Zootaxa 3673 (1): 001–064 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Monograph ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3673.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:2D0CEAB4-5CC6-42B6-8388-FBA7113C87C2 ZOOTAXA 3673 Review and analysis of information on the biology and morphology of immature stages of robber flies (Diptera: Asilidae) D. STEVE DENNIS1,4, JEFFREY K. BARNES2 & LLOYD KNUTSON3 11105 Myrtle Wood Drive, St. Augustine, FL 32086-4838, U.S.A. E-mail: [email protected] 2Department of Entomology, University of Arkansas, 319 Agriculture Building, Fayetteville, AR 72701, U.S.A. E-mail: [email protected] 3Salita degli Albito 29, 04024 Gaeta (LT), Italy E-mail: [email protected] 4Corresponding author Magnolia Press Auckland, New Zealand Accepted by N. Evenhuis: 6 May 2013; published: 17 Jun. 2013 D. STEVE DENNIS, JEFFREY K. BARNES AND LLOYD KNUTSON Review and analysis of information on the biology and morphology of immature stages of robber flies (Diptera: Asilidae) (Zootaxa 3673) 64 pp.; 30 cm. 17 Jun. 2013 ISBN 978-1-77557-200-8 (paperback) ISBN 978-1-77557-201-5 (Online edition) FIRST PUBLISHED IN 2013 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/ © 2013 Magnolia Press All rights reserved. No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing. This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use. ISSN 1175-5326 (Print edition) ISSN 1175-5334 (Online edition) 2 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. Table of contents Abstract . 3 Introduction . 3 Results . 5 Summary of Subfamilies . 5 Discussion . 8 Acknowledgments . 14 References . 15 Tables . 22 Abstract Recent publications on the immature stages of robber flies (Asilidae) are reviewed and listed for the 14 currently recog- nized subfamilies (Asilinae, Bathypogoninae, Brachyrhopalinae, Dasypogoninae, Dioctriinae, Laphriinae, Leptogastri- nae, Ommatiinae, Phellinae, Stenopogoninae, Stichopogoninae, Tillobromatinae, Trigonomiminae, and Willistonininae). For the Phellinae there are only limited data on the pupal cases, and for the Tillobromatinae there are only limited data on the eggs (potential oviposition site in soil) and pupal cases. The recent literature is compared with pertinent pre-1972 pub- lications. Key words: Insecta, Diptera, Brachycera, Asiloidea, immatures, eggs, larvae, pupae Introduction Robber flies are one of the largest families of Diptera with approximately 7,000 species (Geller-Grimm 2011a). Despite their widespread occurrence, there is still limited information on the immature stages. Irwin-Smith (1923) listed papers on the biology and morphology of immatures by genus and species. Hennig (1952) provided descriptive data on the immature stages based on two subfamilies and associated tribes: Asilinae (Leptogastrinae, Asilini) and Dasypogoninae (Laphriini, Dasypogonini). Knutson (1972) referred to papers not mentioned in the previous summaries and listed published morphological descriptions and figures for genera and species in four of the five subfamilies recognized by Hull (1962) (Asilinae, Dasypogoninae, Laphriinae, Leptogastrinae). The Megapodinae were not included because of the absence of information on the morphology of immatures. Londt (1994) listed papers on egg morphology, larval behavior and morphology, and pupal morphology of Afrotropical Asilidae using a ten-subfamily classification (Apocleinae, Asilinae, Dasypogoninae, Laphriinae, Laphystiinae, Leptogastrinae, Ommatiinae, Stenopogoninae, Stichopogoninae, Trigonomiminae). Lavigne et al. (2000) listed publications from 1840 to 1976 on the immature stages that were not included in previous papers, and reviewed robber fly oviposition sites, eggs, and larval habitat, food, and feeding. Dennis et al. (2008a) briefly reviewed the egg and larval stages, and provided more detailed information on the pupae, with descriptions of pupal cases based on Nearctic species of four subfamilies (Asilinae, Dasypogoninae, Leptogastrinae, Laphriinae). They also compared subfamily treatments of Asilidae since 1962 that included five (Asilinae, Dasypogoninae, Laphriinae, Leptogastrinae, Megapodinae) to 14 subfamilies (Asilinae, Bathypogoninae, Brachyrhopalinae, Dasypogoninae, Dioctriinae, Laphriinae, Leptogastrinae, Ommatiinae, Phellinae, Stenopogoninae, Stichopogoninae, Tillobromatinae, Trigonomiminae, Willistonininae). Lavigne (2011b) provided a broad review of eggs of robber flies; larval habitat, development, food and feeding; and pupae, taking into consideration 11 subfamilies (Asilinae, Bathypogoninae, Brachyrhopalinae, Dasypogoninae, Dioctriinae, Laphriinae, Leptogastrinae, Ommatiinae, Phellinae, Stenopogoninae, Stichopogoninae). Shortly after Knutson’s (1972) publication, Papavero (1973) proposed an eight-subfamily classification (Apocleinae, Asilinae, Dasypogoninae, Laphriinae, Laphystiinae, Ommatiinae, Stenopogoninae, and Trigonomiminae). An additional four subfamilies (Atomosinae, Dioctriinae, Megapodinae, Stichopogoninae) were added by other authors by the early 2000s. Bybee et al. (2004) presented the first formal analysis of molecular evidence for phylogenetic relationships among the Asilidae and recognized 10 of the 11 subfamilies (Apocleinae, Asilinae, Dasypogoninae, Laphriinae, ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 3 Laphystiinae, Leptogastrinae, Ommatiinae, Stenopogoninae, Stichopogoninae and Trigonomiminae). The Dioctriinae was not included in the analysis. Dikow (2009a ) evaluated a sample of 158 species from 140 genera (of 545 total genera of Asilidae) and proposed a revised phylogenetic classification, inferred from morphological characters of adults, to recognize 14 subfamilies (Asilinae, Bathypogoninae, Brachyrhopalinae, Dasypogoninae, Dioctriinae, Laphriinae, Leptogastrinae, Ommatiinae, Phellinae, Stenopogoninae, Stichopogoninae, Tillobromatinae, Trigonomiminae, Willistonininae). Based on a subsequent total evidence analysis of adult morphological and DNA sequence data (using 77 of the previously evaluated species), Dikow (2009b) concluded that not all higher-level taxa as previously delineated (primarily in his 2009a paper) are monophyletic. The only subfamilies established as monophyletic are Dioctriinae, Laphriinae, Leptogastrinae, Ommatiinae, and Stichopogoninae. Apocleinae and Asilinae were considered paraphyletic with respect to each other and Laphystiinae paraphyletic with respect to Laphriinae. The Dasypogoninae, Stenopogoninae, and Trigonomiminae were considered not monophyletic. The Bathypogoninae, Brachyrhopalinae, Phellinae, Tillobromatinae, and Willistonininae were elevated to new subfamily status based on their morphological distinctness. Table 1 compares Dikow’s (2009b) subfamily classification with other treatments since 1962. In the present study we summarize the literature on the biology and morphology of the immature stages of robber flies (based on Dikow’s (2009b) 14 subfamilies) published primarily since Knutson’s (1972) morphological summary. Materials and Methods The egg, larval and pupal data are evaluated for major characteristics of genera, although for many there is only information on one species. The original researcher’s terminology generally is used, in particular with respect to the eggs. Also, figures of the eggs were examined to see if additional information could be added to the descriptions. A number of researchers obtained information on eggs by dissecting gravid females or from females in museum collections. Information on the internet that only mentions ovipositing and/or often shows photographs of different species ovipositing, is not included. Terminology The following definitions are used in the text, tables, and figures: EGG MORPHOLOGY: Elongate—longer than wide. Globule—a small spherical mass. Nipple—small protuberance or projection. Oval—resembling an egg in shape. Polygon—many sided. Reticulate—covered with a network of lines; net-like. Ridge—long, narrow elevation. Spherical—shaped like a sphere or round. Striations—longitudinal lines. Tubercle—elevated structure that is pimple-like. LARVAL MORPHOLOGY: D—morphological description or descriptive information. F—figure. G—general morphological and/or other information. L1, L2, and L3—larval instars. M—L4 to L7 instars or mature larvae. 4 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. PUPAL MORPHOLOGY: D—morphological description or descriptive information. F—figure. G—general morphological and/or other information. Results General. Publications on the biology and morphology of the immature stages of robber flies, primarily since 1972, are listed in Table 2. The table includes references in which the researchers provided actual and speculative information, in particular on oviposition/egg sites based on the structure of female ovipositors and the habitat of the larvae and pupae (i.e., vegetation, under bark of trees or ground/soil). The information in Table 2 lists references that have general comments on the family, and then the 14 subfamilies with associated tribes, genera, and species. Tables 3 to 5 focus on the subfamilies and genera, and summarize the major biological and