■ Taxonomic and Host Catalogue of the Tachinidae of America North of Mexico

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The following is from pp. 1–15 of:
O'Hara, J.E. and Wood, D.M. 2004. Catalogue of the Tachinidae (Diptera) of America north of Mexico. Memoirs on Entomology, International 18: iv + 410 pp.

Abstract

This taxonomic catalogue lists the Tachinidae (Diptera) of America north of Mexico, including Bermuda and Greenland but not Hawai‘i and the West Indies. The classification is changed significantly from the last regional Tachinidae catalogue published in 1965 and recognizes 4 subfamilies, 46 tribes, 303 genera and 1345 species. New synonyms, new generic combinations and new records are cited in the introduction and noted in the catalogue. Full nomenclatural information is given for nominal genera and species, including primary type data for the latter. Updated distributions are provided for valid species. Taxonomic and nomenclatural literature pertaining to taxa at all hierarchical levels from subspecies to subfamily is referenced under valid names. An index to the names listed in the catalogue is included.

Introduction

Preamble

It has been over 35 years since the last Tachinidae catalogue for America north of Mexico was published (Sabrosky & Arnaud, 1965). During the intervening years there have been many changes to the classification and nomenclature of the Tachinidae of this region, though not a great many new species have been described. Sabrosky & Arnaud (1965) recognized 5 subfamilies (excluding the Rhinophorinae, now considered a distinct family), 69 tribes, 414 genera and 1281 species, whereas we recognize 4 subfamilies, 46 tribes, 303 genera and 1345 species. The classification adopted by us is discussed in more detail in the Classification section below. Distributions have been updated according to the holdings of the Canadian National Collection of Insects (containing about 250,000 pinned specimens of Tachinidae) and post-1965 literature but for practical reasons we were unable to record new distributional data for more than a few species in other North American collections. This catalogue covers literature published up to 1 September 2003.

The habits of tachinid flies are united by a common life history strategy, the parasitization of other arthropods, almost exclusively insects. Chief among the orders parasitized are the Lepidoptera, with the Symphyta (Hymenoptera), Hemiptera and Coleoptera also significantly attacked. Host associations are not reported in this work except for host records of primary types. The reader is referred to Arnaud's (1978) Host-parasite Catalog of North American Tachinidae for further information on this subject. For a general account of tachinid morphology, biology, behavior and classification, see the introductory pages of Wood's (1987) chapter on the Tachinidae in volume 2 of Manual of Nearctic Diptera. Wood's (1987) chapter contains an illustrated key to the tachinid genera of America north of Mexico.

Our catalogue departs most significantly from former catalogues of Tachinidae of America north of Mexico in being the first to include primary type information. It is also the first to include extensive references pertaining to subsequent literature of a systematic or nomenclatural nature. It is hoped that these additional features will increase the utility of the catalogue by responding to the needs of a broad spectrum of users.

Format

We used the software program Platypus® Version 1.1 (CSIRO, Australia) to input catalogue data and programmed a custom output from the underlying Microsoft Access® 97 database to obtain the desired format. Final changes and the index were made in Corel WordPerfect® 8.

This catalogue is arranged hierarchically according to the categories of subfamily, tribe, genus, subgenus, species and subspecies. Subordinate taxa are arranged alphabetically within each category. Synonyms are provided for taxa in the genus and species groups and are listed chronologically. Generic and subgeneric names, be they valid or synonyms, are cited in the same manner and order: taxon name, author, year (with initial if applicable), page, a note if applicable (e.g. an alternate spelling or a subsequent publication of the same taxon name and description), type species, and form of type fixation. Generic names appear in capital letters and valid generic names in bold. Emendations at all taxonomic levels are cited where known but have not been traced to their original authors. Each type species is cited in its original binomen (Recommendation 67B of ICZN, 1999). If a type species name is now considered a synonym of another name, then the valid name is cited in parentheses. The region of origin of the type species of each genus group name is provided in square brackets if the type species does not occur in America north of Mexico.

Species and subspecies are listed by valid name in bold, accompanied by author, date (always without an initial), and known distribution. Listed under each valid name are the available names that pertain to it (i.e. the available name of the valid species and available names of synonyms), each with the accompanying information: original combination (genus and species), author, year (with initial if applicable), page, and primary type information. Primary type information includes status of type(s) (holotype, lectotype, neotype, or syntypes), sex(es), type depository, type locality, and host name if applicable. If a neotype or lectotype was designated then a citation to the designation is given. Each host name is cited in its current combination, followed in square brackets by the name used in the publication cited unless the name has not changed. Additional notes are included as necessary.

Generic and specific synonyms are generally listed only if their names apply to taxa described from America north of Mexico. For instance, Blepharigena Rondani, 1856, with type species Tachina trepida Meigen, 1824, is not listed as a synonym of Athrycia Robineau-Desvoidy, 1830, because A. trepida does not occur in America north of Mexico. Similarly, none of the nine synonyms of Voria ruralis Fallén in the Palearctic region as listed by Herting (1984) is included under V. ruralis because their names are based on Old World type material. Exceptions have been made for familiar names which might cause confusion by their absence. For example, Euphasiopteryx Townsend, 1915, is listed as a synonym of Ormia Robineau-Desvoidy, 1830, even though its type species is from South America because it has been used as a generic name for species in America north of Mexico.

Original literature was checked, except in the very rarest of instances, to ensure the accuracy of dates, titles, pagination, names, type fixations and primary type information. We did not authoritatively review the dating of older works but have largely followed Stone et al. (1965) with changes based on the more recent works of Evenhuis (1989, 1997, 2003) and Thompson et al. (1999).

Early authors did not always state whether their type series consisted of one or more specimens. In such instances we have generally referred to a single known specimen as the holotype unless there is reason to believe that other specimens may have existed or may exist in other collections (following among others Crosskey, 1973, 1974, 1976). One might consider this action to be contrary to Recommendation 73F of the International Code of Zoological Nomenclature (ICZN, 1999), “Avoidance of assumption of holotype,” but we believe that there should be some flexibility in the assumption of holotype vs. syntype. We do not see the necessity, for instance, of treating all of Francis Walker’s Tachinidae types in BMNH as syntypes when we know that the collection has been kept largely intact and that Walker frequently described species from single specimens. Crosskey (1974: 272-275) elegantly argued in favour of an assumption of holotype in a section entitled, “Holotype or lectotype? The status of a single extant type from a type-series of unknown size” in his study of the British Tachinidae of Walker and Stevens, and we are in basic agreement with the views expressed therein. In cases where an author’s description contains an indication but not an explicit statement that a type series comprised more than one specimen – for instance by mention of both sexes, variation, multiple localities or size range – we use the term syntype even if only a single specimen remains in a collection. We use the term “type(s)” when we are unsure whether a type series consisted of more than one specimen and no type specimens were located. Reference to a type number or dissection number in an original description (e.g. Coquillett, 1897, 1898; Townsend, 1912b, 1919c) is treated as a holotype designation, provided a holotype was appropriately labelled (after Sabrosky, 1983; Wood, 1985).

Type-species designations are cited by original designation, monotypy, subsequent designation, or rarely subsequent monotypy. Subsequent designations are of two kinds, either direct (with an explicit designation of a lectotype) or indirect by mention of a primary type (holotype or “type”). The latter kind is termed a lectotype “fixation” herein. The International Code of Zoological Nomenclature (ICZN, 1999) is not precise about whether the mention of a primary type can be accepted as a lectotype fixation in all situations (see Art. 74.6) but there is a well-established precedent for the sort of fixations recognized herein in such works as Arnaud (1963c), Sabrosky & Arnaud (1963), Crosskey (1969, 1971, 1976) and O’Hara (1994, 2002). Most such fixations are attributable to Townsend’s (1936–1942) Manual of Myiology in which a holotype (“Ht”) was cited for all type species regardless of whether a holotype had been designated in the original description. These references to a “Ht” are accepted as lectotype fixations provided a holotype was not originally designated and a single specimen can be distinguished as the lectotype from among others in a type series, based on the information provided by Townsend (see in particular Crosskey, 1969: 88).

A slight change in the International Code of Zoological Nomenclature between the 2nd and 3rd editions (ICZN, 1964; ICZN, 1985) affects the manner in which certain type species designations are cited compared to Sabrosky & Arnaud (1965), Guimarães (1971) and Wood (1985), among others. Those authors considered the combined description of a new genus and a single new species as a type species fixation by original designation if the expression “gen. n., sp. n.” or equivalent accompanied the species name and the description was published prior to 1931. This was based on Article 68(b)(i) of the Code (ICZN, 1964) which stated: “The formula ‘gen.n., sp.n.’, or its exact equivalent, applied before 1931 to only one of the new nominal species included in a newly established nominal genus, is to be interpreted as original designation if no other type-species was designated.” In the next edition of the Code (ICZN, 1985), the words “only one of the new nominal species” were changed to “only one of two or more new nominal species.” The wording of the same Article in the most recent Code (ICZN, 1999; as Article 68.2.1) is very close to that of ICZN (1985). Hence, it is now necessary to treat the combined description of a new genus and a single new species as type fixation by monotypy regardless of whether the expression “gen. n., sp. n.” was used. Where the expression “gen. n., sp. n.” or equivalent was used in association with one of two or more new species, or in association with one new and one or more described species as was done with Dexodes Brauer & Bergenstamm, 1889, and Phasiopteryx Brauer & Bergenstamm, 1889, type fixation by original designation applies. If a new genus name was “proposed for” or “erected for” a single nominal species but not explicitly designated as the type species (as required by the Code for type fixation by original designation), then we treated this situation as type fixation by monotypy.

Taxonomic and nomenclatural literature pertaining to taxa at all hierarchical levels is referenced in the text under valid names. These citations are based on a near exhaustive search of North American tachinid literature and also include many extralimital works that have content relevant to the taxa catalogued here. Literature devoted to ecological, host or distributional data on Tachinidae is not included, nor are most of the pre-1900 general works by such European authors as Bigot, Brauer & Bergenstamm, Fallén, Macquart, Meigen, Robineau-Desvoidy, Rondani, Schiner and Zetterstedt. These early works are excluded because they contain keys, redescriptions and classifications that are now mostly of historical interest. This could also be said of some of works included here, but we have attempted to cover virtually all of the relevant literature produced by North American authors on the Tachinidae of America north of Mexico during the past century. We also include pre-1900 works by certain North American authors, most notably Coquillett, Townsend and Williston. We have noted misidentifications in the literature where these are known to us but we have not otherwise commented on the value or accuracy of the information contained within the cited works, which varies tremendously.

The following format is used for references under valid names. References are listed in chronological order. Under the genus category, an author name does not accompany a generic name unless the nominal genus cited belongs to a different valid genus. Under the species category, an author name does not accompany a specific epithet unless the nominal species belongs to a different valid species. If synonyms are listed for a valid genus or species, then each reference cites the nominal genus or nominal species used by the author. The spelling of a taxonomic name follows that of the cited author regardless of whether the spelling now represents an error or unjustified emendation, or does not agree in gender with the current genus. Names representing errors and emendations are written within quotation marks.

Collections housing primary types

Collections cited in the catalogue are referred to by the acronyms listed below. Most of these collections house primary types of Tachinidae of America north of Mexico but some are cited only for distributional information. We examined all of the primary types in AMNH, CNC, MCZ, MZLU, NHRS and SEMK, almost all of the primary types in BMNH and USNM, and some of the primary types in many of the other major collections including CAS, CUI, MNHN and NHMW. Numerous curators were also helpful in providing information about primary types in their care (see Acknowledgements section).

AMNH American Museum of Natural History, Department of Entomology, New York, New York, USA.
ANSP Academy of Natural Sciences, Department of Entomology, Philadelphia, Pennsylvania, USA.
BMNH Natural History Museum (formerly British Museum (Natural History)), Department of Entomology, London, UK.
CAS California Academy of Sciences, Department of Entomology, San Francisco, California, USA.
CNC Canadian National Collection of Insects, Agriculture & Agri-Food Canada, Ottawa, Ontario, Canada.
CUI Cornell University, Department of Entomology, Ithaca, New York, USA.
DEI Deutsches Entomologisches Institut, Eberswalde, Germany.
EMEC Essig Museum of Entomology, University of California, Berkeley, California, USA.
FSCA Florida State Collection of Arthropods, Florida Department of Agriculture, Gainesville, Florida, USA.
INHS Illinois Natural History Survey Insect Collection, Urbana, Illinois, USA.
ISNB Insitut royal des Sciences naturelles de Belgique, Bruxelles, Belgium.
LSUK Linnaean Society, London, UK [contact BMNH].
MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA.
MHNL Musée d'Histoire Naturelle de Ville de Lille, Lille, France.
MNHN Muséum National d'Histoire Naturelle, Paris, France.
MTEC Montana Entomology Collection, Montana State University, Bozeman, Montana, USA
MZF Museo Zoologico 'La Specola', Florence, Italy.
MZLU Museum of Zoology, Lund University, Lund, Sweden.
MZUT Museo e Instituto di Zoologia Sistematica dell'Università di Torino, Turin, Italy.
NHMW Naturhistorisches Museum Wien, Vienna [Wien], Austria.
NHRS Swedish Museum of Natural History (= Naturhistoriska Riksmuseet), Department of Entomology, Stockholm, Sweden.
NYSM New York State Museum, University of the State of New York, Albany, New York, USA.
OSU Ohio State University, Columbus, Ohio, USA.
RMNH Nationaal Natuurhistorisch Museum (formerly Rijksmuseum van Natuurlijke Historie), Leiden, The Netherlands.
SDSU South Dakota State University, Severin-McDaniel Insect Collection, Brookings, South Dakota, USA.
SEMK Snow Entomological Museum, University of Kansas, Lawrence, Kansas, USA.
SIIS Staten Island Institute of Arts and Sciences, Staten Island, New York, USA.
SMF Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am Main, Germany.
UBC University of British Columbia, Spencer Entomological Museum, Vancouver, British Columbia, Canada.
UCDC R.M. Bohart Museum of Entomology, University of California, Davis, California, USA.
UGG University of Guelph, Department of Environmental Biology, Guelph, Ontario, Canada.
UMMZ University of Michigan, Museum of Zoology, Ann Arbor, Michigan, USA.
UMNH Utah Museum of Natural History, University of Utah, Salt Lake City, Utah, USA.
UMQ Université de Montréal, Département des Sciences Biologiques, Montréal, Québec, Canada.
UNSM University of Nebraska State Museum, Lincoln, Nebraska, USA.
USNM National Museum of Natural History (formerly United States National Museum), Department of Entomology, Smithsonian Institution, District of Columbia, USA.
WSUP Washington State University, James Entomological Collection, Pullman, Washington, USA.
WVU West Virginia University, West Virginia University Arthropod Collection, Morgantown, West Virginia, USA.
ZIN Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia.
ZMHB Museum für Naturkunde der Humboldt Universität zu Berlin, Berlin, Germany.
ZMUC Zoological Museum, University of Copenhagen, Copenhagen, Denmark.
ZMUK Zoologisches Museum, Universität Kiel, Kiel, Germany.
ZMUO Zoological Museum, University of Oslo, Oslo, Norway.

Geographic coverage and distributional data

The geographic coverage of this catalogue is America north of Mexico including Bermuda and Greenland but not Hawai‘i and the West Indies, following Stone et al. (1965).

Distributions are given for each valid species, in so far as they are known. For most species we have cited the distributions of Sabrosky & Arnaud (1965) (as indicated by “S&A, 1965” in the text), followed by additional distributional information obtained from a thorough search of the Tachinidae holdings in the Canadian National Collection (approximately 250,000 specimens) and a few other sources on an ad hoc basis. We generally did not attempt to verify the accuracy of the original Sabrosky & Arnaud (1965) distributions because of the difficulties involved in such a task. Distributions published after Sabrosky & Arnaud (1965) are cited herein in place of the Sabrosky & Arnaud (1965) entries.

Distributional information is generally given in the same format as in Stone et al. (1965), in which the states and provinces that circumscribe a range are cited following as much as possible a northwest to northeast and southwest to southeast pattern. Outlining state or provincial records are cited after the main range and a “?” is placed in front of a questionable record. The province of Newfoundland and Labrador is treated as two separate areas, Labrador and the island of Newfoundland. Records for the Canadian Northwest Territories published prior to 1999 have been divided into Northwest Territories and Nunavut according to the recent territorial change, with old records that cannot be assigned to one territory or the other cited in parentheses as “Northwest Territories.”

Classification

We recognize four subfamilies of the Tachinidae, the Dexiinae, Exoristinae, Phasiinae and Tachininae. We have replaced the name Goniinae, in usage in the New World for some years (e.g. Sabrosky & Arnaud, 1965; Guimarães, 1971; Wood, 1987b), with Exoristinae, of favoured usage in the Old World. Sabrosky (1999) demonstrated that the name Exoristinae has date priority over that of Goniinae.

The Dufouriini are generally regarded as a tribe in the Dexiinae (e.g. Herting, 1984; Tschorsnig, 1985; Wood, 1987b; Shima, 1989; Richter & Wood, 1995; Ziegler, 1998), though in earlier classifications the taxon was often regarded as the subfamily Dufourinae. Recently, Sabrosky (1999) has shown that Dufouriini, dating from Dufouridae Robineau-Desvoidy, 1830, has date priority over that of Dexiini, dating from Dexiariae Macquart, 1834. Before the new Code was published (ICZN, 1999), the discovery of this circumstance would have required that the subfamily containing the Dufouriini and Dexiini be called the Dufourinae, or else an application to the International Commission on Zoological Nomenclature would have been needed to retain usage of the name Dexiinae. However, a provision in the new Code (ICZN, 1999, Article 35.5) permits a family-group name in prevailing usage to be retained even if an older family-group name is found. Hence, our usage of Dexiinae is consistent with the rules of the Code (ICZN, 1999).

Recent authors have placed rhinophorid flies in their own family (e.g. Pape, 1986; Wood, 1987a) and hence they are excluded from this catalogue. Tribal categories have been modified considerably from those of Sabrosky & Arnaud (1965), with significant influence from Herting’s (1984) Catalogue of Palearctic Tachinidae (essentially unchanged in Herting & Dely-Draskovits, 1993). Our classification below the tribal level follows the extensive reorganization of genera and species of America north of Mexico proposed by Wood (1987b) and reviewed in O’Hara & Wood (1998), except for taxonomic and nomenclatural changes published in post-1965 works by various authors. New changes below the tribal level proposed herein are given below.

Excluded species

Extralimital species listed by Sabrosky & Arnaud (1965) as introduced but not established in America north of Mexico are not included in this catalogue. The following species, listed as occurring in America north of Mexico by Sabrosky & Arnaud (1965) or other authors, are also excluded for the reasons given.

Actia panamensis Curran, 1933a: 3, was moved to Siphona s. lat. by O’Hara (1989: 129) and treated as a strictly Neotropical species. Records of this species from California and Texas (Sabrosky & Arnaud,1965: 1062) were based on misidentifications.

Aporia quadrimaculata Macquart, 1846: 297 (also 1846: 169), treated as a valid species of Paraporia Townsend by Sabrosky & Arnaud (1965: 1028) and recorded by them from Arizona, was misidentified and has not been found in America north of Mexico.

Eugymnochaeta equatorialis Townsend, 1912d: 314, treated as a valid species of Chrysotachina Brauer & Bergenstamm by Sabrosky & Arnaud (1965: 1005) and recorded by them from Florida, is a strictly Neotropical species according to O’Hara (2002). The species misidentified as C. equatorialis by Sabrosky & Arnaud (1965) was described as new by O’Hara (2002) and named Chrysotachina longipennis O’Hara.

Exorista rufata Bigot, 1889: 257, treated as a valid species of Bolomyia Brauer & Bergenstamm by Sabrosky & Arnaud (1965: 1084) and recorded by them from Arizona, was misidentified and has not been found in America north of Mexico.

Musca serva Walker, 1853: 349, described from “United States” and treated as an unplaced species of Tachinidae by Sabrosky & Arnaud (1965: 1108), belongs to the Calliphoridae. It was treated as a valid species of Opsodexia Townsend by Shewell (1987). Though unstated in print, Shewell (1987) recognized Musca serva as a senior synonym of Chaetona bicolor Coquillett, 1899, on the advice of DMW who had examined the primary types of both nominal species.

Myocera simplex Bigot, 1889: 266, treated as a valid species of Ptilodexia Brauer & Bergenstamm by Sabrosky & Arnaud (1965: 989) and Guimarães (1971) and recorded from “scattered U.S. records (?error),” was excluded without mention from Wilder’s revision of Nearctic Ptilodexia. The species was described from “Mexico” and we have no reason to believe that it exists in America north of Mexico.

Parahypochaeta heteroneura Brauer & Bergenstamm, 1891: 33 (also 1892: 337), described from “N.-Amerika” and listed by Sabrosky & Arnaud (1965: 1016) as a valid genus and species combination in the tribe Campylochetini, was determined by Sabrosky (1975) to most likely be a strictly Neotropical species.

Phorocera agilis Robineau-Desvoidy, 1830: 132, listed as an introduced and established species of Parasetigena Brauer & Bergenstamm by Sabrosky & Arnaud (1965: 1056), was based on the misidentification of Parasetigena silvestris (Robineau-Desvoidy) (following Mesnil, 1960: 635) according to Herting (1974a: 13-14). Hence, the species listed as Parasetigena agilis by Sabrosky & Arnaud (1965) is called P. silvestris herein.

Solieria borealis Ringdahl, 1947: 50, treated as a valid species of Solieria Robineau-Desvoidy by Richter & Wood (1995: 46) and recorded by them from Yukon, was misidentified and has not been found in the New World. The Yukon specimens recorded as S. borealis by Richter & Wood (1995) have been reidentified as Solieria boreotis (Reinhard).

Tachina fasciata Fallén, 1820: 5, treated as a valid species (and subspecies) of Exorista Meigen by Sabrosky & Arnaud (1965: 1054) and recorded from Greenland, does not occur in the New World according to Morewood & Wood (2002). The species misidentified as E. fasciata fasciata by Sabrosky & Arnaud (1965) was described as new by Wood (in Morewood & Wood, 2002) and named Exorista thula Wood.

Tachina hortulana Meigen, 1824: 330, treated provisionally as a valid species of Aplomya Robineau-Desvoidy by Sabrosky & Arnaud (1965: 1097) and recorded by them from British Columbia to Ontario, was misidentified and has not been found in America north of Mexico. The T. hortulana of Sabrosky & Arnaud was a Nearctic species of Nilea Robineau-Desvoidy.

Tachina leucophrys Wiedemann, 1830: 308, treated as a valid species of Leschenaultia Robineau-Desvoidy by Brooks (1947: 174) and Sabrosky & Arnaud (1965: 1083) and recorded by these authors from most of North America, is a strictly Neotropical species according to Toma & Guimarães (2002). The species misidentified as L. leucophrys by Brooks (1947) and others was described as new by Toma & Guimarães (2002) and named Leschenaultia reinhardi Toma & Guimarães.

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