Zootaxa 3985 (4): 451–490 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2015 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3985.4.1 http://zoobank.org/urn:lsid:zoobank.org:pub:DAF911DF-B76E-44CA-8355-80A9C9866437 The phylogeny of mole crickets (: : Gryllotalpidae)

OSCAR J. CADENA-CASTAÑEDA Universidad Distrital Francisco José de Caldas, Grupo de Investigación en Artrópodos “Kumangui”, Bogotá, Colombia. E-mail: [email protected]

Table of contents

Abstract ...... 451 Resumen ...... 452 Introduction ...... 452 Material and methods ...... 453 Results and discussion ...... 460 Systematics ...... 466 Gryllotalpidae Leach, 1815 ...... 466 Key to subfamilies, tribes and genera of Gryllotalpidae ...... 467 Scapteriscinae Zeuner, 1939 ...... 468 Indioscaptorini n. trib...... 468 Indioscaptor Nickle, 2003 ...... 468 Scapteriscini Zeuner, 1939 stat. rev...... 468 Scapteriscus Scudder, 1868 ...... 468 Neoscapteriscus n. gen...... 469 Gryllotalpinae Leach, 1815 ...... 469 Triamescaptorini n. trib...... 470 Triamescaptor Tindale, 1928...... 470 Gryllotalpellini n. trib...... 470 Gryllotalpella Rehn, 1917 ...... 471 Key to species ...... 471 Gryllotalpella macilenta (Saussure, 1874) ...... 471 Gryllotalpella minor (Bruner, 1916)...... 471 Gryllotalpella lawrencei n. sp...... 472 Gryllotalpella rehni n. sp...... 473 Gryllotalpella tindalei n. sp...... 473 Gryllotalpini Leach, 1815 stat. rev...... 473 Latreille, 1802 ...... 474 Neocurtillini n. trib...... 474 Kirby, 1906 ...... 474 Key to species ...... 474 Neocurtilla claraziana (Saussure, 1874)...... 475 (Perty, 1832) ...... 475 Neocurtilla ingrischi n. sp...... 475 Neocurtilla townsedi n. sp...... 477 Leptocurtilla n. gen...... 477 Leptocurtilla maranona (Scudder, 1875) n. comb...... 480 Leptocurtilla juanmanueli n. sp ...... 480 Leptocurtilla chopardi n. sp...... 481 Acknowledgements ...... 481 References ...... 482

Abstract

The monophyly and phylogenetic relationships of the family Gryllotalpidae were researched. Twenty-six in-group taxa, representing all known genera of Gryllotalpidae were included in a cladistic analysis, based on 89, morphological charac-

Accepted by D. Rentz: 10 Jun. 2015; published: 14 Jul. 2015 451 ters (including genital characters of the males). The different analyses of the resulting data matrix supported the mono- phyly of Scapteriscinae and Gryllotalpinae and its internal group. Subfamilies, tribes and genera of Gryllotalpidae are fully diagnosed, illustrated and keys to their identification are provided. Four tribes are established (Indioscaptorini n. trib. (Scapteriscinae), Triamescaptorini n. trib., Gryllotalpellini n. trib. and Neocurtillini n. trib. (Gryllotalpinae)) and two other are fully delimited (Scapteriscini stat. rev. and Gryllotalpini stat. rev.). Two new genera are described (Neoscap- teriscus n. gen. and Leptocurtilla n. gen.) and as well as seven new species: Gryllotalpella rehni n. sp., G. tindalei n. sp., G. lawrencei n. sp., Neocurtilla ingrischi n. sp., N. townsendi n. sp., Leptocurtilla juanmanueli n. sp. and L. chopardi n. sp. The following nomenclatural changes were made: All species previously placed in Scapteriscus s.l. are transferred to the new genus Neoscapteriscus, except Scapteriscus oxydactilus and S. headsi that are still placed in Scapteriscus, Gryl- lotalpa chilensis reinst. stat. and Leptocurtilla maranona, n. comb. Finally, previous contributions about the phylogenetic relationships of molecrickets are contrasted with the results of this research.

Key words: Scapteriscinae, Gryllotalpinae, Neoscapteriscus, Leptocurtilla, new classification

Resumen

La monofilia y las relaciones filogenéticas de la familia Gryllotalpidae, han sido investigadas. Veinte seis taxones son in- cluidos en el grupo interno, con representantes de todos los géneros de Gryllotalpidae, el análisis se basó en 89 caracteres morfológicos (incluyendo caracteres genitales de los machos). Los diferentes análisis soportan la monofilia de Scapterisci- nae y Gryllotalpinae, incluyendo sus grupos internos. Se realizan y complementan diagnosis para subfamilias, tribus y gé- neros, ilustraciones de los taxones y claves para sus identificaciones son proporcionadas. Cuatro tribus han sido propuestas (Indioscaptorini n. trib., Triamescaptorini n. trib., Gryllotalpellini n. trib. y Neocurtillini n. trib.) y las otras tribus han sido delimitadas (Scapteriscini stat. rev. y Gryllotalpini stat. rev.). Se describen dos nuevos géneros (Neoscapteriscus n. gen. y Leptocurtilla n. gen.) y siete especies nuevas: Gryllotalpella rehni n. sp., G. tindalei n. sp., G. lawrencei n. sp., Neocurtilla ingrischi n. sp., N. townsendi n. sp., Leptocurtilla juanmanueli n. sp. y L. chopardi n. sp. Los siguientes cam- bios nomenclaturales se realizaron: Todas las especies ubicadas previamente en Scapteriscus s. l. se transfieren al nuevo género Neoscapteriscus a excepción de Scapteriscus oxydactylus y S. headsi que aún se ubican en Scapteriscus, Gryllo- talpa chilensis reinst. stat. y Leptocurtilla maranona n. comb. Finalmente se discute las contribuciones previas sobre las relaciones filogenéticas de los grillos topo y se contrastan con los resultados de esta investigación.

Palabras clave: Scapteriscinae, Gryllotalpinae, Neoscapteriscus, Leptocurtilla, nueva clasificación

Introduction

The mole family Gryllotalpidae (Orthoptera: : Gryllotalpoidea) is a monophyletic group (Zeuner, 1939; Sharov, 1968; Desutter-Grandcolas, 2003; Jost & Shaw, 2006) of the clade, (Desutter, 1990) which contains more than 100 species in three subfamilies (two recent and one fossil), 11 genera (six recent and five fossil) (Eades et al., 2015). Mole crickets are distributed mostly over tropical and some temperate regions. The fossil genera recorded in the group are known from the Cretaceous (Martins-Neto, 1991; Perrichot et. al., 2002). Members of family Gryllotalpidae are one of the most distinctive groups in the order Orthoptera, characterized by having the first pair of legs quite clearly adapted to digging (Leach, 1815; Chopard, 1949), tumescent pronotum, antennae which are shorter than the head and pronotum together and hind legs only little specialized for jumping, features developed for fossorial life throughout their evolutionary history (Cadena-Castañeda, 2011). Research in this group has been mostly taxonomical and based on external morphology; some of the characters tend to have phenotypical plasticity if they are not taken in a measurable way (Nickle, 2003; Cadena-Castañeda, 2015). Taxonomical history of the group begins with the description of by Linnaeus (1758), although back then it was described as Gryllus (Acheta) gryllotalpa, indicating that it was related to common crickets. Later, Latreille (1802), described the genus Gryllotalpa and Leach (1815) officially created the family. Description of the remaining genera were made by the following authors: Scudder in 1868 described Scapteriscus (two-dactyls ), making a distinction of mole crickets with two or four dactyls. Kirby (1906), proposed Neocurtilla as a new name for Curtilla, with N. hexadactyla as type species. Likewise, he proposed S. oxydactylus for Scapteriscus as a type species, being it noteworthy to mention that Kirby did not acknowledge recognize the name Gryllotalpa, and kept most species within genus Curtilla. By 1917, Rehn

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