Systematic Entomology (2006), 31, 377–404 DOI: 10.1111/j.1365-3113.2006.00328.x A new family and genus of acalypterate flies from the Neotropical region, with a phylogenetic analysis of Carnoidea family relationships (Diptera, Schizophora) MATTHIAS BUCK Department of Environmental Biology, University of Guelph, Guelph, Ontario, Canada Abstract. The acalypterate family Inbiomyiidae fam.n. (Diptera, Carnoidea) is described for the newly discovered Neotropical genus Inbiomyia gen.n. with its type species I. mcalpineorum sp.n. from Costa Rica. The genus ranges from Guatemala south to French Guiana and Bolivia and includes a total of fourteen undescribed species, ten of which will be described formally in a separate paper. Inbiomyia is distinctive, with characteristic, extremely shortened head with nonfunc- tional ptilinum and reduced chaetotaxy, shortened first flagellomere with very elongate, dorsoapically inserted arista, proboscis with largely separate labellar lobes that point in different directions, mid tibia lacking apicoventral bristle, unusual fusion of male sternites 5–7, reduced male sternite 8, elongate surstyluslike ventral epandrial lobes, cerci absent in both sexes, extremely truncate female genitalia, and large, extremely flattened eggs. The larva of Inbiomyia and its biology are unknown. Inbiomyia occurs mostly in primary lowland rain forest and often is associated with the decaying foliage of fallen trees. Inbiomyiidae belong in the superfamily Carnoidea. The previously doubtful monophyly of the Carnoidea is accepted tenta- tively on the basis of newly established synapomorphies of the male genitalia. Family level relationships of the Carnoidea are analysed quantitatively for the first time based on a matrix of fifty-eight morphological characters. The putative sister group relationship of Inbiomyiidae to the monotypic Australasian family Australimyzidae is supported by several synapomorphies, mostly from the male and female postabdo- men. Family status for the Australimyzidae is confirmed, rejecting previous claims of a sister group relationship (or synonymy) with the Carnidae. The analysis also leads to revised hypotheses of the relationships of Cryptochetidae and Acartophthalmidae, and the paraphyly of ‘Tethinidae’ with regard to Canacidae, suspected by previous authors, is confirmed. Introduction Aulacigastridae) from the Australasian region, and Inbiomyiidae is the first family of Cyclorrhapha to be Marginidae D.K. McAlpine, 1991 (D.K. McAlpine, described from the New World based on a newly discovered 1991a) from the Afrotropical region. However, two new genus of flies. Only three Cyclorrhapha families have been families of lower Brachycera, the Neotropical Ocoidae newly discovered in the past 50 years: Xenasteiidae Hardy, (Asiloidea) and the Nearctic Oreoleptidae 1980 (Hardy, 1980) and Neminidae D.K. McAlpine, 1983 (Tabanomorpha), were described recently based on newly (D.K. McAlpine, 1983) (originally as a subfamily of discovered genera (Yeates et al., 2003; Zloty et al., 2005). The aims of the present paper are as follows: (i) to describe the genus Inbiomyia gen.n. with its type species Correspondence: Matthias Buck, Department of Environmental I. mcalpineorum sp.n.; (ii) to provide a phylogenetic analysis Biology, University of Guelph, Guelph, Ontario, Canada, N1G of its relationships with other families; and (iii) thereby to 2W1. E-mail: [email protected] justify the erection of a separate family. Formal # 2006 The Author Journal compilation # 2006 The Royal Entomological Society 377 378 M. Buck descriptions of ten additional species of Inbiomyia from group of families, and analysis required a broader character Central and South America, including a discussion of set than previously (e.g. J.F. McAlpine, 1989). Exploration of their species-level phylogenetic relationships, will be pub- new characters focused primarily on head morphology lished elsewhere (Buck, in press). Supplementary notes on the (including its appendages) and the male genitalia, yielding morphology of its sister group, the poorly known family several new promising characters of phylogenetic relevance. Australimyzidae, are provided. In the context of the phyloge- The extensive taxon base allowed an evaluation of the distri- netic analysis, the monophyly of related families is reviewed, bution and frequency of newly discovered characters in a including a new hypothesis on relationships within the cana- larger context. Exemplars from most Schizophora families cid–tethinid complex. (including almost all ‘Acalyptratae’ families) were examined (see Supplementary material). In families that are potentially closely related to Inbiomyia, exemplars from every subfamily History of discovery or even genus were examined, except those thoroughly studied by previous authors (i.e. Chloropidae, Andersson, 1977; The first specimen of Inbiomyia (a female of the type Milichiidae, Brake, 2000). species) was collected in 1980 in Monteverde, Costa Rica Although many synapomorphies support the monotypic by W.R.M. Mason of the Canadian National Collection of Australasian family Australimyzidae as the sister group of Insects (Ottawa). The specimen, examined initially by Inbiomyia, the taxa are very dissimilar and their external J.F. McAlpine, is labelled ‘Aulacigastridae n. gen. det. appearance does not hint at a close relationship. Except for J.F. McAlpine 1980’, and keys to either Aulacigastridae Griffiths (1972), who considered Australimyzidae of uncer- (e.g. J.F. McAlpine, 1981b) or does not key properly tain relationships, all previous authors placed Australimyza (e.g. Hennig, 1973). The superficial similarity to in the superfamily Carnoidea or one of its component Aulacigastridae is based on a few shared diagnostic families (Harrison, 1953, 1959; Colless & McAlpine, 1970, characters commonly used in keys (absence of ocellar bris- 1975, 1991; J.F. McAlpine, 1989; Grimaldi, 1997). The tles, presence of vibrissae, two costal breaks, developed phylogenetic analysis of Inbiomyia family relationships anal cell). The affinities of the ‘Monteverde fly’ remained therefore included the families that are assigned currently puzzling and D.K. McAlpine (Australian Museum, to the Carnoidea. Here, the Carnoidea is tentatively Sydney) first suggested that it could represent an unde- accepted as a monophyletic group based on new genitalic scribed family, but no definitive conclusion could be evidence that lends further support for the monophyly of reached in the absence of a male. Head and wing drawings this otherwise poorly defined superfamily. were prepared (K.C. Khoo, Australian Museum, Sydney, unpublished, 1981; see also Fig. 3A), but later the specimen went missing. In 1999, the author discovered, in the University of Guelph collection, four more specimens of Materials and methods different species of Inbiomyia from Peru and Guatemala. It was only in the following year when Khoo’s drawings were Morphology and terminology circulated that the Guelph specimens were recognized as belonging to the same, presumably new, family of flies. Morphological terminology follows J.F. McAlpine Inbiomyia was then rediscovered at the original locality (1981a) with few exceptions. Male and female terminalia, (Monteverde) and at a second Costa Rican locality, where head capsule, antenna, proboscis and legs (tibial organ of thirty-four specimens of both sexes were collected by the Chloropidae and Cryptochetidae) were examined on author. The later rediscovered original specimen (at the specimens cleared in hot 10% KOH and neutralized United States National Museum) is conspecific with subsequently in glacial acetic acid. All dissected parts were the new Costa Rican material. Significant new material kept in glycerine in a microvial with the specimen. A list of belonging to fourteen species from eight Central and South the taxa examined for the morphological study of the geni- American countries has been discovered since then. A talia and head structure is provided as Supplementary phylogenetic study based on fifty-eight morphological char- material. acters confirms D.K. McAlpine’s initial assumption that the taxon indeed represents an undescribed family of Schizophora. Inbiomyia is distinctive, rich in apomorphic characters and Phylogenetic analysis bears no close resemblance to any described acalypterate Schizophora. The relationships of Inbiomyia are difficult to Parsimony analysis of the matrix was performed with ascertain because the genus does not belong in any well- PAUP* 4.0b10 for Windows (Swofford, 2001) using the defined Schizophora superfamily. Its affinities lie within ‘branch and bound’ algorithm. Characters were three poorly defined superfamilies: Carnoidea, Opomyzoidea unweighted. Step matrices were defined for three charac- and Sphaeroceroidea. None of these superfamilies is satisfac- ters. Branch support for each resolved node in the strict torily characterized by apomorphic characters. The search for consensus trees was calculated with PAUP* according to the sister group of Inbiomyia therefore had to focus on a large Bremer (1994). # 2006 The Author Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 377–404 A new family and genus of acalypterate flies 379 Abbreviations of eye covered in dense ommatrichia except around the margins. Eye colour reddish brown. Frons very short, Cs, costal sector: Cs2 is measured between the apices of R1 approximately half head width, distance between front and R2þ3, Cs3 between the apices of R2þ3 and