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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/

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

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 recognized 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 publications.

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.

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 morphological information for the egg (Tables 3 and 4), larval and pupal (Table 5) stages. Table 6 lists the number of genera and species by subfamily for which information on immatures has been published and compares it with the approximate worldwide number of genera in each subfamily. Profiles or shapes of eggs for a number of species of robber flies are shown in Fig. 1. The larva of Choerades gilva (Linnaeus) and pupal case of Mallophora atra Macquart are shown in Figs. 2 and 3, respectively, with their morphological characteristics labeled.Summary of Subfamilies

Subfamily Asilinae Female robber flies in the subfamily Asilinae drop eggs onto the substrate or onto vegetation, or by inserting their ovipositor into the soil, dry cow or horse dung, or rabbit pellets (Asilus crabroniformis Linnaeus); or in vegetation (in particular between the sheath and stem or dry seed heads of grasses); or onto wire fences (i.e., Mallophora ruficauda (Wiedemann); Castelo & Corley 2004a, Castelo et al. 2006). The number of eggs laid varies from 1 to 729 with most from 1 to 20. The largest numbers of eggs are produced by Mallophora spp., Megaphorus guildianus (Williston), and Porasilus barbiellini Curran, which deposit them in clusters and cover them with a soft chalky-white albumin. It seems that the more eggs produced per oviposition event, the smaller the size of the eggs. Eggs in ovipositions with 1 to 20 eggs, ranged in length from 0.80 to 2.80 mm and in width from 0.30 to 0.82 mm. For more extensive ovipositions, with 32 to 390 eggs, the eggs ranged in length from 0.80 to 1.01 mm and in width from 0.22 to 0.38 mm. Eggs are most often white to creamy white, but can be yellowish white, pearly (shiny) whitish to pinkish, yellowish brown, light brown, or light gray. Eggs are most frequently elongate or oblong, but some genera have more or less oval eggs. Asilinae eggs that have been examined with a scanning electron microscope (SEM) have surface features including striations, elevated bodies (nipples, tubercles), crevices, spherical bodies and globules, and reticulate sculpturing with different shapes of polygons. Antipalus varipes (Meigen) is unique in the subfamily Asilinae with eggs that have fine sand grains on them (except on one longitudinal side of the egg) from the sand in which they are oviposited. Aeropyles are present or absent on Asilinae eggs. They may be evenly spread over the surface, except in the area of the micropyle, or more concentrated. They may be located on cone-shaped structures or on flat areas of the chorion. A micropyle has been observed for some, but not all eggs. The immediate area around the micropyle is typically smooth. The egg stage lasts from 2 to 8 days with an average of approximately 6 days. When the larva emerges from the egg, it continues to live in the soil, decaying vegetable matter, or possibly dry horse and cow dung, where it feeds upon larvae in the insect families Chrysomelidae, Curculionidae, Scarabaeidae, and possibly Asilidae. Scarabaeidae larvae have been the most frequently reported prey. The larval stage of Asilinae can be quite long, lasting from 1 to 3 years. The pupal stage is much shorter, lasting from 14 to 70 days.

ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 5 Information on larval morphology has generally been published on the first two instars and mature larvae. For a number of species there are descriptions of pupal cases, although there are not many detailed descriptions. The literature contains figures of both the larvae and pupal cases.

Subfamily Bathypogoninae The only published information on the eggs of the Bathypogoninae is for Bathypogon spp. There are at least seven eggs per oviposition in the soil. The eggs are cream colored and elongate. The eggs have not been examined with a SEM. There is no information on the larvae of Bathypogoninae and only general morphological information on pupae.

Subfamily Brachyrhopalinae Most eggs of Brachyrhopalinae have been oviposited in the soil or debris on the ground, although Brachyrhopala may oviposit on vegetation such as Acacia spp. Cyrtopogon oviposit 2 to 4 eggs that are elongate and creamy white. The eggs range in length from 1.12 to 1.42 mm and width from 0.42 to 0.56 mm. They have not been examined with a SEM. The larval and pupal stages occur in the soil. The larval stage for Brachyrhopala lasts for about 155 days; whereas the pupal stage for Brachyrhopala and Holopogon lasts for 23 to 42 days. Larvae of Brachyrhopala are reported to feed on the larvae of Cerambycidae (Phoracantha semipunctata (Fabricius)). Holopogon larvae feed on Coleoptera and other insect larvae. Some morphological information has been published on the larvae of Cyrtopogon and Heteropogon. Morphological information on the pupae has been published for Ceraturgus, Cyrtopogon, Heteropogon, and Holopogon.

Subfamily Dasypogoninae Female Dasypogoninae oviposit 1 to 11 eggs in the soil, although Leptarthrus brevirostris (Meigen) has been observed to oviposit on a dead branch on the ground. The eggs are creamy white to white, and long-oval to elongate. They range in length from 0.87 to 1.11 mm and width from 0.02 to 0.51 mm. Megapoda has nipples, tubercles and aeropyles on the egg surface; and one micropyle in a circular area that has a thick evaginated rim. Like Antipalus varipes, Dasypogon diadema (Fabricius) has one to six eggs in a “sand ball or cocoon” from the sand in which they are oviposited. Between the eggs and the sand grains is a “silky lining”; whereas, A. varipes oviposits only one egg that is covered with sand grains without a silky lining. The egg stage of D. diadema lasts from 6 to 17 days in the laboratory. Most Dasypogoninae larvae and pupae develop in the soil; those in the tribe Megapodini develop in plant roots and decaying stumps/logs. The larvae are reported to feed on Cerambycidae and Scarabaeidae larvae. There is no information on how long the larval stage lasts; the pupal stage of D. diadema is reported to last 27 days. Morphological information has been published on the larvae of D. diadema. Pupal case descriptions or descriptive information has been published for Comantella, Dasypogon, Diogmites, Megapodini, and Pseudorus.

Subfamily Dioctriinae There is limited information on the immature stages of the subfamily Dioctriinae. The females drop 1 to 18 eggs per oviposition on vegetation or the ground. The reddish brown or dull brown, subspherical to oval eggs are 0.40 to 0.56 mm in length and 0.30 to 0.48 mm in width. Examination of the eggs with a SEM indicates that they have surface sculpturing such as 5- to 7-sided polygons, oval rings, pimples, and elevated ridges. Aeropyles are present inside or outside of rings and may be surrounded by slightly elevated ridges. There is one micropyle in a smooth area that may be surrounded by cup-like structures. Dioctriinae larvae and pupae develop in the soil or decaying wood. Some morphological information has been published on the larvae and pupae of Dioctria.

Subfamily Laphriinae The majority of Laphriinae females deposit 2 to 18 eggs in dead tree trunks or other wood. Andrenosoma fulvicaudum (Say) oviposits primarily on pinyon pine trees (Pinus monophylla Torr. and Frem.) still smoldering

6 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. from fires and the females have sensory organs (“cercal organs”) on their ovipositors that aid in the selection of oviposition sites such as Coleoptera burrows. Cerotainia albipilosa Curran oviposits on vegetation. Londt (1994), based on the morphology of the ovipositor, speculated that some genera might drop eggs onto the ground. Eggs of Laphriinae species are bright amber, brown or reddish brown. They are usually oval, but can be round or elongate. The oval to round eggs range in length from 0.29 to 0.50 mm and width from 0.25 to 0.48 mm; more elongate eggs range in length from 0.75 to 0.93 mm and width from 0.55 to 0.60 mm. Most Laphriinae eggs have surface features consisting of different shapes of polygons, often with raised ridges and reticulate sculpturing. Some also have pimples or spine-like structures on the surface. Most genera have aeropyles, some of which are cone-shaped. There are one or two micropyles located in broad, smooth areas or in narrow areas with low ridges. Fisher (1986) indicated that no completely smooth eggs have been reported for the tribe Andrenosomatini (as Andrenosomini), but some other Laphriinae have smooth eggs. The chorion features or sculpturing begins at the edge of the smooth micropyle area or posterior to it if there is a “corona” of pale, flocculent or scaly material. These features are usually most distinct in the apical half of the egg, with the remaining half patterned or smooth, although some species have eggs with a chorion that is completely covered with sculpturing. Under laboratory conditions at a constant temperature of 25 °C, the egg stage of Andrenosoma atrum (Linnaeus) lasts up to 50 to 56 days (Musso 1978, 1981b, as A. atra). Laphriinae larvae and pupae are usually found in decaying trees and tree stumps, or under scorched bark of pinyon pine trees (Andrenosoma fulvicaudum). The larvae are reported to feed on Coleoptera larvae from the families Buprestidae, Cerambycidae, and Curculionidae. The larval stage for Choerades lasts for up to 2 years. No information was found on the length of the pupal stage. Detailed morphological information is available on a number of species, in particular in the genera Andrenosoma and Laphria. The most detailed descriptions of Andrenosoma larvae are in Musso (1978, 1981b). The most detailed pupal case descriptions with figures are in Dennis & Barnes (2012), Dennis et al. (2008a), and Musso (1981b). There is general descriptive information for other genera.

Subfamily Leptogastrinae Females in the genus Leptogaster drop, singly, 1 to 8 eggs to the ground as they are flying or resting on vegetation. The oval eggs are amber to light yellow-brown or light yellow. They range in length from 0.32 to 0.50 mm and width from 0.25 to 0.39 mm. The surface of the eggs of Leptogaster and Psilonyx is either smooth, or they have varying shapes of polygons without ridges. The eggs of Beameromyia and Tipulogaster are smooth without ridges. Aeropyles have not been observed on Leptogastrinae eggs. There are one or two micropyles in a smooth to elevated area. The egg stage lasts for 9 to 14 days. Leptogaster larvae develop in soil; those of Mesoleptogaster are said to develop in deciduous trees. Morphological information, including figures, is available for Leptogaster larvae and pupae, and Apachekolos pupal cases.

Subfamily Ommatiinae Limited information has been published since 1972 on the immature stages of Ommatiinae. The eggs of Ommatius are dropped onto the ground. Some descriptive information has been published on the pupae, but only Ommatius spp. pupae have been described in detail with figures.

Subfamily Phellinae For the subfamily Phellinae only general morphological information has been published on the pupae.

Subfamily Stenopogoninae Genera of Stenopogoninae deposit 3 to 20 eggs at a time in the soil. The eggs are suboval to elongate, and creamy white to white. The elongate eggs range in length from 0.90 to 2.29 mm and width from 0.52 to 0.77 mm. The suboval eggs of Archilestroides and Taperigna are 0.90 to 1.20 mm long and 0.60 to 0.72 mm wide. Archilestroides eggs have raised bodies of irregular shapes and sizes. The eggs of Taperigna have hexagonal chorionic sculpturing. Both genera have aeropyles spread homogeneously on the egg surface and one or two circular to elongate micropyles in smooth areas.

ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 7 Stenopogon larvae are known to feed on scarab beetle larvae and other insect larvae. The larvae of Grypoctonus in Japan have been described and figured, and Stenopogon spp. pupae have been described and figured.

Subfamily Stichopogoninae Limited information on the immatures of this subfamily has been published since 1972. The eggs are oviposited in the soil, but none has been described in detail. The eggs of Lasiopogon are oval and some species of Stichopogon are believed to not have aeropyles. Lasiopogon larvae and pupae develop in the soil with the latter maturing in 17 days. Stichopogon larvae and pupae also are believed to develop in the soil. There are general descriptions of the pupae for the subfamily, and of the larvae and pupae for Lasiopogon.

Subfamily Tillobromatinae The only information on the immatures of the subfamily Tillobromatinae is speculation on where the eggs are oviposited (in soil) and a general description of the pupae.

Subfamily Trigonomiminae Genera in the subfamily Trigonomiminae drop 3 to 326 eggs onto the soil or possibly onto vegetation. The spherical to slightly oval or oval eggs are creamy white to light or dark brown. They range in length from 0.24 to 0.32 mm and width from 0.16 to 0.32 mm. Damalis eggs have hexagons and tiny spicules on the surface. The eggs of Holcocephala have irregular pentagons and hexagons with thick or tall, thin ridges. Aeropyles are present on the eggs of Holcocephala. Damalis and Holcocephala eggs have one micropyle with the micropyle of the latter in a broad, smooth or floral-like area. The egg stage of Damalis lasts 14 days and the larvae and pupae occur in the soil. First-instar larvae of Damalis have been described and figured. General information on Trigonomiminae pupae has been published.

Subfamily Willistonininae Ablautus rufotibialis Back deposits one white egg per oviposition in the soil. The eggs of this species have not been examined using SEM. There is no information on the larvae and only a general description of Willistonininae pupae has been published.

Discussion

Biological and morphological information on the immature stages of robber flies has been published on 137 genera and 299 species in the 14 subfamilies (Table 6). For the subfamily, Phellinae only general morphological comments have been made on the pupae, and for the Tillobromatinae, there has only been speculation on the oviposition site (in soil) for the eggs and general morphological comments have been made on the pupae.. This is still limited knowledge when one considers that there are around 545 genera and 7,000 species of robber flies worldwide (Dikow 2009a & b, Geller-Grimm 2003, 2011a). Knutson (1972) indicated that only about 2% of the then approximately 5,000 species of robber flies were known in any immature stage. Currently there is information on 25.1% of the genera and 4.3% of the immature stages of the known described species. A comparison of literature on immature robber flies, including the more detailed publications before 1972, indicates the following for each stage of development. Eggs. The method and location (i.e., live or dead vegetation and ground/soil/sand) of robber fly subfamily and genera oviposition, and egg morphology has been discussed in some detail by at least Melin (1923), Hinton (1981), Londt (1994), and Stubbs & Drake (2001). Londt (1994) also speculated on the method and location of oviposition for a number of genera of Afrotropical Asilidae based on the shape of female ovipositors. There are three general types of oviposition with associated female ovipositor morphology, and egg shape and color. Oviposition by random egg dropping: The female ovipositor is unspecialized, blunt or conical, usually not elongate, the cerci and subgenital plate are simple, and the setae do not have any unusual adaptations (e.g., long or stout) (Londt 1994). The usually spherical to oval eggs are dropped in flight (Leptogaster) or while a female is on

8 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. vegetation (Dioctria), and the eggs drop onto the ground or vegetation. Some genera, such as Pamponerus and Rhadiurgus, also have more elongate eggs. The eggs can be yellowish, brownish or whitish. Oviposition in or on dead or live vegetation: A female’s ovipositor is short (Laphria) or broadly truncate and conical, laterally compressed/flattened (Dysmachus and Eutolmus) or occasionally slightly dorsoventrally compressed (Leptarthrus), or elongate conical. According to Londt (1994), if the ovipositor is not obviously elongate and/or laterally compressed the cerci may be elongate or equipped with long setae that serve to guide eggs into the area that they are being oviposited. The eggs are generally elongate to oval and can be brown, red, or white. Oviposition in soil or dung: The female ovipositor can have pointed and/or upturned cerci and/or subgenital plate, and/or the presence of a number of spines (acanthophorites) or setae (Londt 1994). Genera such as Proctacanthus often have very stout spines, although some genera may have spines that are short (Stichopogon). The spines are used to push aside the soil so that the eggs can be laid and then used to sweep over the soil to conceal the oviposition site. The eggs are generally elongate or oblong, and whitish. Musso (1978, 1981b) studied the eggs of a number of species of robber flies and identified three different types: pigmented eggs, eggs without pigmentation, and eggs covered with fine sand grains. Andrenosoma atrum and A. bayardi Séguy eggs are pigmented with a deep brownish-red tinge, and reticular sculpturing consisting of polygons. Machimus fimbriatus (Meigen), M. pilipes (Meigen), M. rusticus (Meigen), and Dystolmus kiesenwetteri (Loew) usually have unpigmented eggs, but sometimes they are light gray or very pale yellow. Only Antipalus varipes was identified as having individual eggs with a thin pellicle made of fine sand grains. Geller-Grimm (2011b) illustrated and described 1 to 6 eggs oviposited by Dasypogon diadema inside “sand balls or cocoons”. Candan et al. (2004a) indicated that Musso’s classification does not work for all of the eggs of species that they and Castillo et al. (1994) studied. Robber flies oviposit into or onto live or dead vegetation, drop eggs onto the ground while resting or in-flight, or oviposit in soil, dry cow/horse dung or rabbit pellets. The Asilinae exhibit all of these methods and locations of oviposition. Brachyrhopalinae, Dasypogoninae, Stenopogoninae, Stichopogoninae, and Willistonininae oviposit in the soil or debris or dead branches on the ground. Dioctriinae, Leptogastrinae, Ommatiinae, and Trigonomiminae drop eggs onto vegetation or the ground. Laphriinae oviposit primarily onto dead trees or tree stumps, but are also reported to drop eggs onto the ground. Londt (1994) speculated that based on the shape of the ovipositor, some genera of Laphriinae may oviposit in dung (e.g., Hoplistomerus). The majority of robber flies deposit less than 20 eggs per oviposition, with many depositing less than 10 eggs. Some Asilinae genera (i.e., Mallophora, Megaphorus, Porasilus) that deposit their eggs in a frothy or chalky-white “case,” oviposit from 32 to 729 eggs. Most eggs of Asilinae are elongate (Fig. 1) and whitish, although some genera such as Neolophonotus may have oval eggs; Colepia and Dolopus may have more brownish eggs; and Neoaratus eggs may be yellowish. Bathypogoninae (i.e., Bathypogon danielsi Lavigne) has elongate, cream colored eggs. Brachyrhopalinae (i.e., Cyrtopogon montanus Loew) eggs are elongate and creamy white. The eggs of Dasypogoninae are elongate to long-oval and creamy white or white. Dioctriinae eggs are subspherical to oval and reddish-brown to dull brown. Laphriinae eggs are mostly oval to elongate, but also can be oval to spherical; their color can be reddish-brown, dull brown, bright amber or have a red tinge. The eggs of Leptogastrinae appear to be more consistent than other subfamilies and are suboval to oval, amber to light yellow-brown or light yellow. Stenopogoninae eggs are suboval to elongate, and creamy white to white. Trigonomiminae eggs are spherical, slightly elongate to oval or oval, and creamy white, or light to dark amber. The eggs of Willistonininae (i.e., Ablautus rufotibialis) are white. There is no information on the shape or color of the eggs of robber flies in the subfamilies Phellinae, Ommatiinae, Stichopogoninae and Tillobromatinae. As robber fly eggs mature, they may darken in color. Irwin-Smith (1923) indicated that the eggs of Neoaratus hercules (Wiedemann) changed from a light creamy color to a yellowish brown. Robber flies that produce elongate eggs generally have eggs that are at least two times as long as wide. Spherical or round eggs have an equal length and width. Oval to sub-oval eggs have slightly longer lengths than widths. Some eggs from genera in all families of robber flies that have been examined with a SEM generally have sculpturing, except for the Brachyrhopalinae (i.e., Cyrtopogon montanus). The sculpturing usually consists of irregular or regular shaped polygons, often with raised edges. Melin (1923) reported Dioctria eggs with “… irregularly located dot-shaped elevations” and Laphria eggs with irregularly shaped polygons. There also are

ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 9 genera in the subfamilies Asilinae, Dasypogoninae, Laphriinae, Stenopogoninae, and Trigonomiminae that have raised or elevated bodies that can be shaped like globules, spheres, nipples, and/or tubercles. There is limited information on aeropyles and their distribution on the eggs of robber flies They may be present or absent on eggs of Asilinae and Laphriinae, and if present on Asilinae eggs, they are concentrated in the central- to-apical region of the egg. According to Hasbenli et al. (2008), aeropyles are widespread across the surface of most Laphriinae eggs. Aeropyles appear to be present on Dasypogoninae eggs and spread homogeneously over an egg’s surface. Aeropyles also are on the eggs of Dioctriinae, most Stenopogoninae, and Trigonomiminae. Most micropyles are in a broad, smooth area on the apex of eggs. On Asilinae eggs there may be one or no micropyles. On Dasypogoninae (Megapoda labiata (Fabricius)), Dioctriinae, and Trigonomiminae eggs there is one micropyle; whereas Laphriinae, Leptogastrinae, and Stenopogoninae eggs have one or two micropyles. Robber fly eggs generally mature in 1 to 18 days depending on environmental conditions. According to Musso (1978, 1981b), under laboratory conditions, the eggs of Andrenosoma atrum hatched in 50 to 56 days. Larvae. Young or neonate Dioctria flavipennis Meigen larvae emerge from their eggs by pressing against the micropylar region, causing it to separate along a thin, weaker eclosion line (Candan et al. 2004a). According to Stubbs & Drake (2001), the larvae of many species emerge through an irregular split in the egg; whereas species in genera such as Dioctria and Laphria that have “hard shelled” eggs emerge through a small circular flap. Irwin- Smith (1923) also observed the larvae of Neoaratus hercules emerging through a circular cap. Larvae that emerge from eggs that were deposited into or onto vegetation generally drop to the ground where the larvae and pupae develop. The exceptions to this appear to be in the genus Brachyrhopala (Brachyrhopalinae), and the tribe Megapodini and Pseudorus distendens (Wiedemann) (both Dasypogoninae). Most Laphriinae are well known for ovipositing mainly on dead trees and stumps, and the larvae and pupae developing there. However, larvae and pupae of Laphystia and Loewinella are reported to develop in soil.

FIGURE 1. Robber fly egg profiles: A. Leptogaster cylindrica (De Geer, 1776), B. Dioctria rufipes (De Geer, 1776), C. Cyrtopogon lateralis (Fallén, 1814), D. Laphria ephippium (Fabricius, 1781), E. Laphria flava (Linnaeus, 1761), F. Choerades marginata (Linnaeus, 1758), G. Choerades gilva (Linnaeus, 1758), H. Laphria gibbosa (Linnaeus, 1758), I. Pamponerus germanicus (Linnaeus, 1758), J. Asilus crabroniformis Linnaeus, 1758, K. Rhadiurgus variabilis (Zetterstedt, 1838), L. Eutolmus rufibarbis (Meigen, 1820), M. Didysmachus picipes (Meigen, 1820), N. Tolmerus atricapillus (Fallén, 1814). After Melin (1923).

10 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. Complete development of larvae generally requires from 1 to 4 years. Mallophora (Asilinae) and Brachyrhopala (Brachyrhopalinae) larvae are reported to develop in 129 and 155 days, respectively. If environmental conditions are too dry, the larval period can be extended during which the larvae do not grow any larger. Researchers have indicated that there are four to seven larval stages or instars (L1 to L7). Clements & Bennett (1969) identified four instars for Mallophora media Clements. Crespo & Castelo (2010) concluded that Mallophora ruficauda (Wiedemann) has five instars prior to the pupal stage. Musso (1978, 1981b) showed that Machimus rusticus (Meigen) has seven instars. The best morphological information is on the first two instars (L1 and L2) and mature larvae (L7). There is a paucity of information on instars L3 through L6. The most complete information is by Musso (1978, 1981b) for Machimus rusticus in a laboratory rearing at 22 to 25ºC. For this species development from L1 through L6 generally was very rapid with each instar lasting a minimum of 4 to 12 days with a total duration of about 40 days, although larvae could remain in stage L6 for up to one year. During stage L7, larval growth almost completely stopped. Many larvae died from parasitic nematodes or when eaten by Scarabaeidae larvae that were presented as a food source. The L7 stage lasted 7 or 8 months when the first signs of pupation were observed.

FIGURE 2. Choerades gilva (Linnaeus, 1758) larva: A. dorsal view, B. ventral view. abss, abdominal segment 8 spiracles (posterior spiracles); ao, anal opening; cc, circular callosities; dcp, dorsal contractile process (wart, welt, pseudopod); lc, lateral callosities; lcp, lateral contractile process; mx, maxilla; mp, maxillary palpus; pths, prothoracic spiracles (anterior spiracles); s, setae; slp, spine-like process; sp, sclerotized plate; vcp, ventral contractile process; vlcp, ventro-lateral contractile processes. After Melin (1923).

ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 11 Irwin-Smith (1923) described the first-instar larvae of Neoaratus hercules. Musso (1979, 1981b) described the first and second instar larvae of Machimus rusticus and indicated that during the second instar the morphological characteristics that have formed will become more prominent through the mature seventh stadium, although some features such as the maxillary palpi and the thoracic and abdominal setae will become shorter. Musso (1978, 1981b) also described and illustrated the tracheal system of newly hatched larvae of Machimus rusticus. He indicated that the prothoracic spiracles are present, but probably not functional, and therefore the larvae are metapneustic. According to Wood (1981), and Foote & Dennis (1991), the respiratory systems of robber fly larvae are functionally amphipneustic, although there are vestigial spiracles on the first seven abdominal segments. Keilin (1944) indicated that an amphipneustic respiratory system only has two pairs of spiracles, one prothoracic and one postabdominal, and occurs in the majority of cyclorrhaphous Diptera. Keys to mature larvae have been provided for only three or four subfamilies by Bromley (1946) and Melin (1923) for Asilinae, Dasypogoninae, Laphriinae, and Asilinae; and by Brindle (1962), Foote & Dennis (1991), Hennig (1952), Séguy (1927), and Smith (1989) for Asilinae, Dasypogoninae, Laphriinae, and Leptogastrinae. These keys have all focused on the morphology of the mandibles and maxillae, callosities (warts, welts, pseudopods) on the abdominal segments, and the last abdominal segment (Fig. 2). Keys to mature larvae of various genera and/or species in the four subfamilies can be found in Brindle (1962), Bromley (1946), Hennig (1952), Krivosheina (1973), Krivosheina & Mamaev (1973, 1975a and b), Malloch (1917), Melin (1923), and Musso (1978). Morphological characteristics used to separate the larvae include length and width; size and shape of the mouthparts and head capsule; distinct/indistinct antennae; thoracic segments, including presence/absence of callosities; setae on thoracic and last abdominal segments; swellings, pseudopods, callosities, lobes or warts on the abdominal segments; and other characteristics of the last abdominal segment. Individual genus and/or species descriptions of mature larvae can be found in Brindle (1968, 1969), Clements & Bennett (1969), Copello (1927, 1942), Krivosheina (1973, 1974), Krivosheina & Mamaev (1973, 1975a and b), Kurkina (1979), Malloch (1917), Melin (1923), and Séguy (1927). There has been considerable discussion on the food of robber fly larvae. Many researchers, such as Melin (1923), believed that they are phytophagous. However, Malloch (1917) and others, commented that they are predaceous on other insect larvae. Knutson (1972) and Lavigne et al. (2000) summarized literature not included in Melin (1923) that confirms that the larvae feed on insect larvae, in particular larvae of Scarabaeidae, commonly known as white grubs. The larval food information in Table 5 indicates that they feed on Coleoptera larvae with a preference for Scarabaeidae, although they also feed on Buprestidae, Cerambycidae, Chrysomelidae, Curculionidae, Tenebrionidae, other insect larvae, and possibly other robber fly larvae (e.g., Stenopogon and possibly Tolmerus). Based on laboratory experiments, Musso (1978, 1983) indicated that Machimus rusticus and M. pilipes first- instar larvae live on the yolk from the egg. The second instar of M. rusticus feeds on substances excreted and exuded by Scarabaeidae larvae. The third through seventh instars feed on Scarabaeidae, Chrysomelidae, and Curculionidae larvae. Also, Stenopogon sp. larvae were observed to feed on each other. Castelo and colleagues [summarized in Castelo & Crespo (2011), and Groba & Castelo (2011)] have studied in detail the host location of Mallophora ruficauda larvae. They found that the first-instar larva cannot orient towards their preferred host larvae, Cyclocephala signaticollis Burmeister (Coleoptera: Scarabaeidae), because the maxillary palpi that bear the sensory organs are not well developed. Second-instar larvae up to 50 days old have well developed maxillary palpi and can orient to and locate the host. Third through fifth instars are less active, and pupation occurs during the fifth stadium. Robber fly larvae orient to the host Scarabaeidae larvae using chemical cues generated in the host larva’s proctodeum or fermentation chamber. Mallophora ruficauda larvae can both detect and discriminate the odors of different potential hosts and can specifically orient to C. signaticollis larvae. Also, M. ruficauda larvae can determine the quality of the potential hosts with respect to parasitism status and can orient to healthy hosts. The chemicals that the robber fly larvae orient to are to be determined. Pupae. Larvae require 12–90 days to pupate, but most take about 14–35 days. When the last -instar larvae are ready to pupate, they move to within approximately 1–10 cm of the soil surface, or in the case of wood dwelling larvae, near an exit hole in the wood that they or another wood boring insect has made. Laphriinae pupae are known to occupy a gallery to the surface of a tree stump or other part of a plant that was previously made by the larvae (Melin 1923). Most pupae appear to emerge during the morning by using the spurs and spines in a rotary movement to work to the surface of the substrate.

12 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. FIGURE 3. Mallophora atra Macquart, 1834, pupal case: A. Thorax and abdominal segments 1–9 (lateral), B. Facial sheath (lateral), C. Facial sheath (ventral), D. Anterior mesothoracic spines (dorsolateral), E. Abdominal segment 1 (posterodorsal), F. Abdominal segment 2 (dorsolateral). aap, anterior antennal process; abs, abdominal spiracles; amsp, anterior mesothoracic spines; dpp, dorsal posterolateral process; ls, labral sheath; ls1, leg sheath 1; ls2, leg sheath 2; ls3, leg sheath 3; ms maxillary sheath; pap, posterior antennal processes; pmc, posterior mesothoracic callosity; prs, proboscidal (=proboscial, olim, incorrect spelling); ps, palpal sheath; pths, prothoracic spiracle; vc, ventral callosity; vpp, ventral posterolateral process; ws wing sheath. After Dennis et al. (2008b).

ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 13 Dennis & Lavigne (1976a) described the emergence of adult Comantella fallei (Back) and Machimus occidentalis (Hine) (as M. sp. either callidus (Williston) or occidentalis)). Pre-1972 pupal case descriptions are generally brief, and many do not include descriptions of comparable morphological characteristics. Publications with keys, genera and/or species descriptions include Brindle (1968), Bromley (1946), Clements & Bennett (1969), Copello (1927, 1942), Malloch (1915, 1916, 1917), Melin (1923), Musso (1978), Oldroyd (1939), and Séguy (1927). Knutson (1972, 1976) provided more detail and a format when he described the pupal cases of Neomochtherus angustipennis (Hine) and Pseudorus distendens, respectively. This format was followed by Cezar & Lamas (2010), Dennis & Barnes (2012), Dennis et al. (2008a & b), Dennis & Knutson (1988), Dennis & Lavigne (1976a), and Weber & Lavigne (2004). Others, such as Daniels (1987), Musso (1978), and Scarbrough & Kuhar (1995) provided less complete descriptions. Dennis & Knutson (1988), and Knutson (1976) provided keys to the pupal cases of five subfamilies (Asilinae, Dasypogoninae, Laphriinae, Leptogastrinae, Megapodinae) of robber flies. A tentative key to the pupal cases of ten of the 14 subfamilies identified by Dikow (2009a & b) for Asilinae, Brachyrhopalinae, Dasypogoninae, Dioctriinae, Laphriinae, Leptogastrinae, Ommatiinae, Stenopogoninae, Stichopogoninae, Willistonininae) was presented by Dennis & Barnes (2011). There are no known pupal case descriptions for four subfamilies (Bathypogoninae, Phellinae, Tillobromatinae, Trigonomiminae). One of the main distinguishing characteristics of pupal cases is the arrangement of the integumental processes on the abdominal segments. Until Dennis et al. (2008a & b), these were designated as bristles, thorns, teeth, or spurs. These authors observed that some of the abdominal processes do not have a constricted base or area of integumental weakness. Other processes are constricted at the base and in a socket-like area of weakened integument that allows them to articulate. Melin (1923) commented that the dorsal processes on at least abdominal segment 1 are movable in the longitudinal axis and serve as levers to help the pupae move through the medium that they are in. Dennis et al. (2008a & b) found that the socketed processes moved easily which led them to believe that the processes on the abdominal segments are a combination of spines and spurs (Fig. 3) as defined by Comstock (1925) and Daly et al. (1998). A spine is a multicellular, rigid, immovable, thorn-like outgrowth of the cuticle that is not separated from it by a joint. A spur is a multicellular moveable process of the cuticle that is connected to the body wall by a joint, and has a socket or area of integumental weakness around its base. Some spines also might be bristle-like, but a bristle is defined as a unicellular macrotrichium or seta connected with nerves and surrounded at the base by a membranous ring or socket called an alveolus (Daly et al. 1998, McAlpine 1981). Typically, abdominal segment 1 on pupal cases has a dorsal transverse row of spurs, and segments 2–7 each have a dorsal transverse row of long spines alternating with short spurs. However, the known described pupal cases of Laphriinae species lack discernible spurs (Dennis et al. 2008a, Dennis & Barnes 2012). There are dorsolateral, lateral (posterior to the spiracles on the first 7 abdominal segments), and ventral bristle-like spines. Abdominal segment 8 has a distinctive arrangement of dorsal, dorsolateral, lateral, and ventral spurs and spines. Depending on the species, one or more of these groups of processes might be absent. On some species the spiracles can be seen on segment 8, but they are often located more dorsally than on the other abdominal segments. Abdominal segment 9 has a combination of dorsal posterolateral processes, ventral posterolateral processes and ventromedial processes, and sometimes a distinct arrangement of tubercles and callosities. Male pupal cases can often be distinguished from female cases by a pair of enlarged midventral callosities or tubercles.

Acknowledgments

We gratefully acknowledge the copies of pertinent literature provided by Drs. Marcela K. Castelo and Robert J. Lavigne. We also thank the anonymous reviewers for their constructive comments, and Neal Evenhuis for his fine editing of the manuscript.

Alcock, J. (1974) Observations on the behavior of Mallophora fautrix Osten Sacken (Diptera: Asilidae). Pan-Pacific Entomologist, 50, 68–72. Andersson, H. (1999) Red-listed or rare invertebrates associated with hollow, rotting, or sapping trees or polypores in the town of Lund. Entomologisk Tidskrift, 120, 169–183. [in Swedish with English summary] Artigas, J.N. & Papavero, N. (1988) The American genera of Asilidae (Diptera): Keys for identification with an atlas of female spermathecae and other morphological details. II. Key to the genera of Dasypogoninae Macquart, with description of new genera and species and new synonymies. Gayana Zoología, 52, 199–260. Baker, N.T. & Fischer, R.L. (1975) A taxonomic and ecologic study of the Asilidae of Michigan. The Great Lakes Entomologist, 8, 31–91. Barnes, J.K. (2008) Review of the genus Ceraturgus Wiedemann (Diptera: Asilidae) in North America north of Mexico. Zootaxa, 1766, 1–45. Barraclough, D.A. & Londt, J.G.H. (1985) Order Diptera. In: Scholtz, C.H. & Holm, E. (Eds.), Insects of Southern Africa, Butterworth, Durban, pp. 283–321. Barriga, T.J.E. (1990) Parasites and larval predators of Cerambycidae and Buprestidae (Coleoptera) of Chile. Revista Chilena de Entomología, 18, 57–59. Borkent, A. & Rotheray, G. (2009) Key to Diptera families – larvae. In: Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M., Woodley, N.E. & Zumbado, M. (Eds.), Manual of Central American Diptera. Vol. 1. National Research Council Press, Ottawa, pp. 157–191. Brindle, A. (1962) Taxonomic notes on the larvae of British Diptera, 10-The Asilidae. Entomologist, 95, 241–247. Brindle, A. (1968) Taxonomic notes on the larvae of British Diptera, No. 25, The larva and pupa of Dioctria rufipes (DeGeer) (Asilidae). Entomologist, 101, 213–216. Brindle, A. (1969) Taxonomic notes on the larvae of British Diptera, No. 26, The presumed larva of Dioctria oelandica (Linnaeus) (Asilidae). Entomologist, 102, 3–6. Bromley, S.W. (1946) Guide to insects of Connecticut. Part VI. The Diptera or true flies of Connecticut. Third Fascicle. Asilidae. Bulletin of the Connecticut State Geological and Natural History Survey, 69, 1–48. Brown, B.V., Marshall, S.A. & Wood, D.M. (2009) Natural history. In: Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M., Woodley, N.E. & Zumbado, M. (Eds.), Manual of Central American Diptera. Vol. 1. National Research Council Press, Ottawa, pp. 51–63. Bueno, V.H.P. (1987) [Biological aspects and daily rhythm of activities of Porasilus barbiellinii predator of the froghopper]. Pesquisa Agropecuaria Brasileira, 22, 903–915. [in Portuguese with English summary] Bullington, S.W. (1986) Two new genera related to Laphria Meigen (Diptera: Asilidae), with revisions of the included species in North America north of Mexico. Unpublished Doctoral Dissertation, University of Wyoming, Laramie, Wyoming, 275 pp. Bullington, S.W. (2011) The Laphriini pages. Available from: http://users.usachoice.net/~swb/laphriini/laphriini.htm (accessed 09 April 2011) Bybee, S.M., Taylor, S.D., Nelson, C.R. & Whiting, M.F. (2004) A phylogeny of robber flies (Diptera: Asilidae) at the subfamilial level: molecular evidence. Molecular Phylogenetics and Evolution, 30, 789–797. http://dx.doi.org/10.1016/S1055-7903(03)00253-7 Candan, S., Suludere, Z., Hasbenli, A., Çagiran, N., Lavigne, R. & Scarbrough, A. (2004a) Ultrastructure of the chorion of Dioctria flavipennis Meigen, 1820 (Diptera: Asilidae: Stenopogoninae) compared with those of 14 asilid species from the mid-Atlantic region of North America. Proceedings of the Entomological Society of Washington, 106, 811–825. Candan, S., Suludere, Z., Kalender, Y. & Eryilmaz, O. (2004b) Ultrastructure of the chorion of Echthistus cognatus (Loew, 1849) (Diptera, Asilidae). Ohio Journal of Science, 104, 93–96. Cannings, R.A. (1993) New synonymy of Rhadiurgus variabilis (Zetterstedt) (Diptera: Asilidae) with notes on morphology, natural history and geographical distribution. The Canadian Entomologist, 125, 337–354. Castelo, M.K. (2003) Comportamiento de localización y patrones de explotación de hospedadores (Coleoptera: Scarabaeidae) por el moscardón cazador de abejas Mallophora ruficauda (Diptera: Asilidae). Tesis doctoral. Laboratorio de Fisiología de Insectos, Departamento de Biodiversidad y Biología Experimental, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Buenos Aires, 169 pp. Castelo, M.K. & Capurro, A.F. (1999) Especificidad y densodependencia inversa en parasitoides con oviposición fuera del hospedador el caso de Mallophora ruficauda (Diptera: Asilidae) en la pampa Argentina. Universida de Belgrano, Serie Programa de Desarrollo Ambiental, 30, 3–18. Castelo, M.K. & Capurro, A.F. (2000) Especificidad y denso-dependencia inversa en parasitoides con oviposición fuera del hospedador: el caso de Mallophora ruficauda (Diptera: Asilidae) en la pampa Argentina. Ecología Austral, Asociación Argentina de Ecología, 10, 89–101. Castelo, M.K. & Corley, J.C. (2004a) Evaluación de la capacidad reguladora del mascardón cazador de abejas Mallophora ruficauda (Diptera: Asilidae) sobre los gusanos blancos del suelo (Coleoptera: Scarabaeidae). Revista de la Investigaciones Agropecuarias, 33, 59–78.

ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 15 Castelo, M.K. & Corley, J.C. (2004b) Oviposition behavior in the robber fly Mallophora ruficauda (Diptera: Asilidae). Annals of the Entomological Society of America, 97, 1050–1054. http://dx.doi.org/10.1603/0013-8746(2004)097[1050:OBITRF]2.0.CO;2 Castelo, M.K. & Corley, J.C. (2010) Spatial density-dependent parasitism and specificity in the robber fly Mallophora ruficauda (Diptera: Asilidae). Austral Ecology, 36, 72–81. http://dx.doi.org/10.1111/j.1442-9993.2009.02013.x Castelo, M.K. & Crespo, J.E. (2011) Host location in parasitoids with active searching larvae: the case of Mallophora ruficauda. In: Weiss, L.E. & Atwood, J.M. (Eds.), The biology of odors: sources, olfaction and response. Nova Science Publishers, Hauppage, New York, pp. 185–206. Castelo, M.K. & Lazzari, C.R. (2002) Orientación hacia el hospedador en larvas del moscardón cazador de abejas, Mallophora ruficauda (Diptera: Asilidae). V Congreso Argentino de Entomología, 394 pp. Castelo, M.K. & Lazzari, C.R. (2004) Host-seeking behavior in larvae of the robber fly Mallophora ruficauda (Diptera: Asilidae). Journal of Insect Physiology, 50, 331–336. http://dx.doi.org/10.1016/j.jinsphys.2004.02.002 Castelo, M.K., Ney-Nifle, M., Corley, J.C. & Bernstein, C. (2006) Oviposition height increases parasitism success by the robber fly Mallophora ruficauda (Diptera: Asilidae). Behavioral Ecology and Sociobiology, 61, 231–243. http://dx.doi.org/10.1007/s00265-006-0254-5 Castillo, E.E., Jerez, V. & Artigas, J.N. (1994) Egg chorionic microsculpture in robber flies (Diptera Asilinae, Dasypogoninae, Laphriinae and Stenopogoninae). Boletin de la Sociedad de Biologia de Concepción, Chile, 65, 107–116. [in Spanish] Cezar, L.A. & Lamas, C.J.E. (2010) Description of the larva and pupal case of Ommatius orenoquensis Bigot (Diptera, Asilidae, Ommatiinae). Revista Brasileira de Entomologia, 54, 28–31. http://dx.doi.org/10.1590/S0085-56262010000100002 Clements, A.N. (2000) A revision of Brachyrhopala Macquart, an Australian Region genus (Diptera: Asilidae). Invertebrate Taxonomy, 14, 77–114. Clements, A.N. & Bennett, F.D. (1969) The structure and biology of a new species of Mallophora Macq. (Diptera, Asilidae) from Trinidad, W.I. Bulletin of Entomological Research, 58, 455–463. http://dx.doi.org/10.1017/S0007485300057217 Clements, D.K. & Skidmore, P. (1998) The autecology of the hornet robberfly Asilus crabroniformis in Wales, 1997. Countryside Council for Wales Contract Science Report, 263, 63 pp. Clements, D.K. & Skidmore, P. (2002) The autecology of the hornet robberfly Asilus crabroniformis in Wales, 1997–1999. Countryside Council for Wales Contract Science Report, 525, 104 pp. Clements, D.K., Skidmore, P. & Denton, J.S. (1999) The autecology of the hornet robberfly Asilus crabroniformis L. in Wales, 1998. Countryside Council for Wales Contract Science Report, 344, 22 pp. Cole, F.R. (1969) The flies of western North America. University of California Press, Berkeley, 693 pp. Comstock, J.H. (1925) An Introduction to Entomology, second edition. The Comstock Publishing Company, Ithaca, New York, 1044 pp. Copello, A. (1927) Biología del moscardón cazador de abejas (Mallophora ruficauda, Wied). Argentina. Ministerio de Agricultura (Buenos Aires), Circular, 699, 1–19. Copello, A. (1942) Moscardón cazador de abejas “Mallophora ruficauda.” Revista Apicultura, Argentina, 19, 4–5; 219, 10–11; 200, 13–14; 221, 14; 222, 12–13. Crespo, J.E. (2011) Ecología u fisiología del comportamiento de localizacíon del hospedador en el parasitoide Mallophora ruficauda (Diptera: Asilidae). Unpublished Ph.D. Thesis, Faculty of Natural Sciences, University of Buenos Aires, Argentina, 135 pp. Crespo, J.E. & Castelo, M.K. (2008) The ontogeny of host-seeking behaviour in a parasitoid Diptera. Journal of Insect Physiology, 54, 842–847. http://dx.doi.org/10.1016/j.jinsphys.2008.03.002 Crespo, J.E. & Castelo, M.K. (2009) Insights to host discrimination and host acceptance behaviour in a parasitoid (Diptera: Asilidae): implications for fitness. Journal of Insect Physiology, 55, 1072–1078. http://dx.doi.org/10.1016/j.jinsphys.2009.08.002 Crespo, J.E. & Castelo, M.K. (2010) Life-history traits in parasitoid dipteran species with free-living and obligate parasitic immature stages. Physiological Entomology, 35, 160–167. http://dx.doi.org/10.1111/j.1365-3032.2010.00727.x Crespo, J.E., Lazzari, C.R. & Castelo, M.K. (2011) Orientation mechanisms and sensory organs involved in host location in a dipteran parasitoid larva. Journal of Insect Physiology, 57, 191–196. http://dx.doi.org/10.1016/j.jinsphys.2010.11.010 Daly, H.V., Doyen, J.T. & Purcell, A.H. (1998) Introduction to Insect Biology and Diversity, second edition. Oxford University Press, New York, 680 pp. Daniels, G. (1983) A new species of Neoaratus Ricardo (Diptera: Asilidae). Journal of the Australian Entomological Society, 22, 233–236. http://dx.doi.org/10.1111/j.1440-6055.1983.tb01884.x

16 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. Daniels, G. (1987) A revision of Neoratus Ricardo, with the description of six allied new genera from the Australian Region (Diptera: Asilidae: Asilini). Invertebrate Taxonomy, 1, 473–592. http://dx.doi.org/10.1071/IT9870473 Dekhtiarev, N.S. (1929) Notes on the scarabaeid beetles attacking vines in Ukraine. Bulletin of Entomological Research, 20, 95–98. http://dx.doi.org/10.1017/S0007485300020976 Dennis, D.S. (1979) Ethology of Holcocephala fusca in Virginia (Diptera: Asilidae). Proceedings of the Entomological Society of Washington, 81, 366–378. Dennis, D.S. (2012) Ethology of Proctacanthus brevipennis (Wiedemann, 1828) (Diptera: Asilidae) in northeastern Florida, U.S.A. Journal of the Entomological Research Society, 14, 91–109. Dennis, D.S. & Barnes, J.K. (2011) Tentative key to robber fly (Diptera: Asilidae) subfamilies based on pupal cases. Zootaxa, 3031, 37–46. Dennis, D.S. & Barnes, J.K. (2012) The pupal case of a Palearctic robber fly, Andrenosoma atrum (Linnaeus) (Diptera: Asilidae). Proceedings of the Entomological Society of Washington, 114, 16–23. http://dx.doi.org/10.4289/0013-8797.114.1.16 Dennis, D.S., Barnes, J.K. & Knutson, L. (2008a) Pupal cases of Nearctic robber flies (Diptera: Asilidae). Zootaxa, 1868, 1–98. Dennis, D.S., Barnes, J.K. & Knutson, L. (2008b). The pupal case of Mallophora atra Macquart (Diptera: Asilidae) from South America. Proceedings of the Entomological Society of Washington, 110, 234–241. http://dx.doi.org/10.4289/0013-8797-110.1.234 Dennis, D.S. & Knutson, L. (1988) Pupal cases of four species of robber flies from South America (Diptera: Asilidae). Annals of the Entomological Society of America, 81, 851–864. Dennis, D.S. & Lavigne, R.J. (1975) Comparative behavior of Wyoming robber flies II (Diptera: Asilidae). University of Wyoming Agriculture Experiment Station Science Monograph, 30, 1–68. Dennis, D.S. & Lavigne, R.J. (1976a) Descriptions and notes on the pupae and pupal cases of ten species of Wyoming robber flies (Diptera: Asilidae). Proceedings of the Entomological Society of Washington, 78, 277–303. Dennis, D.S. & Lavigne, R.J. (1976b) Ethology of Efferia varipes with comments on species coexistence (Diptera: Asilidae). Journal of the Kansas Entomological Society, 49, 48–62. Dennis, D.S. & Lavigne, R.J.(1976c) Ethology of Leptogaster parvoclava in Wyoming (Diptera: Asilidae). Proceedings of the Entomological Society of Washington, 78, 208–222. Dennis, D.S. & Lavigne, R.J. (1976d) Oviposition of Cyrtopogon inversus (Diptera: Asilidae). The Pan-Pacific Entomologist, 52, 288–290. Dennis, D.S. & Lavigne, R.J. (1979) Ethology of Machimus callidus with incidental observations on M. occidentalis in Wyoming (Diptera: Asilidae). The Pan-Pacific Entomologist, 55, 208–221. Dennis, D.S., Lavigne, R.J. & Bullington, S.W. (1986) Ethology of Efferia cressoni with a review of the comparative ethology of the genus (Diptera: Asilidae). Proceedings of the Entomological Society of Washington, 88, 42–55. Denton, J. & Tagg, D. (1995) Observations of oviposition by Asilus crabroniformis L. on Surrey heathland. Larger Brachycera Recording Scheme Newsletter, 13, 4. Dikow, T. (2003) Family group names in the Asilidae (Diptera). Studia Dipterologica, 10, 459–471. Dikow, T. (2009a) Phylogeny of Asilidae inferred from morphological characters of imagines (Insecta: Diptera: Brachycera: Asiloidea). Bulletin of the American Museum of Natural History, 319, 1–175. http://dx.doi.org/10.1206/603.1 Dikow, T. (2009b) A phylogenetic hypothesis for Asilidae based on a total evidence analysis of morphological and DNA sequence data (Insecta: Diptera: Brachycera: Asiloidea). Organisms, Diversity & Evolution, 9, 165–188. http://dx.doi.org/10.1016/j.ode.2009.02.004 Dikow, T. & Geller-Grimm, F. (2004) Information on robber fly phylogeny: introduction with a brief history of former studies. Available from: http://www.geller-grimm.de/genera19.htm (Accessed 9 May 2012) Esipenko, P.A. (1973) [The biology of Leptogaster cylindrica DeG. (Diptera, Asilidae) in the Khabarovsk Territory]. In: Ivliev, L.A. (Ed.), [Entomological Researches in the Far East. Issue 2. Diptera of the Far East]. Trudy-Biologo-pochvennogo- Instituta, Dal’nevostochnyl-Nauchnyl-Tsentr, Akademiya Nauk SSSR, 5, 130–134. [in Russian] Fisher, E.M. (1983) Pilica formidolosa (mosca asesina, robber fly). In: Janzen, D.H. (Ed.), Costa Rican Natural History. University of Chicago Press, Chicago, pp. 755–758. Fisher, E.M. (1986) A reclassification of the robber fly tribe Andrenosomini with a revision of the genus Dasyllis Loew (Diptera: Asilidae). Unpublished Ph.D. Thesis, University of California, Riverside, California, 361 pp. Fisher, E.M. (2009) Asilidae (Robber flies, assassin flies, moscas cazadoras, moscas ladronas). In: Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M., Woodley N.E., & Zumbado, M. (Eds.), Manual of Central American Diptera, Vol. 1. National Research Council Press, Ottawa, pp. 585–632. Foote, B.A. & Dennis, S. (1991) Asilidae (Asiloidea). In: Stehr, F.W. (Ed.), Immature Insects, Vol. 2. Kendall/Hunt Publishing Company, Dubuque, Iowa, pp. 778–780. Geller-Grimm, F. (2003) A world catalogue of the genera of the family Asilidae (Diptera). Studia Dipterologica, 10, 473–526. Geller-Grimm, F. (2004) Key to the subfamilies of Asilidae (Diptera) after Artigas and Papavero, 1988. Available from: http:// www.geller-grimm.de/subfam.htm (Accessed 8 May 2012)

ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 17 Geller-Grimm, F. (2011a) Information on robber flies, morphology, immature stages. Available from: http://www.geller- grimm.de/genera.htm (Accessed 4 November 2011) Geller-Grimm, F. (2011b) Notes on the biology of Dasypogon diadema. Available from: http://www.geller-grimm.de/dasy.htm (accessed 16 April 2011) Goot, V. van der (1969) Roof-en blaaskop-vliegentabel, 2nd ed. Nederlandse Jeugdbond voor Natuurstudie, ‘sGraveland, Netherlands, 20 pp. Green, D.M. (1998) Asilus crabroniformis hornet robber fly. Worcestershire Record, 5, 10. Groba, H.F. & Castelo, M.K. (2011) Chemical interaction between the larva of a dipteran parasitoid and its coleopteran host: a case of exploitation of the communication system during searching behavior. Bulletin of Entomological Research, 102, 1–9. http://dx.doi.org/10.1017/S0007485311000691 Harusawa, K. (2000) [Note on the ecology of Choerades komurae (Matsumura)]. Hana Abu, 10, 97. [in Japanese] Harusawa, K. (2001) [Observations on the oviposition and hatching of Promachus yesonicus Bigot, 1887 (Diptera: Asilidae)]. Hana Abu, 12, 43–46. [in Japanese with English summary] Harusawa, K. (2002) [Observations on the oviposition and hatching of Grypoctonus aino Speiser, 1928 and G. hatakeyamae (Matsumura, 1916) (Diptera: Asilidae)]. Hana Abu, 14, 13–16. [in Japanese] Hampshire Biodiversity Partnership (2001) Hornet robberfly Asilus crabroniformis species action plan. Biodiversity Action Plan for Hampshire, 2, 1–5. Hasbenli, A., Candan, A.S. & Alpay, N. (2006) A new species of Leptogaster Meigen (Diptera: Asilidae) from Turkey with egg and spermatheca structure. Zootaxa, 1267, 49–57. Hasbenli, A., Suludere, Z., Candan, S. & Bayrakdar, F. (2008) Chorionic structure of the eggs of five Laphriinae species (Diptera: Asilidae) from Turkey. Journal of the Entomological Research Society, 10, 47–60. Hauser, M. & Irwin, M.E. (2003) The Nearctic genus Ammonaios Irwin and Lyneborg 1981. Annals of the Entomological Society of America, 96, 738–748. Hayat, R. (1997) Prey of some robber flies (Diptera: Asilidae) in Turkey. Zoology in the Middle East, 15, 87–94. http://dx.doi.org/10.1080/09397140.1997.10637743 Hennig, W. (1952) Die Larvenformen der Dipteren. 3. Teil. Akademie-Verlag, Berlin, 628 pp. Hespenheide, H.A. (1989) Notes on the biology of Efferia wilcoxi (Bromley) (Diptera: Asilidae). Pan-Pacific Entomologist, 65, 375–380. Hinton, H.E. (1981) Biology of Insect Eggs, Vol II. Pergamon Press Ltd., Oxford, 778 pp. Holloway, G. J., Dickson, J.D., Harris, P.W. & Smith, J. (2003) Dynamics and foraging behaviour of adult hornet robberflies, Asilus crabroniformis: implications for conservation management. Journal of Insect Conservation, 7, 127–135. Howe, M. (2003) Hornet robberfly – Asilus crabroniformis. Lowland Grassland Lead Co-ordination Network, Information Bulletin, 5, unnumbered pages 11–12. Hubble, D. (2011) Is it a hornet? No, it’s Britain’s biggest fly. Available from: http://davehubbleecology.blogspot.com/2011/08/ is-it-hornet-no-its-britains-biggest.html (accessed 5 November 2011) Hudson, I. (1988) Oviposition by Machimus atricapillus. Larger Brachycera Recording Scheme Newsletter, 5, 2. Hull, F.M. (1962) Robber flies of the world: the genera of the family Asilidae. United States National Museum Bulletin, 224, 1–907. http://dx.doi.org/10.5479/si.03629236.224 Irwin-Smith, V. (1923) Studies in life histories of Australian Diptera Brachycera. Part II. Asilidae. No. 1. Catalogue of the species of Asilidae of which the earlier stages have been recorded, with literary index, and bibliography of biological literature. Proceedings of the Linnean Society of New South Wales, 48, 368–374. Johnson, P.E. (1976) A preliminary investigation of the behavior of adult Holcocephala abdominalis (Say) (Diptera: Asilidae) in Putnam County, Indiana; with a description of the larvae and eggs. Unpublished M.S. Thesis, DePauw University, Greencastle, Indiana, 108 pp. Keilin, D. (1944) Respiratory systems and respiratory adaptations in larvae and pupae of Diptera. Parasitology, 36, 1–66. http://dx.doi.org/10.1017/S0031182000011975 Knowles, L. & Hawkeswood, T.J. (1985) Records of predation on Australian Buprestidae and Scarabaeidae (Coleoptera) by Asilidae (Diptera). The Coleopterists Bulletin, 39, 354. Knutson, L. (1972) Pupa of Neomochtherus angustipennis (Hine), with notes on feeding habits of robber flies and a review of publications on morphology of immature stages (Diptera: Asilidae). Proceedings of the Biological Society of Washington, 85, 163–178. Knutson, L (1976) Key to subfamilies of robber flies based on pupal cases, with a description of the pupal case of Doryclus distendens (Asilidae: Megapodinae). Proceedings of the Biological Society of Washington, 88, 509–514. Knutson, L.V. & Murphy, W.L. (1990) Insects: Diptera (flies). In: Morse, R.A. & Nowogrodzki, R. (Eds.), Honey Bee Pests, Predators, and Diseases, second edition. Comstock Publishing Associates, Ithaca, New York, pp. 121–134. Kolomiets, N.G. (1975) [Robber flies (Diptera, Asilidae) of Siberia]. In: [Entomophagous Insects of the Far East, vol. 5]. Vladivostok. pp. 47–57. [in Russian] Krivosheina, N.P. (1973) [Larvae and pupae of robber flies from the genus Laphystia Lw. (Diptera, Asilidae)]. Zoologicheskii Zhurnal, 52, 457–459. [in Russian] Krivosheina, N.P. (1974) [Larvae of robber flies (Asilidae, Ctenotini) – root inhabitants of desert plants]. Biologicheski Nauki, 17, 16–21. [in Russian]

18 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. Krivosheina, N.P. (1975) [Specification of insect complexes living in rotting trunks of Manchurian ash trees]. Entomologicheskie issled na Dalnem Vostoke, Novosibirsk, 28, 89–96. [in Russian] Krivosheina, N.P. & Mamaev, B.M. (1973) [Larvae of assassin flies of the tribe Andrenosomini (Diptera, Asilidae) as xylophilous forms]. Biologicheskie Nauki, 16, 20–23. [in Russian] Krivosheina, N.P. & Mamaev, B.M. (1975a) Revision of materials on morphology and ecology of xylophyllic larvae of assassin flies of the Laphriini tribe (Diptera, Asilidae). Communication I. Genus Choerades Walk. Vestnik Zoologii, 2, 66–72. [in Russian] Krivosheina, N.P. & Mamaev, B.M. (1975b) Revision of materials on morphology and ecology of xylophillic assassin fly larvae of

the tribes Laphriini (Diptera, Asilidae). Communication II. Genus Laphria Mg. Vestnik Zoologii, 6, 46–50. [in Russian] Kurkina, L.A. (1979) [The biology of Machimus annulipes (Diptera, Asilidae).] Entomologicheskoe Obozrenie, 58, 57–63. [in Russian; English translation, 1979, Entomological Review, 58, 26–30] Lavigne, R.J. (1972) Ethology of Ablautus rufotibialis on the Pawnee Grasslands IBP site. Journal of the Kansas Entomological Society, 45, 271–274. Lavigne, R.J. (1979) Notes on the ethology of Efferia argyrogaster (Diptera: Asilidae) in Mexico. Proceedings of the Entomological Society of Washington, 81, 544–551. Lavigne, R.J. (1982) Ethology of Neoitamus vittipes (Diptera: Asilidae) in South Australia. Proceedings of the Entomological Society of Washington, 84, 617–627. Lavigne, R.J. (1984) Ethology of Neocerdistus acutangulatus (Diptera: Asilidae) in South. Australia. Proceedings of the Entomological Society of Washington, 86, 422–431. Lavigne, R.J. (1992) Ethology of Neoaratus abludo Daniels (Diptera: Asilidae) in South Australia, with notes on N. pelago (Walker) and N. rufiventris (Macquart). Proceedings of the Entomological Society of Washington, 94, 253–262. Lavigne, R.J. (1999) Bibliography update 1977–1995 for the Asilidae (Insecta: Diptera), including short translations from Japanese and Russian. University of Wyoming Agricultural Experiment Station College of Agriculture Science MonographSM-55, 91 pp. Lavigne, R.J. (2002) Evolution of courtship behaviour among the Asilidae (Diptera), with a review of courtship and mating. Studia Dipterologica, 9, 703–742. Lavigne, R.J. (2006a) A new species of Bathypogon Loew (Insecta: Diptera: Asilidae) from South Australia with notes on its behaviour. Studia Dipterologica, 13, 171–179. Lavigne, R.J. (2006b) Abundance and behaviour of robber flies (Insecta: Diptera: Asilidae) within Monarto Zoological Park. Final Report to the Projects Manager of Monarto Zoological Park, Goeff Brooks, 37 pp. Lavigne, R.J. (2011a) A new species of Cerdistus (Insecta: Diptera: Asilidae) from Monarto Zoological Park, South Australia with notes on its behaviour. The South Australian Naturalist, 85, 76–84. Lavigne, R.J. (2011b) Australian Asilidae: introduction, biology, eggs, larvae, pupae. (under eggs, larvae and pupae) Available from: http://sites.google.com/site/australianasilidae/introduction-2/biology (Accessed 4 November 2011) Lavigne, R.J., Barr, W.F. & Stephens, G.M. (2001) Checklist of the Asilidae of Idaho. The Idaho Entomology Group, 26 pp. Lavigne, R.J. & Bullington, S.W. (1984) Ethology of Laphria fernaldi (Back) (Diptera: Asilidae) in southeast Wyoming. Proceedings of the Entomological Society of Washington, 86, 326–336. Lavigne, R.J. & Dennis, D.S. (1975) Ethology of Efferia frewingi (Diptera: Asilidae). Annals of the Entomological Society of America, 68, 992–996. Lavigne, R.J. & Dennis, D.S. (1979) Notes on the ethology of Proctacanthus nearno (Diptera: Asilidae) in Wyoming. Proceedings of the Entomological Society of Washington, 81, 438–442. Lavigne, R.J. & Dennis, D.S. (1985) Ethology of three coexisting species of Efferia in Mexico (Diptera: Asilidae). Proceedings of the Entomological Society of Washington, 87, 146–160. Lavigne, R.J., Dennis, D.S. & Gowen, J.A. (2000) Asilid literature update 1956–1976, including a brief review of robber fly biology (Diptera: Asilidae). University of Wyoming, College of Agriculture Science Monograph SM-36, 92 pp. Lavigne, R.J. & Holland, F.R. (1969) Comparative behavior of eleven species of Wyoming robber flies (Diptera: Asilidae). University of Wyoming Agriculture Experiment Station Science Monograph, 18, 1–61. Lavigne, R.J. & Young, A. (2008) Distribution and behaviour of Cerdistus margitis(Walker, 1849) (Insecta: Diptera: Asilidae) in South Australia. The South Australian Naturalist, 82, 18–22. Lavigne, R.J., Young, A. & Glatz, R. (2009) Distribution and behaviour of Mauropteron pelago (Walker, 1849) (Insecta: Diptera: Asilidae) in South Australia. The South Australian Naturalist, 83, 69–77. Lawson, F.A. & Lavigne, R.J. (1984) Oviposition and eggs of an Australian robber fly, Neoaratus abludo Daniels (Diptera:

Asilidae). Proceedings of the Entomological Society of Washington, 86, 773–776. Lehr, P.A. (1958) [The biology of a robber fly feeding on acridids]. Doklady Kazakhstan Akademii Selskokhoz (Alma-Ata), 3, 16–23. [in Russian] Lehr, P.A. (1960) [The robber fly Stenopogon heteroneurus Macquart (Diptera, Asilidae), its behavior and feeding]. Trudy Kazakskogonauchno-Issledovatel’skogo Instituta Zashchita Rastenii, 6, 131–146. [in Russian] Lehr, P.A. (1964) [On the nutrition and significance of robber flies]. Trudy Kazakskogonauchno-Issledovatel’skogo Instituta Zashchita Rastenii, 8, 213–244 [in Russian] Lehr, P.A. (1969) An ecological and morphological analysis of the robber flies (Diptera, Asilidae). Communication II. Entomological Review, 48, 341–357.

ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 19 Lehr, P.A. (1977a) [On the robber flies Laphriinae: (Diptera, Asilidae) of middle Asia]. In: The Systematics and Evolution of Diptera Collection of Scientific Works, Academy of Sciences of the USSR. pp. 45–51. [in Russian] Lehr, P.A. (1977b) Robber flies from the subfamilies Atomosiinae and Laphriinae (Diptera, Asilidae) from Central Asia and Kazakhstan. Entomological Review, 56, 123–131. Lehr, P.A. (1986) Ecological and morphological trends in evolution of Eurasian Diptera (Asilidae). Unpublished paper given at the 1st International Congress of Dipterology, Budapest, 19 August 1986, 11 pp. Lehr, P.A. (1996) [Robber flies of subfamily Asilinae (Diptera, Asilidae) of Palearctic: Ecological and morphological analysis, taxonomy and evolution]. Dalnauka, Vladivostok, 184 pp. [in Russian] Lehr, P.A. (1998) Robber flies of the genus Cyrtopogon Loew, 1847 (Diptera, Asilidae) of the Palearctic. Entomological Review, 78, 1042–1066. Lim, K.P., Stewart, R.K. & Yule, W.N. (1981) Natural enemies of the common June beetle, Phyllophaga anxia (Coleoptera: Scarabaeidae) in southern Quebec. Annals of the Entomological Society of Quebec, 16, 14–27. Linsley, E.G. (1960) Ethology of some bee- and wasp-killing robber flies of southeastern Arizona and western New Mexico. University of California Publications in Entomology, 16, 357–392. Lloyd, D. (2001) Dispersal abilities and population structure of the adult hornet robberfly Asilus crabroniformis at Caeau Bwich SSSI. Countryside Council of Wales Contract Science Report, 458, 39 pp. Londt, J.G.H. (1979) Afrotropical Asilidae (Diptera) 3. The tribe Stichopogonini Annals of the Natal Museum, 23, 833–854. Londt, J.G.H. (1986) Gibbasilus arenaceus, a new genus and species from western Cape Province of South Africa (Diptera: Asilidae: Asilinae). Journal of the Entomological Society of Southern Africa, 49, 1–6. Londt, J.G.H. (1987) Notes on the biology and immature stages of Neolophonotus clichaetus Hull in South Africa. Journal of the Entomological Society of Southern Africa, 50, 427–434. Londt, J.G.H. (1989) Afrotropical Asilidae (Diptera) 18. The genus Smeryngolaphria Hermann, 1912 (Laphriinae). Annals of the Natal Museum, 30, 147–157. Londt, J.G.H. (1991) Afrotropical Asilidae (Diptera) 21. Observations on the biology and immature stages of Damalis femoralis Ricardo, 1925 (Trigonomiminae). Annals of the Natal Museum, 32, 149–162. Londt, J.G.H. (1992) Afrotropical Asilidae (Diptera) 22. The genus Scylaticus Loew, 1858 (Stenopogoninae). Annals of the Natal Museum, 33, 95–187. Londt, J.G.H. (1993) From what- to where, when, why and who: habitat characterization of robber flies (Diptera: Asilidae). In: Proceedings of the Ninth Entomological Congress Organized by the Entomological Society of Southern Africa. Johannesburg, 28 June–1 July 1993, p. 68. Londt, J.G.H. (1994) Afrotropical Asilidae (Diptera) 26. Ethological observations, and a possible ecological classification based on habitats. Annals of the Natal Museum, 35, 97–122. Londt, J.G.H. & Tsacas, L. (1987) The genus Congomochtherus Oldroyd, 1970 (Diptera, Asilidae, Asilinae). Annals of the Entomological Society of France (N.S.), 23, 29–42. Malloch, J.R. (1915) Some additional records of Chironomidae for Illinois and notes on other Illinois Diptera. Bulletin of the Illinois State Laboratory of Natural History, 11, 305–363. Malloch, J.R. (1916) A comparison of the pupae of Promachus vertebratus and Promachus fitchii. Bulletin of the Brooklyn Entomological Society, 11, 66–68. Malloch, J.R. (1917) A preliminary classification of Diptera, exclusive of pupipara, based upon the larval and pupal characters, with keys to imagines in certain families. Part I. Bulletin of the Illinois State Laboratory of Natural History, 12, 161–409. Martin, C.H. (1971) A review of the genus Leptopteromyia in the Western Hemisphere. The Pan-Pacific Entomologist, 47, 264–270. McAlpine, J.F. (1981) Morphology and terminology – Adults. In: McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R. & Wood, D.M. (coords.), Manual of Nearctic Diptera, Vol. 1. Research Branch, Agriculture Canada, Monograph, 27, 1–674. Melin, D. (1923) Contributions to the knowledge of the biology, metamorphosis and distribution of the Swedish asilids in relation to the whole family of asilids. Zoologiska Bidrag från Uppsala, 8, 1–317. Moore, K.M. (1963) Observations on some Australian forest insects. 15. Some mortality factors of Phoracantha semipunctata (F.) (Coleoptera: Cerambycidae). Proceedings of the Linnean Society of New South Wales, 88, 221–229. Moore, K.M. (1972) Observations on some Australian forest insects. 25. Additional information on some parasites and predators of longicorns (Cerambycidae: Phoracanthini). Australian Zoologist, 17, 26–29. Mukaigawa, Y. (1920) [Adult emergence of Promachus yesonicus Bigot]. Konchu Sekai (The Insect World), Gifu, Japan, 24, 352–353. [in Japanese] Musso, J.J. (1972) Observations sur les moeurs prédatrices de quelques Asilides [Dipt. Brachycera] de la Basse-Provence. Annals de la Société Entomologique de France, (N.S.) 8, 409–421. Musso, J.J. (1974) Éthologie. – Observations sur le comportement et le régime alimentaire des larves d’Asilides (Diptera, Brachycera). Comptes Rendus de l’Academie des Sciences Paris, 279 (Serie D), 1309–1312. Musso, J.J. (1978) Recherches sur le développement, la nutrition et l’écologie des Asilidae (Diptera-Brachycera). Doctoral Dissertation, Université de Droit, d’Economie et des Sciences d’Aix-Marseille, 312 pp. Musso, J.J. (1981a) Ecologie des Asilidae (Diptera Brachycera) I – Répartition en Basse Provence, périodes de volet longévité imaginable. Ecologia Mediterrana; Revue d’Ecologie Terrestre et Liminique, 7, 99–112.

20 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. Musso, J.J. (1981b) Morphology and development of the immature stages of some robber flies. (Diptera: Brachycera: Asilidae). Entomologia Generalis, 7, 89–104. Musso, J.J. (1983) Nutritive and ecological requirements of robber flies (Diptera: Brachycera: Asilidae). Entomologia Generalis, 9, 35–50. Musso, J.J. & Garcin, J.E. (1975) Recherches préliminaires sur le régime alimentaire des larves d’asilides (Diptera – Brachycera). Bulletin de la Société Zoologique de France, 100, 313–322. Nagatomi, A., Ohishi, H. & Yang D. (2002) Review of the genera of Leptogastrinae (Diptera, Asilidae) through the literature. The Kagoshima University Museum Monographs, 1, 111 pp. Nawa, U. (1899) [Egg mass of Promachus yesonicus Bigot]. Konchu Sekai (The Insect World), Gifu, Japan, 3, 314. [in Japanese] Ohno, M. (1955) [Notes on the predacious insects attacking Chrysomelidae in Japan]. Akitu, 4, 60–65. [in Japanese] Ohishi, H. (1998) [On eggs of Promachus yesonicus (Diptera, Asilidae)]. Hana Abu, 6, 32. [in Japanese] Oldroyd, H. (1939) A note on the pupa of Andrenosoma albopilosumVilln. (Diptera: Asilidae). Proceedings of the Royal Entomological Society of London, Series A, 14, 7–8. http://dx.doi.org/10.1111/j.1365-3032.1939.tb00014.x Oldroyd, H. (1969) Family Asilidae. In: Handbook for the Identification of British Insects. Diptera Brachycera Section (a) Tabanoidea and Asiloidea. 9 (4), 69–93. Papavero, N. (1973) Studies of Asilidae (Diptera) systematics and evolution. I. A preliminary classification in subfamilies. Arquivos de Zoologia (São Paulo), 23, 217–274. Peterson, A. (1977) Larvae of Insects: An Introduction to Nearctic Species, sixth edition. Edward Brothers, Inc., Ann Arbor, Michigan, 416 pp. [Asilidae, pp. 236–237, 282, Figs. D-10 A–G.] Planning Department, Bridgend County Borough Council & David Clements Ecology (2002) A Local Biodiversity Action Plan for Bridgend County Borough, Volume 2, Species and habitat action plans. Bridgend Biodiversity Partnership, Bridgend County Borough, UK, 134 pp. Rabinovich, M. & Corley, J.C. (1997) An important new predator of honey bees – the robber fly Mallophora ruficauda Wiedemann (Diptera, Asilidae) in Argentina. American Bee Journal, 137, 303–306. Rikhter, V.A. (1964) Zoogeographic features of the robber fly fauna (Diptera, Asilidae) of the Caucasus. Entomological Review, 43, 170–176. Rogers, L.E. & Lavigne, R.J. (1972) Asilidae of the Pawnee National Grasslands, in northeastern Colorado. Agricultural Experiment Station, University of Wyoming, Laramie, Science Monograph, 25, 35 pp. Scarbrough, A.G. (1978) Ethology of Cerotainia albipilosa Curran (Diptera: Asilidae) in Maryland: courtship, mating and oviposition. Proceedings of the Entomological Society of Washington, 80, 179–190. Scarbrough, A.G. (1981) Ethology of Eudioctria tibialis Banks (Diptera: Asilidae) in Maryland: reproductive behavior. Proceedings of the Entomological Society of Washington, 83, 432–443. Scarbrough, A.G. & Kuhar, T.P. (1992) Biodiversity of robber flies in Maryland and Delaware: Part I. Subfamily Leptogastrinae. Maryland Naturalist, 36, 13–38. Scarbrough, A.G. & Kuhar, T.P. (1995) Pupal cases of Machimus Loew in Maryland (Diptera: Asilidae). Maryland Entomologist, 4, 15–22. Scarbrough, A.G. & Sipes, G. (1973) The biology of Leptogaster flavipes Loew in Maryland (Diptera: Asilidae). Proceedings of the Entomological Society of Washington, 75, 441–448. Schreiber, E.T. & Lavigne, R.J. (1986) Ethology of Asilus gilvipes (Hine) (Diptera: Asilidae) associated with small mammal burrows in southeastern Wyoming. Proceedings of the Entomological Society of Washington, 88, 711–719. Séguy, E. (1927) Diptères (Brachycères) (Asilidae). Faune de France, 17, 190 pp. Shelly, T.E. (1985a) Ecological comparisons of robber fly species (Diptera: Asilidae) coexisting in a Neotropical forest. Oecologia, 67, 57–70. http://dx.doi.org/10.1007/BF00378452 Shelly, T.E. (1985b) Observations on the reproductive behavior of two Neotropical robber fly species (Diptera: Asilidae). Proceedings of the Entomological Society of Washington, 87, 253–254. Skidmore, P. (1966) Asilidae (Dipt.) of northern England. The Entomologist’s Record, 78, 230–235. Skidmore, P. (1991) Insects of the British cow-dung community. Field Studies Council Occasional Publication, 21, 1–166. Skidmore, P. (1997a) Survey of the insect fauna of the dung of ungulates on Exmoor. Exmoor National Park Internal Report, 46 pp. Skidmore, P. (1997b) The status of the hornet robberfly Asilus crabroniformis in South Wales. Countryside Council for Wales Internal Report, 33 pp. Skidmore, P. & Grayson, A. (1995) Assassin-flies or robber-flies (Diptera: Asilidae) of Yorkshire, with notes on records from other areas of northern England. Naturalist, 120, 143–148. Smith, J. (2000) The hornet robber fly Asilus crabroniformis: land use and livestock grazing regimes at sites in England. English Nature Research Reports, 387, 30 pp. Smith, K.G.V. (1989) An introduction to the immature stages of British flies. Handbook for the Identification of British Insects, 10, 280 pp.

ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 21 Stubbs, A.E. (1997) Observations on egg laying by the robberfly Machimus rusticus (Diptera: Asilidae). Larger Brachycera Recording Scheme Newsletter, 15, 9–10. Stubbs, A.E. & Drake, M. (2001) British soldierflies and their allies. An illustrated guide to their identification and ecology, covering all flies (Diptera) in the families Acroceridae, Asilidae, Athericidae, Bombyliidae, Rhagionidae, Scenopinidae, Stratiomyidae, Tabanidae, Therevidae, Xylomyidae and Xylophagidae. British Entomological and Natural History Society, Reading, 512 pp. Suludere, Z., Candan, S., Kalender, Y. & Hasbenli, A. (2000) Ultrastructure of the chorion of Machimus rusticus (Meigen, 1820) (Diptera, Asilidae). Journal of the Entomological Research Society, 2, 63–71. Teskey, H.J. (1976) Diptera larvae associated with trees in North America. Entomological Society of Canada Memoir, 100, 53 pp. http://dx.doi.org/10.4039/entm108100fv Uffen, R. (1993) Oviposition of Eutolmus rufibarbis. Larger Brachycera Recording Scheme Newsletter, 10, 7. Veen, M. van (1984) [Oviposition by Philonicus albiceps (Meigen) (Diptera: Asilidae)]. Entomologische Berichten, 44, 56. [in Dutch] Webber, S. & Doyle, J. (2007) Meadow Habitat Action Plan. Surrey Biodiversity Partnership, Surrey, UK, 36 pp. Weber, G.S. & Lavigne, R. (2004) Notes on the behaviour of Blepharotes coriarius (Wiedemann, 1830) (Diptera: Asilidae) with a description of the pupal case. Studia Dipterologica, 11, 13–21. Wei, X., Lu, X., Men, Z. & Ma, H. (1990) Field distribution of Promachus yesohicus [Dip.: Asilidae] larvae and their control effect on white grubs. Chinese Journal of Biological Control, 6, 183. Wei, X., Xu, X. & DeLoach, C. Jr. (1995) Biological control of the white grubs (Coleoptera: Scarabaeidae) by larvae of Promachus yesonicus (Diptera: Asilidae) in China. Biological Control, 5, 290–296. http://dx.doi.org/10.1006/bcon.1995.1036 Wood, G.C. (1981) Asilidae. In: McAlpine, J.F., Peterson, B.V., Shewell, G.E., Vockeroth, J.R. & Wood, D.M. (coords.), Manual of Nearctic Diptera, Vol. 1. Research Branch, Agriculture Canada, Monographs, 27, 549–573. Worcestershire Biodiversity Partnership (2008) Hornet robberfly Asilus crabroniformis species action plan. Worcestershire Biodiversity Action Plan 2008 S17 Hornet Robberfly SAP, 6 pp. Yonetsu, A. (1998) [Observations on the oviposition of Promachus yesonicus Bigot]. Hana Abu, 5, 46. [in Japanese] Zuska, J. (1972) Roupci - dravci mezi dvoukřídlými. ( Asilidae - predators among flies). Živa, 20, 221–222.

TABLE 1. Comparison of some subfamily treatments of Asilidae (source Dennis et al. 2008a). Subfamily Hull (1962), Papavero Lehr (1969, Geller-Grimm Artigas & Papavero Dikow Wood (1981) (1973) 1977b, 1996) (2003, 2004), (1988), Bybee et al. (2009b) Dikow (2003) (2004), Dikow & Geller-Grimm (2004) Apocleinae - 1 XXXX - Asilinae X X X X X X Atomosinae - - X - - - Bathypogoninae - - - - - X Brachyrhopalinae - - - - - X Dasypogoninae X X X X X X Dioctriinae - - X X - X Laphriinae X X X X X X Laphystiinae - X X X X - Leptogastrinae X - X X X X Megapodinae X - X - - - Ommatiinae - X X X X X Phellinae - - - - - X Stenopogoninae - X X X X X Stichopogoninae - - - X X X Tillobromatinae - - - - - X Trigonomiminae - X X X X X Willistonininae - - - - - X

1 X, subfamily recognized by author(s); -, subfamily not recognized.

22 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 23 24 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 25 26 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 27 28 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 29 30 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 31 32 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 33 34 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 35 36 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 37 38 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 39 40 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 41 42 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 43 44 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 45 46 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 47 TABLE 3. Biology of eggs of robber flies (“?” indicates that the oviposition location is speculative based on the morphology of the ovipositor; “-” indicates no information). Subfamily/Genus Oviposition Site Number of Eggs Development per Mass Period (days) Asilinae Alcimus Loew, 1848 in soil? - - Antipalus Loew, 1849 in soil - 7–8 in laboratory Antiphrisson Loew, 1849 in soil - - Asilus Linnaeus, 1758 in soil; onto vegetation near soil 1 10–14 surface; vegetation litter; in or onto dry cow and horse dung; mounds of rabbit pellets; sheep dung? Astochia Becker, 1913 vegetation? - - Blepharotes Duncan, 1840 dead gum tree or in soil at tree base multiple eggs deposited - individually Colepia Daniels, 1987 on plant stem; seed heads or ears of 1–9 6–7 wild wheat Congomochtherus Oldroyd, in soil? - - 1970 Dasophrys Loew, 1858 vegetation; trees - - Dicropaltum Martin, 1975 vegetation; between sheaths and stem or 1–2 - in seed head of grass Dolopus Daniels, 1987 dead vegetation; fern frond 1–4 - Dysclytus Loew, 1858 vegetation? - - Dysmachus Loew, 1860 into vegetation; grass spikelets - - Dystolmus Lehr, 1996 into vegetation - 6 in laboratory Echthistus Loew, 1849 cotton batting in laboratory 1 - Efferia Coquillett, 1893 (Efferia in or onto vegetation; in soil 1–90; mostly 2–19 - Group) Eremisca Hull, 1962 in soil - - Eutolmus Loew, 1848 into vegetation; leaf sheaths of grasses 11 - Heligmonevra Bigot, 1858 vegetation? - - Hippomachus Engel, 1927 vegetation; trees 4–6 - Hoplopheromerus Becker, 1925 vegetation? soil? - - Machimus Loew, 1849 vegetation; dropped onto vegetation or 1–7 1–7 in laboratory soil; grass seed head; tree trunks and dead wood; cotton batting in laboratory Mallophora Macquart, 1838 onto vegetation; wire fences 56–729; average 328 in 2–8 egg mass Mauropteron Daniels, 1987 dropped onto vegetation? or soil - - Megaphorus Bigot, 1857 onto vegetation 189–390 - Neoaratus Ricardo, 1913 onto vegetation 20–42 - Neocerdistus Hardy, 1926 in soil 1 - Neoepitriptus Lehr, 1992 in soil; grass spikelets - - (Wiedemann in Meigen, 1820) Neoitamus Osten Sacken, 1878 in vegetation; buds at apices of lower 3- tree branches Neolophonotus Engel, 1925 vegetation; trees; soil? 23–73 5–6 ...... continued on the next page

48 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. TABLE 3. (Continued) Subfamily/Genus Oviposition Site Number of Eggs Development per Mass Period (days) Neomochtherus Osten Sacken, in vegetation? - - 1878 Philodicus Loew, 1847 in soil? - - Philonicus Loew, 1849 in soil - - Pamponerus Loew, 1849 dropped to soil 3 - Porasilus Curran, 1934 onto vegetation 32–266 6–9 Proctacanthella Bromley, 1934 in soil 2 - Proctacanthus Macquart, 1838 in soil 2–14 - Promachus Loew, 1848 in vegetation; dead branch; soil? 2–31 5–15 Rhadiurgus Loew, 1849 vegetation; soil; mosses; pine needles; 1- twigs on soil Satanas Jacobson, 1901 in soil 1–8 6–8 Synolcus Loew, 1858 vegetation; trees - - Triorla Parks, 1968 vegetation 2–19 -

Bathypogoninae Bathypogon Loew, 1851 soil 7+ -

Brachyrhopalinae Brachyrhopala Macquart, 1847 probably vegetation (e.g., Acacia spp.) - - Cyrtopogon Loew, 1847 forest litter; soil 2–4 - Heteropogon Loew, 1847 debris on soil - - Holopogon Loew, 1847 in soil 6–9 - Rhabdogaster Loew, 1858 in soil? - -

Dasypogoninae Comantella Curran, 1923 in soil 1 - Dasypogon Meigen, 1803 in soil 1–6 in “sand balls or 4–17 in laboratory cocoons” Diogmites Loew, 1866 in soil 1–11 - Eucyrtopogon Curran, 1923 in soil? - - Leptarthrus Stephens, 1829 onto vegetation; dead branch on soil - - Pegesimallus Loew, 1858 in soil - -

Dioctriinae Dioctria Meigen, 1803 drop onto soil 1–18 18 Eudioctria Wilcox & Martin, vegetation; in soil 1 - 1941

Laphriinae Afromelittodes Oldroyd & dead wood? - - Bruggen, 1963 ...... continued on the next page

ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 49 TABLE 3. (Continued) Subfamily/Genus Oviposition Site Number of Eggs Development per Mass Period (days) Andrenosoma Rondani, 1856 in dead wood; smoldering trees from - 50–56 in laboratory fires Anypodetus Hermann, 1907 in soil? - - Cerotainia Schiner, 1868 onto vegetation 3 - Cerotainiops Curran, 1930 near base of small shrubs? - - Choerades Walker, 1851 in trees; tree stumps; wood dust beneath 2–12 - pine trees; litter; wood crevices or holes? dropped to soil? Ctenota Loew, 1873 in soil? - - Dasyllina Bromley, 1935 wood crevices or holes? - - Dasyllis Loew, 1851 nest entrance in dead trees or soil of -- Eulaema spp. bees? female observed probing with ovipositor fallen dead branch of Tetragastris panamensis (Engl.) Kuntze Gerrolasius Hermann, 1920 dropped onto soil - - Goneccalypsis Hermann, 1912 dropped onto soil? - - Hoplistomerus Macquart, 1838 dung? in soil? - - Hyperechia Schiner, 1866 in vegetation; dead wood - - Lampria Macquart, 1838 decaying log - - Laphria Meigen, 1803 in dead trees, stumps, dead wood; exit 3- holes of lignicolous insects Tribe Laphriini in dead tree wood, fallen tree trunks - - Laphystotes Oldroyd, 1974 in soil? - - Laxenecera Macquart, 1838 dropped onto soil - - Loewinella Hermann, 1912 dropped onto soil? - - Notiolaphria Londt, 1977 vegetation? dead wood? - - Nusa Walker, 1851 in soil? - - Pogonosoma Rondani, 1856 vegetation? dead wood? - - Proagonistes Loew, 1858 dead wood? - - Prytania Oldroyd, 1974 on soil? - - Smeryngolaphria Hermann, decaying log - - 1912 Storthyngomerus Hermann, wood crevices or holes? - - 1919 Trichardis Hermann, 1906 in soil? - -

Leptogastrinae all dropped onto soil? Ammophilomima Enderlein, dropped onto soil? - - 1914 Beameromyia Martin, 1957 dropped onto soil singly - Dolichoscius Janssens, 1953 dropped onto soil? - - Euscelidia Westwood, 1849 dropped onto soil? - - Lasiocnemus Loew,1851 dropped onto soil - - ...... continued on the next page

50 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. TABLE 3. (Continued) Subfamily/Genus Oviposition Site Number of Eggs Development per Mass Period (days) Leptogaster Meigen, 1803 dropped onto soil 1–8 singly; average 4 9–14 Psilonyx Aldrich, 1923 dropped onto soil singly - Tipulogaster Cockerell, 1913 dropped onto soil singly -

Ommatiinae all dropped onto soil? Ommatius Wiedemann, 1821 dropped onto soil - -

Stenopogoninae all in soil? - - Acnephalum Macquart, 1838 in soil? - - Afroholopogon Londt, 1994 in soil - - Agrostomyia Londt, 1994 in soil - - Ancylorhynchus Berthold in in soil? - - Latrielle, 1827 Gonioscelis Schiner, 1866 in soil? - - Habropogon Loew, 1847 in soil - - Microstylum Macquart, 1838 in soil? - - Ospriocerus Loew, 1866 in soil 7 - Pycnomerinx Hull, 1962 in soil - - Scleropogon Loew, 1866 in soil 3–9 - Scylaticus Loew, 1858 in soil 7–15 - Sisyrnodytes Loew, 1856 in soil? - - Stenopogon Loew, 1847 soil 7–20 -

Stichopogoninae all in soil? Lasiopogon Loew, 1847 in soil - - Stichopogon Loew, 1847 in soil - -

Tillobromatinae Hypenetes Loew, 1858 in soil? - - Lycostommyia Oldroyd, 1980 in soil? - -

Trigonomiminae all dropped onto soil? - - Damalis Fabricius, 1805 dropped onto soil 326 in vial 14 Holcocephala Jaennicke, 1867 onto vegetation? dropped in flight? 3–46 in vial; probably 31–32 deposits individual eggs Oligopogon Loew, 1847 in soil? - - Rhipidocephala Hermann, 1926 dropped onto soil? - -

Willistonininae Ablautus Loew, 1866 in soil 1 -

ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 51 52 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 53 54 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 55 56 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 57 58 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 59 60 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 61 62 · Zootaxa 3673 (1) © 2013 Magnolia Press DENNIS ET AL. ASILIDAE IMMATURES BIOLOGY AND MORPHOLOGY Zootaxa 3673 (1) © 2013 Magnolia Press · 63 TABLE 6. Number of genera and species for which biological and morphological data have been published on the immature stages of robber flies. Subfamily Information on Immature Stages Total Number of Valid Genera Worldwide Genera Species Asilinae 50 121 174 Bathypogoninae 1 1 2 Brachyrhopalinae 6 16 17 Dasypogoninae 11 23 65 Dioctriinae 2 9 11 Laphriinae 31 89 109 Leptogastrinae 12 13 18 Ommatiinae 1 3 8 Phellinae 0 0 3 Stenopogoninae 14 15 108 Stichopogoninae 2 3 11 Tillobromatinae 2 0 3 Trigonomiminae 4 4 11 Willistonininae 1 2 5 Total 137 299 545