World Catalog of the Beach-Fly Family Canacidae (Diptera) WAYNE N. MATH IS I SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? NUMBER 536 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." 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Smithsonian Contributions toZoology, number 536,18 pages, 1992.?All genera and species of the dipterous family Canacidae, more commonly known as beach or surf flies, are cataloged. Included are 113 species and 12 genera that are arranged within a classification of two tribes and three subfamilies. The distribution of each species is given by major zoogeographic region(s) and country(ies) within each region. Information on the natural history, as available in the literature, and depository of primary types are also provided. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Monlastrea cavernosa (Linnaeus). Library of Congress Cataloging-in-Publication Data Matfiis, Wayne N. World catalog of the beach-fly family Canacidae (Diptera) p. cm. - (Smithsonian contributions to zoology ; no. 536) Includes bibliographical references and index. 1. Canacidae?Classification. 2. Canacidae. I. Title. II. Series. QL1.S54 no. 536 [QL537.C25] 1992 591 s-dc20 [595.77M] 92-29981 @ The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48?1984. Contents Page Introduction 1 Format 1 Faunal Treatments 2 Revisionary Treatments 2 Abbreviations 2 Acknowledgments 3 Catalog 3 Literature Cited 14 Index 18 in World Catalog of the Beach-Fly Family Canacidae (Diptera) Wayne N. Mathis Introduction True flies of the family Canacidae, more commonly known as beach or surf flies, are found in temperate and tropical zones throughout the world, usually in association with maritime beaches. The family has comparatively few species, at least by hexapod standards, and except for Wirth's review (1951), the family has never been treated comprehensively. Wirth's paper is now of limited use, especially at the species level, where less than one-third of the present species were treated (32 of 113 species). Most species and several genera have been described since 1951, and few comprehensive treatments are now available except on a regional level (see review of faunal and revisionary papers below). The purpose of this paper is to partially address the lack of comprehensive works through publication of this catalog, which is intended to have worldwide coverage and to include all taxa that have been described. Catalogs or checklists are an indispensable tool for anyone needing an up-to-date reference to a currently accepted name and frequently to other pertinent information such as biblio- graphic and distributional data. This is so because most information is filed under a species' scientific name, which then becomes the key to retrieval of information from the literature. The system, however, is dynamic and subject to interpretation. The taxonomic literature is constantly changing to reflect current work, and some species are known by several names. Thus a complete listing of names, including synonyms, is an important starting point for locating information, whether as the basis for applied and basic research or simply to satisfy a curiosity. Wayne N. Mathis, Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Reviewers: F. Christian Thompson, Systematic Entomology Labora- tory, USDA, in National Museum of Natural History, Smithsonian Institution, Washington, DC. 20560; Willis W. Wirth, Systematic Entomology Lab (retired). The information included in a catalog is usually arranged in a logical and organized format that allows for its convenient and rapid conveyance?in short, a quick and easy storage and retrieval system. The format and amount of information presented varies greatly, however, and these issues have in part led to semantic debates over differences between the terms "checklist" and "catalog" and attempts to obviate the issue through use of a more neutral term, such as database (Cogan et al., 1980; Thompson and Knutson, 1987). My use of the term catalog is intended to convey a more comprehensive treatment, including information on all valid names, synonyms, type species, and deposition of primary types. The bibliographic section includes complete references (author, date, original and most subsequent citations), and distributional and other biotic information, as available in the literature, are also provided. Not all citations that occur in the literature of beach flies are included in this catalog or the bibliography section, especially where I suspect that the species being treated was misidentified, and inclusion would further promulgate inaccurate distribu- tional data. The sequence of taxa, especially at the generic level and above, should not be interpreted to represent a phylogenetic scheme. Indeed, one subfamily, Zaleinae, comprising only three species, is only questionably included in the family and in this catalog. When McAlpine (1982, 1985) proposed Zaleinae he was unsure of its phylogenetic relationship, although he did associate it with the Canacidae. Others have followed that precedent (Mathis, 1989a), mostly for convenience and completeness. The phylogenetic relationships of Zaleinae remain enigmatic, however, and further study may reveal its placement elsewhere, such as with the Tethinidae. FORMAT.?The format I have adopted follows that advo- cated by systematists from the Systematic Entomology Laboratory (United States Department of Agriculture) (Hodges, pers. comm.). Details are illustrated in the following hypothetical examples of generic and species entries (genera Xus and Yus and species albus and zeus). All valid generic and species names are indicated in bold face type. SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Genus Xus Author(s) (number of species in the genus) Xus Author(s), yeanpage. Type species: Xus albus Authors), year, method of type designation.?Author(s), yeanpage [annotation(s)]. Yus Author(s), yeanpage. Type species: Yus zeus Authors), year, method of type designation.?Author(s), yeanpage [annotation(s) such as "synonymy"]. albus Author(s). Geographic distribution by major faunal realm(s): Country (province or state). Yus albus Author(s), yeanpage [primary type(s) and gen- ders) (deposition information); type locality (Country. Province or state: specific locality (annotation(s) such as elevation or habitat)].?Author, yeanpage [annota- tion^)]. Xus zeus Author(s), year: page [primary type(s) and gender(s) (deposition information); type locality (Coun- try. Province or state: specific locality (annotation(s) such as elevation or habitat)].?Author, yeanpage [annotation(s) such as "synonymy"]. Yus zeus.?Author(s), yeanpage [annotation(s) such as "generic combination"]. Within a taxon, the subordinate taxa are listed alphabetically, i.e., genera within a tribe, species within a genus. Although this is the first treatment of the family on a worldwide basis since Wirth (1951), there are several papers that treat beach flies on a regional basis. These may be of interest and use to the reader, and a summary of these and revisionary works are provided here. FAUNAL TREATMENTS (papers listed chronologically under major faunal realms).?Afrotropical: Frey (1958b, fauna of the Cape Verde Islands); Wirth (1960, South African fauna); Mathis and Wirth (1979, Malagasy fauna); Cogan (1980, catalog); Canzoneri (1982, fauna of Sierra Leone); Canzoneri (1987, fauna of the Sudan); Mathis (1988b, fauna of the Seychelles); Mathis and Freidberg (1991, tribe Canacini and subfamily Nocticanacinae). Australasian/Oceanian: Delfinado (1970, fauna of New Guinea); Hardy and Delfinado (1980, Hawaiian fauna); Mathis (1989a, catalog). Nearctic: Wheeler (1952, fauna of the United States); Wirth (1965, catalog); Cole (1969, fauna of Western North America); Wirth (1987, general description and discussion of family, key to nearctic genera). Neotropical: Wirth (1969b, fauna of the Galapagos Islands); Wirth (1975, catalog); Mathis (1989b, fauna of the Caribbean and Gulf of Mexico). Oriental: Delfinado (1975, Sri Lankan fauna); Delfinado and Wirth (1977, catalog). Palearctic: Becker (1926, palearctic fauna); Seguy (1934, fauna of France); Frey (1936,1945,1949,1958a, faunas of the Canary Islands, Azores, Madeira, and Canary Islands respec- tively); Miyagi (1963, Korean fauna); Stackelberg (1970, western palearctic fauna); Cogan (1976, checklist of the British fauna); Mathis (1982a, fauna of Israel); Mathis and Freidberg (1982, review of western palearctic species); Cogan (1984, catalog). REVISIONARY TREATMENTS (papers listed chronologi- cally).?Wirth (1964, Trichocanace Wirth); Wirth (1969a, Canaceoides Cresson); Wirth (1970, the snodgrassii group = Canacea Cresson); Mathis and Wirth (1978, Paracanace Mathis and Wirth); Mathis (1982a, Canace Haliday); Mathis (1982b, Isocanace Mathis); McAlpine (1982, Zale McAlpine = Zalea McAlpine, 1985); Mathis, 1989b, the texensis group of Nocticanace Malloch). ABBREVIATIONS.?To economize on space I have used well-known acronyms for museums, especially to indicate the deposition of a primary type(s), and also for some locality data. These abbreviations are as follows: AM Australian Museum, Sydney, Australia AMNH American Museum of Natural History, New York, New York, USA ANSP Academy of Natural Sciences of Philadelphia, Pennsylvania, USA BBM Bernice P. Bishop Museum, Honolulu, Hawaii, USA BMNH former British Museum (Natural History), collec- tions in The Natural History Museum, London, England CAS California Academy of Sciences, San Francisco, California, USA DEI former Deutsches Entomologisches Institut, collec - tions in the Institut fur Pflanzenschutzforschung, Zweigstelle Eberswalde, Abteilung Taxonomie der Insekten, Eberswalde, Germany DCSA Dipterorum Collectionis Strobl, Admont, Austria HUS Hokkaido University, Sapporo, Japan ICC Instituto Oswaldo Cruz, Rio de Janeiro, Brazil LACM Los Angeles County Museum of Natural History, Los Angeles, California, USA MCV Museo Civico di Storia Naturale de Venezia, Venice, Italy MNHN Museum National d'Histoire Naturelle, Paris, France MRAC Musee Royal de l'Afrique Centrale (Koninklijk Museum voor Midden-Afrika), Tervuren, Bel- gium NMI National Museum of Ireland, Dublin, Ireland NMP Natal Museum, Pietermaritzburg, South Africa NZAC New Zealand Arthropod Collection, Entomology Division, DSIR, Auckland, New Zealand SUJ Saikyo University, Kyoto, Japan SMN Staatliches Museum fur Naturkunde in Stuttgart, Ludwigsburg, Germany TMC Transvaal Museum Collection, Pretoria, South Africa NUMBER 536 UMO University Museum, Oxford University, Oxford, England USNM former United States National Museum, collections in the National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA ZIL Zoological Institute, Lund University, Lund, Swe- den ZMA Instituut voor Taxonomische Zoologie, Zoologisch Museum, Universiteit van Amsterdam, Amster- dam, Netherlands ZMHU Zoologisches Museum, Humboldt Universitat, Berlin, Germany Abbreviations for primary types: HT = Holotype, LT = Lectotype, ST = Syntype, T = Type (the specific status of the primary type is unknown). Abbreviations for states of the USA: CA = California, FL = Florida, MA = Massachusetts, MD = Maryland, ME = Maine, NC = North Carolina, RI = Rhode Island, SC = South Carolina, TX = Texas, VA = Virginia, WA = Washington. Abbreviations for Canadian provinces and territories: BC = British Columbia, NB = New Brunswick, PE = Prince Edward Island. Abbreviations for Mexican states: BCN = Baja California None, BCS = Baja California Sur, QNR = Quintana Roo, TAB = Tabasco. Abbreviations for Australian states: NSW = New South Wales, QLD = Queensland. ACKNOWLEDGMENTS.?The support and encouragement of numerous persons have directly contributed to the production of this catalog, and I gratefully acknowledge and thank them all for their efforts, time, and opinions. I have especially benefited from discussions on format and cataloging procedures from Drs. Brian V. Brown, Curtis W. Sabrosky, Ronald W. Hodges, F. Christian Thompson, Norman E. Woodley, and Amnon Freidberg. An earlier draft of this paper was critically reviewed by Drs. F. Christian Thompson and Willis W. Wirth. These reviews were invaluable, especially as Thompson is intimately acquainted with the literature on Diptera, and Wirth has contributed much to our knowledge of beach flies. The following curators provided access to collections, primary types in particular, and other information that has substantially enhanced the value of this work: Dr. David K. Me Alpine (AM), Dr. Neal L. Evenhuis (BBM), Dr. Paul H. Amaud, Jr. (CAS), Dr. S. Takagi (HUS), Drs. J. Decelle and R. Jocque (MRAQ, Dr. Charles Hogue (LACM), Mr. Silvano Canzoneri (MCV), Dr. Loic Matile (MNHN), Dr. James O'Connor (NMI), Dr. Brian Stuckenberg (NMP), Dr. Hugo Andersson (ZIL), Dr. H. Schumann (ZMHU). Catalog Family CANACIDAE Jones (113 species) Canacenae Jones, 1906:170, 198 [as a subfamily of Ephydri- dae, incorrect formation of the family-group name]. Type genus: Canace Haliday, 1837. Canaceidae.?Hendel, 1916:297 [incorrect formation of the family-group name].?Wirth, 1951:245-275 [revision]. Canacidae.?Enderlein, 1935:235.?Mathis, 1982b:l-29 [classification]. Subfamily CANACINAE Jones (32 species) As above. Canaceinae.?Hendel, 1913:93 [as a subfamily of Ephydridae, incorrect formation of the family-group name]. Canacinae.?Enderlein, 1914:326 [as a subfamily of Ephy- dridae].?Malloch, 1933:4 [as a subfamily of Ephy- dridae]..?Mathis, 1982b:2 [as a subfamily of Canacidae, phylogeny]. Tribe Canacini Jones (5 species) As above. Canacini.?Mathis, 1982b:3 [as a tribe of Canacinae]. Genus Canace Haliday (5 species) Canace Haliday in Curtis, 1837:281 [published in synonymy; first made available by use in Haliday, 1839:411; see Thompson and Mathis, 1981]. Type species: Ephydra nasica Haliday, 1839, by subsequent monotypy (Hali- day, 1839:411).?Loew, 1860:29 [review]; 1874:76 [review].?Schiner, 1863:268 [review].?Rondani, 1875:176, 169 [review].?Becker, 1896:245 [review]; 1905:215 [catalog]; 1926:106 [review].?Wirth, 1951:259 [review].?Stackelberg, 1970:363 [key].? Cogan, 1980:694 [Afrotropical catalog]; 1984:125 [pal- earctic catalog].?Mathis, 1982a: 58 [review, figures of d* and 9 terminalia, heads].?Mathis and Freidberg, 1991:71-75 [review of Afrotropical fauna]. Ephydra (Canace).?Haliday, 1839:411.?Walker, 1853:268 [review]. actites Mathis. Palearctic: Spain (Canary and Madeira Is- lands). Canace salonitana, in part (misidentification).?Wirth, 1951:264 [review, figure of <$ terminalia]. Canace actites Mathis, 1982a:58 [HT 9 (USNM 76783); Spain. Canary Islands: Teneriffe; figures of head, thorax, SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY