Systematics of New World Curtonotum Macquart (Diptera: Curtonotidae)

Home , Acalyptratae, Curtonotidae, Curtonotum, Diastatidae

ZOOTAXA

3079

JOHN KLYMKO1 & STEPHEN A. MARSHALL2 1Atlantic Canada Conservation Data Centre, P.O. Box 6416, Sackville, New Brunswick, Canada E4L 1G6. E-mail: [email protected] 2School of Environmental Sciences, University of Guelph, Guelph, Ontario, Canada N1G 2W1. E-mail: [email protected]

Magnolia Press Auckland, New Zealand

Accepted by D. Bickel: 4 Aug. 2011; published: 28 Oct. 2011 JOHN KLYMKO & STEPHEN A. MARSHALL Systematics of New World Curtonotum Macquart (Diptera: Curtonotidae) (Zootaxa 3079) 110 pp.; 30 cm. 28 Oct. 2011 ISBN 978-1-86977-801-9 (paperback) ISBN 978-1-86977-802-6 (Online edition)

FIRST PUBLISHED IN 2011 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

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ISSN 1175-5326 (Print edition) ISSN 1175-5334 (Online edition)

Abstract ...... 4 Introduction ...... 4 Phylogenetic relationships...... 5 Material and methods ...... 6 Phylogenetic discussion...... 8 Taxonomy ...... 18 Curtonotum Macquart, 1844: 350 (193) ...... 18 Key to New World species of Curtonotum ...... 25 Species accounts ...... 27 The Curtonotum anus species group ...... 27 Curtonotum floridense Klymko & Marshall, sp. n...... 27 Curtonotum helvum (Loew) ...... 32 The Neotropical Clade (Curtonotum sensu stricto) ...... 37 Curtonotum nigrum Klymko & Marshall, sp. n...... 37 The Curtonotum bathmedum species complex ...... 39 Curtonotum bivittatum Klymko & Marshall, sp. n...... 39 Curtonotum bathmedum Hendel ...... 43 Curtonotum magnum Malloch ...... 45 Curtonotum adusticrus Klymko & Marshall, sp. n...... 48 The Curtonotum murinum species complex ...... 50 Curtonotum apicale Hendel ...... 52 Curtonotum trypetipenne Hendel ...... 55 Curtonotum impunctatum Hendel ...... 58 Curtonotum curtispinum Klymko & Marshall, sp. n...... 61 The Curtonotum vulpinum species complex ...... 64 Curtonotum brunneum Klymko & Marshall, sp. n...... 64 Curtonotum pantherinum (Walker) ...... 67 The Curtonotum taeniatum species complex ...... 71 Curtonotum flavisetum Klymko & Marshall, sp. n...... 72 Curtonotum taeniatum Hendel ...... 75 Curtonotum tumidum Enderlein ...... 80 Curtonotum hendelianum (Enderlein) ...... 83 Curtonotum scambum Klymko & Marshall, sp. n...... 86 Curtonotum gracile Klymko & Marshall, sp. n...... 88 Curtonotum desperatum Klymko & Marshall, sp. n...... 91 Curtonotum atlanticum Klymko & Marshall, sp. n...... 93 Curtonotum hunkingi Klymko & Marshall, sp. n...... 97 Curtonotum papillatum Klymko & Marshall, sp. n...... 99 Curtonotum punctithorax Fischer ...... 102 Species incertae sedis ...... 106 Curtonotum hendeli Malloch ...... 106 Curtonotum simplex Schiner ...... 107 Acknowledgements ...... 108 References ...... 108

The New World species of the curtonotid genus Curtonotum Macquart are reviewed, and all species outside the vulpinum and murinum species complexes (as defined below) are revised. Descriptions and illustrations are provided for 24 species, including 13 newly described species: C. adusticrus sp. n., C. atlanticum sp. n., C. bivittatum sp. n., C. brunneum sp. n., C. curtispinum sp. n., C. desperatum sp. n., C. papillatum sp. n., C. gracile sp. n., C. hunkingi sp. n., C. flavisetum sp. n., C. floridense sp. n., C. nigrum sp. n., and C. scambum sp. n. Curtonotum nigripalpe Hendel is proposed as a new junior synonym of C. hendelianum (Enderlein). A key to these New World species is presented and the phylogenetic relationships between them are discussed. Lectotypes are designated for C. tumidum Enderlein, C. bathmedum Hendel, C. taeniatum Hendel, C. trypetipenne Hendel, C. impunctatum Hendel, 1913, nec impunctatum Hendel, 1932 and C. apicale Hendel. Curtonotum perplexum nom. n., is given as the replacement name for C. impunctatum Hendel, 1932 nec impunctatum Hendel, 1913. Two species complexes are left untreated at the species level (the C. murinum species complex, including C. murinum Hendel, C. coriaceum Hendel, C. perplexum nom. n., and C. decumanum Bezzi; and the C. vulpinum species complex including C. vulpinum Hendel and C. fumipenne Hendel). Both complexes are included in the phylogenetic analysis and key.

Key words: Acalyptratae, Ephydroidea, identification key, Nearctic, Neotropical, new species, phylogeny, taxonomy

Introduction

The family Curtonotidae is a widespread group of acalyptrate flies superficially resembling drosophilids (Figures 6–13) and previously treated as a subfamily of Drosophilidae by Hendel (1917, 1928, 1932), Malloch (1930), Stur- tevant (1921) and Curran (1933, 1934). Enderlein (1914, 1917) treated this group as a subfamily of Ephydridae, but Duda (1924) and Okada (1960, 1966) treated Curtonotum Macquart and related genera as part of the Diastatidae. In 1934 Duda placed them in their own family (Curtonotidae) and this has been widely accepted in later works (Hack- man, 1960; Hennig, 1973; Tsacas, 1974, 1977; Kirk-Spriggs, 2008). The family Curtonotidae is defined by the following combination of characters: subcosta complete, ending in costa; costa with humeral and subcostal breaks; cells bm and dm confluent; aedeagus large and C-shaped with an anteroventrally directed distiphallus; and vagina with a large membranous anteroventral pouch originating poste- rior to the ventral receptacle (McAlpine, 1989: 1486–1487; Meier et al., 1997) (for complete diagnosis see Mar- shall et al., 2010). Four curtonotid genera are currently recognized, Curtonotum, Cyrtona sensu lato Séguy, Axinota van der Wulp, and Tigrisomyia Kirk-Spriggs, and 66 species have been described worldwide (not including the 13 newly described herein). At least 50 new Afrotropical species (mostly Cyrtona sensu lato) await descrip- tion (Kirk-Spriggs, 2008). All New World Curtonotidae are in the genus Curtonotum. Curtonotum as currently defined is a probably paraphyletic group most diverse in the tropics, but occurring in every zoogeographic region except the Australasian/Oceanic Region and Antarctica. As discussed below, Neotropical and southwestern Nearc- tic Curtonotum species comprise a distinct monophyletic group including the type species of the genus and here referred to as Curtonotum sensu stricto. Curtonotum in the broader sense, here referred to as Curtonotum senso lato, is probably paraphyletic. According to Kirk-Spriggs (pers. com.), Cyrtona is also paraphyletic and due to be split into multiple genera, Cyrtona in the current broad sense is here referred to as Cyrtona sensu lato. The habits of Curtonotidae are little known. Larvae of the Afrotropical and Palaearctic Curtonotum simile Tsacas and the Afrotropical C. sahaliense Tsacas develop in locust and grasshopper egg pods (Greathead, 1959; Kirk-Spriggs, 2008). The Nearctic C. helvum has been reared from decaying grasshopper egg pods under labora- tory conditions (Meier et al., 1997). A specimen of an undescribed species in the C. murinum species complex (as defined herein) from Monsoon Valley, Tingo Maria, Peru, was “dug [as a puparium] from dirt in association with Philanthus nests” (E.I. Schlinger and E.S. Ross [CASC]) where it possibly developed as a kleptoparasite on the paralyzed bees with which the nest was stocked. Tsacas (1977) briefly reviewed the biology of Afrotropical Cur- tonotum species, noting the habit of roosting in hollow trees, a variety of mammal burrows (including those of warthogs, porcupines, and anteaters), and under shaded overhangs to avoid heat and rain. One Afrotropical species, C. quinquevittatum Curran, has been observed in the hundreds in recently abandoned warthog burrows (Pollock, 2002). Zimbabwean Cyrtona sensu lato adults are found under overhangs along rivers in the dry season and in adjacent vegetation in the rainy season (Pollock, 2002). The Palaearctic and Oriental species Curtonotum anus (Meigen) is known to take shelter under plants (Duda, 1934) and is common along the shaded and muddy shoreline