NEW FOSSIL LACEWINGS and ANTLIONS (INSECTA, NEUROPTERA) from the LOWER CRETACEOUS CRATO FORMATION of BRAZIL by FEDERICA MENON* and VLADIMIR N

Home , Nemopteridae, Nymphidae, Osmylidae

[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 extremely 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. Legs not preserved. Abdomen incomplete, 4 mm long as preserved. Head, Gumillinae is considered here to be a subfamily of thorax and abdomen strongly compressed. Exoskeleton of most Osmylidae because its venation generally agrees well with portions of head, thorax and abdomen not preserved. that of this family, and to be the sister group of all other Forewing elongate, 25.5 mm long, c. 6.5 mm wide; apex sub- osmylids because of the clear differences between them acute. Trichosors prominent, restricted to apical portion of wing. (see diagnosis above). The loss of the basal sinuous cross- Costal space 0.6 mm maximum width, narrow basally, slightly vein r-m in the hindwing and the absence of the outer dilated near proximal third, then narrowed towards apex. Sub- gradate series of crossveins in both wings, shared by costal veinlets simple, straight, nearly perpendicular to Sc in Gumillinae with Stenosmylinae (the former) and Porismi- basal two-thirds of costal space, increasingly more oblique dis- nae (the latter) might be convergent. tad; veinlets of Sc+R1 closely spaced, strongly oblique. Subcostal space narrow, dilated near fusion of Sc, R1. Sc, R1 fused 4 mm from wing apex. Sc+R1 entering margin before wing apex. Rs originating near wing base. Stem of Rs zigzagged, forked api- Genus NUDDSIA gen. nov. cally, entering margin slightly before wing apex. R1 space wider than costal space, 0.8 mm, strongly narrowed apically, with 23 Type and only species. Nuddsia longiantennata sp. nov. more or less regularly spaced crossveins proximad fusion of Sc, R1, one distad (left wing). Rs with six branches, distal branch Derivation of name. After Dr John Nudds, Head of the Palaeon- deeply forked. Origin of Rs1 at approximately one-third wing tology Research Group at the University of Manchester. Gender length, c. 8 mm from base (left wing); origins of Rs1–Rs3 widely feminine. spaced, other three branches restricted to apical portion of wing. Radial crossveins numerous, regularly spaced (except for apical Diagnosis. May be distinguished from other genera of the portion of space, where crossveins rare), not forming gradate subfamily by the following combination of character series. Fork of M considerably distal to origin of Rs but much states: (1) antennae nearly 2.5 times as long as forewing proximal to origin of Rs1. MA straight, with four pectinate, obli- [not more than 1.5 times in Epiosmylus, Gumilla]; (2) M que branches distally. MP basally straight, parallel to MA, con- in forewing forked highly proximal to origin of Rs1 figuration of distal portion somewhat irregular with 2–3 [slightly distal in Gumilla]; (3) distal half of CuA in fore- branches, most proximal of which long, dichotomously wing arched [parallel to hind margin in Epiosmylus]. branched. Fork of Cu not preserved, apparently near wing base. Configuration of distal portion of CuA rather irregular, dichotomously branched. CuP poorly preserved; distal half clearly Remarks. Nuddsia gen. nov. represents the first fossil arched; short, simple distalmost branches preserved. Anal veins record of this family from South America. Interestingly, not preserved. Crossveins rather regularly spaced, except for the single living genus of this subfamily (Gumilla) also most distal portion of radial space, where crossveins are scarce. occurs in Brazil, and the wing venation of both genera is Gradate series of crossveins not detected. quite similar. Hindwing elongate, 24.5 mm long, 5.5 mm wide, with sub- acute apex. Trichosors restricted to apical portion. Nygmata not detected. Costal space very narrow, 0.5 mm wide basally Nuddsia longiantennata sp. nov (left wing), slightly dilated in pterostigmal region. Subcostal Text-figures 1–2 veinlets simple, perpendicular to Sc in basal half of costal space, increasingly more oblique distad; veinlets of Sc+R1 closely spaced, strongly oblique. Subcostal space very narrow for Derivation of name. Latin, longus, long, and Medieval Latin most of its length, slightly dilated near fusion of Sc, R1. Sc, R1 antenna, sail yard (translation of Greek keraia, insect feeler, fused at 3.5 mm from apex. Sc+R1 entering margin slightly yard-arm), in reference to the extremely long antennae of this before apex. Rs originating near wing base. R1 space 0.8 mm species. wide medially, strongly narrowed apically; with 21 more or less regularly spaced crossveins before fusion of Sc, R1, long hypo- Holotype. SMNS 66000 ⁄ 263. An almost complete specimen stigmal cell after. Rs strongly zigzagged, forked apically, enter- exposed in ventral aspect. ing margin at apex, with 6–7 branches, mostly zigzagged, with deep marginal forks. Origin of Rs1 at approximately one-third ´ Type locality and horizon. Brazil, Ceara, Araripe Basin (Chapada wing length. Radial, medio-radial crossveins numerous, regu- do Araripe), vicinity of Nova Olinda; Crato Formation (Nova larly spaced, not forming gradate series. Basal crossveins m-r Olinda Member), Lower Cretaceous (Upper Aptian). 152 PALAEONTOLOGY, VOLUME 51

TEXT-FIG. 1. Nuddsia longiantennata gen. et sp. nov. A, photograph, and B, line drawing of the holotype, SMNS 66000 ⁄ 263. Scale bar represents 10 mm. not detected, probably absent. Origin of M, its division into branched (two branches). CuA long, zigzagged, parallel to hind MA and MP not preserved; MA straight for entire preserved margin, pectinately branched, with at least 12 rather short, length, pectinately branched distally, with ﬁve branches, most simple branches. Crossveins in medial, medio-cubital spaces proximal of which forked twice. MP zigzagged, with three dis- numerous, more or less regularly spaced. CuP, anal veins not tal branches, most proximal of which long, pectinately preserved. MENON AND MAKARKIN: CRETACEOUS FOSSIL LACEWINGS AND ANTLIONS FROM BRAZIL 153

Family PALAEOLEONTIDAE Martins-Neto, 1992 Ascalaphidae]; (2) origin of Rs situated near wing base in both wings [removed from base in Myrmeleontidae, 1992 Palaeoleontinae Martins-Neto, p. 810 (as a subfamily Araripeneuridae, Ascalaphidae, Nemopteridae, Babinskaii- of Myrmeleontidae). dae]; (3) presectorial crossveins absent in both wings 1997 Palaeoleontidae; Dobruskina et al., p. 93. [present in Myrmeleontidae, Ascalaphidae, Nemopteridae, Babinskaiidae]; (4) ‘oblique vein’ present in forewing Type genus. Palaeoleon Rice, 1969, from the Albian–Cenomanian [absent in Nymphidae]. of Labrador, Canada.

Revised description of venation. In forewing, origin of Rs close to Revised diagnosis. Large myrmeleontoids (forewing 36– wing base; without presectorial crossveins; Rs1 convex, profusely 70 mm long) with relatively long antennae, unusually branched distally; single well-developed median longitudinal fold dense crossvenation in the majority of genera, easily dis- present from wing base to apex; fork of M far removed from wing tinguished from other myrmeleontoid families by the fol- base (Palaeoleon, Baisopardus); MA strongly concave, simple; lowing combination of character states: (1) antennae basal part of MP represented by ‘oblique vein’, distal MP fused long, ﬁliform, neither dilated nor clavate distally [dilated with CuA; Cu divided into CuA, CuP close to wing base; CuA, distally in Myrmeleontidae, Araripeneuridae; clavate in MP+CuA convex, both parallel to MA; MP+CuA, MP+CuA1 and

TEXT-FIG. 2. Nuddsia longiantennata gen. et sp. nov., holotype, SMNS 66000 ⁄ 263. A, left forewing. B, right forewing. C, left hindwing. D, right hindwing. Scale bar represents 5 mm. 154 PALAEONTOLOGY, VOLUME 51 its basalmost branch form large triangular area characteristic of (appearing as a fold), which is often accompanied by a higher myrmeleontoids; CuP relatively short, pectinately branch- false longitudinal vein formed by the bending of branches ing, not fused with 1A; 1A from short, nearly simple to long, of Rs and the crossveins connecting them, called the ‘ante- pectinately branched. In hindwing, origin of Rs close to wing base, rior Banksian line’ (Tillyard 1916; Krivokhatsky 1998). without presectorial crossveins; Rs1 profusely branched distally; This fold will be called herein ‘the anterior Banksian fold’ M forked very close to wing base; MA simple; MP with long as its homology with the distal part of the median flexion pectinate branches; Cu forked close to wing base; CuA parallel to hind margin basally, arched, pectinately branched distally; CuP line of species with the anteroposteriorly asymmetric wings short, close, parallel to hind margin, pectinately branched; anal is unclear. In Parapalaeoleon magnus, the median flexion area very restricted by short simple veins. line appears as a true flexion line running (as in other Neuroptera with anteroposteriorly asymmetric wings: Included genera. Baisopardus Ponomarenko, 1992a (Lower Cre- Wootton 2002, fig. 4B) between MA and Rs, then Rs1, ter- taceous, Lower Valanginian, Baissa, Transbaikalia, Russia; Crato minating far distal to the origin of the anterior Banksian Formation; 3–4 species); Parapalaeoleon gen. nov. (Crato Forma- fold, which begins slightly distal to Rs1 (i.e. distal part of tion; monotypic); Neurastenyx Martins-Neto and Vulcano, 1997 the median flexion line and proximal part of the anterior (Crato Formation; monotypic); Paraneurastenyx Martin-Neto, Banksian fold are divergent). In other species (e.g. Palaeo- 1998 (Crato Formation; monotypic); Palaeoleon (Lower–Upper leon ferrogeneticus, Baisopardus cryptohymen) in which the Cretaceous, Albian–Cenomanian, Labrador, Canada; monotypic); origin of Rs1 is situated very near the origin of Rs, the Samsonileon Ponomarenko, in Dobruskina et al. 1997 (Upper median flexion line crosses Rs1 near its base and continues Cretaceous, Lower Turonian, Israel; monotypic); Metahemerobius Makarkin, 1990a (Upper Cretaceous–Palaeogene, Maastrichtian– further as a concave fold towards the wing apex. In these Danian, Antibes, Siberia, Russia; monotypic). species the median flexion line proximally and the anterior Banksian fold distally form a single straight line, called Remarks. The diagnosis of this family provided by here the median longitudinal fold. Basally, this line Dobruskina et al. (1997) and Heads et al. (2005) is par- appears to be vein-like in compression fossils, as in P. fer- tially erroneous, being based on misinterpretation of the rogeneticus. Such secondary vein elements are frequently venation in more poorly preserved specimens, particularly reformed from membrane (Kukalova´-Peck 1991). concerning M and CuA. The new material provides the opportunity to clarify this. According to these authors, Fusion of MP and CuA in the forewing. This fusion has ‘MP’ in the forewing is forked into ‘MP1’ and ‘MP2’ in apparently occurred independently many times in the the proximal portion of the wing, with ‘MP1’ profusely evolution of the Neuroptera. It is found in a few genera branched distally, and ‘MP2’ simple. However, this fork is of the extant Hemerobiidae (Nusalala Nava´s, 1913, Micro- not detected confidently in any species. The well-pre- mus Rambur, 1842), Dilaridae (Nallachius Nava´s, 1909), served venation of the forewing of Parapalaeoleon magnus and in a Lower Cretaceous psychopsid-like genus from (Text-fig. 3C) shows that this ‘MP1’ is actually Rs1. The the Baissa locality (Makarkin, unpublished). In two speci- homology of Rs1 in all species is unequivocal: this vein is mens of the extant Gerstaeckerella chilensis (Hagen, 1859) always convex, and is followed posteriorly by a strongly (Mantispidae) examined in this study, MP and CuA are concave simple vein (we interpret this as MA). Compari- fused abnormally for short distance at the crossvein 2m- son of Palaeoleon ferrogeneticus Rice, 1969 and Baisopar- cu (which is consequently lost) in the manner that occurs dus cryptohymen Heads et al., 2005 shows that the in Parapalaeoleon magnus. Interestingly, CuA in Gers- structure of Rs2 is also similar in these species: convex taeckerella Enderlein, 1910 (as in other mantispids) is basally and concave distally. short. In Mesochrysopidae, all intermediate stages are Examination of photographs of these specimens reveals present, from MP and CuA widely spaced to touching that what the authors interpreted as the proximal part of (Makarkin and Menon 2005; Nel et al. 2005). The fullest ‘MP1’ (Heads et al. 2005, fig. 3A) and ‘M1’ (Rice 1969, fig. fusion of these veins, however, occurs in the forewing of 2) is actually the proximal part of a longitudinal vein-like the myrmeleontoid lineage (Palaeoleontidae, Araripeneu- fold running between Rs and M. Its course is completely ridae, Myrmeleontidae, Ascalaphidae, Nemopteridae, and coincident with that of the median flexion line in some probably Babinskaiidae), in which MP resembles a cross- insects with anteroposteriorly symmetric wings, such as Si- vein (‘oblique vein’) often not distinguished from true alis Latreille, 1802 (Wootton 2002, fig. 3A), running longi- crossveins (see discussions in Tillyard 1916, 1918; Com- tudinally from the wing base to the wing apex between the stock 1918; Adams 1996). The venation of P. magnus radial and medial vein systems and distally crossing the (Text-figs 3B, 4B) rather clearly demonstrates that this branches of Rs (Wootton 2003, fig. 5). The wings of many full fusion might have resulted from touching or short extant Myrmeleontidae that have anteroposteriorly asym- fusion of a long pectinate MP and short CuA, almost metric wings appear to possess the distal part of this line exactly in the way predicted by Tillyard (1916, fig. 8b; his MENON AND MAKARKIN: CRETACEOUS FOSSIL LACEWINGS AND ANTLIONS FROM BRAZIL 155

‘Cu2’ is, however, actually CuA1). Moreover, MP and Palaeoleon araripensis was the first species of this family CuA in the forewing of Paraneurastenyx ascalaphix Mar- to be described from the Crato Formation (Martins-Neto tins-Neto, 1998 are almost touching but not fused (Mar- 1992). Later, it was transferred to Neurastenyx by Martins- tins-Neto 1998, fig. 1A), as in some Mesochrysopidae Neto (1997), and to Baisopardus by Dobruskina et al. (Makarkin and Menon 2005, fig. 3B). Based on this (1997). The holotype is an almost complete specimen, but hypothesis, it is reasonable to assume that the most prox- lacks the abdomen, and has incomplete hind- and fore- imal pectinate branch of MP+CuA (i.e. MP+CuA1) repre- wings, which are overlapped pairwise. Ponomarenko (in sents the distal portion of CuA (Text-fig. 3B); the next Dobruskina et al. 1997) noted that the hindwing venation branch, MP+CuA2 (fused with MP+CuA1 distally in this of the type specimen was partially incorrectly drawn by specimen), might have originated from the most proximal Martins-Neto (1992, p. 811, fig. 5a): the proximal branches branch of MP (MP1), and other branches of MP+CuA all of his CuA [actually MP, see above] belongs to CuP [actu- originate from MP. Some species of the extant Myr- ally CuA]; therefore, the configuration of MP and CuA are meleontidae show a remarkable reversal of the conditions similar to those of the type species (see also below). of these veins, indirectly supporting this hypothesis. In Baisopardus cryptohymen is represented by a beautifully the Australian Mjoberbia fulviguttata Esben-Petersen, 1918 preserved, complete, single specimen. The species has a and Protoplectron pallidum Banks, 1910, MP and CuA are hindwing shape and venation similar to those of the type not fused, and CuA has few branches (New 1985, figs species, B. banksianus, so we find it quite reasonable that it 263, 290, but see intraspecific variation, e.g. fig. 265). is assigned to Baisopardus. The single major difference between these species is that the former possesses closely Notes on generic and specific composition of Palaeoleonti- spaced subcostal crossveins, which the latter lacks. The dae. Palaeoleon was erected for P. ferrogeneticus, repre- interpretation of the venation in the original description sented by a single, well-preserved, apical two-thirds of a and figures of Heads et al. (2005, fig. 3) is partially incor- wing. We examined photographs of the holotype and rect (see reasoning above) and should be emended. In the found that the original description and figures of Rice forewing, their MA is Rs1 (basally) and Rs2 (distally), (1969, p. 3, figs 1, 3; pl. 1) appear to be fairly adequate, MP1 is the median flexion line (basally) and Rs1 (distally), although the venation could possibly be interpreted dif- MP2 is MA, CuA is MP+CuA, CuA1 is MP+CuA, CuA2 is ferently: Rice’s M1 is the median flexion line (basally) MP+CuA1; in the hindwing, their MA is Rs1 (basally) and and Rs2 (distally), M2 is Rs1, Cu1 is MA, Cu2 is MP (see Rs2 (distally), MP1 is the median flexion line (basally) and reasoning above). Rs1 (distally), MP2 is MA, CuA1 is MP, CuA2 plus CuP is According to Martins-Neto (1997) the genus Neuraste- CuA, 1A is CuP (see above for reasoning). nyx includes N. gigas Martins-Neto and Vulcano, 1997 All wings of ‘Neurastenyx’ polyhymnia are so strongly (type species), N. polyhymnia Martins-Neto, 1997 and N. overlapped that few details of the venation may be clearly araripensis (Martins-Neto 1992). All of these species were identified (Martins-Neto, 1997, fig. 7). Its systematic posi- placed in Baisopardus (see below), along with Baisopardus tion, therefore, may possibly be determined confidently gigas, by Heads et al. (2005) who consequently synonym- only after close re-examination of the type. The assign- yzed these two genera. However, in our opinion, Neurast- ment of this species to Baisopardus (as well as to Neurast- enyx is valid and consists of N. gigas only. The holotype enyx) should be considered tentative, based on the of the type species is poorly preserved (Martins-Neto and general impression of its venation rather than on analysis Vulcano 1997, fig. 7), but it possesses a combination of of particular character states. the character states that distinguish the species from other Samsonileon and Metahemerobius are apparently closely palaeoleontids: large size (forewing length about 70 mm) related genera; this is indicated by their similar forewing and very widely spaced crossveins. venation, in particular, MP+CuA1 and CuP run parallel Baisopardus is considered to include four species and very close for a considerable distance, and 1A is (Heads et al. 2005). The type species, B. banksianus Pon- long and pectinately branched. Unfortunately, both gen- omarenko, 1992a, is known from two isolated hindwings, era are represented by incomplete wings, lacking basal the holotype and paratype, from the Lower Cretaceous of portions in known specimens. Makarkin’s (1990a, fig. 5) Baissa, Transbaikalian Russia (Ponomarenko 1992a). The designations of the veins of Metahemerobius kalligram- genus was included by Dobruskina et al. (1997) in this mus Makarkin, 1990a are partially incorrect: his MA is family with some doubt, whereas Heads et al. (2005) Rs2, anterior branch of MP is Rs1, posterior branch of listed it among the genera of Palaeoleontidae with cer- MP is MA, CuA is MP+CuA, CuP is MP+CuA1, A1 is tainty. In the original description, however, the venation CuP. The venation of Samsonileon fragmentatus Pon- was misinterpreted by Ponomarenko (1992a, p. 49, fig. omarenko, in Dobruskina et al. 1997 is interpreted simi- 5A): his ‘MA’ is our Rs1, ‘MP’ is MA, ‘CuA’ is MP, ‘CuP’ larly here: Ponomarenko’s MA is Rs1 (basally) and the is CuA, ‘1A’ is CuP. anterior Banksian fold or Rs2 (distally), MP1 is probably 156 PALAEONTOLOGY, VOLUME 51 the median flexion line (basally) and Rs1 (distally), CuA Holotype. SMNS 66000 ⁄ 268. An incomplete specimen exposed is MP+CuA, CuP and A1 are CuP and 1A respectively. in lateral aspect, including head, thorax with appendages, and Cretoleon Ponomarenko, 1992b is a monotypic genus right forewing. represented by a forewing of C. acanthoclysoides Pono- marenko, 1992b from the Lower Cretaceous (Aptian) Type locality and horizon. As for Nuddsia longiantennata above. deposits of Bon-Tsagan, Mongolia (Ponomarenko 1992b). It was considered possibly to belong to Palaeoleontidae by Diagnosis. As for the genus. Ponomarenko (in Dobruskina et al. 1997), and listed among the genera of this family by Heads et al. (2005). It is Description. Head 4 mm long. Compound eyes 2.5 mm in diameter. Antennae 25 mm long, composed of 71 preserved segments: tentatively excluded from the family here because it differs scape not preserved; first visible segment wider than other flagel- in the following features of the forewing: the costal space is lar segments, which are square basally, transverse distad. Thorax broad; Rs originates relatively distant from the wing base; not complete (9 mm long as preserved), compressed, exoskeleton the anterior Banksian fold is absent; the subcostal veinlets mostly not preserved; prothorax apparently comparatively short. in the pterostigmal region are very closely spaced and sim- Foreleg: coxa 2 mm long; femur rather stout, 6 mm long; tibia ple; crossveins are rare in the distal portion of the wing. stout, 5 mm long, armed with apical spine; tarsus 4 mm long, five-segmented, distalmost segment approximately as long as four basal segments together; claws very long, 2 mm. Midleg: coxa Genus PARAPALAEOLEON gen. nov. 2 mm long; femur 6 mm long; tibia 4.5 mm long, apical spines not detected; tarsus as in forelegs. Hind legs not preserved. Type and only species. Parapalaeoleon magnus sp. nov. Forewing widest at mid length, 67–68 mm long (estimated), c. 20 mm wide. Costal space narrow, 1.5 mm wide in basal part, slightly narrowed toward fusion of Sc, R1, strongly expanded Derivation of name. Greek, para, near, and Palaeoleon, a generic beyond this. Subcostal veinlets simple, perpendicular to Sc in name. Gender masculine. basal half of costal space, oblique, forked distally. Veinlets of Sc+R1 long, very oblique, usually dichotomously forked, con- Diagnosis. May be distinguished from other genera of the nected by occasional crossveins not forming a regular gradate family by the combination of the following forewing char- series. Sc and R1 fused at 12 mm from wing apex, entering wing acter states: (1) origin of Rs1 far removed from origin of margin apparently well after apex (apical portion of wing not Rs [near origin of Rs in Baisopardus, Palaeoleon]; (2) Rs1 preserved). No crossveins in subcostal space. Pterostigma absent shallowly branched, highly distal to the Banksian fold or not discernible. Stem of Rs straight, not zigzagged for entire [deeply branched, at most slightly distal to the Banksian length; its origin situated near wing base, inclined at a very acute fold in Baisopardus, Palaeoleon]; (3) maximum width of angle to R1. No presectorial crossveins detected. R1 space most wing at mid-point [in distal portion in Baisopardus and dilated at middle portion, strongly narrowed basad and distad; in proximal portion in Palaeoleon]. 85 very closely spaced crossveins proximal to fusion of Sc, R1. Long hypostigmal cell absent. Rs with 13 branches, somewhat sigmoid in configuration, branched dichotomously distally. Ori- Remarks. The new genus is most similar to Baisopardus gin of Rs1 located 21 mm from origin of Rs; branches of Rs and Palaeoleon; other genera are very distant from it. dichotomously branched distally; Rs1 branched much distal to Parapalaeoleon and Baisopardus both occur in the Crato Banksian fold. Rs2, Rs3 appear fused. Median flexion line dis- Formation and are clearly distinguished from each other tinct, running between MA, Rs, then Rs1 from wing base to Rs3. (see diagnosis above). In general, all three genera differ in Anterior Banksian fold distinct, beginning slightly distal to Rs1, the shape of the wing: wide basally with a somewhat elon- directed to wing apex (somewhat posterior to it), unaccompa- gated apex in Palaeoleon, narrower basally and widest at nied by false longitudinal vein formed by bending of branches of mid-length in Parapaleoleon; maximum width after the Rs and crossveins connecting them. Radial crossveins very mid-wing in Baisopardus. The wing shape of Baisopardus numerous, not forming gradate series. Origin of M not pre- araripensis, however, is more similar to that of Parapalaeo- served, probably not fused with R at wing base. Fork of M leon magnus, than to other species of Baisopardus, but it rather distinct, with origin of MA forming shallow angle with stem of M, and MP (‘oblique vein’) stouter than crossveins in possesses other character states of Baisopardus, e.g. Rs1 median space. MA simple, with only shallow marginal fork, originates near the base of Rs and is deeply branched. straight, curved posteriorly distally. ‘Oblique vein’ well developed, located slightly proximal to most proximal branch of MP+CuA. CuA, MP+CuA strongly convex, both parallel to MA, Parapalaeoleon magnus sp. nov. dense crossveins for entire length between these and MA. Text-figures 3–4 MP+CuA with 12 branches, of which most proximal branch (MP+CuA1) pectinately branched; next branch (MP+CuA2) Derivation of name. Latin adjective, magnus, large, in reference fused with it; MP+CuA3 with deep fork; other branches shal- to the large size of the species. lowly forked. All branches of MP+CuA connected by numerous MENON AND MAKARKIN: CRETACEOUS FOSSIL LACEWINGS AND ANTLIONS FROM BRAZIL 157

TEXT-FIG. 3. Parapalaeoleon magnus gen. et sp. nov., holotype, SMNS 66000 ⁄ 268. A, photograph of the specimen. B, close-up of forewing showing fusion of MP and CuA (the names of the veins are partly hypothetical; see text for detail). C, B portion of forewing showing median ﬂexion line (mﬂ) and anterior Banksian fold (bf). Scale bars represent 10 mm.

TEXT-FIG. 4. Parapalaeoleon magnus gen. et sp. nov., holotype, SMNS 66000 ⁄ 268. A, anterior part of body. B, forewing. Scale bars represent 5 mm.

crossveins; near posterior margin crossveins connected by addi- Family MESOCHRYSOPIDAE Handlirsch, 1906 tional crossveins forming short irregular pseudo-longitudinal veins. Crossveins between CuA (also MP+CuA1), CuP numer- Genus TRIANGULOCHRYSOPA Nel et al., 2005 ous, dense. CuP concave basally, rather short, with shallow marginal fork, seven simple branches, crossveins between them rare. Type species. Triangulochrysopa sanzi Nel et al., 2005, by original 1A not fused basally with CuP. Two short unbranched anal veins designation. preserved (1A, 2A). 158 PALAEONTOLOGY, VOLUME 51

Diagnosis. Prothorax somewhat elongated. Forelegs Type locality and horizon. As for Nuddsia longiantennata above. rather short, probably raptorial. Forewing: MP touching (or almost touching) CuA; venation in radial to cubital Diagnosis. Triangulochrysopa formosa may be distin- spaces highly reticulated. Hindwing triangular in shape guished from T. sanzi by the following forewing charac- with distinct tornus, rather long (0.67 times forewing ters: forewing more elongate, length ⁄ width ratio 4.0 [3.3 length); apex of wing acute; MP appears to be continua- in the holotype of T. sanzi], pterostigma distinct [absent tion of M; MA arising from M at right angle or nearly (or not detectable) in T. sanzi], MP touching CuA [con- so. nected by short crossvein in T. sanzi], 1A with two branches [one in T. sanzi]. Included species. Triangulochrysopa sanzi (Lower Cretaceous, Bar- remian, Las Hoyas, Spain, La Huerguina Formation), T. formosa Description. Head oval, wider than long, 4.5 mm long, 2.6 mm sp. nov. (Lower Cretaceous, Upper Aptian, Araripe Basin, Brazil, wide with large compound eyes, each 1 mm in diameter. Anten- Crato Formation). nae incomplete, 13 mm long (estimated), filiform; scape not preserved, flagellum composed of at least 40 equal segments. Remarks. Triangulochrysopa is most similar to the Lower Prothorax elongate, 3.5 mm long, 2.3 mm wide; mesothorax and Cretaceous genera Allopterus Zhang, 1991 and Karenina metathorax oval, robust, 8 mm long, 4 mm wide. Legs only Martins-Neto, 1997 in the structure of the pronotum, partly preserved: fragmentary coxae on all three pairs of legs; and its foreleg and forewing venation. It may be easily right foreleg: femur rather stout, 3.5 mm long, tibiae 3.7 mm long; disarticulated tarsus (belonging probably to left midleg) distinguished from these by hindwing size, shape and with five cup-shaped segments (total length 3 mm), bearing two venation: e.g. the hindwing is short and broad, c. 0.4 of distinct claws. Abdomen composed of nine segments, 22 mm forewing length, with rounded apex and reduced venation long. in Allopterus, and relatively narrow, without tornus, 0.6– Forewing: elongate with sub-acute apex, 44 mm long, 11 mm 0.8 of forewing length, with different venation (in partic- wide (length ⁄ width ratio 4.0). Costal space narrow, slightly ular, Rs inclined at a more acute angle to R1, with Rs1 dilated towards pterostigma. Subcostal veinlets proximal to originating more distally) in Karenina (Makarkin and pterostigma simple (one forked, obviously an anomaly), 36 in Menon 2005; Nel et al. 2005). number, perpendicular to Sc or inclined to apex, becoming According to Nel et al. (2005), Triangulochrysopa (with increasingly inclined, closely spaced to pterostigmal region; ante- Allopterus, Karenina and Armandochrysopa Nel et al., rior tip of most basal subcostal (humeral) veinlet inclined 2005) belongs to the family Allopteridae in the superfam- towards base. Veinlets of Sc, Sc+R1 distal to pterostigma mostly forked, connected by 1–2 crossveins, forming 2–3 rows of cells ily Chrysopoidea. They proposed the following phyloge- between C, Sc+R1. Pterostigma dark coloured, 3.5 mm long, netic relationship: (Cratochrysa Martins-Neto, 1994 + covering seven cells. No crossveins in subcostal space detected. (Liassochrysidae + (Paralembochrysa Nel et al., 2005 + Sc, R1 fused distal to pterostigma. Sc+R1 entering wing margin ((Allopteridae + Mesochrysopidae + Tachinymphidae + at apex. Stem of Rs straight, zigzagged distally; its origin 4.5 mm Mesotermes Haase, 1890) + (Limaiidae + Chrysopi- from wing base. R1 space dilated at origin of Rs1, 1.3 mm dae))))). We have previously regarded Mesotermes and (wider than width of costal space), narrowed towards apex, with the genera of Allopteridae, Mesochrysopidae and Tachi- 28 crossveins; Rs with 20 zigzagged branches not forked before nymphidae as forming the single, monophyletic family marginal forks, each with 1–2 crossveins connected to branches Mesochrysopidae (Makarkin and Menon 2005). We are of fork. Venation in radial space highly reticulated, not forming still of the opinion that the elevation of these groups distinct gradate series. In radial space two longitudinal conver- (and Limaiidae) to family rank is unjustified. gent folds present. No crossvein between stem of Rs, M. Origin of M at 3 mm from wing base. M moderately convex, divided into MA, MP at 8 mm from wing base and at relatively acute angle (c. 45 degrees). MA smooth, slightly convex, entering mar- Triangulochrysopa formosa sp. nov. gin well before wing mid-point, with marginal fork. MP zig- Text-figures 5–6 zagged, without marginal fork, with two long zigzagged branches. Crossveins in medial space not arranged in regular ser- 2001 Unnamed neuropteran; Bechly et al. p. 55, fig. 45. ies. Basal crossvein m-cu located slightly distal to origin of M, oblique, long. Cu not fused with R basally, divided into CuA, Derivation of name. Latin adjective, formosus, graceful, beautiful, CuP distal to crossvein m-cu. CuA strongly convex, straight, with reference to the appearance of the holotype. only slightly bent at MP touching, ending in very shallow marginal fork and with single short branch (it is difficult to identify Holotype. SMNS 66000 ⁄ 271. A complete well-preserved speci- these with certainty as the venation in medial and cubical spaces men exposed in ventral aspect, with all wings wide open; disar- is highly reticulated; CuA could be also strongly zigzagged in ticulated fragments of legs are visible along the abdomen in the distal half, with two branches having each wide marginal fork). matrix. CuA touching MP. CuP short, with two short branches. Basal MENON AND MAKARKIN: CRETACEOUS FOSSIL LACEWINGS AND ANTLIONS FROM BRAZIL 159

TEXT-FIG. 5. Triangulochrysopa formosa sp. nov. A, photograph, and B line drawing of the holotype, SMNS 66000 ⁄ 271. Scale bar represents 10 mm. crossvein cua-cup oblique, long, situated slightly proximal to distinct gradate series. M dividing into MA, MP at 4 mm from meeting point of MP, CuA; distal crossvein cua-cup oblique, wing base; MP appears to be a continuation of M, MA arising rather short, situated far distal to meeting point of MP and from M nearly at right angle. MA smooth, only slightly arched, CuA. At least one distal crossvein between CuP, 1A. 1A, 2A with marginal fork entering wing margin at mid-point (15 mm nearly parallel to hind margin; 1A with one branch; 2A not from wing base). MP zigzagged, divided into two long branches. branched. 3A not detected. CuA strongly convex basally, touching MP at nearly half of its Hindwing: subtriangular in shape with somewhat pointed length, strongly zigzagged distally, with seven short branches. apex, 28 mm long, 8.5 mm maximum width. Costal space Venation between MA, CuA strongly reticulated (especially dis- equally narrow throughout length (0.5 mm). Subcostal veinlets tally, making it impossible to discriminate with certainty longi- simple, forming 31 cells proximal to pterostigma. Pterostigma tudinal veins from crossveins). CuP, anal veins not detected. short (1 mm long), covering three cells, dark coloured. Sc and R1 fused distal to pterostigma. No crossveins in subcostal space Remarks. Both species of the genus, T. sanzi and T. for- detected. Origin of Rs at 5.7 mm from wing base. R1 space mosa, are very similar to each other. T. sanzi was 1 mm at its widest point, with 17 crossveins; hypostigmal cells described from ﬁve specimens from the Barremian of Las short. Stem of Rs zigzagged, with 12 zigzagged branches, simple Hoyas, Spain, which differ somewhat in size (forewing (distal branches) or with shallow, wide marginal fork (proximal length ranges from 27.5 to 39.2 mm) and venation. Nel branches). Crossveins in radial space numerous, not forming 160 PALAEONTOLOGY, VOLUME 51

TEXT-FIG. 6. Triangulochrysopa formosa sp. nov., holotype, SMNS 66000 ⁄ 271. A, left forewing. B, left hindwing. Scale bar represents 5 mm.

et al. (2005) considered these differences insufficient to REFERENCES separate them; however, some may be significant at a specific level. For example, the hindwing venation of the ADAMS, P. A. 1969. A new genus and species of Osmylidae specimens 92 ⁄ 2 ⁄ 3 (Nel et al. 2005, fig. 7.5) and LH-18572 (Neuroptera) from Chile and Argentina, with a discussion of (Nel et al. 2005, fig. 7.4) of T. sanzi is very different; they planipennian genitalic homologies. Postilla, 141, 1–11. are 19.2 mm and 21.6 mm long respectively, whereas the —— 1977. Taxonomy of United States Leucochrysa (Neuroptera: hindwing of the holotype is 31.6 mm long (unfortunately, Chrysopidae). Psyche, 84, 92–102. this wing was not figured). The hindwing ⁄ forewing length —— 1996. Venational homologies and nomenclature in Chrysopi- dae, with comments on the Myrmeleontoidea (Insecta: Neuro- ratio ranges from 0.55 to 0.66 in these paratypes, and is ptera). 19–30. In CANARD, M., ASPO¨ CK, H. and 0.95 in the holotype. The photograph of the holotype MANSELL, M. W. (eds). Pure and applied research in neur- does not reveal such long hindwings; the measurement opterology. Proceedings of the Fifth International Symposium on should probably be 21.6, not 31.6 mm. In any case, after Neuropterology. Privately printed, Toulouse (France), 414 pp. this discovery of another very similar species from a ARCHIBALD, S. B. and MAKARKIN, V. N. 2006. Tertiary South American locality distant from southern Europe, all giant lacewings (Neuroptera: Polystoechotidae): revision and material previously identified as T. sanzi should be description of new taxa from western North America and revised. Denmark. Journal of Systematic Palaeontology, 4, 119–155. BANKS, N. 1910. Myrmeleonidae from Australia. Annals of the Acknowledgements. We thank Dr Gu¨nter Bechly and Dr Gerd Entomological Society of America, 3, 40–44. Dietl (SMNS) for allowing us to study the material we have BECHLY, G., HAAS, F., SCHAWALLER, W., SCHMAL- described; Dr Andre´ Nel (Museum national d’Histoire naturelle, FUSS, H. and SCHMID, U. 2001. Ur-Geziefer – Die faszini- Paris) and Prof. Dr Rafael Martins-Neto (Universidade Federal erende Evolution der Insekten. Stuttgarter Beitra¨ge zur de Juiz de Fora, Brazil) for providing us with advance copies of Nuturkunde, Serie C, 49, 1–96. their papers in press; Prof. Dr Ulrike Aspo¨ck and Dr Harald BERTHOU, P. Y. 1994. Relationship between the ostracod Schillhammer (Naturhistorisches Museum Wien) for providing fauna and the oxic or anoxic character of the Aptian–Albian the photograph of the holotype of Gumilla adspersa; Jean strata of the Araripe basin (NE Brazil). Proceedings of the 14th Dougherty (Geological Survey of Canada, Ottawa, Canada), and Sedimentological Congress, 14, G11. Bruce Archibald (Museum of Comparative Zoology, Harvard COMSTOCK, J. H. 1918. The wings of insects. Comstock University) for providing the photograph of the holotype of Pal- Publication Company, Ithaca, NY, 430 pp. aeoleon ferrogeneticus; Mr Robert Loveridge (University of Ports- DOBRUSKINA, I. A., PONOMARENKO, A. G. and mouth) for providing the photograph of the holotype of RASNITSYN, A. P. 1997. Fossil insects found in Israel. Baisopardus chryptohymen; Bruce Archibald for helpful com- Paleontologicheskii Zhurnal, 1997 (5), 91–95. [In Russian]. ments and for suggestions on the English; James Jepson (Univer- DOLIN, V. G., PANFILOV, D. V., PONOMARENKO, sity of Manchester) for correcting our English; Dr Gu¨nter Bechly A. G. and PRITYKINA, L. N. 1980. Fossil insects of the and Dr Rafael Giois Martins-Neto for reviewing the manuscript; Mesozoic. Naukova Dumka, Kiev, 135 pp. [In Russian]. and Prof. David Batten (University of Manchester) for editorial ENDERLEIN, G. 1910. Klassifikation der Mantispiden nach work. The visit to Stuttgart by FM was funded by the Deutscher dem Material des Stettiner Zoologischen Museums. Stettiner Akademischer Austausch Dienst (DAAD). Entomologische Zeitung, 71, 341–379. MENON AND MAKARKIN: CRETACEOUS FOSSIL LACEWINGS AND ANTLIONS FROM BRAZIL 161

ESBEN-PETERSEN, P. 1918. Results of Dr. E. Mjo¨berg’s (Lower Cretaceous), with taxonomic treatment of the family. Swedish scientific expeditions to Australia 1910–1913. 18. Neu- Cretaceous Research, 26, 801–812. roptera and Mecoptera. Arkiv fo¨r Zoologi, 11(26), 1–37. —— —— 2007. First record of fossil rapismatid-like Ithonidae HAASE, E. 1890. Bemerkungen zur Palaeontologie der Insec- (Insecta, Neuroptera) from the Lower Cretaceous Crato ten. Neues Jahrbuch fu¨r Mineralogie, Geologie und Palaöntolo- Formation of Brazil. Cretaceous Research, 28, 743–753. gie, 2 (1), 1–33. MARTILL, D. M. 1993. Fossils of the Santana and Crato For- HANDLIRSCH, A. 1906–08. Die fossilen Insekten und die Phy- mations, Brazil. Field Guide to Fossils No. 5. The Palaeonto- logenie der rezenten Formen. Ein Handbuch fu¨r Palaöntologen logical Association, London, 159 pp. und Zoologen. Engelmann, Leipzig, ix + 1430 pp., 51 pls. —— and FREY, E. 1995. Colour patterning preserved in Lower HAGEN, H. A. 1859. Beitrag zur Kenntniss der Neuropteren. Cretaceous birds and insects: the Crato Formation of N. E. Stettiner Entomologische Zeitung, 20, 405–412. Brazil. Neues Jarhbuch fu¨r Geologie und Palaöntologie, Monats- HEADS, S. W., MARTILL, D. M. and LOVERIDGE, R. F. hefte, 1995, 118–128. 2005. An exceptionally preserved antlion (Insecta, Neuroptera) MARTINS-NETO, R. G. 1992. Neuro´pteros (Insecta, Plani- with colour pattern preservation from the Cretaceous of Bra- pennia) da Formaçaõ Santana (Cretaćeo Inferior), Bacia do zil. Palaeontology, 48, 1409–1417. Araripe, nordeste do Brasil. VII. Palaeoleontinae, nova sub- KRIVOKHATSKY, V. A. 1998. Antlions (Neuroptera, Myr- famı´lia de Myrmeleontidae e descriçaõ de novos ta´xons. Revi- meleontidae) of the Palaearctic Region (morphology, classifi- sta Brasileira de Entomologia, 36, 803–815. cation, zoogeography). Unpublished PhD thesis, Zoological —— 1994. Neuro´pteros (Insecta, Planipennia) da Formaçaõ Institute, Russian Academy of Sciences, St. Petersburg, 364 pp. Santana (Cretaćeo Inferior), Bacia do Araripe, nordeste do [In Russian]. Brasil – IX – Primeiros resultados da composiçaõ da fauna e KRU¨ GER, L. 1913. Osmylidae. Beitra¨ge zu einer Monographie descriçaõ de novos ta´xons. Acta Geologica Leopoldensia, 17 der Neuropteren-Familie der Osmyliden. III. Literatur und (39 ⁄ 1), 269–288. Katalog. Stettiner Entomologische Zeitung, 74, 193–224. —— 1997. Neuro´pteros (Insecta, Planipennia) da Formaçaõ San- —— 1915. Osmylidae. Beitra¨ge zu einer Monographie der Neur- tana (Cretaćeo Inferior), Bacia do Araripe, nordeste do Brasil. opteren-Familie der Osmyliden. VIII. Anhang II. Stettiner En- X. Descriçaõ de novos ta´xons (Chrysopidae, Babinskaiidae, tomologische Zeitung, 76, 60–87. Myrmeleontidae, Ascalaphidae e Psychopsidae). Revista Uni- KUKALOVA´ -PECK, J. 1991. Fossil history and the evolution versidade de Guarulhos, Se´rie Cieˆncias Exactas e Technolo´gicas, of hexapod structure. 144–179. In NAUMANN, D. (chief 2 (4), 68–83. ed.). The insects of Australia, Volume 1. Second edition. Mel- —— 1998. Neuro´pteros (Insecta, Planipennia) da Formaçaõ bourne University Press, Carlton, 752 pp. Santana (Cretaćeo Inferior), Bacia do Araripe, nordeste do LAMBKIN, K. J. 1988. A re-examination of Lithosmylidia Riek Brasil. XI. Descriçaõ de novos ta´xons de Myrmeleontidae from the Triassic of Queensland with notes on Mesozoic (Paleoleontinae e Pseudonymphinae). Revista Universidade de ‘osmylid-like’ fossil Neuroptera (Insecta: Neuroptera). Mem- Guarulhos, Se´rie Cieˆncias Biolo´gicas e da Sau´de, 3 (5), 38– oirs of the Queensland Museum, 25, 445–458. 42. LATREILLE, P. A. 1802. Histoire naturelle, generale et particu- —— 2000. Remarks on the neuropterofauna (Insecta, Neuro- liere de Crustaces et des Insectes. Volume 3. Dufart, Paris, 467 pp. ptera) from the Brazilian Cretaceous, with keys for the LEACH, W. E. 1815. Entomology. 57–172. In BREWSTER, identification of the known taxa. Acta Geolo´gica Hispanica, 35, D. (ed.). Edinburgh Encylopaedia 9 (1). Edinburgh. 97–118. LINNAEUS, C. 1758. Systema natura per regna tria naturae —— 2003. The Santana Formation paleoentomofauna reviewed. secundum classes, ordines, genera, species, cum characteribus, Part I – Neuropteroida (Neuroptera and Raphidioptera): sys- differentiis, synonymis, locis. Tenth edition. Laurentius Salvius, tematics and phylogeny, with description of new taxa. Acta Holmiae, 824 pp. Geologica Leopoldensia, 25 (55) (for 2002), 35–66. MAISEY, J. C. (ed.) 1991. Santana fossils: an illustrated atlas. —— 2005. New Neuroptera (Nymphidae and Araripeneuridae) Tropical Fish Hobbist Publications, Neptune City, NJ, 459 pp. from Santana Formation (Lower Cretaceous, Araripe basin, MAKARKIN, V. N. 1990a. New lacewings (Neuroptera) from northeast Brasil). GAEA, 1, 5–10. the Upper Cretaceous of Asia. 63–68. In AKIMOV, I. A. —— and VULCANO, M. A. 1997. Neuro´ pteros (Insecta, (ed.). News of faunistics and systematics. Naukova Dumka, Planipennia) da Formaçaõ Santana (Cretaćeo Inferior), Bacia Kiev, 184 pp. [In Russian]. do Araripe, Nordeste do Brasil. VIII. Descriçaõ de novas taxas —— 1990b. A new fossil genus and species of Osmylidae from de Myrmeleontidae, Ascalaphidae e Nemopteridae. Revista the Lower Cretaceous of East Siberia (Neuroptera). Deutsche Universidade de Guarulhos, Se´rie Cieˆncias Biolo´gicas e da Sau´de, Entomologische Zeitschrift, 37, 101–103. 2 (5), 64–81. —— and ARCHIBALD, S. B. 2003. Family affinity of the MENON, F., MARTINS-NETO, R. G. and MARTILL, genus Palaeopsychops Andersen with description of a new spe- D. M. 2005. A new Lower Cretaceous nymphid (Insecta, Neu- cies from Early Eocene of British Columbia (Neuroptera: roptera, Nymphidae) from the Crato Formation of Brazil. Polystoechotidae). Annals of the Entomological Society of Amer- GAEA, 1, 11–15. ica, 96, 171–180. NAVA´ S, L. 1909. Monografıá de la familia de los Dila´ridos —— and MENON, F. 2005. New species of the Mesochrysopi- (Ins. Neur.). Memorias de la Real Academia de Ciencias y Artes dae (Insecta, Neuroptera) from the Crato Formation of Brazil de Barcelona, 3, 7, 619–671, 2 pls. 162 PALAEONTOLOGY, VOLUME 51

——1912. Insectos neuro´pteros nuevos o poco conocidos. Mem- In CAMPOS, D. A., VIANNA, M. S. S., BRITO, P. M. orias de la Real Academia de Ciencias y Artes de Barcelona, 3, and BEURLEN, G. (eds). Atas do I Simposia sobre a Bacia 10, 135–202. do Araripe e Bacias Interiores do Nordeste, Crato, 14–16 Junho —— 1913. Ne´vropte`res. 69–77, 1 pl. In: Mission du Service Geó- de 1990. Departmento Nacional de Produçaõ Mineral 10 graphique de l’Armeé pour la mesure l’un arc de me´ridien e´qua- Distritio (DNPM), Universidade Regional do Cariri torial en Ame´rique du Sud sous le controˆle scientifique de (URCA) e Sociedade Brasileira de Paleontologia (SBP), Crato, l’Acade´mie des Sciences 1899–1906. Tome 10, Fascicle 1. Paris. 405 pp. NEL, A., DELCLOS, X. and HUTIN, A. 2005. Mesozoic RAMBUR, J. P. 1842. Histoire naturelle des insects. Neùropte`res. chrysopid-like Planipennia: a phylogenetic approach (Insecta: Fain et Thunot, Paris, xviii + 534 pp., 12 pls. Neuroptera). Annales de la Socie´te´ Entomologique de France, REN, D. and YIN, J. 2002. A new Middle Jurassic species of 41, 29–69. Epiosmylus from Inner Mongolia, China (Neuroptera: Osmyli- NEW, T. R. 1985. A revision of the Australian Myrmeleontidae dae). Acta Zootaxonomica Sinica, 27, 274–277. [In Chinese, (Insecta: Neuroptera). I. Introduction, Myrmeleontini, Proto- English summary]. plectrini. Australian Journal of Zoology, Supplementary Series, RICE, H. M. A. 1969. An antlion (Neuroptera) and a stonefly 104, 1–90. (Plecoptera) of Cretaceous age from Labrador, Newfoundland. OSWALD, J. D. 1993. Revision and cladistic analysis of the Geological Survey of Canada, Department of Energy, Mines and world genera of the family Hemerobiidae (Insecta: Neuroptera). Resources, Paper, 68-65, iv + 11 pp., 2 pls. Journal of the New York Entomological Society, 101, 143–299. SELDEN, P. A. 2003. A new tool for fossil preparation. The PANFILOV, D. V. 1980. New representatives of lacewings Geological Curator, 7, 337–339. (Neuroptera) from the Jurassic of Karatau. 63–68, 7 pls. In TILLYARD, R. J. 1916. Studies in Australian Neuroptera. No. DOLIN, V. G., PANFILOV, D. V., PONOMARENKO, I. The wing-venation of the Myrmeleontidae. Proceedings of A. G. and PRITYKINA, L. N. Fossil insects of the Mesozoic. the Linnean Society of New South Wales, 40 (for 1915), 734– Naukova Dumka, Kiev, 135 pp. [In Russian]. 750. PONOMARENKO, A. G. 1980. Superorder Myrmeleontidea —— 1918. Studies in Australian Neuroptera. No. 5. The Latreille, 1802. 84–99, pl. 3, fig. 11. In ROHDENDORF, structure of the cubitus in the wings of the Myrmeleontidae. B. B. and RASNITSYN, A. P. (eds). The historical develop- Proceedings of the Linnean Society of New South Wales, 43, ment of the class Insecta. Trudy Paleontologicheskogo Instituta, 116–122. 175, 269 pp. [In Russian]. WALKER, F. 1853. List of the specimens of neuropterous insects —— 1992a. Neuroptera (Insecta) from the Lower Cretaceous of in the collection of the British Museum. Part II. British Transbaikalia. Paleontologicheskii Zhurnal, 1992 (3), 43–50, Museum, London, pp. 193–476. Moscow. [In Russian: English translation, Paleontological WEDMANN, S. and MAKARKIN, V. N. 2007. A new genus Journal, 1992, 26 (3), 56–66]. of Mantispidae (Insecta: Neuroptera) from the Eocene of —— 1992b. New lacewings (Insecta, Neuroptera) from the Germany, with a review of the fossil record and palaeobio- Mesozoic of Mongolia. 101–111, pls 27–29. in GRUNT, geography of the family. Zoological Journal of the Linnean Soci- T. A. (ed.). New taxa of fossil invertebrates of Mongolia. Trans- ety, 149, 701–716. actions of the Joint Soviet-Mongolian Paleontological Expedi- WOOTTON, R. J. 2002. Design, function and evolution in the tion, 41. Nauka Press, Moscow, 142 pp. [In Russian]. wings of holometabolous insects. Zoologica Scripta, 31, 31–40. —— 2003. On some Neuroptera (Insecta) from Upper Jurassic —— 2003. Wings. 1186–1192. In RESH, V. H. and CARDE, Solnhofen Limestone. Annals of the Upper Silesian Museum R. T. (eds). Encyclopedia of insects. Academic Press, London, (Entomology), 12, 87–100. 1266 pp. PONS, D., BERTHOU, P. Y. and CAMPOS, D. A. 1991. ZHANG, J. 1991. A new family of Neuroptera (Insecta) from Quelques observations sur la palynologie de l’Aptien Supe´rieur the late Mesozoic of Shandong, China. Science in China, Series et de l’Albien du Bassin d’Araripe (N. E. du Bre´sil). 241–252. B, 34, 1105–1112.