Micropezidae Loew 1862

• Main Author: Lonsdale, Owen
• Format: Text
• Language: unknown
• Published: Zenodo 2020
• Subjects: Biodiversity; Taxonomy; Animalia; Arthropoda; Insecta; Diptera; Micropezidae; Antarctic; Argentina; Argentine; Burrows; Canada; Chandler; Ferro; Foote; Indian; Merz; Meunier; New Zealand; Norway; Ortiz; Pacific; Rotten; Antarc*; Antarctica; Elephant Seal; Southern Elephant Seal
• Online Access: https://dx.doi.org/10.5281/zenodo.3679563; https://zenodo.org/record/3679563

Micropezidae Loew, 1862 (Figs 312–394, 421–422) Type genus: Micropeza Meigen 1803: 276, in Loew (1862: 38) [see Sabrosky (1999) for discussion]. Type species of genus: Musca corrigiolata Linnaeus, 1767: 995, by monotypy. Micropezidae is the most diverse family of Nerioidea with about 700 described species (Marshall, 2012) and it is certain that many more await discovered based on the productivity of recent revisionary work. Species occur globally except for Antarctica and New Zealand, with most diversity to be found in tropical regions. Five subfamilies are presently accepted within a monophyletic Micropezidae—Calycopteryginae, Calobatinae, Eurybatinae, Micropezinae and Taeniapterinae—reflecting the most recent classification of D.K. McAlpine (1975, 1998). Micropezidae are relatively elongate and gracile, with long, narrow legs, and sometimes colourful patterning. While these aspects are modestly developed in Calobatinae, which is more typically acalyptrate in appearance, they can be exceptionally exaggerated in the other subfamilies, especially in the length of the mid and hind legs, which may far exceed the length of the body (see Marshall (2016: fig. 43, 2017: fig. 97)). A number of species mimic Hymenoptera, especially Ichneumonidae, and lineages in several subfamilies have independently converged on ant-mimicry (see Marshall (2016: figs 40, 41), sometimes including the development of a petiolate and sometimes nodular abdomen. The latter is well-illustrated in the apterous Australian Badisis ambulans McAlpine (Eurybatinae). Wing loss is also seen in Calycopteryx moseleyi Eaton (Figs 328–333), the sole member of the subfamily Calycopteryginae , where vestiges of the wing and halter are still evident. This species, restricted to the Kergulen and Heard Islands in the south Indian Ocean, is adapted to life on windswept islands, and is atypically drab, robust and stout. The more “modestly” proportioned subfamily Calobatinae (Figs 312–316) differs from other Micropezidae in lacking strong dorsal setae on the mid and hind tibiae, in having stronger apical tibial setae, more evenly distributed setae on the katepisternum, and wider abdominal sternites. The subfamily is Holarctic in distribution. Members of the distinct subfamily Micropezinae (Figs 317–321) are exceptionally narrow, gracile and dark in appearance. The head is also elongate, cross-vein bm-m is absent, and there are no fronto-orbital setae. Most species occur in the diverse Micropeza , which is mostly Neotropical but also Holarctic in distribution, but three species are also known from Cryogonus Cresson, restricted to Chile and Argentina. The west Palaearctic Micropeza corrigiolata has been discovered in Canada (Hoebecke & Wheeler, 1994) and South Africa (Barraclough, 1996), where it was likely introduced via agricultural trade since the larvae are known to occur in the root nodules of commercially important legumes. The subfamily Eurybatinae is almost entirely Oriental and Australian in distribution, but one monotypic genus is known from Costa Rica (Marshall, 2002), and one is known from Mauritius and Réunion (Barraclough, 1992a). One genus of Eurybatinae is so far known only from males, Anaeropsis Bigot; it is the only stalk-eyed nerioid and the only non-micropezine without fronto-orbitals (see D.K. McAlpine (1975)). The monophyly of the subfamily is “rather weakly supported” (D.K. McAlpine, 1998), and ongoing research suggests that its tribes Metopochetini (Fig. 327) and Eurybatini should be treated as separate subfamilies (Jackson et al ., in manuscript; Yusof & Marshall, in manuscript). Taeniapterinae (Figs 322–326) is the most diverse subfamily of Micropezidae in terms of both genera and species. It is found in all biogeographic regions with highest diversity in the Neotropics. The subfamily is characterized by a fan-like clustering of setae posteriorly on the katepisternum, sometimes anteriorly shifted ocelli, sometimes a laterally compressed epandrium, a lack of surstyli, elongate fore coxae (also Eurybatinae) and sometimes a pointed anal cell (also Eurybatinae) that can be exceptionaly long in some species (D.K. McAlpine, 1998). Regional catalogues of Micropezidae are available in Steyskal (1965b) [Americas north of Mexico]; Aczél (1949c) [Neotropical Region]; Steyskal (1968a) [Americas south of USA]; Marshall et al. (2016) [Colombia]; Soós (1984a) [Palaearctic Region]; Steyskal (1980b) [Afrotropical Region]; Steyskal (1977c) [Oriental Region]; Evenhuis (1989a) [Australasia/Oceaniania]. The North American fauna has been treated or summarized by Cresson (1938), Merritt (1971, 1972), Merritt & James (1973), Merritt & Peterson (1976) and Steyskal (1987a). Treatments of the Palaearctic fauna have been provided by Czerny (1930a), Greve & Nielsen (1991), Roháček & Barták (1990) and Ozerov (1987). In the last few decades, work on the Afrotropical fauna include genus and regional revisions by Barraclough (1992b, 1993c, 1996) and Marshall (2014, 2017, 2019). Treatments of the Australasian fauna were provided by Steyskal (1947, 1952), who reviewed some species from the Solomon Islands and described Australasian specimens deposited in the USNM. Aczél (1959) treated the Micropezidae of Micronesia. D.K. McAlpine reclassified the family-level groups of Micropezidae and reviewed the Australasian Eurybatinae (D.K. McAlpine, 1975), and later revised the fauna of Australia (D.K. McAlpine, 1998). Li et al. (2015) revised the Oriental Cothornobata . Early works on the Neotropical fauna include Cresson (1930), Hennig (1934b, 1935a, b) and Aczél (1949b, 1951). Contemporary revisionary work of Neotropical taxa is being primarily developed by S.A. Marshall and colleagues: Marshall (2002, 2004 a, b, 2010, 2011, 2013, 2014, 2015, 2016), Marshall & Jackson (2014), Ferro & Marshall (2018). Fossils of five species in four genera of Micropezidae are listed in Evenhuis (1997): Calobata rottensis (Statz) [compression fossil, Oligocene, Germany]; Micropeza prompta Meunier [Copal fossil, Tanzania, Pleistocene / Ho- locene]; Rainieria sp. [amber, Dominican Republic, Oligocene/Miocene]; Electrobata myrmecia Hennig [amber, Baltic Region, Eocene / Oligocene]; E. tertiaria (Meunier) [amber, Baltic Region, Eocene / Oligocene]. D.K. McAl- pine (1998) suggested that both named Electrobata species might be best treated within different genera, which would certainly seem warranted based on their divergence in chaetotaxy, venation and sclerite shape. Evenhuis (1997) further notes that other, possibly undescribed species are mentioned in the literature from Baltic amber, Chiapas amber and Sicilian amber, and a specimen of Micropezidae in Dominican amber is figured in Grimaldi & Engel (2005). An additional “ Electrobata spec.” from Baltic amber was examined in Hennig (1967). Tschirnhaus & Hoffeins (2009) also list additional taxa from Baltic amber that they did not formally describe, but included in a key to Baltic Amber Acalyptratae. One fossil genus and species was previously recognized for Cypselosomatidae— Cypselosomatites succini Hennig. The species was described from Baltic amber by Hennig (1965), who considered it basal to the remainder of the family. D.K. McAlpine (1966) noted that the species more closely approached Micropezidae in morphology, but no reclassification was proposed until a later date when he formally treated it as Micropezidae (D.K. McAlpine, 1998). Biology . Adults of Micropezidae are often encountered on horizontal surfaces such as low foliage and logs, usually in sunspots, and sometimes near moving or standing water, including temporary pools (Merritt & James, 1973; Roháček & Barták, 1990; Marshall, 2010, 2012). Many species occur at a variety of elevations, mostly lowlands, but some species and genera are found only at high-altitudes, including Neotropical Mesoconius (see Marshall (2015)). North temperate species mostly occur in moist wooded areas, but also meadows, marshes and grasses along water (Merritt & James, 1973; Roháček & Barták, 1990; Roháček, 2012b); Micropeza corrigiolata has been collected off of grass and vegetation in moist, shaded areas, and in meadows and fields containing legumes (Barraclough, 1996; Hoebeke & Wheeler, 1994). In Australia, species of Metopochetus were mostly collected in rainforests, similar to Australian Cothornobata and Crepidochetus , as well as on standing Eucalyptus trees and saplings (D.K. McAlpine, 1998). Mimegralla australica Hennig appears to prefer disturbed habitats (D.K. McAlpine, 1998). Neotropical species are known from numerous habitats, including rainforests, edges and disturbed habitats such as cacao plan- tations, but also native forests (Marshall, 2010, 2013). Some taxa appear to require pristine habitats that have had little to no disturbance, such as Mesoconius , which is particularly restricted in its habitat requirements (Marshall, 2010). South African taeniapterines occur in sunlit areas, open woodlands, disturbed areas including suburban areas or cultivated gardens, and cool, shaded, humid forests, sometimes at edges (Barraclough, 1996). Adults can be locally abundant on carrion, artificial honeydew and decaying fruit (Barraclough, 1996; Marshall, 2010, 2012, 2013), but also on ripe or damaged fruit (Oosetrbrook, 1998). Mimegralla albimana Doleschall was considered an opportunistic feeder on a decomposing pig carcass (Chin et al ., 2011). Many taxa are also very frequently found at mammal or bird dung, sometimes in large numbers (Merritt & James, 1973; Marshall, 2010, 2012), and small dung baits of are a reliable means of collection, although some taxa do not appear to be attracted to this substrate, including Tenthes Cresson, Metasphen Frey and some Grallipeza (Taeniapterinae) (Marshall, 2010, 2013). Feeding on insects is known in some adult Micropezidae, especially Calobatinae, which may be at least facultatively predaceous (Marshall, 2012), with observations of specimens feeding on aphids and small, mostly nematocerous Diptera (Colyer & Hammond, 1968). Taeniaptera lasciva (Fab.) is noted as being a predator on adult Diatraea saccharalis (Fab.), the sugarcane moth borer (Bennett & Alam, 1985). Mimicry of Hymenoptera appears to be common among Micropezidae, modeling species of Formicidae, Ichneumonidae and sometimes Pompilidae (Marshall, 2010). While the benefits of ant-mimicry are well documented and known to have evolved independently at least 70 times (Mclver & Stonedahl, 1993), the relative benefits of parasitoid mimicry are yet to be determined, although it would appear to be sufficient given the number of presumed mimics, especially within Taeniapterinae. Many Ichneumonidae, including Ophioninae and Tryphoninae, are capable of stinging, with some being quite painful, and species of several subfamilies are known to release a pungent, possibly protective odour when disturbed (Quicke, 2013). These and other Ichneumonidae are also capable of using their ovipositor to keep distance from an enemy and may engage in aggressive behaviour such as biting. Mimics of Ichneumonidae have entirely to partially bright white fore tarsi and are typically found on foliage or usually horiozontal bark (Berg, 1947), waving one or both extended fore legs in front of them, with these legs having the movement and appearance of the model’s antennae (Hennig, 1935b; Barraclough, 1996; D.K. McAlpine, 1998; Marshall, 2010). Individuals of one of these mimics, Ptilosphen viriolatus Enderlein, were observed in “sleeping aggregations”, wherein sleeping individuals faced the petiole, and slowly and continuously waved their fore legs in a manner similar to that seen while awake (Ortiz, 2001). Barraclough (1996) noted this leg-waiving was sometimes seen between individuals of the same species, although the role of this behaviour and their sex was not determined. Courtship behaviour appears to have an important role in reproduction. Behaviour of the ant mimic Cardiacephala arthriticus (Wiedemann) was recorded by Wheeler (1924) in detail, wherein a male on the top of a large leaf fended off rival males and engaged females to convince them to mate. Females appeared to chase away males until the males convinced them to mate by presenting regurgitated fluid as a gift while “dancing”. The provision of fluid gifts and other courtship activities continued until mating was completed. Marshall (2012) noted that the oral exchange of fluids is also common in Taeniaptera and photographed such a transfer between a pair of T. trivittata Macquart. Barraclough (1996) theorized that similar exchanges of nuptial gifts would be present in those Taeniapterinae with lateral swellings anteriorly in the male pleural membrane such as Mimegralla . He suggested that the swellings had a glandular function similar to that seen in Tephritidae, where the swollen structures were associated with trophallaxis during courtship. Conversely, the author noted an alternative interpretation by D.K. McAlpine, who thought that the swellings were instead “evaporative areas associated with the release of a pheromone”. Other components of taeniapterine copulatory behaviour may involve “kissing” behaviour, not necessarily associated with the exchange of nuptial gifts, along with “stilting and stroking” actions (Marshall, 2012). Female display to attract a mate was observed in the taeniapterine Ptilosphen tetrastigma (Schiner), where a white banded abdomen and positioning of banded white forelegs were used (Marshall, 2012). Oviposition in wood has been recorded for a number of Taeniapterinae and Eurybatinae, where eggs may be laid in cracks or irregularities in the wood or bark surface, sometimes including the openings of beetle burrows (Marshall, 2012). A male Grammicomyia Bigot was observed on a patch of fallen wood that was suitable for oviposition, where he waited for a female to arrive while defending the patch from other males (Marshall, 2012). Much remains to be discovered of micropezid larval life history, but individuals appear to be generalist saprophages in a variety of habitats, with a preference for moist, rotting plant matter, most often including wood (with a minority in roots), grass and fruit, but also dung. Calobatinae have been reared from heaps of decaying vegetation, including grass (Teskey, 1972; Ferrar, 1987), and Russian Calobatella petonella (L.), which is known to overwinter in the soil, was reared from sewage tanks and pig dung (Lobanov, 1960). In Eurybatinae, Li et al . (2015) suggested that larval development in rotting wood was likely widespread, noting that females from five genera were observed ovipositing on that substrate, including taxa photographed by Marshall (2012). Larvae of the unusual Badisis ambulans develop at the bottom of the cup leaves of Cephalotus follicularis (the Albany pitcher plant) (Yeates, 1992; Marshall, 2012), but this host specialization appears to be atypical for the family (D.K. McAlpine 1998). The likely ant model of B. ambulans has been observed on the same host plant (D.K. McAlpine, 1998). In Taeniapterinae, larvae are mostly known from varied rotting media, especially plants: decaying vegetative material for Mimegralla coeruleifrons (Hennig, 1936c); decaying sugar cane cuttings for Taeniaptera lasciva (Cresson, 1938); the surrounding pulp of Metroxylon sagu (true Sago palm) seeds for M. albimana striatosafciata (Berg, 1947); decaying fruit of Myrianthus arborea for M . gowdeyi (Frey) and the palm tree Borassus flabellifera for M. respondens (Walker) (Verbeke, 1951). Albuquerque (1972) was able to rear Scipopus belzebul (Schiner) under lab conditions from rotting banana and human feces. Rotten wood in trees, sometimes noticeably attacked by fungus, has served as substrate for numerous species: rotting Liriodendron tulipfera for Calobatina geometroides (Cresson) (Wallace, 1969); rotting wood of Erythrina caffra for Cephalosphen conifrons (Bigot) (Hennig, 1936c); standing, rotting Ulmus americana suffering fungal attack for Rainieria brunneipes (Cresson) (Steyskal, 1942); beneath the bark of dead trees for M. a. albimana (Bohart & Gressitt, 1951) and M. albimana striatosafciata (Enderlein) (Berg, 1947); dead banana wood for M. albimana galbula (Bohart & Gressitt, 1951), T. lasciva (Fab.), T. annulata (Fab.) and Plocoscelus conifer (Hendel) (Fischer, 1932). Adult occurrence of Rainieria calceata (Fallén) on fallen Fagus sp. (beech) was tentatively taken as an indicator of larval habitat by Chandler (1975a). Numerous species in other genera have also been observed ovipositing in a variety of decaying plant material, including wood, rotting stems and Typha stems (see Marshall (2010, 2012)). Species of Mimegralla (Taeniapterinae) are also known to be primary invaders of live plants. Mimegralla coeruleifrons (Macquart) —otherwise known as the “rhizome fly”—attacks ginger ( Zingiber officinale ) (Steyskal, 1964) and turmeric ( Curcuma longa ), tunneling in the outer and inner regions of the rhizome (Ghorpade et al. , 1988). The host plant experiences yellowing and drying of the leaves and the main shoot, and is opened to invasion by a number of disease-causing fungi and nematodes (Ghorpade et al. , 1988). Steyskal (1964) noted that records cited by Hennig (1952a, b) on Curcuma sp. (wild arrowroot) may also be M. coeruleifrons , and that other larvae found in North America on intercepted ginger from China were possibly Mimegralla , perhaps M. albimana galbula (Osten Sacken). In Micropezinae, the sometimes pestiferous Micropeza corrigiolata might breed in compost heaps (Chinery, 1986), but larvae are known to occur on live legumes, specifically, in the fresh root nodules of Pisum arvense L., Medicago sativa L. and Trifolium pratense L.; the nodule is hollowed out from the inside leaving an empty shell, after which the larva burrows 30 cm into the soil to overwinter (Müller, 1957). Calycopteryx mosleyi (Calycopteryginae) eggs were found on and under leaves on Pringlea antiscorbutica R. Br. (“Kergulen cabbage”). Larvae were found mining shallow pits in the roots of the plant, and were recovered among numerous other substrates, including under stones, other vegetation including Azorella sp., and the soil un- der decomposing animal matter, including the carcass of a “sea elephant” (presumably the Southern elephant seal, Mirounga leonine (Linnaeus)); puparia were recovered from moss and rotting seaweed (Womersley, 1937). Ferrar (1987) suggested that this relatively broad diet may not necessarily include P. antiscorbutica , which is absent from one island upon which the fly occurs. Immature stages. Most descriptions of immature stages of Micropezida : Published as part of Lonsdale, Owen, 2020, Family groups of Diopsoidea and Nerioidea (Diptera: Schizophora) - Definition, history and relationships, pp. 1-177 in Zootaxa 4735 (1) on pages 39-45, DOI: 10.11646/zootaxa.4735.1.1, http://zenodo.org/record/3670762 : {"references": ["Loew, H. (1862) Monographs of the Diptera of North America. Part I. Smithsonian Miscellaneous Collections, 6 (1), i-xxiv + 1 - 221. https: // doi. org / 10.5962 / bhl. title. 1718", "Meigen, J. W. (1803) Versuch einer neuen Gattungs-Eintheilung der europaischen zweiflugligen Insekten. Magazin fur insektenkunde, 2, 259 - 281.", "Sabrosky, C. W. (1999) Family-Group Names in Diptera: An annotated catalog. MYIA, 10, 1 - 576.", "Linnaeus, C. (1767) Systema naturae. 12 th Edition. 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