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Home , Acalyptratae, Argyra, Atrichomelina, Calliphoridae, Canto a Baja California, Dictya

The Marsh of (Di p t e ra: Sc iom yzi dae) by T. W. Fisher and R. E. Orth The Marsh Flies of California (Diptera: ) BULLETIN OF THE CALIFORNIA SURVEY VOLUME 24

THE MARSH FLIES OF CALIFORNIA (Diptera: Sciomyzidae)

T. W. FISHER AND R. E. ORTH

UNIVERSITY OF CALIFORNIA PRESS BERKELEY LOSANGELES LONDON BULLETIN OF THE CALIFORNIA INSECT SURVEY

Editorial Board: Ted Case, John Chemsak, John Doyen, Henry Hespenheide, T. A. Miller, John Pinto, Rudolph Pipa, Jerry A. Powell, Arthur Shapiro, Robbin Thorp

VOLUME 24 Issue Date: January 1983

UNIVERSITY OF CALIFORNIA PRESS BERKELEY AND LOS ANGELES

UNIVERSITY OF CALIFORNIA PRESS, LTD. LONDON, ENGLAND

ISBN 0-520-09665-7 LIBRARY OF CONGRESS CATALOG CARD NUMBER 82-1 3428 01983 BY THE REGENTS OF THE UNIVERSITY OF CALIFORNIA PRINTED IN THE OF AMERICA

LIBRARY OF CONGRESS CATALOGING IN PUBLICATION DATA

Fisher, T. W. The marsh flies of California.

(Bulletin of the California Insect Survey; v. 24) 1. Sciomyzidae-Classification. 2. -Classification. 3. Insects-California-Cldsification. I. Orth, R. E. 11. Title. 111. Series QL475.C3C3 VO~.24 [QL537.S365] 595.709794s 82-13428 ISBN 0-520-09665-7 r595.7741 Contents

Acknowledgments, vii Antichaeta borealis Foote, 29 Antichaeta robiginosa Melander, 29 Introduction 1 Antichaeta testacea Melander, 29 Antichaeta vernalis Fisher and Orth, 30 Biology 5 Evolution of Malacophagy, 6 Meigen, 1803: 277, 30 Biological Control of Vector Snails, 6 Dictya fontinalis Fisher and Orth, 31 Ecological Considerations, 7 Dictya inchCurran, 3 1 Natural Enemies of Sciomyzidae, 7 Dicwa montana Steyskal, 31 Mollusk Hosts of Marsh Flies, 8 Form “A”, 32 Sciomyzid Habitats in California, 11 Form “Alturas”, 32 Collection and Preparation Methods, 12 Form “Intermediate”, 32 Form “so. Cal.”, 33 15 Dictya texensis Curran, 33 Key to the American Genera of Sciomyzidae North of , 19 Genus Dictyacium Steyskal, 1956: 78, 34 Dictyacium firmum Steyskal, 34 Tribe Sciomyzini, 20 Genus Cresson, 1920:40, 20 Genus Elpiva Meigen, 1838: 365, 35 Atrichomelina pubera (Loew), 20 EIgiva connexa (Steyskal), 35 Genus Robineau-Desvoidy, 1830: 695, 21 solicita (Harris), 35 Pherbellia californica Orth, 21 Genus Hoplodictya Cresson, 1920: 67, 35 Pherbellia griseola (FallBn), 22 Hoplodictya acuticornis (Wulp), 36 Pherbellia idahoensis Steyskal, 22 Pherbellia melanderi Steyskal, 23 Genus Robineau-Desvoidy, 1830: 684, 36 nana (Fallen), 23 Limnia boscii (Robineau-Desvoidy) , 36 Pherbellia oregona Steyskal, 24 Limnia inopa (Adams), 37 Pherbellia jmrallela (Walker), 24 Limnia pubeseens (Day), 37 maculata (Cresson), 24 Limnia severa Cresson, 38 Pherbellia subtilis Orth and Steyskal, 25 Genus Hendel, 1900: 333, 38 Pherbellia trabeculata (Loew), 25 Renocera brevis (Cresson), 38 Pherbellia vitalis (Cresson), 26 Genus Latreille, 1804: 196, 38 Genus Lioy, 1864: 1012, 26 Sepedon bifidaSteyskal, 39 Pteromicra pectorosa (Hendel), 26 Spedon borealis Steyskal, 39 Pteromicra siskiyouensis Fisher and Orth, 27 Sepedon capelleiFisher and Orth, 39 Genus Fallen, 1820: 11, 27 Sepedon firscipennisfirscipnnis Loew, 40 Sciorqyza simpkxFallBn, 27 Sepedon pacifica Cresson, 40 Sciomyza varia (Coquillett), 28 Sepedon spintpes americana Steyskal, 41 Tribe Tetanocerini, 28 Genus Dumiril, 1800: 439, 41 Genus Antichaeta Haliday, 1838: 187, 28 Tetanocera ferrugirtea FallCn, 42

V vi CONTENTS

Tetanocera latifibula Frey, 42 Loew, 45 Tetanocera IoewiSteyskal, 43 Tetanocera soror Melander, 45 Tetanocera mesopora Steyskal, 43 Tetanocera vicina Macquart, 45 Tetanocera obnrsifibulaMelander, 43 Literature Cited, 47 Tetanocera plebeia Loew, 44 Ph&S, 5s Tetanocera plumosa Loew, 44 Acknowledgments

Persons from UCR who have provided assistance CDFA California Department of Food and in the laboratory or field are M. Badgley, C. Carter, Agriculture, Sacramento (M. Wasbauer) D. Clarke, J. Hall, M. Hansel, K. Lamb, R. CNC’ Canadian National Collection, Ottawa Medved, S. Nagahashi, E. Oatman, A. Strawn and CU‘ Cornell University, Ithaca, New York S. Swanson. Determinations of mollusks were pro- (C.0. Berg) vided by D. w. Taylor, Tiburon Center for KSU Kent State University, Kent, Ohio (B. Environmental Studies, Tiburon, CA, B. Roth, A. Foote) CAS, and the late W. 0. Gregg. To learn of and to LACM Los Angeles County Museum of Natural gain access to suitable habitats we consulted ranch- History, Los Angeles, California (C.L. ers, personnel of Abatement Districts, Hogue, R. J. Snelling) U.S. Forest Service, county Agricultural Commis- MCZ6 Museum of Comparative Zoology, sioners, California Department of Fish and Game. Harvard University, Cambridge, Their counsel and assistance is very much appreci- Massachusetts ated. The generous advice and critique throughout MNHNP’ Museum National d’Histoire Naturelle, this study of L. V. Knutson and G. C. Steyskal, Paris, France USDA, c/o U. S. National Museum of Natural His- NHMV’ Naturhistorisches Museum, Vienna, tory, C. 0. Berg, CU, and J. C. Hall, UCR, is espe- Austria cially appreciated. NRS Naturhistoriska Riksmuseet, Stockholm, The drawings were prepared by R. E. Orth and Sweden the microphotography was done by M. E. Badgley. OSDA Oregon State Department of Agriculture, Salem Oregon State University, Corvallis Sources of Material osu (P. Oman, Jack Lattin) This survey is based on approximately 24,000 PADA Pennsylvania Department of specimens examined by one or both authors, Agriculture, Harrisburg (K.R. Valley) including, when possible, confirmatory examination UCB University of California, Berkeley (J. A. of specimens recorded in the literature. The litera- Powell, E. I. Schlinger) ture search relating to this paper terminated Janu- UCD University of California, Davis (R. 0. ary 31, 1982. The assistance and cooperation of Schuster) individuals and institutions as indicated by the fol- UCR University of California, Riverside (S. lowing codes are gratefully acknowledged. Super- Frommer) script indicates the number of California for UI’ University of Idaho, Moscow (W. F. which an institution is a type depository. In the Barr) text, material collected by the authors is indicated UK’ University of Kansas, Manhattan (G. W. by F&O. Byers) AMNH2 American Museum of Natural History, USNM” U.S. National Museum of Natural New York History (L. V. Knutson, G. C. Steyskal) ANSP3 Academy of Natural Sciences, wsu Washington State University, Pullman Philadelphia (W.J. Turner) BMNH’ British Museum of Natural History, ZMUH’ Zoological Museum, University of London Helsinki, Finland CAS6 California Academy of Sciences, San ZMB Zoologisches Museum, Berlin, Germany Francisco (P. H. Amaud, Jr.) vii INTRODUCTION

A preliminary survey of Sciomyzidae in Califor- Our study of this group of insects began in 1962 nia by Fisher (1966) listed 31 species plus 10 taxa as part of University of California Agricultural suspected of being undescribed. This bulletin com- Experiment Station research project #2037, now piles available information on the taxonomy, biol- titled “Biological Control of Non-Marine ogy and geographical distribution of 49 species in Mollusks. ” 13 genera of sciomyzid flies known in California Adults of these rather slow-flying Diptera are and on 4 forms of DicQa montana. Also treated are seen resting on emergent rushes or grasses along 5 species from Oregon as reported by Fisher and the margins of flowing or standing fresh water. Orth (1975b), 2 from Nevada, and 1 from Arizona, They often perch with the head directed downward, which because of their proximity may be ultimately and with their enlarged hind legs and habit of sit- found in the state. The text also includes comments ting with the head held higher than the tip of the on the ecology and habitats of sciomyzids, their abdomen, present a -like aspect. Adults potential as biological control agents of snail inter- of the less hygrophilous, or mesic marsh flies, such mediate hosts of certain man- and -attacking as certain species of Limnia and Tetanocera, are trematodes, development of the malacophagous commonly found many meters from free water. habit in Diptera with particular reference to the Pinned specimens representing 12 genera found in Sciomyzidae and their mollusk hosts, and collection California (Renocera not shown) are shown in and preparation methods. Plate 1. Worldwide there are approximately 600 described Adults of the Sciomyzidae are dis- species of marsh flies. Steyskal (1965b) lists 144 tinguished from other acalyptrate Diptera by the species in 21 genera in North America north of following characters: oral vibrissae absent, post Mexico. Currently about 167 species are recognized vertical bristles diverging, costa entire, subcosta and 23 are Holarctic. Of the 49 species listed for complete, one or more tibiae with preapical bristles. California, 10 are Holarctic, and only one- Sepdon Plates 4-6 further identify these and other charac- bifida Steyskal -is restricted to California and adja- ters commonly used to identify marsh flies. ‘Body cent , Norte, Mexico. Table 1 lists length within the family ranges from 2 to 12 mm. genera and numbers of species in North America. Body color may be pale yellowish-brown, to gray to Knutson et al. (1976) lists 74 species in 23 genera black. from the poorly collected Americas south of the According to Hennig (1973: 551, the Sciomyzidae U.S.A. An excellent summary of studies involving together with the , , the Sciomyzidae worldwide is that by Berg and , Rhopalomeridae, and Coleopidae Knutson (1978). comprise the superfamily of the Commonly known as marsh, or swale, flies and acalyptrate Diptera. The suprageneric classification in older entomological literature as Tetanoceridae of the Sciomyzidae by Steyskal (1965a) recognizes or Tetanoceratidae, members of the family the subfamilies Salticellinae, Phaeomyiinae, Hut- Sciomyzidae occur in lakes (margins), ponds, tonininae, Helosciomyzinae,* and Sciomyzinae. swamps, marshes, vernal pools, wet pasture or Only the Sciomyzinae occur in North America, with mesic habitats where the larvae are obligate feeders both of its tribes represented as shown in Table 1. on mollusks. ‘These -killing sciomyzoids were removed from the family Sciomyzidae by Barnes (1980).

1 2 BULLETIN OF THE CALIFORNIA INSECT SURVEY

TABLE 1. SCIOMYZIDAE IN AMERICA NORTH OF MEXICO

To tal Species number of occurring species in Calif.

SCIOMYZINI A trichomelina 1 Colobaea 1 Oidematops 1 Pherbeilia 32 (7)? Ptwomlcra 14 (3) Sciomyza 4 (2)

TETANOCERINI Antichaeta 8 4 Dictyo 22 4 Dic(yacium 2 1 Elgiva 2 (1) 2 2 0 Hedria 1 0 Hoplodictya 5 1 Limnia 17 4 Pherbecta 1 0 Poecilographa 1 0 Renocera 6 1 Sepedomerus 1 0 Sepedon 18 6 Tetanocera 29 (10) 10 (5) 1 0 116 (11) 33 (5) Total Species 169 (23) 49 (10)

?Number of Holarctic species shown in parenthesis ( ). Boyes et al. (1969, 1972) summarized cytological Berg (1962, HopEOdiclya; 1966, Sepedon); Bratt et (karyological) and morphological features of larvae al. (1969, PherbeUrh); Foote (1959, Sciomyza; and adults which characterize the primitive 1961a, Tetamcera);Kaczynski et al. (1969, Perilim- Sciomyzini and the derived Tetanocerini. The pres- nia and South American Shannonrh); Knutson ence of a highly specialized sperm pump, or (1966, Antichaeta). ?cochleate vesicle,? prompted Steyskal and Knut- According to a list compiled by L. V. Knutson son (1975) to conclude that Ethiopian, Oriental, (correspondence) more than 40 entomologists and Australasian Sepedon and certain related genera worldwide are currently involved in sciomyzid are the most highly evolved of all Sciomyzidae. studies. Table 2 presents a generalized morphological com- By way of perspective from an approximate total parison of the two tribes. Species of Renocera and of 300 papers by 102 authors on all aspects of the Antichaeta show intermediate characteristics in the Sciomyzidae, 38% and 37% of the papers are con- pharyngeal sclerite. Plate 7 depicts characteristics of cerned with Nearctic or Palearctic species respec- immature Sciomyzini and Tetanocerini. tively, followed by Ethiopian with 7%, and Neo- Keys to the immature stages of many species of tropical, Australian, and Oriental with 6% each. Sciomyzidae include those published by Neff and MARSH FLIES OF CALIFORNIA 3

TABLE 2. GENERALIZED MORPHOLOGICAL COMPARISONS OF TRIBES SCIOMYZINI AND TETANOCERINI AS BASED ON NORTH AMERICAN SPECIES.

ADULT Sciomyzini Tetanocerini

Head not greatly modified greatly modified

Chaetotaxy poorly differentiated well differentiated

Midfrontal stripes poorly developed well developed

Male postabdomen strongly asymmetrical; symmetrical; one pair with 2 pairs surstyli surstyli well developed

Integument

pigmentation absent present.

spinule patches present absent

Accessory Teeth absent present

Pharyngeal Sclerite

anterodorsal bridge present absent

window in dorsal cornua present absent

indentation between cornuae deep shallow

*Absent in tcrmtrial species. The leading American taxonomic specialists on R. Valley, Pennsylvania Department of Agriculture, the Sciomyzidae worldwide are L. V. Knutson and Harrisburg, PA, New World Diciya. Specialists G. C. Steyskal, Systematic Laboratory, active in Palearctic species are K. Elberg, Institute U.S. Department of Agriculture, c/o U.S. National of Zoology and Botany, Academy of Sciences, Museum of Natural History. American specialists Tartu, Estonian S.S.R.; R. Rozkoh9, Depart- active in certain subgroups include B. A. Foote, ment of Biology of and Man, J. E. Purk- Department of Biological Sciences, Kent State yne University, Brno, Czechoslovakia, and A. University, Kent, Ohio; T. W. Fisher and R. E. Freidberg, Tel Aviv University, Israel. Orth, mainly western North American species; K.

BIOLOGY

Biologies of more than 200 of the approximately embryos. The second and third larvae feed 600 named sciomyzid species worldwide are par- on juveniles or adults of Succineidae and other tially to well known. Among the detailed studies snails. As reported by Foote et al. (1960) Atri- which have been published are those by Berg chomelinu puberu (Loew) is a versatile feeder. (19621, Bratt et al. (19691, Fisher and Orth (19641, Depending on circumstances, its larvae may exhibit Foote (1959, 1963, 1971, 1973, 1976, 19771, Foote , predatory, and saprophagous capabilities et al. (19601, Kaczynski et al. (19691, Knutson in response to available food and intensity of (1966, 1970a, b, 19731, Knutson and Abercrombie intraspecific competition. Larvae of species which (19771, Knutson and Berg (1963, 1964, 19711, attack terrestrial mollusks typically lack float hairs Knutson et al. (1965, 1967, 1970, 19751, Knutson and tend to exhibit greater host specificity than and Valley (19781, Neff and Berg (1961, 1962, aquatic larvae. Nearly all sciomyzid larvae, when 19661,Rozkohj. (19671, Rozkohf and Knutson feeding, maintain air contact through the caudal (19701, Trelka and Berg (19771, Trelka and Foote spiracles, but, as reported by Foote (1971, 1976) (19701. the larvae of Hedria miwta Steyskal and Renoceru (3 Sciomyzid larvae progress through three spp.) have the unusual habit of descending below and are obligate feeders on a wide variety of terres- the water surface in to prey on submerged trial and freshwater pulmonate snails plus certain snails, and sphaeriid clams, respectively. The latter prosobranchs (Hydrobiidae) and pelecypods are attacked by Knutsoniu Ifneutu (Fall&) in Europe (). Puparia may be formed within the according to Knutson (1970b1. This habit also has shell of the host, may be free floating with been reported in Dictya spp. by Valley and Berg upturned posterior spiracular tubes, or may be (1977). Fisher and Orth (1969b) reported that Dic- formed in nearby moist litter. tyo Jbntinulis F&O could not complete development Most Tetanocerini behave as overt predators, on , a pulmonate snail, but did so when first and while developing each larva kills and instar fed on hydrobiid snails in the genera consumes several snails. However, some aquatic Fontelicella, Lithoglyphus, or Tvoniu. Fascinating larvae and most of the terrestrial larvae of Tetano- restrictive host associations also were reported by cerini and most species of Sciomyzini exhibit res- Foote (19761 for three species of Renoceru (aquatic trictive host selection, and some properly may be Tetanocerini) that prey on submerged sphaeriid termed parasitoid in habit since development is clams, and Foote (19771 published the first biology completed on a single host individual. As reported on Oidetnutops (Sciomyzini), which is restricted to by Knutson et al. (19671 larvae of certain species of its host land snail, Stenobemu. Species of Tetano- Pherbelliu and Colobaea excrete a calcareous sep- cera (terrestrial Tetanocerini) which feed on lima- tum prior to pupating and seal themselves within cid were reported by Foote (19631, Knutson the shell of the host snail. Larvae of many et al. (19651, Knutson (1970b1, Trelka and Foote sciomyzid species may feed saprophagously, and (19701, Trelka and Berg (19771. larvae of some species change feeding habits from Such diversity of life style, obligate feeding selective to less selective as the stadia change. For habit, and habitat selection, evidence remarkable example, females of Antichaeta oviposit on egg degrees of adaptive radiation and provide reason clusters of their succineid snail hosts, and the first enough for continued study in this already inten- instar maggots feed on the developing snail sively worked insect family.

5 6 BULLETIN OF THE CALIFORNIA INSECT SURVEY

EVOLUTION OF MALACOPHAGY was a parasitoid of a land snail, Helix pi so^. It was Berg (19531, however, who first conclusively The record has not revealed when the demonstrated that sciomyzid fly larvae-six species actual connection between sciomyzid flies and their in five genera of his initial study-were capable of mollusk hosts occurred. The stage was set over a attacking, killing, and consuming several species of very long geologic time span for this relationship to fresh water snails. Berg (1964b), basing his opinion evolve, because the first land snails appear in on 143 species reared at Cornel1 University, Devonian deposits of the Paleozoic Era, 350 million hypothesized that the Sciomyzidae probably evolved years ago. Subsequently, certain of the pulmonate from a general similar in habit to Atri- gastropods entered the freshwater habitat and chomlina pu&era (Loew) , which gradually became the ancestral stock from which modern developed the habit of feeding on stranded still freshwater pulmonate snails evolved. The earliest living snails in moist situations. Divergence then evidence of modern insects appears in Permian may have given rise to both the aquatic predators deposits of the late Paleozoic Era, or 215 million and terrestrial . Rozkdnf (19671, report- years ago. Berg and Knutson (1978:250) ?regard ing on the dipterous fauna of central Europe, stated the species of fossil ?Sciomyzidae? described in that apart from the general malacophagous family Sciomyza and Tetanocera from the Oligocene and Sciomyzidae (51 species verified) there are only 10 Miocene of Western Europe and Western North malacophagous larvae in four other America by authors [such Cockerel1 early as families- (21, (21, Sar- (190911 as species inquirende. Hennig (1965, 1969) cophagidae (51, and (1). Trelka and described six species of Sciomyzidae from Baltic Foote (1970:893) suggest a possible evolution of Amber [lower Oligocene1 in considerable those species of Tetanmra which feed on slugs detail.. . .? Thus, considering the time span beginning with feeding on hygrophilous snails, involved, it seems reasonable to presume that gradually shifting to slugs in the same moist habi- malacophagy was in place by the early Cenozoic tat, then shifting (as in T. chra Loew) to Era, or 60 million years ago, and probably occurred species found mainly in more mesic habitats. before that during the Cretaceous period of the late Mesozoic Era which was characterized by great swamps and cooling climates. BIOLOGICAL CONTROL OF VECTOR SNAILS The phenomenon of malacophagy in Diptera has In their review of problems associated with the been considered by several entomologists. Schmitz biological control of medically important snails, Bay (1917) made one of the first attempts to deal with et al. (1976) correctly cited the continuing lack of biological relationships between flies and snails. interdisciplinary communication and cooperation Other workers such as Bequaert (1925) and John- among qualified experts on biological control son (1929) suspected that certain snails were methods and specialists on trematodes and snails as actually killed by sarcophagid fly larvae, and even the main reason for the paucity of research effort earlier Keilin (19191-based on his masterful study applied to date in this important field. of the calliphorid, cognata-proposed a sys- The potential use of sciomyzid flies for biological tem for classifying Diptera associated with gastro- control of snail vectors of trematode-caused pod mollusks. Lundbeck (1923) reared 26 species diseases was recognized by Berg (1953, 1964a), of sciomyzid flies from puparia, 3 of which were Berg et al. (19551, and Neff (1964). bray (1964) found in snail shells, and illustrated the puparia of abandoned his project on use of sciomyzid flies to Colobaea bifasciella Falldn and Ctenulus pectoralis control tomntosa, the snail vector of Zetterstedt within the shells of their respective livestock liverfluke in Australia. His findings indi- aquatic snail hosts Lymnaea [= Fossarkzl truncatuh cated that the main predator fly, Dichaetophora hen- and vortex. Seguy (1950: 459-461) de& Kertk, could not dominate so ecologically presumed that Diptera oviposited only on sick or diverse a host. Lynch (1965) reported that in the dead mollusks although his Fig. 195 illustrated cer- Mt. Lofty Ranges and the Murray Lakes regions of tain tetanocerid parasitoids of European snails. South Australia, Dichaetophora biroi (Kertk) was Mercier (1921) provided the ktclue that a considered to be a major predator of L. tomentosa. sciomyzid fly, Salticelb fasciala Meigen, probably He agreed with bray?s findings and reported this MARSH FLIES OF CALIFORNIA 7

fly was effective in reducing localized populations of Beaver (1972; 1974a, b) studied comparative biolo- pulmonate snails during periods of receding water gies and inter- and intraspecific competition of levels, but was ineffective in standing or slowly several species of British Sciomyzidae. Geckler flowing water six inches or more in depth. Fontana (1971) in a laboratory study reported that larvae of (19721, in another Australian study, concluded that Sepedon tenuicornis Cresson consumed 8.1 f 2.5 the larvae of D. biroi do not discriminate between aquatic snails ( ancepd of varying sizes healthy or infected L. tomentosa and that infected with an approximate volume of 544 f 256mm3 snails are not more vulnerable than non-infected before pupation. Capture-recapture studies by snails to predation by larvae of D. biroi. Arnold (1976) with Sepedon f: fuscipennis at Cor- Berg (1971, 1973) reviewed literature pertaining ne11 University showed that adults can overwinter to biological control of snail-borne diseases and and that diapause is triggered by diminishing day decried the lack of interest in exploring non- length in the fall. chemical means of vector control. A non-technical In Thailand a study by Bhuangprakone and treatment of the potential of sciomyzid flies in the Areekul (1973) involved feeding tests in which biological control of snail-borne diseases is that by nine species of aquatic snails were offered to larvae Knutson (1976). of Sepedon plumbella Wiedemann. These authors, So far the few attempts in the field to utilize without published supporting data, stated, “Out- these flies as biological control agents of snails breaks of the diseases (fascioliases) occur in areas which serve as intermediate hosts of human schis- where the insect has not been found.” tosomes or livestock liverflukes have met with In the rice paddy ecosystem of Japan, Nagatomi inconclusive results. One of the more encouraging and Kushigamachi (19651, Nishida and Torii reports was by Davis (1974) who stated that “in (19701, Momoi et al. (1979, and Yano (1975) the state of Hawaii on the islands of Hawaii, Maui, observed that the eggs of Sepedon sauteri Hendel and Oahu, slaughterhouse records indicate a down- [ = aenescens Wiedemannl served as alternate ward trend in the percentage of infested livers hosts for Trichogramma spp., which parasitize eggs between 1966 and 1972 and this strongly suggests of rice stem borers, very serious pests of rice. that the established sciomyzids Sepedon macropus [ = Sepedomerus macropus (Walker)] and S. sauteri NATURAL ENEMIES OF SCIOMYZIDAE Hendel [ - Sepedon aenescens Wiedemannl [ini- tially released in 1959 and 1967, respectively], may Exposed eggs and puparia are especially vulner- be exerting considerable pressure on liverfluke able to attack. Aquatic larvae are subject to attack snails.’’ [Our annotations in brackets.] Berg and from a wide range of predators including Hydra, Knutson (1978:253) state that H. K. Nakao in planaria, , Odonata, Coleoptera, and fish correspondence (November 18, 1976) reported a (particularly Gambusia), and the slow moving or similar decline on Kauai between 1972 and 1976. flying adults are easily seized by toads, frogs, spid- ers, Odonata, robber flies, etc. The list of parasitic insects which emerge from ECOLOGICAL CONSIDERATIONS sciomyzid puparia is substantial, and most are Laboratory and field studies on population polyphagous. Five families of parasitic Hymenop- dynamics operating in predator-prey relationships of tera (Braconidae, Diapriidae, , certain aquatic Sciomyzidae, mainly with Sepedon f: Pteromalidae, Trichogrammatidae) are collectively firscipennis Loew, and snails have been published by noted by Foote et al. (1960), Neff and Berg (19661, Eckblad and Berg (1972), and Eckblad (1973). Bratt et ai. (1969), and Knutson and Abercrombie Their studies indicated that since snails in the (1 977). Recorded genera include Braconidae: genus Lymnaea spend more time at the surface, Aphaereta, Aspilota, Phaenocerpa; Ichneumonidae: they are more likely to be attacked by the typically A tractodes, Eriplanus, Mastrus (?I,Mesoleptus, surface feeding sciomyzid larvae than are species of Phygadeuon; Pteromalidae: Eupteromalus, Spalan- snails which typically remain submerged. It was gia; Diapriidae: Phaenopria; Trichogrammatidae: further shown that the presence of small juvenile Trichogramma. Neff and Berg (19661, also report snails is critical to the survival and development of virus and bacterial diseases of Sepedon puparia, and first instar sciomyzid larvae in those species studied. hymenopterous predators of puparia and adult flies. 8 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Eggs and larvae of most species of marsh flies mollusks suspected of being natural hosts such as were rarely encountered in the field in California. those observed with sciomyzid eggs attached to the The eggs of Spedon paciEca Cresson are easily shell or body, or whose empty shells contained observed but of the several hundred we have seen sciomyzid puparia. The remainder are known hosts, and removed to the laboratory for observation, or those observed in nature actually behg fed upon none was parasitized. In 1963-64, we reared an by sciomyzid larvae. Their names are not followed undescribed uniparental ichneumonid , Phyga- by any qualifying initials in the table. Natural host deuon sp., from puparia of S. pacifica collected at associations are lacking for 19 of the California Vail Lake, Riverside Co., and several successive species of marsh flies. Certain of these have been all-female generations were cultured on house fly reared in the laboratory by Berg?s students at Cor- puparia in the laboratory. ne11 University on snail genera represented in Cali- Detailed biologies of two diapriid , Tri- fornia and are followed by ?L? in Table 3. chopria popei (Muesebeck) which attacks pupae of In general, sciomyzid larvae do not successfully aquatic Sciomyzidae, and T. atrkhomelinae Muese- attack mature land or freshwater snails in excess of beck which attacks pupae of terrestrial Sciomyzidae, 20 mm width or length. However, juveniles of were published by O?eil1 (1973). these snails are subject to attack, especially by second or third instar sciomyzid larvae. The critical importance of eggs and early juveniles of certain MOLLUSK HOSTS OF MARSH FLIES mollusks to the development of newly hatched There appears to be an abundance of host mol- sciomyzid larvae was mentioned earlier. lusks in California which might serve as food for By no means intended as a complete list, the fol- larvae of sciomyzid flies. A check list of California lowing references will provide a general background freshwater mollusks by Taylor (1981) contains 91 on North American non-marine . Those ?described forms.? According to Roth (1972) Cali- which contain treatment of certain species found in fornia has a rather large endemic fauna of terres- California are followed by an asterisk (*I: Bequaert trial mollusks of which 127 species, largely because and Miller (1973)+, Burch (1960, 1962; 1975a, b*), of their restricted numbers or locations, may even- Harman and Berg (1971)*, Hanna (19661, Malek tually be considered endangered. In addition, (19621, Patterson (1970, Taylor (1966, 1970, Hanna (1966) lists 36 terrestrial and 5 freshwater 1975, 1981)*. mollusk species as introductions into western North America. Mollusk Behavior Our field notes replete with ?associated? are With the possible exception of coastal northern species of flies and snails, but accurate are linkages California, habitats and periods of activity of most yet to be learned for most California species. Shells native land snails are closely synchronized with of snails commonly encountered in California are California?s irregular rainfall which must occur in shown in Plate 2. sufficient quantity to soak into the ground (as at the The excellent and detailed host records cited by base of boulders or rock slides) deep enough to Bratt, Eckblad, Foote, Knutson, and Neff Berg, reach and consequently activate dormant snails. were, in most instances, to the species level, but Therefore, these mollusks are not likely targets of since virtually all of these are eastern U.S. (i.e., attack by sciomyzid flies. Certain snails such east of the Mountains) we elected to cite as Rocky species of Vespericoka occur in damp habitats along only their genera in our Table 3. desert canyon streams and possibly could be hosts The listing of mollusk genera only, as in Table 1 to certain mesic oriented sciomyzids in the genera of Foote (19771, appears to offer reasonably accu- Limnia and Tetanocera. rate representation of natural host associations, Succineid (amber) snails are very common since it is doubtful that any sciomyzid larvae are throughout the state and seasonally can be species specific in nature. extremely abundant on wet mud banks of flowing In our Table 3 the term ?probable host? refers or standing water. The tetanocerine genus to mollusks known to be associated in nature with Antichuetu contains several species known to attack these flies and which in laboratory feeding tests amber snails and their eggs in nature. appear to be seiected by the larvae. These include TABLE 3. SCIOMYZIDAE FOUND IN CALIFORNIA AND THEIR KNOWN OR PROBABLE MOLLUSK HOSTS IN NATURE

SCIOMYZINI Atrkhomefinapuka AphNC Foote et a]., 1960 Helisoma Physa Lymnaea Physa virgata F&O tenuis Pherbellia californica NI Rherbellia grimla Lymnaea palusrris Bratt et al., 1969 Pherbellia idahoemis Helisoma P Bratt et a]., 1969 Lymnaea P phym Planorbella tenuis F&O Lymnaea palustris Physa virgata Pkbellia melanderi NI Pherbellia nana nana Gyraulus Bratt et al., 1969 Lymnaea Physa Pherbellia oregona NI Pherbellia parallela Lymnaea Bratt et al., 1969 Physa Physa virgata F&O Pherbellia schoenherri maculata Catinella Bratt et al., 1969 Oxyioma Succinea Pherbellia subtilis Lymnaea Bratt et al., 1969 Pherbellia Pabeculata Helisoma Bratt et al., 1969 Physa Phnorbula NC Planorbella tenuis F&O Succinea calif&rniensis Pherbellia vitalis Helisoma Bratt et al., 1969 Lymnaea Physa Pianobula NC Prerornicra pectorosa NI Pteromicra sisklyouensis NI Sciomyza simplex Lymnaea lom ma Foote, 1959 Sciomyza varia Lymnaea Knutson, 1962

TETANOCERINI Antichaeta borealis Owloma (eggs) Robinson and Foote, 1978 Antichaeta robiginosa NI Antichaeta testacea Owloma (eggs) P Fisher and Orth, 1964 Succinea P Antichaeta vernalis NI F&O Dicw fbntinafis Lithogbphus tlrrbinformis P F&O Dicw incisa NI Dicva montana Lymnaea palustris F&O Pbso virgoto TABLE 3. SCIOMYZIDAE FOUND IN CALIFORNIA AND THEIR KNOWN OR PROBABLE MOLLUSK HOSTS IN NATURE

Dictya texensis Physa virgata F&O Dictyacium Jirmum NI Elgiva connexa NI Elgiva wlicita Lymnaeidae Knutson and Berg, 1964 Physidae Hoplodictva acuticornis Succinea Fisher & Orth, 1972b Limnia boscii NI Limnia inopa NI Limnia pubescens NI Limnia severa NI Renocera brevis Foote, 1976 Sepedon bifida Heiisoma Neff and Berg, 1966 Phya Lymnaea palustris F&O Planorbella tenuis Sepedon borealis Lymnaea P Neff and Berg, 1966 Physa P Sepedon capellei N1 F&O Sepedon fuscipennisfuscipennis Helisoma Berg, 1953 Lymnaea Lymnaea palustris Eckblad and Berg, 1972 Physa integra NC Sepedon pac&a Planorbella tenuis F&O Physa virgata Lymnaea palustris Sepedon spini?s americana Heiisoma Neff and Berg, 1966 Physa Tetanocera ferruginea Helisoma Foote, 1961a Lymnaea palustris Tetanocera latifibula Physa L Foote, 1961a Tetanocera loewi Helisoma L Foote, 1961a Lymnaea palustris L Physa L Tetanocera mesopora NI Tetanocera obtusifibula Lymnaea L Foote, 1961a Tetanocera plebeia Deroceras laeve Trelka and Foote, 1970 Deroceras reticulatum Wloma Tetanocera plumosa Helisoma L Foote, 1961a Lymnaea palustris L Physa L Tetanocera robusta Gyraulus L Foote, 1961a Helisoma L Lymnaea L Physa L Tetanocera soror Physa L Foote, 1961a Tetanocera vicina Gyraulus L Foote, 1961a Lymnaea L

NOTE: FBO Fisher & Orth, unpublished rearing; L - Laboratory mor& only; NC = does not occur in California; NI - no information; P 9 probable hosts. MARSH FLIES OF CALIFORNIA 11

Although we have no direct proof, it is probable SCIOMYZID FLY HABITATS IN CALIFORNIA that Teronoceru plebem Loew attacks slugs in Cali- The classification of aquatic plant habitats pro- fornia as it does in the eastern U.S. and Europe. posed by Mason (1957: 2-5) could serve as a model Behavior of aquatic snails is governed by various for categorizing sciomyzid fly habitats. However, aspects of their free water habitats. Water and sub- two problems are inherent in nearly all interpreta- strate temperatures appear to be parameters that tions of sciomyzid habitat designations. (1) The override other physical factors such as light and diurnal movement of adult flies may place them chemistry within broad limits. many meters from their oviposition sites or host At the onset of winter in fairly stable bodies of mollusks. (2) General designations such as lake, water falling temperatures cause aquatic snails to stream, meadow, etc., can be misleading, because it become inactive. Those remaining exposed on the is the more narrowly defmed subhabitat that is of surface of the substrate become easy prey for immi- specific interest. grating water fowl, but that segment of the popula- In general, grassy banks of streams or lakes, tion which has become attached to submerged sedge meadows with substantial growth of Eleo- rocks, limbs, etc., below the mud line is safe from ckris spp. or Jurtcm spp. are indicators of poten- attack. In the spring with rising water and bottom tially good collecting sites for sciomyzid flies. The mud temperatures these snails become active and presence of easily discernible aquatic or terrestrial form the source of the extremely rapid build up of mollusks does not guarantee good fly collecting. the snail population commonly witnessed early in Conversely, the apparent absence of mollusks does the year after water fowl have emigrated. not preclude the presence of sciomyzids. These cir- In seasonally wet streams, ponds, or farm reser- cumstances could reflect hodpredator population voirs, etc., the fantastic ecological diversity of snails fluctuations, or merely that the flies had moved to is demonstrated. To survive the snails must cope more amenable resting areas during the heat of the with not only seasonal drying but with scouring and day. flooding during heavier than usual periods of pre- In nearly all California lake situations two areas cipitation. a body of water begins to recede, As stand out as most productive of adult Sciomyzidae: many snails burrow deep into the mud, probably (1) banks of streams where they flow into the lake, secrete epiphragms the soil dries further, and as and (2) below the foot of the dam where seepage remain inactive until the ground is again flooded or outflow may create a permanent marsh-like habi- and and the rising water substrate temperatures tat. In these situations, 'TLpha, Scupus, Eleocharis, reach approximately In situation mechan- 50°F. this and cress afford resting places for adult isms which trigger auto-exhumation may include Hyubco~l, flies. The submerged portions of such emergent widely fluctuating temperatures in shallow the vegetation along with aquatic plants such as water, or changing total salts content of the water. Chara, Potomgeton, tape grasses, By burrowing deep the snails assured of at Myriop@llum, as are well as benthic objects provide substrates for least some moisture for the later summer and fall aquatic snails. Along the lake shore, embayments months, and at that depth will also be protected may afford protection from wind induced wave from mild scouring during the winter months. This action, thus permitting aquatic snails to adhere to interpretation probably is accurate in streams of low submerged fallen limbs, rushes, close to the gradient which collect silt. We have dug 18" into rocks, shore. several out stream beds where we knew high dried The banks of rivers and canals did not prove to populations existed indicated by the presence of as be good collecting sites. Overflow areas may exposed shells, and our knowledge of the site, and become marsh or pond habitats of fair stability and have unearthed very few At the Santa Ana snails. the upper reaches and tributaries may offer good River in Riverside, PIanorbeh Helbmal has I- meadow or stream bank habitats. act been observed in the of emerging from the mud Stream habitats are highly variable in California. bottom following rain. Yet digging in that spot ear- of many streams, or sections thereof, are dry lier revealed no snails. Even such seemin&ly safe Beds during the summer and fall months. This is often habitats can be totally scoured out during periods of true of streams below 2,500 feet elevation in south- heavy runoff, and the habitat may be many years ern California and below 1,500 feet in the eastern rebuilding. 12 BULLETIN OF THE CALIFORNIA INSECT SURVEY foothills of the Central Valley. Above 4,000 feet in a particular site. If the vegetation was wet from elevation, or latitudes north of 36", substantial dew or rain the D-Vac was the only collecting volume of water may flow the year around. method to yield undamaged flies. The D-Vac was Seeps and springs can create habitats that are especially useful in situations where it was difficult suitable for some aquatic snails, e.g., P&m, Lym- to swing a net, such as short vegetation (especially naea, and hydrobiids, as well as terrestrial species. when windy) and dense rushes, or grasses beneath The same type of habitat can occur in or along thin low branches of shrubs or trees. It was common trickles of water that emanate from such spring practice with two people collecting for both the D- areas. The fringes of such areas may be classed as Vac and the net to be used simultaneously. In this mesic since free water is not in evidence. event, the sweep net was frequently everted into Mountain meadow habitats can be the result of the D-Vac while it was still running. agricultural or forest management practices, or may The malaise trap was tried, but the return in represent natural undisturbed situations. Large sciomyzids was not encouraging. These flies are expanses of alpine sedge meadows early in the rarely taken in light traps. season are not highly productive of flies; yet as they The wearing of hip boots proved to be the most dry out in late summer and fall, the green practical way to thoroughly sample most sites. They "islands" that remain can offer extremely permitted one to concentrate more on where the productive collecting. net was swinging than on where to step, and were a Of necessity much of the sampling occurred in convenience when kneeling in shallow water or roadside ditches and along streams, or stream beds, mud when searching for mollusks and immature over which roads passed. Near urban areas and in sciomyzids. areas of heavy crop agriculture, we learned not to Preparation of flies consisted of placing two or expect great success in collecting sciomyzid flies, three bags of field collected material loosely within even if the habitats looked right and mollusks were a 10" x 25" plastic bag. Approximately 5 ml of ethyl present. We suspect the lack of flies in these cases acetate were poured on the muslin collar of one of was due to insecticides used for mosquito or the bags and the plastic bag sealed shut for approxi- nuisance insect abatement and/or presence of mately 15 minutes to kill the insects. Placing the insecticides in run-off water from crop areas. plastic bag over a lamp shade or in the sun for a Examples of some of the more productive sites few minutes greatly speeded the action of the fumi- are shown in Table 4. Also indicated is the number gant. The material was then sorted in small lots of Sciomyzini and Tetanocerini recorded from each under a dissecting microscope, and sciomyzid flies site. Photos of certain of these plus other sites can were pinned and placed in Schmitt boxes with be seen in Plate 3. detailed head labels per site.

Mollusks COLLECTION AND PREPARATION METHODS Land snails and slugs were collected in the sam- ple area during a hands-and-knees search, turning Flies over debris, shallowly buried rocks, etc. In early Our most used sampling method involved aerial morning collections mollusks were sometimes on sweep nets. The contents were emptied into a hold- damp vegetation and fell into the sweep net. Shells ing bag. Large bits of debris, large insects, and of commonly collected snails are shown in Plate 2. especially spiders were disposed of, and the bag Preparation of terrestrial mollusks involved first lightly sprinkled with water and placed in a chilled drowning them in water-usually overnight. Adding picnic ice chest pending preparation the evening of approximately 5% alcohol greatly speeded the nar- the same day. cotizing process. It is important to exclude an air A gasoline-powered D-Vac* suction collector was pocket from the killing jar. Mollusks can then be used, particularly if the nets turned up some placed in 1W formaldehyde for a few hours before interesting material, or if the nets did not yield transferring to 70% alcohol. To be properly materia) we had reason to thiik should have been prepared, the foot should be nearly fully extended. *Manufacturedby D-Vac Co., P.O.Box 2095, Riverside, CA 92506. MARSH FLIES OF CALIFORNIA 13

TABLE 4. SCIOMYZIDAE OF CALIFORNIA: SPECIES COLLECTED AT SELECTED SITES IN REPRESENTATIVE BIOTYPES.

Coastal Interm. G.Bas.’ Montane‘

N* C* S’ N C S N C S N C S

SCZOMYZINI Atrichomelina pubera Pherbe~iiagri~eoia”~

nana nana subtilis oregona parallela Propages schoenherri maculata trabsculata vitalis

siskfyouensis sciom siplex varia

ETANOCER INZ Antichaeta boreal&*6 rob&inosa

vernalis Dicw fontinalis incls~’.~ mntana “A” ‘‘Alturas”

“in&rtned. ”

“so. Cal. ” texensis Dic@acium $rtmd6 Elgiw connexa 14 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Coastal' Interm.2 G.b.' Montane'

N' N' C'S' N C S N C S N C S MARSH FLIES OF CALIFORNIA 15

NOTES TO TABLE 4:

1. Coastal influence: N - Del Norte County./ Crescent City, I5 m (50 feet) elev. C * Sonoma Co.; Valley Ford, 30 m (100 feet) elev. S - Orange Co./ San Juan Cr., 76 m (250 feet) elev. 2. Intermediate climate: N - Mendacino Co.; 2 mi NIWillits, 405 m (1,330 feet) elev. C 9 Kern Co.; Lake Isabellr, 808 m (2.650 feet) elev. S = Riverside Co.; Santa Ana Riv., Riverside, 259 m (850 feet) elev. 3. Great Basic influence: N - Modoc Co.; Alturas, 1,311 m (4,300 feet) elev. C - Inyo Co.; Deep Springs Lake, 1524 m (5,000 feet) elev. S = Riverside Co.; Lake Hemet, 1,372 m (4.500 feet) elev. 4. Montane influence: N = Nevada Co.; Boca Springs, 1829 m (6,000 feet) elev. C - Alpine Co.; Heenan Lake, 2,134 m (7,000 fedelev. S - San Bernardino Co.; Cienega Seca, 2,362 m (7.750 feet) elev. 5. Highly restricted distribution. Collected at only one locality, or at 2-4 localities within a 5-10 mile radius. 6. Not collected at the 12 localities used in this Table. 'N = northern counties; C - central counties; S * southern counties.

Aquatic mollusks were gathered in two ways: (1) TAXONOMY hand picking from the substrate, or (2) vigorously dunking bunches of submerged plants to which Introduction mollusks were adhering in and out of a pail full of water. The plants and debris were removed and the Early in the study G. C. Steyskal expressed the water slowly poured from the bucket, leaving the opinion that the area of California north of 39' lati- snails in a compact group which could be later tude would likely yield material of greater interest sorted to species or poured en masse into a jar. The in terms of new or rare species. He was right. We relaxing of aquatic snails is accomplished by adding then arbitrarily designated counties lying roughly small amounts of propylene phenoxytol or menthol between 39-42' as Northern counties, those crystals to water in the collecting jar. Our resuits between 36-39' as Central counties, and those varied from poor to very good. When the snails between 32-36' as Southern counties. were thought to be narcotized, either a little for- Map 1 shows the counties included in each area. maldehyde was added to the jar or the water was Table 5 indicates the total number of species poured off and replaced with 70% alcohol. With iso- recorded per county. The Great Basin eco-climate is lated populations of mollusks malacologists want loosely interpreted. Perhaps "influence" is a more the collecting site described to the exact quarter accurate word. Common sagebrush (Artemisia tri- section. dentaiu Nutt.) was present. Other than the fact we were looking specifically In southern California we were able to make for sciomyzid flies, a reason why we appear to be monthly collections at four localities between 1962 the major, sometimes sole, collectors of the rare and 1966. These localities were: (1) Santa Ana species is that we made use of the D-Vac collector. River, Riverside; (2) Lake Hemet; (3) San Juan Much of the sciomyzid material from California Creek, 3 miles east of San Juan Capistrano; and (4) other than ours was collected by staff and students Big Bear Lake (Boulder Bay). During the course of from UCB, UCD, and UCR. A major contributor study the character of these sites was drastically was Dr. Paul Arnaud, CAS. altered by periods of excessive cold, drought, or precipitation, such that year to year comparisons could not be interpreted with confidence. Collec- tions in central and northern counties occurred mainly during the summer. Monthly collections in a few of those localities could be highly desirable in 126 124 122 120 118 116 114 NORTHERN I I I I 1 I COUNTIES

42- l2 CENTRAL COUNTIES 14 -4 1 11 34 ALAMEDA 21 NAPA 4 28 ALPINE 23 SACRAMENTO 13 27 AMADOR 41 SAN BENITO 7 31 CALAVERAS 32 SAN FRANCISCO 8 29 CONTRA COSTA 30 SAN JOAQUIN 3 24 EL DORAW 33 SANMATEO 18 45 FRESNO 36 SANTACLARA 19 48 INYO 35 SANTA CRUZ 40- lo 46 KINGS 26 SOLANO 17 44 IIADERA 20 SONOMA - 41 6 25 MARIN 37 STANISLAUS 2 47 TULARE 15 38 TUOLUMNE 9 22 YOLO 5 16

38 - - 3t

36 -

SOUTHERN COUNTIES

58 IMPERIAL 50 KERN 54 34 LOS ANGELES - 55 ORANGE 56 RIVERSIDE 51 SAN BERNARDINO 57 SAN DIEGO 49 SAN LUIS OBISPO 52 SANTABARBARA 53 VENTURA

32 - - 32 1 Map 1. California Counties. MARSH FLIES OF CALIFORNIA 17

TABLE 5. SCIOMYZIDAE OF CALIFORNIA: Numbers of species and forms recorded per county, and dominant eco-climate in areas sampled.

Northerncounties. c* i m gb 5 San Francisco X (39-42" approx.) 5 SanMateo X 4 Napa X 4 Stanislaus X 30 Siskiyou X 3 S8KltaCm X 29 Modoc X 2 Madera xx 29 Shasta X X 2 Solan0 xx 24 X Plumas 1 SanBenitO X 22 Mendociao X X 1 San Joaquin X 19 Nevada X 0 X 19 Sierra X Ki 18 Lassen X 14 Placer X X Southerncounties c i m gb 13 Dei Norte X (3 2-36" approx . 10 Humboldt X 9 Butte X 13 Riverside xx 7 Tehama X X 13 Saa Bernardino xx 7 Trinity X 11 Orange X 6 Glenn X 10 SanDiego xx 5 Colusa X 9 Los Angeles X 5 Lake X 9 SanLuisObispo x x 4 Yuba X 8 Kern xx 0 Sutter X 8 SantaBarbara xx 4 Ventura xx Centralcounties c i m gb 3 Imperial X (36-39" approx.) *C = coastal; i - interior; m - montane; gb - 21 Mono xx Great Basin. 18 Inyo X xx 16 Alpine X 13 Tulare X X 12 Marin X 12 Santa Clara X X 11 El Dorado X X 11 Fresno X X 11 Sonoma X X 10 Monterey X X 9 Tuolumne X 8 Amador X 7 Alameda X X 7 Sacramento X 7 Merced x 6 Yo10 X 5 Calaveras X X 5 Contra Costa X X 5 Mariposa X X 18 BULLETIN OF THE CALIFORNIA INSECT SURVEY

TABLE 6. SCIOMYZIDAE OF CALIFORNIA: Summary of collection data to September 1977.

Number of Counties Recorded

Northern Central Southern Total Day collected Elevation (meters) 39-42? 36-39? 32-36? Years (1 9) (29) (10) (58) Earliest Latest Spanned Low High

SCIOM YZINI A trichomelina pubera 13 14 7 34 19-11 7-XI1 1914 1974 3 2164 Pherbellia caljfornica 5 2 0 7 3 1-111 4-VI11 1947 1968 2 1494 Pherbellia griseola 1 0 0 1 16-VI 8-VI11 1968 1976 1262 - Pherbella idahoensis 8 2 0 10 23-IV 9-IX 1947 1974 76 1561 Pherbellia melanderi 1 0 0 1 9-VI - 1966 - 1311 1463 Pherbellia nana nana 19 20 8 48 10-111 22-XI1 1896 1976 3 2134 Pherbellia oregona 2 0 0 2 10-VI 5-VI11 1965 1969 3 15 Pherbellia parallela 12 12 10 34 14-1 27-XI1 1921 1976 3 2329 P. schoenherri maculata 6 3 0 9 26-IV 22-IX 1966 1976 1067 2789 Pherbellia subtilis 2 3 2 7 16-111 12-VI1 1966 1970 2 2408 Pherbellia trabeculata 0 0 4 4 13-11 13-IX 1962 1968 61 2057 Pherbellia vitalis 9 13 6 28 25-11 2 1-1x 1908 1975 -55 2362 Pteromicra pectorosa 1 0 0 1 11-VI - 1966 - 15 - Pteromicra siskiyouensis 3 0 0 3 6-VI 6-VI11 1965 1973 808 1561 Sciomyza simplex 2 1 0 3 9-VI 21-IX 1965 1974 1219 1561 Scioniyza varia 1 2 0 3 25-V 23-VI11 1966 1969 30 1920

TETANOCERINI Antichaeta borealis 1 0 0 1 10-VI1 - 1968 - 1414 - Antichaeta robginosa 3 3 0 6 17-1 10-VI1 1963 1974 146 1524 Antichaeta testacea 7 7 6 20 27-1 15-XI 1954 1974 8 2134 Antichaeta vernalis 2 0 0 2 23-IV 8-VI 1966 1968 405 1494 Dictya fontinalis 4 1 0 5 22-IV 22-IX 1951 1974 30 1798 Dictya incisa 0 0 1 1 24-IV - 1935 - 305 - Dictya montana ?A? 15 13 0 28 3-11 20-x 1937 1976 6 2621 D. montana ?Alturas? 5 0 0 5 8-VI 2 1-1x 1955 1974 933 1494 D. montana ?intermed? 7 13 2 22 10-111 1s-x 1948 1974 9 1387 D. montana ??so.Cal.? 3 10 10 23 1-I 3 1-XI1 1910 1975 -58 2530 Dictya texensis 0 0 6 6 8-1 27-XI1 1957 1976 -55 2362 Dictyacium Jirmum 1 0 0 1 24-VI 1-VI11 1973 1976 1219 - Elgiva connexa 1 2 0 3 6-VI 24-VI11 1965 1972 1311 2164 6 1 0 7 6-VI 22-IX 1965 1974 15 1561 Hoplodictya acuticornis 9 14 7 28 9-1 27-XI1 1908 1974 3 1646 Limnia boscii 0 1 0 1 11-VI1 7-IX 1967 1968 1295 - Limnia inopa 8 12 0 20 15-V 6-XI 1935 1973 305 2377 Limnia pubescens 2 0 0 2 17-VI 31-VI1 1913 1976 809 1219 Limnia severa 10 10 1 21 15-V 7-IX 1918 1975 2 2743 Renocera brevis 1 0 0 1 20-VI1 - 1966 - 1250 - Sepedon bfida 4 7 9 20 14-1 27-XI1 1917 1975 3 2362 Sepedon borealis 8 6 0 14 7-VI 22-IX 1918 1976 930 2377 Sepedon capellei 9 5 1 14 14-IV 16-X 1922 1976 73 2682 Sepedon f: fuscipennis 2 0 0 2 8-VI 22-IX 1966 1967 1067 1506 Sepedon pacifica 16 22 9 47 16-1 13-XI 1907 1976 3 3048 MARSH FLIES OF CALIFORNIA 19

TABLE 6. SCIOMYZIDAE OF CALIFORNIA: Summary of collection data to September 1977.

Number of Counties Recorded Northern Central Southern Total Day collected Elevation (meters) 39-42“ 36-39“ 32-36” Years (19) (29) (10) (58) Earliest Latest Spanned Low High

S. spinipes americana 1 0 0 1 9-VI1 22-IX 1966 1968 1506 - Tetanoceraferruginea 2 1 0 3 10-VI 21-IX 1966 1974 1219 1875 Tetanocera latifibula 7 1 0 8 7-VI 21-IX 1949 1974 1334 1783 Tetanocera loewi 1 0 0 1 11-VI 5-VI11 1966 1968 18 - Tetanocera mesopora 6 0 0 6 9-VI 2 1 -1x 1949 1972 1067 1561 Tetanocera obtusifibula 8 3 0 11 16-V 30-IX 1918 1975 2 1524 Tetanocera plebeia 7 1 0 8 8-VI 12-IX 1910 1976 3 1524 Tetanocera plumosa 14 14 2 30 1-V 4-x 1930 1976 152 2743 Tetanocera robusta 3 0 0 3 9-VI 21-IX 1966 1975 9 1768 Tetanocera soror 5 2 0 7 9-VI 3-VI11 1948 1976 1219 1920 Tetanocera vicina 8 0 0 9 8-VI 21-IX 1947 1975 808 1768 Total number of Sciomyzini 15 10 6 Tetanocerini 33 22 10 48 32 16

order to better understand seasonal activity of KEY TO THE AMERICAN GENERA OF marsh flies and their mollusk hosts. Warm springs SCIOMYZIDAE NORTH OF MEXICO areas such as Hot Creek (Mono Co.), Bishop (Inyo 1. Propleuron with a strong bristle above base of Co.), around Alturas (Modoc Co.), all in areas typi- fore coxa, if bristle inconspicuous, first seg- cally of cold, sub-freezing winters, would be of ment of fore tarsus whitish contrasting with great in terest. remaining segments. SCIOMYZINI ...... 2 Table 6 indicates extremes of collection data Propleuron without strong bristle above base of from all of the material we examined as well as cer- fore coxa; first segment of fore tarsus not tain records from the literature and specialists in strongly contrasting with remaining seg- correspondence. ments. TETANOCERINI...... 7 Our keys to genera and species are designed as 2. Fore tibia with 2 preapical bristles ...... 3 aids to identification rather than as indicators of Fore tibia with 1 preapical bristle ...... 4 natural classification. 3. Aristal hairs long, black; face without a central protuberance in upper half. (Holarctic) ...... Certain species found in the western U.S. are ...... Sciomu FallCn included in keys by Cresson (19201, Melander Aristal hairs dense, short, white; face with a (19201, Foote (1961~1,and Cole (1969). central protuberance in upper half. (Nearctic). In the narrative for each species is given the ori- ...... Oidemotops Cresson ginal name; publication containing original descrip- 4. Vein a, not attaining wing margin; body length tion; type locality; sex of holotype; depository; under 2.5 mm. (Holarctic) ..... Colobaea Zetterstedt synonymy; geographic range and California records. Vein a, attaining wing margin; body length usu- Under “Remarks” are summarized only the results ally over 2.5 mm ...... 5 of our efforts which is thus intended to indicate the 5. Propleural bristle fine and thin; center of pro- comparative abundance of the species, general habi- pleuron, nearly entire mesopleuron and ster- nopleuron, and center of pteropleuron with tat notes, etc. “Discussion” attempts to interpret or expand on certain facts mentioned in “Remarks.” fine Iongish hairs; fore coxa without bristles. 20 BULLETIN OF THE CALIFORNIA INSECT SURVEY

(Nearctic and Neotropical) ...... 18 ...... Abichomelinu Cresson 18. One fronto-orbital bristle; face with central Propleural bristle long and stout; 1 or more of black spot. (Holarctic and Neotropical).: ...... the above pleural regions without hehairs, ...... Diem Meigen fore coxa with at least 1 bristle ...... 6 Two fronto-orbital bristles; face without central 6. Predominately shining black species; frons usu- black spot. (Nearctic and Neotropical)...... ally entirely shining. (Holarctic) ....Pmmicm Lioy ...... Hopl~~wuCresson Species of yellow, brown, or gray color, never 19. Aristal hairs long, plumose; wing densely reticu- shining black; frons not wholly shining. late. (Holarctic) ...... Euthycera Latreille (Cosmopolitan)...... Pherkllia Robineau-Desvoidy Aristal hairs moderately short or pubescent; 7. Vallar bristles present ...... 8 wing densely reticulate or streaked with dark Vallar bristles absent ...... 1 1 shadings ...... 20 8. Arista with sparse blackish hairs ...... 9 20. Face with central protuberance in upper half; Arista with whitish hairs. (Holarctic) ...... &tal hairs moderately short, plumose; wing ...... Limnia Robineau-Desvoidy densely reticulate. (Nearctic) ...... 9. Nearly entire front between the orbitals convex ...... Dic@aciumSteyskal and glossy; 3 dorsocentrals; body yellow with Face without central protuberance in upper half; large black spots. (Nearctic) ...... aristal hairs minute or pubescent, not plu- ...... poecilographu Melander mose; wing not reticulate. but streaked with A large part of front dull; 1 or 2 dorsocentrals dark shadings. (Holarctic) ...... Elgiw Meigen ...... IO 10. Midfrontal stripe pruinose; prosternum with hairs. (Holarctic) ...... Trvpetopdwa Hendel TRIBE SCIOMYZINI Midfrontal stripe shining; prosternum bare. (Nearctic) ...... PkkmSteyskal Genus Atrichomelina Cresson, 1920:40 11. Ocellar bristle absent; postocellar bristle well- developed or lacking...... 12 AtrichomelinaCresson, 1920...... Ocellar bristle present; postocellar bristle The single species, A. pubem (Loew), may be well-developed ...... 14 characterized as follows. Propleural bristle hairlike; 12. Postocellar bristle well-developed; midfemur fore basitarsus white and other tarsal segments with one or more distinctly larger anterior black; fore coxa without bristles; all thoracic pleu- setae near midlength of femur...... 13 rites haired but without bristles. Widespread Postocellar bristle absent; midfemur without distinctly larger anterior setae near midlength throughout California. Male terminalia as in Plate of femur. (Nearctic and Neotropical)...... 8, Figs. 1, 2...... Sepe&merus Steyskal 13. Frons with 2 fronto-orbital bristles; posterior Atrichomlina pubera (Loew) crosmein bisinuate. (Nearctic)...... H&ia Steyskal Map 2; Plate 8, Figs. 1, 2 with only 1 Frons fronto-orbital bristle; poste- Sciotnyza prbacr Loew, 1862:106-107; Middle States; holotype rior crossvein arcuate. (Cosmopolitan)...... m, MCZ no. 13208...... sepedon Latreille Atrichomlina puh; Cresson, 192040 (NW to Achaetomlina 14. Hind tibia with 2 strong preapical bristles. in key, p. 30). (Holarctic) ...... Antirhaeta Haliday Hind tibia with only 1 strong preapical bristle .... 15 Geogmphic range: CANADA:Alberta to Quebec. Mmax Dis- 15. Second antennal segment less than 113 length trito Federal and Chiapis. USA: coast to coast. of third segment. (Holarctic) ...... Renocero Hendel Second antennal segment at least 112 length of Calfornia records (Map 2): widespread, recorded in 34 coun- third segment...... 16 ties. 16. Arista with long blackish hairs 17 ...... Remarks: We examined material from 10 the Arista with whitish hairs ...... 19 of 17. Wing mostly hyaline, may have some spots or 35 states where the species has been reported. In bands; body uniformly yellowish to reddish California we collected 414 m, 622 f which often brown; mesopleuron bare; face without cen- were in drying habitats with stranded yet living tral black spot. (Holarctic) ...... Tetanoceru Dum6ril planorbid, physid, and/or lymnaeid snails. Adult Wing not mostly hyaline, heavily marked with habitus as shown in Plate 1, Fig. 1. Head profile as blackish reticulations in Plate 5, Fig. 1. MARSH FLIES OF CALIFORNIA 21

Plate 9, Figs. 3, 4 ...... P. mlanakriSteyska1 D. Aristal hairs medium length. Wing length: 5 males, 4.6-5.0 mm, average 4.9; 5 females, 4.6-5.2 mm, average 4.9. Humboldt and Del Norte Co.; Pacific Coast. Male terminalia as in Plate 10, Figs. 3, 4 ...... P. oregona Steyskal Mesopleuron entirely bare ...... 4 4. Wing distinctly patterned, apical part of wing darkened with crossbands ...... 5 Wing not patterned, at most anterior margin and crossveins clouded ...... 6 5. Cell R, with 3 to 7 crossbars; 1st section of 4th vein with a stump vein posteriorly; blackish orbito-antennal spot well developed. Male ter- minalia as in Plate 12, Figs. 1, 2 ...... P. mbecuhta (hew) Cell R, without crossbars (preapical) crossband only, vein M without stump vein; orbito- antennal spot not well developed. Male ter- minalia as in Plate 10, Figs. 1, 2 ...... P. nana nana (Falldn) 6. Vein R, distinctly surpassing level of ta; arista bare, at most minutely pubescent. Male ter- minalia as in Plate 11, Figs. 4-6 ...... L_.-I -'-*"JJ I,, ", u ___--I" ,. It ,. In ...... P. subrilis Orth & Steyskal Map 2. California distribution of Atrichonrelina putwa Vein R, not surpassing level of tu; arista with (Loew) short blackish hairs ...... 7 7. Pteropleuron with a cluster of moderate sized Genus Pherbellia Robineau-Desvoidy, 1830: 695 bristles; upper edge of sternopleuron with Pherbellia Robineau-Desvoidy, 1830 ...... several short hairs and 2 or 3 much longer 1. Frons with median stripe extending 2/3 or more and stronger bristles. Male terminalia as in of the distance from anterior ocellus to frontal Plate 11, Fig. 1...... P. cafiBrnicaOrth margin ...... 2 Pteropleuron with several small bristles and 2 Frons without distinct median stripe or with one longer and heavier bristles; upper edge of ster- less than 2/3 as long as distance from ocellus nopleuron with short hairs only ...... 8 to frontal margin ...... 3 8. Orbito-antennal spot well developed; fore femur, 2. Mesopleuron bare; 2 parafrontal bristles; wing tibia, and tarsus blackish; anterior margin of cells with single rows of blackish spots. Male frons not cinereous. Male terminalia as in terminalia as in Plate 11, Figs. 2, 3 ...... Plate 12, Figs. 3, 4 ...... P. vitalis (Cresson) ...... P. schoenherri macuhta (Cresson) Orbito-antennal spot lacking; fore femur, tibia Mesopleuron setulose; 1 parafrontal bristle; wing and tarsus yellowish to brownish; anterior margin cells not spotted. Male terminalia as in Plate of frons bright cinereous. Not yet discovered in 10, Figs. 5, 6 ...... P. parallela (Walker) California. Known from Ormsby County, 3. Mesopleuron with setulae along posterior margin Nevada, adjacent to the California-Nevada (P.fuscipes group) border. Male terminalia as in Plate 8, Figs. 3, 4.... A. Aristal hairs short. Wing length: 5 males, ...... P. argyra Verbeke 4.6-4.9 mm, average 4.9; 5 females, 4.6-5.2 mm, average 4.9. Shasta Co. Male terminalia Pherbellia caljfornicaOrth as in Plate 8, Figs. 5, 6 ...... P. griseola (Fallin) Map 3; Plate 11, Fig. 1 B. Aristal hairs long plumose. Wing length: 10 males, 5.1-6.0 mm, average 5.7; 10 females, Plterbellia californica Orth, 1982: 23-37; Willits, Mendocino Co., 5.6-6.8 average 6.2. Male terminalia as California; holotype m., USNM. mm, in Pherbellia propagesSteyska1, 1966:37-38, in part. Plate 9, Figs. 1, 2 ...... P. idahoensis Steyskal C. Aristal hairs moderately long. Wing length: 8 Geogmphic range: USA: north of 38" latitude, California, males, 4.6-5.6 mm, average 5.2; 8 females, Oregon, Washington. 4.9-5.9 mm, average 5.4. Male terminalia as in 22 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Cul@rnia records (Map 3): EL DORADOCO.: Echo Lake. HUM- BOLDT Ca: Orick. MARIN Co.: Bolinas, San Quentin Point. Tomales. MEND~CINOCo.: 7 mi (11.3 km) north Hopland, Ukiah, Willits. PLUMASCa: Crescent Mills, Rock Creek. SHASTA Co.: Anderson. Sisuuou Ca: Fort Jones.

- -.- - &- .- _L .* ." 1_1 Map 4. California distribution of Pherbellia idahoensis Steyskal Remarks: We examined material from California

. -- -- ~ .-- - -- __ wt..,,+J and Oregon. In California we collected 30 m, 29 f . ,u I at Little Bear Flat in shaded mesic grassy areas on Map 3. California distribution of Pkrkllia caluornica Orth the east side of the meadow. P. griseola, P. idahoensis, P. mehnderi, and P. oregona are in the Remarks: We examined material from California P. jkcips group of Steyskal (19611, and can be and Oregon. In California we collected 102 f, 224 separated with certainty only by examination of m. The largest numbers came from 2 mi (3.2 km) male terminalia. Head profile as in Plate 5, Fig. 3. north of Willits, and 7 mi (11.3 km) north of Hop- land. P. propages occurs east of 105" longitude. Pherbellia idahoensis Steyskal Map 4; Plate 9, Figs. 1, 2 Pherbellia griseola (FalMn) Pherbellia idahoensis Steyskal, 1961:411; Moscow, Latah Co., Map 5; Plate 8, Figs. 5, 6 Idaho; holotype m, USNM. Sciomyza griseolu Fall& 1820: 14; Scandinavia; cotypes m, f, NRS no. 401/66, slides no. 6126-128. Geographic rarge: CANADA:southern British Columbia. USA: Sciomyza fuseipes Macquart, 1835: 407; Elberg and Rozkoh;ljr California north of 36" latitude, Idaho, Oregon, Montana, Utah, 1978: 51. Washington.

Melina griseola; Melander, 1920: 314. California records (Map 4): BUTTECO.: Honcut. LAKECo.: Pherbellia griseola; Steyskal, 1954~:55; 1961:411 (redescribed). Clear Lake Oaks. LASSENCO.: Smith Flat Reservoir. MENDWINO CO.: Willits. Mom CO.: Alturas, Hackamore Reservoir. PLUMAS Geographic range: EUROPE.CANADA and USA: widespread Co.: Crescent Mills. SANTACLARA Co.: Stanford Lake. SHASTA Mills. SISKIYOUCO.: Grass Lake. TULARECo.: north of 40" latitude. Co.: Fall River Woodlake. California records (Map 5): SHASTACO.: 2 mi (3.2 km) south of Pondosa (Little Bear Flat), 4140 feet (1262 m) elevation, V1- Remarks: With the exception of a single speci- 16-1969, VI-24-1976. VII-8-1970, VII-25-1972, VIII-1-1973, men from Silver Creek State Park, Klamath Co., VIII-8-1968 (F&O). Oregon, we examined material only from California MARSH FLIES OF CALIFORNIA 23

- -,^I.*-. [L__l__d-__L__--- _-__ Z%4] _LA_.- - ,. ,I, /. Y, 9. ," I" , I" m ,* "# Y. u ", * .* I. .,s .. I" Map 5. California distribution of Pherkllkz mhnderi Map 6. California distribution of Pherbeilia nana nana Steyskal e; and Pkrbeliia griseola (FallCn) A (FallCn) where we collected 47 f, 46 m. Most of the speci- Pherbellia nana nana (Falldn) mens were from Grass Lake, Siskiyou Co. Map 6; Plate 10, Figs. 1, 2 This is the largest Pherbellia in California and Sciom mm Fall6n. 1820: 12 Scandinavia, cotypes m, f, NRS. was collected by sweeping aquatic vegetation Mehm (Gmpha@nd mm; Cresson, 1920:47. emerging from shallow water along shores of lakes Pherbellm mm; Steyskal, 1956 13. or ponds. Pk&tl& (Ckroceml nuna nam Elberg, 1918:208. Geogruphic range: EUROPE ASIA MEXICO:Distrito Federal. Pherbellia melanderi Steyskal NORTHAMERICA: Alaska, British Columbia to Quebec, south to Map 5; Plate 9, Figs. 3, 4 Florida, west through Texas to California and Oregon. Pherbe//ia melunderi Steyskal, 1963: 117; Three Forks, Gallatin Co., Montana; holotype rn, USNM. California records (Map 6): widespread, in all but the deserts, recorded from 48 counties. Geographic range: USA: California, Colorado, Montana, Ore- gon, Utah. Remarks: We examined material from 10 of the 41 states and provinces where it has been reported. Cal$ornlu records (Map 5): MODOCCO.: Alturas, Cedarville, In California we collected 555 f, 869 m. Davis Creek, VI-9,lO-1966(F8rO). Discussion: This is the most ubiquitous sciomyzid in North America. An Asian subspecies n. Remarks: We examined material from Califor- fly P. reticulata (Thornson, 18681, was designated by nia, Colorado, Oregon, and California Utah. In we Elberg (1978:208). Further study may justify addi- collected 5 f, 6 Visits to the Modoc Co. sites m. tional subspecific assignment(s) of North American and others in that general area later in the summer material. and later in other summers in other years yielded no flies of this species. Therefore we think P. melanderi is an early season species. Adults superficially resemble P. griseola, being of same coloration, but are slightly larger. 24 BULLETIN OF THE CALIFORNIA INSECT SURVEY

- py IL ~ L_ . ..%+A I ~ _-_-,,. ". sm I <* - Map 7. California distribution of Pherbellia oregorta Map 8. California distribution of Pherbellia parallela Steyskal (Walker) Pherbellia oregona Steyskal Geographic range: Nearctic to Neotropical. CANADA.COSTA Map 7; Plate 10, Figs. 3, 4 RICA. MEXICO.USA: widespread coast to coast. Pherbellia oregona Steyskal, 1961:411-412; Forest Grove, Wash- California records (Map 8): widespread, recorded from 34 ington Co., Oregon; holotype m, UI. counties.

Geographic range: USA: California, Oregon, Washington. Remarks: We examined material from 11 of the 32 states and provinces where it has been reported. Califbrnia records (Map 7): DELNORTE CO.: Crescent City, 50 feet (15 m) elevation, VI-1 1-1965, VI-1 1-1966, VIII-5-1968 In California we collected 828 f, 1315 m. Largest (F&O). High Prairie Creek, Hwy 101, 20 feet (6 m) elevation, numbers were collected along margins of lakes, VIII-5-1968 (F&O). HUMBOLDTCo.: Little River State Beach, reservoirs, ponds. Adult habitus as shown in Plate sea level, VI-10-1965 (F&O). 1 mi (1.6 km) south of Orick, VI- 1, Fig. 2. 11-1965 (F&O). Samoa Dunes, VI-10-1969 (J. Powell, CIS). Pherbellia schoenherri maculata (Cresson) Remarks: We examined material from California Map 9; Plate 11, Figs. 2, 3 and Oregon. In California we collected 4 f, 8 m. In our 6 collections P. oregona occurred with 7 other Musca punclaia Fabricius, 1794: 341, preocc. Poda 1761. species of marsh flies, comprising approximately 6% Sciomyza schoenherri FallCn, 1826: 13. Sciompa monilis Meigen, 1830: 17. of the total. Pherbellia wrnalis Robineau-Desvoidy, 1830: 696. Melina (Graphomyzina) maculata Cresson, 1920: 48; Illinois; Pherbellia parallela (Walker) holotype m, ANSP no. 6222. Map 8, Plate 10, Figs. 5, 6 Melina schoenherri; Melander, 1920:316. Pherkellia schoenkrri; Steyskal, 1949: 177. Sciomyza pamllela (Walker) 1853a (1852k401; USA; holotype Pherbellia scknherri macukria, Steyskal, 1965b: 686. m, BMNH. Pherbelim punciata; Elberg and Rozkoini, 1978:Sl (in part). Sciomyza humilis Loew, 1876:330. Pherbellk? schoenherrc Rozko&;ljr. 1981: 179. Pherbellia humilis. Steyskal, 1963: 122. Pherbellia parallela; Knutson, 1981: 336. Geographic range: NORTH AMERICA:Coast to coast north of 37" latitude to Alaska and Yukon, east to Newfoundland. MARSH FLIES OF CALIFORNIA 25

i

y_ __ _~___ _. _-_. -__ -- --% - . --+J I&-- * UI ,_1 Y "I-- I Map 9. California distribution of Pherbellia schoenherri Map 10. California distribution of Pherbellia subrilis macularu (Cresson) Orth and Steyskal

Cal&rrnkr records (Map 9): ALPINECa: Hope Valley, Wood- Cal&rnm records (Map IO): ALPWE CO.: Hope Valley, Wood- fords. INYOCo.: 12 mi (19.2 km) west of Big Pine. Onion Valley. fords. FRESNOCa: Sequoia Lake. ~NYOCO.: Big Pine. MARIN LASSEN Co.: 2 mi (3.2 km) west of Hallelujah Junction, Clear Ca: Bolinas. MENDOCWOCa: Willits. NEVADACO.: Boca Spring. Creek. Mom CO.: Alturas, Lily Lake 7 mi (11.2 km) east of RIVERSIDECa: Lake Hemet. SAN BERNARDINOCo.: Little Pine Creek. MONOCo.: Bridgeport Lake Dam, Hot Creek, Cienega Seca. Mono Lake, Upper Fish Slough. NEVADACo.: Boca Spring, Hobart Mills. SHASTACa: Pondosa (Little Bear Flat). SIERRA Remarks: We examined material from 7 of the Co.: Sattley, Sierraville. SIsKlYou Ca: Mt. Shasta (town). 16 states and provinces where the species has been TULARECO.: 9OOO feet (2743 m) elevation (no locality named). reported. In California we collected 29 f, 49 m by Pherbellia s. maculata is Nearctic; P. s. schoenherri sweeping vegetation along fresh flowing springs and is Palaearctic. streams. Remarks: This species has been reported from 30 states and provinces. We examined material Pherbellia trabeculata (Loew) from California, Nevada, Oregon, and Utah. In Cal- Map 11; Plate 12, Figs. 1, 2 ifornia we collected 81 f, 120 m, mainly in the Sciomyza trabeculata Loew, 1872: 100; Texas; holotype m, MCZ eastern portion of the state. no. 13210. Sciomyza srrigata Malloch, 1914: 324 (not Wulp, 1897). Pherbellia subtilis Orth and Steyskal Melina trabeculara; Melander, 1920: 3 15. Map 10; Plate 11, Figs. 4-6 Pherbellia trabeculara; Steyskal, 1965b: 687.

Pkkllia subtilis Orth and Steyskal, 1980:284, in Orth et a/., Geographic mrrge: NORTHAMERICA: south of approximately 1980; Willits, Mendocino Co., California; holotype m, allo- 34" latitude, California to Texas, south through Mexico (includ- type, USNM no. 75550. ing Bqia California) into Guatemala. Pher6eIlia obscura; Bratt, et al., 1%973 (in part); (not Ring- dahl, 1948). California records (Map 11): ORANGECO.: San Juan Capis- ; Steyskal, 1965b: 687, (not FallCn, 1820); trano (San Juan Creek). RIVERSIDECO.: Lake Hernet, Vail Lake. Palaearctic. SANBERNARDINO CO.: Big Bear Lake (Boulder Bay). SANDIEOO Geogruphic range: widespread in western North America Co.: Palomar Mountain (Doane Pond), Laguna (Mt. presumed), north of 33" latitude, not Alaska. Alpine (Sacatara). 26 BULLETIN OF THE CALIFORNIA INSECT SURVEY

__ _-_ ---_ ~ IL.... - u ... - .... Map 11. California distribution of Pherkllia trabeculata Map 12. California distribution of Pkrbellia vitalis (Loew) (Cresson) Genus Pteromicra Lioy, 1864: 1012 Remarks: We examined material from 7 of the 12 states in the U.S. and Mexico where the species Pteromicra Lioy, 1864 ...... has been reported. In California we collected 110 f, 1. Frontovertex shining yellowish to brown; face 132 m. sub-shining yellowish brown; thoracic pleura yellowish to dark brown; fore femur of female almost entirely black; fore femur of male with Pherbellia vitalis (Cresson) distal half blackened. Male terminalia as in Map 12; Plate 12, Figs. 3, 4 Plate 12, Fig. 5-7 ...... P. pectorosa (Hendel) Melina viralis Cresson, 1920: 43; Berkeley Hills, Alameda Co., Frontovertex shining black, bordered anteriorly California; holotype m, ANSP no. 6221. by a yellowish margin; face shining black, Melina palustris Melander, 1920: 315. tinged with brown; thoracic pleura blackish; Pherbellia vitalis; Steyskal, 1956: 73. distal third of fore femur black. Male ter- minalia as in Plate 13, Figs. 1-3 ...... Geographic range: NORTHAMERICA: Alaska, east to Quebec, ...... P. siskiyouensis Fisher & Orth south to New Jersey, west through New Mexico to southern California. Pteromicra pecturosu (Hendel) California records (Map 12): widespread, in all but the Map 13; Plate 12, Figs. 5-7 deserts, recorded in 28 counties. Dichrochira pectorosa Hendel, 1902: 61; Berlin, Germany; cotypes m, f, NHMV. Remarks: We examined material from 8 of the PIerOmicm peciomsa; Kertesz, 1915: 116; Rozkoh9 and Knut- 27 states where the species has been reported. In son, 1970 1439; lectotype m. California we collected 389 f, 779 m. Geographic range: EUROPENORTH AMERICA: north of 41" lati- tude; Pacific coast to New York.

California record (Map 13); DEL NORTECO.; 1 to 3 mi (1.6- 4.8 km) north of Crescent City, 50 feet (16 m) elevation, VI- 11-1966 (F&O). LZ VINXOdI?V3 do =I?d HSXVR INIIIIa30NV.LS.L 3ElIX.l MARSH FLIES OF CALIFORNIA 29

terminalia as in Plate 15, Figs. 4-9 ...... A. testa- Melander 3. Third antennal segment usually tinged with black on apical half, occasionally only lightly tinged with brown; aristal hairs, long, sparse, longest hairs more than 0.12 mm; sides of whitish-pruinose to testaceous. Male ter- minalia as in Plate 14, Figs. 7-9; Plate 15, Figs. 1-3 ...... A. robiginosa Melander Third antennal segment at most lightly tinged with brown; aristal hairs, short, longest hairs less than 0.07 mm; sides of thorax of female mostly pruinose-cinereous blue, usually less pronounced io male. Male terminalia as in Plate 16, Figs. 1-6 ...... A. vernalis Fisher & Orth

Antichaeta borealis Foote Map 18; Plate 14, Figs. 1-6 Anikhuetu boreulis Foote, 1%1b:161; 8 mi (12.9 km) north of Sandpoint, Bonner Co., Idaho; holotype m, CU.

Geogruphic mnge: USA: California, Idaho, Montana, New York, Ohio. I L -_-. ._ L. cis u I,, t* I. Cul@rnm record (Map 18): Mowc Ca: 3 mi (4.8 km) north of Eagleville, 4640 feet (1414 m) elevation; VI1-10-1%8 (F&O). Map 17. California distribution of Antichaeta festaceu Melander Remarks: We examined material from Califor- nia, Montana, and Ohio. The only known specimen from California is a female from an unshaded boggy gently sloping pasture.

Antichaeta robiginosa Melander Map 16; Plate 14, Figs. 7-9; Plate 15, Figs. 1-3 Anrkhueia roblginosu Melander, 1920317; Three Forks, Gallatin Co., Montana; holotype f, USNM. Fisher and Orth, Antichaeta testacea Melander 1971a: 36 (redescribed). Map 17; Plate 15, Figs. 4-9 Geogruphic runge: USA: California, Montana, Oregon, Wash- Anrkheru iesmcea Melander, 1920: 31% Gallatin Co., Montana; ington. holotype m, USNM.

California recor& (Map 16): INYO Ca: Buckhorn Springs, Geographt mnge: Mutlco: Baja California Norte. USA: Cali- SO00 feet (1524 m) elevation, IV-26-1966 (FdtO). MARMCO.: fornia, Idaho, Montana, New Mexico, Oregon, South Dakota, Novato, 40 feet (12 m) elevation, I-17-1%3 (P. H. Arnaud). Utah. MEND~CINOCo.: Hopland, 600 feet (183 m) elevation, V-25- 1967, V-12-1966 (FBrO); Willits, 1400 feet (427 m) elevation, California records (Map 17): widespread, 20 counties. VI-12-1966 (F&O). MODW Co.: Alturas, Eagleville, Likely, Wil- low Ranch, 4300-4700 feet (1311-1433 m) elevation, VI-6 to Remarks: We examined material from Califor- VII-IO, 1966-1974 (F&O). SHASTACo.: Redding, 480 feet (168 nia, Oregon, and New Mexico. California col- m) elevation, V-24-1967 (F&O). TULARECO.: Woodlake, 550 In we feet (168 m) elevation, V-24-1947 (N. W. Frazier). lected 123 f, 188 m. Biology reported by Fisher and Orth (1964); eggs are deposited on egg masses of Remarks: We examined material from California mainly succineid snails; first instar larvae feed on and Oregon. In California we collected a total of 5 snail embryos; summer pupal diapause at Riverside. f, 10 m in partially shaded to unshaded grassy Collected in open to partially shaded stream and meadows with flowing water present. We suspect meadow habitats. Adult habitus as shown in Plate succineid snails (eggs) are the primary host. 1, Fig. 5. Head profile as in Plate 5, Fig. 5. BULLETIN OF THE CALIFORNIA INSECT SURVEY

process of epandrium with short broad lobe, surstylus in ventral view with lateral bulge and blunt tip. Wing length: IS males, 4.15-4.7 mm, average 4.41; 11 females, 5.05-5.55 mm, average 5.23. Wing length data on D. nmrthewsi, from Steyskal (1960:36). Not yet discovered in California. Known from Parker, Arizona (Valley and Berg 1977:41). Plate 18, Figs. 5-8 ...... D. mnhewsiSteyskal Second antennal segment wholly pruinose, not longer than high; parafrontal spots small or lacking; typical group (Steyskal, 1954b) ...... 2 2. Ventral process of hypandrium with a pretermi- nal point ...... 3 Ventral process of hypandrium without a preter- minal point ...... 4 3. Ventral process of hypandrium short and strongly bent; preterminal point small. Wing length: 10 males, 4.4-5.0 mm, average 4.7; 10 females, 5.2-5.9 mm, average 5.6. Male and female ter- minalia as in Plate 18, Figs. 1-4 ....LJ.inciscurran Ventral process of hypandrium not strongly bent; preterminal point large. Wing length: 10 males, 4.6-5.2 mm, average 5.0; 10 females, 5.3-5.9 mm, average 5.7. Not yet discovered Map 18. California distribution of Anrichoeca wrmlis in California. Known from Arizona, Nevada, Fisher and Orth 0; and Antichuefa borealis and Oregon. Male and female terminalia as in Foote A Plate 17, Figs. 1-4 ...... D. expans Steyskal 4. Surstylus with dorsal tip blunted; ventral process Antichaeta vernulir Fisher and Orth of hypandrium robust and long. Wing length: Map 18; Plate 16, Figs. 1-6 10 males, 4.4-5.1 mm, average 4.9; 10 females, 5.2-6.0 mm, average 5.6. Found in Anricheta wrnaiis Fisher and Orth, 1971a:38; 2 mi (3.2 km) southern California. Male and female ter- north of Willits, Mendocino Co., California; holotype m, allotype, CAS no. 10208. minalia as in Plate 20, Figs. 5-8 ...... D. texensis Curran Geographic runge: USA: California, Idaho, Oregon, Washington. Surstylus with dorsal tip angulate; ventral process of hypandrium not robust and of moderate Ca/&rniO recotds (Map 18): MENDOCINOCa: 2 mi'(3.2 km) length. Found statewide or in northern north of Willits, 1330 feet (404 m) elevation, IV-23,241968 California 5 (FStO). PLUMASCa: Creek at Hwy 36,4900 feet (1494 ...... Rock m) 5. Terminal portion of ventral process of hypan- elevation, VI-8-1966 (FBO). drium flattened. Wing length: 10 males, 5.1- Remarks: We examined material from California 5.6 mm, average 5.3; 10 females, 6.0-6.7 mm, and Oregon. In California we collected 3 f, 2 m. average 6.2. Found in northern California. The collection site 2 mi north of Willits was a nar- Male and female terminalia as in Plate 17, row weed-choked ditch with flowing water, and a Figs. 5-8 ...... D. fontinalis Fisher 8~Orth Terminal portion of ventral process of hypan- nearby large pasture. drium not flattened or only very slightly so. Occurs statewide...... D. Montana Steyskal Genus Dicwu Meigen, 1803: 277 Dk@aMeigen, 1803 ...... Forms of D. montunu Steyskal indicated as follows. 1. Second antennal segment shining on outer upper A. Male and female terminalia as in Plate 19, Figs. half or more, longer than high; large, deep 1-4. Wing length: 10 males, 4.6-5.3 mm, black parafrontal spots present ...... average 5.0; 10 females, 5.6-6.3 mm, average ...... abnormis group (Steyskal, 1954b) 5.9 ...... D. mntana "A" Ventral process of hypandrium short and B. Male and female terminalia as in Plate 19, Figs. straight, with obliquely truncate tip; ventral 5-8. Wing length: 10 males, 3.9-4.4 mm, , MARSH FLIES OF CALIFORNIA

average 4.2; 10 females, 4.4-5.2 mm, average 4.8 ...... D. montana “Alturas” C. Male and female terminalia as in Plate 20, Figs. 1-4. Wing length: 10 males, 4.5-5.2 mm, average 4.9; 10 females, 4.9-6.1 mm, average 5.6 ...... D. montana ‘‘so. Cal.” D. Male and female Terminalia composite of “so. Cal.” and “A.” No illustrations. Wing length: 10 males, 4.5-5.2 mm, average 4.9; 10 females, 5.5-6.0 mm, average 5.8 ...... Dicw montana “intermed.”

Dicoa JbntinalisFisher and Orth Map 19; Plate 17, Figs. 5-8 Dicw fontirrcrlis Fisher and Orth, 1%9b:222; Boca Spring, Nevada Co., California; holotype m. allotype, CAS NO. 10207.

Geographic mrtge: USA: California, Nevada, Oregon, Wash- ington.

calgornm recordT (Map 19): MARINCO.: Inverness, Iv-

*“1“.,+-! L._ - -. 22,24,26-1951 (P. D. Ashlock, CDA). MENDOCMOCO.: 14 mi I L _- ---i- - - _L_ ,” *I . n- * .. (22.5 km) west of Willits, VI-30-1951 (W. C. Bentinck). NEVADACo.: Boca Spring, 5900 feet (1798 m) elevation, IX-22- Map 19. California distribution of Dictya fontinalis 1966 (FBtO); Nevada City, 2500 feet (762 m) elevation, VI-13- Fisher and Orth 0; and Dictya incisu Curran 1965 (T. W. Fisher). SHASTA CQ: Cassel, VII-5-1955 (J. W. A MacSwain). TRINITYCQ: Waldorff Ranch, near Big Bar, VII-3- 1964 (J. C. Borden). Remarks: We have examined material from Sonora and New Mexico. The California record was Remarks: We examined material from all four provided by Dr. Karl Valley (PADA) who exam- states. In California we collected 27 f, 59 m. The ined Melander’s single male specimen in the Boca Spring and Nevada City sites were flowing USNM collection. We have twice collected spring areas in meadows. Biology by Fisher and sciomyzid flies at Afton Junction where the Mojave Orth (1969b). The natural host of first instar larvae River surfaces about 30 mi (48 km) west of Baker, is presumed to be Lithog&phus turbini@rmis (Tryon, but the only Dicwu taken was montana, form “so. 1865) (Hydrobiidae), an operculate prosobranch Cal.” mollusk which appears to be restricted to flowing Diem montana Steyskal fresh shallow (

Dk~uincisu Curran Geographic range: CANADA:Alberta, British Columbia, Map 19; Plate 18, Figs. 1-4 Saskatchewan. MEXICO:Baja California. USA: Arizona, Califor- nia, Colorado, Idaho, Nevada, Oregon, Utah, Washington. DI@W incisa Curran, 1932:6; Great Zuni River, Arizona; holo- type rn, AM”. Steyskal, 1954b:530 (redescribed). California records (Maps 20-23): The species is recorded from 55 of the 58 counties. Geographic mnge: MEXICO:Nuevo Leon, Sonora (Hermosillo, Irnuris). USA: Arizona, California, Kansas, Nevada, New Mex- Remarks: We examined material from all of the ico, Utah, Washington. above listed states and provinces, and over 4000 California record (Map 19): SANBERNARDINO Co.: Baker, specimens from California. Adult habitus as shown IV-24-1935 (A. L. Melander). in Plate 1, Fig. 6. Head profile as in Plate 5, Fig. 6. 32 BULLETIN OF THE CALIFORNIA INSECT SURVEY

- i- -__ - -.-. IL- ., w Y/ .? I? - .-. -? ~ =?~*.4~ I Y I? t. Map 20. California distribution of Dictya montana Map 21. California distribution of DicQd montuna Steyskal, clinal form ?A? Steyskal, clinal form ?Alturas?

Discussion: We consider D. montana a polytypic FORM?A? species which can be segregated into four forms by Map 20; Plate 19, Figs. 1-4 differences in size, male and female terminalia, and Gzlgornia records: widespread north of 36? latitude in 28 north- often by patterns of distribution. The forms are ern and central counties. referred to as ?A?, ?Alturas?, ?Intermediate (Intermed.)?, and ?southern California (so. Cal.)?. Remarks: ?A? does not occur in southern Cali- In the laboratory reciprocal cross breeding occurs, fornia. It is the only form of D. montam we have being of limited to good success depending on the seen in the area of Lake Tahoe for 100 mi (161 combinations attempted, as reported by Fisher and km) to the north, 50 mi (80 km) to the west, 80 Orth (1969b). Knowing this, we hesitate to create mi (129 km) to the south, and to the east and discreet species or subspecies designations for each north of Lake Tahoe in southern Washoe Co., form. All four forms are known to occur in Oregon. Nevada. 324 f, 643 m, or 30% of the D. montana In addition, we have seen form ?so. Cal.? from we collected, were assigned to form ?A.? Nevada, Washington, and Baja California Norte, and form ?Alturas? from Colorado, Idaho, FORM?Alturas? Nevada, and Utah. Map 21; Plate 19, Figs. 5-8 Morphological criteria: Main check points on the Calgornia records: north of 40? latitude in 5 counties in the male postabdomen are the protuberance on the northeastern part of the state. ventral margin of the epandrium and the ventral process of the hypandrium. Females usually can be Remarks: This form co-exists with ?A,? but not distinguished by the nature of the concave posterior with forms ?Intermed.? and ?so. Cal.?. ?Alturas? margin of abdominal sternite VIII, and by male is the smallest of the four forms and superficially association. may therefore be confused with D. umbroides Cur- ran, a typically boreal species. 38 f, 90 m, or 4% of D. montana we collected, were assigned to form ?Alturas.? 33

- I u-;, & ---~ i.. ,* /,. .L *,,U.“.llr ,A, Map 22. California dis’tribution of Dicw mvnmna Map 23. California distribution of Dk@a mvntana Steyskal, clinal form “Intermediate” Steyskal, clinal form “southern California” FORM“so. Cal.” Map 23; Plate 20, Figs. 1-4 Valley and Berg (1977) show D. umbroides at Califwnb rerords (Map 23): widespread in 23 counties, mainly Goose Lake, Oregon. Following exchange of deserts and coastal areas. material, drawings, and correspondence, Dr. Valley concurs in assignment of that material to D. mon- Remarks: Form “so. Cal.” is the only form of tuna form “Alturas.” D. montana south of the Tehachapi Mountains. “A” and “so. Cal.” coexist at Orick (Humboldt FORM“Intermediate” Co.), Olema-Bolinas area (Marin Co.), Bishop Map 22; not figured (Inyo Co.), and Furnace Creek in Death Valley (Inyo Co.). It also has been collected Santa Calfirnia recod: widespread in 22 counties mainly in interior of on the Great Central Valley espacially approaching foothills of Catalina Island, 450 feet (137 m) elevation, VIII- Sierra Nevada. 25-1964 (D. W. Ricker, UCR). 511 f, 1201 m, or 55% of the D. montana we Remarks: “Intermediate” is a form in which the collected, were assigned to form “so. Cal.”. male genitalia are subtly different from either “A” or “so. Cal.”. It is the most common form in the D&ya texemis Curran Great Central Valley where in the north Map 24; Plate 20, Ergs. 5-8 (Sacramento Valley) it tends towards “A” and in Dicw ramsisCurran, 1932:6; Texas; holotype m, AMNH. the south (San Joaquin Valley) towards “so. Cal.”. “Intermediate” also occurs in the Owens Valley Gmg9hic range: MEXICD:Tamaulipas. USA: widespread, (Inyo Co.) and in coastal valleys from San Luis coast to coast. Obispo northward to Mendocino Co. Not montane. This form is nearly allopatric to form “A,” occupy- Culfornia records (Map 24): IMPERIAL Co.: Calexico, Heise Springs, Colorado River (Imperial Dam, Laguna Dam, Palo ing an essentially Upper Sonoran ecological area. Verde, Ripley, Sportsman’s Paradise). LOS ANG~ESCO.: Big 110 f, 256 m, or 11% of D. montanu we col- Tujunga Canyon, Los Angeles International Airport, Mon- lected, were assigned to form “Intermediate.” tebello, San Gabriel River (Whittier Narrows). ORANGECa: 34 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Map 24. California distribution of Diqa tacensis Cur- Map 25. California distribution of E&a mnnexa ran *; and Renocera brevis (Cresson) A (Steyskal) *; and Dic&wcium firmum Steyskal A Sand Canyon Reservoir, San Juan Capistrano (San Juan Creek), U.C at Irvine. RIVERSIDECO.: Corona, Indio. Lake Hernet, Mecca, Pedley, Oasis, Riverside (Santa Ana River), Temecula, Bartle, Siskiyou Co. Male terminalia as in Plate 21, Vail Lake. SAN BERNARDINOCa: Little Cienega Seca, Jenks Figs. 1-3. Lake, Lake Gregory, Morongo Valley. SANTA BARBARACa: Goleta (Atascadero Creek). Dictyacium JirmumSteyskal Remarks: We examined material from 9 of the Map 25, Plate 21, Figs. 1-3 29 states where it has been reported. In California Dicryacium firmum Steyskal, 1956: 79; Waskesiu Lake, this species occurs south of the Transverse Ranges Saskatchewan, Canada; holotype rn, allotype, CNC. usually below 1500 feet (457 m) elevation where it Geographic range: CANADA: Manitoba, Ontario, often occurs with D. montana form “so. Cal.”, but Saskatchewan. USA: California, Colorado, Idaho, Michigan, usually in numbers approximating 2Ooh or less of Nebraska, West Virginia. the D. montana population at a given site. We collected 78 f, 197 m of D. texensis in Cali- Calfirnia record (Map 25): SISKIYOUCO.: Bartle (McIntosh fornia. This species has not been reported from San Ranch), 4000 feet (1219 m) elevation, VI-27-1973, VII-31-1973, Diego County. VIII-1-1973, VI-24-1976 (F&O). Remarks: Discovery in California of this species Genus Di;ctyocium Steyskal, 1956: 78 was unexpected and constitutes a significant range Dictyacium Steyskal, 1956 ...... extension. We collected 22 specimens in an open to Only one species, D. $mum Steyskal, occurs in shaded sedge meadow near a beaver pond on the California. It may be characterized as follows: Face McIntosh Ranch, about 1 mi (1.6 km) east of Bar- with a black central protuberance; parafrontal and tle. The largest number was taken in 1973. Unsuc- orbito-antennal spots large and black; arista1 hairs cessful attempts were made on VI-24,25-1974 and white, moderately short plumose; mesonotum and VI-25-1975. The next specimen, a single male, was abdomen with numerous brown spots; wings taken in 1976. Adult habitus as shown in Plate 1, densely reticulate. In California known only from Fig. 7. Head prome as in Plate 6, Fig. 1. MARSH FLIES OF CALIFORNIA 35

Discussion: The Bartle area is notel for its several unique species of insects and accordingly is frequented by collectors. We have not seen D. firmum from California in collections other than ours. Unless it was an atypically early warming summer, we feel this species has a short flight period which probably peaks from mid-July to mid- August at Bartle.

Genus Elgiva Meigen, 1838: 365 Elgiva Meigen, 1838 ...... 1. Wing without brownish spot around apex of vein R, fore femur with small to moderately developed bristles on the ventral surface. Male terrninalia as in Plate 21, Figs. 4-5 ...... E. conmSteyskal Wing with brownish spot around apex of vein R,; fore femur with well-developed bristles on the ventral surface. Male terminalia as in Plate 21, Figs. 6-9 ...... E. solicita (Harris)

Eigiva connexa (Steyskal) Map 25; Plate 21, Ags. 4, 5 Hedroneurn conma Steyskal, 1954c: 60, Matanuska Valley, Map 26. California distribution of E&iw solicita Alaska; holotype m, allotype, USNM no. 51609. (Hams) Elgiiw mnmSteyskal, 1%5b:690. Hedronewa Ma; Steyskal, 1954: 62; 1%5b: 690. Geographic range: western NORTHAMERICA north of 37" lati- Elpiva solicita; Rozkdshy, 1981: 180. tude. Alaska to Ontario, south to Colorado, west to eastern Cali- fornia. Geographic range: EUROPE. NORTHAMERICA: north of 37" lati- tude. California records (Map 25): ALPINECa: Heenan Lake. Mom Ca: Alturas. Eagleville, Likely, Willow Ranch. MONO California records (Map 26): LASSENCO.: Bieber, Madeline, Co.: Bridgeport, Coleville. Topaz. Smith Flat Reservoir, Susanville. Mom CO.: Alturas, Cedar- ville. Fort Bidwell, Lake City, Hackamore Reservoir, Likely, Remarks: We examined material from 11 of the Willow Ranch. PLUMASCO.: Chester, Quincy. SHASTA CO.: 18 states and provinces where it has been reported. Cassell, Cayton, Hat Creek (Hwy 299). SIERRACO.: Sierraville. SISK~YOOCa: Bray, Grass Lake, Grenada, Meiss Lake. SONOMA In California we collected 23 f, 37 m. Open to par- Ca: Petaluma. tially shaded boggy pastures in the Topaz area yielded most of the specimens. Adult habitus as Remarks: We examined material from 13 states shown in Plate 1, Fig. 8. Head profile as in Plate 6, and provinces. In California we collected 110 f, 95 Fig. 2. m. Approximately 75% of our material came from the Alturas area. Occurrence in the Coast Range Eigiva solicita (Harris) area is represented by a single female labeled Map 26; Plate 21, Figs. 6-9 Petaluma, VI-14-1966(D. Loukonen, CDFA). Musca solirinis Harris, 1780: 1 16. Europe. BMNH ?. Musca rufa Panzer, 1798: 17'. (Primary homonym). Genus Ifoplodiqya Cresson, 1920: 67 Te/anwera /im/a Day, 1881: 88 Steyskal, 1965b:690 (Primary Cresson, 1920 homonym). HoplodicQa ...... &iw su&l/iKloet and hincks, 1945:391. Only one species, Hoplodic@a acuticornir (Wulp) occurs in California. It may be characterized as fol- 'Current consensus of American and European specialists is that lows. Vallar bristles absent; well-developed ocellar Panzer's type specimen has been lost. and post-ocellar bristles; face immaculate; second antennal segment at least one-half length of third 36 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Genus Limnia Robineau-Desvoidy, 1830: 684 Limnia Robineau-Desvoidy, 1830 ...... 1. Brown cell R, of wing with clear spots extending to costal margin; prosternum with hairs ...... 2 Brown cell R, of wing without clear spots extending to costal margin; prosternum without hairs ...... 3 2. Male terminalia as in Plate 22, Figs. 3-6 ...... L. hcii (Robineau-Desvoidy) Male terminalia as in Plate 23, Figs. 3-6. Not yet discovered in California. Known from southern Oregon, Idaho, New Mexico and Utah ...... L. sandowlensisFisher & Otth 3. Second antennal segment with 4 or more strong bristles on the anterior dorsal edge; frons approximately as wide as long; prescutellar bristles approximately 1/2 to 2/3 as long as scutellum. Male terminalia as in Plate 23, Figs. 1, 2 ...... L. pukscens (Day) Second antennal segment with 2 strong bristles on the anterior dorsal edge ...... 4 4. Frons approximately as wide as long; prescutel- lar bristles long as scutellum. Male ter- L!-- -- -.A.--_---_---. ___-_L.iE-dI as I "I x -11 "2 b* .* ,,' .- .. minalia as in Plate 23, Figs. 7, 8 ...... Map 27. California distribution of Hop~icwam- ...... severa Cresson ticornis (Wulp) Frons distinctly longer than wide; prescutellar bristles variable, generally vestigial to 1/2 segment; arista with black hairs; wing heavily length of scutellum. Widespread north of 36" marked with blackish spots and reticulations; one latitude. Male terminalia as in Plate 22, Figs. strong pre-apical bristle on hind tibia; body brown- 7-9 ...... L. inopa (Adams) ish with black to brownish spots and elongate mark- ings; widespread throughout California. Limnia boscii (Robineau-Desvoidy) Hoplodictya acuticornis (Wulp) Map 29; Plate 22, Figs. 3-6 Map 27, Plate 22, Figs. 1, 2 boscii Robineau-Desvoidy, 1830: 690; Carolines; mutilated holotype in Tetamra acutkwnis Wulp, 1897:358; "N. Sonora," cotype MNHNP. series, BMNH. Ternnorem combinata Loew, 1859 295. Limnia inopo; of authors, e.g., Steyskal, 1965b (not Adams, Hopiodictya spinicornis; Cresson, 1920; Steyskal, 1%5b (in part) (not Loew, 1866). 1904). Limnia Cresson, 1920: 75. Hoplodictya acuticornis; Fisher and Orth, 1972b: 178; lectotype combinofo; Limn& boscii; Steyskal, 1965b3691; Fisher and Orth, 1975a: 125; m, BMNH (redescribed; the authors believe that the type Steyskal et ai., 1978: 9 (redescribed). was collected in Cochise Co. or Graham Co., Arizona). Geographic range: CANADA:British Columbia, Ontario. USA: Geographic range: NORTHAMERICA: Kansas northward and Nebraska, Illinois, and Texas west to Pacific coast. eastward to Atlantic coast; also Utah and California.

Calfornia records (Map 27): widespread, 28 counties. Cali/ornia record (Map 29): MONOCO.: Upper Fish Slough [lo mi (16 km) north of Bishop], 4250 feet (1295 m) elevation, 1 m Remarks: We examined material from 13 of the IX-7-1967 (F&O), 1 f VII-11-1968. 17 states and provinces where it has been reported. In California we collected 268 f, 710 m. Remarks: We examined material from 10 states. H. urnticorn& is associated with hygrophilous Discussion: Most of the localities recorded for L. terrestrial snails in the family Succineidae (amber boscii lie east of 100" longitude. The habitat at snails). Adult habitus as shown in Plate 1, Fig. 9. Upper Fish Slough is a permanent open- to lightly- Head profile as in Plate 6, Fig. 3. shaded freshwater marsh which is fed by large MARSH FLIES OF CALIFORNIA 37

Map 29. California distribution of Limnia pubescens (Day) 0; and Limnia boscii (Robineau- Desvoidy) A flowing springs of cool water. Nearby low dry hills summer in northern counties. Usually collected in define the marsh on the east and west. A portion of shaded grassy areas close to springs or streams. the marsh south of the collection area has been set aside as a refuge of the Desert Pup Fish. Until Limnia pubescens (Day) recently were seasonally allowed to graze this Map 29; Plate 23, Figs. 1, 2 area. Creation of the refuge and elimination of sheep should enhance the stability of this habitat. Temtnxem pubescens Day, 1881:86; Washington Territory; m and f cotypes; CNC. Limnm pubescens; Cresson, 1920:77; Fisher & Orth, 1975a: 125; Limnia inopa (Adams) Steyskal et al., 1978: 24,28. Map 28, Plate 22, Figs. 7-9 Ceogmphic range: CANADA:British Columbia. USA: Califor- Tetanocwo inopa Adams, 1904: 448; Washington Territory; holo- nia, Idaho, Oregon, Washington. type m, UK. Limnia cosmltk kvimsralis var. Melander, 1920: 323. Cal@rniu mrds (Map 29): SHASTACa: Cayton (Valley), Limnia bwvimstalis; Steyskal, 1965b: 691. 3050 feet (930 m) elevation, VI-21-1972 to VI-24-1976, VI-26- Limnia boscii; in part, of authors, i.e., Steyskal, 1965b (not 1974, VI-27-1973, VII-25-1972, VIII-1-1973 (F&O); Hat Creek Robineau-Desvoidy. 1830). at Hwy 299, 2650 (808 m) elevation, VI-21-1972. V1-26-1973, Limnm inopa; Fisher and Orth, 1975a: 123 (redescribed); Steys- VI-26-1974. VU-8-1970, VII-25-1972 (F&O); Hat Creek (Hwy kal et al., 1978 14. 89) 3260 feet (994 m) elevation, VII-25-1972 (F&O). SISKIYOU Ca: Bartle (McIntosh Ranch), 4OOO feet (1219 m) elevation, Geographic range: CANADA:Alberta, British Columbia. USA: VII-31-1973 (FBO). TRINITYCa: Carrville, VI-17/VI1-2-1913 (E. California, Idaho, Montana, Oregon, Washington. C. Van Dyke, CAS). Caifornia records (Map 28): north of 36" latitude in 20 central and northern counties. Remarks: We examined material from Califor- nia, Oregon, and Washington. In California we col- Remarks: We examined material from 5 states lected 33 f, 59 m. and provinces. In California we collected 28 f, 37 In the Cayton and Bartle areas we collected adult m. This species was not collected prior to mid- L. pubescens in four discrete colonies which 38 BULLETIN OF THE CALIFORNIA INSECT SURVEY

by Melander in 1942, and we collected a single male in 1972 in a partly shaded marsh near the Melanders’ former summer cabin. Adult habitus as shown in Plate 1, Fig. 10. Head prosle as in Plate 6, Fig. 4.

Genus Renocera Hendel, 1900: 333 Renocera Hendel, 1900...... Only one species, Remcera brevis (Cresson), occurs in California. It may be characterized as fol- lows. Humeral bristle present; prosternum with setae; one fronto-orbital bristle. In California known only from Bartle, Siskiyou Co. (1 speci- men). Male terminalia as in Plate 23, Fig. 9.

Renera brevis (Cresson) Map 24; Plate 23, Fig. 9 Chaetomacem brevis Cresson, 192058; New York; holotype m, USNM. Renocem eynthifornrisMelander, 1920:319; Foote, 1976: 122. Renocera bergiSteyska1, 1954~64,1965b:692. Remcera brevis; Steyskal, 1%5b:692; Foote, 1976: 122.

Map 30. California distribution of LImniu severcl Cres- Geographic range: USA: Alaska, California, Colorado, Idaho, son Michigan, New Mexico, New York, Oregon, Washington.

Cafgornia mrd (Map 24): SI%IYOUCO.: 1 mi (1.6 km) occupied several square meters of drier appearing northwest of Bartle, 4100 feet (1250 m) elevation, VII-20-1966 sub-habitats within the surrounding larger moist to (P. Rude, UCB). wet typical sciomyzid habitats. Remarks: We examined material from Califor- Limnia severa Cresson nia, New Mexico, and Oregon. The California Map 30; Plate 23, Figs. 7, 8 record is a single male. Head profile as in Plate 6, Limnia unguicornis var. severa Cresson, 1920: 80; Cayton, Shasta Fig. 5. Co., California; holotype m, CAS. Discussion: We have not collected R. brevis on Limnia samtogensisvar. sem; Melander, 1920: 324. several attempts in the Bartle area, including Limnia saramgensis var. aresMelander, 1920: 324. Rude’s “1 mi NW” locality. Larvae of all known Limtzia armipes; Steyskal, \%5b: 691. Limnia sewra; Steyskal, 1%5b:692; Fisher and Orth, species of this genus are obligate feeders on finger- 1971b: 164; 1975a:694 (redescribed); Steyskal et al., nail clams. Therefore when attempting to collect 1978: 30. Renocera, stabile waters became first priority areas of search. Geagmphic range: CANADA:British Columbia. USA: Arizona, California, Nevada, Oregon, Washington. Genus Sepedon Latreille, 1804: 196 Calfarnia records (Map 30): 20 northern and central counties, Sepedon Latreille, 1804...... occurring lo sea level in north coastal counties, otherwise mon- 1. Supraspiracular convexity of metathorax without tane. black hairs; medifacies without fine black hairs. Male terminalia in Plate 25, 3-5. 4 as Figs. Remarks: We examined material from states ...... S. /irxipnnis&wipennis Loew and provinces. In California we collected 253 f, Supraspiracular convexity of metathorax with 390 m. black hairs; medifacies with or without fine The southernmost record is Barton Flats at 6200 black hairs ...... 2 feet (1890 m) elevation in the San Bernardino 2. Medifacies without !be black hairs; legs yellow- Mountains. We saw a single specimen taken there ish; frons with parafrontal black spots. Male MARSH FLIES OF CALIFORNIA

terminalia as in Plate 26, Figs. 1-3 ...... S. spinipes americana Steyskal Medifacies with fine black hairs ...... 3 3. Robust species; wing length from 7.0 to 8.9 mm; second antennal segment distinctly compressed. Male terminalia as in Plate 25, Figs. 6-10 ...... S. paciiica Cresson Small species; wing length from 3.6 to 5.5 mm; second antennal segment slender, less compressed ...... 4 4. Hind femur of both sexes simple; hind tibia more or less evenly arcuate; abdomen fre- quently almost black with bluish reflections. Male terminalia as in Plate 24, Figs. 4, 5 ...... S. borealis Steyskal Hind femur of male emarginate ventrally; hind femur of female simple; hind tibia distinctly more curved in distal third, abdomen brown, with little more than a trace of bluish reflection ...... 5 5. Frons with no more than a trace of parafrontal black spots. Male terminalia as in Plate 24, Figs. 1-3 ...... S. b@& Steyskal Frons with distinct parafrontal black 6 - spots ...... I&----- 3------, --E'"34 J I __ I. 1 .u 6. Frons with large parafrontal and orbito-antennal velvety black spots. Not yet discovered in Cal- Map 31. California distribution of s;epedon b@& Steys- ifornia. Known from Prospect, Jackson Co., kal e; and Sepedonj?tsc@nnis&sc@nnisLoew Oregon, approximately 50 miles north of A California-Oregon border. Male terminalia as in Plate 25, Figs. 1, 2 ...... Sepedon borealis Steyskal ...... S. cuscu&nssis Fisher & Orth Frons with small parafrontal and orbito-antennal Map 32; Plate 24, Figs. 4, 5 blackish spots. Male and female terminalia as SeNdon borealis Steyskal. 1951: 283; Yale, Idaho; holotype in Plate 24, Figs. 6-10 ...... S. cup#eiFisher & Orth m, allotype, USNM no. 60906.

Sepdon bifida Steyskal Geographic range: NORTHAMERICA: transcontinental, north of 37" latitude. Map 31; Plate 24, Figs. 1-3 Sepedon b$daSteyskal. 1951:280; Tejon Canyon, Kern Co., Cal- Col@rnia records (Map 32): eastern montane areas north of ifornia; holotype m, allotype, CAS. 36" latitude.

Geogmphic range: MEXICO:Baja California Norte (San Vin- Remarks: We examined material from 11 of the cente, Santo Domingo). USA: California. 28 states and provinces where the species has been reported. In California we collected 54 f, 92 m Colifornm nmwds (Map 31): widespread in central and south- ern counties. No records north of 40" latitude. mainly in wet alpine meadows. This species is dark brown (nearly black), the Remarks: In California we collected 548 f, 1143 darkest of the Sepedon species in California, and m. The species is especially abundant along banks the smallest. of streams or ponds choked with emergent aquatic vegetation such as water cress, brass buttons, water Sepedon capelleiFisher and Orth monkey flower, spike rush. Below 1500 feet (457 Map 33; Plate 24, Figs. 6-10 m) adults are active in all months. Commonly Sepedon capeiki Fisher and Orth. 196%: 157; Bishop, Inyo Co., taken in the Transverse Ranges, and the Great California; holotype m. allotype, CAS no. 10166. Central Valley. Head profile as in Plate 6, Fig. 6. G'eographic range: USA: Caliiornia, Idaho, Nevada, Oregon, Utah, Washington. 40 BULLETIN OF THE CALIFORNIA INSECT SURVEY

k--7 la*--%- :. Map 32. California distribution of Sepedon borealis Map 33. California distribution of Sepedon capellei Steyskal Fisher and Orth

Califrnia recordr (Map 33): widespread in eastern and north- Remarks: We examined material from 26 of the ern Caiifornia usually in montane areas over 3000 feet (914 rn) 38 states and provinces where the species has been elevation and north of 36" latitude. reported. In California we collected 3 f, 11 m. One Remarks: We examined material from the 6 was from Crescent Mills, the remainder from Sier- states. In California we collected 466 f, 791 m, raville. mostly in open alpine meadows. Not spotted on the Discussion: Terminalia and row of long hairs on map was one female tentatively determined as this hind tibia appear to be uniform in S. J: firscipnnis species from Mint Canyon, San Gabriel Mountains, throughout its range except in southeastern U.S. Los Angeles Co., California, VII-1-1965 (W. E. We have seen specimens from several localities in Simmonds, collector). that area with terminalia unlike those illustrated by Steyskal (1951, Fig. 101, for his S. f- floridensis Sepedon JirscipennisJirscipennis Loew which occurs in Louisiana and Florida. Map 31; Plate 25, Figs. 3-5 Sepedon paclfica Cresson Sepedon fuscipennis Loew, 1859: 299; Washington, D.C.; cotypes Map 34; Plate 25, Figs. 6-10 m, f, MCZ. Sepedon J fuscipennis; Steyskal, 1951 (1950): 290. Sepedon pacifica Cresson, 1914:457; Redwood Canyon, Marin Co., California; holotype rn, ANSP no. 6076; Fisher and Geographic range: NORTHAMERICA: widespread north of 36" Orth 1972a: 8 (redescribed). latitude. Sepedon praemiosa; in part, of authors, e.&, Steyskal 1951:279; 1965b:693 (not Giglio-Tos, 1893). Calfornia records (Map 31): PLUMASCO.: 0.5 mi (0.8 km) south of Crescent Mills, 3500 feet (1067 m) elevation, VI-8- Geographic range: CANADA:British Columbia, Saskatchewan. 1966 (F&o).SIERRA Ca: 2 to 3.5 mi (3.2 to 5.6 km) west of MExIco: Baja California Norte. USA: Washington to North Sierraville, 4940 feet (1506 m) elevation, VIII-23-1967, IX-22- Dakota, south to Iowa, west to southern California. 1966 (FBO). California records (Map 34): widespread, records from all but 5 counties. MARSH FXIES OF CALIFORNIA

- 2 a, , Lp

Map 34. California distribution of Sepedon pacfica Map 35. Caliiornia distribution of TeramDcera fmtwginea Cresson (Falldn) e; and *dim spinipes amricana Steyskai A Remarks: We examined material from 15 states and provinces. In California we collected 1679 f, 1921 m. Adults collected in all months in southern Remarks: We examined material from of the California. No records from the deserts except 6 21 states and provinces where the species has been along the western margins. Adult habitus shown as reported. In California we collected 15 f, 11 m in in Plate 1, Fig. 11. open unshaded boggy alpine sedge meadows. Discussion: The Colorado desert apparently func- tions a natural barrier to separate pc&a as S. Genus Tefanocera Dum&& 1800: 439 from S. praemiosa which occurs in eastern Arizona. The two species overlap in Colorado and Utah. TefunoceruDum&& 1800 ...... Black parafrontal spots occur in a low percent of 1. Posterior surface of middle femur with more specimens seen from Ravendale, Smith Flat Reser- than one bristle toward apex, usually three; voir (Lassen Co.), 2 mi (3.2 km) west of prosternum with setae. Male and female ter- Bridgeport (Mono Co.), Indian Tom Lake (Siskiyou minalia as in Plate 30, Figs. 1-4 ...... and Strawberry (Tuolumne (20.1...... T. robusm Loew Co.), Posterior surface of middle femur without bris- tles or with but a single bristle near apex; &@on spinipes americana Steyskal prosternum bare ...... 2 Map 35; Plate 26, Figs. 1-3 2. Posterior surface of middle femur with a single Sepedon SpinipesScopoli. 1763:342. Vienna. Palearctic. bristle near apex ...... 3 Sepedon spines americam Steyskal, 1951 (1950): 277; Hamburg, Posterior surface of middle femur without bris- Livingston Co., Michigan; holotype m, USNM. tles near the apex ...... 8 3. Parafacial hairs extending well above midpoint Geographik range: NORTHAMERICA: Coast to coast north of between lower margin of eye and base of 39" latitude. antennal socket; aristal hairs dense ...... 4 Calflornm recordF (Map 35): SIERRA Ca: 1.6-3.5 mi (2.6-5.6 Parafacial hairs not reaching midpoint between km) west of Sierraville; 4940 feet (1506 m) elevation, VII-9- lower margin of eye and base of antennal 1968, VII-18-1967,VII-23-1967,1X-22-1966 (F&O). socket; aristal hairs sparse to dense ...... 5 42 BULLETIN OF THE CALIFORNIA INSECT SURVEY

4. Vein M with one or more stump veins. Male Aristal hairs sparse; orbito-antennal spot at and female terminalia as in Plate 31, Figs. most faint and not black ...... 12 5-8 ...... T. vicina Macquart 12. Hairs of anterior front at least as long as the Vein M usually lacking stump veins. Male and diameter of an ocellus; second segment of female terminalia as in Plate 29, Figs. 6-9 ...... arista blackish. Male and female terminalia as ...... T. plumosa Loew in Plate 26, Figs. 6-9 ...... T. ferruginea Fallen 5. Aristal hairs especially dense and black; third Hairs of interior front shorter than the diameter antennal segment usually blackish at inser- of an ocellus; second segment of arista yel- tion of arista; orbito-antennal spot usually lowish to brownish. Not yet discovered in large and black. Male and female terminalia California. Known from Reedsport, Douglas as in Plate 28, Figs. 6-8; Plate 29, Fig. 1 ...... Co., Oregon (approximately 120 miles north ...... T. obtuslfibula Melander of California-Oregon border). Terminalia as Aristal hairs moderately dense to sparse; third in Plate 26, Figs. 4, 5 ...... T. bergisteyskal antennal segment not blackish at insertion of arista; orbito-antennal spot not particularly Tetanoceraferruginea Falldn large ...... 6 Map 35; Plate 26, Figs. 6-9 6. Bristles on posterior surface of fore femur usu- ally at least as long as width of femur. Male Tetanocera ferruginea Fallin, 1820: 9; Scandinavia; cotypes rn, f; and female terminalia as in Plate 28, Figs. NRS; Melander, 1920: 327, PI. XXX; Steyskal, 1954~:67. Teranocem ~mngukzrk 1861: 1862: 122; Steyskal, 2-5 mesopora Steyskal Loew, 344, ...... T. 1%5b: 694. Bristles on posterior surface of fore femur Chaetowmcera femcgina; Cresson, 1920:64. short; about one-half width of femur...... 7 Tetanomra lnuonensis Steyskal, 1938: 6; 1954~:67. 7. Orbito-antennal spot brownish. Male and female terminalia as in Plate 27, Figs. 8-10; Plate Geographic range: EUROPENORTH AMERICA: widespread 28, Fig. 1 ...... T. loewi Steyskal north of 35" latitude. Orbito-antennal spot blackish. Male and female terminalia as in Plate 27, Figs. 4-7 ...... Calflornia records (Map 35): eastern California north of 38" ...... T. latitibuia Frey latitude. M~DOCCO.: Alturas, Eagleville, Tulelake. MONOCO.: 8. Hind femur with posterodorsal bristle opposite Bridgeport, Topaz. S~SKIYOUCo.: Meiss Lake. or nearly opposite to apical anterodorsal bris- tle ( T. plebeia will sometimes fit this descrip- Remarks: We examined material from 6 of the tion). Vein M with stump veins. Not yet 21 states and provinces where the species has been discovered in California. Known from reported. In California we collected 17 f, 18 m in Klamath Fails, Klamath Co., Oregon; open to partially shaded sedge meadows. approximately 20 miles north of California- Oregon border. Terminalia as in Plate 30, Tetanocera latifibulaFrey Figs. 5-7 ...... T. rotundicornis Loew Map 36; Plate 27, Figs. 4-7 Hind femur usually without posterodorsal bristle ...... 9 Tetanocera &@h&Frey, 1924 51; Europe; cotypes m, f series; 9. Midfrontal stripe broadly triangular. Not yet ZMUH; Steyskal, 1%5b:6W. Teranocera ksperaSteyska1, 1959: 71; 1%5b: 694. discovered in California. Known from Camp Sherman, Jefferson Co., Oregon (central Geographic range: EUROPE ASIA (SIBERIA).Western NORTH Oregon). Terminalia as in Plate 27, Figs. 1-3. AMERICA:north of 35" latitude...... T. fiscinervisfitterstedt (syn. unicolor Loew) Midfrontal stripe more or less parallel sided ...... 10 California records (Map 36): mainly near the eastern boun- 10. Midfrontal stripe broad; cell AM with dark dary of the state north of 38' latitude. cloud; medifacies of female shining. Male and female terminalia as in Plate 29, Figs. Remarks: We examined material from 10 of the 2-5 ...... T. pkbeia LWW 14 states and provinces where the species has been Midfrontal stripe narrow to moderately wide; reported. In California we collected 110 f, 226 m, cell AM without dark cloud; medifacies of mainly in open unshaded or sparsely shaded grassy female not shining ...... 11 meadows and marshes. 11. Aristal hairs dense; orbito-antennal spot black. Male and female terminalia as in Plate 31, Figs. 1-4 ...... T. soror Melander MARSH RIES OF CALIFORNIA 43

--J --J

It - - - _-___-_A--.u."..d I .. -x, "3 --m -u I" I. 1 >" Map 36. California distribution of Tetanocem htUibulo Map 37. California distribution of Tetanocera mesopora Frey e; and Tetanocera loewisteyskal A Steyskal

Tetanocera loewi Steyskal Geagmphic range: NORTHAMERICA: north of 39" latitude; Map 36; Plate 27, Figs. 8-10; Plate 28, Fig. 1 California to Northwest Temtories, east to Newfoundland, south to Illinois, west to California. Tefanocera kwiSteyskal, 1959: 68, Deerfield Township, Lapeer Co., Michigan; holotype m, allotype, USNM. Cal@rnm mrak (Map 37): LASSEN CO.: Bieber, Hallelujah Junction (Long Valley cresk), Ravendale. Mom Co.: Altum, Geographic mnge: CANADA:British Columbia, Ontario. USA: Cedarville, Eegieville, Lake City, Likely, Fort Bidwell, Willow California, Indiana, Michigan, Minnesota, New York. Ohio, Ranch. PLUMASCO.: Crescent Mills. SHASTACO.: Cassell, Cay- Oregon, Washington, Wisconsin. ton, Hat Creek (Hwy 299). SIERRACa: Sierraville. SISKIYOUCO.: Bray, Fort Jones, Grass Lake. Co/i/ornm records (Map 36): DEL NOR= Ca: 3 mi (4.8 km) north of Crescent City, VI-11-1965, VI-11-1966, Vill-5-1968 Remarks: We examined material from 13 of the (FdrO). 23 states and provinces where this species is known to occur. In California we collected 81 f, 88 m from Remarks: We examined material from Califor- 1965 to 1970. Approximately 30% were collected in nia, Oregon, and Washington. In California this the Likely-Alturas area of Modoc Co. in ditches species occurs north of 40" latitude. We collected a containing water and abundant emergent vegetation total 11 15 m by sweeping low grasses and of f, along (old) Hwy 395, elevation approximately 4400 other low growing plants in drying depressions feet (1341 m) elevation. beneath tall sparse shade along Lake Earl Drive north of Crescent City. Tetanocera obtusBbuIa Melander Map 38; Plate 28, Figs. 6-8; Plate 29, Fig. 1 Tetanocera mesopora Steyskai Tetamxem obrusifibukr Melander, 1920: 328; MI. Constitution Map 37; Plate 28, Figs. 2-5 (OmIsland), Washington; holotype m, USNM. Ternnacera mesapam Steyskal, 1959 70; Walden, Colorado; holo- Geographic mnge: CANADA:British Columbia, Quebec(?). type m, allotype, USNM. USA: California, Idaho, Oregon, Washington. 44 BULLETIN OF THE CALIFORNIA INSECT SURVEY

I... ..,a k - -i VI I SI .* ,. ---?,-----?!I Map 39, California distribution of Tetanocera plekia Loew

Cal$ornia records (Map 38): north of 37" latitude fairly widespread coastally from San Francisco Bay area northward. Not in Sacramento Valley or extreme northeastern part of state. Rewks: We examined material from 11 of the Remarks: We examined material from Califor- 39 states and provinces where this species has been nia, Oregon, Washington. In California we collected reported. In California we collected 27 f, 31 m from 36 f, 55 m from 1966 to 1975. Usually 1-4 speci- 1965 to 1975. mens were collected per stop. The largest single Discussion: Tetanocera plebeia was collected collections were taken 2 mi (3.2 km) north of Boli- mainly in rather mesic situations a few meters from nas, Marin Co., IX-12-1967, 13 feet (4 m) eleva- free water. We presume its larvae attack slugs in tion, around clumps of Juncus in a drying meadow, California, as in the eastern U.S. and 0.5 mi (0.8 km) south of Crescent Mills, Plu- mas Co., 3500 feet (1067 m> elevation, VI-8-1966, Tetanocera plurnosa Loew in a drying shaded ditch along Hwy 89. Map 40; Plate 29, Figs. 6-9 Tetanocera plumosa Loew, 1847:ml; Alaska; holotype f, MCZ Tetanocera pkbeia Loew no. 13221. Map 39; Plate 29, Figs. 2-5 Tetomm nancme Brimley. 1925: 75; Steyskal 1%5b:694; 1%5c:445. Tetunocem plebem hew, 1862: 120. Middle states (D.C. area); Tetammplum$era Steyskal, 1965c:445; Wulp, 1897: 359. holotype m. MCZ no. 13220. Geographic range: NORTHAMERICA: widespread north of 36" Geographic range: EUROPE.NORTH AMERICA: north of 37" latitude. latitude from California to North Carolina, including Alaska. Ca/&rnia rccordF (Map 40): widespread in 30 counties includ- recordr Ca/&rnia (Map 39): DEL NORTECa: Crescent City, ing the Great Central Valley. Smith River. MARINCa: Olema. MEND~CINOCa: Inglenook Fen, 5 mi (8 km) north Fort Bragg. PLACERCO.: Tahoe (City?). PLUMASCo.: Chester, Quincy. SHASTA CQ; Cayton (Valley), 2 Remarks: We examined material from 17 of the mi (3.2 km) south of Pondosa. SIERRACO.: Sattley. SISKIYOU 37 states and provinces where this species has been Co.: Battle, Bray, Gazelle. reported. In California we collected 382 f, 585 m MARSH FLIES OF CALIFORNIA

.. c 2 I _*. ___ .4 - --

------AL - .I--__u --L?*** I u %1 w . /I , _---,. I*. Map 40. California distribution of Tetanocera ptumosa Map 41. California distribution of Tetanocera robustn Loew Loew mostly at elevations in excess of 4000 feet (1219 Tetanocera soror Melander m). Adult habitus as shown in Plate 1, Fig. 12. Map 42; Plate 31, Figs. 1-4 Head profile as in Plate 6, Fig. 7. South of 36" wing Tetamem soror Melander, 1920: 328; Mica, Washington; holo- may exhibit stump veins. type m, USNM. Tetanocera robusta Loew Geographic range: CANADA:Alberta. USA: California, Idaho, Map 41; Plate 30, Figs. 1-4 Montana, Nevada, Oregon, Washington. TemmKura robusta Loew, 1847: 197; Europe; cotypes rn, f; ZMB. Calenm records (Map 42): ALPINECa: Hope Valley. EL Frey 1924:47; K. C. V. Smith, 1957:70, lectotype rn. DORALWCo.: Fallen Leaf Lake Road. NEVADACo.: Boa. Boca Tetanocera papilli/ero Melander 1920: 329; Steyskal, 1965b: 694. Spring, Hobart Mills, Sagehen Creek. PLUMASCo.: Quincy. SHASTA Ca: 2 mi (3.2 krn) south of Pondosa. SIeRRA CO.: Geographic range: EUROPE NOR^ AMERICA:north of 37" lati- Independence Lake, Sattley, Webber Lake. SEKIYOUCa: Bartle. tude, from northern California to Newfoundland. Remarks: We examined material from California Calfirnm records (Map 41): DEL NORTECa: Crescent City. and Oregon. In California we collected 96 f, 100 m LASSEN Ca: Madeline. MomCa: Alturas. Cedarville, Davis Creek, Eagleville, Fandango Pass, Fort Bdwell, Lake City, over 50% of which came from 2 mi (3.2 km) south Likely, Lily Lake (7 mi (1 1.2 km) east of Pine Creek]. Tulelake. of Pondosa (Little Bear Flat), 4140 feet (1262 m) elevation, and about 20% from the Bartle area Remarks: We examined material from 10 of the (McIntosh Ranch), 4000 feet (1219 m) elevation. 20 states and provinces where the species has been reported. In California we collected 24 f, 28 m from 41" latitude northward. 46 BULLETIN OF THE CALIFORNIA INSECT SURVEY

IL.-;,-_-- -~.. >*. - -%-"---J,' 1 (I In *I Map 42. California distribution of Tetonocem soror Map 43. California distribution of Terunocera vicinu Melander Macquart

Tetanocera vicina Macquart Map 43; Plate 31, Figs. 5-8 Tetanocera vicinu Macquart, 1843 (1842): 337: Philadelphia, Pennsylvania; holotype f, MNHNP.

Geographic range: EUROPE.NORTH AMERICA:widespread north of 36" latitude. California to North Carolina.

California records (Map 43): LASSENCO.: Bieber. MODOCCO.: Alturas, Eagleville, Lily Lake 17 mi (11.2 km) east of Pine Creek]. NEVADACo.: Sagehen Creek. PLACERCo.: Martis Creek. PLUMASCO.: Quincy. SHASTA CO.: Cayton, Hat Creek, Hatchet Mountain Summit, Pondosa (Little Bear Flat), Shingletown. SIERRACO.: Sattley, Sierraville. SBKIYOU Co.: Bray, Fort Jones, Gazelle, Grass Lake. Remarks: We examined material from 10 of the 34 states and provinces where the species has been reported. In California we collected 38 f, 36 m north of 39". This species has been confused with T. plumosa which also may have stump veins in the wing. Literature Cited

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Fontana, P. G. Giglio-Tos, E. 1972. Larvae of Dichemphoru biror’ (Kertisz, 1901) 1893. Diagnosi di nuovi generi e di nuove specie di (Diptera: Sciomyzidae) feeding on Lymnueu Ditteri. IX. Turin Univ. Mus. -1. ed Anat. tomenmsu (Pfeiffer, 1885) snails infected with Comp., Bol. 8(158):1-14. Fasciolu hepatica L. Parasitology 640):89-93. Haliday, A. H. Foote, B. A. 1838. New British insects indicated in Mr. Curtis’ 1959. Biology and life history of the snail-killing Guide (concl.). Ann. Nat. Hist. (1839) flies belonging to the genus Scioqw Fallin 2: 183-190. (Diptera: Sciomyzidae). Ann. Ent. SOC. Hanna, G. D. Amer. 52(1):31-43. 1966. Introduced mollusks of western North Amer- 1961a. Biology and immature stages of the snail- ica. Calif. Acad. Sci., Occasional Papers No. killing flies belonging to the genus Tetono- 48, 108 pp. wru (Diptera: Sciomyzidae). Ph.D. thesis. Harman, W. N. and C. 0. Berg Cornell Univ. (L. C. Card No. Mic. 62-105) 1971. The freshwater snails of central New York, pp. 1-190. with illustrated keys to the genera and 1961b. A new species of Anrichueto Haliday, with species. Search l(4): 1-68. Cornell Univ. notes on other species of the genus (Diptera: Harris, M. Sciomyzidae). Proc. Ent. Soc. Wash. 1780. An exposition of English Insects with curious 63(3): 161-164. observations and remarks, wherein each 1961c. The marsh flies of Idaho and adjoining areas Insect is particularly described; its Parts and (Diptera: Sciomyzidae). Amer. Midl. Nat. Properties considered; the different Sexes 65(1): 144-167. distinguished, and the Natural History faith- 1963. Biology of slug-killing Tetanoceru (Diptera: fully related. Decads 111-V, 73-166 pp., 21- Sciomyzidae). Proc. North Central Branch 50 PIS. London. Ent. SOC.Amer. 18: 97. Hendel, F. 1971. Biology of Hedriu mkta (Diptera: Sciomyzi- 1900. Untersuchungen fiber die europaischen Arten dae). Ann. Ent. Soc. Amer. 64(4):931-941. der Gattung Tetunoceru im Sinn Schiner’s. Z. 1973. Biology of Pherbellia prefixa (Diptera: B. Ges. Bd. L.:319-358. Sciomyzidae), a parasitoid-predator of the 1902. Revision der paliiarktischen Sciomyziden operculate snail Valuuru simeru (: (Dipteren-Subfamilie). K.-K. Zoo1.-Bot. ). Ent. Soc. Wash., Proc. Gesell. Wien., Abhandl. 2(1): 1-94. 75(2): 141-149. Hennig, W. 1976. Biology and larval feeding habits of three 1965. Die Acalyptratae des baltischen Bernsteins species of Renoceru (Diptera: Sciomyzidae) und ihre Bedeutung ffir die Erforschung der that prey on fingernail clams (Mollusca: phylogenetischen Entwicklung dieser Sphaeriidae). Ann. Ent. Soc. Amer. Dipteren-Gruppe. Stuttgart Beitr. Natur- 69(1): 121-133. kunde 145: 1-215. 1977. Biology of Oidemmps jhugieus (Diptera: 1969. Neue iibersicht iiber die aus dem baltischen Sciomyzidae), a parasitoid enemy of the land Bernstein bekannten Acalyptratae. Stuttg. snail Stenommu hirsutum (Mollusat: Polygyri- Beitr. Naturkd. 209: 1-42. dae). Proc. Entomol. Soc. Wash. 79(4):609- 1973. Diptera (Zweifliigler). Handb. Zool. 4(2) 619. 2/31: 1-200, W. de G., Berlin. Foote, B. A., S. E. Neff, and C. 0.Berg Johnson, C. W. 1960. Biology and immature stages of Airichomelina 1929. Diptera destroying snails. Psyche 36(2): 106. pukra (Diptera: Sciomyzidae). Ann. Ent. Kaczynski, V. W., J. Zuska, and C. 0. Berg SOC.Amer. 53(2): 192-199. 1969. Taxonomy, immature stages, and bionomics Frey, R. of the South American genera Perilimnia and 1924. Die nord palaarktischen Tetonocero-Arten Shannon& (Diptera: Sciomyzidae). Ann. Ent. (Diptera: Sciomyzidae). Notulae Entomologi- Soc. Amer. 62(3):572-592. cae IV: 47-53. Keilin, D. Geckler, R. P. 1919. On the life-history and larval anatomy of 1971. Laboratory studies of predation of snails by MelMu cognura Meigen parasitic in the snail larvae of the marsh fly, Sepedon tenuicorais Helicella (Heliomunes) virgam Da Costa, (Diptera: Sciomyzidae). Canad. Ent. with an account of the other Diptera living 103(5): 638-649. upon mollusks. Parasitol. 11 (38~4):430-455. 50 BULLETIN OF THE CALIFORNIA INSECT SURVEY

KertBsz, K. 191 5. A Magyar Birodalom Sciomyzidai. Allattani Knutson, L. V., R. Rozkos’ni, and C. 0. Berg Koezleminyek. Vol. 14: 116. 1975. Biology and immature stages of Pherbina and Kloet, G. S. and W. D. Hincks Psacudina (Diptera: Sciimyzidae). Acta Sci. 1945. A check list of British insects. 483 pp. Nat. Brno 90): 1-38. Arbroath, Scotland. [Sciomyzidae, pp. 390- Knutson, L. V., J. W. Stephenson, and C. 0. Berg 392.1 1965. Biology of a slug-killing fly, Tetanocera ehtn Knutson, L. V. (Diptera: Sciomyzidae). Proc. Malacol. SOC. 1962. Snail-killing sciomyzid flies. The Cornell London 36(4): 213-220. Plantations 17(4): 59-63. 1970. Biosystematic studies of fasciata 1966. Biology and immature stages of malaco- (Meigen), a snail-killing fly (Diptera: phagous flies: Antichaeta analis, A. otriseta, Sciomyzidae). Trans. Roy. Ent. SOC.London A. brevipnnis, and A. oblivioscr (Diptera: 122(3):81-100. Sciomyzidae). Trans. Amer. Entom. SOC. Knutson, L., G. C. Steyskal, J. Zuska, J. Abercrombie 92: 67-101. 1976. Family Sciomyzidae; Fasc. 64:l-24. IN A 197Oa. Biology and immature stages of Tetanwa pal- catalogue of the Diptera of the Americas lidivenrris, a parasitoid of terrestrial snails south of the United States. Museo de Zoolo- (Diptera: Sciomyzidae). Ent. Scand. l(2): 81- gia, Univ. de Sfo Paulo. 89. Knutson, L. and K. Valley 1970b. Biology of snail-killing flies in Sweden (Dip- 1978. Biology of a neotropical snail-killing fly, tera: Sciomyzidae). Ent. Seand. l(4): 307-314. Sepedonea isthmi (Diptera: Sciomyzidae) . 1973. Biology and immature stages of Corermcera Proc. Entomol. Soc. Wash. 80(2): 197-209. murginata F., a predator of terrestrial snails Latreille. P. A. (Diptera: Sciomyzidae). Ent. Scand. 1804.. Tableau mgthodique des Insectes. Pp. 129- 4(2): 123-133. 200. IN Sociit6 de Naturalistes et 1976. Sciomyzid flies: another approach to biologi- d’Agriculteurs, Nouveau dictionnaire cal control of snail-borne diseases. Insect d‘histoire naturelle, appliqut aux arts, princi- World Digest 3(4): 13-18. palement a I’agriculture et a I’Qconomie 1981. New combinations and synonymies in rurale et domestique. Vol. 24, [sect. 31: Palearctic and Nearctic Sciomyzidae (Dip- Tableaux m6thodiques d‘histoire naturelle, tera). P~oc.Ent. Soc. Wash. 83(2): 332-338. 238 pp., 5 pls. Paris. Knutson, L. and J. Abercrombie Lioy, P. 1977. Biology of Antichueta melanosom (Diptera: 1864. I ditteri distribuiti second0 un nuovo metodo Sciomyzidae), with notes on parasitoid di classihzione naturale [cont.]. I. R. 1st. Braconidae and lchneumonidae (Hymenop- Veneto di Sci., Let. ed Arti, Atti ser. 3, tera). Proc. Ent. SOC.Wash. 790): 111-125. 9: 499-518, 569-604, 719-771, 879-910, 989- Knutson, L. V. and C,0. Berg 1027, 1087-1126, 1311-1352 fcont.1. 1963. Biology and immature stages of a snail-killing Lmw, H. fly, Hyahrnya dorsalis (Fabricius) (Diptera: 1847. Wissenschaftliche Mittheilungen Ueber Sciomyzidae). Proc. Roy. Ent. Soc. London Tetanocera ferruginea und die ihr verwandten (A) 38(4-6):45-58. Arten. Stettiner Entomologishe Zeitung 1964. Biology and immature stages of snail-killing 8: 194-202. flies: the genus E@ua (Diptera: Sciomyzi- 1859. Die nordamerikanishen Arten der Gattung &e). Ann. Entomol. SOC.Amer. 57(2):173- Temnocera und &pedon. Wiener Entomol. 192. Monatschrift 3 (10): 289-300. 1971. The malacophagous flies of Norway (Diptera: 1861. Diptera. Americae septentrionalis indigena. Sciomyzidae). Norsk. Ent. Tidsskr. 18: 119- Centuria 1-5. 134. 1862. On the North American Sciomyzidae. Mono- Knutson, L. V., C. 0. Berg, L. J. Edwards, A. D. Bratt, graphs of the Diptera of North America. Part and B. A. Foote I. pp. 103-128. [edited by Osten-Sacked. 1967. Calcareous septa formed in snail shells by Smiths. Misc. Coll. 6(l):no. 141. larvae of snail-killing flies. Science 1866. Diptera Americae septentrionalis indigena. 156(3774): 522-523. Centuria sexta. Berlin. Ent. Ztschr. 10: 1-54. Knutson, L. V., S. E. Neff, and C. 0. Berg 1872. Diptera Americae septentrionalis indigena. 1967. Biology of snail-killing flies from Africa and Vol. 2: [Centuriae 6-10], 300 pp. [Republica- southern Spain (Sciomyzidae: Sepdon). tion of Berlin. Ent. Ztschr. 10: 1-54, 18661. Parasitology 57: 487-505. MARSH FLIES OF CALIFORNIA 51

1876. Beschreibung neuer amerikanischen Dip- Nagatomi, A. and K. Kushigamachi teren. Ztschr. f”ur die Gesam. Naturw. 1965. Life history of Sepedon sauteri (Diptera: 48:317-340. Sciomyzidae). Kontyu 33(1): 35-38. Lundbeck, W. Neff, S. E. 1923. Some remarks on the biology of the 1964. Snail-killing Sciomyzid flies: application in Sciomyzidae, together with the description of biological control of gastropod vectors of a new species of Crenulus from Denmark. trematodes. Congr. Intern. Assoc. of Vidensk. Medel. Dansk Naturhist. For., Theoretical and Applied Limnology. Kobenhaven, 76: 101-109. 15(2):933-939. Lynch, J. J. Neff, S. E. and C. 0. Berg 1965. The ecology of Lymnaea tomentosa (Pfeiffer 1961. Observations on the immature stages of Pro- 1885) in Southern Australia. Australian Jour. rodic~a honduranu (Diptera: Sciomyzidae). -1. 13~461-473. Bull. Brooklyn Ent. Soc. 61(2): 46-56. Macquart, J. 1962. Biology and immature stages of Hoplodicwa 1835. Histoire naturelle des Insectes. Dipthres, spinicornisand H. setosa. Trans. Amer. Ent. Tome deuxieme. Diptera, Vol. 2, 703 pp. IN Soc. 88(2):77-93. Roret, N.E. (ed.) Collection des suites H 1966. Biology and immature stages of malaco- Buffon. Paris. phagous Diptera of the genus Sepedon 1843. Dipthres exotiques nouveaux ou peu connus. (Sciomyzidae). Agric. Exp. Sta. Virginia SOC.Roy. des Sci., Agric., Arts. Lille, MCm. Polytechnic Inst., Bull. 566, pp, 1- 1 13. (1842): 162-460. Nishida, T. and T. Torii Malek, E. A. 1970. A handbook of field methods for research on 1962. Medical malacology. Laboratory guide and rice borers and their natural enemies. Intern. notes. Burgess Publ. Co., Minneapolis. 154 Biolog. Prog., Handbook No. 14. Blackwell PP- Sci. Publ., Oxford. 130 pp. Malloch, J. R. ONeill, W. L. 1914. Synonymical notes on North American 1973. Biology of Trichopia pope; and T. atrichomel- Sciomyzidae. Canad. Entom. 46 (9): 323-324. inue (: Diapriidae), parasitoids Mason, H. L. of the Sciomyzidae (Diptera). Ann. Ent. SOC. 1957. A flora of the marshes of California. Univ. of Amer. 666): 1043-1050. California Press, Berkeley and Los Angeles; Orth, R. E. 878 pp. 1982. Five new species of Pherbellia Robineau- Meigen, J. W. Desvoidy, subgenus Oxytaenia Sack, from 1803. Versuch einer neuen Gattungseintheilung der North America (Diptera: Sciomyzidae). Proc. europaischen zweifliigeligen Insekten. Mag. f. Entomol. SOC.Wash. 84(1):23-37. Insektenkunde 2: 259-281. Orth, R. E., G. C. Steyskal and T. W. Fisher 1830. Systematische Beschreibung der bekannten 1980. A new species of Pherbella Robineau- europaischen zweifliigeligen Insekten. Vol. Desvoidy subgenus Chetocera Robineau- 6: 1-401. Hamm. Desvoidy from North America (Diptera: 1838. Systematische Beschreibung der bekannten Sciomyzidae). Proc. Entomol. SOC. Wash. europaischen zweifliigeligen Insekten. Vol. 82(2): 284-292. 7: 1-434. Hamm. Panzer, G. W. F. Melander, A. L. 1798. Faunae insectorum germanicae initiae oder 1920. A review of the Nearctic Tetanoceridae. Ann. Deutschlands Insekten. Heft 54: 1-24. Ent. SOC.Amer. 13(3):305-332. Niirnberg. Mercier, L. Patterson, C. M. 1921. Dipthres de la cote du Calvados. IIe Liste. 1971. Taxonomic studies of the land snail family Annales de la Soci6t6 Entomologique Bel- Succineidae. Malacol. Review 4: 131-202. gique; Vol. 61, pp. 162-164. Ringdahl, 0. Nomoi, S., C. Watanabe, K. Yano, and K. Yasumatsu 1948. Bemerkungen zu schwedischen Sciomyziden. 1975. Revision of rice stem borers, their parasites, Opuscula Entomol. 13(2): 52-54. and the family Sciomyzidae in South and Robineau-Desvoidy, J. B. East Asia. Chapter 8 IN Jap. IBP Synthesis, 1830. &ai sur les Myodaires. [Paris] Inst. de Vol. 7, “Approaches to biological control,” France, CI. des Sci. Math. et Phys., Acad. K. Yasumatsu and H. Mori, editors, pp. 69- Roy. des Sci., Mim. present& par divers 80. Savans [ser. 21, 2: 1-813. 52 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Robinson, W. H. and B. A. Foote 1954b. The American species of the genus Dictya 1978. Biology and immature stages of Antichaeta Meigen. Ann. Ent. SOC. Arner. 47(3):511- borealis (Diptera: Sciornyzidae), a predator of 539. snail eggs. Proc. Entomol. Soc. Wash. 19%. The Sciomyzidae of Alaska (Diptera). Proc. 80(3) :388-396. Entomol. SOC. Wash. 56(2): 54-71. Roth, B. 1956. New species and taxonomic notes in the fam- 1972. Rare and endangered land mollusks in Cali- ily Sciomyzidae (Diptera: Acalyptratae). fornia. Inland Fisheries Administrative Mich. Acad. Sci., Arts, and Letters, Papers Reports No. 72-10 (May). State of Calif. 41: 73-87. Dept. of Fish and Game. [mimeograph copy]. 1959. The American species of the genus Tetano- Rozkoh;lP, R. cera Dumkril (Diptera: Sciomyzidae). Mich. 1967. Zur Morphologie und Biologie der metamor- Acad. Sci., Arts, and Letters, Papers 4453- phosestadien mitteleuropaischer Sciomyziden 91. (Diptera). Acta Sci. Nat. Brno., 1: 117-160. 1960. New North and Central American species of Rozko&, R. Sciomyzidae (Diptera: Acalyptratae). Proc. 1981. A new name and some new synonyms of Entomol. SOC. Wash. 62(1):33-43. Palaearctic Sciomyzidae (Diptera). Ent. 1961. The North American Sciomyzidae related to Swd. 12: 177-180. Pherbelllia jkiipes (Maquart) (Diptera: Rozkoini, R. and L. V. Knutson Sciomyzidae). Mich. Acad. Sci., Arts, and 1970. Taxonomy, biology, and immature stages of Letters, Papers 46: 405-415. palearctic Ptemmicra, snail-killing Diptera 1963. Taxonomic notes on Sciomyzidae (Diptera: (Sciomyzidae). Ann. Entomol. Soc. Amer. Sciomyzidae). Mich. Acad. Sci., Arts, and 63 (5): 1434-1 459. Letters, Papers 48: 113-125. Schmitz, H. 1965a. The subfamilies of Sciomyzidae of the world 191 7. Biologische Beziehungen zwischen Dipteren (Diptera: Acalyptratae). Ann. Ent. SOC. und Schnecken. Biologisches Zentralblatt Amer. 58(4): 593-594. 37: 24-43. 1965b. Family Sciomyzidae. Pp. 685-695. IN Stone, Scopoli, J. A. A., et al., A catalogue of the Diptera of 1763. Entomologia carniolica exhibens insecta car- America north of Mexico. Agr. Res. Sew., nioliae indigene et distributa in ordines, gen- U. S. Dept. Agric. Handbook 276, 1696 pp. era, species, varietates metholo Linnaeana. 1965c. Notes on types of some species described in 421 pp. Vindobonae [-Vienna]. Sciomyza and Tetanocera by Loew, Walker Seguy, E. and van der Wulp (Diptera: Sciomyzidae, 1950. La biologie des diptkres. Encyclopedie ento- , , ). Studia Ent. mologique XXVI, Series A, 609 pp. Paul 8(1-4): 445-448. Lechevalier, Paris. 1966. The nearctic species of Pherbelliu Robineau- Smith, K. C. V. Desvoidy, subgenus Oaytizenia Sack (Diptera: 1957. Selection of a lectotype of Tetamera robusta Sciomyzidae). Mich. Acad. Sci., Arts, and Loew (Diptera: Sciomyzidae). Entom. Letters, Papers 51:31-38. Monthly Mag. 93:70-71. 1973. Pwromicra inermis Steyskal a synonym of Steyskal, G. C. Sciomyza mria (Coquillett). Proc. Entomol. 1938. New and Tetanoceridae (Dip- Soc. Wash.. 75(1):83. tera) from North America. Ocors. Papers of Steyskal, G. C., T. W. Fisher, L. Knutson, and R. E. the Museum of Zoology, Univ. Mich. Press, Orth NO. 386: 1-10. 1978. Taxonomy of North American flies of the 1949. New Diptera from Michigan (Stratiomyidae, genus Limnia (Diptera: Sciomyzidae). Univ. Sarcophagidae, Sciomyzidae). Mich. Acad. Calif. Publ. in Entomology. Vol. 83:l-48, 5 Sci., Arts, and Letters, Papers (1947). plates. 33: 173-180. Steyskal, G. C. and L. Knutson 1951. The genus Sepedon Latreille in the Americas 1975. The cochleate vesicle, a highly specialized (Diptera: Sciomyzidae). Wassman Jour. Biol. device for sperm transfer in male sciomyzid (1950) 8(3): 271-297. flies. Ann. Ent. SOC. Amer. 68(2):367-370. 1954a. The genus Ptemmicra Lioy (Diptera: Taylor, D. W. Sciomyzidae) with especial reference to the 1966. Summary of North American Blancan non- North American species. Mich. Acad. Si., marine Mollusks. Malacologia 4(1):1-172. Arts, and Letters, Papers 39:257-269. (1953 1970. West American freshwater Mollusca, I: meeting) Bibliography of Pleistocene and recent MARSH FLIES OF CALIFORNIA 53

species. San Diego SOC. Natl. Hist., Mem. 4, Sciomyzidae). Ann. Ent. SOC. Amer. pp. 1-73. 63(3): 877-895. 1975. Index and Bibliography of Late Cenozoic Valley, K. and C. 0.Berg Freshwater Mollusca of Western North 1977. Biology, immature stages, and new species of America. Claude W. Hibbard Memorial, snail-killing Diptera of the genus Dicrya Volume 1; Papers on paleontology: no. 10. (Sciomyzidae). Search Agriculture (Entomol- Contribution number 43, The Pacific Marine ogy; Ithaca, 18) Vol. 7(2):1-45. Cornell Station, Univ. of the Pacific, Dillon Beach, Univ., N.Y. Calif. 94929; 384 pp. Walker, F. 1981. Freshwater mollusks of California: a distribu- 1853. Insecta Saundersiana Vol. 1; Diptera, Part tional checklist. Calif. Fish and Game. IV:401 (1852). 67(3): 140-163. Wulp, van der, F. M. Thomson, C. G. 1897. Biologia Centrali-Americana. Diptera. 11. x + 1868. Kongliga svenska fregatten Eugenies resa 489 pp., 13 plates. [Sciomyzinae pp. 354-3601 omkring jorden, etc. 11. Zool. I, Insecta, Dip- Editors: Godman, F. D. and 0. Salvin; Lon- tera: 443-614. don. April, 1888 to May, 1903. Trelka, D. G. and C. 0.Berg Yano, Koji 1977. Behavioral studies of the slug-killing larvae 1975. Bionomics of Sepedon sphegeus (Fabricius) of two species of Temnoceru (Diptera: (Diptera: Sciomyzidae). Chapter 11, p. 85, Sciomyzidae). Proc. Entomol. SOC. Wash. IN JIBP Synthesis Vol. 7, “Approaches to 79(3) ~475-486. biological control,” Univ. Tokyo Press, Trelka, D. G. and B. A. Foote Yasumatsu, K. and H. Mori, editors. 1970. Biology of slug-killing Tefunoceru (Diptera:

PLATES 56 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 1 : Adult males of California sciomyzine genera.

Figs. 1-4, Sciomyzini. Figs. 5-12, Tetanocerini. Index = 2 mm. Fig. 1. Atrichomlinu puberu (Loew). California; San Bernardino Co.; Lake Gregory; 4520 feet (1378 m) elevation; X-6-1962 (T. W. Fisher). Fig. 2. Pherbelliu purullelu (Walker). California; San Diego Co.; Loveland Reservoir; 1355 feet (413 m) elevation; VI-22-1964 (T. W. Fisher & R. E. Orth). Fig. 3. Pteromicru siskiyouensisFisher and Orth. California; Siskiyou Go.; Grass Lake, Hwy 97; 5122 feet (1561 m) elevation; V1-17-1969 (R. E. Orth). Fig. 4. Sciomyzu simpkxFa1lin. California; Modoc Co.; 2 mi (3.2 km) south of Alturas; 4300 feet (1311 m) elevation; VI-10-1966 (T. W. Fisher and R. E. Orth). Fig. 5. Anrichuefu fesfuceu Melander. California; Mono Co.; 2 mi (3.2 km) north of Topaz; SO00 feet (1524 m) elevation; VII-I 1-1966 (T. W. Fisher & R. E. Orth). Fig. 6. Dicrya montunu Steyskal. California; Humboldt Co.; 2 mi (3.2 km) south of Orick; 10 feet (3 m) eleva- tion; VI-12-1966 (T.W. Fisher & R. E. Orth). Fig. 7. Dic~ciumfirmumSteyskaI.California; Siskiyou CO.; Bartle, Hwy. 89; 4000 feet (1219 m) elevation; VIII-1-1973 (T. W. Fisher & R. E. Orth). Fig. 8. E&vu connexo (Steyskal). California; Mono Co.; 3.1 mi (5 km) north of Topaz; 5200 feet (1585 m) elevation; VI-22-1972 (T. W. Fisher & R. E. Orth). Fig. 9. Hoplodic&u ucuricornis (Wulp). California; San Diego Co.; Doane Pond, Palomar State Park; 4646 feet (1416 m) elevation; IX-14-1965 (T. W. Fisher & R. E. Orth). Fig. 10. Limnia severu Cresson. California; Shasta Co.; Cayton, 1 mi (1.6 km) east of Hwy. 89; 3050 feet (930 m) elevation; VI-16-1969 (R. E. Orth). Fig. 11. SepedonpcificaCresson. Cali- fornia; Inyo co.; 5 mi (8 km) north of Lone Pine; 3800 feet (1158 m) elevation; V-17-1966 (T. W. Fisher & R. E. Orth). Fig. 12. Tetanoceru plumoosa Loew. California; Alpine Co.; 0.5 mi (0.8km) west of Heenan Lake; 7000 feet (2134m) elevation; VII-20-1967 (R. E. Orth). MARSH FLIES OF CALIFORNIA 57

1 3

4 6

7 8 9

- 10 11 12 58 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 2: Freshwater and hygrophilous terrestrial snails found in California. Fig. 1. L-R. Lymnaeu srugtmlis appressu Say, 1821. Modoc Co.; Likely, canal on Van Lon Ranch; IX-21-1966.Radix auricularia (Linnaeus, 1758). San Bernardino Co.; Big Bear Lake; VI-26-1963. Lymnaea (Srugnicola) pulustris-group. Riverside Co.; Riverside, Santa Ana River; IX-5- 1962. Plunorkllu (Pierosomu) tenuis Dunker, 1850. Riverside Co.; Norco, Silver Lakes; IX-26-1962. Fig. 2. L-R. Physa uirgutu Gould, 1855. Riverside Co.; Rancho California, Temecula Creek; VI-29-1965.Pseudosuc- cinea colume//u (Say, 1817). Riverside Co.; Norco, Silver Lakes; IX-26-1962.O~/otm sillimani (Bland, 1865). Riverside Co.; Riverside, Santa Ana River; IX-5-1962. Succineu culiforniensis Fischer and Crosse, 1878. Riverside CO.; Rancho California, Vail Lake; VI-29-1965.

BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 3: California collecting sites. Fig. 1. Lake shore. Modoc Co.; Lily Lake, 8 mi (12.9 km) east of Hwy 395; 6700 feet (2042 m) elevation; 1968. Fig. 2. Marsh. San Bernardino Co.; Barton Flats, marsh adjacent to South Fork of Santa Ana River; 6200 feet (1890 m) elevation; 1969. Fig. 3. Flowing spring. Nevada Co.; Boca Spring; 5900 feet (1798 m) elevation; 1966. Fig. 4. Mesic. Siskiyou Co.; Bartle, McIntosh Ranch; 4000 feet (1219 m) elevation; 1973. Fig. 5. Wet pasture. Mendocino Co.; 2 mi (3.2 km) north of Willits; 1330 feet (405 m) elevation; 1969. Fig. 6. Stream. Orange Co.; San Juan Creek; 200 feet (61 m) elevation; 1964. MARSH FLIES OF CALIFORNIA 61

5 6 62 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 4: Characters used in the identification of adult Sciomyzidae. Fig. 1. Pherbeliu paralkkz (Walker), 1852. Male. Side view. Napa Co.; 0.5 mi (0.8 km) east of Younlville; 110 feet (34 m) elevation; VII-9-1965 (T. W. Fisher). Fig. 2. Pher&//iu griseolo (Falldn), 1820. Frontal view. Shasta Co.; 2 mi (3.2 km) south of Pondosa, Hwy 89; 4140 feet (1262 m) elevation; VI-16-1969 (R. E. Orth). fob, fronto-orbital bristles (parafrontals); iv, inner vertical bristle; ob, ocellar bristle; ov, outer vertical bristle; pv, postvertical bristles. Fig. 3. Tetunoceru uicirm (Maquart), 1843. Wing. Shasta Co.; 1 mi (1.6 km) west of Hatchet Mountain Summit; 4200 feet (1280 m) elevation; VII-8-1970 (F&O). al,* = anal veins 1 and 2; C = costa; cell AM - apical media cell; cell cu - cubital cell; cell dc = discal cell; cell R,, = radial cells 1-5;cu = cubital vein; M = media vein; RI-5= fadial veins 1- 5; Sc = subcosta; smp = stump vein; ta = radio-medial crossvein; tp = medial crossvein. Fig. 4. Sciomyzu simplex (Fallh), 1820. Fore tibia and tarsus. Siskiyou Co.; Grass Lake, Hwy 97; 5122 feet (1561 m) elevation; VI-10-1966 (F&O). pa, preapical bristles; tb, tibia. Fig. 5. Pherbe//ia pudlelu (Walker), 1852. Antero-lateral view of thorax. Santa Clara Co.; Llagas Creek; 280 feet (85 m) elevation; IV-25-1968(F&O). cxI, fore coxa; n, notum; PI,, propleura; pl, mesopleura; ppb, preapical bristle. Fig. 6. LimnM pubexens (Day), 1881. Female. Postero-lateral view of thorax. Shasta Co.; Hat Creek, Hwy 299; 2650 feet (808 m) elevation; VI-21-1972(F&O). pt, pteropleura; ssc, supraspiracular convex- ity; vb, valar bristle. Fig. 7. sepedon cujdlleiiisher and Orth, 1969. Male. Hind leg. Nevada; Douglas CO.;Hwy 395, 2 mi (3.2 km) north of junction with Hwy 3, 5500 feet (1676 m) elevation; V-4-1972 (R. E. Orth). cxp hind coxa; fm, femur; tb, tibia. Fig. 8. Anrichem kestaceu Melander, 1920. Hind tibia and tarsus. Mendocino Co.; Ten Mile River, 0.5 mi (0.8 km) east of Hwy 1; 5 feet (1.5 rn) elevation; VI-12-1966(FBtO). pa, preapical bristles; tb, tibia. MARSH JWES OF CALIFORNIA 63

,sc PI ,c F ,cell R1

n 5

Pl I' c: 'tp PP 64 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 5: Head profiles representative of sciomyzine genera found in California. Figs. 1-4, Sciomyzini; Figs. 5-13, Tetanocerini. Fig. 1. Atrichomelinu puberu (Loew), 1862. Male. Siskiyou Co.; Grass Lake, 5122 feet (1561 m) elevation; VIII-6-1965 (F&O). Fig. 2. Scioqyzu simp/exFall6n, 1820. Male. Modoc Co.; 2 mi (3.2 km) south of Alturas, 4300 feet (131 1 m) elevation; VII-23-1967 (F&O). Fig. 3. Pherbe//iu griseolu (Falldn), 1820. Male. Oregon; Klamath Co.; Klamath Game Refuge, 4540 feet (1384 m) elevation; VII-7-1968 (F&O). Fig. 4. Pteromi- cru sisk#ouensisFisher and Orth, 1966. Male. Siskiyou Co.; Grass Lake, 5122 feet (1561 m) elevation; VI-17-1969 (R. E. Orth). Fig. 5. Antichuetu testuceu Melander, 1920. Female. Oregon; Josephine Co.; 0.5 mi (0.8 km) south of Cave Junction, 1200 feet (366 m) elevation; VI-21-1969(R. E. Orth). Fig. 6. Dicm montunu Steyskat, 1954. Male. Nevada; Churchill Co.; 9 mi (14.4 km) southeast of Fallon; 40oO feet (1219 m) elevation; VII-24-1972 (F&O). MARSH FLIES OF CALIFORNIA 65

1.0 mm 66 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 6: Head profiles representative of sciomyzine genera found in California. Fig. 1. Dicmcium firmum Steyskal, 1956. Male. Siskiyou Co.; Bartle; 4OOO feet (1219 m) elevation; VII-31-1973 (F&O). Fig. 2. EIgivu conmu (Steyskal), 1954. Male. Modoc Co.; 3 mi (4.8 km) north of Eagleville; 4640 feet (1414 m) elevation; VII-10-1968 (F&O). Fig. 3. Hop/odic~uucuticornis (Wulp), 1897. Male. Nevada; Douglas CO.; Hwy 395, 2 mi (3.2 km) north of junction with Hwy 3, 5500 feet (1676 m) elevation; V-4-1972 (R. E. Orth). Fig. 4. Limnia sewem Cresson, 1920. Male. Oregon; Jackson Co.; north of Eagle Point, 1330 feet (405 m) elevation; VI-19-1972(F&O). Fig. 5. Renoceru brevis (Cresson), 1920. Male. Oregon; Clackamas Co.; Mt. Hood Natl. Forest; 3380 feet (1030 m) eleva- tion; V1-18-1969 (R. E. Orth). Fig. 6. Seph b~&Steyskal, 1951. Male. Amador Co.; Drytown; 700 feet (213 m) elevation; IX-9-1968(R. E. Orth). Fig. 7. Tefunoceru phrmosu Loew, 1847. Male. San Bernardino Co.; Barton Flats; 6200 feet (1890 m) elevation; VII-9-1969(F&O). MARSH FLIES OF CALIFORNIA 67

1

2

1.0 mm

7 68 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 7: Characters used in the identification of larval Sciomyzidae. Figs. 1-3. Dictya montanu Steyskal, 1954. TETANOCERINI. Laboratory reared third instar larva. Fig. 1. Lateral view. Fig. 2. Cephalopharyngeal skeleton. dc, dorsal cornua; pi, pharyngeal indentation; vc, ventral cornua. Fig. 3. Poste- rior spiracular disc. fh, float hairs. Figs. 4-6. Pherbelliu purullelu (Walker), 1852. SCIOMYZINI. Laboratory reared third instar larva. Fig. 4. Lateral view. sp, spinule patch. Fig. 5. Cephalopharyngeal skeleton. dw, dorsal “window”; vw, ventral “window”. Fig. 6. Poste- rior spiracular disc. MARSH FLIES OF CALIFORNIA 69

1 0.5 mm

2 I 1 0.4 mm 3 W 0.4 mm

7 0.5 mm

5 I I 0.4 mm 0.4 mm 70 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 8: Terminalia of Sciomyzidae found in or near California.

Figs. 1 and 2. Anichomlina ~K&U (Loew), 1862. Male. California; Siskiyou Co.; Grass Lake; 5122 feet (1561 m) elevation; VI-10-1965 (FBCO). Postabdomen, inverted. Fig. 1. Sinistral view. Fig. 2. Posterior view. Figs. 3 and 4. Pherbellu urgyra Verbeke, 1967. Male. Canada; N.W.T.;Hay-River; IX-10-1932(0. Bryant). Postab- domen, inverted. Fig. 3. Sinistral view. Fig. 4. Posterior view. Figs. 5 and 6. Pherbellia griseora (FallBn), 1820. Male. California; Shasta Co.;2 mi (3.2 km) south of Pondosa, Hwy 89; 4140 feet (1262 m) elevation; VI-16-1969(R. E. Orth). Postabdomen, inverted. Fig. 5. Sinistral view. Fig. 6. Poste- rior view. I MARSH FLIES OF CALIFORNIA 71

2 1

A 72 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 9: Terminalia of Sciomyzidae found in or near California. Figs. 1 and 2. Pherbellia idahoensis Steyskai, 1961. Male. California; Santa Clara Co.; Stanford Lake, Stanford University campus; 250 feet (76 m) elevation; VI-8-1965 (T.W. Fisher). Postabdomen, inverted. Fig. 1. Sinistral view. Fig. 2. Posterior view. Figs. 3 and 4. Pherbellio me/anderiSteyskal, 1963. Male. California; Modoc Co.; 2 mi (3.2 km) south of Alturas, 4350 feet (1326 m) elevation; VI-10-1966 (F&O). Postabdomen, inverted. Fig. 3. Sinistral view. Fig. 4. Posterior view. MARSH EIES OF CALIFDRNIA 73 - 0.4 mni

2 74 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 10: Terminalia of Sciomyzidae found in or near California. Figs. 1 and 2. Pherbellia nana nuna (FallCn), 1820. Male. California; Mendocino Co.; 3 mi (4.8 km) north of Willits. Outlet Creek; 1350 feet (411 m) elevation; VI-21-1969 (R. E. Orth). Postabdomen, inverted. Fig. 1. Sinistral view. Fig. 2. Posterior view. Figs. 3 and 4. Pher&//iu oregona Steyskal, 1961. Male. California; Del Norte Co.; 3 mi (4.8 km) north of Crescent City; 40 feet (12 m) elevation; VI-11-1966 (FBtO). Postabdomen, inverted. Fig. 3. Sinistral view. Fig. 4. Posterior view. Figs. 5 and 6. Pherbe//ia para/k/u (Walker), 1852. Male. California; Mono Co.; Fish Slough; 4200 feet (1280 m) elevation; V-23-1968(Fdc.0). Postabdomen, inverted. Fig. 5. Sinistral view. Fig. 6. Posterior view. MARSH nIES OF CALIFORNIA 75

I I 0.4 mm

1

0.4 mm

6 16 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 11: Terminalia of Sciomyzidae found in or near California. Fig. 1. Pher&//iu culijbrnica Orth, 1982. Male. California; Mendocino Co.; 2 mi (3.2 km) north of Willits; 1380 feet (421 m) elevation; V-24-1967 (F8tO). Postabdomen, inverted. Sinistral view. Figs. 2 and 3. Pherbe//ia sehoenherri maculata (Cresson), 1920. Male. California; Modoc Co.; 8.3 mi (13 km) west of Alturas; 4300 feet (1300 m) elevation; VII-19-1967(R. E. Orth). Postabdomen, inverted. Fig. 2. Sinistral view. Fig. 3. Posterior view. Figs. 4-6. Pherbe//iu subri/isOrth and Steyskal, 1980. Male. California; San Bernardino Co.; Little Cienega Seca; 7700 feet (2347 m) elevation; VI-16-1968 (FStO). Postabdomen, inverted. Fig. 4. Sinistral view. Fig. 5. Posterior view. Fig. 6. Ventral view. MARSH FLIES OF CALIFORNIA 77 n

3 78 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 12: Terminalia of Sciomyzidae found in or near California. Figs. 1 and 2. Pherbe//ia trakulam (Loew), 1872. Male. California; Riverside Co.; Lake Hemet; 4500 feet (1372 rn) elevation; VI-25- 1968 (FBtO). Postabdomen, inverted. Fig. 1. Sinistral view. Fig. 2. Posterior view. Figs. 3 and 4. Pherbelfia vitalis (Cresson), 1920. Male. California; Riverside Co.; Rancho California, Vail Lake; 1400 feet (427 m) elevation; V-25-1965 (F8cO). Postabdomen, inverted. Fig. 3. Sinistral view. Fig. 4. Posterior view. Figs. 5-7. Pteromicra pectorosa (Hendel), 1902. California; Del Norte Co.; 3 mi (4.8 km) north of Crescent City; 40 feet (12 m) elevation; VI-11-1966 (F&O). Postabdomen, inverted. Fig. 5. Male. Sinistral view. Fig. 6. Male. Posterior view. Fig. 7. Female. Ventral view. MARSH FLIES OF CALIFORNIA 79

I I / 1 0.4 mm 2

L 0.4 mni 4

I 0.4 mm 7 0.4 mm 80 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 13: Terminalia of Sciomyzidae found in or near California. Figs. 1-3. Pteromicra siskiyouensis Fisher and Orth, 1966. California; Siskiyou Co.; Grass Lake; 5122 feet (1561 m) elevation; VIII-6-1965 (F&O). Postabdomen, inverted. Fig. 1. Male. Sinistral view. Fig. 2. Male. Posterior view. Fig. 3. Female. Ventral view. Figs. 4 and 5. Sciomyza simplex Fallen, 1820. Male. California; Modoc Co.; Likely, north of Pit River; 4400 feel (1341 m) elevation; VIII-23-1%7 (F&O). Postabdomen, inverted. Fig. 4. Sinistral view. Fig. 5. Posterior view. Figs. 6 and 7. Sciomyzu vuria (Coquillett), 1904. Male. California; Modoc Co.; 1 mi (1.6 km) south of Alturas; 4300 feet (1311 m) elevation; V11-19-1967 (R. E. Orth). Postabdomen, inverted. Fig. 6. Sinistral view. Fig. 7. Posterior view. MARSH FLIES OF CALIFORNIA 81

0.4 mm

7 82 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 14: Terminalia of Sciomyzidae found in or near California. Figs. 1-6. Anticbaeta borealisFoote, 1961. Figs. 1-4. Male. Ohio; 4.5 mi (7.2 km) east of Kent; IV-16-1968 (B. A. Foote). Postabdomen, inverted. Fig. 1. Posterior view. Fig. 2. Sinistral view. Fig. 3. Dextral view. Fig. 4. Fifth ven- trite. Figs. 5 and 6. Female. California; Modoc Co.; 3 mi (4.8 km) north of hgleville; 4640 feet (1414 m) elevation; VII-10-1968(F&O). Fig. 5. . Fig. 6. Ventral view of terminalia. Figs. 7-9. Antichaeta robiginosa Melander, 1920. Male. California; Shasta Co.; 5 mi (8 km) northwest of Anderson; 480 feet (146 m) elevation; V-24-1967 (FBtO). Postabdomen, inverted. Fig. 7. Posterior view. Fig. 8. Sinistral view; as, anterior surstylus. Fig. 9. Dextral view. MARSH FLIES OF CALIFORMA 83

1

5

4 0.4 mm 6

as as 84 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 15: Terminalia of Sciomyzidae found in or near California. Figs. 1-3. Anfichuera robiginosa Melander, 1920. Fig. 1. Male. California; Shasta Co.; 5 mi (8 km) northwest of Anderson; 480 feet (146 m) elevation; V-24-1967 (F&O). Fifth ventrite. Fig. 2. Male. California; Mendocino Co.; 2 mi (3.2 km) north of Willits; 1330 feet (405 m) elevation; VI-12-1966 (R. E. Ofth). Antenna. Fig. 3. Female. California; Modoc Co.; 2 mi (3.2 km) south of Alturas; 4300 feet (1311 m) elevation; VI-6-1967 (F&O). Ventral view of ter- minalia. Figs. 4-9. Antichaera testacea Melander, 1920. Figs. 4-7. Male. California; Riverside Co.; Rancho California, Vail Lake; 1400 feet (427 m) elevation; 1V-21-1966 (T.W. Fisher). Postabdomen, inverted. Fig. 4. Posterior view. Fig. 5. Sinistral view. Fig. 6. Dextral view. Fig. 7. Fifth ventrite. Fig. 8. Female. California; San Luis Obispo CO.; San Luisito Creek; 3 mi (4.8 km) southeast of Morro Bay; 75 feet (23 m) elevation; VII-29-1964 (T. W. Fisher). Antenna. Fig. 9. Female. California; Riverside Co.; Lake Hemet; 4500 feet (1372 m) elevation; IV-14-1966 (FBiO). Ventral view of ter- minalia. MARSH FLIES OF CALIFORNIA 85

,. --

1 2 3

5

0.4 mm

8

7 86 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 16: Terminalia of Sciomyzidae found in or near California. Figs. 1-6. Antichaeta wrmlis Fisher and Orth, 1971. California; Mendocino Co.; 2 mi (3.2 km) north of Willits; 1330 feet (405 m) elevation. Figs. 1-3. Male. Postabdomen, inverted. IV-23-1969(F&O). Fig. 1. Posterior view. Fig. 2. Sinistral view. Fig. 3. Dextral view. Fig. 4. Female. Antenna. IV-11-1967(F&O). Figs. 5 and 6. IV-23-1969(F&O). Fig. 5. Male. Fifth ventrite. Fig. 6. Female. Ventral view of terminalia. MARSH FLIES OF CALIFORNIA 87

4

I I 0.4 mm

5 6 88 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 17: Terminalia of Sciomyzidae found in or near California. Figs. 1-4. Dictya expansu Steyskal, 1938. Kansas; Montgomery Co.; 2 mi (3.2 km) northwest of Liberty; IV/14- 1911963 (N. & B. Marston). Figs. 1 and 2. Male. Postabdomen, inverted. Fig. 1. Dextral view. Fig. 2. Anterior view of hypandrium. Figs. 3 and 4. Female, Fig. 3. Internal view of sternites VI1 and VI11 to show apodemes. Fig. 4. Sternites VI-x. Figs. 5-8. Dictyu finrinulisFisher and Orth, 1969. Figs. 5 and 7. Male. California; Nevada Co.; Boca Spring; 5900 feet (1798 m) elevation; VI-7-1966 (F&O), Postabdomen, inverted. Fig. 5. Dextral view; ep, epandrium; hy, hypan- drium; ss, surstylus. Fig. 7. anterior view of hypandrium. Figs. 6 and 8. Female. California; Shasta Co.; Cassel; 3150 feet (960 m) elevation; VII-5-1955 (J. W. MacSwain, UCB). Fig. 6. Internal view of sternites VI1 and VI11 to show apo- demes. Fig. 8. sternites VI-X. MARSH FLIES OF CALIFORNIA 89

X

1x

VI11

VI1

7 s 90 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 18: Terminalia of Sciomyzidae found in or near California. Figs. 1-4 Dictya incisucurran, 1932. Figs. 1 and 2. Male. Mexico; Sonora; 1 mi (1.6 km) north of Imuris; V-11-1953 (E. I. Schlinger & R. C. Bechtel, UCB). Postabdomen, inverted. Fig. 1. Dextral view. Fig. 2. Anterior view of hypan- drium. Figs. 3 and 4. Female. Arizona; Cochise Co.; 5 mi (8 km) west of Portal; 5400 feet (1646 m) elevation; VIII-7- 1958 (P. Opler). Fig. 3. Internal view of sternites VI1 and VI11 to show apodemes. Fig. 4. Sternites VI-X. Figs. 5-8. Dicwa mutthewsiSteyska1, 1960. Figs. 5 and 6. Male. Arizona; Cochise Co.; 5 mi (8 km) west of Portal; 5400 feet (1646 m) elevation; VIII-7-1958 (P. Opler). Postabdomen, inverted. Fig. 5. Dextral view. Fig. 6. Anterior view of hypandrium. Figs. 7 and 8. Female. Mexico; 2 mi (3.2 km) northwest of Puebla; IV-25-1953 (R.C. Bechtel & E. I. Schlinger, UCB). Fig. 7. Internal view of sternites VI1 and VI11 to show apodemes. Fig. 8. Sternites VI-X. MARSH FLIES OF CALIFORNIA 91

4 3

0.4 mm 7 92 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 19: Terminalia of Sciomyzidae found in or near California. Figs. 1-8. Dictya montonu Steyskal, 1954. Figs. 1-4. Form “A”. Figs. 1 and 2. Male. California; Mendocino Co.; Ukiah; 590 feet (180 m) elevation; IV-7-1964 (T. W. Fisher). Postabdomen, inverted. Fig. 1. Dextral view. Fig. 2. Anterior view of hypandrium. Figs. 3 and 4. Female. California; Lassen Co.; 2 mi (3.2 km) west of Hallelujah Junc- tion; 4800 feet (1463 m) elevation; VI-27-1949 (D. Cox). Fig. 3. Internal view of sternites VI1 and VI11 to show apo- demes. Fig. 4. Sternites VI-X. Figs. 5-8. Form “Alturas”. California; Modoc Co.; 1 mi (1.6 krn) south of Alturas; 4300 feet (1311 m) elevation; IX-21-1966 (F&O). Postabdomen, inverted. Fig. 5. Male. Dextral view. Fig. 6. Male. Anterior view of hypandriurn. Fig. 7. Female. Internal view of sternites VI1 and VI11 to show apodemes. Fig. 8. Female. Sternites VI-X. MARSH FLIES OF CALIFORNIA 93

2

1 J 4 3

6

0.4 mm 7 x 94 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 20: Terminalia of Sciomyzidae found in or near California.

Figs. 1-4. Dicw ~OR~URUSteyskal, 1954. Form “so. Cal.”. Figs. 1 and 2. Male. California; Orange Co.; San Juan Creek; 200 feet (61 m) elevation; XII-27-1962 (R. E. Orth). Postabdomen, inverted. Fig. 1. Dextral view. Fig. 2. Ante- rior view of hypandrium. Figs. 3 and 4. Female. California; San Diego Co.; Scissors Crossing; 2280 feet (695 m) eleva- tion; IV-5-1966(T. W. Fisher). Fig. 3. Internal view of sternites VI1 and VI11 to show apodemes. Fig. 4. Sternites VI-X. Figs. 5-8. Dicw texensiscurran, 1932. Figs. 5 and 6. Male. California; Riverside Co.; 2 mi (3.2 km) southeast of Oasis; -180 feet (-55 m) elevation; 111-23-1967 (FStO). Postabdomen, inverted. Fig. 5. Dextral view. Fig. 6. Anterior view of hypandrium. Figs. 7 and 8. Female. Arizona; Phoenix, El Canto Park; XII-6-1962(T. W.Fisher). Fig. 7. Inter- nal view of sternites VI1 and VI11 to show apodemes. Fig. 8. Sternites VI-X. MARSH FLIES OF CALIFORNIA 95

2

3

0.4 mm 7 96 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 21 : Terminalia of Sciomyzidae found in or near California. Figs. 1-3. Dicryaciumjrmum Steyskal, 1956. Male. California; Siskiyou Co.; Bartle, McIntosh Ranch: 4000 feet (1 219 m) elevation; VII-23-1973 (F&O). Postabdomen, inverted. Fig. 1. Sinistral view. Fig. 2. Protandrium. Fig. 3. Abdomi- nal sterna. Figs. 4 and 5. Elgiva connexcI (Steyskal), 1954. Male. California; Modoc Co.; 3 mi (4.8 km) north of Eagleville; 4600 feet (1402 m) elevation; VII-24-1967(F&O). Postabdomen, inverted. Fig. 4. Sinistral view. Fig. 5. Posterior view. Figs. 6-9. E&va solicita (Harris), 1780. Male. California; Sierra Co.; 3.5 mi (5.6 km) northwest of Sierraville; 4900 feet (1494 m) elevation; IX-22-1966 (F&O). Postabdomen, inverted. Fig. 6. Sinistral view. Fig. 7. Hypandrium, sinis- tral view. Fig. 8. Hypandrium, dextral view. Fig. 9. Posterior view. MARSH FUES OF CALIFORNIA 97

7

9 98 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 22: Terminalia of Sciomyzidae found in or near California. Figs. 1 and 2. Hoplodic/jw ucuticwnis (Wulp), 1897. Fig. 1. Male. California; Orange Co.; San Juan Creek; 200 feet (61 m) elevation; VIII-24-1965 (R. E. Orth). Postabdomen, inverted, dextral view. Fig. 2. Female. California; Riverside Co.; Santa Ana River, Riverside; 820 feet (250 m) elevation; 111-8-1963 (FBO). Ventral view of terrninalia. Figs. 3-6. Limniu boscii (Robineau-Desvoidy), 1830. Male. California; Mono Co.; Fish Slough, IO mi (16 km) north of Bishop; 4250 feet (1295 rn) elevation; IX-7-1967 (FBEO). Postabdomen, inverted. Fig. 3. Sinistral view. Fig. 4. Pro- tandrium (terminal process), dextral view. Fig. 5. Protandrium, anterior view; VI, sixth sternite. Fig. 6. Prosternum; ant, anterior; PO, posterior. Figs. 7-9. Limnb inopo (Adams), 1904. Male. Fig. 7. California; Alpine Co.; Heenan Lake; 7000 feet (2134 m) elevation; VI-13-1966 (FBEO). Postabdomen, inverted. Sinistral view. Figs. 8 and 9. California; Modoc Co.; Cedar Pass Campground; 5800 feet (1768 m) elevation; VIII-8-1968 (F&O). Fig. 8. Protandrium, anterior view. Fig. 9. Abdominal sterna. MARSH FLIES OF CALIFORNIA 99

0.4 mm

1

2

6

9 100 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 23: Terminalia of Sciomyzidae found in or near California. Figs. 1 and 2. Limn& pukscens (Day), 1881. Male. Oregon; Jackson Co.; Touvelle State Park; 8 mi (12.9 km) north of Medford; 1350 feet (411 m) elevation; V-21-1960 (D. R. Smith). Postabdomen, inverted. Fig. 1. Sinistral view. Fig. 2. Protandrium, anterior view. Figs. 3-6. Litnnia san&vaknsis Fisher and Orth, 1978. Male. New Mexico; Sandoval Co.; East Fork Jemez River, Las Conchas Campground; 8300 feet (2530 m) elevation; VIII-13-1967 (R. E. Orth). Postabdomen, inverted. Fig. 3. Sinis- tral view. Fig. 4. Protandrium, anterior view. Fig. 5. Protandrium (terminal process), dextral view. Fig. 6. Prosternum. Figs. 7 and 8. Limn& seuera Cresson, 1920. Male. California; Inyo Co.; Lake Sabrina; 9OOO feet (2743 m) elevation; IX-6-1967(FBO). Postabdomen, inverted. Fig. 7. Protandrium, anterior view. Fig. 8. Sinistral view. Fig. 9. Renocera brevis (Cresson), 1920. Male. Oregon; Clackamus Go.; 0.5 mi (0.8km) west of Government Camp; 3600 feet (1097m) elevation; VII-9-1970(F&O). Postabdomen, inverted, sinistral view. MARSH FLIES OF CALIFORNIA 101

fl 0.4 mm

6 102 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 24: Terminalia of Sciomyzidae found in or near California. Figs. 1-3. Sepedon bifdu Steyskal, 1951. Male. California; Riverside Co.; Lake Hemet; 4500 feet (1372 m) elevation; IV-22-1965(T. W. Fisher). Fig. 1. Postabdomen, inverted, sinistral view. Fig. 2. Postabdomen, ventral view. Fig. 3. Hind femur and tibia, sinistral view. Figs. 4 and 5. Sepdon boreulisSteyska1, 1951. Male. California; Nevada Co.; Boca Spring; 5900 feet (1798 m) eleva- tion; VIII-3-1%5 (F&O). Fig. 4. Postabdomen, inverted, sinistral view. Fig. 5. Postabdomen, ventral view. Figs. 6-10. Sepedon cupeki Fisher and Orth, 1969. California; Inyo Co.; 2.5 mi (4 km) west of Bishop; 4250 feet (1295 m) elevation; VIII-1-1965(FBrO). Figs. 6-9.Male. Fig. 6. Postabdomen, inverted, sinistral view. Fig. 7, Postab- domen, ventral view. Fig. 8. Hind femur and tibia, sinistral view. Fig. 9. Aedeagus, sinistral view. Fig. 10. Female. Ven- tral view of terminalia. MARSH FLIES OF CALIFORNIA 103 104 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 25: Terminalia of Sciomyzidae found in or near California. Figs. 1 and 2. Sepedon cascadensisFisher and Orth, 1974. Male. Oregon; Hood River Co.; 0.5 mi (0.8 km) south of Sherwood Campground, Mt. Hood National Forest; 3180 feet (969 m) elevation; VI-18-1969 (F&O). Fig. 1. Postabdo- men, inverted, sinistral view. Fig. 2. Hind femur and tibia, sinistral view. Figs. 3-5. Sepedon~~i~nnisfuscipennisLoew,1859. Figs. 3 and 4. Male. California; Sierra Co.; 2 mi (3.2 km) west of Sierraville; 4940 feet (1506 m) elevation; VIII-23-1967 (FBtO). Postabdomen, inverted. Fig. 3. Sinistral view. Fig. 4. Posterior view. Fig. 5. Female. Oregon; Klamath Co.; Klamath Game Refuge; 4540 feet (1384 m) elevation; VIII-7- 1968 (F&O). Ventral view of terminalia. Figs. 6-10. Sepedon pacifica Cresson, 1914. Male. Fig. 6. Colorado; Fort Collins; IV-16-1958 (N.Marston). Postabdo- men, inverted, sinistral view. Figs. 7-10. California; San Diego Co.; Doane Pond, Palomar State Park; 4646 feet (1416 m) elevation; IX-14-1965 (F&O). Fig. 7. Aedeagus, sinistral view. Fig. 8. Aedeagus, oblique anterior view from direc- tion of arrow in Fig. 7. Fig. 9. Hypandrium, sinistral view. Fig. 10. Hypandrium, viewed from direction of arrow in Fig. 9. MARSH FLIES OF CALIFORNIA 105

.I Q, -.. __ _.-- 7 9 10 106 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 26: Terminalia of Sciomyzidae found in or near California. Figs. 1-3. Sepedon spinipes americuna Steyskal, 1951. California; Sierra Co.; 2 mi (3.2 km) west of Sierraville; 4940 feet (1506 rn) elevation; VIII-23-1967 (F&O). Figs. 1 and 2. Male. Postabdomen, inverted. Fig. 1. Sinistral view. Fig. 2. Posterior view. Fig. 3. Female. Ventral view of terminalia. Figs. 4 and 5. Tetanocera bergisteyskal, 1954. Male. Oregon; Douglas Co.; 7 mi (11 km) east of Reedsport, Umpqua River; 20 feet (6.1 rn) elevation; VIII-6-1968 (F&O). Postabdomen, inverted. Fig. 4. Sinistral view. Fig. 5. Posterior view. Figs. 6-9. Tetanoceraferruginea FallBn, 1823. Figs. 6, 7 and 9. Male. California; Modoc CO.; 2 mi (3.2 km) south of Alturas; 4300 feet (1311 m) elevation; VI-10-1966 (FBtO). Postabdomen, inverted. Fig. 6. Sinistral view. Fig. 7. Poste- rior view. Fig. 9. Protandrium, anterior view. Fig. 8. Female. California; Modoc Co.; 5 mi SE Tulelake; 4000 feet (1219 m) elevation; VI-10-1966 (F&O). Ventral view of terminalia. MARSH FLIES OF CALIFORNIA 107

3

0.4 mm 108 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 27: Terminalia of Sciomyzidae found in or near California. Figs. 1-3. Te#moceraficscinervis(Zetterstedt), 1838. Male. Oregon; Hood River Co.; 0.5 mi (0.8 km) south of Sher- wood Campground, Mt. Hood National Forest; 3180 feet (969 m) elevation; VI-18-1969 (R. E. Orth). Postabdomen, inverted. Fig. 1. Sinistral view. Fig. 2. Protandrium, anterior view. Fig. 3. Posterior view. Figs. 4-7. Tetanocera lotifibuh Frey, 1924. Figs. 4 and 5. Male. California; Modoc Co.; 2 mi (33.2 km) south of Alturas; 4300 feet (1311 m) elevation; VIII-5-1965 (FgiO). Postabdomen, inverted. Fig. 4. Sinistral view. Fig. 5. Poste- rior view. Figs. 6 and 7. California; Modoc Co.; Hackamore Reservoir; 4600 feet (1402 rn) elevation; VI-10-1966 (F&O). Fig. 6. Male. Protandrium, anterior view. Fig. 7. Female. Ventral view of terminalia. Figs. 8-10. Tetonocera foewiSteyskal, 1959. Male. California; Del Norte Co.; 3 mi (4.8 km) north of Crescent City; 50 feet (15 m) elevation; VI-14-1%5 (R. E. Orth). Postabdomen, inverted. Fig. 8. Sinistral view. Fig. 9. Posterior view. Fig. 10. Protandrium, anterior view. MARSH FLIES OF CALIFORNIA 109

3

0.4 mm

\ 7 p-..__. 9 110 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 28: Terminalia of Sciomyzidae found in or near California. Fig. 1. Tetanocera loewisteyskal, 1959. Female. California; Del Norte Co.; 3 mi (4.8 km) north of Crescent City; 50 feet (15 m) elevation; VI-14-1965 (F&O). Ventral view of terminalia. Figs. 2-5. Tetanocera mesopora Steyskal, 1959. California; Plumas Co.; 0.5 mi (0.8 km) south of Crescent Mills; 3500 feet (1067 m) elevation; VI-8-1966(F&O). Figs. 2-4.Male. Postabdomen, inverted. Fig. 2. Sinistral view. Fig. 3. Pos- terior view. Fig. 4. Protandrium, anterior view. Fig. 5. Female. Ventral view of terminalia. Figs. 6-8. Tetanocera obfusifibula Melander, 1920. Male. California; Plumas Co.; 0.5 mi (0.8 km) south of Crescent Mills; 3500 feet (1067 m) elevation; VI-8-1966 (F&O). Postabdomen, inverted. Fig. 6. Posterior view. Fig. 7. Protan- drium, anterior view. Fig. 8. Sinistral view. MARSH FLIES OF CALIFORNIA 111

L 0.4 mm 112 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 29: Terminalia of Sciomyzidae found in or near California. Fig. 1. Tetanocera obmsifibula Melander, 1920. Female. California; Plumas Co.; 0.5 mi (0.8 km) south of Crescent Mills; 3500 feet (1067 m) elevation; VI-8-1966 (F&O). Ventral view of terminalia. Figs. 2-5. Tetanocera plebeiu Loew, 1862. California; Plumas Co.; Meadow Valley, Schneider Creek; 3800 feet (1 158 m) elevation; VI-8-1966 (F&O). Figs. 2-4. Male. Postabdomen, inverted. Fig. 2. Sinistral view. Fig. 3. Posterior view. Fig. 4. Protandrium, anterior view. Fig. 5. Female. California; Sierra Co.; northwest of Sattley, Turner Ranch; 5000 feet (1524 m) elevation; VII-9-1968 (F&O). Ventral view of terminalia. Figs. 6-9. Tetanocera plumosa Loew, 1847. Figs. 6-8. Male. California; San Bernardino Co.; Little Cienega Seca; 7750 feet (2362 m) elevation; VII-9-1969 (F&O). Postabdomen, inverted. Fig. 6. Sinistral view. Fig. 7. Posterior view. Fig. 8. Protandrium, anterior view. Fig. 9. Female. California; San Bernardino Co.; Barton Flats, Seven Oaks Road; 6200 feet (1890 m) elevation; VII-23-1969 (R.E. Orth). Ventral view of terminalia. MARSH FLIES OF CALIFORNIA 113 114 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 30: Terminalia of Sciomyzidae found in or near California. Figs. 1-4. Tetonoceru robusto Loew, 1847. Figs. 1, 3, and 4. Male. California; Del Norte Co.; Crescent City; 30 feet (9 m) elevation; VI-11-1966(F&O). Postabdomen, inverted. Fig. 1. Sinistral view. Fig. 3. Posterior view. Fig. 4. Pro- tandrium, anterior view. Fig. 2. Female. California; Modoc Co.; 4 mi (6.4 km) south of Davis Creek; 4800 feet (1463 m) elevation; VI-9-1966(FbiO). Ventral view of terminalia. Figs. 5-7. Tetonoceru rofundicornisLoew, 1861. Male. Nebraska; Lincoln Co.; 2 mi (3.2 km) east of North Platte; 2800 feet (853 m) elevation; VII-24-1970(F&O). Postabdomen, inverted. Fig. 5. Sinistral view. Fig. 6. Posterior view. Fig. 7. Protandrium, anterior view. MARSH FLIES OF CALIFORNIA 115

6

, I 0.4 mm 116 BULLETIN OF THE CALIFORNIA INSECT SURVEY

PLATE 31: Terminalia of Sciomyzidae found in or near California. Figs. 1-4. Tetunoceru sororMelander, 1920. Figs. 1-3. Male. California; Shasta Co.; 2 mi (3.2 km) south of Pondosa, Little Bear Flat; 4140 feet (1262 m) elevation; VIII-8-1968 (F&O). Postabdomen, inverted. Fig. 1. Sinistral view. Fig. 2. Posterior view. Fig. 3. Protandrium, anterior view. Fig. 4. Female. Oregon; Wasco Co.; Hwy 26, 1 mi (1.6 km) south of Hwy 216 junction; Mt. Hood National Forest; 3200 feet (975 m) elevation; VI-20-1969 (F&O). Ventral view of terminalia. Figs. 5-8. Tetanocera vicina Macquart, 1843. Figs. 5-7. Male. California; Plumas Co.; Meadow Valley, Schneider Creek; 3800 feet (1158 m) elevation; VI-8-1966 (F&O). Postabdomen, inverted. Fig. 5. Sinistral view. Fig. 6. Posterior view. Fig. 7. Protandrium, anterior view. Fig. 8. Female. California; Siskiyou Co.; 15 mi (24 km) southwest of Gazelle; 4180 feet (1274 m) elevation; VI-11-1966 (F&O). Ventral view of terminalia. MARSH FLIES OF CALIFORNIA 117

9

1

5

0.4 ni in