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Manual of North American Agromyzidae (Diptera, Schizophora), with revision of the fauna of the “Delmarva” states
expand article infoOwen Lonsdale
‡ Agriculture & Agri-Food Canada, Ottawa, Canada
Open Access

Abstract

Тhe agromyzid (Diptera: Schizophora: Agromyzidae) fauna of America north of Mexico is described in the first part of this publication, including a genus key and discussions on morphology, life history and classification. The second part is a species-level revision of the family in the “Delmarva” states of the United States of America, that is, of the District of Columbia and the surrounding states of Delaware, Maryland, Virginia and West Virginia. The fauna of this region includes 156 species.

This study presents 346 new state and provincial records and 23 new country records, two of which are new continental records (Agromyza abiens Zetterstedt and A. apfelbecki Strobl). Liriomyza endiviae Hering is no longer considered to occur in North America. Fifteen species are newly described: Agromyza echinalis sp. nov., Melanagromyza brunkei sp. nov., M. eoflacensis sp. nov., M. glyptos sp. nov., M. rutella sp. nov., Ophiomyia capitolia sp. nov., O. cuprea sp. nov., O. galiodes sp. nov., O. heleios sp. nov., O. kalia sp. nov., O. laticolis sp. nov., Cerodontha (Poemyza) ungula sp. nov., Phytomyza avicursa sp. nov., P. catenula sp. nov., and P. winkleri sp. nov. Four new species-level synonyms and one genus-level synonym are provided: Agromyza marmorensis Spencer syn. nov. is included as a synonym of A. aristata Malloch; Melanagromyza fastosa Spencer, syn. nov. is included as a junior synonym of Ophiomyia tiliae (Couden); Melanagromyza verbesinae Spencer is considered a synonym of M. vernoniana Steyskal; Phytomyza ranunculoides Spencer, syn. nov. is included as a junior synonym of Phytomyza loewii Hendel; the genus Liomycina Enderlein, syn. nov. is included as a junior synonym of Phytobia Lioy. Ophiomyia ultima (Spencer), comb. nov. is recombined from Melanagromyza. Euhexomyza albicula Spencer, stat. reinst., comb. nov. is resurrected from synonymy with E. winnemanae (Malloch). New host records are given.

Keywords

Agromyzidae, manual, Nearctic, new species, recombination, revision, synonymy

Introduction

As an agriculturally and ecologically important group, it is critical to understand the systematics, biology, distribution, and impact of species in the family Agromyzidae, a ubiquitous and diverse acalyptrate family of true flies (Diptera: Schizophora). While comprehensive sources on the North American fauna, such as Spencer (1969, 1987) and Spencer and Steyskal (1986b), already discuss these topics, important studies and discoveries have appeared since the time of those publications and it is necessary to provide updates.

Scope of the present study

At higher taxonomic levels, the present work examines those subfamilies and genera occurring in Canada and the continental United States (that is, the Nearctic Region north of Mexico), providing diagnoses and brief discussions on their relationships, classification, life history and diversity. To facilitate generic identification, an identification key is provided, accompanied by illustrations of the highly informative and diagnostic male genitalia, and photographs of live and preserved specimens. Discussions include the results of recent revisionary and phylogenetic studies, including redefinitions of Liriomyza Mik (Lonsdale 2011, 2017), Phytomyza Fallén (Winkler et al. 2009), Nemorimyza Frey (Zlobin 1996) and members of the Ophiomyia genus group (Lonsdale 2014), among others.

The economic importance of the family was overviewed by Spencer (1973). An update on many of the most agriculturally significant species, including their hosts and methods of control, is currently being prepared (Lonsdale et al., in manuscript) and need not be discussed here.

At the species level, this mostly collection-based study revises the fauna of the “Delmarva” states (discussed below), and is intended as a regional snapshot of the family. It will contribute to a more accurate understanding of the group when used alongside other comprehensive regional works such Griffiths’ series on the boreal Phytomyza, Lonsdale and Scheffer’s (2011) revision of the Nearctic holly leaf miners, Boucher’s Nearctic revisions (2002, 2004, 2008), Frick’s (1956) Calycomyza Hendel revision, Lonsdale’s (2011, 2017) Liriomyza revisions, Shi and Gaimari’s (2015) revision of the Californian Melanagromyza Hendel, the primarily northeastern United States surveys of Eiseman et al. (2018–2021), the catalogues compiled by Frick (1952a, 1959, 1965), and of course, Spencer (1969) and Spencer and Steyskal (1986b). The latter two works revised all Agromyzidae known to occur in Canada and Alaska, and the lower continental United States, respectively, and have served as two of the most useful stand-alone resources for the family but need updating.

More broadly, the author hopes that this manual will generate more interest in this relatively difficult and neglected family in North America and facilitate easier identifications, thereby encouraging collectors to gather more field data, preserve more specimens, and explore parts of North America where agromyzids are largely unknown and undescribed. This is an exciting prospect considering that relatively little is known of the North American Agromyzidae outside of California and the northeast, and even in these regions, many new discoveries continue to accumulate. Furthermore, while the morphology of most species in the family can indeed often be accurately characterised as small, somewhat monotonous in appearance (at least externally) and sometimes difficult to work with, it is hoped that the photographs and illustrations given here portray the hidden charm of these animals, especially when considering the often striking and elaborate male genitalia.

The “Delmarva” States

The name “Delmarva”, as used here, is an older collective term referring to the District of Columbia and the surrounding states of Delaware, Maryland, Virginia, and West Virginia (Figs 5, 6). Its modern usage is restricted to the Delmarva Peninsula, flanked by the Delaware and Chesapeake Bays. Defining characteristics of these states (and District), including geography, are discussed in Mathis and Foster (2007) and Murphy et al. (2018). These authors also provide interesting historical context, including background on a series of papers examining the dipteran fauna of the greater Washington, D.C. area that eventually became treatments of individual families in the Delmarva states. The present work is a continuation of this series.

Agromyzidae of the Delmarva states

All species of Agromyzidae verified as being present in the Delmarva states are here keyed, described and illustrated. It is likely that the fauna of the region is much larger than represented however, if one extrapolates the known distribution of other northeastern species and their host plants, and considers the likelihood of there being many more species new to science yet to be discovered. Recent publications have illustrated that undescribed taxa are still in abundance in the northeast despite the long history of research there. Future discoveries will also certainly include taxa previously known from more distant localities that represent either recent introductions of invasive species, or previously unknown populations of more widely distributed species. These assumptions are supported by comparison to the more diverse and recently revised fauna of Hungary, initiated by Papp and Černý (2016), although unlike here, that work includes unconfirmed species predicted to be in the country. Hungary is slightly more northern in position compared to the Delmarva states and slightly smaller in size (ca. 93,030 km2, versus 99,420 km2). Since this work excludes species not yet known to occur in the region, reference to the broader Nearctic treatments of Spencer (1969) and Spencer and Steyskal (1986b) is suggested in case an identification is uncertain; reference to host plant catalogues (such as Spencer 1990 and Benavent-Corai et al. 2005) and non-Nearctic taxonomic treatments is also recommended.

In total, this study presents 346 new state and provincial records and 23 new country records, including species previously known only from Europe, Canada, and the western or southern United States.

Fifteen species are newly described: Agromyza echinalis sp. nov., Melanagromyza brunkei sp. nov., M. eoflacensis sp. nov., M. glyptos sp. nov., M. rutella sp. nov., Ophiomyia capitolia sp. nov., O. cuprea sp. nov., O. galiodes sp. nov., O. heleios sp. nov., O. kalia sp. nov., O. laticolis sp. nov., Cerodontha (Poemyza) ungula sp. nov., Phytomyza avicursa sp. nov., P. catenula sp. nov., P. winkleri sp. nov.

Four new species-level synonyms and one genus-level synonym are provided: Agromyza marmorensis Spencer syn. nov. is included as a synonym of A. aristata Malloch; Melanagromyza fastosa Spencer syn. nov. is included as a junior synonym of Ophiomyia tiliae (Couden); Melanagromyza verbesinae Spencer is considered a synonym of M. vernoniana Steyskal; Phytomyza ranunculoides Spencer syn. nov. is included as a junior synonym of Phytomyza loewii Hendel; the genus Liomycina Enderlein syn. nov. is included as a junior synonym of Phytobia Lioy. Ophiomyia ultima (Spencer) comb. nov. is recombined from Melanagromyza. Euhexomyza albicula Spencer stat. reinst., comb. nov. is resurrected from synonymy with E. winnemanae (Malloch). New host records are provided.

The total number of known species in the Delmarva states is now 156, representing nearly 18% of the 879 species known from the Nearctic north of Mexico (Eiseman, pers. comm.). The Agromyzinae is better represented, with ~ 25% of known species, with Agromyza being especially well-represented at ~ 43% of known species. The Phytomyzinae fauna represents ~ 15% of known species, with Aulagromyza Enderlein (27.3%), Calycomyza (30.7%), Nemorimyza (100%) and Phytobia (41.0%) being best represented, and Liriomyza (8.9%) and Phytomyza (11.3%) being much more weakly represented.

Two genus-level groups occurring the Nearctic are not known from the Delmarva states as of yet. The first is the Cerodontha Rondani subgenus Phytagromyza Hendel, which mines in Poaceae. It is known from two western Nearctic species, C. (Ph.) frankensis Spencer (Fig. 100) and C. (Ph.) pusilla Zlobin, and one Palaearctic and eastern Canadian species, C. (Ph.) flavocingulata (Strobl). These species are keyed in Zlobin (1997). Secondly, species of Pseudonapomyza Hendel occur further to the north and west in North America, but discovery of this genus in the Delmarva states would not be surprising. Pseudonapomyza presently contains at least 100 species worldwide, almost entirely in the Old World, but there are likely dozens more (Černý 1992, 2008; Zlobin 2002a, 2005a; Černý and Zlobin 2008; Boucher 2004). Boucher (2004) last revised the New World fauna, recognising four species. All of these occur in the Nearctic, with P. asiatica Spencer extending into Guadeloupe and Venezuela.

Although the inspiration for this revision was Mathis and Foster’s (2007) revision of the Canacidae (Diptera) of the Delmarva states, and discussions with Wayne Mathis, the motivation came from curating the immense acalyptrate collection at the Museum of Natural History at the Smithsonian (USNM), which contains vast amounts of historical material from these states and district. Strangely, this collection was largely ignored by Spencer and Steyskal (1986b) during their revision of the Agromyzidae of the lower United States, using it mostly to examine type specimens designated by earlier researchers. Spanning more than a century of collection, the USNM is likely the largest repository of Nearctic Agromyzidae, and certainly the most extensive collection from the mid-Atlantic states, populated quite heavily through the efforts of C.H. Curran, K.E. Frick, S.W. Frost, G. Hevel, A.L. Melander, W. Steiner, G. Steyskal, and W.W. Wirth, among others. As the author wished to take advantage of this heavily sampled fauna, and to examine all agromyzid genera in a work of reasonable geographic scope for his first study of the family, a revision of the Delmarva fauna seemed appropriate. The author’s later work on Agromyzidae at the Canadian National Collection of Insects, Arachnids and Nematodes (CNC) provided much additional material complementary to this study, and specimens from that institution also predominate here.

Table 1.

Total number of species for each genus of Agromyzidae in the Nearctic north of Mexico and the Delmarva states.

Subfamily Genus # Nearctic species north of Mexico # species in Delmarva states
Agromyzinae Agromyza 58 25
Euhexomyza 6 1
Japanagromyza 8 1
Melanagromyza 80 16
Ophiomyia 89 18
Phytomyzinae Amauromyza 23 3
Aulagromyza 11 3
Calycomyza 39 12
Cerodontha 73 17
Haplopeodes 5 1
Liriomyza 157 14
Metopomyza 6 1
Nemorimyza 2 2
Phytobia 16 5
Phytoliriomyza 27 6
Phytomyza 275 31
Pseudonapomyza 4 0
TOTAL 879 156

Natural history

The larvae of all Agromyzidae feed inside the living tissue of plants, with species reared from more than 140 families across the monocots, dicots, horsetails, ferns, and liverworts (Hering 1957, 1966; Spencer 1990; Scheffer et al. 2007). At least one species may also occur on a hornwort, with a parasitized puparium tentatively identified as Phytomyza recovered from Nothocerus aenigmaticus (Gilme, 2017). The non-vascular thallophytes and mosses are not known to be hosts, nor are the vascular gymnosperms with the exception of unsubstantiated Phytobia larval traces in fossil and extant tree trunks (Süss 1979), one record of Tropicomyia atomella (Malloch) in leaves of Gnetum (Sasakawa and Pong 1988), and presumed leaf mines of Liriomyza schmidti (Aldrich) on the cycad Zamia (Eiseman et al. 2019).

Feeding habits among taxa are diverse, with many species leaf-mining or boring inside stems, roots, twigs, and trunks, but others are known to mine in seeds, seed pods and flower heads, and species across a number of genera induce galls. Herbaceous annual plants are disproportionately affected by Agromyzidae compared to other similarly feeding phytophages such as the leaf-mining Lepidoptera, which dominate on perennials (Hespenheide 1991). Most species for which larval habits have been discovered are those that mine leaves because the mines produced are often readily observed, and the mines of some may be visible from a distance. Especially conspicuous mines can be produced by species such as Calycomyza flavinotum (Frick), for example, where numerous larvae might mine across the entire surface of large Arctium leaves. Conversely, those species that remain hidden inside thicker, opaque structures are generally overlooked, and as such, many host relationships for these remain to be discovered. An exhaustive and invaluable compendium of these flies and their hosts was provided by Spencer (1990), with an updated list of host genera later provided by Benavent-Corai et al. (2005). Eiseman (2019) provides complete host species lists for all known leaf- and stem-mining species in North American, as well as records of observed mines of unknown agromyzid species.

Since plant hosts are still unknown for many species, and it is becoming apparent that many species once thought to be faithful to a single species or genus of host are actually more varied in their diet, it is not recommended that identifications be made using host alone. Adult male dissections or molecular sequence data are crucial for confident identification, and additional details of larval host usage, mine shape and frass pattern are also useful for contributing to a diagnosis (Eiseman and Lonsdale 2018). While some agromyzids may truly be restricted to a single plant host, it is likely that most can be generally characterised as oligophagous, being limited to multiple hosts in related families or genera. The apparent monophagy of many species will eventually prove to be an artifact of incomplete observations, with a growing body of data revealing that additional rearing records will almost certainly lead to novel host records for many species (Scheffer et al. 2021; Eiseman et al. 2018–2021). The extent to which this is true, however, can only be accurately characterised after more targeted taxonomic and rearing studies, and should prove to be a rich field of study.

Among the agromyzid genera, many lineages have successfully speciated on specific plant taxa, especially on diverse groups such as grasses (e.g., Cerodontha, Metopomyza Enderlein, the Pseudonapomyza atra group, the Agromyza bispinata group, others) and herbaceous Asteraceae (e.g., at least 1/2 of non-Cerodontha phytomyzines). Within these more specialised groups, however, there appear to be periodic but major shifts to entirely different plant families or orders followed by bursts of speciation (Griffiths 1980; Scheffer and Wiegmann 2000). This was supported by Scheffer et al. (2007) at the genus level, who found host use to be quite varied, with at least ten genera occurring on both monocots and dicots, and non-angiosperm feeders derived from angiosperm-feeding stock.

True polyphagy is uncommon in the Agromyzidae, with only an estimated 16 species occurring on a wide diversity of host families (Spencer 1990). As would be expected, many of these occur on a variety of agriculturally significant crops and can sometimes be quite pestiferous and difficult to control, as populations easily accumulate on weeds in and around growing areas or alternate crops. They are also often readily dispersed to other countries through trade, hidden within plant tissue and potting soil. A number of these, especially those in the genus Liriomyza, appear to have originated in North and Central America, but other Old World species are similarly problematic.

Larval movement of agromyzid leaf miners within the host is characteristic and can be used to differentiate them from other leaf miners such as the microlepidoptera, as discussed in Eberle (1967) and Dempewolf (2005): the larva lies on its side and sweeps its body dorsally and ventrally to shave off successive rows of cells.

Mine location and shape, and the pattern by which frass is deposited in the mine, is often characteristic for a given species on a particular host during a certain part of the year, which may allow for tentative field identification (Nowakowski 1962; Dempewolf 2005; Lonsdale and Scheffer 2011). This is not true for all species, however, with similar mines of different species sometimes occurring on the same host plant or population of plants, which has been particularly problematic in the early diagnosis of some pest species. An especially thorough discussion on this topic is provided in Eiseman (2019). The main categories of leaf mine pattern are linear/serpentine and blotch, although there are many variations and irregularities, and a single larva may produce different mines over time, sometimes with later blotch mines obscuring the linear original. Leaf mines are usually also associated with specific parts of the leaf, such as the midrib, margin, lateral veins or even the petiole. The usage of specific cell layers also varies between species, but a single larva may not necessarily be restricted to a single layer over successive instars. Upon pupariation, the larva may remain in the mine or slightly removed from it within the leaf, it may first emerge and drop to the ground, or it may pupariate partially to entirely emerged from the mine, anchoring itself with enlarged hind spiracles, strong threads, or a hardened frass deposit. Seed and stem miners typically pupariate within the host where feeding occurs (Spencer 1987). A few species switch from leaf to stem mining, such as the pestiferous Ophiomyia phaseoli (Tryon), or from stem to seed mining. If a species is gall-forming, the shape and location of the gall is characteristic (Dempewolf 2005). Phytobia species are found under the bark of trees where they feed on young xylem, tissue that in the past was incorrectly referred to as “cambium”, as discussed in Ylioja et al. (1998); their feeding produces characteristic “pith flecks” that are also called “medullary spots”.

Eggs are introduced into host tissue by female oviposition, with the modified terminal segments of the female abdomen specialised for creating a hole in the host and guiding the egg into the appropriate position among the cell layers. The female may produce similar additional punctures in the host to release sap on which both sexes feed (Ferrar 1987; Aukema et al. 1996). These feeding punctures may be abundant and cover the entire leaf surface, with some species such as Phytomyza horticola (Goureau) producing up to 150 punctures per leaf, despite leaving only a few eggs behind (Hill 1986). In Agromyza frontella (Rondani), pheromones left by the female around the egg insertion point, possibly a product of the digestive tract, reduces competition as an oviposition-deterrent (McNeil and Quiring 1983). The larva emerges from the egg after a few days, and feeding may last from a single week to ten months, dependant on the species, host and temperature (Spencer 1987; USDA 2008). Some species such as Phytomyza ilicis Curtis feed well into the winter (Eber 2004). Species with multiple generations per year tend to produce more rapidly developing larvae, and overwintering larvae may arrest growth to resume activity when favourable conditions return (Spencer 1987). Several forms of conspecific competition have been observed for A. frontella on the host, including larval cannibalism, larvae pre-empting competitors by mining the leaf first, and adult feeding on a leaf to preclude larval mining (Quiring and McNeil 1984ac).

Parasitoids are highly effective in controlling agromyzid population numbers, often with high mortality rates that are frequently discouraging to those attempting to rear the flies. Commonly encountered families are Eulophidae, Figitidae, Braconidae, and Pteromalidae, which emerge from the host puparium, but less commonly the larva. Eiseman (2019) has provided an overview of these parasitoids and their relationships for North American taxa. The preservation of these wasps in agricultural settings is essential, as they often suppress pests to levels not requiring additional intervention. In settings where pesticides are used, misapplication may easily affect wasp populations disproportionately, resulting in the collapse of these natural controls followed by pest outbreaks, unintentionally increasing damage, and reducing crop yield and value (Weintraub and Horowitz 1995; Hidrayani et al. 2005).

Classification of Agromyzidae

Family-level relationships

While it is relatively simple to diagnose the family, the assignment of derived or synapomorphic states is problematic due to the uncertainty involved with establishing outgroups, and there is little consensus on sister group relationships. Minor similarities in wing venation, chaetotaxy and genitalia have historically allied the Agromyzidae to other acalyptrates, but these simple characters are of equivocal polarity and can be easily interpreted as plesiomorphic.

The Agromyzidae is currently treated as a member of the Opomyzoidea (McAlpine 1989), at least as a close relative of the Clusiidae (Griffiths 1972b; Frick 1952a; Spencer 1987, 1990), but it is becoming increasingly evident that this superfamily is a “dumping ground” for unplaced families of more generalised morphology, at least in part. As an example, the monogeneric Acartophthalmidae was treated as sister to Clusiidae in McAlpine’s (1989) Clusioinea on the basis of a generalised habitus and a handful of putative synapomorphies (an angulate extension on the pedicel is the only character of note and this has been independently derived in other Acalyptratae). Acartophthalmidae has recently been supported as Carnoidea using quantitative phylogenetic methods, subsequently removing it from the Opomyzoidea (Buck 2006). Most recently, Winkler et al.’s (2010) molecular study of the Opomyzoidea provided little evidence for any family-level relationships, with the exception of weak support for Agromyzidae + Neurochaetidae, a subset of periscelid genera within an expanded Aulacigastridae, and relatively strong support for Xenasteiidae + Australimyzidae (Carnoidea). There are additional other relationships among the Asteoinea supported to varying degrees by different character subsets (a summary of these is provided in Winkler et al. 2010), but the homoplasious nature of these characters and the degree to which they have been reinterpreted in the literature precludes any confident statements about common ancestry.

In an effort to identify any possible synapomorphic morphological characters shared by the opomyzoid families, the author examined representatives of all extant genera, excluding some asteiid, odiniid, teratomyzid and Afrotropical taxa; additional data on Palaearctic Anthomyzidae were taken from Roháček (2006). Except for the putatively ancestral characters of chaetotaxy and wing venation that characterise most Opomyzoidea, almost no additional external or genitalic characters were found that would serve to support relationships between these families, with two major exceptions.

The first of these is Anthomyzidae + Opomyzidae, the monophyly of which received modest support in Roháček (2006). This contrasts uncertain results regarding these two families in Winkler et al. (2010), however, and there are possible conflicting characters of the female genitalia (Kotrba and Baptista 2002). The second is an association between the Agromyzidae and the predominantly Australian Fergusoninidae, which is supported by a relatively substantial number of female genitalic and larval characters. Fergusoninidae is a similarly phytophagous family of small-bodied species that was at one time even considered to be Agromyzidae (Malloch 1924b). However, placement of this family in the superfamily Nerioidea was well-supported by Lonsdale (2020) in a quantitative analysis of the nerioid and diopsoid families using morphological data sets (external, male genitalic and female genitalic) and numerous outgroup families, including Agromyzidae. Putative similarities to Agromyzidae were supported as superficial and there is much disagreement in the structure of the male genitalia. The molecular analyses of Bayless et al. (2021) provide conflicting placement of Fergusoninidae, again suggesting closer association to Agromyzidae in a “cleft pedicel clade”. In their preferred phylogeny, Agromyzidae + Odiniidae is sister to Fergusoninidae + Carnidae, which still supports the similar female and larval characters of Agromyzidae and Fergusoninidae as convergent.

Putative family synapomorphies

In his discussion of the family, McAlpine (1989) listed a number of derived characters to define the Agromyzidae: the larvae feed in living plant tissue with an associated adaptation of the mouthparts, with “mandibles toothed and angularly positioned in relation to hypopharyngeal sclerite and hypopharyngeal and tentoropharyngeal sclerites fused” (Figs 29–31); the anterior larval spiracles lie close together dorsally on the thorax (visible in puparium, Fig. 27); the cells connected to the larval fat bodies contain calcium carbonate crystals; tergite 6 and sternite 5 are enlarged in the male; the male pregenital sclerites (sternites 6–8) are reduced, usually forming a small symmetrical dorsal band; the phallus is “extremely complex” and [at least ancestrally] contains two gonopores; female segment 7 forms a large oviscape with an anterodorsal apodeme (Figs 1113); the membrane between female segments 7 and 8 is densely covered with anteriorly directed denticles (Fig. 19); female segment 8 is modified into one pair of serrated egg-guides; the female cercus has a group of apical trichoid sensilla (Figs 16–18).

Additional external adult characters added here include a relatively large and subshiny ocellar triangle surrounding the ocellar tubercle (Fig. 78) (secondarily reduced or obscured in several lineages, especially Phytomyzinae), and possibly the presence of only one pair of lateral scutellar setae (absent in some Cerodontha). Elaborating on the structure of the internal components of the genitalia (Figs 139–142): the pregonite is absent; the distiphallus is discrete and capsule-like with basal insertion of the ejaculatory duct (often with basal differentiation of the segment to produce a “mesophallus”); the basiphallus is segmented basally to produce a separate cylindrical structure termed the “phallophorus”; the basiphallus is distally composed of one pair of long lateral plates flanking the ejaculatory duct; the epiphallus is a segmented, folding structure beneath the phallophorus.

Generic classification

The genera of Agromyzidae have traditionally been divided between two subfamilies, the Agromyzinae and the Phytomyzinae, which follows the same system originally devised by Fallén (1823a, b) for his “agromyzides” and “phytomyzides”. This system has recently been corroborated using morphological (Dempewolf 2001) and molecular (Scheffer et al. 2007) data. The adults of Phytomyzinae are characterised by plesiomorphic wing venation similar to that of other Opomyzoidea: the subcostal vein is either incomplete distally or it reaches the costa as a thin fold that remains separate from R1; R1 itself is sometimes slightly expanded apically, but it is almost always narrow and straight (Fig. 3). Conversely, the subcosta is fused with R1 distally in the Agromyzinae and the apex of R1 is almost always expanded and sinuate (Fig. 4). This venation is relatively consistent within subfamilies, although a number of Phytobia (Phytomyzinae) have converged upon the state otherwise typical of the Agromyzinae, and some Agromyza Fallén have the derived character appearing very subtly. Regarding larval morphology, the Phytomyzinae (Figs 24, 25, 30, 31) is characterised by a reduced lower arm on the cephalopharyngeal skeleton while the Agromyzinae (Fig. 29) is characterised by two strong arms (Spencer 1987). A variably developed medial longitudinal sclerotised band on the ejaculatory apodeme is here also found throughout the Agromyzinae (Fig. 141), although it only frequently appears in species of the Ophiomyia genus group, being indistinct or secondarily lost in many species of Agromyza and Japanagromyza Sasakawa.

Since inconsistencies in the subfamilies’ defining characters were brought to light by von Tschirnhaus (1971) it has been speculated that the lineages of Agromyzidae cannot be forced into two distinct halves. As a solution, Spencer (1990) provided an alternate system of classification that encompasses four separate genus groups: the Penetagromyza group, which is equivalent to Agromyzinae, and the Napomyza, Phytobia and Phytoliriomyza groups, which together are equivalent to the Phytomyzinae. Little to no evidence was provided in support of these groups or the relationships within them, and their use in classification here appears to be limited, but they are nonetheless mentioned below as they relate to generic relationships.

Subfamily Agromyzinae

Although there are a number of highly characteristic lineages within Agromyzinae, most of the subfamily has relatively non-descript and uniform colouration, venation and often chaetotaxy. To compound this, species are often described on the basis of slight morphometric changes in the male genitalia and subtle variations in external morphology, making species identification problematic.

Agromyzinae can be divided into three relatively distinct clades. The first is the Ophiomyia genus group, an aggregate of morphologically similar and frequently confused genera that includes Ophiomyia Braschnikov (Figs 6483), Melanagromyza (Figs 52–63), Euhexomyza Lonsdale (Figs 4448) and the Old World Tropicomyia Spencer. Lonsdale (2014) re-characterised this group and its genera, and aside from sharing a generally similar habitus, he found species to have a large, bulging lunule, insertion of the antenna below the midpoint of the head, brown halters (with very few exceptions) and no prescutellar acrostichal setae (also absent in some Japanagromyza). Internally, the subepandrial sclerite is divided into two well-sclerotised plates that almost meet at a point medially, the inner lobe of the hypandrium has partially separated into an arched, band-like sclerite, the postgonite is sometimes reduced to absent, modified or fused to other structures, and the mesophallus is shifted to form a small distoventral lobe on the distiphallus (with some exceptions). Species are mostly internal borers, with leaf mining largely restricted to Tropicomyia. Tropicomyia is a small genus of minute, brown leaf mining species with a disproportionate number of polyphages. While the genus is likely not monophyletic and in need of revision (Lonsdale 2014), most species appear to belong to a core lineage that produces irregular mines in the upper epidermal layer that appear “silvery” (Spencer 1973).

The second clade is made up of the genus Agromyza (Figs 37–43), which is often paler and more slender compared to other Agromyzinae, and it is best diagnosed by the presence of a distinct saw-like stridulatory mechanism along the fused lateral margins of the first and second tergites in both sexes (Fig. 38). All species also have one pair of prescutellar acrostichal setae and three or more dorsocentral setae (Fig. 10), although a number of grass-feeding species have only two well-developed pairs.

The last clade is composed solely of Japanagromyza (Figs 49–51), which was redefined by Lonsdale (2013) in his revision of the African species. The genus shares a combination of features characteristic of both Agromyza and the Ophiomyia genus group, leading some authors to treat it as an “intermediate” of the two. Like Agromyza, it often has prescutellar acrostichal setae and the mesophallus is basal to the distiphallus (not ventral). Japanagromyza and a minority of Agromyza also have an inner-distal comb of setae on the hind tibia (von Tschirnhaus 1991). Like the Ophiomyia genus group, Japanagromyza lacks the stridulatory file on the abdomen, it is quite dark in colour and sometimes iridescent, and it has a reduced number of dorsocentrals. Furthermore, while most species leaf-mine like Agromyza, Sasakawa (2010) revealed that the genus presented a broader range of habits otherwise characteristic of the Ophiomyia genus group. Based on the relatively high degree of variation in these characters, and because of the unusual diversity in genitalic features seen across Japanagromyza species globally, it seems unlikely that the genus is monophyletic (Lonsdale 2013).

Subfamily Phytomyzinae

The Phytomyzinae represents a comparatively heterogenous group that varies widely in external and male genitalic morphology, contrasting the often uniform species of Agromyzinae. A genus-level phylogeny of the subfamily was produced by Dempewolf (2001), but portions of his proposed relationships were challenged by Scheffer et al. (2007) and still remain partially unresolved. Despite this, several groups of genera appear to be moderately well-supported.

The first of these is what Spencer (1990) called the “Napomyza group”, and it is largely comprised of the diverse genus Phytomyza (Figs 131–138), which is characterised by proclinate orbital setulae in combination with a costa that extends to vein R4+5, M1 is weak or spectral, and crossvein dm-m is absent or situated basally. There is also slight to strong costalisation of the wing veins. The classification of this genus was recently advanced by Winkler et al. (2009), who provided the most robust quantitative analysis of Phytomyza to date. Winkler et al. (2009) maintained Napomyza and Ptochomyza Hering as subgenera of Phytomyza because they are readily diagnosable, they were both supported as monophyletic, and were both recovered near the base of Phytomyza sensu lato. Winkler et al. (2009) also re-synonymised Chromatomyia Hardy with Phytomyza, supporting the findings of a number of historical studies. An argument for retention of Chromatomyia was proposed by von Tschirnhaus (2021), but reanalysis of the evidence by Lonsdale and Eiseman (2021) revealed that synonymy of the genus was well-founded and should be maintained for a number of reasons: Chromatomyia renders Phytomyza paraphyletic, it is unlikely to be monophyletic itself (although proof is not definitive), larval characters in support of the genus are homoplastic, and male genitalic characters supporting the genus are impractical for use, frequently misdiagnosed or misinterpreted, and also possibly homoplastic.

Other genera in the Napomyza group are poorly represented or unknown in the New World. These resemble Phytomyza in having the costa extend to R4+5, and many species also exhibit costalisation of the wing veins, but the orbital setulae are erect to reclinate or absent. The smallest of these genera is Gymnophytomyza Hendel, which has two Palaearctic species that were treated by Zlobin (1999); species lack the proepisternal seta and there are records of leaf mining in Galium. Aulagromyza and Pseudonapomyza are differentiated by characters of wing venation and their monophyly is yet to be confidently established. Most Aulagromyza are dark-bodied species (Fig. 88), but the A. populicola group (Fig. 89) is made up of bright yellow species that feed on Salicaceae and may not be related, as discussed by Zlobin (2007c) and Winkler et al. (2009). Pseudonapomyza is divided into two groups, the most abundant and widespread of which is the P. atra group (Zlobin 2002a) (Figs 130, 833–839), which has an angulate first flagellomere (with at least one exception), the male mesophallus and distiphallus are distinctive, and larvae feed on Poaceae (Zlobin 2002a). The P. acanthacearum group may be paraphyletic (Zlobin 2002a), and it is restricted to Old World tropical regions, where species feed on Acanthaceae, Amaranthaceae, Asteraceae (Černý 2008) and Bignoniaceae (Madire et al. 2010).

The remaining Phytomyzinae were considered by Spencer (1990) to belong to either the “Phytoliriomyza group” or the “Phytobia group”. There is no strong morphological or molecular support for the Phytoliriomyza group (Scheffer et al. 2007), but a subset of six genera with tubercle-like setae on the posterodistal margin of the epandrium and surstylus is possibly monophyletic: Calycomyza (Figs 9098), Haplopeodes Steyskal (Fig. 120), Liriomyza (Figs 107117), Metopomyza (Figs 118, 119), Phytoliriomyza Hendel (Figs 126–129) and Selachops. These genera were recovered together in Scheffer et al. (2007), but not in Dempewolf (2001), although results in the latter were discussed as possibly being attributable to the inclusion of homoplastic features. Another likely monophyletic lineage within this group is Liriomyza, Haplopeodes and some Phytoliriomyza, where the number of tubercles on epandrium and surstylus is reduced to one, although these structures are sometimes secondarily lost, modified or duplicated. In Metopomyza and the remaining Phytoliriomyza there is usually a discrete comb or otherwise ordered arrangement of tubercles, contrasting the dense, scattered tubercles seen in Calycomyza, Selachops and the unusual L. quadrisetosa (Malloch) (Figs 616–620). Only the small Palaearctic Selachops is not known in the Nearctic (Zlobin 1984); the host of S. flavocinctus Wahlberg was recently revealed to be Carex acuta (Cyperaceae) (Kostromina et al. 2016).

Within this group characterised by tubercle-like setae on the external male terminalia, there is gathering consensus that at least some genera are artificial groupings (Dempewolf 2001; Scheffer et al. 2007). As such, there is a strong need for boundaries to be re-evaluated so that genera can be reconstructed as entities that are both monophyletic and easily diagnosable using external characters. For example, if it is supported that Phytoliriomyza is indeed paraphyletic, it will have to be divided into at least two groups if other genera such as Metopomyza and Liriomyza are to be retained. However, while some lineages of Phytoliriomyza can be diagnosed externally (such as that with an entirely black first flagellomere and that with an apically brown halter and setose eyes), others are inseparable from species in related genera without male dissection, reducing potential generic utility. Dissection is currently also necessary for confident diagnosis of Haplopeodes and most Liriomyza, given that characters of colour, chaetotaxy and wing venation are overlapping and frequently convergent. Some progress has been made to provide natural boundaries for Liriomyza, such as including Galiomyza Spencer as a junior synonym, but consistently useful external diagnostic characters are still lacking and much additional work needs to be done (Lonsdale 2017). Lonsdale (2017) discussed the polyphyly and variant morphology of Galiomyza species in detail before synonymizing it, but Papp and Černý (2017) resurrected it soon thereafter, unusually noting that they preferred to retain it in order to avoid reorganising their as yet unpublished chapter on it, which was to appear in print soon after the synonymy appeared; an unclarified statement on the monophyly of Galiomyza was also mentioned. It is recommended that the synonymy of Galiomyza be retained until scientific evidence is provided to the contrary. The position of the New World genus Haplopeodes is uncertain, although some have proposed it to be an offshoot of Liriomyza (Spencer 1990) or at least a close relative of it (Steyskal 1980). Determining Haplopeodes’ ancestry has proven to be difficult, however, due to an extreme reduction in size and other features that may have been phylogenetically informative, especially those of the male genitalia.

The positions of the other phytomyzine genera have not been confidently resolved and their relationships remain uncertain (Dempewolf 2001; Scheffer et al. 2007). Molecular data weakly supports Cerodontha (Figs 99–106) as sister to the rest of the Phytomyzinae, and there is variable support for a lineage encompassing those genera without tubercles on the surstylus and epandrium, including those in the “Napomyza group”. Of these genera, Amauromyza Hendel (Figs 84–87), Nemorimyza (Figs 121–123) and Phytobia (Figs 124, 125) belong to what Spencer (1990) called the “Phytobia group”. Spencer (1990) considered these genera to be basal in the family because they share a large body size and a supposedly plesiomorphic feeding mode (Nowakowski 1964; Spencer 1990), that is, stem-boring, even though all Nemorimyza and some Amauromyza are leaf-miners. Both of these historically held assumptions of evolutionary progression were found to be erroneous by Scheffer et al. (2007), who instead supported leaf mining as the ancestral state.

The status of two genera placed by Spencer (1990) in the Phytoliriomyza group is uncertain. Genitalic illustrations of Xeniomyza Meijere in Spencer (1990: figs 237, 238) strongly suggest that the genus should be synonymized with Liriomyza. The type species is known from Amaranthaceae (Spencer 1990), and the two Palaearctic species are characterised by loss of dm-m and costalisation of the wing veins (Zlobin 1979). The monotypic Pseudoliriomyza Spencer (host unknown) is known from Tanzania and Indonesia (Spencer 1966d) and has some similarity to Liriomyza, but additional specimens are required for further study. The genus is characterised by loss of vein dm-m, the fourth dorsocentral, the surstylus and the epandrial spine, and there are some genitalic similarities to Amauromyza in the structure of the hypandrium and the ejaculatory apodeme.

Materials and methods

The majority of material examined for this study is deposited in the Smithsonian Institution, National Museum of Natural History, Washington, D.C., USA (USNM), and the Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Canada (CNC). Additional material was examined from, or is deposited in the following institutions:

BLTJ Biological Laboratory, Yazu High School, Tottori Pref., Japan;

BPBM Bernice P. Bishop Museum, Honolulu, Hawaii, USA;

CASC California Academy of Sciences, San Francisco, California, USA;

CFS Canadian Forest Service, Edmonton, Alberta, Canada;

CSCA California State Collection of Arthropods California, USA;

CUIC Cornell University, Ithaca, New York, USA;

DEBU University of Guelph Insect Collection, Guelph, Ontario, Canada;

INHS Illinois Natural History Survey, Champaign, Illinois, USA;

MCZ Museum of Comparative Zoology, Cambridge, Massachusetts, USA;

MNHN Museum National D’Histoire Naturelle, Paris, France;

NHMUK The Natural History Museum, London, United Kingdom;

NMW Naturhistorisches Museum Wien, Vienna, Austria;

NRS Naturhistoriska riksmuseet, Stockholm, Sweden;

OUMNH Oxford University Museum of Natural History, Oxford, United Kingdom;

RNH Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands;

SMNS Staatliches Museum für Naturkunde, Stuttgart, Germany;

UDCC University of Delaware, Newark, Delaware, USA;

UZMH Finnish Museum of Natural History, University of Helsinki, Finland;

VPIC Virginia Tech, Blacksburg, Virginia, USA;

WVDA West Virginia Department of Agriculture, Charleston, West Virginia, USA;

ZIL Russian Academy of Sciences, Zoological Institute, St. Petersburg, Russia;

ZMHU Museum für Naturkunde der Humboldt-Universität, Berlin, Germany;

ZMUC Zoological Museum, University of Copenhagen, Denmark;

ZMUN Zoological Museum, University of Oslo, Norway.

For the following keys and species descriptions, an emphasis was placed on adult description and diagnosis, especially with regards to the highly derived components of the male terminalia, as the majority of material examined was collected as adults with little or no associated host or habitat data. Host genera are provided only, with plant species listed when only a few are known. While external morphology is of course also useful, findings strongly support Nowakowski (1962), who noted that the external characters usually used to separate closely related agromyzid species, such as wing vein ratios, body size, fronto-orbital number, and subtle colour characters, are often of “negligible taxonomic value”. Many of these external characters were here found to vary subtly to widely within species previously thought to be morphologically conserved. This is of particular significance considering the frequent use of these characters in historical identification keys, which often only provide a single character at each couplet.

Adult terminology and definitions are outlined below in the family description. Terminology for external structures largely following Cumming and Wood (2017), except where noted. Genitalic terminology mostly follows Lonsdale (2020), which expands on Cumming and Wood (2017) by discussing structures of the terminalia peculiar to families of Acalyptratae that are not otherwise considered in the earlier work. A few deviations are made here for structures of the hypandrial complex specific to the Agromyzidae, wherein the terminology presented by Steyskal (1969) is provided: specifically, the phallophorus, proepiphallus, and metepiphallus. Gena height is measured vertically at the lowest point of the eye. Dorsocentral setae are numbered starting from the posterior-most seta because setae number often varies anteriorly. Rows of acrostichal setulae are counted at the transverse suture. New host records, and new country/province/state records are indicated by an asterisk.

Most specimens are pinned, and either air-dried or prepared in a critical-point drier. Some voucher material collected by D. Smith or S. Scheffer, excluding most specimens designated as types, was retained in 95% ethanol for potential future study. Abdomens of dissected specimens were initially prepared by dissolving the soft tissue in hot potassium hydroxide, followed by washing in demineralised or deionised water, neutralisation in glacial acetic acid, and further washing, but later dissections used hot lactic acid exclusively. Thorough washing and neutralisation of the potassium hydroxide still leave minute traces of the chemical, which continue to dissolve pigment and weaken sclerotised tissue, and it is not recommended for use. Genitalia were stored in glycerine in microvials pinned with the specimen. Most illustrations were prepared by the author, excluding several figures scanned from the literature, which are indicated in the text and in the acknowledgements. The sizes of illustrated structures are shown in correct proportion to each other, with the epandrium reduced due to space constraints; some structures were not illustrated if they were damaged, could not be found (ejaculatory apodeme), or were uninformative for diagnosis within the genus.

Results

Agromyzidae – adult diagnosis

Adults generally small-bodied, wing length usually 1.5–3.5 mm, but as small as 0.5 mm and as large as 6.5 mm. Postocellar setae diverging. Frons usually with inclinate anterior and reclinate posterior fronto-orbitals. Orbital setulae usually present. Vibrissa present. Arista dorsobasal (Figs 1, 2). Ocellar tubercle surrounded by often large and subshiny ocellar triangle (usually less distinct than in Chloropidae) (Fig. 78), but sometimes reduced or with triangle absent. Lunule distinct, sometimes large. Scutellum with one pair of apical setae and almost always with one pair of well-developed laterobasal setae. Costa with break at end of subcostal vein, which is complete or nearly complete (Figs 3, 4), but sometimes fused to R1 if that vein is apically sinuous; humeral break absent. Vein CuA2 present, enclosing anal cell. Dorsoapical tibial setae absent. Basiphallus segmented into basal phallophorus and usually one pair of distal plates flanking duct. Distiphallus sometimes segmented to produce basal mesophallus. Female with oviscape distinct, dark, and heavily sclerotised (Figs 7, 1113); membrane between female segments 7 and 8 densely covered with denticles; sternite 8 modified into one pair of serrated egg-guides.

A number of other small-bodied acalyptrate families are similar to Agromyzidae, but differ as follows. Chloropidae can be similarly small-bodied and stout with a large ocellar triangle, but this family lacks the anal cell and oviscape, and has a sharp propleural carina. Anthomyzidae are slender and sometimes pale with a vibrissa and larger ocellar triangle, superficially similar to some genera such as Cerodontha, and vein R1 is sinuous apically, similar to Agromyzinae, but in this family, a ctenidial spine is almost always present on the fore femur, the anepisternum is usually bare (with at least one large seta in Agromyzidae), the subcostal vein is clearly fused to R1 apically, and there is no stout oviscape. Odiniidae and Clusiidae have similar chaetotaxy, but preapical tibial setae are usually present, the large oviscape is absent, and Odiniidae have an incomplete subcosta; some Clusiidae do not have these tibial setae, but all species have a projection on the outer- and sometimes inner-distal margin of the pedicel. Other taxa with a large, conspicuous oviscape and complete subcosta, such as Tephritoidea, usually have strongly patterned wings and lack a vibrissa; Fergusoninidae are bright yellow with an anteriorly flattened head with small antennae concealed in deep recesses, a small face, and a very large parafacial and lunule.

Agromyzidae – adult description

General: Wing length small, usually 1.5–3.5 mm (sometimes as small or as large as 0.5 mm and 6.5 mm). Colour varying from almost entirely black or brown to yellow, but many with some degree of pattern that may be more of less typical of a genus (for example, species of Liriomyza and Calycomyza are more conservatively patterned than Phytomyza and Phytoliriomyza); dark species of Agromyzinae sometimes with metallic shine (many Melanagromyza and some Ophiomyia and Japanagromyza); often subshiny to heavily brown or grey pruinose, uncommonly glossy.

Head: (Figs 1, 2) Frons with frontal vitta usually soft, but sometimes better sclerotised, sometimes with several setulae bordering lunule (common in Agromyza and Phytobia); fronto-orbital plate (fused frontal and orbital plates) and variably sized ocellar triangle either indistinct from vitta, more sclerotised, or delimited by difference in texture; ocellar triangle variably sized, sometimes reaching lunule, sometimes reduced and restricted to small, raised ocellar tubercle bearing ocelli. Lunule usually distinct; small soft and sunken, to large heavily sclerotised and bulging. Antenna porrect, first flagellomere rarely angled ventrally; arista subbasal, usually pubescent to bare, rarely with rays more elongate. Eye upright to distinctly oblique; bare to microtrichose. If parafacial continues under eye onto gena, band-like region referred to as “cheek”, and remaining ventral region is referred to as “jowl” (“subgena” in Cumming and Wood 2017). Face shallowly concave, often with broad lateral depressions to receive antennae, sometimes with medial carina that may have a central swelling or “bulb”. Gena usually straight or angled posteriorly, uncommonly deeper or produced anteriorly (many Ophiomyia with gena strongly projecting). Clypeus broad and U-shaped, posteromedial margin sometimes emarginate and anterior margin sometimes truncated (Ophiomyia). Proboscis normal, uncommonly elongate; palpus normally narrow, small, and cylindrical, rarely enlarged or swollen.

Chaetotaxy: In remaining consistent with names occurring in most agromyzid publications, terminology here largely follows that outlined in McAlpine (1981) in the Manual of Nearctic Diptera, with the fronto-orbital setae (usually four pairs, rarely less than three, sometimes five or more) divided into two classes: the inclinate anterior fronto-orbitals are treated as the inferior orbital setae (ori), and the reclinate/slightly inclinate posterior fronto-orbitals are treated as the superior orbital setae (ors); the two sets are most distinct when viewed anteriorly, and are sometimes difficult to differentiate. Pedicel small, margins shallowly rounded, with one dominant dorsomedial seta. Postocellar setae (one pair) divergent, arising from immediately behind ocellar tubercle. Ocellar setae (one pair) divergent and arising from ocellar tubercle at, or behind level of anterior ocellus. Vibrissae (one pair) distinct; multiple vibrissae sometimes present and fused into pointed “fasciculus” or “horn” in male (some Ophiomyia). Orbital setulae usually in one or two rows, sometimes in several rows, rarely absent, reclinate to proclinate, sometimes inclinate or exclinate; mostly or entirely arising from orbital plate lateral to fronto-orbital setae. Two pairs of divergent vertical setae arising from orbital plate near posterior margin of frons, divided into inner vertical seta and outer vertical seta.

Thorax: Sclerites labelled in Figures 8, 10, typical of adult Schizophora.

Chaetotaxy: (Figs 8, 10) One postpronotal seta. Usually two notopleural setae, uncommonly one. One presutural and two postsutural supra-alar setae (sometimes mistakenly termed the intra-alar); one or two postsutural intra-alar setae (sometimes mistakenly termed the intrapostalar, but these may be distinguished in other families of Diptera as illustrated in Cumming and Wood (2017: figs 41, 42) by being on a discrete swelling in the extreme posterolateral corner of the scutum, the “postalar callus”, which is absent in Agromyzidae; default designation of the posterior-most intra-alar as the post-alar is not justified). Usually two to four dorsocentral setae (fourth seta near or anterior to transverse suture), with additional setae sometimes extending presuturally. One pair of prescutellar acrostichal setae present or absent; acrostichal setulae usually present in 2–10 rows (measured at level of transverse suture), but sometimes absent, becoming sparser and more irregular posteriorly, disappearing at or behind level of posterior dorsocentral. Scutellum with one pair of apical setae and usually one pair of anterolateral setae (absent in two subgenera of Cerodontha); scutellum otherwise bare, but Palaearctic Selachops with dorsum densely setose. One strong, upcurved proepisternal seta (absent in Gymnophytomyza). At least one strong anepisternal and katepisternal seta, but sometimes with additional smaller setae posteromarginally on anepisternum and mediodorsally on katepisternum (mostly Agromyzinae).

Legs: Short, slender, fore and mid legs not widely separated. Segments labelled in Figures 7, 8.

Chaetotaxy: Posteroventral margin of femora with row of strengthened setae (sometimes indistinct); posterior margin often with scattered setae. Fore tibia sometimes with one posteromedial seta (most Japanagromyza and some Melanagromyza, Nemorimyza and Phytobia). Mid tibia sometimes with one or two posteromedial setae, rarely three. Hind tibia sometimes with inner-distal “cleaning comb” (von Tschirnhaus 1991). Tibiae without dorsal preapical setae; at least mid tibia usually with small ventroapical seta.

Wing: (Figs 3, 4) Usually clear, rarely whitish or with faint to more distinct clouding, uncommonly patterned (more frequent in Neotropical species). Subcostal break present, humeral break absent; costa extending to R4+5 or M1+2; subcosta sometimes attaining costa, either as a vein or as a fold if the vein proper ends within the costal cell, but sometimes the subcosta is fused to R1 apically. Crossvein dm-m usually near midpoint of wing, sometimes shifted basally or absent; crossvein r-m present; vein bm-m rarely absent (some Pseudonapomyza). M1 and M4 sometimes faint to spectral; CuA+CuP usually faint to spectral, not reaching wing margin. Anal cell (cell cua) present, closed by vein CuA. Calypter (= squama of previous authors) prominent, semi-circular, and fringed with long hairs. Halter white, yellow, black, or bi-coloured.

Female abdomen: (Figs 1123) Tergites and sternites 1–6 well-developed and setose, sternites narrower; tergites 1 and 2 fused. Segment 7 with tergite and sternite completely fused, enclosing spiracles; forming non-retractable, well-sclerotised, and usually elongate and partially shiny oviscape (= ovipositor sheath of Sasakawa and Basalglied of Hendel) that is usually dark brown to black but uncommonly paler; tergite 7 anteromedially with large, hollow, keel-like apodeme that projects into segment 6; remainder of abdomen past segment 7 usually retracted within oviscape, with membranous region past oviscape. Elongate membranous region between segments 7 and 8 (= segment 8 of Sasakawa) with two pairs of sclerotised bands; dorsal and ventral membrane past these bands densely covered with anteriorly directed denticles used to drill into plant tissue. Tergite 8 absent; sternite 8 divided into one pair of short egg guides with shape and arrangement of posteromedial sensilla variable; used for scraping out hole for egg positioning. Segment 9 absent (McAlpine 1981: 44). Segment 10 with tergite and sternite narrow, sometimes fused and often variably modified, terminating in one pair of short, rounded setose cerci that bear stout apical trichoid sensilla. One pair of usually subspherical spermathecae, often with basal invagination at point of insertion to spermathecal duct (reinforced with minute spiralled sclerotisations, excluding smooth apex), which connects spermathecae to elongate, membranous genital chamber. Ventral receptacle recurved with variably shaped apex.

Male abdomen: (Figs 139–142, 147–150, 446–451) Tergites and sternites 1–6 well-developed and setose, sternites narrower; tergites 1 and 2 fused; sternite 6 often large and broadly emarginated posteriorly. Male pregenital segments largely reduced to sternite 8, which is narrow, transverse, and often symmetrical and setose; sternites 6 and 7 absent or occasionally represented by lightly sclerotised regions anterior to epandrium ventrolaterally; tergites 7 and 8 absent. Epandrium (= tergite 9) dorsal and dome-like with slightly thickened anterodorsal margin that often produces one pair of raised points laterally to facilitate articulation with hypandrium. Surstylus (one pair) usually present, setose, variably shaped, sometimes with small tubercle-like setae on inner surface; base articulates with, or is variably fused to inner-ventral surface of epandrium or subepandrial sclerite. Cercus (one pair) variably sized, usually setose, sometimes with short spines on inner surface. Subepandrial sclerite (= sternite 10, or ventral epandrial sclerite of some authors) variably shaped, resting between inner surface of epandrium and hypandrial complex; usually plate-like with one pair of well-sclerotised dorsolateral arms (= bacilliform sclerites); sometimes medially divided, often with a pair of setae or spine-like processes, and/or one pair of paler ventral processes that may become thickened and enlarged (Fig. 490) (see Cerodontha section below for discussion of the structure called “Langfortsatz” by previous authors). Hypandrium (= sternite 9) U- or V-shaped with narrow transverse inner-basal lobe (“pregonite” of some authors, including Frick 1952a) bearing two or three apical setae that may be separate or weakly connected to the main body (e.g., Ophiomyia genus group); articulates with external components of genitalia via posterobasal “arms”. Pregonite absent (sections of postgonite or hypandrium sometimes interpreted as “pregonite”, such as by Papp and Černý 2015, 2020). Postgonite (one pair) usually long and narrow with one (but sometimes zero or several) seta and often small outer-medial sockets, usually slightly upturned and medially cleft apically; articulated with, surrounded by, or fused to inner lobe of hypandrium; subovate and plate-like in some Agromyzinae excluding Agromyza, but sometimes absent or possibly fused to epiphallus. Phallapodeme (= aedeagal apodeme of some authors) long and rod-like with extensive muscular attachment to project phallus from abdomen. Ejaculatory duct originates in ejaculatory apodeme and terminates in the distiphallus (“distal section” of Griffiths 1972a), the last section of the phallus that is sometimes divided into an apical distiphallus proper and a basal mesophallus. The distiphallus appears to be ancestrally bifid in the family, being subtly to conspicuously split in most Phytomyzinae with the paired tubules united or atrophied in a number of lineages (including species of Xenophytomyza, Aulagromyza, Haplopeodes, Phytomyza, etc.) or concealed in a basal enclosure (many Liriomyza, Pseudonapomyza and Phytomyza parvicella (Coquillett)); the paired tubules of the distiphallus are usually less obvious or entirely unified in most Agromyzinae (but see Figs 148, 256, for example), with almost all representatives of the Ophiomyia genus group appearing to have a single exit pore (Lonsdale 2014: fig. 38); in some Phytomyza, the dorsobasal margin of the distiphallus branches into a sclerotised structure called the “supporting sclerite” (Fig. 752). Basiphallus (“basal section” of Griffiths 1972a) divided into two segments, the basiphallus proper (“basal sclerites or paraphalli” of Griffiths 1972a, “endophallus” of Guglya 2013, “mesophallus” of Papp and Černý 2015), which is one or two plates (rarely more) laterally flanking ejaculatory duct before it enters distiphallus, and the basal phallophorus, a tubular or ring-like basal segment through which the duct enters; phallophorus allows for articulation between phallapodeme and remaining distal components of phallus, is fused to the epiphallus and sometimes the basiphallus, and is sometimes flanked by one pair of sclerites that may be derived from the epiphallus (some Ophiomyia) or the phallapodeme (most Phytomyza). Hypophallus (= ”medial lobe” of Griffiths 1972a, “trough-like sclerite” of Griffiths 1974 and “wedge-shaped sclerite” of Griffiths 1980) a lobate membranous region ventrally near apex of basiphallus that is variably sclerotised and/or expanded; if present, sclerites may articulate or fuse with basiphallus or (less commonly) disti/mesophallus; sclerotisations may consist of one pair of plates or rods that may meet each other ventrally and/or reach basiphallus dorsally; sclerite sometimes present as single small medial bar that may end in hair-like structures (many Liriomyza) and/or widen to form flat, irregular plate (as in some Calycomyza). Paraphallus (“paramesophalli” of Griffiths 1972a) (one pair) small, sclerotised, and usually plate or rod-like, usually directed ventrally to ventrodistally; positioned ventrolaterally at base of distiphallus / mesophallus; possibly homologous to transverse sclerite wrapped around the posterobasal surface of distiphallus / mesophallus in Agromyza. Epiphallus divided into two parts, a smaller, darker proepiphallus (“epiphallus” of Frick 1952a), which articulates with phallophorus, and a wider, flatter metepiphallus (“aedeagal hood” of Frick 1952a), which articulates/fuses to proepiphallus and postgonite and/or hypandrium; components of epiphallus lie along each other in repose and fold outwards during copulation to extend phallus, but folding mechanism abandoned in most Agromyzinae, with reduced proepiphallus and usually enlarged and elaborated metepiphallus separated by membranous space. Ejaculatory apodeme composed of large, flat sclerotised blade with asymmetrical base, and smaller, clear basal sperm pump that often has transverse sclerotisation (= pileus ejaculatorius of Steyskal).

Notes on male genitalic homology. Regarding structural homology, a number of points must be clarified. Firstly, as the basalmost component of the phallus meeting the phallapodeme, the phallophorus is clearly homologous with the “basiphallus” of other Schizophora. The term “phallophorus” is here retained, and treated as the basal segment of the basiphallus proper. As the structure is present and often distinct in taxa throughout the family, and it has had wide usage in the literature, the term has utility. It is not entirely certain whether the agromyzid “basiphallus” originates from the same sclerite as the phallophorus, but in assuming this homology, continuity is maintained with the existing literature, and confusion is avoided by eliminating the need to rename structures. The alternative interpretation would be to consider the agromyzid “basiphallus” a novel, secondary sclerotisation of the lateral phallic membrane, but this explanation is not more parsimonious and it does not have the above advantages. Papp and Černý (2015) partially agree with this interpretation, although their “basiphallus” consists solely of the phallophorus; what they call the “mesophallus” is the basiphallus of this study and other works.

Both the phallophorus and folding epiphallus appear to have counterparts in other acalyptrate Diptera and may be phylogenetically informative, but the homology and polarity of these are uncertain. Both structures appear to aid in projecting the phallus from the abdomen for copulation, contrasting the lever and fulcrum method employed by families such as Clusiidae, or the method employed by Tephritoidea, etc., but functional requirements may make these prone to convergence.

Lastly, the position and shape of the hypophallic sclerites in the ventral region distal to the basiphallus varies widely between species, with numerous intermediate forms. Some complex forms have led some authors (e.g., Griffiths 1972a, 1974, 1980) to invent novel terminology for the sclerites present. Noting this variation, and the absence of any sclerotised structures in some taxa, there is the possibility of repeated loss and gain of analogous structures that may be difficult to interpret. Until a more thorough analysis of the structures in this region is made to establish homology, the single term “hypophallus” will be applied to the membranous lobe in this region, including any sclerotised structures found thereupon.

Agromyzidae – immature description

Egg: Small and clear with shape ovate to reniform; surface texture smooth to reticulate. End bearing small micropyle often broader.

Larva: (Figs 25–31) Typically maggot-like, cylindrical, soft and with ends tapered; Phytobia species often longer and narrower (Greene 1914, 1917; Spencer 1987); form varying very little between species with different feeding habits such as leaf-mining and stem-boring (Dempewolf 2005). Mandibles usually asymmetrical; firmly fused at base with movement along the same axis (Spencer 1987); each with as few as one or two accessory hooks, which may be duplicated and variable in shape; often becoming reduced and strengthened in boring species (Dempewolf 2005). Hypopharyngeal sclerite short, linking mandibles to tentopharyngeal sclerite, which has one pair of “arms” (or “cornus”); Agromyzinae with dorsal and sometimes ventral arms of tentopharyngeal sclerite encompassing small “window” that is open posteriorly in dorsal arm (Fig. 29); Phytomyzinae sometimes with window in ventral arm only (Figs 24, 25, 30, 31; Spencer 1987). Anterior and posterior spiracles with at least three or five (but sometimes many more) slit-like openings in clusters of lobes/bulbs that may be narrow and reduced or strongly projecting and expanded, especially in species that use spiracles to anchor puparium to host plant surface (Figs 35, 36); posterior spiracles sometimes with “hook” (Fig. 34); anterior spiracles closely spaced dorsally (Figs 24, 27) (Ferrar 1987; Spencer 1987).

Puparium: (Figs 26–36) Surface smooth and shiny; colour usually pale yellow to reddish brown or dark orange (Figs 26–28), but sometimes white or deep black with metallic shine (e.g., Cerodontha), with overwintering puparium dark brown to black. Shape slender or flattened to more compact and barrel-shaped; segmentation often clearly evident, minute sculpturing or pits sometimes present. Puparial lid with additional split along the midline in species of Ophiomyia genus group (compare Fig. 33 to Fig. 32; Dempewolf 2001).

Key to the north American genera of Agromyzidae

1 Subcostal vein well-developed along entire length and fused to vein R1 before reaching costa (Fig. 4). Vein R1 sinuous for a short distance just before reaching costa, and sometimes thickened (relatively straight in some Agromyza, which has a distinct stridulatory file on lateral margin of tergite 2). Vein R4+5 ending as close, or closer to wing apex compared to vein M1. Lunule often with medial furrow, well-sclerotised and flat to bulging (recessed in some Agromyza and some poorly preserved material) Agromyzinae (2)
Subcostal vein either ending freely in cell or continuing as a fold or atrophied vein to the costa (Fig. 3). Vein R1 straight, not deviated before reaching costa, infrequently thickened apically (excluding some Phytobia). Lateral margin of tergite 2 never with stridulatory file. Vein M1 usually closer to wing apex compared to vein R4+5 (except in Phytobia and some Nemorimyza). Lunule without medial furrow (except in some Phytobia), often soft, and usually flat or recessed (infrequently bulging) Phytomyzinae(8)
2 Prescutellar acrostichal seta present (Figs 10, 51); halter usually white, at least apically 3
Prescutellar acrostichal seta absent and halter entirely dark (apically white in Ophiomyia maculata Spencer) 4
3 Lateral margin of tergite 2 and fused tergite 1 with straight, thin, and well-sclerotised stridulatory file (Fig. 38). Usually at least three conspicuous dorsocentral setae (Fig. 10). Fore tibia without medial seta. Frons sometimes soft and pale; body never with metallic shine Agromyza Fallén
Lateral margin of tergites 1 and 2 weakly sclerotised and sometimes irregular in outline, never with stridulatory file. Only two well-developed dorsocentral setae (Fig. 51), frons well-sclerotised, rounded and dark, and gena shallow (Figs 49, 50) (J. rutiliceps with four dorsocentrals, a projecting frons and gena ¼ height of eye); fore tibia usually with medial posterolateral seta; body sometimes with metallic shine Japanagromyza Sasakawa (in part)
4 Clypeus with anterior margin truncated anteromedially (Figs 71, 73); sometimes narrowed (Figs 77, 80), often with conspicuous corners anteriorly and arms parallel. Sometimes with one or more of a pronounced facial keel that may have a medial bulb, an anteriorly produced gena, and multiplicated vibrissae forming a “fasciculus” or “horn” (Figs 75–77) Ophiomyia Braschnikov
Clypeus rounded anteromedially; if slightly truncated (some Melanagromyza), then with one or more of the following characters: dorsal setulae on eye; one posteromedial seta on fore tibia; more than 1 posteromedial seta on mid tibia; entire thorax and abdomen strongly metallic. Gena sometimes angled anteriorly (M. buccalis, Fig. 53), but never forming a point; vibrissa single, simple 5
5 Fore tibia with small to well-developed posteromedial seta (Fig. 121) 6
Fore tibia bare posteromedially 7
6 Halter white. Costa extending to M1. Only two developed dorsocentral setae (Fig. 51). Ocellar triangle only reaching level of posterior fronto-orbital seta; frons and lunule smooth with minute pruinosity; orbital plate not raised (Figs 49, 50). Eye bare or with few small, inconspicuous ommatrichia. Epandrium without spine. Basiphallus asymmetrical; often with left and right sides divided (Figs 255, 256). Mesophallus (or if absent, insertion of ejaculatory duct) basal to distiphallus; often forked or membranous. Distiphallus without paired ventrobasal tubules Japanagromyza Sasakawa (in part)
Halter brown. Costa extending to M1 or (uncommonly) R4+5. Usually two dorsocentral setae, uncommonly three or four. Ocellar triangle exceeding level of posterior fronto-orbital seta; frons variable, lunule smooth; orbital plate distinct and often raised or bulging (Figs 52–57). Eye often conspicuously with ommatrichia dorsally, particularly in males. Epandrium usually produced into small, but conspicuous spine posteroventrally (Fig. 267). Basiphallus symmetrical; ring- (Fig. 304) or U-shaped (Fig. 256). Mesophallus inserted ventromedially into distiphallus as a cylindrical chamber (Figs 264, 265). Distiphallus with paired ventrobasal tubules, often flanking mesophallus Melanagromyza Hendel (in part)
7 Thorax (and sometimes head and abdomen) sometimes with green (Fig. 60), blue or coppery metallic sheen. Eye often conspicuously with ommatrichia, particularly in males. Orbital plate usually small, not much visible laterally, but sometimes prominent as below. Mid tibia with 1 or 2 posteromedial setae. Postsutural intra-alar setae unequal in length with anterior seta stronger. Epandrium usually with posteroventral spine (Fig. 267). Phallophorus simple, cylindrical (Figs 264, 265). Basiphallus symmetrical; ring-like or U-shaped. Distiphallus with paired ventrobasal tubules, often flanking mesophallus Melanagromyza Hendel (in part)
Thorax without metallic sheen (Figs 4448). Eye bare. Orbital plate bulging, forming prominent ring with parafacial. Mid tibia without posteromedial setae. Postsutural intra-alar setae approximately subequal. Epandrium without spine as below (Fig. 250). Phallophorus produced anteroventrally into a large pouch. Basiphallus asymmetrical. Distiphallus never with paired basal tubules Euhexomyza Lonsdale
8 Lunule conspicuous, at least as high as pedicel, well-sclerotised and usually convex (even if partially concealed by pronounced orbital plates) (Figs 99–105); ranging from subcircular to long and narrow, but if shallower than a semicircle, lunule very large and broad (some Dizygomyza). Orbital setulae sometimes only reclinate in part, usually entirely erect to reclinate; vein dm-m always present. Distiphallus often long, robust and bifid (sometimes secondarily fused or more membranous) with apex usually swollen or bell-shaped (Figs 493, 494, 511, 512, 543, 544) Cerodontha Rondani (9)
Lunule weakly sclerotised, slightly concave and usually shallow; if appearing higher than pedicel, then either orbital setulae entirely proclinate (Phytomyza and some Phytoliriomyza) or epandrium with rows of conspicuous tubercle-like setae (some Phytoliriomyza). Distiphallus various 15
9 First flagellomere pointed (Fig. 106) or angled anterodorsally. Scutellum with lateral scutellar setae absent (i.e., only one pair of setae present apically) 10
First flagellomere rounded anteriorly. Scutellum with two pairs of setae (some global Poemyza rarely with one pair, but lunule very high and narrow) 11
10 First flagellomere produced as a thin point anterodorsally (Fig. 106); point uncommonly reduced, some non-Nearctic species with point absent and segment simply angulate. Body with conspicuous yellow patches C. (Cerodontha) Rondani
First flagellomere angulate anterodorsally. Body brown to black (Fig. 104) C. (Xenophytomyza) Frey
11 Costa only extending to vein R4+5 C. (Phytagromyza) Hendel
Costa ending at vein M1 12
12 Prescutellar acrostichal seta present. Lunule slightly higher than wide, well-sclerotised, smooth, gradually tapering to a point dorsally, and light brown to brown with even covering of very short velvety pubescence (Fig. 99) C. (Butomomyza) Nowakowski
Prescutellar acrostichal seta usually absent, but uncommonly weak to well-developed. If lunule higher than wide, then not as above. Lunule uncommonly approaching state as defined above, but if so (e.g., C. (D.) scirpivora Spencer), then lunule smoothly rounded above and male first flagellomere densely haired 13
13 Lunule narrow, either higher than wide or appearing higher than wide with sides obscured beneath prominent orbital plates (Figs 101, 102); surface often shiny and minutely textured C. (Poemyza) Hendel
Lunule as wide, or wider than high; surface smooth with indistinct to prominent pubescence 14
14 Lunule flat or curved, meeting anterior margin of frons, well-sclerotised (Fig. 105). Head usually entirely grey to dark brown. Male first flagellomere often enlarged and sometimes with long, conspicuous hairs; antennal bases often distinctly separated C. (Dizygomyza) Hendel
Lunule bulging, usually projecting higher than sunken anterior margin of frons, weakly sclerotised (Fig. 103). Lunule bright yellow with frons sometimes also yellow. Male first flagellomere not larger than those of females and with hairs moderately sized; antennal bases closely approximated C. (Icteromyza) Hendel
15 Vein R4+5 as close, or closer to wing apex than vein M1 (Figs 689, 690). Lunule often silvery or with shiny pubescence (Fig. 124). Usually large and stout-bodied. Subepandrial sclerite stout, with halves broadly fused, and usually with single large spine (pronounced in Nemorimyza) (Figs 670, 684) 16
Vein M1 closer to wing tip than vein R4+5 (Figs 429, 693, 836). Lunule often bare or pubescent, rarely silvery. Usually small and slender-bodied. Subepandrial sclerite various, but u ncommonly as above 17
16 Fore tibia sometimes with posteromedial seta (Fig. 121); halter sometimes with dark spot. Surstylus with base elongate, strongly projecting dorsally into epandrium (Figs 662, 663). Basiphallus composed of two wide interlocking plates; apices flanking hypophallus narrow and curved (Figs 665, 666). Distiphallus stout, with abruptly widened base. Common Holarctic species (N. posticata (Meigen)) with male abdomen posterior to tergite 3 or 4 yellow Nemorimyza Frey
Fore tibia rarely with posteromedial setae. Halter entirely white. Surstylus small, lobate (Figs 683–685). Male abdomen usually entirely dark, never as above. Basiphallus not as above; narrowed with halves fused basally to form a stalk (Fig. 681). Distiphallus usually narrower than above, at least towards base Phytobia Lioy
17 Costa extending to vein M1 (Fig. 463) 18
Costa extending to vein R4+5, or slightly past (Fig. 435) 24
18 Two distinct dorsocentral setae, with anterior two pairs highly reduced and barely longer than surrounding setulae (few global species with three or four setae developed). Head and notopleuron pale yellow with antenna and most of remaining notum contrastingly dark (some exceptions outside of Nearctic) (Figs 9098). Epandrium and surstylus each with dense clusters of tubercles (Figs 446–448) Calycomyza Hendel
Three or more pronounced dorsocentral setae; if only two present, colour never as above. Arrangement of tubercles on epandrium usually much reduced (Figs 621, 626), arranged in rows (Fig. 710) or otherwise not as above (but see Liriomyza quadrisetosa – Figs 616–620) 19
19 Halter white to entirely black, but never white with apical surface brownish. Distiphallus usually black, dense, and surrounded by minutely spinulose membrane (Figs 424, 426). Gena usually 1/3–1/2 height of eye (Fig. 87), but sometimes shallow (Fig. 85). Ejaculatory apodeme large, bowl-shaped sclerotisation on sperm pump (Fig. 418) Amauromyza Hendel
Halter white, sometimes with apical surface of halter brown. Distiphallus without spinulose membrane. Gena usually < 1/3 eye height, uncommonly higher. Sperm pump uncommonly as above, usually with much smaller, ovate or band-like sclerotisation (Figs 464, 472, 531, 588) that may be vestigial to absent (Fig. 659) 20
20 Scutellum yellow centrally and orbital plate brown and slightly to conspicuously pronounced, contrasting with the weakly sclerotised central vitta (also usually brown). Broad, sclerotised ocellar triangle and orbital plates joined to form sharp angles with the outline of an “M”. Surstylus with at least one, and epandrium with two rows of tubercle-like setae (Figs 654–657) Metopomyza Enderlein
If orbital plate dark brown as above, then not pronounced and scutellum entirely brown. If ocellar triangle and orbital plate joined along posterior margin of frons, then broadly joined by an intervening space (i.e., connecting angle broadly rounded or squared). Male genitalia infrequently as above (some Phytoliriomyza) 21
21 Apical surface of halter light brown (Fig. 127). Orbital setulae proclinate. Eye with minute ommatrichia. Side of frons with light silvery pubescence (Fig. 126). Scutellum entirely brown or yellowish medially Phytoliriomyza Hendel (in part)
Halter entirely white. Orbital setulae upright to reclinate, sometimes absent; if proclinate (some Phytoliriomyza) then epandrium with dense row of tubercle-like setae. Eye bare or with few scattered ommatrichia. Side of frons not reflective as above. Scutellum usually distinctly yellow medially 22
22 With one of the following characters: frons with slight reflective pruinosity anterolaterally; orbital setulae proclinate; first flagellomere slightly elongate and black; scutellum and large portions of scutum matt grey; epandrium and surstylus with rows of tubercle-like setae (Fig. 705) Phytoliriomyza Hendel (in part)
Frons never with reflective pruinosity. First flagellomere yellow to dark brown, never black. Scutellum usually subshiny to shiny (sometimes pruinose) and with central yellow stripe (sometimes reduced to absent). Epandrium and surstylus usually with one or two tubercle-like setae, but if with more, these never arranged in a discrete row 23
23 Vein dm-m often present (Fig. 652). Male abdomen sometimes with anterolateral stridulatory organ in membrane (Fig. 107). Phallus with at least two distinct sclerotised segments (Figs 593, 594). Apical section of ejaculatory duct pigmented and abruptly dilated Liriomyza Mik
Vein dm-m often absent (Fig. 581). Male abdomen never with stridulatory organ as above. Phallus small and membranous with base sclerotised (Figs 579, 580). Apical section of ejaculatory duct clear and of equal width along length Haplopeodes Steyskal
24 Orbital setulae proclinate (Fig. 133). Phallophorus often flanked by one pair of rod-like sclerites (Figs 803, 804) Phytomyza Fallén s. l. (25)
Orbital setulae reclinate or upright (as in Fig. 2). Phallophorus without paired lateral sclerites 26
25 Vein dm-m present; usually in line with, or slightly basal to r-m, but sometimes slightly distal to r-m as below. Scutum mostly dark with grey pruinosity (Fig. 131). Distiphallus undivided apically, with apical sclerotised ring or tubule connected to dorsal sclerotisation on shaft P. (Napomyza) Westwood
Vein dm-m usually absent; if present, then positioned slightly distal to r-m (Fig. 435). Scutum variably coloured and tomentose (Figs 132–138). Distiphallus usually divided apically; if undivided and with apical ring, then ring continuous with ventral sclerotisation along shaft P. (Phytomyza) Fallén
26 Vein r-m basal to dm-m (Fig. 88); if dm-m absent, then r-m positioned far from vein bm-m and colouration bright yellow (Fig. 89). Vein M sclerotised far basal to vein r-m. First flagellomere rounded apically, sometimes with dorsomedial margin forming a slightly raised angle Aulagromyza Enderlein
Vein dm-m and basal section of M absent, with r-m and bm-m nearly forming a straight transverse line (Fig. 833) [some Old World species with vein r-m distal to vein dm-m, with dm-m enclosing a very small cell between it and bm-m]. Colour always black (Fig. 130). First flagellomere with projecting anterodorsal angle [Old World species outside P. atra group with segment rounded] Pseudonapomyza Hendel

Taxonomic treatment

Agromyzinae

Agromyza Fallén

Agromyza Fallén, 1810: 21. Type species: reptans Fallén 1823, by subsequent designation [Rondani 1875: 168]. Described without named species. Westwood 1840: 151 [invalid design. of nigripes Meigen 1930 as type]; Rondani 1856: 121 [designation of aeneoventris Fallén as type (now Melanagromyza)], 1875: 168 [designation of reptans as type]; Frick 1952a: 370 [synonymy]; Spencer 1969: 29; Spencer and Steyskal 1986a: 183 [proposed fixation of reptans as type], 1986b: 57; ICZN 1988: 76 [fixation of reptans as type].

Calyptomyza Hardy, 1850: 486. Type species atra Hardy, 1850 [= Agromyza lathryi Hendel, 1923], by monotypy (Bland 2000; synonymy).

Adromyza. Misspelling. Meigen 1830: viii.

Domomyza Rondani, 1856: 121 [incorrectly treated as Agromyzidae]. Type species: cincta Rondani, 1956 (= Cacoxenus indagatorLoew 1858), by monotypy. Sidorenko 1996: 21 [petition to suppress Domomyza – senior synonym of Cacoxenus Loew, 1858 (Drosophilidae)]; ICZN 1997: 195 (suppression of genus name).

Mesonevra Lioy, 1864: 1312. Type species: Agromyza mobilisMeigen 1830, by monotypy. Syn. Frick (1952a).

Cecidomyiaceltis Patton, 1897: 247. Type species: Cecidomyiaceltis desertaPatton 1897: 247, by original designation. Syn. von Tschirnhaus (2016).

Stomacrypolus Enderlein, 1936a: 178 [nomen nudum – no type species designated].

Stomacrypolus Enderlein, 1936b: 42 [attributes to Enderlein 1936a]. Type species: Agromyza ambigua Fallén, 1823, by original designation. Frick 1952a [synonymy not explicit]. Spencer and Martinez 1987: 255 [listed as “error (Enderlein 1936b)”].

Both sexes of Agromyza have a dark, straight, well-sclerotised file along the lateral margin of the fused first and second tergites, making it an easily recognised genus. The second tergite itself is also produced posteriorly along the lateral margin, forming a shallow, yet distinct angle (Figs 37, 38). Species in this genus also have one pair of prescutellar acrostichals and three or more dorsocentrals, although several, including A. bispinata Spencer, A. parilis Spencer and their relatives usually have only two that are immediately apparent. Internally, most species have a transverse sclerite wrapped around the posterobasal surface of the mesophallus (which is often fused to the distiphallus and possibly homologous to the paraphallus), but this is absent in some taxa such as A. varifrons Coquillett and A. vockerothi Spencer. There is also no basal sclerotisation on the sperm pump (also absent in many Japanagromyza), there is a haired membrane extending from the posterior face of the hypandrial lobe (sometimes indistinct), and most of those species with a transverse sclerite on the distiphallus also have a unique phallus (Figs 168, 169): the two sclerites of the basiphallus are approximated basally, divergent distally and the apex is abruptly elbowed inwards; the S-shaped distiphallus (including the fused mesophallus) is usually capsule-like with a ventral subbasal, crack-like fissure for entry of ejaculatory duct. This phallus type is found in numerous species, including those in the A. nigripes group, a diverse, predominantly Poaceae-feeding clade (Hendel 1931; Griffiths 1963; Spencer 1990: 353).

For the purposes of species-level identification, the morphology (including shape, colour, and hairiness) of the first flagellomere is quite important, and if these are missing from the specimen it is recommended that the male genitalia be dissected.

Key to the Delmarva species of Agromyza

1 Palpus yellow (apex infuscated in A. apfelbecki). Face white to light brown, rarely brown. First flagellomere usually light yellow, sometimes with outer surface lightly infuscated 2
Palpus brown or darker. Face light to dark brown. First flagellomere orange to dark brown 8
2 At least 6 dorsocentral setae and 5 fronto-orbital setae. Wing length at least 4.0 mm. Epistoma large, distinct A. apfelbecki Strobl
No more than 4 large dorsocentrals and four fronto-orbital setae. Wing length 3.4 mm or less. Epistoma narrow 3
3 Body, excluding most of head and legs, yellow. Thorax shiny. Mid tibia with 2 or 3 posteromedial setae. First flagellomere with apical patch of slightly longer, paler hairs. Wing length 2.7–3.4 mm. Eye 6.1–9.7 × higher than gena. Surstylus indistinct, appearing as broadly rounded ventral margin of epandrium; with tubercle-like setae (Fig. 180). Cercus only with unmodified setae. Distiphallus terminating in a large, dark bulb (Figs 181, 182). Basiphallus consisting of 1 pair of lateral bands A. diversa Johnson
Body, excluding most of head, dark brown. Thorax densely to lightly pruinose. Mid tibia without posteromedial setae. First flagellomere with all hairs similar. Wing length usually 1.7–2.4 mm; 3.4 mm in A. deserta. Eye 1.6–3.9 × higher than gena. Surstylus ending in triangular point; without tubercle-like setae. Cercus with tubercle-like setae along inner face. Distiphallus a flat, curved sclerite. Basiphallus consisting of a single band 4
4 Thorax with dense grey pruinosity. Calypter margin and hairs white. Legs light yellow. Parafacial visible laterally, continuing under eye as distinct cheek. Ca. 8 rows of acrostichal setulae. Eye 1.6–2.7 × higher than gena. Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.9. Mesophallus several ×longer than distiphallus (males unknown for A. pallidiseta). Ejaculatory apodeme minute and subcylindrical 5
Thorax with light dusting of pruinosity. Calypter margin and hairs brown. Legs dark brown with tarsi and knees paler. Parafacial not visible laterally; cheek not strongly delimited. Four rows of acrostichal setulae. Eye 3.5–3.9 × higher than gena. Ultimate and penultimate sections of vein M4 of equal length. Mesophallus shorter than distiphallus. Ejaculatory apodeme large and fan-shaped 7
5 Setae yellow. Ocellar seta 1/2 length of postocellar A. pallidiseta Malloch
Setae dark brown. Ocellar seta as long, or nearly as long as postocellar 6
6 Wing length 2.0–2.4 mm. Legs yellow with mid and hind coxae at least partially brown (Fig. 37). Presutural dorsocentral not much longer than surrounding setulae. Distiphallus straight (Figs 165, 166) A. aristata Malloch
Wing length 3.4 mm. Legs brown with apex of femora and base of tibiae yellow. Presutural dorsocentral distinct. Distiphallus curved A. deserta (Patton)
7 First flagellomere entirely light yellow, or with dark dorsobasal spot. Mid and hind tibiae dark brown with base yellow, and fore tibia yellow with centre brownish. Male cercus thin along length (Fig. 237) A. varifrons Coquillett
First flagellomere yellow with outer face lightly infuscated on distal 2/3. Tibiae dark brown with only base of fore tibia yellowish. Male cercus broad and V-shaped (Figs 186, 187) A. fission Eiseman & Lonsdale
8 Five or 6 fronto-orbital setae, with only one ors (some A. virginiensis with posterior ori appearing as ors). Fronto-orbital plate and parafacial not projecting. Centre of frons always brownish orange. Distiphallus long, thin, tubular, and bifid 9
Four fronto-orbital setae on one or both sides of frons, with two ors, but if 5 fronto-orbitals present (A. aprilina), fronto-orbital plate and parafacial projecting. Centre of frons usually dark brown (pale in A. aprilina). Distiphallus undivided 10
9 At least apical 1/2 of first flagellomere brown. Anterior margin of buccal cavity sharply angulate. Distiphallus strongly curved, forming a semicircle (Figs 147, 148) A. ambrosivora Spencer
First flagellomere entirely orange, with dark spot around base of arista, or with distal 1/2 darker. Anterior margin of buccal cavity straight or slightly rounded, at most with midpoint slightly produced. Distiphallus nearly straight (Fig. 151) A. virginiensis Spencer
10 First flagellomere black; elongate to slightly enlarged and with anterodorsal angle. Body dark brown to black. Calypter margin and hairs white. Distiphallus relatively short 11
First flagellomere dark brown or paler; circular to elongate oval, with apex broadly rounded; if flagellomere black and with anterodorsal point (some A. kincaidi), then segment not elongate, body paler, calypter margin and hairs usually yellow to brown, and distiphallus large and clavate 12
11 Costa extending just past vein R4+5. Five fronto-orbital setae. Sides of ocellar triangle straight. Anterodistal margin of epandrium deeply incised before surstylus (Figs 157, 158). Surstylus acutely triangular and directed anteriorly. Distiphallus tapered to apex, which is directed distally (Fig. 160) A. aprilina Malloch
Costa extending to vein M1+2. Four fronto-orbital setae (some Canadian material with five). Sides of ocellar triangle slightly concave. Epandrium without deep emargination as above. Surstylus relatively small and lobate (Fig. 143). Distiphallus parallel-sided with apical 1/2 slightly angled dorsally (Fig. 146) A. albipennis Meigen
12 Four strong dorsocentral setae, presutural seta nearly as long as preceding seta and nearly as long as acrostichal seta. Thorax covered with brownish grey pruinosity, strongly contrasting brown to golden-brown setae. Mid tibia with 1 posteromedial seta. Basiphallus consisting of a single sclerite. Distiphallus and separate mesophallus simple and plate-like 13
Three to 6 dorsocentral setae, strongly decreasing in height anteriorly (anterior setae often barely larger than surrounding setulae). Thorax usually shiny, but sometimes with light pruinose dusting. Setae usually dark brown, but sometimes brown or (rarely) yellow. Mid tibia with two posteromedial setae. Basiphallus consisting of 2 sclerites. Distiphallus (including fused mesophallus) with transverse dorsobasal band; bulb-like or long and bifid 14
13 Calypter hairs brown. Antenna entirely dark brown. Mid and hind tibiae brown with apices yellow (base sometimes also yellow). Distiphallus longer than wide, flat, and curved; short, cylindrical with longer ventral plate (Figs 248, 249) A. vockerothi Spencer
Calypter hairs usually white (only brown in holotype). Base of first flagellomere paler in colour, usually distinctly orange. Mid and hind tibiae yellow, sometimes with hind tibia brownish medially. Distiphallus very short, mesophallus dense and thick apically, and with long, thin, flat “tail” (Figs 191, 192) A. isolata Malloch
14 First flagellomere usually at least 1.3 × longer than pedicel (larger in males); shape circular (most males) or elongate oval (some males and most females); with elongate pale hairs present and restricted to distal margin in females, but often extending to basal 1/2 or basal 1/3 of segment in males (Figs 41, 42); long-haired section without sharply delimited boundary, and with hairs relatively more sparse than in “tufted” state described below. Only males can be identified further 15
First flagellomere not much longer than pedicel, with shape ovate; longer hairs, if present, restricted to discrete, sharply delimited apical tuft that may form a minute spot (Figs 39, 40) or encompass the distal 1/3 of the segment 17
15 Surstylus with 2 tubercle-like setae of variable size; small and lobate, directed anteriorly and barely visible laterally (Figs 170–173) A. bispinata Spencer
Surstylus with 3 or more tubercle-like setae, rarely 2; subtriangular in outline and distinct laterally 16
16 Surstylus with 3–5 tubercle-like setae (rarely 2) (Figs 226–228). Margin of surstylus, seen laterally, bulging at base of tubercles A. tacita Spencer
Surstylus with numerous tubercle-like setae (Figs 232, 233). Margin of surstylus, seen laterally, relatively straight A. echinalis sp. nov.
17 If present, pale tuft of hairs on first flagellomere encompassing entire distal 1/3 of segment, or nearly as high as segment and teardrop-shaped. Scutum with light pruinosity. Four or 5 dorsocentrals, often with 1 presutural. Distiphallus cup-shaped with central tuft of hairs (strongly enlarged in A. canadensis) 18
If present, pale tuft of apical hairs on first flagellomere circular to ovate in outline, and ranging in size from 1/2 height of first flagellomere to only as wide as base of arista. Scutum usually shiny to subshiny. Number of dorsocentrals variable, but often with only 2 well-developed setae. Distiphallus capsule-like and usually bell-shaped in outline 19
18 Three postsutural dorsocentral setae and one sutural to presutural dorsocentral (European specimens with 1 or 2 additional dorsocentrals). Buccal cavity curving onto face. Parafacial entirely visible laterally. Frons orange-brown with fronto-orbital plate dark brown. Distiphallus as long as mesophallus, and with distal 1/2 flared (Figs 141, 142). Mesophallus with dorsal collar A. abiens Zetterstedt
Four postsutural dorsocentral setae and sometimes 1 sutural to presutural dorsocentral (Fig. 10). Margin between face and buccal cavity abruptly angled. Parafacial barely visible laterally (Fig. 8). Sides and centre of frons the same colour. Distiphallus much enlarged, ring-shaped (Figs 177, 178). Mesophallus without collar A. canadensis Malloch
19 Costa extending to vein R4+5. Wing length 3.6–4.3 mm. Distiphallus longer than basiphallus and dark (Figs 196, 197) A. kincaidi Malloch
Costa extending to vein M1+2. Wing length < 2.6 mm. Distiphallus shorter than basiphallus and usually pale 20
20 Five dorsocentral setae, with presutural dorsocentral as long as preceding setae, and nearly as long as acrostichal seta. Calypter hairs white to light brown in male (female unknown). Surstylus with four spines (Fig. 222). Distiphallus broad, short, and very dark (Figs 223, 224). Dark, sclerotised hypophallus present A. soka Eiseman & Lonsdale
Only 2 or 3 well-developed postsutural dorsocentrals, with third seta (if present) usually much shorter than acrostichal. Calypter hairs sometimes brown in male, usually white in female. Surstylus with > 4 spines. Distiphallus pale and bell-shaped in ventral view. Hypophallus absent 21
21 Frons and most of gena orange-brown, or at least distinctly paler than scutum; fronto-orbital plate and posterior region of frons variably dark brown (frons appearing entirely dark in poorly preserved specimens). Lunule light brown to yellow. Frons buckled anteriorly, with anterior region visible in lateral view. Distiphallus bulging subapically 22
Frons, gena and lunule entirely dark brown, with frons evenly curved to meet face. Sides of distiphallus relatively straight or concave subapically 23
22 Fronto-orbital plate always dark brown to level of anterior or posterior ors, and sometimes dark along most of lateral margin. Distiphallus strongly swollen distally, almost circular in shape (viewed ventrally) (Fig. 214) A. parvicornis Loew
Fronto-orbital plate usually brownish orange around base of all fronto-orbital setae, but sometimes as above. Distiphallus only gradually swollen subapically, with distal section only slightly wider than basal section (viewed ventrally) (Fig. 215) A. proxima Spencer
23 Antenna dark brown with base of first flagellomere sometimes orange. Basiphallus produced distolaterally (i.e., apex split); without membranous lateromedial lobes (Figs 219, 220). Distiphallus narrowest medially A. pudica Spencer
Antenna dark brown, but if first flagellomere paler, then entirely concolourous on outer face. Basiphallus not produced distolaterally; with 1 pair of membranous lateromedial lobes. Distiphallus narrowest apically 24
24 Apical tuft of pale hairs present on apex of male first flagellomere. Calypter hairs white to brown. Mesophallus with 1 pair of small, lateral membranous lobes at midpoint. Distiphallus relatively pale, not angled dorsally and with apical section (seen ventrally) irregular in outline (Figs 203, 204) A. parca Spencer
Apical tuft of hairs absent from male first flagellomere. Calypter hairs dark brown. Mesophallus produced laterally into 1 pair of large, triangular, basally sclerotised wings. Distiphallus relatively dark, slightly angled dorsally and with distal section (seen ventrally) evenly tapered (Figs 208, 209) A. parilis Spencer

Species descriptions

Unless otherwise stated, species are characterised in part as follows: body and setae dark brown with ocellar triangle always darker; gena highest posteriorly; notum subshiny; stridulatory organ present; one pair of well-developed acrostichal setae present; acrostichal setulae in eight to ten rows; costa extending to vein M1+2; abdomen dark brown.

Agromyza abiens Zetterstedt

Figs 139–142

Agromyza abiens Zetterstedt, 1848: 2747. Hendel 1931: 147 [as synonym of rufipes Meigen]; Spencer 1963d: 3 [stat. reinst.], 1972: 35, 1976: 88.

Agromyza echii Kaltenbach, 1860: 217. Spencer 1963d [synonymy].

Description

Wing length 3.6–4.1 mm (♂), 4.3 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.7. Eye height divided by gena height: 3.6–5.3 (up to 10.0 in German material examined). First flagellomere slightly elongate with distal margin covered with tuft of dense, pale hairs. Fronto-orbital plate and parafacial slightly produced, curving under eye as distinct cheek. Ocellar triangle hemispherical in outline. Epistoma relatively large to narrow. Thorax and abdomen with light greyish pruinosity.

Chaetotaxy : Three ori, two ors. Four to six dorsocentrals (four in Nearctic specimen), slightly decreasing in size anteriorly (more so presuturally). Mid tibia with two posteromedial setae.

Colouration : Body dark brown with halter white and gena and parafacial light brown; lunule, pedicel, scape and frons (between narrow fronto-orbital plates) brownish orange; first flagellomere orange with deeper brown tint past base, sometimes mostly brown; palpus brownish; posterior 1/3–1/2 of frons sometimes darker; ocellar triangle dark brown, distinct, slightly larger than tubercle; notopleuron slightly paler than rest of notum. Calypter margin and hairs white. Examined European material with paler parts of head lighter orange, parafacial and posterior parts of frons darker, thorax reddish, palpus reddish orange and lunule yellow. Wing veins light brown; coxae and femora paler brown with apices of latter yellow; tibiae and tarsi light brownish orange.

Genitalia : (Figs 139–142) Surstylus entirely fused to epandrium and with several tubercle-like setae on inner face (concentrated posteriorly). Hypandrium produced at apex. Postgonite broad and flat apically with inner-medial swelling. Phallophorus elongate and narrow, interlocking and partially fused to basiphallus. Left 1/2 of basiphallus short, arising apically from phallophorus; right 1/2 of basiphallus broad basally and with thin curved apical section; apices of both halves abruptly converging. Hypophallus small and membranous with lightly sclerotised medial section. Mesophallus not much longer than wide, rounded and slightly flattened basally, and with several basal, lateral and anteromedial extensions; distiphallus separate, not much longer than wide, cup-like with internal fringe of hairs and basal 1/2 more abruptly narrowed. Ejaculatory apodeme asymmetrical, with distal margin irregular and medial rib offset and annulated.

Hosts

BoraginaceaeAegonycho, Amsinckia (?), Anchusa, Asperugo, Borago, Brunnera, Buglossoides, Cerinthe, Cynoglossum, Echium, Lappula, Lindelofia, Lithospermum, Lycopsis, Myosotis, Nonea, Omphalodes, Onosma, Pentaglottis, Pulmonaria, Podonosma, Pulmonaria, Solenanthus, Symphytum (Spencer 1976; Benavent-Corai et al. 2005; Ellis 2021).

Distribution

USA: VA*. Europe.

Type material

Syntypes [abiens]: Sweden. Scania ad Tranas, Esperod, Ostra Torp. [Not examined – one ST labeled as lectotype (Spencer, 1963d)]

Syntypes [echii]: Germany [not given]. [Not examined]

Material examined

England. Hampstead, 26.vi.1954, em. 2.viii.[19]54, CNC352640 (1♀, CNC). Germany. Thuringen: H. Buhr, mine an Cynoglossum officinale, vii.1964, Hering, CNC352639, No. 2034 (1♂ 1♀ [with puparia], CNC). Europe. [illegible], 20.vii.1953, em. 11.viii.[19]53, CNC352638 (1♂, CNC), [illegible] viii.1908 (2♂ 1♀, USNM). USA. VA: Strausberg, 21.ix.1980, W.H. Rowe (1♂, USNM).

Comments

The Virginia male recorded here is the first Nearctic instance of this otherwise European species, which feeds on a wide range of Boraginaceae. The irregular, uneven basiphallus, the distinct mesophallus and the cup-like distiphallus with an inner fringe of hairs are characteristic. Externally, this species is relatively large with a white male calypter, it has five fronto-orbitals, a relatively pale head and the apical third of the first flagellomere is densely covered in pale hairs. Morphology is incredibly similar to Agromyza pseudoreptans Nowakowski, which also occurs in Europe and North America and may occur in the Delmarva states, but the distiphallus is distinctly longer than wide (not as long as wide) and is similarly constricted dorsoapically (not broadly domed, as illustrated in previous works).

Agromyza albipennis Meigen

Figs 34, 143–146

Agromyza albipennis Meigen, 1830: 171. Spencer 1969: 32; Spencer and Steyskal 1986b: 262; Černý et al. 2020: 194.

Agromyza dubitata Malloch, 1913a: 311. Frick 1953: 68 [as synonym reptans Fallén], 1957: 199 [as synonym nigripes Meigen]. Spencer and Steyskal 1986b [syn.].

Agromyza albo-hyalinata Zetterstedt, 1848: 2742. Griffiths 1963 [synonymy].

Agromyza albipennis fennica Griffiths, 1963: 128. Spencer 1976 [synonymy].

Description

Wing length 2.1–2.9 mm (♂), 2.5–3.5 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.8. Eye height divided by gena height: 3.6–5.6. First flagellomere relatively elongate with dorsoapical corner slightly pointed; velvety with small apical tuft of dense pale hairs (only distinct when viewed anteriorly). Fronto-orbital plate and parafacial slightly pronounced, curving under eye as cheek. Ocellar triangle reaching level of anterior ors, sides concave.

Chaetotaxy : Two ors, two or three ori (if three, anterior seta smaller). Four postsutural dorsocentrals, anterior two much shorter. Mid tibia with two posteromedial setae.

Colouration : Body black with halter white and wing veins paler and fore knee sometimes very narrowly yellow. Calypter margin and hairs white. Faint greenish shine sometimes evident, mostly on abdomen.

Genitalia : (Figs 143–146) Surstylus broad and triangular, entirely fused to epandrium, margin darkly pigmented, setae clustered at apex and inner margin with several medial to subapical tubercle-like setae. Inner lobe of hypandrium with furrows leading to base of setae. Postgonite small, flat, lobe-like. Halves of basiphallus converging to point of fusion at base; apices abruptly converging; right 1/2 with long membranous extension at midpoint. Distiphallus of “nigripes-type” (capsule-shaped with subbasal opening for entry of ejaculatory duct, pronounced dorsobasal collar, and medial convolution), with sides parallel medially, one pair of elongate parallel dark ventromedial swellings, and a slightly broadened apical margin with inner surface minutely spinulose; slightly angled dorsally.

Hosts

PoaceaeTriticum aestivum (only known North American host), Agrostis, Arrhenatherum, Brachypodium, Bromus, Calamagrostis, Dactylis, Deschampsia, Festuca, Glyceria, Hordeum, Milium, Phalaris, Phalaroides (Ellis 2021), Phleum, Poa, Secale, Setaria. In Europe, most commonly occurring on Phalaria arundinacea, Hordeum and Poa (Spencer and Steyskal 1986b; Benavent-Corai et al. 2005).

Distribution

Canada: AB, BC, MB, NB*, NL, NS*, NT, NU*, ON, QC, SK*, YT*. USA: AK, CA, CO, DC*, IA, IL, MA, MD*, MI, NC*, NY, OH, PA*, SC*. Widespread in Palaearctic (Černý et al 2020).

Type material

Syntype [albipennis]: Austria [not given] (1♀, NMW). [Not examined].

Syntype [dubitata]: USA. MA: Beverly (1♀, USNM; two females reported in Spencer and Steyskal 1986b).

Paratypes examined

[dubitata]: Canada. QC: Beaulieu, Ile de Montreal, 14.vii.1906 (1♀, USNM), Ottawa [illegible], Beaulieu, 20.vii.1912, “13” (1♀, USNM), Cottage Beaulieu, Beaulieu, 16.vii.1906, “15” (1♀, USNM), Cottage Beaulieu, Beaulieu, 21.viii.1906, “19” (1♀, USNM), Cottage Beaulieu, Beaulieu, 10.vi.1906, “21 0” (1♀, USNM). USA. MA: Beverly, 23.viii.1969 (1♀, USNM).

Syntypes [albo-hyalinata]: “Ovasa ad Esperod Scaniae… Dania” (? ZIL). [Not examined].

Holotype [albipennis fennica]: Finland. Messuby (1♂, UZMH). [Not examined].

Additional material examined

Canada. AB: Black Foot Hills, 9.viii.1940, A.R. Brooks, CNC352861 (1♀, CNC), Lethbridge, 27.vi.1923, H.L. Seamans, CNC352860 (1♀, CNC), Banff Natl. Pk., Lk. Louise, 1402 m, 7.vii.1955, R. Coyles, CNC352693 (1♀, CNC), Banff, Jasper Hwy, Sunwapta Pass, 2011 m, 7.vii.1955, R. Coyles, CNC352692 (1♀, CNC), Elkwater, 11.vi.1956, CNC352652 (1♂, CNC), Onefour, 49°6'0"N, 110°24'0"W, 4.vi.1955, J.R. Vockeroth, CNC352653, CNC352654 (2♂, CNC), Elk Island N.P. Wood Bison Trail wetland alongside aspen forest, lots of tall grass, 53.5666°N, 112.8514°W, 722 m, BIOBus, 1.vii.2012, BIOUG06731-D05 (1♂, CNC), Waterton Lakes NP, Highway 6 pulloff east of 2 Flags Lookout montane forest, douglas fir and lodgepole pine (mostly conifer with aspen/birch understory), 49.065°N, 113.7781°W, 1562 m, BIOBus, 11.viii.2012, BIOUG06502-C07 (1♀, CNC), BC: Liard Hot Spg. Mi496 Alaska Hwy, 457 m, 9–10.vii.1959, E.E. MacDougall, CNC352698 (1♀, CNC), Moosehorn Lake, 58°10'0"N, 132°7'0"W, 1371 m, 25.vii.1960, R. Pilfrey, CNC352658 (1♂, CNC), Summit L., Mi392 Alaska Hwy, 1402 m, 16.vii.1959, E.E. MacDougall, CNC352699 (1♀, CNC), Terrace, 10.vi.1960, 5.vi.1960, C.H. Mann, CNC352655, CNC352656, CNC352657, CNC352697 (3♂ 1♀, CNC), Vernon, 3.ix.1931, R.D. Bird, CNC352651, 2s37, IV 31 (1♂, CNC), Bobson, 6.v.1952, H.R. Foxlee, CNC352862 (1♀, CNC), Oliver, 20.v.1923, C.B. Garrett, CNC352863 (1♀, CNC), Vernon, 31.vii.1937, H. Leech, CNC352864 (1♀, CNC), Victoria, 20.v.1919, W. Downes, CNC352844 (1♂, CNC), MB: Minnedosa, 5mi N, 8.vii.1958, R.L. Hurley, CNC352690 (1♀, CNC), Shilo, 5mi SW, 28.v.1958, R.B. Madge, CNC352663 (1♂, CNC), Western MB, Riding Mountain Nat. Pk. – North Escarpment Trailhead wet meadow/marsh, 50.674°N, 99.65°W, 726 m, J. Straka, J. Crossey, 15.viii.2008, 08BDIP-1973 (1♀, CNC), Aweme, 26.vi.1916, N. Criddle, CNC352842 (2♂, CNC), 28.viii.1917, CNC352843 (1♂, CNC), NB: Kouchibouguac N.P., 8.vii.1977, J.F. McAlpine, Code-6020N, CNC352688, CNC352689 (2♀, CNC), NL: Gros Morne; Gros Morne Mountain Hiking Trail, 49.5657°N, 57.8324°W, 39 m, J. Crossey, R. Labbee, A. Smith, M. Zhang, 15.vii.2009, 09BBEDI-0938 (1♀, CNC), St. John’s, Agric. Exp. Sta., 12.vii.1967, 15.vii.1967, J.F. McAlpine, CNC353033, CNC353041, CNC353042 (2♂ 1♀, CNC), NS: Kentville, 22.viii.1912, from tortricid larva on apple, [illegible], CNC352650 (1♂, CNC), NT: Ft. McPherson, 19.vii.1957, R. Hurley, CNC352695 (1♀, CNC), NU: [N.W.T.] Muskox L., 64°45'0"N, 108°10'0"W, 2.viii.1953, J.G. Chillcott, CNC352696 (1♀, CNC), ON: Almonte, 18.v.1951, J.F. McAlpine, CNC352679 (1♀, CNC), Hailerbury, 12.viii.1947, G.S. Walley, CNC352646 (1♂, CNC), Marmora, 1.vi.1952, 13.v.1952, J.C. Mitchell, J.R. Vockeroth, CNC352674, CNC352675, CNC352676 (3♀, CNC), Maynooth, 25.v.1951, J.F. McAlpine, CNC352677 (1♀, CNC), Minnedosa, 5mi N, 8.vii.1958, R.L. Hurley, CNC352691 (1♀, CNC), Normandale, 42°42'0"N, 80°19'0"W, 27.v.1956, J.R. Vockeroth, CNC352645 (1♂, CNC), 29.v.1956, J.R. Lonsway, CNC352678 (1♀, CNC), Ottawa, 8.vii.1952, G.E. Shewell, CNC352647 (1♂, CNC), Port Hope, 25.v.1925, N.K. Bigelow, CNC352680 (1♀, CNC), Port Severn, 3mi N, 18.v.1959, J.G. Chillcott, CNC352681 (1♀, CNC), Putnam, 26.vi.1925, G.S. Walley, CNC352644 (1♂, CNC), St. Williams, 42°40'0"N, 80°25'0"W, 23.v.1956, J.R. Vockeroth, CNC352649 (1♂, CNC), Turkey Pt., 42°39'0"N, 80°21'0"W, 25.v.1956, J.R. Vockeroth, CNC352683 (1♀, CNC), Brockville, 5.viii.1903, W. Metcalfe, CNC352847 (1♀, CNC), Cottage Beaulieu, 16.vii.1906, 145, CNC352854 (1♀, CNC), Mer Bleue, 10.v.1938, G.E. Shewell, CNC352834 (1♂, CNC), Norway Point, Lake of Bays, 31.vii.1919, J. McDunnough, CNC352851 (1♀, CNC), Orillia, 18.vii.1923, C.H. Curran, CNC352848 (1♀, CNC), 17.viii.1923, CNC352850 (1♀, CNC), Ottawa, 15.vii.1938, A. Brooks, CNC352830 (1♂, CNC), Ottawa, 11.ix.1947, G.E. Shewell, CNC352829 (1♂, CNC), Ottawa, A. Brooks, 5.vii.1938, CNC352852 (1♂, CNC), Ottawa, 27.vii.1912, Beaulieu, CNC352853 (2♂ 2♀, CNC), Pt. Ryerse, 1.vi.1939, G.E. Shewell, CNC352835 (1♂, CNC), St. Johns, Little Montreal River, 9.vii.1937, G.E. Shewell, CNC352855 (1♀, CNC), Simcoe, 8.vi.1939, G.E. Shewell, CNC352849 (1♀, CNC), Simcoe, G.E. Shewell, 14.vi.1939, CNC352831 (1♂, CNC), Simcoe, G.E. Shewell, 22.vi.1939, CNC352832 (1♂, CNC), Simcoe, G.E. Shewell, 29.v.1939, CNC352833 (3♂ 1♀, CNC), Metcalfe, 2mi N, 10.vi.1982, B.E. Cooper, CNC353037 (1♂, CNC), Metcalfe, 25.v.1983, B.E. Cooper, CNC353038 (1♂, CNC), Ottawa, damp second-growth Acer-Betula wood, 15.viii.2003, 20.vii.1974, 29.vi.1991, J.R. Vockeroth, CNC353032, CNC353035, CNC353036 (2♂ 1♀, CNC), Port Severn, 3mi N, black spruce bog, 18.v.1959, J.G. Chillcott, CNC353039, CNC353034 (2♂, CNC), QC: Ile de Montreal, 7.vii.1906, Beaulieu, CNC352837 (1♂, CNC), Old Chelsea, 13.vi.1961, J.R. Vockeroth, CNC353040 (1♂, CNC), St. Johns, Little Montreal River, 9.vii.1937, G.E. Shewell, CNC352836 (1♂, CNC), Wakefield, 9.vii.1946, G.E. Shewell, CNC352838 (1♂, CNC), Woburn, 19.vi.1923, C.H. Curran, CNC352859 (1♀, CNC), Aylmer, 16.vii.1959, C.H. Mann, CNC352684 (1♀, CNC), Farnham, 5.vi.1963, J.R. Vockeroth, CNC352642, CNC352682 (1♂ 1♀, CNC), Hull, 20.vi.1956, J.R. Vockeroth, CNC352685 (1♀, CNC), Indian House L., 20.vii.1954, 27.vi.1954, 27.vii.1954, R. Coyle, R. Coyles, W.R. Richards, CNC352648, CNC352686, CNC352687 (1♂ 2♀, CNC), Knowlton, Bolton Pass, 243 m, 5.vi.1963, J.R. Vockeroth, CNC352641 (1♂, CNC), Mt. Orford, 365 m, 5.vi.1963, J.R. Vockeroth, CNC352643 (1♂, CNC), Abbotsford, G. Shewell, 29.v.1936, CNC352839, CNC352858 (1♂ 1♀, CNC), Abbotsford, 10.viii.1937, G.E. Shewell, CNC352840 (1♂, CNC), Abbotsford, G. Shewell, 20.viii.1936, CNC352841 (1♂, CNC), 30.v.1936, CNC352857 (1♀, CNC), Abbotsford, 20.v.1931, J.B. Maltais, CNC352856 (3♂ 3♀, CNC), SK: Christopher L., 27.viii.1948, A.R. Brooks, CNC352705, CNC352706 (2♀, CNC), Val Marie, 49°15'0"N, 107°44'0"W, 5.vi.1955, 9.vi.1955, J.R. Vockeroth, CNC352659, CNC352662, CNC352664, CNC352665, CNC352666, CNC352700, CNC352701, CNC352702, CNC352703, CNC352704, CNC352707 (5♂ 6♀, CNC), Grasslands National Park, Visitor’s Center grass cattle pasture, 49.246°N, 107.732°W, 812 m, J. Cossey, N. Jeffery, J. Straka, 14.vii.2008, 08BBDIP-2742 (1♂, CNC), Saskatoon, 12.vii.1940, 22.vii.1939, 25.v.1926, 25.viii.1923, 28.ix.1925, sf. wheat, A.P. Robinson, K.M. King, King, 13288; D679; ‘41, 1328B; D676; ‘41, 1328B; D677, 1328B; D678, 16410- 3N9B, 16423 1229B; D348, 16425 26BS5; D350, 63AN, D663, CNC352845, CNC352846, CNC352865, CNC352866, CNC352867, CNC352868, CNC352869, CNC352870, CNC352871 (2♂ 7♀, CNC), YT: Otter Lake, 1219 m, 15.vii.1960, J.E. H. Martin, CNC352694 (1♀, CNC). USA. AK: Anchorage, 18.vi.1951, 3.vii.1951, IDEMA Illustration, R.S. Bigelow, CNC352668, CNC352720 (1♂ 1♀, CNC), Fairbanks, 13.vi.1952, J.B. Bartley, CNC352669 (1♂, CNC), Mackenzie Delta, Reindeer Depot, 30.vi.1948, J.R. Vockeroth, CNC352661 (1♂, CNC), Umiat, 13.vii.1959, 24.vii.1959, 5.vii.1959, 6.vii.1959, 7.vii.1959, 7.viii.1959, 9.viii.1959, IDEMA Illustration, J.E.H. Martin, R. Madge, CNC352660, CNC352667, CNC352673, CNC352712, CNC352713, CNC352714, CNC352715, CNC352716, CNC352717, CNC352718, CNC352719 (3♂ 8♀, CNC), CO: Walden, 11.viii.1965, G.F. Knowlton (1♂, USNM), Doolittle Ranch, 9800', Mt. Evans, 9.vii.1961, C.H. Mann (1♂, USNM), Doolittle Ranch, Mt. Evans, 2987 m, 3.viii.1961, 9.vii.1961, C.H. Mann, CNC352670, CNC352708 (1♂ 1♀, CNC), Boulder, 5500', 5.vi.1961, B.H. Poole (1♀, USNM), Mt. Evans, 3535 m, 11.vii.1961, C.H. Mann, CNC352711 (1♀, CNC), Mt. Evans, 3566 m, 22.vii.1961, B.H. Poole, CNC352710 (1♀, CNC), Nederland, 2529 m, 5.vii.1961, J.G. Chillcott, CNC352672 (1♂, CNC), Jackson Co., Rabbit Ears Pass, 7.vii.1961, J.G. Chillcott, CNC352671, CNC352709 (1♂ 1♀, CNC), DC: Theo Roosevlt Id, 4.vi.1977, W.N. Mathis (1♂, USNM), MA: Boston, June (1♂ 1♀, USNM), Concord, 19.vii.1961, marsh, W.W. Wirth (2♂ 3♀, USNM), Forest Hills, 21.ix.1913, A.L. Melander (2♀, USNM), MD: Colesville, 4.vii.1976, W.W. Wirth (2♀, USNM), Colesville, 14.vi.1975, Malaise trap, W.W. Wirth (1♀, USNM), NC: Macon Co., Highlands, Lake Ravenel, 7.vi.1986, Malaise trap, W.W. Wirth (1♂, USNM), NY: Ithaca, 15.viii.1926, A.L. Melander (1♂, USNM), Geneva, 28.v.1914, A.L. Melander (1♂, USNM), Long Island, Cold Spring Harbor, A.L. Melander (1♂ 2♀, USNM), Allegany State park, 28.v-3.vi.1963, mossy woods, W.W. Wirth (1♀, USNM), PA: Mineral Spr., 5.ix.1927, A.L. Melander (1♂, USNM), Chester Co., Avondale, Stroud Res. Ctr., 28.ix.2006, K. Styer (1♀, UDCC), SC: Black Falls, 7.viii.1953, A.L. Melander (1♀, USNM).

Comments

See comments for Agromyza aprilina.

Agromyza ambrosivora Spencer

Figs 147–151

Agromyza ambrosivora Spencer, 1969: 35. Spencer and Steyskal 1986b: 64; Scheffer et al. 2007: 770; Scheffer and Lonsdale 2018: 85; Eiseman and Lonsdale 2018: 7.

Description

Wing length 2.3–2.7 mm (♂), 2.7–3.9 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.5–0.7. Eye height divided by gena height: 2.9–3.6. First flagellomere slightly longer than wide with apex broadly rounded; without tuft of pale hairs. Fronto-orbital plate and parafacial slightly projecting. Ocellar triangle broadly rounded and short, not extending much past ocelli. Buccal cavity subquadrate with anteromedial margin emarginate.

Chaetotaxy : Four or five thinner ori with anterior seta sometimes shorter; one ors. Four postsutural dorsocentrals, decreasing in length anteriorly. Mid tibia with two posteromedial setae. Female sometimes with strong additional seta between acrostichal seta and posterior dorsocentral.

Colouration : Body predominantly brown, halter white. Gena (excluding ventral margin), parafacial and frons (excluding posterolateral corner, posterior margin and ocellar triangle) light brownish orange. Scape and pedicel yellow; first flagellomere orange with distal 1/2–2/3 brownish. Lunule orange with yellowish pilosity. Face brown. Palpus and clypeus dark brown. Tarsi yellow (slightly darker on posterior legs), fore tibia light brown and knees narrowly yellow. Calypter margin and hairs white. Female with frons darker (sometimes entirely brown or with anterior margin lighter), gena darker and fore tibia darker medially.

Genitalia : (Figs 147–151) Surstylus triangular, entirely fused to epandrium; with apical setae and medial tubercle-like setae. Hypandrium long and thin with narrow apical point. Postgonite lobate, directed apically. Phallophorus produced apically and fused to base of basiphallus. Halves of basiphallus widely bowed with bases broadly fused; elongate and narrow with margins irregular. Distiphallus very long, thin, dark, bifid, and strongly curved into semicircle at split; base of distiphallus with transverse dorsal band and paired ventral sclerites. Hypophallus large, membranous, and directed apically. Ejaculatory apodeme dark and very reduced in size.

Variation : Female from CA differs as follows: wing length 4.1 mm; anterior (of four) ori small; only three dorsocentrals; eye height divided by gena height 2.5; tibiae yellow to base and apex; gena mostly pale.

Hosts

AsteraceaeArtemisia (?), Ambrosia artemisiifolia, A. trifida and probably A. douglasiana (Spencer and Steyskal 1986b; Benavent-Corai et al. 2005); Helianthus annuus (Eiseman and Lonsdale 2018).

Distribution

Canada: ON. USA: CA, CO, MA, MD, NY, PA, VA*.

Type material

Holotype: Canada. ON: Pelee, em. 31.vii from leaf-mines on Ambrosia artemisifolia, leg. 8.vii.1967, CNC352722 (1♂ [with puparium], CNC).

Paratypes: Canada. ON: Pelee, K.A. Spencer, mine Ambrosia artemisifolia, 15.vii.1967, em. 1–10.viii.1967, CNC352723-CNC352744 (10♂ [5 with puparia], 12♀ [8 with puparia], CNC).

Additional material examined

Canada. ON: Belleville, P. Harris, vii.1969, em. 22.ix.1969, CNC352721 (1♀ [with puparium], CNC), Ottawa, damp second-growth Acer-Betula woods, 8.viii.1993, J.R. Vockeroth, CNC353060 (1♂, CNC). USA. CA: Carbon Canyon, 14.vi.1977, on Artemesia d. (1♀, USNM), CO: Chaffee Co., Poncha Springs, South Arkansas River, 8.vii.2015, C.S. Eiseman, Helianthus, em. 28.vii-1.viii.2015, #CSE1872, CNC654328–654332 (1♂ 4♀, CNC), MA: Worcester Co., Sturbridge, Leadmine Rd., 6.vii.2013, em. 20.vii.2013, C.S. Eiseman, ex Ambrosia artemisiifolia, #CSE725, CNC392679, CNC392680 (2♀, CNC), MD: Plummers Isl., 30.v.1913, R.C. Shannon (1♀, USNM), College Park, 7.vii.1935, C.T. Greene (1♂, USNM), Pr. William Co., Woodbridge, 26.vii.1968, J.W. Adams (1♂, USNM), Montgomery Co., Colesville, 26.vi.1977, Malaise trap, W.W. Wirth (2♂, USNM), 30.vi.1977 (1♂, USNM), Colesville, W.W. Wirth, 4.vii.1976 (2♀, USNM), 11.vii.1974 (2♂, USNM), 24.vii.1974 (1♂, USNM), 28.vii.1976 (2♂, USNM), 1.viii.1976 (2♂, USNM), 4mi SW of Ashton, 1.ix.1981, Malaise trap, G.F. and J.F. Hevel (1♀, USNM), Bethseda, G. Steyskal, 6.vii.1970 (2♂, USNM), 7.vii.1970 (1♂, USNM), Allegeny Co., Little Orleans, Little Orleans campground, 4.vi.1999, sweeping, C.R. Bartlett (1♂, UDCC), nr. West Mifflin, Coal Valley Rd. #2, 5.vii.1997, sweeping, C.R. Bartlett (1♀, UDCC), PA: Chester Co., Pottstown, Warwick County Park, 10.viii.2014, ex. Ambrosia trifida, em. 2.ix.2014, C.S. Eiseman, #CSE1371, CNC384846 (1♀, CNC), VA: Great Falls, 21.vi.1931, A.L. Melander (1♂, USNM), Alexandria, “viii-5”, J.M. Aldrich (1♀, USNM), Fairfax Co., Turkey Run Park, nr. mouth of Turkey Run, 38°57.9'N, 7°09.4'W, Malaise trap, 18–30.v.2007, D.R. Smith (1♀, USNM), Turkey Run Park, 0.3 km W mouth Turkey Run, 38°58'N, 77°09.6'W, Malaise trap, D.R. Smith, river 14–17.v.2006 (1♂, USNM), river trap, 17–24.v.2006 (2♀, USNM), Great Falls Park, swamp trail, 38°59.4'N, 77°15.2'W, Malaise trap, trap #2, 18.iv.2.v.2007, D.R. Smith (1♀, USNM).

Comments

Agromyza ambrosivora is an easily recognised species with one pronounced ors and four or five thinner ori. The phallus forms a long, dark bifid tubule, which is very similar to that found in A. virginiensis (Fig. 151): these two species can be separated externally using the characters mentioned in the key. A third related species, A. rudbeckiana Scheffer and Lonsdale, a leaf-miner of Heliopsis and Rudbeckia (Asteraceae), has a more evenly arched distiphallus that is only split near the apex, and unlike A. ambrosivora, there are two ors and three ori (not one and four) (Scheffer and Lonsdale 2018).

Agromyza apfelbecki Strobl

Figs 152–155

Agromyza abiens var. Apfelbecki Strobl, 1902: 504.

Agromyza andalusiaca Strobl, 1906: 380. Hendel 1920 [synonymy].

Agromyza Apfelbecki. Hendel 1920: 117. Hendel 1931: 107.

Agromyza apfelbecki. Spencer 1966c: 172; Papp 1984: 264; Spencer 1990: 253.

Description

Wing length 4.0–4.1 mm (♂), 4.4–4.8 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.4–0.7. Eye height divided by gena height: 2.1–2.4. Ocellar triangle small and rounded. First flagellomere small and rounded; without apical tuft of pale hairs. Fronto-orbital plate slightly projecting and parafacial strongly projecting, partially to strongly continuing under eye as cheek. Wing slightly pointed apically with apex between M1 and R4+5. Thorax with light to dense grey pruinosity.

Chaetotaxy : Three to seven ori (two in Spanish female); two ors. Six or seven dorsocentrals, two or three presutural, decreasing in size anteriorly. Mid tibia without posteromedial setae.

Colouration : Body mostly dark brown. Head yellow with back of head, occiput, clypeus, mentum and ventral margin of gena dark brown, distal 2/3 of first flagellomere brownish, venter of face light brown and frons slightly darker anteriorly. Knees yellowish orange; tibiae and tarsi paler than femora with tibiae slightly darker medially. Scutum with nearly imperceptible metallic green shine in VA female. Calypter margin and hairs white to dark brown. Halter white, rarely with small lateral spot and ventral margin of knob brown (VA female). Abdomen dark brown with indistinct (VA) or strong coating of grey pruinosity, except on brownish orange male terminalia (sometimes with dark dorsomedial spot on epandrium). European specimens often with fronto-orbital plate, face, apex of palpus, parafacial and ring under eye mostly dark, tibiae often yellowish or more orange on distal 1/3, and epistoma usually larger.

Genitalia : (Figs 152–155) Epandrium relatively broad and shallow, with suture between it and surstylus barely evident. Surstylus slightly curved anteriorly and with numerous small, pointed, tubercle-like setae along inner-basal/medial surface. Phallophorus tapered distally and partially fused to flat, pale sclerites of basiphallus, which are weakest and irregular along dorsal margin, with right sclerite partially separated from darker, curved apical section. Hypophallus sac-like and membranous with partial sclerotisation along anteromedial surface. Distiphallus nearly cylindrical with irregularly sclerotised margins not entirely meeting ventrally; twisted past midpoint. Ejaculatory apodeme minute.

Hosts

AsteraceaeCarduus, Cirsium, Cynara (Spencer 1990).

Distribution

USA: VA*. France, Italy, Malta, Germany, Spain, Croatia (Papp and Černý 2015). Turkey. Chile (Valparaiso, La Cruz; introduced) and Argentina (Valladares 1998; Çöl et al. 2006).

Type material

Syntypes [Apfelbecki]: Yugoslavia. Zadar (♂♀, coll. Strobl). [Not examined]

Syntypes [andalusiaca]: Spain. “Southern Spain” [= Algreciras?] (?, coll. Strobl). [Not examined]

Material examined

France. Perpignan, 10.x.1959, mine Cynara carduuculus, em. 24.v.1959, K.A. Spencer, CNC352745 (1♂ [with puparium], CNC). Germany[?]. “Agrom. Apfelbecki Str., det. Hendel” (1♀, USNM). Spain. Barcelona, “prat”, 27.iii.1960, K.A. Spencer (1♀, USNM), CNC352746 (1♀, CNC), En Alcachofe, La Cruz, Valpo, Nac. 5.ix.1959, N. Hichins (5♂ 2♀ 1?, USNM). USA. VA: Fairfax Co., Turkey Run Park, 0.3 km W mouth Turkey Run, 38°58'N, 77°09.6'W, Malaise trap, 29.iii-25.iv.2007, D.R. Smith (1♀, USNM).

Comments

While difficult to differentiate from a number of Palaearctic taxa, Agromyza apfelbecki is quite distinct from Nearctic Agromyza because it is exceptionally large (wing length at least 4.0 mm), there are numerous dorsocentrals (at least six), the epistoma is large and pronounced, the head is predominantly pale (including the palpus and face) and the halter is sometimes maculated. Agromyza apfelbecki is one of only three Agromyza known to feed on Asteraceae, and is distributed primarily around the Mediterranean where it often occurs on globe artichoke (Spencer, 1990).

The above Virginia record is the first known occurrence of this species in the Nearctic, where it was likely introduced accidentally on its host plant. As discussed by Dempewolf (2004), Agromyza apfelbecki can be a major pest on artichoke if populations become large enough, particularly to younger plants; the damage caused by larval feeding can vary significantly between years (Spencer 1973), possibly due to climatic conditions, as frost can severely increase mortality among larvae (Ricchello 1928).

Agromyza aprilina Malloch

Figs 156–160

Agromyza aprilina Malloch, 1915c: 359. Frison 1927: 192 [lectotype design.]; Frick 1957: 199 [as synonym subnigripes Malloch], 1959: 358; Spencer 1969: 36; Spencer and Steyskal 1986b: 262 [stat. reinst.].

Description

Wing length 2.2–2.5 mm (♂), 2.5–3.2 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.8. Eye height divided by gena height: 2.2–3.4. First flagellomere slightly longer than high with rounded dorsoapical point (pronounced to relatively indistinct), without apical tuft of pale hairs. Fronto-orbital plate and parafacial projecting (more so anteriorly), continuing under eye as cheek. Ocellar triangle longer than wide and subshiny. Costa extending slightly past R4+5. Smaller females with vein dm-m incomplete.

Chaetotaxy : Two ori; three ors (or vice versa), sometimes appearing as four ori and one ors. One presutural dorsocentral and three or four postsutural dorsocentrals, decreasing in length anteriorly. Mid tibia with two posteromedial setae.

Colouration : Body dark brown with halter white and gena and frons (aside from fronto-orbital plate and ocellar triangle) reddish, fore knee yellow and tarsi dirty yellow. Calypter margin and hairs white. Female from “Turkey Run, headquarters” with light green metallic shine on thorax, abdomen and pleuron (faintest).

Genitalia : (Figs 156–160) Anterodistal margin of epandrium deeply incised before surstylus. Surstylus triangular, fused to epandrium with suture evident; inner-basal margin with several small tubercle-like setae. Hypandrium slightly produced at apex, inner margin of lobe thickened with several setulae. Postgonite small and lobe-like with thin plate-like base. Phallus relatively small, narrow, and weakly sclerotised. Phallophorus tapering apically, fused to halves of basiphallus, which converge to base. Halves of basiphallus with internal fold apically, with distal portion sharply directed inwards; lateral membrane broad, twisted past midpoint. Distiphallus of "nigripes-type” (capsule-shaped with subbasal opening for entry of ejaculatory duct, pronounced dorsobasal collar, and medial convolution); nearly carinate ventromedially, distal section with elongate medial split.

Host

Unknown.

Distribution

Canada: AB, BC, MB, NB*, ON, QC, SK, YT*. USA: IL, MD*, NC*, NH, VA*.

Type material

Lectotype: USA. IL: Cottonwood Grove, Urbana, 16–20.iv.1915 (1♀, INHS). [Not examined]

Paralectotype examined

USA. IL: Urbana, 20.iv.1915, Cottonwood, CNC352757, Type No. 2725 (1♀, CNC).

Additional material examined

Canada. AB: Banff, C.B.D. Garrett, 8.vi.1922, CNC352760 (1♀, CNC), MB: Ninette, J.F. McAlpine, 21.v.1958, CNC352778 (1♀, CNC), 9.v.1958, CNC352755 (1♂, CNC), R.B. Madge, 9.v.1958, CNC352776 (1♀, CNC), Shilo, 5mi SW, J.F. McAlpine, 28.v.1958, CNC352777 (1♀, CNC), NB: Kouchibouguac N.P., Hanley and Cooper, 23.v.1977, Code-5113Q, CNC352754, CNC352774, CNC352775 (1♂,2♀, CNC), W.P. Hanley, 20.v.1977, Code-5098B, CNC352753 (1♂, CNC), ON: Bell’s Cor[ner]., J.F. McAlpine, 8.v.1951, CNC352766 (1♀, CNC), Marmora, J.F. McAlpine, 23.iv.1952, CNC352751, CNC352768, CNC352769 (1♂ 2♀, CNC), 24.iv.1952, CNC352752, CNC352767 (1♂,1♀, CNC), 25.iv.1952, CNC352772 (1♀, CNC), 28.iv.1952, CNC352770 (1♀, CNC), 29.iv.1952, CNC352773 (1♀, CNC), 9.v.1952, CNC352771 (1♀, CNC), Maynooth, J.F. McAlpine, 24.v.1951, CNC352764 (1♀, CNC), Metcalfe, B.E. Cooper, 14.v.1983, CNC353020, CNC353022 (2♂, CNC), 25.v.1983, CNC353021, CNC353024 (2♂, CNC), 27.iv.1983, CNC353019, CNC353025, CNC353026 (3♂, CNC), 3.v.1983, CNC353027 (1♂, CNC), 30.iv.1983, CNC353018, CNC353023 (2♂, CNC), 7.v.1994, CNC353028–353031 (4♂, CNC), Ottawa, 28.iv.1955, J.R. Vockeroth, CNC352765 (1♀, CNC), Woodroffe, J.G. Chillcott, 30.iv.1952, CNC352763 (1♀, CNC), QC: Abbotsford, G.E. Shewell, 27.iv.1936, CNC352747, CNC352748, CNC352750, CNC352758 (3♂,1♀, CNC), 27.iv.1966, CNC352749 (1♂, CNC), Shewell, 15.v.1936, CNC352759 (1♀, CNC), SK: Saskatoon, A.R. Brooks, 28.iv.1949, CNC352756 (1♂, CNC), 9.v.1949, CNC352762 (1♀, CNC), YT: Dawson, 14 mi E, P.J. Skitsko, 6.viii.1962, 396 m, CNC352761 (1♀, CNC). USA. MD: Montgomery Co., Plummers Island, 38°58'N, 77°10'W, Malaise trap, lower trap, 30.iii-22.iv.2006, D.R. Smith and J.W. Brown (1♀, USNM), Cabin John Bridge, 23.iv.1914, R.C. Shannon (1♂, USNM), Plummers Isl., R.C. Shannon, 8.iv.1914 (4♂ 3♀, USNM; 2♂ 2♀, CNC), 12.iv.1914 (1♂, USNM), 5.iv.1914 (1♂, USNM), 3.iv.1914 (1♂ 1♀, USNM), Plummers Island, Rock Run, 25.iii.1914, R.C. Shannon (1♂, USNM), nr. Plummers Isl., 28.iii.1915, R.C. Shannon (2♂, USNM), NC: Smokies, Andrews Bald, 9.vii.1914, A.L. Melander (1♂, USNM), VA: Fairfax Co., Turkey Run Park, nr. mouth of Turkey Run, 38°57.9'N, 7°09.4'W, Malaise trap, 29.iii-25.iv.2007, D.R. Smith (4♂ 1♀, USNM), Turkey Run Park, nr. headquarters bldg. 38°57.7'N, 77°08.9'W, Malaise trap, 29.iii-17.iv.2007, D.R. Smith (1♀, USNM), Great Falls Park, swamp trail, 38°59.4'N, 77°15.2'W, Malaise trap, trap #1, 18.iv-2.v.2007, D.R. Smith (1♀, USNM).

Comments

The surstylus and epandrium of Agromyza aprilina are characteristic, readily differentiating it from similar species. In the Delmarva states it can be most easily confused for the relatively common A. albipennis, but the ventral margin of the first flagellomere is flatter in this latter species, there are four fronto-orbitals (not five), the parafacial is not as pronounced, the setae are darker and the costa extends to vein M1+2. The shape of the basiphallus and distiphallus also differ (see Figs 145, 146). While the first flagellomere can appear less pointed in some material, particularly poorly preserved specimens, it is never entirely rounded as noted in Spencer (1969).

Agromyza aprilina can also be potentially mistaken for Agromyza nigripes Meigen, but the latter has darker tarsi, the costa ends at vein M1+2, and the male genitalia (based on the dissection of Welsh male in the USNM) are much stouter with slight differences in the shape of the basiphallus and distiphallus. The species A. ambigua Fallén (North America, Europe), A. kincaidi Malloch (North America; Figs 196, 197) and A. conjuncta Spencer (widespread in Europe) can also be mistaken for this species, but the characteristic phallus, surstylus and epandrium of A. aprilina allows for definitive identification. Other Nearctic Agromyza with a pointed first flagellomere and a shortened costa are restricted to the west coast and have a larger, darker distiphallus (Spencer and Steyskal 1986b).

Agromyza aristata Malloch

Figs 37, 161–166

Agromyza aristata Malloch, 1915b: 13. Frick 1957: 199; Spencer 1969: 38; Spencer and Steyskal 1986b: 59; Eiseman and Lonsdale 2018: 8; Eiseman et al. 2021: 5.

Phytagromyza aristata. Frick 1952a: 416.

Agromyza ulmi Frost, 1924: 54. Frick 1952a: 375. Frick 1959 [synonymy].

Agromyza marmorensis Spencer, 1969: 48. Syn. nov.

Description

(Fig. 37). Wing length 2.0–2.1 mm (♂), 2.3–2.4 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.9. Eye height divided by gena height: 1.9–2.7. First flagellomere subcircular, without apical tuft of pale hairs. Fronto-orbital plate (more so anteriorly) and parafacial strongly projecting and distinctly continuing under eye as cheek. Ocellar triangle relatively small with corners rounded. Palpus relatively ovate.

Chaetotaxy : Two ori; two ors. Ocellar seta thinner than postocellar, and sometimes shorter, but never reaching 1/2 length. Three postsutural dorsocentrals, decreasing in length anteriorly; possibly two more anteriorly that are not much longer than surrounding setulae. Acrostichal seta not much longer than surrounding setulae. Four scattered rows of acrostichal setulae. Mid tibia without posteromedial setae.

Colouration : Head predominantly light yellow; first flagellomere yellow; back of head brown above foramen and with grey pruinosity dorsally; ocellar triangle dark brown; posterolateral corner and posterior margin of frons brownish, with spot extending laterally to just behind level of anterior ors; clypeus light brown; face white. Thorax dark brown, covered with grey pruinosity, halter white. Calypter margin and hairs white to yellow. Legs yellow with mid and hind coxae brown, at least in part. Abdomen dark brown; epandrium yellow or with orange to brown tint with dark brown dorsal spot; if epandrium yellow, sternite 8 sometimes also yellow with medial spot.

Genitalia : (Figs 161–166) Surstylus fused with epandrium with suture obliterated, shape triangular (nearly equilateral), directed inwards, with several setae along posterior margin. Cercus relatively broad and flat with inner surface covered with pointed tubercle-like setae. Hypandrium setulose, slightly sinuate, and inner lobe with medial desclerotisation. Postgonite short. Phallophorus fused to single sclerite of basiphallus, which originates on left side and is twisted dextrally. Hypophallus membranous with faint left lateral sclerotisation. Mesophallus long, narrow, dark, and flat, lying along ventral margin of membrane; distiphallus short, flat and wrapped ventrally (nearly cylindrical), shortest dorsally. Ejaculatory apodeme minute and finger-like.

Hosts

CannabaceaeCeltis laevigata, C. occidentalis, C. pallida (leaf mine only); UlmaceaeUlmus alata (leaf mine only), U. americana, U. rubra (Eiseman et al. 2021).

Distribution

Canada: AB, NB*, ON, QC*. USA: IL, IN, IA, KS*, MI*, NY, OH, PA, VA, VT; known from leaf mines in AL, AR, CO, CT, FL, GA, MA, MD, MN, ND, NJ, TN, TX, WI.

Type material

Holotype [aristata]: USA. IL: Havana, Gleason’s Sand Dune, 30.iv.1914, C.A. Hart and J.R. Malloch (HT ♀, INHS). [Not examined]

Paratypes examined

[aristata]: USA. IL: St. Joseph, Salt Fork, 3.v.1914, Type No. 2726, CNC352790 (1♀, CNC), PA: Arendtsville, 27.iv.1923, S.W. Frost (3♂, USNM).

Syntype [ulmi]: USA. PA: Arendtsville, 22.iv.1923, S.W. Frost (1♂, USNM; type No. 50033).

Holotype [marmorensis]: Canada. ON: Marmora, 7.v.1952, J.R. Vockeroth (HT ♂, CNC).

Additional material examined

Canada. AB: Onefour, on wild cherry blossom, 2.vi.1956, O. Peck, CNC353069, CNC353072, CNC352809 (2♂,1♀, CNC), NB: Curventon, 19.v.1959, W.W. Moss, C-2, CNC352810 (1♀, CNC), ON: Almonte, 18.v.1951, J.F. McAlpine, CNC352788, CNC352803–352806 (1♂,4♀, CNC), Marmora, 16.v.1952, J.C. Mitchell, CNC352786 (1♂, CNC), 30.iv.1952, J.F. McAlpine, CNC352802 (1♀, CNC), Marmora, 28.iv.1952, J.F. McAlpine, CNC352787 (1♂, CNC), Metcalfe, 14.v.1994, B.E. Cooper, CNC353066 (1♂, CNC), 17.v.1994, CNC353067 (1♂, CNC), 7.v.1994, CNC353068 (1♂, CNC), Midland, swampy wood, 26.v.1959, J.G. Chillcott, CNC353071 (1♂, CNC), Nippising, 27.ii.1959, #83, S58-1-1381-01, CNC352779 (1♂, CNC), Ottawa nr. Uplands Airport, 22.v.1990, J.M. Cumming, CNC353064 (1♂, CNC), Ottawa, damp second-growth Acer-Betula wood, 1.vi.1995, J.R. Vockeroth, CNC353063 (1♂, CNC), 28.v.1995, CNC353070 (1♂, CNC), at sap on Acer stump, 10.v.1991, J.R. Vockeroth, CNC353062 (1♂, CNC), bleeding elm, 22.v.1952, J.F. McAlpine, CNC352791 (1♀, CNC), 28.v.1951, J.F. McAlpine, CNC352794 (1♀, CNC), 5.v.1952, B. Hartley, CNC352783 (1♂, CNC), J.G. Chillcott, CNC352783–352795, CNC352780, CNC352792 (2♂,3♀, CNC), 8.v.1952, J. Chillcott, CNC352782, CNC352785, CNC352796–352800 (2♂,5♀, CNC), Pt. Pelee, 3.vi.1929, G.S. Walley, CNC352801 (1♀, CNC), QC: Hemmingford, 10.v.1931, J.B. Maltais, CNC352807 (1♀, CNC), Old Chelsea, 8.v.1938, G.E. Shewell, CNC352808 (1♀, CNC). USA. IL: White Heath, 30.iv.1916 (1♀, USNM), Champaign, 28.iv.1953, J.F. McAlpine, CNC352789 (1♂, CNC), IN: Lafayette, J.M. Aldrich, 4.v.1916 (1♂, USNM), 11.v.1916 (1♀, USNM), KS: Manhattan, C.W. Sabrosky, 27.iv.1934 (1♀, USNM), 17.vi.1934 (1♀, USNM), Lawrence, Nat. Hist. Res., 27.iii.1954, J.G. Chillcott, CNC352781 (1♂, CNC), 28.v.1956, J.G. Chillcott, CNC352811–352818 (8♀, CNC), MI: Detroit, 24.v.1935, G. Steyskal (1♂, USNM), E Lansing, C. Sabrosky, 2.vi.1934 (1♀, USNM), 29.iv.1942 (1♀, USNM), NC: Scotland Co., Laurinburg, St. Andrews University, 18.iv.2017, T.S. Feldman, Celtis laevigata, em. ~17.iv.2018, #CSE4418, CNC1144090 (1♀, CNC), OK: Payne Co., Mehan, 36.014339°N, 96.996744°W, 5.iv.2016, em. by 11.iv.2017, M.W. Palmer, ex Ulmus rubra, #CSE3448, CNC939912 (1♂, CNC), VA: Chain Bridge, 23.iv.1922, J.R. Malloch (1♀, USNM), Fairfax Co., Great Falls Park, quarry, 38°59.1'N, 77°14.8'W, Malaise trap, D.R. Smith, 18–23.iv.2007 (2♀, USNM), 13–24.v.2007 (1♀, USNM), VT: USA. Vermont: Chittenden Co., South Burlington, Winooski Gorge, 16.v.2015, C.S. Eiseman, Ulmus rubra, em. 24.iii-6.iv.2016, #CSE2264, CNC654497–654499 (3♀, CNC).

Comments

The dissected paratype of Agromyza aristata discussed by Spencer (1969) has been examined and the genitalia are identical to those of the male illustrated here, not as illustrated in Spencer (1969). The unusual illustration in Spencer (1969) appears to belong instead to a non-type male in the USNM from Illinois that has had the phallus broken off near the base of the mesophallus. This mistake in interpreting the male genitalia led Spencer to describe A. marmorensis in the same revision, stating that while "Not satisfactorily distinguishable from A. aristata” based on external characters, the differences in the genitalia were significant enough to separate them. Since it is now clear that the genitalia of the two species are also identical, A. marmorensis is here included as a junior synonym of the senior A. aristata.

The new records for NB, QC, MI and KS represent new adult records, but these were previously suspected from leaf mines (Eiseman et al. 2021).

Agromyza bispinata Spencer

Figs 38, 41, 42, 167–174

Agromyza bispinata Spencer, 1969: 39. Spencer and Steyskal 1986b: 263; Eiseman and Lonsdale 2018: 8.

Description

(Figs 38, 41, 42). Wing length 2.2–2.4 mm (♂). Female unknown [see discussion below]. Length of ultimate section of vein M4 divided by penultimate section: 0.5–0.7. Eye height divided by gena height: 4.2–6.0. Male first flagellomere large (usually at least 30% longer than pedicel, but not much longer than pedicel in a minority of specimens) and varying in shape from elongate and subovate to (more commonly) circular; at least distal 1/2 (usually distal 2/3) covered with long hairs. Ocellar triangle relatively small with corners slightly rounded. Fronto-orbital plate slightly projecting.

Chaetotaxy : Two ori; two ors. Three dorsocentrals, decreasing in length anteriorly with anterior seta not much longer than surrounding setulae. Mid tibia with two posteromedial setae.

Colouration : Body primarily dark brown with halter white. Base of first flagellomere and distal margin of pedicel orange; orange region sometimes either extending to basal 1/3 of first flagellomere or strongly reduced (particularly if spines on surstylus reduced). Gena sometimes paler, excluding ventral margin. Calypter white with margin sometimes yellowish or slightly brown, and hairs light brown. Base of fore tibia paler or segment paler overall. Tarsi yellow to orange-brown.

Genitalia : (Figs 167–173) Surstylus small, lobate, margin darkly pigmented; with two or three medial setae and two large posterior spines; spines sometimes minute and rounded and distiphallus more elongate with relatively pronounced internally haired medial section; surstylus barely visible when viewed laterally and with basal suture obliterated. Hypandrium relatively narrow and tapered apically with membranous window in lobe. Postgonite relatively small and upcurved. Halves of basiphallus converging to, and overlapping at base; with small mediolateral membranous lobe on each sclerite; twisted medially and broad apically, with converging ventrodistal points. Distiphallus of “nigripes-type” (capsule-shaped with subbasal opening for entry of ejaculatory duct, pronounced dorsobasal collar, and medial convolution); relatively narrow and pale with sides parallel, being only slightly wider apically.

Variation : Phallus sometimes elongate with pronounced medial section with densely spinulose inner surface and elongate medial membranous projection on right sclerite of mesophallus (Fig. 174); if so, spines on epandrium reduced to absent. Possibly a new species, but “typical” males show intermediate degrees of spine reduction on epandrium that resemble the state seen in these other males.

Host

PoaceaeElymus hystrix (Eiseman and Lonsdale 2018).

Distribution

Canada: ON, MB*. USA: CT*, GA, IA, MD, NC, NH*, NY, PA*, UT, VA, WV*.

Type material

Holotype: Canada. ON: Simcoe, 5.vi.1939, G.S. Steyskal (1♂, CNC).

Paratypes : Canada. ON: Midland, 20.vii.1955, J.G. Chillcott, CNC352821 (1♂, CNC), Pt. Ryerse, 1.vi.1939, G.E. Shewell, CNC352822, (1♂, CNC), Simcoe, 5.vi.1939, G.E. Shewell, CNC352819 (1♂, CNC).

Males examined

Canada. MB: Ninette, 28.vii.1958, “sq. fringe darker spines on surs. larger aed. larger”, J.G. Chillcott, CNC352820 (1♂, CNC). USA. CT: Redding, 11.vi.1929, A.L. Melander (1♂, USNM), Rabin Co., 13.vii.1957, W.R. Richards, CNC352824 (1♂, CNC), IA: Winneshiek Co., 43°25'55.97"N, 92°0'34.78"W, 16.vii.2015, C.S. Eiseman, Elymus hystrix em. 7.viii.2015 , #CSE1973, CNC564662 (1♂, CNC), MD: Bethseda, 30.v.1980, G.C. Steyskal, swept ex. Viola papilionaceae (2♂, USNM), Bethseda, 14.v.1981, G.C. Steyskal (1♂, USNM), 11.viii.1981 (1♂, USNM), 14.viii.1981 (2♂, USNM), 16.viii.1981 (1♂, USNM), 1.viii.1981 (3♂, USNM), 17.v.1969 (1♂, USNM), 7.viii.1981 (2♂, USNM), 13.ix.1981 (1♂, USNM), 27.v.1972 (1♂, USNM), Plummers Isl., 19.vi.1913, R.C. Shannon (1♂, USNM), Montgomery Co., Dickerson, 14.vii1974, G.A. Foster (13♂, USNM), 4mi SW of Ashton, G.F. and J.F. Hevel, 16.viii.1986 (1♂, USNM), 24.vii.1982 (1♂, USNM), 27.viii.1981 (1♂, USNM), Cabin John, 20.vi.1931, A.L. Melander (1♂, USNM), NC: Macon Co., Wayah Gap, 3800', 29.vii.1957, J.G. Chillcott (1♂, USNM), Jackson Co., Cherokee, 609 m, 25.vii.1957, CNC352823 (1♂, CNC), NH: Bretton Wda, 1.vii.1936, A.L. Melander (1♂, USNM), PA: Allegeny Co., Little Orleans, Little Orleans campground, 6.vi.1998, sweeping, C.R. Bartlett (1♂, UDCC), VA: Falls Church, Holmes Run, 26.vi.1961, light trap, W.W. Wirth (1♂, USNM), Shenandoah, Big. Meadows, A.L. Melander, 1.vii.1939 (1♂, USNM), 2.vii.1939 (2♂, USNM), 5.vii.1939 (1♂, USNM), Great Falls, 9.vii.1926, A.L. Melander (1♂, USNM), Glencarlyn, 2.vi.1925, J.R. Malloch (1♂, USNM), Fairfax Co., Dead Run, 22.vi.1916, R.C. Shannon (1♂, USNM), Fairfax Co., Turkey Run Park, nr. mouth of Turkey Run, 38°57.9'N, 7°09.4'W, Malaise trap, 18–30.v.2007, D.R. Smith (1♂, USNM), WV: Parkersburg, 21.vi.1970, G. Steyskal (1♂, USNM), Morgan Co., nr. Great Cacapon, 1.ix.1984, G.F. and J.F. Hevel (1♂, USNM), Hardy Co., Lost River St. Pk., 1–14.viii.1960, K.V. Krombein (1♂, USNM).

Females examined

USA. CT: Colebrook, 28.viii.1941, A.L. Melander (1♀, USNM), MD: Coleville, W.W. Wirth, 21.v.1977 (1♀, USNM), 11.v.1977 (1♀, USNM), NC: Macon Co., Wayah Bald, 1066 m, 13.vii.1957, W.R. Richards, CNC352828 (1♀, CNC), Macon Co., Wayah Gap, 16.vii.1957, J.G. Chillcott, CNC352827 (1♀, CNC), NH: Franconia Notch, 8.vii.1931, J.M. Aldrich (1♀, USNM), NY: Ithaca, 31.v.1914, A.L. Melander (3♀, USNM), TN: Smokies, Chimneys, 25.vi.1941, A.L. Melander (1♀, USNM), VA: Falls Church, 18.vii.1960, W.W. Wirth (1♀, USNM), Shenandoah, Big Meadows, A.L. Melander, 3.vii.1939 (1♀, USNM), 2.vii.1929 (1♀, USNM), Shenandoah, Lewis falls, 3.vii.1939, A.L. Melander (1♀, USNM), Fairfax Co., Dead Run, 28.vii.1915, R.C. Shannon (1♀, USNM), Shenandoah Co., Mt. Jackson, 25.v.1962, J.G. Chillcott, J.R. Vockeroth, CNC352825, CNC352826 (2♀, CNC).

Comments

Agromyza bispinata belongs to a complex of species defined by an enlarged and long-haired male first flagellomere, whose species appear to be separable only on the basis of male genitalic morphology. The females of these species, which are currently indistinguishable from one another, differ from the males in having an elongate oval first flagellomere with the long pale hairs restricted to the distal margin; the examined females are listed in the material examined section for this species. The sparse, long hairs on the first flagellomere of these species (Figs 41, 42) should not be confused with the discrete apical tuft of hairs characteristic of many other Agromyza species, including A. canadensis, A. kincaidi, and A. pudica (see Figs 39, 40). This complex currently includes A. echinalis in the east Nearctic, A. hockingi Spencer in Canada and the western United States (but expected in the eastern United States), and the widespread A. tacita and A. bispinata. There is a relatively large amount of genitalic variation within the latter two species, including distiphallus morphology in A. bispinata and epandrial spine number in A. tacita, which suggests the presence of cryptic taxa. The presence of only two epandrial spines, which is likely the derived state, currently defines A. bispinata within this group, but more rigorous analysis of the clade as a whole is required to better delineate species.

Agromyza canadensis Malloch

Figs 1, 2, 7–10, 175–178

Agromyza canadensis Malloch, 1913: 299. Frick 1952a: 372; Spencer 1969: 39, 1990: 198; Spencer and Steyskal 1986b: 59; von Tschirnhaus 1993: 512 [suspected to be conspecific with A. pseudorufipes Nowakowski].

Description

(Figs 1, 2, 7–10). Wing length 3.1–3.9 mm (♂), 3.3–4.0 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.5–0.7. Eye height divided by gena height: 4.2–7.3. First flagellomere small and rounded; pale tuft of apical hairs varies from teardrop-shaped to encompassing distal 1/3 of segment. Ocellar triangle relatively small with corners rounded.

Chaetotaxy : Three or two ori; two ors. Five dorsocentrals (sometimes small fifth seta indistinct), decreasing in size anteriorly. Mid tibia with two posteromedial setae.

Colouration : Setae brown. Body predominantly brown with yellowish tint or yellow with brownish tint, with thorax darker and halter white. First flagellomere dark brown to brown and scape and pedicel yellow to brownish; face and lunule dirty yellow in ON specimens; gena (excluding ventral margin), parafacial (excluding inner margin) and anterior margin of postgena light brown in female; postpronotum and notopleuron yellowish in part; lateral and sometimes apical margins of scutellum and postsutural scutum yellow in females and some males. Pleuron paler brown with venter darker and posterior region yellowish; metanotum yellow with brown mottling and dark brown mediotergite; males sometimes darker. Calypter margin and hairs white. Legs yellow with faint brownish tint; tarsi sometimes paler and femora sometimes slightly darker. Abdomen brownish yellow, paler in female with dorsum sometimes brownish and oviscape brown.

Genitalia : (Figs 175–178) Surstylus indistinct, fused to epandrium; inner surface with many small, rounded tubercle-like setae not reaching anterior corner. Hypandrium relatively narrow with apex pointed; inner lobe small, rounded and with large separate medial sclerite closely associated with postgonite. Postgonite large and flat with partial fusion to hypandrial band; with sparse hairs on outer face. Halves of basiphallus flat, wide and poorly sclerotised; apices broad, converging past internal subapical fold. Distiphallus very large, resembling a thick band with anterior and posterior margins strongly curled outwards; basal portion with small, shallow lateral sculpturing. Ejaculatory apodeme well-developed basally, but reduced, thin and pale apically.

Hosts

BoraginaceaeCynoglossum, Mertensia (Benavent-Corai et al. 2005).

Distribution

Canada: ON, QC*. USA: CA, VA*.

Type material

Holotype: Canada. ON: Cottage Beaulieu, Beaulieu, 14.viii.1906 (1♀, USNM).

Additional material examined

Canada. ON: Hawkeston, 28.vi.1927, C.H. Curran, CNC353079 (1♂, CNC); Mer Bleu[e], 5mi E Ottawa, 9.vi.1966, D.D. Munroe, CNC353082 (1♂, CNC), Ottawa, Montfort Hosp., 17.vii.1993, J.R. Vockeroth, CNC353074 (1♂, CNC), Ottawa, 18.viii.1924, G.S. Walley, CNC353081 (1♀, CNC), damp second-growth Acer-Betula woods, 17.viii.2000, J.R. Vockeroth, CNC353075 (1♀, CNC), 30.v.1958, J.R. Vockeroth, CNC353078 (1♂, CNC), 30.vi.1958, J.R. Vockeroth, CNC353080 (1♂, CNC); damp second-growth Acer-Betula woods, 6.viii.1996, J.R. Vockeroth, CNC353073 (1♂, CNC), QC: Burnett, 13.viii.1977, J. O’Hara, CNC353076 (1♀, CNC), Old Chelsea, 11.vi.1959, J.R. Vockeroth, CNC353077 (1♀, CNC). USA. VA: nr. Plummers Island, 20.v.1914, R.C. Shannon (1♂, USNM), Giles Co., Ripplemead, White Riverbend Pk., 37°19'N, 80°41'W, 9.v.2006, S.A. Marshall (1♀, DEBU), Fairfax Co., Great Falls Park, quarry, 38°59.1'N, 77°14.8'W, Malaise trap, D.R. Smith, 24.iv-2.v.2007 (4♀, USNM), 3–10.v.2007 (12♀, USNM).

Comments

Agromyza canadensis is highly similar to the Palaearctic A. pseudorufipes Nowakowski, especially with regards to the derived structure of the phallus. It is also a borage feeder on Myosotis, Podonosma and Trigonotis (Benavent-Corai et al. 2005). Von Tschirnhaus (1993) suggested that the two species could be conspecific, but additional study is required.

Agromyza deserta (Patton)

Cecidomyiaceltis deserta Patton, 1897: 247. Spencer 1990: 54.

Agromyza deserta. von Tschirnhaus 2016: 153; Scheffer and Lonsdale 2018: 85.

Description

(von Tschirnhaus 2017: figs 6–9). Wing length 3.4 mm (♂). Female unknown. Length of ultimate section of vein M4 divided by penultimate section: 0.4. Eye height divided by gena height: 4.5. Ocellar triangle not much larger than tubercle. Orbital plate narrow, inner margin weakly defined. Epistoma present, margin straight, as long as width of first flagellomere. Ventral margin of gena relatively straight, shallow anteriorly and deep posteriorly. Scutum densely matt. Costa extending to M1. Tibiae without medial setae

Chaetotaxy : Ocellar and postvertical setae strong. Two ori (three ori on right side), decreasing in length anteriorly; two ors. Minute interfrontals anteromedially on frons. Four dorsocentral setae on left side, five on right, decreasing in length anteriorly. Acrostichal seta strong. Acrostichal setulae in six irregular rows. Two or three anepisternal setae, four or five katepisternals.

Colouration : Setae dark brown. Mostly dark brown. Face, parafacial, gena and postgena (to point above midpoint of eye) and mouthparts yellowish white with dark brown line along venter of gena; lunule and antenna pale yellow with arista brown; frons light yellow with brownish/orange tint, with ocellar triangle and posterolateral corner of frons dark brown, faded brownish region around ocellar triangle, with brownish stripe along orbital plate almost to base of posterior ori, becoming indistinct anteriorly. Apices of femora and bases of tibiae narrowly yellow. Halter white with stem faintly brownish anteriorly. Calypter margin and hairs brown.

Genitalia : (von Tschirnhaus 2017: figs 1–3, 5) Cercus large, bent on outer margin and straight on inner; inner-distal surface with numerous pointed tubercle-like setae. Surstylus fused to epandrium, indistinct, without tubercle-like setae. Hypandrium subtriangular; apex rounded with narrow, pointed apodeme that may be 2 × as long as wide; inner lobe with basal section narrowing to point of attachment to thicker, angled distal section bearing row of several sockets, one of which has a moderately long seta. Postgonite with minute setulae in irregular double row; shape presumably similar to that of A. aristata. Phallophorus narrow, width < 1/3 length. Basiphallus dorsally fused to phallophorus on narrow dorsal/left lateral sclerite, which is apically truncated; right lateral sclerite much narrower, but as long as left sclerite. Paraphallus and hypophallus absent. Ejaculatory duct becoming broader along length of basiphallus, thick along mesophallus. Mesophallus a single band-like ventral sclerite partially fused to distiphallus; nearly as long as basiphallus, strongly curved to form a shallow semicircle. Distiphallus very shallow, band-like, and ventrally curved, obliquely angled. Ejaculatory apodeme unknown.

Host

CannabaceaeCeltis occidentalis (von Tschirnhaus 2016).

Distribution

USA: CT, NY, WV.

Type material

Holotype: USA. CT: Orange, gall on twig of Celtis occidentalis, W.H. Patton. [1 gall, Lost]

Other material

USA. WV: Morgan County ca. 6 mi NW Hedgesville, 39°37'N, 78°03'W, ex twig swelling gall Celtis occidentalis, coll. 6-v-1998, em. 15-iii-1999, R. J. Gagné (1♂, USNM). [not examined]

Comments

The adult of this species was thoroughly described in von Tschirnhaus (2016), which also provided a discussion on related species, host usage, and treatment in the literature.

Agromyza diversa Johnson

Figs 179–182

Agromyza diversa Johnson, 1922: 26. Frick 1953: 58; Spencer 1969: 41; Spencer and Steyskal 1986b: 59; Eiseman and Lonsdale 2018: 9.

Liriomyza diversa. Frick 1952a: 402.

Description

Wing length 2.9–3.3 mm (♂), 2.9–3.8 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.8. Eye height divided by gena height: 6.1–-9.7. First flagellomere sometimes slightly longer than high; with teardrop-shaped tuft of pale hairs at apex.

Chaetotaxy : Two ori; two ors. Three or four dorsocentrals, decreasing in length anteriorly. Mid tibia with two or three posteromedial setae.

Colouration : Setae black. Body yellow, except as follows: head mostly dark brownish with yellow tint, with mouthparts light yellow, lunule yellow, fronto-orbital plate (narrow) sometimes yellowish to yellow, gena (excluding ventral margin) and face light brown, and ocellar tubercle sometimes yellowish around base of ocellar and postocellar setae; postpronotum, notopleuron and scutellum pale yellow/whitish; halter white; female with oviscape dark brown and face yellow. Calypter margin and hairs white.

Genitalia : (Figs 179–182) Surstylus not distinguishable from epandrium; inner face of surstylus flat and covered with numerous small, rounded tubercle-like setae that do not extend to anterior corner. Postgonite relatively large, flat, and broad. Halves of basiphallus long and narrow, with right sclerite bent at midpoint; apices not bent inwards. Hypophallus convoluted and with scalloped apical fringe and irregularly sclerotised base. Mesophallus subcylindrical, with sides bulging and with transverse dorsal band; closely associated with distiphallus, which is bulging medially, truncated apically and with subapical constriction.

Host

UrticaceaeLaportea canadensis.

Distribution

Canada: ON. USA: IA, IL, IN, IN, MA, MD*, NC, NY, OH, TN, VA, VT.

Type material

Holotype: USA. MA: Chester, 7.viii.1912, C.W. Johnson (1♀, MCZ).

Paratype examined

USA. IN: Lafayette, viii.1912, J.M. Aldrich, Paratype No. 50034 U.S.N.M. / CNC Type No. 16287, CNC352872 (1♀, CNC).

Additional material examined

Canada. ON: Simcoe, 26.vi.1939, G.E. Shewell, CNC352873 (1♀, CNC). USA. OH: Delaware Co., Sunbury, Monkey Hollow Rd., 13–15.ix.2014, ex. Laportea canadensis, em. 16–24.x.2014, C.S. Eiseman, #CSE1432, CNC384847, CNC384848 (1♂ 1♀, CNC), IA: Ames, Pammel Woods, W.H. Robinson, 24.vi.1970 (2♂, USNM), 9.viii.1969 (1♂, USNM), 19.vi.1970 (1♂, USNM), Ames, Pammel Woods, 11.vi.1970, R.M. Miller (1♀, USNM), Ames, 1.viii.1969, W.H. Robinson (1♂, USNM), Allamakee Co., 3mi ESE Waterville, 43°11.0'N, 91°14.1'W, 2.viii.1960, J. Laffoon (1♂, USNM), Boone Co., Ledges State Park, 14.vii.1971, R.M. Miller (1♀, USNM), IL: University of Illinois Woods, 2.vii.1945, “X 526” (1♂, USNM), 15.vii.1945, “X 527” (1♂, USNM), MD: Montgomery Co., Dickerson, 14.vii.1974, G. Foster (1♂, USNM), OH: Delaware Co., Sunbury, Monkey Hollow Rd., 13–15.ix.2014, em. 16–24.x.2014, C.S. Eiseman, ex Laportea canadensis, #CSE1432, CNC384847, CNC384848 (1♂ 1♀, CNC), NC: Graham Co., Poplar Cove, 2500', 35°21.3'N, 83°56.1'W, 20.vi.1958, J. Laffoon (1♀, USNM), NY: Ithaca, 15.vi.1932 (1♀, USNM), TN: Davidson Co., Nashville, West Meade Waterfall (36.094655, -86.913252), 21.viii.2017, em. 15–17.ix.2017, C.S. Eiseman, ex Laportea canadensis, #CSE4273, CNC939958–939960 (1♂ 2♀, CNC), VA: Fairfax Co., Dead Run, 28.vii.1915, R.C. Shannon (1♀, USNM).

Agromyza echinalis sp. nov.

Figs 232–235

Description

Externally as described for A. bispinata except as follows: Wing length 2.5–2.6 mm (♂). Female unknown. Length of ultimate section of vein M4 divided by penultimate section: 0.6. Eye height divided by gena height: 4.2. First flagellomere always enlarged in male, broadly ovate, slightly longer than high and covered with long hairs past base. Additional ori sometimes present on one side of frons. Wing veins yellow. Antenna orange with brownish tint, mostly brownish past base of first flagellomere; frons tinted with orange; fore tibia yellow with centre brownish.

Genitalia : (Figs 232–235) Surstylus large and triangular with numerous long spines along inner margin and with one spine at apex; basal suture obliterated, inner surface flat and easily viewed posteriorly. Hypandrium and postgonite as described for A. bispinata. Halves of basiphallus with medial membranous process that is longer and truncated on right side. Distiphallus of “nigripes-type” (capsule-shaped with subbasal opening for entry of ejaculatory duct, pronounced dorsobasal collar, and medial convolution); ventral curve with dark truncated elbow; ventral surface with medial longitudinal suture; sides shallowly rounded and internally spinulose subapically.

Host

Unknown.

Distribution

USA: MD, TN, VA.

Etymology

Gr., referring to the characteristic spiny surstylus.

Type material

Holotype: USA. TN: Clarksville, 13.v.1936, E.W. Howe (1♂, USNM).

Paratypes: USA. MD: nr. Plummers Isl., 6.vi.1914, R.C. Shannon (1♂, CNC), VA: Fairfax Co., Dead Run, 19.vi.1915, R.C. Shannon (1♂, USNM).

Comments

The heavily spinulose surstylus of Agromyza echinalis is the primary defining feature that distinguishes it from other species with an enlarged first flagellomere. As in A. tacita (Figs 226–228) and A. hockingi (Colorado, Texas, Utah; Spencer and Steyskal 1986b: figs 411–413), the surstylus is triangular and clearly visible laterally, but these other two species have a slightly darker and more rounded surstylus with a maximum of five stout spines.

Agromyza fission Eiseman & Lonsdale

Figs 183–187

Agromyza fission Eiseman & Lonsdale, 2018: 9.

Description

(from Eiseman and Lonsdale 2018). Wing length 2.3 mm (♂). Female unknown. Length of ultimate section of vein M4 divided by penultimate section: 1.0. Eye height divided by gena height: 3.9. First flagellomere small and rounded, without pale tuft of hairs. Fronto-orbital plate slightly projecting (more so anteriorly). Ocellar triangle small and rounded. Thorax with light pruinosity.

Chaetotaxy : Two ori; two ors. Three strong dorsocentrals, anterior seta 2/3 length of second. Mid tibia without posteromedial setae.

Colouration : Setae black. Body mostly dark brown. Head yellowish orange with first flagellomere infuscated on distal 2/3, back of head and occiput dark brown, frons dark brown behind level of hind fronto-orbital, face white and clypeus brown to dark brown. Calypter margin and hairs brown. Halter white. Tarsi, base of fore tibia and apex of fore femur yellow.

Genitalia : (Figs 183–187) Surstylus small, lobate, setulose, and slightly angled anteriorly; fused to anteroventral margin of epandrium with suture partially evident. Cercus broad and emarginate apically; inner surface covered with numerous small tubercle-like setae. Postgonite small and rounded with shallow inner lobe. Single sclerite of basiphallus fused to phallophorus on left side, twisted dextrally; distal margin forming small transverse sclerite. Mesophallus small and flat. Distiphallus composed of a single dark, flat, curved sclerite (shortest dorsally) nearly forming a complete tube; lateral margins fringed and not meeting ventrally.

Host

CannabaceaeCeltis occidentalis (Eiseman and Lonsdale 2018).

Distribution

USA: IA, MD, OK, WI.

Type material

Holotype. IA: Allamakee Co., Red Oak Prairie (43°14'13.43"N, 91°7'8.58"W), 16.vii.2015, em. 2.viii.2015, C.S. Eiseman, ex Celtis occidentalis, #CSE1925, CNC564711 (1♂, CNC).

Paratypes. IA: same collection as holotype, CNC564712 (1♂, CNC), MD: Plummers Isl., 23.v.1914, R.C. Shannon (1♂, USNM), OK: Payne Co., Mehan, 36.014339°N, 96.996744°W, 5.iv.2016, em. 20– 22.iv.2017, M.W. Palmer, ex Celtis occidentalis, #CSE3529, CNC939918, CNC939919 (1♂ 1♀, CNC); WI: Buffalo Co., Alma, S1287 State Road 88, 17.vii.2015, em. 3.iv.2016, C.S. Eiseman, ex Celtis occidentalis, #CSE2311, CNC634841 (1♂, CNC).

Comments

Although largely similar to Agromyza varifrons, the male cerci are highly derived and diagnostic: they are strongly widened apically and deeply cleft, with small, tubercle-like setae covering the inner surface.

Agromyza isolata Malloch

Figs 188–192

Agromyza isolata Malloch, 1913: 306. Frick 1952a: 373; Spencer and Steyskal 1986b: 264; Eiseman and Lonsdale 2018: 11.

Agromyza albitarsis Meigen. Misidentification. Frick 1959: 353.

Agromyza populoides Spencer, 1969: 52. Spencer and Steyskal 1986b [synonymy].

Description

Wing length 2.3 mm (♂), 2.5–2.7 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.7. Eye height divided by gena height: 4.8–7.5. First flagellomere small and rounded; without pale tuft of apical hairs. Ocellar triangle only indicated laterally as subshiny regions.

Chaetotaxy : Two ori; two ors. Four strong dorsocentrals, one presutural. Mid tibia with one posteromedial seta.

Colouration : Setae golden-brown, distinct from scutum. Scutum with brownish grey pruinosity. Calypter margin and hairs white (brown in isolata holotype). Body predominantly brown with halter white; base of first flagellomere and most of pedicel distinctly orange, entirely dark in isolata holotype; BC, CA, and CO material with first flagellomere dark, with basal margin slightly lighter; tarsi, tibiae (hind tibia brownish medially) and apices of femora yellow. Wing veins light brown.

Genitalia : (Figs 188–192) Surstylus broad, not distinct from epandrium and with a number of medial tubercle-like setae along inner-distal margin on posterior 1/2. Hypandrium relatively narrow with long, tapered tip. Postgonite small and dome-like. Phallophorus elongate and produced on left side, fusing with single sclerite of basiphallus. Basiphallus short, shallowly curved and restricted to dorsal surface. Mesophallus curved in cross-section, dark, and with long, bifid, tapering “tail”, with bases either separated or fused; distiphallus short, dark, and cap-like. Ejaculatory apodeme relatively small, blade pale.

Host

SalicaceaePopulus spp., Salix spp. (Eiseman and Lonsdale 2018).

Distribution

Canada: AB, BC*, ON, NS*, QC, SK. USA: CA, CO, MN [leaf mine only], PA, VA*, VT, WA.

Type material

Holotype [isolata]: USA. CA: Eureka, “22.5”, H.S. Barber (1♀, USNM; type No. 17573).

Holotype [populoides]: Canada. SK: Regina, em. 23.viii.1965, [ex leaf mine on Populus deltoides ♀ X P. balsamifera ♂ (Spencer, 1969)], ex Gracillarid sp., RRD W65, Ex 2616(10), G.N. Still (1♂ [with puparium], CNC).

Paratypes examined

[populoides]: Canada. ON: Grand Bend, 20.vii.1939, G.E. Shewell, CNC352879 (1♂, CNC), Ottawa, em. 28.vii.1967, mine Pop. Bals. 10.vii.1967, CNC352880 (1♀ [with puparium], CNC), SK: same data as holotype CNC352875–352878, CNC352881-CNC3528884 (4♂ 4♀ [seven with puparia], CNC).

Additional material examined

Canada. BC: Terrace, grass, clover and buttercups, 7.vi.1960, R. Pilfrey, sweeping, CNC353092 (1♂, CNC), Wasa Lk., em. 29.vii.1959, “59-6779-01”, “Populus trichocarpa”, “R’rd”, CNC353083, CNC353084 (2♂ [with puparia], CNC), NS: S. Harbour Bch., PG962943, 6.vii.1983, J.R. Vockeroth, CNC353091 (1♂, CNC), ON: Grand Bend, 20.vii.1939, G.E. Shewell, CNC353089 (1♂, CNC), Ottawa, 15.ix.1997, J.R. Vockeroth, CNC353090 (1♂, CNC), 16.ix.1952, CNC353088 (1♀, CNC). USA. CO: Doolittle Ranch, Mt. Evans, 2987 m, 3.viii.1961, C.H. Mann, CNC353085 (1♂, CNC), J.G. Chillcott, CNC353087 (1♀, CNC), Summit L. Flats, Mt. Evans, 3901 m, 24.vii.1961, C.H. Mann, CNC353086 (1♂, CNC), VA: Falls Church, reared 25.v.1916, Populus, host: underside tentminer, J.J. deGryse (1♂ 2♀, USNM), VT: Chittenden Co., South Burlington, Winooski Gorge, 29.vi.2014, ex. Populus tremuloides, em. 15.vii.2014, C.S. Eiseman, #CSE1176, CNC384839–384841 (3♂, CNC), Chittenden Co., Williston, Mud Pond, 28.viii.2016, C.S. Eiseman, Populus grandidentata, em. 18–19.ix.2016 , #CSE3000, CNC654500–654504 (2♂ 3♀, CNC).

Agromyza kincaidi Malloch

Figs 39, 40, 193–197

Agromyza kincaidi Malloch, 1913: 285. Frick 1952a: 372 [as synonym of ambigua Fallén]; Spencer 1969: 45; Spencer and Steyskal 1986b: 264.

Description

(Figs 39, 40). Wing length 3.6–4.0 mm (♂), 3.9–4.3 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.7–0.8. Eye height divided by gena height: 2.9–3.2. First flagellomere small and rounded with small apical tuft of pale hairs. Ocellar triangle 1/2 length of frons, shape equilateral or more tapered apically. Buccal cavity subquadrate and clypeus broad. Costa only extending to R4+5. Thorax with light pruinosity.

Chaetotaxy : Two ori; two ors. Four postsutural dorsocentrals, one presutural, decreasing in length anteriorly. Mid tibia with two posteromedial setae.

Colouration : Body dark brown with halter white. Gena slightly paler posteriorly. Fore tarsus, apex of fore femur and base of tibiae yellowish. Wing veins light brown. Calypter margin and hairs white. Female fronto-orbital plate slightly paler and less pruinose.

Genitalia : (Figs 193–197) Surstylus not strongly differentiated from epandrium, which has deep desclerotised anteroventral emargination; with numerous posterior and medial tubercle-like setae and with rounded inner-distal point bearing several small setae. Hypandrium gently arched, not produced apically; inner lobe largely membranous and with two furrows ending in one setula each. Postgonite small, flat and lobate. Halves of basiphallus folded inwards near apex; right sclerite shorter and fused to enlarged distal margin of phallophorus. Distiphallus of “nigripes-type” (capsule-shaped with subbasal opening for entry of ejaculatory duct, pronounced dorsobasal collar, and medial convolution); distal portion dark and extremely elongate with wide ventral furrow (dark medial ridge within furrow); inner surface with pronounced spinulose ridge; dorsal surface covered by dark membranous sheath. Ejaculatory apodeme broad and well-developed with distal margin pale and rib pronounced.

Host

Unknown – likely a grass-feeder, possibly on Bromus purgans (Spencer and Steyskal 1986b).

Distribution

Canada: AB, BC, MB, NL, NS, YT. USA: AK, CA, CO, IA*, NC*, NY*, TN, UT.

Type material

Holotype: USA. AK: Juneau, 25.vii.1899, Kincaid (1♀, USNM; type No. 15565).

Additional material examined

Canada. AB: Banff, Johnston Canyon, 1432 m, 18.vii.1962, K.C. Herrmann, CNC352900, CNC352937 (1♂,1♀, CNC), 30–31.vii.1962, CNC352938 (1♀, CNC), Banff, 19.vii.1922, E. Hearle, CNC353102 (1♂, CNC), Calgary, 20 mi W, Jumping Pd. Cr., 9.viii.1962, K.C. Hermann, CNC352901 (1♂, CNC), Wate[illegible], 15.vii.1923, H.L. Seamans, CNC352903 (1♂, CNC), BC: Bowser, 8.vi.1955, R. Coyles, CNC352936 (1♀, CNC), Kleanza Cr., 14 mi E Terrace, 4.vii.1960, C.H. Mann, CNC352942 (1♀, CNC), Mt. Thornhill nr. Terrace, 1066 m, in boggy seepage area, 14.vii.1960, J.G. Chillcott, CNC352933 (1♀, CNC), Pr. Rupert, Ledum-Kalmia bog, 4.iv.1960, R. Pilfrey, CNC352935 (1♀, CNC), 4.vi.1960, C.H. Mann, CNC352896, CNC352897 (2♂, CNC), Robson, 24.v.1950, H.R. Foxlee, CNC352928 (1♀, CNC), Summit Lake, Mi392 Alaska Hwy., 1280 m, 31.vii.1959, R.E. Leech, CNC352898 (1♂, CNC), Summit Lake, 1280 m, 21.vii.1959, R.E. Leech, CNC352899 (1♂, CNC), Terrace, Airport area, 4.viii.1960, C.H. Mann, CNC352934 (1♀, CNC), Terrace, 11.viii.1960, W.R. Richards, CNC352895 (1♂, CNC), 67 m, 15.viii.1960, B. Heming, CNC352929 (1♀, CNC), 67 m, 8.vi.1960, W.W. Moss, CNC352932 (1♀, CNC), 19.vii.1960, C.H. Mann, CNC352931 (1♀, CNC), 20.vii.1960, W.R. Richards, CNC352885 (1♂, CNC), 31.iv.1960, R.J. Pilfrey, CNC352894 (1♂, CNC), 31.v.1960, C.H. Mann, CNC352930 (1♀, CNC), 31.vi.1960, J.G. Chillcott, CNC353099 (1♂, CNC), MB: Churchill, 14.vii.1952, E.F. Pope, CNC352939 (1♀, CNC), Churchill, 58°46'N, 94°10'W, 5.viii.1952, J.G. Chillcott, project#NBP, CNC_Diptera59120 (1 ex, CNC), Deer River, mile 473 m Hudson Bay Ry., 3.viii.1952, J.G. Chillcott, CNC352886 (1♂, CNC), Int. Peace Gardens, Turtle Mtn. For. Res., 7.viii.1958, J.G. Chillcott, CNC352940 (1♀, CNC), NL: Bell Is., 4–7.viii.1967, J.F. McAlpine, CNC353108 (1♂, CNC), Lab., Cartwright, 20.vii.1955, E.F. Cashman, CNC352941 (1♀, CNC), St. John’s, Agric. Exp. Sta., 1.viii.1967, J.F. McAlpine, CNC353111–353113, CNC353117 (4♂, CNC), 15.vii.1967, J.F. McAlpine, CNC353126 (1♂, CNC), 16.vii.1967, J.F. McAlpine, CNC353124, CNC353110 (1♂,1♀, CNC), on Ranunculus, 18.vii.1967, J.F. McAlpine, CNC353103–353107 (5♂, CNC), Abies balsamea, 19.vii.1967, J.F. McAlpine, CNC353109 (1♀, CNC), 21.vii.1967, J.F. McAlpine, CNC353131, CNC353135 (2♂, CNC), 24.vii.1967, J.F. McAlpine, CNC353119 (1♀, CNC), 26.vii.1967, J.F. McAlpine, CNC353133 (1♂, CNC), 27.vii.1967, J.F. McAlpine, CNC353128 (1♂, CNC), 29.vii.1967, J.F. McAlpine, CNC353120, CNC353129 (1♂,1♀, CNC), 3.viii.1967, J.F. McAlpine, CNC353114–353116 (3♂, CNC), 30.vii.1967, J.F. McAlpine, CNC353121, CNC353125 (1♂,1♀, CNC), 31.vii.1967, J.F. McAlpine, CNC353130 (1♂, CNC), 9.viii.1967, J.F. McAlpine, CNC353118, CNC353122, CNC353123, CNC353127, CNC353132, CNC353134, CNC353136 (4♂,3♀, CNC), NS: Lockeport, Cranberry I., 24.vii.1958, J.R. Vockeroth, CNC352902 (1♂, CNC), YT: Ogilvie Mts., North Fork Crossing, 1.vii.1962, P.J. Skitsko, CNC352893 (1♂, CNC), Rampart House, 67°25'N, 140°59'W, 17.vii.1951, J.E.H. Martin, project#NBP, CNC_Diptera109349 (1♀, CNC), Whitehorse, 24.viii.1959, R. Madge, CNC352926, CNC352927 (2♀, CNC). USA. AK: Anchorage, 18.vii.1951, R.S. Bigelow, CNC352915 (1♀, CNC), Cold Bay, on tundra, 18.viii.1952, W.R.M. Mason, CNC352892 (1♂, CNC), Curry, 29.vi.1952, W.R. Mason, CNC352914 (1♀, CNC), King Salmon, Naknek, 13.vii.1952, J.B. Hartley, CNC352921, CNC352923, CNC352925 (3♀, CNC), 16.viii.1952, CNC352919 (1♀, CNC), 3.vii.1952, CNC352917 (1♀, CNC), 31.vii.1952, CNC352924 (1♀, CNC), 5.viii.1952, CNC352916 (1♀, CNC), 6.vii.1952, CNC352918, CNC352920, CNC352922 (3♀, CNC), Mile 315 Richard. Hwy., 8.vi.1951, W.R. Mason, CNC352913 (1♀, CNC), Naknek, 18.vii.1952, J.B. Hartley, CNC352910, CNC352911 (2♀, CNC), 8.vii.1952, J.B. Hartley, CNC352907 (1♀, CNC), W.R. Mason, CNC352912, CNC352908, CNC352909 (3♀, CNC), CO: Doolittle Ranch, Mt. Evans, 2987 m, 22.vii.1961, J.G. Chillcott, CNC352905 (1♀, CNC), 3.viii.1961, W.R.M. Mason, CNC352888 (1♂, CNC), B.H. Poole, CNC352906 (1♀, CNC), Echo L., Mt. Evans, 3230 m, 25.vii.1961, C.H. Mann, CNC352904 (1♀, CNC), IA: Ames, 10.v.1947, A.R. Brooks, CNC352943 (1♀, CNC), NC: Gt. Smokies, Clingmans Dome, A.L. Melander, 18.vii.1941 (4♂ 3♀, USNM), 19.vii.1941 (4♂, USNM), 21.vi.1941 (1♂, USNM), Smokies, Andrews Bald, 9.vii.1941, A.L. Melander (2♂ 1♀, USNM), Smokies, Forney Ridge, 26.vi.1941, A.L. Melander (1♂, USNM), Clingman’s Dome, Grt. Sm. Mt. Nat. Park, 6.viii.1957, C.J. Durden, CNC352891 (1♂, CNC), 2011 m, 22.viii.1957, J.G. Chillcott, CNC352887 (1♂, CNC), NY: White Face Mt., 4000', 14.vii.1958, A.L. Melander (1♂, USNM), TN: Indian Gap to Clingman’s Dome, Gr. Sm. Mt. Nat. Park Tenn., 1584–2011 m, 6.viii.1957, J.G. Chillcott, CNC352889, CNC352890 (2♂, CNC).

Comments

Agromyza kincaidi is highly similar in appearance to the European and Nearctic A. ambigua Fallén (Spencer and Steyskal 1986b: figs 367–369), which has a highly similar phallus, an anterodorsally pointed first flagellomere (sometimes very strongly pointed in A. ambigua), a costa that extends just past R4+5 and dark colouration. While never directly compared, differentiation between the two has historically relied most heavily on the colour of the calypter margin and hairs, which are supposedly brownish in A. kincaidi and bright white to yellow in A. ambigua (Spencer 1969); vein colour does not appear to be yellowish in any A. ambigua examined, as noted in Spencer and Steyskal (1986b); examination of specimens further reveals calypter margin/hair colour to range from yellowish to brownish, and is even white in some material identified by Spencer. The limits of these two species should be re-evaluated in the future.

Agromyza pallidiseta Malloch

Agromyza pallidiseta Malloch, 1924: 192. Frick 1952a: 375; Spencer and Steyskal 1986b: 58.

Description

Wing length 2.3–2.4 mm (♀). Male unknown. Length of ultimate section of vein M4 divided by penultimate section: 0.6. Eye height divided by gena height: 1.6. Head as described for A. aristata except parafacial not as strongly projecting and ocellar seta 1/2 length of postocellar seta.

Chaetotaxy : As described for A. aristata, except there are four dorsocentrals (one presutural), not three.

Colouration : As described for A. aristata except as follows: setae yellow; first flagellomere yellow with basal 1/3 very pale; posterolateral corner of frons only dark to level of posterior (not anterior) ors.

Variation : Florida specimen (head mostly collapsed) with ocellar seta ~ 2/3 length of postocellar and fronto-orbital plate brown to base of anterior fronto-orbital.

Host

Unknown – possibly Celtis occidentalis (Ulmaceae).

Distribution

Canada: ON*. USA: DC, FL*.

Type material

Holotype: USA. DC: Rock Creek Park, 29.v.1922, J.R. Malloch (1♀, USNM).

Additional material examined

Canada. ON: Ottawa, Arboretum, adult on Celtis occidentalis, 27.v.2017, O. Lonsdale, CNC799468–799471 (4♀, CNC). USA. FL: Torreya St. Park, 29.iv.1952, O. Peck, on wood mud flat, CNC352944 (1♀, CNC).

Comments

The grey pruinose notum, yellow setae and head of Agromyza pallidiseta make it an easily recognised, albeit uncommon, species in eastern North America, known only from the holotype, one Florida female and four Ontario females. The Ontario females were taken as adults on Celtis occidentalis.

The Californian female listed as Agromyza pallidiseta in Spencer and Steyskal (deposited in the USNM) is clearly not conspecific with the holotype, as it is 2 × as large (4.6 mm), has a longer ocellar seta and brown setae.

Agromyza parca Spencer

Figs 198–204

Agromyza parca Spencer in Spencer and Steyskal 1986b: 265; Eiseman and Lonsdale 2018: 11; Eiseman et al. 2021: 6.

Description

Wing length 2.5–2.7 mm (♂), 2.4–2.7 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.8. Eye height divided by gena height: 3.5–6.7. First flagellomere small and rounded; males with small tuft of apical hairs (as wide as base of arista in specimens with entirely white calypter, and several times width of base of arista in males with darker calypter hairs); females with dark calypter also with wide tuft of hairs on first flagellomere. Ocellar triangle short and equilateral in female, longer in male. Scutum shiny to subshiny.

Chaetotaxy : Two ori (sometimes small to well-developed additional anterior ori on one side); two ors. Two well-developed dorsocentrals and one or two much smaller setae anteriorly. Mid tibia with two posteromedial setae.

Colouration : Body dark brown with halter white, lunule and gena (excluding ventral margin) paler, tarsi dirty yellow and tibiae sometimes yellow at base. Wing veins light brown. Calypter margin white with hairs brown to white.

Genitalia : (Figs 198–204) Surstylus with weak basal suture and numerous small medial and apical tubercle-like setae on inner margin. Inner lobe of hypandrium broad with several furrows leading to empty sockets. Postgonite small and lobate. Both sclerites of basiphallus with thin mediolateral membranous lobes; apices typically incurved. Distiphallus of “nigripes-type” (capsule-shaped with subbasal opening for entry of ejaculatory duct, pronounced dorsobasal collar, and medial convolution); distiphallus split ventromedially with dark subbasal swelling and central ridge; with spinulose inner structures and minute anterodorsal scales; seen ventrally, sides nearly parallel or segment narrowest apically (mostly in specimens with white calypter hairs).

Host

PoaceaeDichanthelium clandestinum, D. scoparium, Glyceria canadensis, G. striata (Eiseman et al. 2021).

Distribution

USA: CT*, DC*, IA, MA, MD*, NC, NH*, NJ*, NY*, TN.

Type material

Holotype: USA. NC: Mitchell Co., Roan Mtn., 6200', 13.viii.1957, J.G. Chillcott, CNC352945 (1♂, CNC).

Paratype: USA. NC: Same data as holotype, CNC352946 (1♀, CNC).

Additional material examined

USA. CT: Putnam Park, 20.vii.1939, A.L. Melander (1♂, USNM), Redding, 3.vi.1935, A.L. Melander (2♀, USNM), DC: 11.vi.1926, J.M. Aldrich (1♂, USNM), IA: Allamakee Co., Footbridge Farm, 22.vii.2018, J. van der Linden, Poaceae, em. by 14.viii.2018, #CSE4946, CNC1643675–1643677 (2♂ 1♀, CNC), MA: Concord, 17.vii.1961, W.W. Wirth (2♂ 1♀, USNM), Lavale, 9.v.1970, G. Steyskal (1♂, USNM), Hampshire Co., Pelham, Arnold Rd., 2.vii.2013, C.S. Eiseman, ex Dichanthelium clandestinum em. 20–22.vii.2013, #CSE724, CNC392666, CNC392667 (1♂,1♀, CNC), em. 20–23.iv.2014, #CSE1089, CNC384728, CNC384729 (2♀, CNC), Plymouth Co., West Bridgewater, Maple St., 15.viii.2013, C.S. Eiseman, ex Dichanthelium clandestinum em. 2–5.ix.2013, #CSE866, CNC392683–392687 (2♂ 3♀, CNC), MD: Colesville, W.W. Wirth, 4.vi.1977 (1♂, USNM), 1.viii.1976 (1♀, USNM), Bethseda, G. Steyskal, 12.ix.1981 (1♂, USNM), 14.v.1981 (1♀, USNM), Plummers Isl., at light, 3.viii.1915, R.C. Shannon (1♂, USNM), nr. Plummers Isl., 14.v.1915, R.C. Shannon (1♀, USNM), Montgomery Co., Carderock Park, 18.v.1989, M.J. and R. Molineaux (1♀, USNM), NC: Chatham Co., Haywood, 5.vi.1986, G.C. Steyskal (1♂, USNM), Durham Co., Durham, 17-Acre Wood Preserve, 36°1'27.82"N, 78°55'29.73"W, 8.v.2017, T.S. Feldman, Dichanthelium, em. 4.v.2018, #CSE4482, CNC1135677, CNC1135678 (1♂ 1♀, CNC), Scotland Co., Laurinburg, St. Andrews University, 10.v.2017, T.S. Feldman, Dichanthelium, em. 6–14.v.2018, #CSE4502, CNC1144099, CNC1144100 (1♂ 1♀, CNC), Wake Co., Morrisville, Lake Crabtree County Park, 35°50'37.77"N, 78°47'43.02"W, 6.vi.2018, T.S. Feldman, Dichanthelium scoparium, em. 25.vi.2018, #CSE4695, CNC1135686 (1♂, CNC), NH: White Mts., Stinson Lake, 23.vii.1961, W.W. Wirth (1♀, USNM), NJ: Lakehurst, 1.vi.1962, J.E. Puleston (1♂, USNM), NY: Bear Mt., 30.v.1941, A.L. Melander (1♀, USNM).

Tentatively identified

USA. NC: Scotland Co., Laurinburg, St. Andrews University, 3.v.2017, T.S. Feldman, Dichanthelium, em. 6.v.2018, #CSE4503, CNC1144101, CNC1144102 (2♀, CNC)

Comments

Agromyza parca is currently defined by an evenly dark antenna, minute dorsoapical spinules within the distiphallus and small membranous lateral lobes on the basiphallus. There is a slight amount of variation within the species, possibly indicating the presence of a cryptic taxon. Those specimens with an entirely white calypter (i.e., hairs not brown) have the pale tuft of hairs on the first flagellomere that is no wider than the base of the arista in the males and absent in the female, the eye is 3.5–5.9 × higher than the gena (not 5.2–6.7 × higher) and the distiphallus is usually slightly constricted apically (Fig. 198).

Agromyza parilis Spencer

Figs 205–209

Agromyza parilis Spencer in Spencer and Steyskal 1986b: 266.

Description

Wing length 2.5–2.7 mm (♂), 3.2–3.4 (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.7. Eye height divided by gena height: 2.5–2.8. First flagellomere slightly longer than high and broadly rounded, without apical tuft of pale hairs. Ocellar triangle relatively small and rounded.

Chaetotaxy : Two ori; two ors. Two well-developed dorsocentrals and one smaller anterior seta. Mid tibia with two posteromedial setae.

Colouration : Body dark brown with halter white, and gena and antenna sometimes slightly paler, with first flagellomere orange on inner-basal 1/3 and tarsi slightly paler. Calypter white with hairs dark brown; hairs sometimes yellowish in females.

Genitalia : (Figs 205–209) Surstylus narrow and rounded, entirely fused to epandrium and with numerous medial tubercle-like setae. Hypandrium with broadly rounded subtriangular apex; inner lobe with membranous base and inner-distal emargination. Postgonite small and lobate. Halves of basiphallus interlocking at base and with inner margins dark; outer margin weakly sclerotised, wide, wing-like and pointed; distal section distinct with long processes flanking distiphallus. Hypophallus lobate with pointed medial process. Distiphallus of “nigripes-type” (capsule-shaped with subbasal opening for entry of ejaculatory duct, pronounced dorsobasal collar, and medial convolution); distiphallus with dark enlarged sclerite above insertion of ejaculatory duct; distal section dark and elongate with medial suture and subapical texturing; transverse dorsobasal band produced ventrally. Ejaculatory apodeme with narrow rounded blade with numerous striations, long stem, and stout base.

Host

Unknown – likely Poaceae (Spencer and Steyskal 1986b).

Distribution

USA: TN, VA*.

Type material

Holotype: USA. TN: Hamilton Co., East Ridge, 9.v.1952, G.S. Walley, CNC352947 (1♂, CNC).

Material examined

USA. VA: Fairfax Co., Turkey Run Park, nr. mouth of Turkey Run, 38°57.9'N, 7°09.4'W, Malaise trap, D.R. Smith, 25.v-6.vi.2006 (1♂ 4♀, USNM), 18–30.v.2007 (3♂, USNM), Turkey Run Park, 0.3 km W mouth Turkey Run, 38°58'N, 77°09.6'W, Malaise trap, D.R. Smith, 17–24.v.2006 (4♂ 1♀, USNM), river, 14–17.v.2006 (4♂ 5♀, USNM), Great Falls Park, quarry, 38°59.1'N, 77°14.8'W, Malaise trap, 18–24.v.2007, D.R. Smith (1♂ 3♀, USNM).

Comments

While externally similar to a number of small dark eastern Agromyza, the first flagellomere of A. parilis is entirely dark on the outer face and there is no apical tuft of pale hairs in either sex. The male genitalia are most distinctive, however, and should be examined for confident identification.

Agromyza parvicornis Loew

Figs 210–214

Agromyza parvicornis Loew, 1869: 49. Frick 1952a: 373, 1957: 199 [lectotype designation]; Spencer 1969: 51, 1973: 252; Spencer and Steyskal 1986b: 67.

Description

Wing length 2.5–3.2 mm (♂), 2.6–2.9 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.7. Eye height divided by gena height: 2.1–3.6. First flagellomere small and rounded with minute tuft of pale apical hairs. Fronto-orbital plate projecting and anterior 1/2 of frons soft and buckled.

Chaetotaxy : Two ori (sometimes three on one side); two ors. Two dorsocentrals, sometimes with very small third seta present anteriorly. Two posteromedial setae on mid tibia.

Colouration : Body mostly dark brown with orange tint, first flagellomere brown with base orange (more extensive on inner surface); frons brownish orange with ocellar triangle, posterior margin and fronto-orbital plate to level of anterior or posterior ors dark brown, sometimes dark laterally along entire margin of eye; lunule yellowish; parafacial beige and gena (excluding ventral margin) light brown; halter white; fore knee and tarsus yellowish. Calypter white with hairs light brown; uncommonly dark brown.

Genitalia : (Figs 210–214) Surstylus visible laterally as small lobe; inner surface with more than a dozen tubercle-like setae. Hypandrium and postgonite as described for A. bispinata. Halves of basiphallus converging to base where both sclerites are fused to each other and phallophorus; with internal fold past midpoint and apices converging as broad triangular lobes. Distiphallus of “nigripes-type” (capsule-shaped with subbasal opening for entry of ejaculatory duct, pronounced dorsobasal collar, and medial convolution); base of distiphallus narrow, distal portion broadly rounded, widest subapically and with internal spinulose processes. Ejaculatory apodeme well-developed with central rib visible near base; blade pale.

Hosts

PoaceaeEchinochloa crus-galli, Panicum miliaceum, Zea mays and probably other cereals (Spencer and Steyskal 1986b).

Distribution

Canada: BC, ON, QC (Zhu et al. 2004). USA: "Widespread, present in most states” (Frick 1959). Argentina, Cuba, Guadeloupe, Puerto Rico, Dominican Republic, Saint Vincent (Valladares 1998; Martinez and Etienne 2002).

Type material

Lectotype: USA. DC [not given]: “Loew coll.”, parvicornis m. (1♂, MCZ).

Material examined

Canada. BC: Milner, 12.vii.1953, G.J. Spencer, CNC352949 (1♂, CNC), ON: Ottawa, Fletcher Wildlife Gardens, 45°23'7"N, 75°42'11.01"W, 9.vi.2013, O. Lonsdale, CNC1144190 (1♂, CNC), Ottawa, 8.vii.1925, F. Ide, CNC352950 (1♂, CNC), Point Pelee, 8.ix.1954, C.D. Miller, CNC352954, CNC352955 (2♀, CNC), Simcoe, 26.vi.1939, G.E. Shewell, CNC352952 (1♂, CNC), 29.vi.1939, CNC352953 (1♂, CNC), St. Lawrence Is. Nat. Park, Grenadier I. Centre, 26.vi.1975, H.C.W. Walther, Code 1-2406-25, CNC352951 (1♂, CNC), Picton, 10.vii.1970, J.F. McAlpine, CNC353093 (1♂, CNC), QC: Ile de Montreal, 17.vi.1966, Beaulieu, CNC352956 (1♀, CNC). USA. AZ: Sunnyside Canyon, Huachua Mts., 9.vii.1940, D.E. Hardy (1♂, USNM), DE: Wilmington, 31.vii.1951, D.F. Bray (1♂, USNM), Sussex Co., Dewey Beach, 21.xiii.2003, sweeping, K. Bennett (1♂, UDCC), IN: Lafayette 13.[?].1915, J.M. Aldrich (1♀, USNM), MA: Concord, vii.1960, E.H. Wheeler, reared ex. Zea mays (2♂ 3♀, USNM), Franklin Co., Northfield, 276 Old Wendell Rd., 42°38'48.74"N, 72°25'32.15"W, 31.viii.2017, C.S. Eiseman, Zea mays, em. 2–23.vi.2018, #CSE4580, CNC1135703–1135712 (5♂ 5♀, CNC), MD: Montgomery Co., Colesville, Malaise trap, W.W. Wirth, 7.viii.1975 (1♀, USNM), 14.vi.1973 (1♂, USNM), 20.viii.1975 (1♂, USNM), Bethseda, 6.viii.1967, G. Steyskal (1♂, USNM), Colesville, 24.vii.1974, W.W. Wirth (1♂, USNM), 4mi SW of Ashton, Malaise trap, G.F. and J.F. Hevel, 3.ix.1981 (1♂, USNM), 1.ix.1981 (2♂, USNM), NY: L.I. Veg. Res. Fm, Riverhead, at light, 1–7.viii.1938, 30.viii.1938, 7–20.viii.1938, CNC352957, CNC352958, CNC352959, CNC352960, CNC352961, CNC352962, CNC352963, CNC352964, CNC352965, CNC352966, CNC352967, CNC352968 (11♂ 1♀, CNC), TN: Nashville, G.G. Ainslie, Webster No. 9467C (1♂ [with puparium], USNM), VA: Chain Bridge, 14.v.1924, J.R. Malloch (1♀, USNM).

Comments

Agromyza parvicornis and A. proxima are highly similar in appearance and develop in some of the same plant genera, and can only be reliably differentiated on the basis of the relative width of the distiphallus and length of the basiphallus. It is possible that A. proxima represents a less common eastern morphological variant of the other, but in the absence of corroborating evidence, these two are maintained here as separate.

Agromyza proxima Spencer

Fig. 215

Agromyza proxima Spencer, 1969: 52. Spencer and Steyskal 1986b: 67.

Description

Wing length 2.2–2.4 mm (♂), 2.2–2.5 (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.7. Eye height divided by gena height: 2.8–3.4. First flagellomere small and rounded, sometimes with very small tuft of pale hairs apically. Scutum shiny.

Chaetotaxy : Two ori (sometimes three on one side); two ors. Two well-developed dorsocentrals with smaller third seta anteriorly. Hind tibia with two posteromedial setae.

Colouration : As described for A. parvicornis, except fronto-orbital plate usually brownish orange around base of all fronto-orbital setae.

Genitalia : (Fig. 215) Genitalia as described for A. parvicornis except as follows: basiphallus short, only slightly longer than distiphallus; distal section of distiphallus narrow, with sides only slightly bulging subapically.

Variation : MD male from Anne Arundel Co. with distal 2/5 of first flagellomere covered with pale hairs; genitalia as described above, but surstylus relatively small and spherical; distal 2/3 of first flagellomere dark and pedicel orange distally; frons dark brown. VA female with relatively large ovate distoventral patch of pale hairs on first flagellomere; body entirely dark with pedicel slightly paler, calypter entirely white, notopleuron and postpronotum reddish, and wing veins whitish on basal 1/2. The female is only tentatively allied with the MD male on the basis of similar dark colouration and the increased size of the pale haired patch on the first flagellomere.

Hosts

PoaceaeDichanthelium sp, Echinochloa walteri, Panicum dichotomiflorum (Scheffer and Lonsdale 2018, Spencer and Steyskal 1986b).

Distribution

Canada: MB. USA: DE*, FL, MA*, MD*, NY, VA(?).

Type material

Holotype: USA. FL: Sweetwater, Tamiami Canal (1♂, USNM).

Paratypes: Canada. MB: 2 mi N Forrest, Populus balsamifera stand around slough, 19.vii.1958, J.G. Chillcott, CNC352969 (1♂ [head missing], CNC). USA. FL: Sweetwater, 3.viii.1963, ex. Echinochloa walteri, leg. 23.vii.1963, K.A. Spencer (1♂ 1♀ [same pin, with puparia], USNM).

Additional material examined

USA. DE: Newark, 30.vii.1974 (1♂, UDCC), Stanton, 8.viii.1951, D.F. Bray (1♂, USNM), MA: Greenfield, 6.[?].1914, A.L. Melander (1♂, USNM), Wayland, 28.[?], J.J. Pratt, LT (1♂, USNM), MD: Bethseda, 12.ix.1981, G.C. Steyskal (1♀, USNM), Colesville, 15.viii.1975, Malaise trap, W.W. Wirth (1♂, USNM), Anne Arundel Co., nr. Edgewater, Smithsonian Envir. Res. Centre, 38°53'N, 76°33'W, 10.vii.1993, G.F. Hevel (1♂, USNM), VA: Sherando Lake, 10 mi SW Waynesboro, 25.vi.1970, L.V. Knutson (1♀, USNM).

Agromyza pudica Spencer

Figs 216–220

Agromyza pudica Spencer in Spencer and Steyskal 1986b: 267; Eiseman and Lonsdale 2018: 12, 2019: 3.

Description

Wing length 2.4–2.6 mm (♂), 2.0–2.6 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.7–0.9. Eye height divided by gena height: 4.8–4.9. First flagellomere small and ovate with small tuft of apical hairs in male. Ocellar triangle small and rounded (smaller in holotype). Scutum shiny.

Chaetotaxy : Two ori (sometimes three on one side); two ors. Three dorsocentral setae with anterior seta reduced. Mid tibia with two posteromedial setae.

Colouration : Body dark brown with orange tint; first flagellomere dark brown with faint orange tint towards base sometimes evident; lunule paler; parafacial sometimes with orange tint; halter white; tarsi and base of fore tibia yellow. Wing veins light brown. Calypter white with hairs usually brown (white in some females).

Genitalia : (Figs 216–220) Genitalia largely as described for A. parvicornis, except as follows: basiphallus long and well-sclerotised, halves overlapping at base, weakened point present past midpoint and apex split with dark ventral branch; distiphallus narrow, with broad medial constriction and with elongate ventral sulcus below distomedial ridge; ejaculatory apodeme small with blade pale.

Host

PoaceaeDichanthelium spp. (Eiseman and Lonsdale 2018, 2019).

Distribution

Canada: ON*. USA: AR, CT*, DC*, GA, MA, MD*, MN, NC, OH, OK, NY*, SC*, VA*.

Type material

Holotype: USA. AR: Garland Co., Hot Springs, 15.v.1979 (1♂, USNM).

Paratypes examined

USA. GA: Rabun Co., Rabun Bald, 1280 m, 16.vii.1957, J.G. Chillcott, CNC352972 (1♀, CNC), Rabun Bald, 914 m, 14.vii.1957, J.G. Chillcott, CNC352971 (1♂, CNC), NC: Pisgah Forest, Looking Glass Pk., 19.vii.1957, “K.A. Spencer, col.”, W.R. Richards, CNC352970 (1♂, CNC).

Additional material examined

Canada. ON: Algonquin Pk., 45°50'0"N, 77°38'0"W, 19.vii.1991, J.R. Vockeroth, CNC353094 (1♂, CNC), Metcalfe, 2mi N, 15.v.1982, B.E. Cooper, CNC353095 (1♂, CNC), 10.vi.1982, CNC353097 (1♂, CNC), 25.v.1982, CNC353096 (1♂, CNC), Ottawa, damp second-growth forest in Acer-Betula wood, 27.vi.1989, J.R. Vockeroth, CNC353098 (1♂, CNC). USA. CT: Waterton, 5.vi.1931, A.L. Melander (1♀, USNM), Putnam Park, A.L. Melander, 18.vii.1939 (1♂, USNM), 20.vii.1939 (1♂, USNM), 24.vii.1939 (1♀, USNM), Redding, A.L. Melander, 26.vi.1929 (1♂, USNM), 1.vi.1929 (1♀, USNM), 10.vi.1929 (1♀, USNM), 8.vi.1930 (1♂ 2♀, USNM), 11.vi.1929 (1♀, USNM), 27.vii.1938 (1♀, USNM), DC: 11.vi.1926, J.M. Aldrich (2♂, USNM), Chain Bridge, 8.v.1928, J.M. Aldrich (1♂, USNM), Washington, 17.viii.1913, A.L. Melander (2♂, USNM), MA: Concord, 17.vi.1961, W.W. Wirth (1♂, USNM), Franklin Co., Northfield, 276 Old Wendell Rd., 4.vii.2016, C.S. Eiseman, Dichanthelium acuminatum ssp. fasciculatum, em. 21.vii.2016, #CSE2787, CNC654200–654202 (1♂ 2♀, CNC), MD: Bethseda, 30.v.1980, G.C. Steyskal (1♀, USNM), Plummers Isl., 3.vi.1914, R.C. Shannon (1♂, USNM), Montgomery Co., 4mi S of Ashton, 24.vii.1982, G.F. and J.F. Hevel (2♀, USNM), NC: Scotland Co., Laurinburg, St. Andrews University, 18.v.2016, T.S. Feldman, Dichanthelium em. 13.vi.2016 , #CSE2572, CNC634813–634815 (3♂, CNC), Durham Co., Durham, 17-acre Wood Preserve, 15.v.2016, T.S. Feldman, Dichanthelium clandestinum, em. 4.vi.2016, #CSE2542, CNC654303–654306 (4♂, CNC), Scotland Co., Laurinburg, St. Andrews University, 11.v.2016, T.S. Feldman, Dichanthelium scoparium, em. 3.vi.2016, #CSE2540, CNC653948, CNC653949 (1♂ 1♀, CNC), NY: Lk. George, 26.vii.1929, A.L. Melander (1♀, USNM), Yonkers, 3.vi.1927, A.L. Melander (1♀, USNM), 5.vi.1927 (1♀, USNM), Peekskill, 11.vi.1927, A.L. Melander (1♂, USNM), Bear Mt., 31.v.1941, A.L. Melander (1♂, USNM), OH: Hocking Co., South Bloomingville, Deep Woods Farm, 5.viii.2016, C.S. Eiseman, Dichanthelium clandestinum, em. 20–22.viii.2016, #CSE2925, CNC654480–654482 (1♂ 2♀, CNC), SC: Beaufort Co., Fripp Island, 26.ix.1973, G.C. Steyskal (1♂, USNM), Highland Fall, 6.vii.1941, A.L. Melander (1♀, USNM), Cobleskill, 11.viii.1970, L. Knutson (1♀, USNM), VA: Glencarlyn, 23.v.1925, J.R. Malloch (1♂, USNM), Falls Church, Holmes Run, 24.viii.1960, light trap, W.W. Wirth (1♀, USNM), Rosslyn, 25.vi.1913, R.C. Shannon (1♀, USNM), Fairfax Co., Great Falls Park, swamp trail, 38°59.4'N, 77°15.2'W, Malaise trap, trap #1, 3–7.v.2007, D.R. Smith (1♂, USNM).

Comments

Agromyza pudica is recorded here in Canada, D.C., and five states for the first time.

Agromyza soka Eiseman & Lonsdale

Figs 221–224

Agromyza soka Eiseman & Lonsdale, 2018: 13.

Description

(from Eiseman and Lonsdale 2018). Wing length 2.3–2.5 mm (♂), 2.3–3.0 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.7. Eye height divided by gena height: 6.9–9.2. Ocellar triangle relatively small and rounded. First flagellomere small and nearly circular or slightly longer than high, with nearly indistinct apical tuft of pale hairs. Notum pruinose.

Chaetotaxy : Two or three ori (anterior seta small if present); two ors. Ocellar and postvertical setae subequal to outer vertical seta. Five dorsocentrals, strongly decreasing in length anteriorly. Eight irregular rows of acrostichal setulae. Mid tibia with one (male paratype) or two posteromedial setae.

Colouration : Setae dark brown with light brown reflection. Body predominantly dark brown (female darker) with light pruinosity; antenna dirty orange with distal 1/2 of first flagellomere infuscated (more so dorsally) in holotype, entirely brown in paratypes, with antenna entirely dark brown in female; frontal vitta, gena and postgena paler; apices of fore or all femora narrowly yellow; tarsi yellow; fore tibia light brown, fading to yellow at base. Calypter white with hairs brown. Halter white.

Genitalia : (Figs 221–224) Surstylus not distinct from epandrium, barely visible laterally, flat on inner surface and with four large spines. Cerci narrow and convergent. Hypandrium broad with thick arch, small apical process, and large inner lobe with two distal setae and several minute basal pits. Postgonite lobate and downturned. Proepiphallus and metepiphallus strongly reduced, flattened. Phallophorus elongate on left side. Halves of basiphallus strongly diverging from, and partially fused to phallophorus; lateromedially with lightly sclerotised membranous lobe; apex folded inwards, with pointed basal process and elongate distal process that is fused to mesophallus. Hypophallus broad, flat, and heavily sclerotised; apically split in dissected NC male. Mesophallus cylindrical, dark, basally rounded, slightly longer than wide, fused to distiphallus. Distiphallus broad, black, ventrally tilted and cup-like with constricted opening enclosing haired inner process. Ejaculatory apodeme well-developed with blade paler, no medial rib evident.

Host

FabaceaeRobinia pseudoacacia, Wisteria floribunda.

Distribution

USA: CT, NC, VA.

Type material

Holotype: USA. VA: nr. Plummers Isl., 20.v.1914, R.C. Shannon (1♂, USNM).

Paratypes: USA. CT: Hartford Co., East Hartford, Two Rivers Magnet Middle School, 4.vi.2016, em. 27–29.iv.2017, C.S. Eiseman, ex Robinia pseudoacacia, #CSE3574, CNC939943–939946 (3♂ 1♀, CNC), NC: Scotland Co., Laurinburg, St. Andrews University, 24.iv.2015, em. 16–18.iii.2016, T.S. Feldman, ex Robinia pseudoacacia, #CSE2248, CNC653954, CNC653955 (1♂ 1♀, CNC), 10.iv.2017, Robinia pseudoacacia, em. 22.iv.2018, #CSE4423, CNC1135665 (1♂, CNC), 4.iv.2016, em. 18.iv– 3.v.2017, T.S. Feldman, ex Wisteria floribunda, #CSE3518, CNC939744–939747 (1♂ 3♀, CNC).

Agromyza tacita Spencer

Figs 225–231

Agromyza tacita Spencer, 1969: 59. Spencer and Steyskal 1986b: 66.

Description

(Fig. 225). Externally as described for A. bispinata except as follows: Wing length 2.3–3.3 mm (♂). Length of ultimate section of vein M4 divided by penultimate section: 0.5–0.8. Eye height divided by gena height: 2.7–4.8. Small additional ori present in male with five spines on surstylus. First flagellomere sometimes paler with orange tint or predominantly orange with distal 1/2 infuscated; always enlarged in male. Postpronotum sometimes light brown or with orange tint. Wing veins brown to yellow. Fore tibia of MD male yellow with centre brownish.

Genitalia : (Figs 226–231) Surstylus fused to epandrium, easily viewed laterally, with dark anterior lobe and dark pronounced protuberances at base of marginal spines (three to five in number). Hypandrium, postgonite and ejaculatory apodeme as described for A. bispinata. Left sclerite of basiphallus overlapping base of right sclerite, which is fused to phallophorus; both halves with lateromedial membranous projection, and with weak connection to incurved distal section. Distiphallus of “nigripes-type” (capsule-shaped with subbasal opening for entry of ejaculatory duct, pronounced dorsobasal collar, and medial convolution); ventral curve with truncated sclerotised elbow; distal section with bulging, internally spinulose medial section.

Variation : Dissected paratype with two spines on surstylus (as in A. bispinata), but surstylus and remaining genitalia otherwise identical to holotype.

Variation : female paratypes: Wing length 2.9–3.1 mm. Length of ultimate section of vein M4 divided by penultimate section: 0.4. Eye height divided by gena height: 8.7–11.8. First flagellomere smaller, ovate; base always orange, reminder orange to brown. Anterior 1/2 of frons sometimes orangish.

Host

Unknown – likely Poaceae (Spencer and Steyskal 1986b).

Distribution

Canada: AB*, MB[?], NB*, ON, QC. USA: MD*, MO, NH*, NY*, UT*, VA*.

Type material

Holotype: Canada. ON: Ottawa, 17.vi.1946, G.E. Shewell, CNC352973 (1♂, CNC).

Paratypes: Canada. MB: Int. Peace Gardens, Turtle Mtn. For. Res., 7.viii.1958, J.G. Chillcott, CNC Type No. 10355, CNC352975 (1♀, CNC), ON: Ottawa, 15.vii.1957, J.E.H. Martin, CNC Type No. 10355, CNC352977 (1♀, CNC), 20.vi.1957, J.G. Chillcott, CNC Type No. 10355, CNC352974 (1♂, CNC), QC: Hull, 5.vi.1959, J.R. Vockeroth, CNC Type No. 10355, CNC352976 (1♀, CNC), Old Chelsea, summit of King Mt., 350 m, 11.vi.1959, J.R. Vockeroth, CNC Type No. 10355, CNC352978 (1♀, CNC).

Additional material examined

Canada. AB: Elkwater L., 21.vii.1956, O. Peck, CNC353101 (1♂, CNC), NB: Kouchibouguac N.P., 11.vii.1977, J.F. McAlpine, Code-6035C, CNC352990–352992 (3♂, CNC), 12.vii.1977, J.F. McAlpine, Code-6040H, CNC352989 (1♂, CNC), 13.vii.1977, G.A. Calderwood, Code-5599I, CNC352993 (1♂, CNC), 30.vi.1977, J.R. Vockeroth, Code-5456V, CNC352986, CNC352987, CNC352981–352985 (7♂, CNC), 6.vii.1977, G.A. Calderwood, Code-54900, CNC352988 (1♂, CNC), 9.vii.1977, J.F. McAlpine, Code 6024R, Code-6024R, CNC352979, CNC352980 (2♂, CNC), ON: Ottawa, on leaf of Philadelphus coronarius, 13.vii.1959, J.R. Vockeroth, CNC353100 (1♂, CNC). USA. MD: Plummers Isl., 14.vi.1913, R.C. Shannon (1♂, USNM), NH: Jefferson Notch, 30.vii.1961, creek margin, W.W. Wirth (1♂, USNM), NY: Bear Mt., 31.v.1941, A.L. Melander (1♂, USNM), UT: Cache Co., Green Canyon, 23.vii.1964, Malaise trap, W.J. Hanson (1♂, USNM), VA: Shenandoah, Big Meadows, 3.vii.1939, A.L. Melander (1♂, USNM), Fairfax Co., Dead Run, 9.vi.1915, R.C. Shannon (1♂, USNM).

Comments

See comments for Agromyza bispinata. The female paratypes of A. tacita cannot be confidently assigned to this species, which is mostly defined by male genitalic characters, and their placement should be re-evaluated when the boundaries of species in the A. bispinata complex can be better delimited.

Agromyza varifrons Coquillett

Figs 236–242

Agromyza varifrons Coquillett, 1902: 189. Frick 1952a: 374; Spencer 1969: 59; Spencer and Steyskal 1986b: 269; Scheffer and Lonsdale 2018: 85.

Description

Wing length 1.7–2.2 mm (♂), 1.8–2.3 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.9–1.0. Eye height divided by gena height: 3.5–10.0. Arista pubescent. First flagellomere small and rounded, without pale tuft of hairs. Fronto-orbital plate slightly projecting (more so anteriorly). Ocellar triangle small and rounded.

Chaetotaxy : Two ori; two ors. Orbital setulae in one row. Ocellar and postvertical setae subequal to fronto-orbitals. Two strong dorsocentral setae, with second seta ~ ¾ length of first seta; much weaker third dorsocentral not much larger than acrostichal seta. Acrostichal seta ~ 2 × length of setulae. Acrostichal setulae in 8 scattered rows, tapering posteriorly to level of first dorsocentral. Mid tibiae without posteromedial setae.

Colouration : Setae dark brown. Antenna yellow to orange; first flagellomere darker orange with basal margin paler and with small brown curved spot around base of arista posteriorly and along outer surface. Frons yellow with posterolateral corner and posterior margin dark brown; orbital plate darker, brownish orange behind anterior ors, becoming darker posteriorly, and with narrow brownish line continuing anteriorly along eye margin beside ori; ocellar spot rounded, dark brown, slightly larger than tubercle; remaining posterior 1/2 or less of frontal vitta brownish orange, becoming darker posteriorly, sometimes with this dark region restricted to large circle around ocellar spot. Lunule light yellow. Gena yellow with ventral brownish line. Parafacial yellow. Face whitish yellow. Mouthparts yellow; clypeus with brown tint that is darker laterally. Postgena, back of head and remainder of body mostly dark brown; body subshiny. Apex of fore femur yellowish orange for length equal to width of femur apex; apex of mid femur narrowly paler brown apically; tibiae sometimes paler brown with base and apex yellowish, and mid tibia sometimes with yellow mottling medially; tarsi yellow, becoming more brown tinted on posterior legs; wing veins brown; halter yellow with orange-brown tint on stem; calypter margin and hairs brown.

Variation : Posterior 1/2 of frons brownish in holotype and FL specimen. IL male with first flagellomere entirely light yellow, head brighter and scutum shiny.

Genitalia : (Figs 236–242) Epandrium as long as high; fused to small, inwardly directed surstylus. Cercus large, well-developed, venter of inner surface covered with tubercle-like setae. Hypandrium with large apical apodeme and narrow basal arms; medially with weakly sclerotised region connected to L-shaped inner lobe bearing row of minute setae. Postgonite in lateral view with short, flat globose body with extremely long tail; inner surface with apically setose shelf. Phallophorus elongate cylindrical, apicodorsal margin irregularly sclerotised to base of basiphallus. Basiphallus with small left basolateral plate partially fused to large dorsal plate that is strongly curved ventrally at base and with medial and apical lobes that wrap along right lateral surface of shaft. Ejaculatory duct widening to mesophallus, which is round, bulbous, and clear with thick triangular sclerotisation ventrally. Distiphallus composed of one pair of upturned “wings” that nearly meet ventrally and are very narrowly connected dorsally; each wing with small, dark, smooth base expanding into densely haired, distally pointed lobes. Ejaculatory apodeme weakly sclerotised, with broad slanted base and short stem grading into thick blade with faint apical striations; sperm pump with slight basal sclerotisation.

Hosts

CannabaceaeCeltis laevigata. Possibly Ulmus Americana (Ulmaceae) (Eiseman 2021).

Distribution

Canada: ON, QC. USA: AK, DC, FL, IA, IL, KS, MS, NY, PA, TX, VA*.

Type material

Holotype: USA. DC: “District of Columbia” (1♀, USNM).

Additional material examined

USA. FL: Hialeah, em. 10.v.1963, mine Celtis laevigata, 22.ix.1963 (1♂ 1♀ [same pin, with puparia]), Gainesville, Newman’s Lake, 26.iv.1952, O. Peck, CNC352994 (1♂, CNC), Hialeah, 22.ix.1963, mine Celtis laevigata, Em. 5.x.1963, [K.A. Spencer], CNC352996 (1♂ 1♀ [with puparia], CNC), IL: [illegible], 15.v.1919 (1♀, USNM), NY: Suffolk Co., Stony Brook Village, larva coll. 19.vi.1993, S.J. Scheffer, 93-187 (1♂[illustrated], USNM) , eclosed 9.vii.1993 (1♂, USNM), eclosed 8.vii.1993 (1♀, USNM), eclosed 10.vii.1993 (3♂, USNM), eclosed 11.vii.1993 (1♀, USNM), elcosed 17.vii.1993 (1♂, USNM), TX: Kerrville, 4.iv.1959, J.F. McAlpine, CNC352995 (1♂, CNC), Welder Wildlife Ref. nr Sinton, 19–23.iii.1965, J.G. Chillcott, CNC352997 (1♀, CNC), VA: Fairfax Co., Turkey Run Park, nr. mouth of Turkey Run, 38°57.9'N, 7°09.4'W, Malaise trap, 26.iv-2.v.2007, D.R. Smith (3♀, USNM), Turkey Run Park, 0.3 km W mouth Turkey Run, 38°58'N, 77°09.6'W, Malaise trap, D.R. Smith, 18–30.v.2007 (1♀, USNM), 29.iii-25.iv.2007 (1♂, USNM), Great Falls Park, quarry, 38°59.1'N, 77°14.8'W, Malaise trap, D.R. Smith, 24.iv-2.v.2007 (3♂ 1♀, USNM), 10–17.v.2007 (1♂, USNM), 3–10.v.2007 (1♂, USNM).

Comments

This species is clearly allied to Agromyza aristata, being similar in external and male genitalic morphology. The anterior portion of the head is mostly pale, the cercus is covered with small tubercle-like setae along the anterior surface, and the mesophallus is a ventral band separate from a curved, plate-like distiphallus. The most immediate difference in the genitalia is the densely haired distiphallus and short mesophallus, but the hypophallus has an apical apodeme, the basiphallus is also strongly curved and irregular in outline, the postgonite broadly lobate (lateral view) and the ejaculatory apodeme is well-developed. Externally, the pale portions of A. aristata are light yellow, not orange to yellowish, and the legs are extensively yellow.

Agromyza virginiensis Spencer

Fig. 151

Agromyza virginiensis Spencer, 1977: 237. Spencer and Steyskal 1986b: 64.

Description

Wing length 3.2 mm. Length of ultimate section of vein M4 divided by penultimate section: 0.6. Eye height divided by gena height: 3.4. First flagellomere longer than wide with rounded apex; apical margin possibly with longer, paler hairs. Fronto-orbital plate and parafacial slightly projecting. Ocellar triangle broadly rounded and short, not extending much past ocelli. Buccal cavity subquadrate.

Chaetotaxy : Four ori; one ors (not two as reported by previous authors). Four postsutural dorsocentrals, decreasing in length anteriorly. Mid tibia with two posteromedial setae.

Colouration : Body predominantly brown with halter white. Gena (excluding ventral margin), parafacial and frons (excluding posterolateral corner, posterior margin and ocellar triangle) light brownish orange, with fronto-orbital plate lightly pigmented posteriorly to base of posterior ori. First flagellomere, scape and pedicel orange, sometimes with dark spot at base of arista. Lunule orange. Face brown. Palpus and clypeus dark brown. Calypter margin and hairs white. Legs dark brown with knees yellow and tibiae and tarsi with orange tint (brighter on tarsi).

Genitalia : (Fig. 151) Surstylus triangular with basal suture obliterated; with minute apical setae and medial tubercle-like setae. Hypandrium relatively long and narrow with minute apical point. Postgonite very broad, flat, and lobate. Halves of basiphallus long with ventral margin weakly defined, apices hooked, and right sclerite with broader base. Distiphallus very long, cylindrical, dark and with distal 2/5 bifid and not strongly curved. Ejaculatory apodeme small and thin, without apical blade.

Variation : Canadian material differs as follows: wing length 3.0–3.2 mm (♂), 3.4–3.8 mm (♀); eye height divided by gena height 5.1–10.6; first flagellomere with relatively discrete ovate pilose patch; 2 ors and 2–3 ori; first flagellomere brownish on distal and dorsal margins anterior to arista base; posterior 1/3 of frons much darker; fronto-orbital plate with ill-defined brown margin along length; parafacial dark brown; distiphallus only split on distal 1/10.

Host

Unknown – adult collected on Phyllocarpus (Fabaceae).

Distribution

Canada: ON*. USA: VA.

Type material

Holotype: USA. VA: Great Falls, vi.1922, Banks (1♂, MCZ).

Additional material examined

Canada. ON: Orleans, Chapel Hill, at flowers of Phyllocarpus, 23.vi.1994, J.R. Vockeroth, CNC353052 (1♀, CNC), Ottawa, Damp second-growth Acer-Betula woods, 12.vii.1994, J.R. Vockeroth, CNC353053 (1♀, CNC), 15.vi.1998, CNC353059 (1♀, CNC), 16.viii.1992, CNC353046 (1♂, CNC), 17.vi.2003, CNC353056 (1♀, CNC), 2.vii.1993, CNC353044 (1♂, CNC), 24.vii.2000, CNC353049 (1♂, CNC), 29.vi.1994, CNC353058 (1♀, CNC), 29.vi.1997, CNC353047 (1♂, CNC), 29.vii.1992, CNC353045 (1♂, CNC), 30.vi.1991, CNC353054 (1♀, CNC), 30.vii.1993, CNC353051 (1♀, CNC), 6.viii.1993, CNC353043, CNC353057 (1♂,1♀, CNC), 7.viii.1993, CNC353050 (1♂, CNC), 7.viii.2000, CNC353055 (1♀, CNC), 9.vii.2000, CNC353048 (1♂, CNC).

Comments

The Canadian material varies slightly from the holotype, being more similar in external appearance to Agromyza ambrosivora, although the straight (not strongly curled) distiphallus would seem to preclude this option, and the anterior margin of the buccal cavity is relatively straight (more pointed in A. ambrosivora). The phallus of these Canadian specimens further differs from the holotype in that only the apex of the distiphallus is cleft, but these external and phallic differences will be tentatively treated as within-species variation until additional specimens can be measured.

The phallus is similar to that of a Nearctic and European leaf miner on Urtica (Urticaceae), Agromyza reptans (Spencer and Steyskal 1986b: figs 395, 396). The distiphallus of both is straight and apically split, but that of A. reptans is much shorter, paler, and with the apical split represented by one pair of separate, darker cylindrical arms (not the ends of a single, continuous, narrow tube), barely longer than wide and stouter than the base.

Agromyza vockerothi Spencer

Figs 243–249

Agromyza vockerothi Spencer, 1969: 60. Spencer and Steyskal 1986b: 269; Eiseman and Lonsdale 2018: 14.

Description

Externally as described for A. isolata except as follows: Wing length 2.4–2.7 mm (♂), 2.4 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.8. Eye height divided by gena height: 3.8–6.5. Sometimes three ori on one side of frons. Overall colour slightly darker, with antenna dark brown and calypter margin and hairs always brown; mid and hind tibiae brown with apex and sometimes base yellowish. First flagellomere sometimes appearing slightly angulate and with some longer apical hairs.

Genitalia : (Figs 243–249) Surstylus not distinct from epandrium (which has desclerotised distal emargination), and with several setae and tubercle-like setae on inner margin. Hypandrium long and narrow, sides nearly parallel and with tapered apical process. Postgonite small, dome-like with several outer setulae. Phallophorus atrophied on right side, produced and slightly setulose on left side, fused to right sclerite of basiphallus, twisting around shaft; left sclerite of basiphallus narrow, curving around venter at base. Basiphallus broad and fringed basally with dextral twist. Hypophallus composed of weak, long narrow band. Mesophallus short, cylindrical, with longer ventral plate. Distiphallus short, subcylindrical with ventral break, and fringed along distal and ventral margins.

Host

RosaceaeRubus.

Distribution

Canada: AB, BC, ON, NS. USA: CT*, DC*, MA, MD*, NC, NH*, NY*, PA*, TN, VA*.

Type material

Holotype: Canada. NS: Shelburne, 10.viii.1958, J.R. Vockeroth, Type No. 10356, CNC352998 (1♂, CNC).

Paratypes examined

ON: Maynooth, 22.vi.1953, J.F. McAlpine, CNC353000 (1♂, CNC), Ottawa, 5.vi.1946, G.E. Shewell, CNC353001, CNC353002 (2♂, CNC), Golden Lake, 3.ix.1956, J.R. Vockeroth, CNC353003 (1♀, CNC), QC: Old Chelsea, 18.viii.1961, J.R. Vockeroth, CNC352999 (1♂, CNC).

Material examined

Canada. BC: 20 km E Pemberton, 14.viii.1991, A. Borkent, CNC353014, CNC353015 (2♂, CNC), Hagensborg, 12.vii.1992, A. Borkent, CNC353016 (1♂, CNC), 42 km NE Hope, 2.viii.1991, A. Borkent, CNC353015 (1♂, CNC), 32 mi SW of Terrace, 9.vii.1960, J.G. Chillcott, CNC353017 (1♂, CNC), ON: St. Lawrence Is. Nat. Park, Aubrey Island, 4.ix.1976, W. Reid, Code 4618-R, CNC353006 (1♂, CNC), St. Lawrence Is. Nat. Park, Cedar Island, 31.viii.1976, W. Reid, Code 4586-L, CNC353007 (1♀, CNC), St. Lawrence Is. Nat. Park, McDonald Is., 14.vi.1976, 6.ix.1976, A. Carter, W. Reid, Code 4092-J, Code 4633-G, CNC353005, CNC353008 (1♂ 1♀, CNC), St. Lawrence Is. Nat. Park, Thwartway Is., 24.vii.1976, W. Reid, Code 4198-L, CNC353009 (1♀, CNC). USA. CT: Kent Fall, 28.viii.1940, A.L. Melander (1♂, USNM), Redding, 28.v.1939, A.L. Melander (1♂, USNM), DC: Washington, “v-17”, J.M. Aldrich (1?, USNM), MA: Franklin Co., Northfield, Crag Mountain, 11.x.2013, ex. Rubus (blackberry), em. 27.iii.2014, C.S. Eiseman, #CSE1030, CNC384794 (1♂, CNC), MD: Glen Echo, 21.viii.1921, J.R. Malloch (1♂, USNM), Colesville, 1.v.1977, W.W. Wirth (1♀, USNM), Montgomery Co., Dickerson, 14.vii.1974, G.A. Foster (1♀, USNM), NC: Highlands, Wilson’s Gap, 944 m, 12.v.1957, J.R. Vockeroth, CNC353011 (1♀, CNC), Mitchell Co., Roan Mtn, 1889 m, 13.viii.1957, J.G. Chillcott, CNC353010 (1♀, CNC), Wyah Bald, 13.vi.1957, C.J. Durden, CNC353012 (1♀, CNC), NH: White Mts., Stinson Lake, 23.vii.1961, W.W. Wirth (1♂, USNM), NY: Allegany St. Pk., 28.v-3.vi.1963, stream margin, W.W. Wirth (1♂, USNM), Orleans Co., Albion, 11.vi.1963, Burma woods, W.W. Wirth (1♂, USNM), Monroe Co., Braddock Bay, 12.vi.1963, near marsh, W.W. Wirth (1♂, USNM), PA: Mineral Spr., 5.ix.1927, A.L. Melander (2♂, USNM), Monroe Co., Mt. Pocono, 28.vii.1985, G.F. and J.F. Hevel (1♂, USNM), TN: Smokies, Chimneys, A.L. Melander, 25.vi.1941 (1♀, USNM), 20.vi.1941 (1♂, USNM), Indian Gap to Clingman’s Dome, 5200–6600', 6.viii.1957, J.G. Chillcott, CNC353004 (1♂, CNC), VA: Luray, 24.vi.1923, A.L. Melander (1♂, USNM), Fairfax Co., Turkey Run Park, nr. mouth of Turkey Run, 38°57.9'N, 7°09.4'W, Malaise trap, river trap, 17–24.v.2006, D.R. Smith (1♀, USNM).

Euhexomyza Lonsdale

Euhexomyza Lonsdale, 2014: 497. Type species Melanagromyza simplicoidesHendel 1920, by original designation.

Euhexomyza was described by Lonsdale (2014) for a lineage of mostly north-temperate Salicaceae-feeding species where the larva forms an ovate gall in the cortex of a twig. These species were previously treated as part of Hexomyza Enderlein, but Lonsdale (2014) found the genus to be a polyphyletic dumping ground for a miscellany of gall-forming species, with the morphologically disparate type species belonging to Ophiomyia. He subsequently included the name Hexomyza as a junior synonym of Ophiomyia.

Species are brown and non-metallic (Fig. 44), they have a rounded clypeus and lack a prescutellar acrostichal. The fronto-orbital plate and parafacial are pronounced, forming a strong ring around the eye, the postsutural intra-alars are subequal (anterior seta never strong), and only the mid tibia sometimes has a lateromedial seta. The male genitalia (at least of the north temperate Salicaceae-feeders) are typified by a distiphallus that has the base broad and rounded and the apex tapered, and the phallophorus has a unique ventral “pouch” emerging from the distal margin (Fig. 250; Lonsdale 2014: fig. 62).

Species description

Euhexomyza winnemanae (Malloch)

Figs 47, 48

Agromyza winnemanae Malloch, 1913a: 314. Shewell 1953: 465.

Melanagromyza winnemanae. Frick, 1952a: 380, 1959: 367.

Hexomyza winnemanae. Spencer & Steyskal, 1986b: 250.

Euhexomyza winnemanae. Lonsdale, 2014: 503.

Description

(Figs 47, 48). Wing length 1.9 mm (♂♀). Length of ultimate section of vein M4 divided by penultimate section: 1.2. Eye height divided by gena height: 8.6. Ocellar triangle large, glossy outside of tubercle and sharply pointed anteriorly. First flagellomere broadly rounded. Clypeus broadly rounded anteriorly. Distance between crossveins as long as dm-m. Costa not extending much past R4+5. Lunule broad, very shallow. Base of antennae broadly separated. Face flat. Fronto-orbital plate narrow.

Chaetotaxy : One strongly incurved ori; three ors; setae decreasing in width and length anteriorly. Ocellar and postocellar setae well-developed. Orbital setulae reclinate. Two well-developed dorsocentrals. Acrostichal setulae in eight rows. Posteromedial tibial setae absent.

Colouration : Calypter and wing entirely white with veins brownish (costa darker). Halter brown. Body entirely brown to dark brown with gena and centre of frons paler.

Genitalia : Unknown.

Host

Unknown – likely Salix (Salicaceae).

Distribution

Canada: ON. USA: MD. Gall attributed to this species observed in AB (Spencer, 1969).

Type material

Holotype [winnemanae]: USA. MD: Plummers Island, 27.vi.1909, W.L. McAtee (1♀, USNM).

Holotype [albicula]: Canada. ON: Ottawa, 7.iv.1948, ex. gall on Salix sp., G.E. Shewell (1♂, CNC).

Paratype [albicula]: Canada. ON: Dryden, 12–13.vi.1960, Kelton and Whitney (1♀, CNC).

Comments

Euhexomyza albicula (Spencer) stat. reinst., comb. nov. (Figs 45, 46, 250) was included as a junior synonym of H. winnemanae by Spencer and Steyskal (1986b), but following examination of the types of both species, both are recognised here as distinct, and the former is resurrected from synonymy. Aside from having white wings and a similar body size, the two species are strikingly dissimilar. Euhexomyza albicula differs as follows: eye height divided by gena height 2.0–2.1; eye relatively small and broadly surrounded by fronto-orbital plate, parafacial and occiput when seen laterally; fronto-orbital plate broad and bulging (more so medially); two ors, two ori (not much longer than ocellar seta); ocellar triangle broad, rounded apically and subshiny; first flagellomere smaller, ovate and with longer hairs; lunule semi-circular and bulging; clypeus broader medially; ocellar and postocellar setae not much longer than ocellar tubercle; scutellum with second small pair of distolateral scutellar setae well-developed and with additional seta on left side in female; veins white, excluding costa, subcosta and base of Rs; calypter margin and hairs brown; body subshiny and light brown to brown (not dark brown).

One additional pair of lateral scutellar setae is also present in the northeastern North American species Euhexomyza salicis (Malloch), which has the costa extending to vein M1+2 (not R4+5).

Japanagromyza Sasakawa

Japanagromyza Sasakawa, 1958: 138. Type species: Agromyza duchesneae Sasakawa, 1954, by original designation. Spencer 1969: 62; Spencer and Steyskal 1986b: 53; Lonsdale 2013: 445.

Geratomyza Spencer in Spencer and Stegmaier 1973: 140. Spencer 1984 [synonymy].

Japanagromyza is species-poor in North America north of Florida, but the species J. viridula (Coquillett) is relatively abundant from the east coast to Alberta, Kansas and Arizona, and the unusual J. rutiliceps (Melander) is known from Montana to California. The genus, which was redefined in Lonsdale (2013), has one pair of acrostichal setae and/or a lateromedial seta on the mid tibia. Agromyza also has developed acrostichal setae, but it differs in having a stridulatory file. Those Japanagromyza without acrostichals are superficially similar to Melanagromyza, but species in the latter genus have a longer ocellar triangle, more pronounced fronto-orbital plates, and often dorsally setose eyes.

The majority of species occurring in the New World can be characterised as follows: colour black (or with iridescence); only two pairs of widely spaced dorsocentrals; knob of halter white (base of knob and stem sometimes variably brown, and occasionally entirely dark, as in J. brooksi Spencer from Ontario); fore tibia with lateromedial seta (absent in J. brooksi and a number of non-Nearctic species); surstylus often long and narrow; male cercus very large and often bearing large tubercle-like setae (similar structures are found in some Agromyza); hypandrium with long, narrow, ventrally curving apodeme apically; phallus long, narrow and clear (morphology much more varied globally), and with ejaculatory apodeme sometimes very thin or atrophied. The unusual J. rutiliceps (Melander) from the western United States differs in most of these features, and its placement is uncertain.

Japagromyza viridula (Coquillett)

Figs 49–51, 251–256

Agromyza viridula Coquillett, 1902: 189. Frick 1952a: 374, 1959: 359.

Japanagromyza viridula. Spencer, 1966a: 3, 1969: 63; Spencer and Steyskal 1986b: 55; Scheffer et al. 2007: 770; Eiseman and Lonsdale 2018: 16.

Description

(Figs 49–51). Wing length 2.7–3.2 mm (♂), 3.1–3.2 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.7. Eye height divided by gena height: 8.0–8.3. Ocellar triangle faintly outlined. Fronto-orbital plate and ocellar tubercle subshiny. Lunule height 1/2 width, silvery pruinose; lunule and face broad and well-sclerotised with face flat. Clypeus broadly rounded. Notum and head with light dusting of pruinosity. Arista virtually bare.

Chaetotaxy : All setae well-developed and setulae short and sparse. Two ori; two ors (widely separated). Two postsutural dorsocentrals. Ten rows of acrostichal setulae; acrostichal setula present. Postocellar and ocellar setae well-developed. Fore tibia with one lateral seta; mid tibia with two posteromedial setae.

Colouration : Setae black (setulae on ocellar tubercle sometimes golden-brown). Body predominantly dark brown; thorax (less so on pleuron) and abdomen with metallic green reflection, and epandrium faintly to distinctly reddish brown. Wing veins brown. Calypter and halter white.

Genitalia : (Figs 251–256) Surstylus fused to epandrium; suture obliterated; very narrow, apex rounded, nearly as high as epandrium; inner surface with patches of numerous tubercle-like setae distally and basally; twisted, inner face visible posteriorly. Hypandrium large, elongate with very long, stout, apical process; inner lobe bare, narrow and transverse. Postgonite broad, ovate, flat, and bare. Phallophorus small, fused to basiphallus. Basiphallus a single plate dorsal to right lateral. Hypophallus slightly shorter than basiphallus, dark, linear. Distiphallus very long, pale, and relatively straight, composed of longer flagellum on left side and shorter flagellum on right side, connected by thin membrane; often longer than illustrated. Mesophallus indistinct. Ejaculatory apodeme narrow, blade reduced with very shallow basolateral process; sperm pump unsclerotised with minutely spinulose ventral surface.

Hosts

FagaceaeQuercus spp., Castanea mollissima (Eiseman and Lonsdale 2018).

Distribution

Canada: NB*, NS, ON, QC*. USA: DC, GA, IN, KS, MA, ME, NC, OK, PA, SC, TN, VA. Puerto Rico.

Type material

Holotype: USA. DC: “June”, “Collection Coquillett” (1♂, USNM; Type No. 6660).

Paratypes examined

USA. DC: “DC” (1♀, USNM), MA: Beverly, 29.vi.1876, Burgess (1♀, USNM). Puerto Rico. “Porto Rico”, Aguadilla, i.1899, A. Busck (1♀, USNM).

Additional material examined

142♂ 97♀ 1?, CNC, USNM. Canada. NB: Kouchibouguac N.P., 8.vi.1977, J.R. Vockeroth, Code – 5222V, Code – 52336, CNC358580, CNC358581, CNC358582 (3♀, CNC), 9.vii.1977, J.F. McAlpine, Code – 6023Q, CNC358583 (1♀, CNC), QC: Rigaud, 11.vi.1981, J.R. Vockeroth, CNC358579 (1♂, CNC).

Comments

Superficially, Japanagromyza viridula can be mistaken for species of Melanagromyza because of its metallic green colouration, but it can be easily separated by the single pair of stout prescutellar acrostichal setae.

There was an error produced in the Japanagromyza key in Spencer and Stegmaier (1973) that was perpetuated in Spencer and Steyskal (1986b) and Wiegmann (1991), leaving the only Delmarva species unidentifiable. Japanagromyza viridula was differentiated from U.S. congeners with prescutellar acrostichal setae (J. inaequalis (Malloch), J. aequalis Spencer and J. polygonivora Wiegmann (= J. polygoni Spencer) by having the “mesonotum predominantly black”. Upon examination of the types of all of these species deposited in the USNM, however, there appears to be little, if any difference in the colour of the notum. Instead, J. viridula can be diagnosed by the arista, which is virtually bare (hairs not as long or longer than width of central filament) by examination of the male genitalia, or sometimes by the setulae on the ocellar tubercle, which are black or golden-brown (always black in the remaining species, which are only found in and south of Florida).

Melanagromyza Hendel

Melanagromyza Hendel, 1920: 114. Type species: Agromyza aeneoventris Fallén, 1823, by original designation. Frick 1952a: 375; Spencer 1969: 64; Spencer and Steyskal 1986b: 18; Lonsdale 2014: 495; Shi and Gaimari 2015: 10.

Limnoagromyza Malloch, 1920: 147. Type species: diantherae Malloch, 1920, by original designation. Frick 1952a [synonymy].

Many Melanagromyza have a noticeably metallic green, blue, or coppery sheen on the abdomen (not including the black oviscape), which is also often present on the notum, but this feature should not be used as diagnostic in and of itself as a metallic sheen is also found in a handful of other Agromyzinae. This sheen is usually very faint in these other taxa, but some Japanagromyza species, including the relatively abundant eastern species J. viridula, are strikingly green. Conversely, those Melanagromyza with little or no metallic colouration can be easily mistaken for some Euhexomyza and Ophiomyia, although dorsally setulose (not bare) eyes, a wider fronto-orbital plate and a broadly rounded clypeus will reveal their generic affiliation.

Externally, the Delmarva Melanagromyza are best diagnosed by an absence of characters found in other local Agromyzinae: there are no prescutellar acrostichals (present in Agromyza and Japanagromyza) or a stridulatory file on syntergite 1+2 (Agromyza); the clypeus is usually broadly rounded (apically truncated in all Ophiomyia), but if the anterior margin is straight, the lateral corners are rounded; the eye is usually setulose dorsally (always bare or very sparsely short-setulose in Ophiomyia); the gena is never strongly produced anteriorly, there is never a vibrissal fasciculus, and the facial keel is never prominent if present (many Ophiomyia).

The male genitalia are most characteristic of the genus: the basiphallus is short, symmetrical and U-shaped or ring-like; the epandrium sometimes has a small posterodistal spine behind the surstylus (not always visible in lateral view); the ventrobasal surface of the distiphallus has one pair of thin tubules flanking the mesophallus (unconfirmed in some non-Nearctic species).

Key to the Delmarva Melanagromyza

1 Calypter margin and hairs dark brown. Eye always bare or with sparse setulae 2
Calypter margin and hairs white. Eye almost always setulose dorsomedially 3
2 Fronto-orbital plate and parafacial broad and easily viewed laterally. Gena highest posteriorly. Four ori. Abdomen with light coppery tint. Wing length 2.8–3.6 mm. Basiphallus forming complete ring (Figs 269, 270). Distiphallus broad and stout M. burgessi (Malloch)
Fronto-orbital plate and parafacial narrow, inconspicuous laterally. Gena highest at midpoint. Two ori. Abdomen with green metallic sheen. Wing length 1.9–2.3 mm. Basiphallus U-shaped (Figs 291, 292). Distiphallus thin and gracile M. minimoides Spencer
3 Gena at its highest near anterior margin of eye (Figs 52, 53). Distiphallus with dark distolateral plate and prominent dorsomedial process M. buccalis Spencer
Gena highest near midpoint of eye. Distiphallus not as above 4
4 Four fronto-orbital setae; anterior 2 pairs (ori) very widely spaced (Fig. 57). Surstylus sometimes with elongate tubercle-like setae (Figs 318, 319). Proepiphallus narrow, straight and V-shaped. Basiphallus open ventrally M. virens group (5)
Usually more than 4 fronto-orbital setae, at least on one side; distance between anterior setae not much larger than distance between any other pair of setae. Tubercle-like setae on surstylus all short. Proepiphallus rounded laterally, upcurved anteriorly and usually with separate halves. Basiphallus sometimes forming a ring 10
5 Head (seen laterally) broadest near middle, i.e., without bulging frons. Fronto-orbital plate relatively narrow, slightly wider behind midpoint. Seen laterally, distiphallus gradually tapering apically and with 1 pair of posteromedial bulges/carinae dorsomedially (Fig. 287) M. matricarioides Spencer
Head (seen laterally) usually broadest dorsally, with fronto-orbital plate bulging anterodorsally. Fronto-orbital plate usually swollen medially and narrowed anteriorly. Distiphallus abruptly widened on basal 1/2 and without clear posteromedial bulge 6
6 Wing length 1.8–2.6 mm. Fronto-orbital plate often 1/3 or 1/4 width of frons (Fig. 56). Distiphallus relatively short and squat 7
Wing length 2.5–3.5 mm. Fronto-orbital plate swollen to only 1/4 width of frons (Fig. 54), never 1/3. Distiphallus more elongate 8
7 Fronto-orbital plate (seen dorsally) swollen medially, usually at least 1/3 width of frons (sometimes 1/4 width). Male orbital setulae conspicuously long and bushy compared to setulae on eyes (Fig. 57). Wing length 1.8–2.4 mm. Surstylus narrow and with elongate tubercle-like setae (Figs 318, 319). Mesophallus longer, held slightly away from distiphallus. Distiphallus slender apically (Figs 322, 323) M. virens (Loew)
Fronto-orbital plate swollen to only ¼ width of frons, never 1/3. Male orbital setulae slightly longer than ommatrichia, sometimes barely so, relatively sparse. Wing length 1.8–2.6 mm. Surstylus broader and with only short tubercle-like setae at apex (Fig. 297). Mesophallus slender, held close to distiphallus. Distiphallus bulging laterally along distal surface (Figs 299, 300) M. splendida Frick
8 Notum and abdomen with green shine that may sometimes also be partially coppery. Distal edge of surstylus ~ 1/3 height of epandrium. Basiphallus overlapping base of distiphallus (Figs 326, 327). Large, dark distoventral plate of distiphallus (viewed laterally) continuing past point of insertion of mesophallus M. virginiensis Spencer
Notum with green to bluish tint. Abdomen green (M. vernoniae bronze-green in IN and blue in MN). Distal edge of surstylus at least 1/2 height of epandrium. Basiphallus separate from distiphallus, if only slightly. Dark distoventral plate of distiphallus (viewed laterally) never continuing past point of insertion of mesophallus 9
9 Orbital setulae much longer than ommatrichia, relatively bushy. Notum with blue tint that may become greenish posteriorly (entirely green in IN). Distiphallus gradually narrowing apically; distal section straight (seen laterally); basal section smooth; with inner spinulose patch. Phallophorus longer than wide (Figs 306, 307) M. vernoniae Steyskal
Orbital setulae short, as long as ommatrichia. Notum with faint green tint. Abdomen shiny green. Distiphallus only wide near base, with remainder long and curved (seen laterally); basal section strongly convoluted; inner spinulose patch strongly reduced to absent. Phallophorus higher than long (Fig. 312) M. vernoniana Steyskal
10 Large and stout-bodied; wing length 3.0–5.0 mm. Three or 4 ori, rarely 2. Abdomen often with blue metallic shine, at least posteriorly. Distiphallus stout and broad. No space between apex of basiphallus and base of distiphallus 11
Smaller and more slender; wing length 2.8–3.4 mm. Two or 3 ori. Abdomen blue, green, or coppery. Distiphallus small and slender. Membranous space between basiphallus and base of distiphallus distinct 12
11 Ocellar tubercle with thin strip of setulae extending to base of postocellar. Eye bare. Three dorsocentral setae. First flagellomere as high as gena. Postpronotum with large orange spots anteriorly and laterally. Presutural scutum with faint, narrow pruinose strips medially. Distiphallus broad and rounded with dorsomedial process; internal surface minutely tuberculate; ventrolateral tubules stout and broadly looped; outer surface not sculptured (Figs 275, 276). Basiphallus broad medially, and thin and ill-defined laterally. Metepiphallus without spines M. diantherae (Malloch)
Setulae restricted to anterior 1/2 of ocellar tubercle. Eye setulose. Two or 3 dorsocentral setae. First flagellomere narrower than gena. Postpronotum without orange spots. Scutum without pruinose strips. Distiphallus longer than wide with stout base and long ventromedial plate; with one pair of spinulose internal structures; ventrolateral tubules nearly indistinct laterally; with sculptured lateral surface (Figs 283, 284). Metepiphallus with spines M. glyptos sp. nov.
12 Distiphallus separated from basiphallus by much more than height of basiphallus (Figs 257, 258) M. angelicae (Frost)
Distiphallus separated from basiphallus by less than height of basiphallus 13
13 Wing length 2.8–3.4 mm. Mid tibia with 2 posteromedial setae. Apex of distiphallus sharply upturned (Figs 303, 304). Basiphallus forming complete ring M. subvirens (Malloch)
Wing length 2.1–2.8 mm, sometimes up to 3.1 mm. Mid tibia with 1 or 2 posteromedial setae. Distiphallus not upcurved apically. Basiphallus U-shaped 14
14 Surstylus relatively small and angled with posterior corner slightly produced. Ventromedial tubules on distiphallus shallowly to strongly looping below main body of distiphallus (Figs 293, 294). Distiphallus separated from phallophorus by height of basiphallus; sides of distiphallus flared outwards towards apex when viewed ventrally, and gradually narrowed apically when viewed laterally. Distal margin of ejaculatory apodeme pointed M. rutella sp. nov.
Surstylus more robust with distal margin broad and straight with posterior corner shallow. Ventromedial tubules on distiphallus held closely against distiphallus. Distiphallus separated from phallophorus by nearly 2 × height of basiphallus; abruptly narrowed on distal 1/2 (seen laterally); shape gently tapering (viewed ventrally). Distal margin of ejaculatory apodeme rounded 15
15 Wing length 2.6–2.7 mm. Ocellar triangle nearly reaching lunule or ending at medial ori. Eye height divided by gena height 5.3. Notum and abdomen faintly greenish. Mid tibia with 1 medial seta. Epandrium deeper dorsally (Fig. 277). Distiphallus (viewed laterally) with strong medial bulge before a strong apical constriction (Fig. 279) M. eoflacensis sp. nov.
Wing length 2.6–3.0 mm. Fronto-orbital plate ending at anterior ors or posterior ori. Eye height divided by gena height 3.8–4.8. Notum more evenly and strongly greenish, and abdomen with very strong green shine. Mid tibia with 1 long and 1 shorter medial seta. Epandrium shallower dorsally (Fig. 261). Distiphallus (viewed laterally) more broadly bulging medially and very gently sloping to apex, not sharply constricted (Fig. 263) M. brunkei sp. nov.

Species descriptions

Melanagromyza angelicae (Frost)

Figs 257–260

Agromyza angelicae Frost, 1934: 40. Frick 1952: 377, 1959: 362; Steyskal 1981: 38; Spencer and Steyskal 1986b: 24.

Description

Wing length 2.8–3.2 mm (♂), 3.2–3.9 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.9. Eye height divided by gena height: 3.1–5.6. Clypeus broadly rounded. Ocellar triangle and fronto-orbital plate subshiny, ill-defined, extending slightly past posterior ori. Fronto-orbital plate slightly visible laterally (more pronounced in male). Gena rounded, highest behind midpoint of eye.

Chaetotaxy : Four ori with anterior seta strongly reduced to absent, and posterior seta sometimes appearing as third ors; two ors. Orbital setulae erect, in two, sometimes three irregular rows, sometimes with several reclinate, but becoming proportionately more proclinate anteriorly; widely spaced from eye margin. Eye pilose dorsomedially. Two dorsocentral setae. Acrostichal setulae in eight irregular rows. Mid tibia with two posteromedial setae.

Colouration : Body, including halter, dark brown in base colour. Gena, parafacial, distal margin of pedicel and inner-distal margin of fronto-orbital plate paler. Notum with faint greenish metallic shine (faded anteriorly). Calypter margin and hairs white. Abdomen metallic green.

Genitalia : (Figs 257–260) Epandrium shallow dorsally, with small posterodistal spine. Surstylus ~ 3/5 length distal region of epandrium, with broad distal margin shallowly angled; with several dense, irregular rows of tubercle-like setae on inner surface. Metepiphallus with several small spines on thickened venter and broad spinulose ridge on thickened lateral section. Proepiphallus short and U-shaped. Basiphallus U-shaped with dorsum broad, separated from mesophallus by several times its own height. Distiphallus relatively small with base not exceeding that of mesophallus; ventrolateral tubules small, inconspicuous; swollen basal section longer than flatter plate-like distoventral region, which has sides gradually narrowed.

Host

Angelica atropurpurea, possibly other Angelica spp. (Spencer and Steyskal 1986b).

Distribution

USA: DE*, MD*, NY, OH.

Type material

Holotype: USA. NY: Ithaca, [reared as a stem borer on Angelica atropurpurea], A.S. Mills (1♂, USNM). [Not examined]

Paratypes examined

. USA. NY: Same data as holotype, 26.iii.1926 (1♀, USNM), 27.iii.1926 (1♂, USNM), 29.iii.1926 (1♀, USNM), 16.iv.1926 (1♂ 1♀, USNM), 17.iv.1926 (2♂, USNM), 21.iv.1926 (1♀, USNM), [no date] (2♂, USNM).

Additional material examined

USA. DE: Wilmington, 9.vi.1974 (2♂, UDCC), MD: Cabin John, 20.vi.1931, A.L. Melander (1♂, USNM), Montgomery Co., Bethseda, 28.iv.1968, G. Steyskal (1♂, USNM), 4mi SW of Ashton, 3.vi.1984, G.F. and J.F. Hevel (1♂, USNM), OH: 3 mi. E Streetboro, 20.iii.1970, Biol Note No. 17-1, D. Witwer (1♂ [with puparium], USNM), Biol Note No. 17-6 (1♂ [with puparium], USNM), Biol Note No. 17-11 (1♂ [with puparium], USNM), Biol Note No. 17-12 (1♀ [with puparium], USNM), Biol Note No. 17-13 (1♂ [with puparium], USNM).

Comments

Melanagromyza angelicae is only known from the eastern United States. Previous records from the western United States and Europe have been determined to belong to other Melanagromyza (Spencer and Steyskal 1986b).

Similar described species with a wide space between the basiphallus and meso/distiphallus include Melanagromyza hicksi Steyskal, collected in Ontario and New York and reared fom Althaea rosea; this species is very robust-bodied, has four or five ori, a gena 1/5–1/4 eye height, a mesophallus that projects further basally past the base of the distiphallus, and a distiphallus that is more strongly tapered apically (Steyskal 1981: fig. 1). Melanagromyza lomatii Steyskal is known from Oregon and was reared from Lomatium nudicaule, and is of a similar size with a similarly high gena; this species has four or five ori, the facial carina is slightly pronounced and wide, the notum and abdomen have a faint bluish shine, the calypter margin and hairs are brown, and the distiphallus is relatively short and stout, and while strongly tapered in lateral view, it is broad and truncated in ventral view (Steyskal 1981: fig. 4). Melanagromyza panacis Steyskal is known from Indiana and Ohio, and has been reared from Panax quinquefoilus; it is slightly smaller (2.5–2.8 mm), the phallus is proportionally shorter, and the distiphallus is slightly larger and stouter with slightly less space between its base and the slightly stouter basiphallus.

Melanagromyza brunkei sp. nov.

Figs 261–263

Description

As described for M. eoflacensis, except as follows: three ori; wing length 2.6–3.0 mm (♂); fronto-orbital plate ending at anterior ors or posterior ori; eye height divided by gena height 3.8–4.8; head very dark brown, without slightly paler brown regions excluding very narrow inner margin of fronto-orbital plate; notum more evenly greenish and with matching, but brighter abdomen; VA male with additional medial dorsocentral on left side as long as anterior dorsocentral; mid tibia with one long and one shorter seta posteromedially; dorsomedial bulge on distiphallus less prominent, and seen laterally, gradually tapering to apex, not abruptly narrowed; seen ventrally, distiphallus with stouter, more prominent narrow plates flanking distomedial tube (Figs 262, 263).

Etymology

The specific name is a patronym for the collector of the holotype.

Host

Unknown.

Distribution

USA: IN, VA.

Type material

Holotype: USA. VA: Giles Co., Ripplemead, Rte 460 bridge, 37°19'43"N, 80°40'48"W, 11–25.v.2008, A. Brunke, debu00304024 (1♂, DEBU).

Paratype : USA. IN: Vincennes, v-6, swept from wint. wheat (1♂, USNM).

Comments

Melanagromyza brunkei is difficult to differentiate from M. eoflacensis, but there are subtle external differences and the genitalia are clearly different when viewed side-by-side, with the profile of the distiphallus being especially diagnostic (see key).

Melanagromyza buccalis Spencer

Figs 52, 53, 264–268

Melanagromyza buccalis Spencer, 1969: 67. Spencer and Steyskal 1986b: 243; Shi and Gaimari 2015: 10; Eiseman and Lonsdale 2018: 17.

Description

(Figs 52, 53). Wing length 1.9–2.4 mm (♂), 2.2–2.5 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.9. Eye height divided by gena height: 3.9–5.6. Gena relatively broad and curved anteriorly with highest point anterior to midpoint of eye. Clypeus slightly truncated to rounded anteromedially. Face with narrow, shallow ridge. Fronto-orbital plate sometimes slightly to moderately swollen medially, narrowed anteriorly and with setae slightly inset (similar to species in M. virens group). Ocellar triangle short, nearly equilateral and with anterior point open.

Chaetotaxy : Two to four ori, but if only two ori, then ori sometimes widely spaced as in M. virens group; two ors. Orbital setulae short, in two to three irregular rows, erect to reclinate with inner row partially inclinate. Eye pilose dorsomedially. Two dorsocentral setae. Acrostichal setulae in eight irregular rows. Mid tibia with two posteromedial setae.

Colouration : Body, including halter, dark brown in base colour. Ocellar triangle and fronto-orbital plate sometimes light brown. Scutum brown with sometimes indistinct greenish shine. Calypter margin and hairs white. Abdomen metallic green, or less commonly, coppery-green.

Genitalia : (Figs 264–268) Epandrium with small posterodistal spine. Metepiphallus with several clustered spine-like ventral projections and numerous narrow lateral ridges. Proepiphallus V-shaped. Basiphallus U-shaped. Distiphallus separated from phallophorus by slightly less than length of basiphallus; base of distiphallus and mesophallus level; distiphallus narrow, long, with sides parallel, with short, thick dorsomedial lobe, and one pair of internal spinulose patches; distolateral plate forming conspicuous, dark oblique band. Ejaculatory apodeme subovate with stem reduced and margin clear.

Host

Unknown, but Spencer and Steyskal (1986b) speculated Asteraceae, and Shi and Gaimari (2015) reported one instance of rearing from an unknown legume in California; some adults collected around a variety of families including Apiaceae, Salicaceae, Rutaceae, Hydrangeaceae, Rosaceae and Asteraceae (Shi and Gaimari 2015; Eiseman and Lonsdale 2018), and Baccharis (Asteraceae) from Maryland.

Distribution

Canada: MB*, NB*, ON, QC. USA: AZ, CA, CO, DE*, GA*, IA*, IL*, IN*, MA, MD, MO, NC*, NH*, NJ*, NM*, NY, PA*, TN*, VA, WV*.

Type material

Holotype: Canada. QC: Lake Bernard, 7.viii.1938, G.E. Shewell, CNC258290 (1♂, CNC).

Paratypes: Canada. NS: Truro, 8.vii.1913, CNC Type No. 10360, CNC358296 (1♂, CNC), ON: Mer Bleue, 3.vi.1938, A.R. Brooks, CNC358295 (1♂, CNC), Osgoode, 18.vi.1964, J.R. Vockeroth, CNC358292, CNC358322 (1♂,1♀, CNC), Ottawa, 2mi E, Cyrville Road, 31.v.1965, B.V. Peterson, CNC358321 (1♀, CNC), Ottawa, CEF, on larch, 3.viii.1962, J.F. McAlpine, CNC358302 (1♂, CNC), Ottawa, 20.vi.1954, D. Cobb, CNC358298, CNC358303, CNC358305, CNC358309, CNC358312 (5♂, CNC), 25.viii.1908, J.M. Fletcher, CNC358299 (1♂, CNC), 7.x.1947, G.E. Shewell, CNC358307 (1♂, CNC), 9.vi.1962, J.R. Vockeroth, CNC358300, CNC358311 (2♂, CNC), Simcoe, 15.vi.1939, G.E. Shewell, CNC358301 (1♂, CNC), 19.vi.1939, CNC358319 (1♀, CNC), 20.vi.1939, CNC358310 (1♂, CNC), 22.vi.1939, CNC358308 (1♂, CNC), 23.vi.1939, CNC358320 (1♀, CNC), 4.vi.1939, CNC358306 (1♂, CNC), Spencerville, 14.viii.1939, Hammond, en copulae, CNC358316 (2♂♀, CNC), QC: Wakefield, 20.vi.1946, G.S. Walley, CNC358291 (1♂, CNC), Abbotsford, 22.vi.1937, G. Shewell, CNC358293 (1♂, CNC), Hull, 25.ix.1923, C.H. Curran, CNC358294 (1♂, CNC), Knowlton, 1.viii.1929, L.J. Milne, en copulae, CNC358314 (2♂♀, CNC), L’Assumption, 7.viii.1936, Shewell, CNC358304, CNC358313, CNC358315, CNC358318 (2♂ 4♂♀, CNC), Lac Bernard, 7.viii.1938, G.E. Shewell, CNC358317 (1♀, CNC).

Additional material examined

Canada. MB: Aweme, 20.viii.1917, N. Criddle, CNC358381 (1♀, CNC), 27.viii.1917, CNC358382 (1♀, CNC), NB: Pokeshaw, 47°47'N, 65°14'W, 27–30.vi.2011, S.E. Brooks, Malaise trap, CNC423058 (1♂, CNC), Saint-Jacques NB Botanical Garden, 47°26'20"N, 68°23'39"W, 27.vii.2013, O. Lonsdale, CNC358505 (1♀, CNC), ON: Ottawa, 20.vi.1954, D. Cobb, CNC358325 (1♂, CNC), St. Lawrence Is. Nat. Park, Grenadier I. Centre, 14.viii.1975, R.J. McMillan, Malaise trap, Code 2-263D, CNC358379 (1♀, CNC), 21.viii.1975, Code 2-278S, CNC358380 (1♀, CNC), Thornhill, 30.v.1964, J.R. Vockeroth, CNC358326 (1♂, CNC), QC: St. Ann-Perade, 6.viii.1930, A.L. Melander (1♂, USNM), Montreal, 8.viii.1950, A.L. Melander (1♂, USNM). USA. AZ: Chiricahua Mts., S.W.R.S., 5400', 30.iv.1979, K.N. Barber (2♂, DEBU), CA: Victorville, 16.v.1955, W.R.M. Mason, CNC358467 (1♂, CNC), “UpStaAnaRiv”, Cienaga, 28.v.1948, J.L. Sperry (1♀, USNM), Vacaville, 19.iv.1949, A.T. McClay (1♀, USNM), Davis, 14.vi.1953, E.I. Schlinger (1♂, USNM), Toulumne Co., Summit Sonora Pass, 9.viii.1948, sweeping Salix eastwoodiae Ckll., Lot No. 108-1, K.E. Frick (3♂, USNM), Summit Sonora Pass, 10.viii.1948, Lot No. 116-1, K.E. Frick (1♀, USNM), San Diego Co., Warmer Spr., 30.viii.1955, E.I. Schlinger (1♂, USNM), Berkeley, 9.vii.1917, J.M. Aldrich (1♂, USNM), 8.vii.1917 (2♂, USNM), Hemet Lake, 13.vi.1961, 500', J.G. Chillcott, CNC358533 (1♂, CNC), CO: Boulder, Valmont Butte, 1615 m, 1.vi.1961, J.R. Stainer, CNC358378 (1♀, CNC), Idaho Springs, 5mi SW, 2621 m, 27.v.1961, J.G. Chillcott, CNC358324 (1♂, CNC), Mt. Vernon Cn. nr. Golden, 2194 m, 31.vii.1961, C.H. Mann, CNC358468 (1♂, CNC), DC: Washington, “C 1 Aug 1956”, P.H. Arnaud Jr. (1♂, USNM), Washington, “viii.5”, J.M. Aldrich (1♀, USNM), Washington, 17.viii.1915, A.L. Melander (1♂ 1♀, USNM), Chain Bridge, 8.v.1928, J.M. Aldrich (1♀, USNM), DE: Dover, 24.vi.1939, A.L. Melander (1♂, USNM), Newark, 31.v.1974 (1♀, UDCC), Newark, 25.viii.1974, D. Buntin (1♀, UDCC), GA: Pine Mt., 1mi North, 12.vii.1957, W.R. Richards, CNC358347 (1♀, CNC), Rabun Bald, 9.viii.1957, W.R. Richards, CNC358355–358357 (3♀, CNC), Rabun Co., 13.vii.1957, W.R. Richards, CNC358354 (1♀, CNC), Warwoman Crk., 4.vi.1957, W.R. Richards, CNC358358 (1♀, CNC), IA: Ames, 19.vi.1947, A.R. Brooks, CNC358461 (1♀, CNC), 28.vi.1947, CNC358458 (1♂, CNC), IL: Champaign, 1.vi.1953, J.F. McAlpine, CNC358460 (1♀, CNC), 22.ix.1956, CNC358459 (1♂, CNC), IN: LaFayette, vi-10, J.M. Aldrich (1♂, USNM), vii-10 (1♂, USNM), x-14 (1♀, USNM), MA: New Bedford, 30.viii.1896, Mass, A.L. Melander (1♂, USNM), Horse Neck Beach, 8.viii.1896, Hough, A.L. Melander (1♂, USNM), Dennisport, C. Cod, 1.viii.1964, J.R. Vockeroth, CNC358535 (1♀, CNC), Franklin Co., Northfield, 276 Old Wendell Rd., 20.viii.2016, mating on Erigeron annuus, #CSE2916, CNC654071, CNC654072 (1♂ 1♀, CNC), MD: Plummers Isl., 20.vii.1913, R.C. Shannon (1♀, USNM), Talbot Co., McDaniel (Wades Point), 19–21.ix.1986, Malaise trap in salt marsh with flowering Baccharis, W.E. Steiner (2♀, USNM), Fairfax Co., Dead Run, 14.v.1914, R.C. Shannon (1♀, USNM), Montgomery Co., Colesville, W.W. Wirth, 27.vii.1976 (1♀, USNM), 28.vii.1976 (2♀, USNM), Bethseda, C.W. Sabrosky, 24.vii.1961 (1♀, USNM), Bethseda, G.C. Steyskal, 10.vii.1970 (1♂, USNM), 16.vii.1979 (1♂, USNM), 24.viii.1975 (4♂ 1♀, USNM), 4mi SW of Ashton, G.F. and J.F. Hevel, 1.ix.1981, Malaise trap (1♂ 1♀, USNM), 24.vii.1982 (3♀, USNM), 15.viii.1982 (2♀, USNM), 19.v.1985 (1♀, USNM), 29.v.1986 (2♀, USNM), 31.v.1986 (1♀, USNM), 5.vi.1986 (1♀, USNM), P.G. Co, Camp Springs, Malaise trap, G.F. Hevel, 28.viii.1979 (1♀, USNM), 6.ix.1979 (1♀, USNM), 14.ix.1979 (1♀, USNM), 15.ix.1979 (2♀, USNM), NC: Highlands, 3800', 21.v.1957, J.R. Vockeroth, CNC358534 (1♂, CNC), Cashiers, 12.vii.1957, W.R. Richards, CNC358351–358353 (3♀, CNC), Clingman’s Dome, 5.viii.1957, W.R. Richards, CNC358350 (1♀, CNC), Highlands, Horse Cove, 30.vii.1957, W.R. Richards, CNC358349 (1♀, CNC), Highlands, Little Bear Pen Mt., 5.viii.1957, W.R. Richards, CNC358348 (1♀, CNC), Highlands, 14.vii.1957, W.R. Richards, CNC358331 (1♂, CNC), 16.viii.1957, CNC358359, CNC358360 (2♀, CNC), 1158 m, 17.viii.1957, J.G. Chillcott, CNC358344 (1♀, CNC), 18.vii.1957, J.G. Chillcott, CNC358339 (1♂, CNC), 1158 m, 21.v.1957, J.R. Vockeroth, CNC358328 (1♂, CNC), 21.vii.1957, W.R. Richards, CNC358327, CNC358329, CNC358330 (3♂, CNC), 1158 m, 21.viii.1957, J.G. Chillcott, CNC358343 (1♀, CNC), W.R. Richards, CNC358361–358366 (6♀, CNC), 22.viii.1957, W.R. Richards, CNC358367–358370 (4♀, CNC), 1158 m, 23.viii.1957, J.G. Chillcott, CNC358338 (1♂, CNC), W.R. Richards, CNC358371 (1♀, CNC), 27.viii.1957, W.R. Richards, CNC358372, CNC358373 (2♀, CNC), 29.vii.1957, W.R. Richards, CNC358332 (1♂, CNC), 1158 m, 3.vi.1957, J.R. Vockeroth, CNC358345, CNC358346 (2♀, CNC), 5.viii.1957, J.G. Chillcott, CNC358333–358337, CNC358340–358342 (5♂ 3♀, CNC), Macon Co., Wayah Gap, 1249 m, 29.vii.1957, J.G. Chillcott, CNC358374 (1♀, CNC), NH: Gorham, 15.vii.1958, J.R. Vockeroth, CNC358462 (1♀, CNC), Fabyan, 30.vi.1936, A.L. Melander (1♂, USNM), NJ: Hemlock Falls, Aug, A.L. Melander (1♀, USNM), NM: Grant Co., ca. 20 mi n. Silver City, 32°57'N, 108°10'W, 7100', meadow, creek, 11.viii.2007, J.D. King, CNC358528 (1♂, CNC), NY: Trudeau, 15.vii.1977, S.W.T. Batra (1♂, USNM), Long Island, Cold Spring Harbor, July, A.L. Melander (1♂, USNM), PA: Susquehanna Co., Rushboro, 12.v.1964, J.G. Chillcott, CNC358465 (1♂, CNC), Chester Co., Toughkenamon, Stroud Res. Ctr., 39°51'37", 75°46'58", 14.ix.2007, E. Lake (2♂, UDCC), nr. Toughkenamon, Stroud Res. Ctr., N39 51 37.2, W75 46 58.2 (1♀, UDCC), 14.ix.2007, C.R. Bartlett (1♀, UDCC), Oxford, 3.ix.2000, Malaise trap, R.L. Snyder (1♀, UDCC), TN: Knoxville, 20.v.1957, J.R. Vockeroth, CNC358375 (1♀, CNC), VA: Augusta Co., Reed’s Gap, 792 m, 7.iv.1962, J.R. Vockeroth, CNC358466 (1♂, CNC), Blacksburg, 28.v.1962, J.G. Chillcott, CNC358376 (1♀, CNC), Montgomery Co., Christiansburg, ex flowers Umbelliferae, 21.vi.1962, J.G. Chillcott, CNC358377 (1♀, CNC), Montomery Co., Longshop, 548 m, 29.v.1962, J.R. Vockeroth, CNC358323 (1♂, CNC), Arlington 4-mile Run, 18.v.1977, W.N. Mathis (1♀, USNM), Veitch, 9.v.1912, J.R. Malloch (1♀, USNM), Great Falls, “vi-1”, G.E. Quinter (1♀, USNM), Falls Church, Holmes Run, W.W. Wirth, 27.vii.1960 (1♂, USNM), 6.vi.1961, light trap (1♂, USNM), Fairfax Co., Alexandria, Four Mile Run, 6.ix.1976, W.N. Mathis (1♂, USNM), Page Co., 7mi W of Lunay, 8.vii.1978, G.F. Hevel (2♀, USNM), WV: Greenbrier Co., Charmco, 6.ix.1983, G.F. and J.F. Hevel (1♂, USNM), WV: Thermopolis, 30.viii.1940, A.L. Melander (1♂, USNM).

Comments

Melanagromyza buccalis is a relatively abundant species most easily recognised by its angled gena, which is highest anterior to the midpoint of the eye. The distiphallus (dark distal outer casing and one pair of dark dorsolateral projections) is also diagnostic. Some specimens, even within collecting events, differ from others in being relatively dark or with a swollen fronto-orbital plate that may be accompanied by widely spaced ori, similar to the state observed in the M. virens group.

Melanagromyza burgessi (Malloch)

Figs 269–271

Agromyza burgessi Malloch, 1913a: 323.

Melanagromyza burgessi. Frick 1952: 378 [as synonym M. lappae (Loew)], 1953: 69, 1959: 363; Spencer and Steyskal 1986b: 244; Shi and Gaimari 2015: 14.

Melanagromyza malefica Spencer, 1981: 46. Spencer and Steyskal 1986b [synonymy].

Description

Wing length 2.7–3.1 mm (♂), 2.9–3.6 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.7. Eye height divided by gena height: 2.2–3.0. Gena strongly angled, highest posteriorly. Fronto-orbital plate and parafacial projecting and conspicuous, obscuring base of antenna (seen laterally) and continuing under eye as cheek; fronto-orbital plate well-developed, setae slightly inset. Clypeus strongly bowed laterally with anterior margin shallowly rounded to slightly truncated medially. Ocellar triangle subshiny and ill-defined.

Chaetotaxy : Four or five ori; two ors. Ocellar setulae short and slightly proclinate, with setulae on posterior 1/3 sometimes reclinate. Orbital setulae short, in three rows, reclinate. Eye bare. Two dorsocentral setae. Acrostichal setulae in ten irregular rows. Three very strong anepisternal setae. Mid tibia with two posteromedial setae.

Colouration : Body, including halter, dark brown in base colour, sometimes with faint blue or greenish shine evident on notum. Parafacial, antenna and gena sometimes paler. Calypter margin and hairs brown. Abdomen with light coppery, greenish, or bluish shine (only coppery observed in this study).

Genitalia : (Figs 269–271) Epandrium with small posterodistal spine. Surstylus rounded with short setae along inner-distal margin and shorter tubercle-like setae in two to three irregular rows on inner surface. Metepiphallus very dark with serrated ventral ridges and two stout coalescing lateral ridges. Proepiphallus thick and globular. Basiphallus forming complete ring. Distiphallus separated from phallophorus by ~ 1.5 × length of basiphallus; distiphallus and mesophallus with bases level; distiphallus with long spinulose internal structures and thin distal projection subequal in length to basal section of distiphallus, which is relatively flat and abruptly ending at midpoint (seen ventrally), with only narrow distoventral plate continuing apically. Ejaculatory apodeme well-developed, typical of Melanagromyza, with blade relatively narrow.

Host

Unknown.

Distribution

USA: CA, CO, IL, IN, KS, MA, MD*, MI, ND, NY.

Type material

Holotype [burgessi]: USA. MA: Beverly, 2.vi.1876, Burgess (1♀, USNM; type No. 15685).

Paratypes [burgessi]: USA. ND: Tower City, 5.vi.1906, G.I. Reeves, Webster No. 3122 (2♀, USNM), CO: “Colo”, “1563” (1♀, USNM).

Holotype [maelifica]: USA. CA: San Diego Co., La Mesa, 23.iii.1962, P.A. Rude (1♂, CAS). [Not examined]

Additional material examined

USA. IN: Lafayette, “v-20”, swept from wint. wheat (1♀, USNM), Evansville, 7.v.1914, swept from wint. wheat, J.M. Aldrich (1♀, USNM), KS: Manhattan, 4.v.1932, D.A. Wilbur (1♀, USNM), Manhattan, C.W. Sabrosky, 20.iv.1934 (3♂, USNM), 25.iv.1934 (1♀, USNM), MD: Montgomery Co., 4mi SW of Ashton, 27.v.1984, G.F. and J.F. Hevel (1♂, USNM), Bethseda, 3.vi.1972, G.C. Steyskal (1♀, USNM), MI: St. Joseph, 30.v.1938, C.W. Sabrosky (1♂, USNM).

Melanagromyza dianthereae (Malloch)

Figs 272–276

Limnoagromyza dianthereae Malloch, 1920: 147.

Melanagromyza dianthereae. Frick 1952a: 378, 1959: 364; Spencer and Steyskal 1986b: 22.

Description

Wing length 3.4–3.9 mm (♂), 4.4–5.0 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.8. Eye height divided by gena height: 2.8–3.4. Male first flagellomere slightly enlarged, rounded, with longer hairs past base. Fronto-orbital plate and parafacial visible laterally, broadly rounded, and continuing under eye as very narrow cheek; fronto-orbital plate well-developed, setae slightly inset. Ocellar triangle long and narrow with anterior point open. Gena relatively short, slightly angled upwards anteriorly. Clypeus broadly rounded with anterior margin slightly thickened. Scutum with faint, thin longitudinal pruinose stripes medially.

Chaetotaxy : Three or four ori; two ors (one in CNC female paratype). Orbital setulae in three irregular rows, reclinate to erect, sometimes proclinate anteriorly. Ocellar setulae erect. Eye bare. Three dorsocentrals, with anterior seta close to second and ~ 3/5 length. Acrostichal setulae in ten irregular rows. Mid tibia with two posteromedial setae.

Colouration : Body, including halter, dark brown in base colour. Head nearly black and with greenish tint. Notum with greenish shine; anterolateral margin lighter brown with postpronotum particularly pale and with yellowish spots. Calypter margin and hairs white. Abdomen brown with green shine, sometimes becoming more blue posteriorly or with colour most pronounced medially and on posterior 1/2 of tergites.

Genitalia : (Figs 272–276) Epandrium without small posterodistal spine. Surstylus shallow and subtriangular, with long marginal setae and several shorter, stouter setae. Hypandrium broad and stout with lobe thick. Metepiphallus smooth; lateral portion nearly divided into separate sclerite and with two crossing ridges. Proepiphallus minute. Basiphallus U-shaped with ends tapered and dorsum very broad and ill-defined with thick transverse medial ridge. Distiphallus large, bulbous, close to phallophorus, tapered apically; with long, thin dorsomedial process; ventrolateral tubules strongly arched, level with mesophallus; internal surface shallowly textured and outer surface minutely tuberculate dorsally.

Hosts

AcanthaceaeJusticia americana (formerly treated as Dianthera).

Distribution

USA: DC*, IL, IN, MD, VA*.

Type material

Holotype: USA. IL: Muncie, 15.viii.1917, Frison and Malloch (1♀, INHS). [Not examined]

Paratypes examined

USA. IN: Lafayette, 19.viii.1916, J.M. Aldrich, swept from Dianthera wildcat Cv. (1♂, USNM), swept from Dianthrea americana (1♂, USNM), Lafayette, on Dianthera americana, 2.vi.1915, J.M. Aldrich, CNC356825, CNC358427 (1♂ 1♀, CNC).

Additional material examined

USA. DC: Rock Creek Park, 18.vii.1933, J.M. Aldrich (1♂ 2♀, USNM), Rock Creek Park, 14.viii.1931, on Dianthera americana (1♂, USNM), IN: Lafayette, J.M. Aldrich, swept from Dianthera americana, 11.v.1915 (1♂, USNM), 11.vi.1915 (3♂, USNM), 18.vi.1915 (2♂ 2♀, USNM), 21.viii.1916 (1♂, USNM), Lafayette, J.M. Aldrich, 18.viii.1916 (1♂, USNM), 10.vi.1915 (4♂ 5♀, USNM) 11.vi.1915 (1♂, USNM), vi-iv.1915 (1♀, USNM), 21.viii.1916 (1♀, USNM), Lafayette, on Dianthera 26.vi.1915 (2♂, USNM), 29.vi.1916 (1♂, USNM), “vii-14” (1♂, USNM), Lafayette, A.L. Melander, “vi-2” (3♂ 1♀, USNM), MD: Plummers Isl., 6.vii.1963, G. Steyskal (1♂ 1♀, USNM), Plummers Isl., K.V. Kromberin, 3.viii.1962 (1♂, USNM), 11.vii.1962 (4♂ 1♀, USNM), 9.vi.1963 (7♂ 2♀ 1?, USNM), 5.viii.1962 (1♂, USNM), 7.viii.1962 (1♂ 1♀, USNM), 1.ix.1962 (3♂ 6♀, 2?, USNM), 31.vii.1962 (2♂, USNM), VA: Fairfax Co., Potomac River at Scott Run, 7.vi.1955, C.W. Sabrosky (2♀, USNM).

Melanagromyza eoflacensis sp. nov.

Figs 277–280

Description

Wing length 2.6–2.7 mm (♂). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.9. Eye height divided by gena height: 5.3. Clypeus broadly rounded. Ocellar triangle and fronto-orbital plate subshiny to slightly matt with triangle reaching level of mid ori or nearly reaching lunule (tapering anteriorly), and fronto-orbital plate slightly visible laterally. Gena rounded, highest behind midpoint of eye.

Chaetotaxy : Three ori (sometimes four on one side); two ors. Orbital setulae in two irregular rows, erect to reclinate. Eye pilose dorsomedially, hairs relatively dense. Two dorsocentral setae. Acrostichal setulae in eight irregular rows. Mid tibia with one posteromedial seta (shorter than width of tibia).

Colouration : Body, including halter, dark brown in base colour. Gena, parafacial, distal margin of pedicel and inner-distal margin of fronto-orbital plate paler. Notum faintly greenish (faded anteriorly), and femora sometimes also slightly reflective. Calypter margin and hairs white. Abdomen with bluish (MD-Bethseda), greenish (IN, PA) or coppery shine (MD-Plummers Isl.).

Genitalia : (Figs 277–280) Epandrium with small posterodistal spine. Metepiphallus with ventral surface dark and with short serrated ridge; lateral margin strongly ridged and nearly separate. Proepiphallus narrow and V-shaped with medial desclerotisation. Hypandrium stout, subtriangular and with small apical process. Basiphallus short and U-shaped; space between basiphallus and distiphallus separated by slightly less than height of basiphallus (slightly more in Lewiston, PA male, illustrated here). Base of mesophallus slightly exceeding that of distiphallus. Distiphallus with spinulose pad along inner-dorsal margin; with thicker basal section ending in bulge before much narrowed apical 1/2; base of distiphallus in IN male straighter and slightly narrower. Ejaculatory apodeme similar to that of M. subvirens, with distal margin darker in holotype.

Etymology

The specific name indicates similarity to M. osoflacensis Spencer, while indicating is relative eastern distribution (eos – Gr. for east).

Host

Unknown.

Distribution

USA: IN, MD, PA, VA.

Type material

Holotype: USA. MD: Montgomery Co., Bethseda, 3.vi.1972, G. Steyskal (1♂, USNM).

Paratype: USA. IN: LaFayette, 10.v.1915, swept from grass, J.M. Aldrich (1♂, USNM), MD: Plummers Isl., 20.iv.1921, H.S. Barber (1♂, USNM), PA: Lewiston, 7.vi.1940, A.L. Melander (1♂, USNM).

Additional material examined

USA. VA: Blacksburg, 2100', 1.vi.1962, J.G. Chillcott (1♂[only wing and genitalia remaining], CNC).

Comments

Melanagromyza eoflacensis can be partially diagnosed by a single posteromedial seta on the mid tibia (not two), but similar species (M. angelicae, M. osoflacensis, M. panacis, M. subvirens, M. brunkei) may be easily misidentified as this taxon, particularly if one or both of the mid tibiae are damaged. As such, the genitalia should always be examined for verification. The phallus of M. eoflacensis is distinct in having a space between the basiphallus and distiphallus that is nearly equal to the height of the basiphallus, the mesophallus only slightly projects past the base of the distiphallus, and seen laterally, the distiphallus has a thick dorsobasal section ending in a pronounced bulge before it abruptly narrows (similar to the state seen in M. angelicae).

The terminalia of Melanagromyza angelicae (Figs 257–260) differ in having a much longer distance between the basiphallus and the base of the distiphallus. Melanagromyza subvirens (Figs 301–304) differs in having a ring-like basiphallus and a flared membranous carina anterodorsally, and the wing is slightly larger. The phalli of M. osoflacensis and M. panacis (Spencer and Steyskal 1986b: figs 35, 36) are more similar, but the former species has very little space between the basiphallus (with posteromedial notch) and the more gradually tapered distiphallus, the fronto-orbital plate is slightly more pronounced and the ocellar triangle is shinier; the latter species has a slightly longer distance between the basiphallus and the base of the distiphallus and it is stouter-bodied but with a similar wing length (2.5–2.8 mm) and the posterior two ori are more closely spaced. The abdomen in these species is also always shiny green, with that of M. subvirens sometimes also bluish. Also see comments for M. brunkei.

The genitalic illustration of this species was erroneously provided for Melanagromyza osoflacensis Spencer in Shi and Gaimari (2015), provided by the present author, at a period when he was not aware of the identity of this new species.

The Virginia male is only tentatively included as most of the body is missing (only one wing is left on the pin) and the genitalia are preserved on a mini-prep slide on an angle that does not allow for fully confident identification, but it appears to agree with the above description.

Melanagromyza glyptos sp. nov.

Figs 54, 55, 281–284

Description

(Figs 54, 55). Wing length 3.6–3.9 mm (♂), 3.1–3.4 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.7. Eye height divided by gena height: 2.4–4.9. Gena angled dorsally on anterior 1/2, highest near midpoint. Fronto-orbital plate and parafacial broadly visible laterally, continuing under eye as distinct cheek; fronto-orbital plate well-developed, setae slightly inset. Clypeus narrowed and slightly truncated medially, sides bowed outwards. Sides of ocellar triangle shallowly concave. Parafacial and ocellar triangle subshiny.

Chaetotaxy : Three or four ori, sometimes with slightly larger gap between anterior two setae; two ors. Orbital setulae dense, in several irregular rows, reclinate to erect. Ocellar setulae erect. Eye broadly pilose dorsomedially. Third dorsocentral (variable from slightly longer than setulae to 2/3 length of second dc) sometimes present directly in front of second dorsocentral in male. Acrostichal setulae in eight irregular rows. Mid tibia with two posteromedial setae.

Colouration : Body, including halter, dark brown in base colour. Scutum shiny with faint greenish reflection. Calypter margin and hairs white. Abdomen (excluding tergite 1) sometimes coppery, with light blue shine usually evident posteriorly.

Genitalia : (Figs 281–284) Epandrium with small posterodistal spine. Hypandrium with apex strongly produced as apodeme. Metepiphallus with serrated ridges, one pair of longer ventral spines, and several overlapping lateral ridges. Proepiphallus V-shaped, narrowly divided medially and with apex upcurved. Basiphallus U-shaped with sides diverging and tips irregular in outline. Distiphallus separated from phallophorus by less than height of basiphallus; high, with spinulose internal structures, apex abruptly truncated, sides subparallel, and with short, thin bifid ventroapical plate; lateral surface of distiphallus with raised plate bearing minute scales and ridges; mesophallus projecting basally from distiphallus and far exceeding basal margin of distiphallus.

Variation: Male from Plummers Island differs as follows: two ori; frons with several longitudinal wrinkles on each side of ocellar triangle; wing length 3.5 mm; length of ultimate section of vein M4 divided by penultimate section 0.5; eye height divided by gena height: 4.8; abdomen with light blue shine; lateral margin of basiphallus less sculptured; lateral plate on dorsal lobe of distiphallus less pronounced, without medial scales. Several specimens from Turkey Run with wing length 3.0–3.2 mm (♂), 3.4 mm (♀), abdomen and notum distinctly green with apex of abdomen bluish, and phallus slightly smaller and paler.

Host

Unknown.

Distribution

USA: MD, SC, VA.

Etymology

Gr. glyptos for carved, referring to sculptured outer lateral surface of distiphallus.

Type material

Holotype: USA. NC: Buncombe Co., 4 km SW Black Mtn., 21–27.1986, W.E. Steiner, Malaise trap, mixed deciduous and hemlock forest nr. small stream (1♂, USNM).

Paratypes: USA. MD: Montgomery Co., Plummers Island, 38°58'N, 77°10'W, Malaise trap, lower trap, 24.iv-7.v.2006, D.R. Smith and J.W. Brown (1♂ 1♀, USNM; 1♀, CNC), Plummers Isl., 11.vii.1915, R.C. Shannon (1♂, USNM). VA: Shenandoah, Lewis Falls, 4.vii.1939, A.L. Melander (1♂, USNM), Fairfax Co., Turkey Run Park, nr. mouth of Turkey Run, 38°57.9'N, 7°09.4'W, Malaise trap, D.R. Smith, 29.iii-25.iv.2005 (1♂ 5♀, USNM; 1♂, CNC), 26.iv-2.v.2007 (12♂ 9♀, USNM), river, 14–17.v.2006 (2♀, USNM), Turkey Run Park, nr. headquarters bldg. 38°57.7'N, 77°08.9'N, Malaise trap, 29.iii-17.iv.2007, D.R. Smith (2♂ 1♀, USNM; 1♂, CNC), Great Falls Park, swamp trail, 38°59.4'N, 77°15.2'W, Malaise trap, 24.iv-2.v.2007, D.R. Smith (1♂, USNM; 1♀, CNC), Shenandoah, N Park Pinnacles, 19.vii.1952, W.W. Wirth (1♂, USNM).

Comments

Melanagromyza glyptos and M. diantherae are superficially similar, being large and stout-bodied with a blue metallic shine and more than five fronto-orbital setae. The two species are otherwise quite different in both outward and genitalic morphology, and can be easily diagnosed using the characters listed in the key.

Melanagromyza matricarioides Spencer

Figs 285–288

Melanagromyza matricarioides Spencer, 1969: 72. Plakidas 1982: 3.

Description

Wing length 2.3 mm (♂). Female unknown (see variation). Length of ultimate section of vein M4 divided by penultimate section: 0.7. Eye height divided by gena height: 5.7. Gena highest behind midpoint of eye. Clypeus rounded, relatively thin and with arms slightly converging apically. Fronto-orbital plate and ocellar triangle subshiny. Fronto-orbital plate well-developed, relatively narrow, broadest behind midpoint of eye but not bulging, with swelling very shallow and indistinct; setae inset. Ocellar triangle narrow with anterior point open. Parafacial projecting dorsally.

Chaetotaxy : Two broadly separated ori; two ors. Orbital setulae in two irregular rows, becoming sparser posteriorly, erect to slightly proclinate. Ocellar setulae erect. Eye pilose in small spot dorsomedially. Two dorsocentral setae. Acrostichal setulae in eight irregular rows. Mid tibia with two posteromedial setae.

Colouration : Body, including halter, dark brown in base colour. Inner margin of fronto-orbital plate (more so anteriorly) and anterior corner of ocellar triangle sometimes light brown. Notum with faint greenish shine posteriorly (blue-green in holotype). Abdomen with green shine (bluer in holotype). Gena, notopleuron, postpronotum, and parafacial sometimes paler. Wing veins pale. Calypter margin and hairs white.

Genitalia : (Figs 285–288) Epandrium with small posterodistal spine. Surstylus narrow and downturned with discrete group of elongate, apical tubercle-like setae (not as figured in Spencer 1969). Hypandrium broad, short, and thick with long apical process. Metepiphallus with shallow spine and serrated ridge ventrally, and with narrow plate with several coalescing ridges laterally. Proepiphallus V-shaped and upcurved. Basiphallus U-shaped, arms relatively thick. Distiphallus separated from phallophorus by less than height of basiphallus; bases of distiphallus and mesophallus level; distiphallus with dorsomedial swelling, internal surface smooth or with remnants of spinulose internal structure (holotype), and with thick distoventral plate and dorsal processes comprising slightly narrowed apex.

Variation : Of the two non-type specimens mentioned in the original description, only the female could be located. This specimen differs as follows: wing length 2.6 mm; eye height divided by gena height 5.1; eye with smaller patch of setulae dorsally adjoining eye; fronto-orbital plate more abruptly bulging at midpoint to ~ ¼ width of frons, setulae in up to three irregular rows; abdomen with green shine. Possibly not conspecific, although most other possible local species to which it may belong in the M. virens group have the head longer towards the dorsum, and none except this species are known from Anthemis.

Hosts

Asteraceae subfamily AsteroideaeMatricaria matricariodes, Anthemis sp., A. cotula, Rudbeckia laciniata[?].

Distribution

Canada: ON. USA: MD, PA[?].

Type material

Holotype: Canada. ON: Ottawa, em. 4.vii.1955, ex. “stem mine in pineapple weed” [Matricaria matricarioides], leg. G. Lewis, reared J.F. McAlpine, CNC358428 (1♂, CNC; Type No. 10366).

Additional material examined

Canada. ON: Ottawa, 25.vii.1962, J.R. Vockeroth, em. 26.vii.1962 from cultivated Anthemis (1♀, CNC). USA. MD: Ft. Detrich, 29.vi.1976, SWT Batra, AC-1 ex. Anthemis cotula L. (1♂, USNM).

Comments

The Maryland male presents the second record of Melanagromyza matricarioides in the United States following Plakidas (1982), who reared specimens from Rudbeckia laciniata in western Pennsylvania. Multiple adults from one or more Pennsylvania localities were recorded from 26 August to 3 October. The specimens were originally identified by G. Steyskal, and it is unknown why they were excluded from Spencer and Steyskal (1986b); only one specimen could be relocated, and examination has found this male to be Melanagromyza virginiensis (see below).

This species belongs to the Melanagromyza virens group, revealed by the widely spaced ori, and by the surstylus, which is incredibly similar to that of M. virens itself, although the long palisade-like setae are arranged in a much more discrete apical cluster. The distiphallus is slightly more compressed dorsoventrally (not teardrop-shaped) and has one pair of characteristic shallow carinae along the dorsolateral margin.

Melanagromyza minimoides Spencer

Figs 289–292

Melanagromyza minimoides Spencer, 1966b: 13. Spencer and Stegmaier 1973: 45; Spencer and Steyskal 1986b: 32; Scheffer et al. 2007: 770; Shi and Gaimari 2015: 51; Monteiro et al. 2019: 157; Eiseman et al. 2021: 10.

Melanagromyza radicicola Steyskal, 1981: 40. Spencer and Steyskal 1986b: 246. Syn. Spencer (1990).

Description

Wing length 1.9–2.1 mm (♂), 2.1–2.3 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.7. Eye height divided by gena height: 5.0–6.3. Gena angulate, highest near midpoint of eye. Clypeus broadly rounded. Fronto-orbital plate and ocellar triangle mostly ill-defined; ocellar triangle rounded anteriorly, extending to region between ors, subshiny. Fronto-orbital plate slightly visible to not visible when head viewed laterally; well-developed but relatively narrow compared to most congeners, slightly subshiny, setae slightly inset.

Chaetotaxy : Two ori; two ors. Orbital setulae in two irregular rows, erect to reclinate. Eye bare or with few minute, sparsely arranged hairs. Ocellar and postocellar setae well-developed, but thin. Two dorsocentral setae. Acrostichal setulae in eight irregular rows. Mid tibia with two posteromedial setae.

Colouration : Body, including halter, dark brown in base colour. Gena slightly pale. Notum greenish, sometimes becoming bluish anteriorly. Calypter margin and hairs brown. Abdomen metallic green, sometimes coppery.

Genitalia : (Figs 289–292) Epandrium with small posterodistal spine. Surstylus small and rounded with tubercle-like setae clustered along inner and inner-marginal surfaces. Apex of hypandrium extremely elongate and tapered, subtriangular with base shallow and broadly arched; inner lobe attached to remainder of hypandrium by lightly sclerotised bridge. Metepiphallus broad and box-like with serrated ridges ventrally and irregular ridges laterally. Distiphallus separated from phallophorus by ~ 1.5 × height of basiphallus; mesophallus produced slightly past base of distiphallus; distiphallus small and ovate in ventral view with thin spinulose internal pads; distoventral plate vestigial, continuing as elongate, irregular duct that often has small, faint subapical sclerotised spot. Ejaculatory apodeme typical of Melanagromyza, blade relatively narrow and slightly pointed apically.

Hosts

AsteraceaeBorrichia frutescens, Helenium flexuosum, Helianthus annuus, Heliopsis helianthoides, Melanthera nivea, Rudbeckia laciniata, Symphyotrichum simmondsii, Verbesina encelioides, V. virginica. CucurbitaceaeCucurbita foetidissima. UrticaceaeUrtica dioica, U. gracilis subsp. holosericea (Eiseman et al. 2021).

Distribution

USA: AR, CA, FL, IA, MD, MO*, MT*, NM*, OH, TX. Argentina. Bolivia. Brazil. Dominican Republic. Guadeloupe. Mexico. Venezuela. Uruguay.

Type material

Holotype [minimoides]. USA. FL: Hialeah, 10–14.ii.1964, ex. seeds of Verbesina sp., leg. 2.ii.1964, C.E. Stegmaier (1♂, USNM [on card with female paratype]).

Paratype [minimoides]. USA. FL: Same data as holotype (1♀, USNM), Miami Dodge Is., 2.iii.1966, ex seeds Borrichia frutescens, em. 4.iii.1966, K.A. Spencer, CNC358429 (1♂, CNC).

Holotype [radicicola]. USA. MD: Bethseda, xii.1979, G.C. Steyskal, “emerged indoors from nettle st.” (1♂, USNM).

Additional material examined

Mexico. Durango: 20.ix.1936, C.S. Rude (1♂, USNM). USA. FL: Hialeah, 15.xi.1964, C. Stegamier, ex. Verbesina (1♂ 1♀, USNM), Key Largo, beach on NE, 13.xii.1985, A.L. Norrbom, reared ex. flower of Borrichia frutescens (L.) DC (14♂ 14♀ [seven with puparia], USNM), Key Largo, Pennekamp State Park, 13.xii.1985, A.L. Norrbom, reared ex. flower head of Borrichia frutescens (L.) (1♂, USNM), Big Pine Key, 13.xii.1985, ex. flower head of Borrichia frutescens (L.) (1♂ 2♀, USNM), Dade Co., Naranja, 12.xii.1985, A.L. Norrbom, reared ex flower head of Bidens alba v. radiata (1♂, USNM), IA: Winneshiek Co., Decorah, Will Baker Park, 43°18'9.12"N, 91°48'42.15"W, 10–13.viii.2017, J. van der Linden, Rudbeckia laciniata, em. by 16.viii.2017, #CSE4945, CNC1643663–1643667 (2♂ 3♀, CNC), MD: Montgomery Co., Bethseda, G. Steyskal, 31.viii.1970 (1♀, USNM), 30.viii.1970 (1♂, USNM), reared ex. flower heads of Heliopsis helianthoides, 21.viii.1970 (2♂ 7♀ 8puparia, USNM), 24.viii.1970 (4♂ 3♀, USNM), 26.viii.1970 (1♂, USNM), 27.viii.1970 (3♂ 1♀, USNM), 28.viii.1970 (2♀, USNM), 30.viii.1970 (1♂ 3♀, USNM), 31.viii.1970 (1♀, USNM), 2.ix.1970 (3♀, USNM), 9.ix.1970 (1♂ 3♀, USNM), 26.ix.1970 (2♀, USNM) reared ex. flower heads of Rudbeckia lacinata, 13.viii.1970 (1♀, USNM), 24.viii.1970 (1♀, USNM), 28.ix.1970 (1♂, USNM), 1.ix.1970 (3♂, USNM), 2.ix.1970 (1♀, USNM), 6.ix.1970 (2♂, USNM), 8.ix.1970 (1♂ 1♀, USNM), 9.ix.1970 (2♂ 1♀, USNM), 10.ix.1970 (3♂ 1♀, USNM), 11.ix.1970 (1♂ 1♀, USNM), 14.ix.1970 (1♂ 3♀, USNM), 16.ix.1970 (2♂ 1♀, USNM), 17.ix.1970 (2♂ 7♀, USNM), 20.ix.1970 (1♀, USNM), viii-ix.1971 (1♀ 7puparia, USNM), 13.viii.1974 (puparia, USNM), 23.viii.1974 (1♀, USNM), 31.viii.1971 (2♀, USNM), 2.ix.1971 (6♀, USNM), 4.ix.1971 (1♀, USNM), 3.ix.1972 (1♀, USNM), 11.ix.1972 (1♀[with puparium], USNM), 1.ix.1974 (7♀[with puparia], USNM), 12.ix.1974 (12♀, USNM), MO: St. Louis, 6.28, reared ex Helianthus pitcheriana, Webster Grve, No. 33125, Satterthwaite (5♂ 9♀[gel capsule], USNM), MT: St. Louis Webster Grvs, No. 22195, “7.30”, reared from Heliopsis, Satterthwaite Collector (2♂ 3♀, USNM), NM: Roosevelt Co., Portales, 4000ft, Malaise trap, 23–26.viii.1993, O’Hara and Jorgensen, debu00128110 (1♂, DEBU), TX: Hidalgo Co., Weslaco, ii.1975, C.E. Rogers, larva in Helianthus annuus (3♂ 2♀, USNM).

Melanagromyza rutella sp. nov.

Figs 293–296

Description

Wing length 2.5–2.7 mm (♂), 2.4–2.8 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.7. Eye height divided by gena height: 2.1–5.9. Clypeus broadly rounded. Ocellar triangle and fronto-orbital plate subshiny. First flagellomere narrow, not broadly rounded below. Fronto-orbital plate and parafacial projecting with portion visible laterally 1/2 height of first flagellomere; fronto-orbital plate well-developed, setae inset. Ocellar triangle subshiny to matt, sometimes appearing to end at posterior ori, but if anterior margin ill-defined, then appearing to continue to lunule. Gena rounded, highest behind midpoint of eye.

Chaetotaxy : Three ori (four on one side of IN male); two ors. Orbital setulae short, in up to four irregular rows, erect to reclinate. Eye pilose dorsomedially. Two dorsocentral setae. Acrostichal setulae in eight to ten irregular rows. Mid tibia with one or (more commonly) two posteromedial setae.

Colouration : Body, including halter, dark brown in base colour. Notum greenish (faded anteriorly); femora sometimes also slightly reflective. Calypter margin and hairs white. Abdomen with greenish shine, sometimes becoming slightly coppery/reddish.

Genitalia : (Figs 293–296) Posterodorsal margin of epandrium slightly produced; posteroventral spine well-developed. Surstylus 1/2 height of epandrium, with up to three rows of tubercle-like setae along inner distal margin. Basiphallus short, U-shaped, and extending slightly past base of distiphallus. Base of mesophallus extending past that of distiphallus; distiphallus with paired ventrolateral tubules shallowly raised from base of distiphallus, or prominent, separate and strongly looped (as illustrated); with dorsoapical margin strongly flared laterally; venter with strong divided carina. Ejaculatory apodeme well-developed with base dark, becoming paler distally on stem; blade large, spade/diamond-shaped, weakly sclerotised but with defined medial rib and strong marginal band (less so laterally) well-defined.

Host

Unknown.

Distribution

USA: IN, VA.

Etymology

L. rutrum for spade, shovel, referring to the shape of the large ejaculatory apodeme.

Type material

Holotype: USA. VA: Fairfax Co., Great Falls Park, quarry, 38°59.1'N, 77°14.8'W, Malaise trap, D.R. Smith, 24.iv-2.v.2007 (1♂, USNM).

Paratypes: USA. IN: La Fayette, 11.v.1916, J.M. Aldrich (1♂ 2♀, USNM), VA: Same collection as holotype (1♂ 2♀, USNM; 2♀, CNC), 3–10.v.2007 (3♀, USNM).

Comments

Melanagromyza rutella is dark (i.e., without a slightly paler gena, shoulders, etc.) with a white calypter and three ori on both sides of the frons. The gena is also often relatively high, the blade of the ejaculatory apodeme is nearly diamond-shaped, the distiphallus is stout and strongly flared apically, the ventrolateral tubules of the phallus are distinct and sometimes broadly looped, and the distiphallus has a single internal rectangular pad with numerous transverse rows of small spines.

There is a resemblance to Melanagromyza urticae Eiseman and Lonsdale (Eiseman et al. 2021: figs 16–19, 111–116), especially with regards to the male genitalia, but this other Iowa species has two ori, three ors, the surstylus is not produced slightly posteriorly, the mesophallus is inserted almost basally on the distiphallus (the ventrolateral tubules are more pronounced in M. rutella), the dorsal chamber of the distiphallus is stouter and almost parallel-sided in ventral view (not constricted medially) and higher in lateral view, and the tubule emerging from the dorsal chamber apically is longer.

Melanagromyza splendida Frick

Figs 297–300

Melanagromyza splendida Frick, 1953b: 207. Spencer 1981: 59; Spencer and Steyskal 1986b: 25; Shi and Gaimari 2015: 72.

Description

Wing length 1.8–2.6 mm (♂), 1.9–2.6 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.5–0.7. Eye height divided by gena height: 3.9–6.9. Deepest portion of gena 1/3–1/2 distance from anterior margin. Fronto-orbital plate not projecting, sometimes slightly visible in lateral view; strongly swollen in males, 25–28% frons width, narrowing anteriorly; narrower in females, 16–23% frons width; setae inset along inner margin of plate. Clypeus broadly rounded. Ocellar triangle slightly longer than wide.

Chaetotaxy : Two ori with anterior seta strongly displaced anteriorly; two ors. Eye extensively pilose, hairs sparse excluding dense dorsal patch. Orbital setulae erect to reclinate, slightly proclinate anteriorly (mostly inner row), in up to three or four irregular rows; setulae longer in male. Two dorsocentral setae. Acrostichal setulae in eight irregular rows. Mid tibia with two posteromedial setae.

Colouration : Body, including halter, dark brown in base colour. Fronto-orbital plate sometimes whitish along anteromedial margin. Notal colour ranging from brilliant green to dull green with coppery tint. Calypter margin and hairs white. Abdomen metallic green, sometimes bronze (Austin, TX).

Genitalia : (Figs 297–300) Epandrium with posterodistal spine. Surstylus < 1/2 height of epandrium, distal margin slightly rounded and angled, with several irregular rows of tubercle-like setae along inner-distal surface. Hypandrium subtriangular, long with thin, tapered distal process. Proepiphallus dark and V-shaped. Metepiphallus broad and dark with several sublateral tubercles and lateral ridges. Basiphallus U-shaped. Space between distiphallus and phallophorus less than height of basiphallus. Distiphallus relatively globular with distinct transverse fold laterally on distal 1/3 that curves towards venter; base exceeding that of mesophallus; inner surface only with clusters of shallow bumps (no spines); with short narrow tubule emerging apically below short plate; seen ventrally, width of distiphallus varying from relatively slender (as in illustration) to stouter around middle.

Variation : One AZ male (dissected) with fronto-orbital plate swollen to 1/3 width of frons (ie. similar to M. virens). Dissected Newark (DE) male, possibly a separate species, differs as follows: dark distoventral process on distiphallus highly reduced, distiphallus slightly more quadrate in outline (seen ventrally), and surstylus not downturned apically, but with long tubercule-like setae typical of M. matricariodes and M. virens; clypeus distinctly truncated apically and orbit slightly less swollen medially.

Hosts

AsteraceaeAmbrosia, Bidens, Centaurea*, Cineraria*, Coreopsis*, Erechtites, Flaveria, Gaillardia, Galinsoga*, Gnaphalium, Helianthus, Hymenoxis*, Lactuca, Parthenium, Tagetes, Zinnia. CucurbitaceaeCucurbita. ApiaceaeApium, Daucus. NC record of feeding on potato requires verification.

Distribution

USA: AZ, CA, DE*, FL, HI, IL, IN*, MD*, MI[?], MO*, NC*, NY[?], SC[?], TX*. Argentina*. Bahamas. Chile. Jamaica. Mexico.

Type material

Holotype: USA. HI: Kamuela, 5.xii.1950, ex. celery (1♂, USNM).

Paratype: USA. HI: same data as holotype (1♀, USNM).

Additional material examined

Argentina. Salta: Rt. 34, 4 km S Metan, i.1983, Galinsoga parviflora, tunneling inside stems (1♂ 1♀, USNM), Jujuy: Rt. 9, 1 km S Maimara, 31.xii.1982, Hymenoxis robusta, tunneling inside stem (1♀, USNM). Bahamas. Bimini Isl., 22–31.i.1968, light trap, G.M. Stokes (2♂, USNM). Chile. Valdivia: Santiago, 4.v.1978, 78.3999, reared ex. sunflower stalk (1♂, USNM). USA. AZ: Yuma, 20.v.1968, D.M. Tuttle (2♂ 2♀, USNM), CA: Ortega Hiwy., near summit, 11.vi.1944, A.L. Melander (1♂, USNM), San Simeon, 31.viii.1945, A.L. Melander (1♀, USNM), Orange Co., Fullerton, 13.ii.1969, R.D. Goeden and D.W. Ricker, insectary reared on Ambrosia psilostachya Decandolle (2♂, USNM), Santa Barbara Co., Santa Barbara, 22.iv.1969, R.D. Goeden and D.W. Ricker, insectary reared on Ambrosia psilostachya (2♂ [with puparia], USNM), San Bernardino Co., Highland, 2.v.1969, R.D. Goeden and D.W. Ricker, insectary reared on Ambrosia acanthicarpa Hook. (1♀ [with puparium], USNM), Cucamonga, 20.v.1969, R.D. Goeden and D.W. Ricker, insectary reared on Ambrosia acanthicarpa Hook. (1♀ [with puparium], USNM), Salinas, 1.xi.1930, Bred from [illegible]ule (1♂, USNM), L.A. Co., Westwood Hills, 3.xi.1938, ex Cineraria stems acc. 94 (1♂, USNM), DE: Lum’s Pond, 17.vii.1974, D. Buntin (2♂, USNM), Newark, D. Buntin, 19.viii.1974 (2♂, USNM), 10.vii.1975 (1♂, USNM), 9.vi.1975 (1♂, USNM), FL: Dade Co., Naranja, 12.xii.1985, A.L. Norrbom (1♂, USNM), Highlands Co., Archibold Biological Sta., 24.iii.1969, S.W. Frost, CNC358443 (1♂, CNC), Highland Co., Archibold Biol. Sta., 24.iv.1967, B.V. Peterson, CNC358464 (1♂, CNC), HI: Honolulu, 6.v.1929, ex Cornflower (Centaurea), J.F. Illingworth (2♂, USNM), 7.v.1929 (1♂ 1♀, USNM), 8.v.1929 (1♀, USNM), IL: Champaign, tomato leaves, 12.ix.1957, J.F. McAlpine, CNC358451 (1♂, CNC), Alhambra, 24.vi.1937, reared from sunflower, destroying stem, Satterthwaite (4♂ 10♀, USNM), IN: Lafayette, J.M. Aldrich, “vii-21” (1♂, USNM), “x-14” (1♂, USNM), 4.ix.1916 (1♂, USNM), “vii-26” (1♂, USNM), “6-27” (1♂, USNM), “v-23” (1♂, USNM), “v-27” (2♂, USNM), [no date] (1♂, USNM), MD: College Park, 2.viii.1931, C.T. Greene (1♂, USNM), NC: Chadboura, 27.v.1910, feeding on potato, E.G. Smyth (1♂, USNM), MO: Webster Groves, 17.v.1932, Helianthus tuberosa, Satterthwaite (1♀, USNM), [locality not provided, likely Webster Groves], 32165b, Helianthus tuberosa (1♂, USNM), TX: Galveston, vi.1900, A.L. Melander (1♂, USNM), Mexico, in carrot, Laredo, Tx., 31255, 18.i.1943, Lot No. 43-1075 (1♂, USNM), 31457, 13.ii.1943, Lot No. 43-1590 (1♂ 1♀, USNM), Mexico, in carrot root, El Paso, 33818, Lot No. 42-6728 (1♀, USNM).

Comments

The short, squat distiphallus is highly characteristic of this species, especially the bulging laterodistal region that appears as a transverse fold. Male dissection is recommended for confident identification.

While Melanagromyza splendida has previously been recorded as far north as New York and Michigan in the United States, almost all of the northern males identified by previous authors dissected here have been M. virginiensis. The remaining specimens for which previous Michigan, New York, and South Carolina records of M. splendida were based (Spencer and Steyskal 1986b) have not been found, and it is suspected that these might also be misidentifications of M. virginiensis.

Several Texas records, which may represent interceptions from Mexico, note carrot as host. When considering the economic significance of carrots, the widespread occurrence of the fly, and the absence of previous rearing records, it is possible that the labels are in error or Daucus is not a preferred host.

Melanagromyza subvirens (Malloch)

Figs 301–304

Agromyza subvirens Malloch, 1915a: 105.

Melanagromyza subvirens. Frick, 1959: 366; Spencer and Steyskal 1986b: 26.

Description

Wing length 2.8–3.2 mm (♂), 3.4–3.5 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.5–0.6. Eye height divided by gena height: 5.0–5.5. Gena highest at ~ anterior 1/3, angled upwards anteriorly, with distinct cheek. Fronto-orbital plate well-developed, slightly visible laterally, setae slightly inset. Clypeus broadly rounded. Fronto-orbital plate and ocellar triangle slightly less matt than surrounding frons. Ocellar triangle reaching second ori from rear.

Chaetotaxy : Five to six fronto-orbitals with posterior one or two reclinate (ors). Ocellar setulae short, reclinate, in two irregular rows. Eye sparsely pilose dorsomedially, sometimes slightly denser in male. Anterior genal seta sometimes vibrissa-like. Two dorsocentral setae; MD male with third dorsocentral on left side. Acrostichal setulae in eight irregular rows. Mid tibia with two posteromedial setae.

Colouration : Body, including halter, dark brown in base colour; female darker, blackish. Scutellum with faint greenish shine in male, female sometimes with bluish tint. Calypter margin and hairs white. Abdomen with green or bluish green shine, and females sometimes with strong blue shine only. Slightly reddish, at least on antenna and gena, and sometimes more widespread.

Genitalia : (Figs 301–304) Epandrium with small posterodistal spine. Surstylus ~ 1/2 length of epandrium venter (epandrium dorsum shallower); inner surface with several irregular rows of tubercle-like setae. Hypandrium subtriangular with slightly produced apex. Metepiphallus with serrated ventral and lateral ridges. Proepiphallus V-shaped. Basiphallus forming complete ring with ventrodistal margin ill-defined. Distiphallus separated from phallophorus by height of basiphallus; mesophallus slightly exceeding base of distiphallus; distiphallus relatively flat, with internal spinulose pads, strongly upturned and apically widened dorsoapical membranous process; ventrolateral tubules short. Ejaculatory apodeme well-developed with short stem and narrow rib slightly exceeding apex of broad, pale, ovate blade.

Host

Unknown.

Distribution

USA: IA, IL, MD*, NC*, PA, VA.

Type material

Holotype: USA. IL: St. Joseph, 17.v.1914, C.A. Hart and J.R. Malloch (1 ♀, INHS). [Not examined].

Paratype: USA. IL: St. Joseph, 17.v.1914, Salt Fork (1♀, USNM).

Material examined

USA. IL: Oregon, 21.vi.1917, en copulae; CNC358452 (1♂ 1♀, CNC), MD: Laurel, 1.vi.1965, Malaise trap, CNC358454, CNC358455, CNC358456 (3♀, CNC), nr. Plummers Isl., 2.v.1915, R.C. Shannon (1♀, USNM), Montgomery Co., Bethseda, 26.v.1968, G. Steyskal (1♂, USNM), NC: Wilson’s Gap, 944 m, 25.v.1957, J.R. Vockeroth, CNC358453 (1♂, CNC).

Melanagromyza vernoniae Steyskal

Figs 305–308

Melanagromyza vernoniae Steyskal, 1981: 41. Spencer and Steyskal 1986b: 23.

Description

Wing length 3.1–3.5 mm (♂), 2.8–3.4 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.7. Eye height divided by gena height: 4.6–5.5. Clypeus broadly rounded. Male eye with short, scattered setulae covering dorsomedial region of eye (setulose region relatively narrow compared to M. vernoniana) that may sometimes be relatively dense; female eye virtually bare. Fronto-orbital plate slightly widened at middle, only slightly more than one quarter width of frons at widest point; subshiny; bulging anteriorly and slightly visible laterally, sometimes continuing as ring below eye. Ocellar triangle relatively narrow and parallel-sided, reaching level of posterior ori.

Chaetotaxy : Two ori with anterior seta strongly displaced anteriorly (one IN male with additional seta between standard ori that is weak on left side and well-developed on right); two ors with anterior seta slightly inclinate. Orbital setulae with up to three irregular, scattered rows, generally erect, slightly reclinate anteriorly and proclinate posteriorly and along outer row; relatively long and bushy, considerably longer than ommatrichia (more so in males). Two dorsocentral setae. Acrostichal setulae in eight irregular rows. Katepisternum with one to several short setae clustered dorsomedially. Mid tibia with two posteromedial setae.

Colouration : Body, including halter, dark brown in base colour; pedicel and scape sometimes slightly paler brown. Notum with slight bluish shine that may be slightly greenish, especially on posterior third of scutum and on scutellum. Calypter margin and hairs white. Abdomen with strong metallic green shine. MN male with abdomen blue. IN specimens with notum greenish (not blue) and abdomen greenish coppery in males and green to blue in females.

Genitalia : (Figs 305–308) Surstylus (entirely fused to epandrium) with relatively long, flat distal margin ~ 2/3 length of distal section of epandrium (dorsum of epandrium shallower) that is slightly produced at anterior corner; inner surface with two (posteriorly) to four (anteriorly) rows of short, thickened setae, of which a few at slightly produced corner of surstylus are elongate, although state not as exaggerated as seen in M. virens. Hypandrium extremely long and produced apically and with longitudinal basal ridge; inner lobe irregular in outline and weakly sclerotised. Metepiphallus with small spines medially and with coalescing ridges laterally. Proepiphallus V-shaped. Basiphallus U-shaped with corners at right angles and arms parallel. Mesophallus and distiphallus level at base. Distiphallus with basal section ~ 2/3 length of segment, relatively narrow, not bulging dorsally, with well-developed internal spinulose pads; ventrolateral tubules relatively long, narrow, recessed.

Host

AsteraceaeVernonia noveboracensis.

Distribution

USA: DC, IN*, MN*.

Type material

Holotype [vernoniae]: USA. DC: 18.x.1969, D. Anderson, Vernonia noveboracensis, emg. indoors after refrigeration 19.xii.1969 (1♂, USNM).

Paratypes [vernoniae]: USA. DC: Washington, E Bank CandO Canal nr. Ariz. Ave., reared from larva in stem Vernonia noveboracensis, D.M. Anderson, 23.i.1970 (1♂, USNM), 28.iii.1970 (2♂, USNM).

Additional material examined

USA. DC: Washington, E Bank CandO Canal nr. Ariz. Ave., 14.x.1969, D.M. Anderson, Reared from larvae in stem Vernonia noveboracensis, 23.ii.1970 (3♀, USNM), Washington, E Bank CandO Canal nr. Ariz. Ave., 28.iii.1970, D.M. Anderson, Reared from larvae in stem Vernonia noveboracensis (1♀, USNM), IN: LaFayette, J.M. Aldrich, v-23 (1♂ 1♀, USNM), v-27 (6♂ 3♀, USNM), v-28 (1♂, USNM), iv-27 (3♂, USNM), vi-10 (1♂, USNM), vi-22 (1♀, USNM), vi-23 (1♂, USNM), vii-8 (1♀, USNM), 6-27 (1♂, USNM), MN: New Ulm, 30.v.1916, J.M. Aldrich (1♂, USNM).

Melanagromyza vernoniana Steyskal

Figs 309–317

Melanagromyza vernoniana Steyskal, 1981: 41. Spencer and Steyskal 1986b: 23.

Melanagromyza verbesinae Spencer in Spencer and Steyskal 1986b: 248. Syn. nov.

Description

Wing length 2.3–2.8 mm. Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.8. Eye height divided by gena height: 4.1–5.0. Clypeus broadly rounded. Pilosity on male eye dense and relatively broad dorsally, covering most of dorsal third of eye. Fronto-orbital plate slightly widened at middle, not exceeding 1/5 width of frons; subshiny; reaching level of posterior ori.

Chaetotaxy : Two ori with anterior pair of setae widely separated from second pair; two ors. Orbital setulae in up to three irregular rows, reclinate to slightly lateroclinate with inner row partially proclinate; as long as eye setulae, and slightly darker and much less densely arranged compared to eye setulae. Two dorsocentral setae. Acrostichal setulae in eight irregular rows. Mid tibia with two posteromedial setae.

Colouration : Body, including halter, dark brown in base colour; pedicel and portions of frons sometimes slightly paler. Notum with faint greenish to bluish shine; indistinct in poorly preserved specimens. Calypter margin and hairs white. Abdomen with strong metallic green shine, sometimes bronze.

Genitalia : (Figs 309–313) Epandrium with small posterodistal spine. Surstylus one 1/2 length of epandrium or slightly less; slightly produced, elongate; inner surface with several irregular rows of tubercle-like setae, posterodistal corner with several of these longer, posteriorly directed (not as pronounced as state seen in M. virens and M. matricariodes). Metepiphallus with serrated ventral ridges and ventrally coalescing lateral ridges. Proepiphallus very dark and V-shaped. Phallophorus short. Basiphallus U-shaped with minute irregularities in lateral margin. Distiphallus separated from phallophorus by slightly more than height of basiphallus; base of mesophallus produced slightly past base of distiphallus; distiphallus with broad, short base and elongate distal section that is slightly upcurved when seen laterally and elongate oval when seen ventrally; distiphallus sometimes with one pair of short subtriangular plates emerging anterodorsally from swollen basal section.

Variation : See comments section.

Host

AsteraceaeHelianthus annuus*, H. tuberosus*, Helenium autumnale*, Verbesina alternifolia, Vernonia noveboracensis.

Distribution

Canada: ON*. USA: DC, IL*, IN*, MD, MO*, OH, OK*, TN, TX, VA*, VT*, WI*.

Type material

Holotype [vernoniana]: USA. MD: Cropley, 20.x.1968, D.M. Anderson, emg. Ex. stem Vernonia noveboracensis (1♂, USNM).

Paratype [vernoniana]: USA. DC: Washington, E bank CandO Canal nr. Ariz. Ave., 28.iv.1970, D.M. Anderson, reared from larva in stem Vernonia noveboracensis (1♂, USNM).

Holotype [verbesinae]: USA. OH: Jennings Woods, 6.0mi NE Ravenna, 21.iii.1970, ex. stem Verbesina alternifolia (1♂, USNM).

Paratypes examined

[verbesinae]: USA. TN: East Ridge, 6.v.1952, G.S. Walley, CNC358449 (1♂, CNC), TX: Kerrville, swept ex meadow, 18.iv.1959, J.F. McAlpine, CNC358448 (1♂, CNC).

Additional material examined

Canada. ON: Wellington Co., nr. Arkell, 43°33'18"N, 80°10'16"W, trail beside river, 24.vi.2010, O. Lonsdale, CNC358506 (1♂, CNC). USA. IL: Urbana, in Nat. Res. Mus., 5.v.1957, J.F. McAlpine, CNC358450 (1♂, CNC), Hinsdale, 25.x.1929, H[elianthus] tuberosus, Webster Grve, Satterthwaite, No. 29308818 (1♂, USNM), No. 29308817 (1♂, USNM), Hinsdale, [illegible], 25.x.1929, Webster Grvs, No. 29310a, [illegible], 8.i.1930, Satterthwaite (1♀, USNM), Chicago, 6.vi.1903, A.L. Melander (1♂, USNM), IN: Lafayette, 27.x.1915, reared from sunflower pith, Cage No. C1490, Satterthwaite (1♂, USNM), MD: Montgomery Co., Bethseda, 23.v.1970, G. Steyskal (1♂, USNM), MO: Black Jack, 28.x.1930, Webster Grove, Helianthus tuberosa, Satterthwaite, [illegible], No. 40319g, 13.iv.1931 (1♀, USNM), No. 30419a, 20.iv.1931 (1♂, USNM), Web Groves Sta., dead at cold room window, 14.iv.1933 (3♂ 4♀[gel capsule], USNM), 20.iv.1933 (2♀[with 2 Braconidae in gel capsule], USNM), 21.iv.1933 (1♀[with Braconidae in gel capsule], USNM), OK: Murray Co., Sulphur, Chickasaw Rec. Area, 4.vi.1979, S.andJ. Peck, prairie vegetation (1♂, DEBU), MO: Cross Keys, 1.xii.1932, dead 3.ii.1933, Webster Grv, No. 32S11, R.B. Swain (1[unemerged from puparium], USNM), Webster Grvs, No. 32052C, Helianthus annuus, laboratory garden, [date illegible], dead 11.v.1932, Satterthwaite (1♂, USNM), Webster Groves, Helianthus annuus, 23.iii.1930, No. 30012, issued 1.iv.1930, R.C. Lange (1♂, USNM), 16.iii.1931, No. 31025C, issued 14.iv.1931 (1♂, USNM), Webster Groves, sunflower, iss. [?].iii.1933, No. 33034R, Satterthwaite (1♂, USNM), iss. 4.iv.1933, 26.iii.1933 (1♂ 2♀, USNM), 26.iii.1933, No. 33039f, iss. 3.iv.1933 (1♂ 2♀, USNM), iss. 5.iv.1933, No. No. 33039f, 26.iii.1933 (1♂, USNM), iss. 30.iii.1933, No. 330392, 26.iii.1933 (1♂, USNM), Webster Groves, 26.vii.1929, reared from sunflower, issued 3.viii.1929, dead 3.viii.1929, No. 29162, Satterthwaite (1♂, USNM), Webster Groves, sunflower, issued 12.viii.1929, dead 8.13, No. 29161, 26.vii.1929, Satterthwaite (1♂, USNM), Webster Groves, 9.vi.1929, sunflower, Satterthwaite (1♀, USNM), 21.viii.1928 (1♀, USNM), Maplewood, 24.vii.1930, issued 22.viii.1930, Webster Grvs, No. 30246b, Satterthwaite (1♂, USNM), issued 25.viii.1930, No. 30246a (1♀, USNM), East St. Louis, [?], 9.ix.1929, sunflower, Webster Grvs, No. 2925290, issued ix.1930, J.B. Sahan (1♀, USNM), Cross Keys, Artichoke, Webster Grvs, R.B. Swain, 1.xii.1932, iss. 14.iii.1933, No. 32497p (1♂, USNM), 1.xii.1932, iss. 4.iii.1933, No. 32497t (1♂, USNM), 1.xii.1932, iss. 1.iv.1933, No. 32497aa (1♂, USNM), 1.xii.1932, iss. 29.iii.1933, No. 32497d (3♂ 1♀, USNM), 1.xii.1932, iss. 1.iv.1933, No. 32497ab (1♂, USNM), 1.xii.1932, iss. 5.iv.1933, No. 324970 (1♀, USNM), 1.xii.1932, iss. ?3.iv.1933, No. 32497h (1♀, USNM), 1.xii.1932, iss. 2.20, No. 33003 (1♀, USNM), Florrisant, Artichoke, Webster Grvs, 2.xii.1932, iss. 12.iv.1933, No. 32495 (1♀, USNM), Black Jack, Artichoke, Webster Grvs, R.B. Swain, 18.x.1932, iss. [illegible], No. 32476 (1♂, USNM), 1.xii.1932, iss. 31.iii.1933, No. 32497f (1♂, USNM), 1.xii.1932, iss. 13.iv.1933, No. 32446 (1♀, 6♂ 2♀[gel capsule], USNM), VA: Batesville, M.L. Bobb, 6.vi.1940 (1♂, VPIC), 26.iv.1939 (1♂, VPIC), 18.v.1940 (1♀, VPIC), 30.iv.1940 (1♂, VPIC), Fairfax Co., Great Falls Park, quarry, 38°59.1'N, 77°14.8'W, Malaise trap, 3–10.v.2007, D.R. Smith, CNC358447 (1♂, CNC), Charlottesv., 30.xi.1930, Webster Grvs, No. 30480s, Helianthus tuberosa, [illegible], 14.iv.1931, Dr. Phillips (1♀, USNM), 20.xi.1930, No. 30480h (1♀, USNM), VT: Plainfield, 24.vi.1980, B.V. Peterson, CNC358504 (1♂, CNC), WI: Grant Co., Thomas Wet. Prairie, 30.ix.1997, A.H. Williams, Helenium autumnale, [host plant] stripped of leaves and inflorescences, put into sterile containers over sterile soil and netted with hosiery, outdoors until 2.iii.1998 when caged in lab, em. 4–15.iv.1998, CNC934525–934529 (3♂ 2♀, WIRC).

Comments

While never directly compared in the literature, male terminalia and external morphology reveal strong similarities between Melanagromyza vernoniana and M. verbesinae, especially the phallus, which includes a distiphallus that is long and curved with the basal section abruptly swollen and globose with suspended ventrolateral tubules.

The examined Melanagromyza verbesinae paratypes have a faint greenish tint on the notum and a relatively broad, truncated surstylus, and were apparently used for the underlying original concept of the species. The wing length is also 2.4–2.8 mm, not 2.1 mm as noted in original description. Hosts are unknown for paratypes. Examination of the genitalia has also revealed that the basal section of the distiphallus is produced into one pair of weakly sclerotised, subtriangular dorsal plates. These genitalic features are also seen in the non-type material reared from “artichoke” (Figs 314–317) and some specimens reared from Helianthus annuus. The M. vernoniana types (ex Vernonia) and the M. verbesinae holotype (ex Verbesina) have a faint bluish tint on the notum, a slightly shallower epandrium, a narrower, rounded surstylus, and no dorsal plates arising from the basal globose section of the distiphallus; wing length is 2.3–2.8 mm.

Aside from these type specimens, additional material has been examined exhibiting overlap in genitalic morphology between the two extremes outlined above. Some males have well-developed to reduced dorsal plates on the distiphallus and a narrower surstylus; specimens without these dorsal plates on the distiphallus sometimes have a stouter truncated surstylus, such as those reared from Helianthus tuberosus and one male from H. annuus. Colouration is also not as distinct as previously considered. The green or blue shine is faint and difficult to discern, and while some specimens are clearly greener or bluer compared to others, intermediate colouration is sometimes evident. The difference in colouration, however, is much less than that seen in other Melanagromyza species (although usually this is manifest on the abdomen, not the notum) and may be insignificant. The continuum of states seen in the genitalia, the similarities in colour and body length (which are not as disparate as previously considered), and the strong similarities in other external morphological features support the synonymy of these two species; this is compounded by the especially strong similarities between the M. verbesinae holotype and the type material of M. vernoniana. Rearing records of both extreme genitalic types from Helianthus further support synonymy.

Some of the newly recorded specimens examined here were reared from “artichoke” at Webster Groves. This may refer to Cynara scolymus, the “globe artichoke”, but it is more likely that it refers to Helianthus tuberosus, otherwise known as “Jerusalem artichoke”, also studied at Webster Groves. The Helenium-reared specimens were first reported as Melanagromyza sp. in Williams (1999).

Melanagromyza virens (Loew)

Figs 4, 56, 57, 62, 63, 318–324

Agromyza virens Loew, 1869: 46.

Melanagromyza virens. Frick, 1952a: 380, 1957: 200 [lectotype designation], 1959: 367; Spencer 1969: 76; Spencer and Stegmaier 1973: 51; Spencer and Steyskal 1986b: 25; Scheffer et al. 2007: 770; Shi and Gaimari 2015: 85.

Melanagromyza heterothecae Spencer, 1966b: 10. Syn Spencer (1969).

Description

(Figs 4, 56, 57, 62, 63, 324). Wing length 1.8–2.3 mm (♂), 2.4–2.6 mm (♀); female lectotype and paralectotype 2.9 mm and 3.2 mm, respectively. Length of ultimate section of vein M4 divided by penultimate section: 0.6. Eye height divided by gena height: 4.6–6.5. Gena usually deepest at 1/3 distance from anterior margin, or slightly behind. Clypeus broadly rounded. Ocellar triangle narrow, reaching level of posterior ori, sides concave. Fronto-orbital plate widest medially, broadest behind middle; slightly visible laterally, sometimes more so anteriorly; very broad, each plate usually at least 1/3 width of frons, sometimes nearly touching medially and sometimes slightly narrower in female; less commonly as narrow as 1/4 width of frons. Fronto-orbital plate and most of ocellar triangle subshiny. Frons and eye bulging anterodorsally.

Chaetotaxy : Two widely spaced ori; two ors. Orbital setulae relatively dense, long and bushy, especially in males where setulae usually exceed length of setulae on eye; in up to three or four rows; mostly lateroclinate, reclinate to erect, inner rows partially proclinate; brownish yellow. Eye narrowly pilose dorsomedially in male, nearly bare in female; relatively sparse on both. Two dorsocentral setae. Acrostichal setulae in eight irregular rows. Mid tibia with two posteromedial setae.

Colouration : Body, including halter, dark brown in base colour. Notum with light greenish shine, sometimes bluish. Calypter margin and hairs white. Abdomen (excluding tergite 1) strongly coppery or metallic green, but sometimes hints of both colours present; sometimes bluish in male or with stronger metallic blue shine in female.

Genitalia : (Figs 318–323) Epandrium with regularly developed to slightly elongate posterodistal spine. Surstylus narrow, produced and angled posteroventrally, with two or three irregular rows of tubercle-like setae that sometimes extend along entire length of surstylus or extend nearly to base in one to two rows; apical setae very elongate and either clustered apically or (more frequently) extended in irregular row to midpoint of surstylus. Basiphallus slightly separated from phallophorus, U-shaped with dorsal section straight; overlapping or nearly overlapping distiphallus. Mesophallus inserted before midpoint of distiphallus; base exceeding that of distiphallus. Distiphallus teardrop-shaped with ventrodistal surface dark, plate-like and with margins slightly to strongly sinuate (especially pronounced in illustrated male from Chicago); internal spinulose structure well-developed; ventrolateral tubules small, recessed.

Variation : (Figs 62, 63) Lectotype female, DC male and Falls Church, VA male with fronto-orbital plate only 1/4 width of frons, and consequently with fewer rows of setulae; males with orbital setulae as long as setulae on eye. Female lectotype with seta on hind tibia on outer surface at basal 1/3 and abdomen with unusually pronounced blue metallic shine.

Hosts

AsteraceaeEupatorium capillifolium, Heterotheca subaxillaris.

Distribution

Canada: BC, ON, NB*, QC. USA: DC, DE*, FL, IL, ID, MA, MD, NC*, NJ, NY*, PA, SC, TN*, VA*.

Type material

Lectotype [virens]: USA. PA: “Penn”, “Loew coll”, “virens m.” (1♀, MCZ; type No. 15703).

Holotype [heterothecae]: USA. FL: Hialeah, 18.vii.1962, ex. stem of Heterotheca subaxillaris (Lam.) Britt. And Rusby, C.E. Stegmaier (1♂, originally deposited in USNM). [Missing]

Additional material examined

Canada: NB: Sainte-Anne-de-Kent, 46°34'N, 64°47'W, 28–29.vii.2013, O. Lonsdale, CNC358532 (1♂, CNC). USA. DC: Washington, 17.viii.1913, A.L. Melander (1♂, USNM), Washington, E bank CandO canal nr. Ariz. Ave., 23.i.1970, D.M. Anderson, reared from larva in stem Vernonia noveboracensis, 25.ii.1970 (1♂, USNM), Washington, 20.i.1969, D.M. Anderson, ex. stem of Eupatorium rugosum [illegible], emerged indoors 5.iii.1969 (1♀, USNM), Washington, E Bank CandO Canal nr. Ariz. Ave., reared from larva in stem Vernonia noveboracensis, D.M. Anderson, 23.i.1970 (1♂, USNM), 28.iii.1970 (2♂ 1♀, USNM), 14.x.1969, reared 23.ii.1970 (3♀, USNM), Cropley, 20.x.1968, D.M. Anderson, ex. stem Vernonia noveboracensis, emerged indoors 16.ii.1969 (1♂, USNM), DE: Gumboro, 2.viii.1952, C. Sabrosky (1♂, USNM), Rehoboth, “4/8/41”, A.L. Melander (1♂, USNM), FL: Highlands Co., Archbold Bio. Sta., 13–19.iv.1970, W.W. Wirth (1♀, USNM), Cape Sable, 31.iii.1953, W.R.M. Mason (1♀, CNC), IL: Chicago, A.L. Melander, CNC358432 (1♂, CNC), MD: Glen Echo, 30.vii.1922, J.R. Malloch (1♂, USNM), Plummers Isl., 20.vii.1913, J.R. Malloch (1♂, USNM), NC: Carteret Co., Atlantic Beach, 3–4.ix.1986, G.F. and J.F. Steyskal (1♂ 1♀, USNM), Mongomery Co., 4mi SW of Ashton, Malaise trap, 26.v.1981, G.F. and J.F. Hevel (1♂, USNM), NY: Long Island, Cold Spring Harbor, A.L. Melander (1♀, USNM), Ithaca, 31.v.1914, A.L. Melander (1♂, USNM), Cld. Sp. Harb., L.I., A.L. Melander, CNC358430 (1♂, CNC), PA: Spring Br., 9.v.1945, DDT Expt (1♂, USNM), TN: Cocke Co., Twin Creeks Uplands Res. Lab, Great Smokey Mtn. N.P., 35°41.2'N, 83°30'W, 1900', 27.v.1999, S.D. Gaimari (1♂, USNM), VA: Shenandoah, Big Meadows, 3.vii.1939, A.L. Melander (1♂, USNM), Fairfax Co., Dead Run, 5.v.1915, R.C. Shannon, CNC358431 (1♀, CNC), Falls Church, 11.iv.1923, W,. Middleton (1♂, USNM).

Comments

Nearctic Melanagromyza with a medially swollen fronto-orbital plate were treated as either M. virens or M. splendida in most of the previous literature. Spencer later described M. virginiensis in Spencer and Steyskal (1986b), and although he did not note the presence of this character in the description or key, a distinctly widened fronto-orbital plate is indeed present. A fronto-orbital plate ¼ the width of the frons is also found in M. vernoniae, M. vernoniana, M. walleyi Spencer, and several undescribed Nearctic species.

In addition to the swollen fronto-orbital plates, Melanagromyza virens was defined by the possession of a distinct surstylus that was narrow, bent and with several long, stout apical setae, a character that led Spencer (1969) to include M. heterothecae as a junior synonym. Unfortunately the type specimens of M. heterothecae cannot be located, but the illustrations of the phallus in the original description (Spencer 1966b) agree with the structure of dissected M. virens from Florida to Canada, and the synonymy is here maintained. While the surstylus is partially diagnostic of the species, a similar (or identical) surstylus is found in other species, including M. matricarioides, which was incorrectly illustrated in Spencer (1969), and M. walleyi Spencer from Tennessee. The phallus of this latter species (Spencer and Steyskal 1986b: figs 44, 45) is narrower, widest past the middle and abruptly constricted subapically, and the internal spinulose structure is smaller.

The Albertan Melanagromyza bidenticola Sehgal (Sehgal 1971: figs 26–31) is also similar to M. virens, largely agreeing in external morphology and phallic structure. This species, however, is larger (2.8–3.0 mm ♂, 3.1–3.3 mm ♀), the ocellar triangle is longer, the eye is 4.9–5.9 × higher than the gena, the scutum is slightly more metallic and the overall pigment on the body is darker. With regards to the male genitalia, the surstylus is wider and not downturned, although the posterodistal setae are slightly thicker, the basiphallus is slightly thicker and the mesophallus is slightly longer (although this difference may be negligible).

Melanagromyza virginiensis Spencer

Figs 58, 59, 325–328

Melanagromyza virginiensis Spencer in Spencer and Steyskal 1986b: 248.

Description

(Figs 58, 59). Wing length 3.1–3.2 mm (♂), 2.8–3.3 (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.7. Eye height divided by gena height: 3.7–5.9. Clypeus broadly rounded. Ocellar triangle slightly longer than wide. Fronto-orbital plate ¼ width of frons, widest near middle. Fronto-orbital plate and ocellar triangle subshiny.

Chaetotaxy : Two widely separated ori; two ors. Eye extensively pilose, with hairs sparse excluding dense dorsal patch. Orbital setulae erect to slightly proclinate, and mostly lateroclinate. Two dorsocentral setae. Acrostichal setulae in eight irregular rows. Mid tibia with two posteromedial setae.

Colouration : Body, including halter, dark brown in base colour. Notum with light bluish tint anteriorly (often absent) and more intense greenish shine posteriorly. Calypter margin and hairs white. Female anepisternum with greenish tint. Abdomen (excluding tergite 1) metallic green.

Genitalia : (Figs 325–328) Epandrium with small posterodistal spine. Surstylus 1/2-length of epandrium and with several rows of tubercle-like setae on inner surface. Hypandrium subtriangular with short apical process. Metepiphallus smooth ventrally and with short ridge on separate lateral sclerite. Proepiphallus narrow and weakly sclerotised. Basiphallus U-shaped and relatively long. Distiphallus separated from phallophorus by height of basiphallus; base of distiphallus extending slightly past base of closely held mesophallus; large, stout, and well-developed with especially long, broad, high distal section. Ejaculatory apodeme large and well-developed.

Hosts

AsteraceaeRudbeckia laciniata*.

Distribution

KS*, MA*, MD*, NH*, NY*, PA*, VA.

Type material

Holotype: USA. VA: Montgomeny Co., Blacksburg, 28.v.1962, J.G. Chillcott, CNC358457 (1♂, CNC).

Additional material examined

USA. KS: Lawrence, J.M. Aldrich (1♂, USNM), MA: Boston, May, A.L. Melander (1♀, USNM), MD: Glen Echo, 28.v.1919, J.M. Aldrich (1♀, USNM), Plummers Isl., at light, 28.iv.1914, R.C. Shannon (1♀, USNM), Chain Bridge, 12.ix.1913, R.C. Shannon (1♀, USNM), Montgomery Co., Bethseda, 3.vi.1972 (5♂, USNM), 28.v.1986 (1♀, USNM), 3.vi.1986 (2♀, USNM), 6.ix.1981 (1♂, USNM), Howard Co., Fulton, 21.v.1989, M.J. and R. Molineaux, J.E. Creeden (1♂, USNM), NC: Carteret, Co., Atlantic Beach, 3–4.ix.1984, G.F. and J.F. Hevel (1♀, USNM), NH: Franconia Ntch, 8.vii.1931, A.L. Melander (2♀, USNM), NY: Ithaca, 31.v.1914, A.L. Melander (1♂ 2♀, USNM) , New York City, “vCortlndPk”, 15.v.1926, A.L. Melander (1♀, USNM), PA: Pittsburg, McCandless Township, Allegheny Co., J. Plakidas, 20.ix.1978, “fly larva feed on gall tissue” (1♂, USNM), VA: Glencarlyn, 12.vi.1920, J.R. Malloch (1♂, USNM), Glencarlyn, 21.viii.1929, emg., “8-9.29”, Rudbeckia gall, J.C. Bridwell (1♂, USNM), nr. Plummers Isl., 19.x.1914, R.C. Shannon (1♀, USNM), Montgomery Co., Bethseda, 3.vi.1972, G. Steyskal (2♀, USNM), Talbot Co., McDaniel (Wades Point), 19–21.ix.1986, Malaise trap, salt marsh with flowering Baccharis, W.E. Steiner (1♀, USNM), Northampton Co., Kiptopeke, 4–6.x.1986, W.E. Steiner et al. (1♂ 1♀, USNM).

Comments

Melanagromyza virginiensis has widely separated ori, as in all other species of the M. virens group, and the fronto-orbital plate is only ¼ the width of the frons. The phallus is most diagnostic, with the distiphallus being especially long and stout-bodied with the distolateral margins nearly parallel and the ventrolateral tubules pronounced.

While the host species is not mentioned on any of the specimen labels, the male collected by Plakidas was reared from Rudbeckia laciniata (Plakidas 1982). It is interesting that this Melanagromyza atypically induces a gall, as apparent from both Plakidas’ specimen and the reared specimen from Virginia.

Ophiomyia Braschnikov

Ophiomyia Braschnikov, 1897: 40 [as subgenus of Agromyza]. Type species: Agromyza maura Meigen, 1838 [misidentified as curvipalpis Zetterstedt], by monotypy. Hendel 1920: 114 [as genus]; Frick 1952a: 375; Spencer 1964: 775, 1969: 81; Spencer and Steyskal 1986b: 37; Černý 1994: 455; Lonsdale 2014: 486.

Tylomyza Hendel, 1931: 181 [as subgenus of Ophiomyia]. Type species: Madiza pinguis Fallén, 1820, by original designation. Enderlein 1936a: 179 [as genus]. Syn. Spencer (1964a).

Stiropomýza Enderlein, 1936a: 179. Type species: Phytomyza aeneonitens Strobl, 1893, by monotypy. Syn. Frick (1952a) [not explicit].

Siphonomyza Enderlein, 1936a: 179. Type species: Agromyza proboscidea Strobl, 1900, by monotypy. Syn. Frick (1952a) [not explicit].

Aulomyza Enderlein, 1936a: 179. Type species: Melanagromyza longilingua Hendel, 1920, by monotypy. Frick 1952a: 375 [as synonym of Melanagromyza]. Syn Spencer (1966a) [as synonym of Ophiomyia].

Siridomyza Enderlein, 1936a: 179. Type species: Ophiomyia madizina Hendel, 1920 [=A. nasuta Melander], by monotypy. Syn. Frick (1952a) [as synonym of Tylomyza, not explicit].

Solenomyza Enderlein, 1936a: 179. Type species: Melanagromyza rostrata Hendel, 1920, by monotypy. Frick 1952a: 375 [as synonym of Melanagromyza]. Syn Spencer (1966a) [as synonym of Ophiomyia].

Stirops Enderlein, 1936a: 179 [nomen nudum – no type species designated].

Stirops Enderlein, 1936b: 42 [attributed to Enderlein 1936a]. Type species: Ophiomyia submaura Hering, 1926, by original designation [Enderlein 1936b: 42]. Syn. Frick (1952a) [not explicit].

Triopisopa Enderlein, 1936a: 179 [nomen nudum – no type species designated].

Triopisopa Enderlein, 1936b: 42. Type species: Agromyza simplex Loew, 1869, by original designation. Frick 1952a: 375 [as synonym of Melanagromyza]. Syn. Spencer (1966a) [as synonym of Ophiomyia].

Hexomyza Enderlein, 1936a: 179 [nomen nudum – no type designation].

Hexomyza Enderlein, 1936b: 42 [attributed to Enderlein (1936a)]. Type species: Melanagromyza sarothamni Hendel, 1923, by original designation. Hendel 1936: 570 [as synonym of Melanagromyza – followed by Frick (1952a)]; Spencer 1966a: 38, 1969: 79; Spencer and Steyskal 1986b: 34. Syn. Lonsdale (2014).

Carinagromyza Sasakawa, 1954: 23. Type species: Carinagromyza heringi Sasakawa, 1954 [= Ophiomyia sasakawaiSpencer and Martinez 1987], by original designation. Spencer and Martinez 1987 [synonymy].

Penetagromyza Spencer, 1959: 253. Type species: Penetagromyza aloes Spencer, 1959, by original designation. Spencer 1990: 390, 1991: 57. Syn. Lonsdale (2014).

Kleinschmidtimyia Spencer, 1986: Spencer, 1986: 249. Type species: Melanagromyza pisi Kleinschmidt, 1961, by original designation. Spencer 1990: 391. Syn. Lonsdale (2014).

Ophiomyia is a diverse, widespread genus sometimes mistaken for other Agromyzinae such as Melanagromyza and Euhexomyza, but many species exhibit unusual diagnostic external features and varied male genitalic morphology. All examined Nearctic species and virtually all global species, can be diagnosed by an apically truncated clypeus, as discussed in Lonsdale (2014). While the clypeus may sometimes be bulging or broadly rounded, the anterolateral margins are angulate, not rounded. The clypeus is usually also very thin and elongate, with the arms frequently bowed outwards, being particularly pronounced in those species with an anteriorly produced gena. Although several Melanagromyza such as M. buccalis and the new species M. glyptos approach the derived state seen in Ophiomyia, their relationship with the rest of Melanagromyza is revealed by dorsally pilose eyes, ventrolateral tubules on the distiphallus and a symmetrical basiphallus.

Other useful diagnostic characters of Ophiomyia are as follows: if present, there is only a single posteromedial seta on the mid tibia (usually two in other Agromyzinae); the calypter is usually brown marginally (white in most other Agromyzinae); there is usually a medial vertical carina separating the antennal bases and the centre of this carina usually also has a medial swelling that is spindle-shaped to subspherical. The gena is also often produced anteriorly, at least slightly, and can be strongly produced with an apical fasciculus. This fasciculus is an aggregation of multiplicated vibrissae that are variably fused to produce a “horn”. In many species, the inner lobe of the hypandrium is differentiated into two distinct sclerites (but see O. simplex): a setulose, arched sclerite (also found in Melanagromyza and Euhexomyza) and a flat subovate sclerite. The proepiphallus is also expanded laterally to form one pair of upcurved, strongly pigmented lobes (plate-like in O. simplex), the metepiphallus usually has one pair of ventromedial spines (not multiple spines, as is characteristic of most Melanagromyza), and the sclerites of the basiphallus are usually fused basally with the left sclerite atrophied. Lastly, the distiphallus is usually somewhat asymmetric, being slightly twisted sinistrally, and there are no paired ventrolateral tubules (characteristic of Melanagromyza).

Key to the species of Delmarva Ophiomyia

1 Orbital setulae proclinate (Figs 64, 65). Three pairs of dorsocentral setae. Male without ors (Fig. 65); female with one or two ors (Fig. 64). Distiphallus with minutely tuberculate bilobed distal section (Figs 379, 380). Metepiphallus with multiple pairs of ventromedial spines O. nasuta (Melander)
Orbital setulae reclinate. Two pairs of dorsocentral setae. Two pairs of ors. Distiphallus not as above. Metepiphallus almost always with one pair of ventromedial spines 2
2 Costa extending just past vein R4+5. Gena highest behind midpoint of eye (Fig. 70). Fronto-orbital plate and parafacial shiny, prominent, and easily viewed laterally. Bands of basiphallus twisted counter-clockwise (Figs 392, 393). Inner setulose band of hypandrium fused to lobe (Fig. 390) O. simplex (Loew)
Costa extending to vein M1+2. Gena highest anterior to midpoint of eye. Fronto-orbital plate and usually parafacial barely visible laterally. Bands of basiphallus either fused and collar-like basally, or with right band produced. Inner band of hypandrium separate from lobe 3
3 Clypeus broad and nearly parallel-sided along length (Fig. 71). Face with weak carina dividing antenna, if present (Fig. 71). Gena not produced, vibrissal angle > 90° (Fig. 72). Male without vibrissal fasciculus. Distiphallus small, bulb-like (Fig. 331). Basiphallus basal and collar-like 4
Clypeus usually slightly to strongly narrowed anteriorly (Fig. 77). Face with carina (sometimes indistinct) with medial swelling below antennal base (Fig. 75). Gena at least slightly projecting, vibrissal angle 90° or less (Fig. 76). Male with fasciculus. Distiphallus relatively dark, large, and globular/ovate in shape (Figs 358, 387). Basiphallus produced on right side 7
4 Gena entirely receding along anterior margin. Distiphallus darker and subcylindrical, without flagellum 5
Gena slightly angled on anterior margin, not immediately vertical or receding (similar to Melanagromyza spp.). Distiphallus pale and bulb-like with terminal clear flagellum 6
5 Eight rows of acrostichal setae. Wing length 2.5 mm. Distiphallus broad and parallel-sided apically (Fig. 406) O. ultima (Spencer)
Six rows of acrostichal setae. Wing length 2.0–2.3 mm. Distiphallus and mesophallus narrow, linear and with bell-shaped apex (Figs 353, 354) O. kalia sp. nov.
6 Eye 5.0–6.7 × higher than gena. Two ori. Fronto-orbital plate slightly visible laterally. Distiphallus relatively small and spade-shaped, with small distolateral sclerotisations and no ventral elaboration of membrane; apical flagellum directed distally (Figs 330, 331) O. abutilivora Spencer
Eye 2.5–4.3 × higher than gena. Often three ori, at least on one side, with orbital seta between ori variably developed. Fronto-orbital plate and parafacial projecting, with parafacial continuing under eye as distinct cheek. Distiphallus rounded, with large ventral carina, and often with pronounced lateral "wings”; apical flagellum directed ventrally (Figs 401, 402) O. tiliae (Couden)
7 Facial bulb with extensive shiny section on dorsal 1/2; sides of carina above bulb usually parallel-sided basally. Ocellar triangle shiny, less commonly subshiny. Abdomen with faint metallic shine (often difficult to see). Mid tibia with single posteromedial seta 8
Facial bulb and ocellar triangle not shiny (shiny in some O. coniceps, but these with very pronounced genal process that is longer than high, and male with very short, wide fasciculus); sides of facial carina above bulb immediately divergent. Abdomen brown to glossy black, rarely coppery (O. cuprea). Mid tibia with or without posteromedial seta 10
8 Eye 4.0–6.0 × higher than gena. Ocellar triangle attaining anterior margin of frons as thin line. Surstylus narrow, ~ 1/3 length of epandrium. In profile, distiphallus widest subapically (Figs 358, 359) O. kwansonis Sasakawa
Eye 8.0–8.4 × higher than gena. Ocellar triangle not attaining anterior margin of frons, and usually not distinct anterior to ors. Surstylus ~ 1/2 length of epandrium. In profile, distiphallus widest near base 9
9 Three ori sometimes present on one or both sides of frons. Facial bulb smooth or only slightly furrowed (Fig. 78). Phallus with strong distoventral membranous keel. Basiphallus short, not reaching mesophallus. Distiphallus with inner surface tuberculate to weakly spinulose distally, and more strongly spinulose basally; shape subovate (seen ventrally); dorsum smoothly sloping to apex (seen laterally) (Figs 365, 366) O. labiatarum Hering
Only two ori. Facial bulb with strong medial furrow. Phallus with weak, sac-like distoventral membrane. Basiphallus reaching past base of mesophallus. Distiphallus with medial lobe strongly spinulose internally; shape elongate with base tapered (seen ventrally); distal 1/2 abruptly flattened (seen laterally) (Figs 411, 412) O. virginiensis Spencer
10 Body brown to light brown with fronto-orbital plate and ocellar triangle (excluding tubercle) white/beige. Frons with anteromedial furrow. Wing veins whitish. Eye nearly as wide as high, with long axis nearly vertical. Basiphallus extensively sclerotised along right margin, strongly bent ventrally, and membrane thickened and lightly sclerotised distoventrally (Fig. 346) O. galiodes sp. nov.
Body dark brown with parts of frons sometimes slightly paler, but never whitish. Frons evenly rounded anteriorly. Wing veins light brown to brown. Eye distinctly higher than wide, with long axis often diagonal. Phallus not as above 11
11 Five to six rows of acrostichal setulae. Abdomen brown with weak coppery luster. Distiphallus relatively dark and elongate with distal 1/2 tapering and left-distal surface minutely tuberculate; with long, dark, upcurved and pointed posteromedial spur (Figs 344, 345) O. cuprea sp. nov.
Eight to ten rows of acrostichal setulae. Abdomen brown to black. Distiphallus not as above; if with posteromedial spur, then spur short, straight, and blunt 12
12 Three ori. Genal angle 70°–90°. Mid tibia with single weak posteromedial seta. Notopleuron and postpronotum reddish or eye 2.8 × higher than gena. Distiphallus widest at midpoint and broadly rounded (viewed ventrally), with surface strongly ridged and furrowed (Figs 386, 387) O. sexta Spencer
Two ori. Genal angle often < 60° or 45°, rarely 90° (O. heleios). Mid tibia usually without posteromedial setae. Notopleuron and postpronotum the same colour as the rest of the scutum and gena not particularly high. Shape of distiphallus variable, but not as above 13
13 Vibrissal fasciculus shorter than length of genal process (Figs 75–77). Parafacial prominent. Mesophallus with small dark transverse band across point of insertion with distiphallus; distiphallus with small to large lobe projecting from left ventral margin (Figs 340, 341) O. coniceps (Malloch)
Fasciculus longer than genal process. Parafacial barely visible laterally. Mesophallus without transverse band; distiphallus without lobe as above 14
14 Genal process short, with dorsal margin straight (not concave), forming a 60°-90° angle (as in Fig. 79). Distiphallus with strong ventromedial sclerotisation (broad to thin and band-like); inner surface of distiphallus lightly spinulose and outer surface smooth 15
Genal process of variable length, but dorsal margin curving to a narrow point (as in Fig. 76). Distiphallus not excessively sclerotised distoventrally; either inner surface covered with numerous stout spines or outer surface minutely tuberculate apically 16
15 Genal process forming a slightly < 90° angle. Wing length 2.5 mm. Notum dark brown with light green reflection (reflection absent on postpronotum and notopleuron). Base of femora paler, yellow to base ventrally on mid and hind legs. Calypter white with hairs brown. Distiphallus with thick distoventral cluster of ridges (Figs 349, 350) O. heleios sp. nov.
Gena strongly produced, forming an approximate 45° angle. Wing length 1.7 mm. Notum and femora dark. Calypter margin and hairs brown. Distiphallus with thin distoventral ridge (Fig. 336) O. capitolia sp. nov.
16 Scutum bare medially behind anterior dorsocentral. Surstylus with 4 apical tubercle-like spines (Fig. 395). Distiphallus relatively thin and C-shaped in lateral view; posterodorsal margin with tail-like extension (Fig. 397) O. texana (Malloch)
Scutum setulose medially behind anterior dorsocentral, at least partially. Surstylus with numerous apical spines. Distiphallus stout and irregular in lateral view; without tail-like process 17
17 Wing length 2.2–2.3 mm. Mid tibia without posteromedial setae. Facial bulb narrow and elongate, not much wider than remainder of keel. Distiphallus pale, subquadrate (ventral view), densely spinulose on inner surface (Figs 368, 369) O. laticolis sp. nov.
Wing length 1.8–2.0 mm. Mid tibia with one posteromedial seta. Facial bulb broadly rounded. Distiphallus longer than wide, with short, but distinct tubule emerging dorsoapically; inner surface smooth (Figs 375, 376) O. maura (Meigen)

Species descriptions

Unless otherwise stated, the following species are assumed to be as follows: Body, including halter, brown. Orbital setulae reclinate. Notum subshiny with two dorsocentrals and eight to ten rows of acrostichal setulae. Costa extending to vein M1+2. Hypandrium differentiated into two distinct sclerites: a setulose, arched sclerite and a flat subovate sclerite. Proepiphallus expanded laterally to form one pair of upcurved, strongly pigmented lobes.

Ophiomyia abutilivora Spencer

Figs 71, 72, 329–331

Ophiomyia abutilivora Spencer in Spencer and Steyskal 1986b: 250; Eiseman et al. 2021: 16.

Description

(Figs 71, 72). Wing length 2.0–2.3 mm (♂♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6. Eye height divided by gena height: 5.0–5.7. Facial carina present with narrow medial bulb. Gena slightly angled forward, highest near midpoint of eye. Clypeus slightly tapering apically. Fronto-orbital plate very narrow, ocellar triangle nearly extending to posterior ori. Notum shiny.

Chaetotaxy : Orbital setulae reclinate. Male vibrissal fasciculus absent. Two ori; two ors. Mid tibia with one posteromedial seta.

Colouration : Body, including halter dark brown with gena paler. Wing veins yellow to brown. Calypter margin and hairs brown.

Genitalia : (Figs 329–331) Inner posterodistal margin of epandrium with rows of minute tubercles. Distal margin of surstylus broad and truncated with rows of pointed tubercle-like setae on inner surface. Metepiphallus with subapical spine. Basiphallus with slender dorsoventral arms, partially fused to phallophorus. Hypophallus membranous, carinate. Distiphallus small and spade-shaped (broad basally and strongly tapered apically); apex with long, tapered flagellum that has base surrounded by light sclerotisations; mesophallus narrow, barely wider than duct. Ejaculatory apodeme stout, well-developed, with blade relatively narrow.

Hosts

MalvaceaeAbutilon theophrasti, A. permolle. Adults taken from Sida cordifolia (Spencer and Steyskal 1986b) and possibly Alcea.

Distribution

USA: DE*, FL, IA, MD*, MN, MS; stem mines only in IL and WI (Eiseman et al. 2021).

Type material

Holotype: USA. MN: Dakota Co., Rosemount, Agricultural Research Station, 13.ix.1978, R.N. Andersen and R. Ralston (1♂, USNM).

Paratypes examined

. USA. FL: Broward Co., Ft. Lauderdale, 14.iii.1980, H.E. Walker, Fl. Abutilon permolle, MN: Rosemount, 13.ix.1987, ex. Abutilion theophrasti (1♂ 3♀, USNM), MS: Stoneville, 22.x.1979, K.E. Frick, 79-25, ex. stem gall Abutilion theophrasti (4♂ 1♀, USNM), Merigold, 22.x.1979, K.E. Frick, 79-25, ex. stem gall Abutilion theophrasti (1♂ 2♀, USNM).

Additional material examined

USA. DE: Newark, 6.ix.1960, hollyhock, P. Burbutis (2♂, USNM), Newark, “8/10/1953” (1♂, USNM), IA: Winneshiek Co., Decorah, Trout Run Trail, 43°18'5.04"N, 91°48'6.60"W, 10–13.viii.2017, J. van der Linden, Abutilion theophrasti, em. by 26.vii.2017, #CSE4940, CNC1643668, CNC1643669 (1♂ 1♀, CNC), MD: Montgomery Co., 4mi SW of Ashton, 6.ix.1981, Malaise trap, G.F. and J.F. Hevel (1♂, USNM).

Comments

The distiphallus of Ophiomyia abutilivora is highly reduced and lobate, with the apex continuing distally as a long membranous flagellum. A similar phallus is found in the larger O. tiliae, but the clypeus of this species is nearly rounded, the distiphallus appears pear-shaped when viewed ventrally, the apical flagellum points ventrally and there is one pair of membranous "wings” ventral to the distiphallus. Other putative Nearctic relatives (based on the genitalic figures in Spencer and Steyskal 1986b and Spencer 1969) are more western in distribution, including the Californian species O. bernardensis Spencer, O. shastensis Spencer, O. jacintensis Spencer, and O. yolensis Spencer, and possibly O. monticola Seghal from northern Canada and Alaska.

Ophiomyia capitolia sp. nov.

Figs 332–336

Description

Wing length 1.7 mm (♂). Female unknown. Length of ultimate section of vein M4 divided by penultimate section: 0.8. Eye height divided by gena height: 6.2. Eye slightly angled diagonally. Facial carina stout, slightly more narrow than stout bulb. Gena shallowly produced, projecting at 60° angle with margins straight. Clypeus narrow and truncated. Buccal cavity narrowed anteriorly with anterior margin straight. Distance between crossveins more than length of dm-m. Ocellar triangle and fronto-orbital plate (narrow) subshiny and ill-defined.

Chaetotaxy : Male with vibrissal fasciculus ~ 1/2 length of gena. Two ori; two ors. Mid tibia with one posteromedial seta.

Colouration : Body, including halter dark brown. Wing veins brown. Calypter margin and hairs brown.

Genitalia : (Figs 332–336) Metepiphallus small, pale and with one pair of ventromedial spines. Epiphallic lobes clear with dark floating sclerite. Halves of basiphallus broadly fused at base with remainder of left sclerite absent; right sclerite relatively short and truncated with long distoventral process reaching venter. Mesophallus continuing distally as thick ridge that bifurcates apically; distiphallus pale and globular with inner surface minutely spinulose. Ejaculatory apodeme with long dark stalk and several annulations on striated blade; sperm pump with weak sclerotisation.

Host

Unknown.

Distribution

USA: DC.

Etymology

The specific name refers to the fact that the type was collected in the United States capitol.

Type material

Holotype: USA. DC: Washington, 17.viii.1913, A.L. Melander (1♂, USNM).

Comments

Ophiomyia capitolia is one of the smaller Delmarva Ophiomyia with a male wing length of 1.7 mm. This species also has a distinct genal process, a long fasciculus and a relatively distinct wrinkle under the eye within the cheek. Only examination of the male genitalia can provide reliable identification of this species (see key).

Ophiomyia coniceps (Malloch)

Figs 75–77, 82, 337–341

Agromyza coniceps Malloch, 1915a: 107.

Ophiomyia coniceps. Frick 1952a: 382; Spencer 1969: 85, 1981: 75; Spencer and Steyskal 1986b: 46; Shi, Chunyan and Gao 2015: 59; Eiseman and Lonsdale 2018: 19.

Description

(Figs 75–77, 82). Wing length 1.7–1.9 mm (♂), 2.4 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.9–1.0. Eye height divided by gena height: 6.8. Facial carina distinct with large medial bulb; carina as broad as bulb dorsally with sides diverging. Eye angled diagonally. Gena strongly produced anteriorly, forming an ~ 45° angle. Parafacial also strongly produced, visible laterally, and blending into gena. Fronto-orbital plate narrow. Clypeus strongly produced and narrowed anteriorly, with apex slightly expanded; sides bowed laterally, widest near base. Sides of ocellar triangle slightly concave; triangle subshiny but sometimes glossier. Crossveins separated by length of dm-m or less.

Chaetotaxy : Male vibrissal fasciculus short and upcurved, not longer than genal process. Two ori (not three, as stated in Spencer and Steyskal 1986b); two ors. Mid tibia without posteromedial setae.

Colouration : Body, including halter dark brown. Fronto-orbital plate and ocellar triangle reddish in DE male. Calypter margin and hairs dark brown. Abdomen sometimes with faint coppery/golden shine. Wing veins brown.

Genitalia : (Figs 337–341) Surstylus broad, rounded, short, slightly produced ventrally, inner-distal surface with two to three rows of tubercle-like setae. Hypandrium pointed apically, sides slightly bowed, basal arms sinuate. Postgonite typical, narrow with dorsal 1/2 weakly attached and dorsally directed. Phallophorus dark and constricted at base, narrow, venter longer, swollen and produced. Basiphallus with arms united at base, left arm short, weak, and ill-defined, right arm long, reaching level of mesophallus. Mesophallus cylindrical, clear, inserted ventromedially into distiphallus. Distiphallus, large, mostly empty with medial shelf-like process that is slightly angled apically; somewhat ovate in ventral view with basal 1/2 narrower and slightly longer; inner surface minutely textured basally, with longer spinules medially, apically with shallow tubercles and striations to left side and longer rounded spinules to right side; mostly open dorsally; left ventrodistal margin produced as characteristic lobe that may be modestly to strongly developed.

Host

AsteraceaeAntennaria plantaginifolia and possibly other Antennaria; Sonchus asper (Eiseman and Lonsdale 2018).

Distribution

Canada: BC, MB, ON, QC, SK. USA: CA, DE*, IN, LA, MA, OK, UT, VA*; possibly also AL, CT, IA, WI (Eiseman and Lonsdale 2018).

Type material

Holotype: USA. UT: Salt Lake, 14.viii.1914, ex. Sonchus asper, P.H. Timberlake (1♀, USNM; type No. 193904).

Additional material examined

Canada. MB: Aweme, 2.vi.1916, N. Criddle, J.M. Aldrich (1♂, USNM). USA. DE: Newark, 1.vii.1974 (1♂, USNM), IN: Evansville, 7.v.1914, J.M. Aldrich (1♂, USNM), LA: Lake Charles, 9.vi.1917, J.M. Aldrich (1♂, USNM), MA: Hampshire Co., Granby, Mt. Norwottuck, 27.iii.2012, em. 11.iv–24.iv.2012, C.S. Eiseman, ex Antennaria plantaginifolia (5♂ 3♀, CNC), OK: Payne Co., Mehan, 36.014339°N, 96.996744°W, 10.xii.2015, em. 15.i.2016, M.W. Palmer, ex Antennaria plantaginifolia, #CSE2214, CNC653970–653972 (2♂ 1♀, CNC), 12.xii.2015, em. by 12.i.2016, M.W. Palmer, ex Antennaria plantaginifolia, #CSE2216, CNC653988 (1♂, CNC), 14.i.2016, em. by 10–16.ii.2016, M.W. Palmer, ex Antennaria plantaginifolia, #CSE2237, CNC654000–654002 (3♂, CNC), 27.ii.2017, em. 18.iii– 15.vi.2017, C.S. Eiseman, ex Antennaria plantaginifolia, #CSE3229, CNC939913–939917 (3♂ 2♀, CNC), VA: Luray, 24.vi.1933, A.L. Melander (1♂, USNM), Page Co., 7mi W of Lunay, 8.vii.1978, G.F. Hevel (1♂, USNM), Shenandoah, Big Meadows, 3.vii.1939, A.L. Melander (1♂, USNM).

Comments

Ophiomyia coniceps will key to O. apta Spencer in Spencer and Steyskal (1986b: figs 224–226) because the number of ori in that work is incorrectly recorded as three, not two. This can no longer be directly verified from the holotype, however, since the head is missing, but the original illustration and description in Malloch (1915) clearly indicates two ori. Ophiomyia coniceps has the most pronounced genal process of any Delmarva species, a strongly projecting parafacial and most important for diagnostic purposes, a very short and sometimes wide vibrissal fasciculus that is usually sharply pointed. The male genitalia are similar to those of other Ophiomyia with a fasciculus, but the outline of the distiphallus, the longer inner-medial spinules, and especially its small left ventrodistal lobe are unique.

Ophiomyia cuprea sp. nov.

Figs 342–345

Description

Wing length 2.2 mm (♂). Female unknown. Length of ultimate section of vein M4 divided by penultimate section: 0.9. Eye height divided by gena height: 9.3. Facial carina relatively long below, nearly as wide as bulb and with diverging margins above. Clypeus narrow. Gena slightly projecting, forming almost a 60° angle. Ocellar triangle subshiny around ocelli. Fronto-orbital plate thin and subshiny. Buccal cavity narrowed anteriorly with anterior margin straight. Distance between crossveins more than length of dm-m.

Chaetotaxy : Male with vibrissal fasciculus ~ 2/3 length of gena and strongly upcurved. Two ori on right side, three on left; two ors. Setae on mid tibia not visible, possibly absent. Five to six rows of acrostichal setulae.

Colouration : Body, including halter dark brown. Calypter margin and hairs brown. Abdomen with weak coppery shine. Veins light brown.

Genitalia : (Figs 342–345) Surstylus ~ 1/2 length of epandrium, rounded marginally and with tubercle-like setae on inner surface. Metepiphallus pale with one pair of ventromedial spines. Epiphallic lobe clear with dark sclerite. Halves of basiphallus broadly fused at base with remainder of left sclerite absent; right sclerite apically broad and truncated. Distiphallus dark and well-sclerotised with distinct basal and distal sections – basal section high, tapered to base, minutely spinulose internally and with internal dorsomedial hook; distal section shallower, with left margin receding and minutely tuberculedridged, and right 1/2 longer, internally spinulose and ill-defined apically.

Host

Unknown.

Distribution

USA: MD.

Etymology

The specific epithet is derived from the Latin for coppery, referring to the colour of the abdomen, which has a weak metallic lustre.

Type material

Holotype: USA. MD: nr. Plummers Isl., 5.v.1915, R.C. Shannon (1♂, USNM).

Comments

The head (particularly the gena) and genitalia of Ophiomyia cuprea are similar to those of the Californian O. delecta Spencer (Spencer and Steyskal 1986b: figs 271, 272), but O. delecta differs in having eight rows of acrostichal setulae (not five to six), a black (not coppery) abdomen and a facial bulb that is much narrower than the pedicel (Spencer 1981). Furthermore, based on Spencer’s illustration, the distiphallus of that species is more broadly rounded basally and atrophied apically. Spencer (1981) also describes the similar O. definita Spencer (later provided the new name O. subdefinita Spencer; Spencer and Steyskal 1986b: figs 268, 269) in his Californian revision, but the fasciculus is straight (not curved), the abdomen is also black and the distiphallus is much paler; the number of rows of acrostichal setulae are not mentioned, but it is indicated that there are also eight.

Ophiomyia galiodes sp. nov.

Figs 346, 347

Description

Wing length 2.2 mm (♂), 1.9–2.2 (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–1.0. Eye height divided by gena height: 6.2–9.8. Facial carina (wide dorsally) and bulb distinct. Parafacial, seen laterally, broadly rounded and projecting past anterior margin of gena. Gena slightly produced anteriorly, forming an angle of ca. 70° and with dorsal and ventral margins straight, not curved. Clypeus broadest posteriorly; apex very narrow, produced. Buccal cavity narrowed anteriorly with margin straight. Ocellar triangle concave subapically. Frons slightly furrowed medially. Crossveins separated by more than length of dm-m.

Chaetotaxy : Vibrissal fasciculus well-developed, slightly upcurved. Two ori; two ors. Mid tibia without posteromedial setae.

Colouration : Body, including halter dark brown, but relatively pale compared to congeners. Fronto-orbital plate and outer margin of ocellar triangle beige. Wing veins white. Calypter margin light brown with hairs brown.

Genitalia : (Figs 346, 347) Surstylus slightly < 1/2 length of epandrium, with several marginal tubercle-like setae. Metepiphallus pale and largely desclerotised with one pair of small ventromedial spines. Proepiphallus pigmented medially, darker ventrally. Halves of basiphallus broadly fused at base with remainder of left sclerite absent; right sclerite relatively short and ill-defined; strongly bent ventrally (possibly an artifact of preservation). Distiphallus short, strongly spinulose internally and with base extending past that of mesophallus; membrane distal to phallus relatively broad, flat, and lightly sclerotised.

Host

RubiaceaeGalium.

Distribution

USA: MD.

Etymology

The specific name indicates the similarity of this species to its possible sister taxon, Ophiomyia galii, which also feeds on Galium.

Type material

Holotype: USA. MD: Beltsville, 16.iv.1975, S.W.T. Batra, ex. Galium sp. (1♂, USNM).

Paratypes: USA. MD: Same collection as holotype (1♀, USNM; 1♀, CNC).

Comments

The phallus of Ophiomyia galiodes (only illustrated in lateral view due to inflexibility of the structure) is similar to that of O. laticolis (Figs 368, 369), but the distiphallus is narrower, larger and closer to the apex of the basiphallus. Furthermore, the eye is larger, rounder and bulging anteriorly, and the shoulders, fronto-orbital plate and ocellar triangle are pale, although these colour characters may be an artifact of preservation.

Ophiomyia galiodes is also similar to the European O. galii Hering, which is the only other Ophiomyia known from Galium (Spencer, 1990). Their chaetotaxy, wing length and wing vein proportions are nearly the same, and they both have a ventrally curved hypandrium and similar distiphallus. The gena of O. galii is produced at a 45° angle (not 70°) (Spencer 1972), the basiphallus is longer and less spinulose, and the mesophallus is longer, narrower and extends past the base of the distiphallus (Papp and Černý 2015: fig. 152).

Ophiomyia heleios sp. nov.

Figs 348–350

Description

Wing length 2.5 mm (♂). Female unknown. Length of ultimate section of vein M4 divided by penultimate section: 0.9. Eye height divided by gena height: 7.5. Facial carina long and narrow with small subspherical bulb at midpoint. Gena not strongly produced, forming slightly < 90° angle. Clypeus narrow with posterior arms curved outwards. Ocellar triangle shiny black, longer than wide and tapering anteriorly.

Chaetotaxy : Orbital setulae reclinate. Male vibrissal fasciculus long, narrow, and slightly upcurved. Two ori; two ors. Mid tibia with one small posteromedial seta.

Colouration : Body, including halter dark brown, with gena paler, ocellar triangle black, notum (excluding postpronotum and notopleuron) with faint green reflection, and femora paler on basal 3/5 with mid and hind femora yellowish ventrally on basal 2/5. Calypter white with hairs dark brown.

Genitalia : (Figs 348–350) Surstylus narrow with few inner-marginal tubercle-like setae. Distal margin of hypandrium long and tapered. Metepiphallus very small and with one pair of spines. Halves of basiphallus very ill-defined apically, slightly fused at base and with left side shorter. Base of distiphallus strongly exceeding base of mesophallus, with short dorsal margin (curled inwards and with several very short inner spines) that is atrophied on left side; distoventral margin of distiphallus well-sclerotised medially, produced as apically converging ridges. Ejaculatory apodeme atrophied to base on one side, clear along margins and with well-developed medial rib.

Host

Unknown.

Distribution

USA: VA.

Etymology

Gr. “of a marsh, dwelling in a marsh”, referring to the collection locality of the type.

Type material

Holotype: USA. VA: Fairfax Co., Great Falls Park, swamp trail, 38°59.4'N, 77°15.2'W, Malaise trap, trap #2, 8.ix-11.v.2006, D.R. Smith (1♂, USNM).

Comments

Ophiomyia heleios can be differentiated from Nearctic congeners by a white calypter margin (although the hairs are still typically dark), a greenish notum, a short gena, a slightly atypical ejaculatory apodeme and an unusually atrophied distiphallus that has a thick distoventral sclerotisation medially.

Ophiomyia kalia sp. nov.

Figs 351–355

Description

Wing length 2.1 mm (♂), 2.0–2.3 (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.7–1.0. Head largely collapsed in holotype. Eye height divided by gena height: 5.0–6.6. Frons entirely pruinose and ocellar triangle indistinct; gena rounded, not produced anteriorly and relatively narrow along entire length. Face with weak carina, barely dividing antennae. Clypeus with sides parallel.

Chaetotaxy : Orbital setulae reclinate. Male vibrissal fasciculus absent. Two ori (sometimes three on one side); two ors. Mid tibia without posteromedial setae. Six rows of acrostichal setulae.

Colouration : Body, including halter dark brown, with gena slightly paler. Wing veins brown. Calypter margin and hairs brown.

Genitalia : Surstylus 1/2 length of epandrium, broadly rounded and with tubercle-like setae along inner-distal margin. Metepiphallus very small with serrated ridges and spines. Proepiphallus with dark patches in lateral extensions. Basiphallus curved in cross-section, ventrally curved and receding dorsomedially. Mesophallus broad, basal to distiphallus (not inserted ventromedially to distiphallus). Distiphallus straight, cup-like with internal folds. Ejaculatory apodeme large with blade pale and ovate, with pronounced medial rib.

Host

Unknown.

Distribution

USA: VA.

Etymology

The specific epithet is Greek for bird nest, referring to the nest from where the holotype was collected.

Type material

Holotype: USA. VA: Long Bridge, 22.v.1913, ex. birds nests Pun grackle, R.C. Shannon (1♂, USNM).

Paratypes: USA. VA: Fairfax Co., Turkey Run Park, nr. mouth of Turkey Run, 38°57.9'N, 7°09.4'W, Malaise trap, D.R. Smith, 18–30.v.2007 (1♂ 2♀, USNM; 2♀, CNC), 14–17.v.2006 (8♀, USNM; 2♂, CNC).

Comments

The straight, highly simplified distiphallus of this species is characteristic, as is the basally inserted mesophallus, the large, pale ejaculatory apodeme, and narrow gena. Also see comments for Ophiomyia abutilivora.

Ophiomyia kwansonis Sasakawa

Figs 356–359

Ophiomyia kwansonis Sasakawa, 1961: 355. Shiao and Wu 1999: 344; Matsumoto and Sasakawa 2006: 20; Williams and Steck 2011: 7, 2014: 421; Shi, Chunyan and Gao 2015: 61.

Description

Wing length 2.0–2.5 mm (♂♀). Length of ultimate section of vein M4 divided by penultimate section: 0.8–1.0. Eye height divided by gena height: 4.0–6.0. Facial carina distinct; bulb narrow, slightly wider than carina, with medial longitudinal furrow and dorsal 1/2 shiny. Clypeus strongly narrowed anteriorly. Anterior portion of buccal cavity narrowed, not much wider than clypeus. Gena slightly produced anteriorly, forming an ~ 80° angle. Notum subshiny. Ocellar triangle with sides convex, nearly meeting past midpoint and attaining anterior margin of frons. Fronto-orbital plate narrow and shiny, slightly widening around base of fronto-orbitals.

Chaetotaxy : Male vibrissal fasciculus thick, slightly upcurved and not longer than gena. Two ori; two ors. Mid tibia with small posteromedial seta past midpoint.

Colouration : Body, including halter dark brown to black, with nearly indistinct metallic sheen on abdomen, and with fronto-orbital plate and ocellar triangle paler. Wing veins brown. Calypter margin and hairs grey.

Genitalia : (Figs 356–359) Surstylus narrow with several scattered rows of tubercle-like setae at apex and inner-distal surface. Metepiphallus with single pair of posteromedial spines. Basiphallus sclerotised basally on dorsobasal and left lateral surfaces, and with long right-lateral process that is widened at apex and attaining level of base of distiphallus. Distiphallus with blunt subconical base that extends posteriorly past narrow mesophallus and is internally spinulose on ventral and left ventrolateral surfaces; widest near midpoint, distally with spinulose right lateral chamber and thin distomedial chamber with sclerotised ventral surface and irregular internal folds. Ejaculatory apodeme relatively narrow with distal margin of blade reduced, pale and irregular.

Host

AsphodelaceaeHemerocallis fulva.

Distribution

USA: AL, AR, CA, CT, DE, FL, GA, IL, IN, KA, KY, LA, MA, MD, MI, MO, MS, NC, NH, NY, OH, OK, PA, SC, TN, TX, VA, WI, WV. Japan, Taiwan. Slovenia.

Type material

Holotype: Japan. Honshu: Kyoto, Shimogamo, on Hemerocallis fulva kwanso, 21.v.1956, Sasakawa (1♂, Osaka Museum of Natural History). [Not examined]

Additional material examined

USA. MD: Anne Arundel Co., Davidsonville, 7.vii.2011, on Hemerocallis sp., G.L. Williams (1♂ 11♀ [in alcohol], CNC; 1♂ 1♀, CSCA).

Comments

Originally described from Japan as a leaf miner in the ornamental daylily Hemerocallis fulva kwanso, this fly is a recent invasive in North America (Williams and Steck 2011, 2014), and will soon likely occur in most regions where daylily is grown. Life history and immature stages are discussed in Sasakawa (1961) and Williams and Steck (2011). The only other agromyzid known to occur on Hemerocallis is Liriomyza hemerocallis Iwasaki, also from Japan, but this species feeds on seeds, not within leaves (Iwasaki 1993).

Ophiomyia labiatarum Hering

Figs 78–80, 360–366

Ophiomyia labiatarum Hering, 1937: 509. Spencer 1969: 87, 1976: 67; Spencer and Steyskal 1986b: 51; Černý 2018: 122.

Description

(Figs 78–80). Wing length 2.1–2.4 mm (♂), 2.4–2.5 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.7–1.0. Eye height divided by gena height: 8.0–8.4 (4.0 in one dissected VA male from Turkey Run). Facial carina and medial bulb distinct; carina as wide as bulb dorsally or slightly narrower, with sides usually parallel and never strongly diverging; bulb usually strongly to shallowly furrowed or heart-shaped. Clypeus narrowed and produced anteriorly. Anterior margin of buccal cavity narrowed and straight or rounded. Eye slightly angled diagonally. Gena slightly produced, forming an angle slightly < 90°. Ocellar triangle and fronto-orbital plate subshiny. Notum shiny.

Chaetotaxy : Male vibrissal fasciculus distinct, nearly as long as gena, but only forming a discrete point in male with three ori on both sides. Two ori (sometimes three on one side, rarely on both sides); two ors. Mid tibia with one posteromedial seta.

Colouration : Body, including halter dark brown. Abdomen sometimes with faint coppery/golden shine. Wing veins brown. Calypter margin and hairs dark brown.

Genitalia : (Figs 360–366) Metepiphallus with one pair of ventromedial spines. Ventral subapical membrane around phallus pronounced and keel-like (less developed in male with three ori on both sides). Basiphallus with sclerotised dorsobasal plate and right lateral band. Base of mesophallus and distiphallus level. Distiphallus with small transverse sclerite at base of mesophallus; distiphallus subovate (slightly flattened apically) with inner basal margin spinulose and inner distal margin with shallow bumps; apex ill-defined.

Hosts

LamiaceaeCalamintha, Clinopodium, Galeopsis, Lamium, Leonurus, Nepeta, Prunella, Salvia, Satureja, Scutellaria, Stachys (Spencer 1976; Spencer and Steyskal 1986b; Benavent-Corai et al. 2005). AsteraceaeSolidago canadensis*.

Distribution

Canada: AB, NB*, ON, QC. USA: DE*, IN, MD*, PA*, VA*. Europe, Cyprus, Egypt, Israel, Turkey (Černý, 2018).

Type material

Lectotype: Germany. Meckelenburg (1♂, ZMHU). [Not examined]

Material examined

Canada. AB: Wabamun, caught on Solidago, 1–3.vii.1966, K.A. Spencer, CNC358537 (1♂, CNC), NB: Kouchibouguac N.P., 13.vii.1977, J.F. McAlpine, IDEMA illustration, “L. Yuzyk July 1981”, code-6042J, CNC358542–358544 (2♂,1♀, CNC), 23.v.1977, Hanley and Cooper, code-5113Q, CNC358545 (1♀, CNC), 9.vii.1977, J.F. McAlpine, code-6023Q, CNC358546 (1♂, CNC), ON: Midland, swampy woods, balsam poplar, 2.v.1959, J.G. Chillcott, CNC358551 (1♂, CNC), Ottawa, 22.vii.1954, W.R.M. Mason, CNC358550, CNC358552 (1♂ 1♀, CNC), St. Lawrence Is. Nat. Park, Thwartway Is., 15.vii.1976, A. Carter, code 4118-J, CNC358547 (1♂, CNC), 21.vii.1976, W. Reid, code 4174-N, CNC358549 (1♀, CNC), 24.vii.1976, W. Reid, code 4198-L, CNC358548 (1♀, CNC), QC: Beech Grove, 23.vi.1951, J.F. McAlpine, CNC358539, CNC358540 (2♂, CNC), Harrington Lk., Gatineau Pk., 7.vi.1954, E.E. Sterns, CNC358541 (1♂, CNC). France. [Lower Normandy]: Verson b. Caen, 4880, 9.viii.1942, Dr. H. Buhr. Nr., mine an Stachys recta CNC358538 (1♂, CNC). USA. DE: Newark, spring 1974, R.W. Rust (1♂, USNM), IN: Lafayette, 6.vii.1915, reared from catnip, issued13.vii.1915, Satterthwalt (1♂, USNM), Lafayette, swept from grass, “v-17”, J.M. Aldrich (1♂, USNM), Lafayette, “iv-6”, swept from wint. wheat (1♂, USNM), MD: Kent Is., “35-13-4”, Seek and Nixon (1♂, USNM), College Park, 28.v.1939, C.T. Greene (1♂, USM), Beltsville, B.H. Braun, i.1972 (1♂, USNM), i.1972, ex. Solidago stems (2♂, USNM), i.1972, reared ex. Eurosta solidaginis gall (1♂ 2♀, USNM), Beltsville, 12.iv.1972, em. Indoors, B.H. Braun, ex. stem Solidago canadensis (1♂ 1♀, USNM), Montgomery Co., Clarksburg, Little Bennett Reg. Park, 21.ix.1990, W.E. Steiner, M.J. and R. Molineaux (1♂, USNM), PA: Kimblesville, spring 1974, R.W. Rust (1♂, USNM), VA: Fairfax Co., Turkey Run Park, nr. mouth of Turkey Run, 38°57.9'N, 7°09.4'W, Malaise trap, 18–30.v.2007, D.R. Smith (1♂, USNM).

Comments

Ophiomyia labiatarum, a polyphagous Lamiaceae feeder also found on Solidago with a relatively northern distribution, is somewhat variable with regards to head morphology (although the facial bulb is always shiny dorsally), but the male phallus can be readily diagnosed.

Ophiomyia laticolis sp. nov.

Figs 367–371

Description

Wing length 2.2–2.3 mm (♂). Female unknown. Length of ultimate section of vein M4 divided by penultimate section: 0.7–0.8. Eye height divided by gena height: 9.0 (non type). Eye angled diagonally. Facial carina thin along length with narrow medial bulb. Gena shallowly produced, forming a 45° angle. Clypeus produced and abruptly narrowed apically with apex truncated. Buccal cavity narrowed anteriorly with anterior margin nearly straight. Distance between crossveins more than length of dm-m. Ocellar triangle and fronto-orbital plate (narrow) subshiny. Head damaged.

Chaetotaxy : Male with vibrissal fasciculus ~ 1/2 length of gena. Two ori (slightly thinner); two ors. Mid tibia with one weak posteromedial seta.

Colouration : Gena relatively high, flat and pale in holotype. Body, including halter dark brown with frons darker. Wing veins brown. Calypter margin and hairs brown.

Genitalia : (Figs 367–371) Epandrium broad, rounded, narrower in lateral view and slightly shorter to dorsum. Surstylus fused to epandrium, 1/2 length of distal region of epandrium; apex narrower, rounded; inner surface with two scattered rows of tubercle-like setae. Metepiphallus small and pale with one pair of ventromedial spines. Halves of basiphallus broadly fused at base; left sclerite short, weakly sclerotised and ill-defined, and right sclerite longer, truncated and with slight dextral curve. Distiphallus strongly spinulose on inner surface but with few other internal elaborations; slightly flattened and subquadrate in ventral view.

Host

Unknown.

Distribution

USA: MD, VA.

Etymology

The specific name compounds the Latin for wide (latus) and penis (colis).

Type material

Holotype: USA. MD: Montgomery Co., Bethseda, 28.iv.1968, G. Steyskal (1♂, USNM),.

Paratype: USA. VA: Fairfax Co., Turkey Run Park, 0.3 km W mouth Turkey Run, 38°58'N, 77°09.6'W, Malaise trap, river trap, 17–24.v.2006, D.R. Smith (1♂, USNM).

Comments

The new species Ophiomyia laticolis is highly similar to O. ambrosia Spencer from California (Spencer and Steyskal 1986b: figs 232–234), but the genitalia appear to be slightly smaller, the base of the mesophallus is nearly level with the base of the distiphallus, and the sides are severely flared, almost forming one pair of lateral points. Externally, the gena of O. ambrosia is slightly flatter and paler, but this may be an artifact of preservation; the eye is 5.6 × higher than the gena, and wing length is similar (2.2 mm).

The genitalia of this species also closely resemble those of Ophiomyia duodecima Spencer (Quebec; Spencer, 1969: figs 134–136), in that the distiphallus is subquadrate in ventral view. The distiphallus of this Canadian species, however, is 2 × as large, the mesophallus is small and gracile, the left sclerite of the basiphallus is larger and not basally fused to the right sclerite (a possible artifact of illustration) and the gena is much higher (1/4 eye height).

Ophiomyia maura (Meigen)

Figs 372–376

Agromyza maura Meigen, 1838: 7.

Agromyza curvipalpis Zetterstedt. Misidentification, in part. Frost 1924: 41.

Ophiomyia bicornis Kaltenbach, 1869: 195; Hendel 1920: 130 [as synonym of curvipalpis], 1931: 188 (as synonym maura).

Agromyza affinis Malloch, 1913: 317; Hendel 1920: 130 [as synonym of curvipalpis], 1931: 188 (as synonym of maura).

Ophiomyia maura. Hendel 1920: 130 [as synonym of curvipalpis], 1931: 188 [also includes O. pulicaria Braschnikow, A. curvipalpis and Agromyza texana Frost as synonyms]; Frick 1952: 382, 1959: 370; Sasakawa 1961: 358; Spencer 1969: 89; Papp and Černý 2015: 323; Scheffer and Lonsdale 2018: 86; Eiseman and Lonsdale 2018: 21.

Ophiomyia curvipalpis. Hendel 1920: 130.

Ophiomyia asteris Kuroda, 1954: 82. Sasakawa 1961 [synonymy].

Description

Wing length 1.8–2.0 mm (♂) 2.0–2.1 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.7–1.1. Eye height divided by gena height: 6.2–7.8. Eye slightly oblique, but still relatively round. Arista short pubescent. Ocellar triangle nearly reaching level of posterior ori. Ocellar plate narrow. Facial carina well-developed, medially with smooth, subshiny, ovate bulb; carina above bulb as wide as bulb, with shallow medial groove continuing onto lunule; lunule height subequal to width of carina dorsally. Genal process nearly as long as high, forming an approximate 60° angle. Clypeus with arms bowed inwards anteriorly, meeting at small subquadrate anteromedial extension that is shallowly concave anteriorly and with one pair of minute anterolateral points. Crossveins separated by length of dm-m or slightly less.

Chaetotaxy : Male vibrissal fasciculus thick and upcurved, ~ 2/3 length of gena. Two ori (sometimes one additional ori present on one side); two ors. Two dorsocentral setae, anterior seta ~ 4/5 length of posterior. Acrostichal setulae in six scattered rows. Mid tibia with one or no posteromedial setae.

Colouration : Setae black. Body dark brown, including halter. Calypter, including margin, light brown; hairs dark brown; ocellar triangle and ocellar plate paler brown.

Genitalia : (Figs 372–376) Metepiphallus pale and narrow with one pair of ventromedial spines. Epiphallic lobes clear with dark sclerite. Epandrium shallow, venter slightly curved anteriorly, fused to surstylus. Surstylus small, short, subtriangular, with tubercle-like setae in three rows that narrow to one anteriorly. Hypandrium subtriangular with pointed apex and bowed basal arms. Phallophorus with narrower, darker base, short dorsum, and higher, longer venter; seen ventrally, venter with one pair of dark ridges. Basiphallus with weak, short left lateral arm connected basally to longer, better-defined right lateral arm that does not reach level of mesophallus. Mesophallus narrow, cylindrical, inserted ventromedially into larger distiphallus; base of mesophallus approximately level with base of distiphallus. Base of distiphallus (overlapping mesophallus) with broad, laterally bulging anterior section that abruptly narrows to a small rounded point apically; basal section with minute divots towards apex and several small internal spinules in right anterolateral section; basal section with broad opening dorsally bearing characteristic large flared membranous fold that encompasses broad, curved, strongly produced plate; distal section of distiphallus with distinct sclerotised medial tubule that is flanked ventrally by one pair of weakly differentiated and minutely textured hemispheres. Ejaculatory apodeme with short, narrow stem emerging from wide asymmetrical base that continues onto pale, irregularly sclerotised blade that is atrophied to one side; sperm pump with dark transverse bar ventrally.

Host

AsteraceaeAster, Callistephus, Doellingeria (?), Erigeron, Eurybia, Euthamia, Oclemena, Solidago, Symphyotrichum (Benavent-Corai et al. 2005; Eiseman and Lonsdale 2018; Ellis 2021). Possibly Eupatorium (Spencer 1990).

Distribution

Canada: NB*, QC*. USA: CT, DE*, MA, MD*, ME, NY, VA*, VT; possibly also CO, MN, OH (Eiseman and Lonsdale 2018). Europe. Northeastern China. Turkey. Japan. Canary Islands. Oman. Yemen

Type material

Holotype [affinis]: USA. MD: Glen Echo, 3.vi.1898, R.P. Currie (1♀, USNM).

Holotype [asteris]: Japan. Honshu: Kantō, Yokohama, 24.v.1938, leaf miner in Aster indicus L., M. Kuroda (1♀, BLTJ).

Syntypes [bicornis]: Germany. [not given]. (Type data unknown)

Syntypes [maura]: Germany. [not given]. (Types lost).

Additional material examined

Estonia. Tallinn, 18.vii.1995, M. v.Tschirnhaus, ex Solidago (1♂, CNC). Canada. NB: Kouchibouguac National Park, 46°48'49.90"N, 64°55'40.02"W, 22.v.1977, Hanley and Cooper, Code – 5111O, CNC758931 (1♀, CNC), 23.v.1977, Code – 5113Q, CNC758930 (1♂, CNC), QC: Old Chelsea, 45°30'0.40"N, 75°48'52.80"W, 30.v.1952, J.F. McAlpine, CNC758932 (1♂, CNC). USA. CT: Litchfield Co., Canaan, 21.vii.2015, em. 29.vii.2015, C. Vispo, ex Solidago canadensis, #CSE2168, CNC564684 (1♀, CNC), DE: Newark, 1.vii.1974 (1♂, USNM), MA: Berkshire Co., Lenox, Parsons Marsh, 20.vi.2017, em. 24.vi.2017, C.S. Eiseman, ex Solidago patula, #CSE3853, CNC939713 (1♂, CNC), Franklin Co., Montague, Montague Plains Wildlife Management Area, 8.vi.2017, em. 21.vi.2017, C.S. Eiseman, ex Solidago arguta, #CSE3843, CNC939720 (1♂, CNC), Northfield, 276 Old Wendell Rd., 10.x.2016, em. 19.iv.2017, C.S. Eiseman, ex Eurybia divaricata, #CSE3524, CNC939673 (1♂, CNC), 2.vii.2017, em. 15.vii.2017, ex Euthamia graminifolia, #CSE3956, CNC939661 (1♂, CNC), Hampshire Co., Pelham, Butter Hill, 21.vi.2013, em. 24.vi.2013, C.S. Eiseman, ex Solidago caesia, #CSE591 (2♀, CNC), South Hadley, near Lithia Springs Reservoir, 11.v.2016, em. 2.vi.2016, C.S. Eiseman, ex Solidago arguta, #CSE2536 (1♀, CNC), Middlesex Co., Shirley, 42.556117, -71.613381, 3.viii.2017, em. 9.viii.2017, C.S. Eiseman, ex Solidago gigantea, #CSE4095, CNC939727, CNC9397278 (1♂ 1♀, CNC), Nantucket Co., Nantucket, Gardner Farm, 13.vi.2013, em. 24.vi.2013, C.S. Eiseman, ex Solidago latissimifolia, #CSE600 (1♀, CNC), Nantucket, Squam Swamp, 12.vi.2013, em. 26–30.vi.2013, C.S. Eiseman, ex Solidago latissimifolia, #CSE609, CNC384817, CNC384818 (1♂ 1♀, CNC), ME: Knox Co., Camden, Bald Mountain, 5.x.2013, C.S. Eiseman, ex. Oclemena acuminata em. 20.iii.2014, #CSE1011, CNC384784 (1♂, CNC), VA: Montgomery Co., Christiansburg, 2.vi.1962, J.G. Chillcott (1♂, CNC), VT: Chittenden Co., South Burlington, Winooski Gorge, 29.vi.2014, ex. Solidago flexicaulis, em. 10.vii.2014, C.S. Eiseman, #CSE1167, CNC384885 (1♂, CNC).

Comments

While relatively indistinct externally, the male genitalia of Ophiomyia maura are more diagnostic: the epiphallic lobes are clear with a dark basal sclerite; the distiphallus is relatively large and well-sclerotised with a dark anterodorsal tubule, it has a transverse dorsomedial “shelf” that is only slightly curved, and there are two minutely tuberculate distoventral hemispheres; the phallophorus has two sclerotised ridges that are obvious when viewed ventrally. It is quite similar in North America to O. carolinensis Spencer, O. parda Eiseman and Lonsdale and O. quinta Spencer, all of which now have small indicators on the distiphallus that reveal identity: O. carolinensis has a small basal protuberance on the distiphallus (Spencer and Steyskal 1986b: figs 260, 261) and a darker, broader, more distinct anterodorsal process emerging from the distiphallus, and O. parda and O. quinta have an inner-medial plate projecting dorsally that is long and more strongly curved (Eiseman and Lonsdale 2018: figs 254, 255). Also see discussion in Eiseman and Lonsdale (2018).

Ophiomyia nasuta (Melander)

Figs 64–66, 377–382

Agromyza curvipalpis Zetterstedt. Misidentification, in part. Melader 1913: 251.

Agromyza maura var. nasuta Melander, 1913: 260.

Agromyza youngi Malloch, 1914: 312. Frick 1952a: 384 [as synonym of pinguis Fallén]. Frick (1959) [as synonym of nasuta Melander].

Ophiomyia madizina Hendel, 1920: 130. Spencer 1964a [synonymy].

Tylomyza madizina. Hendel 1931: 185. Frick 1952a: 385, 1959: 372

Siridomyza madizina. Enderlein 1936a: 179.

Tylomyza nasuta. Frick, 1952a: 384 [as synonym of pinguis Fallén], 1957: 201 [lectotype designation], 1959: 372.

Ophiomyia nasuta. Spencer, 1964: 789, 1969: 91, 1976: 71, 1990: 261; Spencer and Steyskal 1986b: 42; Scheffer et al. 2007: 770; Černý 2018: 122; Eiseman and Lonsdale 2018: 23.

Description

(Figs 64–66, 377). Wing length 2.0–2.2 mm (♂), 2.1–2.5 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.8–1.0. Eye height divided by gena height: 5.4–7.4. Face short, with carina broad and diverging above large, semispherical bulb. Gena stout and produced anteriorly; highest subapically. Clypeus slightly narrowed anteriorly (more pronounced in female). Anterior margin of buccal cavity broad. Anterior angle of ocellar triangle long and tapered, nearly reaching anterior margin of frons. Frons minutely pitted with ocellar triangle and fronto-orbital plate shiny. Fronto-orbital plate and parafacial slightly projecting and continuing under eye as distinct cheek. Body shiny. Distance between crossveins approximately as long as dm-m.

Chaetotaxy : Orbital setulae proclinate and relatively long and dense. Male vibrissal fasciculus absent. Two ori; one or two ors present in female, absent in male. Three dorsocentral setae, slightly decreasing in length anteriorly. Mid tibia with one small posteromedial seta.

Colouration : Body, including halter dark brown to black. Calypter margin and hairs dark brown. Wing veins light brown to brown, sometimes whitish.

Genitalia : (Figs 378–382) Surstylus 1/2-length of epandrium at base, narrowing apically with apex broadly rounded; height approximately equal to length at base; with irregular clusters of tubercle-like setae apically and anteriorly on inner surface. Metepiphallus small with three pairs of medial and basal spines. Phallophorus slightly elongate with constricted base. Halves of basiphallus broadly fused at base with short, ill-defined left lateral sclerite and slightly longer right lateral sclerite. Distiphallus with base extending past base of mesophallus; minutely tuberculate dorsally and apically, inner surface minutely spinulose laterally near base, with small dorsomedial hook and distal section divided into two hemispheres. Ejaculatory apodeme with well-developed stalk, blade with striations and base with thin membranous tubule.

Host

AsteraceaeTaraxacum officinale.

Distribution

Across northern North America from YT to QC, south to northern CA, CO, NM and NC (Eiseman and Lonsdale 2018). Europe, Japan, Turkey, Kazakhstan (Černý 2018).

Type material

Lectotype [nasuta]: USA. WA: Kamiac Butte, 1.vi.1912 (1♂, USNM).

Paralectotypes [nasuta]: USA. ID: Troy, 14.vi.1908 (1♂, USNM), WA: Pullman (4♂, USNM). Austria.“Styria” (1♂, USNM).

Holotype [youngi]: USA. NY: Albany, 28.iv.1913, D.B. Young (1♂, NYSM). [Not examined].

Syntypes [madizina]: “Austr., Germ.” (23♂♀, NMW, USNM). [USNM ♂ and ♀ examined].

Additional material examined

Canada. BC: nr. Golden Rest Area, rt. 1, 20.vi.2001, C.R. Bartlett (1♀, UDCC), ON: Ottawa, 45°19'1.20"N, 75°43'12"W, 90 m, 7.vi.2016, J.E. O’Hara, Malaise trap, CNC629668, CNC629688 (2♀, CNC), SK: Duck Lake, 15.iv.1924, K.M. King, CNC358553 (1♀, CNC). USA. CT: Redding, 10.v.1930, A.L. Melander (2♀, USNM), DE: Pike Creek, 25.vi.1975, D. Buntin (1♀, UDCC), Bridgeville, 8.viii.195, H.E. Milliron, on peppers (1♀, UDCC), New Castle Co., Newark, Thorn Lane beside RR tracks, 18.vii.1993, field sweep, D.S. Chang (1♀, UDCC), UD Newark Farm, 39°40'14.81"N, 75°44'54.89"W, 18.ix.2007, sweep net, M. Frye (1♀, UDCC), Newark, UD Farm, 5.x.1997, W.P. Brown, sweep net (1♀, UDCC), Newark, UofD Gardens, 3.viii.2007, T. Cooper (1♀, UDCC), Sussex Co., Millville, Route 26, 5.x.1997, M.J. Harrison (1♀, UDCC), ID: Lafayette, 30.iv.1915, swept from grass, J.M. Aldrich (1♀, USNM), IL: Chicago, A.L. Melander (1♀, USNM), MA: Franklin Co., Northfield, 276 Old Wendell Rd., 5.v.2016, C.S. Eiseman, Taraxacum officinale, 20–21.v.2016 , #CSE2476, CNC654194 (1♂, CNC), MD: Temple Hills, 17.vii.1978, G.F. Hevel (1♀, USNM), Lavale, 9.v.1970, G. Steyskal (1♂, USNM), Montgomery Co., 4mi SW of Ashton, G.F. and J.F. Hevel, 18.iv.1987 (1♀, USNM), 25.iv.1987 (1♀, USNM), 15.viii.1982 (1♂ 2♀, USNM), 17.vii.1978 (1♂, USNM), 18.iv.1987 (1♂, USNM), Colesville, 31.vii.1975, W.W. Wirth (2♂, USNM), Colesville, 31.viii.1973, Malaise trap, W.W. Wirth (1♂ 2♀, USNM), P.G. Co., Temple Hills, 17.vii.1978, G.F. Hevel (2♂, USNM), Camp Springs, 8.vii.1979, G.F. Hevel (2♀, USNM), Oxon Hill, G.F. and S. Hevel, 22.vii.1978 (1♀, USNM), 24.vii.1978 (1♀, USNM), Frederick Co., 1mi N, 4.7mi W of Point of Rocks, 11.vii.1982, G.F. and J.F. Hevel (1♀, USNM), MI: Chippewa Co., 3.vi.1937, R.andK. Dreisbach (1♀, USNM), Ingham Co., E Lansing, 22.vii.1963, F.E. Giles (1♀, USNM), E. Lansing, C. Sabrosky, 3.vi.1937 (1♀, USNM), 29.v.1937 (1♀, USNM), NM: Cloudcroft, 26.v.1964, J.F. McAlpine, CNC358586 (1♂, CNC), NY: Ithaca, A.L. Melander, 28.v.1937 (1♀, USNM), 31.v.1913 (1♀, USNM), White Plains, 1.v.1909, Bueno, A.L. Melander (1♀, USNM), Geneva, 28.v.1914, A.L. Melander (1♂ 5♀, USNM), Chazy, 8.viii.1930, A.L. Melander (1♀, USNM), OH: Oxford, Mallot’s Lawn, 12.x.1978, B.A. Steinly (1♀, USNM), PA: Chester Co., Toughkenamon, 39°51'37, 75°46'58, 14.ix.2007, E. Lake Stroud Res. Ctr. (1♀, UDCC), Bucks Co., Pineville, 24.v.1969, J.W. Adams (1♀, USNM), TN: Gt. Smokies N.P., Chimneys, 20.vi.1941, A.L. Melander (1♀, USNM), Sevier Co., Rte. 441, 3mi NW NC/TN border, Great Smokey Mt. N.P., 35°38.3'N, 83°27.8'W, 4500', S.D. Gaimari, 27.v.1999 (1♀, USNM), VA: Pulaski, 7.v.1979, G. Steyskal (4♂ 1♀, USNM), Montgomery Co., Mill Cr., Rt. 785 NE Blacksburg, 30.iv.1978, C.M. and O.S. Flint (1♀, USNM), Giles Co., 10 km NW Blacksburg, Bald Knob, 18.v.1997, S.A. Marshall (1♂, DEBU), Butt Mtn., 8 km NW Blacksburg, 21.v.2005, S.A. Marshall (1♂, DEBU).

Comments

Ophiomyia nasuta is a widespread and relatively commonly encountered species that can be diagnosed quite readily using external characters: the facial carina is short with the bulb large and semispherical, the gena apically produced and truncated, there is no male fasciculus, there are three dorsocentral setae, the ocellar setulae are dense and proclinate, and there is sexual dimorphism in number of ors.

Ophiomyia sexta Spencer

Figs 383–387

Ophiomyia sexta Spencer, 1969: 98. Spencer and Steyskal 1986b: 257.

Description

Wing length 2.1–2.4 mm (♂), 2.5 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.8–0.9. Eye height divided by gena height: 2.5–4.2. Facial carina stout with sides diverging above, thin below; bulb well-developed, sometimes with shallow medial furrow. Gena shallowly produced, forming 70°–80° angle. Clypeus produced and strongly narrowed apically. Buccal cavity narrowed anteriorly with anterior margin small and straight. Distance between crossveins as long as dm-m or less. Ocellar triangle and fronto-orbital plate shiny. Excluding VA specimen, fronto-orbital plate and parafacial strongly projecting.

Chaetotaxy : Male with vibrissal fasciculus ~ 1/2 length of gena. Three ori; two ors; sometimes only one ors or with two ori on one side only. Mid tibia with one weak distomedial seta.

Colouration : Body, including halter dark brown, but sometimes portions of ocellar triangle and fronto-orbital plate paler. Wing veins brown. Calypter margin and hairs brown.

Genitalia : (Figs 383–387) Metepiphallus small with one pair of ventromedial spines and several free basal sclerites. Epiphallic lobes with pale basal pigmentation. Halves of basiphallus fused at base, with left sclerite shorter, pale, and ill-defined, and right sclerite well-developed. Distiphallus extremely wide, bilobed and ridged, resembling a brain in ventral view; base high, and distal ridged portion strongly flattened with pale dorsomedial process. Ejaculatory apodeme with stalk short, blade pale and subovate with strong medial rib.

Host

Unknown.

Distribution

Canada: AB, MB, QC, NT. USA: CO, IL*, MS*, VA*.

Type material

Holotype: Canada. AB: Cypress Hills, 25.vi.1966, K. Spencer (1♂, CNC).

Paratypes examined

Canada: MB: 2mi W Stockton, spruce-sand community, 20.v.1958, J.F. McAlpine (1♂, CNC), NT: Hay River, 15.vii.1959, P.R. Erlich (1♂, CNC), QC: Harrington Lake, Gatineau Park, 31.v.1954, E.E. Sterns (1♂, CNC), Kingsmere, 12.v.1958, J.G. Chillcott (1♀, CNC). USA. CO: Flagstaff Co., Boulder, 5800[ft], 10.vi.1961, C.H. Mann (1♂, CNC).

Additional material examined

USA. CO: Boulder, Flagstaff Cn., 1767 m, 10.vi.1961, C.H. Mann, CNC358559 (1♂, CNC), IL: nr. Forest City, forest, 21.v.1953, J.F. McAlpine, CNC358560–358562 (3♂, CNC), MS: Winston Co., Noxubee Nat. Wildlife R., Tripletts, 33°16'N, 88°51'W, pasture road, 19.v.2013, J.M. Cumming, CNC358563 (1♂, CNC), VA: Veitch, 9.vi.1912, J.R. Malloch (1♂, USNM).

Comments

Compared to other Ophiomyia with a fasciculus, O. sexta has a relatively short genal process, three ori and quite closely spaced crossveins. Internally, the incredibly large distiphallus is characteristic, being brain-shaped in ventral view and dorsoventrally compressed.

Ophiomyia simplex (Loew)

Figs 69, 70, 388–393

Agromyza simplex Loew, 1869: 46.

Melanagromyza simplex. Hendel 1920: 128; Frick 1952a: 379, 1959: 366.

Triopisopa simplex. Enderlein 1936a: 179.

Ophiomyia simplex. Spencer 1966a: 55, 1969: 98; Spencer and Steyskal 1986b: 39; Zlobin 2005b: 177.

Hexomyza simplex. von Tschirnhaus 2000: 113; Papp and Černý 2015: 180.

Description

(Figs 69, 70). Wing length 2.1–2.4 mm (♂), 2.3–2.5 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 1.0–1.2. Eye height divided by gena height: 4.5–5.4. Facial carina thin and shallow with bulb absent. Gena strongly recessed anteriorly, highest behind midpoint of eye. Fronto-orbital plate and parafacial strongly projecting and continuing under eye as distinct cheek; parafacial shiny. Ocellar triangle shiny. Clypeus narrow with sides parallel to slightly bowed. Buccal cavity broad. Distance between crossveins narrow. Costa extending just past R4+5.

Chaetotaxy : Male vibrissal fasciculus absent. Two or three ori; two ors. Mid tibia with one posteromedial seta.

Colouration : Body, including halter dark brown and shiny. Calypter margin and hairs dark brown.

Genitalia : (Figs 388–393) Epandrium with shallow ventromedial point. Surstylus small and lobate/subtriangular, folded inwards just below midpoint of anterior margin of epandrium, and with scattered, marginal tubercle-like setae. Hypandrium relatively broad with sides parallel at base and arms meeting posteriorly; with stout apical process; additional circular sclerite absent. Metepiphallus short with one pair of basal spines and one pair of basolateral arms. Halves of basiphallus separate and broadly twisted dextrally. Mesophallus broad, inserted slightly off base of distiphallus towards its venter. Distiphallus cup-shaped with ventromedial suture continuing onto mesophallus; with one pair of ventral processes with inner margin shallowly spinulose. Ejaculatory apodeme asymmetrical and relatively thin distally, with dark, well-developed stem and medial rib.

Host

AsparagaceaeAsparagus officinalis. Adults have been collected on potato and beans.

Distribution

Canada: BC*, ON, QC. USA: Widespread. Europe.

Type material

Holotype: USA. “Middle States” (1♂, destroyed).

Material examined

Germany. Berlin, from C. Schirmer (1♂ ♀, USNM), Crossen a.O., 22.v.1932, Hering, Mine an Asparagus officinalis No. 3871, CNC358570 (1♂, CNC). Canada. BC: Armstrong, collected from asparagus, 18.v.1976, 76-535, CNC358571 (1♂, CNC), ON: Ottawa, 25.v.1941, G. Matthewman, CNC358565–358569 (2♂ 3♀, CNC), Simcoe, 2.vi.1989, G.E. Shewell, CNC358564, CNC358572–358574 (2♂,2♀, CNC), Vineland, 20.vi.1937, G.E. Shewell, CNC358577 (1♀, CNC), QC: Abbotsford, 2.ix.1936, G.E. Shewell, CNC358576 (1♀, CNC), 24.vii.1936, CNC358575 (1♀, CNC). USA. CA: Lakeside, 4.iv.1944, Asparagus (1♂ 1♀, USNM), DC: “DC” (1♂, USNM), DE: Rising Sun, 12.v.1953, on asparagus, D. MacCreary (1♂, UDCC), Georgetown, 17.v.1955 (1♂ 2♀, UDCC), Stanton, 7.viii.1951 (1♂ 1♀, UDCC), GA: Tifton, 24.ix.1896, A.L. Melander (1? [head and abdomen missing], USNM), IA: Pleasant Valley, 29.vii.1931, H.M. Harris (1♀, USNM), IL: Chicago, A.L.Melander (1♂, USNM), MD: Cabin John, 10.v.1897 (2♂, USNM), Chillum, 6.iii.1914, collected on Asparagus (1♂, USNM), Colesville, 20.viii.1975, Malaise trap, W.W. Wirth (1♂, USNM), College Park, “6-29”, C.T. Greene (2♂, USNM), College Park, 10.viii.1914, collected on potato, W.H. White (1♂, USNM), College Park, “6-29”, C.T. Greene (1♀, USNM), MI: E Lansing, 1.vi.1929, on asparagus, R.W. Pettit (2♂ 1♀, USNM), Grand Rapids, 13.iv.1931, C.W. Sabrosky (1♂ 1♀, USNM), NY: Ithaca, A.L. Melander (2♂ 2♀, USNM), No. 2, Williamsville, Swp. 1, beans, 25.vii.1964 (1♂, UDCC).

Comments

Ophiomyia simplex is one of the few Ophiomyia with an anteriorly recessed gena, a strongly shiny and pronounced fronto-orbital plate and parafacial, a costa that extends only slightly past R4+5, a basiphallus composed of one pair of twisted bands and a bell-shaped distiphallus.

This species was transferred to Hexomyza by von Tschirnhaus (2000), which was rejected by Zlobin (2005), but again considered Hexomyza by Papp and Czerny (2015). The boundaries and definition of Hexomyza was considered by Lonsdale (2014), who found the genus to be polyphyletic and the type species embedded within Ophiomyia; as a result, Hexomyza was subsequently included as a junior synonym of Ophiomyia, and the new genus Euhexomyza Lonsdale was erected for the remaining related orphaned species. Ophiomyia simplex appears to share none of the defining characters of Euhexomyza, except for the high lunule, pronounced orbital plate and parafacial and smaller posteriormost intra-alar, which commonly occur throughout the genus group, including Ophiomyia. There are no convincing adult or larval characters that would ally this stem miner with the gall-forming species previously considered Hexomyza, especially among the highly dissimilar male genitalia.

Ophiomyia texana (Malloch)

Figs 394–398

Agromyza texana Malloch, 1913: 319. Hendel 1931: 194 (as synonym of proboscoidea Strobl).

Ophiomyia texana. Frick 1952a: 383; Spencer and Steyskal 1986b: 259.

Ophiomyia shiloensis Spencer, 1969: 98. Spencer and Steyskal 1986b [synonymy].

Ophiomyia arguta Spencer in Spencer and Stegmaier 1973: 169. Spencer and Steyskal 1986b [synonymy].

Ophiomyia modesta Spencer, 1981: 94. Spencer and Steyskal 1986b [synonymy].

Description

Wing length 1.8–1.9 mm (♂♀). Length of ultimate section of vein M4 divided by penultimate section: 0.8. Eye height divided by gena height: 11.0. Facial carina slender, distinct, with slender, smooth, matt facial bulb; carina not expanding immediately above bulb. Eye angled diagonally. Genal process with a broad, relatively short base approximating 60°, but apex much narrower, extending as a fine point. Clypeus strongly extended anteriorly as narrow process. Ocellar triangle with lateral margins straight to slightly concave, ending between ori and ors, but with sides produced as shallow grooves appearing sometimes to extend to lunule. Anterior margin of buccal cavity narrowed and straight. Notum shiny; bare between dorsocentrals. Crossveins separated by slightly more than length of dm-m.

Chaetotaxy : Male vibrissal fasciculus thick and upcurved, pointed, at least 2/3 length of gena. Two ori; two ors. Region between dorsocentrals smooth or with only a few anteromedial setulae. Mid tibia without posteromedial setae.

Colouration : Body, including halter dark brown. Gena and inner margin of fronto-orbital plate sometimes light brown. Wing veins dirty white to brown. Calypter margin and hairs dark brown.

Genitalia : (Figs 394–398) Surstylus narrow with few apical tubercle-like setae. Proepiphallus with lateral lobes clear with base lightly pigmented. Left sclerite of basiphallus very short, pale, and ill-defined apically, separate from right sclerite; right sclerite elongate, exceeding base of mesophallus. Base of mesophallus extending far past base of distiphallus excluding distinctive long basal tail-like process; distiphallus internally spinulose with paired ventral sclerotised ridges, subovate in ventral view and chevron-shaped in lateral view; with small dorsomedial spine. Ejaculatory apodeme with very large, pale blade past small flattened stem.

Host

BrassicaceaeRorippa, Descurainia. DE specimen collected on “spider bush”, likely as an adult.

Distribution

Canada: MB. USA: CA, DE*, TX. Bahamas. Reported by Frick (1959) as widespread from MI to WA.

Type material

Holotype [texana]: USA. TX: Brownsville, “bred from roripa”, 27.i.1909, McMillan and Marsh (1♂, USNM; type No. 15582).

Holotype [arguta]: Bahamas. Eleuthera Is.: Hatchet Bay, nr. Alicetown, 2.iv.1953, E.B. Hayden and L. Giovannoli (1♂, AMNH). [Not examined]

Holotype [modesta]: USA. CA: Ventura Co., Point Mugu State park, 2.iv.1977, K.A. Spencer (1♂, CAS). [Not examined]

Holotype [shiloensis]: Canada. MB: 5mi SW of Shilo, 22.vii.1958, J.G. Chillcott (1♂, CNC).

Paratypes [shiloensis]: Canada. MB: Churchill, 3.ix.1945, R. Richards (1♂, CNC), Treesbank, 17.viii.1958, J.G. Chillcott (1♂, CNC).

Additional material examined

USA. DE: Newark, 12.ix.1960, spider bush, D.F. Bray (1♂, USNM).

Comments

Although Ophiomyia texana can be partially recognised by the long upcurved fasciculus, a relatively small, slender, smooth, matt facial bulb and a scutum that is largely without medial setulae between the dorsocentral setae, examination of the male phallus is essential for confident identification. The distiphallus is chevron-shaped in profile, the inner surface is sparsely but evenly spinulose, and the base has a tail-like process.

The Delaware record presented here significantly expands the northeastern range of this widespread but uncommon taxon.

Ophiomyia tiliae (Couden)

Figs 399–404

Agromyza tiliae Couden, 1908: 35.

Melanagromyza tiliae. Frick 1952a: 380, 1957: 200 [lectotype designation].

Melanagromyza fastosa Spencer, 1969: 67. Syn. nov.

Hexomyza tiliae. Spencer 1973: 299.

Ophiomyia fastosa. Spencer and Steyskal 1986b: 255.

Ophiomyia tiliae. Spencer and Steyskal 1986b: 260.

Description

Wing length 1.9–2.7 mm (♂), 2.7–3.3 mm (♀). Length of ultimate section of vein M4 divided by penultimate section: 0.6–0.8. Eye height divided by gena height: 2.5–4.3. Fronto-orbital plate varying from slightly visible laterally to relatively pronounced; parafacial sometimes produced, continuing under eye as broad cheek. Gena relatively high and shallowly rounded, only slightly angled anteriorly to produce shallow ventromedial angle. Facial carina indistinct to absent, but antennal bases always slightly separated; bulb slender, not strongly pronounced, usually appearing as slight swelling below antennal bases. Clypeus stout and parallel-sided. Ocellar triangle and fronto-orbital plate shiny. Crossveins narrowly separated.

Chaetotaxy : Male vibrissal fasciculus absent. Two to three ori (orbital setula between ori variably strengthened); two ors. Mid tibia with one posteromedial seta.

Colouration : Body, including halter dark brown with gena paler. Wing veins light brown. Calypter margin and hairs brown. Abdomen sometimes with very faint, almost indistinct metallic lustre – in NY male, tint greenish with tergites 1–3 bluish.

Genitalia : (Figs 399–404) Externally as described for O. abutilivora, excluding texturing on epandrium. Phallophorus strongly compressed laterally. Basiphallus ring-shaped and with adjoining dorsobasal membrane lightly sclerotised. Distiphallus subspherical with anterior and ventral margins thick; ventrally bordered by light transverse sclerite (upcurved and pointed at sides; highly faded in tiliae holotype, likely resulting from method of preservation as other specimens reared from Tilia with this sclerite moderately to well-developed), long thin membranous flagellum (pointed ventrally) and large carinate, membranous hypophallus.

Variation : VPIC male with clypeus appearing broadly rounded (actually bulbous medially), but anterior corners still present. Cheek only evident anteriorly. Two ori, closely spaced anteriorly. Anteroventral membrane on phallus reduced, with “wings” of distiphallus barely evident.

Host

MalvaceaeTilia americana, Tilia sp.

Distribution

Canada: AB, ON*, QC. USA: CO, IL, IN, MA (Eiseman and Charney 2010), MO, NY*, PA, VA.

Type material

Lectotype [tiliae]: USA. MO: Jennings, 4.iv.1907, T.