6(1):1-8, jan/jun 2010 © Copyright 2010 by Unisinos - doi: 10.4013/gaea.2010.61.01 New neuropteran (, Palaeoleontidae, Araripeneuridae and ) from the Santana Formation, Early NE Brazil

Rafael Gioia Martins-Neto Universidade Federal do Ceará (UFC), Campus Cariri/FUNCAP/Sociedade Brasileira de Paleoartropodologia – SBPr. Av. Tenente Raimundo Rocha, s/n, Cidade Universitária, 63000-000, Juazeiro do Norte, CE, Brazil. [email protected] (Deceased in August 6, 2010)

Viviane Zeringóta Rodrigues Centro de Ensino Superior de Juiz de Fora – CES/JF, Programa de Pós-graduação em Biologia, Universidade Federal de Juiz de Fora – UFJF, Campus Universitário Martelos, 36036-900, Juiz de Fora, MG, Brazil. [email protected]

ABSTRACT This paper aims to present the results of a taxonomic study with the from the Crato Member, Santana Formation, Lower Cretaceous from Araripe Basin, Ceará, Brazil. Based on these results, it is proposed the following new taxa: Nuddsia repatriata n. sp. (Osmylidae), Neurastenyx conani n. sp. (Palaeoleontidae), Diegopteryx raptorius n. gen. et n. sp. (Araripeneuridae, Cratopteryxinae n. subfam.), and Putzneura parcimoniosa n. gen. et n. sp. (Psychopsidae). The new taxa attests the success of the Neuroptera in equatorial areas of Gondawana during the end of , similar to that reached in Northern Hemisphere, which is valid mainly to the Psychopsidae from Brazil, that show big-size and conspicuous colour patterns, intense pubescence, and it is associated to other endemic forms.

Key words: Neuroptera, Santana Formation, Araripe Basin, Lower Cretaceous, Brazil.

RESUMO NOVOS NEURÓPTEROS CRETÁCEOS (OSMYLIDAE, PALAEOLEONTIDAE, ARARIPENEURIDAE E PSYCHOPSIDAE) DA FORMAÇÃO SANTANA, NE DO BRASIL. Este artigo apresenta resultados de um estudo taxonômico de fósseis de insetos do Membro Crato da Formação Santana, Bacia do Araripe, Ceará, Brasil. Com base nos resultados, os seguintes novos táxons são propostos: Nuddsia repatriata n. sp. (Osmylidae), Neurastenyx conani n. sp. (Pal- aeoleontidae), Diegopteryx raptorius n. gen. et n. sp. (Araripeneuridae, Cratopteryxinae n. subfam.) e Putzneura parcimoniosa n. gen. et n. sp. (Psychopsidae). Os novos táxons descritos atestam o sucesso dos Neuroptera nas áreas equatoriais do Gondwana no fi nal do Mesozóico, semelhante àquele que possuíam no Hemisfério Norte. Para a assembleia do Brasil, destacam-se os Psychopsidae, por seu grande tamanho e presença de cor, pubescência e sua associação a um conjunto de outras formas endêmicas de Neuroptera.

Palavras-chave: Neuroptera, formação Santana, Bacia do Araripe, Cretáceo Inferior, Brasil.

INTRODUCTION Martins-Neto and Rodrigues, 2009). The partially rescued here and redescribed, present contribution provides new forms being housed in Brazilian Institutions. The Neuropterans are one of the to that described, including a giant form commonest orders of fossil insects in of Neuroptera. The Araripe neurop- MATERIAL AND METHODS terms of named species in the laminated terofauna reveals again a great diversity, limestones of Araripe Basin, with sev- close to that which today characterizes The samples here focused consist of eral morphologic, paleoecologic, and the extant groups, mainly those included four selected slabs recently collected in the paleoethologic aspects being focused in Osmylidae and Psychopsidae. Some expositions along Nova Olinda-Santana on previous works (Martins-Neto, 1991, few Neuroptera taxa described by foreign do Cariri road, 4 km from the municipal 1992, 1994, 1997, 1998, 2005, 2006; authors based on material keep out from district of Nova Olinda, in the well-known Martins-Neto and Vulcano, 1989, 1997; Brazil and which holotypes are fortunately laminated limestone from the Crato New neuropteran insects from the Santana Formation, NE Brazil

Member, Santana Formation, Late in age (Figure 1A-B). The biostratigraphy of the Santana Formation was previously proposed by Coimbra et al. (2002). The terminology used here is based on Martins-Neto (2002). The abbrevia- tions cited in the text are: MA, anterior media; MP, posterior media; RA, anterior radial; RP, posterior radial; rp-ma, cross vein connecting RP with MA and ScP, posterior subcosta.

SYSTEMATIC PALEONTOLOGY

Order NEUROPTERA Olivier, 1789 OSMYLIDAE Leach, 1815 Subfamily GULLIMINAE Navás, 1912

Nuddsia Menon and Makarkin, 2008

Type species. Nuddsia longiantennata Menon and Makarkin, 2008, housed at the Staatliches Museum für Naturkunde Stuttgart, Germany, under the number SMNS 66000/263.

Nuddsia repatriata Martins-Neto n. sp. (Figures 2A-C, 6A)

Etymology. Referring to the conditions that sample and specimen are repre- sented in Brazilian institution. Holotype. Sample MPSC I - 1889, housed at the Paleontological Museum Figure 1. A. Geological context from Araripe Basin (drawing modified from Vianna from Universidade Regional do Cariri and Neumann, 2002); stratigraphic units and age from Assine (1992); Ponte and Appi (1990) and Ponte and Ponte-Filho (1996). B. General aspect of the quarry and from (URCA), at Santana do Cariri municipal- the limestones beds that furnish the fossil samples. ity, Ceará, Brazil. Type locality. Road between Nova Olin- da and Santana do Cariri, 4 km from the town of Nova Olinda, Ceará State, Brazil. protuberant compound eyes (1.25 mm base, after slightly sinous distally, and Type stratum. Limestone levels, Crato long and 1 mm wide). Prothorax trap- strongly curved at the distal part, reach- Member, Santana Formation. ezoidal, 1.6 mm long (anterior margin ing the apical area after the apex; radial Age. Aptian (late Early Cretaceous). wider than the posterior one). The cell (rb) as long as the subsequent Diagnosis. Fore wing 27 mm long and mesothorax is robust and wider than one; twenty radial cells, heterogeneous; 7 mm wide. The M origin is close to the head and abdomen. Abdomen long (10 hipostigmal cell (h) long and narrow; six wing base; MP origin at the same level mm long) and relatively narrow (2.8 secondary branches, the distal two deeply of the RP origin. CuA reach the anal area mm maximal width, at the distal part multi-branched. The space between RA after the mid length of the wing, with (Figure 2C). Fore wing length 27 mm and RP is relatively wider than both cos- secondary branches relatively long and and 7 mm wide, with narrow costal area, tal and subcostal area. M origin (Figure distally dichotomous. The area between filled by short, perpendicular, some they 2B) at the wing base, forking at the same CuA and anal margin is three times wider Y-shapped, small veinlets, forming a mo- level of the RP origin to the wing base; than the MP/CuA one. saic of heterogeneous cells. ScP slightly MA long, slightly sigmoid at the proxi- Description. Head width twice length curved, distally fused with RA. RP origin mal part, straight at the distal one, with (1.6 mm long and 3.3 mm wide); large close to the wing base, zigzagged at its a long distal dichotomy; MP long, quite

2 Rafael Gioia Martins-Neto and Viviane Zeringóta Rodrigues

multi-branched. MP1 with five long distal secondary branches, distally with small dichotomies; the oblique vein is present (o) MP2+CuA not reaching the mid length of the wing. CuP, short, zigzagged, distally fused to MP2+CuA trougth a cross vein (cua-cup). Anal area with a very small and protuberant anal field. Description. Fore wing 58 mm long and 13 mm wide. Costal area relatively nar- row (little wider than the subcostal area), slightly widening toward the apex, filled by pectinated transverse undichotomous veinlets. ScP distally fused to RA. RP origin very close to the wing base: nine secondary branches, the seven distal ones (adopting RP1 as the most distal Figure 2. Nuddsia repatriata Martins-Neto, n. sp., holotype (MPSC-I-1889). one, will be RP1 to RP7) sigmoid and A. Fore wing; B. Base of the fore wing. C. Body detail of the same specimen. all just distally with a small dichotomy; Scale bar: 5 mm. RP8 deeply dichotomous as well as RP9; RP8 origin very far from RP7, close to parallel to MA, also with a long distal Family PALAEOLEONTIDAE the wing base. MA long, distally curve, dichotomy. CuA reach the anal margin Martins-Neto, 1992 unbranched, undichotomous; MP1 quite after the mid length of the wing and with parallel to MA with six relatively long, at least ten long, transverse secondary Neurastenyx Martins-Neto distally dichotomous secondary branch- branches, with the preserved distal part and Vulcano, 1997 es; MP2+CuA1 relatively short, reaching of at least two dichotomous ones. The the anal area before the mid-length wing, area between CuA and the anal margin is Type species. Neurastenyx araripensis with five transversal secondary branches, three times wider than the MP/CuA one. Martins-Neto and Vulcano, 1997, by three of which distally dichotomous. Discussion. Nuddsia repatriata n. sp. original designation. Species included Oblique vein (o) is conspicuous at the differs from Nuddsia longiantennata Me- also Neurastenyx gigas Martins-Neto and level of the most proximal MP2+CuA1 non and Makarkin (2008) by having a Vulcano, 1997; Neurastenyx polyhymnia secondary branch. CuP very short, re- greater wing length, CuA reaching the Martins-Neto, 1997; Neurastenyx cryptohy- stricted to the wing base, connected to anal margin after the wing mid length men n. comb.; and Neurastenyx conani n. sp. MP2+CuA1 throught two transversal (before in N. longiantennata) and CuA cross veins, one at the point of the be- secondary branches distally dichoto- Neurastenyx conani Martins-Neto n. sp. ginning of the last proximal MP2+CuA1 mous. The area between CuA and the (Figures 3A, 6B) secondary branch and the distal extrem- anal margin is three times wider than ity of o, and another parallel and above it. the MP/CuA one in N. repatriata n. sp. Etymology. Alluding to the anti-hero CuP with four short secondary branches, (as wide as in N. longiantennata). Ad- Conan the Barbarian, created by Robert the most proximal longer and curved. ditionally, N. repatriata n. sp. exhibits E. Howard. A very small and protuberant anal area. two Y-shaped cross veins at the costal Holotype and only specimen. Iso- Conspicuous color pattern is present in and subcostal area, quite all RP second- lated fore wing (CPCA 3575; Figure the whole wing. ary branches multi-branched (the last 6B), housed at Centro de Pesquisas da Discussion. Neurastenyx conani n. sp. two distal ones deeply dichotomous) Chapada do Araripe – CPCA/DNPM, shares with the known Neurastenyx and CuA with at least two secondary Crato municipality, Ceará State, North- species the presence of a conspicuous branches distally dichotomous. N. lon- east Brazil. oblique vein (o), big-sized fore wing, giantennata has not preserved the wing Type locality, type stratum, and age. short CuP, Banksian-line on the RP sec- base, so the M origin is unknown. In N. The same as Nuddsia repatriata n. sp. ondary branches, RP proximal secondary repatriata n., sp. the MP origin occurs at Diagnosis. Big-sized neuropteran, fore branch close to the wing base, and M the level of RP origin (generic diagno- wing with more than 50 mm long and origin at the wing base. sis). The N. repatriata n. sp. abdomen presents a conspicuous color pattern. Fore N. conani n. sp. differs from N. ararip- is at least two times longer than the wing with the first proximal RP second- ensis Martins-Neto and Vulcano, 1997 for N. longiantennata one. ary branch close to the wing base, distally having the oblique vein aligned with the

Gaea - Journal of Geoscience, vol. 6, n. 1, jan/jun 2010, p. 1-8. 3 New neuropteran insects from the Santana Formation, Early Cretaceous NE Brazil most proximal MP2+CuA branch. In N. essential for diagnostic characters, was (2005) however not drawn, a relatively araripensis (Figure 3E) this oblique vein lacking (Ponomarenko, 1992). Other short cubital area and a multi-branched ends after this branch. N. conani n. sp. is form of Baisopardus, the species B. bank- proximal secondary branch of RP, also distinct this aspect is from N. poly- sianus Ponomarenko, 1992, from Russian characteristics that definitively remove hymnia (Figure 3G), which has the oblique Early Cretaceous, was based on two B.cryptoneura from Baisopardus and in- vein ending before the most proximal isolated hind wings, in which is missing cludes it on Neurastenyx. MP2+CuA. Those two species are also the conspicuous oblique vein, the MP is N. conani, the new species above distinct from N. conani by a very nar- clearly not fused to CuA and the CuP described shares with N. cryptoneura n. row space between MA and MP2+CuA that the characteristics is straight (Figure comb. an identical colour pattern (not (medial area), and a reduced number of 2B). All these morphological parameters present in the others known species, like MP2+CuA secondary branches (four in are absent from the Neurastenyx species, Baisopardus banksianus Ponomarenko). N. polyhymnia and six in N. conani n. sp.). indicating that Baisopardus cryptoneura can- However it differs in a set of distinctive The N. gigas Martins-Neto, 1997 differs not belong to that , showing also characteristics: fore wing greater (58 of N. conani for having a notably elongate that the Neurastenyx species cannot be mm long and 13 mm wide in N. conani body and big-sized wing length. synonymized to Baiosopardus as proposed n. sp.), and ScP and RA not proximally by Heads et al. (2005). fused (at least not at the same level as Neurastenix cryptoneura n. comb. Heads et al. (2005) indicate the in N. cryptoneura). If such a kind of fu- Baisoparduas cryptoneura Heads et al., 2005 presence of a subcostal area filled by sion exists in N. conani it could be too several pectinated cross veins (Figure much before of the most RP proximal Heads et al. (2005), when describes 2C), inexistent in the especimen here secondary branch and quite at the same Baisopardus cryiptoneura, from the same described, and that exhibits only a wing level where it occurs in N. cryptoneura, Crato Member, synonymies all the corrugation at this area. For otherwise, apart from others not preserved ones Neurastenyx species to the Russian ge- B. cryptoneura shares with Neurastenyx the in N. cryptoneura as the CuP zigzagged, nus Baisopardus Ponomarenko, based presence of a conspicuous cross vein the very small anal area with a protuber- apparently on an complete material, but on the fore wing, possible to see on the ant anal field and cua-cup connecting in which the basal part of the wings, published photograph of Heads et al. the distal part of CuP at MP2+CuA (however not present on Baisopardus banksianus). These set of distinctive characteristic allow us to segregate N. conani and N. cryptoneura, elevating to five the known species of Neurastenyx. Parapalaeoleon Menon and Makarkin, 2008 (Figure 3D) and Paraneurastenyx Mar- tins-Neto, 1998 (Figure 3F) are also distinct for the discussed Neurastenix species.

Family ARARIPENEURIDAE Mar- tins-Neto, 2002 Subfamily CRATOPTERYXINAE n. subfam.

Type genus. Cratopteryx Martins-Neto and Vulcano, 1989, designated here. Other included genus: Diegopteryx n. gen. Diagnosis. Mid-sized araripeneurid with fore wing length between 14 to Figure 3. A. Neurastenyx conani Martins-Neto n. sp., holotype (CPCA 3575). 25 mm. Antenna long (30 to 40% of B. Baisoptera banksianus Ponomarenko, 1992 (partially reproduced from Ponomarenko, the fore wing length), distally clavated. 1992); C. “Baisoptera” cryptoneura Heads et al., 2005 (partially reproduced from Intraradial cells (cir.) not conspicuously Heads et al., 2005); D. Parapalaeoleon magnus Menon and Makarkin, 2008 (partially reproduced from Menon and Makarkin, 2008); Neurastenyx araripensis Martins-Neto greater than the subsequent one. and Vulcano, 1997 (partially reproduced from Martins-Neto and Vulcano, 1997); Discussion. Cratopteryxinae n. subfam. F. Neurastenyx ascalaphyx Martins-Neto, 1998 (partially reproduced from Martins- differ of the closest subfamily Araripe- Neto, 1998); G. Neurastenysx polyhymnia Martins-Neto, 1997 (partially reproduced neurinae Martins-Neto, 2002 for having from Martins-Neto, 1997). Scale bars: 5 mm. a relatively small intraradial cell (notably

4 Rafael Gioia Martins-Neto and Viviane Zeringóta Rodrigues elongated in Araripeneurinae) allied to long and 6 mm wide, with costal area rela- origin occurs at R, far from MP origin a notably long antenna, smaller than the tively narrow, widening toward apex, filled (at the wing base in both C. robertosan- mesothorax in Araripeneurinae. by several veinlets, several ones Y-shapped, tosi and C. nemopteroides); CuA forked in and the distal ones dichotomous. ScP CuA1 and CuA2, at the same level of Diegopteryx Martins-Neto n. gen. distally fused to RA; ScP base conspicu- MA origin (unforked, straight and paral- ously sigmoid; subcostal area with only lel to MP and MA in both C. robertosantosi Type species. Diegopteryx raptorius one transverse basal cross vein (scp-ra). and C. nemopteroides); CuP transverse, Martins-Neto n. sp., here designated. RP origin close to the wing base; radial distally fused to CuA2 (parallel to CuA in Etymology. Alluding to Diego, from basal cell (rb) very small, smaller than the both C. robertosantosi and C. nemopteroides). Spanish etymology, the chosen, and subsequent ones; second and third radial pteryx, wing. cell the biggest and wider, occupying circa Family PSYCHOPSIDAE Handlirsch, Diagnosis. Hind wing with costal vein- 1/3 of the wing lenght; hipostigmal cell (h) 1906 lets Y-shapped. ScP base sigmoid, distally elongate, wider at its proximal part; ten sec- fused to RA. Radial basal cell (rb) very ondary RP branches, some of them with a Putzneura Martins-Neto gen. n. small, followed by two subsequent cells small dichotomy. MA originates at R, close notably long. MA origin at R, far from to the wing base. MP origin at wing base. Type species. Putzneura parcimoniosa MP origin. CuA forked in CuA1 and CuA forked in CuA1 and CuA2, relatively Martins-Neto n. sp. CuA2, at the same level of MA origin. short, reaching the anal margin at less than Etymology. Latinized from the Ger- CuP transverse, distally fused to CuA2. 1/3 of the wing length. CuP short, distally many putz, adornment, and from putz Discussion. Diegopteryx greatly differs fused to CuA2. or putzgrila, Portuguese colloquial speech from its closest related genus Cratop- Discussion. Diegopteryx raptorius n. sp. from Mooca, district of São Paulo mu- teryx Martins-Neto and Vulcano, 1989, greatly differs from all other close related nicipality, São Paulo State, Brazil, that sharing the relatively long and clavate Araripe neuropterans for having a set of means a big, splendid thing, and neura, antenna (longer than the wing width or autapomorphies in the hind wing mor- from Neuroptera: so, an ornamented and the thorax length), not found in other phology, like the radial basal cell (rb) very great Neuroptera. known Araripe myrmeleontoid genera; small, followed by two subsequent cells Diagnosis. Big-sized psychopsid neu- by having a forked CuA, and CuP distally notably long, greater than those found ropteran (vena triplica present) with fore fused to CuA2; rb notably small and two in Cratopteryx robertosantosi Martins-Neto wing longer than 80 mm; costal veinlets central radial cells very great, occupying and Vulcano, 1989, as well as in C. ne- pectinated just distally dichotomously circa 1/3 of the hind wing length. mopteroides Martins-Neto, 2002. The MA branched; ScP distally fused to RA, and

Diegopteryx raptorius Martins-Neto n. sp. (Figures 4, 7A-B)

Etymology. Alluding to the raptor habit of the forelegs. Holotype and only specimen. Housed at Centro de Pesquisas da Chapada do Araripe – CPCA/DNPM, Crato munici- pality, Ceará State, Northeast Brazil, un- der the number CPCA 3576a and 3576b. Type locality, type stratum, and age. The same from the previous discussed forms. Diagnosis. The main morphology is the same of the genus. Fore wing 25 mm long and 8 mm wide; hind wing 21 mm long and 6 mm wide; antenna 9 mm long, distally clavate. Intensely pubescent legs. Description. Distally clavate, monoliform antenna, 9 mm long. Robust and promi- nent thorax, and legs covered by relatively Figure 4. Diegopteryx raptorius Martins-Neto n. sp., holotype (CPCA 3576). A. Hind long and dense setae. Fore wing 25 mm wing detail. B. Fore wing apex detail; C. Antenna detail; D. Hind leg detail; E. Middle leg detail. Scale bar: 5 mm. long and 8 mm wide. Hind wing 21 mm

Gaea - Journal of Geoscience, vol. 6, n. 1, jan/jun 2010, p. 1-8. 5 New neuropteran insects from the Santana Formation, Early Cretaceous NE Brazil

Figure 5. A-B. Putzneura parcimoniosa Martins-Neto, n. sp., holotype (CPCA 3577): A. distal part of the fore wing detail; B. base of the preserved wing of the same specimen; C. Pulchroptilonia espatifata Martins-Neto, 1997, drawn from the supplementary material RGMN-T-011 (not furnished in Martins- Neto, 1997). Scale bar: 5 mm. costal area distally relatively narrow. RP, MA, and MP secondary branches densely and deeply dichotomous, but not conspicuously pectinated. Intense cross veins pattern, sometimes aligned, absent from the distal part of subcostal area between ScP and RA. Discussion. The Family Psychopsidae Figure 6. A. Nuddsia repatriata Martins-Neto, n. sp., holotype (MPSC-I-1889); has extent representatives in Australia, B. Neurastenyx conani Martins-Neto n. sp., holotype (CPCA 3575). Scale bar: 5 mm. South Africa, and South Asia, apart from more than a dozen fossil genera, especially concentrated in the Mesozoic of Asia. last one also known for the Lower Cre- The first record for a Brazilian fossil The fossil record begins in , taceous of Japan. Arctopsychops Makarkin, genus is Pulchroptilonia Martins-Neto, when it is found in Australia, with Trias- 1994, for Russian one. 1997 (not Pulchroptionia as misspelled by sopsychops Tillyard and Protopsychops Til- For the European is known Jepson et al., 2009), described to the same lyard (New, 1988) and at Virginia, United Apeirophlebia Handlirsch, 1908, Liassopsy- Santana Formation here discussed. The States (Fraser et al. 1996). Angaropsychops chops Bode, 1953 and Archepsychops Til- Putzneura n. gen. here designed greatly Martynova is known for the Upper Juras- lyard, 1919. Other genera are also known differs from Pulchroptilonia Martins-Neto sic of Transbaikalia (Martynova, 1949), for the Baltic amber (McLeod, 1970), and in having ScP distally fused to RA, coast- Baisopsychops Makarkin, 1992, to the Lower for the Asian Miocene (Makarkin, 1991). al area relatively narrow before ScP/RA Cretaceous of Siberia, Embaneura Za- Recently, Jepson et al. (2009) describes fusion, no trace of trichosors, macro or lessky, 1953, Grammopsychops Martynova, four England Lower Cretaceous psychop- microtrichia, and intense cross vein pat- 1954, and Kagapsychops Fujiyama, 1978 to sid genera: Valdipsychops, Cretapsychops, tern, alternating aligned and unaligned the Upper Cretaceous of Kazakhstan, the Micropsychops, and Psichopsites. transverse cross veins rows. Additionally,

6 Rafael Gioia Martins-Neto and Viviane Zeringóta Rodrigues

Holotype. An isolate fore wing (CPCA 3577), housed at Centro de Pesquisas da Chapada do Araripe – CPCA/DNPM, Crato municipality, Ceará State, North- east Brazil. Type locality, type stratum, and age. As for Nuddsia repatriata n. sp.. Diagnosis. As for the genus. Description. Apical and triangular fore wing fragment, 52 mm long and 42 mm wide. ScP distally fused to RA. Vena triplica present. Preserved costal area narrow, filled by undichotomous pectinate until close the ScP/RA fu- sion, slightly longer and with small distal dichotomy after it. RP secondary branch deeply dichotomous, part of them reaching the costal area close to the SCP/RA fusion, others encompass- ing the entire acuminate apex. Intense cross veins pattern, sometimes aligned, but absent at subcostal area, distal part of the area between ScP and RA, at least in the preserved wing.

FINAL COMMENTS

The Psychopsidae is an from a well-successful group that appar- ently does not suffer with the stressing conditions from Cretaceous/Paleogene boundary. They appear at Triassic times in Gondwana areas, with forms of con- troversial affinities, and from what there is no consensus about its real relations with the family. In addition, it diversifies during the Jurassic, mainly in Northern Figure 7. A-B. Diegopteryx raptorius Martins-Neto n. sp., holotype (CPCA 3564), part (A), and counterpart (B), respectively; C. Putzneura parcimoniosa Martins-Neto, Hemisphere and become still more n. sp., holotype (CPCA 3577). Scale bar: 5 mm. abundant in Cretaceous. Like the forms here described, the Psychopsidae-like taxa are characterized Putzneura is probably the biggest known here described consistent and distinct by big-sizes, a profuse, close enervation, Psychopsidae, with a preserved apical of all other known psychopsids, recent conspicuous color patterns, dense covering fragment of the fore wing (52 mm long or , and the second biggest Neu- by trichosors, and macro and microtrichia. and 42 mm wide), that judging the ScP/ roptera (as well as Insecta) from Araripe Ethologically the group is also pecu- RA distal fusion, represent less of the (the first is Makarkinia adamsi Martins- liar due to its extremely wide costal area, the mid-length, so attaining 95-105 mm Neto, 1997). which allows to mimetize an enlarged tho- if complete. Another parameter, the rax while resting. The intense pubescency length/width ratio in complete known Putzneura parcimoniosa Martins-Neto n. sp. and color pattern probably make them wings of the family indicates a rate be- (Figures 5A, 7C) unattractive for their possible predators. tween 2.5 to 3. Taking into account this Other interesting aspect in the pres- parameter the wing length of Putzneura Etymology. Latinized from parcimoni- ence of Psychopsidae at Cretaceous could measure nearly 113 to 127 mm. All ous, allusive to the fragmentation state levels of Araripe Basin is its bigger size those characters make the new species that does not reveal all of its morphology. when compared to those insects known

Gaea - Journal of Geoscience, vol. 6, n. 1, jan/jun 2010, p. 1-8. 7 New neuropteran insects from the Santana Formation, Early Cretaceous NE Brazil at the same time in Northem Hemi- HEADS, S.W.; MARTILL, D.M.; LOVERIDGE, MARTINS-NETO, R.G. 2006. Insetos fósseis sphere. In Brazil they are accompanied R.F. 2005. An exceptionally preserved ant- como bioindicadores em depósitos sedimen- lion (Insecta, Neuroptera) with colour pattern tares: um estudo de caso para o mesozóico sul- by some endemic Neuroptera groups, as preservation from the Cretaceous of Brazil. americano. Revista Brasileira de Zoociências, Makarkiniidae, in an apparent sympatry Palaeontology, 48(6):1409-1417. 8(2):155-183. with , , Myr- JEPSON, J.E.; MAKARKIN, V.N.; JARZEM- MARTINS-NETO, R.G.; RODRIGUES, V.Z. meleontidae, , , BOWSKI, E.A. 2009. New lacewings (Insec- 2009. New Neuroptera (Insecta, Osmylidae ta: Neuroptera) from the Lower Cretaceous and Mesochrysopidae) from the Santana For- Berothidae, and Ithonidae. Wealden supergroup of Southern England. mation, Lower Cretaceous of northeast Bra- Cretaceous Research, 30(2009):1325-1338. zil. Gæa Journal of Geoscience, 5(1):15-20. ACKNOWLEDGEMENTS MAKARKIN, V.N. 1991. Miocene Neuroptera MARTINS-NETO, R.G.; VULCANO, M.A. from the North Caucasus and Sikhote- 1989. Neurópteros (Insecta, Planipennia) da Alin Mts. Paleontologicheskiy Zhurnal, Formação Santana (Cretáceo Inferior) Bacia RGMN thanks Placido Cidade Nu- 1991(1):57-68. do Araripe, Nordeste do Brasil. II - Superfa- vens and Antonio Alamo Feitosa Sa- MARTINS-NETO, R.G. 1991. Sistemática dos mília . Revista Brasileira raiva from Regional do Cariri University Ensifera (Insecta, Orthopteroida) da Forma- de Entomologia, 33(2):367-402. ção Santana, Cretáceo Inferior do Nordeste MARTINS-NETO, R.G.; VULCANO, M.A. (URCA) and Paleontological Museum do Brasil. Acta Geologica Leopoldensia, 1997. Neurópteros (Insecta, Planipennia) from Santana do Cariri, as well as Artur 32(14):3-162. da Formação Santana (Cretáceo Inferior). Andrade and José Betimar Figueira MARTINS-NETO, R.G. 1992. Neurópteros Bacia do Araripe, Nordeste do Brasil. VIII (CPCA/DNPM), by the facilities and (Insecta: Planipennia) da Formação Santana descrição de novos taxa de Myrmeleontidae, (Cretáceo Inferior), Bacia do Araripe, Nor- Ascalaphidae e Nemopteridae. Revista da support given by their institutions. deste do Brasil. VII. Palaeoleontinae, nova Universidade de Guarulhos, Série Ciências Thanks also are due to Andre Nel subfamília de Myrmeleontidae e descrição Biológicas, 2(5):64-81. (MNHN), and Oscar Florencio Gallego de novos táxons. Revista Brasileira de En- McLEOD, E.D. 1970. The Neuroptera of the (UNNE), by suggestions and comments tomologia, 36(4):803-815. Baltic Amber. I. Ascalaphidae, Nymphidae, MARTINS-NETO, R.G. 1994. Neurópteros and Psychopsidae. Psyche, 77(2):147-180. on this paper. Special thanks goes to Gæa (Insecta, Planipennia) da Formação Santana MENON, F.; MAKARKIN, V.N. 2008. New - Journal of Geoscience and its plesiomor- (Cretáceo Inferior), Bacia do Araripe, Nor- fossil lacewings and (Insecta, phic relative, Acta Geologica Leopoldensia, deste do Brasil. IX - Primeiros resultados Neuroptera) from the Lower Cretaceous where some of RGMN works were da composição da fauna e descrição de Crato Formation of Brazil. Palaeontology, novos taxa. Acta Geologica Leopoldensia, 51:149-162. published and that now accompany 39(1):269-288. NEW, T.R. 1988. The Psychopsidae (Insecta: me in a special scientific moment, the MARTINS-NETO, R.G. 1997. Neurópteros Neuroptera) of Australia and the Oriental hundredth published paper. To Tania (Insecta, Planipennia) da Formação San- region. 2(7):841-883 Lindner Dutra, a special friend in this tana (Cretáceo Inferior), Bacia do Araripe, PONOMARENKO, A.G. 1992. Neuroptera (In- Nordeste do Brasil. X - Descrição de novos secta) from the Lower Cretaceous of Trans- trajectory, and to those colleagues who taxa (Chrysopidae, Babinskaiidae, Myrme- baikalia. Paleontological Journal, 3:43-50. have given me support, both in life and leontidae, Ascalaphidae e Psychopsidae). PONTE, F.C.; APPI, C.J. 1990. Proposta de in the “battle” to protect the Araripe fos- Revista da Universidade de Guarulhos, Série revisão da coluna litoestratigráfica da Bacia sils, my deep gratitude. This is a special Ciências Exatas e Tecnológicas, 2(4):68-83. do Araripe. In: CONGRESSO BRASI- MARTINS-NETO, R.G. 1998. Neurópteros LEIRO DE GEOLOGIA, 36, Natal, Anais, moment when I finally have hope that (Insecta, Planipennia) da Formação San- 1:211-226. this important heritage could be studied tana (Cretáceo Inferior), Bacia do Araripe, PONTE, F.C.; PONTE-FILHO, F.C. 1996. by our local students. Nordeste do Brasil. XI - Descrição de novos Estrutura geológica e evolução tectônica táxons de Myrmeleontidae (Palaeoleontinae e da Bacia do Araripe, Brasil. Departamento Pseudonymphinae). Revista da Universidade Nacional da Produção Mineral, Delegacia do REFERENCES Guarulhos, Série Ciências Biológicas e da Ministério de Minas e Energia, Pernambuco Saúde, 3(5):38-42. e Ceará, 68 p. ASSINE, M.L. 1992. Análise estratigráfica da MARTINS- NETO, R.G. 2002. The Santana VIANNA, M.S.S.; NEUMANN, V.H.L.M. 2002. bacia do Araripe, Nordeste do Brasil. Revista Formation paleoentomofauna reviewed. Membro Crato da formação Santana, Chapa- Brasileira de Geociências, 22(3):289-300. Part I - Neuropteroida (Neuroptera and da do Araripe, Ceará: riquíssimo registro da COIMBRA, J.C.; ARAI, M.; CARREÑO, A.L. Raphidioptera): systematic and phylogeny, fauna e flora do Cretáceo. In: C. SCHOBBE- 2002. Biostratigraphy of Lower Cretaceous with description of new taxa. Acta Geologica NHAUS; D.A. CAMPOS; E.T. QUEIROZ; microfossils from the Araripe Basin, north- Leopoldensia, 25(55):35-66. M. WINGE; M.L. BERBERT-BORN (ed.), eastern Brazil. Geobios, 35(6):687-698. MARTINS- NETO, R.G. 2005. New Neuroptera Sítios geológicos e paleontológicos do Brasil. FRASER, N.C.; GRIMALDI, D.A.; OLSEN, (Nymphidae and Araripeneuridae) from San- Brasília, DNPM, p. 113-120. P.E.; AXSMITH, B. 1996. A Triassic Lager- tana Formation (Lower Cretaceous, Araripe stätte from eastern North America. Nature, Basin, northeast Brazil). Gæa Journal of Submitted on: 20/03/2010 380:615-619. Geoscience, 1(1):5-10. Accepted on: 22/06/ 2010

8