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Empidoidea Higher Classification
Higher classification and phylogeny of Dolichopodidae
Systematic position and monophyly of Dolichopodidae
The Dolichopodidae are classified in the superfamily Empidoidea, the sister group to the Cyclorrhapha (Cumming et al. 1995; Collins and Wiegmann 2002), along with the paraphyletic "Empididae." The traditional Dolichopodidae (i.e. Dolichopodidae s.str.) are part of a clade that includes the genera traditionally placed in the "empidid" subfamily Microphorinae (i.e. the microphorine and parathalassiine genera) (Cumming et al. 1995; Grimaldi and Cumming 1999; Cumming and Brooks 2002). The monophyly of this clade is supported by the orientation of the male genitalia and pregenital segments 7 and 8, which are rotated and lateroflexed to the right (Cumming et al. 1995); the possession of extensions of the subepandrial sclerite that reach beyond the base of the phallus to fuse with the hypandrium of the male genitalia (Cumming et al. 1995); and by the long, rod-shaped lower metapleural arm of the thoracic skeleton (Ulrich 1984, 1990).
The subfamily Microphorinae as defined by Chvála (1981, 1983, 1986, 1987, 1988), including the tribes Microphorini and Parathalassiini, is paraphyletic with respect to the Dolichopodidae s.str. The Dolichopodidae s.str. form a well-supported monophyletic group with the parathalassiine genera supported by the possession of an incomplete or evanescent crossvein bm-cu (Chvála 1983; Ulrich 1991; Grimaldi and Cumming 1999; Cumming and Brooks 2002), as well as several features of the mouthparts (i.e. clypeal ridge perpendicular and broad, lacinia of mouthparts absent, palpus short, six geminately sclerotized pseudotrachea) (Sinclair and Cumming 2006), and thorax (i.e. broad ventral portion of presternum, intersegmental ridge between the meso- and metapleuron forming two pockets, upper metapleural arm rod-shaped) (Ulrich 1990). Some preliminary cladistic analyses of Microphorinae + Dolichopodidae s.str. (Cumming and Brooks 2002; Shamshev and Grootaert, 2002) indicate that the parathalassiine genera may also be paraphyletic with respect to the Dolichopodidae s.str. As such, the proposals of Cumming and Sinclair (2000), Ulrich (2003, 2004, 2005) and Sinclair and Cumming (2006) to include the microphorine and parathalassiine genera within Dolichopodidae seem well justified.
Recently, Ulrich (2003, 2004, 2005) has proposed a revised ranking system for the family group names of Dolichopodidae necessitated by his inclusion of the microphorine and parathalassiine genera within the family. In Ulrich’s proposed classification, three subfamilies of Dolichopodidae are recognized: Microphorinae, Parathalassinae and Dolichopodinae, of which the latter is equivalent to the traditional Dolichopodidae (i.e. Dolichopodidae s.str.). Correspondingly, Ulrich (2003, 2004, 2005) treats the traditional subfamilies of the Dolichopodidae s.str. as tribes.
Monophyly of Dolichopodidae s.str.
Woodley (1989) listed two synapomorphies supporting the monophyly of the Dolichopodidae s.str. The first is a reduction in wing venation characterized by the first branch of the radial vein originating near the wing base just below the humeral crossvein, a feature that is also present in microphorines, and by the position of the r-m crossvein which is located near the wing base. The second synapomorphy is the presence of paired tooth-like structures of the mouthparts termed the epipharyngeal armature (Cregan 1941; McAlpine 1981; Satô 1991). These structures are unique to the dolichopodids and are presumably utilized in tearing the bodies of their prey. The mouthparts of dolichopodids are also characterized by the apomorphic possession of an enlarged epipharyngeal carina which projects vertically into the head capsule (Cregan 1941; McAlpine 1981; Satô 1991, Sinclair and Cumming (2006). The monophyly of Dolichopodidae s.str. is further supported by the possession of an hypopygial foramen which is positioned on the left lateral side of the male genital capsule (Sinclair and Cumming 2006).
Classification and phylogeny of Dolichopodidae s.str.
Fundamental works on the classification and systematics of Dolichopodidae s.str. include Loew (1864), Lundbeck (1912), Becker (1917-1918, 1922a,b, 1923), Stackelberg (1930, 1933, 1934, 1941, 1971), Parent (1938), Negrobov and Stackelberg (1971-1977), Robinson (1964, 1975), Negrobov (1977-1979), d’Assis Fonseca (1978), and Robinson and Vockeroth (1981). Although new species of dolichopodids continue to be described at relatively rapid rate, it has long been recognized that there are many problems with the higher-level classification of the family and a comprehensive review of world subfamilies and genera is badly needed (Robinson and Vockeroth 1981). Most dolichopodid subfamilies have not been studied in a phylogenetic context on a world scale and many have uncertain limits and are questionably monophyletic (e.g., Diaphorinae, Hydrophorinae, Peloropeodinae, Rhaphiinae, Sympycninae). Furthermore, virtually nothing is known about the phylogenetic relationships of the genera within these subfamilies.
Several subfamily classifications have been proposed for the dolichopodids over the past 150 years (Lioy 1863-1864; Schiner 1864; Aldrich 1905; Kertész 1909; Becker 1917-1918, 1922a,b, 1923; Robinson 1970a,b; Ulrich 1981; Negrobov 1986). The earliest classification was that of Lioy (1863-1864), whose concept of the Dolichopodidae included the Scenopinidae (Famiglia Scenopiniti), Lonchopteridae (Famiglia Lonchopteriti), Platypezidae (Famiglia Platypeziti) and Pipunculidae (Famiglia Cephalopsiti) in addition to the dolichopodids (which he divided into two groups: Famiglia Hydrophoriti and Famiglia Medeteriti). Lioy’s classification was apparently ignored by subsequent workers. Several months following Lioy (1863-1864), Schiner (1864) published his catalogue of European Diptera, in which he recognized four subfamilies of Dolichopodidae (Rhaphinae, Dolichopinae, Hydrophorinae and Diaphorinae). Aldrich (1905) was the first to treat the Nearctic fauna and recognized 12 subfamilies, including Sciapodinae (as Agonosominae), Diaphorinae, Rhaphiinae, Sympycninae, Neurigoninae, Xanthochlorinae, Thinophilinae, Medeterinae, Hydrophorinae, Plagioneurinae, Aphrosylinae and Dolichopodinae (as Dolichopinae). Kertész (1909), subsequently followed Schiner’s (1864) classification in his catalog of the Palaearctic species, but also recognized Aldrich’s New World subfamily Plagioneurinae. Lundbeck (1912) adopted the classification of Kertész (1909) in his treatment of the Danish fauna, but considered Aldrich’s (1905) system to be a more natural arrangement of the genera.
Becker’s series of monographs of the of the world fauna of Dolichopodidae (Becker 1917-1918, 1922a,b, 1923) provided the most comprehensive treatment of the family available at the time and remains the foundation of modern dolichopodid systematics. Becker recognized 11 subfamilies including: Dolichopodinae, Plagioneurinae, Hydrophorinae, Aphrosylinae, Medeterinae, Rhaphiinae, Neurigoninae, Diaphorinae, Stolidosomatinae, Sympycninae (as Campsicneminae) and Sciapodinae (as Chrysosomatinae). Becker’s system was similar to that of Aldrich (1905); however, he did not recognize Xanthochlorinae, and divided the genera assigned to that group among Rhaphiinae (Achalcus Loew), Sympycninae (Chrysotimus Loew and Xanthochlorus Loew) and Diaphorinae (Xanthina Aldrich). Becker also synonymized the Thinophilinae within the Hydrophorinae, and placed Teuchophorus Loew and Campsicnemus Haliday in the Sympycninae, Eutarsus Loew and Peloropeodes Wheeler in the Rhaphiinae, and Argyra Macquart, Nematoproctus Loew and Coeloglutus Aldrich in the Diaphorinae. Becker’s subfamily classification was widely adopted by most subsequent workers (Stackelberg 1930, 1933, 1934, 1941, 1971; Parent 1938; Foote et al. 1965; Dyte 1975; Dyte and Smith 1980).
Robinson (1970a,b) proposed a revised subfamily classification based on the Nearctic and Neotropical fauna arguing that Becker’s (1917-1918, 1922a,b, 1923) treatment put too much emphasis on extreme forms and that not enough attention was given to intermediate members of the family. Robinson (1970a) also criticized Becker and followers for lumping most of the smaller dolichopodids into ill-defined subfamilies such as the Sympycninae. Robinson’s classification recognized 14 subfamilies and incorporated characters of the mouthparts (Cregan 1941), genitalia (Buchmann 1961) and immature stages (Dyte 1967). Robinson (1970a) dismantled Becker’s concept of the Rhaphiinae and split it into three subfamilies, namely Peloropeodinae, Systeninae (containing the single genus Systenus Loew) and a more restricted Rhaphiinae (composed solely of the nominal genus Rhaphium Meigen). Robinson (1970a,b) also recognized the Xanthochlorinae of Aldrich (1905) (exclusive of Chrysotimus which he placed in the subfamily Peloropeodinae), erected Enliniinae for the genera Enlinia Aldrich and Harmstonia Robinson, and synonymized Aphrosylinae within the Hydrophorinae. Although no phylogenetic hypothesis was provided, Robinson (1970a) suggested close relationships between Rhaphiinae and Diaphorinae, Systeninae and Medeterinae, Medeterinae and Neurigoninae, and Stolidosomatinae and Sympycninae.
Ulrich (1981) modified Robinson’s (1970a,b) classification to encompass the world fauna and recognized 10 subfamilies. Ulrich expanded Systeninae to include Achalcus and Xanthina from Robinson’s Xanthochlorinae, as well as Epithalassius Mik and Euxiphocerus Parent. Ulrich also added Stolidosomatinae, Peloropeodinae and Xanthochlorus to Sympycninae, and expanded Rhaphiinae to include Plagioneurus Loew (i.e. Plagioneurinae), Argyra, Keirosoma Van Duzee, Nematoproctus, Pseudargyra Van Duzee and Somillus Brèthes from Robinson’s Diaphorinae, as well as Pinacocerus Van Duzee from Robinson’s Sympycninae. In addition, Ulrich transferred several Old World genera into Sympycninae (i.e. Bathycranium Strobl, Coracocephalus Mik, Eucoryphus Mik, Mastigomyia Becker, Nurteria Dyte and Smith, Rhynchoschizus Dyte) and into Dolichopodinae (i.e. Anasyntormon Parent, Colobocerus Parent, Halaiba Parent, Katangaia Parent, Phalacrosoma Becker, Syntomoneurum Becker, Vetimicrotes Dyte). Ulrich’s (1981) classification was subsequently adopted (with some modifications) by Bickel and Dyte (1989). Recently, Brooks (2005) redefined the limits of the Dolichopodinae and excluded Colobocerus, Katangaia, Pseudohercostomus Stackelberg and Vetimicrotes.
Negrobov (1986) rejected Ulrich’s (1981) system and proposed a classification similar to that of Robinson (1970a,b) which recognized 14 subfamilies and included tribal classifications for Diaphorinae, Dolichopodinae, Hydrophorinae, Medeterinae and Sympycninae. Negrobov relegated Robinson’s Stolidosomatinae to tribal status within the Sympycninae and erected the subfamily Coeloglutinae for the Neotropical genera Coeloglutus and Neotonnoiria Robinson, which Robinson placed in the Neurigoninae. Bickel (1998) questioned the validity of the Coeloglutinae and considered it a synonym of Neurigoninae. More recently, Naglis (2001) assigned this group tribal status (Coeloglutini) within the Neurigoninae and recognized two additional tribes, i.e. Dactylomyiini and Neurigonini (Naglis 2002a,b, 2003a,b). Like Robinson (1970a,b), Negrobov (1986) recognized a monogeneric subfamily concept of Systeninae; however, Bickel (1986) included Systenus within Medeterinae. Negrobov’s (1991) Palaearctic catalogue followed his 1986 classification.
Since Negrobov’s (1986) classification, two additional subfamilies have been erected, i.e. Babindellinae (Bickel 1987) and Achalcinae (Grootaert and Meuffels 1997). Bickel (1987) erected Babindellinae for the Australian genus Babindella Bickel, which is characterized by the possession of a symmetrical male postabdomen. The Achalcinae was erected following the discovery of the genus Scepastopyga Grootaert and Meuffels from Papua New Guinea. Grootaert and Meuffels (1997) also included Achalcus and Xanthina (from Robinson’s Xanthochlorinae and Ulrich’s Systeninae) within the Achalcinae and suggested a close relationship to the Sciapodinae and Medeterinae. Pollet (2005) described a fourth genus of Achalcinae, Australachalcus Pollet. The recently published catalog and checklist of the Dolichopodidae of America north of Mexico (Pollet et al. 2004; Brooks et al. 2005) followed a classification similar to Robinson (1970a) and Negrobov (1987, 1991). However, Achalcus was placed in the Achalcinae (sensu Grootaert and Meuffels 1997) instead of in Xanthochlorinae (Robinson 1970a) or Enliniinae (Negrobov 1987, 1991), and Systenus was placed in the Medeterinae instead of Systeninae (sensu Bickel 1986). Nematoproctus was included in the Rhaphiinae sensu Negrobov (1987, 1991), rather than in the Diaphorinae.
Despite the various classifications proposed for the Dolichopodidae, most of the differences of opinion have focused on certain subfamilies, (e.g., Sympycninae, Rhaphiinae) which have been repeatedly redefined. In contrast, the classification and limits of other subfamilies (e.g., Sciapodinae, Dolichopodinae) have remained relatively stable.
Phylogenetic relationships and possible origins of Dolichopodidae s.str.
Besides passing comments on the affinities of certain dolichopodid subfamilies (Robinson, 1970a; Grootaert and Meuffels 1997), very little has been written about their phylogenetic relationships. Most published discussions regarding subfamily relationships are based primarily on the morphology of the male hypopygium and the pregenitalic segments (Negrobov 1986; Bickel 1994). There are two main morphological forms of the dolichopodid hypopygium, the “encapsulated” condition and the “pedunculate” condition. In the encapsulated condition (e.g., Diaphorinae, Sympycninae, Plagioneurinae) the hypopygium (when at rest) is partially covered by tergite 6 and segment 7 acts as a short lever to roll the genital capsule ventrally and anteriorly (Negrobov 1986; Bickel 1987). In the pedunculate condition (e.g., Sciapodinae, Medeterinae, Neurigoninae, Babindellinae, Dolichopodinae) the hypopygium is permanently exserted (although it may rest partially enfolded by the venter of the preabdomen) and segment 7 is modified into a peduncle or external arm on which the genital capsule is supported (Negrobov, 1986; Bickel, 1987). The pedunculate condition is also present in the microphorine and parathalassiine genera (Chvála 1986, 1987, 1988; Ulrich, 1988), and is part of the groundplan of the Dolichopodidae s.l., whereas the encapsulate hypopygium has been derived within the Dolichopodidae s.str., perhaps several times.
Recently, Ulrich (2004) hypothesized that the Dolichopodidae s.str. descended from marine costal inhabitants. Ulrich argued that the common ancestor of the parathalassiines (which are almost exclusively marine littoral), and the Dolichopodidae s.str., invaded the sea coast from the ancestral terrestrial habitat of the microphorines. Ulrich further suggested that marine littoral habitats are still occupied by a basal grade of Dolichopodidae s.str., which have traditionally been placed in the subfamily Hydrophorinae. The remaining Dolichopodidae s.str. abandoned the marine habitat in favor of a return to terrestrial and freshwater habitats, assuming independent reinvasions of the sea coast by several species in various subgroups. To date, Ulrich’s (2004) hypothesis is yet to be tested by a rigorous phylogenetic analysis.
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First published on the Internet on 18 November 2005