Studies of Gymnomyzinae (Diptera: Ephydridae), IV: A Revision of the Shore-Fly Genus Hecamede Haliday WAYNE N. MATH IS m wtu. SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? NUMBER 5 4 1 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." This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Folklife Studies Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world of science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. Robert McC. Adams Secretary Smithsonian Institution S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y ? N U M B E R 5 4 1 Studies of Gymnomyzinae (Diptera: Ephydridae), IV: A Revision of the Shore-Fly Genus Hecamede Haliday Wayne N. Mathis SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1993 A B S T R A C T Mathis, Wayne N. Studies of Gymnomyzinae (Diptera: Ephydridae), IV: A Revision of the Shore-Fly Genus Hecamede Haliday. Smithsonian Contributions to Zoology, number 541, 46 pages, 64 figures, 1 table, 1993.?Species of Hecamede, which now number 12, are revised. New synonyms are as follows (the junior synonym is listed first): Hecamede grisescens Becker is conspecific with Notiphila albicans Meigen, Hecamede qffinis Canzoneri and Meneghini is conspecific with Hecamede brasiliensis Cresson, Hecamede lacteipennis Lamb is conspecific with Notiphila granifera Thomson, and Hemicyclops maculipleuris de Meijere is conspecific with Hecamede planifrons de Meijere. The following new species are described (type localities are noted in parentheses): Hecamede (Hecamede) africana (Kenya. 100 km N Mombasa), H. (H.) australis (Australia. New South Wales: Deewhy), H. (H.) maritima (Egypt. Sinai: Nabk), H. (H.) socotra (South Yemen. Socotra: R?s Sh6ab), H. (Soikia) tomentosa (Nigeria. Lagos), H. (S.) bocki (Australia. New South Wales: Deewhy). The monophyly of Hecamede within the tribe Hecamedini is demonstrated, and a hypothetical phytogeny of the included species is generated from a matrix of 23 characters. From this phytogeny, a classification of two subgenera {Hecamede and Soikia) is recognized, and in the subgenus Hecamede, two species groups (the planifrons group and the albicans group) are distinguished. All known species of Hecamede and its outgroups are found in the Old World, and the probable origin of the genus also occurred there, specifically within the basin of the Indian Ocean. The two species that occur in the New World appear to have arrived there as part of range expansion. No species occur along the Pacific coast of the New World. Maps, keys to subgenera and species, and illustrations (scanning electron micrographs and line drawings) are provided to assist in the identification of the species. 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 Montastrea cavernosa (Linnaeus). Library of Congress Cataloging-in-Publication Data Mathis. Wayne N. Studies of Gymnomyzinae (Diptera, Ephydridae). (Smithsonian contributions to zoology ; nos. 522,541) Includes bibliographical references. Contents: 2. A revision of the shore fly subgenus Pseudochecamede Hendel of the genus Allotrichoma Becker 4. A revision of the shore fly genus Hecamede Haliday. 1. Ephydridae?Classification. I. Title. II. Series. QL1.S54 no. 522 591s 91-14038 [QL537.E7] 595.77'4 ? 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 Methods 2 Acknowledgments 2 Tribe Hecamedini Mathis 3 Key to Genera of the Tribe Hecamedini 3 Genus Hecamede Haliday 4 Key to Subgenera of Hecamede 5 Subgenus Hecamede Haliday 5 Key to Species of the Subgenus Hecamede 6 The albicans Group 7 Hecamede (Hecamede) africana, new species 7 Hecamede (Hecamede) brasiliensis Cresson 8 Hecamede (Hecamede) albicans (Meigen) 11 Hecamede (Hecamede) australis, new species 16 Hecamede (Hecamede) granifera (Thomson) 18 Hecamede (Hecamede) inermis Malloch 24 Hecamede (Hecamede) maritima, new species 26 Hecamede (Hecamede) socotra, new species 28 The planifrons Group 29 Hecamede (Hecamede) planifrons (de Meijere) 29 Subgenus Soikia Canzoneri and Meneghini 31 Key to Species of the Subgenus Soikia 32 Hecamede (Soikia) nuda Wirth 32 Hecamede (Soikia) tomentosa, new species 35 Hecamede (Soikia) bocki, new species 37 Phylogenetic and Biogeographical Considerations 39 Characters Used in the Phylogenetic Analysis 40 Literature Cited 44 in FIGURE 1.?Frontispiece, habitus of Hecamede (Soikia) nuda Wiith. Studies of Gymnomyzinae (Diptera: Ephydridae), IV: A Revision of the Shore-Fly Genus Hecamede Haliday Wayne N. Mathis Introduction Among valid shore-fly genera, which now number more than 110, only 13 were named before Hecamede Haliday (1837), the subject of this revision. The initial naming of Hecamede, however, was not without controversy, and the early nomencla- tural history that resulted is rather convoluted. Haliday (1837) first proposed the genus in an addendum to a book on British entomology that John Curtis, an English contemporary, published. Authorship of Hecamede is thus ambiguous, with the possibility that either Haliday or Curtis was the author. Contemporaries, including Curtis, all considered Haliday as the author, even though Curtis unquestionably published the book. Although valid arguments, including precedence, could be made for either author, Thompson and Mathis (1981) elected to follow the obvious intention of the two principals and have credited authorship to Haliday. That precedent is continued here. Further compounding the nomenclatural status of Hecamede was its proposal in synonymy, a fact that was overlooked by most subsequent workers and which makes its availability and validation, and the dates of either, a consideration that is subject to a rule that is seldom used (ICZN, 1985, Article 11, Section e). Although Hecamede was initially listed without a diagnosis or description of any kind, which would normally make the name unavailable, its listing was as a synonym. According to the rules, a name first proposed as a synonym becomes available if prior to 1961 it had been adopted as the name of a taxon or treated as a senior homonym. Haliday (1839:221,224) fulfilled these requirements two years after he first listed Hecamede as a synonym by including it in a paper on British Hydromyzidae (= Ephydridae) where it was treated as a Wayne N. Mathis, Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. valid subgeneric name under the genus Notiphila Fallen, included in a key, and also given a separate diagnosis. In this same paper, Haliday (1839:224) also cited Notiphila albicans Meigen as the only included species, and this species became the type species of Hecamede by monotypy. Even though Hecamede became available in 1839, the rules (Section e) further state that such a "name dates from its first publication as a synonym," or in this case 1837. Thus by an encumbered process Hecamede was given birth into the world of zoological nomenclature in 1837 and conferred retroactive legitimacy in 1839. As an aside to the issues pertaining to publication of a name in synonymy is Haliday's listing of Notiphila albicans Meigen as an available species name under Hecamede. Most synonyms that Haliday introduced in the 1837 list did not include an available species, but where one was included, that species can serve as an indication (sense of the code, ICZN), and before 1931 would make the associated generic name available (ICZN, 1985, Article 12(b)(5)). Thus the inclusion of N. albicans further substantiates the validity of the Hecamede from the date when it was first listed in 1837. Despite nomenclatural issues that are debated from time to time (Thompson and Mathis, 1981), the status of Hecamede as a zoological taxon has remained stable from its inception and has been universally recognized by subsequent workers. Although the generic concept of Hecamede has remained unchanged, the genus has never been revised on a world basis nor within the context of a phylogenetic framework at the specific, generic, or tribal levels. As part of this revision, the genus is recharacterized, its monophyly confirmed within the phylogenetic framework of the newly established tribe He- camedini (Mathis, 1991), and the phylogenetic relationships among the included species are elaborated. The coverage of this revision is worldwide. This revision is a continuation of studies on the tribe SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Hecamedini that are intended to better characterize the tribe and elucidate the relationships between the component lineages. Genus-group taxa of this tribe that have recently been revised are: Eremotrichoma Giordani Soika (as a subgenus of Allotrichoma; Mathis, 1986a), Elephantinosoma Becker (Mathis and Deeming, 1987), Diphuia Cresson (Mathis, 1990), and the subgenus Pseudohecamede Hendel of the genus Allotrichoma Becker (Mathis, 1991). Only Allotrichoma, sensu stricto, the most speciose and widespread genus of the tribe, remains to be revised. METHODS.?The methods used generally in this study were explained previously (Mathis, 1986a, 1990, 1991). Because specimens are small, all less than 4 mm, study and illustration of the male terminalia required use of a compound microscope. To better assure effective communication about structures of the male terminalia, the terminology of other workers in Ephydridae (see references in Mathis, 1986b) has been adopted. Usage of these terms, however, should not be taken as an endorsement of them from a theoretical or morphological view over alternatives that have been proposed (Griffiths, 1972; Me Alpine, 1981). Rather, I am deferring to tradition until the morphological issues are better resolved. The terminology for structures of the male terminalia is provided directly on Figures 13 and 14 (Hecamede albicans) and is not repeated for comparable illustrations of other species. Two venational ratios are commonly used in the descriptions and are defined here for the convenience of the user (ratios are averages of three specimens). 1. Costal vein ratio: the straight line distance between the apices of R2+3 and R^j/distance between the apices of R, and R2+3* 2. M vein ratio: the straight line distance along M between crossveins (r-m and dm-cu)/distance apicad of crossvein dm-cu. The phylogenetic analysis was performed with the assistance of Hennig86 (copyrighted), a computerized algorithm that produces cladograms on the basis of parsimony. Before performing the analysis, the character data were arranged in transformation series and then polarized primarily using outgroup procedures. ACKNOWLEDGMENTS.?Although this study was based primarily on specimens in the National Museum of Natural History, numerous others were borrowed, particularly type specimens of the species previously described. To my colleagues and their institutions listed below who loaned specimens, I express my sincere thanks. Without their cooperation this study could not have been completed. AMNH ANIC ANSP American Museum of Natural History, New York, New York (Dr. David A. Grimaldi and Mr. Julian Stark) Australian National Insect Collection, CSIRO, Canberra, Australia (Dr. Donald Colless) Academy of Natural Sciences of Philadelphia, Pennsylvania (Drs. Jon K. Gelhaus and Donald Azuma) BBM Bernice P. Bishop Museum, Honolulu, Hawaii (Dr. Neal L. Evenhuis) BMNH British Museum (Natural History), London, Eng- land (Drs. Adrian Pont and Brian Pitkin) CNC Canadian National Collection, Ottawa, Canada (Dr. J.R. Vockeroth) MBP personal collection of M. Bartdk, Prague, Czech Republic (formerly Czechoslovakia) MNHN Museum National d'Histoire Naturelle, Paris, France (Dr. Loic Matile) MNRJ Museu Nacional, Rio de Janeiro, Brazil (Dra. Mdrcia S. Court) NMW Naturhistorisches Museum, Vienna, Austria (Dr. Ruth Contreras Lichten berg) NMWL National Museum of Wales, Cardiff, Wales, United Kingdom (Mr. John C. Deeming) NRS Naturhistoriska Riksmuseet, Stockholm, Sweden (Dr. Per Inge Persson) NZAC New Zealand Arthropod Collection, Auckland, New Zealand (Dr. Beverly A. Holloway) TAU Tel Aviv University (Zoology Department), Tel Aviv, Israel (Dr. Amnon Freidberg) USNM former United States National Museum, collections in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. UZMC Universitetets Zoologiske Museet, Copenhagen, Denmark (Drs. Thomas Pape and Leif Lyne- borg) WSU Washington State University, Pullman, Washing- ton (Dr. Richard Zack) ZMA Instituut voor Taxonomische Zoologie, Zoologisch Museum, Universiteit van Amsterdam, Nether- lands (Dr. Th. H. van Leeuwen) Ms. Hollis B. Williams prepared the distribution maps, and the line illustrations were carefully rendered by Mrs. Elaine R.S. Hodges. The frontispiece was produced in carbon dust by Mrs. Natalia A. Florenskaya (St. Petersburg, Russia). Mrs. Susann Braden and Ms. Victoria Godwin assisted with the preparation of the scanning electron micrographs. Drs. F.C. Thompson and C.W. Sabrosky reviewed portions of the manuscript that deal with nomenclature. For reviewing a complete draft of this paper I thank Drs. G.W. Courtney, A.L. Norrbom, and R.V. Peterson. I am also grateful to Dr. David Challinor, former Assistant Secretary for Research, Smith- sonian Institution, and Dr. Stanwyn G. Shetler, Deputy Director, National Museum of Natural History, for financial support to conduct field work through grants from the Research Opportunity Fund. Dr. J.W. Ismay made special effort to collect shore flies in Papua New Guinea; his efforts are gratefully appreciated. Dr. Curtis W. Sabrosky kindly hand NUMBER 541 carried the holotypes of H. qffinis and Soikia argentata from MNHN to Washington, D.C., and Dra. Marcia S. Couri sent me information on the holotype of H. brasiliensis. This is contribution number 352 from the Caribbean Coral Reef Ecosystems (CCRE), Smithsonian Institution, which is partly supported by a grant from the Exxon Corporation. Tribe Hecamedini Mathis Hecamedini Mathis, 1991:2. Type genus: Hecamede Haliday, 1837. DIAGNOSIS.?Head: Arista with 3 to 5 dorsally branching rays, longer 2 or 3 rays subequal, inserted toward aristal base; compound eye bare of microsetulae or the latter very sparse. Thorax: Usually with a gray to silvery stripe on thorax from postpronotum through ventral portion of notopleuron; anterior supra-alar seta lacking; posterior notopleural seta inserted at distinctly elevated position, especially as compared to anterior seta; anepisternum usually two toned, dorsal portion concol- orous with mesonotum, ventral portion gray; anepisternum with 2 subequal setae inserted along posterior margin. Wing: venation of wing generally pale colored; vein R ^ elongate, section III much shorter than section II; apical section of vein M longer than section between crossveins r-m and dm-cu; alula wide, width subequal to that of costal cell. Abdomen: Male terminalia: Pregonite either lacking or fused indistinguishably with postgonite; subepandrial sclerite lack- ing; postgonite generally elongate and bearing few setulae, usually only 2 are conspicuous. Key to Genera of the Tribe Hecamedini 1. Oral opening large, gaping; only reclinate fronto-orbital seta present; anteroventral margin of face essentially flat, at same level with rest of oral margin; clypeus broad; katepistemal seta lacking Elephantinosoma Becker Oral opening narrow; usually both a reclinate and proclinate fronto-orbital seta present; anteroventral margin of face emarginate with narrow clypeus exposed within facial emargination; katepistemal seta usually present 2 2. Scutellum bearing 3 marginal setae; postgenal margin sharp; gena high, over xli eye height Hecamede Haliday Scutellum bearing 2 marginal setae; postgenal margin rounded; gena short, less than l/2 eye height 3 3. Palpus mostly yellow; prescutellar acrostichal setae greatly reduced or absent; 1 katepistemal seta; face with 1 large lateral seta Eremotrichoma Giordani Soika Palpus blackish; 1 pair of prescutellar acrostichal setae, well developed; 2 katepistemal setae, the 2nd seta smaller and inserted below larger seta; face with 2 or more large lateral setae 4 4. Color generally black; microtomentum sparse, subshiny to shiny Diphuia Cresson Color generally gray to brown; microtomentum dense, generally appearing dull (Genus Allotrichoma Becker) 5 5. Median facial carina above facial prominence distinct, high, acute; presutural seta lacking (except in A. slossonae); proboscis geniculate, label la lanceolate, elongate, nearly equal to length of mediproboscis; face carinate above medial prominence Subgenus Pseudohecamede Hendel Median facial carina above facial prominence shallow; presutural seta present; labella broad, fleshy, shorter than mediproboscis 6 6. Clypeus microtomentose, usually gray; 5th tergum of male long and tubular, cerci of male terminalia elongate and with irregular, species-specific shape Subgenus Allotrichoma Becker Clypeus bare, black; 5th tergum of male short, l/2 length of 4th tergum; cerci normally developed, oval to lenticular "The atrUabrae Group" [see Mathis (1991) for information regarding the recognition and naming of this species group] SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Genus Hecamede Haliday Hecamede Haliday in Curtis, 1837:281 [published in synonymy; first made available by use of Haliday, 1839:221, 224; type species: Notiphila albicans Meigen, by monotypy].?Loew, 1860:12-13 [review].?Schiner, 1863:245 [description].?Becker, 1896:119-121 [discussion, description]; 1926:18- 19 [description, revision of palearctic species].?Cresson, 1925:228-229 [treated in subfamily Gymnopinae (= Gymnomyzini), notes on species of Old World].?Malloch, 1933:11 [discussion].?Harrison, 1959:219 [de- scription].?Wirth, 1965:737 [nearctic catalog]; 1968:5 [neotropical ca- talog].?Papp, 1975:25 [description].?Cogan and Wirth, 1977:325 [Orien- tal catalog].?Cogan, 1980:657 [Afrotropical catalog]; 1984:130-131 [palearctic catalog].?Canzoneri and Meneghini, 1983:231 [diagnosis].? Mathis, 1989a:641 [Australasian/Oceanian catalog]. Notiphila (Hecamede).?Walker, 1853:254 [review]. DESCRIPTION.?Small to medium-sized shore flies, length 1.30 to 3.50 mm, thinly to mostly densely microtomentose. Head: Wider than high; frons wider than high, at least posterior portion (from anterior ocellus posteriad) densely microtomentose, usually gray, anterior x/i frequently mostly red to mostly yellow and thinly microtomentose; ocellar setae lacking but usually setulae present; 1 to 3 intrafrontal setae, inserted laterad or in front of anterior ocellus; a reclinate fronto-orbital seta (sometimes with lateroclinate orientation); proclinate fronto-orbital seta variable; pseudopostocellar setae present, development variable; both inner and outer vertical setae present; ocelli arranged to form isosceles triangle, with distance between posterior pair slightly greater than between anterior ocellus and either posterior ocellus. Antenna exerted; pedicel with well-developed, proclinate, dorsal seta; aristal length subequal to combined length of pedicel and flagellomere 1, bearing 3 or 4 dorsal rays, with basal rays longer than apical 1-2. Face between antennae and antennal grooves shallowly but sharply carinate; antennal grooves moderately well devel- oped; face becoming more prominent below level of facial grooves, distinctly conically protrudent; facial setae in 1 vertical series near parafacial; facial setae arising from small, shiny tubercles; anteroventral facial margin shallowly emargi- nate; clypeus protrudent through ventral facial emargination. Eye irregularly round to subovate; apparently bare of mi- crosetulae (using a stereomicroscope). Gena high, at least l/i eye height, usually bearing 1 to several genal seta; posterior margin of gena acutely angulate. Palpus pale; labella broad, fleshy, shorter than mediproboscis. Thorax: Mesonotum usually darker than pleural region, usually tan to brown; chaetotaxy variable, setae usually dark colored, arranged in setal tracks as follows: rows of acrostichal setulae variable; dorsocentral track terminated posteriorly with larger seta; intra-alar setulae irregularly seriated; presutural seta variable; 1 postpronotal seta; 1 postalar seta; 3 scutellar setae well developed with sparse, scattered setulae on scutellar disc; 2 notopleural setae, insertion of posterior seta elevated above level of anterior seta; 2 anepisternal setae along posterior margin; katepisternal setae variable. Wing mostly hyaline to slightly milky white, veins mostly pale yellow; venation as follows: vein R2+3 extended beyond level of crossvein dm-cu, 3rd costal section about '/3 length of 2nd section; alular marginal setulae short, less than l/2 alular height. Legs: tibiae pale, mostly yellow to yellowish orange, less densely micro- tomentose than femora. Abdomen: Tergum 5 of male not generally visible from dorsal view, retracted within 4th; 4th tergum long, length subequal to combined length of 2nd and 3rd. Male terminalia as follows: symmetrical; epandrium and cerci well developed, separate; surstylus well developed, elongate, shape variable with species; gonite (= postgonite + fused pregonite ?) triangular to lunate; aedeagus simple, tubular, lightly scle- rotized, no distinct basiphallus or distiphallus, apex usually membranous; ejaculatory apodeme lacking; aedeagal apodeme evenly curved to angulate, better developed toward attachment with hypandrium; hypandrium well developed, a lightly sclerotized plate that is a shallowly concave receptacle for the male genitalia. DISTRIBUTION AND BIOGEOGRAPHIC CONSIDERATIONS.? Species of the genus Hecamede occur primarily on maritime coasts throughout most temperate and tropical regions of the world, sometimes in great abundance. A major departure from this generalization is the apparent absence of any species along the Pacific coast of the Western Hemisphere. Even that coastline may soon be invaded (see "Natural History" section of//, brasiliensis). Although Hecamede was proposed early in the nomenclatu- ral history of shore flies and its known species are geographi- cally widespread, the genus has never included many species. Moreover, most species now included in the genus have widespread distributions, typical of many coast-inhabiting Ephydridae. One species occurs on both sides of the Atlantic Ocean and another throughout the Indian and much of the Pacific oceans. Even those species that are more localized are relatively widespread, such as the coasts of Australia or those of Madagascar and the southeastern coast of Africa, and except for H. socotra and possibly H. africana, none is known to be a narrow endemic. The former is known only from the island of Socotra, and the latter is described from specimens collected recently in East Africa, and although presently known from a single locality, sampling for this species along the east coast of Africa is woefully incomplete. Collecting was attempted at only two or three sites in Kenya. NATURAL HISTORY.?All instars occur primarily on and/or in highly organic microhabitats that are associated with beaches and other maritime habitats. Ardo (1957:184) suggested that H. albicans is confined "wholly to the marine shore dune ecosystem." Apparently the larvae and adults are general scavengers, perhaps secondarily on bacteria that are associated with concentrated organic debris. Haliday (1839:224) appropri- ately described this medium as "marine rejectamenta." The debris frequently accumulates in the strand zone, where it often decays, providing a good habitat and food source for many beach and shore flies. Adults also feed on pollen and nectar NUMBER 541 from various flowering plants that occur near or on beaches. The number of generations per year is apparently determined by climate and is dependent upon availability of adequate food and habitat. Ardo (1957) suggested two generations for H. albicans in northern Europe, but several generations are possible in Connecticut (Steinly, 1992) and the tropics. What little is known about the immature stages of Hecamede is reported in the following papers (further details are cited in the treatment of the appropriate species): Bohart and Gressitt (1951; description and discussion of the adult and figure of the puparium of H. granifera (as H. persimilis) from Guam, the Solomons, and Guadalcanal), Tenorio (1980; reared and described the adults, third-instar larvae, and puparia of H. granifera (as H. persimilis), which was found on all major islands of the Hawaiian Islands), Norrbom (1983; description and illustrations of the third-instar larva and the puparium of//. albicans; discussion of feeding habits as a scavenger on dead horseshoe crabs, Limulus polyphemus L.), and Steinly (1992; description and illustrations of the third-instar larva and puparium of H. albicans; discussion of feeding habits as a scavenger on decaying horseshoe crabs, putrefying blue mussels (Mytilus edulis L.), and common spider crabs (Libinia emarginata Leach)). In addition to the papers just noted, A. Valdenberg (1985) completed a Master's thesis on the macrofauna of the Mediterranean beach at Ma'agan Michael, Israel, which includes information and illustrations on the natural history and immature stages of//, albicans. DISCUSSION.?Hecamede is easily distinguished from other shore-fly genera of the tribe Hecamedini by the following synapomorphic characters: gena high, usually over one-half height of eye and in some species nearly equaling it; face conically protrudent, frequently pointed and with point bare of microtomentum; postgenal margin sharp; acrostichal setae arranged in four to six irregular rows; scutellum with 3, rarely 4, pairs of marginal setae, these not arising from tubercles; scutellar disc with numerous, scattered setulae; katepisternum with 1 large seta and numerous smaller setae anteriad of larger seta; fore femur slightly to conspicuously swollen, especially when compared with femora of middle and hind legs; tibiae pale, although sometimes invested with white to gray micro- tomentum; wing membrane lacteous; and wing veins pale colored. Key to Subgenera of Hecamede Facial prominence bare at apex, shiny; proclinate fronto-orbital seta present, well developed; intrafrontal seta present. Acrostichal setae in about 6 irregular rows; katepisternal seta well developed Hecamede Haliday Facial prominence microtomentose; proclinate fronto-orbital seta either weakly developed or lacking; intrafrontal setae lacking. Acrostichal setae in 4 more or less regular rows; katepisternal seta lacking Soikia Canzoneri and Meneghini Subgenus Hecamede Haliday Hecamede Haliday, 1837:281 [see generic synonymy]. Hemicyclops de Meijere, 1913:66 [type species: Hemicyclops planifrons de Meijere, 1913, by monotypy (preoccupied, Boeck, 1873, Crustacea)].? Cogan and Wirth, 1977:325 [Oriental catalog].?Mathis, 1989a:641 [synon- ymy with Hecamede]. DESCRIPTION.?Small to medium-sized shore flies, length 1.15 to 3.10 mm; setae generally well developed and numerous. Head: Area surrounding ocellar triangle and fronto-orbits densely microtomentose, anterior portion of frons more thinly microtomentose, frequently reddish; proclinate fronto-orbital seta 1 or 2; intrafrontal setae 1, inserted about midway between anterior ocellus and anterior margin of frons, insertions more approximate than ocellar setae. Antenna usually pale, yellow to yellowish orange, flagellomere 1 sometimes mostly black (//. planifrons) or slightly blackish anterodorsally near insertion of arista. Conical facial prominence bare at apex, shiny, size variable; clypeus microtomentose. Thorax: Mesonotum either mostly light brown to faintly golden or bronzish or mostly gray; pleural areas mostly gray but sometimes with a wide, faintly gold stripe through dorsum of anepisternum; chaetotaxy generally well developed; acrosti- chal setae arranged in about 6 irregular rows; prescutellar acrostichal setae usually longer than other acrostichal setae and more widely set apart; presutural and prescutellar acrostichal setae usually evident but variable; katepisternal seta well developed. Abdomen: Male terminalia as follows: 5th tergum nor- mally retracted within 4th, anterior 3A to 7/s more lightly sclerotized than posterior portion, and covered with numerous, evenly scattered spicules, from dorsal view with anterior margin broadly bifid, posterior 'A to Vs a sclerotized band bearing 8 to 11 setae along posterolateral margin; epandrium a dorsolateral band, gradually enlarged posteriorly; surstylus long and usually narrow with apex slightly expanded and bearing setulae (shape, however, varies depending on species); cercus long and narrow, length subequal to or greater than that of surstylus, apex bearing a few long setulae; gonites with basal extension that connects over aedeagus, forming a gonal arch, usually pointed ventrally; aedeagus cylindrical, becoming smaller apically, curved basally, thereafter straight SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY DISTRIBUTION.?Of the two subgenera of Hecamede, this is by far the most widespread, with species occurring along temperate and tropical beaches along the Indian, much of the Pacific, and both sides of the Atlantic oceans. Only the western coast of the Western Hemisphere is apparently uninhabited. DISCUSSION.?Hecamede, sensu stricto, is apparently the relatively more derived subgenus and is characterized by several synapomorphies (see description above and "Phylo- genetic Consideration" p. 39) that confirm its monophyly. It also has the most species (9), and these, for the most part, closely resemble each other and are fairly easily clustered into two monophyletic lineages, the albicans and planifrons species groups. When de Meijere (1913:66) proposed the genus Hemicy- clops he was unaware that this name was preoccupied (Boeck, 1873; Crustacea) nor did he appreciate the interspecific variation within Hecamede, the genus with which Hemicyclops was compared for diagnostic purposes. This study, which included an examination of the features that de Meijere used to characterize Hemicyclops, has revealed that the characters he used reflect interspecific variation that is encompassed within the concept of Hecamede sensu stricto. Thus, I am treating Hemicyclops as a junior, preoccupied synonym of Hecamede that will not need a replacement, genus-group name. Key to Species of the Subgenus Hecamede 1. Proclinate fronto-orbital setae 2; gena with 1 black seta near middle 2 Proclinate fronto-orbital seta 1; gena either with a few to several black setae, scattered, or bare of setae 3 2. Clypeus dark colored; only apical tarsomere dark colored; posteriorproclinate fronto-orbital seta inserted distinctly anteriad of reclinate seta (distance between subequal to width of median ocellus) H. brasiliensis Cresson Clypeus with anterior surface mostly yellow; apical 2 tarsomeres dark colored; posterior proclinate fronto-orbital seta aligned essentially with reclinate seta. . . H. africana, new species 3. Fore femur bearing a distinct row of short, slightly stout setae along anteroventral margin 4 Fore femur lacking a distinct row of setae along anteroventral margin 5 4. Gena conspicuously shorter than eye height and bearing scattered setae; 1 pair of large, intrafrontal setae inserted in front of ocellar setae; flagellomere 1 mostly yellow, at most faintly black dorsally, pedicel mostly yellowish orange, sometimes mostly black dorsally H. granifera (Thomson) Gena almost as high as eye height and lacking setae (a few pale, fine setulae are present, however); intrafrontal seta either lacking or greatly reduced; flagellomere 1 blackish orange laterally, more orangish medially, pedicel with dorsal surface blackish, otherwise reddish orange (The planifrons Group) H. planifrons de Meijere 5. Fore basitarsus with 2 or 3 black setae inserted toward base of posteroventral surface 6 Fore basitarsus lacking black setae at base of posteroventral surface 7 6. Tibiae, especially the fore tibia, mostly grayish, only apices of fore tibia mostly yellow (Australia) H. australis, new species Tibiae mostly yellow, similar to basal tarsomeres (east and west coasts of the North Atlantic) H. albicans (Meigen) 7. Mesonotum darker, golden brown to brown; 5th tergum of /////////S// 1 I I I I i i 1 r Y / ^ / / / / / / ) J Y / / / / / } ^ / / / / ^ ^ \ ~ \ fnvxyAM FIGURE 64.?Hypothetical phylogeny for species of Hecamede. Numbers correspond with those used in the text. 15. Anepisternal setulae. 0 = present, especially on dorsal '/3; 1 = reduced and/or lacking. 16. Katepistemal seta. 0 = well developed; 1 = absent. 17. Adornment of anteroventral surface of fore femur. 0 = unadorned, lacking a distinct row of setae along anteroven- tral margin; 1 = bearing a distinct row of short, slightly stout setae along anteroventral margin. 18. Adornment of posteroventral surface of fore femur. 0 = 42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Species ancestor nuda (Africa) nuda (Oman) tomentosa bocki planifrons africana brasiliensis granifera maritima socotra inermis albicans australis TABLE 1.?Matrix Head (1-10) 1000000100 2101000202 2101000202 2201000202 2101000201 1102011011 0211111001 0211111001 1111111101 1111111101 1111111101 1111111101 1111111101 1111111101 of Character Data. Thorax (11-21) 00000011000 01101112031 01101112031 01101112001 01101110000 l??10001010 10010011000 10010011000 10010001000 10010011000 10010011000 10010011000 10010011100 10010010120 Abdomen (22-23) 00 02 02 03 05 04 01 01 11 11 11 11 11 11 bearing 5 to 7 setae on apical '/2-2/3; 1 = lacking setae along this surface; 2 = setae along this surface greatly enlarged. This character is coded to reflect a transforma- tion series where both derived states arose independently from the ancestral state; thus the coding for this character is the same that was done for character 1. 19. Fore basitarsus. 0 = lacking black setae at base of posteroventral surface; 1 = bearing 2 or 3 black setae inserted toward base of posteroventral surface. 20. Tibial coloration. 0 = mostly yellow, similar to basal tarsomeres, at most with thin investment of mostly white microtomentum; 1 = fore tibia reddish orange, usually invested with gray to dark gray microtomentum; 2 = especially the fore tibia, mostly grayish, only apices of fore tibia mostly yellow; 3 = fore tibia dark brown medially, mostly yellow at apices. This character was made nonadditive, as polarization beyond that indicated by the outgroup could not be determined. 21. Setae at base of hind basitarsus. 0 = mostly yellow; 1 = 2 to 4 black setae inserted near base on anteroventral surface. ABDOMEN 22. Length of cerci. 0 = subequal to height of epandrium; 1 = much longer than epandrium. 23. Shape of surstylus in lateral view. 0 = only slightly longer than wide; 1 = long and moderately narrow, posterior margin straight, anterior margin sinuous, narrow at middle, then slightly to moderately enlarged apically, length as long as epandrial height in lateral view; 2 = long and very narrow, curved anteriorly, pointed apically; 3 = long and narrow, curved anteriorly, rounded apically; 4 = broadly expanded and rounded apically; 5 = long, moderately wide, parallel-sided. This character was made nonadditive, as polarization beyond that indicated by the outgroup could not be determined. The matrix resulting from these data is shown above. From this matrix, using the implicit enumeration procedure of Hennig86, a single, most parsimonious tree resulted (Figure 64) that has a length of 36 steps, an overall consistency index of 0.94, and a retention index of 0.96. The analysis of each character is given in Table 2. The two basal lineages or sister groups, which are here recognized as subgenera, are the first division of the cladogram within the genus Hecamede. One lineage of this sister-group relationship, the subgenus Soikia, comprises but three species; the other, the subgenus Hecamede, includes the remaining species of the genus. The latter assemblage, Hecamede sensu stricto, is further divided into the two clades that are denoted here as the planifrons and albicans species groups. The planifrons group has only two species, both from the Oceanian Region, and the albicans group has eight species that occur throughout much of the Old World and along portions of the eastern coast of the New World. With the exception of the albicans species group, the cladogram is well resolved, and corroboration of the basal relationships is substantiated by numerous synapomorphies. The degree of resolution and abundance of evidence gives one confidence that the relationships at this level on the cladogram are reasonably accurate. Certainly the evidential requirements for the lineages giving rise to the two basal divisions are well met, as are those for the basal lineages of the two species groups within Hecamede, sensu stricto. The largely unresolved condition among species in the albicans group is problematic, however. Indeed, the species comprising this group are very similar externally, and except for H. africana and H. brasiliensis, which comprise a monophyletic lineage that is the sister group to the remainder of the albicans species group, no additional synapomorphies were discovered that would further resolve the relationships among these species. Although two species, H. albicans and H. australis, are linked as sister NUMBER 541 43 species in the analysis and cladogram, the evidence for this relationship is weak (the occurrence of the black seta on the ventral surface of the hind basitarsus is apparently quite variable), indicating a questionable sister-group relationship. The Old World (including Oceania), with 12 species, has a comparatively diverse fauna of Hecamede than does the New World, which has but two species. Moreover, the latter two species, H. albicans and H. brasiliensis, also occur on the Old World. The distributions of these two species and the sister-group relationship between H. brasiliensis and the East African endemic H. africana, lead one to suspect that both species are adventive to the New World. If this is the case, it would make the name of H. brasiliensis somewhat of an anomaly. Although occurring in Brazil, this species may have dispersed there from Africa, where it occurs along with its sister species, H. africana. Recent evidence suggests that H. brasiliensis may be susceptible to being transported and introduced, probably passively, as was noted previously in the "Natural History" section under that species (p. 11). It is not surprising, therefore, that it now ranges from the west coast of Africa to the tropical coasts of the Atlantic seaboard of the New World (Brazil to Belize) and finally to the islands of the Galapagos Archipelago. As both Eremotrichoma, the putative sister group of Hecamede, and all of the known species of Hecamede occur in the Old World, it is likely that the lineage that gave rise to these taxa originated there. Based on the present-day distributions of the included species, especially those that arose from basal lineages, a reasonable case could be advanced for an origin of the genus (at least its basal most lineage) to have occurred along the maritime coasts somewhere within the basin of the Indian Ocean. Thereafter, through a succession of speciation events associated with range expansion (dispersal), division (vicariance), and isolation, that lineage became subdivided into at least the 12 species that now comprise the genus. The two species that now occur in the New World, probably dispersed there from Old World origins through subsequent range expansion or introductions. In summary, the genus Hecamede is a monophyletic assemblage of 12 known species within the tribe Hecamedini. The included species are arrayed, according to the proposed phylogeny, into a classification of two subgenera, Soikia (three species) and Hecamede (nine species). In the latter subgenus there are two species groups, the planifrons group, with one species, and the albicans group, with eight species. The relationships between species of the albicans group are at present largely unresolved. The sublineages of the genus just elaborated are also monophyletic. The genus probably had its origin in the Old World, perhaps within the basin of the Indian Ocean where there is greatest species diversity and where taxa from all major lineages within the genus and the outgroup are represented. Literature Cited Adachi, M. 19S2. New Records and Name Changes in Hawaiian Ephydridae. Pro- ceedings of the Hawaiian Entomological Society, 14{3):353-354. Ardd, P. 1957. Studies in the Marine Shore Dune Ecosystem with Special Reference to the Dipterous Fauna. Opuscula Entomologica, supplement, 14: 1-25S, 82 figures, 9 tables. Becker, Th. 18%. Dipterologische Studien IV: Ephydridae. Berliner Entomologische Zeitschrifi. 41(2):91-276. 1903. Aegyptische Dipteren. 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Diptera Scandinaviae; Disposita et Descripta, 12:4547-4942. Lund. 1860. Diptera Scandinaviae; Disposita et Descripta, 14:6191-6609. Lund. REQUIREMENTS FOR SMITHSONIAN SERIES PUBLICATION Manuscripts intended for series publication receive substantive review (conducted by their originating Smithsonian museums or offices) and are submitted to the Smithsonian Institution Press with Form SI-36, which must show the approval of the appropriate authority designated by the sponsoring organizational unit. Requests for special treatment?use of color, foldouts, case-bound covers, etc.?require, on the same form, the added approval of the sponsoring authority. Review of manuscripts and art by the Press for requirements of series format and style, completeness and clarity of copy, and arrangement of all material, as outlined below, will govern, within the judgment of the Press, acceptance or rejection of manuscripts and art. 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