New species of the Mesochrysopidae (Insecta, Neuroptera) from the Crato Formation of Brazil (Lower Cretaceous), with taxonomic treatment of the family

doi:10.1016/j.cretres.2005.05.009

Cretaceous Research

Volume 26, Issue 5, October 2005, Pages 801-812

https://doi.org/10.1016/j.cretres.2005.05.009 Get rights and content

Abstract

Two new species of the genus Karenina Martins-Neto belonging to the Mesozoic family Mesochrysopidae (K. leilana sp. nov. and K. longicollis sp. nov.) are described from the Nova Olinda Member, the lowest unit of the Lower Cretaceous Crato Formation, northeast Brazil. A detailed definition (diagnosis and description) of this family is provided, based on those genera most similar to the type genus, Mesochrysopa Handlirsch. The genera Tachinymphes Ponomarenko, Siniphes Ren and Yin, Allopterus Zhang, Karenina Martins-Neto and Mesascalaphus Ren et al. are added to it, whereas the genera Osmylites Haase (= Nymphoides Panfilov), Chrysoleonites Martynov, Microsmylus Panfilov and Liassochrysa Ansorge and Schlüter are excluded. Allopteridae is regarded as a synonym of Mesochrysopidae. The phylogenetic position of the family is discussed. The occurrence of Mesochrysopidae in South America demonstrates that its widespread distribution included Gondwana.

Introduction

The Mesochrysopidae is an extinct Mesozoic family whose status and composition were unclear hitherto. Unfortunately, this is a common feature of almost every extinct higher taxon of Neuroptera, and characterises the generally unresolved state of the systematics of the order. A more-or-less detailed definition of this family has not been provided previously; the type genus has not been re-examined since the classic work of Handlirsch (1906–1908). Therefore, the opinion of some authors on the heterogeneity (paraphyly) of this group was quite reasonable (Willmann and Brooks, 1991, Nel and Henrotay, 1994, Makarkin, 1997). Until now, it was treated either as a separate family (e.g., Adams, 1956, Martynova, 1962, Makarkin, 1990, Makarkin, 1997, Carpenter, 1992, Makarkin and Archibald, 2003, Ponomarenko, 2003) or as a subfamily of the Chrysopidae (e.g., Adams, 1967, Schlüter, 1982, Schlüter, 1984, Séméria and Nel, 1990, Martins-Neto, 2000, Martins-Neto, 2003).

The family was erected by Handlirsch (1906–1908) for the two monotypic genera, Mesochrysopa Handlirsch, 1906 and Mesotermes Haase, 1890 from the Upper Jurassic of Solnhofen, Germany. Handlirsch also assumed that Pseudomyrmeleon Handlirsch, 1906, represented by one poorly preserved single specimen, also from Solnhofen, may belong to this family. Martynov (1927) described another Upper Jurassic genus, Mesypochrysa Martynov, 1927, from the southern Kazakhstan locality of Karatau, and placed it in the Mesochrysopidae. Later, Panfilov (1980) assigned five genera to it, also described from Karatau (Chrysoleonites Martynov, 1925, Aristenymphes Panfilov, 1980, Macronympha Panfilov, 1980, Microsmylus Panfilov, 1980 and Nymphoides Panfilov, 1980). Ansorge and Schlüter (1990) added the new genus Liassochrysa Ansorge and Schlüter, 1990 from the Lower Jurassic of Dobbertin, Germany, and Nel and Henrotay (1994) described Protoaristenymphes Nel and Henrotay, 1994 from the Lower Jurassic of Luxembourg. Recently, Ponomarenko (2003) synonymized Nymphoides and Osmylites Haase, 1890, and considered Osmylitidae to be a synonym of Mesochrysopidae.

In this paper we propose a detailed definition of the Mesochrysopidae as a monophyletic group based on those genera most similar to the type genus Mesochrysopa Handlirsch, 1906, excluding from it some genera, namely Osmylites (= Nymphoides), Chrysoleonites, Microsmylus, and Liassochrysa, and adding others: Tachinymphes Ponomarenko, 1992, Siniphes Ren and Yin, 2002, Allopterus Zhang, 1991, Karenina Martins-Neto, 1997, and Mesascalaphus Ren et al., 1995. The family Allopteridae is regarded as a synonym of Mesochrysopidae, whereas placing Osmylitidae and Mesochrysopidae in synonymy is not justified.

The family was hitherto known only from the Mesozoic of Eurasia. Our study shows, however, that it also occurs in the Lower Cretaceous of South America, being represented in the Brazilian Crato Formation by the genus Karenina with three species, two of which are new. Descriptions of the latter are provided below.

Section snippets

Material and methods

We examined two specimens for this study, found in one of the small quarries or stone yards in the Nova Olinda municipality; the precise locality is not known. Preparation was carried out using an aeroneedle (Selden, 2003) to remove minor amounts of matrix obscuring portions of the fossils. Drawings were made with a camera lucida attached to an Olympus SZH stereomicroscope, and digital photographs were taken with a Sony DCS-717 camera at 2560 × 1920 pixel resolution or a D1X digital camera

Stratigraphy and depositional setting

The Crato Formation is a local stratigraphic unit of the Brazilian non-marine Cretaceous, extending over the Araripe sedimentary basin (Araripe Plateau), in the states of Cearà, Pernambuco and Piauí, northeast Brazil, about 7° south of the Equator. The most important outcrops are in the eastern part of the Araripe Plateau, especially near the towns of Crato, Santana do Cariri and Nova Olinda (see fig. 1 in Martins-Neto, 1992). The stratigraphy of the entire Araripe Basin, and of the Crato unit

Systematic palaeontology

Order: Neuroptera Linnaeus, 1758

Family: Mesochrysopidae Handlirsch, 1906 (syn. Allopteridae Zhang, 1991, syn. nov.)

Diagnosis. Medium-sized to large, graceful neuropterans of somewhat myrmeleontoid appearance, with relatively narrow wings and body. The following combination of forewing character states is diagnostic: (1) trichosors absent (apomorphy, shared with several other families); (2) costal space narrow (apomorphy, shared with several other families); (3) not or only slightly expanded

Discussion

The genus Karenina from the Crato Formation was previously assigned to Ascalaphidae (Martins-Neto, 1997). Indeed, the mesochrysopid affinity of the type species (K. breviptera) is not obvious, based on the description of the holotype, because of incomplete preservation. The two new species described here provide character states, particularly of venation of the fore and hindwings, that clearly confirm the mesochrysopid affinity of this genus. The forewing venation of Karenina is most similar to

Acknowledgements

We thank Dr. David M. Martill (University of Portsmouth) for allowing us to study the specimens; Prof. Rafael Gioia Martins-Neto (Sociedade Brasileira de Paleoartropodologia, Ribeirao Preto, Brazil) for providing the photograph of the holotype of Karenina breviptera; Christian Neumann (Museum für Naturkunde, Humboldt-Universität zu Berlin) for providing the photograph of the holotype of Liassochrysa stigmatica; Dr. Jörg Ansorge (Ernst-Moritz-Arndt Universität, Greifswald) and Dr. Günter Bechly

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nov. and Myrmeleontoidea. However, Ponomarenko (1992) and Makarkin and Menon (2005) suggested that the chrysopoid lineage (Mesochrysopidae and Chrysopidae) and myrmeleontoid lineage might have had a common ancestor, and form a monophylum. So, there is a probability that this condition may be a synapomorphy of the two lineages.
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