Jeffrey M. Cumming

Homology and phylogenetic implications of male genitalia in Diptera - Eremoneura

Homologies of male genitalic structures in the Eremoneura (Empidoidea + Cyclorrhapha) are examined and implications for the phylogenetic relationships of the included families are discussed in light of other characters. A revised epandrial hypothesis for the evolution of male genitalia within the Eremoneura is presented, based on comparison of male genitalic features throughout the Brachycera, and a periandrial hypothesis is rejected. Ground plan modifications of the genitalia of Eremoneura include complete sclerotization of the subepandrial membrane along its length to form a subepandrial sclerite, formation of bacilliform sclerites, loss of the lateral ejaculatory processes of the sperm pump, development of a deeply emarginate epandrium, and fusion of the hypandrium with the gonocoxites. Ground plan apomorphies of the Cyclorrhapha, including Opetiidae, consist of a sperm pump separated from the base of the phallus, circumversion of the genitalia, loss of the gonocoxal apodemes, and development of surstyli. The ground plan of the Empidoidea is characterized by the apomorphic loss of gonostyli, and the development of a sperm pump with a lever-like ejaculatory apodeme. Surstyli have developed independently in several lineages of Empidoidea, parallel to their development in Cyclorrhapha, as have losses of the gonocoxal apodemes. The above characters support the monophyly of the Eremoneura and also indicate that the Empidoidea and Cyclorrhapha are sister groups, as opposed to alternative hypotheses that suggest the Empidoidea is paraphyletic with respect to the Cyclorrhapha. The monophyly of Cyclorrhapha, exclusive of Opetiidae, is supported by the presence of a phallapodeme. The Lonchopteridae and Phoroidea are united partially on the basis of a similar reduction of pregenital sclerites in the male. The Phoroidea is characterized by the apomorphic loss of gonostyli and abdominal spiracle 7 in the male. The Syrphoidea and Schizophora are hypothesized to share a synapomorphic lever-like phallapodeme. The Syrphoidea is characterized by a right-side deflexion of the hypopygium, whereas the Schizophora possess gonostyli that are adducted against the hypandrium, and circumversion that is completed entirely within the puparium.

CATALOG OF THE DOLICHOPODIDAE (DIPTERA) OF AMERICA NORTH OF MEXICO

Abstract A catalog of the American species of Dolichopodidae north of Mexico is provided. Fifty-five valid genera and 1288 valid species and 3 subspecies in 14 subfamilies are listed. Compared to the North American catalog by Foote et al. (1965), 224 species are added, and there are 15 replacement names, 84 synonyms, 5 species removed from synonymy, and 76 transfers, as well as 10 species included in the previous catalog that are removed from the new list. Furthermore, 13 additional genera and 3 generic synonymies are included. Nomenclatural changes proposed in this catalog consist of 1 new generic synonymy, 4 new synonyms, 2 new names, 40 new combinations, and 1 species newly removed from synonymy. Additional nomenclatural changes and problems are discussed. All validly named taxa described before February 2004 are included. For each genus, the species list is preceded by notes on its distribution, biology, and ecology as well as a list of pertinent keys and revisions, when such information is available. For each valid species name, the list includes the author, year of publication, page number of original description, type depository and locality, and distribution in North America by state or province and occurrence in other realms, as well as taxonomic notes wherever applicable. Additional distribution records from the literature have been added. All known synonyms of genera and species as well as errors, emendations, and misidentifications are included. A bibliography of 492 publications is provided as well as an index to over 1838 included names. Statistics and comments on the history of North American dolichopodid research are presented. The diversity and distribution of Nearctic Dolichopodidae and their relationships with the fauna of other realms is tabulated and discussed.

Brachyceran Diptera (Insecta) in Cretaceous ambers, Part IV, Significant New Orthorrhaphous Taxa

Abstract Thirteen species of basal Brachycera (11 described as new) are reported, belonging to nine families and three infraorders. They are preserved in amber from the Early Cretaceous (Neocomian) of Lebanon, Albian of northern Spain, upper Albian to lower Cenomanian of northern Myanmar, and Late Cretaceous of New Jersey USA (Turonian) and Alberta, Canada (Campanian). Taxa are as follows, with significance as noted: In Stratiomyomorpha: Stratiomyidae (Cretaceogaster pygmaeus Teskey [2 new specimens in Canadian amber], Lysistrata emerita Grimaldi & Arillo, gen. et sp. n. [stem-group species of the family in Spanish amber]), and Xylomyidae (Cretoxyla azari Grimaldi & Cumming, gen. et sp. n. [in Lebanese amber], and an undescribed species from Spain). In Tabanomorpha: Tabanidae (Cratotabanus newjerseyensis Grimaldi, sp. n., in New Jersey amber). In Muscomorpha: Acroceridae (Schlingeromyia minuta Grimaldi & Hauser, gen. et sp. n. and Burmacyrtus rusmithi Grimaldi & Hauser gen. et sp. n., in Burmese amber, the only definitive species of the family from the Cretaceous); Mythicomyiidae (Microburmyia analvena Grimaldi & Cumming gen. et sp. n. and Microburmyia veanalvena Grimaldi & Cumming, sp. n., stem-group species of the family, both in Burmese amber); Apsilocephalidae or near (therevoid family-group) (Kumaromyia burmitica Grimaldi & Hauser, gen. et sp. n. [in Burmese amber]); Apystomyiidae (Hilarimorphites burmanica Grimaldi & Cumming, sp. n. [in Burmese amber], whose closest relatives are from the Late Jurassic of Kazachstan, the Late Cretaceous of New Jersey, and Recent of California). Lastly, two species belonging to families incertae sedis, both in Burmese amber: Tethepomyiidae (Tethepomyia zigrasi Grimaldi & Arillo sp. n., the aculeate oviscapt of which indicates this family was probably parasitoidal and related to Eremochaetidae); and unplaced to family is Myanmyia asteiformia Grimaldi, gen. et sp. n., a minute fly with highly reduced venation. These new taxa significantly expand the Mesozoic fossil record of rare and phylogenetically significant taxa of lower Brachycera.

Male terminalia of Diptera (Insecta): a review of evolutionary trends, homology and phylogenetic implications

The male terminalia character system in Diptera is reviewed. The phylogenetic implications of apomorphic changes are traced on published cladograms. New synapomorphies include: anteroventral parameral apodeme for the Tipulomorpha; parameral sheath encompassing desclerotized aedeagus for Neodiptera (exclusive of Axymyiidae); endoaedeagus for Xylophagomorpha + Tabanomorpha. Apystomyiidae are classified as the sister group to the Eremoneura based on four synapomorphies (lateral ejaculatory processes absent, subepandrial sclerite extending from base of hypoproct to phallus, bacilliform sclerites extending to tips of the epandrium and surstyli functionally developed, but not articulated) and lack of eremoneuran synapomorphies (i.e., loss of gonostyli, presence of postgonites and phallic plate). The Diptera sperm pump with a functional ejaculatory apodeme is a possible autapomorphy of Diptera, exclusive of Nymphomyiidae and Deuterophlebiidae. Internal details of the male terminalia of Sylvicola and Mycetobia (Anisopodidae), Hilarimorpha (Hilarimorphidae) and Apystomyia (Apystomyiidae) are newly illustrated and homologies of the aedeagus, paramere and sperm pump of the Tipuloidea are clarified.

The Neotropical genera Macrostomus Wiedemann and Porphyrochroa Melander (Diptera, Empididae, Empidinae)

Macrostomus ferrugineus (Fabricius, 1805), the type-species of Macrostomus Wiedemann, 1817 and Porphyrochroa palliata (Coquillett, 1902), the type-species of Porphyrochroa Melander, 1927 are redescribed and figured from the types. Both Neotropical genera are considered sistergroups and their synapomorphies are presented. The species belonging to both genera, and those Neotropical species remaining in the heterogeneous group Rhamphomyia Meigen, 1822 are listed. A key to genera is provided, 25 species are transferred to Porphyrochroa: P. abdominalis (Bezzi, 1905) comb. nov., P. argyrina (Bezzi, 1909) comb. nov., P. barueri (Smith, 1962) comb. nov., P. carrerai (Smith, 1962) comb. nov., P. catarinae (Smith, 1962) comb. nov., P. cyanogaster (Wheeler & Melander, 1901) comb. nov., P. digitata (Smith, 1962) comb. nov., P. distinctipennis (Smith, 1962) comb. nov., P. divisa (Smith, 1962) comb. nov., P. dolichocera (Bezzi, 1905) comb. nov., P. fasciventris (Curran, 1931), P. furcifer (Wheeler & Melander, 1901) comb. nov., P. galactodes (Bezzi, 1909) comb. nov., P. juri (Smith, 1962) comb. nov., P. micrargyra (Bezzi, 1909) comb. nov., P. monstrosa (Bezzi, 1909) comb. nov., P. mundurucu (Smith, 1962) comb. nov., P. mura (Smith, 1962) comb. nov., P. orthoneura (Bezzi, 1905) comb. nov., P. palliata (Coquillett, 1902), P. penicillata (Bezzi, 1909) comb. nov., P. perpulchra (Bezzi, 1909) comb. nov., P. pulchriventris (Bezzi, 1905) comb. nov., P. rotundipennis (Bezzi, 1905) comb. nov., P. seticauda (Smith, 1963) comb. nov., P. variseta (Smith, 1962) comb. nov. and P. wiedemanni (Smith, 1962) comb. nov. A lectotype is designated for P. palliata (Coquillett, 1902).

The New World genera of Parathalassiinae (Diptera: Empidoidea: Dolichopodidae s.l.), with new species of Thalassophorus and Eothalassius

Abstract The parathalassine genera Thalassophorus Saigusa, Eothalassius Shamshev and Grootaert, and Chimerothalassius Shamshev and Grootaert are recorded from the New World for the first time. Thalassophorus arnaudi Brooks and Cumming sp. nov. is described from specimens collected at coastal localities in British Columbia, Oregon, and California, and represents the second known species in the genus, the type species being T. spinipennis Saigusa, known only from Rishiri Island in Hokkaido Prefecture, Japan. Detailed illustrations of the male genitalia of T. spinipennis are provided. Eothalassius borkenti Cumming and Brooks sp. nov. is described from specimens collected along the Pacific coast of Costa Rica, bringing the total number of described species of this former Southeast Asian genus to three, with one probable additional described species in the Mediterranean region. The genus Chimerothalassius, which was previously known from a single New Zealand species, is recorded from the island of Dominica, based on a female specimen plus a slide-mounted wing of an undescribed species. A new undescribed parathalassiine genus is also recorded from Chile, based on limited material of two undescribed species. A key to the six genera of Parathalassiinae in the New World, including Parathalassius Mik and Microphorella Becker, is given, as are some preliminary remarks on the limits and phylogenetic relationships of the parathalassiine genera.

Systematic revision of Nearctic species of Achalcus Loew (Diptera: Dolichopodidae) with comments on their phylogeny, ecology and zoogeography

The species of the genus Achalcus Loew from the Nearctic Region are revised and a key to males and females is provided. Achalcus utahensis (Harmston & Miller) and A. oregonensis (Harmston & Miller) are redescribed, and A. californicus sp.n., A. dytei sp.n. and A. similis sp.n. are described. Together with eight of the nine known Palaearctic species of the genus, they represent a monophyletic species group, which is characterized by the presence of five pairs of dorsocentral bristles and the incorporation of the epandrial setae onto the shaft of the epandrial lobe. Like their Palaearctic congeners, most Nearctic species occur in damp habitats, such as coastal forests and riparian vegetation. Most specimens were collected in the Pacific coastal states from British Columbia to California, but one species occurs in Utah, and one of six species which were not described because males are lacking, was discovered in Ontario.

Revision of the Nearctic Species of Heleodromia (Diptera: Empidoidea: Brachystomatidae)

Abstract The Nearctic species of the brachystomatid genus Heleodromia Haliday are revised. The following six species are recognized: Heleodromia boreoalpina Saigusa, H. chillcotti Sinclair sp. nov., H. cranehollowensis Cumming and Coovert sp. nov., H. irwini Wagner, H. pullata (Melander), and H. woodi Brooks sp. nov. The following new Palearctic distribution records are also reported: H. boreoalpina (Russian Far East), H. irwini (Germany, Russian Far East), and H. immaculata Haliday (Russian Far East). A key to adults, illustrations of male terminalia, and known distributions are included.

Revision of the genus Macrostomus Wiedemann (Diptera, Empididae, Empidinae). IV. The amazonensis species-group.

Six new species of Macrostomus Wiedemann are described, namely M. acreanus sp. nov. from Brazil (Acre state), M. amazonensis sp. nov. from Guyana and Brazil (Roraima, Amazonas, Pará, Rondônia, and Mato Grosso states), M. albicaudatus sp. nov. from Brazil (Roraima, Amazonas, and Pará states), M. paraiba sp. nov. from Brazil (Paraíba state), M. trombetensis sp. nov. from Brazil (Pará state) and M. xavieri sp. nov. from Guyana and Brazil (Amazonas state). The six species are all treated in the M. amazonensis species-group, which is defined on the basis of one pair of ocellar setae and no supra-alar postsutural setae in combination with characters of the male and female terminalia. A key to the included species is presented and the geographical distributions of the species are mapped.

Revision of the Empis subgenus Enoplempis Bigot, east of the Rocky Mountains (Diptera: Empididae).

The Empis subgenus Enoplempis Bigot, east of the Rocky Mountains of North America is revised. A total of 19 species are recorded from this region including seven new species: E. (En.) amytis Walker, E. (En.) appalachicola Sinclair sp. nov., E. (En.) arthritica Melander, E. (En.) ctenonema Melander, E. (En.) enodis Melander, E. (En.) gladiator Melander, E. (En.) gulosa Coquillett, E. (En.) loripedis Coquillett, E. (En.) montywoodi Brooks sp. nov., E. (En.) nodipoplitea Steyskal, E. (En.) nuda Loew, E. (En.) pectinata Sinclair sp. nov., E. (En.) penicillata Brooks sp. nov., E. (En.) prodigiosa Cumming sp. nov., E. (En.) snoddyi Steyskal, E. (En.) stenoptera Loew, E. (En.) tridentata Coquillett, E. (En.) vockerothi Cumming sp. nov., E. (En.) volsella Sinclair sp. nov. The following new synonymies are designated: E. (En.) longipes Loew, E. (En.) longeoblita Steyskal, E. (En.) deterra Walley = E. (En.) amytis; E. (En.) cacuminifer Melander = E. (En.) gulosa. Lectotypes are designated for the following species: E. (En.) arthritica, E. (En.) cacuminifer, E. (En.) ctenonema, E. (En.) enodis, E. (En.) gladiator, E. (En.) loripedis, E. (En.) stenoptera and E. (En.) tridentata. A key to eastern species is presented and distributions illustrated. The form of nuptial gift presentation displayed within this group, including the use of balloons (with or without prey) and unwrapped prey are indicated for species when known.