Phylogenetic Relationships of Tachinid Flies in Subfamily Exoristinae Tachinidae: Diptera) Based on 28S Rdna and Elongation Factor-1A
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Systematic Entomology *2002) 27,409±435 Phylogenetic relationships of tachinid flies in subfamily Exoristinae Tachinidae: Diptera) based on 28S rDNA and elongation factor-1a JOHN O. STIREMAN III Department of Ecology and Evolutionary Biology,University of Arizona,Tucson,U.S.A. Abstract. The phylogenetic relationships within the largest subfamily of Tachi- nidae,Exoristinae,were explored using nucleotide sequences of two genes *EF-1 a and 28S rDNA). A total of fifty-five and forty-three taxa were represented in the analyses for each gene,respectively,representing forty-three genera. Neighbour joining,parsimony and maximum likelihood inference methods were employed to reconstruct phylogenetic relationships in separate analyses of each gene,and parsimony was used to analyse the combined dataset. Although certain taxa were highly mobile,phylogenetic reconstructions generally supported recent clas- sification schemes based on reproductive habits and genitalia. Generally,the monophyly of Tachinidae and Exoristinae was supported. Tribes Winthemiini, Exoristini and Blondeliini were repeatedly constructed as monophyletic groups, with the former two clades often occupying a basal position among Exoristinae. Goniini and Eryciini generally clustered together as a derived clade within Exoristinae; however,they were never reconstructed as two distinct clades. These results suggest that the possession of unembryonated eggs is plesiomorphic within the subfamily and that there may have been multiple transitions between micro- type and macrotype egg forms. Introduction 1987; Williams et al.,1990; Eggleton & Belshaw,1993), and the wide variety of mechanisms by which they attack Tachinidae is generally regarded as a relatively recent, them *O'Hara,1985). These oviposition strategies include actively radiating clade of parasitic flies *Crosskey,1976). planidial larvae that seek out hosts,minute eggs that are Some authorities believe it may be the most species-rich ingested by hosts and hatch in the gut,and membranous family of Diptera *Crosskey,1980),although currently it eggs that hatch shortly after deposition on a host,near a is outnumbered in described species *c. 8200 according to host or on a host's food plant. Cantrell & Crosskey,1989) by Tipulidae. Tachinids are Despite the taxonomic,morphological and ecological found on all continents and most major islands,being diversity that tachinids display,the family has been subject absent from only very high latitude polar regions and to almost no modern analyses of the relationships between some small oceanic islands *Wood,1987). All species with taxa *though see O'Hara,1989). In this paper,I present a known life histories are internal *endo-) parasitoids of other molecular-based phylogenetic reconstruction of one sub- insects and arthropods. In this regard,they often play family of Tachinidae,Exoristinae * Goniinae of Sabrosky significant roles as natural enemies of herbivorous insects & Arnaud,1965),with reference to patterns of host use and in ecological communities *Sheehan,1994; Stireman,2001). oviposition strategy. The taxonomic diversity of tachinids is matched by the immense range of hosts they attack *at least fourteen orders of arthropods,from sawflies to spiders; Ferar, Tachinidae It is quite likely that Tachinidae represent a monophyletic group based on a number of characteristics,including the Correspondence: Department of Ecology and Evolutionary presence of a swollen subscutellum,the reduced mandibles Biology,310 Dinwiddie Hall,Tulane University,New Orleans, and fusion of the labrum to the cephalopharyngeal skeleton LA 70118,U.S.A. E-mail: [email protected] in first-instar larvae,and most notably the parasitic lifestyle # 2002 The Royal Entomological Society 409 410 John O. Stireman III that all known species exhibit *Wood,1987; Pape,1992). combination of several character states,none of which is However,relationships between taxa within Tachinidae exclusive of all other Tachinidae *Crosskey,1973). Perhaps are ambiguous. Four subfamilies are generally recognized: the most consistent shared character of the group is the Phasiinae,Dexiinae,Tachininae and Exoristinae *Fig. 1). presence of a haired prosternum,but there are several Of these subfamilies,only Dexiinae can clearly be defined tachinines that exhibit this trait *e.g. Hystricia; Wood, by a synapomorphy *i.e. hinged aedeagus,Tschorsnig,1985; 1987),in some exoristine species it can be variable,e.g. Wood,1987). Each of the remaining subfamilies can only be Chetogena *personal observation),and it may be plesio- defined on the basis of a series of characters,often with morphic. Exoristinae are probably the most taxonomically many exceptions *Tschorsnig,1985; Wood,1987),and their difficult group of Tachinidae due to the great diversity of monophyly is uncertain at best. morphologically similar species. All authorities of the family appear to agree that the A provisional morphology-based phylogeny of Exoris- `taxonomy of the family is difficult and confused' *Sabrosky tinae and other tachinid subfamilies is shown in Fig. 1. & Arnaud,1965),they are `one of the most difficult families This `consensus' phylogeny is based on the classifications of Diptera in which to make practical identifications' and discussions of Herting *1984),Crosskey *1976), *Crosskey,1976) and there is `no consistent classificatory Tschorsnig *1985) and Wood *1987). Some of these more scheme within the family' *Wood,1987). The current state recent classificatory schemes *e.g. Herting,1984) place con- of tachinid systematics and taxonomy can best be summar- siderable emphasis on the oviposition strategy and genital ized with this quote from Wood's *1987) comprehensive morphology as well as on host associations,in contrast to treatment of Nearctic Tachinidae: the aforementioned reliance on the arrangement of setae. `. Descriptions of new taxa,even today,sometimes Two well defined tribes,Acemyiini *which attack Orthoptera) neglect to mention unique features,but instead,present and Siphonini,have been placed alternatively in Exoris- further combinations or recombinations of the same tinae or Tachininae by various authors *e.g. Herting, limited suites of characters. Furthermore,these char- 1960; Sabrosky & Arnaud,1965,respectively). The acter suites are usually not correlated with internal systematic position of Acemyini remains uncertain structure,life cycle,larval characters,or other charac- *Fig. 1),but most modern authors now place Siphonini ter states . Such convergence,multiplied many times in Tachininae *O'Hara,1989; Andersen,1996). The over because of the vast number of superficially similar `microtype' Exoristinae,which attack hosts by laying species,has resulted in a long history of incorrect minute eggs on the host's food plant that is subsequently identifications and misunderstanding of relationships, ingested,are presumed to form a monophyletic group a trend that still continues . even the most recent classi- based on this remarkable oviposition habit *Goniini). fications still rely mainly on arrangements of bristles These forms were previously *Townsend,1936±1941; and probably contain few monophyletic taxa.' Sabrosky & Arnaud,1965) distributed among several In the past two decades,there has been a dramatic mixed macrotype-microtype tribes due to their diverse improvement in our understanding of the composition of appearance,and their relationships are not well under- tachinid genera and the relationships of species therein *e.g. stood. The composition of and relationships between a Tschorsnig,1985; Wood,1985,1987; Barraclough,1992; large assemblage of macrotype genera,Eryciini,are not O'Hara,1994; O'Hara & Wood,1998),but there are still well agreed upon and are left largely unresolved in the few rigorous phylogenetic treatments of the deeper relation- provisional tree. Blondeliini appear to form a clade based ships between genera,tribes or subfamilies *though see on host use *a predominance of Chrysomelidae and other treatments of Siphonini in O'Hara,1989; Andersen,1996). coleopteran hosts) and a number of chaetotaxic and wing venation characteristics,including a small prealar bristle relative to the first postsutural dorsocentral bristle *probably Exoristinae sensu Herting, 1984) plesiomorphic),divergent subapical bristles,and R 45 and M ending near the wing tip,but the exact compos- The phylogenetic reconstruction performed in this ition is still rather uncertain *Wood,1985). Exoristini and study is focused on the tribal and generic relationships in Winthemiini,which tend to be rather well defined and subfamily Exoristinae *this name is considered to have relatively consistent in composition between authors,are priority over Goniinae according to Sabrosky,1999). Exor- placed basally due to their habit of laying unincubated istinae comprise about one-half of all described tachinid eggs,which is thought to be the plesiomorphic state for species and the vast majority of species that are regularly the family *Wood,1987). recorded as parasites of insect pests *Crosskey,1976,1980). In the present study,I evaluated the current understand- Members of this subfamily attack hosts belonging to at ing of relationships between tribes and genera within least eight orders of insects,including adult Coleoptera, Exoristinae based upon morphological characters *Herting, Phasmida and Blattaria *O'Hara,1985),but the majority 1984; Tschorsnig,1985; Wood,1987). By employing a of species use larval Lepidoptera as hosts. Although Exor- modern numerical phylogenetic