[Palaeontology, Vol. 51, Part 1, 2008, pp. 149–162] NEW FOSSIL LACEWINGS AND ANTLIONS (INSECTA, NEUROPTERA) FROM THE LOWER CRETACEOUS CRATO FORMATION OF BRAZIL by FEDERICA MENON* and VLADIMIR N. MAKARKIN *School of Earth, Atmospheric and Environmental Sciences, University of Manchester, Williamson Building, Oxford Road, Manchester M13 9PL, UK; e-mail: [email protected] Institute of Biology and Soil Sciences, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok 690022, Russia; e-mail: [email protected] Typescript received 8 August 2006; accepted in revised form 22 January 2007 Abstract: Remarkable new fossil taxa of Neuroptera from on a new interpretation of venational characters evident on the laminated limestone of the Crato Formation, north-east the exceptionally well-preserved specimen of P. magnus; vein Brazil, are described: Nuddsia longiantennata gen. et sp. nov. homologies are determined and the fusion of MP and CuA is (Osmylidae, Gumillinae), the first fossil record of this family discussed. The genus Triangulochrysopa has been known pre- from South America, Parapalaeoleon magnus gen. et sp. nov. viously only from the Lower Cretaceous of Las Hoyas, Spain. (Palaeoleontidae), and Triangulochrysopa formosa sp. nov. (Mesochrysopidae). A diagnosis of Gumillinae is provided; Key words: Araripe, Crato Formation, Nova Olinda Mem- Epiosmylidae is considered to be a synonym of this subfam- ber, Neuroptera, Osmylidae, Palaeoleontidae, Mesochrysopi- ily. A revised diagnosis of Palaeoleontidae is provided, based dae, taxonomy. The Crato Formation is well known for the diversity and gart, Germany. No fossil record of Osmylidae was known quality of its fossil insects; three-dimensional features and hitherto from South America. colour pattern are often preserved (Martill and Frey 1995; Heads et al. 2005). The formation ranges across the Ara- ripe Plateau in the states of Ceara`, Pernambuco and Pia- MATERIAL AND METHODS uı`, north-east Brazil. It is composed of a series of finely laminated limestones that accumulated at the bottom of a All specimens described here are from the lowest member lagoon during the initial phase of the opening of the of the Crato Formation. They were probably collected by Atlantic Ocean (Martill 1993). Its precise age is somewhat workers in one of the small mines or stone yards around unclear: it is generally considered to be Late Aptian–Early Nova Olinda, although the exact locality is unknown. Albian, 110–120 Ma (Berthou 1994), and has been dated They are preserved as limonitic replacements after pyrite as Late Aptian on the basis of palynological data (Pons (Martill and Frey 1995). The specimens were prepared et al. 1991). The formation is subdivided into three mem- on-site by the collectors. An aeroneedle was used, where bers: Nova Olinda, Barbalha and Jamacaru. The basal necessary, to remove any residual sediment and dust Nova Olinda Member is the most fossiliferous: insects are (Selden 2003). found together with other arthropods, plants and wood, Drawings were made using a camera lucida attached to fish, pterosaurs and other vertebrates (Maisey 1991). a Wild stereomicroscope; photographs were taken with a The order Neuroptera is one of the most significant Sony DCS-717 digital camera and a D1X digital camera and diverse groups of insects found in the Crato Forma- attached to a Wild M8 stereo-zoom microscope. tion, with 55 species having been described (Martins-Neto 2000, 2003, 2005; Heads et al. 2005; Makarkin and Menon 2005, 2007; Menon et al. 2005; Nel et al. 2005). A SYSTEMATIC PALAEONTOLOGY vast amount of material in a variety of museums remains unstudied. We describe here two new genera and three Wing venation terminology follows Comstock (1918), with a few new species belonging to the families Osmylidae, Pala- exceptions in accordance with current usage in neuropterology eoleontidae and Mesochrysopidae, based on specimens (see Archibald and Makarkin 2006; Wedmann and Makarkin housed at the Staatlisches Museum fu¨r Naturkunde Stutt- 2007). Terminology of wing spaces follows Oswald (1993) ª The Palaeontological Association doi: 10.1111/j.1475-4983.2007.00740.x 149 150 PALAEONTOLOGY, VOLUME 51 Venation abbreviations used in the text and figures are as fol- Karatau, Kazakhstan). Another species of this genus lows: 1A–3A, anal veins; bf, anterior Banksian fold; Cu, cubitus; (E. panfilovi Ren and Yin, 2002) was added later from the CuA, anterior cubitus; CuA1, most proximal branch of anterior Middle–Upper Jurassic of China (Ren and Yin 2002). cubitus; CuP, posterior cubitus; M, media; MA, anterior branch Since its establishment, Epiosmylidae has been generally of media; mfl, median flexion line; MP, posterior branch of treated as either a synonym of Osmylidae (e.g. Pono- media; MP1, most proximal branch of MP; R, radius; R1, first marenko 1980; Makarkin 1990b; Ren and Yin 2002) or a branch of radius; Rs, radial sector; Rs1, most proximal branch of taxon whose status is unclear, ‘either an isolated branch radial sector; Sc, subcosta. within Osmylidae or else a separate family’ (Makarkin Institutional abbreviation. SMNS, Staatliches Museum fu¨r and Archibald 2003, p. 175). Recently, Ponomarenko Naturkunde Stuttgart, Germany. (2003) described another osmylid-like species with extre- mely long antennae, ‘Nymphites cf. lithographicus’, from the Upper Jurassic of Solnhofen, Germany, and assigned Class INSECTA Linnaeus, 1758 it to the Nymphitidae. However, the photograph and Order NEUROPTERA Linnaeus, 1758 drawings of this specimen provided indicate the possibil- Family OSMYLIDAE Leach, 1815 ity that it belongs to Osmylidae (possibly to Gumillinae Subfamily GUMILLINAE Nava´ s, 1912 because of its long antennae); its venation is most similar to that found in this family, and its relation to the true 1912 Gumillini Nava´s, p. 191 (as a ‘tribe’ of Osmylidae). Nymphites lithographicus Handlirsch, 1906 is unclear, as 1988 Gumillinae Lambkin, p. 455 (as a subfamily of its holotype is very poorly preserved. Osmylidae). The subfamily Gumillinae has not been formally 1980 Epiosmylidae Panfilov, in Dolin et al., p. 100, syn. defined. The ‘tribe’ Gumillini was created by Nava´s nov. (1912) within Osmylidae for the genus Gumilla Nava´s, including the two extant Brazilian species, G. adspersa Type genus. Gumilla Nava´s, 1912 (Recent). Nava´s, 1912 and G. longicornis (Walker, 1853), which should perhaps be considered synonyms (Adams 1977). Diagnosis. Medium-sized osmylids that may be distin- Nava´s (1912) grouped all other osmylids in the ‘tribe’ guished from other species of the family by the following Osmylini; thus the actual rank of these ‘tribes’, according combination of character states: (1) antennae exceedingly to Nava´s (1912), could be considered at the subfamily elongate, much longer than forewing [much less than level. Gumilla was subsequently excluded from the family forewing length in other subfamilies]; (2) scape strongly by Kru¨ger (1913, 1915), then included again by Adams enlarged [moderately enlarged in other osmylids]; (3) (1969, 1977). Adams (1969, p. 2) mentioned that this basal sinuous crossvein r-m in hindwing absent, character genus is highly aberrant and ‘is too poorly known for state shared with Stenosmylinae [present in other subfam- meaningful discussion’. He correctly concluded that Nav- ilies]; (4) outer gradate series of crossveins absent in both a´s’ (1912, fig. 24) drawing of the forewing is reasonably wings, as in Porisminae [present in other subfamilies]; (5) accurate, and provided some additional diagnostic fea- trichosors limited at most to apical part of wings [nor- tures of the genus (Adams 1977). Unfortunately, the char- mally present at least in distal half of wings in other os- acters of the male terminalia are largely unknown, beyond mylids]. the short description of Nava´s (1912, p. 57) (‘cercis cylin- dricis, pilosis, brevibus, externe convexis’), and they have Included genera. Gumilla (Recent, Brazil; one or two species); never been illustrated. Epiosmylus Panfilov, in Dolin et al. 1980 (Upper Jurassic, Kara- It is reasonable to assume that Epiosmylidae and tau, Kazakhstan, Karabastau Formation; Middle–Upper Jurassic, Gumillinae might be the same, as they share some impor- Daohugou, Inner Mongolia, China; two species); Nuddsia gen. nov. (Lower Cretaceous, Upper Aptian, Araripe Basin, Brazil, tant character states (see diagnosis, above), even though Crato Formation; one species). Epiosmylus and Gumilla, their type genera, differ quite considerably by the configuration of MA, MP, CuA and Remarks. Three taxa with extremely long antennae, at CuP in the forewing, which are straighter with branching least 1.5 times longer than the forewing length, are known that is fairly regular in the former and strongly irregular within Neuroptera. These include the extant Chrysopidae, in the latter (see also note of Lambkin 1988, p. 455). in the majority of Apochrysinae and a few genera of Examination of the photograph of the holotype of G. ad- Chrysopinae, and the Osmylidae, in the subfamily Gumil- spersa reveals that its forewing venation is somewhat linae and the fossil Epiosmylidae. anomalous (the venation of right and left wings is The family Epiosmylidae was erected to accommodate strongly asymmetrical). The forewing venation of Nuddsia Epiosmylus longicornis Panfilov, 1980 (Upper Jurassic of (e.g. configuration of MP, CuA, CuP) is also irregular to MENON AND MAKARKIN: CRETACEOUS FOSSIL LACEWINGS AND ANTLIONS FROM BRAZIL 151 some extent (Text-fig. 2), providing additional support Diagnosis. As for the genus. for placing it in Gumillinae. The hindwing venation of all species (including the holotype of G. adspersa and the Description. Head twice as wide as long (1 mm long, 2 mm new species) does not differ in its main character states. wide); large compound eyes. Antennae extremely long (60 mm), Ocelli are absent in the extant Gumilla (present in all filiform; scape very large, bulky; pedicel elongate, rounded, larger other osmylids) but are not detectable in fossil taxa than flagellar segments. Prothorax nearly quadrate, c.1mm because of poor preservation or as the head is missing. long. Mesothorax not clearly visible, apparently wide.