Studies of Ephydrinae
(Diptera: Ephydridae), VII:
Revision of the Genus
Setacera Cresson
WAYNE N. MATHIS
J
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? NUMBER 350
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S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y ? N U M B E R 350
Studies of Ephydrinae
(Diptera: Ephydridae), VII
Revision of the Genus
Setacera Cresson
Wayne N. Mathis
SMITHSONIAN INSTITUTION PRESS
City of Washington
1982
A B S T R A C T
Mathis, Wayne N. Studies of Ephydrinae (Diptera: Ephydridae), VII: Re-
vision of the Genus Setacera Cresson. Smithsonian Contributions to Zoology, number
350, 57 pages, 138 figures, 1982.?The genus Setacera Cresson is revised on a
world basis. Setacera is shown to be a monophyletic lineage within the tribe
Ephydrini, closely related to Ephydra Fallen. Like most ephydrines, the im-
mature stages of Setacera inhabit floating algal mats, primarily in fresh-water
environments. A hypothetical phytogeny for the genus is proposed in which
the species are arranged in five species groups. The cladistic relationships
between these groups are indicated by morphological character evidence.
Three new species are described: S. freidbergi (Israel and Iran), S. jamesi (coast
of California, Oregon, and Washington), and S. trichoscelis (Ecuador). Four
synonyms are newly proposed, all of Ephydra breviventris Loew: S. fluxa Miyagi,
E. glabra Meijere, E. laeta Hendel, and S. pedalis Cresson. Keys to species
groups and species, illustrations, and distribution maps are provided.
OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded
in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea
cavernosa (Linnaeus).
Library of Congress Cataloging in Publication Data
Mathis, Wayne N.
Studies of Ephydrinae (Diptera: Ephydridae)
(Smithsonian contributions to zoology ; no. 285, 295, 303, 325, 350)
Includes bibliographies.
Contents: 1. Revisions of Parascatella Cresson and the triseta group of Scatella Robineau?
Desvoidy / Wayne N. Mathis and Guy E. Shewell?2. Phylogeny, classification, and
zoogeography of nearctic Lamproscatella Hendel / Wayne N. Mathis?[etc.]?7. Revision of
the genus Setacera Cresson / Wayne N. Mathis.
Supt. of Docs, no.: SI 1.27:350
1. Ephydridae. I. Shewell, Guy E. II. Wirth, Willis Wagner. III. Title. IV. Series: Smith-
sonian contributions to zoology ; no. 285, etc
QL1.S54 no. 285, etc. [QL537.E7] 591s [595.77'4] 78-606062 AACR2
Contents
Page
Introduction 1
Methods 2
Acknowledgments 2
Tribe EPHYDRINI Zetterstedt 3
Genus Setacera Cresson 4
Key to New World Species of Setacera 9
Key to Old World Species of Setacera 10
The micans Group 11
1. Setacera atrovirens (Loew) 12
2. Setacera micans (Haliday) 15
The breviventris Group 17
3. Setacera breviventris (Loew) 18
4. Setacera multicolor (Soika) 22
5. Setacera viridis Miyagi 23
The aurata Group 25
6. Setacera aurata (Stenhammar) 26
The trina Group 27
7. Setacera freidbergi, new species 28
8. Setacera meneghinii Canzoneri 31
9. Setacera trina Collin 32
The aldrichi Group 34
10. Setacera aldrichi Cresson 34
The pacifica Group 36
11. Setacera durani Cresson 38
12. Setacera jamesi, new species 40
13. Setacera needhami Johannsen 42
14. Setacera pacifica (Cresson) 45
15. Setacera pilicornis (Coquillett) 49
16. Setacera trichoscelis, new species 53
Literature Cited 55
FIGURE 1.?Setacera freidbergt, habitus.
Studies of Ephydrinae
(Diptera: Ephydridae), VII
Revision of the Genus
Setacera Cresson
Wayne N. Mathis
Introduction
As a continuing endeavor towards a generic
revision and catalog of the subfamily Ephydrinae,
this study of the genus Setacera Cresson was un-
dertaken. Although my initial intention was to
revise the New World species only, it became
necessary to expand the coverage to a world-wide
basis to understand and characterize the genus
properly. This has delayed publication of the
New World portion and has markedly increased
logistical problems, particularly in dealing with
collections from the Old World, but from the
perspective of my overall goal, the revision is
substantially more valuable.
Setacera is a comparatively new generic name
(Cresson, 1930). Species described previously
were included in the genus Ephydra Fallen, a
practice still preferred by a few authors (Soika,
1956; Dahl, 1959; Nartschuk, 1970). Until now
no comprehensive study of the genus has been
available, and species descriptions were either
published singly or as parts of faunal studies of
limited geographic scope. Only two major faunal
reviews are available. Becker's (1926) monograph
of the palaearctic Ephydridae included the de-
Wayne N. Mathis, Department of Entomology, National Museum of
Natural History, Smithsonian Institution, Washington, D.C. 20560.
scribed species of that region as segregates of the
genus Ephydra, and Sturtevant and Wheeler's
(1954) synopsis of North American species in-
cluded a review of Setacera.
Prior to the present study, 18 names were
available. Of these, 13 are now considered to be
valid, and five are relegated to junior synonymy.
With description of three new species, the genus
now comprises 16 species, and each major faunal
realm is represented by at least one species. The
New World fauna is richest, with nine species,
and North America alone has eight. Only two
species are known to occur in more than one
major faunal realm: S. micans (Haliday) is holarc-
tic, and S. breviventris (Loew) is found in the
Palaearctic, Afrotropical, Oriental, and Austra-
lian regions. Setacera is primarily a northern tem-
perate genus, with only three species occurring
below the Equator, and two of these, S. breviventris
and S. multicolor (Soika), are found in austral
temperate areas as well.
The life history of the genus is virtually un-
known. References are few and scattered, and the
immature stages of only two species have been
described (Johannsen, 1935; Foote, 1982). (The
immature stages of "E. micans" were discussed by
Beyer (1939), although I suspect that he was
dealing with a species of Ephydra.)
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
My objective with this revision is to treat all
known species in a comprehensive study, with
focus also on intrageneric relationships. The tribal
and subfamilial framework will be dealt with
elsewhere.
METHODS.?The methods and descriptive for-
mat used generally in this study were explained
in parts I-IH and V-VI of the Ephydrinae series
(Mathis and Shewell, 1978; Mathis, 1979b, 1980,
1982; Mathis and Wirth, 1981; Mathis and Simp-
son, 1981), part I of the Notiphilinae series
(Mathis, 1979a), and elsewhere (Mathis, 1979c).
The following procedures or characters, however,
need further explanation.
Velvety tomentose band: At the vertex of the head
and immediately adjacent to each compound eye
is an area that is densely tomentose, appearing
velvety. The extent and shape of this area is a
good character, especially for characterizing spe-
cies groups within Setacera.
Facial angle: Angle formed by anterior surface
of face (prefrons) and dorsal surface as viewed
from a lateral aspect.
Male 5th tergal ratio: Tergal length/tergal
width. Both measurements are maximum dis-
tances as viewed from a dorsal aspect.
Female ventral receptacle: Use of characters of
this structure is of recent development (Clausen
and Cook, 1971; Clausen, 1973, 1977; Mathis,
1979a, 1980), and it is appropriate to formalize
the terminology for specific parts. The receptacle
can be generally divided into two major parts,
the operculum and the extending process. The
operculum is the usually simple, hatlike structure
situated at the dorsum. Directions are arbitrarily
selected, operculum dorsad, although they agree
with the illustrated aspect most often published
(see references listed previously). The extending
process, the remainder of the receptacle, exhibits
considerable diversity of structure, hence the ter-
minology for the various parts. Extending ven-
trally from the operculum and ending at the
juncture with the ventral, curved portion, is usu-
ally a necklike process that I call the cervix. The
ventral portion, usually curved, is designated as
the corpus. The shape of the various parts of the
extending process, their relative size, etc., are
excellent characters, used extensively in this paper
to characterize higher taxa, species groups, etc.
ACKNOWLEDGMENTS.?Numerous persons and
institutions have cooperated to make this study
possible. I wish to express my appreciation for
their consideration, especially to the curators and
their respective institutions for the loan of speci-
mens (an asterisk indicates collections from which
type specimens were borrowed).
AMNH American Museum of Natural History, New
York (Dr. Pedro W. Wygodzinsky)
ANIC Australian National Insect Collection, CSIRO,
Division of Entomology, Canberra, Australia
(Dr. D. H. Colless)
ANSP* Academy of Natural Sciences of Philadelphia
(Dr. Daniel Otte)
BMNH British Museum (Natural History), London,
England (Mr. Brian H. Cogan)
CAS California Academy of Sciences, San Francisco
(Dr. Paul H. Arnaud, Jr.)
CNC Canadian National Collection, Ottawa, On-
tario, Canada (Dr. J. R. Vockeroth)
CU Cornell University, Ithaca, New York (Dr. L.
L. Pechuman)
DEI former Deutsches Entomologisches Institut, col-
lections in the Institut fur Pflanzenschutzfor-
schung, Zweigstelle Eberswalde, Abteilung
Taxonomie der Insekten, Eberswalde, Ger-
many (DDR) (Dr. Giinter Morge)
DLD personal collection of Dr. D. L. Deonier, Ox-
ford, Ohio
HNHM Hungarian Natural History Museum, Buda-
pest, Hungary (Dr. L. Papp)
HU* Museum fur Naturkunde, Humbolt Universi-
tat, Berlin, DDR (Dr. H. Schumann)
HUS* Hokkaido University, Sapporo, Japan (Dr. S.
Takagi)
ITZA* Instituut voor Taxonomische Zoologie, Zoolo-
gisch Museum, Universiteit van Amsterdam,
Amsterdam, Netherlands (Dr. Th. H. van
Leeuwen)
KSU Kent State University, Kent, Ohio (Dr. B. A.
Foote)
KU University of Kansas, Snow Entomological Mu-
seum, Lawrence, Kansas (Dr. George W.
Byers)
MCSNV Museo Civico di Storia Naturale de Venezia,
Italia (Dr. Silvano Canzoneri)
MRYAC* Musee Royal de I'Afrique Centrale, Tervuren,
Belgium (Drs. R. Jocque and J. Decelle)
NUMBER 350
NMW Naturhistorisches Museum, Wien, Austria (Dr.
Ruth Contreras-Lichtenberg)
NRS Naturhistoriska Riskmuseet, Stockholm, Swe-
den (Dr. Per Inge Persson)
UCD University of California, Davis (Dr. Robert O.
Schuster)
UCR University of California, Riverside (Mr. Saul I.
Frommer)
UMN University of Minnesota, St. Paul (Dr. Philip J.
Clausen)
UMO University Museum, Oxford University, Ox-
ford, England (Dr. M.W.R. de V. Graham)
USNM* former United States National Museum collec-
tions in the National Museum of Natural
History, Smithsonian Institution
WNM personal collection of Dr. Wayne N. Mathis,
College Park, Maryland
WSU M. T. James Insect Collection, Washington
State University, Pullman (Dr. William J.
Turner)
ZIL Zoological Institute, Lund University, Lund,
Sweden (Dr. Hugo Andersson)
Miss Hollis B. Williams prepared all of the
maps and organized the locality data; Mr. L.
Michael Druckenbrod, Ms. Amy Bartlett, and
Ms. M. Ryan rendered the habitus illustrations;
the frontispiece was done by Mr. George L. Ven-
able. The manuscript was typed by Ms. Noreen
Connell and was critically reviewed by Drs. Willis
W. Wirth, B. A. Foote, and Wayne E. Clark. I
also wish to thank Dr. S. Dillon Ripley, Secretary
of the Smithsonian Institution, for financial sup-
port to conduct field work through a Fluid Re-
search Grant.
Tribe EPHYDRINI Zetterstedt
EPHYDRINI Zetterstedt, 1837:48 [as the "family" Ephydri-
nae].?Wirth and Stone, 1956:45 [first use formally as a
tribe].
DIAGNOSIS.?Specimens of Ephydrini may be
distinguished from other Ephydridae by the fol-
lowing combination of character states.
Adults: Mesofrons subquadrate, becoming
slightly wider posteriorly, with shiny, metallic
luster; frequently with convergent intrafrontal
bristles inserted near anterior margin of meso-
frons; dorsum of interfoveal carina usually shiny,
concolorous with mesofrons; fronto-orbital bris-
tles lateroclinate; facial setae along oral margin
usually dense and long; dorsocentral bristles 5
pairs, anterior 1 or 2 pairs presutural, although
sometimes weak; intrapostalar bristle well devel-
oped, at least equal to one-half length of postalar
bristle; supra-alar bristle well developed, sub-
equal to postalar bristle; notopleuron sparsely
setulose; propleuron setulose; prosternum setose,
usually more evident along posterior margin near
forecoxae; mesopleuron with 1 large bristle near
middle along posterior margin, several smaller
bristles or setae may also be present; hind coxal
strap setose; pulvilli rudimentary or lacking; tar-
sal claws shallowly curved and usually elongate.
Third-Instar Larva: Mouthhooks not joined to-
gether basally, each mouthhook spatulate and
dentate marginally; anterior spiracles with 2-8
marginal papillae; posterior spiracles borne dis-
tally on bifid, rectractile respiratory tube, tube
one-third to one-sixth total body length; spira-
cular caps each bearing 4 spiracular openings (or
series of openings), openings slitlike, oval, each
bordered basally by hydrofuge interspiracular
process; segments 5-12 with ventral prolegs bear-
ing crochet-like spines in well-defined rows; dorsal
patterns composed of flattened spines usually pre-
sent; if prolegs and dorsal patterns absent, then
spiracular openings subdivided and spiracular
caps elongate.
DISCUSSION.?Larvae of most Ephydrini are
easily recognized by their elongate respiratory
tube, ventral prolegs, and dorsal spine patterns.
The larvae of Dimecoenia are exceptional but can
be distinguished by the shape of their mouth
parts, the unique structure of the posterior spira-
cles, and their habitat distribution (salt marshes).
The monophyly of this tribe is well established,
being based on the following synapotypies:
1. Setal vestiture of prosternum: in members of this tribe, the
prosternum is setose, especially ventrally and posteriorly
around the coxal cavities and usually more extensively. The
generalized condition in the family is for the prosternum to
be bare of setae.
2. Hind coxal strap: the hind coxa has a strap that extends
around the posterior side. This strap bears four or five setae
in members of Ephydrini. Elsewhere in the family it is bare.
3. Pulvilli: with few exceptions in the family, the pulvilli
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
are evident as conspicuous pads beneath the tarsal claws. In
members of Ephydrini, however, the pulvilli are either ru-
dimentary or are lacking entirely.
4. Tarsal claws: the tarsal claws are shallowly curved and
are usually elongate in members of Ephydrini. The gener-
alized condition is for claws to be conspicuously curved and
short.
5. Larval prolegs: with the exception of a secondary loss in
Dimecoenia, larvae of Ephydrini have prominent, ventral
prolegs that bear crochets. These structures are an adapta-
tion to the algal-mat habitat of the immatures of these flies
and assist in grasping the substrate. The secondary loss of
prolegs in larvae of Dimecoenia apparently occurred as the
latter shifted back to a mud-shoreline habitat. Larvae of
Dimecoenia have creeping welts, similar to those of other mud-
inhabiting Ephydridae (Mathis and Simpson, 1981).
6. Habitat of immatures: the generalized habitat for the
subfamily Ephydrinae is probably shoreline mud. This is the
habitat of most species of Scatellini and of the subfamily
Parydrinae; however, members of Ephydrini have adapted
to algal mats on the surface of both lentic and lotic water
systems.
Although Ephydrini is undoubtedly monophy-
letic, its companion tribe Scatellini is not. Ephy-
drini is but one of several monophyletic lineages
arising from the ancestral lineage that now com-
prises the concept of Scatellini (Mathis, 1979c,
1980). The actual sister group of Ephydrini, sensu
stricto, probably gave rise to the Paracoenia-Calo-
coenia group of genera. This latter lineage plus
Ephydrini, as here delimited, is characterized by
the following character states (some have become
modified secondarily):
7. Number of dorsocentral bristles: although other genera of
the subfamily Ephydrinae sometimes have five pairs of
dorsocentral bristles (e.g., Nottocoenia Mathis and Austrocoenia
Wirth), the anterior pair (or pairs) is weakly developed. Only
in members of Ephydrini are there five well-developed pairs
(the anterior pair is presutural; specimens of Cirrula gigantea
have the anterior four pairs of dorsocentral bristles weakly
developed, a condition I interpret to be secondary).
8. Development of intrapostalar bristle: in most species of the
family, the intrapostalar bristle is either lacking or is very
much reduced, less than one-half the length of the postalar
bristle. In members of this lineage, the intrapostalar bristle
is frequently as long.
9. Setal vestiture of propleuron: throughout most of the family
this pleural region is bare of setae (although frequently it is
thinly to densely tomentose). In members of this lineage,
there are numerous setulae that are generally conspicuously
evident.
A more detailed account of the tribes and
genera of Ephydrinae will be forthcoming in a
generic revision that is now in progress.
Genus Setacera Cresson
Setacera Cresson, 1930:116 [type-species: Ephydrapacifica Cres-
son, by original designation); 1935:346-349 |new species
for Nearctic Region].?Loew, 1860:35 37 [review of pa-
laearctic species, as Ephydra (in part)].?Schiner, 1863:261
[review of Austrian species].?Becker, 1896:217, 219 [re-
view of palaearctic species, as Ephydra (in part)]; 1905:209
[catalog of palaearctic species, as Ephydra (in part)];
1926:75 [review of palaearctic species, as Ephydra
(in part)].?Seguy, 1934:434, 435 |review of French spe-
cies, as Ephydra (in part)].?Wirth and Stone, 1956:472
[review of Californian species].?Siurtevant and Wheeler,
1954:201-204 [review of nearctic species].?Collin,
1963:147 149 [review of British species].?Wirth,
1965:754, 755 [catalog of nearctic species]; 1968:24 [cata-
log of neotropical species].?Miyagi, 1966:138 140;
1977:83, 84 [review of Japanese species].?Cole, 1969:402
[synopsis of Western North American species].?Nart-
schuk, 1970:387 |review of European species in Russia, as
Ephydra (in part)].?Dahl, 1974:186 [notes on Scandina-
vian species].?Papp, 1975:107-109 [review of Hungarian
species].?Cogan and Wirth, 1977:338 [catalog of oriental
species].?Cogan, 1980:668 [catalog of afrotropical spe-
DIAGNOSIS.?Cruciate intrafrontal bristles lack-
ing or weakly developed, not conspicuous; fronto-
orbital bristles 2 pairs, lateroclinate; 3rd antennal
segment with prominent bristle inserted laterally,
just below aristal insertion; arista with subpectin-
ate to pectinate branching rays along dorsum
from between basal one-half to two-thirds of ar-
istal length; fronto-orbits shiny with metallic lus-
ter, concolorous with mesofrons; dorsum of inter-
foveal facial carina nearly flat, sloping very grad-
ually; anterior facial ridge projecting markedly
forward in many species, from which anterior
surface of face extends ventrally at nearly right
angle to oral margin, face receding at obtuse
angle to oral margin in other species; genal bristle
short, usually less than one-half length of arista;
dorsocentral bristles 5 pairs (1 + 4); posthumeral
bristles 1 pair, development variable, but gener-
ally well developed; structures of male terminalia
symmetrical, complicated in most species by ad-
NUMBER 350
dition of several secondary processes and prongs,
especially gonite and hypandrium; epandrium
elongate; surstyli well developed, generally fused
basomedially, frequently with ventrally project-
ing lateral arms, see species group and species
descriptions for additional details; female termin-
alia quite variable, see species group and species
descriptions; female ventral receptacle with op-
erculum as high as wide, broadly rounded dor-
sally, wider than high, somewhat angulate, sub-
trapezoidal, or almost entirely lacking.
DESCRIPTION.?Moderately small to large shore
flies, length 2.46 to 5.67 mm; generally dark
colored but with considerable grayish tomentose
vestiture, subshiny to shiny dorsally, becoming
more subdued, usually grayish or slightly oliva-
ceous laterally and ventrally.
Head: Wider than high from cephalic view;
slightly longer than high in profile. Frons subrec-
tangular to trapezoidal, wider than long; meso-
frons subrectangular, slightly wider than long,
shiny with metallic luster, setose, especially ante-
riorly, sculpturing inconspicuous posteriorly, be-
coming granulose anteriorly; ocellar triangle
equilateral or isosceles, very slightly raised in
relief from mesofrons, finely tomentose, dull, fre-
quently concolorous with dull portion of para-
frons; parafrons dull, finely tomentose, generally
dark colored except for shiny fronto-orbits, which
are concolorous with mesofrons. Ocellar bristles
1 pair, large, proclinate, divergent, inserted be-
hind level of median ocellus; postocellar bristles
small, 1-2 pairs, inserted directly behind ocellar
bristles; fronto-orbital bristles 2 pairs, large, with
1-2 smaller pairs of setae inserted alternately with
larger bristles; vertical bristles 2 pairs, large, inner
bristle inserted anteromediad of alignment of
outer bristle and fronto-orbital bristles; spacing
between each fronto-orbital and between inner
vertical bristle about equidistant; postocular setae
approximately subequal to each other, all small.
Antenna mostly black, unicolorous, dull, tomen-
tose to micropubescent; 3rd antennal segment as
long as or longer than combined length of first 2
segments; rounded apically, bearing a prominent
bristle laterally, inserted just below arista; arista
thickened basally, gradually tapered to apex, with
several subpectinate to pectinate dorsal branches
along basal one-half to two-thirds. Face broadly
arched, protruding, interfoveal facial carina
prominent, dorsum of interfoveal carina subshiny
to shiny, sparsely tomentose, nearly concolorous
with mesofrons, surface nearly flat but inclined
anteroventrally; antennal foveae well impressed,
shiny, like mesofrons, or dull, tomentose; face
below transverse facial ridge extended ventrad
perpendicular to oral margin or receding at ob-
tuse angle; facial setae along margin more prom-
inent, especially along dorsal slope; vestiture of
face densely tomentose, grayish to almost silvery
white, coloration along ridge frequently brownish
to golden brown; oral margin slightly emarginate
at middle from cephalic view. Eye usually slightly
wider than high, suboval, oriented at slight to
obvious oblique angle to oral margin; gena wide,
eye-to-cheek ratio at least 1 : 0.34; genal bristles
usually prominent, 1 pair; remaining postocular
area uniformly setose. Oral opening large, gap-
ing; mouth parts large, well sclerotized, premen-
tum bowl-shaped, narrow, dull, finely tomentose,
concolorous with posterior portion of gena,
sparsely setose; maxillary palp 2-3 times longer
than wide, setose.
Thorax: Dark colored, dorsum subshiny or
shiny, becoming darker and shinier posteriorly;
pleural areas dull, tomentose, becoming progres-
sively lighter toward venter. Chaetotaxy as fol-
lows: no prominent acrostichal bristles, small se-
tae, when present, arranged in 2 rows; dorsocen-
tral bristles 5 pairs (1 + 4), large, posteromost
pair displaced laterally from alignment of others;
posthumeral bristles generally well developed, 1
pair, sometimes reduced or lacking; presutural
bristles 1 pair; supra-alar bristles 1 pair; postalar
bristles 1 pair; intrapostalar bristle 1 pair, at least
one-half length of postalar bristle; scutellar bris-
tles 2 pairs, lateral, apical pair much longer;
humeral bristles usually 1-2 pairs, usually more
weakly developed; notopleural bristles 2 pairs;
mesopleural bristles 1 pair plus 2-3 pairs of me-
dium-sized setae around larger mesopleural bris-
tle and several uniform, scattered, smaller setae;
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
sternopleural bristles 1 pair, inserted toward pos-
terodorsal corner, several smaller setae mainly
around larger bristle; other pleural sclerites bare,
except for patches of dense tomentosity, as on
supraspiracular convexity. Halter pale, yellowish.
Legs mostly dark, concolorous with pleural areas
but with some lighter areas toward apices; males
often with tufts of hairs on coxae or toward apices
of tibiae; basitarsus almost as long as combined
length of remaining tarsomeres; tarsal claws rel-
atively long, with very little curvature; pulvilli
reduced or lacking. Wing hyaline to very lightly
infumated; costal vein extending to M1+2.
Abdomen: Generally unicolorous, subshiny to
shiny, not so dark as posterior portion of meso-
notum or scutellum. Female with 6-7 visible
segments; male with 5, often 5th triangular, some-
times longer than wide, or trapezoidal, truncate
apically. Male terminalia symmetrical; cereal
cavity within epandrium near dorsum; surstyli
fused medially, broadly attached at ventral mar-
gin of epandrium; aedeagus bulbous; gonite
highly modified, considerably reduced or bearing
well-sclerotized, acutely pointed processes similar
to those of hypandrium; hypandrium generally
with lateral, paired symmetrical processes, degree
of sclerotization variable, usually weak towards
middle. Female ventral receptacle longer than
wide; operculum as high as wide, rounded dor-
sally; extending process more or less j-shaped.
GEOGRAPHIC DISTRIBUTION.?Few shore fly gen-
era are as widespread as Setacera, and in the tribe
Ephydrini, Setacera is by far the most widely
distributed, with species occurring in all major
faunal realms.
NATURAL HISTORY.?The immature stages of
Setacera closely resemble others of the tribe Ephy-
drini, and Johannsen (1935) considered them to
be the most highly specialized of the family. Like
other ephydrines, larvae of Setacera have long,
terminal respiratory tubes and eight pairs of
short, conical, abdominal prolegs that bear cro-
chets. The last pair of prolegs is the largest, and
the crochets are opposable to those of the other
prolegs. Johannsen (1935) figured the cephalo-
pharyngeal skeleton of S. needhami Johannsen, a
species described inadvertently from the imma-
ture stages (Cresson, 1935).
Unlike most ephydrines, members of Setacera
primarily inhabit fresh-water environments, al-
though there are occasional records of species
from salty habitats, and S. pacifica (Cresson) is
regularly found in association with alkaline wa-
ters (Foote, 1982). Johannsen (1935) reportedly
reared a specimen of S. atrovirens (Loew) from a
puparium collected in a brine pool near Ithaca,
New York, and Karl (1930) and Beyer (1939)
stated that S. micans is halophilous. Subsequent
authors, Frey (1948) and Dahl (1959), however,
have discredited these earlier observations and
have found S. micans to be a fresh-water species.
I suspect that the earlier authors' observations
were based on misidentifications (see species
treatment of S. micans, page 15).
Most species seem to prefer lentic aquatic sys-
tems, especially where a layer of floating algae
has accumulated on the water's surface. This is
the typical habitat of most species of Ephydrini,
and their crochet-bearing prolegs are apparently
an adaptation to this habitat, allowing movement
through and attachment to the algae.
PHYLOGENY.?The species now included in Se-
tacera were previously placed in the genus Ephydra
Fallen, and some recent authors still prefer the
precedent of Ephydra, with Setacera as an included
subgenus (Soika, 1956; Dahl, 1959). Setacera is
indeed closely related to Ephydra, as evidenced by
the similarity of adults and immatures of both
genera. Setacera, however, can be consistently dis-
tinguished in both sexes from all other genera of
Ephydrini, and its monophyly corroborated by
the following synapotypies (numbers accompany-
ing discussion of characters correspond with those
on the cladogram):
10. 3rd antennal segment seta: aside from the arista, there are
usually no other large structures emanating from the 3rd
antennal segment. Specimens of Setacera, however, have a
large seta inserted just below the aristal insertion on the
lateral surface.
11. Vertico-orbits: within the tribe Ephydrini, the vertico-
orbits are generally either shiny or densely tomentose and
grayish, appearing dull. This area, in specimens of Setacera,
is uniquely invested with a dense patch of tomentum that
NUMBER 350
appears velvety. Velvety areas occur elsewhere in a few
species of the tribe (parafrons in Cirrula gigantea Cresson;
frons and orbits in Ephydra auripes Aldrich) but not in the
specific area as described for Setacera.
12. Genal seta: this seta is usually very prominent, arising
below the eye. Although this seta is still larger than surround-
ing ones in specimens of Setacera, its comparative size is
smaller, and for convenience, I have compared it with the
length of the arista.
13. Cruciate intrafrontal bristles: although some species of the
tribe Ephydrini do not have these bristles, most genera have
at least a few species in which they occur. Consequently, my
interpretation of the general ground plan of the tribe is for
their presence, and their lack in Setacera is apparently unique.
14. Prescutellar acrostichal setae: as with the preceding char-
acters, these setae are generally present in Ephydrini. I know
of no specimens of Setacera, however, where they are present,
and I interpret this apparent loss to be synapotypic.
I have arranged the genus into six species
groups and have attempted to show cladistic
relationships between these by forming nested
clusters as sister lineages could be identified (Fig-
ure 2). The genus is arranged into two major
clusters, comprising either two or four species
groups. The character evidence to corroborate
the monophyly of the species groups and their
intrageneric relationships are as follows.
The first major cluster of lineages includes the
micans and breviventris species groups. The mono-
phyly of this stem lineage and of its component
species groups is characterized by the following
characters:
15. Length of 5th abdominal tergum of male: in most ephy-
drines, this tergum is as long as either the 3rd or the 4th
terga and frequently almost as long as wide, occasionally
longer. In males of these two species groups this tergum is
conspicuously shorter than either the 3rd or 4th, a synapo-
typic character for this lineage.
The micans group is monophyletic, with char-
acter evidence as follows:
17. Configuration of aedeagus: the aedeagus among most
ephydrines is nearly as wide as long, and its apex is broadly
rounded. In males of the micans group the aedeagus is three
to four times longer than wide, and its apex is acutely
pointed. I interpret this to be an synapotypic character,
being unique to this species group.
18. Configuration of the female ventral receptacle: generally the
operculum of this structure is nearly as high as the extending
process, sometimes more so, and its width is subequal to its
FIGURE 2.?Hypothetical phylogeny of species groups of
Setacera.
height. In females of the micans group the operculum is
relatively small, both its width and height, especially as
compared to the size of the extending process (see illustra-
tions of included species).
The sister group of the micans group is the
breviventris group. The monophyly of the latter is
established as follows:
19. Development of posthumeral bristle: one of the apotypic
characters of the tribe Ephydrini is the well-developed
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
posthumeral bristle. For most ephydrines this bristle is as
strong as the presutural bristle. In members of the breviventris
group, however, the posthumeral bristle is comparatively
weak and is considerably smaller than the presutural bristle,
if it is existent at all.
The second major lineage comprises the re-
maining species groups and is divided into two
sublineages, each of which gives rise to two species
groups. The monophyly of the second major lin-
eage is established as follows:
16. Vestiture of 3rd and 4th sterna of male: typically these
sterna bear a few scattered, generally small setae. In males
of this lineage, however, the unique condition exists of a
dense patch of large setae, usually more conspicuous toward
the posterior portion of the sternum. I interpret this character
to be apotypic.
The monophyly of the first stem lineage, which
gives rise to the aurata and trina species groups, is
established as follows:
20. Distance between cerci and 9th sternal bristles of female: the
distance between the ventral margin of the cerci and the
base of the 9th sternal bristles in females is generally not
greater than the height of the cerci. In females of this lineage,
the 9th sternal bristles are situated farther ventrad, making
the distance between them and the cerci conspicuously
longer than the cereal height. This character is unique to
this lineage.
21. Length of 8th stemites of female: generally, each sternite
is five or more times longer than wide, but in females of this
lineage, the length is at most three to four times its width,
an apotypic character.
22. Length of 8th tergites of female: this character is somewhat
correlated with character no. 20. The 8th tergites in females
of this lineage are very long and partially account for the
ventral position of the 9th stemites. This is best seen by
comparing the figures of the female terminalia accompany-
ing the species treatments. The longer 8th tergites are an
apotypic character.
The aurata and trina groups are the two species
groups of this sublineage. The monophyly of the
aurata group is established as follows:
24. Configuration of epandrium: although it is not uncommon
for the epandrium to have appendages of various sorts, this
is the only lineage to have a bluntly rounded, parallel-sided,
posterolateral process arising from each side. This unique
condition is interpreted to be apotypic.
The trina species group is characterized, and its
monophyly is established, as follows:
25. Configuration of the male gonite: this is a difficult character
to assess, as the generalized condition is not known. The
configuration of the gonite in males of this group is unique,
however, and for the present I am interpreting it as an
apotypic character.
26. Configuration of epandrium (see no. 24): although the
shape of the epandrium, particularly of the ventral surstyli,
differs quite markedly throughout the genus, that of this
group is uniquely similar. The ventrolateral angles are ex-
planate and slightly recurved (see illustrations accompany-
ing species treatments). This character is apparently apo-
typic for this group.
The sister group of the stem lineage of the
aurata and trina groups is the stem lineage that
gives rise to the aldrichi and pacifica groups. This
lineage is characterized and its monophyly is
corroborated by the following:
23. Vestiture of antennal fovea: for most ephydrines, the
antennal foveae are invested with tomentosity and give the
appearance of being relatively dull as compared with the
dorsum of the interfoveal carina. The latter and the meso-
frons are usually bare of tomentosity and have a metallic
luster. The antennal foveae of these groups are shiny, with
a metallic luster and color similar to the interfoveal carina.
As this character is unique to members of these groups, I
interpret it to be apotypic.
The aldrichi group is the sister group of the
pacifica group, and it is characterized, and its
monophyly established, by:
27. Configuration of aedeagus (see no. 17): as before, the
aedeagus is typically broadly rounded apically and almost
as wide as long. Males of the aldrichi group have a somewhat
pointed aedeagus that I interpret to be apotypic.
28. Configuration of epandrium (see nos. 24, 26): males of the
aldrichi group have an anteroventral, digitiform process,
apparently a unique condition, and one that I interpret to
be apotypic.
The monophyly of the pacifica group is estab-
lished by:
29. Configuration of vertico-orbits (see no. 11): in Setacera this
band is more or less broad and usually has a subanterior
swelling, but in members of the pacifica group this band is
very narrow and is sometimes difficult to detect. The nar-
rowed aspect of this character is interpreted to apotypic.
30. Configuration of female ventral receptacle (see no. 18): for
most ephydrines, the operculum is typically wider than high.
For females of the pacifica group, however, the height is
subequal to its width, an apotypic character.
In addition to the characters just enumerated
NUMBER 350 9
to distinguish Setacera and the included species secondarily sexually dimorphic features. Males of
groups, Ephydra is characterized by the presence these species bear prominent hair-tufts of varying
of three lateroclinate, fronto-orbital bristles. Seta- length at the tibial apices and often on the coxae.
cera has only two. The extent and length of the tufts, or their ab-
DISCUSSION.?Some members of Setacera have sence, are excellent species-level characters.
Key to New World Species of Setacera
1. Arista with dorsally branching rays at most slightly longer than aristal
width at base 2
Arista with dorsally branching rays conspicuously longer than aristal width
at base, nearly twice 4
2. Antennal foveae concolorous with dorsum of interfoveal carina, shiny;
vertico-orbits mostly shiny, concolorous with fronto-orbits; facial prom-
inence in profile angulate, abruptly rounded; 5th tergum of male longer
than 4th, about as wide at base as long 12. S.jamesi, new species
Antennal fovea pollinose, whitish gray, contrasting with dorsum of inter-
foveal carina; vertico-orbits appearing velvety, contrasting with shiny
fronto-orbits; facial prominence in profile gently rounded, obtuse; 5th
tergum of male considerably shorter than 4th, much wider at base than
long 3
3. Surstyli, in posterior view, more widely separated medially along apical
half; lateral margins curved, not conspicuously angulate; with medial,
shallowly pointed process between surstyli 1. S. atrovirens (Loew)
Surstyli, in posterior view, closely opposed medially along apical half,
almost touching; lateral margins angulate; lacking medial process be-
tween surstyli 2. S. micans (Haliday)
4. Facial prominence in profile with dorsal slope longer than height of lower
portion of face; 5th tergum of male broadly truncate, width of posterior
margin more than half basal width 10. S. aldrichi Cresson
Facial prominence in profile with dorsal slope short, less than height of
lower portion of face; 5th tergum of male rounded or truncate, if
truncate, width of posterior margin not over half basal width 5
5. Supraspiracular convexity with a papilla-like prominence toward antero-
dorsal margin; epandrium with a medial longitudinal sulcus
14. S. pacifica (Cresson)
Supraspiracular convexity evenly rounded, lacking any distinct promi-
nence; epandrium at most with only faint indication of a sulcus 6
6. Midtibia of male lacking long hairs along posterodorsal surface subequal
in length to tibial width near apex; femaje with 6 visible abdominal
segments from dorsal view 15. S. pilicornis (Coquillett)
Males with a "patch of long hairs along posterodorsal surface subequal in
length to tibial width near apex of midtibia; females with 7 visible
abdominal terga from dorsal view 7
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
7. Midfemur of male lacking posteroventral row of long setae equal in length
to greatest width of femur; 3rd and 4th abdominal sterna of male with
dense patch of well-developed, conspicuous setae posteromedially; cerci
of female not unusually prominent, height greater than length
11. S. durani Cresson
Midfemur of male with posteroventral row of 4-7 long setae near base
subequal in length to greatest width of femur; 3rd and 4th abdominal
sterna of male lacking dense patch of well-developed setae postero-
medially; cerci of female conspicuously prominent, length equal to
height 8
8. Fused surstyli distinctly wider at basal one-third; lateral margins of basal
two-thirds of surstyli broadly and more or less evenly rounded (western
North America) 13. S. needhami Johannsen
Fused surstyli as wide apically, or nearly so; lateral margins of basal two-
thirds of surstyli somewhat angulate (Peru)
16. S. trichoscelis, new species
Key to Old World Species of Setacera
1. Male 5th abdominal tergum shorter than 3rd or 4th tergum; male 3rd and
4th sternum without dense patch of stout setae toward posterior margin;
base from which spinelike setae of female ovipositor arise not farther
from cerci than length of cerci 2
Male 5th abdominal tergum as long as or longer than 4th tergum; male
3rd and 4th sternum with dense patch of stout setae toward posterior
margin; base from which spinelike setae of female ovipositor arise more
distant from cerci than length of cerci 5
2. Posthumeral bristle strong, larger than largest humeral bristle and generally
subequal to posterior notopleural bristle; anterior margin of face in
profile shallowly arched and forming a more or less right angle with oral
margin; front tibia mostly dark colored; larger species (northern
Europe) 2. S. micans (Haliday)
Posthumeral bristle weaker or absent, if present not stronger than largest
humeral bristle; anterior margin of face in profile straight and forming
an obtuse angle with oral margin; front tibia with basal one-half
yellowish in most specimens; smaller species 3
3. Surstyli acutely corniform apically, median surface not conspicuously setose
(Old World, except for northern Europe) . . . . 3. S. breviventris (Loew)
Surstyli bluntly rounded apically, not corniform, median surface more or
less densely setose 4
4. Width of gap separating surstyli about equal to length of surstyli (Mada-
gascar, South Africa) 4. S. multicolor (Soika)
Width of gap separating surstyli less than length of surstyli (Japan,
Korea) 5. S. viridis Miyagi
5. Surstylus in profile with slender, parallel-sided, apically truncate, lateral
process that projects anteroventrally; gonite broad throughout most of
its length, lacking secondary process (northern Europe)
6. S. aurata (Stenhammar)
NUMBER 3f)0 11
Surstylus in profile with posteriorly curved, ventrolateral process that is
subapically enlarged; gonite conspicuously narrowed apically and with
secondary process 6
6. Gonite with secondary process triangular, broad basally, tapered at more
or less constant angle to apex 9. S. trina Collin
Gonite with secondary process more bandlike than triangular, taper at base
gradual, becoming more abrupt toward apex 7
7. Surstyli in posterior view with acutely pointed process between; surstylus
with recurved, lateral apex distinctly explanate, rounded; gonite with
secondary process broader, about twice as long as wide
8. S. meneghinii Canzoneri
Surstyli in posterior view with truncate process between; recurved, surstylus
with lateral apex less flared and more angulate; secondary gonite with
process narrower, width 3 times length ... 7. S. freidbergi, new species
The micans Group
SPECIES INCLUDED.?Setacera atrovirens (Loew);
S. micans (Haliday).
DIAGNOSIS.?Specimens of the micans group
may be distinguished by the following combina-
tion of characters: antennal foveae densely to-
mentose, contrasting distinctly with subshiny to
shiny dorsum of interfoveal carina; arista with
longest dorsally branching rays about equal to
aristal width at base; vertico-orbits with velvety
tomentose band moderately wide, widest suban-
teriorly at vertex; posthumeral bristle well devel-
oped, subequal to posterior notopleural bristle,
distance between it and presutural bristle nearly
equal to that between notopleural bristles; su-
praspiracular convexity evenly convex; legs and
coxae of males lacking dense patches or rows of
setae; fore- and midtibiae mostly dark colored,
concolorous with femora, only tibial-femoral ar-
ticulation pale; 5th tergum of male rounded ap-
ically, apical width less than length, length less
than that of 3rd or 4th tergum, anteroventral
corner lacking process; 3rd and 4th sterna of male
lacking dense patch of setae toward posterior
margin.
Male Terminalia: Epandrium lacking median
sulcus, greatest width subdorsally in posterior
view, dorsum rounded if complete, frequently
weak or lacking above cereal cavity; surstyli
broadly fused basally, apical one-half separated,
apices tapered more in posterior view, evenly so
in lateral view, more setose submedially in pos-
terior view; aedeagus 3-4 times longer than wide,
tapered gradually to apex, apex acutely pointed,
relatively simple; gonite lacking processes or
prongs, simple, posterodorsal portion angulate,
angle of about 90? in lateral view; hypandrium
poorly sclerotized, formed by 2 symmetrical
plates, more or less fused anteriorly.
Female Terminalia: 7th tergum complete; 8th
tergites 2-3 times higher than wide, widest at
venter in lateral view, venter broadly rounded,
tapered dorsally to dorsal point, anterior margin
more or less straight, posterior margin rounded
and tapered; 8th sternites moderately elongate,
5-6 times longer than wide; 9th sternum well
sclerotized, more or less quadrate although irreg-
ular, compact, lacking lateral or dorsal exten-
tions; 9th sternal bristles approximate but not
aligned, more or less clumped; distance between
bristle insertions and venter of cerci at most equal
to height of cerci, usually less; cerci higher than
wide, posterior margin rounded in lateral view;
female ventral receptacle with operculum greatly
reduced, less than width of extending process,
papilla-like; extending process broadly j-shaped,
lacking marked distinction between cervix and
corpus.
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
DISTRIBUTION.?Holarctic; northern North
America above 38? north latitude and generally
east of the Rocky Mountains and European Pa-
laearctic Region north of 50? north latitude.
DISCUSSION.?Although this species group is
easily distinguished, the included species are pres-
ently separable only by examining the male ter-
minalia. In Europe there is no problem, as there
is only one species occurring there, but in North
America, females are identified tentatively either
from being associated with identified males and/
or from distributional data.
1. Setacera atrovirens (Loew)
Figures 3-12
Ephydm atrovirens Loew, 1862:169.?Osten Sacken, 1878:203
[nearctic catalog].?Aldrich, 1905:629 [nearctic catalog].
Setacera atrovirens.?Cresson, 1930:116 [listing].?Johannsen,
1935:53 [description of immature stages].?Wirth,
1965:755 [nearctic catalog, in part].
Setacera atroviems [sic].?Sturtevant and Wheeler, 1954:202,
203 [key and locality data].
DESCRIPTION.?Medium-sized to moderately
large shore flies, length 3.5 to 4.7 mm; dorsum
dark olivaceous brown to green, becoming duller,
grayer ventrally.
Head (Figures 3-5): Head width-to-height ra-
tio averaging 1 : 0.68; frons width-to-length ratio
averaging 1 : 0.48; mesofrons, fronto-orbits, and
dorsum of interfoveal carina deep bluish green to
mostly bluish, dorsum of interfoveal carina fre-
quently sparsely tomentose; dorsal surface of face
shorter than height of anterior surface; angle
formed by anterior surface and dorsal surface
obviously obtuse, about 135?; anterior surface of
face usually lightly yellowish brown, especially
dorsally, sometimes becoming almost whitish ven-
trally. Eye height-to-width ratio averaging
1 : 0.95; eye-to-cheek ratio averaging 1 : 0.40.
Thorax: Costal vein ratio averaging 1 : 0.29;
Mi+2 vein ratio averaging 1 : 0.79.
Abdomen: Male terminalia (Figures 6-9) with
epandrium usually poorly sclerotized dorsally if
at all evident; surstyli with lateral margin more
or less evenly rounded, not angulate, median
surfaces separated, distance approximately equal
to surstylar width in posterior view, with pointed
projection medially between surstyli toward base.
Female terminalia (Figures 10, 11) as in species-
group description.
TYPE MATERIAL.?The male lectotype, herein
designated, is labeled "Mittel St. [mid-Atlantic
states of the United States]/Loew Collection]./
atrovirens/50/Type 11179 [red]." The lectotype
is in the Museum of Comparative Zoology, Har-
vard University, Cambridge, Massachusetts, type
number 11179.
OTHER SPECIMENS EXAMINED.?CANADA. Nova
Scotia: Petite Riviere, 2 Jul 1935, J. McDunnough
(1$; CNC). Ontario: Black Sturgeon Lake, 28 Jun
1962 (1$; CNC); Grand Bend, 8 Jul 1939, G. E.
Shewell (1(5; CNC); Guelph, 1 Nov 1955, J. R.
Vockeroth (1$; CNC); Kingston, 10 Jul 1938, A.
FIGURES 3-5.?Setacera atrovirens: 3, right antenna, lateral aspect; 4, right vertex and vertico-
orbits of head, dorsal aspect; 5, right vertico-orbit showing velvety tomentosity, dorsal aspect.
NUMBER 350 13
FIGURES 6-11.?Setacera atrovirens: 6, male terminalia, posterior aspect; 7, same, lateral aspect; 8,
surstyli, posterior aspect; 9, internal male genitalia, lateral aspect; 10, female terminalia, lateral
aspect; 11, female ventral receptacle, lateral aspect.
L. Melander (1$; USNM); Ottawa, 25 Apr-30
Jul, 1922-1965, A. R. Brooks, C. H. Curran,
J.E.H. Martin (15c?, 49$; ANSP, CNC); Ottawa,
Mer Bleue, 26 May 1923, C.H. Curran (19;
ANSP). Quebec: Abbotsford, 1 Jul-26 Sep, 1935-
1936, G. E. Shewell (29; CNC); Old Chelsea, 9
Oct 1955, J. R. Vockeroth (1<5; CNC). UNITED
STATES. Delaware: Sussex Co., Rehoboth, 25 Jun
1939, A. L. Melander (19; USNM). District of
Columbia: A. L. Melander (Id; USNM). Illinois:
Mason Co., Havana, river shore, 16 Nov 1913
(19; ANSP). McHenry Co., 21 Aug 1903, 1927,
A. L. Melander (4c?, 29; ANSP, USNM). Indiana:
LaGrange Co., Pigeon River Fish and Wildlife
Area, 30 Jul 1977, M. Minno (1(5, 29; USNM).
Iowa: Boone Co., Boone, 1.5 mi W, 4 Aug 1960,
D. L. Deonier (19; DLD); Ledges State Park, 30
Jul 1971, R. M. Miller (Ic5; USNM); Little Wall
Lake, 14 Jul-22 Sep, 1960-1962, D. L. Deonier
(14<5, 139; DLD). Greene Co., Spring Lake Rec-
reational Reserve, 19 Sep 1961, D. L. Deonier
(59; DLD). Hamilton Co., Goose Lake, 2 Jul
1961, D. L. Deonier (Ic5; DLD). Louisa Co., Lake
Odessa, 9 Aug 1960, D. L. Deonier (1(5, 49; DLD).
Winneshiek Co., Siewers Spring State Park near
Decorah, 9 Sep 1961, D. L. Deonier (Id, 19;
DLD). Kansas: Douglas Co., 900 ft (19; KU).
Maine: Penobscott Co., Old Town, Lake Pushaw,
1 Aug 1966, W. W. Wrirth (Ic5; USNM). Michigan:
Cheboygan Co., 10 Aug 1932, J. D. Nottingham
(1(5; KU). Ingham Co., East Lansing, 27 Jun
1936, C. Sabrosky (19; ANSP). Livingston Co.,
E. S. George Reserve, 24 Jul 1943, G. C. Steyskal
(Ic5, 19; USNM). Minnesota: Clearwater Co.,
Itasca State Park, 15 Jun-3 Aug 1937, A. R. Barr,
D. Keith, K. C. Kim, M. E. Rueger (13(5, 439;
UMIN). Itasca Co., Grand Rapids, 30 Jun 1966,
B. C. Ahlm (1(5, 89; UMIN). La Sueur Co., 28
Aug 1923, W. E. Hoffmann (19; UMIN). Polk
Co., Crookston, 27 Jun 1937, D. G. Denning (19;
UMIN). Ramsey Co., St. Paul, 22 April 1922, W.
E. Hoffmann (19; UMIN). Waseca Co., Waseca,
3 Jul-11 Sep, 1941-1967, R. E. Carlson, H. T.
Peters (Ic5, 29; UMIN). Montana: Lake Co., Pol-
son, 2.3 mi E, 24 Jul 1973, B. A. Foote (2c5, 19;
KSU). Nebraska: Cherry Co., Pelican Lake, 2 Jun
1969, W. W. Wirth (29; USNM). New York: Jef-
ferson Co., Picton Island, Clayton, 18 Aug 1960,
B. Heineman (19; AMNH). Tompkins Co., Key-
den Lake, 8 Aug 1961, D. L. Deonier (4c5, 29;
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
DLD); Ithaca, 4 Sep 1973 (29; CU); McLean
Reserve, 29 Mar 1962, L. V. Knutson (1$; CU).
North Dakota: Burleigh Co., Long Lake, 4 Jun
1969, W. W. Wirth (2$; USNM). Ramsey Co.,
Devils Lake, 5 Jun 1969, W. W. Wirth (1$;
USNM). Ohio: Portage Co., Kent, 9 Apr 1969, R.
Miller (16; KSU); Kent, 1 mi E, 11 Sep 1972, B.
A. Foote (406\ KSU); Kent, 4.5 mi E, 1 May
1969, W. Eastin (1$; KSU); Ravenna, 15 mi E,
28 Sep-11 Oct 1971, B. A. Foote (16\ 39; KSU).
Wayne Co., Rittman, 0.5 mi S, 18 Sep 1969, B.
A. Foote (1(5, 2$; KSU). South Dakota: Hand Co.,
Burdette, 20 Jul 1937, R. H. Beamer (19; KU).
Wisconsin: Dane Co., 20 Jul 1900 (29; USNM).
DISTRIBUTION (Figure 12.)?East central North
America, primarily around the Great Lakes, be-
tween 55? and 105? west longitude and 38? and
49? north latitude.
NATURAL HISTORY.?Foote (1979; 1982) has
recently published detailed studies on the life
history and immature stages of S. atrovirens. In
particular, he reported the feeding preferences of
this species, using both field and laboratory stud-
ies. Most of the information to follow is para-
phrased from his observations and analysis.
The life cycle of S. atrovirens takes approxi-
mately 25 days and is divided among the stadia
as follows:
Egg incubation
Larval period (3 instars)
Pupal period
Adult preoviposition period
2.3 days
10 days
7 days
6-9 days
The fly apparently overwinters as an adult,
probably in a state of reproductive diapause.
During the warm season, from May to September
in the northern states, four or more generations
are produced.
In the field, Foote discovered eggs in floating
mats of primarily blue-green algae. Along the
marshy shores of Flathead Lake, Montana, the
fly preferred growths of Nostoc, but near Kent,
Ohio, they were usually found in mats of Oscilla-
toria or a mixed mat of Anabaena and Spirogyra.
Foote's laboratory experiments showed that lar-
vae of this species would develop readily in algae
of the genera Anabaena, Cylindrospermum, Lyngbya,
Nostoc, and Navicula. In contrast, however, larvae
were apparently unable to use species oiAnacystic,
Gloeocapsa, and Chlorella. Interestingly, utilization
experiments between two species of Oscillatoria
clearly showed that larvae of S. atrovirens can
discriminate between the two algal species. Foote
found that 80% of the larvae reached the pupal
stage when fed 0. tenuis, but no larvae formed
FIGURE 12.?Distribution map of Setae era atrovirens.
NUMBER 3")() 15
puparia in cultures of 0. chalybea. From larval gut
dissections, Deonier (1972) found that the algae
ingested by larvae of S. atrovirens included about
one-half diatoms and one-half other algae.
In part, larval age also determined food pref-
erence. Newly hatched larvae appeared to prefer
small unicells, whereas older larvae utilized a
broader range of algal species, frequently tri-
chomes of blue-green algae. Larvae can feed com-
pletely submerged, but return to the water's sur-
face periodically for respiration.
Just prior to forming puparia, larvae left the
algal colonies and sought out stems or narrow
leaves of aquatic macrophytes, to which they
became attached using the last prolegs. Puparia
were frequently formed below the water's surface,
and submergence apparently had no effect on
subsequent development.
Foote observed several successful attacks on
Setacera larvae by larvae of water-scavenger bee-
tles (Hydrophilidae), which are probably the pri-
mary predator of the larval stages. Foote also
reared a chalcidoid parasitoid from a few field-
collected puparia.
REMARKS.?Females of this species are not pres-
ently separable from those of S. micans and are
identified only by direct association with a male
or by distributional data. Males are distinguish-
able by the shape and position of their terminalia,
particularly the surstyli.
2. Setacera micans (Haliday)
FIGURES 13-22
Ephydra micans Haliday, 1833:175.?Nartschuk, 1970:387
[key].
Setacera micans.?Cresson, 1930:116.?Collin, 1963:147 [lec-
totype designation, figure of male terminalia].?Dahl,
1974:186 [distribution, figure of male terminalia].?Papp,
1975:108 [distribution, figure of male terminalia].
Setacera atrovirens of North American authors [misidentifica-
tion in part].?Sturtevant and Wheeler, 1954:203 [re-
view].?Wirth, 1965:755 [nearctic catalog].
DESCRIPTION.?Medium-sized to moderately
large shore flies, length 3.9 to 4.5 mm; generally
dark olivaceous brown to green dorsally, pleural
areas and venter becoming lighter, duller grayer.
Head (Figure 13): Head width-to-height ratio
averaging 1 : 0.62; frons width-to-length ratio
averaging 1 : 0.48; mesofrons, fronto-orbits, and
dorsum of interfoveal carina deeply greenish blue
to mostly greenish; dorsal surface of face shorter
than anterior surface; angle formed by anterior
and dorsal surfaces obtuse, approximately 120?;
face mostly silvery whitish, sometimes dorsum of
anterior surface darker, lightly brownish to
golden. Eye height-to-width ratio averaging
1 :1 ; eye-to-cheek ratio averaging 1 : 0.42.
Thorax: Costal vein ratio averaging 1 : 0.29;
M1+2 vein ratio 1 : 0.88.
Abdomen: Male terminalia (Figures 14-17)
with epandrium usually complete dorsally
around cerci although very narrow; surstyli in
posterior view with lateral margins angulate to-
ward apices, median margins approximate, much
closer than surstylar width, lacking basal median
pointed projection. Female terminalia (Figures
18, 19) as in species-group description.
TYPE MATERIAL.?The male lectotype, desig-
nated by Collin (1963), is labeled "Holywood/
Lectotype." I have not examined the lectotype,
and the label data I have quoted were taken from
Collin (1963). The lectotype is presumably in
Haliday's collection, National Museum of Ire-
land, Dublin.
OTHER SPECIMENS EXAMINED.?ENGLAND. But-
ten, Issoy, 11 Jul 1908, Verrill-Collin collection
(1?; UMO). Surrey, Kew, 4 Aug 1868 (Id;
UMO). NETHERLANDS. Ankeveen, 1943, D. Piet
(1$; ITZA). Hilversum, 3 Mar 1912, Meijere (1$;
ITZA). SWEDEN. (1$; NRS). Ostergotland (Id, 39;
NRS). Stockholm: (4d, 59; NRS). Skane: (19; NRS).
Lilla Viken, Hoor, 3 Jul-14 Aug 1954, R. Dahl
(29; ZIL). Lomma N flytveg (Kivik), 24 Aug
1954, R. Dahl (Ic5, 29; ZIL). Fyleoset, Ystad, 17
Jun 1954, R. Dahl (19; ZIL). CANADA. Manitoba:
Aweme, 13 Jun-17 Oct, 1922-1927, N. Criddle,
H. A. Robertson, R. M. White (Id, 109; CNC);
Birtle, 21 Jul 1928, R. D. Bird (1<5; AMNH);
Brandon, 17-21 Jul 1958, J. G. Chillcott (36, 49;
CNC); Churchill, 2 May-30 Aug, 1918-1952, C.
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
19
FIGURES 13-20.?Setacera micans: 13, head, lateral aspect; 14, male terminalia, posterior aspect;
15, same, lateral aspect; 16, surstyli, posterior aspect; 17, internal male genitalia, lateral aspect;
18, female terminalia, posterior aspect; 19, same, lateral aspect; 20, female ventral receptacle,
lateral aspect.
20
D. Bird, Bryant, W. R. Richards (59; CAS,
CNC); Douglas, 2 mi E, 27 Jul 1958, J. G.
Chillcott (1$; CNC); Treesband, 18 Oct 1927, N.
Criddle (56, 39; CNC); Winnipeg Beach, 2 Jul
1923, A. J. Hunter (Id; CNC). New Brunswick:
Chamcook, Glebe Road, 26 Jun 1965, G. E.
Shewell (1$; CNC). Northwest Territories: Yellow-
knife, 19 May-11 Jul, 1949-1953, E. F. Cashman,
J. G. Chillcott (26, 29; CNC, USNM). Saskatche-
wan: Saskatoon, 12 Jul-24 Oct, 1924-1948, K. M.
King, J. R. Vockeroth (ld\ 3$; CNC). Yukon
Territory: Rampart House, 5-20 Jun 1951, J.E.H.
Martin (16, 52; CNC, USNM); Takhini Hot
Springs, 2400 ft, 16 Aug 1962, P. J. Skitsko (\6;
CNC). UNITED STATES. Alaska: Greater Anchorage
Area Borough, Anchorage, Eagle River Flats, 9
May 1948, F. S. Blanton (16\ 19; USNM); Lower
Yukon River, 3-19 Jul 1951, C. O. Berg (16, 29;
CU); Tonsina, 18 May 1954, W. C. Frohne (16;
USNM).
DISTRIBUTION (Figures 21, 22).?Holarctic.
Northern North America between 49? and 68?
north latitude (Canada and Alaska) and between
94? and 149? west longitude. Northern Europe
in the Palaearctic Region, with confirmed distri-
bution in the British Isles and Baltic countries
but probably occurring more widely into Siberia.
Collin (1963) recorded this species from the
following counties in England and Wales (mostly
unverified): Glamorganshire, Herefordshire,
Huntingdonshire, Norfolk, Suffolk, and Surrey,
from June to September.
vv;i/#?/?^:?>:--
FIGURE 21.?Distribution map of Setacera micans in Europe.
NUMBER 350 17
FIGURE 22.?Distribution map of Setacera micans in North America.
NATURAL HISTORY.?Considerable confusion
exists in the literature with regard to the natural
history of this species, due apparently to misiden-
tifications. Whereas some authors reported the
species to be halophilous (Karl, 1930; Beyer,
1939), others found no preference toward salty
environs (Frey, 1948; Dahl, 1959). To the con-
trary, the latter found this species in association
with a Lemna pool biotope. As none of the North
American records of this species is from salty
habitats, I suspect that the first authors were
dealing with a species of the closely related genus
Ephydra, of which several species are halophilous
(Wirth, 1971, 1975).
Thieneman (in Beyer, 1939) suggested that the
adult season for S. micans is during the autumn.
Dahl (1959), however, found adults more com-
monly during the "height-of-summer."
REMARKS.?Cresson (1930) apparently misi-
dentified this species in his review of the subfam-
ily Ephydrinae in the Naturhistorischen Mu-
seum, Vienna. In his discussion of this species,
Cresson mentioned that the male terminalia are
turgid. In fact, the male terminalia of true S.
micans are unique among Old World species of
this genus in not being turgid; rather, they be-
come smaller in size toward their apices. The
other character states Cresson cited would char-
acterize most of the other Old World species of
Setacera, which leaves me without clues as to
which species Cresson identified as S. micans. I
would guess, however, that he had specimens of
either S. aurata or S. freidbergi, or perhaps a com-
bination of both. The epandrium and surstyli in
males of both of the latter species are turgid,
particularly near their apices.
The breviventris Group
SPECIES INCLUDED.?Setacera breviventris (Loew);
S. multicolor (Soika); and S. viridis Miyagi.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
DIAGNOSIS.?Specimens of the breviventris group
may be distinguished by the following combina-
tion of characters: antennal fovea densely tomen-
tose, contrasting distinctly with subshiny to shiny
dorsum of interfoveal carina; arista with longest
dorsally branching rays at most slightly longer
than aristal width at base; vertico-orbits with
velvety tomentose band narrow but conspicuous,
slightly enlarged subanteriorly; posthumeral bris-
tle lacking or weakly developed, if present dis-
tance between it and presutural bristle equal to
about one-half distance between notopleural bris-
tles; supraspiracular convexity evenly convex;
legs of male lacking conspicuous tufts of rows of
large setae; foretibia, and to a lesser degree mid-
tibia, with basal one-half pale; 5th tergum of
male truncate posteriorly and short, apical width
greater than length, length less than either 3rd or
4th terga; 3rd and 4th sterna of male lacking
dense patch of stout setae toward posterior mar-
gin.
Male Terminalia: Epandrium subrectangular
in posterior view, lateral margins more or less
parallel sided, dorsum very broadly rounded to
subtruncate, with median sulcus extending from
cereal cavity ventrally to just before base of sur-
styli; surstyli, in posterior view, formed as 2 ven-
trally extending arms or projections of various
shapes, with U-shaped pocket between; gonite
with extending process, narrow to moderately
wide; aedeagus rounded apically; hypandrium
usually with median and lateral prongs.
Female Terminalia: 7th tergum complete, oc-
casionally with sclerotization weakened dorsolat-
erally; 8th tergite subtriangular in lateral view,
height less than cereal height; 8th sternite mod-
erately long, length subequal to height of 8th
tergite; 9th sternite dorsoventrally compressed,
wider than high, sclerotization limited to 3 coni-
cal bases from which bristles arise; 9th sternal
bristles with lateral and median bristles sepa-
rated, lateral bristle inserted singly on conical
base, median bristles (1 from each side) approxi-
mate, both inserted on a median base; cercus in
lateral view with posterodorsal cleft and with
anteroventral projection; female ventral recepta-
cle with operculum wider than high, and extend-
ing process longer than height of operculum.
DISTRIBUTION.?This strictly Old World species
group is widely distributed in all of the included
zoogeographic regions. Setacera breviventris is ap-
parently the most widespread species of the genus.
NATURAL HISTORY.?Unlike the other species
groups, the species of the breviventris group are
apparently more commonly encountered along
the banks of lotic aquatic systems, especially small
streams with exposed, mud banks. The specimens
I collected were all found in such habitats.
DISCUSSION.?Although the species group is eas-
ily recognized in both sexes from assessment of
external characters, its component species are
difficult to distinguish without reference to char-
acters of the male terminalia. The female termin-
alia, like external features, are apparently reliable
only at the species-group level, making their iden-
tification problematic without direct association
with males. Distribution data are not totally re-
liable, as some of the species are at least partially
sympatric.
3. Setacera breviventris (Loew)
FIGURES 23-29
Ephydra breviventris Loew, 1860:37.
Ephydra laeta Hendel, 1913:99. [New synonym.]
Ephydra glabra Meijere, 1916:272. [New synonym.]
Setacera breviventris.?Cresson, 1930:117 [review].?Collin,
1963:147 [review].
Setacera pedalis Cresson, 1930:117. [New synonym.]
Setacera fluxa Miyagi, 1966:139; 1977:83 [review]. [New syn-
onym.]
Setacera glabra.?Cogan and Wirth, 1977:338 [oriental cata-
log]-
Setacera laeta.?Gogan and Wirth, 1977:338 [oriental cata-
log]-
DESCRIPTION.?Moderately small to medium-
sized shore flies, length 2.46 to 3.74 mm; dorsum
generally olivaceous green with some brownish to
grayish coloration, becoming duller, grayish to
whitish gray ventrally.
Head: Head width-to-height ratio averaging
1 : 0.65; frons width-to-length ratio averaging
1 : 0.49; mesofrons, fronto-orbits deeply bluish to
NUMBER 350 19
greenish blue, dorsum of interfoveal carina con-
colorous or frequently more greenish; dorsal sur-
face of face shorter than height of anterior sur-
face; angle formed by dorsal surface and anterior
surface moderately angulate, approximately
105?; face with anterior surface silvery white,
frequently facial angle darker, golden to yellowish
brown. Eye height-to-width ratio averaging
1 : 1.1; eye-to-cheek ratio averaging 1 : 0.44.
Thorax (Figure 23): Costal vein ratio averag-
ing 1 : 0.30; M1+2 vein ratio averaging 1 : 0.75.
Abdomen: Male terminalia (Figures 24, 25)
with surstyli in posterior view with lateral and
median margins irregular, apices with shallowly
mucronate, obliquely oriented point, slightly up-
turned in lateral view; gonite in lateral view with
moderately wide posterior process, sides very shal-
lowly sinuate, apex mucronate, median prong
narrow, parallel sided until just before apex, apex
acutely pointed, curved anteroventrally, anterior
process shorter than others, truncate, postero-
ventral angle with 2-3 long setae, situated medi-
ally to other 2 processes; aedeagus rounded in
lateral view, very broadly rounded in dorsal view;
hypandrium with 2 pairs of prongs, lateral pair
stouter and sinuate, median pair very slightly
curved, more slender. Female terminalia (Figures
26-28) as in species group description; female
ventral receptacle with extending process shal-
lowly curved along inner curvature of corpus.
TYPE MATERIAL.?The male lectotype of Ephy-
dra breviventris Loew, herein designated, is labeled
"268 [handwritten]/Coll. H. Loew/14473/Ephy-
dra breviventris m. [handwritten]/Typus [red]/
LECTOTYPE Ephydra breviventris Loew by W.
N. Mathis [handwritten; red]." The lectotype is
in the Zoologisches Museum, Humboldt-Univer-
sitat, Berlin, Germany [DDR]. The lectotype is
pinned directly and is in good condition. As Loew
(1860) did not specify a holotype specimen, I
have designated the only male from Loew's type
series as the lectotype. Loew's original description
indicated that both male and female specimens
were in the type series, although how many spec-
FIGURES 23-28.?Setacera breviventris: 23, thorax, dorsal aspect; 24, male terminalia, posterior
aspect; 25, same, lateral aspect; 26, female terminalia, lateral aspect; 27, cerci, posterior aspect;
28, female ventral receptacle lateral aspect.
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
imens of each sex was not cited. I assume that the
specimens, other than the lectotype, have been
lost or have lost their identity as being part of
Loew's original type series. Fortunately, the re-
maining male specimen is in good condition and
is easily identified.
The male lectotype of Ephydra laeta Hendel,
herein designated, is labeled "Tainan Formosa
H. Sauter 11. 09. (9 Nov 1909)/TYPUS [red]/
Hendel det./Ephydra laeta Hendel/LECTO-
TYPE Ephydra laeta Hendel by W. N. Mathis
[handwritten; red]." The lectotype is in the Na-
turhistorisches Museum, Vienna, Austria. The
specimen is double mounted (minute nadel in
foam block) and is in only fair condition, being
teneral, missing several setae, and both wings
removed (the left wing is glued to the form block).
The abdomen has been removed and dissected
(total length of specimen before dissection 2.97
mm); the structures are in an attached microvial.
Hendel (1913) did not mention a type series nor
did he specify a holotype specimen in the original
description. Because several specimens of this spe-
cies exist that bear label data indicating that they
were collected during the same expedition from
which Hendel obtained the specimen(s) he de-
scribed, it is appropriate to select a lectotype to
avoid any confusion as to which specimen the
name should be associated. Accordingly, from
that series I have selected the only known male,
which is also the only specimen to bear a "TY-
PUS" label. The "TYPUS" label may have been
placed on the pin by Hendel.
The male holotype of Ephydra glabra de Meijere
is labeled "Batavia [Jakarta] VIII 07 [Aug 1907]
Jacobson [handwritten; collector]/Ephydra gla-
bra det. de Meijere Type [scientific name and
"Type" handwritten; black bordered]/HOLO-
TYPE [red with black border]/HoloTYPE Ephy-
dra glabra de Me. 6 det B. H. Cogan 1971.
[species' name handwrittenj/Setacera glabra de
Meij det B. H. Cogan 1971. [species' name hand-
written]." The holotype is in the Instituut voor
Taxonomische Zoologie, Zoologisch Museum,
Amsterdam, Netherlands. The specimen is double
mounted (minute nadel in foam block) and is in
good condition.
The male holotype of Setacera pedalis Cresson is
labeled "Austr[ia]. inf. Wien 9-68 Mik/TYPE
no. 6372 Setacera PEDALIS 6 E. T. Cresson, Jr,
[red]." The holotype is in the Academy of Natural
Sciences of Philadelphia, type 6372. The speci-
men is double mounted (minute nadel in cork
block) and is in good condition. Cresson's original
description also lists two male and nine female
paratypes from "Austria, Alte Sammlung." One
of the latter specimens is in the Academy and is
labeled "ALLO-TYPE 6372 Setacera Pedalis 9
[red]."
The male holotype of Setacera fluxa is labeled
"Fukui 26, Aug. 1963 [yellowj/japonia Honshu
I[chiro]. MIYAGI [yellow]/-type Setacera fluxa
I. Miyagi [name and author handwritten; red]."
The holotype is in the Entomological Institute,
Hokkaido University, Sapporo, Japan. The spec-
imen is double mounted (minute nadel) and is in
good condition. There is considerable disagree-
ment between the label data accompanying the
presumed holotype and paratypes and the pub-
lished data in Miyagi's (1966) paper. According
to the published data, the type-locality and date
are "Wajima, Ishikawa-ken, Honshu, 27-vi-63."
The same type of error is encountered in the
paratype series. I have been assured by the cura-
tor of the insect collection of the Hokkaido Uni-
versity, Dr. S. Takagi, that the specimens I was
sent for study are the holotype and paratypes,
which would mean that the publication con-
tained several errors of transcription.
OTHER SPECIMENS EXAMINED.?ANGOLA. Provin-
cia do Cuanza Sul: Chachoeiras, 20 mi SW Gabela,
18-19 Mar 1972 (1$; BMNH). AUSTRALIA. Austra-
lian Capital Territory: Black Mountain, 2 Jan-30
Dec, 1955-1968, I.F.B. Common (306\ 43?;
ANIC); Fyshwick, 29 Oct-24 Nov, 1961-1966, P.
W. Geier, R. Pilfrey (ld\ 1$; ANIC, USNM). New
South Wales: Gunnedah, 26 Dec 1964, M. I. Niti-
kin (16\ 1$; BMNH). Queensland: Cunnamulla,
Nov 1937 (26, 29; USNM); Gilruth Plains, Cun-
namulla, J. H. Ricks (39; USNM); Ingham, 9
Aug 1960, K. L. Harley (19; USNM); Marlbor-
NUMBER 350 21
FIGURE 29.?Distribution map oi Setacera breviventris.
ough, 63 mi N, 9 May 1955, K. R. Norris (16*;
USNM). Western Australia: Cannington, 30 Jan
1935, K. E. Norris (1$; USNM). AUSTRIA. Klos-
terneueberg (16*, 1$; NMW, USNM); "Alte
Sammlung" (16*, 1$; ANSP, USNM). Kreyen-
berg, 22 May 1911-1912, Jentschoufu, Schant
(2?; DEI). BANGLADESH. Rajshahi, 1-6 Feb 1917
(16*; USNM). EGYPT. Cairo, Nov (16*; USNM).
Faiyum, Mar (1$; USNM). GREECE. Macedonia:
Struma Vail., Jun 1934, Shannon and Hadjini-
calaou (16; USNM). GUAM. 18 Aug 1938, R. G.
Oakley (\6; USNM). HUNGARY. Hodmegzova-
sarhely, 27 Jul 1963, fenycsapda (1$; HNHM);
Szaszka, 22 Jul 1899, Kristen (1$; HNHM); Tar-
hos, 25 Aug 1963, fenycsapda (1(5; HNHM). IN-
DIA. Amdrah Pradesh: Hyderabad, 28 Oct-4 Nov
1971, J. C. Deeming, A. C. Pont (1$; BMNH);
Tirupati, 132 m elevation, 18 Apr 1962, D. Q.
Cavagnaro, E. S. Ross (16*; USNM). Karnataka:
Mudigere, 19 km W, W. N. Mathis, A. Freidberg
(96, 16$; USNM). Union Territory of Delhi: New
Delhi, 5-30 Nov, 1967-1968, K. E. Gibson (126*,
29$; USNM). INDONESIA. Java: Batavia [Jakarta],
1907, Jacobson (1<5; ITZA). Sumatra: Fort de
Kock, 920 m elevation, 1925, E. Jacobson (3$;
BMNH). IRAN. Bisotun Kermanshah: Camp 26,
24-25 Jun 1964, J. Neal (26*; USNM). Gilan:
Camp 30, Bandar Pahlavi, 10-12 Jul 1964, J.
Neal (166\ 19$; USNM). ISRAEL. Mt. Carmel, 22
Aug 1941 (1$; USNM). ITALY. Lombardy: Pa via,
15 Aug 1895 (26, 1$; HNHM, USNM). Bagni di
Tivoli, 27 Oct 1974, W. Rossi (16; MCSNV).
JAPAN. Honshu: Fukui, 26 Aug 1963,1. Miyagi (16*;
HUS); Kyoto, Taiza, 1 Aug 1963, I. Miyagi (3$;
HUS); Omae-zaki, 22 Jul 1963, I. Miyagi (26*;
HUS); Wada Fukui, 27 Jun 1963, I. Miyagi (26*,
1$; HUS). KENYA. Taveta Forest, Aug 1947, M.
Steele (16*; BMNH). NEPAL. Birganj Lothar, near,
450 ft elevation, 3 Sep 1967 (1$; BMNH); Go-
davari, 5000 ft elevation, 5-15 Aug 1967 (56*, 2$;
BMNH); Simra Adhabhar, near, 600 ft elevation,
25 Aug 1967 (2$; BMNH). NIGERIA. Lake Chad,
5 mi W, oxbow lake near Komadugu, Yobe River,
12 Apr 1967, J. C. Deeming (16*; BMNH); Zaria,
Sumaru, 6 Feb-6 Mar, 1967-1972, J. C. Deeming
(16*, 4$; BMNH). PHILIPPINES. Luzon: Mt. Makelk-
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
elen, Baker (36*, 1$; USNM). SOLOMON ISLANDS.
Guadalcanal: Honiara, 8-16 Nov 1953, J. D. Brad-
ley (76*, 10$; BMNH). SRI LANKA. Anudradhapura:
Wilpattu, Hunuwilagama, Wildlife Society Bun-
galow, 200 ft elevation, 10-19 Mar 1970, D.
Davis, W. Rowe (19; USNM); Padaviya, Irriga-
tion Bungalow, 180 ft elevation, 27 Feb-9 Mar
1970, D. Davis, W. Rowe (46*, 3?; USNM). Col-
ombo: Ratmalana, 28-29 Feb 1968, T. F. Halstead
(36\ 119; CAS). Jaffna: Chundikkulam Sanctuary,
25 ft elevation, 7 Nov 1976, G. F. Hevel, R. E.
Dietz (19; USNM). Mannar: Mannar, 4 mi NW,
100 ft elevation, 3 Nov 1976, G. F. Hevel, R. E.
Dietz (16*, 29; USNM). TAIWAN. Tainan, Feb
1909, H. Sauter (59; NMW, USNM). THAILAND.
Bangkok, Huay Kwang, Huai Khwang Sathani,
Aug-Sep 1962, J. Scanlon (16; USNM); Bang-
kok, Bang Pho, Aug-Sep 1962, J. Scanlon (16*;
USNM); Bangkok, Plukchit, Aug-Sep 1962, J.
Scanlon (19; USNM); Bangphra, Cholburi, Oct
1962, J. Scanlon (36, 29; USNM). VIET NAM. Long
Bihn, 1969, W. H. Pierce (16*; USNM).
DISTRIBUTION (Figure 29).?This is the most
widespread species of the genus, occurring in four
of the major zoogeographic regions of the Old
World: Australian, Oriental, Palaearctic, and Af-
rotropical. The distribution falls between 7? and
162? east longitude and between 37? south lati-
tude and 57? north latitude.
NATURAL HISTORY.?I have collected this spe-
cies in southern India along the exposed and
muddy banks of small streams. Although lentic
aquatic systems occurred in the same area, the
species occurred more commonly along lotic sys-
tems.
REMARKS.?As indicated in the synonymy, sev-
eral names have been found to be junior syn-
onyms of S. breviventris. In each case, I have ex-
amined the appropriate type material to confirm
the conspecificity of the included taxa.
4. Setacera multicolor (Soika)
FIGURES 30-35
Ephydra (Setacera) multicolor Soika, 1956:128.
Setacera multicolor.?Cogan, 1980:668 [afrotropical catalog].
DESCRIPTION.?Moderately small shore flies,
length 2.86 to 2.94 mm (based on 2 females);
dorsum generally dark, olivaceous green with
some greenish and grayish coloration, becoming
duller and grayer ventrally.
Head (Figure 30): Head width-to-height ratio
averaging 1 : 0.59; frons width-to-length ratio
30
32
31 34
FIGURES 30-34.?Setacera multicolor: 30, head, lateral aspect; 31, male terminalia, posterior
aspect; 32, same, lateral aspect; 33, female terminalia, lateral aspect; 34, female ventral
receptacle, lateral aspect.
NUMBER 3")0 23
averaging 1 : 0.48; mesofrons, fronto-orbits, and
dorsum of interfoveal carina concolorous, deeply
bluish with some greenish tinges; dorsum of face
shorter than anterior surface; angle formed by
dorsal surface and anterior surface moderately
angulate, approximately 105?. Eye height-to-
width ratio averaging 1 : 1.06; eye-to-cheek ratio
averaging 1 : 0.42.
Thorax: Costal vein ratio averaging 1 : 0.29;
M1+2 vein ratio averaging 1 : 0.90.
Abdomen: Male 5th tergum with anteroventral
angle drawn out to form posteriorly curved, nar-
row projection; male terminalia (Figures 31, 32)
with surstyli in posterior view with apices
rounded, ovate, setose along median margin,
pocket formed between surstyli moderately wide,
width subequal to depth; gonite with a posterior
wide, blunt projection, broadly rounded apically,
median projection mostly fused with anterior one,
forming narrow, posterior process of about one-
half length of anterior process, anterior process
long, moderately narrow, more or less parallel
sided, rounded apically; aedeagus wider than
long, broadly rounded dorsally; hypandrium with
2 pairs of prongs, posteromedian pair approxi-
mate, short, less than one-half length of other
pair, anterolateral pair robust, curved apically
posteriorly. Female terminalia (Figures 33, 34) as
in species group description; female ventral recep-
tacle with inner surface of corpus curvature com-
pressed, angulate.
TYPE MATERIAL.?The female holotype is la-
beled "HOLOTYPUS [light orange with sub-
marginate black border]/MUSEE DU CONGO
KatangarMulongo 12-111-1926 Dr. H. Schoute-
den/HOLOTYPUS Ephydra multicolor n. sp. A.
G[iordani]. S[oika]. [handwritten; red]." The
male allotype has the same locality label data as
the holotype. Both are in the Musee Royal de
l'Afrique Centrale, Tervuren, Belgium. The holo-
type is double mounted (minute nadel in foam
block) and is in fair condition (several setae of the
head are missing). Soika's original description
states that the holotype is a male, but the speci-
men labeled "HOLOTYPUS" is definitely a fe-
FIGURE 35.?Distribution map of Setacera multicolor.
male. Fortunately the allotype is a male that I
dissected to confirm its identity.
OTHER SPECIMENS EXAMINED.?MADAGASCAR.
Sangoritelo, 11 May 1965, Blgor (1<J; BMNH).
SOUTH AFRICA. Cape of Good Hope, Nature Re-
serve, 7 Mar 1968, P. J. Spangler (let; USNM).
TANZANIA. Mpanda, Jun 1960, N. Leleup (29;
MRYAC, USNM). ZAIRE. Kivu: Kavivira (Uvira),
Dec 1954, G. Marlier (let, 19; MRYAC).
DISTRIBUTION (Figure 35).?Afrotropical; be-
tween 3? and 35? south latitude and between 18?
and 44? east longitude.
NATURAL HISTORY.?See species group treat-
ment.
REMARKS.?Both species of Setacera occurring
in the Afrotropical Region are members of the
breviventris group, S. multicolor being the southern
counterpart of S. breviventris.
5. Setacera viridis Miyagi
FIGURES 36-42
Setacera viridis Miyagi, 1966:138; 1977:83 [review].
DESCRIPTION.?Moderately small to medium-
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
sized shore flies, length 2.93 to 3.52 mm; generally
olivaceous dark brown to greenish blue dorsally,
becoming grayer and duller ventrally.
Head (Figure 36): Head width-to-height ratio
averaging 1 : 0.65; frons width-to-length ratio
averaging 1 : 0.48; mesofrons, fronto-orbits, and
dorsum of interfoveal carina metallic green,
bluish green, or mostly bluish. Face in profile
much higher than wide, anterior surface and oral
surface forming an obtuse angle; facial angle
moderately angulate, approximately 100?; face
with anterior surface mostly light golden to brown
white, ventrally nearly silvery white. Eye height-
to-width ratio averaging 1 : 1 ; eye-to-cheek ratio
averaging 1 : 0.36.
Thorax (Figure 37): Costal vein ratio averag-
ing 1 : 0.25; Mi+2 ratio averaging 1 : 0.74.
Abdomen: Male 5th tergum with anteroventral
angle formed into narrow, posteriorly curved,
acutely pointed projection; male terminalia (Fig-
ures 38-40) with surstyli in posterior view short,
rounded apically, median margin conspicuously
more setose than lateral margin, with dorsome-
dian, angulate notch, apices tapered evenly to
acutely pointed apex in lateral view; gonite with
posterior projection broad and truncate, width
twice length, median and anterior projections
united basally, forming 2 pointed projections ap-
ically, posterior margin curved anteriorly on ap-
ical half, deeply U-shaped emargination formed
between apical points; aedeagus broadly rounded
in posterior view, angulate apically in lateral
view; hypandrium with I pair of prongs, arising
from anterior one-third along median margin,
curved anteriorly. Female terminalia (Figures 41,
42) as in species group description; female ventral
receptacle with inner curvature of corpus mod-
erately compressed.
TYPE MATERIAL.?The male holotype is la-
beled "Sapporo: 1961-vi-10/Japonia Hokkaido
I. Miyagi/-type Setacera viridis I. Miyagi '[19]66
[name, author, and date handwritten; red]." The
holotype is in the Entomological Institute, Hok-
kaido University, Sapporo, Japan. The specimen
is double mounted (minute nadel) and is in good
condition. Of the paratypes Miyagi (1966) listed,
I have examined only three males and one female,
all with the same locality data as the holotype.
Other paratypes Miyagi listed are from the fol-
lowing localities, JAPAN. Hokkaido: Numata, 14
Jun 1961 (26, 29; HUS); Oki-Tokachi, 12 Jun
1961 (46\ 5$; HUS); Mashike, 24 Jul 1964 (Id,
4$; HUS); Yakijirijima, 12 Aug 1961 (26, 3$;
HUS); Abashiri, 25 Jul 1962, S. Takagi (16;
FIGURES 36-42.?Setacera viridis: 36, head, lateral aspect; 37, thorax, dorsal aspect; 38, male
terminalia, posterior aspect; 39, same, lateral aspect; 40, internal male genitalia, lateral aspect;
41, female terminalia, lateral aspect; 42, female ventral receptacle.
42
NUMBER 350 25
HUS); Toyotomi, Wakkanai, 2 Jul 1961 (26, 3$;
HUS).
OTHER SPECIMENS EXAMINED.?JAPAN. Hok-
kaido: Tomamae, 12 Aug 1961,1. Miyagi (1<5, 3$;
HUS). Honshu: Wada Fukui, 27 Jun 1963, I.
Miyagi (16*; HUS). KOREA. Seoul [near], Jun
1955, light trap (16\ 1$; USNM).
DISTRIBUTION.?Eastern Palaearctic; Japan
(Hokkaido and Honshu Islands) and Korea.
NATURAL HISTORY.?Miyagi (1966) reported
that this species occurs on the shores of fresh-
water lakes and pools where no alkalinity exists.
REMARKS.?Setacera viridis is like other species
of the breviventris group in that I can distinguish
only male specimens to species. The females listed
above were directly associated with males.
Although S. viridis is closely related to S. brevi-
ventris and apparently has similar habitat prefer-
ences, Miyagi (1966) usually did not find these
two species together in Japan. All of his collec-
tions of this genus from Hokkaido were S. viridis,
while those of Shikoku, Kyushu, and the Ryukyu
Islands were S. breviventris. Only on Honshu
(Wada Fukui) were the two species encountered
together.
The aurata Group
SPECIES INCLUDED.?Setacera aurata (Stenham-
mar).
DIAGNOSIS.?Specimens of the aurata group may
be distinguished by the following combination of
character states: antennal foveae densely tomen-
tose, contrasting distinctly with subshiny to shiny
dorsum of interfoveal carina; longest dorsally
branching rays of arista at most slightly longer
than aristal width at base; vertico-orbits with
velvety tomentose band moderately wide, con-
spicuous, lacking subanterior enlargement;
posthumeral bristle well developed, subequal to
posterior notopleural bristle, distance between it
and presutural bristle about equal to that be-
tween notopleural bristles; supraspiracular con-
vexity evenly convex; legs lacking conspicuous
rows or tufts of larger setae; fore- and midtibiae
dark, concolorous with femora, at most with tib-
ial-femoral articulation pale; 5th tergum of male
subtruncate apically, length approximately equal
to that of 4th, much longer than apical width,
anteroventral margin not produced into narrow
process; 3rd and 4th sterna of male with dense
patch of stout setae toward posterior margin.
Male Terminalia (Figures 44-47): Epandrium
lacking median furrow, greatest width subdorsal
in posterior view, dorsum subtruncate; surstylus
in lateral view with a parallel-sided, lateral pro-
jection that is subtruncate apically and oriented
anteroventrally, lateral margin in posterior view
with subbasal lateral swelling on lateral process,
ventromedian process with lateral margin setose;
gonite irregularly subtriangular, higher than
wide, posterior margin very shallowly emarginate
along basal half, thereafter tapered, rounded to
acutely pointed apex, anterior margin more or
less straight to just before apex, apical one-fourth
or less curved inwardly more abruptly to form
point, basal anterior margin with blunt short
anterior projection, setulose; aedeagus broadly
rounded apically; hypandrium lacking prongs or
narrow process, formed by 2 nearly semicircular
plates, each poorly sclerotized.
Female Terminalia (Figures 48, 49): 7th tergum
complete; 8th tergite long and moderately nar-
row, 3 times higher than wide, more or less par-
allel sided, venter and dorsum evenly and nar-
rowly rounded; 8th sternites short, 3 to 4 times
longer than wide; 9th sternum oriented vertically,
generally poorly sclerotized, especially dorsally,
narrowly U-shaped in posterior view, each lateral
arm setulose basally, more so along median mar-
gin; 9th sternal bristles evenly spaced, approxi-
mate, median 2 aligned slightly more ventrally
than lateral ones, base from which bristles arise
protruded posteroventrally, distance between
bristle insertions and ventral margin of cerci
greater than cereal height; cercus in lateral view
higher than wide, posterior margin broadly
rounded; female ventral receptacle with opercu-
lum comparatively small, wider than high, shal-
lowly papilionate dorsally, extending process
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
44
FIGURES 43-49.?Setacera aurata: 43, head, lateral aspect; 44, male termtnalia, posterior aspect;
45, male terminalia, lateral aspect; 46, internal male genitalia, lateral aspect; 47, aedeagus,
lateral aspect; 48, female ventral receptacle, lateral aspect; 49, female terminalia, lateral aspect.
49
long, cervix about one-half total length, corpus
compressed.
DISTRIBUTION.?See treatment of only included
species (page 27).
6. Setacera aurata (Stenhammar)
FIGURES 43-50
Ephydra aurata Stenhammar, 1844:167.?Zetterstedt, 1846:
1810 [review].?Nartschuk, 1970:387 [key].
Ephydra micans [in part] of authors [not Haliday, 1833:175].
?Loew, 1860:36 [review].?Schiner, 1863:261 [review].
?Becker, 1896:218 [review]; 1905:209 [palaearctic cata-
log]; 1926:75 [review].?Dahl, 1959:111 [review].
Setacera aurata.?CoWin, 1963:148 [review].?Dahl, 1974:186
[distribution, figure of male terminalia].?Papp, 1975:109
[distribution, figures of head, male and female termina-
lia].?Karnecka, 1980:421 [list, Czechoslovakia].
Setacera micans [in part] of authors [not Haliday, 1833:175].
?Canzoneri, 1978:28, 29 [figure of male terminalia].
DESCRIPTION.?Medium-sized to moderately
large shore flies, length 3.52 to 4.76 mm; generally
olivaceous brown with some faint greenish to
bluish coloration dorsally, becoming grayer and
duller ventrally.
Head (Figure 43): Head width-to-height ratio
averaging 1 : 0.63; frons width-to-length ratio
averaging 1 : 0.48; mesofrons, fronto-orbits, and
dorsum of interfoveal carina with metallic luster,
blue to bluish green, dorsum of interfoveal carina
also with sparse tomentose vestiture, slightly
duller; dorsal surface of face shorter than anterior
surface, angle formed by these surfaces obtuse,
about 110?, anterior surface mostly silvery white,
often with some yellowish to yellowish brown
coloration dorsally; eye height-to-width ratio
averaging 1 : 1 ; eye-to-cheek ratio averaging
1 : 0.43.
Thorax: Legs, except for tibial-femoral artic-
ulation mostly dark colored, gray to blackish,
articulation pale. Costal vein ratio averaging
1 : 0.26; MJ+2 vein ratio averaging 1 : 0.90.
Abdomen: Male and female terminalia (Fig-
ures 44-49) as in species group description.
TYPE MATERIAL.?The male lectotype, desig-
nated by Dahl (1959) as a "holotype," is labeled
"[a green diamond, white on underside]/Holoty-
pus aurata Stenh R Dahl [species epithet and "R
Dahl" handwritten, red]." The lectotype is dou-
ble mounted (clipped pin in plastic tube), is in
poor condition (lower portion of head and nu-
merous setae are missing, wings are torn, terminal
segments of abdomen have been removed, dis-
sected, and are in an attached microvial), and is
in the Zetterstedt collection, Zoological Institute,
NUMBER 350 27
Lund, Sweden (the specimen is temporarily in
the custody of Dr. Richard Dahl, Helsingborg,
Sweden).
OTHER SPECIMENS EXAMINED.?ENGLAND. Essex:
Pitsea, 24 Sep 1926, Richards (39; USNM). Nor-
folk: Ring Mere, north of Thetford, 5 Aug 1953
(1<5; UMO); Roudham Heath, 1 Sep 1937 (1$;
UMO). FRANCE. La Panne, 8 Aug 1923, M.
Goetghebuer (16*, 1$; USNM). NETHERLANDS.
Amsterdam, Mar-Oct, Meijere (26, 29; ITZA).
Diemen, 9 Jul 1905, Meijere (1$; ITZA). Hilver-
sum, 5 Mar-16 Oct, 1907-1911, Meijere (2d\ 49;
ITZA). Zerbrug, Feb 1908, Gillary (1$; ITZA).
NORWAY. Bergen, 2 Mar 1952, H. Wiering (1$;
ITZA). SWEDEN. Ostergb'tland: (1 6*; NRS). Scania:
Fyleoset, Ystad, 17 Jun-8 Jul, 1951-1954, R.
Dahl (46, 59; ZIL); Lilla Viken, Hoor, 3 Jul 1954,
R. Dahl (2?; ZIL). N. Tvarminne strand, 19 Jul
1960, R. Dahl (16; USNM). GERMANY (DDR).
Berlin (Jungfhd) (19; NRS). w. GERMANY.
Borkum, Jul 1895 (19; NRS). FINLAND. Jakobstad
(Pietarsaari), 1952, Sotra (16; USNM). Locality
unspecified (66, 39; HU, ITZA, NMW, ZIL).
Collin (1963) also recorded this species from
the following counties in England and Wales:
Cambridgeshire, Essex, Glamorganshire, and
Norfolk. Dahl (1974) examined specimens of S.
aurata from the following localities in Sweden
(unverified and all collected by him). Shane: Fors-
lovsholm, 20 Jul 1971; Hoor, 1 Aug 1966; Ystad,
Fylean, 8 Jul 1954, Lomma, 24 Aug 1954. Blek-
inge: Karlshanm, 3 Aug 1968. Karnecka (1980)
listed this species from Bohemia and Moravia in
Czechoslovakia.
DISTRIBUTION (Figure 50).?Palaearctic. North-
ern Europe from Norway and European Russia
south to France and Hungary.
REMARKS.?Except for Zetterstedt (1846) and
a few recent workers (see synonymical bibliog-
raphy), most authors have confused this species
with S. micans. The misidentification began when
Loew (I860) listed S. aurata as a synonym of S.
micans, a precedent that was followed by several
European authors (see synonymy). Dahl (1959)
further complicated the misidentification of Loew
and others by stating that he had "examined the
FIGURE 50.?Distribution map of Setacera aurata.
male genitalia of the holotype of that species [S.
aurata] in Stenhammar's Collection. The exami-
nation revealed the identity with micans." Dahl
could not have studied the "holotype" of S. aurata,
as Stenhammar did not designate one; however,
Dahl's statement is sufficient for a lectotype des-
ignation. Collin (1963) correctly identified both
species, designated a lectotype for S. micans, illus-
trated the male terminalia of a probable syntype
of S. aurata from Stenhammar's collection, noted
several differences between the male terminalia
of S. micans and S. aurata, and specified the identity
of specimens in several collections he had exam-
ined from Scandinavia. Dahl (1974) corrected his
earlier misidentification and cited additional lo-
cality data for S. aurata; he also figured a surstylus
of male S. micans and S. aurata. Canzoneri (1978),
however, perpetuated the confusion with these
two species by publishing figures of S. aurata
under the name of "5". micans."
This species is distinguished from congeners by
the characters noted in the species group diag-
nosis.
The trina Group
SPECIES INCLUDED.?Setacera freidbergi, new spe-
cies; 5. meneghinii Canzoneri; S. trina Collin.
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
DIAGNOSIS.?Specimens of the trina group may
be distinguished by the following combination of
characters: antennal foveae densely tomentose,
contrasting distinctly with subshiny to shiny dor-
sum of interfoveal carina; arista with longest
dorsally branching rays at most slightly longer
than aristal width at base; vertico-orbits with
velvety tomentose band moderately narrow but
conspicuous, slightly enlarged subanteriorly at
vertex; posthumeral bristle moderately well de-
veloped, subequal to largest humeral bristle to
slightly weaker than posterior notopleural bristle,
distance between it and presutural bristle slightly
more than one-half that between notopleural bris-
tles; supraspiracular convexity evenly convex;
legs of male lacking conspicuous tufts or rows of
large setae; fore- and midtibiae with up to basal
one-third pale, although frequently considerably
less; 5th tergum of male subtruncate apically,
length approximately equal to that of 4th, ante-
roventral margin not produced into narrow pro-
cess; 3rd and 4th sterna of male with dense patch
of stout setae toward posterior margin.
Male Terminalia: Epandrium lacking median
sulcus, greatest width subdorsal in posterior view,
dorsum subtruncate; surstyli broadly fused with
ventral margin of epandrium, lateral margin in
posterior view rounded, gradually tapered in-
wardly to just before apices, thereafter recurved
laterally, apex in lateral view slightly flared pos-
teriorly; gonite in lateral view with posterior and
anterior process, anterior process with angulate
margin, curved apically, inner surface of curva-
ture setulose; aedeagus broadly rounded apically;
hypandrium formed by 2 narrowly triangular,
poorly sclerotized plates, lacking prongs or ex-
tended processes, attached anteriorly to 5th ster-
num.
Female Terminalia: 7th tergum complete; 8th
tergites long and narrow, over 3 times higher than
wide, widest subdorsally, dorsum angulate, nar-
rowest at midheight, from there to venter gradu-
ally becoming wider, venter evenly rounded; 8th
sternites short, about 3 times longer than wide;
9th sternum oriented vertically, generally poorly
sclerotized, broadly U-shaped in posterior view,
as high as wide, each dorsally extending arm
slightly turgid, lateral margin rounded; 9th ster-
nal bristles evenly spaced, approximate, and more
or less aligned horizontally, base from which bris-
tles arise not protruding; distance between bristle
insertion and ventral margin of cerci greater than
cereal height; cercus in lateral view higher than
wide, posterior broadly rounded; female ventral
receptacle with operculum comparatively small,
about twice as wide as high, broadly trapezoidal
in shape, extending process comparatively long,
cervix less than one-half length, corpus moder-
ately compressed.
DISTRIBUTION.?The composite distribution of
the trina group occupies an area from Great Brit-
ain through Germany, Italy, and Israel to Iran.
Nearly all specimens I have examined are from
type-localities only.
DISCUSSION.?The monophyly of this species
group is well established. The unique shape of
the gonite of the male terminalia, especially the
anterior process, is an apotypic character. The
anterior process is narrow and recurved apically,
and the inner curvature is setulose.
External differences between the species of the
trina group are slight, and I have relied on features
of the male and female terminalia to determine
the identity of specimens. Because most species
are known only from topotypical specimens, the
extent and limits of intraspecific variation are
essentially unknown.
7. Setacera freidbergi, new species
FIGURES 1, 51-66, 70
DESCRIPTION.?Medium-sized to moderately
large shore flies, length 3.37 to 4.31 mm; generally
greenish, sometimes faintly greenish brown to
gray dorsally, becoming distinctly grayer and
duller ventrally.
Head (Figures 51, 59-61): Head width-to-
height ratio averaging 1 : 0.68; frons width-to-
length ratio averaging 1 : 0.49; metallic luster of
mesofrons, fronto-orbits, and dorsum of interfov-
eal carina bluish to very slightly greenish blue;
NUMBER 330 29
FIGURES 51-58.?Setacera freidbergi: 51, head, lateral aspect; 52, thorax, dorsal aspect; 53, internal
male genitalia from type-locality; 54, internal male genitalia from Iran; 55, abdomen of male,
dorsal aspect; 56, male terminalia, posterior aspect; 57, surstyli from type locality, posterior
aspect; 58, surstyli from Iran, posterior aspect.
dorsal surface of face shorter than height of an-
terior surface; facial angle formed by anterior
surface and dorsal surface about 110?; face with
anterior surface yellowish brown, golden, to
mostly whitish dorsally, becoming amost entirely
whitish ventrally; eye height-to-width ratio aver-
aging I : 0.82; eye-to-cheek ratio averaging
1 : 0.44.
Thorax (Figures 52, 62-64): Fore- and midti-
biae with basal one-third to two-thirds pale,
otherwise grayish blue to faintly grayish green.
Costal vein ratio averaging 1 : 0.28; Mi+2 vein
ratio averaging 1 : 0.91.
Abdomen (Figure 55): Male terminalia (Fig-
ures 53, 54, 56-58) with apices of recurved sutstyli
forming distinct angulate point (Figure 56); gon-
ite in lateral view nearly parallel sided over most
of length, slender, taper on apical one-fourth
more abrupt (Figure 53). Female terminalia as in
Figures 65, 66; female ventral receptacle with
cervix very gradually widened toward corpus;
junction of cervix with corpus gradual; corpus
moderately compressed, more so than in S. trina.
TYPE MATERIAL.?The male holotype is la-
beled "ISRAEL: Nahal Tut (5km E. Fureidis) 19
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURES 59-66.?Setacera freidbergi: 59, ventral portion of head, lateral aspect; 60, left vertex and
vertico-orbits of head, dorsal aspect; 61, left vertico-orbit showing velvety tomentosity, dorsal
aspect; 62, notopieuron and surrounding area of thorax, lateral aspect; 63, propleuron, lateral
aspect; 64, hind coxal strap bearing setae, lateral aspect; 65, female terminalia, ventral aspect;
66, same, lateral aspect.
?
NUMBER 350 31
May 1980 Mathis and Freidberg collectors." Al-
lotype female and 34 paratypes (156\ 199) are
labeled with the same locality data as the holo-
type. The holotype is double mounted (minute
nadel in plastic elastomer base) and is in excellent
condition. The holotype, allotype, and most par-
atypes are in the National Museum of Natural
History, Smithsonian Institution, USNM 76884.
Two male and two female paratypes are in the
insect collection of Tel Aviv University.
OTHER SPECIMENS EXAMINED.?IRAN. Kerman-
shah Province: Bisotun Camp 26, 24-26 Jun 1964,
John Neal (66, 89; USNM).
DISTRIBUTION (Figure 70).?Near East of the
Old World between 35? and 48? east latitude
and between 32? and 35? north longitude.
NATURAL HISTORY.?Dr. Amnon Freidberg
and I collected the topotypical series from a small
algae-covered pond that was fed mostly from
runoff irrigation water. As with most Setacera
species, specimens were nearly impossible to col-
lect by normal sweeping, and our series was col-
lected by lowering a net over the top of a few flies
as they rested on the water's surface. The speci-
mens were abundant on the pond's surface and
were not prone to fly far when disturbed.
ETYMOLOGY.?This species is named to honor
Dr. Amnon Freidberg, who helped collect the
type series, and who has promoted the study of
Diptera from the Near East.
REMARKS.?The specimens from Iran are very
similar to those from the type-locality, although
the shape of the gonite consistently differs slightly.
I am of the opinion that the two populations are
conspecific and that the differences represent in-
traspecific variation of allopatric populations. A
subspecific name could be applied, but no advan-
tage is gained by so doing.
This species is closely related to the other two
known species of the trina group, and I can distin-
guish them only after examination of the male
terminalia.
8. Setacera meneghinii Canzoneri
FIGURES 67-70
Setacera meneghinii Ganzoneri, 1978:28.
DESCRIPTION.?Moderately large shore flies,
length 4.0 to 4.5 mm; generally olivaceous brown
with some bluish to greenish tinges dorsally, be-
coming grayer and duller ventrally (Canzoneri,
1978; the male paratype I examined was approx-
imately 3.9 mm but lacked the terminal segment
of the abdomen).
Head (Figure 67): Head width-to-height ratio
1 : 0.58; frons width-to-length ratio 1 : 0.48;
metallic luster of mesofrons and fronto-orbits
deeply greenish blue, dorsum of interfoveal carina
bronzish green; dorsal surface of face shorter than
anterior surface; angle formed by anterior surface
and dorsal surface obtuse, about 120?; face
brownish gold dorsally, becoming mostly whitish
ventrally. Eye width-to-height ratio 1 :1 ; eye-to-
cheek ratio 1 : 0.44.
Thorax: Fore- and midtibiae with basal one-
fourth or less pale. Costal vein ratio 1 : 0.26; M1+2
vein ratio 1 : 0.92.
Abdomen: Male terminalia (Figures 68, 69)
with apices of recurved surstyli rounded laterally,
not pointed (Figure 69) in posterior view; gonite
in lateral view moderately wide, tapered gradu-
ally toward apex until just before apex, thereafter
abruptly tapered to form acutely pointed apex.
Females unavailable for study.
TYPE MATERIAL.?The male holotype is la-
68
67
FIGURES 67-69.?Setacera meneghinii: 67, head, lateral aspect;
68, internal male genitalia, lateral aspect; 69, surstyli, pos-
terior aspect.
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 70.?Distribution map of Setacera trina (filled circles); S. meneghinii (star); S. freidbergi
(diamonds).
beled (based on the label data of a male parato-
potype) "RIETI Sorgenti Solfuree 14-VIII-58
[14 Aug 1958] A. Giordani Soika." The allotype
female and a male paratype are from the same
locality and were collected on the same date. The
female paratype from Bagni di Tivoli (27 Oct
1974, W. Rossi) was misidentified and is a female
of S. breviventris. The type series is in the Museo
Civico di Storia Naturale de Venezia, Italy.
DISTRIBUTION (Figure 70).?Presently known
only from the type locality, ITALY. Rieti Province:
Rieti.
REMARKS.?My treatment of this species is
based on minimal data. Aside from the original
description, I have examined one male parato-
potype.
This species seems to be intermediate between
S. freidbergi and S. trina, as assessed by the shape
of the gonite.
9. Setacera trina Collin
FIGURES 70-74
Setacera trina Collin, 1963:148.?Karnecka, 1980:421 [list,
Czechoslovakia].
Ephydra trina.?Nartschuk, 1970:387 |key].
DESCRIPTION.?Moderately large shore flies,
length 4.21 mm (1 female); generally olivaceous
brown with some greenish coloration dorsally,
becoming grayer and duller ventrally.
Head (Figure 71): Head width-to-height ratio
averaging 1 : 62; frons width-to-length ratio av-
eraging 1 : 0.47; metallic luster of mesofrons and
fronto-orbits greenish blue to mostly bluish, dor-
sum of interfoveal carina lighter colored, sub-
shiny, mostly bronzish green; dorsal surface of
face shorter than height of anterior surface; facial
angle formed by anterior surface and dorsal sur-
face about 110?; face silvery white to light brown-
ish white, slightly darker dorsally; eye height-to-
width ratio averaging 1 : 1 ; eye-to-cheek ratio
averaging 1 : 0.43.
Thorax: Tibiae mostly dark colored, only fem-
oral-tibial articulation pale. Costal vein ratio av-
eraging 1 : 0.29; Mi+2 vein ratio averaging 1 : 1 .
Abdomen: Male terminalia (Figure 72) with
gonite in lateral view with posterior projection
broadly triangular, apical angle about 45?. Fe-
NUMBER 350 33
74
FIGURES 71-74.?Setacera trina: 71, head, lateral aspect; 72, male terminalia of holotype, lateral
aspect; 73, female ventral receptacle, lateral aspect; 74, female terminalia, lateral aspect.
male terminalia as in Figure 74; female ventral
receptacle (Figure 73) with cervix parallel sided,
C-shaped corpus moderately compressed, curva-
ture at junction with cervix abrupt.
TYPE MATERIAL.?The male lectotype, herein
designated, is labeled with a round disk, on the
underside of which is written "Snailwell [Cam-
bridge] 4.7.11 [4Jul 1911]/Setacera trina 6 TYPE
[handwritten]/LECTOTYPE Setacera trina Col-
lin 6* by W. N. Mathis [handwritten, red]." The
lectotype is in the University Museum, Oxford
University, Oxford, England. The specimen is
double mounted (minute nadel in foam block)
and is in good condition, although some of the
tarsomeres of the left legs are missing. The ab-
domen has been removed and dissected; the male
terminalia are mounted in balsam on a celluloid
rectangle that is pinned separately from the rest
of the body.
OTHER SPECIMENS EXAMINED.?GREAT BRITAIN.
Norfolk: Roudham Heath, 28 Sep 1947 (1$;
UMO). GERMANY. Bay em: Dachau (\6; USNM).
DISTRIBUTION (Figure 70).?In addition to the
localities listed for the specimens I examined,
Collin (1963) recorded this species from the fol-
lowing counties in Great Britain: Cambridge-
shire, Devonshire, Hampshire, Herefordshire,
Hertfordshire, Norfolk, Suffolk, and Surrey. Kar-
necka (1980) recently recorded this species from
Bohemia and Moravia in Czechoslovakia. Al-
though specimens are known only from Great
Britain, Czechoslovakia, and Germany, I suspect
that the species will be found to occur more
widely in northern Europe. This is the northern-
most species of the trina group.
REMARKS.?In the discussion section of the
original description, Collin (1963:148) com-
mented briefly on some, perhaps all, of the spec-
imens of the type series. His discussion included
one sentence that probably referred to a type,
although it is not clear whether Collin was refer-
ring to a type in the nomenclatural sense or to a
particular type of male. Collin wrote: "The draw-
ings have been made from the type of male taken
at Snailwell (Cambs.) on 4th July, 1911 " I
have designated a lectotype for S. trina to elimi-
nate any confusion regarding the status of its
supposed type.
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
The width of the male gonite at its base is the
largest within the species group and serves to
distinguish this species from closely related con-
geners.
The aldrichi Group
SPECIES INCLUDED.?Setacera aldrichi Cresson.
DIAGNOSIS.?Specimens of the aldrichi group
may be distinguished by the following combina-
tion of characters: antennal foveae sparsely to-
mentose, subshiny, concolorous and with same
luster as dorsum of interfoveal carina; arista with
longest branching dorsal rays about twice aristal
width at base; vertico-orbits with velvety tomen-
tose band narrow but conspicuous, with suban-
terior enlargement; posthumeral bristle well de-
veloped, subequal to posterior notopleural bristle,
distance between it and presutural bristle about
equal to that between notopleural bristles; su-
praspiracular convexity evenly convex; fore- and
midtibiae dark, concolorous with femora, at most
with immediate tibial-femoral articulation pale;
5th tergum of male broadly truncate to slightly
emarginate, length greater than either 3rd or 4th
terga, apical width greater than or equal to
length, anteroventral corner with narrow, long,
process, curved posteriorly, truncate apically; 3rd
and 4th sterna of male lacking dense patch of
stout setae toward posterior margin.
Male Terminalia: Epandrium with median sul-
cus from cereal cavity to base of surstyli, subrec-
tangular, dorsum subtruncate, lateral margins
nearly parallel sided, slightly wider dorsally in
posterior view; surstyli not fused basomedially,
separation nearly equal to width of 1 surstylus at
base, basal two-thirds flared laterally subapically,
acutely pointed; gonite lacking pronglike pro-
cesses, broadly protruding posteriorly, posterior
margin irregularly sinuate, with anteroventro-
medial truncate projection bearing short,
rounded, lateral process; aedeagus broad basally,
acutely pointed posteroapically; hypandrium
with 2 anterior prongs, lateral prong over twice
length of median one, median prong short, both
acutely pointed, median area of hypandrium with
sclerotization less well developed.
Female Terminalia: 7th tergum incomplete,
with 2 lateral tergites; 8th tergites about 3 times
higher than wide in lateral view, swollen and
widest at midheight, dorsum and venter narrowly
rounded; 8th sternites short, at most 3 times
longer than wide; 9th sternites vertically oriented
with 1 ventral projection horizontal, bearing
larger bristles; 9th sternal bristles approximate,
irregularly aligned horizontally several smaller
setae dorsad of larger bristles; cercus higher than
wide, with anterodorsal projection, posterior mar-
gin subtruncate to broadly rounded; female ven-
tral receptacle with operculum wider than high,
broadly rounded dorsally, extending process with
cervix about as long as corpus, lateral margin
sinuate, corpus curvature compressed.
DISTRIBUTION.?See species treatment, page 36.
NATURAL HISTORY.?See species treatment,
page 36.
DISCUSSION.?This species group is the sister
lineage of the pacifica species group. This relation-
ship and the monophyly of the species group are
shown in Figure 2, with accompanying character
evidence.
10. Setacera aldrichi Cresson
FIGURES 75-82
Setacera aldrichi Cresson, 1935:348.?Sturtevant and Wheeler,
1954:202, 203 [key and locality data].?Wirth and Stone,
1956:472 [key].?Wirth, 1965:754 [nearctic catalog].?
Cole, 1969:402 [locality data].
DESCRIPTION.?Medium-sized to moderately
large shore flies, length 3.84 to 4.92 mm; dorsum
darkly olivaceous brown to mostly greenish, be-
coming grayer, duller ventrally.
Head (Figure 75): Head width-to-height ratio
averaging 1 : 0.64; frons width-to-length ratio
averaging 1 : 0.48; mesofrons and fronto-orbits
deeply blue to greenish blue, dorsum of interfov-
eal carina more greenish blue, slightly lighter
than mesofrons; shape of face sexually dimorphic,
NUMBER 350 35
FIGURES 75-81.?Setacera aldrichi: 75, head, lateral aspect; 76, male terminalia, lateral aspect;
77, same, posterior aspect; 78, abdomen, dorsal aspect; 79, surstyli, posterior aspect; 80, female
ventral receptacle, lateral aspect; 81, female terminalia, lateral aspect.
dorsal surface of male longer than height of an-
terior surface, subexplanate, anterior facial ridge
broadly rounded, antennal foveae only slightly
lower than dorsum, female with dorsal surface
about equal to anterior surface, not subexplanate,
antennal foveae inclinate; facial angle about
110?; anterior surface of face golden brown dor-
sally, becoming silvery white ventrally. Eye
height-to-width ratio averaging 1 : 0.98; eye-to-
cheek ratio averaging 1 : 0.43.
Thorax: Midtibia of male with dense patch of
short setae ventroapically; costal vein ratio aver-
aging 1 : 0.28; Mi+2 vein ratio averaging 1 : 0.83.
Abdomen (Figure 78): Male terminalia (Fig-
ures 76, 77, 79) as in species group description.
Female terminalia (Figures 80, 81) as in species
group description.
TYPE MATERIAL.?The male holotype is la-
beled "Potlatch Ida[ho] VI. 20. 07. [20 Jun 1907;
J. M. Aldrich collector]/c5 TYPE No. 6515 Seta-
cera ALDRICHI 6 E. T. Creson, Jr, [red]." The
holotype is in the Academy of Natural Sciences
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
of Philadelphia, ANSP 6515. Cresson's original
description lists a female paratopotype that is also
in the Academy's collection. The holotype speci-
men is directly pinned and is in good condition.
OTHER SPECIMENS EXAMINED.?CANADA. British
Columbia: Clayton, 9 Aug 1917, A. L. Melander
(26*; USNM). UNITED STATES. California: Mono Co.,
Mammoth Lakes, 29 Jul 1940, L. C. Kuitert (19;
SU). Idaho: Latah Co., Moscow, J. M. Aldrich
(16; USNM); Moscow, Lake Merton, 2 Mar
1911, J. M. Aldrich (1$; USNM); Potlach, 20
Jun 1907, J. M. Aldrich (16\ 19; ANSP, USNM).
Nez Perce Co., Lewiston, 22 Jun 1968, R. Sanders
(Id; WSU). Oregon: Grant Co., Seneca, 4.8 mi S,
17 Jun 1972, W. N. Mathis (56, 5$; USNM).
Union Co., Ladd Canyon, 14 mi S LaGrande,
4250 ft, pond, 30 Jul 1977, R. S. Zack, E. J. Davis
(36; USNM). Utah: Rich Co., Garden City, 9 Aug
1967, G. F. Knowlton (\6; USNM). Washington:
Clallam Co., Sequim, 25 Aug 1910 (1$; ANSP).
Grant Co., O'Sullivan Dam, 4-5 Apr 1956, H. G.
Davis (16; WSU). Whitcam Co., Steptoe Canyon,
10 mi SW Pullman, 19 Feb 1977, W. J. Turner
(Ic3; USNM). Wyoming: Lincoln Co., Kemmerer,
14 Aug 1950, A. H. Sturtevant (16\ 1$; USNM).
Teton Co., Yellowstone Park Lake, 18 Jul 1923,
A. L. Melander (16*; ANSP).
DISTRIBUTION (Figure 82).?Western North
America between 110? and 128? west longitude
and 37? and 53? north latitude.
NATURAL HISTORY.?I have collected specimens
of S. aldrichi by sweeping emergent vegetation
near the shore of a small backwater area of a
mountain stream. Label data accompanying bor-
rowed specimens that I examined indicate that
the species also occurs on or near lakes and ponds.
FIGURE 82.?Distribution map of Selacera aldrichi.
The pacifica Group
SPECIES INCLUDED.?Setacera durani Cresson; S.
jamesi, new species; S. needhami Johannsen; S. pa-
cifica (Cresson); S. pilicornis (Coquillett); and S.
trichoscelis, new species.
DIAGNOSIS.?Specimens of the pacifica group
may be distinguished by the following combina-
tion of characters: antennal foveae sparsely to-
mentose, subshiny, nearly concolorous with dor-
sum of interfoveal carina and with similar metal-
lic luster; vertico-orbits with velvety tomentose
band very narrow, inconspicuous; posthumeral
bristle well developed, subequal or slightly weaker
than posterior notopleural bristle, distance be-
tween it and presutural bristle slightly less than
that between notopleural bristles; fore- and mid-
tibiae mostly dark and concolorous with femora,
at most with tibial-femoral articulation pale; 5th
tergum of male longer than either 3rd or 4th,
narrowly to bluntly rounded, width at apex much
less than length; 3rd and 4th sterna of male
usually with dense patch of stout setae toward
posterior margin, secondarily reduced in some
species (S. needhami and S. trichoscelis).
Male Terminalia: Epandrium subtriangular in
posterior view, much narrower dorsally than ven-
NUMBER 350 37
trally, dorsum narrowly rounded, becoming grad-
ually broader, truncate ventrally, lacking median
sulcus (S. pacifica with superficially indication);
cerci almost as wide as high in lateral view, with
anteroposterior orientation; surstyli broadly fused
basally (best seen in posterior view), usually with
lateral and sometimes with median projections
(shape diagnostic at species level), apically curved
anteriorly in lateral view; gonite with pronglike
projections, conformation and arrangement dif-
fering with species; aedeagus bluntly rounded
apically; hypandrium with 1-2 projections, at
least partially and usually mostly well sclerotized.
Female Terminalia: 7th tergum variable; 8th
tergite comparatively short, almost as wide as
high in some species, shape of dorsum in lateral
view varying with species; 8th sternites elongate,
4 or more times longer than wide; 9th sternites
vertically oriented, projecting posteriorly, well
sclerotized, forming 2 conical projections; 9th
sternal bristles borne at apex of 9th sternal pro-
jections, 2 large bristles on each, these approxi-
mate, often difficult to distinguish; female ventral
receptacle with operculum as high as wide, sub-
trapezoidal to dome-shaped in lateral view, ex-
tending process with cervix as long as corpus,
more or less paralled sided, juncture of cervix and
corpus indicated on inner surface of curvature by
small, lateral indentation, forming budlike pro-
jection in lateral view, curvature of corpus wide,
open.
DISTRIBUTION.?New World. Except for 51. tri-
choscelis, this species group is North American,
mostly occurring west of the 100? meridian. An
obvious exception is S. pilicornis, which occurs in
southeastern United States and in Mexico.
NATURAL HISTORY.?Adults are generally as-
sociated with lentic aquatic systems, usually
where algal mats have accumulated on the wa-
ter's surface. Only the larva and puparium of S.
needhami have been described (Johannson, 1935).
DISCUSSION.?The characterization of the paci-
fica group and the evidence to establish its mon-
ophyly was elaborated earlier (see "Phylogeny,"
page 6). Herein (Figure 83) I have arranged the
species comprising the pacifica group into three
FIGURE 83.?Hypothetical phylogeny of species of ihe pacifica
species group.
primary lineages: (1) the stem lineage giving rise
to S. pilicornis, which is the sister group to the
remaining species; (2) the stem sublineage giving
rise to S. trichoscelis, S. needhami, and S. jamesi,
which comprise the sister group of sublineage
number three; and (3) the stem sublineage giving
rise to S. durani and 5. pacifica, which comprise the
sister group of sublineage number two. Character
evidence to establish these lineages is as follows.
Stem lineage number one is characterized, and
its monophyly is established, as follows:
31. Configuration of male terminalia: the unique shape of the
male terminalia of this species is autapotypic.
32. Configuration of female terminalia: see discussion of no. 31
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
(see figures accompanying species treatment).
The stem lineage giving rise to sublineages two
and three (see discussion above), the remaining
species of the pacifica group, is characterized, and
its monophyly is established, as follows:
33. Vestiture offorecoxa of male: throughout the subfamily,
the forecoxae are typically invested with some setulae, espe-
cially along the lateral margins. In males of this lineage,
however, the setae are longer, appearing like long pile. I
interpret this character to be apotypic.
The second sublineage is characterized, and its
monophyly is established, as follows:
34. Epandrium: a median furrow is the typical and therefore
the relatively plesiotypic condition. The lack of such a furrow
in males of this group is apotypic.
35. Configuration of cerci of female: the female cerci are
generally higher than wide, th<: plesiotypic condition. In
females of this sublineage, the cerci are as wide as high, an
apotypic character.
36. Vestiture of gonite of male: the presence of two long,
posterior setae on the gonite is a feature unique to this
sublineage. These setae are not present in the other lineages
of the genus.
The third sublineage is characterized, and its
monophyly is established, as follows:
37. Configuration of male lerminalia: males of this group have
a median projection, apparently a unique, apotypic feature.
The second sublineage is further divided into
two groups. The first is characterized, and its
monophyly is established, as follows:
38. Vestiture of mid- and hind tibiae: typically there are a few
setae on the mid- and hind tibiae, but males of this sublin-
eage have apicoventral tufts of long setae, an apotypic
character.
39. Vestiture of midfemur of male: males of these two species
have a short row of distinct setae along the venter of the
midfemur. Because this is apparently a unique condition, I
interpret it to be apotypic.
The second lineage of the second sublieage is
characterized, and its monophyly is established,
as follows:
40. Size: this is the largest species of the genus, apparently
an apotypic character.
11. Setacera durani Cresson
FIGURES 84 -90
Setacera durani Cresson, 1935:348.?Sturtevant and Wheeler,
1954:202, 203 (key and locality data].?Wirth and Stone,
1956:472 [key].?Wirth, 1965:754 [nearctic catalog].?
Cole, 1969:402 [locality data].
DESCRIPTION.?Medium-sized to moderately
large shore flies, length 3.45 to 4.95 mm; generally
mostly gray to bluish green, dark, becoming
grayer and duller ventrally.
Head (Figure 84): Head width-to-height ratio
averaging 1 : 0.61; frons width-to-length ratio
averaging 1 : 0.48; mesofrons, fronto-orbits, and
dorsum of interfoveal carina deeply greenish blue;
arista with longest dorsally branching rays about
twice greatest aristal width at base; dorsal surface
of face explanate laterally (best seen from dorsal
view), antennal fovea only slightly lower than
dorsum of interfoveal carina; facial angle approx-
imately 90?; angle formed by anterior surface
and oral margin obtuse; anterior surface of face
mostly silvery white, facial angle brownish to
olivaceous. Eye height-to-width ratio averaging
1 : 1.1; eye-to-cheek ratio averaging 1 : 0.52.
Thorax: Supraspiracular convexity evenly
convex; forecoxa of male with dense patch of
long, slender setae, especially posteroventrally;
midtibia of male with dense patch of short setae
apicoventrally. Costal vein ratio averaging
1 : 0.27; Mi+2 vein ratio averaging 1 : 0.79.
Abdomen: Male 5th tergal ratio averaging
1 : 0.66, anteroventral angle lacking process. Male
terminalia (Figures 85-87): surstyli in posterior
view with lateral margins broadly rounded, lat-
eral projections blunted pointed, 2 U-shaped
emarginations ventrally, separated by median
process that is parallel sided basally, tapering
apically to acutely pointed apex, this process in
lateral view curved posteriorly; gonite lacking
pronglike process but with acutely pointed,
broadly based posterior process at about mid-
height; aedeagus folded back on itself, broadly
rounded apically; hypandrium with lateral mar-
gins well sclerotized becoming weaker toward
median, median formed into pocket, with an
NUMBER 350 39
FIGURES 84-89.?Setacera durani: 84, head, lateral aspect; 85, 3rd and 4th sterna of male, ventral
aspect; 86, surstyli, posterior aspect; 87, male terminalia, lateral aspect; 88, female terminalia,
lateral aspect; 89, female ventral receptacle, lateral aspect.
anteromedial and posterolateral prong, postero-
lateral prong arising from broad base, thereafter
slender, tapered to acutely pointed apex, antero-
medial prong longer, gradually tapered from base
to apex. Female terminalia (Figures 88, 89): 7th
tergum complete; 8th tergites with anterior mar-
gin rounded, posterior margin shallowly emargin-
ate, ventral margin broadly rounded, posterior
portion slightly higher than anterior portion; 8th
sternites elongate, 5-6 times longer than greatest
width; cercus higher than wide, posterior margin
evenly rounded; female ventral receptacle (Figure
89) with operculum subtrapezoidal but with
rounded lateral margins, otherwise as in species
group description.
TYPE MATERIAL.?The male holotype is la-
beled: "Los Angeles R[iver]. [Los Angeles Co.]
California]. Aug. 15, 1916 V. Duran, Coll/cV
TYPE No. 6516 Setacera DURANI E T Cresson,
Jr. [name handwritten, red]." The holotype spec-
imen is double mounted (minute nadel in card-
board base), is in excellent condition, and is in
the Academy of Natural Sciences in Philadelphia,
ANSP 6516. Cresson's original description listed
a female paratopotype; this specimen is also in
the ANSP.
OTHER SPECIMENS EXAMINED.?MEXICO. Baja
California Norte: San Vicente, 20 Sep 1941, Ross,
Bohart (1(5; CAS); Santo Domingo, 5.7 mi E
Hamilton Ranch, 22 Apr 1963, H. B. Leech, P.
H. Arnaud, Jr. (46, 19; CAS, USNM); Tijuana,
60 km S, 26 Jun 1950, A. L. Melander (Id;
USNM). UNITED STATES. Arizona: Cochise Co., Por-
tal, Southwestern Research Station, 5-9 Jun
1972, W. W. Wirth (1(5; USNM). Yuma Co.,
Alamo Canyon, 7 May 1960, C. E. Benson (Id;
KSU). California: Alameda Co., Oakland, 26 May
1915, M. C. Van Duzee (1$; CAS). Contra Costa
Co., Danville, 12 Jul 1951, F. X. Williams (1<5;
CAS). Imperial Co., Calexico, 15 mi E, 5-6 Jun
1961, H. F. Howden (3$; CNC). Inyo Co., Mojave
Desert, Lovejoy Spring, 10 May 1944, A. L. Me-
lander (2(5; USNM). Kern Co., Red Rock Can-
yon, 30 Apr 1950 (16, 19; USNM). Los Angeles
Co., Los Angeles, 15 Aug 1915 (1?; ANSP); Tu-
munga Canyon, 3 May 1950, A. Wheeler (1$;
USNM). Napa Co., Moskowite Corner, 5.5 km
NW Capeel Creek, P. H. Arnaud, Jr. (1$; CAS).
Riverside Co., Cathedral Canyon, 3 Apr 1945, A.
L. Melander (1$; USNM); Palm Springs, Palm
Canyon, 26 Nov 1944, A. L. Melander (1(5;
USNM). San Bernardino Co., Big Bear Lake, 26
Jul-14 Sep, 1932-1934, A. J. Basinger, J. D.
Beamer (1(5, 1$; CAS, USNM); Helendale, 18
May 1955, W. R. Richards (11(5, 15$; CNC);
Victorville, 20 May 1955, W. R. Richards (Ic5;
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 90.?Distribution map oiSetacera durani (filled circles)
and S.jamesi (diamonds).
CNC); Wrightwood, SW Victorville, 30 Apr
1950, A. H. Sturtevant (66; USNM). San Diego
Co., Apr 1915, M. C. VanDuzee (1$; CAS);
Julian, 5 Mar 1945, A. L. Melander (16*; USNM);
San Luis Rey Campground, 28 Jun 1968, P. H.
Arnaud, Jr. (1<3; CAS). San Luis Obispo Co.,
Harmony, 25 Sep 1938, M. Cazier (16, 1$;
AMNH). Stanislaus Co., Del Puerto Road, Frank
Raines Park, 1 Nov 1969, T. W. Davis (76, 27?;
CAS). Oregon: Sherman Co., Moro, 10 Aug 1950,
O. Sutton (1$; USNM).
DISTRIBUTION (Figure 90).?Western North
America between 108? and 122? west longitude
and 30? and 46? north latitude. The known
distribution of the species forms a backwards J,
beginning in north central Oregon, extending
southward through California and eastward
across southern Arizona.
NATURAL HISTORY.?Practically nothing is
known about the ecology or immatures of this
species. Foote (1982) collected adults from an
algal mat that had formed in a small sewage-
polluted stream near Patagonia, Arizona.
REMARKS.?This species and S. pacifica are sister
species, as evidenced by the joint possession of a
median, triangular projection between the sur-
styli. Specimens of S. durani are separable exter-
nally from those of S. pacifica by the evenly
rounded supraspiracular convexity and by the
distinct tuft of setae at the apex of the midtibia.
12. Setacera jamesi, new species
FIGURES 90-98
DESCRIPTION.?Large shore flies, length 5.06 to
5.67 mm; generally brown to olivaceous brown or
gray, becoming grayer and duller ventrally.
FIGURE 91.?Setacera jamesi, head, lateral aspect.
NUMBER 350 41
Head (Figure 91): Head width-to-height ratio
averaging 1 : 0.67; frons width-to-length ratio
averaging 1 : 0.47; mesofrons and fronto-orbits
deeply blue to slightly greenish blue; dorsum of
interfoveal carina mostly greenish to slightly
bluish green; arista with longest dorsally branch-
ing rays short, at most slightly longer than aristal
width at base; dorsal surface of face broadly
explanate (best seen in dorsal view), dorsum of
interfoveal carina and antennal foveae almost on
same horizontal plane; dorsal surface of face
shorter than anterior surface; facial angle at
about 90?; angle formed by anterior surface and
oral margin obtuse in lateral view, anterior sur-
face receding to oral margin; eye height-to-width
ratio averaging 1 : 1.14; eye-to-cheek ratio aver-
aging 1 : 0.57.
Thorax: Supraspiracular convexity evenly
convex; forecoxa of male with longer, more nu-
merous setae anteromedially and posteriorly, to
a lesser extent on midcoxa; midtibia of male with
dense patch of slightly longer setae posteroapi-
cally; costal vein ratio averaging 1 : 0.26; Mi+2
vein ratio averaging 1 : 0.88.
Abdomen: Male 5th tergal ratio averaging
1 : 0.89; 5th tergum of male lacking anteroventral
projection. Male terminalia (Figures 92-94): sur-
styli in posterior view (Figure 94) subtriangular,
broadly fused basally lateral margins narrowing
to ventral apex at right angles to each other,
lacking distinct lateral projections, bluntly
rounded and curved anteriorly in lateral view;
gonite with ventrolateral prong, curved medially,
acutely pointed, 2 long, posterior setae (longer
than gonal prong), broadly attached to aedeagus
dorsally; aedeagus folded back on itself, broadly
rounded apically; hypandrium pocket shaped,
lateral margin overlaid, with 2 medial, cruciate,
well sclerotized prongs. Female terminalia (Fig-
ures 95-98): 7th tergum complete; 8th sternite
subtriangular, anterior margin rounded, posterior
margin shallowly concave, posteroventral corner
slightly projecting; only slightly higher than wide;
8th sternites about 4 times longer than wide; 9th
sternites almost twice as long as high; cercus
higher than long, with subquadrate posterodorsal
projection; female ventral receptacle (Figure 96)
92
FIGURES 92-98.?Setacera jamesi: 92, internal male genitalia, lateral aspect; 93, male terminalia,
lateral aspect; 94, surstyli, posterior aspect; 95, cerci of female terminalia, lateral aspect; 96,
female ventral receptacle, lateral aspect; 97, female 8th tergite and 9th sternite, lateral aspect;
98, female 8th tergite, sternites, and 9th sternite, ventral aspect.
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
with lateral margin of operculum slightly flared
ventrally, cervix comparatively shorter than other
species, lacking distinct indentation at juncture
of cervix with corpus.
TYPE MATERIAL.?The male holotype is la-
beled: "Orick [Humboldt Co.] CAL[ifornia] 18
Sep. [19]34 ALMelander/HOLOTYPE Setacera
jamesi Mathis [handwritten, red]." Allotype fe-
male and one male paratype are labeled with the
same label data as the holotype. Other paratypes
are as follows. Oregon: Lincoln Co., Boiler Bay, 9
Mar 1930, J. Wilcox (1?; USNM). Washington:
Pacific Co., Ocean Park, 18 Aug 1950, M. T.
James (36\ 2?; USNM, WSU). The holotype
specimen is in excellent condition (directly
pinned) and is in the National Museum of Nat-
ural History, Smithsonian Institution, USNM
76885.
DISTRIBUTION (Figure 90).?Coast of western
North America between 41? and 48? north lati-
tude.
NATURAL HISTORY.?The immatures and ecol-
ogy of the species are presently unknown. The
apparent association of this species with coastal
aquatic systems may indicate a tolerance toward
salty environs and, perhaps, a preference for it.
ETYMOLOGY.?The specific epithet jamesi hon-
ors Dr. M. T. James, Washington State Univer-
sity, for his continuing contribution to dipterol-
ogy-
REMARKS.?The sister group of & jamesi com-
prises S. needhami and S. trichoscelis, as indicated
earlier (see "Discussion," page 37). Externally,
specimens of S. jamesi are separable from either of
its closely related congeners by its larger size,
shorter aristal rays, and more normally shaped
female cerci. The male terminalia are also dis-
tinctive.
13. Setacera needhami Johannsen
FIGURES 99-108
Setacera needhami Johannsen, 1935:53.?Cresson, 1935:347
(description of adult].?Sturtevant and Wheeler,
1954:202, 203 [key and locality data].?Wirth and Stone,
1956:472 |key].?Wirth, 1965:755 [nearctic catalog].?
Cole, 1969:402 [locality data].
DESCRIPTION.?Moderately small to moder-
ately large shore flies, length 2.98 to 4.54 mm;
dorsum generally dark, brown, olivaceous green
to grayish green, becoming more subdued, lighter,
grayer ventrally.
Head (Figure 99): Head width-to-height ratio
99
100
FIGURES 99-104.?Selacera needhami: 99, head, lateral aspect; 100, male terminalia, posterior
aspect; 101, same, lateral aspect; 102, abdomen of female, lateral aspect; 103, female terminalia,
lateral aspect; 104, female ventral receptacle, lateral aspect.
NUMBER 350 43
FIGURES 105-107.?Setacera needhami: 105, left midfemur and midtibia, posterior aspect; 106, left
midfemur, posterior aspect; 107, apex of left midtibia, posterior aspect.
averaging 1 : 0.69; mesofrons and fronto-orbits
deeply dark blue to greenish blue, dorsum of
interfoveal carina mostly greenish; frons width-
to-length ratio averaging 1 : 0.59; arista with
longest branching rays nearly twice greatest aris-
tal width; dorsal surface of face with antennal
fovea inclined at distinctly lower level than dor-
sum of interfoveal carina; dorsal surface of face
shorter than anterior surface in profile; facial
angle moderately angulate, about 110?; angle
formed by anterior surface and oral margin ob-
tuse, anterior surface receding. Eye height-to-
width ratio averaging 1 : 1.02; eye-to-cheek ratio
averaging 1 : 0.37.
Thorax: Supraspiracular convexity evenly
convex; fore- and midcoxae of male normal; mid-
femur of male (Figures 105, 106) with 4-5 long,
well-developed setae ventrally toward base; mid-
tibia of male (Figures 105, 107) with large, dense
patch of long setae anteroventrally to ventrally;
hind tibia of male with small patch of long setae
anteroventrally; costal vein ratio averaging
1 : 0.28; M1+2 vein ratio averaging 1 : 0.84.
Abdomen: Male 5th tergal ratio averaging
1 : 0.62; 5th tergum of male lacking anteroventral
process. Male terminalia (Figures 100, 101): sur-
styli broadly based in posterior view, lateral mar-
gins rounded to just before apex, thereafter more
or less parallel sided, short, apex wide and shal-
lowly emarginate, lateral projections very shal-
low; gonite with slender dorsal and ventral
prongs, both curved medially, with 2 long poste-
riorly directed setae inserted at about midheight
in lateral view; aedeagus folded back on itself;
hypandrium very poorly sclerotized medially, lat-
erally and posteriorly well sclerotized, with 2 long,
well-developed anteromedial prongs, tapered to
acutely pointed apex. Female terminalia (Figures
102-104): 7th tergum complete; 8th tergites more
or less subquadrate in lateral view, dorsum shal-
lowly sinuate, posterior margin concave, poster-
oventral corner more projected, nearly as high as
wide; 8th sternites elongate, 5-6 times longer
than width; cercus wider than high in lateral
view, posterior margin shallowly rounded; female
ventral receptacle (Figure 104) with dorsum of
operculum bearing shallow, median bump, lat-
eral margins rounded, otherwise as in species
group description.
TYPE MATERIAL.?The male lectotype, herein
designated, is labeled: "Laguna Canyon Orange
Co. California]. [handwritten]/Shore of Pond J.
G. Needham Sept. 1922 [handwritten]/TYPE
No. 6574 Setacera NEEDHAMI 6 E T Cresson,
Jr. [red]/LECTOTYPE Setacera needhami Jo-
hannsen by W. N. Mathis [handwritten, red]."
The lectotype is in the Academy of Natural Sci-
ences of Philadelphia, Pennsylvania. The speci-
men is double mounted (glued to a paper point)
and is in good condition, although the right front
leg and the apical tarsomeres of the middle left
leg are missing.
OTHER SPECIMENS EXAMINED.?UNITED STATES.
Arizona: Yuma Co., Horsetank, 2 Dec 1962, E. I.
Schlinger (1$; UCR). California: Alameda Co.,
Berkeley, 4 Oct 1947, W. W. Wirth (26", 29;
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
USNM); Jewell Lake, 4 Oct 1947, W. W. Wirth
(26*, 59; USNM). Colusa Co., Maxwell, 12 Aug
1954, A. A. Grigarick (16\ 19; USNM). Contra
Costa Co., Jewell Lake, 4 Oct 1947, W. W. Wirth
(36*, 19; USNM). Humboldt Co., Trinidad, 18
Sep 1934, A. L. Melander (Id; USNM). Inyo Co.,
Independence, 13 mi NE, 10 Mar 1964, J. D.
Birchim (16*; CAS). Kern Co., Poso Creek, 18 Jun
1947, B. Brookman (1$; USNM). Riverside Co.,
Temecula, 25 May 1965, T. W. Fisher (1$; UCR).
Orange Co., Laguna Beach, J. G. Needham (2
larvae, 1 puparium; CU); Laguna Canyon, J. G.
Needham (39; ANSP). Sacramento Co., Sacra-
mento, 8 Jun-24 Sep, 1920-1933, H. H. Keifer,
C. M. Packard (6d\ 99; USNM); Citrus Heights,
11-15 Jun 1967, Keuter (29; CAS). San Bernar-
dino Co., Wright wood, 30 Apr 1950 (56*, 49;
USNM). San Diego Co., San Diego, 12-13 May
1916, H. G. Dyar (26; ANSP, USNM). Shasta
Co., Redding, 7 Nov 1970, T. R. Haig (19;
USNM). Tehama Co., Corning, 1/2 mi S, 23 Sep
1973, W. N. Mathis (16\ 19; USNM). Yolo Co.,
Davis, 30 Jan-21 Oct, 1932-1966, J. S. Buckett,
A. A. Grigarick, M. E. Irwin, W. H. Lange, H. R.
Moffitt, E. I. Schlinger (46, 79; USNM). Nevada:
Washoe Co., Pyramid Lake, 24 Jun 1927, E. P.
Van Duzee (16*; CAS). Oregon: Benton Co., Cor-
vallis, 29 Mar-20 Jun, 1960-1972, J. Capizzi, W.
N. Mathis (26*; USNM); Corvallis, 12 mi S, 12
Sept 1972, W. N. Mathis (16*, 29; USNM); Oak
Creek Lab, 9 Sep 1972, W. N. Mathis (16*;
WNM); Willamette River, 12 Oct 1971, W. N.
Mathis (26*, 29; USNM). Linn Co., Peoria, 2 mi
S, 24 Oct 1971, W. N. Mathis (16*, 19; KSU,
USNM). Polk Co., Corvallis, 14 mi N, 24 May
1972, W. N. Mathis (29; USNM). Umatilla Co.,
Cold Springs, 11 Apr 1954, J. J. Davis, M. T.
James (19; USNM). Utah: Davis Co., Farming-
ton, 13 Feb 1934, W. L. Thomas (16*; ANSP).
Weber Co., Hooper, 20 Oct 1933, T. A. Rowe
(19; ANSP). Washington: Franklin Co., Kahlotus
Lake, 11 May 1951, J. J. Davis (19, 1 puparium;
USNM). Grant Co., Soda Lake near O'Sullivan
Dam, Columbia Wildlife Refuge, 19 Mar 1977,
R. S. Zack (246*, 119; USNM, WSU). Pacific Co.,
Ocean Park, 18 Aug 1950, M. T. James (16*;
WSU). Whitman Co., Almota, 23 Mar 1977, R.
S. Zack (16*; USNM).
ZOOGEOGRAPHIC DISTRIBUTION (Figure 108).?
Western North America between 111? and 125?
west longitude and 32? and 48? north latitude.
NATURAL HISTORY.?I have collected adults of
this species at numerous localities in association
with lentic aquatic systems. Presumably the im-
mature stages also occur in these habitats.
REMARKS.?The authorship of S. needhami is
properly credited to O. A. Johannsen (1935)
(Cresson, 1935:347; Wirth, 1965:755), who pub-
lished a brief diagnosis of the larva and puparium
of this species. Johannsen's diagnosis, however,
was published without the intention of it being a
new species description, and Cresson's (1935) be-
lated description of the adult was not published
until nine months later. The publication of Jo-
hannsen's description of the immature stages be-
fore Cresson's treatment of the adult is unfortu-
nate and has resulted in the ambiguous usage of
FIGURE 108.?Distribution map of Setacera needhami.
NUMBER 350 45
this name today. Until now, the status of Johan-
nsen 's species was unsatisfactory, being based on
the immature stages that are still not recogniz-
able. Indeed, larvae and puparia of Ephydra and
Setacera are still not always separable at the ge-
neric level.
If the range of S. needhami (based on adults)
were isolated, there would be no problem. This,
however, is not the case. In southern California,
the range of a second species, S. durani, is broadly
sympatric with that of S. needhami, and the possi-
bility exists of finding immatures of both species
at the same locality, hence the potential confu-
sion. To promote stability and to avoid the pos-
sibility of further confusion, I have herein desig-
nated an adult male specimen collected by Need-
ham as the lectotype. The adult male of S. need-
hami is easily recognized, making application of
the specific name unambiguous.
According to the current rules of zoological
nomenclature (ICZN, 1964), a lectotype is to be
designated from one of the syntypes, the latter
being defined as "all the specimens of the type-
series . . . " when no holotype has been designated
(art. 73c, ICZN). The rules also define a "type-
series" as "all the specimens on which its author
bases the species, except any that he refers to as
variants, or doubtfully associates with the nomi-
nal species, or expressly excludes from it" (art.
72B, ICZN).
My reasoning for considering adult specimens
as part of the type series is as follows. Because
Johannsen's diagnosis of S. needhami was not in-
tended as a new species description, the content
and format of his description were incomplete.
Johannsen did not designate a holotype, nor did
he clearly state which specimens were included in
the type series; however, he specifically stated:
"Some specimens of adults, in not very good
condition, were determined as Setacera needhami by
E. T. Cresson (Jr.). They were reared from larvae
and puparia collected by Dr. J. G. Needham at
Laguna Beach, California [the same larvae and
puparia Johannsen then described]." The adult
specimens are the imagos of the same individuals
that Johannsen described as larvae and puparia
(Johannsen had the exuviae) and are part of the
type series by direct association. In addition, Jo-
hannsen made no statement that would exclude
the adult specimens; to the contrary, he expressly
associated them as "reared from larvae and pu-
paria . . . ." Moreover, as stated by Johannsen
(cited previously), adult specimens, not the im-
matures, were sent to Cresson for identification.
As the describer, Johannsen clearly based his
concept of S. needhami on these adults, which by
definition became part of the type series from
which the lectotype was selected.
This species and S. trichoscelis are sister species
but are separable by rather subtle characters of
the male terminalia and by allopatry. As the
characters of the male terminalia are consistent
within the sampling I examined, I have recog-
nized the appropriate populations as distinct spe-
cies. No intermediates are known.
14. Setacera pacifica (Cresson)
FIGURES 109-125
Ephydra pacifica Cresson, 1925:167.
Setacera pacifica.?Cresson, 1930:116.?Sturtevant and
Wheeler, 1954:202, 203 [key and locality data].?Wirth
and Stone, 1956:472 [key].?Wirth, 1965:755 [nearctic
catalog].?Cole, 1969:402 [locality data].
DESCRIPTION.?Moderately large to large shore
flies, length 4.09 to 5.12 mm; dorsum generally
green to olivaceous brown, becoming grayish
green to green ventrally, duller.
Head (Figures 109, 116-118): Head width-to-
height ratio averaging 1 : 0.66; frons width-to-
length ratio averaging 1 : 0.49; mesofrons, fronto-
orbits, and at least posterior portion of dorsum of
interfoveal carina deeply bluish to greenish blue,
anterior portion of interfoveal carina sometimes
more greenish; arista with longest dorsally
branching rays nearly twice aristal width at base;
dorsal surface of face moderately explanate, an-
tennal foveae only slightly lower and inclined
than dorsum of interfoveal carina, nearly horizon-
tal; dorsal surface of face shorter than anterior
surface, facial angle about 95?; angle formed by
46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
114 115
FIGURES 109-115.?Setacera pacifica: 109, head, lateral aspect; 110, thorax, dorsal aspect; 111,
surstyli, posterior aspect; 112, internal male genitalia, lateral aspect; 113, male terminalia,
lateral aspect; 114, female terminalia, lateral aspect; 115, female ventral receptacle, lateral
aspect.
anterior surface and oral margin; eye height-to-
width ratio averaging, 1 : 0.98; eye-to-cheek ratio
averaging 1 : 0.44.
Thorax (Figures 110, 119-124): Supraspira-
cular convexity with papilla-like, lateral projec-
tion toward anterodorsal portion (Figures 123,
124); fore- and midcoxae of male (Figure 121)
with dense patches of long setae, especially ven-
trally and posteroventrally; front basitarsus of
male with long, fine, pale setae ventrally; mid-
and hind tibiae of male with slightly more dense
patch of setae apicoventrally; costal vein ratio
averaging 1 : 0.28; M1+2 vein ratio averaging
1 : 0.83.
Abdomen: Male 5th tergal ratio averaging
1 : 0.81; 5th tergum of male lacking an antero-
ventral projection. Male terminalia (Figures
111-113): epandrium with shallow, median fur-
row; surstyli, in posterior view, with base rounded
laterally, projecting processes narrow, tapered to
acutely pointed apices, each process curved an-
teriorly in lateral view, median projection trian-
gular, wider at base than more lateral processes,
median projection, in lateral view, situated ante-
rior of processes and slightly curved posteriorly;
gonite with 3 posteriorly projecting prongs, dor-
somedial pair more heavily sclerotized, apex
slightly curved dorsally, median and ventral pairs
along same lateral plane, median prong wider
and shorter, more or less straight, ventral prong
narrow and generally curved ventrally; aedeagus
folded back on itself, rounded apically; hypan-
drium more or less evenly sclerotized, prongs and
extreme lateral margins more heavily sclerotized,
median plate area very shallow, with 2 prongs,
posteromedian pair oriented medially, sub-
apically more abruptly tapered, curved, clawlike,
anterolateral prong evenly tapered, mostly
straight. Female terminalia (Figures 114, 115):
7th tergum incomplete, formed by 2 lateral ter-
gites; 8th tergites higher than wide in lateral view,
dorsum pointed, venter rounded, anterior margin
more or less straight, posterior margin straight to
very shallowly concave; 8th sternites over 6 times
longer than wide; cercus as wide or wider than
high, roughly subquadrate, posterior margin
broadly rounded to subtruncate; female ventral
receptacle (Figure 115) with operculum mostly
rounded dorsally and laterally, otherwise as in
species group description.
TYPE MATERIAL.?The male holotype is la-
beled: "Vancouver 5 IV[Apr]/[19]02 B[ritish]
Colombia] R J. Crew/<3/TYPE No. 6344 Ephy-
dra PACIFICA ( 5 E T Cresson, Jr. [red]." The
holotype is in the Academy of Natural Sciences
of Philadelphia, ANSP 6344. The specimen is
pinned directly and is in good condition, although
NUMBER 350 47
FIGURES 116-124.?Setacera pacijica: 116, right vertex and vertico-orbits of head, dorsal aspect;
117, right vertico-orbit showing velvety tomentosity, dorsal aspect; 118, right antenna, lateral
aspect; 119, notopleuron and surrounding area of thorax, lateral aspect; 120, propleuron, lateral
aspect; 121, fore- and midcoxae, anterior aspect; 122, prosternum anterior aspect; 123, posterior
portion of thorax, lateral aspect; 124, supraspiracular convexity with papilla, lateral aspect.
all tarsomeres except for the basitarsus of the
right midleg are missing.
OTHER SPECIMENS EXAMINED.?CANADA. British
Columbia: Oliver, 12 Aug 1953, 1000 ft, D. F.
Hardwick (1<J, 1$; CNC); Oliver, Mclntyre
Creek, 6 Jun 1959, R. E. Leech (1<J; CNC);
Oliver, White Lake, 28 May 1959, L. A. Kelton
(26; CNC); Vancouver Island, Cowichan Island,
12 Jul 1924, A. L. Melander (16, 1$; USNM);
Vernon, 31 Aug 1937, H. Leech (2$; CNC).
Manitoba: Aweme, 4 Oct 1923, N. Criddle (19;
CNC); Whitewater Lake, 4 mi N Whitewater, 30
Jul 1958, R. L. Hurley (Id; CNC). UNITED STATES.
California: Lassen Co., Susanville, 41 mi NW, 17
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Jun 1974 (1$; USNM). San Bernardino Co., Bar-
ton Flat, S Fork Camp, 31 Aug-3 Sep 1944, A. L.
Melander (26*, 29; USNM), Big Bear Lake, 24
May-26 Jul, 1932-1966, R. H. Beamer, A. L.
Melander, R. E. Orth (36*, 39; KU, UCR,
USNM), Upper Santa Ana River, 7-11 Oct 1946,
J. L. Sperry (46, 19; USNM). Tehama Co., Battle
Creek Campground, S fork, 12 Aug 1977, P. H.
Arnaud, Jr. (16*; CAS). Colorado: Jefferson Co.,
Foxton, 18 Oct 1881 (1$; ANSP). Larimer Co.,
Fort Collins, 23 May-21 Aug, 1926-1941, R. H.
Beamer, M. T.James (16\ 1$; KU, USNM). Iowa:
Story Co., Ames, Izaak Walton League Lake, 23
Apr 1962, D. L. Deonier (16*; DLD). Michigan:
Kalamazoo Co., Gull Lake Biological Station, 22
Jul 1963, R. L. Fischer (16*; USNM). Montana:
Lake Co., Ronan, 3 mi S, 30 Jun-17 Aug 1971,
B. A. Foote (66*, 14$; KSU, USNM), Ronan, 3.2
mi S, 8 Aug 1972, W. N. Mathis (16*; USNM),
Ronan, 5 mi S, 1-20 Jul 1973, B. A. Foote (76*,
5$; KSU). Sheridan Co., Medicine Lake, 9 Jun
1969, W. W. Wirth (1$; USNM). Nebraska: Cherry
Co., Big Alkali Lake, 2 Jun 1969, W. W. Wirth
(2?; USNM). New Mexico: Catron Co., Glenwood,
1 Jun 1972, W. W. Wirth (1$; USNM). Dona
Ana Co., Las Cruces, 14 Jun 1917, J. M. Aldrich
(16*; USNM). Hildalgo Co., Cienega Lake, 15 mi
N Rodeo, 30 Jul 1965, H. B. Leech (16*; USNM).
North Dakota: Burleigh Co., Long Lake, 4 Jun
1969, W. W. Wirth (16*; USNM). Oregon: Harney
Co., Burns, 20 Jul 1962, K. Goeden (26*, 2$;
USNM), Crane Hot Springs, 8 Mar-24 May
1975, W. N. Mathis (16*, 1$; USNM), Harney
Lake, south shore, 16 Jun 1972, W. N. Mathis
(22; USNM). Klamath Co., Klamath Falls, 31
May 1964, J. Schuh (26*; WSU). Lake Co., Ana
Reservoir, 24 Sep 1971, W. N. Mathis (16*;
USNM), Summer Lake, 4.8 mi N, 11 Jul 1974,
W. N. Mathis (36*, 39; USNM). South Dakota:
Mellette Co., Little White River, 4 Jun 1969, W.
W. Wirth (16*; USNM). Utah: Piute Co., Kings-
ton, 3 Jul 1938, G. F. Knowlton (16*, 1$; ANSP).
Utah Co., Goshen Pond, 1 Feb-27 Apr,
1968-1969, W. N. Mathis (176*, 79; USNM).
Washington: Benton Co., Prosser, Irrigation Exper-
iment Station, 17 Aug 1957, W. Cone (19; WSU).
Grant Co., Soda Lake near O'Sullivan Dam,
Columbia Wildlife Refuge, 19 Mar 1977, R. Zack
(26*, 29; USNM, WSU); O'Sullivan Dam, 4-5
May 1956, H. G. Davis (16*, 19; WSU). San Juan
Co., Olga, 26 Jul 1909 (19; ANSP). Whitman
Co., Almota, 3 Mar 1977, W. J. Turner (19;
WSU), Pullman, 25 Mar-11 Sep, 1971-1976, W.
J. Turner, R. S. Zack (16*, 19; WSU). Wyoming:
Bighorn Co., Shell Creek, S Shell Canyon, 24 Jun
1964, H. B. Leech (19; CAS).
DISTRIBUTION (Figure 125).?North America,
primarily in the West, between 85? and 124? west
longitude and 31? to 51? north latitude. This
species is the most widespread of the pacifica
group.
NATURAL HISTORY.?Like most species of this
species group, specimens of S. pacifica occur mostly
in association with lentic aquatic systems but
with one major difference?there appears to be a
preference toward alkaline water (Foote, 1982). I
have also collected several adults from algal mats
on the surface of the pooled effluent of thermal
springs in Oregon (Crane Hot Spring) and Utah
(Goshen Ponds).
Foote's (1982) study of S. atrovirens has extensive
notes on S. pacifica. Like the former species, a
generation can be completed in 25 to 30 days
during the warm season, with divisions among
stadia as follows:
Egg incubation
Larval period (3 instars)
Pupal period
Adult preoviposition period
2.3 days
10 days
7 days
6-9 days
Foote studied this species at highly alkaline
ponds near Ninepipes Wildlife Refuge, south of
Ronan, Montana. The ponds had large quantities
of carbonate and bicarbonate ions, and the pH
averaged above nine.
Both eggs and larvae were abundant in floating
algal mats comprised mostly of Anabaena and
Oscillatoria. Larvae were also found in a mat of
primarily RhyzocIonium, a filamentous green alga,
the diatom genera Navicula and Synedra, and the
desmid genera Closterium, Cosmarium, and Stauras-
trum. Cells of Cosmarium, among others, were found
in the gut of 2nd- and 3rd-instar larvae.
NUMBER 350 49
FIGURE 125.?Distribution map of Setacera pacifica.
As in S. atrovirens, the puparia of S. pacifica are
attached by their hindmost prolegs to pieces of
aquatic macrophytes and occasionally to fila-
ments of algae. Also like S. atrovirens, the puparial
attachment was frequently below the water's sur-
face, as much as 10 cm.
REMARKS.?Together with S. durani, this species
is the sister lineage of the group comprising S.
jamesi, S. needhami, and S. trichoscelis. Specimens of
S. pacifica are separable from those of closely
related congeners by the papilla-like projection of
the supraspiracular convexity, the setation of the
arista and legs, and by the conformation and
position of the male terminalia.
15. Setacera pilicornis (Coquillett)
FIGURES 126-133
Ephydra pilicornis Coquillett, 1902:184.
Setacera pilicornis.?Sturtevant and Wheeler, 1954:202, 204
[key, locality data, and discussion of synonymy].?Wirth,
1965:755 [nearctic catalog]; 1968:24 [neotropical catalog].
Setacera knabi Cresson, 1935:346 [synonymy by Sturtevant
and Wheeler, 1954:204].
DESCRIPTION.?Medium-sized to moderately
large shore flies, length 3.34 to 4.05 mm; dorsum
mostly grayish green to blue with some brownish
coloration, becoming more subdued, grayer ven-
trally.
Head (Figure 126): Head width-to-height ra-
tio averaging 1 : 0.57; frons width-to-length ratio
averaging 1 : 0.54; mesofrons and fronto-orbits
deeply greenish blue to mostly green, dorsum of
interfoveal carina usually more lightly colored,
mostly greenish; arista with longest dorsally
branching rays nearly twice aristal width at base;
dorsal surface of face with antennal foveae dis-
tinctly inclined from dorsum of interfoveal carina,
not subexplanate; dorsal surface in profile shorter
than anterior surface; facial angle about 115?;
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
126
FIGURES 126, 127.?Selacera pilicornis: 126, head, lateral aspect; 127, thorax, dorsal aspect.
angle formed by anterior surface of face and oral
margin obtuse; anterior surface of face with dorsal
ridge usually golden brown, thereafter becoming
silvery white ventrally. Eye height-to-width ratio
averaging 1 : 1.1; eye-to-cheek ratio averaging
1 : 0.34.
Thorax (Figure 127): Posthumeral bristle
weak, at most slightly stronger than largest hu-
meral bristle; supraspiracular convexity evenly
convex; coxae and legs of male lacking conspicous
tufts or dense patches of longer setae; costal vein
ratio averaging 1 : 0.23; M1+2 vein ratio averaging
1 : 0.81.
Abdomen: Male 5th tergal ratio averaging
1 : 0.69; 5th tergum of male lacking anteroventral
projection. Male terminalia (Figures 128-130):
epandrium broadly rounded dorsally; surstyli in
posterior view with lateral margin tapering grad-
ually, straight, to apex of projecting arms, each
arm curved laterally apically, no median process
toward venter, with 1 long posterior seta; aedea-
gus usually folded back on itself, rounded api-
cally; hypandrium well sclerotized laterally, weak
medially, with 2 posterior prongs, lateral pair
acutely pointed, wider at base, tapered more
steeply, median prong more slender, curved pos-
teriorly, more parallel sided. Female terminalia
(Figures 131, 132): 7th tergum complete; 8th
narrow, tapered ventrally, widest subdorsally, an-
terior margin shallowly concave, posterior margin
rounded; 8th sternites elongate, over 6 times
longer than wide; cercus in lateral view wider
than high, posterior margin broadly rounded,
with distinct, separate sclerite between cercus and
8th tergite; female ventral receptacle (Figure 131)
with operculum rounded dorsally and laterally,
otherwise as in species group description.
TYPE MATERIAL.?The male holotype of senior
synonym is labeled: "BISC [anye]. BAY,
FL[orida]A.[Mrs. Annie T.] Slosson Collector/
Type No 6645 U.S.N.M. [red]." The specimen is
double mounted (glued to a paper point) and is
in good condition. The male holotype of the
junior synonym is labeled: "Miami, 23. II. 12 [23
Feb 1912] Fl[orid]a/Fredk Knab Collector/67
TYPE No. Setacera KNAB 6 E. T. Cresson, Jr,
[pink]/Type No 51098 USNM [red]." This spec-
imen is double mounted (glued to a paper point)
NUMBER 350 51
130
132
FIGURES 128-132.?Setacera pilicomis: 128, male terminalia, lateral aspect; 129, internal male
genitalia, lateral aspect; 130, surstyli, posterior aspect; 131, female ventral receptacle, lateral
aspect; 132, female terminalia, lateral aspect.
and is in good condition. Both holotypes are in
the National Museum of Natural History, Smith-
sonian Institution.
OTHER SPECIMENS EXAMINED.?MEXICO. Ta-
basco: Villahermosa, 6 Aug 1964, P. J. Spangler
(3<5, 59; USNM). UNITED STATES. Florida: Alachua
Co., Austin Lary Forest, 5 Jul 1969, L. A. Hetrach
(16; USNM); Gainesville, 29 Apr-11 Dec,
1958.-1972, H. R. Dodge, R. P. Esser, F. W.
Mead, J. W. Perry (7(5, 10$; USNM). Broward
Co., Fort Lauderdale, 19 Dec 1960, G. F. Spencer
(16, 49; USNM). Charlotte Co., Punta Gorda, 12
Apr 1952, J. R. Vockeroth (1?; CNC). DadeCo.,
Homestead, Subtropical Experimental Station, 2
May 1967, B. V. Peterson (1$; CNC); Miami, 23
Feb 1912, F. Knab (39; ANSP). Highland Co.,
Archbold Biological Station, 14 Apr-1 Jun,
1960-1967, B. V. Peterson, S. W. Frost, J. G. and
B. L. Rosen (56, 49; AMNH, CNC, USNM).
Hillsborough Co., Plant City, 15 Aug 1930, J. O.
Nottingham (26\ 39; KU, USNM); Tampa, 21
Apr 1961, L. A. Kelton (16*; CNC). Lee Co., Fort
Myers, 26 Dec 1953, B. H. Judd (19; USNM).
Polk Co., Fort Meade, 13 Aug 1930, P. W. Oman
(\6, 19; KU, USNM). Taylor Co., Williams
Landing, 21-30 Jun 1967, R. Smith (19; USNM).
Georgia: Liberty Co., St. Catherines Island, 21-23
Apr 1978, R. W. Matthews (356, 679; USNM).
Louisiana: Lafourche Parish, Cut Off, 9-11 Jul
1975, V. A. Brou (16, 29; USNM). Iberville Par-
ish, Sunshine, 8-15 Aug 1972, V. A. Brou (\6,
59; USNM). Orleans Parish, New Orleans, 28
Jan-28 Nov, 1942-1974, V. A. Brou, D. G. Den-
ning, E. H. Metzler (36, 89; DLD, USNM). St.
Charles Parish, Norco, Bonnet Carre Spillway, 20
Feb 1972, E. H. Metzler (36; DLD). St. Helena
Parish, Fluker, 10 mi SW, 13 Apr 1972, E. H.
Metzler (26, 39; DLD). St. John the Baptist
Parish, Edgard, 2 Mar-24 Jun 1972, V. A. Brou
(26, 19; USNM). Mississippi: Jackson Co., Van-
cleave Road, 15 Jun 1962, D. L. Deonier (1<5;
DLD). South Carolina: Beaufort Co., Hilton Head
Island, 12 Jul 1965, H. F. Howden (19; CNC).
DISTRIBUTION (Figure 133).?Southeastern
North America, especially the Gulf Coast states
(South Carolina to Louisiana), and southcentral
and Gulf coastal Mexico between 80? and 100?
west longitude and 18? and 33? north latitude.
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 133.?Distribution map of Setacera pilicornis.
NATURAL HISTORY.?Numerous specimens of
this species were collected as prey of sphecid
wasps on St. Catherines Island, Georgia. The
wasps were being studied behaviorally and eco-
logically by Dr. R. W. Matthews and students
(University of Georgia). Both sexes were selected
as prey and in about equal numbers.
REMARKS.?When Sturtevant and Wheeler
(1954) reviewed the nearctic species of Setacera,
they tentatively listed S. knabi Cresson as a junior
synonym of S. pilicornis. Their uncertainty
was due to the fact that the type specimen of S.
knabi, stated to be in the USNM, could not be
located for comparison. The type was subse-
quently located by Drs. Selwyn S. Roback and
Willis W. Wirth among specimens at the Acad-
emy of Natural Sciences of Philadelphia that
were not returned to the lending institution fol-
lowing the untimely death of E. T. Cresson, Jr. I
have examined the types of both species and can
confirm their conspecificity and consequently the
status of S. knabi as a junior synonym. Only one
species of Setacera is known to occur in Florida.
Although Coquillett's original description of
this species indicated only a single male specimen,
which automatically becomes the holotype, the
collection at the Academy of Natural Sciences of
Philadelphia contains four additional specimens
labeled as paratypes with Cresson's characteristic
blue labels. These specimens cannot be paratypes,
however, as Coquillett's description specifically
listed only one male specimen.
In the phylogenetic scheme of this species
group, S. pilicornis is the sister species of the re-
NUMBER 350 53
maining species in the group, being separable
from them by the lack of shaggy-appearing setae
on the forecoxae of males and by the unique
characters of the male and female terminalia.
16. Setacera trichoscelis
 y new species
FIGURES 134-138
DESCRIPTION.?Medium-sized to moderately
large shore flies, length 3.24 to 4.21 mm; mostly
subdued, pollinose, light brown to grayish green
except for dorsum which is subshiny to shiny.
Head (Figure 134): Head width-to-height ra-
tio averaging 1 : 0.70; frons width-to-length ratio
averaging 1 : 0.49; frons and fronto-orbital plate
distinctly shiny with metallic blue to greenish
blue reflections; frons with sparsely scattered,
pilose hairs; ocelli arranged in isosceles triangle,
distance between posterior pair shorter than be-
tween median ocellus and either posterior ocellus.
Antenna unicolorous, blackish brown, dull, to-
mentose with grayish vestiture; 3rd segment
slightly longer than combined length of 1st and
2nd; 2nd segment with some greenish color ven-
trally. Face mostly silvery white except dorsal
shelf; antennal fovea and interfoveal space with
metallic reflections similar in color to mesofrons;
area immediately surrounding base of antenna
with golden brown tomentosity; lower portion of
face receding; facial angle approximately 90?
from profile view; length of lower portion longer
than distance between base of antenna to facial
angle. Eye width-to-height ratio averaging
1 : 0.92, oval, oriented at oblique angle to plane
of epistoma. Gena wide, eye-to-cheek ratio aver-
aging 1 : 0.43; concolorous with face, becoming
duller posteriorly and with some grayish green to
olivaceous coloration.
Thorax (Figure 135): Mesonotum mostly
brown to greenish brown; anterior portion more
subdued, pollinose, grayish brown, becoming
darker and with more shiny, metallic green, pos-
teriorly; scutellum with mostly greenish to bron-
zish blue metallic luster shining through darker-
brown vestiture. Mesopleuron, pteropleuron, and
sternopleuron mostly concolorous, grayish golden
brown to olivaceous; forecoxa shiny gray; hypo-
pleuron mostly gray to lightly olivaceous in color.
Femora mostly gray but with some bluish or light
brown coloration, becoming darker apically; ti-
biae mostly concolorous with femora but with less
grayish coloration; tarsi yellowish orange but
with some blackish coloration; legs of males with
tufts of long hairs near apices of mid- and hind
tibiae; posteroventral surface of mid femur with
FIGURES 134, 135.?Setacera trichoscelis: 134, head, lateroblique aspect; 135, thorax, dorsal aspect.
54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
138
FIGURES 136-138.?Selacera Irichoscelis: 136, male terminalia, lateral aspect; 137, internal male
genitalia, lateral aspect; 138, surstyli, posterior aspect.
distinct row of bristles, which become larger to-
ward base. Costal vein index averaging 1 : 0.30;
M1+2 vein index averaging 1 : 0.83.
Abdomen: Dorsum mostly unicolorous, brown
with metallic bluish green to green luster; tergum
of 1st segment more pollinoise, grayish, other
terga becoming darker and more shinier poste-
riorly but not as dark as dorsum of scutellum. 5th
tergum of male longer than wide, nearly as long
as combined length of 3rd and 4th segments;
lateral margins tapering gradually to subtruncate
apex. Epandrium of male terminalia triangular,
becoming wider toward venter; surstyli attached
to ventral margin of epandrium, broadly fused to
form subrectangular plate with ventrolateral,
rounded processes; 5th sternum loosely attached
to posteroventral corners of epandrium, anterior
ends produced into gently curved, well-scle-
rotized, sickle-like processes; see figures of termi-
nalia (Figures 136-138) for further details.
TYPE MATERIAL.?The male holotype is la-
beled: "Yaguarcocha, 3 km N. Ibarra, 1950 m,
Imbabura, Ecuad[or], 8-9. VI. 65 [8-9 Jun
1965], L. Pena/Holotype Setacera trichoscelis
Mathis [handwritten; red]." Female allotype and
36 paratypes (106*, 26$) are labeled with the same
label data as the holotype. The holotype, allotype,
and most of the paratypes are in the Canadian
National Collection, Ottawa, type 15239. Two
pairs of male and female paratypes are in the
National Museum of Natural History, Smithson-
ian Institution.
ETYMOLOGY.?The species epithet trichoscelis is
of Greek derivation and is a combination of tricho
and scelis ("hair" and "leg" respectively). The
name refers to the tufts of long hairs on the mid-
and hind legs of males.
DISTRIBUTION.?This species is known only
from the type-locality.
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