New Orthoptera Stenopelmatoidea and Hagloidea (Ensifera) from the Santana Formation (Lower Cretaceous, Northeast Brazil) with Description of New Taxa

Home , Caelifera, Ensifera, Gryllotalpoidea, Prophalangopsidae, Stenopelmatoidea, Welwitschiaceae

New Orthoptera Stenopelmatoidea and Hagloidea (Ensifera) from the Santana Formation (Lower Cretaceous, Northeast Brazil) with description of new taxa

Rafael Gioia Martins-Neto Sociedade Brasileira de Paleoartropodologia – SBPr. Centro de Ensino Superior de Juiz de Fora – CES/JF. Programa de Pós-graduação em Ecologia, Universidade Federal de Juiz de Fora – UFJF, Campus Universitário Martelos, 36036-900, Juiz de Fora, MG, Brazil. [email protected]

ABSTRACT This work presents the results of a taxonomic study carried out on three specimens of fossil Orthoptera from the Crato Member laminated limestone, lowest unit of the Santana Formation (Lower Cretaceous), Araripe Basin. All fossils were found in Pedra Branca Quarry, near Santana do Cariri and Nova Olinda municipalities in Ceará State, northeast Brazil. The following new taxa are proposed: Brauckmannia groeningae Martins-Neto, n. gen. et n. sp. (Brauckmanniidae Martins-Neto, n. fam.), Euclydes ramosfernandesi n. gen. et n. sp. (Mimnermidae: Euclydesinae Martins-Neto, n. subfam.), and Prezottophlebia helbae Martins-Neto, n. gen. et n. sp. (Hagloidea: Prezzottophlebiidae Martins-Neto, n. fam.). The meaning of their morphological characters (robustness and body shape) and their presence in the laminated limestone is discussed like a contribution to the past distribution of Ensifera and to the paleoecological inferences.

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

RESUMO Este trabalho apresenta o estudo taxonômico de três espécimes de ortópteros fósseis, provenientes dos níveis de calcários laminados do Membro Crato, unidade inferior da Formação Santana (Cretáceo Inferior), bacia do Araripe. Os fósseis provêm do afloramento da Mina Pedra Branca, nas proximidades dos municípios de Santana do Cariri e Nova Olinda, no Estado do Ceará, nordeste do Brasil. Os seguintes novos táxons são propostos: Brauckmannia groeningae Martins-Neto, n. gen. et n. sp. (Brauckmanniidae Martins-Neto, n. fam.), Euclydes ramosfernandesi n. gen. et n. sp. (Mimnermidae: Euclydesinae Martins-Neto, n. subfam.) e Prezottophlebia helbae Martins-Neto, n. gen. et n. sp. (Hagloidea: Prezzottophlebiidae Martins-Neto, n. fam.). O significado dos caracteres morfológicos (a forma do corpo e sua robustez) e de sua presença entre os níveis de calcários laminados é discutido como uma contribuição para a distribuição pretérita de Ensifera e para a paleoecologia do grupo.

Palavras-chave: Ensifera, insetos, Formação Santana, bacia do Araripe, Cretáceo Inferior, Brasil.

INTRODUCTION MATERIAL AND METHODS Gorochov (1995). Abbreviations cited in the text: MA, anterior media; R, radial; RA, an- The Orthoptera and Neuroptera The material consists of three selected terior radial; RP, posterior radial; ScA, ante- are the most common orders of insects slabs collected in Pedra Branca Quarry, rior subcosta; ScP, posterior subcosta. from the Araripe Basin in terms of located at Nova Olinda-Santana do Cariri collected specimens and number of road, 4 km from the municipal district of SYSTEMATIC PALEONTOLOGY named species (Martins-Neto, 1991, 1995, Nova Olinda and was originated from the 2006). Orthopterans are mainly laminated limestone from the Crato Order ORTHOPTERA Olivier, 1789 represented by Grylloidea, among the Member, lowermost unit of the Santana Suborder ENSIFERA Chopard, 1920 Ensifera, and by Locustopsoidea, among Formation, Araripe Basin (Ponte and Superfamily STENOPELMATOIDEA the Caelifera. Some groups are especially Appi, 1990), considered Upper Aptian in Burmeister, 1838 rare such as Hagloidea, Tettigonioidea, age (Ponte and Ponte-Filho, 1996; and Gryllotalpoidea. The present study is Coimbra et al., 2002). Brauckmanniidae, n. family a contribution and addition to past The adopted terminology is based in research made by the author, and reveals Martins-Neto (1991) and the taxonomic Diagnosis. Big sized ensiferan, new forms of orthopterans. insertions follows Martins-Neto (1991) and macropterous, stunt, body length around

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40 mm. All tibia robust, smaller than Brauckmannia groeningae n. sp. IV sickle-like, distally curved, without ter- femora and tarsi longer than tibia. Fore, (Figure 1) minal claws, and intensely pubescent. mid and hind tarsi four segmented and all Fore, mid and hind tarsi four segmented articles with a notably developed pulvilli. Etymology. In honor to Elke Gröning and all articles with a notably developed Segment IV of all tarsi modified in a stickle- (Clausthal Institute, Germany). pulvilli. No evidence of spines or apical like structure, without terminal claws. Holotype and only specimen. RGMN- spurs in the tibia. No evidence of cerci. Discussion. Within Stenopelmatoidea, 500, housed at Sociedade Brasileira de Wings poorly preserved, longer than the only Cooloolinae Rentz (Mimnermidae Paleoartropodologia (Rua Arnaldo body. Those characters make Brauckmannia Brunner-Wattenwyl) shares some Vitaliano, 150, 14091-220, Ribeirão Preto, groeningae n. sp. one of the biggest known apomorphic characteristics with SP, Brazil). Cretaceous orthopteran. Brauckmanniidae, such as the modification Type locality. Pedra Branca Quarry, Nova of the fore legs (tibia short and robust, Olinda-Santana do Cariri road, 4 km from Family MIMNERMIDAE Brunner- tarsi four-segmented) and body stunt. the municipal district of Nova Olinda, Wattenwyl, 1888 Brauckmanniidae n. fam. differs from all Ceará State, Brazil. Euclydesinae, n. subfamily known Stenopelmatoidea in having all Type stratum. Laminated limestone level, three pair of legs modified (tibia shorter Crato Member, lowermost unit of the Diagnosis. Big sized ensiferans with than femur, tarsi longer than tibia and with Santana Formation, Araripe Basin. body length around 23 mm. Females with well-developed pulvilli). Age. Aptian, Lower Cretaceous. long setiform ovipositor, longer than the Diagnosis. As for the genus. body. Fore wing with wide costal area Brauckmannia n. gen. Description. Big sized orthopteran, toward the apex, narrow at the wing base; macropterous, stunt, with body length ScA short and ScP reaching the apical area; Type species. Brauckmannia groeningae, n. around 40 mm, probably male (no RA unbranched; RP three-branched and sp., designated here. evidence of ovipositor preserved). Head MA two-branched; RP origin close to the Etymology. In honour to Carsten and twice wider than long, 9 mm wide and 4 wing base; conspicuous ma-mp. Hind Brigitte Brauckmann (Clausthal University, mm long. Pronotum seliform little wider wing with not well-developed anal fan and Germany). and shorter than the head. Fore femur costal area homogeneously narrow. Diagnosis. As for the family, with tibia (Figure 1B.1), 12.6 mm long and 2.6 mm Discussion. Euclydesinae n. subfam. is without spines or apical spurs. wide, with 0.20 of robustness index similar to the Zeuneropterinae Kevan & Discussion. The most similar known genus (RI=width/length). Fore tibia 6.6 mm Wighton, in the general wing morphology, is Cooloola Rentz (Cooloolinae) in having long and 2.4 mm wide (RI = 0.36), and specially in having MA two-branched and the fore and hind pair of legs modified. no evidence of tympanums. Mid tibia the presence of ma-mp. Euclydesinae n. However, Brauckmannia n. gen. greatly (Figure 1B.2), 7.1 mm long and 3 mm subfam. differs from Zeuneropterinae in differs in having also the mid pair of legs wide (RI = 0.42). Mid tarsus length, 9.6 having ScA short (relatively long in modified, longer tarsal elements (short in mm. Hind femur 25 mm long and 6.5 Zeuneropterinae), ScP long, reaching the Cooloola), macropterous (micropterous in mm wide (RI = 0.26). Hind tibia (Figure apical area (the costal margin circa 1/3 of Cooloola), and virtually no spines, apical spurs 1B.3) 13.8 mm long and 3 mm wide (RI the wing apex), RA unbranched (multi- and distal claws (present in Cooloola). = 0.21). Hind tarsus 14.5 mm with digit branched in Zeuneropterinae).

Figure 1. Brauckmannia groeningae Martins-Neto, n. sp. A. Holotype; B. 1, fore leg; 2, mid leg; 3, hind leg. Scale bar: 5 mm.

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Additionally Euclydesinae n. subfam. to the mid length of wing (close to the abdomen as wide as the thorax. exhibits a notably distally wide costal area wing base in Euclydes n. gen.). Kevania Ovipositor long, lance-like, 33 mm long. (narrow in Zeuneropteridae). The new Martins-Neto, originally interpreted as Fore wing 32 mm long and 13 mm wide subfamily differs also from other Prophalangopsinae (Martins-Neto, 1991), (Figure 2B.1), after pushing back (the Mimnermidae subfamilies by its notably and later as Hagloidea incertae sedis costal area is preserved and folded along long and setiform ovipositor, normally (Gorochov, 1995), is now transferred to ScP). Costal area very narrow at the wing shorter and blade-like in the known the Euclydesinae n. subfam., within base and widening toward the apex, filled subfamilies. Mimnermidae. Another genus, by several cross veins. ScA short, close to Cratohaglopsis Martins-Neto (1991), also the wing base, and ScP long, sinuous, Euclydes n. gen. described for the Crato Member, reaching the apex. RA sinuous and although poorly preserved, also exhibits unbranched, parallel to ScP. RP origin Type species. Euclydes ramosfernandesi, n. a lance-like ovipositor and wing venation close to the wing base, originating three sp., designated here. pattern, which could permit its inclusion long secondary branches: RP1 divergent Etymology. Generic and specific names in this new subfamily. However Euclydes of RP2, and RP3 parallel to RP2. MA in memoriam of the Brazilian naturalist, n. gen. differs from Cratohaglopsis Martins- origin at the wing base. MA1 curved, Euclydes Ramos Fernandes. Neto, 1991 by having a hind wing with convergent to the RP3 extremity. MP Diagnosis. As for the subfamily. notably narrower radial sector area, RA apparently two-branched and ma-mp Discussion. The closest known without secondary branches (three in conspicuous at the wing base. Several Zeuneropterinae genera, Zeuneroptera Cratohaglopsis) and RP with only one cross-veins in the whole wing. Hind wing Kevan and Wighton and Albertoilus secondary branch (six in Cratohaglopsis). with a narrow and homogeneous costal Kevan and Wighton, from the Canadian area (Figure 2B.2). ScA and ScP as for Paleocene (Kevan and Wighton, 1981), Euclydes ramosfernandesi, n. sp. the fore wing, but ScP reaching the apical originally placed in Prophalangopsidae, (Figure 2) area boundary. RA unbranched and RP were subsequently placed in the apparently two-branched. MA two- Mimnermidae by Gorochov (1995). Holotype and only specimen. RGMN- branched and MA1 and MA2 long and Euclydes n. gen. greatly differs from both 501, housed at Sociedade Brasileira de distally convergent. mentioned genera, as previously Paleoartropodologia (SBPr). discussed (see subfamily discussion). A Type locality, type stratum, and age. The Superfamily HAGLOIDEA third genus, Kevania Martins-Neto, 1991, same for Brauckmannia groeningae n. sp. Handlirsch, 1906 described from the same sediments from Description. Body stunt, 23 mm long, Family Prezottophlebiidae, n. fam. the Araripe Basin, exhibits a very similar proventriculum preserved (Figure 2B.3). hind wing but differs in having RA and Big and prominent eyes. Pronotum Diagnosis. Fore wing with costal RP multi-branched and RP origin close trapezoidal, shorter than the head, and margin narrow and radial area wide. RA

Figure 2. Euclydes ramosfernandesi Martins-Neto n. sp. A. Holotype; B. 1, fore wing; 2, hind wing; 3, body detail. Scale bar: 5 mm.

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distal branches long, branching at the branching of RP at mid length, close to Prezotophlebia Martins-Neto, n. gen. wing mid length. Presence of intercalary the apex in the other families, presence veins. of intercalary veins that is absent in the Etymology. In honor to the Brazilian Discussion. Differs from all known other families, and by a wide radial field, ethologist Fábio Prezoto. Hagloidea families by the unique location normally narrow and homogeneous in Type species. Prezotophlebia helbae, n. sp., of R branching of RA and of secondary the other ones. designated here.

Figure 3. Prezottophlebia helbae Martins-Neto, n. sp. A. Holotype; B. Wing detail; C. 1, head leg; 2, fore leg; 3, mid leg; 4, detail of abdomen; 5, fore wing. Scale bar: 5 mm.

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Diagnosis. As for the family. Martins-Neto (1991) attributed two FINAL COMMENTS new species Kevania araripensis and Prezotophlebia helbae, n. sp. Cratohaglopsis santanaensis to the The insect fauna described until the (Figure 3) Hagloidea, both transferred here to the moment to Crato Member, Santana Stenopelmatoidea. Furthermore, Formation, are revealing an unexpected Etymology. In honor to the ethologist illustrates another specimen (Martins- diversity in the Lower Cretaceous times to Helba Helena Santos Prezoto. Neto, 1995, fig. 64) that also was attributed northeast Brazil, presently dominated by Holotype. RGMN-511, housed at Socieda- to Hagloidea. This form, represented by Orthoptera (Ensifera and Caelifera), as well de Brasileira de Paleoartropodologia (SBPr). a bizarre neuropteran, is assigned to the as Neuroptera. The present contribution Type locality, type stratum and age. genus Rafaelia by Nel et al. (2005), recently reveals that this dominance also includes As for Brauckmannia groeningae n. sp. renamed to Rafaeliana Nel et al. (2006) and bizarre forms of Stenopelmatoidea Description. Mid-sized Orthoptera, body motivates the creation of a new family of members and new elements belonging to length 28 mm, macropterous, with neuropterids, Rafaelidae (Nel et al. 2005). poorly know groups, such as relatively small triangular head (Figures 3A Finally, a wing fragment described as Gryllotalpoidea (to be described). Another and 3C.1). Pronotum trapezoidal, larger Phasmomimella? araripensis (Martins-Neto, interesting aspect regards to the use of than the head. Abdomen stunt, with the 1991) shows to be very similar to the apical those insects in paleoecology, showing last four segments compressed (Figure area of the fore wing from Rafaeliana and groups morphologically as distinct as 3C.4). Fore leg relatively small (Figure 3C.2), could be also assigned to this genus. Euclydesinae and cearagryllyds exhibiting with femur 10 mm long and 1 mm wide, As a result, the Hagloidea is only similar habitus, expressed in the ovipositor tibia 6 mm long and 1 mm wide, intensely represented now in Araripe basin by the shape and length (notably long and pubescent and with just one small tarsal new genus and specie Prezotophlebia helbae narrow), body shape and robustness. This segment preserved. Mid leg (Figure 3C.3), described here. On the other hand, this could represent a strategy in relation to the 12 mm long and 2 mm wide, with 7 mm family and other related forms strongly occupation of the ecological niche. Like was long and 1.5 mm wide, smooth tibia and dominate over Grylloidea in the Asian suggested also in previous works, under only two small tarsal segments preserved. Lower Cretaceous (Gorochov, 2001a). For the soil lives a diverse community of Hind femur 20 mm long and 4 mm wide the England Lower Cretaceous, the burrowing crickets (Gryllotalpoids and (RI = 0.20) with a smooth tibia. Fore wing frequency of these taxa is more or less closely related species) and on the leaves, a 40 mm long and 6 mm wide (Figures 3B intermediate (Gorochov, 2001b), and as diverse community of grasshoppers and 3C.5), costal area narrow, filled by discussed here, rare in the Brazilian Lower (especially Locustopsidae). pectinate cross veins, with ScP running Cretaceous. directed toward to the costal area, RA This clearly reflects geographical and/ ACKNOWLEDGEMENTS origin about 1/3 of the wing base, or ecological differences, as first pointed sinuous with two sigmoid secondary by Gorochov (2001b). For this author Thanks are due to Federica Menon branches, distally dichotomous. Area this could be due to a humid tropical (Manchester) for the English corrections, between ScP and RA with intercalary climate present in Brazil, in opposition comments and suggestions, and to branches. RP slightly sigmoid, distally to the drier temperate climate existing in Carsten and Brigitte Brauckmann multibranched, and with at least three Siberia-Mongolia, and intermediate (Clausthal University, Germany), by the secondary branches preserved. MA two- within these extremes in England. collaboration and friendship. This branched, distally dichotomous and with Nevertheless, a hot, dry, tending to arid research was supported by the Centro de at least one intercalary vein preserved. climate has been widely suggested in the Ensino Superior de Juiz de Fora – CES/ Unbranched MP, parallel to the MA literature during the Crato´s time at JF, Grant 228.18.08.2006. branches. CuA three-branched. Dense Araripe Basin (Lima, 1983; see also cross-vein pattern. Martins-Neto, 2006 to a revision). The REFERENCES new data here presented gives support to REVIEW OF THE HAGLOIDEA the more recent studies made with the CARVALHO, I. 2000. Geological AND PHASMOMIMOIDEA other elements of the Early Cretaceous environments of dinosaur footprints in FROM THE BRAZILIAN life in the basin, that proposed a hot, but the intracratonic basins of northeast LOWER CRETACEOUS not so arid climate, to this time in the Brazil during the Early Cretaceous interior areas of northeast Brazil (Carva- opening of the South Atlantic. Cretaceous Recent studies are given a great lho, 2005; Dilcher et al., 2005) or at least, Research, 21:255-267 contribution in the knowledge of the maintain some dubiety about the dry COIMBRA, J.C.; ARAI, M. and diversity and taxonomic insertion of the conditions in the surrounding CARREÑO, A.L. 2002. Biostratigraphy Lower Cretaceous insects, mainly in environments of the Araripe paleolake of Lower Cretaceous microfossils from northeast Brazil. (Kunzmann et al., 2006). the Araripe Basin, northeast Brazil.

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Geobios, 35(6):687-698. WILDE, W. 2006. Gymnosperms from the new extraordinary neuropterid family DILCHER, D.L.D.; BERNARDES-DE- Early Cretaceous Crato Formation (Brazil). from the Lower Cretaceous Crato OLIVEIRA, M.E.; PONS, D. and II. Cheirolepidiaceae, Fossil Record - Formation of Brazil: a new insect order? LOTT, T.A. 2005. Welwitschiaceae from Mitteilungen aus dem Museum für (Insecta, Neuropterida). Cretaceous the Lower Cretaceous of northeastern Naturkunde, 9:213-225. Research, 26:845-852. Brazil. American Journal of Botany, LIMA, M. 1983. Paleoclimatic reconstruction NEL, A.; GARROUSTE, R.; BECHLY, G.; 92:1294-1310. of the Brazilian Cretaceous based on POHL, B. and ESCUILLIÉ, F. 2006. GOROCHOV, A.V. 1995. System and palynological data. Revista Brasileira de Rafaeliana, a replacement generic name evolution of the Suborder Ensifera Geociências, 13:223-228. for Rafaelia Nel et al., 2005 (Orthoptera). Proceedings of the Zoological MARTINS-NETO, R.G. 1991. Sistemáti- (Neuropterida). Brèves communications Institute, 260(I-II):1-223, 1-212. ca dos Ensifera (Insecta, Orthopteroida) Bulletin de la Société Entomologique de GOROCHOV, A.V. 2001a. The most da Formação Santana, Cretáceo Inferior France, 111(2):190. interesting finds of orthopteroid insects do Nordeste do Brasil Acta Geologica PONTE, F.C. and APPI, C.J. 1990. Propos- at the end of the 20th century and a new Leopoldensia, 32(14):3-162. ta de revisão da coluna litoestratigráfica recent genus and species. Journal of MARTINS-NETO, R.G. 1995. Comple- da Bacia do Araripe. In: CONGRESSO Orthoptera Research, 10(2):353-367. mentos ao estudo sobre os Ensifera BRASILEIRO DE GEOLOGIA, 36, GOROCHOV, A.V. 2001b. Preliminary (Insecta, Orthopteroidea) da Formação Natal, Anais, 1:211-226. notes on the history of South American Santana, Cretáceo Inferior do Nordeste PONTE, F.C. and PONTE-FILHO, F.C. Ensifera (Orthoptera). Acta Geologica do Brasil. Revista Brasileira de 1996. Estrutura geológica e evolução Leopoldensia, 24(52/53):81-86. Entomologia, 39(2):321-345. tectônica da Bacia do Araripe, Brasil. KEVAN, D.E. McE. and WIGHTON, D.C. MARTINS-NETO, R.G. 2006. Insetos fós- Departamento Nacional da Produção 1981. Paleocene orthopteroids from seis como bioindicadores em depósitos Mineral, 4º e 10º Distritos Regionais. South Central Alberta, Canada. Canadian sedimentares: um estudo de caso para o Delegacias do MME em Pernambuco e Journal of Earth Sciences, 18(12):1824- mesozóico sul-americano. Revista Bra- Ceará, 68 p. 1837. sileira de Zoociências, 8(2):155-183. KUNZMANN, B.; MOHR, A.R.; BER- NEL, A.; BECHLY, G.; GARROUSTE, R.; Submetido em 27/02/07 NARDES-DE-OLIVEIRA, M.E. and POHL, B. and ESCUILLIÉ, F. 2005. A Aceito em 15/05/07

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