GENUS TRICHOMYIA HALIDAY IN CURTIS, 1839

Trichomyia Haliday in Curtis, 1839: 745.

Type species: Trichomyia urbica Haliday in Curtis, 1839 (by monotypy).

Species included: Trichomyia acanthostylis Quate, 1996; Trichomyia amazonensis Araújo & Bravo, 2012; Trichomyia andina Bejarano, Pérez-Doria & Sierra, 2010; Trichomyia ancyropenis Duckhouse, 1978; Trichomyia anira Araújo & Bravo, 2016; Trichomyia annectens Duckhouse, 1978; Trichomyia annae Bravo, 2001; Trichomyia antiquaria Quate, 1961; Trichomyia aquita Araújo & Bravo, 2016; Trichomyia araguaensis Araújo & Bravo, 2016; Trichomyia armata Barretto, 1954; Trichomyia atlantica Araújo & Bravo, 2012; Trichomyia axeli Stebner & Solórzano Kraemer, 2014; Trichomyia bahiensis Araújo & Bravo, 2016; Trichomyia barbata Duckhouse, 1978; Trichomyia barrettoi Barretto, 1954; Trichomyia batu Quate, 1962; Trichomyia bifalcata Duckhouse, 1978; Trichomyia bifurcata Araújo & Bravo, 2016; Trichomyia biloba Quate, 1999; Trichomyia biuncata Duckhouse, 1978; Trichomyia botosaneanui Wagner, 1993; Trichomyia bou Bravo, 1999; Trichomyia brachypenis Duckhouse, 1978; Trichomyia braziliensis Satchell, 1956; Trichomyia brevicornis (Loew, 1850); Trichomyia brevitarsa (Rapp, 1945b); Trichomyia brochata Quate, 1957; Trichomyia buceras (Loew, 1845); Trichomyia buchholzi Wagner & Masteller, 1996; Trichomyia bulbosa Duckhouse, 1978; Trichomyia caelibata Quate, 1965; Trichomyia caipora Araújo & Bravo, 2016; Trichomyia californica Wagner, 1980; Trichomyia capixaba Araújo, dos Santos, Bravo & de Carvalho, 2017; Trichomyia carenata Araújo & Bravo, 2016; Trichomyia carlestolrai Wagner, 2001; Trichomyia cauga Bravo, 1999; Trichomyia cerdosa Araújo & Bravo, 2016; Trichomyia cetrae Araújo & Bravo, 2016; Trichomyia cinthiae Araújo & Bravo, 2016; Trichomyia cirrata Coquillett, 1902; Trichomyia clavellata Quate, 1996; Trichomyia colligata Araújo & Bravo; Trichomyia colosensis Pérez-Doria, Hernández & Bejarano, 2010; Trichomyia complexa Duckhouse, 1965; Trichomyia complicata Araújo & Bravo, 2016; Trichomyia conchulata Araújo & Bravo, 2016; Trichomyia concinna Meunier, 1905; Trichomyia confusa Araújo & Bravo, 2016; Trichomyia congoensis Satchell, 1956; Trichomyia contigua Duckhouse, 1978; Trichomyia coronula Duckhouse, 1978; Trichomyia coutinhoi (Barretto, 1954); Trichomyia crassicornis (Meunier, 1905); Trichomyia crinita Araújo & Bravo, 2016; Trichomyia crucis Duckhouse, 1978; Trichomyia danieli Bravo, 2001; Trichomyia declivivena Quate, 1963; Trichomyia decora Meunier, 1905; Trichomyia dentata Araújo & Bravo, 2016; Trichomyia discalis Quate, 1963; Trichomyia distincta Meunier, 1905; Trichomyia divaricata Duckhouse, 1978; Trichomyia divergens Duckhouse, 1978; Trichomyia dolichakis Quate, 1996; Trichomyia dolichopogon Alexander, Freitas & Quate, 2001; Trichomyia dolichostylis Duckhouse, 1978; Trichomyia dolichotrix Quate, 1999; Trichomyia duckhousei Wang, Zhang & Azar, 2011; Trichomyia eatoni Satchell, 1956; Trichomyia elongata Araújo & Bravo, 2016; Trichomyia fairchildi Vargas & Diaz Nájera, 1953; Trichomyia falcata Quate & Quate, 1967; Trichomyia fergusoni Duckhouse, 1965; Trichomyia festiva Bravo, 2001; Trichomyia flinti Wagner & Masteller, 1996; Trichomyia fluminensis Bravo, 2001; Trichomyia formosula Meunier, 1905; Trichomyia furtiva Quate & Quate, 1967; Trichomyia fusca Satchell, 1950; Trichomyia gabia Araújo, dos Santos, Bravo & de Carvalho, 2017; Trichomyia glomerosa Quate, 1963; Trichomyia grossa Araújo & Bravo, 2016; Trichomyia gustavoi Araújo, dos Santos, Bravo & de Carvalho, 2017; Trichomyia hardeggensis Omelková & JeŽek, 2012; Trichomyia hawaiiensis Quate, 1954; Trichomyia hileiana Araújo & Bravo, 2016; Trichomyia hispida Araújo & Bravo, 2016; Trichomyia horrida Araújo & Bravo, 2013; Trichomyia humerosa Duckhouse, 1978; Trichomyia iarae Bravo, 2001; Trichomyia imarui Araújo & Bravo, 2012; Trichomyia incomplexa Duckhouse, 1965; Trichomyia incrustabilis Araújo & Bravo, 2016; Trichomyia inedita Araújo & Bravo, 2016; Trichomyia inermis Barretto, 1954; Trichomyia inopis Duckhouse, 1978; Trichomyia intricata Quate, 1996; Trichomyia itabunensis Bravo, 2002; Trichomyia itocoae Tokunaga & Komyo, 1955; Trichomyia ituberensis Araújo & Bravo, 2016; Trichomyia ivani Bravo, 2001; Trichomyia jezeki Araújo & Bravo, 2012; Trichomyia jugabilis Duckhouse, 1978; Trichomyia juxta Duckhouse, 1978; Trichomyia kostovi JeŽek, 1990; Trichomyia lamasi Araújo, Bravo & de Carvalho, 2017; Trichomyia lengleti Lak, Azar, Nel, Néraudeau & Tafforeau, 2008; Trichomyia leei Duckhouse, 1965; Trichomyia lobata Araújo & Bravo, 2016; Trichomyia longa Araújo & Bravo, 2016; Trichomyia longicornis (Loew, 1850); Trichomyia longiseta Araújo & Bravo, 2016; Trichomyia lyrata Duckhouse, 1978; Trichomyia malaya Quate, 1962; Trichomyia malichyi Wagner, 1982; Trichomyia manacapurensis Araújo & Bravo, 2016; Trichomyia mariensis Araújo & Bravo, 2016; Trichomyia masneri Wagner, 1999; Trichomyia mecocerca Quate, 1963; Trichomyia mendesi Araújo & Bravo, 2016; Trichomyia menezesi Araújo, dos Santos, Bravo & de Carvalho, 2017; Trichomyia mineira Araújo & Bravo, 2016; Trichomyia minima Withers, 2004; Trichomyia mishi Bravo, 1999; Trichomyia muiraquita Araújo & Bravo, 2018; Trichomyia myrmecophila Araújo & Bravo, 2015; Trichomyia nebulicola Ibañez-Bernal, 2004; Trichomyia noctivolata Quate & Quate, 1967; Trichomyia nocturna Bravo, 2001; Trichomyia nortensis Araújo & Bravo, 2016; Trichomyia notata Araújo & Bravo, 2016; Trichomyia nova Meunier, 1905; Trichomyia nuda (Dyar, 1926); Trichomyia oahuensis Quate, 1954; Trichomyia onorei Bravo, 2002; Trichomyia paenefalcata Duckhouse, 1978; Trichomyia palauensis Quate, 1959; Trichomyia palmata Duckhouse, 1978; Trichomyia pantanensis Araújo, Bravo & de Carvalho, 2017; Trichomyia paraensis Araújo & Bravo, 2016; Trichomyia parafalcata Duckhouse, 1978; Trichomyia paranoctivolata Duckhouse, 1978; Trichomyia parvula Szabó, 1960; Trichomyia pedicilata Satchell, 1956; Trichomyia pedrabranquensis Bravo, 2001; Trichomyia pintoi Santos & Leite, 2012; Trichomyia piricornis Freeman, 1949; Trichomyia pitinguensis Araújo & Bravo, 2016; Trichomyia plumata Bravo & Araújo, 2013; Trichomyia pollex Duckhouse, 1978; Trichomyia procera Meunier, 1905; Trichomyia propinqua Duckhouse, 1978; Trichomyia pseudoannae Araújo & Bravo, 2018; Trichomyia pseudodactylis Quate, 1996; Trichomyia pseudosilvatica Araújo & Bravo, 2016; Trichomyia ptilotis Quate, 1996; Trichomyia pua Araújo & Bravo, 2016; Trichomyia pulchra Meunier, 1905; Trichomyia puntarenas Araújo & Bravo, 2016; Trichomyia quadrispinosa Duckhouse, 1978; Trichomyia quatei Bravo, 2001; Trichomyia queirozi Bravo, 2002; Trichomyia quimbaya Bejarano, Pérez-Doria & Sierra, 2009; Trichomyia ramalhoi Bravo, 2001; Trichomyia ransangi Quate, 1965a; Trichomyia rawlinsi Wagner, 1999; Trichomyia reducta Duckhouse, 1978; Trichomyia repanda Duckhouse, 1978; Trichomyia ribeiroi Araújo & Bravo, 2016; Trichomyia richardi Stebner & Solórzano Kraemer, 2014; Trichomyia riodocensis Alexander, Freitas & Quate, 2001; Trichomyia rondonensis Araújo & Bravo, 2016; Trichomyia risaraldensis Bejarano, Pérez-Doria & Sierra, 2009; Trichomyia saga Bravo, 2000; Trichomyia satterlmairi Wagner & Masteller, 1996; Trichomyia saurotis Quate, 1996; Trichomyia sequoiae Quate, 1955; Trichomyia serrajiboiensis Bravo, 2001; Trichomyia sertaneja Araújo & Bravo, 2012; Trichomyia silvatica Bravo, 2002; Trichomyia singularis Quate & Quate, 1967; Trichomyia sinuosa Araújo & Bravo, 2016; Trichomyia smithi Quate, 1963; Trichomyia spinicauda Araújo & Bravo, 2016; Trichomyia spinosa Araújo & Bravo, 2016; Trichomyia squamosa (Enderlein, 1937); Trichomyia stangae Araújo & Bravo, 2016; Trichomyia stephani Beran, Doczkal, Pfister & Wagner, 2010; Trichomyia styloryncha Curler & Moulton, 2010; Trichomyia sulbaianensis Bravo, 2002; Trichomyia tanypenis Duckhouse, 1978; Trichomyia teimosensis Bravo, 2002; Trichomyia tenera Meunier, 1905; Trichomyia triaina Duckhouse, 1978; Trichomyia triangularis Quate, 1996; Trichomyia travassosi (Barretto, 1954); Trichomyia trifida Quate, 1965a; Trichomyia trifilis Quate, 1965a; Trichomyia tritruncula Quate, 1996; Trichomyia trivialis Quate & Quate, 1967; Trichomyia trukensis Quate, 1959; Trichomyia truncata Araújo & Bravo, 2016; Trichomyia uncinata Duckhouse, 1978; Trichomyia urbica Haliday in Curtis, 1839; Trichomyia vargasi (Barretto, 1954); Trichomyia vazi (Barretto, 1954); Trichomyia wasmanni (Holmgren, 1905); Trichomyia wirthi Quate, 1955; and Trichomyia xaniostylis Quate, 1996.

Monophyly: Characters in support of the monophyly of the genus are as follows: oval shape of the dorsal border bounded by the postoccipital margin and opistosomal suture limited by the postmentum (12-2); flagellomeres located mediobasally in flagellomeres (28-0); cercus acuminated (60-1).

Diagnosis: Distance between antennal sockets short, less than one-third the width of the antennal socket. Dorsal border bounded by the postoccipital margin and oval or subtriangular opistosomal suture. Palpus with four or three segments; when four, the first and second with a small proximal area fused. Antennae with the first flagellomere approximately the same length as the second flagellomere; all flagellomeres connected asymmetrically in most species. Ascoids long, showing approximately the same length as or being longer than the flagellomere, and located near the flagellomere base. Scutum with alveoli concentrated on the margins and absent in the central region. In all species, male terminalia bearing a projection at the apex of the cercus with a more accuminate shape.

Description: Male. Head subcircular and eyes without an ocular bridge. Distance between antennal sockets short, less than one-third of the antennal socket width. Supraocular bristles in a single row. Two or three supracervical bristles, and the dorsal border bounded by the postoccipital margin and an opistosomal suture, oval or piriform. Labella globose and irregular with a swollen apex or elongated with an acuminate apex. Palpus with three or four segments; when with four, the first two segments are completely or partly fused. Scape cylindrical, approximately the same size as the spherical or cylindrical pedicel. The first flagellomere is nearly as long as the second flagellomere, with a pyriform or fusiform shape, and the insertion point between them is asymmetrical. Ascoids longer or of the same length as the flagellomere, arranged in an S or C shape and positioned mediobasally. Scutum with alveoli concentrated at the margins and absent in the central region, except in T. nuda and T. incrustabilis. Anepimeron with alveoli, except in T. nuda, T. piricornis, T. urbica and T. kostovi. Wing with the radial fork closer to the apex than the apex of CuA 2, with some variation. Alveoli in tergites II – VI randomly distributed or concentrated in the apical region. Owing to the wide variations in male genitalia at a specific level, it is not possible to identify many patterns for the genus. There is great variation in the shapes of the epandrium, cercus and gonostylus. Species with a projection at the apex of the cercus are usually acuminated. The gonostylus is little or highly sclerotized, and most species have an arm in the gonocoxite, which can vary in number, direction and shape. The hypandrium and gonocoxite are fused.

Female: The head, antenna and wing are similar to those in males. Spermathecal ducts are sclerotized and annulated. Females of most species belonging to this genus have not been described, and given their diversity and lack of studies, the characteristics of female genitalia are not listed here for the present genus.

Distribution: Worlwide distribution, with records in Australia, Austria, Belgium, Brazil, Bulgaria, Caribbean, China, Colombia, Costa Rica, Czech Republic, Denmark, Dominican Republic, England, France, Germany, Greece, Hawaii, Hungary, Indonesia, Jamaica, Japan, Madagascar, Malaya, Mexico, Micronesia, The Netherlands, Norway, New Guinea, New Zealand, Panama, Peru, Philippines, Poland, Puerto Rico, Romania, Russia, Slovakia, Spain, Suriname, Sweden, the USA and Venezuela.

Comments: T h e r e d u c t i o n o f s e g m e n t s i n the maxillary palp of Trichomyia is well illustrated in phylogeny. In Phlebotominae and Bruchomyiinae, the maxillary palp shows five segments. Reduction to four segments is observed in species of Psychodinae and Sycoracinae and in some species of Trichomyiinae, and reduction to three segments is observed only in species of Trichomyiinae (Bravo, 2000). Based on Psychodinae fossils, Azar & Nel (2003) proposed that there is a gradation between species from five to four palp segments, because they observed a weak groove in the segments of the palp, which probably resulted from the fusion of the two segments. The phylogeny of Trichomyiinae proposed here (Fig. 8) indicates sequential reduction from four to three segments in Trichomyia. First, segments one and two of the maxillary palp were partly fused, with the first segment overlapping the second one (17-1). Further reduction to three segments occurred posteriorly (16-0).