated but large wings, ScP crossing through nodus, nodal Cr vertical and subnodus subvertical, bases of veins RP3/4 and IR2 midway between arculus and nodus show that this fossil is a sieblosiid (Nel, 1986; Nel and Escuillié, 1992, 1993; Nel and Paicheler, 1994; Nel et al., 1997; Riou and Nel, 1995). Miostenolestes nov. gen. strongly differs from gen. and sp. A and Oligolestes in having its vein ScP clearly crossing through nodus (apomorphy). Other differences with Oligolestes grandis (Statz, 1935) are as follows: “pterostigma of Miostenolestes nov. gen. shorter and covering less cells (six instead of eight in O. grandis)”;

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“wings narrower”; “cubito-anal area with only one row of cells, instead of three in O. grandis”, and “more postnodal cross-veins (24 instead of 18)” (Statz, 1935; Schmidt, 1958; Fischer, 1974: Fig. 2). Miostenolestes nov. gen. also differs from Sieblosia in the following characters: “more postnodal cross-veins”; number and structure of secondary longitudinal veins in areas between RP3/4 and IR2, between RP2 and IR1, and between IR1 and RP1, number of rows of cells in cubito-anal area (see below the new diagnosis of Sieblosia). Miostenolestes nov. gen. shares with Oligolestes, Sieblosia and Paraoligolestes, unlike Parastenolestes and Stenolestes, the plesiomorphic character state “IR1 not strongly curved”. These four genera also differ from Stenolestes in their “fore and hind wing discoidal cells elongate, longer than broad”. Miostenolestes, Sieblosia and Oligolestes differ from the clade (Paraoligolestes + (Parastenolestes + Stenolestes)) as follows: “part of CuA distal of nodus level shorter than proximal part”; “presence of two or more rows of cells between RP3/4 and IR2 close to posterior wing margin” (plesiomorphy, but more precisely two long secondary longitudinal veins in Sieblosia, relatively short in Oligolestes, and two rows of cells for a distance of three cells in Miostenolestes). But this last character is reversed in Stenolestes ronzonense (Maneval, 1936) and Stenolestes iris Scudder, 1895. Miostenolestes differs from Paraoligolestes in the “oblique cross-vein ‘O’ in a basal position, only two cells distal of base of RP2”. Genus Sieblosia Handlirsch, 1906 stat. rest. Type species: Sieblosia jucunda (Hagen, 1858) stat. rest. Diagnosis: Pterostigma basally recessed, with a long area between RA and RP1 distal of pterostigma (autapomorphy); IR1 not curved, three long secondary longitudinal veins between RP2 and IR1, part of CuA distal of nodus level longer than proximal part, discoidal cell long, four rows of cells in cubito-anal area, nodal Cr and subnodus perpendicular to RP or nearly so, base of first secondary longitudinal vein between MA and RP3/4 opposite nodus (autapomorphy); 18 postnodal cross-veins; one long zigzagged secondary vein between IR1 and RP1 just distal of pterostigma. Geological settings: Oligocene, Sieblos, Germany. Remark: The location of the holotype of S. jucunda is unknown. It is probably lost (Fischer, 1974).

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Discussion: Hagen (1858) originally described this fossil in the genus Heterophlebia. Handlirsch (1906) transferred it to his monotypic genus Sieblosia, type genus of the family Sieblosiidae Handlirsch 1906. Fischer (1974) re-analysed the description and the figures of Hagen. Nel (1986) synonymised Sieblosia with Stenolestes. But S. jucunda lacks the main synapomorphies of (Parastenolestes + Stenolestes) or of Stenolestes alone, i.e. “IR1 distinctly curved”, “discoidal cell not elongate”, “more than 20 postnodal cross-veins”, “part of CuA distal of nodus level shorter than proximal part”. We have included S. jucunda in the phylogenetic analysis of the sieblosiid genera. It falls in a trichotomy with Oligolestes and the clade that comprises all the other sieblosiid genera (see the most parsimonious cladogram of the sieblosiid genera). This ambiguous situation is due to the conflict between the plesiomorphic character state “area between RP3/4 and IR2 with two secondary longitudinal veins”, absent in other Sieblosiidae, and the apomorphic character state “more than two secondary longitudinal veins between RP2 and IR1”, which is shared by Sieblosia and Stenolestes. Genus Paraoligolestes Nel and Escuillié, 1993. Type species: Paraoligolestes miocenicus Nel and Escuillié, 1993. Other species included: P. stavropolensis nov. sp. Paraoligolestes stavropolensis nov. sp. Fig. 7. Material: Holotype PIN 254/2171 (a complete wing), paratype PIN 254/2202 (a fragment of a wing base), Palaeoentomological Laboratory at the Academy of Science of Russia, Moscow. Etymology: After the locality Stavropol. Diagnosis: Closely related to P. miocenicus, the main differences being as follows: wing larger (42.5 mm long instead of 40.0 mm in P. miocenicus), more numerous postnodal cross-veins (25 instead of 21), and presence of only two rows of cells in cubito-anal area instead of three in P. miocenicus. Geological settings: Middle Miocene, Stavropol, Stavropol Province, North Caucasus, Russia. Description: Wing hyaline, 42.5 mm long, 9.1 mm wide; distance from base to arculus about 3.8 mm; from arculus to nodus 6.1 mm, from nodus to pterostigma 25.0 mm, from

Fig. 7. P. stavropolensis nov. sp., holotype PIN 254/2171, wing (scale bar represents 2 mm). Fig. 7. P. stavropolensis nov. sp., holotype PIN 254/2171, aile (l’échelle représente 2 mm).

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pterostigma to apex 4.8 mm; nodus in a very basal position; pterostigma long and broad, 4.3 mm long, 1.0 mm wide, covering four cells; pterostigmal brace absent; only three primary antenodal cross-veins Ax0, Ax1 and Ax2 present, Ax2 aligned with arculus, distance between Ax1 and Ax2 2.1 mm; discoidal cell unicellular, 1.6 mm long and 0.6 mm wide, rather long and narrow, not broadened in its distal part, and with its distal side MAb not parallel with basal side, basal side of discoidal cell 0.5 mm long, anterior side 1.1 mm long, posterior side 1.6 mm long, MAb 0.8 mm long; ScP crossing through nodus, nodal cross-vein Cr nearly perpendicular to RA and subnodus Sn only slightly oblique; 25 postnodal cross-veins, not aligned with the 27 postsubnodal crossveins; bases of RP3/4 and IR2 between arculus and nodus, that of RP3/4 closer to arculus than to nodus, 2.0 mm distal of arculus, that of IR2 closer to nodus, 2.7 mm distal of arculus; base of RP2 nine cells and 8.2 mm (PIN 254/2171) or seven cells (PIN 254/2202) distal of subnodus; base of IR1 four cells and 3.0 mm distally; oblique cross-vein “O” six cells and 4.7 mm distal of base of RP2; vein CuP 0.4 mm distal of base of AA; cubito-anal area rather narrow, with two rows of cells between CuA and posterior wing margin; CuA reaching posterior wing margin very distally, 3.1 mm distal of nodus level; part of CuA basal of nodus level 5.1 mm long, part of CuA distal of nodus level 4.3 mm long; area between MP and posterior wing margin broad, with five rows of cells; postdiscoidal area narrow, 0.8 mm wide; area between MA and RP3/4 distally very broad; area between RP3/4 and IR2 narrow, 0.8 mm wide; IR2 slightly zigzagged in its distal fourth; four secondary longitudinal veins in area between IR2 and RP2; basal part of IR1 straight and distal part slightly curved; two secondary longitudinal veins in area between RP2 and IR1; one row of cells in area between IR1 and RP1; one row of cells in area between RP1 and RA distal of pterostigma. Discussion: Its shortly petiolated but large wings, ScP crossing through nodus, nodal Cr vertical and subnodus subvertical, bases of veins RP3/4 and IR2 midway between arcu-

Fig. 8. S. fasciata nov. sp., holotype PIN 254/3192 (scale bar represents 5 mm). Fig. 8. S. fasciata nov. sp., holotype PIN 254/3192 (l’échelle représente 5 mm).

lus and nodus show that P. stavropolensis is a sieblosiid. It shares with the group [Miostenolestes + (Paraoligolestes + (Parastenolestes + Stenolestes))] the character state “ScP crossing through nodus”. P. stavropolensis shares with the clade (Paraoligolestes + (Parastenolestes + Stenolestes)) the two characters “part of CuA distal of nodus level shorter than proximal part” and “only one row of cells between RP3/4 and IR2, even close to posterior wing margin”. It differs from Parastenolestes and Stenolestes and it is very similar to P. miocenicus in its discoidal cell distinctly longer than broad, and its IR1 not strongly curved. It also shares with this last species the oblique vein in a very distal position. Genus Stenolestes Scudder, 1895. Type species: S. iris Scudder, 1895. Other species included: S. ronzonense (Maneval, 1936), Stenolestes coulleti Nel and Papazian, 1986, Stenolestes falloti (Théobald, 1937), Stenolestes fischeri Nel, 1986, Stenolestes camoinsi Nel, 1986, Stenolestes hispanicus Nel, 1991, Stenolestes dauphinensis Nel et al., 1997, Stenolestes belligaudi Nel et al., 1997, Stenolestes andancensis Riou and Nel, 1995, S. fasciata nov. sp., S. cerestensis nov. sp. Stenolestes fasciata nov. sp. Figs. 8 and 9. Material: Holotype PIN 254/3192 (a complete wing), Palaeoentomological Laboratory at the Academy of Science of Russia, Moscow. Etymology: After the presence of a dark zone crossing the distal two-thirds of wing. Diagnosis: Presence of a dark zone crossing the distal twothirds of wing; CuA well prolonged distal of nodus level; numerous (35) postnodal cross-veins; four rows of cells in cubito-anal area; two rows of cells between IR1 and RP1 just distal of pterostigma; base of RP2 nine cells distal of subnodus; no long zigzagged secondary veins in distal fourth of area between RP3/4 and IR2; seven rows of cells between RP2 and IR1; presence of two rows of cells in areas below pterostigma and distal of pterostigma.

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Fig. 9. S. fasciata nov. sp., holotype PIN 254/3192, wing (scale bar represents 2 mm). Fig. 9. S. fasciata nov. sp., holotype PIN 254/3192, aile (l’échelle représente 2 mm).

Geological settings: Middle Miocene, Vishnevaya Balka creek, Stavropol, Stavropol Province, North Caucasus, Russia. Description: Wing 45.4 mm long, 10.4 mm wide, hyaline, except for presence of a dark zone crossing its distal two-thirds; distance from base to arculus about 5.5 mm; from arculus to nodus 6.2 mm, from nodus to pterostigma 22.4 mm, from pterostigma to apex 4.1 mm; nodus in a very basal position; pterostigma long and broad, 4.7 mm long, 1 mm wide, covering 14 cells, some of then being disposed in two rows; pterostigmal brace absent; only three primary antenodal crossveins Ax0, Ax1 and Ax2 present, Ax2 nearly aligned with arculus, distance between Ax1 and Ax2 2.2 mm; discoidal cell unicellular, 1.7 mm long and 1 mm wide, rather short and broad, broadened in its distal part, with its distal side MAb not parallel with basal side, basal side of discoidal cell 0.6 mm long, anterior side 0.9 mm long, posterior side 1.1 mm long, MAb 1.1 mm long; ScP crossing through nodus, nodal cross-vein Cr and subnodus Sn perpendicular to RA; 35 postnodal cross-veins, not aligned with the 37 postsubnodal crossveins, some of them being in two rows; bases of RP3/4 and IR2 between arculus and nodus; RP3/4 closer to arculus than to nodus, 1.5 mm distal of arculus; IR2 closer to nodus, 3.4 mm distal of arculus; base of RP2 nine cells and 6.8 mm distal of subnodus; base of IR1 four cells and 2.6 mm distally; oblique cross-vein “O” four cells and 1.9 mm distal of base of RP2; vein CuP 0.8 mm distal of base of AA; cubitoanal area rather broad, with three rows of cells between CuA and posterior wing margin; CuA reaching posterior wing margin very distally, 4.5 mm distal of nodus level; part of CuA basal of nodus level 5.2 mm long, part of CuA distal of nodus level 5.5 mm long; area between MP and posterior wing margin broad, with seven rows of cells; postdiscoidal area narrow, 1.1 mm wide; area between MA and RP3/4 distally very broad, with six long secondary longitudinal veins; area between RP3/4 and IR2 narrow, distally 0.6 mm wide; IR2 not zigzagged; four secondary longitudinal veins in area between IR2 and RP2; basal part of IR1 slightly zigzagged and distal part strongly curved opposite distal half of pterostigma; two secondary longitudinal veins and seven rows of cells in area between RP2 and IR1; two rows of cells in area between IR1 and RP1; one to two rows of cells in area between RP1 and RA distal of pterostigma.

Discussion: Its shortly petiolated but large wings, ScP crossing through the nodus, nodal Cr and subnodus vertical, bases of veins RP3/4 and IR2 midway between arculus and nodus show that this fossil is a sieblosiid. It has the main characters of the genus Stenolestes, viz. ScP crossing through nodus, part of CuA distal of nodus level shorter than basal part, more than 20 postnodal cross-veins, IR1 with a strong curve, area between MA and RP3/4 with more than four long secondary longitudinal veins, discoidal cell rather short and broad, more than three rows of cells in area between IR1 and RP2, cells of cubito-anal area not elongate (Nel and Escuillié, 1993; Nel and Paicheler, 1994). S. fasciata nov. sp. can be separated from S. fischeri in its CuA prolonged distal of nodus level (Nel, 1986; Nel et al., 1997). S. dauphinensis has its base of RP2 only two cells distal of subnodus (Nel et al., 1997). S. belligaudi has distinctly fewer postnodal cross-veins (24 instead of 35 in S. fasciata nov. sp.), it has four rows of cells in the cubito-anal area, and its part of CuA distal of nodus level is distinctly shorter than in S. fasciata nov. sp. S. falloti has only 28 postnodal cross-veins but its wing is longer (48 mm long instead of 43 mm in S. fasciata nov. sp.) (Nel and Paicheler, 1994). S. coulleti differs from S. fasciata nov. sp. as follows: CuA reaching posterior wing margin opposite or just distal of nodus level, three rows of cells between IR1 and RP1 just distal of pterostigma, instead of two in S. fasciata nov. sp., five rows of cells in cubito-anal area. S. hispanicus differs from S. fasciata nov. sp. as follows: base of RP2 only five cells distal of subnodus, instead of nine cells in S. fasciata nov. sp., CuA reaching posterior wing margin opposite nodus (Nel, 1991). S. andancensis differs from S. fasciata nov. sp. as follows: postnodal cross-veins less numerous (23 instead of 35), subnodus of normal obliquity, presence of only four rows of cells between RP2 and IR1 instead of seven (Riou and Nel, 1995). S. camoinsis differs from S. fasciata nov. sp. as follows: four to five rows of cells in cubito-anal area, instead of three in S. fasciata nov. sp., only four rows of cells between RP2 and IR1, base of RP2 four to five cells distal of subnodus. S. ronzonense differs from S. fasciata nov. sp. in the presence of two rows of cells in postdiscoidal area at nodus level, and the presence of six long secondary longitudinal veins in area between RP2 and IR1, instead of two in S. fasciata nov. sp. (Maneval, 1936; Nel, 1986; Nel and Paicheler, 1994). S. iris

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differs from S. fasciata nov. sp. as follows: presence of two long zigzagged secondary veins in distal fourth of area between RP3/4 and IR2 (Scudder, 1895). The wing base of S. iris is unknown but other characters of distal halves are similar in S. iris and S. fasciata nov. sp. Stenolestes (?) adygeianensis nov. sp. Fig. 10. Material: Holotype PIN 4705/1, Palaeoentomological Laboratory at the Academy of Science of Russia, Moscow. Etymology: After Adygeia Republic. Diagnosis: Wing with a large black zone in its distal twothirds (limits shown in Fig. 10 by dotted lines); oblique crossvein “O” in a very distal position; numerous, 26 postnodal cross-veins; presence of long secondary longitudinal veins in areas between MA and RP3/4, RP3/4 and IR2, IR2 and RP2, but not in postdiscoidal area; seven rows of cells in area between RP2 and IR1; two rows of cells between IR1 and RP1; cubito-anal area broad, with three rows of cells; part of CuA distal of nodus level shorter than proximal part. Geological settings: Lower Oligocene, Belaya river upstream Abadzekhskaya village, North Caucasus, Adygeia Republic, Russia. Description: Wing apex and base not preserved; basal part of wing hyaline, part between nodus and pterostigma black, wing about 38.7 mm long, 10.4 mm wide; distance from base to arculus unknown, probably about 5.3 mm; from arculus to nodus 7.7 mm, from nodus to pterostigma 17 mm, from pterostigma to apex 4.9 mm; nodus in a basal position; pterostigma long and broad, 4 mm long, 0.8 mm wide, covering probably about 10 cells; pterostigmal brace absent; only Ax2 Ax2 preserved, aligned with arculus, discoidal cell unicellular, 1.9 mm long and about 0.6 mm wide, rather long and narrow, not broadened in its distal part, and with its distal side MAb not parallel with basal side, anterior side of discoidal cell 0.9 mm long, MAb 1 mm long; extreme apex of ScP poorly preserved, nodal cross-vein Cr slightly oblique and subnodus Sn perpendicular to RA and RP; 26 postnodal crossveins, not aligned with the 31 postsubnodal cross-veins, the most distal postsubnodal cross-veins disposed in two rows

opposite pterostigma; bases of RP3/4 and IR2 between arculus and nodus; RP3/4 closer to arculus than to nodus, 1.9 mm distal of arculus; IR2 closer to nodus, 3.8 mm distal of arculus; base of RP2 four cells and 3 mm distal of subnodus; base of IR1 three cells and 2.3 mm distally; oblique cross-vein “O” in a very distal position, 15 cells and 5.2 mm distal of base of RP2; cubito-anal area with three rows of cells between CuA and posterior wing margin; CuA reaching posterior wing margin distally, 2.7 mm distal of nodus level; part of CuA basal of nodus level 6.4 mm long, part of CuA distal of nodus level 3.4 mm long; area between MP and posterior wing margin very broad, with 12 rows of cells; postdiscoidal area narrow, 1 mm wide, but with a six-cells long secondary longitudinal vein near posterior wing margin; area between MA and RP3/4 distally very broad, with six long secondary longitudinal veins between them, the base of the most basal one being opposite base of RP2; area between RP3/4 and IR2 relatively broad, with one to two long secondary longitudinal veins between them, distal of level of base of RP2, 1 mm wide; IR2 with a double slight curve; two visible secondary longitudinal veins in area between IR2 and RP2; basal part of IR1 zigzagged, part basal of pterostigma straight, but distal part not preserved; seven rows of cells and five to six secondary longitudinal veins in area between RP2 and IR1; two rows of cells in area between IR1 and RP1. Discussion: Its very large wings, ScP probably crossing through nodus, nodal Cr subvertical and subnodus vertical, bases of veins RP3/4 and IR2 midway between arculus and nodus show that Stenolestes (?) adygeianensis is a sieblosiid, related to the clade [Sieblosia and Miostenolestes and (Paraoligolestes + (Parastenolestes + Stenolestes))]. It shares with the clade (Paraoligolestes + (Parastenolestes + Stenolestes)) the character “part of CuA distal of nodus level shorter than proximal part”, but it does not share the character state “only one row of cells between RP3/4 and IR2, even close to posterior wing margin”. It differs from Miostenolestes, Paraoligolestes, and Parastenolestes in the “presence of more than two long secondary longitudinal veins between RP2 and IR1”. This last character is present in Stenolestes and Sieblosia. It differs from Sieblosia in having the part of CuA distal of nodus

Fig. 10. Stenolestes (?) adygeianensis nov. sp., holotype PIN 4705/1, wing (scale bar represents 2 mm). Fig. 10. Stenolestes (?) adygeianensis nov. sp., holotype PIN 4705/1, aile (l’échelle représente 2 mm).

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level distinctly shorter than proximal part. Nevertheless, it shares with Sieblosia the presence of very long secondary veins between MA and RP3/4. But this last character is convergently present in S. ronzonense. Thus, we provisionally attribute this fossil to the genus Stenolestes. Nevertheless, this attribution will need further confirmation after the discovery of its distal part of IR1 and of its wing base. Stenolestes (?) adygeianensis nov. sp. greatly differs from the other Stenolestes spp., except S. ronzonense in the great number of cells and secondary longitudinal veins in the distal two-thirds of the wing. It shares with S. ronzonense the presence of numerous secondary veins in areas between RP3/4 and MA, RP3/4 and IR2, and between IR2 and RP2. But S. ronzonense has also a secondary longitudinal vein in the postdiscoidal area, unlike S. (?) adygeianensis nov. sp. Also, the oblique cross-vein “O” is in a very basal position in S. ronzonense, unlike S. (?) adygeianensis nov. sp. Stenolestes cerestensis nov. sp. Figs. 11–15. Material: MNHN-LP-R 63846, Laboratory of Palaeontology, Muséum National d’Histoire Naturelle, Paris, France (a nearly complete specimen with the four wings in connection with the thorax). Etymology: After the type locality Céreste. Diagnosis: Wings hyaline, moderately long, about 38 mm long; 15–17 postnodal cross-veins; only four cells in area between RP and IR2, basal of RP2 and distal of subnodus level; CuA short, ending opposite nodus level; three to four rows of cells in cubito-anal area; three to four rows of cells in area between RP2 and IR1; curve of IR1 moderate; oblique cross-vein “O” in a distal position.

Fig. 12. S. cerestensis nov. sp., holotype in coll. Patrick Brisac, distal part of abdomen (scale bar represents 10 mm). Fig. 12. S. cerestensis nov. sp., holotype in coll. Patrick Brisac, partie distale de l’abdomen (l’échelle représente 10 mm).

Geological settings: Oligocene, Stampian, Céreste, Vaucluse, France. Description: Wings hyaline, hind wing 37.7 mm long, 8.0 mm wide; distance from base to arculus about 4.5 mm;

Fig. 11. S. cerestensis nov. sp., holotype in coll. Patrick Brisac (scale bar represents 10 mm). Fig. 11. S. cerestensis nov. sp., holotype in coll. Patrick Brisac (l’échelle représente 10 mm).

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Fig. 13. S. cerestensis nov. sp., holotype in coll. Patrick Brisac, last abdominal segments (scale bar represents 1 mm). Fig. 13. S. cerestensis nov. sp., holotype in coll. Patrick Brisac, derniers segments abdominaux (l’échelle représente 1 mm).

from arculus to nodus 8.1 mm, from nodus to pterostigma 17.3 mm, from pterostigma to apex about 4.0 mm; pterostigma long and broad, 4.9 mm long, 0.8 mm wide, covering seven cells; only three primary antenodal cross-veins Ax0, Ax1 and Ax2 present, Ax2 nearly aligned with arculus, distance between Ax1 and Ax2 2.0 mm; discoidal cell unicellular, 1.5 mm long and 1.0 mm wide, rather short and broad, broadened in its distal part but longer than that of fore wing, with its distal side MAb nearly parallel with basal side, basal side of discoidal cell 0.7 mm long, anterior side 1.2 mm long, posterior side 1.5 mm long, MAb 1.0 mm long; ScP crossing through nodus, nodal cross-vein Cr of normal obliquity; subnodus Sn perpendicular to RA; 17 postnodal cross-veins, not aligned with the 16 postsubnodal cross-veins; bases of RP3/4 and IR2 between arculus and nodus; RP3/4 closer to

arculus than to nodus, 2.7 mm distal of arculus; IR2 closer to nodus, 5.0 mm distal of arculus; base of RP2 four cells and 4.5 mm distal of subnodus; only four cells in area between RP and IR2, basal of RP2 and distal of subnodus level; base of IR1 four cells and 3.3 mm distally; oblique cross-vein “O” seven cells and 4.8 mm distal of base of RP2; vein CuP 0.8 mm distal of base of AA; cubito-anal area 1.8 mm wide, with three rows of cells between CuA and posterior wing margin; CuA reaching posterior wing margin very basally, nearly opposite nodus level; area between MP and posterior wing margin broad, with eight to nine rows of cells; postdiscoidal area narrow, 1.0 mm wide; area between MA and RP3/4 distally very broad, with two main secondary longitudinal veins; area between RP3/4 and IR2 narrow, distally 1.1 mm wide, but with a long secondary longitudinal vein between them; IR2 zigzagged in its distal end; three secondary longitudinal veins in area between IR2 and RP2; basal part of IR1 slightly zigzagged and distal part rather weakly curved opposite distal half of pterostigma; two secondary longitudinal veins and three rows of cells in area between RP2 and IR1; two rows of cells in area between IR1 and RP1; one row of cells in area between RP1 and RA distal of pterostigma; fore wing very similar to hind wing, except in the following points: fore wing 37.5 mm long, 8.3 mm wide; distance from base to arculus about 4.5 mm; from arculus to nodus 7.7 mm, from nodus to pterostigma 17.5 mm, from pterostigma to apex 4.6 mm; nodus in a very basal position; pterostigma long and broad, 4.3 mm long, 1.0 mm wide, covering only four cells; pterostigmal brace absent; discoidal cell 1.25 mm long and 1.25 mm wide, distinctly shorter and broader than that of hind wing,

Fig. 14. S. cerestensis nov. sp., holotype in coll. Patrick Brisac, hind wing (scale bar represents 2 mm). Fig. 14. S. cerestensis nov. sp., holotype in coll. Patrick Brisac, aile postérieure (l’échelle représente 2 mm).

Fig. 15. S. cerestensis nov. sp., holotype in coll. Patrick Brisac, fore wing (scale bar represents 2 mm). Fig. 15. S. cerestensis nov. sp., holotype in coll. Patrick Brisac, aile antérieure (l’échelle représente 2 mm).

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only 15 postnodal cross-veins, not aligned with the 17 postsubnodal cross-veins; base of IR1 two cells and 2.4 mm distal of that of RP2; cubito-anal area 2.5 mm wide, with four rows of cells between CuA and posterior wing margin; area between RP3/4 and IR2 narrower than in hind wing, with no secondary longitudinal vein between them; distal part of IR1 more strongly curved opposite distal half of pterostigma; two secondary longitudinal veins and four rows of cells in area between RP2 and IR1. Abdomen complete, first segment very short, 3.5 mm long, second 6.0 mm long, third and fourth 6.5 mm long, fifth 7.0 mm long, sixth 4.5 mm long, seventh and eight 2.5 mm long. Only three apical abdominal appendages apparently visible, i.e. a very small median one with a median furrow and two slightly longer lateral appendages, 0.7 mm long, which could correspond to the “epiproctophoran” epiproct and paraprocts. Discussion: This fossil can be attributed to the genus Stenolestes for the same reasons as for S. fasciata nov. sp. It differs from all the other Stenolestes spp., except S. belligaudi, S. hispanicus, and S. fischeri, in its short CuA, ending opposite nodus level and low number of postnodal cross-veins. In particular, S. coulleti from the same locality has five to six rows of cells in cubito-anal area and more than 28 postnodal crossveins. S. falloti has a longer CuA, extending well distal of nodus level. The wings of S. fischeri are completely dark blue and distinctly longer (44 mm long instead of 38 mm in S. cerestensis nov. sp.). The wings of S. belligaudi are hyaline but as long as those of S. fischeri. The preserved structures of S. hispanicus are very similar to those of S. cerestensis nov. sp., except for the presence of seven cells in the area between RP and IR2, basal of RP2 and distal of the subnodus level, instead of four. 3. Phylogenetic analysis of the “sieblosiid-like” genera Nel and Escuillié (1993) proposed a phylogenetic analysis of the Sieblosiidae, including Oligolestes. We have undertaken a new analysis of the “sieblosiid-like” genera (Sieblosiidae plus gen. and sp. A and Oligolestes). The chosen outgroups are the genera Lestes Leach, 1815 (Oligocene to Recent, Zygoptera: Lestidae), Dysagrion Scudder, 1878 (Eocene, Zygoptera: Dysagrioninae) and Frenguellia Petrulevicˇius and Nel, 2003 (Eocene, ?Epiproctophora: Frenguelliidae), because they show strong similarities of different kinds with the Sieblosiidae in their wing venations. The choice of outgroups that would be closely related to the Sieblosiidae is problematic because the exact affinities of the Sieblosiidae in Odonata remain uncertain (Fleck et al., 2004). The analysis is based on 13 unordered and equally weighted characters of wing venation (see Tables 1 and 2). We rejected several other characters of the wing venation because they are variable within the sieblosiid genera (position of oblique vein “O” of specific value in the Sieblosiidae, variable in the genus Stenolestes (Nel and Paicheler, 1994), number of supplementary veins between RP3/4 and MA).

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Table 1 List of characters Liste des caractères 1. Vein ScP... Not crossing through the nodus. 0 Crossing through the nodus. 1 2. Subnodus... Of pronounced normal obliquity. 0 Nearly vertical, vertical or of inversed obliquity. 1 3. Discoidal cell... Longer than wide. 0 Broader than long. 1 4. Angle between MAb and MP + CuA... Very sharp, between 45° or less. 0 Opened, more than 45° and less than 90°. 1 5. Part of CuA distal of nodus level... Longer than proximal part of CuA. 0 Shorter than proximal part of CuA. 1 6. Cubito-anal area... With one row of cells. 0 With two rows of cells or more. 1 With more than three rows of cells. 2 7. Base of RP2... Close to subnodus (four cells or less). 0 Distal of subnodus (more than four cells). 1 8. Area between RP3/4 and IR2... With one to two long secondary longitudinal veins. 0 Without secondary longitudinal veins. 1 9. Number of postnodal cross-veins... Less than 15. 0 More than 15. 1 10. IR1... Strongly zigzagged and/or poorly defined. 0 Well-defined, weakly zigzagged but without a strong curve. 1 With a strong curve. 2 11. Number of secondary longitudinal veins between RP2 and IR1... Two or less. 0 More than two. 1 12. Pterostigma... Relatively short, covering less than six cells. 0 Long, covering six cells or more. 1 13. General shape of wing... Not very broad, with a long petiole. 0 Very broad, with a short petiole. 1

The analysis was made using the software Paup 4.0b10 for PC, using the branch and bound option. The choice and order of introduction of the outgroup(s) do not perturb the result of the analysis. The resulting most parsimonious tree is 21 steps long, with a consistency index CI 0.66, CI excluding uninformative character 0.63, retention index RI 0.65, and RC 0.43 (see Fig. 16). Gen. and sp. A, Oligolestes and the Sieblosiidae fall in the same clade, supported by the characters states “base of RP2 well distal of subnodus”, and “pterostigma long, covering six cells or more”. The monophyly of the clade (Oligolestes + Sieblosiidae) is supported by the character state “angle between MAb and MP + CuA very opened”. The clade [Sieblosia and Miostenolestes and (Paraoligolestes + (Parastenolestes + Stenolestes))] is supported by the character states “ScP crossing through nodus”. But the exact structure of the nodus is uncertain in Sieblosia, even if, after the figures in Hagen (1858), ScP probably crossed through the nodus, as in Stenolestes. We have chosen to code the state “1” for this character

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Table 2 List of characters and taxa Liste des caractères et des taxons Taxa/characters Lestes Dysagrion Frenguellia Gen. and sp. A Oligolestes Paraoligolestes Miostenolestes nov. gen. Sieblosia Parastenolestes Stenolestes

1 0 0 0 0 0 1 1 1 1 1

2 0 1 1 0 1 1 1 1 1 1

3 0 1 ? 0 0 0 0 0 0 1

in Sieblosia in the present analysis. But if this character is coded “unknown” (or “0”) for Sieblosia, the analysis gives nine (or seven) minimal cladograms (Length 21 steps, CI 0.66, CI excluding uninformative characters 0.63, RI 0.63, RC 0.42), with a clade that comprises all the Sieblosiidae plus Oligolestes and gen. and sp. A, but also with a polytomy between Oligolestes, Sieblosia, Miostenolestes, Paraoligo-

4 0 1 0 0 1 1 1 1 1 1

5 0 0 0 1 0 1 0 0 1 1

6 0 1 1 1 1 1 0 1 1 1

7 0 0 0 1 1 1 1 1 1 1

8 0 0 1 1 1 1 1 0 1 1

9 0 1 1 1 1 1 1 1 1 1

10 0 1 1 1 1 1 1 1 2 2

11 0 0 0 0 0 0 0 1 0 1

12 0 0 0 1 1 1 1 1 1 1

13 0 1 1 1 1 1 1 1 1 1

lestes, and (Parastenolestes + Stenolestes) in their strict consensus tree, because of the conflicts affecting other characters. The clade (Paraoligolestes + (Parastenolestes + Stenolestes)) is supported by the synapomorphy “part of CuA distal of nodus level shorter than proximal part”, but also present in gen. and sp. A. The clade (Parastenolestes + Stenolestes) is supported by the character state “IR1 with a strong curve”.

4. Palaeobiogeographic remarks The present discoveries show that the Sieblosiidae are not restricted to the Western Europe but were also present and diverse in the Oligocene and Miocene of Caucasus. This region was connected to Western Europe during the Eocene. The Sieblosiidae are still unknown in the Cenozoic of the Asiatic part of Russia and in North America. If the Cenozoic odonatofauna of Siberia is still very badly known, it is not the case for the North American one. Interestingly, there was an epicontinental sea crossing through Russia at the limit between Europe and Asia during the Eocene. It could have acted as a boundary, preventing the arrival of the Sieblosiidae in Asia and North America in the Upper Eocene and Lower Oligocene. Later, the opening of the Bering Detroit in the Miocene also acted as a barrier between Asia and North America for this group of poor flyers.

Acknowledgements

Fig. 16. Most parsimonious cladogram of the sieblosiid genera. Fig. 16. Cladogramme le plus parcimonieux des genres de Sieblosiidae.

Many thanks to Mr. Gabriele Stroppa of Pesaro, Italy, for the loan of the specimen of gen. and sp. A, and its deposit in the future Museum of Pesaro, and to Mr. Patrick Brisac of Malataverne, France, for the photography, loan, and future deposit of the type specimen of S. cerestensis nov. sp. in the Museum of Paris. We are also greatly in debt towards the late Dr. Vladimir Zherikhin, Palaeoentomologist at the Academy of Science of Moscow.

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