A Phylogenetic Study of the Tribe Dryxini Zatwarnicki (Diptera: )

WAYNE N.MATHIS and TADEUSZ ZATWARNK

W9\ I

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A Phylogenetic Study of the Tribe Dryxini Zatwarnicki (Diptera: Ephydridae)

Wayne N. Mathis and Tadeusz Zatwarnicki

Smithsonian Institution Press Washington, D.C. 2002 ABSTRACT Mathis, Wayne N., and Tadeusz Zatwarnicki. A Phylogenetic Study of the Tribe Dryxini Zat- warnicki (Diptera: Ephydridae). Smithsonian Contributions to Zoology, number 617, 101 pages, 154 figures, 2 tables, 2002.—The shore- tribe Dryxini is revised, including a cladistic analysis at the generic level, and now includes eight genera. Two of the genera, Omyxa and Papuama, are new, as are their respective type species. Of the remaining six genera, the spe- cies are revised for Dryxo Robineau-Desvoidy, Corythophora Loew, Oedenops Becker, and the subgenus Phaiosterna Cresson of the genus Loew. In addition, the species of Afro- limna Cogan, Oedenopiforma Cogan, and the limbata group (Paralimna) are reviewed. This study revealed the following synonyms: two genus-group names: Karema Cresson (1929) =Corythophora Loew (1862), and Cyphops Jaennicke (1867) =Dryxo Robineau-Des- voidy (1830); and seven species-group names: Karema loewella Cresson (1929) =Corytho- phora longipes Loew (1862), Cyphops fasciatus Jaennicke (1867) and Dryxo spreta Osten Sacken (1882) =Dryxo lispoidea Robineau-Desvoidy (1830), Paralimna ligabuei Canzoneri (1987) ^Paralimna madecassa Giordani Soika (1956), Oedenops aurantiacus Giordani Soika (1956) and Oedenops flavitarsis Miyagi (1977) =Oedenops isis Becker (1903), and Paral- imna {Phaiosterna) vidua Giordani Soika (1956a) =Ephydra bicolor Macquart (1851). Six new species are described in four genera (type locality in parentheses): Dryxo brahma (Sri Lanka. Colombo: Negombo), D. freidbergi (Cameroon. Kribi (beach), Rt. N7), D. india (India. Nedungadu), Omyxa scuta (Iran. 40 km SE Minab), Paralimna (Phaiosterna) longiseta (Dominican Republic. Azua: near Pueblo Viejo (18°24.8'N, 70°44.7'W)), and Papuama ismayi (Papua New Guinea. Central Province: Daramouka Village). The cladistic analysis was based on 45 morphological characters and resulted in nine most parsimonious cladograms of 55 steps with consistency and retention indices of 0.83 and 0.83, respectively. The tribe is divided into four basal sublineages in the strict consensus cladogram. The first sublineage comprises a single genus (number of species indicated in parentheses), Afrolimna (2), which is Afrotropical in distribution. The second sublineage likewise includes a single genus, Paralimna (>85), including Phaiosterna as a subgenus. Paralimna currently has greater species diversity than the rest of the tribe combined; it is pantropical, with numerous species ranging into subtropical regions. The third sublineage comprises three genera: Dryxo (9), Corythophora (2), and Omyxa (1), with Corythophora as the sister group to Dryxo and Omyxa. Genera of this sublineage occur only in the Old World, with greatest species diversity in Africa. The three genera of the fourth sublineage are Papuama (2), Oedenops (3), and Oede- nopiforma (3). In this sublineage, Papuama is the sister group to Oedenops and Oedenopi- forma. Oedenops is also pantropical and subtropical in distribution, but Papuama occurs in the Australasian/Oceanian and Oriental regions, and Oedenopiforma occurs in the Old World, pri- marily Africa and Australia.

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Library of Congress Cataloging-in-Publication Data Mathis, Wayne N. A phylogenetic study of the tribe Dryxini Zatwamicki (Diptera:Ephydridae) / Wayne N. Mathis and Tadeusz Zatwarnicki. p. cm. - (Smithsonian contributions to zoology ; no. 617) Includes bibliographic references (p. 99). 1. Ephydridae. I. Zatwarnicki, Tadeusz. II. Title. III. Series.

QL1.S54 no. 617 [QL537.E7] 590 s-dc2l [595.774] 2001049798

© 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 1 Acknowledgments 2 Systematics 3 Tribe DRYXINI Zatwarnicki 3 Key to Genera of DRYXINI 4 Genus Afrolimna Cogan, new status 5 Key to Species of Afrolimna Cogan 6 1. Afrolimna carolinika (Cogan), new combination 7 2. Afrolimna keiseri (Cogan), new combination 8 Genus Corythophora Loew 10 Key to Species of Corythophora Loew 11 3. Corythophora flavipes (Cogan), new combination 11 4. Corythophora longipes Loew 13 Genus Dryxo Robineau-Desvoidy 16 Key to Species of Dryxo Robineau-Desvoidy 19 5. Dryxo brahma, new species 20 6. Dryxo digna Osten Sacken 22 7. Dryxo freidbergi, new species 24 8. Dryxo India, new species 26 9. Dryxo lispoidea Robineau-Desvoidy 27 10. Dryxo margaretae Cogan 30 11. Dryxo nudicorpus Miyagi 32 12. Dryxo ornata (Macquart) 34 13. Dryxo woodi Cresson 38 Omyxa, new genus 40 14. Omyxa scuta, new species 41 Genus Oedenopiforma Cogan 42 Key to Species of Oedenopiforma Cogan 44 15. Oedenopiforma argentea (Cogan) 45 16. Oedenopiforma madecassa (Giordani Soika) 47 17. Oedenopiforma uniseta (Malloch), new combination 49 Genus Oedenops Becker 51 Key to Species of Oedenops Becker 52 18. Oedenops afrus Wirth 53 19. Oedenops isis Becker 54 20. Oedenops nudus (Coquillett) 57 Genus Paralimna Loew 59 Key to Subgenera of Paralimna Loew 60 Subgenus Paralimna Loew 61 The limbata Group 63 Key to Species of the limbata Group 64 21. Paralimna {Paralimna) limbata Loew 64 22. Paralimna {Paralimna) reticulata Cogan 65 Subgenus Phaiosterna Cresson 67 Key to Species of Phaiosterna Cresson 68 23. Paralimna {Phaiosterna) bicolor (Macquart) 69

in IV SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

24. Paralimna {Phaiosterna) decipiens Loew 72 25. Paralimna {Phaiosterna) fusca Bock 76 26. Paralimna {Phaiosterna) lineata de Meijere 78 27. Paralimna {Phaiosterna) longiseta, new species 81 28. Paralimna {Phaiosterna) obscura Williston 83 Papuama, new genus 88 Key to Species of Papuama 89 29. Papuama calva (Bock), new combination 90 30. Papuama ismayi, new species 90 Phylogenetic Considerations 94 Characters Used in the Phylogenetic Analysis 95 General 95 Head 95 Thorax 95 Abdomen 96 Analysis, Results, and Conclusions 97 Literature Cited 99 FIGURE 1.—Frontispiece of Dryxo brahma, new species (Kadaimparu. Sri Lanka, $).

A Phylogenetic Study of the Tribe Dryxini Zatwarnicki (Diptera: Ephydridae)

Wayne N. Mathis and Tadeusz Zatwarnicki

Introduction The origin of a taxon is not always reliably indicated by its present diversity and/or abundance, and classifications that are Like other tribes of the shore-fly family Ephydridae, Dryxini not based on phylogenetic relationships can likewise be mis- Zatwarnicki occurs in both the Old and New Worlds, and a few leading. Thus, conducting research on the phylogeny of Dryx- of the included genera, such as Paralimna Loew and Oedenops ini has applications that transcend simply knowing the rela- Becker, are equally widespread. Most genera of Dryxini, how- tionships among the component lineages or providing the basis ever, are found only in the Old World, and the Afrotropical Re- for a classification that facilitates the storage and retrieval of gion is especially rich in genera and species. Conversely, no information. genus of this tribe occurs exclusively in the New World, even Dryxini, which Zatwarnicki (1992) first proposed, is the at the subgeneric level. most recent tribe in the nomenclatural history of shore . It Despite being relatively widespread, comparatively diverse, comprises all the taxa formerly included in the tribe Notiphilini and often abundant, there are no studies on the phylogenetic re- Bigot, with the exception of the genus Fallen. In the lationships within the tribe, and most of the included genera most recent world catalog of shore flies (Mathis and Zatwar- nicki, 1995), five genera and 101 species were included in have never been revised. To remedy in part this gap in our un- Dryxini, with the vast majority, more than 85 species, being in derstanding, we initiated this phylogenetic study of Dryxini. the genus Paralimna. Although Dryxini is moderately diverse, Our objectives are to provide a phylogeny and classification at we know that Paralimna alone is far richer in species than the generic level and to present species revisions for Dryxo present numbers would indicate. Here we describe six new spe- Robineau-Desvoidy, Corythophora Loew, Oedenops, and the cies, three in Dryxo, one in Omyxa, one in Paralimna subgenus Phaiosterna Cresson of the genus Paralimna. The {Phaiosterna), and one in Papuama. Two of these genera, species of Afrolimna Cogan and Oedenopiforma Cogan, and of Omyxa and Papuama, are also newly described. the limbata group {Paralimna), are also reviewed. Genera that are revised at the species level are provided with a "Historical Review" as a subsection under the generic treatment. Wayne N. Mathis, Department of Entomology, National Museum of METHODS.—The descriptive terminology, with the excep- Natural History, Smithsonian Institution, Washington, DC 20560- tions noted in Mathis (1986), Mathis and Zatwarnicki (1990a), 0169, USA. Tadeusz Zatwarnicki, Department of Zoology, University and below, follows that published in the Manual of Nearctic of Agriculture, ul. Cybulskiego 20, 50-205 Wroclaw, Poland. Diptera (MeAlpine, 1981). Although many specimens of Review Chairperson: John M. Burns, Department of Entomology, Dryxini are among the largest in the Ephydridae, study and il- National Museum of Natural History, Smithsonian Institution, Wash- lustration of the male terminalia required use of a compound ington, DC. 20560-0169, USA. microscope. For most structures of the male terminalia, we Reviewers: Philip J. Clausen, Department of Entomology, University have followed the terminology used by other workers in Ephy- of Minnesota, St. Paul. Minnesota 55018, USA. Stephen D. Gaimari, dridae (see references in Mathis, 1986, and Mathis and Zatwar- Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0169, USA. Allen L. nicki, 1990a, 1990b), for example, the terminology for sursty- Norrbom, Systematic Entomology Laboratory/USD A, c/o Smithsonian lus, which in Dryxini is divided into a presurstylus (surstylus) Institution, Washington, DC 20560-0168, USA. and a postsurstylus (clasper). Zatwarnicki (1996) has suggested SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY that these structures correspond with the pregonostylus and the AM Australian Museum, Sydney, Australia (Daniel J. Bickel, David K. postgonostylus and that the subepandrial plate is the same as McAlpine; including collections formerly housed at the School of Public Health and Tropical Medicine, University of Sydney) the medandrium. The terminology for structures of the male AMNH American Museum of Natural History, New York, New York, USA terminalia is provided directly on Figures 6-8 and is not re- (David A. Grimaldi) peated for comparable illustrations of other species. ANIC Australian National Collection, Canberra, Australia (David Alternative spellings for some localities are cited in paren- K. Yeates) thesis, especially for locality names that were transliterated into ANSP Academy of Natural Sciences, Philadelphia, Pennsylvania, USA (Jon K. Gelhaus and Donald F. Azuma) English. States of the United States in the "Distributions" sec- BAR personal collection of M. Bartak (University of Agriculture, Praha, tions are abbreviated as follows: AK=Arkansas, AZ=Arizona, Czech Republic) CA=California, FL=Florida, GA=Georgia, IA=Iowa, BMNH The Natural History Museum (formerly the British Museum (Natu- IL=Illinois, LA=, MO=Missouri, MS=Mississippi, ral History)), London, England (Brian Pitkin and John E. Chainey) BPBM Bernice P. Bishop Museum, Honolulu, Hawaii, USA (Neal L. NJ=New Jersey, OK=Oklahoma, TN=Tennessee, TX=Texas, Evenhuis) UT=Utah. CAS California Academy of Sciences, San Francisco, California, USA The species descriptions are composite and are not based (Paul H. Arnaud, Jr.) solely on the holotypes. Four head and two venational ratios CNC Canadian National Collection, Ottawa, (James E. O'Hara used in the descriptions are defined below (all ratios are aver- and Bruce Cooper) CU Cornell University, Ithaca, New York, USA (James K. Liebherr) ages of three specimens (the largest, the smallest, and one DEI Deutsches Entomologisches Institut, Eberswalde, Germany (Frank other)). Menzel and Joachim Ziegler) HNHM Hungarian Natural History Museum, Budapest, Hungary (Laszlo 1. Frons width-to-length ratio is the frons width divided by the Papp) frons length. Length is measured from the anterior margin HUS Hokkaido University, Sapporo, Hokkaido, Japan (Masaki Suwa of the frons to the posterior margin of the posterior ocelli; and lshiro Miyagi) width is measured at the level of the anterior ocellus. INHS Illinois Natural History Survey, Champaign, Illinois, USA (Donald W. Webb) 2. Face width-to-height ratio is the narrowest width between IRSN Institut Royal de Sciences Naturelles, Bruxelles, Belgium (Patrick the eyes, divided by the facial height. Grootaert) 3. Gena-to-eye ratio is the genal height measured at the maxi- KU Snow Entomological Museum, University of Kansas, Lawrence, mum eye height, divided by the eye height. Kansas, USA (Steve Ashe and George W. Byers) 4. Eye width-to-height ratio is the eye width divided by the MCV Museo Civico di Storia Naturale de Venezia, Venice, Italy (Enrico Ratti and Leone Rampini) eye height, where both measurements are the longest dis- MCZ Museum of Comparative Zoology, Harvard University, Cambridge, tances taken with the eye oriented laterally. Massachusetts, USA (Philip D. Perkins) 5. Costal-vein ratio is the straight-line distance between the MNHN Museum National d'Histoire Naturelle, Paris, France (LoTc Matile) MRAC Musee Royal de rAfrique Centrale (Koninklijk Museum voor apices of veins R2+3 and R4+5 divided by the distance be- Midden-Afrika), Tervuren, Belgium (Eliane De Coninck) tween the apices of veins Rj and R2+3- NMNH National Museum of Natural History, Smithsonian Institution, 6. M-vein ratio is the straight-line distance along vein M be- Washington, D.C., USA tween crossvein dm-cu and r-m, divided by the distance api- NMSA Natal Museum, Pietermartzburg, South Africa (David A. Barra- cad of crossvein dm-cu. clough) NMSAC National Museum, Prague, Czech Republic (Jan Jezek) The phylogenetic analysis was performed with the assis- NMW Naturhistorisches Museum, Wien, Austria (Ruth Contreras-Licht- tance of Hennig86°, a computerized algorithm that produces enberg) cladograms by parsimony. Character data were polarized pri- NNIC Namibian National Insect Collection, State Museum, Windhoek, Namibia (Ashley H. Kirk-Spriggs) marily using outgroup procedures. We did not include autapo- NRS Naturhistoriska Riskmuseet, Stockholm, Sweden (Thomas Pape) morphies in the cladistic analysis (they were made inactive) NSF Naturmuseum Senckenberg, Frankfurt, Germany (W. Tobias) because that would have skewed the consistency and retention TZ personal collection of Tadeusz Zatwarnicki, Department of Zool- indices, but we listed them on the cladogram and included ogy, University of Agriculture, Wroclaw, Poland UMO University Museum, Oxford University, Oxford, England (John W. them as part of generic treatments and phylogenetic consider- Ismay) ations to document the monophyly of the lineages, particularly USNM collections of the former United States National Museum, now in at the generic level. the NMNH ACKNOWLEDGMENTS.—Although most specimens for this UZMC Zoologisk Museum, Copenhagen, Denmark (Verner Michelsen and study, including the primary types, are in the National Museum Rudolf Meier) ZIL Zoological Institute, Museum of Zoology and Entomology, Lund of Natural History, Smithsonian Institution, numerous others University, Lund, Sweden (Roy Danielsson) were borrowed, particularly type specimens of the species pre- ZMA Instituut voor Taxonomische Zoologie, Zoologisch Museum, Uni- viously described. To our colleagues and their institutions versiteit van Amsterdam, Amsterdam, Netherlands (Ben Brugge) listed below who loaned specimens, we express our sincere ZMHU Zoologisches Museum, Humboldt Universitat, Berlin, Germany (Marion Kotrba, Hubert Schumann, and Hella Wendt) thanks. Without their cooperation this study could not have ZMUM Zoological Museum, Moscow University, Moscow, Russia (Marina been completed. G. Krivosheina) NUMBER 617

Tadeusz Zatwarnicki prepared the distribution maps, Susann seta; posterior notopleural seta at same level as anterior seta; G. Braden and Walter R. Brown assisted with the scanning ventral anepisternal seta elongate, twice length of dorsal seta (a electron microscopy, Victor Krantz produced the final photo- synapomorphy for Notiphilini+Dryxini); costa elongate, ex- graphs, and Elaine R.S. Hodges and Young T. Sohn produced tended to vein M (costa short, extended only to vein R4+5 in the habitus illustrations. We also thank Jaroslaw Kama and Notiphilini; a synapomorphy for Notiphilini); midtibia with Miroslaw Solowski for the life-like habitus of Dryxo. For re- prominent, erect, extensor setae on dorsal surface (a synapo- viewing a draft of this paper, we thank Philip J. Clausen, James morphy for Notiphilini+Dryxini); abdominal tergites fasciate F. Edmiston, Stephen D. Gaimari, Allen L. Norrbom, and Terry (a synapomorphy for Notiphilini+Dryxini); male terminalia A. Wheeler. We are also grateful to David Challinor (former with surstylus divided into presurstylus (surstylus) and postsur- assistant secretary for research, Smithsonian Institution), Stan- stylus (clasper); presurstylus with apex angulate and bifurcate; wyn G. Shetler (former deputy director of the NMNH), Anna subepandrial plate reduced (a synapomorphy for Notiphilini+ K. Behrensmeyer (former associate director of the NMNH), Dryxini); pre- and postgonite reduced and/or lacking (structure and David Pawson (former associate director of the NMNH) remaining may represent fused and/or reduced pre- and post- for financial support to conduct field work and study primary gonite); and hypandrium connected basally with postsurstylus, types through grants from the Research Opportunity Fund. not with epandrium. Funding from a Short-term Visitors Grant provided T. Zatwar- DESCRIPTION.—Small to very large shore flies of the sub- nicki with assistance for a month-long visit at the Smithsonian family , body length 1.65-11.20 mm, general body Institution to conduct the research that in part resulted in this facies robust. paper. Field work in the Seychelles was facilitated through a Head: Fronto-orbital setae reclinate and/or proclinate; grant from Helmut Hollmann, benefactor for research on shore reclinate fronto-orbital seta 1, well developed, inserted me- flies. We gratefully acknowledge his generous assistance. diad of proclinate setulae; proclinate fronto-orbital setae mod- Travel to and from the Seychelles was largely provided by Brit- erately to greatly reduced, usually 2, inserted laterad of recli- ish Airways, and we are grateful to them for generously sup- nate seta or lacking; pseudopostocellar seta greatly reduced or porting this research. lacking, divergent to widely divergent if present; both inner Field work in the West Indies was funded largely by grants and outer vertical setae usually well developed, sometimes in- from the Biodiversity Program (Biological Surveys and Inven- ner vertical weakly developed; ocellar seta stronger than tories) of the NMNH (Lynne R. Parenti, former chair, George weakly developed pseudopostocellar seta. Pedicel lacking R. Zug, chair). Field work in Guyana was supported by the prominent, elongate setae; arista bearing numerous long, dor- Smithsonian Institution's Biological Diversity of the Guianas sally projected, hair-like branches. Face wide, transversely Program (publication number 40; Vicki A. Funk, Director; arched, usually projected anteriorly from antennal base; ven- Carol L. Kelloff, Coordinator). tral facial margin flat or very shallowly emarginate; facial se- This is contribution number 594 from the Caribbean Coral tae, if present, inserted near lateral margins, usually in vertical Reef Ecosystems Program, Smithsonian Institution, which is series somewhat parallel with parafacial setae, setae inclinate, partly supported by a grant from the Exxon Corporation. frequently reduced in size or lacking; clypeus short, wide, band-like. Eye round to slightly oval with slight to conspicu- System atics ous oblique orientation; appearing bare of interfacetal setulae. (We have discovered, using scanning electron microscopy, Tribe DRYXIM Zatwarnicki that most if not all shore flies have at least some interfacetal Dryxini Zatwarnicki, 1992:85 [type genus: Dryxo Robineau-Desvoidy, setulae.) Gena high; genal seta frequently reduced in size, in- 1830].—Mathis and Zatwarnicki, 1995:115-127 [world catalog]. serted near ventral margin; subcranial cavity moderately large DIAGNOSIS.—Dryxini is the sister group of Notiphilini, and and cavernous. the tribe is distinguished from Notiphilini and other shore-fly Thorax: Acrostichal setae generally reduced, sometimes tribes by the following combination of characters: ocellar seta lacking, in 2-8 regular or irregular rows if present; only pre- stronger than the weak pseudopostocellar seta; reclinate fronto- scutellar acrostichal setae well developed, insertion moder- orbital seta well developed, proclinate setae reduced or lacking; ately widely apart; dorsocentral setae usually 4 (1+3), ante- eye appearing bare; face wide, transversely arched, and gener- rior setae sometimes lost secondarily, leaving 1 posterior seta; ally projected anteriorly; facial setae in 1 or 2 serial rows run- posteriormost dorsocentral seta inserted laterad of alignment ning more or less parallel to parafacial; gena high (secondarily of dorsocentral track; presutural supra-alar seta usually short in some species); subcranial cavity moderately large and present (secondarily lost in Corythophora, Dryxo, and cavernous; dorsocentral setae 4 (1+3; secondarily reduced in Omyxa); postsutural supra-alar seta usually present, well de- some genera of Dryxini (Corythophora, Dryxo, and Omyxd) (3 veloped (secondarily lost in Dryxo freidbergi (described setae in Notiphilini, 1+2; a synapomorphy for Notiphilini); herein) and Papuama ismayi (described herein)); postalar seta prescutellar acrostichal setae present, well developed; postsu- 1, well developed; scutellum with 2 setae inserted along lat- tural supra-alar setae strong, longer than posterior notopleural eral margins; disc of scutellum with few to numerous setulae; 4 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

postpronotal seta 1 or 2, sometimes lost secondarily {Dryxo uama). Male terminalia: epandrium inverted U-shaped; presur- margaretae Cogan, D. digna Osten Sacken, D. lispoidea Rob- stylus lobate or L-shaped; postsurstylus setulose with apex bent ineau-Desvoidy, and Omyxa (described herein)), larger seta and bifurcate; subepandrial plate reduced; pregonite and post- posteroclinate, 2nd seta, if present, shorter, inclinate; no- gonite reduced and/or lacking (remaining structure, which may topleural setae usually 2 (secondarily reduced to 1 in Corytho- be a composite of pre- and postgonites, termed a gonite); hy- phora, Dryxo, and Omyxa), posterior seta at same level as an- pandrium frequently concave, connected basally with postsur- terior seta; anepisternum usually with 2 large setae along stylus, not epandrium. posterior margin, ventral seta usually elongate, sometimes DISCUSSION.—In addition to the synapomorphies listed ear- twice length of dorsal seta, extended posteriorly to level of Her (see tribal diagnosis) and those in the "Phylogenetic Con- halter (dorsal seta secondarily reduced or lacking in Corytho- siderations" section (beginning on page 94) that indicate a sis- phora and in some species of Dryxo). Wing generally hyaline ter-group relationship for Dryxini and Notiphilini, Dryxini is to faintly infuscate, patterned in some species of Paralimna; further characterized by the following autapomorphies that es- costal vein elongate, extended to vein M; vein R2+3 long, ex- tablish the monophyly of the tribe: (1) gena high (secondarily tended past midpoint of 2nd costal section. Midtibia with 1-4 short in some species of Paralimna); (2) face wide, trans- prominent, erect, extensor setae along dorsal surface; tarsal versely arched, and generally projected anteriorly; (3) male claws regularly developed, relatively short, shallowly curved; terminalia with surstylus divided into a presurstylus (surstylus) pulvillar pads present, normally developed. Halter generally and a postsurstylus (clasper); presurstylus with apex angulate whitish yellow. and bifurcate; (4) pregonite and postgonite reduced and/or Abdomen: Tergites usually with fasciate pattern (second- lacking; (5) hypandrium connected basally with postsurstylus, arily unicolorous in Oedenops, Oedenopiforma, and Pap- not with epandrium.

Key to Genera of DRYXINI 1. Notopleuron bearing 1 large seta [Figures 1, 19]; presutural supra-alar seta lacking [Figures 19, 47]; mid- and hindfemora moderately long to very long, subequal to abdomen length [Figure 1] 2 Notopleuron bearing 2 setae [Figure 5]; presutural supra-alar seta usually present [Fig- ures 81, 92, 118] (lacking in Papuama and in one species of Oedenops); mid- and hindfemora normally developed, much shorter than abdomen 4 2. Ocellar seta present, although short, inserted slightly in front of anterior ocellus [Fig- ures 17, 18]; reclinate fronto-orbital seta present [Figure 18]; anepisternum bearing 1 well-developed seta along posterior margin; vein Rj bare along dorsum; R stem vein lacking setulae; crossvein dm-cu normally developed, nearly straight, forming acute inner angle with vein M (southern Afrotropical) Corythophora Loew Ocellar seta lacking [Figures 45, 46]; reclinate fronto-orbital seta lacking [Figure 46]; anepisternum bearing 2 or 3 thin, long, hair-like setae along posterior margin; vein Rl bearing several setulae along dorsum; R stem vein basad of humeral crossvein bearing several pale, thin setulae on ventral surface; crossvein dm-cu moderately long to long, sinuous 3 3. Arista bearing 7-9 long dorsal hairs [Figures 54, 55]; katepisternum lacking row of slender setae along dorsal margin, and katepisternal seta reduced; crossvein dm-cu shallowly sinuous, generally forming angle with adjacent margin of wing; mid- and hindfemora normally developed, much shorter than length of abdomen (India, Iran, Oman) Omyxa, new genus Arista bearing 12 or more long dorsal hairs [Figures 45, 46]; katepisternum bearing row of slender setae near dorsal margin, and katepisternal seta usually well devel- oped (secondarily reduced or absent in some species); crossvein dm-cu sinuous, long, generally running parallel with adjacent margin of wing; mid- and hindfemora elongate, subequal to length of abdomen (Afrotropical, Australian, Oriental) Dryxo Robineau-Desvoidy 4. Katepisternal seta absent or very weakly developed 5 Katepisternal seta present, usually well developed (sometimes pale) 6 NUMBER 617

5. Arista bearing 3-5 hairs [Figure 80] (Afrotropical, Australian (Queensland), Nearctic (southern), Neotropical, Oriental, Palearctic (Egypt, Israel, Japan)) Oedenops Becker Arista bearing 8 or more hairs (Australasian, Oriental) Papuama, new genus 6. Long facial setae 2 or 3, length subequal to combined length of pedicel and 1st flagel- lomere, if 2 setae, these well separated, dorsal seta at about midheight, ventral seta closer to oral margin than to dorsal seta. R stem vein bearing 1-3 (usually 2) setulae on dorsum (Afrotropical, Australian, Oriental) Oedenopiforma Cogan Long facial seta 1 (if other long setae present, setae not as long or as separated as above, these usually arranged in a somewhat vertical series of short setulae). R stem vein lacking setulae 7 7. Forefemur lacking row of closely set, very short, somewhat blunt, tooth-like spines along anteroventral surface; anterior proclinate fronto-orbital seta larger than poste- rior seta; face, gena, anepisternum, anterior surface of tibiae, and basolateral surface of scutellum not silvery microtomentose as in combination below; forefemur lack- ing long setae or setae not as below (pantropical with occasional extensions into temperate zone) Paralimna Loew Forefemur bearing anteroventral row of very short, stout, tooth-like setae; proclinate fr- onto-orbital setae greatly reduced and subequal, setula-like; face, gena, anepister- num, anterior surface of tibiae, and basolateral surface of scutellum densely, silvery microtomentose; forefemur bearing 4 or 5 long setae along apical V2 of posteroven- tral surface, length approaching twice width of femur (Afrotropical) Afrolimna Cogan

Genus Afrolimna Cogan, new status flagellomere moderately broadly rounded; arista bearing 13 or 14 long hairs along length of dorsal surface. Face usually bear- Afrolimna Cogan, 1968:322 [as a subgenus of Paralimna; type species: Para- limna carolinika Cogan, 1968, original designation].—Cogan, 1980:662 ing 1 moderately well-developed facial seta and series of [Afrotropical catalog].—Mathis and Zatwarnicki, 1995:118 [world catalog]. smaller setae or setulae ventrad. Parafacial at anterior margin of eye narrow, width much less than length of 1st flagellomere. DIAGNOSIS.—This genus is distinguished from other genera Gena moderately high, height slightly greater than height of of Dryxini by the following combination of characters: gena 1 st flagellomere. high, height about V3 or more or eye height; 1st fiagellomere short, length at most 1.5 times width; face, gena, anepister- Thorax (Figure 5): Dorsocentral setae 4 (1 +3 or 2+2), num, anterior surface of tibiae, and basolateral surface of well developed, 2nd seta near or at transverse suture, posterior scutellum densely, silvery microtomentose; forefemur bearing pair displaced laterally; only prescutellar acrostichal seta well 4 or 5 long setae on apical V2 of posteroventral surface, their developed and well separated, other acrostichal setulae incon- length approaching twice width of femur; forefemur of both spicuous, in numerous irregular rows; presutural supra-alar sexes bearing row of short, tooth-like setulae apically along seta well developed, subequal in length to anterior dorsocen- anteroventral surface (very inconspicuous in A. keiseri tral seta; postpronotal seta well developed; notopleural setae 2, (Cogan)); male terminalia (Figures 6-8) with presurstylus an- posterior seta much shorter, about V2 length of anterior seta; gulate, elbowed in posterior view; aedeagal apodeme in lateral anepisternum bearing 2 large setae along posterior margin, view with prominent, distinctive, broad keel and long, nearly dorsal seta shorter, sometimes up to V2 length of ventral seta; parallel-sided medial projection; and hypandrium angulate in katepisternal seta well developed. Posterior margin of scutel- lateral view. lum truncate, nearly flat. R stem vein lacking setulae; cross- DESCRIPTION.—Medium-sized to moderately large shore vein dm-cu regularly developed, nearly straight, about 3 times flies, body length 3.40-4.30 mm. longer than apical section of vein CuA], and at distinct angle Head (Figures 3, 4): Frons shallowly arched anteroven- with posterior margin of wing. Forefemur bearing comb-like trally, not projected forward, sparsely setulose; ocelli in equi- row of very short, stout, spine-like setulae along apical V2 of lateral triangle; both inner and outer vertical setae well devel- anteroventral surface (very inconspicuous in A. keiseri); oped; reclinate fronto-orbital seta, ocellar seta, and para- foretibia of male lacking several long, slender setae at apex on vertical seta well developed; proclinate fronto-orbital setae 2, ventral surface; forebasitarsus lacking row of long, slender, anterior seta much better developed, subequal to posterior no- pale setulae inserted along anterior surface; midtibia with 3 topleural seta, posterior seta much reduced, setula-like. First dorsal extensor setae (subapically, subbasally, and near mid- SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY die); mid- and hindfemora normally developed, much shorter apex in lateral view (Figures 8, 11); lateral aedeagal process than abdomen. shorter than or subequal to aedeagus in lateral view, apex ei- Abdomen: Coloration variable with species, but lacking a ther tapered to acute point or broadly rounded; hypandrium distinctly fasciate pattern. Male terminalia: epandrium broadly angulate, moderately shallow. and uniformly U-shaped in posterior view (Figures 6, 9), lat- DISTRIBUTION (Figure 2).—Afrotropical: , Nige- eral arms slightly wider; cercus moderately broad to broad ria, Sudan, Zaire, Zimbabwe. ventrally, pointed dorsally; presurstylus angulate in posterior DISCUSSION.—Afrolimna was first described as a subgenus view (Figures 6,9), elbowed, medially oriented arm slightly to within Paralimna (Cogan, 1968, 1980; Mathis and Zatwar- much wider than vertical portion, medial surface truncate to nicki, 1995). Our cladistic analysis and determination that concave; postsurstylus in lateral view curved, slightly widened Afrolimna has a closer relationship with Corythophora, Dryxo, medially, thereafter narrowed, apex shallowly bifurcate; and Omyxa than with Paralimna alters its phylogenetic posi- aedeagal apodeme in lateral view hemispherical with short tion within Dryxini, and thus, we accord it generic status. dorsal and ventral arms, anteromedial margin of apodeme As implied by the generic name, the genus is known only bearing distinctive, long, nearly parallel-sided projection from the Afrotropical Region, where there are two described longer than width of keel, prominent keel irregularly rounded; species for which we provide the following key, diagnoses, and aedeagus wider basally, gradually narrowed to broad, angulate illustrations.

Key to Species of Afrolimna Cogan 1. Notopleuron dusted with silvery microtomentum; wing of male immaculate, hyaline 1. A. carolinika (Cogan) Notopleuron dark brown, concolorous with remainder of mesonotum; wing of male with dark spot at level of crossvein dm-cu in cells X\ and ^+3 2. A. keiseri (Cogan)

FIGURE 2.—Distribution map for Afrolimna (stippled). NUMBER 617

1. Afrolimna carolinika (Cogan), new combination keel and long, almost parallel-sided medial projection as long as width of keel; aedeagus horn-shaped in lateral view (Figure FIGURES 3-8 8), wide basally, tapering to posteriorly oriented, pointed apex; Paralimna (Afrolimna) carolinika Cogan, 1968:322; 1980:662 [Afrotropical lateral aedeagal process as long as aedeagus in lateral view, catalog].—Mathis and Zatwamicki, 1995:118 [world catalog]. broadly and unevenly rounded apically; and hypandrium angu- DIAGNOSIS.—This species is distinguished from Afrolimna late, moderately deep. keiseri by the characters indicated in the key and by the fol- TYPE MATERIAL.—The holotype male of Paralimna caro- lowing characters from the male terminalia (Figures 6-8): linika Cogan is labeled "Holo-type [round label with red mar- epandrium uniformly U-shaped in posterior view (Figure 6), gin]/Chipinda Pools Tsetse Fly Ops. S. Rhodesia [Zimbabwe] width even throughout; cercus greatly widened ventrally, Lower Lundi R. 22.X.1960 R. Goodier 1329 [all data except pointed anteromedially; presurstylus angulate in posterior "Tsetse Fly Op. S. Rhodesia" and "19" of the year date are view (Figure 6), elbowed, medially oriented arm much wider handwritten]/Pres by R.Goodier. B.M. 1961-72./Paralimna than vertical portion, medial surface concave with dorsome- carolinika sp.n det.B.H.Cogan 1966 [species name and "sp.n" dial angle projected, acutely pointed; postsurstylus in lateral handwritten]." The holotype is pinned directly, is in excellent view curved, slightly widened medially, thereafter narrowed, condition, and is deposited in BMNH. Two female paratypes apex shallowly bifurcate; aedeagal apodeme with prominent (BMNH) bear the same label data as the holotype. Other

FIGURES 3-5.—Afrolimna carolinika (Cogan): 3, head, anterior aspect; 4, same, lateral aspect; 5, mesonotum, dorsal aspect. Scale=0.5 mm. SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

epandrium

cercus T 0.1m1mm

presurstylus (pregonostylus)

aedeagal apodeme (phalloapodeme)

postsurstylus (postgonostylus)

FIGURES 6-8.—Male terminalia of Afrolimna carolinika (Cogan): 6, epandrium, cerci, and presurstyli, posterior aspect; 7, internal male terminalia, ventral aspect; 8, same, lateral aspect. paratypes are as follows: NIGERIA. Zungeru, Feb 1912, J.W.S. 2. Afrolimna keiseri (Cogan), new combination Macfie (29; BMNH). SUDAN. Yirol, 28 Feb 1954, E.T.M. Reid FIGURES 9-11 Paralimna (Afrolimna) keiseri Cogan, 1968:323; 1980:662 [Afrotropical cata- OTHER SPECIMENS EXAMINED.—Afrotropical: ZAIRE. Ka- log].—Mathis and Zarwarnicki, 1995:118 [world catalog]. songo, Aug 1959, P.L.G. Benoit (lef; USNM). DISTRIBUTION.—Afrotropical: Nigeria, Sudan, Zaire, Zim- DIAGNOSIS.—This species is distinguished from Afrolimna babwe. carolinika by the characters indicated in the key and by the fol- NUMBER 617

FIGURES 9-11.—Male terminalia of Afrolimna keiseri (Cogan): 9, epandrium, ccrci, and presurstyli, posterior aspect; 10, internal male terminalia, ventral aspect; 11, same, lateral aspect.

T 0.1mm 1

11

lowing characters from the male terminalia (Figures 9-11): wider basally, gradually narrowed to broad, angulate apex in epandrium broadly and uniformly U-shaped in posterior view lateral view (Figure 10); lateral aedeagal process shorter than (Figure 9), lateral arms slightly wider; cercus moderately broad aedeagus in lateral view, tapered to acute point apically; and ventrally, pointed anteriorly; presurstylus angulate in posterior hypandrium angulate, moderately shallow. view (Figure 9), elbowed, medially oriented arm only slightly TYPE MATERIAL.—The holotype male of Paralimna keiseri wider than vertical portion, medial surface truncate and with Cogan is labeled "Holo-type [round label with red margin]/ 90° dorsomedial angle not projected; postsurstylus in lateral Madagascar [Toliara] Sud Sept-Lacs 100m dct Tulear 13- view curved, slightly widened medially, thereafter narrowed, 16.11.58 B.Stuckenberg." The holotype is double mounted (mi- apex shallowly bifurcate; aedeagal apodeme with prominent nuten in a rectangular block of polyporus), is in good condi- keel, keel irregularly rounded and bearing long, almost paral- tion, and is deposited in the NMSA. Six paratypes {!

OTHER SPECIMENS EXAMINED.—Afrotropical. MADAGAS- gin of wing. Forefemur of male lacking comb-like row of CAR. Toliara: South Berenty Reserve, 20 Apr 1991, A. Freid- short, stout, tooth-like, sometimes flattened setae along apical berg, F. Kaplan (2ef; USNM). V2 of anteroventral surface; foretibia of male lacking several DISTRIBUTION.—Afrotropical: Madagascar. long, slender setae at apex on ventral surface; midtibia bear- ing 4 dorsal extensor setae (basally, subbasally, subapically, and apically); mid- and hindfemora greatly elongate, sub- Genus Corythophora Loew equal to length of abdomen; midtibia bearing 2 ventroapical Corythophora Loew, 1862b: 13 [type species: Corythophora longipes Loew, setae and 1 posteroapical seta; foretarsus with mostly cylin- 1862, monotypy].—Bezzi, 1908a: 195 [synonymy with Dryxo Robineau- drical tarsomeres, not distinctly flattened; forebasitarsus lack- Desvoidy]. ing row of long, slender, pale setulae inserted along anterior Karema Cresson, 1929:182 [type species: Karema loewella Cresson, 1929 surface. {^Corythophora longipes Loew), original designation].—Cogan, 1968: 356-358 [revision]; 1980:661 [Afrotropical catalog].—Mathis and Zatwar- Abdomen: Coloration with distinctly fasciate pattern. Male nicki, 1995:116-117 [world catalog]. [New synonym] terminalia: cercus ovoid to allantoid, bearing numerous setulae along medial margin; presurstylus a narrow, band-like process DIAGNOSIS.—This genus is distinguished from other genera at ventral margin of cercus; aedeagus in lateral view wide ba- of Dryxini, especially Dryxo, which is similar, by the follow- sally, at midlength narrowed to slender process; aedeagal apo- ing combination of characters: ocellar seta present, although deme elongate, with long process extended beyond portion short, inserted slightly in front of anterior ocellus; reclinate bearing keel and with secondary, short process extended exter- fronto-orbital seta present; notopleuron bearing 1 large seta; nally at about midlength; hypandrium wide and robust, lateral presutural supra-alar seta lacking; anepisternum with 1 well- margin with emargination and processes. developed seta along posterior margin; katepisternal seta re- HISTORICAL REVIEW.—Corythophora, as a generic name, duced and lacking row of slender setae along dorsal margin; has been largely overlooked until now. For more than 90 vein R-i bare along dorsum; R stem vein lacking setulae; vein years, Corythophora was treated as a junior synonym of R node bearing 3-5 setulae on dorsum; crossvein dm-cu nor- Dryxo (Bezzi, 1908a). This oversight, which was perpetuated mally developed, nearly straight, forming nearly 90° inner an- in the literature, is typical of interpreting a written description gle with vein M; forefemur of both sexes lacking row of without studying the primary types. Limited or essentially no short, peg-like setulae apically along anteroventral surface; access to the types of Corythophora longipes Loew was un- and mid- and hindfemora elongate, subequal to length of ab- doubtedly the primary impediment to recognizing the identity domen. and thus the nomenclatural precedence of Corythophora. Like DESCRIPTION.—Medium-sized to large shore flies, body Bezzi, Cresson (1929) also did not examine Loew's type se- length 3.40-6.60 mm. ries, and he and all subsequent authors have accepted Bezzi's Head: Frons shallowly arched anteroventrally, not pro- determination that Corythophora was a junior synonym of jected forward, sparsely setulose; ocelli in isosceles triangle, Dryxo. As a result, most previous treatments of the species in- distance between posterior pair shorter than between anterior cluded in Corythophora since Bezzi were published under the ocellus and either posterior ocellus; inner vertical seta and rec- name of Karema (Cresson, 1929; Wirth, 1956, 1960; Cogan, linate fronto-orbital seta well developed; ocellar and paraverti- 1968, 1980; Canzoneri and Raffone, 1987; Canzoneri and cal setae weakly developed or absent. First flagellomere Rampini, 1994; and Mathis and Zatwarnicki, 1995). Of these acutely rounded; arista bearing 10-14 long hairs along length treatments, Cogan's revision (1968) was the most comprehen- of dorsal surface. Face lacking conspicuous facial setae. sive and included illustrations. Parafacial at anterior margin of eye narrow, width much less We borrowed and studied Loew's type and here consider than length of 1st flagellomere. Gena high, height subequal to Corythophora to be the senior synonym of Karema Cresson, as combined length of 1st flagellomere and pedicel. the type species for both generic names, Corythophora lon- Thorax: Anterior dorsocentral setae lacking, only posteri- gipes Loew and Karema loewella Cresson, are conspecific. ormost pair present (0+1); acrostichal setulae poorly devel- DISTRIBUTION (Figure 12).—Afrotropical: Angola, oped, inconspicuous, in 2 rows; presutural supra-alar seta Botswana, Cameroon, Chad, Ethiopia, Ghana, Kenya, Mada- lacking; postpronotal seta well developed; notopleural seta 1; gascar, Malawi, Namibia, Nigeria, Sierra Leone (Northern anepisternum bearing 1 large seta at posterior margin, dorsal Province: Bumbuna), South Africa (Cape Province, Natal, seta lacking; katepisternal seta weakly developed, hair-like. Transvaal), Sudan, Swaziland, Tanzania, Zaire, Zimbabwe. Posterior margin of scutellum broadly rounded. Wing with PHYLOGENETIC RELATIONSHIPS.—Cresson (1929:182) sug- apex of costa at subcostal break bearing 2 short, stout setae; gested that Corythophora (as Karema) is related to Dryxo and vein R| bare along dorsum; R stem vein lacking setulae; vein is a "connectant" between the latter and Paralimna. He also R node bearing 3-5 long, pale setulae on dorsum; crossvein made a comparison with the genus Parydra Stenhammar. Al- dm-cu regularly developed, nearly straight, longer than apical though the latter relationship with Parydra is untenable, being section of vein CuA], and at distinct angle with adjacent mar- based on convergent characters, Cresson's primary suggestion NUMBER 617 11

FIGURE 12.—Distribution map for Corythophora (hatched) and Omyxa (dots).

is confirmed and further documented by our research. Dryxo, nal seta reduced; (3) vein R node bearing 2-3 long, posteriorly with Omyxa as its immediate sister group, forms a lineage that directed setulae on dorsum; (4) adeagal apodeme with long is the sister group to Corythophora. projection extended from near articulation with hypandrium; Unambiguous autapomorphies for Corythophora are (1) ane- (5) midtibia bearing 4 erect setae along dorsum; and (6) poste- pisternum with 1 large seta at posterior margin; (2) katepister- rior margin of scutellum broadly rounded.

Key to Species of Corythophora Loew Femora and tibiae mostly yellowish to ferruginous, fore- and hind femora with some grayish microtomentum dorsally toward base; ocellar seta long, length 2-3 times distance between anterior ocellus and either posterior ocellus; apical scutellar setae nearly parallel sided (Madagascar) 3. C.Jlavipes (Cogan) Femora and tibiae almost entirely black; ocellar seta short, length subequal to distance between anterior ocellus and either posterior ocellus; apical scutellar setae cruciate (widespread Africa) 4. C. longipes Loew

3. Corythophora flavipes (Cogan), new combination angular, blackish brown mesofrons extended broadly to ante- rior margin; frons laterad of mesofrons densely microtomen- FIGURES 13-16 tose with distinct golden brown and whitish gray spots. Ocellar Karema flavipes Cogan, 1968:357; 1980:661 [Afrotropical catalog].—Mathis and Zatwamicki, 1995:116 [world catalog]. seta long, length 2-3 times distance between anterior ocellus and either posterior ocellus, mostly proclinate and slightly di- DIAGNOSIS.—This species is distinguished from its congener vergent; outer vertical seta comparatively long, about % or by the characters indicated in the key. DESCRIPTION.—Moderately large to large shore flies, body more length of inner vertical seta. Antenna rather short, black; length 4.40-6.60 mm. arista bearing 10-12 dorsal rays. Face mostly lightly golden, Head: Head squarish, especially in lateral view. Frons antennal grooves silvery gray. Gena high, silvery gray; gena- wider than long, with distinct, sparsely microtomentose, rect- to-eye ratio 0.51-0.60. 12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

13 T 0.1m1m

15 NUMBER 617 13

FIGURES 13-16 (opposite).—Male terminalia of Corythophora flavipes lows: MADAGASCAR. Antsiranana: Sambirano, Lokobe, (Cogan): 13, epandrium, cerci, and presurstyli, posterior aspect; 14, same, lat- Nossi-Be (6 m), 9-23 Nov 1957, B.R. Stuckenberg (1?; eral aspect; 15, internal male terminalia, ventral aspect; 16, same, lateral aspect. BMNH). Fianarantsoa: Ranomafana, Dec 1955, B.R. Stuck- enberg (19; NMSA). Toamasina: Mananara, Est Ivontaka 1 Thorax: Mesonotum generally gray to black, darker later- (15 m), 10-14 Mar 1958, B.R. Stuckenberg (Id , 2?; BMNH, ally, with distinct stripes in acrostichal and dorsocentral tracks NMSA); Maroansetra, Sahasoa, Fanpanambo (80 m), 26-29 that converge posteriorly; scutellum frequently two toned, apex Mar 1958, B.R. Stuckenberg (ltf, 1?; BMNH); Maroansetra, blackish brown, anterior portion gray to silvery gray anterolat- Navana-Antongil (6 m), 20-25 Mar 1958, B.R. Stuckenberg erally or golden on anterior portion; pleural areas mostly sil- (19; NMSA). Toliara: Ranohira (860 m), 26 Jan^ Feb 1958, very gray to silvery white; anepisternum with distinct, large B.R. Stuckenberg(lrf-,29; BMNH,NMSA); Tulear (=Toliara), golden spot toward posterodorsal angle. Apical scutellar setae Sept-Lacs (100 m), 13-16 Feb, B.R. Stuckenberg (3d"; BMNH, parallel sided. Wing faintly golden brown, mostly hyaline; cos- NMSA). tal-vein ratio 0.24-0.25; M-vein ratio 1.1-1.2. Legs mostly yel- OTHER SPECIMENS EXAMINED.—Afrotropical. MADAGAS- lowish to reddish yellow, with some surfaces darker; femora CAR. Antsiranana: Sambirano, Lokobe, Nossi-Be (6 m), 9-23 reddish yellow with basodorsal areas blackish, especially on Nov 1957, B.R. Stuckenberg (2?; NMSA). Fiana- fore- and hindfemora, and invested with grayish white microto- rantsoa: Ranomafana (7 km W; 900 m), 1 Feb-8 Oct 1988, mentum; posterobasal surface of hindfemur shiny black; tibiae 1990, W.E. Steiner (43 d", 36?; USNM). Mahajanga: Ma- generally reddish yellow, apical portion blackish; tarsi becom- junga (=Mahajanga; km 530), Oct 1949, R.P (1 9; NMSA). ing progressively darker apically, basitarsomere reddish yel- Toamasina: Mananara, Est Ivontaka (15 m), 10-14 Mar 1958, low, apical tarsomere blackish brown. B.R. Stuckenberg (19; NMSA); Maroantsetra, Est Ambodi- Abdomen: Tergite 1 blackish brown at middle, lateral mar- voangy (20 m), 16-20 Mar 1958, B.R. Stuckenberg (39; gins gray; tergite 2 gray but with extensive brown area from NMSA); Maroansetra, Sahasoa, Fanpanambo (80 m), 26-29 middle and extended laterally; tergites 3 and 4 distinctly fasci- Mar 1958, B.R. Stuckenberg (19; NMSA). Toliara: Betroka ate, with wide, dark brown band extended across anterior % or (23°16'S, 46°5'E), 1 Aug 1948, A.R. (Id*; NMSA); South Ber- more of tergite, usually extended more posteriorly at middle, enty Reserve, 20 Apr 1991, A. Freidberg, F. Kaplan (Id"; frequently dark coloration broadly extended to posterior mar- USNM). gin; male tergite 5 entirely blackish brown; female tergite 5 fas- DISTRIBUTION.—Afrotropical: Madagascar. ciate across anterior margin. Male terminalia (Figures 13-16): epandrium in posterior view (Figure 13) broadly inverted U- 4. Corythophora longipes Loew shaped with dorsal margin somewhat truncate and wide, ven- trally extended arms comparatively wide; presurstyli in poste- FIGURES 17-22 rior view (Figure 13) joined medially, forming band-like struc- Corythophora longipes Loew, 1862b: 13.—Cresson, 1929:181 [synonymy with ture at ventral margin of cerci; postsurstylus in lateral view Dryxo ornata (Macquart)]. Dryxo longipes.—Bezzi, 1908b: 187 [generic combination]. (Figure 16) shallowly concave along wide, basal margin, there- Karema loewella Cresson, 1929:182.—Wirth, 1956:388 [list, South Africa after narrowed at midlength to long, curved, slightly tapered (Botswana, Cape Province, Natal, Transvaal)]; 1960:393 [list, Namibia].— process; aedeagus in lateral view (Figure 16) with basal Vz en- Giordani Soika, 1956b:490 [list, Zaire].—Cogan, 1968:357 [revision]; larged, margin toward aedeagal apodeme extended and broadly 1980:661 [Afrotropical catalog].—Canzoneri and Raffone, 1987:70 [list, rounded, and apical Vi greatly narrowed and shallowly curved Kenya].—Canzoneri and Rampini, 1994:245 [list, Sierra Leone].—Mathis anteriorly, aedeagus in ventral view (Figure 15) with lateral, and Zatwamicki, 1995:117 [world catalog]. [New synonym.] rounded lobes subbasally, thereafter gradually tapered apically DIAGNOSIS.—This species is distinguished from its congener to acute point; aedeagal apodeme in lateral view (Figure 16) by the characters indicated in the key. with extended fan-like structure enlarged, broadly rounded; hy- DESCRIPTION.—Medium-sized to large shore flies, body pandrium in ventral view (Figure 15) wide, obtusely pointed at length 3.40-6.0 mm. anterior margin, apex comparatively more narrowly rounded in Head (Figures 17, 18): Head squarish, especially in lateral lateral view (Figure 16). view. Frons wider than long, with distinct, sparsely microto- TYPE MATERIAL.—The holotype male of Karema flavipes mentose, rectangular, blackish brown mesofrons extended Cogan is labeled "Holo-type [round label with red submargin]/ broadly to anterior margin; frons laterad of mesofrons densely Madagascar[.] Est[,] Ambodivoangy 20m[,] dct Maroantsetra microtomentose with distinct golden brown and whitish gray 16-20.111.58 B.Stuckenberg/d7[red triangle]/Karema flavipes spots. Ocellar seta short, length subequal to distance between sp. n. det. B. H. Cogan 1967 [species name and "1967" hand- anterior ocellus and either posterior ocellus, proclinate; outer written]." The holotype is double mounted (minuten in rectan- vertical seta short, about V2 length of inner vertical seta. An- gular block of silicon), is in excellent condition, and is depos- tenna rather short, black; arista bearing 10-12 dorsal rays. Face ited in the MNHN. One paratype (d*; NMSA) bears the same mostly lightly golden, antennal grooves silvery gray. Gena locality label data as the holotype. Other paratypes are as fol- high, silvery gray; gena-to-eye ratio 0.52-0.56. 14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

19 18

FIGURES 17-19.—Corythophora longipes Loew: 17, head, lateral aspect; 18, same, anterior aspect; 19, mesono- tum, dorsal aspect. Scale=0.5 mm.

Thorax (Figure 19): Mesonotum generally dark gray to with wide, dark brown band extended across anterior 1/2-% of black, darker laterally; scutellum frequently two-toned, apex tergite, usually extended more posteriorly at middle, but some blackish brown, anterior portion gray to silvery gray anterolat- specimens with band weaker at middle; male tergite 5 with erally or golden on anterior portion; pleural areas mostly sil- only anterolateral corners dark brown, remainder gray, female very gray to silvery white; anepisternum with some faint, tergite 5 fasciate across anterior margin; some specimens with golden coloration. Apical scutellar setae cruciate. Wing faintly golden to silvery microtomentum on ventral surface of tergite, golden brown, mostly hyaline; costal-vein ratio 0.23-0.28; M- usually adjacent to dark band. Male terminalia (Figures vein ratio 0.95-1.0. Femora and tibiae almost entirely black, 20-22): epandrium in posterior view (Figure 20) U-shaped, only femoral apex and extreme base of tibiae yellowish; ventral with dorsal margin rounded and ventrally extended arms com- surface of femora bare, shiny, dorsal surface moderately paratively narrow; presurstyli in posterior view (Figure 20) not densely microtomentose; tibiae silvery white, extensively and fused medially, appearing as 2 narrow, band-like processes ex- densely microtomentose, especially anterobasal and poster- tended medially, with apices pointed; postsurstylus in lateral obasal surfaces, and appearing reflective; tarsi black, mostly view (Figure 22) long and comparatively narrow, not enlarged covered with moderately dense, gray to whitish gray microto- basally, shallowly curved, apex bilobed; aedeagus in lateral mentum. view (Figure 22) with basal 1/2-% greatly enlarged, margin Abdomen: Tergite 1 dark brown at middle, lateral margins away from aedeagal apodeme extended and broadly rounded, gray; tergite 2 mostly gray; tergites 3 and 4 distinctly fasciate, and apical V2 greatly narrowed and shallowly curved posteri- NUMBER 617 15

FIGURES 20-22.—Male terminalia of Corythophora lon- gipes Loew: 20, epandrium, cerci, and presurstyli, poste- rior aspect; 21, internal male terminalia, ventral aspect; 22, same, lateral aspect.

T 0.1mm 1

20

21

orly, aedeagus in ventral view (Figure 21) shield-like, lateral ure 21) broad with rounded lateral margins, anterior margin margins lacking conspicuous lobes and at midlength gradually concave medially. tapering to bilobed apex (rounded notch at apex); aedeagal TYPE MATERIAL.—The holotype female of Corythophora apodeme with extended fan-like structure only slightly en- longipes Loew is labeled "[South Africa.] 'Caffraria' 446. larged and narrowly rounded; hypandrium in ventral view (Fig- [handwritten]/332. [handwritten]/ Corythophora longipes 16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

[typewritten]/Naturhistoriska Riksmuseet Stockholm Loan no Park (Timbetene Tswiri waterholes; savanna woodland near 798/96 [light bluish green label]." The holotype is directly Skukuza), 9 Dec 1972, B.R. and P. Stuckenberg (29; NMSA); pinned, is in good condition (hindtarsus on left side missing Kruger Park, Tshorwane, Dec 1972, B.R. and P. Stuckenberg and some setae on head missing), and is deposited in the NRS. (19; NMSA); Rio Limpopo (23°00'S, 27°57'E), 25-26 Apr The holotype female of Karema loewella Cresson is labeled 1972 (lcf, 39; BMNH); Waterbert, Jul 1948, O. Beyer (lcf; "[South Africa. Transvaal:] Barberton. 12.8.16 [12 Aug 1916] BMNH). H.K.Munro. [date handwritten; label with black submargin]/ SUDAN. Katire (12 km W), 28 Aug 1980, A.L. Armstrong TYPE 9 Karema LOEWELLA E.T.CressonJr. [red label; spe- (19; USNM). W. Darfur, Jebel Marra, Killing (2134 m), 7 Apr cies name and gender handwritten]/Karema loewella Cresson 1932, M. Steele (2cf; ANSP, BMNH). HOLOTYPE No: 976 [species name, author, and number hand- SWAZILAND. Mbuluzone River, 28 Mar 1984, P. Kelly, J. written in red ink]." The holotype is double mounted (minuten Cory (2cf; BMNH). Mluwla Pools, 17 Mar 1984, P. Kelly, J. in polyporus rectangular block), is in fair condition (several se- Cory, (19; BMNH). Sihoye, Mar 1984, P. Kelly, J. Cory (29; tae of head and thorax, especially on right side, lacking; right BMNH). Sihoye Stream, 29 Mar 1984, P. Kelly, J. Cory (29; wing removed, glued to locality label), and is deposited in the BMNH). NMSA. A female paratype (19; ANSP) bears the same locality TANZANIA. Bukoba, 10 Apr 1912, C.C. Gowdey (lcf; label data as the holotype. BMNH). OTHER SPECIMENS EXAMINED.—Afrotropical. ANGOLA. ZAIRE. Ituri Kasenyi (Lac Albert), 5 Jul 1953, J. Verbeke Cunene: Rocadas, Cunene (river), 19-22 Feb 1972 (6cf, 7?; (lcf; IRSN). Kongold (25°25'S, 23°E), 8 Feb 1974, R. Baker BMNH). Namibe: Mocamedes (2 mi N), 29 Feb 1972 (lcf, (19; BMNH). Matadi (19.3 km N), 28 Jul 1957 (lcf; USNM). 1 ?; BMNH). Without Further Locality: Bruco, 26 Feb-2 Kivu: Rutshuru, Sep-Oct 1934, 1936, Delville (5cf, 13 9; Mar 1972 (17c/, 13?; BMNH, TZ). MRAC, ZMHU). lie Idjwi: Shashi-Busibu, 15 Apr 1953, J. BOTSWANA. River Nata (20°12'S, 26°H'E), 23 Apr 1972 Verbeke (lcf, 29; IRSN); Uvira, Kawezi, 23 May 1958, G. (19; BMNH). River Semowane (20°25'S, 26°23'E), 23-24 Apr Marlier(2cf, 19; MRAC). 1972 (19; BMNH). ZIMBABWE. Balla-Balla, Apr 1931, A. Cutherbertson (19; CAMEROON. West Cameroons: Bolo, 30 Jan 1970, F.H.L. AMNH). Lower Lundi River (Chipinda Pools), 22 Oct 1960, Disney (lcf; BMNH). R.Goodier(19;BMNH). CHAD. W. Goulem, 8-9 Aug 1958, G.B. Popov (lcf; BMNH). DISTRIBUTION.—Afrotropical: Angola, Botswana, Came- ETHIOPIA. Ever, 30 Oct 1968, R. Kano (lcf, 19; BMNH). roon, Chad, Ethiopia, Ghana, Kenya, Malawi, Namibia, Nige- GHANA. Akosombo, 22-25 Jun 1973, L. Knutson (19; ria, Sierra Leone (Northern Province: Bumbuna), South Africa USNM). (Cape Province, Natal, Transvaal), Sudan, Swaziland, Tanza- KENYA. Biretwo (40 km E Eldoret), 12 May 1991, A. Freid- nia, Zaire, Zimbabwe. berg, F. Kaplan (9

FIGURE 23.—Distribution map for Dryxo.

Indonesia (Java, Sulawesi, Sumatra), Philippines (Luzon, noted in the key that follows (see four synapomorphies in the Mindanao, Samar), Sri Lanka, Taiwan, Thailand. Palearctic: first half of the first couplet). In addition, we suggest that the Canary Islands, China, Iran (Baluchistan), Japan (Hokkaido, shape of the presurstylus is similar within the group and is Honshu, Ishikawa, Kyushu, Shikoku), Morocco, Russia (Far probably another synapomorphy. The presurstylus is wide East). (best seen in posterior view), lacks a basomedial arm, and the DISCUSSION.—Species of Dryxo are among the largest of medial margin is irregular, having one or two notches or shal- shore flies and are characterized by many derived character low breaks. states that establish the monophyly of the genus. Synapomor- A second species group, the lispoidea group, also includes phies that we have discovered are as follows: (1) ocellar seta three species: D. digna, D. lispoidea, and D. nudicorpus. This absent; (2) reclinate fronto-orbital seta absent; (3) prescutellar group is characterized by the three characters noted in the key acrostichal setae absent; (4) 6-10 large setae inserted postero- (last half of couplet 5). The presurstylus in this group has a ba- dorsad of katepisternal seta; (5) R stem vein bearing several somedial arm that is especially well developed in D. lispoidea long, pale setulae; (6) short setulae along dorsum of vein Rj; and D. nudicorpus. A well-developed basomedial arm is proba- (7) crossvein dm-cu long, sinuous, oriented parallel with pos- terior margin of wing, forming obtuse inner angle with vein bly a plesiomorphic character, however, as it occurs in other M; and (8) tarsomeres bearing long setae anteroapically and species of Dryxo (D. woodi Cresson), in its sister group, posteroapically, length of setae subequal to width of tarsomere Omyxa, and elsewhere, such as in Paralimna. at apex. The other three species of Dryxo are unplaced in species Although we did not analyze the phylogenetic relationships groups, although D. woodi and D. freidbergi (described herein) among the species of Dryxo rigorously, some patterns are evi- may be closely related, as their adults are quite similar exter- dent and are noted here. The species groups are informally nally and both species occur in the Afrotropical Region. Dryxo designated, meaning that we do not recognize them in a classi- india (described herein) appears to be a comparatively derived fication as subgenera. The ornata group comprises three spe- species, as evidenced by the unusually flattened tarsomeres of cies: D. brahma (described herein), D. margaretae Cogan, and the male and by the shape of the structures of the male termina- D. ornata (Maquart). This group is characterized and its lia (quadrate presurstylus and angulate aedeagal apodeme, es- monophyly is corroborated by several synapomorphies as pecially the keel). NUMBER 617 19

Key to Species of Dryxo Robineau-Desvoidy 1. Tergites 3 and 4 with dark band along posterior margin, at least in middle, band some- times connected laterally with anterolateral spot or spot separate; tergite 1 with me- dial patch of short, dorsally erect setulae; male hindbasitarsus and 2nd hindtarsom- ere bearing long, slender setae dorsally; katepisternal seta well developed, length subequal to notopleural seta (the ornata group) 2 Tergites either with dark band toward anterior margin (at most with very thin dark area along posterior margin) or lacking band; tergite 1 with sparse, inconspicuous, dorsal- ly erect setulae; male hindtarsi lacking long, slender setae dorsally; katepisternal seta usually greatly reduced, much weaker than notopleural seta, or lacking 4 2. Inner vertical seta much reduced, about Vi length of outer vertical seta; male hindbasi- tarsus bearing bush-like, long, slender setae along entire length of dorsum (Oriental) 5. D. brahma, new species Inner vertical seta subequal to or slightly longer than outer vertical seta; male hindbasi- tarsus bearing several long, slender setae toward base, setae becoming shorter and much fewer toward apex 3 3. Postpronotal seta absent; R stem vein bearing about 6 long, thin setulae along posterior margin basad of humeral crossvein (Afrotropical: Madagascar) 10. D. margaretae Cogan Postpronotal seta well developed; R stem vein bearing 2 or 3 long, thin setulae on dor- sum along posterior margin basad of humeral crossvein (Afrotropical) 12. D. ornata (Macquart) 4. Postsutural supra-alar seta greatly reduced or absent; anepisternum lacking moderately long, slender setae along posterior margin (Afrotropical) 7. D. freidbergi, new species Postsutural supra-alar seta well developed; anepisternum bearing 1 or 2 moderately long, slender setae along posterior margin 5 5. Parafacial essentially uniformly concolorous; femora and tibia concolorous, faintly reddish, with sparse whitish microtomentum; mesonotum grayish tan to grayish brown, lacking dark brown spots or areas 6 Parafacial with wide blackish brown to golden brown stripe immediately laterad of an- tennal base, contrasting distinctly with silvery gray remainder of parafacial; femora except apices gray, concolorous with pleural areas, contrasting with yellow tibiae; mesonotum with numerous small, dark brown spots at bases of setae and setulae, with 2 large, somewhat triangular dark brown spots at posterior margin of scutum, and with 2 spots on scutellum around bases of apical scutellar setae (the lispoidea group) 7 6. Foretarsomeres 2-5 of male greatly expanded apically, width of 2nd tarsomere greater than length, narrow basally at attachment with preceding tarsomere; tergites 3-5 of female with dark brown spot anterolaterally; wing of male lacking black spots (Oriental) 8. D. india, new species Foretarsomeres 2-5 of male normally developed, width of 2nd tarsomere far less than length; tergites 3-5 of female generally unicolorous, lacking darkened spot antero- laterally; wing of male with 2 blackened, microtomentose spots, 1 in cell dm at pos- terior margin adjacent to pointed convergence of crossvein dm-cu and vein CuAj, and 1 at posterior margin adjacent to vein Aj +CuA2 (Afrotropical) 13. D. woodi Cresson 7. Dark band on tergites 3-5 lacking narrow, medial extension to posterior margin; post- pronotal seta present; scutellar disc, and in some specimens the frons, lacking a gray, medial, microtomentose stripe (Palearctic) 11. D. nudicorpus Miyagi Dark band on tergites 3-5 with narrow, medial extension to posterior margin; post- pronotal seta absent; frons and scutellar disc mostly dark but with gray, medial, mi- crotomentose stripe 8 20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Katepisternal seta well developed, subequal to notopleural seta; tergites 2-5 gray on ventral side, lacking blackish brown spot; anepisternum mostly gray, at most with brown spot near middle; 1 well-developed anepisternal seta on posterior margin (Oriental) 6. D. digna Osten Sacken Katepisternal seta weakly developed, much thinner and shorter than notopleural seta; tergites 2-5 with blackish brown spot on ventral side; anepisternum with large, wide, transverse, dark brown stripe; 2 weakly developed anepisternal setae along posterior margin (Australasian/Oriental, Oriental) 9. D. lispoidea Robineau-Desvoidy

5. Dryxo brahma, new species as usually short stripes along acrostichal and dorsocentral tracks; scutellum with medial area gray, becoming blackish FIGURES 1,24-26 gray to blackish brown posteriorly and laterally and with dark Dryxo lispoidea of authors, not Robineau-Desvoidy, 1830:787 [misidentifica- brown areas at bases of lateral setae; anepisternum generally tion, in part].—Becker, 1926:93 [review, list, "Persisch-Belutschistan"].— Cogan and Wirth, 1977:332 [in part; Oriental catalog].—Cogan, 1984:144 concolorous with rest of pleural area, sometimes slightly [in part; Palearctic catalog].—Mathis and Zatwarnicki, 1995:116 [in part; darker, lacking maculation in middle. Postsutural supra-alar world catalog]. seta well developed but slightly shorter than postalar seta; ane- pisternal setae 1-3, thinly developed; katepisternal seta well DIAGNOSIS.—This species is distinguished from congeners developed, length subequal to notopleural seta. Wing hyaline; by the following combination of characters: inner vertical seta much reduced, length about V4 that of outer vertical seta; male R stem vein bearing 5-7 short, thin, pale setulae dorsally; vein hindbasitarsus with bush-like, long, slender setae along entire Rj with short, sparse setulae along dorsal length; costal-vein length of dorsum; 2nd hindtarsomere of male bearing long, ratio 0.31-0.33; M-vein ratio 0.65-0.76. Leg color variable; slender setae on dorsal surface; katepisternal seta well devel- femora blackish brown with moderately dense whitish gray mi- oped, length subequal to notopleural seta; 3rd and 4th tergites crotomentum; tibiae blackish brown apically, basal portion yel- with dark band along posterior margin, at least in middle, pos- lowish red, thinly invested with whitish gray microtomentum; terior band sometimes connected laterally with anterolateral tarsi blackish brown dorsally, yellowish orange to orange ven- spot or spot separate; and 1st tergite with medial patch of short, trally, apical tarsomeres becoming blackish gray; forebasitar- dorsally erect setulae. somere of male normally developed, similar to basitarsomeres DESCRIPTION.—Large to very large shore flies (Figure 1), of mid- and hindlegs; foretarsomeres 2-5 of male normally de- body length 7.00-11.20 mm; dorsum of head and thorax gener- veloped, not greatly expanded apically, width much less than ally dark brown with some gray to yellowish gray; legs gener- length; lateroapical setae of foretarsi of male tapered; hindbasi- ally dark colored. tarsus and 2nd tarsomere of male bearing dense patch of long, Head: Generally densely microtomentose. Frons mostly to slender setae along dorsum; tarsomeres of midleg bearing long almost entirely gray to tan, sometimes with some blackish setae lateroapically, aligned with plane of leg, length of setae brown maculations, especially posteriorly as short stripe lat- subequal to width of tarsomere at apex. erad of ocelli and extended anteriad and at vertex laterad; inner Abdomen: Third and 4th tergites with dark band along pos- vertical seta short and thin, length about 1/2-% that of outer ver- terior margin, at least in middle, posterior band sometimes con- tical seta; ocelli arranged in equilateral triangle. Antenna black- nected laterally with anterolateral spot, or spot separate; 1st ish brown, usually with some gray to tan microtomentum dor- tergite with medial patch of short, dorsally erect setulae. Male sally; arista white to tannish white basally, bearing 11-13 terminalia (Figures 24-26): epandrium in posterior view (Fig- dorsal rays. Face, parafacial, gena, and microtomentose portion ure 24) like a broadly pointed arch, narrow dorsally and of clypeus concolorous, whitish gray, contrasting with darker- slightly angulate, arms wider at dorsolateral level, ventrally be- colored frons; face bare except for series of setulae paralleling coming narrower; cercus ovoid, wider subdorsally; presursty- frontal suture; parafacial with transverse, golden brown to lus in posterior view (Figure 24) broadly triangular, length blackish brown stripe or maculation at level of antennal base, nearly twice basal width, dorsomedial angle formed, not pro- thereafter ventrally concolorous with face. Eye slightly higher duced, medial margin irregular, with angulate indentation and than wide. Gena with anterior portion concolorous with face extension; ventral apex rounded, lateral margin very shallowly and parafacial, becoming whiter posteriorly, high, height convex; postsurstylus in lateral view (Figure 26) wide basally, slightly more than V2 eye height; gena-to-eye ratio 0.57-0.63. produced at angle toward aedeagus, setulose lobe just beyond Clypeus with dorsal V2 of medial portion bare. midlength, inner lobe of bilobed apex wider than outer lobe, es- Thorax: Mesonotum generally mottled, gray to tannish sentially bare of setulae; aedeagus in lateral view (Figure 26) gray, especially peripherally and along setal tracks, with bases about equally wide throughout short length, inner surface irreg- of setae blackish brown; scutum with some blackish brown to ularly convex, external surface irregularly concave with angu- blackish gray areas sometimes evident posteriorly; gray areas late indentation, apex truncate; aedeagal apodeme in lateral NUMBER 21

FIGURES 24-26.—Male terminal la of Dryxo brahma Mathis and Zatwarnicki: 24, epandrium, cerci, and presurstyli, posterior aspect; 25, internal male termina- lia, ventral aspect; 26, same, lateral aspect.

24

25

view (Figure 26) with keel moderately shallow and with long, brahma

Nathan (17tf, 119; IRSN); Yelburga (near), 10 Dec 1974, K. brown to blackish brown but with a gray, medial, microtomen- Ghorpade (la-, 19; USNM). Orissa: Pottangi (21 km SE; tose stripe; anepisternum mostly gray, at most with a brown 550 m), 4 Feb 1962, D.Q. Cavagnaro, E.S. Ross (19; CAS). spot near middle; 1 well-developed anepisternal seta along pos- Rajasthan: Rajaputana, Deesa (=Disa, 24°15'N, 72°10'E), terior margin; katepisternal seta well developed, subequal to Apr-7 Nov 1897, 1898, C.G. Nurse (9tf, 2$; BMNH). Tamil notopleural seta; male hindtarsus lacking long, slender setae on Nadu: Anamali Hills, Cinchona, May 1964, T.R.S. Nathan dorsal surface; 1 st tergite with sparse, inconspicuous, dorsally (4tf; USNM); Coimbatore (11°N, 76°58'E), Mar-Oct 1947, erect setulae; dark band on tergites 3-5 with a narrow, medial 1955, 1957, P.S. Nathan (25o", 229; BPBM, USNM); Coim- extension to posterior margin; postpronotal seta absent; and batore (25.8 km NW; 640 m), 8 Mar 1962, D.Q. Cavagnaro, tergites 2-4 gray on ventral side, lacking blackish brown spot. E.S. Ross (39; CAS); , Karubagaram, 19 Jan 1951, DESCRIPTION.—Large to very large shore flies, body length P.S. Nathan (19; IRSN); Karaikal, Kurunebaganan, Apr 1955, 7.20-9.30 mm; generally densely microtomentose, gray, with 1 P.S. Nathan (3d , 39; USNM); Karaikal, Pondicherry (11°56'N, dark brown maculation pattern on dorsum. 79°53'E), Mar 1964, T.R.S. Nathan (lcf; USNM); Madras (19; Head: Generally densely microtomentose. Frons mostly USNM); Nedungadu (10=58^, 79°46'E), 29 Apr, P.S. Nathan dark brown, with a usually thin, gray, medial stripe; dark brown (lcf; USNM); Nilgiri Hills, Cherangade (1067 km), Oct 1950, area laterad of ocellar triangle in V-shaped pattern, with its ver- P.S. Nathan (1

Abdomen: Tergites 1 and 2 mostly densely microtomen- ally; ventral surface entirely gray, densely microtomentose. tose, gray; tergite 1 with faint dark brown area medially; tergite Male terminalia (Figures 27-29): epandrium in posterior view 2 with a brown to blackish brown spot laterally; tergites 3 and 4 (Figure 27) rounded arch-shaped, narrow dorsally, arms be- with wide, dark brown band along anterior margin and with coming wider ventrally; cercus ovoid, wider subventrally; medial projection extended posteriorly to posterior margin; presurstylus in posterior view (Figure 27) narrowly triangular, tergite 5 with dark brown band and shallow extension medi- twice as long as wide, dorsomedial angle narrowly produced,

FIGURES 27-29.—Male terminalia of Dryxo digna Osten-Sacken: 27, epandrium, cerci, and presurstyli, posterior aspect; 28, internal male ter- minalia, ventral aspect; 29, same, lateral aspect.

T 0.1m1m 24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY medial margin irregular, ventral apex narrowly pointed, lateral DESCRIPTION.—Large to very large shore flies, body length margin shallowly and evenly arched; postsurstylus in lateral 5.90-6.45 mm; generally white with some brown coloration view (Figure 29) narrowed basally, becoming wider to subapi- dorsally and with apices of legs yellowish. cal widest level and thereafter apical extension short, parallel Head: Generally densely microtomentose. Frons mostly sided, and broadly truncate apically, in ventral view (Figure 28) silvery white to silver with some golden areas medially, anterior with inner lobe of bilobed apex longer, wider, and essentially of anterior ocellus; length of inner and outer vertical setae about bare of setulae; aedeagus in lateral view (Figure 29) as long as coequal, both reduced and about equal to length of pedicel; wide, inner margin rounded, external margin with short, nar- ocelli arranged either in equilateral or slightly isosceles trian- row, rounded projection; aedeagal apodeme in lateral view gle, usually with distance between posterior ocelli shorter than (Figure 29) with keel very shallow; hypandrium moderately between either posterior ocellus and anterior ocellus. Antenna wide and deep. blackish brown to black; scape and pedicel sometimes with TYPE MATERIAL.—The lectotype male of Dryxo digna Osten considerable fine, grayish to brownish microtomentum, espe- Sacken, here designated to stabilize and make more universal cially dorsally; arista silvery white, bearing 8 or 9 dorsal rays. the use of this name, is labeled "Dryxo digna O.S. cf type Face white to faintly tannish white, bare except for series of set- ulae paralleling frontal suture; parafacial concolorous with face [handwritten]/Philippinen [handwritten]/Coll. O.S. [handwrit- except for a wide, brown to blackish brown stripe immediately ten]/ coll. Oldenberg/Coll. Osten-Sacken/Syntypus [red label; laterad of antennal base. Eye slightly higher than wide. Gena "1972" handwritten on the underside]/Typus [maroon-red label high, anterior portion concolorous with face and parafacial, be- with pencil mark through it]/LECTOTYPE tf Dryxo digna Os- coming whiter posteriorly; gena-to-eye ratio 0.50-0.54. ten Sacken By Mathis & Zatwarnicki [handwritten except for Clypeus mostly bare medially, shiny black, becoming densely "LECTOTYPE" and "By"; label with black submarginal bor- microtomentose laterally, especially along ventral margin. der]." The lectotype is pinned directly, is in good condition (posterior margin of wings torn, thorax cracked around pin), Thorax: Mesonotum with small dark brown spots at bases and is deposited in the DEI. There are also six paralectotypes of setae and setulae, 2 large, somewhat triangular to quadrate (4tf, 2?; DEI) that are here designated. dark brown spots toward posterior margin of scutum, varying amounts of dark brown areas medially and along setal tracks OTHER SPECIMENS EXAMINED.—PHILIPPINES. Luzon. (none consistently expressed), and 2 spots on scutellum at Quezon: Maliao, C.F. Baker (l

FIGURES 30-32.—Male terminalia of Dryxo fre- idbergi Mathis and Zatwarnicki: 30, epandrium, cerci, and presurstyli, posterior aspect; 31, internal male terminalia, ventral aspect; 32, same, lateral aspect.

T 0.1mm 1

(Figure 32) wide basally, becoming narrower to apex, with sub- furcate arms at attachment with hypandrium long; hypandrium apical, angulate indentation, in ventral view with both lobes of moderately wide and deep. bilobed apex of about equal size, inner lobe essentially bare of TYPE MATERIAL.—The holotype male is labeled "CAME- setulae; aedeagus in lateral view (Figure 32) with base very ROON Kribi (beach) Rt. N7 28.29.XI. 1987 [28-29 Nov 1987] wide, immediately narrowed, especially on inner surface, ex- A. FREIDBERG/HOLOTYPE Dryxo freidbergi

NMNH. The allotype female and six other paratypes (3d1, 29; yellowish orange to orange; femora extensively whitish gray USNM) bear the same locality data as the holotype. basally; tarsomeres black apically and laterally, apical tarsom- DISTRIBUTION.—Afrotropical: Cameroon. ere almost entirely black; foretarsomeres 2-5 of male greatly ETYMOLOGY.—The specific epithet, freidbergi, is a genitive expanded apically, width equal to length, narrow basally at at- patronym to honor one of the collectors of this new species tachment with preceding tarsomere; lateroapically, setae of fo- from Africa. retarsomeres of male tapered evenly; hindbasitarsus of male lacking long, slender setae on dorsum; 2nd hindtarsomere of 8. Dryxo india, new species male bearing short setulae, none clustered; tarsomeres of mid- leg bearing long setae lateroapically, aligned with plane of leg, FIGURES 33-35 length of setae subequal to width of tarsomere at apex. DIAGNOSIS.—This species is distinguished from congeners Abdomen: Dorsum mostly grayish white, lacking darkened by the following combination of characters: mesofrons yellow- bands anteriorly or posteriorly; male with slightly darkened ish orange to grayish tan; inner vertical seta subequal in length area anterolaterally on 2nd tergite, this area also bearing denser to outer vertical seta; scape and pedicel reddish, 1st flagello- patch of setulae, those inserted more toward anterolateral mar- mere black; parafacial lacking transverse stripe; mesonotum gin longer, more erect; female with dark brown spot anterolat- generally concolorous with frons, lacking dark spots at bases of erally. Male terminalia (Figures 33-35): epandrium frequently setae and lacking maculation pattern; postsutural supra-alar exposed and conspicuous from dorsal angle, in posterior view seta well developed; anepisternal seta and katepistemal setae (Figure 33) rounded arch-shaped, narrow dorsally and shal- well developed although slender, subequal; legs mostly yellow- lowly emarginate, arms widest subventrally, thereafter ven- ish orange to orange; femora extensively whitish gray basally; trally becoming narrower, bluntly pointed; cercus narrowly tarsomeres black apically and laterally; foretarsomeres 2-5 of hemispherical, dorsal apex rounded, ventral apex pointed; male greatly expanded apically, width equal to length, and nar- presurstylus in posterior view (Figure 33) irregularly quadrate, row at attachment with preceding tarsomere; hindbasitarsus of width dorsally only slightly more than length, dorsomedial an- male lacking long, slender setae on dorsum; 2nd hindtarsomere gle very narrowly produced, apex acutely pointed, medial mar- of male lacking long setulae along dorsum; 3rd and 4th tergites gin sinuous, ventral margin nearly flat, with shallow papilla- lacking dark band; and 1st tergite with medial patch of short, dorsally erect setulae. like point near middle, lateral margin nearly straight; postsur- stylus in lateral view (Figure 35) C-shaped with curved dorsal DESCRIPTION.—Large to very large shore flies, body length extension attached to aedeagus, ventral portion wider and with 7.60-8.70 mm; head and thorax generally tan to whitish tan apex only very shallowly bifurcate; aedeagus in lateral view dorsally, abdomen mostly whitish gray; legs yellowish orange, (Figure 35) with base moderately wide, becoming wider to becoming darker brown apically and whitish basally. widest width subapically, thereafter narrowed immediately to Head: Generally densely microtomentose. Frons yellowish rounded apex; aedeagal apodeme in lateral view (Figure 35) L- orange to grayish tan; inner vertical seta subequal in length to shaped, with keel very shallow, ridge-like, and pointed ven- outer vertical seta; ocelli arranged in isosceles triangle, with distance between posterior ocelli longer than between either troapically, bifurcate arms at attachment with hypandrium posterior ocellus and anterior ocellus. Antenna with scape and short; hypandrium short, moderately wide and deep. pedicel faintly reddish orange with thin dusting of whitish gray TYPE MATERIAL.—The holotype male is labeled "Nedun- microtomentum; 1st flagellomere blackish brown to black; gadu S. India 4-II P.S. Nathan/HOLOTYPE Dryxo india tf arista silvery white, bearing 11-14 dorsal rays. Face, parafa- W.N.Mathis & Zatwarnicki [red label; species name and "& cial, gena, and clypeus concolorous, creamy white, contrasting Zatwarnicki" handwritten]." The holotype is pinned directly, is with darker-colored frons; face bare except for series of setulae in good condition, and is deposited in the NMNH. paralleling frontal suture; parafacial lacking transverse stripe, OTHER SPECIMENS EXAMINED.—INDIA. Karnataka: Shi- dorsal Va slightly tannish; essentially uniformly concolorous moga (568.5 m; trapped by 200 Petromax lamp), River Tunga, with face, lacking stripe. Eye slightly higher than wide. Gena 29 May 1936, P.S. Nathan (5d\ 9?; IRSN); Shimoga, 20 Jan-29 high, height slightly more than V2 eye height, anterior portion May 1936, P.S. Nathan (3?; IRSN). Tamil Nadu: Karaikal concolorous with face and parafacial, becoming whiter posteri- (lO^H 79°45'E), Jul 1956, P.S. Nathan (13d1, 179; BMNH); orly; gena-to-eye ratio 0.56. Clypeus mostly microtomentose, Kurumbagaram, Karaikal, Jul-Dec 1951, 1952, P.S. Nathan medial portion with dorsal V2 or more bare, somewhat shiny. (4d\ 8$; BMNH); Tranquebar (=Tarangambadi, ll°02'N, Thorax: Mesonotum yellowish tan to whitish tan, lacking 79°51'E), Jul-Dec 1952, P.S. Nathan (7d\ 9?; BMNH, IRSN); darkened spots at bases of setae and lacking maculation pat- Walayar Forests (10°51'N, 76°51'E), South Malabar, 5 Aug tern; postsutural supra-alar seta well developed but slightly 1956, P.S. Nathan (2cT, 19; BMNH). shorter than postalar seta; anepisternal and katepistemal setae DISTRIBUTION.—Oriental: India (Karnataka, Tamil Nadu). well developed although slender, subequal in length. Wing hya- ETYMOLOGY.—The specific epithet, india, is a noun in appo- line; R stem vein bearing 4-6 short, thin setulae dorsally; cos- sition and refers to the Indian subcontinent where this species tal-vein ratio 0.40-0.44; M-vein ratio 0.57-0.61. Legs mostly occurs, perhaps exclusively. NUMBER 617 27

URES 33-35.—Male terminal la of Dryxo india Mathis ! Zatwarnicki: 33, epandrium, cerci, and presurstyli, terior aspect; 34, internal male terminalia, ventral ect; 35, same, lateral aspect.

35

9. Dryxo lispoidea Robineau-Desvoidy Dryxo spreta Osten Sacken, 1882:242. [New synonym.] Dryxo digna of authors, not Osten Sacken [misidentification].—Cresson, FIGURES 36-38 1929:182 [synonymy with D. spreta].—Cogan and Wirth, 1977:332 [in part; Dryxo lispoidea Robineau-Desvoidy, 1830:787.—Becker, 1926:93 [review, Oriental catalog].—Mathis and Zatwamicki, 1995:115 [in part; world catalog]. list, "Persisch-Belutschistan"].—Cogan and Wirth, 1977:332 [in part; Orien- tal catalog].—Cogan, 1984:144 [in part; Palearctic catalog].—Mathis and DIAGNOSIS.—Although Cresson (1929) synonymized D. Zatwamicki, 1995:116 [in part; world catalog]. spreta {-D. lispoidea) with D. digna, a precedent followed by Cyphops fasciatus Jaennicke, 1867:368. [New synonym.] subsequent workers, this species is readily distinguished from Dryxo (Cyphops) fasciata.—Cogan and Wirth, 1977:332 [generic combina- tion].—Mathis, 1989:644 [Australasian/Oceanian catalog].—Mathis and Zat- D. digna and other congeners by the following combination of warnicki, 1995:115 [world catalog]. characters: frons and scutellar disc mostly brown to blackish 28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY brown but with a gray, medial, microtomentose stripe; anepis- Abdomen: Tergites 1 and 2 mostly gray, posterior margin ternum with a large, wide, transverse, dark brown stripe; 2 of tergite 2 dark brown and sometimes with medial stripe dark weakly developed anepisternal setae along posterior margin; brown; tergites 2-5 with dark brown band along anterior mar- katepisternal seta usually greatly reduced, much weaker than gin, posterior margin of brown band with medial projection ex- notopleural seta; male hindtarsi lacking long, slender setae on tended to posterior margin; lateral margin gray to whitish gray; dorsal surface; 1 st tergite with sparse, inconspicuous, dorsally ventral surface usually with brown maculation, usually more erect setulae; dark band on tergites 3-5 with narrow, medial ex- evident on tergites 2-5. Male terminalia (Figures 36-38): epan- tension to posterior margin; postpronotal seta absent; and terg- drium in posterior view (Figure 36) rounded arch-shaped, nar- ites 2-5 with blackish brown spot on ventral side. row dorsally, arms wider at dorsolateral level, ventrally becom- DESCRIPTION.—Large to very large shore flies, body length ing narrower; cercus ovoid, wider subventrally, pointed 5.50-9.00 mm; dorsum of head and thorax generally dark dorsomedially; presurstylus in posterior view (Figure 36) brown, with some gray to yellowish gray; legs generally dark deeply bifurcate, each arm long and slender, medial arm ori- colored. ented medially, apex turned dorsomedially and bearing 5 or 6 Head: Generally densely microtomentose. Frons mostly setulae extended ventrally, lateral arm oriented ventrally, grad- dark brown, with some tan and gray coloration; dark brown ually tapering to medially oriented apex; postsurstylus in lat- area in V-shaped pattern, with its vertex posterior and its arms eral view (Figure 38) narrow basally, becoming wider at very broad; dark brown area adjacent to eye near vertex, ex- midlength and with a swelling along anterior margin, margin tended from vertical setae forward to middle-width level of bearing several setulae, postsurstylus thereafter narrowed be- eye; inner vertical seta short and thin, length about 1/2-% that fore becoming wider at ventral margin, ventral margin bifur- of outer vertical seta; ocelli arranged in equilateral triangle. cate with posterior lobe slightly wider, posteroventral margin Antenna blackish brown; arista blackish brown, bearing 12-14 bearing 5 or 6 setulae; aedeagus in lateral view (Figure 38) nar- dorsal rays. Face, parafacial, gena, and clypeus concolorous, row basally, immediately becoming wider, with broad apex whitish gray, contrasting with darker-colored frons; face bare truncate; aedeagal apodeme in lateral view (Figure 38) with except for series of setulae paralleling frontal suture; parafacial keel very shallow, ridge-like, and developed on half attached to with transverse, golden brown to blackish brown stripe or mac- base of aedeagus only; hypandrium moderately wide and deep. ulation at level of antennal base, thereafter ventrally concolor- TYPE MATERIAL.—The neotype of Dryxo lispoidea Robin- ous with face. Eye slightly higher than wide. Gena high, height eau-Desvoidy, here designated, is the lectotype male of Cy- slightly more than Vz eye height; anterior portion concolorous phops fasciatus and bears labels as indicated below for that with face and parafacial, becoming whiter posteriorly; gena-to- specimen. The original type series of Dryxo lispoidea was col- eye ratio 0.62-0.65. lected on the island of Sumatra and apparently has been de- Thorax: Mesonotum generally blackish brown; scutum stroyed (the second author and Volker Hollmann, pers. comm., with some gray areas marginally, along transverse suture, and 1995, after they visited Paris and consulted with L. Matile). As as small, usually short stripes along acrostichal and dorsocen- it is likely that the type specimens represented this species, tral tracks; scutellum with medial stripe and lateral margins based in part on distribution, we are fixing its name as the se- whitish gray to gray except for dark brown areas around base nior objective synonym by designating the lectotype of Cy- of lateral scutellar setae; anepisternum with dark brown spot phops fasciatus as the neotype for Dryxo lispoidea. near middle. Postsutural supra-alar seta well developed but The lectotype male of Cyphops fasciatus Jaennicke, here slightly shorter than postalar seta; anepisternal setae 1 or 2, designated to stabilize and make more universal the use of this weakly developed, thin; katepisternal seta very weakly devel- name, is labeled "Mas[culus]. Wiesb. Java. Fritz [handwritten]/ oped, smaller than anepisternal seta(e). Wing hyaline; R stem von Heyden [collection]/Typus [red label with black border]/ vein bearing 5-7 short, thin, pale setulae dorsally; costal-vein Cyphops fasciatus Jaen[nicke] Java [handwritten, square label ratio 0.41-0.44; M-vein ratio 0.71-0.77. Legs mostly dark col- with black border]/LECTOTYPE d" Cyphops fasciatus Jaen- ored; femora and tibiae blackish brown to brown with moder- nicke By Mathis & Zatwarnicki [handwritten except for "LEC- ately dense gray microtomentum; tarsi yellowish orange to or- TOTYPE" and "By"; label with black submarginal border]." ange ventrally, dorsally gray to yellowish gray, apical The lectotype is pinned directly, is in poor condition (partially tarsomeres becoming blackish gray; forebasitarsomere of male eaten by dermestids), and is deposited in the NSF. normally developed, similar to basitarsomeres of mid- and The lectotype male of Dryxo spreta Osten Sacken, here des- hindlegs; foretarsomeres 2-5 of male normally developed, not ignated to stabilize and make more universal the use of this greatly expanded apically, width much less than length; some name, is labeled "Dryxo spreta O.S. type. [handwritten]/Philip- lateroapical setae of foretarsi of male parallel sided and with pinen Coll. Osten-Sacken/D 419. [handwritten]/spreta O.S. broadly rounded apices; hindbasitarsus of male lacking long, [handwritten]/Becker det./Syntypus [red label; number "1972" slender setae on dorsum; tarsomeres of midleg bearing long se- on the underside]/TYPUS [pink label]/LECTOTYPE

37 38

FIGURES 36-38.—Male terminalia of Dryxo lispoidea Robineau-Desvoidy: 36, epandrium, cerci, and presurstyli, posterior aspect; 37, internal male terminalia, ventral aspect; 38, same, lateral aspect. 30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ginal border]." The lectotype is pinned directly, is in poor con- from each other using external characters as well as structures dition (right eye caved in, thorax cracked toward right of pin, of the male terminalia (see diagnosis and description). left hindleg missing), and is deposited in the DEL There is also Specimens of other species were often misidentified as D. li- a male paralectotype (DEI) that is here designated. spoidea, especially specimens on the Indian subcontinent, OTHER SPECIMENS EXAMINED.—Australasian/Oceanian. where a few species of Dryxo are apparently sympatric. AUSTRALIA. Queensland: Byfield State Forest, 3 Jan 1976, G. Daniels (9

FIGURES 39-41.—Male terminalia of Dryxo margare- tae Cogan: 39, epandrium, cerci, and presurstyli, pos- terior aspect; 40, internal male terminalia, ventral aspect; 41, same, lateral aspect.

pleural area, sometimes slightly darker, lacking maculation in sally, yellowish orange to orange ventrally, apical tarsomeres middle. Postsutural supra-alar seta well developed but slightly becoming blackish gray; forebasitarsomere of male normally shorter than postalar seta; anepisternal setae 1-3, thinly devel- developed, similar to basitarsomeres of mid- and hindlegs; fo- oped; katepisternal seta well developed, length subequal to no- retarsomeres 2-5 of male normally developed, not greatly ex- topleural seta. Wing hyaline; R stem vein bearing 5-7 short, panded apically, width much less than length; lateroapical setae thin, pale setulae dorsally; vein Rj with short, sparse setulae of foretarsi of male tapered; hindbasitarsus and tarsomere 2 of along length; costal-vein ratio 0.29-0.33; M-vein ratio male bearing sparse patch of long, slender setae posterodor- 0.56-0.62. Leg color variable; femora blackish brown with sally; hindtarsomere 2 of male bearing some long setae along moderately dense whitish gray microtomentum; tibiae blackish dorsum; tarsomeres of midleg bearing long setae lateroapically, brown apically, basal portion yellowish red, thinly invested aligned with plane of leg, length of setae subequal to width of with whitish gray microtomentum; tarsi blackish brown dor- tarsomere at apex. 32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Abdomen: Tergites 3 and 4 with dark band along posterior some specimens, frons lacking medial, microtomentose, gray margin, at least in middle, band sometimes connected laterally stripe; katepisternal seta greatly reduced, much weaker than with anterolateral spot, or spot separate; 1st tergite with medial notopleural seta or lacking; postsutural supra-alar seta present, patch of short, dorsally erect setulae. Male terminalia (Figures well developed; anepisternum bearing 2 long, slender setae 39-41): epandrium in posterior view (Figure 39) rounded arch- along posterior margin; legs, especially femora and tibiae, shaped, narrow dorsally and slightly angulate, arms wider at mostly grayish black to black; male hindtarsi lacking long, dorsolateral level, ventrally becoming narrower; cercus ovoid, slender setae on dorsal surface; tergites with band toward ante- wider subventrally; presurstylus in posterior view (Figure 39) rior margin, at most with very thin, dark area along posterior broadly triangular, length about 1.5 times basal width, dorso- margin; tergite 1 with sparse, inconspicuous, dorsally erect set- medial angle narrowly produced, short, medial margin with 2 ulae; tergites 3-5 with prominent dark bands along anterior step-like processes, ventral apex broadly rounded, lateral mar- margin; and dark bands of tergites 3-5 lacking narrow, medial gin very shallowly arched; postsurstylus in lateral view (Figure extension to posterior margin. 41) wide and angulate basally, bearing shallow, setulose lobe DESCRIPTION.—Large shore flies, body length 7.45-7.90 just beyond midlength; inner lobe of bilobed apex wider and mm; generally blackish brown dorsally but with considerable slightly longer than outer lobe, essentially bare of setulae; areas gray. aedeagus in lateral view (Figure 41) with base moderately Head: Generally densely microtomentose. Frons mostly to wide, immediately narrowed, especially on inner surface, ex- entirely brownish black, at most with thin gray stripe anterior ternal surface with recurved, narrow lobe at midlength, apex ir- of anterior ocellus and slightly lighter areas at lateral margin; regularly and relatively broadly rounded; aedeagal apodeme in inner vertical seta short and thin, length about V6 that of outer lateral view (Figure 41) with keel shallow, ridge-like, with vertical seta; ocelli arranged in equilateral triangle. Antenna long, narrow extension toward attachment with hypandrium, blackish brown to black; arista blackish brown to black, bear- bifurcate arms at attachment with hypandrium long; hypan- ing 13-16 dorsal rays. Face, parafacial, gena, and clypeus con- drium somewhat wide and deep. colorous, whitish gray to silvery gray, contrasting with darker- TYPE MATERIAL.—The holotype male is labeled "Holo-type colored frons; face bare except for series of setulae paralleling [round label with red submargin]/Madagascar[.] [Tamatave:] frontal suture; parafacial with transverse, blackish brown stripe Est[,] Sahasoa Fampanambo 80m[,] dct Maroansetra[,] or maculation at level of antennal base, thereafter ventrally 26-29.III.58[,] B.Stuckenberg/Dryxo margaretae sp. n. det. B. concolorous with face, dark coloration merged and concolor- H. Cogan 1967 [species name and "1967" handwritten]." The ous with lateral margins of frons. Eye slightly higher than holotype male is pinned directly, is in excellent condition (right wide. Gena high, slightly more than Vz eye height, anterior por- midleg missing), and is deposited in the MNHN. Two paratypes tion concolorous with face and parafacial, becoming whiter (ld\ 19; NMSA) bear the same locality data as the holotype. posteriorly; gena-to-eye ratio 0.62-0.65. Other paratypes are as follows: MADAGASCAR. Toama- Thorax: Mesonotum generally blackish brown except for sina: Moramanga (=Mamaranga), Perinet (Niagarakely For- gray postpronotum, area just laterad of postsutural supra-alar est), Dec 1955, B.R. Stuckenberg (2cf, 19; BMNH, NMSA). seta, and faint stripes, especially anteriorly, in dorsocentral OTHER SPECIMENS EXAMINED.—Afrotropical. MADAGAS- track; pleural areas uniformly gray; scutellum more or less CAR. Fianaratsoa: Ranomafana (600 m), 18-20 Jan 1990, evenly rounded, not conspicuously truncate; postpronotal seta W.E. Steiner (\

FIGURES 42-44.—Male terminalia of Dryxo nudi- corpus Miyagi: 42, epandrium, cerci, and presurst- yli, posterior aspect; 43, internal male terminalia, ventral aspect; 44, same, lateral aspect.

**!

43 44

tae on dorsum; tarsomeres of midleg bearing long setae lat- dorsolateral level, becoming narrower ventrally; cercus ovoid, eroapically, aligned with plane of leg, length of setae subequal wider subventrally, pointed dorsomedially; presurstylus in pos- to width of tarsomere at apex. terior view (Figure 42) deeply bifurcate, each arm long and Abdomen: Anterior margin of tergites 3-5 brownish black, slender, medial arm bearing 5 or 6 short setulae extended ven- posterior margin of band curved and pointed medially. Male trally, arm oriented medially, apical portion becoming wider, terminalia (Figures 42—44): epandrium in posterior view (Fig- apex turned dorsomedially, lateral presurstylar arm oriented ure 42) rounded arch-shaped, narrow dorsally, arms wider at ventrally, mostly parallel sided but apical portion slightly ex- 34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY panded and narrowly rounded at apex; postsurstylus in lateral long, thin setulae along posterior margin basad of humeral view (Figure 44) narrow basally, becoming wider at midlength crossvein; male hindbasitarsus with several long, slender setae and with a swelling along anterior margin, swelling bearing toward base, setae becoming shorter and much sparser toward several setulae, thereafter slightly narrowed before becoming apex; male hindtarsomere 2 bearing long, slender setae on dor- wider at ventral margin, ventral margin bifurcate with both sal surface; katepisternal seta well developed, length subequal lobes broadly developed and subequal in size, ventrolateral sur- to notopleural seta; tergites 3 and 4 with dark band along poste- face bearing numerous setulae; aedeagus in lateral view (Fig- rior margin, at least in middle, band sometimes connected lat- ure 44) narrow basally, immediately becoming wider, with erally with anterolateral spot, or spot separate; and tergite 1 broad apex truncate; aedeagal apodeme in lateral view (Figure with medial patch of short, dorsally erect setulae. 44) with keel moderately shallow, ridge-like, and developed on DESCRIPTION.—Moderately large to very large shore flies, half attached to base of aedeagus only; hypandrium moderately body length 4.75-8.85 mm; generally blackish brown dorsally wide and deep. but with considerable areas gray. TYPE MATERIAL.—The holotype male is labeled "Jyozankei Head (Figures 45, 46): Generally densely microtomentose. 17-VII-1961/Japonia Hokkaido I, MIYAGIMype Dryxo nudi- Frons mostly to almost entirely gray to tan, sometimes with corpus I. Miyagi [red label; all except for "-type" handwrit- some blackish brown maculation, especially posteriorly as 1 or ten]." The holotype is pinned directly, is in poor condition 2 short, anteriorly directed stripes of variable shape laterad of (damaged in shipment; wings, most legs, and most setae miss- ocelli and from vertex; inner vertical seta subequal or slightly ing; legs and wing fragments glued on a label below speci- longer than outer vertical seta; ocelli arranged in equilateral tri- men), and is deposited in the HUS. angle. Antenna blackish brown, usually with some gray to tan OTHER SPECIMENS EXAMINED.—Palearctic. JAPAN. microtomentum dorsally; arista white to tannish white basally, lshikawa: Oono, Kanazawa, 11 Jun 1964, H. Kurahashi (lex; bearing 11-13 dorsal rays. Face, parafacial, gena, and microto- BMNH). Label data in Japanese kanji that we were unable to mentose-covered portion of clypeus concolorous, whitish gray, read(59;BMNH). contrasting with darker-colored frons; face bare except for se- CHINA. Shan-hai-Kwan (in mountains), 1 Sep 1906, F.M. Th- ries of setulae paralleling frontal suture; parafacial with trans- omson (1 cf, 29; BMNH). verse, golden brown to blackish brown stripe or maculation at RUSSIA. Far East: Juzhnoe Primor'e, Lazovski zapovednik level of antennal base, thereafter ventrally concolorous with (25 km SE L'azo), 31 Jul 1986 (2cf; ZMUM). Primurskiy Kraj, face. Eye slightly higher than wide. Gena high, height slightly Pokrovka (48°3lX 134°55'E), Suj-fun, 2 Aug 1935, A. Mish- more than Vfe eye height, anterior portion concolorous with face chenko (1 ?; ZMUM). Pristan Bulochna, 10 viorst vyshe Ust- and parafacial, becoming whiter posteriorly; gena-to-eye ratio komenogosk, 9 Jul 1914, Kazantseva (29; ZMUM). 0.52-0.56. Clypeus with dorsal Vb of medial portion bare. DISTRIBUTION.—Palearctic: China, Japan (Hokkaido, Hon- Thorax (Figure 47): Mesonotum generally mottled, par- shu, lshikawa, Kyushu, Shikoku), Russia (Far East). tially to mostly dark brown, with some gray to tannish gray ar- REMARKS.—This is the only species of Dryxo that extends eas, especially peripherally and along setal tracks, bases of se- into temperate climates. tae blackish brown; scutum with anterior Vfe extensively gray We continue to interpret, recognize, and treat this species' with brown to blackish brown maculation, latter becoming epithet, nudicorpus, as a noun in apposition, as apparently Miy- much darker with some gray areas sometimes evident, usually agi did when first describing this species. Corpus is a Latin as short stripes along acrostichal and dorsocentral tracks; noun meaning "body." scutellum generally and extensively blackish brown, only me- diobasal, somewhat triangular area whitish gray; anepisternum 12. Dryxo ornata (Macquart) generally concolorous with rest of pleural area, sometimes slightly darker, lacking maculation in middle. Postsutural su- FIGURES 45-50 pra-alar seta well developed but slightly shorter than postalar Blepharitarsis ornatus Macquart, 1844:411. seta; anepisternal setae 1-3, thinly developed; katepisternal Dryxo ornata.—Bezzi, 1908a: 194 [generic combination].—Wirth, 1955:53 seta well developed, length subequal to notopleural seta. Wing [list, Tanzania (Ngaruka)]; 1956:388 [list, Mozambique, South Africa (Natal, hyaline; R stem vein bearing 5-7 short, thin, pale setulae dor- Transvaal), Zimbabwe]; 1960:393 [list, Namibia].—Giordani Soika, 1956b:490 [list, Zaire].—Cogan, 1968:359-360 [revision]; 1980:661 [Afro- sally; vein R] with short, sparse setulae along length; costal- tropical catalog]; 1984:144 [Palearctic catalog].—Can/.oneri and Meneghini, vein ratio 0.32-0.36; M-vein ratio 0.56-0.58. Leg coloration 1969:163 [list, Zaire].—Canzoneri and Rampini, 1994:245 [list, Sierra Le- variable; femora and tibiae generally concolorous, blackish one].—Mathis and Zatwamicki, 1995:116 [world catalog]. brown with moderately dense whitish gray microtomentum; Dryxo ornatus.—Becker, 1926:93 [review, list, Canary Islands]. tarsi blackish brown dorsally, yellowish orange to orange ven- DIAGNOSIS.—This species is distinguished from congeners trally, apical tarsomeres becoming blackish gray; forebasitar- by the following combination of characters: inner vertical seta somere of male normally developed, not swollen, similar to ba- subequal or slightly longer than outer vertical seta; postprono- sitarsomeres of mid- and hindlegs; foretarsomeres 2-5 of male tal seta present, well developed; R stem vein bearing 2 or 3 normally developed, not greatly expanded apically, width NUMBER 617 35

46

FIGURES 45—47.—Dryxo ornata (Macquart): 45, head, lateral aspect; 46, same, anterior aspect; 47, mesonotum, dorsal aspect. Scale=0.5 mm. much less than length; lateroapical setae of foretarsi of male ta- very shallow, setulose lobe just beyond midlength, both lobes pered; hindbasitarsus of male bearing sparse patch of long, of bilobed apex about equal in size, inner lobe essentially bare slender setae posterodorsally; hindtarsomere 2 of male bearing of setulae; aedeagus in lateral view (Figure 50) about equally some long setae along dorsum; tarsomeres of midleg bearing wide throughout length, inner surface irregularly convex, ex- long setae lateroapically, aligned with plane of leg, length of ternal surface irregularly concave with angulate indentation, setae subequal to width of tarsomere at apex. apex truncate; aedeagal apodeme in lateral view (Figure 50) Abdomen: Tergites 3 and 4 with dark band along posterior with keel shallow, ridge-like, with long, narrow extension to- margin, at least in middle, band sometimes connected laterally ward attachment with hypandrium, bifurcate arms at attach- with anterolateral spot or spot separate; 1st tergite with medial ment with hypandrium long; hypandrium somewhat narrow patch of short, dorsally erect setulae. Male terminalia (Figures and shallow. 48-50): epandrium in posterior view (Figure 48) like a broadly TYPE MATERIAL.—The syntypes of Blepharitarsis ornatus, pointed arch, narrow dorsally and slightly angulate, arms which were collected at "D'Afrique, Shubar" (=Sierra Leone?), wider at dorsolateral level, ventrally becoming narrower; cer- were not located and may well not exist. The second author and cus ovoid, wider subventrally; presurstylus in posterior view Volker Hollmann, who visited MNHN in 1995, were unable to (Figure 48) broadly triangular, length about 1.5 times basal find any specimen(s) (pers. comm., 1995). Lacking a primary width, dorsomedial angle broadly produced, short, medial type, we have followed Cogan's concept of this species, as he margin with shallow lobe at midlength, ventral apex broadly was the last revisor. Cogan's concept was and is generally rec- rounded, lateral margin very shallowly arched; postsurstylus ognized (see references in species synonymy), and the species in lateral view (Figure 50) narrow, pointed basally, bearing is easily identified. Thus, we have not deemed it necessary to 36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 48-50.—Male terminalia of Dryxo ornata (Macquart): 48, epandrium, cerci, and presurstyli, posterior aspect; 49, internal male ter- minalia, ventral aspect; 50, same, lateral aspect.

T 0.1mm 1

49

designate a neotype. This is a common and widespread Afro- davala (12.9-16.1 km NW Sa da Bandeira, 27-29 Mar 1972 tropical species and is the only species of Dryxo known thus far (ltf; BMNH). Bruco, 26 Feb-2 Mar 1972 (Id1; BMNH). Lan- from Sierra Leone. dana (=Cacongo; 5°14'S, 12°8'E), Port Congo (19; AMNH). OTHER SPECIMENS EXAMINED.—Afrotropical. ANGOLA. BOTSWANA. River Nata (20°12'S, 26°H'E), 23 Apr 1972 Cuanza Sul: Cachoeiras (20 mi SW Gabela), 18-19 Mar (lcf; BMNH). 1972 (5d\ 119; BMNH). Cunene: Rocadas, Cunene (River), BURUNDI. Burburi (2000 m), 10 Oct 1948, F.J. Francois (Id1, 19-22 Feb 1972 (6d\ 69; BMNH, TZ). Namibe: Rio Giraul 19; IRSN). Lestrade (2d\ 29; MRAC). Rumonge (sur la rive (16.1 km NE Mocamedes), 27-29 Feb 1972 (Id"; BMNH); du Lac Tanganyika; 780 m), 13 Feb 1934, 1949, A. Lestrade, Mocamedes (3.2 km N), 29 Feb 1972 (Id", 29; BMNH); Tun- F.J. Francois (8d\ 79; IRSN, MRAC). Rumongi River, Lac NUMBER 617 37

Tanganyika, 16 Sep 1948, F.J. Francois (10d\ 39; IRSN). NIGER. Zungeru, 1 Feb-15 Mar, 1911, 1912, J.W.S. Macfie Usumbura (780 m), 19 Apr 1953, F. Francois (6o\ 3?; IRSN). (8er, 59; AMNH, BMNH). Alagua, 1912, J.W.S. Macfie (19; CAMEROON. Kribi (route N7, beach), 28-29 Nov 1987, A. BMNH). Shao, 14 Mar 1912, J.W.S. Macfie (19; BMNH). Freidberg, F. Kaplan (lOoT, 179; USNM). RWANDA. Kisenyi (pres emb. river Sebeya), 4 Apr 1953, J. CHAD. Ouli Bangala, Logone, 1 Mar 1966, J.C. Hitchcock, Verbeke (5

ZAIRE. Kivu: Kasongo, Lualaba River, Jul 1959, P.L.G. and in gazetteers available to us, "Nalanda" is located in central Benoit (Id1, 19; MRAC); Kasongo, Pongo River, Oct 1959, Sri Lanka, which would be a new record from that country as P.L.G. Benoit (3 ex; MRAC); Lake Tanganyika, Lueba near well as from the Oriental Region. The specimen (19; ANSP), Baraka, 19 Aug 1927, R. Bois (9

FIGURES 51-53.—Male terminalia of Dryxo woodi Cres- son: 51, epandrium, cerci, and presurstyli, posterior aspect; 52, internal male terminalia, ventral aspect; 53, same, lateral aspect.

T 0.1mm 1

52

erately long, slender setae along posterior margin; katepisternal surface; foretarsus of male unmodified, similar to that of fe- seta usually greatly reduced, much weaker than notopleural male; forebasitarsomere of male normally developed, similar to seta, or lacking. Wing hyaline; costal-vein ratio 0.38-0.41; M- that of female, not conspicuously swollen; lateroapical setae of vein ratio 0.49-0.51; wing of male with 2 blackened, microto- forebasitarsomere of male tapered, sharply pointed apically. mentose spots, 1 in cell dm at posterior margin, adjacent to Abdomen: Dorsum and venter of tergites generally uni- pointed convergence of cross vein dm-cu and vein CuAj, and 1 formly whitish gray to tan and lacking darkened areas, such as at posterior margin, adjacent to vein Aj+CuA2. Femora and bands or spots, except for dark medial area toward anterior of tibia concolorous, faintly reddish, with sparse whitish microto- 5th tergite of male. Male terminalia (Figures 51-53): epan- mentum; male hindtarsi lacking long, slender setae on dorsal drium in posterior view (Figure 51) rounded arch-shaped, nar- 40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY row dorsally, arms widest subapically, thereafter becoming nar- lacking; anepisternum with 2 thin, long, hair-like setae along rowed and moderately pointed at ventral apex; cercus ovoid, posterior margin; katepistemum with thin, hair-like seta, lack- both apices equally narrowly rounded; presurstylus in posterior ing row of slender setae near dorsal margin; vein Rj bearing view (Figure 51) deeply bifurcate, each arm long, medial arm several setulae along dorsum; R stem vein basad of humeral very slender, oriented medially with slight arch basally, thereaf- crossvein bearing several pale, thin setulae on ventral surface; ter straight, bearing patch of setulae extended ventrally toward crossvein dm-cu moderately long, shallowly sinuous, general apex of medial arm, lateral arm thickly developed (compared to orientation at distinct angle with posterior margin of wing, medial arm) oriented ventrally, parallel sided throughout most forming acute inner angle with vein M; forefemur of male and of length, apex slightly narrowed to bluntly rounded ventral female lacking row of short, peg-like setulae apically along an- apex; postsurstylus in lateral view (Figure 53) very narrow ba- teroventral surface; and mid- and hindfemora moderately elon- sally, thereafter gradually expanded to very wide, truncate gate, although not subequal to length of abdomen. apex, apical margin sinuous with distinct papilla-like projec- DESCRIPTION.—Medium-sized to large shore flies, body tion, bearing numerous setulae on midlength swelling along an- length 3.80-5.10 mm. terior margin, lacking bifurcation apically; aedeagus in lateral Head: Frons projected forward as a shield-like, squarish, view (Figure 53) somewhat narrow basally, immediately be- nearly flat plate, densely setulose; ocelli in isosceles triangle, coming wider for most of length, with dorsal margin moder- with distance between posterior pair shorter than distance be- ately deeply incised toward base, narrow apex moderately tween anterior ocellus and either posterior ocellus; both inner pointed; aedeagal apodeme in lateral view (Figure 53) slender and outer vertical setae present, but only outer seta well devel- and shal lowly arched, with keel very shallow, lacking ridge-like oped; reclinate fronto-orbital seta, ocellar seta, and paravertical extension, developed into T-like apex attached to base of aede- seta lacking. First flagellomere with apex moderately rounded; agus, apex attached to hypandrium evenly narrow throughout arista short, stout, bearing 7-9 long hairs along length of dorsal length; hypandrium short, moderately wide and quite deep. surface and at apex. Face mostly bare, bearing row of short, TYPE MATERIAL.—The holotype female (not a male, as indi- pale setulae in shallow groove paralleling parafacial. Parafacial cated in the original publication) of Dryxo woodi Cogan is la- at anterior margin of eye moderately wide, width subequal to beled "Type [round label with red submargin]/704 [handwrit- height of 1st flagellomere. Gena high, height slightly greater ten]/[MALAWI] Nyasaland[,] Cholo R.C.WoodTYPE No. than combined length of 1st flagellomere and pedicel. Dryxo WOODI

^•rT' __ __ ,—^

'' v;','' '!• '!'" '

55 FIGURES 54-56.—Omyxa scuta Mathis and Zatwarnicki: 54, head, anterior aspect; 55, same, lateral aspect; 56, mesonotum, dorsal aspect. 42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY dial, oriented posteromedially then anteromedially, medial gin large, broadly curved, in dorsal view base of aedeagus thinly but deeply incised to anterior ocellus; inner vertical seta much wider than apex, becoming slightly wider to about short, thin, about V2 size of outer vertical seta (these with apex midlength, thereafter tightly recurved, forming a short, lateral apparently broken). Antenna essentially concolorous with ante- process, then angled medially to wider, capitate apex; aedeagal rior margin of frons, 1st flagellomere slightly darker; 1st flagel- apodeme in lateral view (Figure 60) with only apices curved lomere short, length subequal to combined length of scape and posteriorly toward attachments with aedeagus and hypandrium, pedicel; arista extended beyond apex of 1st flagellomere, bear- anterior crest long, irregularly arched; hypandrium in lateral ing 7-10 dorsal hairs, these more curved toward apex. Face view (Figure 60) angulate, anterior % wide, anterior margin concolorous with anterior portion of frons but duller and uni- truncate, in ventral view (Figure 59) deeply incised subbasally, formly colored, dorsal portion of face between antennal bases with narrowly angulate lateral processes. vertical, not projected, thereafter ventrally flared anteriorly to TYPE MATERIAL.—The holotype male is labeled "S. Iran 40 oral margin, generally lacking setae or setulae except bearing km SE Minab 21. 5 1973 [21 May 1973]/Loc. no. 205 Exp. row of short, thin, pale-colored setulae in vertical groove paral- Mus. Nat Praha/Cat. No. 33751 [pink label with black submar- lel to parafacial. Parafacial moderately wide, slightly less than gins; handwritten]." The holotype is double mounted (minuten width of pedicel at apex, concolorous with face; gena high, be- in a small block of cork), is in good condition (some setae coming wider posteriorly from parafacial, also becoming more missing), and is deposited in NMSAC. The allotype female and silvery white, lacking a prominent genal seta; gena-to-eye ratio six paratypes (4cf, 2?; NMSAC, USNM) bear the same locality 0.38. Clypeus band-like, black with gray microtomentum; pal- data as the holotype but with different catalog numbers pus short, mostly black but with some gray microtomentum. (NMSAC 33747-33754). Other paratypes are as follows: Ori- Thorax (Figure 56): Mesonotum generally concolorous ental. INDIA. Tamil Nadu: Coimbatore, Jul 1956, P.S. Nathan with posterior portion of frons, with only very sparse microto- (l

59 60

FIGURES 57-60.—Male terminalia of Omyxa scuta Mathis and Zatwarnicki: 57, epandrium, cerci, and presur- styli, posterior aspect; 58, same, lateral aspect; 59, internal male terminalia, ventral aspect; 60, same, lateral aspect.

Head: Frons shallowly arched anteroventrally, not pro- ocellus and either posterior ocellus; ocellar seta, inner vertical jected forward, sparsely setulose; ocelli in isosceles triangle, seta, and reclinate fronto-orbital seta well developed; procli- distance between posterior pair longer than between anterior nate fronto-orbital setae greatly reduced, subequal in length, 44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY anterior seta not conspicuously larger than posterior seta; ture; aedeagus simple, usually broad basally, apex pointed to paravertical seta well developed, subequal to length of poste- truncate; aedeagal apodeme with conspicuous, moderately rior notopleural seta. First flagellomere broadly rounded; arista deep to deep keel; hypandrium wider than long (best seen in bearing 6-8 long hairs along dorsal surface. Face with 2 or 3 posterior view), shallowly and sometimes irregularly curved in long (subequal to length of large, reclinate seta), inclinate (but lateral view. not cruciate) facial setae on ventral V2 of face and 3 or 4 HISTORICAL REVIEW.—Oedenopiforma was first described smaller setulae interspersed along same vertical alignment as by Cogan (1968) as a subgenus of Paralimna and initially in- larger setae, facial setae and setulae vertically aligned, parallel cluded only two species, O. madecassa (Giordani Soika, with parafacial. Parafacial at anterior margin of eye narrow, 1956a), the type species, and O. argentea, which Cogan de- width much less than length of 1st flagellomere. Gena high, scribed in the same paper. Other than Cogan's (1980) Afrotro- height subequal to combined length of 1 st flagellomere and pical catalog and our world catalog (Mathis and Zatwarnicki, pedicel. 1995), there are no further references to the genus and species. Thorax: Dorsocentral setae 4 (1+3), well developed, pos- Consistent with our catalog, herein we accord generic status to terior pair displaced laterally; acrostichal setulae well devel- this taxon, but we also expand the generic concept to include oped, conspicuous, in about 8 irregular rows; presutural supra- two additional species, Paralimna uniseta Malloch and P. liga- alar seta well developed, subequal in length to anterior, dorso- buei Canzoneri. The latter species is Afrotropical in distribu- central seta; postpronotal seta well developed; notopleural se- tion, like the two originally included species, and P. uniseta is tae 2, posterior seta shorter; anepistemum variable, bearing 1 known thus far only from Australia. Structurally, P. uniseta is large seta along posterior margin and with dorsal seta reduced, more divergent from the other three species (see first couplet of or (usually) bearing 2 setae with apical seta V4-V2 length of key and description of male terminalia). larger, ventral seta; katepistemal seta well developed. Posterior None of the included species is common in collections avail- margin of scutellum truncate, nearly flat. R stem vein bearing 1-3 (usually 2) setulae, these oriented posteriorly; crossvein able to us, and virtually nothing is known about their natural dm-cu regularly developed, nearly straight, longer than apical history. section of vein CuAj, and at distinct angle with posterior mar- DISTRIBUTION (Figure 61).—Australasian/Oceanian: Austra- gin of wing. Forefemur bearing comb-like row of short, stout, lia (New South Wales, Northern Territory, Western Australia). spine-like setulae along apical V2 of anteroventral surface; Afrotropical: Kenya, Madagascar, Namibia, Nigeria, Rwanda, foretibia of male bearing several long, slender setae at apex on South Africa (Transvaal), Sudan, Zaire. Palearctic: Iran. ventral surface; forebasitarsus with row of long, slender, pale DISCUSSION.—Oedenopiforma is characterized by several setulae inserted along anterior surface; midtibia with 3 dorsal synapomorphies that confirm the monophyly of the genus. extensor setae (subapically, subbasally, and near middle); mid- Among derived character states that we have discovered are the and hindfemora normally developed, much shorter than length following: (1) forefemur bearing comb-like row of short, stout, of abdomen. spine-like setulae along apical V2 of anteroventral surface; (2) Abdomen: Lacking pattern of facia. Male terminalia: epan- R stem vein bearing 1-3 (usually 2) setulae; and (3) abdomen drium in posterior view a broad to moderately narrow inverted lacking a pattern of fascia. U, bearing setulae toward posterior margin; cercus ovoid, dor- Specimens of Oedenopiforma are relatively scarce in collec- sum in posterior view slightly narrower than venter, bearing tions, and despite concerted effort over a number of years, in- numerous setulae, especially on medial portion; presurstylus in cluding examination of thousands of specimens of Dryxini posterior view wider than high, bearing few to many short to from major museums throughout the world, we were able to lo- long setulae; postsurstylus slightly curved anteriorly in lateral cate only a few specimens. In this review, we recognize three view, base wider than apex, especially in lateral view, with species of Oedenopiforma, one of which, O. madecassa, has preapical notch, bearing setulae in patches along inner curva- two junior synonyms, one newly proposed herein.

Key to Species of Oedenopiforma Cogan Facial series of setae 3; paravertical setae greatly reduced or lacking. Stem vein bearing 1 long setula, length equal to twice width of vein at level of insertion. Forefemur bearing minute, peg-like setulae anteroventrally (Australia) 17. O. uniseta (Malloch) Facial series of setae 2, these well separated; paravertical setae moderately to well de- veloped (with same orientation as inner vertical seta although smaller). Stem vein bearing 1-3 (usually 2) setulae, length of each about equal to width of vein at level of insertion. Forefemur bearing conspicuous, peg-like setulae anteroventrally 2 NUMBER 617 45

FIGURE 61.—Distribution map for Oedenopiforma (hatched and dots).

2. Face in both sexes yellow to orangish yellow, distinct from grayish yellow, microto- mentose antenna; forefemur of male lacking row of setulae as above; apex of foretib- ia and forebasitarsus of male bearing long, pale setulae (widespread Africa) 15. O. argentea (Cogan) Face and antenna of male appearing velvety, densely microtomentose, dark brown; face of female silvery gray; antenna of female dark brown (Madagascar, South Af- rica, Sudan) 16. O. madecassa (Giordani Soika)

15. Oedenopiforma argentea (Cogan) low, microtomentose antenna; facial series of setae 2, these well separated. Gena-to-eye ratio 0.34-0.35. FIGURES 62-65 Thorax: Stem vein bearing 1-3 (usually 2) setulae dorsally Paralimna (Oedenopiforma) argentea Cogan, 1968:322; 1980:663 [Afrotropi- along posterior margin, length of each about equal to width of cal catalog]. vein at level of insertion; costal-vein ratio 0.35-0.36; M-vein Oedenopiforma argentea.—Mathis and Zatwarnicki, 1995:117 [catalog, gener- ic recombination]. ratio 0.90-0.94. Forefemur of male lacking row of long, fine, pale, ventrally oriented setulae along anteroventral surface; DIAGNOSIS.—This species is distinguished from congeners apex of foretibia and forebasitarsus of male bearing long, pale by the following combination of characters: in both sexes face setulae. yellow to orangish yellow, distinct from grayish yellow, micro- tomentose antenna; facial series of setae 2, these well sepa- Abdomen: Male terminalia (Figures 62-65): epandrium rated; and forefemur of male lacking row of long, fine, pale, wider than high in posterior view (Figure 62), evenly rounded, ventrally oriented setulae along anteroventral surface. semicircular, width even, bearing setulae of equal length to- DESCRIPTION.—Small to moderately small shore flies, body ward posterior margin, in lateral view (Figure 63) much wider length 1.80-2.30 mm. dorsally, thereafter ventrad narrowed abruptly, anterior margin Head: Paravertical seta well developed, about Va-Vfe length sinuous; cercus ovoid, bearing numerous setulae, especially of inner vertical seta. Arista with 7 or 8 dorsal rays. Face yel- medially; presurstylus horizontal tear-drop-shaped, much low to orangish yellow in both sexes, distinct from grayish yel- wider medially, narrowed laterally, bearing several long setu- 46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

0.1mm

64 65

FIGURES 62-65.—Oedenopiforma argentea (Cogan): 62, epandrium, cerci, and presurstyli, posterior aspect; 63, same, lateral aspect; 64, internal male terminalia, ventral aspect; 65, same, lateral aspect.

lae; postsurstylus with truncate to very shallowly arched basal ten]." The holotype is double mounted (pin in a rectangular margin, base twice apical width, bearing 2 or 3 subapical setu- piece of celluloid), is in fair condition (pin fold in much of me- lae at level of preapical notch, bearing 3 or 4 setulae on basal Vb sonotum), and is deposited in BMNH. of inner curvature; aedeagus with apex pointed in lateral view OTHER SPECIMENS EXAMINED.—Afrotropical. KENYA. Lake (Figure 65); aedeagal apodeme with moderately wide and deep Victoria (10 km S Kisumu), 19 Nov 1986, A. Freidberg (3cf, keel, extended processes about equal in length in lateral view 1?;USNM). (Figure 65); hypandrium wider than long, bluntly rounded an- NAMIBIA. Ameib Farm (on vegetation around pools), 31 teriorly, with lateral pointed process at anterolateral corner, Jan-2 Feb 1972 (9d\ 59; BMNH). Kamanjab (43.5 km ESE), thereafter posteriorly parallel sided, posterior margin broadly Otjitambi Farm, 13-15 Feb 1972 (80tf\ 97?; BMNH). emarginate, excavate. NIGERIA. Kaduna: Zaria, Samaru, 21 Feb-1 Nov 1967, TYPE MATERIAL.—The holotype female of Paralimna ar- 1968 (2o"; BMNH). Kwara: New Bussa, 12 Jan 1970, J.T. gentea Cogan is labeled "Pres. by Imp.BurEnt./Holo-type Medler (lcT; USNM). Osun: Ile-Ife, 20 May 1969, J.T. [round label with red margin]/N. Nigeria. Ilorin. 22.2. 191 Dr. Medler(l?;BMNH). J.W.Scott-Macfie./1920-271/Paralimna argentea sp.n. 9 det RWANDA. Kisenyi (pres emb. riv. Sebeya), 4 Apr 1953, Ver- B.H.Cogan 1966 [species name, gender, and "sp.n." handwrit- beke(2d\ 7?;MRAC). NUMBER 617 47

SOUTH AFRICA. Transvaal: Kruger Park (Olifants River Abdomen: Male terminalia (Figures 66-68): epandrium near Balule; riparian woodland), 9 Dec 1972, B.R. and P. about as wide as high in posterior view (Figure 66), dorsum Stuckenberg (19; NMSA). more acutely rounded and more thinly developed than lateral ZAIRE. Elisabethville (light), 17 Dec 1949, C. Seydel (19; arms, bearing several long setulae toward posterior margin; USNM). cercus broadly ovoid, only slightly narrower dorsally, bearing Palearctic. IRAN. Derpehan (12 km E Senderk), 11-12 May numerous long setulae, evenly scattered; presurstylus in poste- 1977 (13 ex; NMSAC). rior view (Figure 66) as a nearly parallel-sided, horizontal bar DISTRIBUTION.—Afrotropical: Kenya, Namibia, Nigeria, bearing tiny setulae along ventral margin, apex only slightly Rwanda, South Africa (Transvaal), Zaire. Palearctic: Iran. narrower than base; postsurstylus in lateral view (Figure 68) REMARKS.—A male from Sri Lanka may also be this spe- with truncate, nearly flat basal margin, base nearly twice apical cies, although the row of short, stout setae along the anteroven- width, bearing 2 or 3 subapical setulae at level of preapical tral surface is not as well developed. Label data for this male notch and bearing 5 or 6 setulae on basal V3 of inner curvature; are Amp. Dist. Maha Oya, 1 May 1980, W.N. Mathis, T. Wijes- aedeagus with apex irregularly truncate in lateral view (Figure inhe, L. Jayawickrema (lcf; USNM). 68); aedeagal apodeme with wide and moderately deep keel, extended processes about equal in length in lateral view (Fig- 16. Oedenopiforma madecassa (Giordani Soika) ure 68); hypandrium in ventral view (Figure 67) wider than long, bluntly rounded V-shaped, apex of posteriorly extended FIGURES 66-68 arms with wide, medial process, posterior margin very broadly Paralimna madecassa Giordani Soika, 1956a:123.—Cogan, 1968:321 [revi- excavate. sion]; 1980:663 [Afrotropical catalog]. TYPE MATERIAL.—The lectotype male of Paralimna made- Oedenopiforma madecassa.—Mathis and Zatwarnicki, 1995:117 [catalog, ge- neric recombination]. cassa Giordani Soika, here designated to stabilize and make Paralimna brunneifacies Giordani Soika, 1956a: 123.—Cogan, 1968:321 [syn- more universal the use of this name, is labeled "HOLOTYPUS onymy]. [pink label with black submarginal line]/COLL. MUS. Paralimna ligabuei Canzoneri, 1987:84.—Mathis and Zatwarnicki, 1995:121 CONGO Madegascar: [Tamatave:] Maroansetraf,] XII-1949 [world catalog]. [New synonym.] [Dec 1949] J. Vadon/HOLOTYPUS Paralimna madecassa DIAGNOSIS.—This species is distinguished from congeners Soika [red label; handwritten except for "HOLOTYPUS"]." by the following combination of characters: face and antenna The lectotype male is double mounted (minuten), is in good of male appearing velvety, densely microtomentose, dark condition (right fronto-orbital seta missing, posterior portion of brown; face of female silvery gray; facial series of setae 2, left wing destroyed), and is deposited in the MRAC. Although these well separated; antenna of female dark brown; mesofrons Giordani Soika (1956a: 123) wrote that the two specimens com- of male dark brown anteriorly, becoming golden brown posteri- prising the type series are females, one of the syntypes that we orly; anterior fronto-orbits concolorous with anterior portion of are here designating as the lectotype is clearly a male. mesofrons, not appearing velvety; scutum of male either The lectotype female of Paralimna brunneifacies Giordani mostly concolorous with scutellum except for golden brown Soika, here designated to stabilize and make more universal the anteromedial area or with disc of scutellum gray, contrasting use of this name, is labeled "HOLOTYPUS [pink label with with dark brown scutum; and forefemur of male usually bear- black submarginal line]/COLL. MUS. CONGO Madegascar: ing row of long, fine, pale, ventrally oriented setulae along an- [Tamatave:] Maroansetra[,] XII-1949 [Dec 1949] J. Vadon/ teroventral surface. HOLOTYPUS Paralimna brunneifacies Soika [red label; hand- DESCRIPTION.—Small to moderately small shore flies, body written except for "HOLOTYPUS"]." The lectotype female is length 1.65-2.40 mm. double mounted (minuten), is in fair condition (right portion of Head: Paravertical seta well developed, length about 1/3-1/2 mesonotum removed, right mid- and hindtarsi missing), and is inner vertical seta. Arista with 6-8 dorsal rays. Face and an- deposited in the MRAC. tenna of male appearing velvety, densely microtomentose, dark The holotype male of Paralimna ligabuei Canzoneri is la- brown; face of female silvery gray; antenna of female dark beled "[Sudan] Kartoum Bare (nilo) 6-xii-80 Leg. Rallo [hand- brown; facial series of setae 2, these well separated. Gena-to- written]/HOLOTYPUS Paralimna ligabuei nov. det. Canzoneri eye ratio 0.26-0.36. S. [red label; species name handwritten]/MUSEO CIVICO DI Thorax: Scutum of male mostly concolorous with scutel- STORIA NATURALE DI VENEZIA Materiale tipico descritto lum except for golden brown anteromedial area. Stem vein Inv. n. 03732 [red label; number handwritten]." The holotype is bearing 1-3 (usually 2) setulae dorsally along posterior margin, double mounted (minuten in a long, rectangular block of poly- length of each about equal to width of vein at level of insertion; porus), is in fair condition (several setae missing; abdomen re- costal-vein ratio 0.41-0.42; M-vein ratio 0.80-0.85. Forefemur moved and dissected, and the parts are in an attached mi- bearing conspicuous, peg-like setulae anteroventrally; forefe- crovial), and is deposited in the MCV. mur of male with row of long, fine, pale, ventrally oriented set- OTHER SPECIMENS EXAMINED.—Afrotropical. MADAGAS- ulae along anteroventral surface. CAR. Antananarivo: Ambatondrazaka, Centre Station Agric, 48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 66-68.—Oedenopiforma madecassa (Giordani Soika): 66, epandrium, cerci, and presurstyli, posterior aspect; 67, internal male terminalia, ventral aspect; 68, same, lateral aspect.

0.1mm

67 68

Alaotra (800 m), 24 Dec 1957, B.R. Stuckenberg (19; SOUTH AFRICA. Transvaal: Kruger Park (Olifants River BMNH). Toliara: Ampanihy (250 m), 16-18 Feb 1958, B.R. near Balule; riparian woodland), 9 Dec 1972, B.R. and P. Stuckenberg (19; BMNH); Morondava (black light near sand Stuckenberg (ltf; NMSA). beach), 28-31 Mar 1990, W.E. Steiner, C. Kremen, V. Razafi- DISTRIBUTION.—Afrotropical: Madagascar, South Africa mahatratra (39; USNM); Ranohira (860 m), 26 Jan-4 Feb (Transvaal), Sudan. 1958, B.R. Stuckenberg (39; BMNH); Saint Augustin (6 m), REMARKS.—This species is obviously sexually dimorphic, 11-13 Feb 1953, B.R. Stuckenberg (2

70 71

FIGURES 69-71.—Oedenopiforma uniseta (Malloch): 69, epandrium, cerci, and presurstyli, posterior aspect; 70, internal male terminalia. ventral aspect; 71, same, lateral aspect. NUMBER 617 51

1 1962, E.S. Ross, D.Q. Cavagnaro (Id ; USNM); Millstream, 7 topleural setae 2, posterior seta shorter than anterior seta; ane- Apr 1971, D.H. Colless (Id"; ANIC). pisternum bearing 2 poorly developed setae along posterior DISTRIBUTION.—Australasian/Oceanian: Australia (New margin, dorsal seta shorter; katepistemal seta poorly developed South Wales, Northern Territory, Western Australia). or absent. Posterior margin of scutellum truncate, nearly flat. REMARKS.—This species and other undescribed species Veins lacking setulae; costal section I lacking long setae; cross- from Australia (represented by single male specimens) appar- vein dm-cu regularly developed, nearly straight, longer than ently form a species group that is distinguished by the charac- apical section of vein CuAj, and at distinct angle with posterior ters in the first couplet of the key to species of Oedenopiforma. margin of wing. Forefemur shallowly excavate along apical V3 The group should be revised when additional specimens are of anteroventral surface, excavation bearing comb-like row of available. short, stout, tooth-like setulae along apical V2 of anteroventral surface; forefemur of male lacking comb-like row of short, stout, slightly flattened setae along apical V2 of anteroventral Genus Oedenops Becker surface; foretibia of male lacking several long, slender setae at Oedenops Becker, 1903:178 [type species: Oedenops isis Becker, 1903, mono- apex on ventral surface; forebasitarsus lacking row of long, typy].—Becker, 1926:94 [review].—Cresson, 1929:180 [assigned to subfam- slender, pale setulae inserted along anterior surface; midtibia ily Notiphilinae (=Hydrelliinae)]; 1947a:37, 46 [generic key; review, Neotro- with 2 dorsal, erect, extensor setae (subapically and subba- pical Region]; 1947b: 106 [generic key, Afrotropical Region].—Curran, sally); mid- and hindfemora normally developed, much shorter 1934:353 [key to nearctic genera].—Seguy, 1951:5 [generic key].—Wirth and Stone, 1956:465,470 [generic key, review, Californian species].—Wirth, than length of abdomen. 1965:748 [Nearctic catalog]; 1968:16 [Neotropical catalog].—Cole, Abdomen: Lacking fasciate pattern. Male terminalia: epan- 1969:395-396 [generic key, list].—Cogan, 1980:662 [Afrotropical catalog]; drium inverted U-shaped, width of arms wider than dorsum in 1984:148 [Palearctic catalog].—Mathis and Zatwamicki, 1995:117-118 posterior view; cercus oval to distinctly wider ventrally; [world catalog]. presurstyli almost touching or fused medially; postsurstylus an- DIAGNOSIS.—This genus is distinguished from other genera gulate anteriorly near middle, apex slightly spatulate, shallowly of Dryxini by the following combination of characters: arista bifurcate; aedeagus in lateral view with posterobasal, curved bearing 3-6 hairs; presutural supra-alar seta present (lacking in projection, usually slightly tapered toward apex, apical margin O. isis); notopleuron bearing 2 large setae (anterior notopleural sometimes uneven; aedeagal apodeme with relatively narrow seta well developed); anepisternal and anterior dorsocentral se- but long keel in lateral view; hypandrium shallowly curved in tae well developed (dorsocentral setae 1+3); katepistemal seta lateral view, relatively short, shallowly developed. absent; forefemur in both sexes bearing row of short, peg-like HISTORICAL REVIEW.—The nomenclatural history of the ge- setulae apically along anteroventral surface; and mid- and nus Oedenops is relatively recent and brief, as the genus was hindfemora normally developed, much shorter than abdomen. described early in this century and there are relatively few spe- DESCRIPTION.—Small to moderately small shore flies, body cies. Becker (1903) first described Oedenops and its then only length 1.70-2.65 mm; generally gray, often strikingly bicol- included species, O. isis, from specimens collected in Egypt. A ored, rusty yellowish on head. year earlier, however, Coquillett (1902) had described the first Head: Frons shallowly arched anteroventrally, not projected species now placed in Oedenops, O. nudus, but in the genus forward, sparsely setulose; ocelli in isosceles triangle, distance Paralimna. Coquillett's species was first collected in the state between posterior pair slightly longer than between anterior of Tabasco, Mexico, and has since been found to occur more ocellus and either posterior ocellus; ocellar seta, inner vertical widely in both North and South America. It remains the only seta, and reclinate fronto-orbital seta well developed; proclinate known species of Oedenops from the New World. Cresson fronto-orbital seta greatly reduced, frequently 1 or both lacking; (1929) recognized that Coquillett's species was closely related paravertical seta greatly reduced or absent. First flagellomere to O. isis and made the generic combination. Wirth (1956) de- broadly rounded; arista bearing 3-5 long hairs, mostly along scribed a second Old World species, O. afrus, from specimens basal V2 of dorsal surface. Face with 1-3 short, inclinate (but not collected in South Africa, and that same year Giordani Soika cruciate) facial setae on ventral Vi of face and 3 or 4 smaller set- (1956a) described O. aurantiacus from specimens collected in ulae interspersed along same vertical alignment as larger setae, Zaire. The latter species, however, is conspecific with O. isis facial setae and setulae vertically aligned, parallel with parafa- and is thus a junior synonym. Giordani Soika was apparently cial. Parafacial at anterior margin of eye narrow, width much unaware of the variability among specimens of O. isis, espe- less than length of 1st flagellomere. Gena high, height subequal cially the pronounced sexual dimorphism. This same confusing to combined length of 1st flagellomere and pedicel. variability may also have been a factor when Miyagi (1977) de- Thorax: Dorsocentral setae 4(1+3), well developed, pos- scribed a second junior synonym of O. isis as O. flavitarsis. Miyagi's species was based on specimens collected in Japan, terior pair displaced laterally; presutural supra-alar seta usually which is rather distant from Egypt. No other species have been well developed, subequal in length to anterior, dorsocentral described in the genus. seta (secondarily lost in O. isis and O. aurantiacus); acrostichal setulae poorly developed, inconspicuous, absent or, if present, Oedenops occurs worldwide in tropical zones and ranges in about 2 irregular rows; postpronotal seta well developed; no- into some temperate climates, such as the southwestern United 52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

States and into Japan. There are relatively few species in the Malaysia, Pakistan, Taiwan, Thailand, Vietnam. Palearctic: genus, but two of these, O. isis and O. nudus, are very wide- Egypt, Israel, Japan (Hokkaido, Honshu, Kyushu, Shikoku). spread, occurring on more than one continent. DISCUSSION.—Considerable variation is exhibited in speci- mens of Oedenops, especially the coloration of females, al- Nothing is known about the immature stages of Oedenops, though the differences expressed are not strictly sexually di- and judging from the few specimens that we have collected or morphic. The variation seems most evident on the face and that were made available to us, adults are seldom collected and mesonotum. The face of males tends to be more intensely col- are relatively scarce in collections and perhaps in nature. Foote ored, such as the yellowish orange to yellow coloration of O. (1995) did report specimens of Oedenops as regularly occur- isis. In females, often from the same locality, the face varies ring in stands of mangrove. from almost as intensely colored, like the males, to being much DISTRIBUTION (Figure 72).—Afrotropical: Cameroon, Ethio- paler and more subdued, frequently whitish gray. pia, Kenya, Madagascar (Toliara), Namibia, South Africa Synapomorphies that we have discovered and that establish (Cape Province, Natal), Zaire. Australasian/Oceanian: Austra- the monophyly of Oedenops are as follows: (1) only 3-6 ari- lia (Queensland). Nearctic: USA (AK, AZ, CA, FL, IA, IL, stal hairs; (2) setae on mesonotum generally reduced or absent; MO, MS, OK, TN, TX, UT). Neotropical: Brazil, Ecuador, (3) katepisternal seta absent; and (4) abdomen lacking fasciate Guatemala, Honduras, Mexico (Chiapas, Tabasco), Peru, Vene- pattern. zuela, West Indies (Barbados, , Dominican Republic, The origins of this genus were likely in the Old World, where Puerto Rico, St. Vincent). Oriental: China, India (Karnataka), most of the species diversity occurs.

Key to Species of Oedenops Becker 1. Presutural supra-alar seta absent; antenna pale yellow in both sexes (Afrotropical, Aus- tralasian/Oceanian, Oriental, Palearctic) 19. O. isis Becker Presutural supra-alar seta present; scape, pedicel, and 1st flagellomere (somewhat) brownish orange, dark gray, or black 2 2. Male frons and dorsal facial region reddish brown, gena golden; female frons, gena, and face grayish yellow; 2 hair-like setae in facial series. Midfemur of male bearing a few scattered setulae along posteroventral margin (Nearctic, Neotropical) 20. O. nudus Coquillett Male frons brown, face and gena mostly silvery white; female with face and gena, par- ticularly latter, silvery gray; 2 or 3 hair-like setae in facial series. Midfemur of male bearing distinct, comb-like row of numerous, short setae along posteroventral margin (Afrotropical) 18. O. afrus Wirth

FIGURE 72.—Distribution map for Oedenops. NUMBER 617 53

18. Oedenops afrus Wirth white to silvery gray; face and gena of female silvery gray. An- FIGURES 73-75 tenna dark gray to black. Presutural supra-alar seta present. Midfemur of male bearing comb-like row of numerous short Oedenops afra Wirth, 1956:387.—Cogan, 1968:355 [revision]. Oedenops afrus.—Cogan, 1980:662 [Afrotropical catalog].—Mathis and Zat- setae along posteroventral margin. warnicki, 1995:117 [world catalog]. Head: Frons of male brown with some faint blackish color- DIAGNOSIS.—This species is distinguished from congeners ation on parafrons, fronto-orbits gray to brownish gray. Scape by the following combination of characters: male frons brown, and pedicel gray, 1st flagellomere blackish gray; arista bearing face and gena mostly silvery white; female with face and gena, 4 or 5 dorsally branching hairs. Face bearing 2 or 3 setae in particularly latter, silvery gray; scape and pedicel and to a de- vertical series; face and gena of male mostly silvery white, face gree 1st flagellomere brownish orange, dark gray, or black; 2 or sometimes with faint yellowish to tannish coloration, espe- 3 hair-like setae in facial series; presutural supra-alar seta cially dorsally; face and gena of female silvery gray. Gena present; and midfemur of male bearing distinct, comb-like row high; gena-to-eye ratio 0.43-0.54. Maxillary palpus black with of numerous short setae along posteroventral margin. whitish gray microtomentum. DESCRIPTION.—Moderately small shore flies, body length Thorax: Mesonotum mostly grayish brown to brown, 2.10-2.35 mm; frons of male brown, face and gena silvery slightly darker through acrostichal tract and posterior portion

FIGURES 73-75.—Male terminalia of Oedenops afrus Wirth: 73, epandrium, cerci, and presurstyli, posterior aspect; 74, internal male terminalia, ventral aspect; 75, same, lateral aspect. 54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY of scutum, becoming more grayish brown to gray laterally; berg (lcf; NMSA). Natal: Ahrens, near Lilani, Apr 1962, pleural areas mostly gray. Presutural supra-alar seta present, B.R. and P. Stuckenberg (19; NMSA). subequal in length to anterior dorsocentral seta. Costal-vein ra- ZAIRE. Kivu: Kavimvira (Uvira; at light), Dec 1954, G. tio 0.33-0.35; M-vein ratio 0.80-0.86. Femora and tibiae gray; Marlier(19;MRAC). basal tarsomeres yellow, other tarsomeres becoming progres- DISTRIBUTION.—Afrotropical: Kenya, Namibia, South Af- sively darker, apical 1 or 2 tarsomeres brownish orange to rica (Cape Province, Natal), Zaire. brown; midfemur of male bearing comb-like row of numerous REMARKS.—This species, widespread in eastern and south- short setae along posteroventral margin. ern Africa, is the sister species to the other two species of Abdomen: Dorsum mostly gray, sometimes with some Oedenops. A synapomorphy is the fringe-like row of closely brownish coloration anteromedially. Male terminalia (Figures set setulae along the posteroventral margin of the male midfe- 73-75): epandrium broadly inverted U-shaped, lateral arms mur. There are no conspicuous color differences between the wider than dorsum in posterior view (Figure 73); cercus tear- sexes. drop-shaped, widened ventrally, narrowed dorsally, dorsal apex pointed in posterior view (Figure 73); presurstylus with long, 19. Oedenops isis Becker narrow, pointed projection medially, ventral margin pointed and FIGURES 76-78 slightly oriented medially, short; postsurstylus in lateral view Oedenops isis Becker, 1903:179; 1926:94 [review, list, Egypt].—Cresson, (Figure 75) with angulate, shallow projection medially along 1929:183 [synonymy with O. nudus]; 1947b: 108 [removed from synonymy anterior surface, apical portion clavate, very shallowly bifur- with O. nudus; list, Egypt (Island Elephantina), Sudan].—Wirth, 1956:388 cate; gonite long, narrow, sinuous in lateral view (Figure 75); [compared with O. afrus Wirth].—Cogan, 1968:355-356 [revision]; aedeagal apodeme with long, moderately wide keel, irregularly 1980:662 [Afrotropical catalog]; 1984:148 [Palearctic catalog].—Mathis and rounded in lateral view (Figure 75); aedeagus with posterobasal Zatwamicki, 1995:117-118 [world catalog]. Oedenops nudus of authors, not Coquillett [misidentification].—Cresson, projection wide, rounded apically, rest of aedeagus slightly 1929:183-184 [in part; discussion, synonymy of O. isis Becker with O. curved, width even, apex acutely pointed and projected; hypan- nudus (Coquillett)]. drium shallow and shallowly curved, anterior margin slightly Oedenops aurantiacus Giordani Soika, 1956a: 123.—Cogan, 1968:355 [revi- narrowed in ventral view, notched medially (Figure 74). sion]; 1980:662 [Afrotropical catalog].—Mathis and Zatwamicki, 1995:117 [world catalog]. [New synonym.] TYPE MATERIAL.—The holotype male of Oedenops afrus Oedenops flavitarsis Miyagi, 1977:46.—Cogan, 1984:148 [Palearctic cata- Wirth is labeled "Port ALFRED [handwritten] South Africa log].—Mathis and Zatwamicki, 1995:117 [world catalog]. [New synonym.] [Cape Province:] 17-3-1953 [17 Mar 1953; day, month, and DIAGNOSIS.—This species is distinguished from congeners by "53" handwritten][,] B.Stuckenberg/tf HOLOTYPE Oedenops the following combination of characters: antenna pale yellow in afra W.W.Wirth [red label; sex and species name handwrit- both sexes; presutural supra-alar seta absent; fore- and midlegs ten]." The holotype is double mounted (minuten in a long, rect- mostly yellow to yellowish orange; and midfemur of male bear- angular block of polyporus), is in good condition (the right ing sparse, short row of setulae along posteroventral margin. wing is missing), and is deposited in the NMNH (USNM DESCRIPTION.—Small to moderately small shore flies, body 62817). length 1.70-2.40 mm. OTHER SPECIMENS EXAMINED.—Afrotropical. KENYA. Lake Head: Antenna pale yellow to orange in both sexes. Gena- Victoria, Kisumu (10 km S), 19 Nov 1986, A. Freidberg (5cf, to-eye ratio 0.38-0.40. Maxillary palpus faintly grayish yellow 69; USNM). to yellow. NAMIBIA. Aus (3.2 km E), Plateau Farm, 14-17 Jan 1972 Thorax: Generally gray; occasionally with broad, yellow- (19; BMNH). Dwyka River, Merweville-Koup Road, 2 Jan ish brown to brown stripe between dorsocentral tracks of setae, 1972 (31cf, 329; BMNH). Fish River Canyon (27°37'S, more evident on posterior Vfe of scutum; anepisternum yellow- 17°36'E), 12-14 Jan 1972 (2d\ 29; BMNH). Helmeringhausen ish gray, this coloration extended onto immediately adjacent (25 mi W), Barby Farm, 17-18 Jan 1972 (19; BMNH). Ka- sclerites in some specimens. Supra-alar seta lacking. Costal- manjab (27 mi ESE), Otjitambi Farm, 13-15 Feb 1972 (29; vein ratio 0.25-0.27; M-vein ratio 0.75-0.80. Color of legs BMNH). Omaruru (53 km ENE), Otikoko Sud Farm, 10-13 variable and somewhat sexually dimorphic; fore- and midleg of Feb 1972 (ltf, 1 9; BMNH). Swakopmund, 26-30 Jan 1972 males and of some females mostly yellow to yellowish orange, (3d*, 149; BMNH). Windhoek, Hoffnung Farm (lakeside vege- only base of forefemur and apical tarsomere sometimes dark- tation), 7 Feb 1972 (1 d", 5 9; BMNH). ened, otherwise femora mostly gray, especially for females; SOUTH AFRICA. Cape Province: Brandkop Area, Calvinia hindfemur with basal Vfe or more gray to blackish gray, apex District, South-West Cape, 14 Oct 1964, B.R. and P. Stucken- yellowish; forefemur with row of short, stout, pale, tooth-like berg (19; NMSA); Dwyka River, Merweville-Koup Island, 2 setulae along anteroventral surface at apical Vfe; midfemur of Jan 1972 (Id1; BMNH); Port St. Johns, 20-25 Nov 1961, B.R. male bearing few scattered setulae along posteroventral margin. and P. Stuckenberg (39; BMNH, NMSA, USNM); Steytlerville Abdomen: Male terminalia (Figures 76-78): epandrium in (7 km N; Groot River; Lucerne field), 30 Oct 1978, R.M. posterior view inverted U-shaped (Figure 76), generally thick Miller (19; NMSA); Wit River Valley, Cambria Area (33°40'S, walled, each arm becoming wider ventrally; cercus hemispheri- 24°34'E), Patensie District, 6 Dec 1967, B.R. and P. Stucken- cal, bearing long setulae, ventral and dorsal apices about NUMBER 617 55

FIGURES 76-78.—Male terminalia of Oedenops isis Becker: 76, epandrium, cerci, and presurstyli, pos- terior aspect; 77, internal male terminalia, ventral aspect; 78, same, lateral aspect.

0.1mm

78

equally rounded; presurstylus in posterior view triangular (Fig- keel; hypandrium moderately concave, anterior margin broadly ure 76), bearing longer setulae along medial margin, somewhat rounded and shallowly pointed in ventral view (Figure 77). fused dorsomedially with ventral margin of cercus; postsursty- TYPE MATERIAL.—The lectotype male of Oedenops isis lus with anteroapical surface shallowly concave, posterior sur- Becker, here designated to stabilize and make more universal face nearly flat, apex capitate, very shallowly bifurcate; aedea- the use of this name, is labeled "[Egypt] Assuan 44554.1.[Jan] gus in lateral view (Figure 78) with relatively long [handwritten]/Typus [red label]/LECTOTYPE cf Oedenops isis posterodorsal projection narrowed and extended apically, rest Becker By Mathis and Zatwarnicki [handwritten except for of aedeagus wide basally, curved and becoming narrower poste- "LECTOTYPE" and "By"; label with black submarginal bor- riorly, apex shallowly concave; gonite rod-like, portion attached der]/Zool.Mus. Berlin [yellow label]." The lectotype is double to hypandrium slightly curved in lateral view (Figure 78); mounted (of the two minuten, the lectotype is mounted on the aedeagal apodeme more or less triangular, with narrow, long minuten nearer the main pin in a rectangular block of pith; the 56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY second, more outside specimen, a female, is here designated a PAKISTAN. Lahore, Jun-Aug 1957, J. Maldonado (lcf; paralectotype), is in good condition (some setae of the mesono- USNM). tum missing or displaced), and is deposited in the ZMHU. The THAILAND. Loei: Chiang Khan Mae Kong, 11 Feb 1999, P. original description (Becker, 1903:179) cited "Insel Philae bei Grootaert (19; IRSN); Na Haeo, 12 Feb 1999, P. Grootaert Assuan" as the type locality. In addition to the lectotype and fe- (39; IRSN). male paralectotype there are five additional female paralecto- TAIWAN. Anping, Sep 1908, H. Sauter (Id1; DEI). types, here designated, that have the same locality data and are VIETNAM. Prenn (15 km from Da Lat; ll°56isr, 108°25'E), deposited in the MNHN, USNM, and ZMHU. 15-19 Oct 1988, O. Mahunka, Vasarhelyi (Id1; HNHM). Ha- The lectotype male (not a female as indicated in the original noi, Song Hong (21°37'N, 104°44'E), 17 Jan 1986, O. publication) of Oedenops aurantiacus Giordani Soika, here des- Mahunka (4tf, 49; HNHM). ignated to stabilize and make more universal the use of this Palearctic. EGYPT. Asswan, Jan (Id1, 59; USNM, ZMHU; name, is labeled "HOLOTYPUS [light orange label with black lectotype and paralectotypes). Helwan, 31 Mar 1964, G. Stey- submarginal border]/COLL. MUS[EE DU]. CONGO Madagas- skal (19; USNM). Elephantine, Reimoser (ltf; ANSP). car: [Tamatave] Maroansetra[,] XII-1949 [Dec 1949][,] J. Va- ISRAEL. Ein Avdat, 29 Mar 1980, A. Freidberg, W.N. Mathis don/HOLOTYPUS Oedenops aurantiacus Soika [species name (29; USNM). and author handwritten]/LECTOTYPE Oedenops aurantiacus DISTRIBUTION.—Afrotropical: Cameroon, Ethiopia, Mada- G. Soika a" By Mathis & Zatwarnicki [all except "LECTO- gascar (Toliara), Namibia, Sudan. Australasian/Oceanian: Aus- TYPE" and "By" handwritten; label with black submarginal tralia (Queensland). Oriental: China, India (Karnataka), Malay- border]." The lectotype is double mounted (minuten in a long sia, Pakistan, Taiwan, Thailand, Vietnam. Palearctic: Egypt, rectangular block of polyporus), is in poor condition (left wing Israel, Japan (Hokkaido, Honshu, Kyushu, Shikoku). missing, right wing folded, most setae of the mesonotum miss- REMARKS.—Considerable variation is evident in facial and ing), and is deposited in the MRAC. Although the lectotype is thoracic coloration. In specimens from some localities, the col- labeled "HOLOTYPUS," Giordani Soika (1956a: 123) recog- oration appears to be sexually dimorphic, with the male face nized a type series of two "? 9," neither of which was designated being deeply yellowish and the female's grayish white to gray. as the holotype, and thus the need for this lectotype designation. At other localities, the female face is also yellowish, varying The holotype male of Oedenops flavitarsis Miyagi is la- from lightly grayish yellow to deeply yellow, quite similar to beled "Japonia Kyushu I. MIYAGI Is. T[s]ushima 6, Aug. that of males. Likewise, the brown stripe between the dorso- 1963/-type Oedenopus [sic] flavitarsis I. Miyagi [red label; all central setae varies from being restricted only to that area to except "-type" handwritten]." The holotype is double mounted having the brown coloration covering much of the mesonotum. (minuten in a rectangular card), is in poor condition (damaged In addition to variation in color, two other factors probably in shipment; wings on small rectangular card below specimen; contributed to the misidentifications and descriptions of the abdomen removed and dissected, with the parts in an attached two junior synonyms. The first was Giordani Soika's mistaken microvial), and is deposited in the HUS. determination of the sex of the lectotype, which is a male, not a OTHER SPECIMENS EXAMINED.—Afrotropical. CAMEROON. female. With some sexual dimorphism evident in this species, Kribi (beach; route N7), 28-29 Nov 1987, A. Freidberg (3cf; mistaking the sex of the type series would hinder the accurate USNM). identification of this species. Although the male of O. auranti- ETHIOPIA. Lake Langano, 13 Dec 1989, A. Freidberg, F. Ka- acus is deeply reddish brown, the structures of the male termi- plan (1

80

FIGURES 79-81 .—Oedenops nudus (Coquillett): 79, head, lateral aspect; 80, same, anterior aspect; 81, mesono- tum, dorsal aspect. Scale = 0.5 mm. 58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY supra-alar seta present, subequal in length to anterior dorsocen- M.R. Wheeler (Id1; USNM). Perry County: Lithium, 5 Sep tral seta. Costal-vein ratio 0.29-0.33; M-vein ratio 0.86-0.90. 1952, M.R. Wheeler (19; USNM). Oklahoma. Kingfisher Femora whitish gray to gray; tibiae mostly gray; basal tarsom- County: Kingfisher, 13 Sep 1952, M.R. Wheeler (Id1; eres yellow, other tarsomeres becoming progressively darker, USNM). Tennessee. Henry County: Paris Landing State Park, apical 1 or 2 brownish orange to brown; forefemur bearing 29 Jul 1954, M.R. Wheeler (19; USNM). Texas. Bastrop short row of closely set, tooth-like setulae anteroventrally County: Bastrop, 4 Nov 1951, M.R. Wheeler (19; USNM). along apical Vi; midfemur of male bearing few scattered setu- Culberson County: Van Horn, 14 Jun 1950, M.R. Wheeler lae along posteroventral margin. (2d", 49; USNM). Travis County: Austin, 14 Jun-27 Oct Abdomen: Dorsum mostly gray, sometimes with some 1901, 1950, M.R. Wheeler (6d\ 159; ANSP, USNM). Utah. brownish coloration anteromedially. Male terminalia (Figures Emery County: Green River, 19 Jul 1988, D. and W.N. 82-84): epandrium in posterior view (Figure 82) inverted U- Mathis(19;USNM). shaped, lateral arms wider than dorsal portion, ventral margin Neotropical. BARBADOS. St. Andrew: Long Pond broadly rounded; cercus in posterior view (Figure 82) ovate, (13°15.1X 59°33.3'W), 10-11 Sep 1996, W.N. Mathis (18d\ ventral margin more broadly rounded than dorsal margin; 49; USNM). 5/. Peter: Six Mens Bay (13°16.5'N, presurstyli nearly touching medially, triangular, slightly longer 59°38.8'W), 12 Sep 1996, W.N. Mathis (Id1; USNM). than wide; postsurstylus with anterior margin angulate medially, BRAZIL. Natal, 5-24 Feb 1943, F.M. Snyder (19; AMNH). dorsal extension narrowed to pointed apex, ventral extension DOMINICA. Layou River mouth, 9 Jan-24 Mar 1965, W.W. truncate, very shallowly bifurcate apically; gonite very narrow, Wirth(74d\439;USNM). with apices slightly curved; aedeagal apodeme triangular in lat- DOMINICAN REPUBLIC. La Vega: Constanza (-16 km SE; eral view (Figure 84), keel long and relatively narrow, exten- 18°50.2'N, 70°40.7'W; 1550 m), 27 Mar 1999, W.N. Mathis sions short; aedeagus with posterobasal projection narrowly de- (Id1; USNM); El Rio (9.5 km E; 19°0.9'N, 70°33.5'W; 980 m), veloped, curved posteriorly, acutely pointed, rest of aedeagus 6 May 1995, W.N. Mathis (12d\ 89; USNM); Rio Camu (3.5 nearly parallel sided, slightly curved, broadly rounded apically; km NW La Vega; 19°13.7'N, 70°35.2'W; 100 m), 10 May hypandrium with very irregular surface in lateral view (Figure 1995, W.N. Mathis (2tf, 29; USNM). Puerto Plata: Rio 84), V-shaped in ventral view, pointed anteriorly (Figure 83). Camu (14 km E Puerto Plata; 19° 11.9% 70°37.4'W), 17 May TYPE MATERIAL.—Holotype male Paralimna nuda Coquillett 1995, W.N. Mathis (2d\ 19; USNM). is labeled "Mexico [handwritten]/Frontera 3.9 Tab[asco]/Coll ECUADOR. Balao Guayas, Dec 1955, J.R. Levi-Castillo (2d\ Townsend/on moist sand at river edge [handwritten]/Type No 19; USNM). 6641 U.S.N.M. [red label; number handwritten]/Paralimna nuda GUATEMALA. Escuintla, Grutas de San Pedro Martir, 10 Aug Coq. [handwritten; label with black submargin]." The holotype 1965, P.J. Spangler (Id1; USNM). is double mounted (minuten in a rectangular block of pith), is in HONDURAS. Puerto Castilla, 28 May 1924, R.H. Painter (19; good condition, and is deposited in the NMNH (USNM 6641). ANSP). OTHER SPECIMENS EXAMINED.—Nearctic. UNITED STATES. MEXICO. Chiapas: Boca de Cielo (17 km S Puerto Arista), Arizona. Cochise County: Chiricahua Mountains, Martyr 18 May 1985, A. Freidberg, W.N. Mathis (Id1, 39; USNM). Dam, 11 Jun 1951, M.R. Wheeler (Id1, 2?; USNM); Fairbank, PERU. Madre de Dios: Manu, Rio Manu (near Romero), 8 13 Jun 1951, M.R. Wheeler (2d\ 29; USNM); San Pedro River Sep 1988, W.N. Mathis (6d\ 39; USNM). (12 km E Sierra Vista; Hwy 90; 124 m), 3 Jun 1991, J. Gelhaus, PUERTO Rico. Fajardo, Las Croabas (Seven Seas Beach; C. Nelson (4

FIGURES 82-84.—Male terminalia of Oedenops nudus (Coquillett): 82, epan- drium, cerci, and presurstyli, posterior aspect; 83, internal male terminalia. ven- tral aspect; 84, same, lateral aspect.

0.1MM

83 84

The obvious common feature to all habitats, fresh water or ma- [key, review of Afrotropical species]; 1948:6-12 [review of Australasian spe- rine, is a sandy substrate, often bare of conspicuous vegetation. cies, key].—Malloch, 1933:11 [review]; 1934b: 196-197 [generic key, Mar- REMARKS.—This is the only species of Oedenops to occur in quesas and Society Islands].—Curran, 1934:347 [key to Nearctic genera].— the New World, where the species is relatively widespread. Wirth and Stone, 1956:465, 470 [generic key, review, California species].— Wirth, 1965:748 [Nearctic catalog]; 1968:14-16 [Neotropical catalog].— Cole, 1969:392, 396 [generic key, review of species from western United Genus Paralimna Loew States].—Cogan and Wirth, 1977:333-335 [Oriental catalog].—Cogan, 1980:663-664 [Afrotropical catalog]; 1984:148 [Palearctic catalog].—Mathis Paralimna Loew, 1862a: 138 [type species: Paralimna appendiculata Loew, and Zatwamicki, 1995:118-127 [world catalog]. 1862 (=Notiphila punctipennis Wiedemann, 1830), monotypy]; 1862b: 13 [re- view, Afrotropical species].—Osten Sacken, 1878:201 [Nearctic catalog].— DIAGNOSIS.—This genus is distinguished from other genera Becker, 1896:114-115 [review, discussion of relationships].—Williston, of Dryxini by the following combination of characters: anterior 1908:308 [key to Nearctic genera].—Hendel, 1914:103-109 [review, Oriental species].—Cresson, 1916:101-124 [revision]; 1946:229 [key to Nearctic spe- proclinate fronto-orbital seta larger than posterior seta; arista cies]; 1947a:37,46 [key, review of Neotropical species]; 1947b:106, 108-118 bearing 8 or more dorsal hairs; facial seta usually 1, if more, 60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY not well separated; notopleuron bearing 2 large setae; anepis- apex on ventral surface; forebasitarsus lacking row of long, ternal and anterior dorsocentral (1+3) setae well developed; slender, pale setulae inserted along anterior surface; midtibia presutural supra-alar seta present (lacking in 1 species ofOede- with 3 dorsal extensor setae (subapically, subbasally, and near nops); katepisternal seta present; forefemur lacking row of basal Vb); mid- and hindfemora normally developed, much short, somewhat blunt spines along anteroventral surface; and shorter than abdomen. mid- and hindfemora normally developed, much shorter than Abdomen: Male terminalia: epandrium broadly and uni- length of abdomen. formly U-shaped in posterior view, lateral arms of similar DESCRIPTION.—Small to large shore flies, body length width; cercus ovoid with rounded apices; presurstylus bar-like, 1.85-7.80 mm. wide and short with extero-lateral appendage directed an- Head: Frons wider than high, shallowly arched anteroven- tero-internal or antero-extemal; presurstyli usually separated, trally, not projected forward, sparsely setulose; ocelli in equi- rarely fused to each other; postsurstylus in dorsal view about 5 lateral or isosceles triangle; pseudopostocellar setulae well de- times as long as wide, apex broadly bifurcate, internal lobe veloped, divergent; paravertical seta moderately well wide and directed ventrally; in lateral view postsurstylus dis- developed, less than Vz length of outer vertical seta; ocellar tinctly sinuous, ventral margin bearing long setae, terminal bi- seta, inner and outer vertical setae, and reclinate fronto-orbital furcation with 2 very long and 2 short setae inside; aedeagus seta well developed; proclinate fronto-orbital setae 2, both elongate, shape in dorsal view triangular or rhomboidal, mostly shorter than reclinate seta, moderately well developed, anterior with lateral appendices with length to % that of aedeagus; in seta about twice length of posterior seta. First flagellomere lateral view aedeagus irregular in shape, mostly tapered anteri- broadly rounded; arista bearing 5 or more long rays along dor- orly with sinuous margins, apex usually acutely pointed and sal surface. Face with 2 or 3 long (subequal to length of large projected; aedeagal apodeme in dorsal view with apex wide, in reclinate seta), inclinate (but not cruciate) facial setae on ven- lateral view triangular; hypandrium in dorsal view broadly tral 1/j of face and with 3 or 4 smaller setulae interspersed along oval, posterior margin incised to about Vb length of hypan- same vertical series as larger setae. Parafacial at anterior mar- drium; in lateral view moderately deep to very deep (height gin of eye narrow, width much less than length of 1 st flagello- equal to length). mere. Gena high, height subequal to combined length of 1st DISTRIBUTION.—The genus Paralimna occurs pantropically, flagellomere and pedicel; genal seta well developed. with a few species extending into temperate areas (Egypt, Ja- Thorax: Mesonotum more or less uniformly setulose; pan, northern Australia, North America). acrostichal setulae in 6-8 irregular rows; 1 pair of large, pre- DISCUSSION.—Paralimna, with 85 species (Mathis and Zat- scutellar, acrostichal setae; dorsocentral setae 4(1+3), well de- warnicki, 1995), includes most of the species diversity of veloped, posterior pair displaced laterally; acrostichal setulae Dryxini, and we know that there are numerous undescribed well developed, conspicuous, in about 6 irregular rows; presu- species in collections and, undoubtedly, undescribed species tural seta well developed, subequal in length to anterior dorso- that await collection. For example, in a faunistic work on the central seta; supra-alar setae 2(1 + 1), presutural seta inserted shore flies of the West Indies, Mathis (in prep.) is describing more mediae! than postsutural seta; postpronotal seta well de- seven new species, which comprise over one-half of the known veloped; notopleural setae 2, posterior seta shorter; anepister- fauna of Paralimna from these islands. num bearing 2 large setae along posterior margin, ventral seta The species diversity of Paralimna is matched by its struc- greatly elongate; katepisternal seta well developed. Posterior tural diversity, and the genus is difficult to characterize. The margin of scutellum truncate, nearly flat. Veins lacking setulae; only synapomorphy we have discovered that establishes the costa long, extended to vein M; crossvein dm-cu regularly de- monophyly of the genus is the lateral aedeagal processes of the veloped, nearly straight, longer than apical section of vein aedeagus. CuAi, and at distinct angle with posterior margin of wing. Herein we revise at the species level only the subgenus Mid- and hindfemora normally developed, not conspicuously Phaiosterna, but we also review the species of the limbata elongate; fore tibia of male lacking several long, slender setae at group.

Key to Subgenera of Paralimna Loew 1. Gena short, height about VA eye height; 1st flagellomere elongate, length more than twice width 2 Gena high, height about Vb eye height or more; 1 st flagellomere shorter than above, length at most 1.5 times width Paralimna Loew, in part 2. Mesonotum (brown medially and through dorsocentral track, yellowish to brownish gray between) and anepisternum (brown dorsally and ventrally, gray between) con- spicuously vittate; crossvein dm-cu and sometimes adjacent veins with halo of brown (the limbata group) Paralimna Loew, in part Mesonotum and anepisternum mostly unicolorous and grayish brown to black; cross- vein dm-cu and rest of wing essentially unicolorous Phaiosterna Cresson NUMBER 617 61

Subgenus Paralimna Loew ally and through dorsocentral track, yellowish to brownish gray between) and anepisternum (brown dorsally and ventrally, gray Paralimna Loew, 1862a: 138 [type species: Paralimna appendiculata Loew, 1862 (=Notiphila punctipennis Wiedemann, 1830), monotypy].—Cresson, between) conspicuously vittate; and crossvein dm-cu and some- 1947a:47-54 [diagnosis, review ofNeotropical species]; 1947b: 108—118 [re- times adjacent veins with halo of brown (the limbata group). view of Afrotropical species, key]; 1948:6-9, 11-12 [review of Afrotropical DESCRIPTION.—Small to large shore flies, body length species, key].—Wirth, 1965:748 [Nearctic catalog]; 1968:14-15 [Neotropi- 1.85-6.30 mm. cal catalog].—Cogan, 1980:663-664 [Afrotropical catalog].—Mathis and Head (Figures 85-89): Ocelli in equilateral triangle; Zatwarnicki, 1995:118-126 [world catalog]. pseudopostocellar setulae well developed, divergent; paraverti- Poecilothorax Becker, 1922:73 [as a genus; type species: Poecilothorax angus- cal seta well developed, subequal in length to anterior procli- tus Becker, 1922 (=Paralimna puncticollis Becker, 1922), monotypy].— nate fronto-orbital seta; ocellar seta well developed, usually Cresson, 1947b: 108 [synonymy]. longer than anterior fronto-orbital seta and subequal to outer DIAGNOSIS.—This subgenus is distinguished from other con- vertical seta, setae well separated, inserted anterolaterad of an- generic subgenera by the following combination of characters: terior ocellus. First flagellomere usually gradually narrowed to- gena short, height about VA eye height; 1 st flagellomere elon- ward apex; length about twice width; arista bearing 9-16 long gate, length more than twice width; mesonotum (brown medi- hairs along dorsal surface. Face usually conspicuously convex,

FIGURES 85-90.—Scanning electron micrographs of Paralimna (Paralimna) fellerae (Belize. St arm Creek Dis- trict: Glover's Reef; scale length in parenthesis; scale for all photographs = Figure 87): 85, head, lateral aspect (0.43 mm); 86, face, anterior aspect (0.30 mm); 87, antenna, lateral aspect (150 urn); 88, ommatidia and occa- sional interfacetal setae of compound eye, lateral aspect (30 urn); 89, enlargement of same, lateral aspect (6.0 urn); 90, mesonotum, dorsal aspect (0.50 mm). 62 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY comparatively well arched, usually bearing 1 (occasionally 2 or face; foretibia of male lacking several long, slender setae at 3) long (subequal to length of large, reclinate seta), inclinate, apex on ventral surface; forebasitarsus lacking row of long, but not cruciate facial seta on ventral V2 of face, if more than 1, slender, pale setulae inserted along anterior surface; midtibia setae approximate; usually 3 or 4 smaller setulae interspersed with 3 dorsal extensor setae (subapically, subbasally, and near along same vertical series as larger setae. Eye with oblique ori- basal Vb). entation. Gena comparatively high, height slightly more than Abdomen (Figures 97-99): Frequently with fasciate pattern Va eye height. with darker band basally. Male abdominal sternites 3 and 4 Thorax (Figures 90-96): Acrostichal setulae well devel- rectangular, longer than wide; sternite 5 narrow basally, be- oped, conspicuous, in about 6 irregular rows; presutural supra- coming wider posteriorly, posterior margin shallowly concave. alar seta well developed, subequal in length to anterior dorso- Male terminalia (Figures 97-99): presurstyli L-shaped, medial central seta. Veins lacking setulae; crossvein dm-cu regularly arms from each side connected just below cerci, ventral arm re- developed, nearly straight, longer than apical section of vein duced, shorter than cercus (Figures 97-99); lateral aedeagal CuAj, and at distinct angle with posterior margin of wing. process enlarged; aedeagal apodeme broad basally and api- Forefemur of male lacking comb-like row of short, stout, cally; hypandrium forming a relatively deep, conically shaped slightly flattened setae along apical Vfe of anteroventral sur- pouch (Figures 98, 99).

FIGURES 91-96.—Scanning electron micrographs of Paralimna (Paralimna) fellerae (Belize. Stann Creek Dis- trict: Glover's Reef; scale length in parenthesis; scale for all photographs • Figure 94): 91, scutellum, dorsal aspect (231 pm); 92, pleural sclerites, lateral aspects (0.50 mm); 93, notopleuron, lateral aspect (167 pm); 94, left foreleg, anterior aspect (250 urn); 95, enlargement of same, anterior aspect (86 jim); 96, left foreleg, poste- rior aspect (0.27 mm). NUMBER 617 63

FIGURES 97-99— Male terminalia of Paralimna (Paralimna) punctipennis (Wiedemann): 97, epandrium, cerci, and presurstyli, posterior aspect; 98, internal male terminalia. ventral aspect; 99, same, lateral aspect.

T 0.1mm 1

98

DISTRIBUTION.—As for the genus. The limbata Group DISCUSSION.—There are now nearly 80 species in this sub- genus, and many additional species remain to be described, DIAGNOSIS.—The limbata group is distinguished from other especially species from the New World. Except for the lim- species of Paralimna, especially species of the subgenus bata group, which had been placed in the subgenus Phaiosterna, by the following combination of characters: gena Phaiosterna (Mathis and Zatwarnicki, 1995), we have not short, height about V* eye height; 1st flagellomere elongate, characterized any of the sublineages within this subgenus. length more than twice height, broadly rounded apically; arista Mathis is now revising the New World species of Paralimna, bearing 12-14 long dorsal rays; mesonotum vittate, broad and that revision will include phylogenetic and descriptive brown stripe medially and through dorsocentral track, yellow- sections that will address these sublineages, as they are ger- ish to brownish gray between brown stripes; anepisternum mane to that fauna. brown dorsally and ventrally, gray between, conspicuously vit- 64 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY tate; wing with crossvein dm-cu and sometimes adjacent veins DISCUSSION.—We have reviewed this species group because surrounded with halo of brown; forefemur bearing comb-like Cogan (1968) recognized the included species to be related to row of short, stout, spine-like setulae along apical V2 of an- if not included in the subgenus Phaiosterna (although Cogan teroventral surface. Male terminalia: epandrium as wide as did not recognize Phaiosterna as a subgenus of Paralimna, he high; cercus well developed, bearing numerous setulae; presur- identified the included the species as a species group in which stylus quite variable, depending on species; postsurstylus he placed the species of Phaiosterna as well as the two we re- somewhat rectangular, deeply bifurcate apically, bearing some- view herein). Here we restrict the characterization of the spe- what vertical row of long setulae; aedeagal apodeme with cies group to include only two Afrotropical species, P. limbata prominent keel, irregularly rectangular to slightly rounded; Loew and P. reticulata Cogan. Although both of these species aedeagus in lateral view with moderately deep notch dorsoba- superficially resemble species of Phaiosterna, the similarities sally, broadly developed over basal %, apex pointed; hypan- probably arose through convergence. drium wider than long, moderately to shallowly depressed. The adults of the limbata group are similar externally, partic- DISTRIBUTION.—Afrotropical: Angola, Cameroon, Ethiopia, ularly in overall shape, but some structures of the male termi- Gambia, Ghana, Kenya, Liberia, Madagascar, Malawi, nalia are markedly different, especially the presurstylus (see Mozambique, Nigeria, Rwanda, Senegal, Sierra Leone, South species descriptions). Other structures of the male terminalia, Africa (Cape Province), Sudan, Tanzania, Uganda, Zaire, Zam- however, such as the postsurstylus, aedeagal apodeme, aedea- bia, Zimbabwe. gus, and gonite, are very similar.

Key to Species of the limbata Group Frons with dark brown medial and orbital stripes but lacking velvety black, microto- mentose squarish area on fronto-orbit; frons lacking black spots adjacent to ocelli; pedicel and 1st flagellomere blackish brown. Wing lacking reticulate pattern in membrane, but crossvein dm-cu with dark brown halo 21. P. limbata Loew Frons with squarish spot at base of reclinate fronto-orbital seta densely microtomen- tose, appearing velvety black; frons with black spots (sometimes slightly shiny) an- terior of anterior ocellus and laterad of posterior ocelli; pedicel tan to grayish yel- low; 1st flagellomere with dorsal V2 dark brown, ventral V2 yellowish. Wing conspicuously reticulate 22. P. reticulata Cogan

21. Paralimna (Paralimna) limbata Loew setulae, especially dorsomedially (Figure 100); presurstylus in posterior view a shallowly curved bar, much wider than high, FIGURES 100-102 bearing few tiny setulae along posterior margin (Figure 100); Paralimna limbata Loew, 1862b: 13.—Becker, 1922:72 [list from Sudan (Renk, Tonga)].—Cresson, 1929:188 [discussion].—Giordani Soika, postsurstylus somewhat rectangular in lateral view (Figure 1956b:495 [list, Zaire].—Cogan, 1968:326 [revision]; 1980:663 [Afrotropi- 102), apex broadly bifurcate, bearing vertical row of setulae, cal catalog].—Canzoneri and Meneghini, 1969:150-151 [list, Zaire].—Can- apical setulae longer; aedeagus in lateral view (Figure 102) zoneri, 1982:57 [list, Sierra Leone]; 1986:68 [list, Sierra Leone]; 1987:83 with moderately deep notch dorsobasally, broadly developed [list, Sudan].—Canzoneri and RarTone, 1987:64 [list, Kenya].—Canzoneri 3 and Rampini, 1990:109 [list, Sierra Leone]; 1994:248 [list, Sierra Leone]. over basal /», apex pointed; aedeagal apodeme with wide, ir- Paralimna (Phaiosterna) limbata.—Cresson, 1947b: 115-116 [review; list, regularly rounded, moderately deep keel, extended process to Zaire].—Mathis and Zatwamicki, 1995:126-127 [world catalog]. base of aedeagus wider; gonite in lateral view (Figure 102) a shallowly curved, small bar, slightly wider toward base of DIAGNOSIS.—This species is distinguished from congeners by the frons having a dark medial and orbital stripe but lacking a postsurstylus; hypandrium wider than long (Figure 101), shal- velvety black, microtomentose squarish area on the fronto-orbit lowly depressed, in lateral view moderately curved (Figure and by the wing lacking a maculation pattern in the membrane. 102). DESCRIPTION.—Head: Frons with dark brown medial and TYPE MATERIAL.—The lectotype male, here designated to orbital stripe but lacking black spots. Antenna with pedicel and stabilize and make more universal the use of this name, is la- 1 st flagellomere blackish brown. beled "[South Africa] 450 [handwritten]/330 [handwritten]/ Thorax: Scutum with dark medial stripe. Wing with cross- Paralymna [sic] limbata/248 66 [pink label]/Riksmuseum vein dm-cu surrounded by dark brown halo. Legs, including Stockholm [light green label]/LECTOTYPE Paralimna limbata basitarsi, black. Loew c? By Mathis & Zatwarnicki [all except "LECTOTYPE" Abdomen: Male terminalia (Figures 100-102): epan- and "By" handwritten; label with black submarginal border]." drium about as wide or wider than high in posterior view, ex- The lectotype is pinned directly, is in poor condition (wings tended arms much wider than dorsum (Figure 100); cercus broken off, antennae and most setae missing), and is deposited wider dorsally, 3-4 times higher than wide, bearing numerous in the NRS. NUMBER 617 65

FIGURES 100-102.—Male terminalia of Paralimna (Paral- imna) limbata Loew: 100, epandrium, cerci, and presurstyli, posterior aspect; 101, internal male terminalia, ventral aspect; 102, same, lateral aspect.

T 0. 1mi 1

102

OTHER SPECIMENS EXAMINED.—Afrotropical. CAMEROON. DISTRIBUTION.—Afrotropical: Angola, Cameroon, Ethiopia, Bambalang (off route Nil; 35 km E Bamenda; 1200 m), 18 Gambia, Ghana, Kenya, Liberia, Malawi, Mozambique, Nige- Nov 1987, A. Freidberg, F. Kaplan (3c?; USNM). Limbe ria, Rwanda, Senegal, Sierra Leone, South Africa (Cape Prov- (shore), 14-15 Nov 1987, A. Freidberg (ld\ 2?; USNM). ince), Sudan, Tanzania, Uganda, Zaire, Zambia, Zimbabwe. ETHIOPIA. Gila River, Mission Station, Sep 1963, M.L. Schmidt (19; USNM). 22. Paralimna (Paralimna) reticulata Cogan KENYA. Lake Bugoria, 26 Aug 1983, A. Freidberg (3d1; USNM). Marmar Springs (S of Maralal), 28 Nov 1986, A. FIGURES 103-105 Freidberg (Id1; USNM). Masai Mara, Keekoro Lodge, 29-30 Paralimna (Paralimna) reticulata Cogan, 1968:327; 1980:664 [Afrotropical Aug 1983, A. Freidberg (Id1; USNM). catalog]. LIBERIA. Grand Cape Mount: Dia, 17 Feb 1953, O. Blicken- Paralimna (Phaiosterna) reticulata.—Math is and Zatwamicki, 1995:127 [world catalog]. staff(2d\39;USNM). MALAWI. Lake Malombe, Nkungulu, 25 Oct 1983, A. Freid- DIAGNOSIS.—This species is distinguished from congeners berg (13 d\ 27 ?; USNM). by the frons having a dark medial and orbital stripe but lacking a MOZAMBIQUE. Luabo, Jan 1956, P. Usher (1 ?; USNM). velvety black, microtomentose squarish area on the fronto-orbit TANZANIA. Mount Wa Mbu (near Lake Manyara Swamp; and by the wing lacking a maculation pattern in the membrane. 900 m), 6 Sep 1992, A. Freidberg (Id1, 39; USNM). Ngoron- DESCRIPTION.—Head: Frons with dark brown medial and goro Reserve, Gate (1500 m), 6 Sep 1992, A. Freidberg (2cf; orbital stripe but lacking black spots. Antenna with pedicel and USNM). 1st flagellomere blackish brown. 66 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Thorax: Scutum with dark medial stripe. Wing with cross- lateral view (Figure 105), apex broadly bifurcate, bearing verti- vein dm-cu surrounded by dark brown halo. Legs, including cal row of setulae, apical setulae longer; aedeagus in lateral view basitarsi, black. (Figure 105) with moderately deep notch dorsobasally, broadly Abdomen: Male terminalia (Figures 103-105): epandrium developed over basal 3A, apex pointed; aedeagal apodeme with about as wide as high in posterior view, extended arms much wide, irregularly rectangular, moderately deep keel, extended wider than more narrowly rounded dorsum (Figure 103); cercus process to base of aedeagus wider and slightly longer; gonite in wide ventrally, 3 times higher than wide, bearing numerous long lateral view (Figure 105) a shallowly curved, small bar; hypan- setulae, especially medially (Figure 103); presurstylus in poste- drium wider than long (Figure 104), shallowly to moderately de- rior view (Figure 103) elongate, length 2-3 times width, pointed pressed, in lateral view moderately curved (Figure 105). apically, medial margin more conspicuously curved, bearing few TYPE MATERIAL.—The holotype male of Paralimna reticu- setulae along curvature; postsurstylus somewhat rectangular in lata Cogan is labeled "holo- type [red margin]/Madagascar-

FlGURES 103-105.—Male terminalia of Paralimna (Paralimna) reticulata Cogan: 103, epandrium, cerci, and presurstyli, poste- rior aspect; 104, internal male terminalia, ventral aspect; 105, same, lateral aspect.

0.1mm 1

104 105 NUMBER 617 67

Ouest [Toliara:] Ranohira[,] 860m[,] 26.1^.11.58 [26 Jan-4 lacking several long, slender setae at apex on ventral surface; Feb 1958][,] B.Stuckenberg/Paralimna reticulata sp. n. det. forebasitarsus lacking row of long, slender, pale setulae in- B.H.Cogan 1966 ["reticulata sp. n." and last number of year serted along anterior surface; midtibia with 3 dorsal extensor handwritten]." The holotype is double mounted (minuten in a setae (subapically, subbasally, and near basal Vb). block of plastic foam), is in excellent condition, and is depos- Abdomen: Frequently with fasciate pattern with darker ited in the NMSA. band basally. Male abdominal sternites 3 and 4 rectangular, OTHER SPECIMENS EXAMINED.—Afrotropical. MADAGAS- longer than wide; sternite 5 narrow basally, becoming wider CAR. Antsiranana: Sambava (30 km W), 13 Apr 1991, A. posteriorly, posterior margin shallowly concave. Male termi- Freidberg, F. Kaplan (3cr, 2 ?; USNM). Toliara: Fort Dauphin nalia: presurstylus L-shaped, medial arms from each side con- (50 km W), 22 Apr 1991, A. Freidberg, F. Kaplan (Id1; USNM). nected just below cerci, ventral arm reduced, length less than DISTRIBUTION.—Afrotropical: Madagascar. cerci; lateral aedeagal process enlarged; aedeagus pointed api- REMARKS.—The reticulate wing of this species is unmistak- cally (Figures 107-112); aedeagal apodeme broad basally and able and facilitates identification. Like the wing, the presursty- apically; hypandrium forming relatively deep, conically lus is markedly different, being two to three times longer than shaped pouch. wide. In P. limbata, the presurstylus is much wider than long DISTRIBUTION (Figure 106).—Afrotropical: Aldabra, Bu- and is bar-like. rundi, Cameroon, Ethiopia, Gambia, Ghana, Kenya, Madagas- car, Namibia, Nigeria, Seychelles (Cousin, La Digue, Mahe, Subgenus Phaiosterna Cresson Praslin), Sudan, Tanzania, Uganda, Yemen, Zaire. Australa- sian/Oceanian: American Samoa, Australia (Coral Sea Islands Phaiosterna Cresson, 1916:104 [type species: Paralimna decipiens Loew, Territory, Northern Territory, Queensland, Western Australia), 1878, original designation].—Cresson, 1947a:54-55 [review of Neotropical Christmas Island, Federated States of Micronesia, Fiji, Guam, species]; 1947b: 115-117 [review of Afrotropical species, key]; 1948:9—11 [review of Australasian species, key].—Wirth, 1965:748 [Nearctic catalog]; Kiribati (Gilbert Islands, Phoenix Islands), Marquesas (Fatu 1968:16 [Neotropical catalog].—Cogan and Wirth, 1977:334 [synonymy Hiva, Nuku Hiva), Northern Marianas, Palau, Papua New with Paralimna].—Cogan, 1980:663-664 [Afrotropical catalog, as the sub- Guinea, Society Islands (Moorea, Raiatea, Tahiti), Solomon genus Paralimna in part].—Mathis and Zatwarnicki, 1995:126-127 [world Islands, Vanuatu, Western Samoa. Oriental: India (Andhra catalog]. Pradesh, Assam, Kerala, Tamil Nadu, West Bengal), Indone- DIAGNOSIS.—This genus is distinguished from other conge- sia (Java, Lesser Sunda Islands, Sumatra), Laos, Malaysia, neric subgenera by the following combination of characters: Nepal, Pakistan, Philippines (Luzon, Panay, Visayan), Sri gena moderately high to high, gena-to-eye ratio 0.19-0.40; 1st Lanka, Taiwan, Thailand, Vietnam. Nearctic: Bermuda, USA flagellomere short, length at most 1.5 times width; mesono- (AK, AZ, CA, FL, GA, LA, MS, NJ, TX). Neotropical: Ar- tum and anepisternum mostly unicolorous and grayish brown gentina, Bahamas, Belize, Bolivia, Brazil (Mato Grosso, Rio to black; and crossvein and veins and rest of wing essentially de Janeiro), Colombia, Costa Rica, Ecuador, Guatemala, Guy- unicolorous. ana, Honduras, Mexico (Jalisco, San Luis Potosi, Sonora, DESCRIPTION.—Small to moderately sized shore flies, body Tamaulipas, Veracruz), Panama, Paraguay, Trinidad and To- length 1.85-3.85 mm; chaetotaxy generally well developed. bago, Venezuela, West Indies (Barbados, Cuba, Dominica, Head: Ocelli in equilateral triangle; pseudopostocellar set- Dominican Republic, Grand Cayman, , Jamaica, Pu- ulae well developed, divergent; paravertical seta well devel- erto Rico, St. Lucia, St. Thomas, St. Vincent, Virgin Islands). oped, subequal in length to anterior proclinate fronto-orbital Palearctic: Egypt. seta; ocellar seta well developed. Antenna black; 1 st flagello- DISCUSSION.—Cresson described Phaiosterna as a subgenus mere broadly rounded, length about twice width; arista bearing of Paralimna, and we have continued to recognize this group 6-8 long hairs along dorsal surface. Face only slightly convex, as a subgenus. Certainly the group is monophyletic, as evi- comparatively flattened, bearing 2 or 3 long (subequal to length denced by numerous synapomorphies. The sister group of of large, reclinate seta), inclinate (but not cruciate) facial setae Phaiosterna is not as well established, although there is some on ventral Vfe of face, and 3 or 4 smaller setulae interspersed external resemblance with species of the limbata group. along same vertical series as larger setae. Eye with vertical ori- The species of Phaiosterna are very similar externally, and entation. Gena comparatively short, height slightly more than there is considerable variation and overlap in such features as width of 1st flagellomere. the amount and pattern of microtomentum, for example. From Thorax: Acrostichal setulae well developed, conspicuous, localities where more than one species occurs, however, the in about 6 irregular rows; presutural supra-alar seta well devel- differences between species seem to be accentuated, perhaps as oped, subequal in length to anterior, dorsocentral seta. Veins a result of character displacement. Otherwise, especially in lacking setulae; crossvein dm-cu regularly developed, nearly populations that are allopatric, the only reliable characters are straight, longer than apical section of vein CuAj, and at distinct those of the male terminalia. Exceptions to this generality are angle with posterior margin of wing. Legs black; forefemur of the two new species. For example, the new species from the In- male lacking comb-like row of short, stout, slightly flattened dian subcontinent has a shiny, mostly bare mesonotum that setae along apical Vz of anteroventral surface; foretibia of male consistently differs from P. lineata, which is more densely mi- 68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 106.—Distribution map for Paralimna (Phaiosterna).

crotomentose. Males of the new species from tropical America, more uniformly dark colored (there is still some distinctly bi- P. longiseta, bear long setulae on the foreleg, a character that colored areas, especially the scutum, but the differences are not consistently distinguished them from congeners. Otherwise, as pronounced as in typical Paralimna); (2) gena secondarily however, accurate identification will frequently require exami- shorter; (3) 1st flagellomere long, length about twice height; nation of structures from the male terminalia. (4) presurstylus reduced; (5) lateral aedeagal process enlarged; The synapomorphies that we have discovered that confirm and (6) hypandrium a relatively deep, conically shaped pocket the monophyly of Phaiosterna are as follows: (1) generally or pouch.

Key to Species of Phaiosterna Cresson New World Species 1. Ventral apex of male foretibia bearing numerous long, slender setulae; male forebasi- tarsus bearing long setulae along posterior surface (Neotropical) 27. P. longiseta, new species Foreleg of male lacking long, slender setulae 2 2. Abdomen subshiny to shiny, sparsely microtomentose; aedeagus in posterior view higher than wide, narrow (Nearctic, Neotropical) 28. P. obscura Williston Abdomen appearing dull, microtomentose, tannish gray; aedeagus in posterior view wider than high; wide (Nearctic, Neotropical) 24. P. decipiens Loew

Old World Species 1. First flagellomere bearing long fringe of whitish setulae along dorsum and dorsal por- tion of apex, length of setulae greater than Vz height of 1st flagellomere. Mesono- tum unicolorous, blackish brown, sparsely microtomentose, subshiny to shiny (Aus- tralasian/Oceanian, Oriental) 25. P.fusca Bock First flagellomere either lacking long fringe of setulae or setulae short, considerably less than V2 height of flagellomere. Mesonotum usually with some evidence of stripes, varicolored, grayish tan to dark brown, densely microtomentose, at most slightly subshiny 2 2. Face grayish yellow, mostly uniform in color, generally lighter-colored species, gray to tan, but frequently with dark brown stripes between dorsocentral setae (Afrotropi- cal, Australasian/Oceanian, Palearctic) 23. P. bicolor (Macquart) Face generally dark brown with gray to yellowish gray areas; usually darker-colored species, pleural regions dark brown (Australasian/Oceanian, Oriental) 26. P. lineata de Meijere NUMBER 617 69

DIAGNOSIS.—This species is distinguished from congeners, especially those of the subgenus Phaiosterna, by the following combination of characters: first flagellomere lacking fringe of long, whitish setulae along dorsum and dorsal portion of rounded apex; apex of 1 st flagellomere evenly rounded; and mesonotum black, dull to subshiny, with moderately dense gray to brown microtomentum, frequently with conspicuous, longitudinal stripes medially. DESCRIPTION.—Small to moderately small shore flies, body length 1.95-2.90 mm; generally dark, whitish gray to brown species, moderately densely to densely microtomentose, dor- sum appearing dull to slightly subshiny, especially mesonotum. Head: Generally grayish brown. First flagellomere bearing long fringe of whitish setulae along dorsum and dorsal portion of apex, setulae longer than Vfe height of 1st flagellomere. Face uniformly colored, grayish yellow, subdued, lacking lighter colored areas on darker background. Gena-to-eye ratio 0.21-0.25. Thorax: Generally whitish gray to brown; mesonotum moderately densely gray to brown microtomentose, appearing dull to very slightly subshiny, usually with conspicuous, darker brown, longitudinal stripes medially, between dorsocentral se- tae. Costal-vein ratio 0.48-0.50; M-vein ratio 0.82-0.89. Fore- leg of male lacking long, slender setulae. Abdomen: Slightly lighter in color than mesonotum, mostly gray; anterior portion of tergites fasciate, darker gray to brown, less microtomentose. Male terminalia (Figures 112-115): epandrium in posterior view inverted U-shaped 111 112 (Figure 113), moderately wide, dorsally slightly narrower than width of lateral arms and with dorsal margin of cereal cavity narrowly rounded; cercus broadly pointed dorsally and nar- FIGURES 107-112.—Ventral view of the aedeagus: 107, Paralimna (Phaio- rowly projected and pointed ventromedially, medial margin sterna) lineala de Meijere; 108, P. (P.) longiseta, new species; 109, P. (P.) concave in posterior view (Figure 113); presurstyli narrowly obscura Williston; 110, P. (P.) decipiens Loew; III, P. (P.)fusca Bock; 112, P. connected medially, forming nearly straight, long band at ven- (P.) bicolor (Macquart). tral margin of cerci, produced ventrolaterally as wide, triangu- lar-shaped projections; postsurstylus in lateral view (Figure 23. Paralimna (Phaiosterna) bicolor (Macquart) 115) with anterior margin wider and swollen medially, bearing several setulae medially, bifurcate apically, anterior lobe longer FIGURES 112-115 and only slightly narrower, posterior lobe in lateral view Ephydra bicolor Macquart, 1851:276. broadly rounded, truncate apically; aedeagal apodeme with Paralimna bicolor.—Wirth, 1975:40-41 [generic combination, lectotype des- keel asymmetrical, more angulate toward hypandrium; aedea- ignation]. gus in posterior view with base subrectangular, apex broadly Paralimna lineata of authors, not de Meijere [misidentification].—Lamb, rounded to truncate medially, in lateral view (Figure 115) nar- 1912:317 [list, Seychelles].—Cresson, 1948:10 [in part; listed as a species in- rowly triangular and with pointed lobe near middle along pos- certae].—Cogan, 1980:663 [Afrotropical catalog].—Canzoneri, 1987:83 terior surface, becoming wider subapically, thereafter narrowed [list, Sudan].—Canzoneri and Raffone, 1987:64 [list, Kenya].—Canzoneri and Rampini, 1990:109 [list, Sierra Leone]; 1994:248 [list, Sierra Leone]. to narrow extension, acutely pointed apically; lateral aedeagal Paralimna (Phaiosterna) lineata in part of authors, not de Meijere [misidentifi- processes robust (Figure 114), parallel sided throughout length, cation].—Mathis and Zatwamicki, 1995:127 [world catalog]. narrowly rounded apically; hypandrium deeply invaginated, Paralimna decipiens of authors, not Loew [misidentification].—Becker, pocket-like (Figures 114, 115). 1903:168-169 [list, Egypt]; 1926:16 [list, Egypt]. TYPE MATERIAL.—The lectotype male (not a female as Phaiosterna (Paralimna) aequalis of authors, not Cresson [misidentifica- stated by Wirth) of Ephydra bicolor Macquart (designated by tion].—Cogan, 1968:325 [revision, Afrotropical Region]. Wirth, 1975:40) is labeled "Holo-type [round label with red Paralimna (Phaiosterna) vidua Giordani Soika, 1956a: 124.—Cogan, borderj/Ephydra bicolor. P. Macq. n. sp. [handwritten]/E. bi- 1968:325 [synonymy with P. aequalis Cresson]; 1980:663 [synonymy with P. color EX COLL. BIGOT/LECTOTYPE

0.1mm

114 115

FIGURES 113-115.—Male terminalia of Paralimna (Phaiosterna) bicolor (Macquart): 113, epandrium, cerci, and presurstyli, posterior aspect; 114, internal male terminalia, ventral aspect; 115, same, lateral aspect.

TOTYPE" and "By"; label with black submarginal border]." DU CONGO" handwritten]/HOLOTYPUS Paralimna vidua A. The lectotype is directly pinned, is in poor condition (right G[iordani]. Soika [red label; handwritten]/Paralimna aequalis wing mostly missing, several setae missing or misoriented, and Cress, det. B.H.Cogan 1967 ["aequalis Cress." and "7" hand- covered by hyphae), and is deposited in the UMO. In Mac- written]." The holotype is double mounted (minuten in a paper quart's paper, the type locality is stated to be "Egypte." card), is in poor condition (both wings torn, part of right wing The holotype female of Paralimna vidua Giordani Soika is missing, generally appearing "rubbed"), and is deposited in the labeled "HOLOTYPUS [light orange label with black submar- MRAC. We have also studied a female paratype (MRAC) that ginal border]/vois autres ex. en alcool [handwritten]/MUSEE bears the same locality data as the holotype. DU CONGO [Zaire] Banana-6-X-1934 [6 Oct 1934] (D.M. OTHER SPECIMENS EXAMINED.—Afrotropical. ALDABRA. Wanson) n. 2 {pris de trous de Crabes) [all except "MUSEE Grande Terre: Cinq Cases, fresh water pool, 23-29 Jan 1968, NUMBER 617 71

B.H. Cogan, A.M. Hutson (4d\ 39; BMNH); Dune Jean-Louis, May 1997, V. Hollmann, W.N. Mathis (15d\ 29; USNM, at light, 13-20 Mar 1968, B.H. Cogan, A.M. Hutson (1?; ZMHU); Intendance (4°47.1'S, 55°3O.1'E), 16 May 1997, BMNH). W.N. Mathis (Id"; USNM); La Retraite (4°35.1'S, 55°27.7'E), BURUNDI. Usumbura (780 m), 10 Apr 1953, F. Francois (19; 18-21 May 1997, V. Hollmann, W.N. Mathis (7d\ 29; USNM, 1RSN). ZMHU); Machabee, 5-6 Apr 1986, W.N. Mathis (3Id", 289; CAMEROON. Kribi (beach), route N7, 28-29 Nov 1987, A. USNM); Point Nord D'est (4°34.5'S, 55°27.7'E), 5 May 1997, Freidberg (2d\ 69; TZ, USNM). Kumba, 6 Oct 1949, H. Old- W.N. Mathis (3d"; USNM); Police Bay (4°48.0'S, 55°31.3'E), royd (19; BMNH). Limbe (shore), 14-15 Nov 1987, A. Freid- 16 May 1997, W.N. Mathis (4d"; USNM); Roche Caiman Bird berg, F. Kaplan (6d\ 99; USNM). Sanctuary (4°38.3'S, 55°28.1'E), 7 May 1997, V. Hollmann, ETHIOPIA. Lake Shala (7°21'N, 39°11'E), 1 Jul 1973, R. W.N. Mathis (12d\ 99; USNM, ZMHU). Praslin: Anse Ker- Baker (29; BMNH). lan Farm (4°18.5'S, 55°41.2'E), 13 May 1997, V. Hollmann, GAMBIA. Bakau (kotu stream, 3 km SW), 23 Feb 1977 (lcT; W.N. Mathis (8d*, 49; USNM, ZMHU); Anse Lazio (4°17.6'S, BMNH). Gunjur (5 km SSW; near beach in oil palm and man- 55°42.1'E), 8-13 May 1997, W.N. Mathis (6d% 29; USNM); grove vegetation), 22 Feb 1977 (19; BMNH). Anse Volbert (4°18.9'S, 55°44.7'E), 8 May 1997, W.N. Mathis GHANA. Neawani, 29 Sep 1921, J.W.S. Macfie (3d1, 39; (Id1; USNM); Baie Ste. Anne, Anse Takamaka (4°19.6'S, BMNH). Ashanti, Obuasi, 3 Apr 1906, W.H. Graham (19; 55°46.3'E), 10-13 May 1997, W.N. Mathis (3d"; USNM); Cote BMNH). D'Or (4°17.6'S, 55°42.1'E), 9 May 1997, W.N. Mathis (7d"; KENYA. Gazi (60 km S Mombasa), 5 May 1991, A. Freid- USNM). berg, F. Kaplan (2d"; USNM). Lake Victoria, Kisumu (10 km Lamb (1912:317) reported this species from Seychelles, 1 S), 19 Nov 1986, A. Freidberg (Ad , 69; USNM). Mombasa (45 Mahe, Port Glaud (beach), 5 Nov 1908, and Cascade (marshy km S), 5 May 1991, A. Freidberg, F. Kaplan (19; USNM). ground near sea level), 20 Feb 1909. MADAGASCAR. Antsiranana: Diego Suarez (30 m), 4-9 SUDAN. Yirol, 28 Mar 1954, E.T.M. Reid (ld\ 29; BMNH). Dec 1957, B.R. Stuckenberg (2d\ 29; BMNH, NMSA); Nosy TANZANIA. Mto Wa Mbu, near Lake Manyara Swamp (900 Be Beach, Ambatoloaka, 4-7 Apr 1992, A. Freidberg, F. Ka- m), 6 Sep 1992, A. Freidberg (19; USNM). plan (2cf; USNM); Nosy Tanikely, 6 Apr 1996, A. Freidberg, F. UGANDA. Katwe, Lac cratere salin, 20 Feb 1954 (19; IRSN). Kaplan (19; USNM). Toamasina: Est Ivontaka, Mananara YEMEN. Kirsh (914 m; on camels near waterholes; 14°37'N, (15 m), 10-14 Mar 1958, B.R. Stuckenberg (29; BMNH, 46°45'E), P.W.R. Petrie (3d1, 39; BMNH). NMSA). Toliara: Saint Augustin (6 m), 11-13 Feb 1953, ZAIRE. Kivu: Kayimvira (Uvira; at light), Feb-Mar 1955, B.R. Stuckenberg (Id", 29; BMNH, NMSA); South Berenty G. Marlier (19; MRAC); Rutshuru (river Kanzarue; 1200 m), Reserve, 20 Apr 1991, A. Freidberg, F. Kaplan (ld\ 19; 15 Jul 1935, G.F. de Witte (19; MRAC). Without Further USNM). Locality: Albert National Park (=Virunga National Park), NAMIBIA. Skeleton Coast Park, Mowe Bay (19°21.51'S, S.L. Edouard: Bitshumbi (925 m), 20-22 Apr 1936, L. Lip- 12°42.29'E), 22 Mar 1998, A.H. Kirk-Spriggs (Id"; NNIC). pens (5d\ 79; MRAC); Kamande (925 m), 1 Oct 1935, L. Lip- NIGERIA. Lake Chad, Mallam Fatori, 13 Apr 1967, J.C pens (Id1; MRAC); Kitembo (925 m), 3-4 Apr 1936, L. Lip- Deeming (29; BMNH). pens (2tf, 19; MRAC); May ya Moto (950 m), 15-16 Nov SEYCHELLES. Cousin: 30 Mar 1952, E.S. Brown (49; 1934, G.F. de Witte (39d\ 559; MRAC); Rwindi (river; 1000 BMNH). La Digue: Grand Anse (4°22.4'S, 55°50.6'E), 12 m), 9 Feb-17 Apr 1936, L. Lippens (12d\ 209; MRAC); Talia May 1997, V. Hollmann, W.N. Mathis (19d\ 179; USNM, (925 m), 7 Apr 1936, L. Lippens (39; MRAC). Ituri Kasenyi ZMHU); La Passe (4°20.8'S, 55°49.8'E), 14 May 1997, V. (Lac Albert), 5 Jul 1953, J. Verbeke (Id", 19; IRSN). Hollmann, W.N. Mathis (5d\ 29; USNM, ZMHU); Petite Anse Australasian/Oceanian. PAPUA NEW GUINEA. Central: Mo- (4°22.4'S, 55°50.6'E), 12 May 1997, V. Hollmann, W.N. reguina, near Cape Rodney (stoney river bank), 23 Jul 1982, Mathis (13 d\ 6 9; USNM, ZMHU). Mahe: Airport, 7-8 Apr J.W.Ismay(ld';USNM). 1986, W.N. Mathis (6d\ 29; USNM); Anse Boileau (4°42.5'S, Palearctic. EGYPT. Birket Qariin, near El Faiyum, Mar (29; 55°28.7'E), 4 May 1997, V. Hollmann (3d1; ZMHU); Anse For- ZMHU). Cairo, Mar (29; ZMHU). Cairo, El Marg (orange or- bans (4°46.7'S, 55°31.5'E), 4-19 May 1997, V. Hollmann, chard), 22 Mar 1996, M. Bartak (3d", 39; BAR). W.N. Mathis (9d% 119; USNM, ZMHU); Anse Royale Bay, DISTRIBUTION.—Afrotropical: Aldabra, Burundi, Cameroon, 6-9 Apr 1986, W.N. Mathis (17d\ 189; USNM); Auberge Ethiopia, Gambia, Ghana, Kenya, Madagascar, Namibia, Nige- D'Anse Boileau (4°42.3'S, 55°28.6'E), 7-18 May 1997, V. ria, Seychelles (Cousin, La Digue, Mahe, Praslin), Sudan, Tan- Hollmann, W.N. Mathis (28d\ 189; USNM, ZMHU); Baie de zania, Uganda, Yemen, Zaire. Australasian/Oceanian: Papua La Mouche (4°44.3'S, 55°29.5'E), 19 May 1997, V. Hollmann, New Guinea. Palearctic: Egypt. W.N. Mathis (Id1, 69; USNM, ZMHU); Beau Vallon, 20 REMARKS.—Our study of specimens of Phaiosterna from Feb-10, 11 May 1965, W.T. Tarns, I.B. Nye (5d; BMNH); Bel the Afrotropical Region indicates a single species, P. bicolor, Ombre (4°37'S, 55°24.9'E), 3 May 1997, W.N. Mathis (9d\ occurring there. The other four names that have been used for 29; USNM); Bel Ombre, Corsaire (4°37.1'S, 55°24.6'E), 3-18 this region (Cogan, 1980) are synonyms or represent misidenti- 72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY fications (see species synonymy). One available name, P. stripes between dorsocentral setae, especially anteriorly. Cos- vidua, was originally determined to be conspecific with P. lin- tal-vein ratio 0.41-0.50; M-vein ratio 0.94-0.97. Ventral apex eata, a valid species that occurs in the Oriental and Australa- of foretibia lacking numerous, long, slender setulae; posterior sian/Oceanian Regions, not the Afrotropical Region. Our con- surface of forebasitarsus lacking scattered long, slender setulae. clusions are based on dissection and study of structures of the Abdomen: Slightly to distinctly lighter in color than me- male terminalia of numerous Afrotropical specimens (Mada- sonotum, mostly brownish gray to brownish black; tergites gascar, Seychelles, and Cameroon). usually fasciate, with anterior portion of tergites darker gray to brownish black, less microtomentose, posterior portion more 24. Paralimna {Phaiosterna) decipiens Loew densely microtomentose, grayer. Male terminalia (Figures 110, 119-121): epandrium in posterior view inverted U-shaped FIGURES 110,116-121 (Figure 119), width about equal throughout length with dorsal Paralimna decipiens Loew, 1878:195.—OstenSacken, 1878:201 [Nearctic cat- margin of cereal cavity broadly rounded; cercus acutely alog].—Becker, 1896:270 [list, Texas].—Coquillett, 1900:259 [list, Puerto pointed dorsomedially and ventrally, medial margin nearly Rico].—Aldrich, 1905:624 [Nearctic catalog].—Jones, 1906:178-179 [key, straight; presurstyli in posterior view (Figure 119) connected catalog].—Thaxter, 1917:672-675 [parasites: Stigmatomyces rostratus Thax- ter, S. paralimnae Thaxter]; 1931:145-146 [parasite: S. gracilior Thaxter].— medially, forming shallowly arched and medially narrowed Wirth and Stone, 1956:470 [list, California]. band at ventral margin of cercus, produced ventrolaterally as Paralimna {Phaiosterna) decipiens.—Cresson, 1916:105 [subgeneric combi- narrow, triangular-shaped projections; postsurstylus in lateral nation], 108 [revision, figure of head]; 1946:230 [review, Nearctic Region]; view (Figure 121) with anterior margin bifurcate apically, ante- 1947a:54-55 [review].—Wirth, 1965:748 [Nearctic catalog]; 1968:16 [Neo- rior lobe narrower and shorter than apically expanded posterior tropical catalog].—Cole, 1969:396 [list, western United States].—Mathis and Zatwarnicki, 1995:126 [world catalog].—Mathis, 1995:633-635 [list, lobe, bearing several setulae medially and 2 longer setulae just Galapagos Islands]; 1997:59-60 [list, Belize, figures of male genitalia]. above bifurcation; gonite short, ovate; aedeagal apodeme with keel symmetrical (Figure 121), attachment with base of aedea- DIAGNOSIS.—This species is distinguished from congeners, gus recurved in direction of keel, attachment with hypandrium especially those of the subgenus Phaiosterna, by the following curved slightly ventrally; aedeagus broadly rounded apically, combination of characters: coloration variable but tending to be thereafter forming a short, narrow point (Figure 120); lateral tan to grayish brown, moderately microtomentose, at most sub- aedeagal processes wide throughout length, broadly rounded shiny; 1st flagellomere lacking fringe, or fringe of long, whit- apically, held close to lateral margins of aedeagus; hypandrium ish setulae along dorsum and dorsal portion of rounded apex much shorter than V2 height of 1st flagellomere; apex of 1st fla- deeply invaginated, pocket-like (Figures 120, 121). gellomere with dorsoapical corner more angulate than poster- TYPE MATERIAL.—The lectotype male of Paralimna decipi- oapical corner; mesonotum mostly uniform, gray to dark ens Loew (designated by Mathis, 1995:633) is labeled "[yellow brown, dull to subshiny, with moderately dense gray to brown square]/Loew Coll./decipiens m[ihi]. [handwritten]/Type microtomentum, consistently darker medially, between dorso- 11136 [number handwritten, red label]/LECTOTYPE Paral- central setae; ventral apex of male foretibia lacking numerous imna decipiens Loew cf By W.N.Mathis [handwritten except long, slender setulae; posterior surface of forebasitarsus lack- for "LECTOTYPE" and "By," label with black submarginal ing scattered long, slender setulae; and aedeagus wider than border]." Loew noted that the type series was from Texas, al- long in dorsal view, somewhat rectangular with distal angles though a locality label bearing this information does not ac- rounded, and in lateral view with distal aspect of sclerotized company the lectotype. The lectotype is pinned directly, is in portion slightly concave and actual apex of aedeagus as a good condition (some setae are missing on the dorsum of the pointed process projecting in an anteroventral angle. head and thorax; part of the abdomen has been removed and DESCRIPTION.—Small to medium-sized shore flies, body dissected, and the parts are in an attached microvial), and is de- length 1.85-3.60 mm; generally dark gray brown to black spe- posited in the MCZ. A female paralectotype (label data similar, cies, moderately densely microtomentose, dorsum appearing head missing) is designated here. more or less uniform in color, dull to subshiny, especially me- OTHER SPECIMENS EXAMINED.—Nearctic. UNITED STATES. sonotum. Arizona. Coconino County: Bill Williams Fork, F.H. Snow Head (Figures 116, 117): Generally gray brown to black. (6cf, 6?; ANSP). Pima County: Kits Peak (near; 32°N, First flagellomere bearing short, generally inconspicuous 111°32'W; 1097 m), Baboquivari Mountains, 7-9 Aug 1916 fringe of whitish setulae along dorsum and dorsal portion of (29; AMNH). Arkansas. Garland County: Hot Springs, 24 apex, length of setulae far less than V2 height of 1 st flagello- Jun, H.S. Barber (Id", 1?; ANSP). California. Del Norte mere. Face gray to grayish brown, microtomentose, appearing County: Crescent City, 19 Apr 1908, M.C. Van Duzee (19; dull, subdued. Gena-to-eye ratio 0.23-0.29. AMNH). Imperial County: Calipatria, 13 Nov 1921, E.R. Thorax (Figure 118): Generally gray brown to black, legs Kalmbach (Id1, 2?; ANSP); Salton Sea, 24 Nov 1921, E.R. darker, black with some grayish microtomentum; mesonotum Kalmbach (1

Sill

116

118

117 FIGURES 116-118.—Paralimna (Phaiosterna) decipiens Loew: 116, head, anterior aspect; 117, same, lateral aspect; 118, mesonotum, dorsal aspect. Scale = 0.5 mm.

23 Feb 1912, F. Knab (lcf; ANSP); Royal Palm Park, 28-29 County: New Orleans, 1 Jan 1932, A.L. Melander (Id1; Jan 1933, 1939, A.L. Melander (Id1, 29; ANSP). Henry ANSP). Mississippi. Harrison County: Pass Christian, 6 Jul County: Clewiston, 20 Jan 1938, A.L. Melander (1 tf; ANSP). 1917; J.M. Aldrich (Id1, 19; ANSP). Oktibbeha County: Manatee County: Bradenton, Mar, M.C. Van Duzee (Id1; Mississippi State, 30 Oct 1925 (lcT; ANSP). New Jersey. Cape ANSP). Marion County: Silver Springs, 2 Apr 1932, A.L. May County: Cape May, 9 Sep 1932, W. Stone (19; ANSP). Melander (Id1; ANSP). Monroe County: Everglades, 25 Jan Texas. Cameron County: Brownsville, 2-13 Jan 1928 (19; 1932, A.L. Melander (ltf, 1 ?; ANSP); Flamingo, 25 Jan 1939, AMNH); Brownsville (pond shore), 29 Nov 1910(19; ANSP); A.L. Melander (3d1, 59; ANSP). Orange County: Orlando, 2 Brownsville (9.7 km N), 20 Jul 1933, C.W. Sabrosky (Id", 19; Feb 1932, A.L. Melander (2d\ 2?; ANSP). Palm Beach ANSP). Galveston County: Dickinson, Jun 1929 (2d\ 29; County: Lake Worth, A.T. Slosson (1 d", 1 9; AMNH). Pinel- ANSP). Travis County: Austin, 7-20 Oct 1899, 1901 (Id1, las County: Saint Petersburg, 15 Feb 1924, E.T. Cresson, Jr. 49; AMNH, ANSP). (3 9; ANSP). Polk County: Lakeland, 10 Nov 1911(1?; Neotropical. BAHAMAS. Andros: Andros Town, 27-28 Feb AMNH). Georgia. Charlton County: Bill's Island, Okefeno- 1966, O.L. Cartwright (Id1, 29; USNM); Mangrove Cay, 26 kee Swamp, Jun 1912 (19; ANSP). Louisiana. St. Charles Apr 1953, E.B. Hayden (Id1, 19; AMNH). Exuma Cays: 74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 119-121.—Male terminalia of Paralimna (Phaiosterna) decipiens Loew: 119, epandrium, cerci, and presurstyli, posterior aspect; 120, internal male terminalia, ventral aspect; 121, same, lateral aspect.

T 0.1m1mm 1

120 121

Stainaard Cay, 13 Jan 1953, E.B. Hayden (W, 39; AMNH). BOLIVIA. La Paz: Yolosa (16°14'S, 67°44.4'W; 1185 m), Rum Cay: near Port Nelson, 16 Mar 1953, E.B. Hayden, L. 18 Mar 2001, W.N. Mathis (3d1; USNM). Giovannoli (1

USNM); Palmarejo (67 km E Havana), 26 Apr 1983, W.N. Los Gemelos (Scalesia forest, 600 m), 1-9 Apr 1989, Stewart Mathis (8cf, 1?; USNM); Puerto Escondido, 26 Apr 1983, B. Peck, Bradley J. Sinclair (lcf, 69; CNC); Puntudo (1 km N, W.N. Mathis (lcf, 2?; USNM). Matanzas: Palpite (2 km Scalesia forest, 650 m), 1-30 Mar 1989, S.B. Peck, Bradley J. NE), 30 Apr 1983, W.N. Mathis (19; USNM); Playa Larga (1 Sinclair (3 9; CNC). km E), 2 May 1983, W.N. Mathis (1?; USNM). Pinar del GRAND CAYMAN. Airport (W end; 19° 17.4^, 81°22.2'W), / Rio: Soroa (22°47.7 N, 83°W), 4-6 Dec 1994, W.N. Mathis 26 Apr 1994, W.N. Mathis (5d\ 29; USNM). Airport (2 km E; (9cf, 19; USNM); Soroa (2 km E), 29 Apr 1983, W.N. Mathis 19° 17. IX 81°21'W), 26 Apr 1994, W.N. Mathis (6cf; USNM). (9

19; USNM). Playa de Guayanilla (18°0.4'N, 66°46.1'W), 19 lapping with P. obscura. Accurate identification may require Sep 1995, D. and W.N. Mathis (5d*; USNM). Punta Jacinto examination of the aedeagus and lateral aedeagal processes. (near Guanica; 17°57'N, 66°52.6'W), 20 Sep 1995, D. and Because specimens of P. decipiens are similar to and fre- W.N. Mathis (Id1; USNM). quently occur sympatrically with those of P. obscura, Mathis MEXICO. Jalisco: Guadalajara, 8 Aug 1903 (Id1; ANSP). (1995) had previously designated a lectotype for the former to Sonora: Imuris, 2 Jul 1952, C. and P. Vaurie (3d1, 29; keep nomenclatural issues at the same level of resolution as our AMNH). Tamaulipas: Tampico, 27 Jul-28 Dec 1908, 1933, understanding of the zoology. C.W. Sabrosky (29; ANSP). Veracruz: Tamos, 28 Jul 1933, 1 C.W. Sabrosky (Id ; ANSP). 25. Paralimna (Phaiosterna) fusca Bock PANAMA. Canal Zone: Ancon, 4 Dec 1909, S.T. Darling (2d\ 29; ANSP). FIGURES 111, 122-124 ST. LUCIA. Dauphin Boguis (1.6 km S Marquis; 14°01'N, Paralimna fusca Bock, 1988:896.—Mathis and Zatwarnicki, 1995:120 [world 60°55'W), 17 Jun 1991, D. and W.N. Mathis (4cf; USNM). Mi- catalog]. coud (13°49'N, 60°54'W), 15 Jun 1991, D. and W.N. Mathis DIAGNOSIS.—This species is distinguished from congeners 1 (3d ; USNM). Soufriere (beach; 13°51TS|, 16°54'W), 11-12 Jun by the following combination of characters: first flagellomere 1991, D. and W.N. Mathis (5<*\ 19; USNM). Sulphur Spring bearing fringe of long, whitish setulae along dorsum and dorsal (13°50'N, 61°03'W), 14 Jun 1991, D. and W.N. Mathis (4o\ portion of rounded apex, setulae longer than V2 height of 1st 39; USNM). flagellomere; apex of 1st flagellomere evenly rounded; and ST. VINCENT. Charlotte: Owia Salt Pond (13°22.5'N, mesonotum black, subshiny to shiny, with sparse brown micro- 1 61°08.5'W), 6 Sep 1997, W.N. Mathis (Id , 19; USNM); Peru- tomentum, lacking conspicuous, longitudinal stripes. / vian Vale (13°10.7 N, 6P08.7'W), 6-8 Sep 1997, W.N. Mathis DESCRIPTION.—Moderately small to medium-sized shore (6

FIGURES 122-124.—Male terminalia of Paralimna (Phaiostema) fusca Bock: 122, epandrium. cerci, and presurst- yli, posterior aspect; 123, internal male terminalia, ventral aspect; 124, same, lateral aspect.

T 0.1mi 1 122

123 124

aedeagal processes narrow throughout length, narrowly PAPUA NEW GUINEA. Central: Yule Island (rocky shore), 1 rounded apically (Figure 123); hypandrium deeply invaginated, Jan 1983, J.W. Ismay (1

USNM); Kirinda, 25 Apr 1980, L. Jayawickrema, W.N. Thorax: Generally gray to dark brown, legs darker; me- Mathis, T. Wijesinhe (17o", 37?; USNM). Western Province. sonotum mottled, moderately densely gray to dark brown mi- Colombo: Colombo, Kalutaluwewa (light trap), 19 Feb 1958, crotomentose, appearing dull to very slightly subshiny, infre- (lcf, 1?; BMNH); Negombo, 7 May 1980, L. Jayawickrema, quently with darker brown, longitudinal stripes medially, W.N. Mathis, T. Wijesinhe (5d\ 3?; USNM). between dorsocentral setae, although consistently darker there. THAILAND. Bangkok: Pratomvan (at light), Aug-Sep 1962, Costal-vein ratio 0.51-0.52; M-vein ratio 0.85-0.94. Foreleg of J.E. Scanlon(19;BMNH). male lacking long, slender setulae. DISTRIBUTION.—Australasian/Oceanian: Australia (Coral Abdomen: Slightly lighter in color than mesonotum, Sea Islands Territory, Northern Territory, Queensland), Christ- mostly grayish brown; anterior portion of tergites fasciate, mas Island, Guam, Papua New Guinea, Vanuatu. Oriental: In- darker gray to brown, less microtomentose. Male terminalia dia (West Bengal), Indonesia (Sumatra), Malaysia, Sri Lanka, (Figures 107, 125-127): epandrium inverted U-shaped in pos- Thailand. terior view (Figure 125), wide, width about equal throughout REMARKS.—Although this species was described and until length, with dorsal margin of cereal cavity broadly rounded; now was known only from Queensland, Australia, we have dis- cercus pointed dorsomedially and ventrally, medial margin sin- covered it to occur widely throughout the Australasian/Ocean- uous; presurstyli in posterior view (Figure 125) connected me- ian and Oriental Regions. dially, forming shallowly arched band at ventral margin of This is the only species of the subgenus Phaiosterna from cerci, produced ventrolaterally as narrow, triangular-shaped the Old World that has a subshiny to shiny dorsum, especially projections; postsurstylus in lateral view (Figure 127) with an- the mesonotum, and lacks any evidence of longitudinal stripes. terior margin swollen medially, bearing several setulae medi- ally, bifurcate apically, anterior lobe narrower and shorter than 26. Paralimna {Phaiosterna) lineata de Meijere posterior lobe in lateral view, posterior lobe broadly rounded apically; aedeagal apodeme with keel asymmetrical, more an- FIGURES 107,125-127 gulate toward hypandrium and more rounded toward base of Paralimna lineata de Meijere, 1908:165.—Bezzi, 1928:154 [list, Fiji].—Mal- aedeagus, attachment with base of aedeagus shallowly re- loch, 1933:11 [list, Marquesas]; 1934a:313 [list, Samoa].—Cresson, 1948:10 curved, attachment with hypandrium curved slightly ventrally; [in part; listed as a species incertae].—Cogan and Wirth, 1977:334 [Oriental catalog].—Bock, 1988:893-894 [revision].—Mathis, 1989:644 [Australa- aedeagus in lateral view (Figure 127) narrowly triangular, sub- sian/Oceanian catalog]. rectangular in ventral view (Figures 107, 126), slightly nar- Phaiosterna aequalis Cresson, 1929:193 [list, Philippines, Taiwan, Viet- rowed basally, otherwise mostly parallel sided, apex broadly nam].—Cogan and Wirth, 1977:334 [synonymy].—Cogan, 1980:663 [syn- rounded to truncate thereafter, forming short, narrow point; lat- onymy]. eral aedeagal process wide throughout length, broadly rounded Paralimna (Phaiosterna) aequalis.—Cresson, 1948:10 [generic combination; list, India, Philippines, Taiwan].—Mathis and Zatwarnicki, 1995:127 [world apically, spatulate (Figure 126); hypandrium deeply invagi- catalog]. nated, pocket-like (Figures 126,127). DIAGNOSIS.—This species is distinguished from congeners, TYPE MATERIAL.—The lectotype male of Paralimna lineata especially those of the subgenus Phaiosterna, by the following de Meijere (designated by Bock, 1988:894) is labeled "[Indo- combination of characters: first flagellomere lacking fringe or nesia.] Java[:] I." 08 [Jan 1908] Semarang. Jacobson./Paral- fringe of long, whitish setulae along dorsum and dorsal portion imna lineata de Meijere Type, [handwritten]/ Lectotype [red la- of rounded apex far shorter than Vfe height of 1st flagellomere; bel]/LECTOTYPE Paralimna lineata o" de Meij. det B.H. apex of 1 st flagellomere with dorsoapical corner more angulate Cogan 1971. [all except "det B.H. Cogan" handwritten]/LEC- than posteroapical corner; and mesonotum mottled, gray to TOTYPE Paralimna lineata de Meijere cr By I.R. Bock, 1988 dark brown, dull to subshiny, with moderately dense gray to ["LECTOTYPE" and "By" in red type; all else handwritten; la- brown microtomentum, consistently darker medially, between bel with black submargin]." The lectotype is double mounted dorsocentral setae. (minuten into venter of specimen and in rectangular block of DESCRIPTION.—Small to medium-sized shore flies, body pith), is in fair condition (setae on right side largely missing), length 2.15-3.0 mm; generally dark, whitish gray to brown and is deposited in the ZMA. Three female paralectotypes bear species, moderately densely to densely microtomentose, dor- the same locality label data as the lectotype. sum appearing mottled, dull to slightly subshiny, especially The holotype male of Phaiosterna aequalis Cresson is la- mesonotum. beled "Fruhstorfer Mittel-Annam [Vietnam]/cf/Phaiosterna Head: Generally grayish brown to dark brown. First flagel- AEQUALIS E.T.Cresson,Jr. [red label; species name hand- lomere bearing short, generally inconspicuous fringe of whitish written]/Collection of the Academy of Natural Sciences of setulae along dorsum and dorsal portion of apex, setulae far Philadelphia (ANSP)." The holotype is double mounted (mi- less than V2 height of 1 st flagellomere. Face mottled, gray to nuten in a rectangular block of foam-like plastic, is in good dark brown, moderately densely microtomentose, appearing condition (right foreleg missing; some setae, especially on left dull, subdued. Gena-to-eye ratio 0.23-0.24. side of head, displaced or missing), and is deposited in ANSP NUMBER 617 79

FIGURES 125-127.—Male terminalia of Paral- imna (Phaiosterna) lineata de Meijere: 125, epan- drium, cerci, and presurstyli, posterior aspect; 126, internal male terminalia, ventral aspect; 127, same, lateral aspect.

T .1m1i

126 127

(6340). There are also four paratypes (lcf, 3?; ANSP, NMW) FEDERATED STATES OF MICRONESIA. Yap: Giliman, 11 Jun that bear the same locality data as the holotype. 1957, C.W. Sabrosky (19; BMNH); Ruul, near Kolonia, 13 Jun OTHER SPECIMENS EXAMINED.—Australasian/Oceanian. 1957, C.W. Sabrosky (2 9; BMNH). Ifalik: Ifalik Atoll, 6 Sep AMERICAN SAMOA. Tutuila: Aunuu Island, 10 Dec 1953, C. 1953 (ltf; BMNH). Kosrae (=Kusaie): Mutunlik, 18 Mar Hoyt (lcf; BMNH); Malae'imiV'ey, 7 Jul 1953, C. Hoyt (lcf, 1953, J.F.C. Clarke (lcT; BMNH). Ponape: Not, Palieij, Jan 1?; BMNH); Pago Pago, 14 Dec 1925, G.H. Hopkins (Id1; BMNH). 1948, N.S. Hurlbut (1 d"; BMNH). Truk Islands: Moen, South AUSTRALIA. Queensland: Cairns (bay shore and puddles), Valley, Mt. Tonaachau, 25 Apr 1949, R.W.L. Potts (19; 19-25 Apr 1957, W.W. Wirth (3d1, 5 9; BMNH). Western BMNH). Australia: Millstream (at light), 7 Apr 1971, D.H. Colless FIJI. Viti Levu: Lautoka, 28 Feb 1919, H. Greenwood (2tf, (2?;ANIC). 19; BMNH); Lautoka (0.5 m), Mar 1976, N.L. Krauss (1 d", 19; 80 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

USNM); Nandi (0-100 m), 5 May 1973, N.L.H. Krauss (2d\ (Id1, 19; BMNH); Madras, Tondiarpet (over open sewage), 13 3?;USNM). Aug, Fletcher (Id", 19; ANSP, BMNH); Kurumbagaram (asso- GUAM. Agana (muddy stream), 25 May 1936, R.L. Usinger ciated with stagnant water), Nov 1951, P.S. Nathan (4tf\ 29; (1?; BMNH). Agat (grass), 20 May 1936, O.H. Sweezy (1?; BMNH). West Bengal: Calcutta, Tollygunge, 29 Sep 1904, BMNH). Inarajan (on rice), 25 Jul 1936, O.H. Sweezy (1?; Brunetti (2tf\ 69; BMNH). Calcutta, Edengard, 22 Dec 1956, BMNH). A.P. Kapur (29; BMNH). Diamond Harbour, 1 Dec 1927, R. KIRIBATI. Gilbert Islands: Marakei Atoll, Dec 1957, Hoogly(19;BMNH). N.L.H. Krauss (19; BMNH). INDONESIA. Java: Djakarta, Aug 1908, Jacobson (Id1; MARQUESAS. Fatu Hiva: Oomoa Valley, 21 Aug 1930, Le ZMA); Wonosobo, May 1909, Jacobson (19; ZMA). (Al- Bronnec (lcf; BMNH). Nuku Hiva: Haaoatupa, 14 Jun 1987, though Cogan labeled these two specimens as paralectotypes, W.N. Mathis (6d", 1 ?; USNM); Taiohae, 9-15 Jun 1987, W.N. the localities are not mentioned in the original description, nor Mathis (8d\ 29; USNM); Toovii (800 m), 10-12 Jun 1987, do the collection dates correspond.) Lesser Sunda Islands. West W.N. Mathis (34d", 129; USNM). Sumbawa: Semongkat (8°35'S, 117°20'E), 10-15 May 1927, NORTHERN MARIANAS. Saipan: Tsutsuuran, 31 Aug 1944, Sunda-Expedition, Rensch (3d1, 19; BMNH); Dompoe D.G. Hall (Id"; USNM). (8°32'S, 118°28'E), 24-25 May 1927, Sunda-Expedition, Ren- PALAU. Babelthuap: Babelthuap (along streams and ca- sch (2d\ 29; BMNH); Soembawa-Besar (8°30'S, 117°26'E), cao), 16 Apr-21 May 1957, C.W. Sabrosky (5d\ 29; BMNH); 24 Apr-2 May 1927, Sunda-Expedition, Rensch (Id1; BMNH). Ulimang, 14 Dec 1947, H.S. Dybas (19; BMNH). LAOS. Ban Theuong (18 km NW Xieng Khouang; 1035 m; at PAPUA NEW GUINEA. Central: Aieme River (river side), 31 light), 2-6 Aug 1960, R.E. Leech (2tf; BPBM, TZ). Bolouens Oct 1982, J.W. Ismay (19; USNM); Brown River Bridge (12 Plateau (16 km S Thateng; 1020 m; at light), 22-24 Jul 1960, km NW), 18 May 1986, J.W. Ismay (4

FIGURES 128-130.—Male terminalia of Paralimna (Phaiosterna) longiseta, new species: 128, epandrium, cerci, and presurstyli, posterior aspect; 129, internal male termina- lia, ventral aspect; 130, same, lateral aspect.

T 0.1mm 1

129 130

forming shallowly arched band at ventral margin of cerci, pro- TYPE MATERIAL.—The holotype male is labeled "DOMINI- duced ventrolaterally as triangular-shaped projections; postsur- CAN RP. Azua: near Pueblo Viejo 18°24.8'N,70o44.7'W stylus in lateral view (Figure 130) with anterior margin swollen 19Mayl998, WNMathis/USNM ENT 00087712 [plastic bar medially, bearing several setulae, bifurcate apically, anterior code label]/HOLOTYPE

W.N. Mathis (2eT; USNM); El Rio (9.5 km E; 19°0.7'N, lacking fringe or fringe of long, whitish setulae along dorsum 70°33.6'W; 980 m), 24 May 1998, D. and W.N. Mathis (6tf; and dorsal portion of rounded apex far shorter than Vfe height of USNM); Rio Camu (3.5 km NW La Vega; 19°13.7'N, 1st flagellomere; apex of 1st flagellomere with dorsoapical cor- 70°35.2'W; 100 m), 10 May 1995, W.N. Mathis (6rf, 1?; ner more angulate than posteroapical corner; mesonotum USNM); Rio Camu (3.5 km NW La Vega; 19°13.8'N, mostly uniform, gray to dark brown, dull to subshiny, with 70°35.2'W; 100 m), 18 May 1998, D. and W.N. Mathis (6d\ moderately dense gray to brown microtomentum, consistently 4 ?; USNM). Peravia: Rio Ocoa (San Jose Ocoa; 18°31TN, darker medially, between dorsocentral setae; ventral apex of 70°30.4'W), 21 May 1998, D. and W.N. Mathis (5tf, 1?; male foretibia not bearing numerous long, slender setulae; pos- USNM). Puerto Plata: Rio Camu (14 km E Puerto Plata; terior surface of forebasitarsus not bearing scattered long, slen- 19°41.9'N, 70°37.5'W), 23 May 1998, D. and W.N. Mathis der setulae; aedeagus narrow in dorsal view, slightly longer (6cT; USNM); Rio Perez (near Imbert; 19°44.1'N, 70°50.2'W), than wide, apical part of sclerotized portion distinctly pointed, 24 May 1998, D. and W.N. Mathis (6tf; USNM). especially evident in lateral view. OTHER SPECIMENS EXAMINED.—Neotropical, BRAZIL. Mato DESCRIPTION.—Moderately small to medium-sized shore Grosso: Capitao Vasconselos, Rio Tuatuari, Upper Xingu flies, body length 2.25-3.50 mm; generally dark, gray brown to Basin, 31 Jul 1957, B. Malkin (18tf, 289; CAS, USNM). black species, moderately densely microtomentose, dorsum ap- HONDURAS. Cortes: Omoa (15°47.8'N, 87°58.4'W), 26 pearing relatively uniform in color, dull to subshiny, especially Sep 1995, D. and W.N. Mathis (3d1, 19; USNM); San Pedro mesonotum. Sula (8 km S; 15°25.7X 88°01.4'W), 25-26 Sep 1995, D. and Head (Figures 131-136): Generally gray brown to black. W.N. Mathis (2tf; USNM). First flagellomere bearing short, generally inconspicuous JAMAICA. 5/. Andrew: Wag Water River, 25 Feb 1969, fringe of whitish setulae along dorsum and dorsal portion of W. W. Wirth (1 cC; USNM). St. Thomas: Yallahs River (mouth; apex, setulae far shorter than V2 height of 1st flagellomere. H^TM, 76°35.6'W), 14 May 1996, D. and W.N. Mathis, H. Face gray to grayish brown, microtomentose, appearing dull, Williams (8c/, 1?; MCV, USNM). subdued. Gena-to-eye ratio 0.24-0.27. MEXICO. San Luis Potosi: Tamazunchale, 23 Nov 1946, Thorax (Figures 137-139): Generally gray brown to black, R.E. Skinner (9tf; CAS, USNM). legs darker, black with some grayish microtomentum; mesono- PUERTO RICO. Rio Hoconuco (18°7.6'N, 67°2.6'W), 20 Sep tum moderately densely gray brown to brownish black, micro- 1995, D. and W.N. Mathis (lcf; USNM). tomentose, appearing dull to subshiny, frequently with lighter DISTRIBUTION.—Neotropical: Brazil (Mato Grosso), Hondu- brown, short to long stripes between dorsocentral setae, espe- ras, Mexico, West Indies (Dominican Republic, Jamaica, Puerto cially anteriorly. Costal-vein ratio 0.48-0.53; M-vein ratio Rico). 0.95-1.0. Ventral apex of foretibia lacking numerous long, ETYMOLOGY.—The specific epithet, longiseta, alludes to the slender setulae; posterior surface of forebasitarsus lacking scat- long, fine setae on the foreleg of males. tered long, slender setulae. REMARKS.—Older specimens tend to be darker, more rusty Abdomen: Slightly to distinctly lighter in color than me- colored and have more setae missing or broken. sonotum, mostly brownish gray to brownish black; tergites We have frequently collected this species sympatrically with usually fasciate, with anterior portion of tergites darker gray to the other two species of the subgenus Phaiosterna that occur in brownish black, less microtomentose than posterior, more the New World. densely microtomentose and grayer portion. Male terminalia (Figures 109, 140-142): epandrium in posterior view inverted U-shaped (Figure 140), wide, width about equal throughout 28. Paralimna (Phaiosterna) obscura Williston length, with dorsal margin of cereal cavity narrowly rounded; FIGURES 109,131-142 cercus with medial margins deeply sinuous, concave dorsally, Paralimna obscura Williston, 1896:391.—Coquillett, 1900:259 [list, Puerto acutely pointed dorsomedially, ventral margin produced ven- Rico].—Aldrich, 1905:624 [Nearctic catalog].—Jones, 1906:178-179 [key, tromedially in posterior view; presurstyli in posterior view catalog]. (Figure 140) at most very narrowly connected medially or not Paralimna (Phaiosterna) obscura.—Cresson, 1916:105 [subgeneric combina- touching, produced ventrolaterally as triangular-shaped projec- tion], 109-110 [revision]; 1946:230 [review, Nearctic Region]; 1947a:54-55 tions; postsurstylus in lateral view (Figure 142) with anterior [review].—Wirth, 1965:748 [Nearctic catalog]; 1968:16 [Neotropical cata- log].—Mathis and Edmiston, 1991:832-834 [discussion, designation of a margin swollen medially, bearing several setulae, bifurcate api- lectotype].—Mathis and Zatwamicki, 1995:127 [world catalog].—Mathis, cally, with anterior lobe short and narrow, posterior lobe 1997:56-59 [list, Belize, scanning electron micrographs, figures of male broadly rounded in lateral view; gonite rod-like, very slightly genitalia]. arched; aedeagal apodeme with keel asymmetrical (Figure 142) DIAGNOSIS.—This species is distinguished from congeners, and only moderately produced, with shallow indentation to- especially those of the subgenus Phaiosterna, by the following ward ventral margin; aedeagus somewhat triangular in lateral combination of characters: specimens tending to be dark brown view (Figure 142), with long, narrow, medial process apically; to black, shiny, sparsely microtomentose; 1st flagellomere lateral aedeagal processes robust (Figure 141), at apex very 84 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 131-139.—Scanning electron micrographs of Paralimna (Phaiosterna) obscura Williston (Belize. Stann Creek District: Wee Wee Cay; scale length in parenthesis; scale for all photographs = Figure 131): 131, head, lateral aspect (0.33 mm); 132, head, anterior aspect (0.30 mm); 133, frons, anterodorsal aspect (176 pm); 134, face, anterior aspect (176 vim); 13S, ommatidia of compound eye, lateral aspect (17.6 um); 136, enlarge- ment of interfacetal seta, lateral aspect (6.0 um); 137, mesonotum, dorsal aspect (0.38 mm); 138, scutellum, dorsal aspect (176 urn); 139, thorax, lateral aspect (0.38 mm). NUMBER 617 85

FIGURES 140-142.—Male terminalia of Paralimna (Phaiosterna) obscura Williston: 140, epandrium, cerci, and presurstyli, posterior aspect; 141, internal male ter- minalia, ventral aspect; 142, same, lateral aspect.

T .1m

140 1

141 142

slightly oriented medially; hypandrium very deeply invagi- piece of cardboard), is in good condition, and is deposited in nated, pocket-like (Figures 141,142). BMNH. There are also 15 paralectotypes as follows: BMNH 1 TYPE MATERIAL.—The lectotype male of Paralimna ob- (4o\ 3?), KU (3

Mathis (5cf; USNM); Speyside (Doctor River; 11°18.2'N, Papuama, new genus 60°31'W), 19 Apr 1994, D. and W.N. Mathis (7cT, 1?; TYPE SPECIES.—Papuama ismayi Mathis and Zatwarnicki, USNM); Speyside (Doctor River; 1 km NW; 11°18'N, by present designation. 60°32'W), 12-13 Jun 1993, W.N. Mathis (5

Key to Species of Papuama 1. Thoracic setae normally developed, dorsocentral setae 1 +3, pre- and postsutural supra- alar setae and anterior notopleural seta generally well developed; scutum and scutel- lum mostly concolorous, yellowish tan to gray (Australia) 29. P. calva (Bock) Thoracic setae weakly developed, dorsocentral setae 0+1, pre- and postsutural supra- alar setae and anterior notopleural seta greatly reduced or lacking; scutum mostly dark brown, contrasting with grayish scutellum (Australasian/Oceanian, Oriental) 30. P. ismayi, new species

FIGURE 143.—Distribution map for Papuama (hatched and dots). 90 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

29. Papuama calva (Bock), new combination brown dorsally and gray to silvery white on the face and pleu- ral areas; legs dark brown; chaetotaxy as in generic description. FIGURES 144-146 Head (Figures 147, 148): Frons with vertex dark brown, Paralimna calva Bock, 1988:895.—Mathis and Zatwarnicki, 1995:119 [world becoming golden brown anteriorly to about level of anterior catalog]. ocellus, thereafter anteriorly gray to silvery white. Antenna DIAGNOSIS.—This species is similar to P. ismayi but is dis- brownish black to black; 1st flagellomere broadly rounded api- tinguished from it and other taxa of Dryxini by the following cally; arista with 8-10 dorsal rays. Face projected anteriorly, combination of characters: ocellar seta present, well developed; forming sloping, narrow shelf between and ventrad of antennal proclinate fronto-orbital setae greatly reduced, at most setulae- bases, color entirely gray to silvery white, appearing blackish like; reclinate fronto-orbital seta present, well developed; ari- gray from some angles; bearing 3 or 4 long, slender setae in a stal hairs 6-8; notopleuron with 2 setae, posterior seta more vertical row laterally and with several shorter setulae between weakly developed; dorsocentral setae well developed (1+3); row of longer setae and parafacial. Gena-to-eye ratio anepisternal setae along posterior margin weakly developed; 0.30-0.33. katepistemal seta present but weakly developed, pale, setulae- Thorax (Figure 149): Mesonotum mostly dark brown with like or occasionally absent. Forefemur lacking row of closely some marginal areas gray to silvery white; scutellum usually set, short, tooth-like setulae along anteroventral margin. Male considerably lighter, more silvery gray, than scutum; anterior terminalia as figured (Figures 144-146). notopleural seta very weakly developed or lacking; dorsocen- TYPE MATERIAL.—The holotype male is labeled "Cas- tral setae (0+1) weakly developed or absent, with only poste- tlereagh R[iver]. Mendooran, N.S.W. 24 March 1971 D.K. rior seta well developed; anepisternal setae along posterior McAlpine/HOLOTYPE [bright red label]." The holotype is margin moderately weakly developed but evident; katepistemal double mounted (glued to a relatively large paper triangle), is seta weakly developed, setula-like, pale. Wing hyaline; costal- in excellent condition, and is deposited in the AM. vein ratio 0.27-0.32; M-vein ratio 0.68-1.0. Coxae, femora, OTHER SPECIMENS EXAMINED.—Australasian/Oceanian. and tibiae mostly blackish gray to silvery gray, usually darker AUSTRALIA. New South Wales: Mendooran, Castlereagh dorsally and on posterior surface; forefemur bearing row of River, 24 Mar 1971, D.K. McAlpine (3d1, 2?; AM). closely set, short, tooth-like setulae along anteroventral mar- DISTRIBUTION.—Australasian/Oceanian: Australia (New gin; tarsi yellowish orange to blackish orange, darker dorsally, South Wales, Queensland). apical tarsomere blackish brown. REMARKS.—We have determined that this species, although Abdomen: Male terminalia (Figures 150-153): epandrium not demonstrating some of the autapomorphies of Papuama is- in posterior view (Figure 150) thick-walled, inverted U- mayi, is similar and related to the latter species. This determi- shaped, in lateral view pointed anterodorsally, sloping down nation is primarily based on characters of the male terminalia. posteriorly to dorsal Vb, thereafter ventrally nearly parallel We suspect that this species is more like, if not the same spe- sided, becoming very slightly wider only ventrally, ventral cies as, the basal taxon that gave rise to this genus and that P. margin with shallow, broadly based point near middle; cercus ismayi is a more derived species, as evidenced by the reduction in posterior view (Figure 150) irregularly oval; presurstylus in chaetotaxy. longer than wide, in posterior view finger-like, curved medi- ally, broadly rounded apically, in lateral view curved posteri- 30. Papuama ismayi, new species orly on ventral Vz, apex pointed, bearing 3 or 4 long setulae medially from basal Vr, postsurstylus (Figures 152, 153) 3-4 FIGURES 147-153 times longer than wide, with medial, short process just before DIAGNOSIS.—This species is similar to Papuama calva but is midheight dorsally, apex shallowly bifurcate, each process of distinguished by the following combination of characters: ocel- bifurcation broadly rounded in lateral view, lateral process lar seta present, well developed; proclinate fronto-orbital setae longer and spatulate, bearing 3 setulae on ventral Vz along pos- greatly reduced, at most setulae-like; reclinate fronto-orbital terior margin and 2 setulae at dorsal Vz along anterior margin, seta present, well developed; arista I hairs 8-10; notopleuron inner process of bifurcation with apex foot-like, posterior por- bearing 1 well-developed seta, anterior seta very weakly devel- tion produced; aedeagus in lateral view (Figure 153) broadly oped or lacking; dorsocentral setae weakly developed or absent foot-like, anteroventral portion broadly projected for short (0+1); anepistemal setae along posterior margin weakly devel- length, projection bearing several short papillae, in ventral oped; katepistemal seta present but weakly developed, pale, view (Figure 152) like 3-pronged fork, medial prong nearly setulae-like; scutum mostly dark brown, contrasting with gray- twice length of lateral prongs; aedeagal apodeme angulate ish scutellum; and forefemur bearing row of closely set, short, (Figure 153) with posterior crest dorsomedially, attachment to tooth-like setulae along anteroventral margin. aedeagus broader and shorter than attachment with hypan- DESCRIPTION.—Small to moderately small shore flies, body drium; hypandrium broadly and shallowly concave, anterior length 1.65-2.60 mm; generally densely microtomentose, dark margin broadly rounded (Figures 152, 153). NUMBER 617 91

FIGURES 144-146.—Male terminalia of Papuama calva (Bock): 144, epandrium, cerci, and presurstyli, posterior aspect; 145, internal male terminalia, ventral aspect; 146, same, lateral aspect.

I 0.1mm

145

TYPE MATERIAL.—The holotype male is labeled "PAPUA block of plastic), is in excellent condition, and is deposited in NEW GUINEA Cen.Prov., Daramouka Vill. 19 Sept 1982 JW the NMNH. The allotype female and seven other paratypes Ismay (riverside)." The holotype is double mounted (minute in (3cf, 4?; USNM) bear the same locality label as the holotype. 92 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

147

149

148

FIGURES 147-149.—Papuama ismayi, new species: 147, head, anterior aspect; 148, same, lateral aspect; 149, mesonotum, dorsal aspect. Scale=0.S mm.

Other paratypes are as follows: PAPUA NEW GUINEA. PAPUA NEW GUINEA. Bainyik, 14 Dec 1963, D.K. Me Alpine Central: LaLoki (river margin), 2 Feb 1986, J.W. Ismay (3a", (5cf, 79; BMNH). Imbia, near Maprik, 19 Dec 1963, D.K. 69; USNM); Woitope (river margin), 9 Jun 1986, J.W. Ismay Me Alpine (Id1, 19; BMNH). (19; USNM); Dauramouku Village (river side), 19 Sep 1982, Oriental. INDONESIA. Sumatra: Fort de Kock (920 m), J.W. Ismay (2cf, 4 9; USNM). North: Gira River near Popon- 1925, E. Jacobson (2tf, 39; BMNH). detta (8°45'S, 148°15'E; bank fast river), 30 Jun 1982, J.W. Is- NEPAL. Chitwan: Sauraha, Dhungari Khola (2 km E; may (17d\ 89; MCV, USNM). Smithsonian Institution Camp; rocky bank), 1 Nov 1985, W.N. OTHER SPECIMENS EXAMINED.—Australasian/Oceanian. Mathis(lcf;USNM). IRIAN JAYA. Siriwini River (Nabire; medium-sized river with THAILAND. Loei: Na Haeo, 12 Feb 1999, P. Grootaert (lcr, gravel and mud banks), 21 Apr 1997, P. Grootaert {!

151

0.1mm

150

152 153

FIGURES 150-153.—Male terminalia of Papuama ismayi, new species: 150, epandrium, cerci, and presurstyli, posterior aspect; 151, same, lateral aspect; 152, internal male terminalia, ventral aspect; 153, same, lateral aspect.

REMARKS.—This species exhibits many apomorphies, espe- Bock. The shape of the presurstylus, in particular, demonstrates cially reduction of setae, that obscures and makes determina- this relationship. Assuming this relationship to be accurate, it is tion of phylogenetic relationships difficult. The structures of apparent that this species then underwent considerable anagen- the male terminalia, however, not only clearly differentiate this esis, resulting in the autapomorphies, such as the reduced chae- species but also demonstrate a close relationship with P. calva totaxy, that characterize this species. 94 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Phylogenetic Considerations ondarily reduced in some genera of Dryxini, whereas in Notiphilini there are 3 setae (1+2), a synapomorphy for The tribe Dryxini, which is one of five tribes now placed in Notiphilini); (4) costa elongate, extended to vein M (the costa the subfamily Hydrelliinae (Mathis and Zatwarnicki, 1995), is short, extended only to vein R4+5, in Notiphilini, a synapo- appears to be most closely related to the tribe Notiphilini (Zat- morphy for Notiphilini); (5) male terminalia with surstylus di- warnicki, 1992). We have identified five synapomorphies (in- vided into a presurstylus (surstylus) and a postsurstylus cluding Zatwarnicki's (1992) characters 20 and 21) that corrob- (clasper) (postsurstylus reduced or fused with epandrium with orate Dryxini's sister-group relationship with Notiphilini: (1) only presurstylus present in Notiphilini, a synapomorphy for ventral anepisternal seta elongate, twice length of dorsal seta Notiphilini); (*6) presurstylus with apex angulate and bifur- (secondarily reduced in a few taxa); (2) midtibia with promi- cate; (*7) pre- and postgonite reduced and/or lacking (the nent, erect, extensor setae along dorsal surface (Zatwarnicki's structure remaining may represent a fused and/or reduced pre- character 20); (3) only reclinate fronto-orbital seta well devel- and postgonite); and (*8) hypandrium connected basally with oped, proclinate seta(e) reduced or lacking; (4) abdominal terg- postsurstylus, not with epandrium. ites fasciate (secondarily reduced in some taxa); and (5) sub- In the presentation on genus-level relationships that follows, epandrial plate reduced (Zatwarnicki's character 21). the characters used in the analysis are noted first. Each charac- The following characters distinguish Dryxini from Notiphil- ter is immediately followed by a discussion to explain its states ini and confirm the tribe's monophyly (synapomorphies are and to provide perspective and any qualifying comments about noted by an "*"): (*1) gena high (secondarily short in some that character. After presentation of the information on charac- species); (*2) face wide, transversely arched, and generally ter evidence, a hypothesis of the cladistic relationships is pre- projected anteriorly (the face in Notiphila is comparatively sented and briefly discussed. The cladogram (Figure 154) is the narrower and much flatter); (3) dorsocentral setae 4(1+3; sec- primary mode to convey relationships; the discussion is to sup-

11 23 37 -Notiphila I I I 15 30 34 41 -D-D-OB—Afrolimna 112 1 29 34 40 43 44 -Paralimna

1111 10 •Paralimna (Phaiosterna)

14 18 20 25 31 34 41 • • • • • p-jl Corythophora 5 9 10 38 42 45 4 11 12 15 16 17 19 22 28 {XHX1IID •• 13 21 27 35 111111 12 1111111 •Dryxo 2 3 5 6 7 24 27 2 2 -o 1 2 1 1 2 1 18 • Omyxa 4 8 20 31 • Oedenops 1 26 32 35 1 2 2 1111 4 18 24 36 2 4 30 33 36 39 -Q • • •— Oedenopiforma -D-O-D- llllll 11 12 D-D—Papuama ismayi 2 1 •Papuama clava

FIGURE 154.—Cladogram depicting hypothetical cladistic relationships among genera of Dryxini (55 steps, con- sistency index 0.92, retention index 0.92) (open squares=ambiguous synapomorphies and/or autapomorphies; solid squares=unambiguous synapomorphies and/or autapomorphies). NUMBER 617 95 plement the cladogram and is intended only to complement the 7 (8). Number of aristal hairs: (0) arista bearing 7-14 dorsal latter. In the discussion of character data, a "0" indicates the hairs; (1) arista bearing 3-6 dorsal hairs (autapomor- state of the outgroup; a "1" or "2" indicates the derived states. phy for Oedenops). All multistate characters (5, 10, 11, 18, 20, 24, 27, 31, 34, 35, 8 (9). Shape of face: (0) face relatively narrow and flat; (1) and 41) were treated as nonadditive (-), and characters 8, 9, 13, face wide and transversely arched (autapomorphy for 14, 21, 23, 29, 32, 37, 38, and 42, which are autapomorphies tribe Dryxini). for various genera or tribes, were made inactive (]) for the 9(10). Height of gena (at ventral margin of eye): (0) rela- analysis and do not figure into the calculation of the consis- tively short; (1) relatively high (autapomorphy for tency index. The numbers used for characters in the presenta- tribe Dryxini); (2) secondarily short in Paralimna tion are the same as those on the cladogram, and the sequence (Phaiosterna) and in the limbata group of Paralimna. is the same as noted in the character matrix (Table 1). The ge- nus Notiphila, which is the only genus included in the tribe Thorax Notiphilini, was the outgroup in our phylogenetic analysis. 1(11). Number of dorsocentral setae: (0) 4 (1+3; the general- ized condition in Dryxini); (1) 3 (1+2) (autapomor- CHARACTERS USED IN THE PHYLOGENETIC ANALYSIS phy for tribe Notiphilini); (2) 1 (0+1; only posterior- (character number in parenthesis) most dorsocentral seta, which is displaced laterally, is present) (apparently homoplasious: synapomorphy for General Dryxo, Omyxa, and Corythophora; autapomorphy for Papuama ismayi). 1 (1). Color of head and thorax: (0) similar in both sexes; (1) 2(12). Presutural supra-alar seta: (0) present; (1) absent (ap- distinctly sexually dimorphic (synapomorphy for parently homoplasious: synapomorphy for Dryxo, Oedenops and Oedenopiforma). Omyxa, and Corythophora; autapomorphy for Pap- uama ismayi). Head 3(13). Prescutellar acrostichal setae: (0) 1 pair present, dis- 1 (2). Development of proclinate fronto-orbital setae: (0) tinct; (1) absent (autapomorphy for Dryxo). present, well developed, anterior seta usually larger 4(14). Shape of posterior margin of scutellum: (0) flat or than posterior seta; (1) absent or weakly developed nearly so; (1) broadly rounded (autapomorphy for (synapomorphy for Dryxo, Omyxa, Oedenops, Oede- Corythophora). nopiforma, and Papuama). 5(15). Setulae on venter of scutellum: (0) absent; (1) present 2 (3). Ocellar seta: (0) present (sometimes reduced); (1) ab- (synapomorphy for Dryxo, Omyxa, and Corytho- sent (synapomorphy for Dryxo and Omyxa). phora). 3 (4). Paravertical setae: (0) well developed, conspicuously 6(16). Number and position of notopleural setae: (0) 2, one in- evident although sometimes not long; (1) indistin- serted toward anterior angle, the other inserted toward guishable from other setulae or lacking (synapomor- the posterior angle; (1)1 seta, inserted centrally (syna- phy for Dryxo, Omyxa, Corythophora, Oedenops, and pomorphy for Dryxo, Omyxa, and Corythophora). Papuama). 7(17). Dorsally curved setula (sometimes weakly developed) 4 (5). Fronto-orbital setae: (0) both proclinate and reclinate near posterodorsal angle of anepistemum: (0) present; fronto-orbital setae well developed, although reclinate (1) absent. seta usually better developed; (1) only reclinate seta 8(18). Number of anepistemal setae: (0) 2, both subequal in well developed, proclinate setae reduced or lacking size (most Ephydridae); (1)2, with ventral seta much (an autapomorphy for the tribe Dryxini); (2) fronto- longer than dorsal seta (synapomorphy for tribes orbital setae absent (synapomorphy for Dryxo and Dryxini and Notiphilini); (2) 1, with dorsal seta Omyxa). greatly reduced (autapomorphy for Oedenopiforma); 5 (6). Width of parafacial: (0) narrow, width much less than (3) bearing 2 or 3 long, thin, hair-like setae along pos- length of 1st flagellomere; (1) wide, width greater terior margin (autapomorphy for Papuama); (4) all se- tae along posterior margin weakly developed (autapo- than length of 1st flagellomere (synapomorphy for morphy for Omyxa); (5) 1, with dorsal seta lacking Dryxo and Omyxa). (autapomorphy for Corythophora). 6 (7). Shape of frons: (0) shallowly arched anteroventrally, not projected forward, sparsely setulose; (1) projected 9(19). Proepisternal seta (stigmatal): (0) present; (1) very forward as a densely setulose, shield-like plate (syna- weakly developed or absent (synapomorphy for pomorphy for Dryxo and Omyxa). Dryxo, Omyxa, Corythophora, and Papuama). 96 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE I.—Matrix of characters and taxa used in the cladistic analysis of Dryxini (numbers for characters and character states correspond with those used in the text). 1111111122222222223333333333444444 Taxon 0000000001 1234567890 2345678901234567890123456789012345 Notiphila 000000000010000001000010000000000000100000000 Dryxo 21101121 101111 10110200210000002001000101 0 0 0 0 0 1 Omyxa 1 1 0 21001 1 14100102001 1000000000 1 0 0 0 0 0 1 Corythophora 0 0 0 0 0 0 210111151101001001001002000 1 0 0 2 0 0 1 Papuama ismayi 0 0 0 0 2 10 0 0 0 0 3 0000000000 0 0 0 0 10 1 1 0 0 0 0 1 Papua ma clava 0 0 0 0 00000003 0000000000 0 0 0 0 10 1 1 0 0 0 0 1 Oedenops 0 0 0 1 0 0 0 0 0 0 0 102000001000121 10 1 1 0 1 1 0 0 0 0 1 Oedenopiforma 0 0 0 0 0 0 0 0 0 0 0 0 200000101000 10 110 1 0 0 1 1 0 0 0 0 1 Afrolimna 0 0 0 0 0 0 0 0 0 0 0 10 0 1 00000000000 10 0 0 2 0 0 0 1 0 0 1 0 0 1 Paralimna 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0000000000 10 0 0 0 1 0 0 0 1 0 I 0 1 1 1 P. (Phaiosterna) 0 0 0 0 1 0 0 0 20000000 0000000000000001000 1 0 1 0 1 1 1

10(20). Katepisternal seta: (0) well developed; (1) reduced males (synapomorphy for Papuama, Oedenops, (autapomorphy for Corythophora); (2) absent (auta- Oedenopiforma, and Afrolimna). pomorphy for Oedenops). 21(31). Erect setae along dorsum of midtibia (a synapomor- 11(21). Vestiture of katepisternum: (0) lacking row of 6-10 se- phy for the tribes Notiphilini and Dryxini): (0) 3 setae; tae posterodorsad of katepisternal seta; (1) bearing (1)4 setae (autapomorphy for Corythophora); (2) 2 row of 6-10 hair-like to large setae posterodorsad of setae (autapomorphy for Oedenops). large katepisternal seta (autapomorphy for Dryxo). 22(32). Setae of tarsomeres: (0) not bearing long setae; (1) 12(22). Costal setae at apex of subcostal break: (0) present, bearing long, anteroapical and posteroapical setae, prominent; (1) rudimentary, very short (synapomor- length of setae subequal to width of tarsomere at apex phy for Dryxo, Omyxa, and Corythophora). (synapomorphy for Oedenops and Oedenopiforma). 13(23). Costal vein: (0) long, extended to vein M; (1) short, extended to vein R4+5 (autapomorphy for tribe Abdomen Notiphilini). 1(33). Color pattern on abdomen: (0) fasciate; (1) unicolor- 14(24). R stem vein: (0) bare of setulae; (1) bearing 1-3 (usu- ous, lacking fasciate pattern (synapomorphy for Pap- ally 2) fine setulae along dorsum (autapomorphy for uama, Oedenops, and Oedenopiforma). Oedenopiforma); (2) bearing several fine setulae on 2(34). Appendages of aedeagus: (0) lacking a lateral aedeagal dorsum (synapomorphy for Dryxo and Omyxa). process; (1) with lateral aedeagal processes fused with 15(25). Vein R node: (0) bare of setulae; (1) bearing 2 or 3 aedeagus (an autapomorphy for the genus Paralimna); long setulae on dorsum (autapomorphy for Corytho- (2) with lateral aedeagal processes adjacent to but sep- phora). arate from aedeagus (synapomorphy for Afrolimna and 16(26). Color of wing membrane: (0) hyaline to faintly infus- apparently for Corythophora). cate, sometimes with distinct pattern; (1) lightly milky 3(35). Shape of apex of aedeagus: (0) with medial projection; white, especially toward base and alula (synapomor- (1) lacking medial projection (synapomorphy for phy for Oedenops and Oedenopiforma). Oedenops and Oedenopiforma); (2) projections re- 17(27). Length and shape of crossvein dm-cu: (0) nearly duced (autapomorphy for Dryxo). straight, relatively short; (1) shallowly sinuous, mod- 4(36). Shape of aedeagus: (0) simple, oval in outline in poste- erately long (autapomorphy for Omyxa); (2) conspicu- rior view, lacking appendages; (1) bearing posterodor- ously sinuous, long (autapomorphy for Dryxo). sal fold (synapomorphy for Oedenops and Papuama). 18(28). Leg length and width: (0) normally developed; (1) 5(37). Condition of pre- and postsurstylus: (0) male termina- long and slender (synapomorphy for Dryxo, Omyxa, lia with surstylus divided into a presurstylus (sursty- and Corythophora). lus) and postsurstylus (clasper); (1) postsurstylus re- 19(29). Vestiture of male forefemur: (0) lacking distinctive duced or fused with epandrium with only presurstylus setal adornment; (1) bearing row of distinctive, fre- present (autapomorphy for tribe Notiphilini). quently flattened and curved setae along anteroventral 6(38). Shape of apex of presurstylus: (0) presurstylus a sim- surface (autapomorphy for Paralimna {Paralimna)). ple to complex structure at ventral margin of epan- 20(30). Vestiture of forefemur: (0) lacking comb-like row of drium, apex not angulate and bifurcate; (1) presursty- short, stout, spine-like setulae; (1) bearing comb-like lar apex angulate and bifurcate (autapomorphy for row of short, stout, tooth-like setulae in males and fe- tribe Dryxini). NUMBER 617 97

7(39). Shape of presurstylus: (0) robust and well sclerotized; TABLE 2.—Analysis of characters based on the cladogram (figure 154) and (1) lobate and weakly sclerotized (synapomorphy for weights (varying between 1 and 10) and status of characters after successive weighing (+ = additive; - = nonadditive; [ = active; ] = inactive). Oedenops, Oedenopiforma, and Papuama). 8(40). Postsurstylus: (0) lacking setae: (1) bearing 2 setae Consistency Retention Character Steps index index Weight Status near apex (synapomorphy for Paralimna sensu stricto 1 1 100 100 10 +[ and Paralimna {Phaiosterna)). 2 2 50 75 3 +[ 9(41). Shape of aedeagal apodeme: (0) lacking an anterome- 3 1 100 100 10 +[ dial projection; (1) with a long, medial, apically blunt 4 2 50 75 1 +[ projection (autapomorphy for Afrolimna); (2) with 5 2 100 100 10 -[ 6 1 100 100 10 +[ long projection extended from near articulation with 7 1 100 100 10 +[ hypandrium (autapomorphy for Corythophora). 8 1 100 100 10 +] 10(42). Condition of gonite: (0) pre- and postgonites distinct 9 1 100 100 10 +] and separate (pregonite usually smaller and bearing 2 10 2 100 100 10 -[ 2 66 66 4 -[ apical setulae); (1) pre- and postgonite reduced and/or 11 12 2 50 66 3 +[ lacking (structure remaining may represent fused and/ 13 1 100 100 10 +] or reduced pre- and postgonite; autapomorphy for 14 1 100 100 10 +] tribe Dryxini). 15 2 50 66 10 +[ 11(43). Position of gonite relative to postsurstylus: (0) gonite 16 1 100 100 10 +[ 17 1 100 100 10 +[ positioned at apical VA-VZ of postsurstylus; (1) gonite 18 3 100 100 10 -[ positioned at apex of postsurstylus (synapomorphy for 19 2 50 75 3 +[ Paralimna sensu stricto and Paralimna {Phaio- 20 2 100 100 10 -[ sterna)). 21 1 100 100 10 +] 22 1 100 100 10 +[ 12(44). Shape of ventral margin of postsurstylus: (0) straight 23 1 100 100 10 +] or shallowly curved; (1) conspicuously sinuous (syna- 24 2 100 100 10 -[ pomorphy for Paralimna sensu stricto and Paralimna 25 1 100 100 10 +[ {Phaiosterna)). 26 1 100 100 10 +[ 27 2 100 100 10 -[ 13(45). Connection of hypandrium posteriorly (basally): (0) 28 1 100 100 10 +[ hypandrium connected posteriorly with epandrium 29 1 100 100 10 +] and aedeagal apodeme; (1) hypandrium connected ba- 30 2 50 75 10 +[ sally with postsurstylus, not with epandrium (autapo- 31 2 100 100 10 -[ 32 1 100 100 10 +] morphy for tribe Dryxini). 33 1 100 100 10 +[ 34 3 66 50 3 -[ 2 100 100 10 -[ Analysis, Results, and Conclusions 35 36 2 50 50 2 +[ Using the implicit enumeration (ie*) option of Hennig86, 37 100 100 10 +] 38 100 100 10 +] which is an exhaustive search, nine most parsimonious trees 39 100 100 10 +[ were generated from the analysis of the 45 characters. The cla- 40 100 100 10 +[ dograms have a length of 55 steps and consistency and reten- 41 100 100 10 -{ tion indices of 0.83 and 0.83, respectively. A strict consensus 42 100 100 10 +] 43 100 100 10 +[ tree (nelsen) was then derived from the nine trees. The consen- 44 100 100 10 +[ sus tree has a polytomy of four lineages arising at the base of 45 100 100 10 +[ the tribe Dryxini, with Notiphila as the sister group. The matrix was then subjected iteratively to successive weighing (xs w, ie*, cc) to determine a character's contribution, or weight, and ter weights, the analysis of the resultant cladogram resulted in to find cladograms supported by the most consistent characters consistency and retention indices of 0.92 and 0.92, respectively. (Carpenter, 1988; Dietrich and McKamey, 1995). The succes- As indicated on the consensus cladogram (Figure 154), the sive weighing stabilized at 426 steps and reduced the number of tribe Dryxini is divided into four basal sublineages that form an trees from nine to six. A consensus of these six trees resulted in unresolved quadritomy. The first basal sublineage is the genus a tree that is identical to the consensus tree from the nine initial Afrolimna, and as would be expected from a basal lineage that trees. The consensus tree (Figure 154), although with unre- comprises a single genus, its relationship to other genera or solved lineages (polytomies), is our cladogram of choice. The groups of genera within Dryxini was the most unstable. Afrol- analysis of the characters for this cladogram and the weights of imna, in the initial nine trees, varied from being the sister group the various characters are given in Table 2. Given these charac- to each of the other lineages to being the sister group to com- 98 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY bined lineages, such as the genera related to Dryxo (Corytho- is known thus far from the Oriental (India) and southeastern phora, Dryxo, and Omyxa) or the genera related to Oedenops Palearctic (Iran, Oman) Regions. The taxa of this sublineage (Oedenops, Oedenopiforma, and Papuama). Afrolimna, which include some of the largest shore flies known, and by their size was described and has been treated as a subgenus within Paral- alone they are attractive material to collect and study. Unfortu- imna, is accorded generic status. Although somewhat subjec- nately, nothing is known about their immature stages. tive, our preferred relationship for Afrolimna is as the sister The fourth basal sublineage within Dryxini includes three group of the genera related to Dryxo, but the only evidence to genera: Papuama (2 species), Oedenops (3 species), and Oede- support this relationship is character 15, which is not compel- nopiforma (3 species). Like Paralimna, Oedenops occurs in ling. As alluded by its name, Afrolimna is known only from the both the Old and New Worlds. The three known species are Afrotropical Region, and only two species are presently quite widespread, with O. isis occurring in the Afrotropical, known. The monophyly of Afrolimna is corroborated by char- Australasian/Oceanian, Oriental, and Palearctic Regions, O. acters 15, 30, 34, and 41. afrus being found only in Africa, and O. nudus occurring only The second basal sublineage comprises Paralimna, includ- in the New World. Oedenops occurs primarily in tropical and ing the subgenus Phaiosterna. Unlike Afrolimna, Paralimna subtropical regions but has range extensions into temperate Ja- varied little in its relationship to the other genera. In six of the pan and North America. nine trees, Paralimna remained a separate and distinct lineage Our review of Oedenopiforma provides a basis for a more that was not closely related to other lineages within Dryxini substantive revision, including description of new taxa, as with two exceptions: (1) as the sister group to Afrolimna or, (2) noted previously. Oedenopiforma is known only from the Old as the sister group to a combined lineage of Afrolimna and the World, and there, primarily from Africa and Australia. The genera related to Dryxo. Paralimna occurs pantropically in the Australian species, O. uniseta, has a disjunct distribution from Old and New Worlds with considerable species diversity (85 the other species in Oedenopiforma, but a specimen from Sri described species), including numerous undescribed species in Lanka represents a link to African species. The Sri Lankan both areas. In our review of Paralimna, we recognized only specimen, however, may represent a new species or is a major two subgenera despite the exceptional species diversity that ex- range extension for O. argentea. Specimens of this genus are its in this genus. Our study of Paralimna is very preliminary, relatively scarce in collections, and the species are represented however, and with description of the numerous undescribed by very few specimens. Like most taxa of Dryxini, the imma- species and investigation of the phylogenetic relationships ture stages are unknown. among these species, a much more elaborated classification for Within Dryxini, most lineages at the generic level or higher, Paralimna will undoubtedly emerge. The monophyly of Para- including all basal lineages, occur exclusively or mostly in the limna is corroborated by characters 34, 40, 43, and 44. Old World tropics and subtropics. We are thus of the opinion The third basal sublineage is Dryxo and related genera that the tribe probably had it origins there, perhaps in Africa. {Corythophora and Omyxa). This sublineage is corroborated by From the Old World, there were independent extensions into several synapomorphies and autapomorphies (characters 4, 11, the New World by species of the genera Paralimna and Oede- 12, 15, 16, 17, 19, 22, and 28), as are the relationships among nops. In the New World, only Paralimna radiated and speci- these three genera. The three genera of this group include only ated to any extent, and there are now numerous species, de- 12 described species: Corythophora (2 species), Dryxo (9 spe- scribed and undescribed, that occur exclusively there. cies), and Omyxa (1 species). Corythophora is at the base of Although our evidence is meager (there is no cladogram of lin- this sublineage and is the sister group to the lineage comprising eages within Paralimna), there is some indication that more Dryxo and Omyxa. Corythophora occurs exclusively in the than one included lineage, such as species of the subgenera Afrotropical Region, and the monophyly of the genus is estab- Phaiosterna and Paralimna, expanded into the New World and lished by six unambiguous autapomorphies (characters 14, 18, did so independently. 20, 25, 31, and 41); both Dryxo and Omyxa are known only Numerous shore-fly taxa remain to be collected, described, from the Old World. The species of Dryxo are relatively more and analyzed. Within Dryxini, we note that Paralimna, in par- widespread, occurring primarily in tropical to subtropical zones ticular, is in need of species-level revision. Even though this within the Afrotropical, Oriental, and Australasian Regions. An genus already includes most of the tribe's species diversity, al- exception is D. nudicorpus, which occurs in temperate areas of most 80%, there are numerous undescribed species that have China, Japan, and eastern Russia. Omyxa, which is monotypic, already been segregated in existing collections. Literature Cited

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Werner 1980. 71: Family Ephydridae. In R.W. Crosskey, editor, Catalogue of the untemommenen zoologischen Expedition nach dem anglo-Agyptis- Diptera of the Afrotropical Region, pages 655—669, 1 map. London: chen Sudan (Kordofan) 1914, VI: Diptera. Denkschriften der Akad- British Museum (Natural History). emie der Wissenschaften in Wien, Mathematisch-Natur- 1984. Family Ephydridae. In A. Soos and L. Papp, editors, Catalogue of wissenschaftliche Klasse. 98(1923):57-82, 6 figures. Palaearctic Diptera, 10:126-176. Amsterdam and Budapest: 1926. 56a Ephydridae und 56b Canaceidae. In E. Lindner, editor, Die Elsevier Science Publishers and Akademiai Kiado. Fliegen der Palaearktischen Region, 6(1):1-115, 134 figures. Cogan, Brian Henry, and Willis Wagner Wirth Bezzi, Mario 1977. Family Ephydridae. In M.D. Delfinado and D.E. Hardy, editors, A 1908a. 6: Simuliidae, Bombyliidae, Empididae, Syrphidae, Tachinidae, Catalogue of the Diptera of the Oriental Region, Volume 3: Subor- Muscidae, Phycodromidae, Borboridae, Trypetidae, Ephydridae, der Cyclorrhapha (Excluding Division Aschiza), pages 321-339. Drosophilidae, Geomyzidae, Agromyzidae, Conopidae. In L. Honolulu: University Press of Hawaii. Schultze, editor, Zoologische und Anthropologische Ergebnisse Cole, Frank Raymond, with E.T. Schlinger einer Forschungsreise im westlichen und zentralen Sudafrika aus- 1969. The Flies of Western North America, xi+693 pages, 360 figures. gefuhrt in den Jahren 1903-1905; Erster Band: Systematik und Berkeley and Los Angeles: University of California Press. Tiergeographie, IV: Insecta (Erste Serie), D: Diptera (I). Denk- Coquillett, Daniel William schriften der Medicinisch-Naturwissenschaftlichen Gesellschaft zu 1900. Report on a Collection of Dipterous from Puerto Rico. 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