(Neuroptera and Raphidioptera): Systematic and Phylogeny, with Description of New Taxa

Acta Geologica Leopoldensia XXV(55): 35-66, 2002

THE SANTANA FORMATION PALEOENTOMOFAUNA REVIEWED. PART I - NEUROPTEROIDA (NEUROPTERA AND RAPHIDIOPTERA): SYSTEMATIC AND PHYLOGENY, WITH DESCRIPTION OF NEW TAXA.

Rafael Gioia Martins-Neto Universidade Guarulhos I Sociedade Brasileira de Paleoartropodologia - SBPr: Rua Arnaldo Vitaliano, 150, apto 81; 14091-220 - Ribeidio Preto - SP, Brasil. E-mail: [email protected]

RESUMO: Vma ampla revisao da paleoentomofauna da Forma~ao Santana e efetuada, abordando nesta primeira parte os Neuropteroida (Neuroptera e Raphidioptera) e os seguintes novos taxa sao propostos: Cratopteryx nemopteroides n. sp., Araripeneura urda n. sp., Cratoalloneura verdandia n. sp., (Neuroptera Myrmeleontoidea Araripeneuridae n. status), Caririchrysa skulda n. sp. (Chrysopidae), Caririraphidia sertaneja n. gen. et n. sp., e Caririraphidia? reticulata n. sp. (Raphidioptera Raphidioidae). 0 genero Caririchrysa Martins-Neto & Vulcano (Chrysopidae) e reestabelecido. Adicionalmente, novos materiais recentemente coletados sao aqui figurados, todos provenientes do nivel de calcario laminado, Membro Crato, unidade inferior da Forma~ao Santana, Bacia do Araripe (Cretaceo Inferior do nordeste do Brasil). 0 genero Lugalla Willmann 1994, como discutido aqui, e considerado sin6nimo de Cratoraphidia Martins-Neto & Nel 1992, representado tanto no Cretaceo Inferior do Brasil quanta no da Asia. Atraves de uma analise filogenetica efetuada, sao discutidas as rela~oes entre os Myrmeleontoidea e a subafamilia Araripeneurinae Martins-Neto e elevada a categoria de familia e a nova tribo Cratoalloneurini n. trib. e proposta.

Palavras-chave: Insetos f6sseis, Neuroptera, Raphidioptera, Forma~ao Santana, Cretaceo Inferior, Brasil.

ABSTRACT: A wide systematic revision is made on the Santana Formation paleoentomofauna, focusing in this first part Neuropteroida (Neuroptera and Raphidioptera). The following new taxa are proposed: Cratopteryx nemopteroides n. sp., Araripeneura urda n. sp., Cratoalloneura verdandia n. sp., (Neuroptera Myrmeleontoidea Araripeneuridae n. status), Caririchrysa skulda n. sp. (Chrysopidae), Caririraphidia sertaneja n. gen. et n. sp., and Caririraphidia? reticulata n. sp. (Raphidioptera Raphidioidae). The genus Caririchrysa Martins-Neto & Vulcano (Chrysopidae) is reestablished. Additionally, newly collected specimens are figured for the first time, all coming from the laminated limestone level, Crato Member, lowermost unit of the Santana Formation, Araripe Basin, Lower Cretaceous, Northeast Brazil. The genus Lugalla Willmann 1994, as discussed here, is considered as a junior synonymy of Cratoraphidia Martins-Neto & Nel 1992, present in the Brazilian Lower Cretaceous as well as in the Asiatic one. After a phylogenetical approach, the Myrmeleontoidea relationship are discussed and the subfamily Araripeneurinae Martins-Neto is rised to family status and a new tribus, Cratoalloneurini n. trib. is proposed. Key words: Fossil insects, Neuroptera, Raphidioptera, Santana Formation, Lower Cretaceous, Araripe Basin, Brazil. 36

Introduction (15%), Amphiesmenoptera (Trichoptera + Lepidoptera), with 12 species (5%), This research presents a taxonomic Blattopteromorha (Blattoptera + Isoptera + study of forty-five new fossil insects taxa, Dermaptera), with 11 speCIes (5%), distributed in the orders and suborders Hymenoptera, with 10 speCIes (4%), Ephemeroptera, Odonatoptera, Ensifera, Antliophora (Diptera + Mecoptera), with 6 Caelifera, Isoptera, Dermaptera, species (3%) and Coleoptera, with 4 species Auchenorrhyncha, Heteroptera, Coleoptera, (2%). About 239 described species for the Hymenoptera, Neuroptera, Raphidioptera, Santana Formation, 167 were described by Trichoptera, Lepidoptera, and Diptera, which the author and collaborators, 6 for other came from laminated limestone of the Brazilian authors and 66 described by foreign uppermost part of the Crato Member, lowest researchers. Those data demonstrates that unit of the Santana Formation (Lower 28% of the described species are not housed Cretaceous), Araripe Basin, near Santana do in Brazil, besides the absolute lack of Cariri and Nova Olinda municipalities (Ceara stratigraphic control, but fortunately, 72% of State, Northeast Brazil), as well as new the described species are housed at the approaches on insect paleobiogeography, country and with full stratigraphic controls, chronostratigraphy, paleoecology, what turns reliable all data which can be paleobiomechanic, taphonomy and extracted of them. phylogeny. A wide systematic revision is The Santana Formation neuropterofauna made on the Santana Formation is composed by fifty-five species distributed paleoentomofauna, beginning in this first part in thirty-three genera. The Neuroptera are with the Neuropteroida (Neuroptera and represented by the families Ascalaphidae, Raphidioptera), being described four new Myrmeleontidae, Psychopsidae, Chrysopidae, species of N europtera and one new genus, Berothidae, Sisyridae, Nemopteridae, two new species, and a new combination of Babinskaiidae, Paleoleontidae, Roeslerianidae Raphidioptera. and Makarkiniidae. Except for the four last, The material newly collected comes all the others possess extant representation. from the Lower Cretaceous of Brazil. Type Raphidioptera is represented by four genera locality: Mina Pedra Branca, Nova Olinda and six species (including a new genus and Santana do Cariri road, 4 km of the municipal two new species described here) and district of Nova Olinda. Type stratum: Megaloptera by a genus and a single species. laminated limestone level, Crato Member, The Brazilian Cretaceous neuropterans lowermost unit of the Santana Formation, exhibit several particularities. Among them Araripe Basin. Upper Aptian, Lower the fact that, until now, just one Cretaceous. Raphidioptera genus is known from another Until this moment, the distribution of place, exhibiting, therefore, a high endemism the Santana Formation paleoentomofauna degree. Babinskaiidae, Paleoleontidae, reveals that the groups better represented are Roeslerianidae and Makarkiniidae are Orthopteroida (Ens ifera + Caelifera + families entirely extinct. During the Phasmatoptera), with 61 described species Cretaceous however, paleoleontinids are (25%), Neuropteroida (Neuroptera + represented in Asia, Canada, Spain and Raphidioptera + Megaloptera), with 55 England. Nemopteridae are absent in the described species (23 %), Hemipteroida current Brazilian fauna, in spite of their (Auchenorrhyncha + Coleorrhyncha + abundance and diversity during the Heteroptera), with 43 described species Cretaceous, with three genera and five (18%), Paleoptera (Odonatoptera + described speCIes. The myrmeleontids Ephemeroptera), with 37 described species dominate the Santana Formation 37 neuropterofauna, with ten genera (35,7%) and Cratoalloneura, just known by the holotype. twenty-five species (48%) and also dominate Among the most collected specimens, are the in number of specimens (53%). In the Santana genera Caririneura (eighteen), Cratoneura Formation Myrmeleontidae Blitterdorffia has (ten), Pseudonymphes (eight), Araripeneura the greatest number of known species (five), (seven), Blittersdorffia (seven), and followed by the genera Caririneura (four), Cratoalloneura (six) (see Table I). Cratoneura (four), Pseudonymphes (four), The geological aspects of the Santana and Araripeneura (three). The genera Formation where extensively focused in Caldasia, Cratoalloneura, Paracaririneura, previous papers (Martins-Neto 1996, 2002). Cratopteyx and Bleyeria are, until this The adopted terminology follows Martins moment, monospecific and, except by Neto (2000).

Table I. The Santana Formation neuropterofauna.

ORDERlSfamily FAMILY SPECIES N

Neuroptera Chrysopidae Limaiinae Limaia conspicua Martins-Neto & Vulcano 1989a 5 Limaia adicotomica Martins-Neto 1997 2 Araripechrysa magnifica M-N & Vulcano 1989a 2 Chrysopidae Mesochrysopinae Caririchrysa criptovenata Martins-Neto & V. 1989a 4 Caririchrysa confusa Martins-Neto & Vulcano 1989a 1* Caririchrysa skulda n. sp. 3 Chrysopidae Cratochrysinae Cratocrhysa willmanni Martins-Neto 1994 2 Cratochrysa sublapsa Martins-Neto & Vulcano 1997 1* Berothidae Araripeberotha martinsi M-N & Vulcano 1990a 1* Caririberothafairchildi M-N & Vulcano 1990a 1* Sisyridae Cratosysirops gonzagai Martins-Neto 1997 1* Psychopsidae Pulchroptilonia spatiJata Martins-Neto 1997 2 Myrmeleontoidea Araripeneuridae Araripeneura regia Martins-Neto & Vulcano 1989b 3+1 Araripeneura gracilis Martins-Neto & Vulcano 1989b 2 Araripeneura urda n. sp. 1* Caldasia cretacea Martins-Neto & Vulcano 1989b 1* Caririneura microcephala M-N & Vulcano 1989b 1* Caririneura damianii Martins-Neto 1992a 6+ 1 Caririneura crassatella Martins-Neto & Vulcano 1997 2+1 Cratoalloneura acuminata Martins-Neto 1992b 4+2 Cratoalloneura verdandia n. sp. 2 Cratoneura longissima Martins-Neto 1992b 5 Cratoneura pulchella Martins-Neto 1992b 2 Cratoneura dividens Martins-Neto 1994 1+3 Paracaririneura priscila M-N & Vulcano 1997 1* Cratopteryx robertosantosi M-N & Vulcano 1989b 1* Cratopteryx nemopteroides n. sp. 1* Myrmeleont. Pseudonymphinae Blittersdorffia pleoneura M-N & Vulcano 1989b 2 Blittersdorffia volkheimeri M-N & Vulcano 1989c 1* Blittersdorffia dicotomica Martins-Neto 1990b 1* Blittersdorffia polyp/usia Martins-Neto 1997 1+ 1 Blittersdorffia pulcherrima M-N & Vulcano 1997 1* Bleyeria nordestina Martins-Neto 1992a 1* Pseudonymphes araripensis M-N & Vulcano 1989b 4 Pseudonymphes ponomarenkoi Martins-Neto 1992a 2 38

Pseudonymphes brunherottae Martins-Neto 1994 1* Pseudonymphes zambonii Martins-Neto 1998 1* Palaeoleontidae Neurastenyx araripensis Martins-Neto 1992b 1* Neurastenyx gigas Martins-Neto & Vulcano 1997 1* Neurastenyx polyhymnia Martins-Neto 1997 1* Paraneurastenyx ascalaphix Martins-Neto 1998 1* Nymphoidea Babinskaiidae Babinskaia pulchra Martins-Neto & Vulcano 1989b 1* Babinskaiaformosa Martins-Neto & Vulcano 1989b 2 Neliana maculata Martins-Neto 1994 3 Neliana impolluta Martins-Neto 1997 1* Ascalaphoidea Ascalaphidae Karenina breviptera Martins-Neto 1997 1* Cratoscalapha electroneura M-N & Vulcano 1997 1* Nemopteroidea Roeslerianidae Roesleriana exotica Martins-Neto & Vulcano 1989b 4 Nemopt. Cratonemopteriginae Cratonemopteryx audax Martins-Neto & V. 1989b 2 Cratonemopteryx robusta M-N & Vulcano 1989b 2 Cratonemopteryx speciosa M-N & Vulcano 1997 1+1 Nemopteridae Krikaiinae Krila pilosa Martins-Neto 1992b 2 Osmyloidea Makarkinidae Makarkinia adamsi Martins-Neto 1992a 1* Raphidioptera Raphidioidea Austroraphidia brasiliensis (Nel, S. & M-N) Will 94 1+2 Arariperaphidia rochai M-N & Vulcano 1990b 2+1 Arariperaphidia sp. 1 Cratoraphidia pulchra Martins-Neto & Nel 1992 1* Caririraphidia sertaneja n. sp 1* Caririraphidia? reticulata n. sp. 1* Megaloptera Gen. et sp. n. Martins-Neto 1999 1* * -just de holotype is known + 1 - supplementary material presented here N - number of known collected specimens

Sistematic Paleontology Nova Olinda municipality, Ceara State, Northeast Brazil. Neuroptera Type stratum: Laminated limestone level, Superfamily Myrmeleontoidea Burmeister Crato Member, lowermost unity of Santana 1839 Formation, Araripe Basin. Family Araripeneuridae n. status Age: Upper Aptian, Lower Cretaceous. Genus Cratopteryx Martins-Neto and Vulcano 1989 Diagnosis: Fore wing circa 15 mm long. Type species Cratopteryx robertosantosi Hipostigmal cell (b) very elongated and Martins-Neto and Vulcano 1989, by original narrow, occupying quite the whole extension designation. of the apical margin. Intraradial cell (cir) small, of the same size as for the subsequent Cratopteryx nemopteroides n. sp. cell. Eight premedial cross veins. Hind wing a (PI. lA; Fig. lA, B) little longer than the fore wing elongated and notably narrow. Etymology: Allusive to the nemopterid aspect of the hind wing. Discussion: C. nemopteroides n. sp. is similar Holotype: MPFT-I-004, Museu de to C. robertosantosi Martins-Neto and Paleontologia Fon;a da Terra, Sao Paulo - SP Vulcano described for the same deposits in Type locality: Pedra Branca Mine, Nova the general habitus, differing however by Olinda-Santana do Cariri road, at 4 K.m from having the MA origin far to the radial basal 39

CuP

Figure 1. A-B) Cratopteryx nemopteroides n. sp., drawn from holotype MPFT-I-004, fore wing (A) and hind wing (B), respectively; C-E) Araripeneura regia Martins-Neto and VuIcano 1989, drawn from supplementary material (MPFT-I-009), fore wing (C), hind wing (D) and body details (E), respectively. Terminology: MA, MP, anterior and posterior Media, respectively; CuA, CuP, anterior and posterior cubitus, respectively; h, hipostigmal cell; phc, pre-hipostigmal cell; ir, intraradial cell; 0, oblique vein. Scale bar 1 mm. cell and eight premedial cells (closer to radial dichotomous at the apical margin. Presence of basal cell and just four premedial cross veins a well-defined pterostigma in the ScP and RA in C. robertosantosi). fusion area. RP origin very close to wing base. Thirteen radial cells, being the basal one Description: (Holotype MPFT-I-004, Figure triangular. Hipostigmal cell notably long and lA, B). Fore wing 14 mm long (Figure lA), narrow of which basal margin is anterior to as preserved, and relatively wide, with ScP and RA fusion and the distal one close to acuminate apex. Costal area relatively wide, boundary of apical/anal margin. Seven narrower at the wing base, widening toward branches of RP, all without marginal the apex, that is twice wider than wing base, dichotomies and four sectorial cells. MA filled by pectinated cross veins, longer and origin far from the RP origin. RP with seven 40 distal secondary branches. Intraradial cell Paulo- SP (cir) relatively small, with similar size of the Type locality, type stratum and age: As for subsequent cell. Eight cross-veins before MA Cratopteryx nemopteroides n. sp. origin. MPI long, basally straight, distally with a single long fork. MP2+CuA parallel Araripeneura urda n. sp. and very close to MPI. Oblique vein (0) very (PI. IC, D; Fig. 2A-C) close to wing base. Six CuA secondary branches, all with distal dichotomies. Just a Etymology: Allusive to Urd, "goddess of the long cross-vein before of the most proximal past" in the Viking mitology. branch of CuA (CuA2). CuP reach obliquely Holotype: MPFT-I-OlO, Museu de the anal margin, straight, a little after the 113 Paleontologia For9a da Terra, Sao Paulo - SP of wing base with, at least, seven long Type locality, type stratum and age: As for secondary branches. Hind wing (Figure IB) Cratopteryx nemopteroides n. sp. long and narrow, a little longer than fore wing. Costal margin relatively wide. Diagnosis: Fore wing circa 13 mm long. Hipostigmal cell present. CuA2 origin anterior to RP origin, close to the wing base. Caririneura damianii Martins-Neto 1992 (PI. lIA) Discussion: The smallest species of the genus, with CuA2 origin anterior to RP Supplementary material: MPFT- 1-007, origin, very close to the wing base (at same Museu de Paleontologia For9a da Terra, Sao level in two other known species). Paulo - SP Additionally, Araripeneura urda n. sp. differs Type locality, type stratum and age: As for by having a greater number of radial cells as Cratopteryx nemopteroides n. sp. well as a greater number of secondary branches ofRP, MA, MPl, MP2+CuAl, and Caririneura crassatella Martins-Neto and CuP. Vu1cano 1997 (PI. lIB) Description: (Holotype MPFT-I-OIO, Figure 2A-C). Fore wing 13.3 mm long (Figure 2A) Supplementary material: MPFT -1-008, and relatively wide, with acuminate apex. Museu de Paleontologia For9a da Terra, Sao Costal area relatively narrow, little wider at Paulo - SP wing apex, filled by pectinated cross veins, Type locality, type stratum and age: As for longer and dichotomous at apical margin. Cratopteryx nemopteroides n. sp. Presence of a well-defined pterostigma in ScP and RA fusion area. RP origin very close Genus Araripeneura Martins-Neto and to wing base. Ten radial cells, basal one being Vu1cano 1989 the greater. Hipostigmal cell narrow, Type species Araripeneura regia Martins rectangular, twice longer than precedent one, Neto and Vulcano 1989, by original and, with basal margin placed at the same designation. level of the ScP and RA fusion. Eight RP branches, all without marginal dichotomies Araripeneura regia Martins-Neto and and three sectorial cells. MA origin close to Vulcano 1989 RP origin, with six distal secondary branches. (PI. IB; Fig. 1C-E) Intraradial cell (cir) long, placed at mid length of wing. MPI long, partially straight, Supplementary material: MPFT-I-009, without distal dichotomy. MP2+CuA parallel Museu de Paleontologia For9a da Terra, Sao and very close of MP. Six CuA secondary 41 branches, all they having distal dichotomies. Five radial cells, rectangular and of similar Just a long cross-vein before most proximal shape and SIze. Body (Fig. 2C) partially branch of euA (CuA2). CuP straight, preserved. reaching obliquely the anal margin the anal margin at mid length of wing, with four long Genus Cratoalloneura Martins-Neto 1992 secondary branches. Hind wing (Figure 2B) Type species Alloneura acuminata Martins long and narrow. Costal margin wider than Neto & Vulcano 1989, by original that of fore wing. Hipostigmal cell present. designation.

CuP ~E

Figure 2. A-C) Araripeneura urda n. sp., drawn from holotype MPFT-I-OIO, fore wing (A), hind wing (B), and body detail (C), respectively; D-F} Cratoalloneura verdandia n. sp., drawn from holotype MPFT-I-013, fore wing (D), hind wing (E), and body detail (F), respectively; G} Caririchrysa skulda n. sp., fore wing drawn from holotype MPFT-I-OI6; H} Blittersdorffia polyplusia Martins-Neto 1997, body detail drawn from supplementary material (MPFT-I-015). Scale bar 1 mm. 42

Cratoalloneura acuminata Martins-Neto Cratoalloneura acuminata Martins-Neto; in (PI. IID, E) second radial cell in Cratoalloneura verdandia n. sp.), and in its banks ian line not Supplementary material: MPFT-I-Oll, well-defined (present and well-defined In Museu de Paleontologia Forya da Terra, Sao Cratoalloneura acuminata Martins-Neto). Paulo - SP (PI. IID), and MPFT-I-012 (PI. lIE). Description: (Holotype MPFT-I-013, Figure Type locality, type stratum and age: As for 2D-F). Fore wing 20 mm long (Figure 2D), as Cratopteryx nemopteroides n. sp. preserved, and relatively wide, with acuminate apex. Costal area relatively narrow, filled by relatively long pectinated Cratoalloneura verdandia n. sp. cross-veins, all not dichotomous. RP origin (PI. IE; Fig. 2D-F) close to wing base. Ten relatively small radial cells. Hipostigmal cell long and notably Etymology: Allusive to Verdandi, "goddess narrow, with basal margin well before ScP of the present" in the Viking mitology. and RA fusion and the distal one close to Holotype: MPFT-I-013, Museu de wing apex. Seven RP branches, all without Paleontologia Forya da Terra, Sao Paulo - SP. marginal dichotomies. MA origin at second Supplementary material: MPFT-I-014, radial cell, with five distal secondary housed in the same institution as above (PI. branches. Intraradial cell (cir) long and IF). narrow. No preserved cross-veins before MA Type locality, type stratum and age: As for origin. MPI long, partially straight, distally Cratopteryx nemopteroides n. sp. with a single and long dichotomy. MP2+CuA parallel and very close to MPI. Eight CuA Diagnosis: Fore wing circa 20 mm long with, secondary branches, all with multiple distal at least, one pre sectorial cross-vein. Banksian dichotomies. Two long cross-vein before of line not well-defined. MA origin at the the most proximal branch of CuA (CuA2). second radial cell. CuP straight, reaching obliquely to anal margin a little after the 113 of wing base, with Discussion: Belongs to the genus at least eight long sigmoid secondary Cratoalloneura Martins-Neto 1992b by branches. Hind wing (Figure 2E) long and having MA origin close to basal radial cell, narrow. Costal margin relatively wide. very far from basal radial cell in Cratoneura Hipostigmal cell present, long and narrow. species, and CuP (long, circa 113 of wing Ten radial cells, the basal one being the base in Cratoalloneura verdandia n. sp., greater. Four pre medial cross veins. CuP short, close to the wing base in the other distally fused with M. Body (Fig. 2F) Cratoneura species). Cratoalloneura partially preserved, with a small head and verdandia n. sp. is similar to Cratoalloneura eyes, pronotum as long as the head and acuminata Martins-Neto 1992b, by having a mesonotum ellipsoid. similar fore wing length (20 mm), MA origin close to basal radial cell, same number of Genus Cratoneura Martins-Neto 1992 radial cells (ten), the same number of Type species Cratoneura longissima Martins secondary branches of RP and MP2+CuAl, Neto 1992, by original designation. and by the CuP length. Cratoalloneura verdandia n. sp. differs however, of Cratoneura dividens Martins-Neto 1994 Cratoalloneura acuminata Martins-Neto, by (Fig. 3) having at least one pre-sectorial cross-vein, MA ongIn (in basal radial cell In Supplementary material: RGMN - T - 010, 43

1IillITl1Ul~ _-[JIJd§ ~~ T~-=~~~ L- B

Figure 3. (sequence I). A-B) Cratoalloneura acuminata Martins-Neto and VuJcano, fore wing (A) and hind wing (B), respectively, drawn from supplementary material; C-O) Cratoneura /ongissima Martins-Neto, intraradial cell (A) and hipostigmal cell (D) detail, drawn from holotype. (sequence II). Cratoneura dividens Martins-Neto, intraradial cell detail (A) and hipostigmal cell detail (B), drawn from supplementary material (RGMN-T-OlO), intraradial cell (C) and hipostigmal cell (D), drawn from holotype (UnG-lT-045); (sequence III). Cratoneura dividens Martins-Neto, intraradial cell detail (A) and hipostigmal cell (B), drawn from supplementary material (RGMN-T-OI2), intraradial cell detail (C), ScP/RA fusion detail (D) and hipostigmal cell detail (E), drawn from supplementary material (RGMN-T-OI3). 44

012 and 013, Martins-Neto Collection, Cratopteryx nemopteroides n. sp. Sociedade Brasileira de Paleoartropodologia - SBPr. Family Chrysopidae Hagen 1866 Type locality, type stratum and age: As for Genus Caririchrysa Martins-Neto and Cratopteryx nemopteroides n. sp. Vu1cano 1989 Type species Caririchrysa criptovenata Genus Blittersdorffia Martins-Neto and Martins-Neto and Vulcano, by original Vu1cano 1989 designation. Type species Blittersdorffia pleoneura Martins-Neto and Vu1cano 1989 Emended diagnosis: Fore wmg with ScP short, reaching the costal margin at mid Blittersdorfjia polyplusia Martins-Neto 1997 length of wing; CuP with two or more (PI. IIC) secondary branches.

Supplementary material: MPFT-I-015, Museu de Paleontologia Forya da Terra, Sao Paulo - SP Type locality, type stratum and age: As for Cratopteryx nemopteroides n. sp.

Family Nemopteridae B Genus Cratonemopteryx Martins-Neto 1992 Type species Cratonemopteryx audax Martins-Neto and Vulcano 1989, by original designation.

Cratonemopteryx speciosa Martins-Neto and Vulcano 1997 (Fig.4A-D)

Supplementary material: RGMN-TI64; Martins-Neto Collection - SBPr Figure 4. A-D) Cralonemopteryx speciosa Martins Type locality, type stratum and age: As for Neto and Vuicano, Fore wing (A), hind wing (B) hind Cratopteryx nemopteroides n. sp. wing apex detail (C) and body detail (D), respectively, drawn from supplementary material; E-F) Roesleriana Genus Roesleriana Martins-Neto and sp. fore wing (E) and hind wing (F), respectively, drawn from supplementary material. Vulcano 1989 Type species Roesleriana exotica Martins Caririchrysa sku Ida n. sp. Neto and Vulcano 1989, by original (PI. IIF: Fig. 2G) designation. Etymology: Allusive to Skuld "goddess of Roesleriana sp. the future" in the Viking mitology. (Fig.4E-F) Holotype: MPFT-I-016, Museu de Paleontologia Forya da Terra, Sao Paulo - SP. Supplementary material: RGMN - T166; Supplementary material: GPII T-1675, Martins-Neto Collection - SBPr hosed III Paleontological Collection, Type locality, type stratum and age: As for Geosciences Institute, Universidade Sao 45

Paulo, USP (Martins-Neto 1992a, PI. IIF, Fig. Description: (Holotype, Fig. 2G). Fore wing 3A) and RGMN-T008, Martins-Neto 14 mm long and 5 mm wide. Costal area Collection, housed in the Sociedade Brasileira relatively wide. ScP short, reaching costal de Paleoartropodologia - SBPr (Martins-Neto margin at mid length of wing. RA long, 1997, PI. IB, Fig. 4). running parallel to costal margin, reaching Type locality, type stratum and age: As for apical area close to wing apex. Fifteen radial Cratopteryx nemopteroides n. sp. cells, basal one being smaller than subsequent. RP origin close to wing base. Diagnosis: Fore wing circa 15 mm long with Humeral vein present. M origin close to RP the M base very short, forming with r-m a origin, forming with r-m a small cell, smaller small cell as small as the basal radial cell. than basal radial cell. Pseudomedia and pseudocubitus well-defined by continuous Discussion: Similar to Mesypochrysa zigzag-like cross-veins. CuA long, parallel to latipennis Panfilov 1980, from the Asiatic MP2. CuP with at least three secondary Jurassic, in the general appearance, differing branches. however by the M morphology, which base is long, forming with r-m a long cell in Mesypochrysa latipennis (plesiomorphic Raphidioptera condition), short, forming with r-m a small Superfamily Raphidioidea sensu Willmann cell, as small as the basal radial cell in 1994 Caririchrysa sku Ida n. sp. (maybe an Family incertae sedis apomorphic condition). Caririchrysa sku Ida n. sp. differs of Caririchrysa criptovenata Caririraphidia n. gen. Martins-Neto and Vu1cano 1989a and Caririchrysa confusa Martins-Neto and Type species: Caririraphidia sertaneja n. sp., Vu1cano 1989a described for the same by present designation. outcrops by the size: fore wing 20 mm long in Etymology: Allusive to the Cariri Valley, C. criptovenata, 9 mm in C. confusa and 14 to which the material came from, and raphidia, 16 in C. skulda n. sp. from Raphidioidea.

Remarks: Martins-Neto and Vu1cano 1989 Diagnosis: Similar to Cratoraphidia Martins proposes the genus Caririchrysa for the Neto and Nel 1992 and Lugalla Willmann species Caririchlysa criptovenata and C. 1994, by having M transverse to the RP confusa. Martins-Neto (1992a), sinonymyzed origin, differing, however, by having m-cu Caririchrysa with Mesypochrysa Martynov perpendicular to M (horizontal, parallel to 1925 alleging several similarities. However a RA in Cratoraphidia as well as Lugalla). short ScP and the presence of CuP secondary branches are characteristics absent in the Discussion: Caririraphidia n. gen. differs of genus Mesypochrysa, justifying the generic all known genera by having M transverse to separation, being re-established the name RP origin, synapomorphy shared with just Caririchrysa for C. criptovenata, C. confusa two genera: Cratoraphidia Martins-Neto and and C. sku Ida n. sp. Nel 1992, of the same deposits, and Lugalla The specific characters of C. skulda n. Willmann 1994, from the Lower Cretaceous sp. are important apomorphies that could have of Mongolia, differing however, by having m a generic status but both holotype of C. cu perpendicular to M, horizontal, parallel to criptovenata and C. conjiJsa don't have the RA in Cratoraphidia as well Lugalla. basal area of the wing preserved. Remarks: The diagnostic characteristics of 46

Cratoraphidia Martins-Neto and Nel 1992 as The intraradial cell is the greater, placed near well as Lugalla Willmann 1994 are identical mid length of wing. Presence of two rows of and, by the nomenclatural priority code, cross-veins in radial sector, all parallel and Lugalla must be treated as new synonymy of oblique to costal margin, forming continuous the genus Cratoraphidia, sister genus of and homogeneous banksian lines. M origin in Caririraphidia n. gen. the RP base, sigmoidal, oblique to costal margin. Four cross-veins connecting RP to M Caririraphidia sertaneja n. sp. (rp-m). M forks in MA and MP circa at level (PI. IIIC; Fig. 5B, G) of intraradial cell. m-cu continuous to M, perpendicular. CuA, as preserved, parallel to Etymology: Allusive to interior, which the M, forking a little before m-cu, close to the material came from. wing base. Holotype: CD-I-l21, Desiree collection, housed in the Museu Nacional, Rio de Janeiro. Type locality: Unknown. Probably from outcrops between the Nova Olinda and Santana do Cariri municipalities, Ceara State, Northeast Brazil. Type stratum: Laminated limestone level of the Crato Member, lowermost unit of the Santana Formations, Araripe Basin. Age: Upper Aptian. Lower Cretaceous.

Diagnosis: Fore wing 17 mm long and 5 mm wide. Prothorax a little longer than the head.

Discussion: Differs of Cratoraphidia pulchra Martins-Neto and Nel 1992, apart the generic .-: characteristics, by having a greater size (Fore - wing 14.6 mm long and 4.3 mm wide in C. pulchra), by having the prothorax as long as the head (prothorax shorter than the head in C. pulchra).

Description: Body elongated and narrow (Figure 5B), circa 17 mm long. Prothorax (5 mm) a little longer than head (4.3 mm). Fore Figure 5. A, H) Caririraphidia? reticulata n. sp., wing (Figure 7G), 17 mm long and 5 mm drawn from holotype RGMN-163, general aspect (A) wide. Costal area relatively wide, with at least and fore wing detail (H), respectively; 8, G) eight pectinated cross-veins, perpendicular to Caririraphidia sertaneja n. sp., drawn from holotype costal margin. ScP slightly curved, reaching CD-I-121, general aspect (8) and fore wing detail (G), respectively; C) Arariperaphidia sp. drawn from costal margin circa 1/3 of t apex. RP long, specimen MPFT-I-006; D-F) Austroraphidia reaching apical margin above apex. RP origin brasiliensis Nel, Semeria and Martins-Neto 1992, circa 1/3 of wing base. Five radial cells, quite drawn from supplementary material MPFT-I-OI7, fore rectangular, the third one being the greater, wing (D), hind wing (E), and pronotum detail (F), longer than second and fourth together. Six respectively; I) Arariperaphidia rochai Martins-Neto and Vulcano 1990, drawn from supplementary material RP branches, all with marginal dichotomies. CD-I-046. Scale bar I mm. 47

Caririraphidia? reticulata n. sp. (five in C. sertaneja n. sp. irregulars in size, (PI. IlIA; Fig. SA, H) being the third one greater than second and fourth together). The placement in the genus Etymology: Allusive to the reticulated is tentative because important diagnostic pattern of the wing venation. characters as M and euA are unavailable in Holotype: RGMN-163, Martins-Neto this specimen as preserved. Collection, housed in the Sociedade Brasileira de Paleoartropodologia - SBPr. Genus Arariperaphidia Martins-Neto and Vulcano 1990 Diagnosis: Prothorax one time and half Type species. Arariperaphidia rochai longer than head. Fore wing 17 mm long and Martins-Neto and Vulcano 1990, by original 4 mm wide. RP 6-branched, parallel and designation. without marginal dichotomies. Eight radial cells, all, except basal one, relatively small. Arariperaphidia rochai Martins-Neto and Three rows of cross-veins in radial sector. V ulcano 1990 (PI. IIIB; Fig. SI) Description: Body elongated and narrow, circa 17 mm long (Figure SA). Head a little Supplementary material: CD-I-046, Desiree longer than wide, 3 mm long and 2.S mm Collection, housed in the Museu Nacional, wide. Eyes relatively large, prominent Rio de Janeiro. laterally, with 1.2 mm in it greater diameter, Type locality: Uncertain, probably came and 0.7 mm, in the smaller. Antenna from the outcrops between Nova Olinda and relatively long, with 4.4 mm. Pronotum Santana do Cariri municipalities, Ceara State. rectangular, 4 mm long and 1.8 mm wide. Northeast Brazil. Fore wing (Figure SH) 17 mm long and 4 mm Type stratum: Stratigraphic pOSItIOn wide. Eight radial cells that, except the basal unknown. Laminated limestone level, Crato one, are relatively small, square to rectangular Member, lowermost unit of Santana in shape. Six RP branches, all they without Fonnation, Araripe Basin. marginal dichotomies. Presence of three rows Age: Upper Aptian. Lower Cretaceous. of parallel cross-veins, all oblique to costal margm, not forming well-defined banksian Remarks: Specimen of similar size of lines. holotype (9.S mm), and identical wing venation. Discussion: Similar to Caririraphidia sertaneja n. sp. by having the same size (fore Arariperaphidia sp. wing and body 17 mm long) and the same (PI. IIIG; Fig. SC) number of RP branches (six), differing however, by having a prothorax relatively Supplementary material (female): MPFT-l- greater than head and RP branches without 006, housed in the Museu de Paleontologia marginal dichotomies (all distally Fon;;a da Terra, Sao Paulo - SP. dichotomous in C. sertaneja n. sp.). Additionally, C. reticulata n. sp. differs of C. Remarks: Female specimen. Body (except sertaneja n. sp., by having three rows of the ovipositor) and fore wing 11 mm long. cross-veins in the radial sector, not forming The preserved venation is similar to well-defined banksian lines (two rows in C. Arariperaphidia rochai Martins-Neto and sertaneja n. sp. and forming two well-defined Vu1cano 1990b. The size slightly greater and banks ian lines) and eight radial cells which, penultimate radial cell relatively greater, and except the basal one, are all relatively small ovipositor, are characteristics that don't 48 permit the placement of the specimen in A. In the case of Santana Formation rochai, but are not enough to propose a new paleoentomofauna, the exemplified steps in species for it. the Plate IV don't reflect transport but maybe variable flottability periods of the specimens, Genus Austroraphidia Willmann 1994 before their final deposition. Invariably, all Type species Raphidia brasiliensis Nel, the individuals already arrived died to the Semeria and Martins-Neto 1990 depositional site, with a minimum transport (very probably by the action of the winds). Austroraphidia brasiliensis Nel, Semeria and Not only the grasshoppers exhibit those Martins-Neto 1990 taphonomic peculiarities, but also the hopes (PI. lliF; Fig. 5E-F) (a regional Brazilian popular name for extant Tettigoniidae. In this case for Elcanidae) and Supplementary material: (female) MPFT-I- several neuropteran groups. The same 017, Museu de Paleontologia Fon;:a da Terra, analysis applied for the grasshoppers can be Sao Paulo - SP (Plate ITIF; Fig. 5E-F); used for Neuroptera Myrmeleontidae (Plate (male?). MPFT-I-018 (PI. ITID-E), part and V). In the sequence I (A-E), it is observed the counterpart, housed at the same institution. same taphonomic peculiarities (variable Type locality, type stratum and age: As for flottability periods) for specimens of the Cratopteryx nemopteroides n. sp. genus Cratoneura observed before the final deposition, and identical steps. In the sequence IT (A-D), the same situation is Taphonomy of Santana Formation Fossil observed now for specimens of the genus Insects and its Reflex in the Paleoecology. Blittersdorjjia. In the Plate VI (Sequence I A C» the same for Neuroptera Chrysopidae, In terms of taphonomic peculiarities, observing in "A", the specimens in natural according the abundance and the quality of post-mortem positioned (wings in rest preservation, four main fragmentation steps position). Although the wings are still in rest can be detected (of five possible), exemplified position, they are dissociated and broken into in the Plate IV by Caelifera. fragments (Sequence IB), but probably for a The preservation in the Step I occurs larger flottability period and subjection to when the specimen is intact, totally small oscillations in the lake, produced articulated and in a natural post-mortem perhaps by waves, rather than transport. In the position (wings in rest position), of that step sequence IT (A-C), the specimens of the genus two sub steps derive: predation, when the Araripeneura/Caririneura are showed specimen is destroyed by another organism, exhibiting the same peculiarities. before or during the deposition, and Some fortuitous cases of "micro fossilisation problems, when the specimen is transport" can be observed in the Figure 6A, destroyed, merely for collection problems. where, in a sample around lOx 5 cm, they are The Step II is characterised by the partial loss concentrated specimens of, at least, eight of the appendices (antennae and members), different arthropod orders, chaotically that in extant grasshoppers it is subject of few preserved, suggesting absence of a hours after death. In the Step III, all the preferential sense of orientation. In this appendices are lost, the head drops and the specific case, the currents of low intensity body begins its deterioration process, should have been responsible for the culminating in the Step IV, when are just accumulation of specimens III Araripe preserved isolated wings. The next step (V) paleo lake boundaries (the sample also would be characterised by the deterioration of contains detritus and plant remains). Also in the wings, until their total fragmentation. that case, all the insects already arrived died 49 to the depositional site (wings in a rest reconstruction of the Santana paleo lake at position). In the Figure 6B, a palaeoecologic Araripe times.

Figure 6. A) evidences of a micro-transport in the erato Member (reproduced from Martins-Neto 2002); B) A paleoecological reconstruction of the Araripe paleolake (reproduced from Martins-Neto 2002). 50

Phylogenetical approach on the Araripe representing Myrmeleontoidea. Neuropterofauna The matrix of data (Table II) was submitted to the program Henning 86, option Several authors discussed the mhenning*; bb* (Farris 1989) interface for nemopterid classification, being considered as Windows (Microsoft) Tree Gardner 2.2, a monophyletic group, although their resulting in a single tree, length 50, ci 74 and relationships with the other families are far of r1 88, not ordered and not weighted (Fig. 9). a consensus. Other authors demonstrate that Nemopteridae sensu lato are close to List of characters Myrmeleontidae + Ascalaphidae (see Aspock 1992). Handlirsh (1906) consider the 1. Oblique vein (0): absent (0), present (1) Kalligramatidae and Tillyard (1925) the Nymphidae could be their closer groups. The presence of an oblique vein IS Holzel (1975), Henry (1978), and Mansell synapomorphic for Nemopteroidea + (1986, 1992) considered Nemopteridae as M yrmeleontoidea. sister group of Ascalaphidae + Nymphidae. Popov (1973) suggested Nemopterinae and 2. Hipostigmal cell (h): present (0), absent (1) Crocinae as having a familial status and Monserrat (1996) formally proposed The presence of a hipostigmal cell is an Nemopteridae and Crocidae as distinct plesiomorphy of Neuropteroidea. The drastic families. reduction of the hipostigmal cell occurs several The Araripe's neuropterofauna is times within Neuropteroidea: in Ascalaphoidea, extremely diverse, containing several groups Nemopteridae, several genera of represented today as chrysopids, berothids, Myrmeleontidae (as Cymotales), Neurastenyx + nemopterids, myrmeleontids, ascalaphids and Paraneurastenyx and Bleyeria, being so psychopsids. The main focus of this homoplastic. The lengthening of the phylogenetical approach is the inner group of hipostigmal cell however occurs two times in a Myrmeleontoidea and Nemopteroidea and it non-homologous way, independently in the relationships with the closer groups (as group Caldasia + Cratoalloneura + Cratoneura Ascalaphoidea). For this propose was selected (where the hipostigmal cell is very long, thirty four characters exclusively of external occupying near 113 of the fore wing length and morphology (mainly wing venation) available notably narrow, narrower than the radial area in extant species as well as in fossils and was before the hipostigmal cell. In one of the used as external groups Psychops (extant) innermost group of Crocidae (Pterocroce + and Pulchroptilonia (fossil) representing Dielocroce + Ajghanocroce + Anacroce, for me Psychopsidae; Osmylops (extant), Dielocrocinae) the hipostigmal cell is also representing Osmylidae (without fossil notably longer but contrary of the mentioned specimens in Araripe); Babinskaia and myrmeleontid group, is as wide as the radial Neliana, representing Babinskaiidae (close to area and the anterior margin of h is before the Nymphidae); Ascalapha (extant), and ScPIRA fusion, placed at mid length of wing, Cratoscalapha (fossil), representing compressing the radial cross-veins. So, Ascalaphidae, Stenorrhachus (extant), hipostigmal cell long and narrow constitutes a Marquettia (fossil) Pastranaia (extant), sinapomorphy for Caldasia + Cratoalloneura + Pterocroce (extant), Roesleriana (fossil), Cratoneura (2"), and hipostigmal cell long, as Cratonemopteryx (fossil) and Krika (fossil), wide as the radial area with the anterior margin representing Nemopteroidea; a extant notably before the ScPIRA fusion, constitutes a myrmeleontids (Dendroleon and Cymotales) synapomorphy for Dielocrocinae (2'). with twelve fossil genera from Araripe, 51

Table II. Matrix of characters.

I 2 3 4 5 6 7 8 9 10 II 12 13 14 IS 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 Grupo 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Psychop. 0 0 0 0 0 0 0 0 0 0 0 0 0 I 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Cratopsvc 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Osmylid 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Neliana 0 0 0 0 0 0 I 0 0 0 0 I I 0 0 I 0 0 0 0 0 I I 0 I 0 0 0 1 0 0 0 0 0 Babinskai 0 0 0 0 0 0 I 0 0 0 0 1 I 0 0 I 0 0 0 0 0 1 1 0 I 0 0 0 1 0 0 0 0 0 Ascalaph 0 I 0 0 0 0 1 0 0 0 0 I 1 0 I 0 0 0 0 0 0 I 0 0 I 0 0 0 0 0 0 0 0 0 Cratoscal 0 I 0 0 0 0 I 0 0 0 0 I I 0 I 0 0 0 0 0 0 1 0 0 I 0 0 0 0 0 0 0 0 0 Karenina 0 I 0 0 0 0 1 0 0 0 0 1 I 0 I 0 0 0 0 0 0 I 0 0 I 1 0 0 0 0 0 0 0 0 Nemopter 1 I 0 0 0 0 1 I 1 0 0 1 1 0 1 0 0 0 1 0 0 1 1 0 I 0 0 0 0 I 0 0 0 0 Crocidae 1 0 0 0 0 0 I 2 1 0 0 I I 0 I 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 I 0 0 Roesleria 1 0 0 0 0 0 0 I I 0 0 1 I 0 1 0 0 0 1 0 0 I 0 0 1 0 0 0 0 0 0 0 0 0 Cratonem 1 0 0 0 0 0 1 I 1 0 0 1 1 0 I 0 0 0 I 0 0 1 0 0 1 0 0 0 0 1 0 0 0 0 Krika 1 0 0 0 0 0 1 1 1 0 0 1 1 0 1 0 I 0 1 0 0 1 0 0 I 0 0 0 0 1 0 0 0 0 Dendrole I 0 0 0 0 0 1 0 0 0 0 1 1 0 I 0 0 0 0 1 0 1 0 0 1 0 1 1 0 0 0 0 0 0 Cvmotal 1 I 0 0 0 0 I 3 0 0 0 I I 0 I 0 0 0 0 0 0 I 0 0 1 0 I I 0 0 0 0 0 0 Neuraste I I 0 0 0 0 0 0 0 0 0 I I 0 1 0 0 0 0 ? 0 0 0 0 I 0 I I 0 0 0 0 I 0 Paraneur I I 0 0 0 0 0 0 0 0 0 I I 0 I 0 0 0 0 ? 0 I 0 0 I 0 I I 0 0 0 0 1 0 Cratopter I 0 0 0 I 0 0 3 0 0 0 I I 0 I 0 0 0 0 0 I I 0 0 I 0 0 0 0 0 0 0 0 0 Pseudony 1 0 0 0 0 0 0 0 0 0 0 1 I 0 I 0 0 0 0 I 0 I 0 0 I 0 I 0 0 0 0 0 I 0 Blittersdor I 0 0 0 0 0 0 0 0 0 0 I I 0 I 0 0 0 0 I 0 I 0 0 I 0 I 0 0 0 0 0 I 0 Bleyeria I 0 0 0 0 0 0 0 0 0 0 I I 0 I 0 0 0 0 ? 0 I 0 0 I 0 I 0 0 0 0 0 I 0 Paracarir I 0 0 0 I 0 0 0 0 0 0 I I 0 I 0 0 I 0 I I I 0 I I 0 0 0 0 0 0 0 0 I Caldasia I 0 0 0 I I 0 0 0 I I I I 0 I 0 0 I 0 ? I I 0 0 I 0 0 0 0 0 1 0 0 0 Araripen I 0 I 0 I 0 0 0 0 0 1 1 I 0 1 0 0 1 0 I 1 1 0 0 I 0 0 0 0 2 0 0 0 0 Cratoallon 1 0 I I 1 1 0 0 0 1 1 I I 0 I 0 0 1 0 I I I 0 0 I 0 0 0 0 0 I 0 0 0 Caririneur I 0 0 0 1 0 0 3 0 0 I 1 1 0 I 0 0 I 0 I I I 0 0 I 0 0 0 0 2 0 0 0 0 Cratoneur I 0 0 I I I 0 0 0 I I ! I 0 I 0 0 I 0 I I 1 0 0 I 0 0 0 0 0 1 0 0 0

3. Fore wing MA origin: far from radial basal branches (0), multibranched (1). cell (0), close to radial basal cell (1) The presence of several MA secondary The MA origin close to radial basal cell branches (six or more) is synapomorphic for occurs independently in Araripeneura and Cratopteryx + Paracaririneura + Cratoalloneura. Araripeneura + Caririneura + Caldasia + Cratoalloneura + Cratoneura. 4. Fore wing apical area: wide (0), notably narrow (1) 6. Fore wing MPl: unbranched (0), multibranched (1 ). The presence of a notably narrowing of the apical area is synapomorphic for For wing with MPI multibranched IS Cratoalloneura + Cratoneura. synapomorphic for Caldasia + Cratoalloneura + Cratoneura. 5. Fore wing MA: with few secondary 52

7. Fore wing pre-sectorial cross veins: absent Hind wing costal area wider than that of (0), present (1). fore wing is synapomorphic for Caldasia + Cratoalloneura + Cratoneura. In the most basal groups of Neuroptera (as Osmylidae), the pre-sectorial cross-veins 11. Fore wing intraradial cell: short (0); long are absent (plesiomorphy). The presence of (1) these cross-veins is sinapomorphic for Babinskaiidae + Ascalaphoidea + A long intraradial cell in the fore wing Nemopteroidea + Myrmeleontoidea, the constitutes a synapomorphy of Araripeneura character reversion occurring independently + Caririneura + Caldasia + Cratoalloneura in Roesleriana (Nemopteroidea), in the group + Cratoneura. Neurastenyx + Paraneurastenyx + Pseudonymphes + Blittersdorffia + Bleyeria 12. Fore wing ScP and RA: distally not fused and In the group Cratopteryx + (0); distally fused (1) Paracaririneura + Araripeneura + Caririneura + Caldasia + Cratoalloneura + ScP and RA distally fused is a Cratoneura, all fossils from Araripe. synapomorphic condition of Osmyloidea + Babinskaiidae + Ascalaphoidea + 8. Hind wing: as long as the fore wing (0), Nemopteroidea + Myrmeleontoidea. longer than fore wing (1) J 3. Fore wing costal margin: wide (0); During a long time this was considered narrow (1) as an important synapomorphy for Nemopteroidea. However this character The narrowing of the fore wing costal occurs independently in other groups of margin is synapomorphic for Babinskaiidae + Myrmeleontoidea as Cymotales (extant Ascalaphoidea + Nemopteroidea + Myrmeleontidae), and in the fossil taxa Myrmeleontoidea. Cratopteryx and Caririneura. Then the character could be multistate as follows: hind 14. Fore wing vena triplica: absent (0); wing slightly longer than fore wing, present (1) homoplastic, occurring independently in Cymotales, Cratopteryx and Caririneura (1); The presence of a vena triplica in the hind wing two times longer than fore wing, fore wing is synapomorphic for Psychopsidae sinapomorphic for Nemopteridae + + Pulchroptilonia. Roesleriana + Cratonemopteryx + Krika (2), and hind wing three times longer than fore 15. Fore wing MP2+CuAl and CuA2: wing, constituting a Crocidae autapomorphy parallel (0); forming a triangular area (1) (3). MP2+CuAl and CuA2 forming a 9. Hind wing: as wide as fore wing (0); triangular area in the fore wing is a notably narrower than the anterior (1) synapomorphic condition for Ascalaphoidea + Nemopteroidea + Myrmeleontoidea. The narrowing of the hind wing is synapomorphic for Nemopteroidea. 16. Fore wing MP2+CuAl: reaching the anal margin (0); reaching the apical margin (1) 10. Hind wind costal area: narrower than that of fore wing (0); wider (1) A long MP2+CuAl in the fore wing, reaching the apical margin is a synapomorphy 53 of Babinskaia + Neliana. this fossil group.

17. Microtrichia: absent (0); present m all 22. Fore wing radial cells: abundant (0); veins (1). reduced (1)

Presence of microtrichia in all veins is The presence of a great number of autapomorphic for Krika. radial cells is a plesiomorphic condition of basal nemopteroid groups as well as 18. Fore wing CuA2: unbranched (0); two Psychopsidae and Osmyloidea. The drastic branched (1) reduction of the number of these cells IS a synapomorphic condition shared by CuA2 two-branched in the fore wing is Babinskaiidae + Ascalaphoidea + synapomorphic for Paracaririneura + Nemopteroidea + Myrmeleontoidea. Within Araripeneura + Caririneura + Caldasia + the mner group of Myrmeleontoidea, Cratoalloneura + Cratoneura. Neurastenyx exhibits a reversion of this character. 19. Hind wing: without distal dilatation (0); with dilatation (1) 23. Fore wing RP origin: close to the wing base (0); far from the wing base (1). The presence of distal dilatations in the hind wmg IS synapomorphic for The RP origin within Neuropteroida is Nemopteridae + Roesleriana + extremely variable, occurring very close to Cratonemopteryx + Krika within the wing base in the basal groups like Nemopteroidea, occurring reversions in few Osmyloidea, and circa 113 in the mean part of genera within Nemopteridae. groups such Myrmeleontoidea and Ascalaphoidea. RP origin notably far from 20. Antennae: long and clavate (0); short and the wing base (after the mid length of wing) clavate (1) occurs two times, independently in Babinskaiidae and in Nemopteridae within Antennae short and clavate IS a Nemopteroidea. homoplastic character within Myrmeleontoidea, occurring independently in 24. Fore wing RP branches: three or more several groups (and at least one extant genus (0); one (1) within Ascalaphidae). All Araripe myrmeleontids shares a short and clavate The drastic reduction of the RP antenna, except Cratopteryx. branches occurs just one time within Myrmeleontoidea, constituting an 21. Fore wing pre-hipostigmal radial cell: autapomorphy of Bleyeria. rectangular (0); trapezoidal (1) 25. Fore wing MP2 and CuAl: unfused (0); All Neuropteroidea have a pre fused (1) hipostigmal radial cell rectangular. The presence of a trapezoidal pre-hipostigmal The fusion of MP2 and CuAl is a radial cell occurs just one time in synapomorphic condition of Babinskaiidae + Myrmeleontoidea, in the group Cratopteryx + Ascalaphoidea + Nemopteroidea + Paracaririneura + Araripeneura + Myrmeleontoidea. Caririneura + Caldasia + Cratoalloneura + Cra ton eura , constituting a synapomorphy of 26. Fore wing ScP and RA: normal (0); 54 drastic thickening (1) synapomorphic for Araripeneura + Caririneura. The drastic thickening of ScP and RA in the fore wing occurs just one time within 32. Hind wing apical venation: present (0); Neuropteroida, constituting an autapomorphic absent (1) condition for Cratoscalapha. The loose of the hind wing apical 27. Fore wing CuA2: long (0); short (1) venation is autapomorphic for Crocidae.

Within the group Ascalaphoidea + 33. Fore wing oblique vein: posterior to Nemopteroidea + Mynneleontoidea, where cubitus fork (0); anterior (1) MP2 and CuAI are partially fused, CuA2 is rather a long vein, oblique to MP2 + CuAI. Oblique vein posterior to cubitus fork is CuA2 short is synapomorphic for typical a synapomorphic condition shared by the mynneleontids (Mynneleontidae + Mynneleontoidea group Neurastenyx + Neurastenyx + Paraneurastenyx + Paraneurastenyx + Pseudonymphes + Pseudonymphes + Blittersdorjjia + Bleyeria). Blittersdorjjia + Bleyeria.

28. Fore wing radial sector: without banksian 34. Fore wing cross-veins: nonnal (0); line (0); with a conspicuous banksian line (1) fonning mosaics (1)

The presence of a conspicuous banksian The presence of a great number of line linking the RP secondary branches is also supplementary cross-veins fonning mosaics a synapomorphy of Mynneleontidae + in the fore wing is autapomorphic for Neurastenyx + Paraneurastenyx + Paracaririneura. Pseudonymphes + Blittersdorjjia + Bleyeria.

29. Fore wing CuP: straight (0); zigzag-like Results (1) The present phylogenetical approach Fore wing zigzag-like is autapomorphic based exclusively on external morphology for Babinskaiidae (Babinskaia + Neliana). does not absolutely solve the relationships of the Neuropteroida outgroups but appears 30/31. Fore wing Cua2 origin: posterior to useful to solve the phylogeny of the inner RP origin (0); anterior to RP origin (1); at the mynneleontids groups. Babinskaiidae is a same level (2) well-defined monophyletic group supported by two synapomorphies (characters 16 and The CuA2 ongm in Ascalaphoidea + 29). The Ascalaphoidea for example are not Nemopteroidea + Mynneleontoidea is supportable by synapomorphies, at least based posterior to RP origin (synplesiomorphic on the characters discussed here. However condition). CuA2 origin anterior to the RP three inner groups of Mynneleontoidea are origin occurs independently two times within well-defined: Nemopteroidea, typical Ascalaphoidea + Nemopteroidea + mynneleontids and araripeneurids. Within Mynneleontoidea: within Nemopteroidea Nemopteroidea, Crocidae and Nemopteridae (Nemopteridae + Cratonemopteryx + Krika) are monophyletic, as well as Roesleria and in the group of Mynneleontoidea (Family Roeslerianidae). Within (Caldasia + Cratoalloneura + Cratoneura). Mynneleontoidea, three distinct groups are CuA2 and RP origin at the same level is confinned: typical Mynneleontidae, including 55 extant taxa (Cymotales and Dendroleon), the Caririneura, Caldasia, Cratoalloneura, and palaeoleontinids (Palaeoleontidae), including Cratoneura. exclusively fossil taxa (Neurastenyx and Paraneurastenyx from Brazilian Lower Cratoalloneurini n. trib. Cretaceous, but with a wide distribution which representatives are known for Cladistic diagnosis: Canadian, Spanish, English, Russian and Fore wing hipostigmal cell notably long Chinese Lower Cretaceous). This group have and narrow; some plesiomorphic characters (great number Fore wing MPl multibranched (more than of radial cells for example) and compound five secondary branches); rather by big-sized species. The third group, Hind wing costal area wider than the fore Pseudonymphidae, includes endemic genera wmg one; from Brazilian Lower Cretaceous and other Fore wing CuA2 origin anterior to the RP genera described for the Asiatic Lower ongm. Cretaceous as well. The inner relationship of these groups are unclear, being supported by Constituent genera: Caldasia, Cratoalloneura few apomorphic characters, but the whole and Cratoneura. group is monophyletic and supported by two synapomorphies (characters 27 and 28). An endemic and monophyletic group of Acknowledgements Myrmeleontoidea is defined as follows: Thanks are due to Dr. Andre Nel Araripeneuridae n. status. (Museum National d'Histoire Naturelle, Paris - Entomologie) by criticism and suggestions Type genus Araripeneura Martins-Neto and on the manuscript. A special thanks also to Vulcano 1989 Museu de Paleontologia Fon;a da Terra, Sao Cladistic diagnosis: Paulo, by the loan of several specimens as Fore wing with MA multibranched (more well as Dr. Alexander Wilhein Armin Kellner than five secondary branches) also by the loan of specimens, now belonging Fore wing pre-hipostigmal cell trapezoidal to Museu Nacional Paleontological Collection, Rio de Janeiro. This contribution Constituent genera: Cratopteryx Martins-Neto is part of my defended doctoral thesis (2002) and Vulcano 1989; Paracaririneura Martins at Universidade do Vale do Rio dos Sinos - Neto and Vulcano 1997; Araripeneura UNISINOS, which receive my special thanks Martins-Neto and Vulcano 1989; Caririneura too. Martins-Neto and Vulcano, 1989; Caldasia Mm1ins-Neto and Vulcano, 1989; Cratoalloneura Martins-N eto 1992 and References Cratoneura Martins-Neto 1994. All from Brazilian Lower Cretaceous. ASPOCK, U. 1992., Crucial points in the phylogeny of the Neuroptera (Insecta). In: CANARD, M., Araripeneurinae sensu stricto ASPOCK, H. and MANSELL, M. W. (Eds.). Current research in Neuropterology. Proceedings Cladistic diagnosis: Fore wing with a notably long intraradial Forth Int. Symp. On Neuropterology, Toulouse, pp. cell 63-73. BORROR, D. 1. and DeLONG, D. M., 1964. An Constituent genera: Araripeneura, introduction to the study of insects. New York, 56

Holt, Rinehart and Winston, Inc. 653 pp. MARTINS-NETO, R. G., 1992b. Neuropteros (Insecta: BURMEISTER, H., 1839. Neuroptera. Handbuch der Planipennia) da Forma<;ao Santana (Cretaceo Entomologie, 2(2): 757-1017. Inferior), Bacia do Araripe, Nordeste do Brasil. VII. CARPENTER, F. M., 1960. Fossil Nemopteridae. Palaeoleontinae, nova subfamilia de Psyche, 66: 20-24. Myrmeleontidae e descri<;ao de novos taxons. FARRIS, J., 1989. Hennig86: a PC-DOS Program for Revta. Bras. Ent., 36 (4): 803-815. Phylogenetic Analysis. Cladistics, 5: 163. MARTINS-NETO, R. G., 1994. Neuropteros (Insecta, HAGEN, H. A., 1866. Die Neuropteren Spaniens ach Planipennia) da Forma<;ao Santana (Cretaceo Ed. Pictet's Synopsis des Neuropteres d'Espagne Inferior), Bacia do Araripe, Nordeste do Brasil. IX - und Dr. Staudingers. Mitteilungen. Stettin. Primeiros resultados da composi<;ao da fauna e Entomol. 27: 281-320. descri<;ao de novos taxa. Acta. Geologica HANDLIRSCH, A., 1906. Die Fossilen Insekten und Leopoldensia, 39(1): 269-288. die Phylogenie der Rezenten Formen. In: MARTINS-NETO, R. G., 1996. New Mayflies Handbuch fur Palaontologie und Zoology. Leipzig, (Insecta, Ephemeroptera) from the Santana Engelmann. 640 p. Formation (Lower Cretaceous). Araripe Basin, HENRY, C. S., 1978. An unusual ascalaphid larva Northeast Brazil, Revista Espanola de (Neuroptera, Ascalaphidae) from Southern Africa, Paleantologia, 11(2): 54-70. with comments on larval evolution within the MARTINS-NETO, R. G., 1997. Neuropteros (Insecta, Myrmeleontoidea. Psyche, 85(2): 265-274. Planipennia) da Forma<;ao Santana (Cretaceo HOLZEL, H., 1975. Revision der NetzflUgler - unter Inferior), Bacia do Araripe, Nordeste do Brasil. X - familie Crocinae (Neuroptera: Nemopteridae). Descri<;ao de novos taxa (Chrysopidae, Entomol. Ger., 2: 44-97. Babinskaiidae, Myrmeleontidae, Ascalaphidae e MANSELL, M. W., 1986. Biogeography and Psychopsidae), Rev. Universidade de Guarulhas, Phylogeny of the Crocinae (Neuroptera: serie Ciencias Exatas e Tecnologicas, 2(4): 68-83. Nemopteridae) In: GEPP, J., ASPOCK, H. and MARTINS-NETO, R. G., 1998. Neuropteros (Insecta, HOLZEL, H. (eds.), Recent research in Planipennia) da Forma<;ao Santana (Cretaceo Neuropterology. Graz: 77-84. Inferior), Bacia do Araripe, Nordeste do Brasil. XI - MANSELL, M. W., 1992. The systematic position of Descri<;ao de novos taxons de Myrmeleontidae the Nemopteridae. In: CANARD, M., ASPOCK, H. (Palaeoleontinae e Pseudonymphinae). Rev. and MANSELL, M. W. (eds.), Current research in Universidade Guarulhas ser. Ciencias Biologicas e Neuropterology. Proc. Fourth International da Saude, 3(5): 38-42. Symposium in Neuropterology. Toulouse, pp. 233- MARTINS-NETO, R. G., 1999. La Paleoentomofauna 241. Brasilefia. Estado actual del conocimiento. Rev. MARTINS-NETO, R. G., 1992a. Neuropteros (Insecta: Entoma!' Argentina, 58(1-2): 71-85. Planipennia) da Forma<;ao Santana (Cretaceo MARTINS-NETO, R. G., 2000. Remarks on the Inferior), Bacia do Araripe, Nordeste do Brasil. V. Neuropterofauna (Insecta, Neuroptera) from the Aspectos filogeneticos, paleoecologicos, Brazilian Cretaceous with keys for the paleobiogeograticos e descri<;ao de novos taxa. An. identification of the known taxa. Acta Geologica Acad. Bras. Ci., 64(2): 117-148. Hispanica, 35(1-2): 97-118. 57

MARTINS-NETO, R. G., 2002. Insetos fosseis como Forma((ao Santana (Cretaceo Inferior). Bacia do bioindicadores em depositos sedimentares: um Araripe, Nordeste do Brasil. VIII descri((ao de

estudo de caso para 0 mesozoico sul-americano. novos taxa de Myrmeleontidae, Ascalaphidae e Sao Leopoldo, RS. Tese de Doutorado. Nemopteridae. Rev. UnG, Serie Ciencias Universidade do Vale do Rio dos Sinos - Unisinos, B iol6gicas, 2(5):64-8l. 214 p. MARTYNOV, A. V., 1925. Contributions to the MARTINS-NETO, R. G. and NEL, A., 1992. Un knowledge of fossil insects from Jurassic beds in nouveau fossile de Raphidioptere de la Formation Turkestan. 2. Raphidioptera (cont.), Orthoptera (s. Santana, Cretace Inferieur del Bresil I.), Odonata, Neuroptera. Izv. Ross. Akad. Nauk., 2: (Neuropteroidea, Raphidioptera) Bull. Soc. Ent. 569-598. France, 97(5): 425-428. McLACHLAN, F. R. S., 1885. On the discovery of a MARTINS-NETO, R. G. and VULCANO, M. A., species of neuropterous family Nemopteridae in 1989a. Neur6pteros (Insecta, Planipennia) da South America, with general considerations Forma((ao Santana (Cretaceo Inferior), Bacia do regarding the family. Trans. En!. Soc. Condo Part Araripe, Nordeste do Brasil. Familia III: 375-879. Chrysopidae. An. Acad. bras. Ci., 60(2): 189-20l. MONSERRA T, V. J., 1996. Larval stages of European MARTINS-NETO, R. G. and VULCANO, M. A., Nemopterinae, with systematic considerations on 1989b. Neur6pteros (Insecta, Planipennia) da the Family Nemopteridae (Insecta, Neuroptera). Forma((ao Santana (Cretaceo Inferior) Bacia do Dtsch. Ent. z., 43(1): 99-12l. Araripe, Nordeste do Brasil. II - Superfamilia NEL, A., SEMERIA, Y. and MARTINS-NETO, R. G., Myrmeleontoidea. Rev. Bras. Ent., 33(2): 367-402. 1990. Un Raphidioptera F6ssile du Cretace MARTINS-NETO, R. G. and VULCANO, M. A., Inferieur du Bresil (Neuropteroidea). Neuroptera 1989c. Neur6pteros (Insecta, Planipennia) da International, 6(1); 27-37. Forma((ao Santana (Cretaceo Inferior), Bacia do ORFILA, R. N., 1955. Un nuevo Nemopteridae Araripe, Nordeste do Brasil. IV - Complementos as americano com una sinopsis de la familia. Rev. Soc. Partes I e II, com descri((ao de novos taxa. An. Ent. Argentina, 17: 29-32. Acad. Bras. 0.,61(3):311-318. PANFILOV, D. V., 1980. New representatives of MARTINS-NETO, R. G. and VULCANO, M. A., lacewings (Neuroptera) from the Jurassic of 1990a. Neur6pteros (Insecta, Planipennia) da Karatau. In. DOLIN, V. G., PANFILOV, D. V. and Forma((ao Santana (Cretaceo Inferior) Bacia do PRITYKINA, L. N. (eds.), Mesozoic fossil insects. Araripe, Nordeste do Brasil. III - Superfamilia Naukowa Dunka, p. 82-111. Mantispoidea. Rev. Bras. Ent., 34(3): 619-625. PETRI, S., 1998. PaleocIimas da Era Mesoz6ica no MARTINS-NETO, R. G. and VULCANO, M. A., Brasil evidencias paleontol6gicas e 1990b. Primeiro Registro de Raphidioptera sedimentol6gicas. Revta. UnG, Serie Geociencias, (Neuropteroidea) na Forma((ao Santana, Cretaceo 3(6): 22-38. Inferior, Bacia do Araripe, Nordeste do Brasil. Rev. POINAR, G. 0. and STANGE, L. A., 1996. A new Bras. Ent., 34(1): 241-249. antlion from Dominican amber (Neuroptera: MARTINS-NETO, R. G. and VULCANO, M. A., Myrmeleontidae). Experientia, 52: 383-386. 1997. Neur6pteros (Insecta, Planipennia) da POPOV, A., 1973. Ober die praimaginalen stadien 58

palaarktisher vertreter der ordnund Neuroptera und versuch einer neuen systematischen gruppierung der familien mit rilcksicht auf ihre morphologischen und 6kologischen besonderheiten. Bull. Ins!. Zoo!. Mus. Sofia, 37: 79-10l. TILL YARD, R. J., 1925. A new species of spoon winged lacewing from Western Australia. J. Royal Soc. West Australia, 12: 1-4. WILMANN, R., 1994. Raphidiodea aus dem Lias und die Phylogenie der Kamelhasfliegen (Insecta: Holometabola). Pa/dont. z., 68(1/2): 167-197. WHITHYCOMBE, C. L., 1923. Systematic notes on the Crocini (Nemopteridae) with descriptions of new genera and species. Trans. Ent. Lond. Pars I, II: 269-287, PI. XII, XIII.

Recebido em 03/04/2003 Revisiio aceita em 15/04/2003 59

K I

Figure 7. Neuroptera representatives. A) Pulchroptilonia spatifata Martins-Neto 1997 (Psychopsidae); B) General extant Osmylidae, reproduced from Borror & DeLong 1964; C) Neliana maculata Martins-Neto 1994 (Babinskaiidae); D) Cratoscalapha electroneura Martins-Neto and Vulcano 1997 (Ascalaphidae); E) Stenorrhachus walkeri McLachlan, modified from McLachlan 1885 (Nemopteridae); F) Marquettia americana Carpenter, modified from Carpenter 1960 (Nemopteridae); G) Pastranaia riojana Orfila, modified from Orfila 1955 (Crocidae), H) Pterocroce storeyi Withicombe, modified from Withicombe 1923 (Crocidae); Roesleriana exotica Martins-Neto and Vulcano 1989 (Roeslerianidae); Cratonemopteryx speciosa Martins-Neto & Vulcano 1997 (Nemopteridae); K) Cymotales dulcis Gerstaecker, reproduced from Poinar & Stange 1996 (Myrmeleontidae).

J-=~ ~~~f~;~~q~, -;~~ttrpY

Figure 8. Neuroptera representatives. A) Neurastenyx araripensis Martins-Neto 1992 (Palaeoleontidae); B) Paraneurastenyx ascalaphix Martins-Neto 1997 (Palaeoleontidae); C) Bleyeria nordestina Martins-Neto 1992 (Myrmeleontidae); D) Cratopteryx robertosantosi Martins-Neto and Vulcano 1989 (Araripeneuridae); E) Pseudonymphes araripensis Martins-Neto and VuIcano 1989 (Pseudonymphidae); F) Blittersdorffia pulcherrima Martins-Neto and Vulcano 1997 (Myrrneleontidae); G) Paracaririneura priscila Martins-Neto and Vulcano 1997 (Araripeneuridae); H) Araripeneura regia Martins-Neto and VuIcano 1989 (Araripeneuridae); Cratoalloneura verdandia n. sp. (Araripeneuridae); J) Cratopteryx nemopteroides n. sp. (Araripeneuridae); K, L) Cratoneura dividens Martins-Neto 1994 (Araripeneuridae). 60

ARARIPENEURIDAE ARARIPENEURINAE ~ s::: CD I S .s:::'" .. s::: ~ III s::: 'tJ 0 S 1'!!'" ~ ~ .s:::: .S!'" .S! :::J ~ .g ~ .. S 1'!!'" CD .g .~ ii 0 ~ :::J s::: I::"! CD o

Figure 9. Phylogeny of the Araripe Neuropterofauna (except Chrysopidae, Berothidae and related groups). 61

Plate I. A) Cratopteryx nemopteroides n. sp., holotype MPFT-I-004; B) Araripeneura regia Martins-Neto and Vulcano 1989, supplementary material MPFT-I-009; C-D) Araripeneura urda n. sp., holotype MPFT-I-OlO, part (C) and counterpart (D), respectively; E-F) Cratoalloneura verdandia n. sp., holotype MPFT-I-OI3 (E) and supplementary material MPFT-I-14 (F). 62

Plate II. A) Caririneura damianii Martins-Neto, supplementary material MPFT-I-007; B) Caririneura crassatella Martins-Neto and Vulcano 1997, supplementary material MPFT-I-008; C) Blittersdorffia polyplusia Martins-Neto 1997, supplementary material MPFT-I-OI5; D-E) Cratoalloneura acuminata Martins-Neto and Vulcano 1989, supplementary material MPFT-Oll CD) and MPFT-I-OI2, respectively; F) Caririchrysa skulda n. sp., holotype MPFT-I-016. 63

Plate III. A) Caririraphidia? reticulata n. sp., holotype RGMN-163; B) Arariperaphidia rochai Martins-Neto and Vulcano 1990, supplementary material CD-I-046; C) Caririraphidia sertaneja n. sp., holotype CD-I-121; D-F) Austroraphidia brasiliensis Nel, Semeria and Martins-Neto 1992, supplementary material MPFT-I-O 17, part (D), counterpart (E), and MPTF-I-O 18 (F); G) Arariperaphidia sp., specimen MPFT-I-006. 64

STE~ II: ~~rtiallost of the· (antenna and members)

PREDATION COLLECTING PROBLEMS

.~ .

..• 'If STEP I

Plate IV. Taphonomic peculiarities on Santana Formation Caelifera, exhibiting several fragmentation steps discussed in the text All specimens are in natural post-mortem position (wings in rest position), indicatives that they came died in the Araripe paieolake. 65

Plate V. Taphonomic peculiarities on Santana Formation Neuroptera. Sequence I (A-E): specimens of genus Cratoneura (Neuroptera Mynneleontidae); Sequence II: specimens of genus Blittersdorffia (Neuroptera Mynneleontidade ). 66

'II

Plate VI. Taphonomic peculiarities on Santana Formation Neuroptera. Sequence I (A-C): specimens of genus Caririchrysa (Neuroptera Chrysopidae); Sequence II: specimens of genus Araripeneura / Caririneura (Neuroptera Myrmeieontidade ).

Bibliography of the Neuropterida

Bibliography of the Neuropterida Reference number (r#): 10702

Reference Citation: Martins-Neto, R. G. 2002 [2003.??.??]. The Santana Formation paleoentomofauna reviewed. Part I -- Neuropteroida (Neuroptera and Raphidioptera): systematic and phylogeny, with description of new taxa. Acta Geologica Leopoldensia (R.S.)25(55):35- 66.

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